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Vertical distribution and biomass of fine roots in three subalpine forest plant associations in southwestern… Nuszdorfer, Friedrich Carl 1982

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VERTICAL DISTRIBUTION AND BIOMASS OF FINE ROOTS IN THREE SUBALPINE FOREST PLANT ASSOCIATIONS IN SOUTHWESTERN BRITISH COLUMBIA by FRIEDRICH CARL NUSZDORFER B . S c , U n i v e r s i t y o f B r i t i s h Co lumbia , 1975 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Department o f Fo res t r y ) We accept t h i s t h e s i s as conforming to the requ i red s tandard THE UNIVERSITY OF BRITISH COLUMBIA October , 1932 © F r i e d r i c h Car l Nuszdor fe r In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y available for reference and study. I further agree that permission for extensive copying of t h i s thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. I t i s understood that copying or publication of t h i s thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of The University of B r i t i s h Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date /? Oj~J^ /TVL C-6 (3/81) i i ABSTRACT Roots smaller than 5 mm in diameter were studied in three t y p i c a l high elevation forested plant associations of coastal southwestern B r i t i s h Columbia. The objectives were to quantify their d i s t r i b u t i o n with depth, examining changes with plant association and time; to examine their relationships with foliage; and to seek correlations with s o i l properties. Coring randomly in three replicates of each plant association was followed by washing of s o i l and organic matter from roots with water in combination with manual sorting. Length and surface area were estimated for a l l samples. Standard a n a l y t i c a l methods were used for determining s o i l physical and chemical properties. S o i l temperature and moisture were measured during the growing season. The peak in mass of both overstory and underst'ory < 2 mm roots was within the upper 10 cm of the s o i l . Overstory 2-5 mm roots peaked between 5 and 30 cm. Understory 2-5 mm roots peaked within 10 cm of the surface of the forest f l o o r . The large v a r i a t i o n between rep l i c a t e s of the plant associations made i t impossible to determine i f changes in root mass with time of sampling were real or due to random v a r i a t i o n . Unsuberized roots of the overstory vegetation contributed very l i t t l e to biomass of £ 5 mm roots. Biomass of £ 5 mm roots of the overstory vegetation varied from 740 to 1320 g m"2, length from 3.8 to 6.5 km m"2, and surface area from 7.3 to 11.9 m2 m*2. The ridgetop association had the most overstory and understory roots and i t s s o i l (including organic horizons) was lowest in t o t a l nitrogen concentration and CEC. The midslope association had intermediate amounts of overstory roots, the lowest amount of understory roots, and i t s s o i l was intermediate in nitrogen concentration and CEC. The receiving association had the lowest amounts of overstory roots, intermediate amounts of understory roots, and i t s s o i l had the highest nitrogen concentration and CEC. The ridgetop ecosystems had the highest r a t i o of root-to-shoot and r a t i o of root-to-foliage plus twig, being approximately double the r a t i o for the receiving association. The midslope association was intermediate. Correlations of root mass with s o i l properties were generally poor. i v TABLE OF CONTENTS Abstract i i L i s t Of Tables v i i L i s t Of Figures x Acknowledgements x i i 1. Introduction 1 2. Literature Review 3 2.1 Depth D i s t r i b u t i o n of Fine Roots 3 2.2 Biomass of Fine Roots and Ratios to Shoots 11 2.3 Length and Surface Area of Fine Roots 22 2.4 S o i l Physical and Chemical Parameters 25 3. The Study Area 26 3.1 Location and Description 26 3.2 Climate and Microclimate 35 3.3 Geology and S o i l s 38 4. Methods 44 4.1 F i e l d 44 4.1.1 Roots 45 4.1.2 S o i l Water 46 4.1.3 S o i l Temperature 48 4.2 Laboratory 49 4.2.1 Root Washing and Separation 49 4.2.2 Root Length and Surface Area 54 4.2.3 Root Biomass 56 4.2.4 S o i l Physical and Chemical Parameters 57 5. Results And Discussion 59 V 5.1 Variation of Roots with Depth '.. 62 5.1.1 Overstory Roots 62 5.1.2 Understory Roots 75 5.1.3 Overstory Compared With Understory 80 5.2 Changes in V e r t i c a l D i s t r i b u t i o n and Root Biomass With Time 82 5.2.1 Overstory Root V e r t i c a l D i s t r i b u t i o n 82 5.2.2 Understory Root V e r t i c a l D i s t r i b u t i o n 84 5.2.3 Overstory Root Biomass 86 5.2.4 Understory Root Biomass 91 5.3 Variation in Root Biomass Between Plots 94 5.4 Variation in Root Biomass Between Plant Associations 96 5.5 Ratio of Overstory Root Biomass to Overstory Above-Ground Biomass 103 5.6 S o i l Physical and Chemical Properties 109 5.6.1 Results of Analysis 109 5.6.2 Correlations With Root Mass-Volume 116 5.6.2.1 Overstory Roots 116 5.6.2.2 Understory Roots 123 5.6.3 Results Summarized by Plant Association .... 126 6. Summary And Conclusions 132 Literature Cited 137 Appendix 1 147 Appendix 2 148 Appendix 3 149 Appendix 4 150 v i Appendix 5 152 Appendix 6 169 Appendix 7 170 v i i LIST OF TABLES Table Page 2.1 D i s t r i b u t i o n of roots < 6.4 mm diameter in forest ecosystems 4 2.2 Biomass and root-to-shoot ra t i o s for roots < 10 mm in diameter in forest ecosystems 12 2.3 Lengths of roots < 6 mm in diameter in forest ecosystems 23 3.1 Characteristics of the overstory component of the sample plots; modified af t e r Krumlik (1979) 31 3.2 Summary of c h a r a c t e r i s t i c s of the three plant associations 36 3.3 Summary of c h a r a c t e r i s t i c s of modal s o i l s based on two p i t s in each of three r e p l i c a t e s of the OPHO THPL and ABAM TSME associations, and two p i t s in two replicates and three p i t s in a t h i r d r e p l i c a t e of the VAME TSME Association 40 5.1 Relationships between length, surface area and mass of roots 60 5.2 P r o b a b i l i t i e s of F-values from ANOVA of l o g 1 0 root mass-volume 63 5.3 Varia t i o n in overstory, suberized < 2 mm root mass-volume with depth for the plant associations . 68 5.4 Variation in overstory 2-5 mm root mass-volume with depth for the plant associations 70 5.5 Effect of mineral contamination of overstory v i i i roots < 2 mm in diameter on mass-volume and biomass estimates for the three plant associations .. 71 5.6 Cumulative percentage of root biomass in the s o i l for the three overstory root categories in the three plant associations 74 5.7 Variation in understory < 2 mm root mass-volume with depth for the plant associations 76 5.8 Variation in understory 2-5 mm root mass-volume with depth for the plant associations 78 5.9 Cumulative percentage of root biomass in the s o i l for the two understory root sizes in the three plant associations 79 5.10 Changes in the percentage of root mass above 30 cm and depth of 90 % of the root mass for the root categories and plant associations 81 5.11 Results of Duncan's Multiple Range Test on transformed (log 1 0(x+1)) root mass-volume (kg n r 3 ) 95 5.12 Comparison between overstory and understory root biomass for the three plant associations 98 5.13 Comparison between overstory and understory root lengths for the three plant associations 99 5.14 Comparison between overstory and understory root surface areas for the three plant associations .... 100 5.15 Ratios of biomass of unsuberized, < 2 mm, 2-5 mm, and £ 5 mm overstory roots to whole-tree biomass for the three plant associations 105 5.16 Ratios of root-to-foliage plus twig biomass for the three plant associations 108 5.17 Bulk density and coarse fragment content of the s o i l s taken from 14-17 September 1977 separated by plant association and depth 110 5.18 Organic matter, nitrogen, and carbon-to-nitrogen rat i o s of the s o i l s taken from 14-17 September 1977 separated by plant association and depth 112 5.19 Hydrogen-ion a c t i v i t i e s and cation exchange capacities of s o i l s taken from 14-17 September 1977 separated by plant association and depth 114 5.20 Volumetric water contents of s o i l s separated by plant association, time of sampling, and depth .... 115 5.21 Temperatures in the s o i l s at 0-10 cm and 10-30 cm depths separated by plant association and time of sampling 117 5.22 Simple correlations between s o i l properties and the OPHO THPL, ABAM TSME, and VAME TSME for the three categories of overstory root biomass 119 5.23 Simple correlations between s o i l properties and the OPHO THPL, ABAM TSME, and VAME TSME plant associations for the two size categories of understory root biomass 124 5.24 Summary of s o i l properties for the OPHO THPL, ABAM TSME, and VAME TSME 127 X LIST OF FIGURES Figure Page 3.1 Location of the sampling s i t e at 49°45 ' N and 123° 03' W 27 3.2 Location of replicates 1, 2, and 3 of the Oplopanaco (horridi) - Thujetum plicatae (0) Association, Abieto (amabilis) - Tsugetum mertensianae (A) Association, and Vaccinio (membranacei) - Tsugetum mertensianae (V) Association 28 3.3 A view of the Oplopanaco (horridi) - Thujetum plicatae Association (abietetosum amabilis subassociation ) 29 3.4 A view of the Abieto (amabilis) - Tsugetum mertensianae Association (variant abieto-tsugetum mertensianae) 32 3.5 A view of the Vaccinio (membranacei) - Tsugetum mertensianae Association 34 4.1 Selection of < 2 mm overstory roots 52 4.2 Understory < 2 mm roots 52 4.3 Example of roots after sorting 53 5.1 Variation in overstory < 2 mm root mass-volume for replicates of the plant associations 65 5.2 Variation in overstory 2-5 mm root mass-volume with depth for replicates of the plant x i associations 66 5.3 Variation in unsuberized root mass-volume with time for the three plant associations 83 5.4 Variation in understory < 2 mm root mass-volume with time for the three plant associations 85 5.5 Variation in overstory root biomass with time for the three plant associations 88 5.6 Variation in understory root biomass with time for the three plant associations 92 5.7 Summary by plant association of biomass, length, and surface area for the various sizes and categories of overstory (0) and understory (U) roots 97 5.8 Relationship between foliage plus twig biomass and overstory < 2 mm suberized plus unsuberized root biomass 107 xi i ACKNOWLEDGEMENTS Contributions of the following were ess e n t i a l to the completion of thi s study and are gr a t e f u l l y acknowledged: Dr. J.P. Kimmins for his d i r e c t i o n and support throughout; P. Courtin for f i e l d and laboratory assistance; Dr. J . Otchere-Boateng and K.M. Tsze for s o i l analysis; Dr. T. Kozak for advice regarding the s t a t i s t i c a l design and subsequent s t a t i s t i c a l analysis; Dr. T.M. Ballard, Dr. T.A. Black, Dr. J..V. Thirgood, Dr. J.H. Bassman, and K.L. Kassay for c r i t i c a l reviews; and C.J. Lowe for typing and word processing. 1 1 . INTRODUCTION Subalpine forest ecosystem structure and function in coastal southwestern B r i t i s h Columbia have been the focus of only one study (Brooke and others 1970). Thus, information about t h i s subject i s limited. U n t i l recently, t h i s lack of knowledge has had no dire c t impact on the forest industry, but as the supply of mature timber at lower elevations has been depleted, logging pressure on subalpine forests has increased. The resulting clearcuts frequently proved d i f f i c u l t to reforest; and i t became obvious that these forests could not be treated in the same manner as those at lower elevations. It also became clear that more information about the structure and function of subalpine forest ecosystems was needed. As a result of the limited information available, research on climax forest ecosystems was started. In the i n i t i a l stages th i s involved a number of investigations concerning above-ground biomass of trees, forest productivity, and nutrient cyc l i n g (Krumlik 1979); the role of above-ground understory vegetation in nutrient c y c l i n g (Yarie 1978); rates of above-ground l i t t e r decomposition; s o i l physical and chemical properties; properties of the s o i l free water; and studies of forest microclimate. The present study was undertaken to examine the below-ground fine root component of these ecosystems. 2 The objectives of t h i s thesis were as follows: 1. To quantify and compare the v e r t i c a l d i s t r i b u t i o n , biomass, length, and surface area of the roots £ 5 mm in diameter of the overstory and understory vegetation in ridgetop, middle-slope, and lower-slope forest ecosystems during one growing season. 2. To determine the relationship between biomass of the ^ 5 mm roots of the overstory vegetation and the above-ground biomass of the overstory vegetation for the ecosystems studied. 3. To determine i f root d i s t r i b u t i o n was correlated with s o i l bulk density, content of coarse fragments, organic matter, nitrogen and phosphorus, s o i l pH, cation exchange capacity, s o i l moisture, or s o i l temperature. A co-worker determined the nutrient content of the overstory roots < 5 mm in diameter. (Courtin, P. 1978. Concentrations of macronutrients in fine roots of a coastal subalpine forest in B r i t i s h Columbia. University of B r i t i s h Columbia 70 p. BSF Thesis.) A l l of the studies mentioned above must also be done in younger stands of various ages i f we are to increase our understanding of the response of subalpine forest ecosystems to s i l v i c u l t u r a i practices. 3 2. LITERATURE REVIEW Investigations of the anatomy, morphology, and physiology of roots and their response to environmental factors such as temperature, l i g h t , moisture, and nutrient a v a i l a b i l i t y have been reviewed by Lyr and Hoffman (1967), and Sutton (1969). Textbooks that deal with the subject are also available (KSstler and others 1968, Carson 1974, Torrey and Clarkson 1975, Russell 1977). Currently, the emphasis in research on roots i s on quantifying their growth and production, a subject reviewed by Hermann (1977). 2.1 Depth D i s t r i b u t i o n of Fine Roots Data on the depth d i s t r i b u t i o n of fine roots are summarized in Table 2.1. The percentage of mass of fine roots per unit area contained in the upper 30 cm of the s o i l and the depth that includes 90 % of the t o t a l mass of fine roots i s presented. These points were chosen a r b i t r a r i l y to allow comparison between the studies. Most s t r i k i n g in Table 2.1 i s the v a r i a b i l i t y in d i s t r i b u t i o n of roots. Percentage of mass above 30 cm varied from 24 to 96. In one case, 90 % of the root mass was above 23 cm while in another i t was above 125 cm. Depth of sampling also varied considerably, to a maximum of 270 cm. Considering the e f f e c t s of species, le v e l s of stand density, stand age, and environmental and edaphic factors on root growth t h i s v a r i a b i l i t y i s not surprising. In addition, some of these Table 2 . 1 . D i s t r i b u t i o n of roots < 6.4 mm diameter i n f o r e s t ecosystems Percent of Depth of Spec ies ( a g e ) , S i t e and /or S o i l S i z e Mass Above 90% o f Depth o f D e s c r i p t i o n , L o c a t i o n , Source 30 cm Mass Sampling Con i fe rous Fores ts - m m - cm Abies f i rma (20), moderate ly mois t < 2 66 83 180 s o i l , Japan , Kar izumi 1976 2-5 63 86 180 Abies - Vacc in ium (mature) , n a t u r a l , 1810 m, Montana, Weaver 1977 < 5 ^ 5 0 / u Abies - Vaccin ium (mature ) , n a t u r a l , c Ac C 7 7n 2360 m, Montana, Weaver 1977 5 4 3 s / / u Chamaecyparis obtusa ( 3 8 ) , p l a n t a t i o n , < 2 83 45 120 moderately moist s o i l , Japan , Kar izumi 1976 2-5 87 36 120 Chamaecyparis p i s i f e r a ( 3 8 ) , moderate ly < 2 88 36 90 mois t s o i l , Japan , Kar izumi 1976 2-5 92 27 90 Table 2.1 cont inued Spec ies ( a g e ) , S i t e and /or S o i l D e s c r i p t i o n , L o c a t i o n , Source S i z e Percent of Mass Above 30 cm Depth of 90% o f Mass Depth of Sampling Con i fe rous Fo res ts — mm — cm Cryptomer ia j a p o n i c a ( 3 4 ) , p l a n t a t i o n , dry s o i l , Japan , Kar izumi 1976 < 2 2-5 72 65 60 75 120 120 Cryptomer ia j a p o n i c a ( 3 4 ) , p l a n t a t i o n , moderately moist s o i l , Japan , Kar izumi 1976 < 2 2-5 56 47 75 83 120 120 Cryptomer ia j a p o n i c a ( 2 9 ) , p l a n t a t i o n , moist s o i l , Japan , Kar izumi 1976 < 2 2-5 49 41 80 102 120 120 L a r i x l e p t o l e p i s ( 4 5 ) , p l a n t a t i o n , moderately mois t s o i l , Japan , Kar izumi 1976 < 2 2-5 86 78 40 48 120 120 P i cea g lauca ( 3 9 ) , p l a n t a t i o n , s i l t loam s o i l , Ma ine, S a f f o r d and B e l l 1972 < 3 96 28 45 P i cea g lauca and Ab ies l a s i o c a r p a (230) , n a t u r a l , sand s o i l , c e n t r a l B r i t i s h Columbia Kimmins and Hawkes 1978. Overs tory component < 6.4 92 23 98 Understory component <6 .4 92 23 98 Table 2.1 cont inued Spec ies ( a g e ) , S i t e and /or S o i l D e s c r i p t i o n , L o c a t i o n , Source S i ze Percent o f Mass Above . 30 cm Depth of 90% of Mass Depth o f Sampling Con i fe rous Fores ts —mm — cm P icea sp . ( 6 7 ) , poor s i t e , Germany, Z b t t l 1964 < 2 79 39 70 P i cea sp . ( 6 8 ) , medium s i t e , Germany, Z o t t l 1964 < 2 80 42 70 P i cea sp . ( 5 7 ) , good s i t e , Germany, Z o t t l 1964 < 2 92 25 70 Pinus d e n s i f l o r a ( 3 5 ) , p l a n t a t i o n , moderately moist s o i l , Japan , Kar izumi 1976 < 2 2-5 58 43 125 120 270 270 Pinus s t robus ( 4 2 ) , moderate ly moist s o i l , Japan , Kar izumi 1976 < 2 2-5 84 76 48 53 90 90 Pinus s p . ( 6 5 - 7 0 ) , p l a n t a t i o n , sand s o i l , Germany, Hausdor fer 1957 Overs tory component < 2 67 43 50 Understory component, (g rass) < 2 93 28 50 Pinus sp . ( 125 ) , p l a n t a t i o n , sand s o i l , Germany, Hausdor fer 1957 Overs tory component < 2 71 41 50 Understory component, (g rass) < 2 89 32 50 Table 2.1 cont inued Percent o f Depth of Spec ies (age ) , S i t e and /o r S o i l S i z e Mass Above 90% of Depth o f D e s c r i p t i o n , L o c a t i o n , Source 30 cm Mass Sampling Con i fe rous Fo res t s Pseudotsuga m e n z i e s i i ( 20 ) , p l a n t a t i o n , g r a v e l l y sandy loam s o i l , southern B r i t i s h Co lumbia , Nnyamah and B lack 1977a Pseudotsuga m e n z i e s i i ( 36 ) , p l a n t a t i o n , sand to sandy loam s o i l , B r i t a i n , Reynolds 1970 Pseudotsuga - Ca lamagros t is (mature) , n a t u r a l , 1830 m, Montana, Weaver 1977 Pseudotsuga - Symphoricarpos (mature) , n a t u r a l , 1650 m, Montana, Weaver 1977 Tsuga canadensis ( 38 ) , moderate ly mois t s o i l , Japan , Kar izumi 1976 -mm- cm (most <2) 81 32 65 < 6 24 75 107 (most <1) < 5 65 63 70 < 5 67 59 70 < 2 66 80 150 2-5 69 74 150 Table 2.1 cont inued Spec ies ( a g e ) , S i t e and /o r S o i l S i z e Percent o f Mass Above Depth o f 90% o f Depth o f D e s c r i p t i o n , L o c a t i o n , Source 30 cm Mass Sampling -mm- cm Deciduous Fo res t s B e t u l a davu r i ca ( 3 2 - 4 0 ) , moderate ly mois t s o i l , Japan , Kar izumi 1976 . < 2 2-5 83 79 47 50 90 90 Be tu la p l a t y p h y l l a v . j a p o n i c a (19 -39 ) , moderate ly mois t s o i l , J a p a n , Kar izumi 1976 < 2 2-5 84 83 45 47 90 90 Fagus s y l v a t i c a n a t u r a l , Germany Meyer and Got tsche 1971 < 2 2-5 80 63 41 48 80 80 L i r i odend ron t u l i p i f e r a (48 ) , n a t u r a l , s i l t loam s o i l , Tennessee, H a r r i s and o thers 1973 < 5 85 38 60 Popu lus , (mature) , 1830 m, Montana, Weaver 1977 < 5 90 30 70 Populus - Symphoricarpos (mature) , 1560 m, Montana, Weaver 1977 < 5 62 56 70 Quercus mongol ica v . g r o s s e s e r r a t a (36 -46 ) , moderately mois t s o i l , J a p a n , Kar izumi 1976 < 2 2-5 73 73 52 53 120 120 00 Table 2.1 cont inued Percent o f Depth o f Spec ies ( a g e ) , S i t e and /or S o i l S i z e Mass Above 90% of Depth of D e s c r i p t i o n 30 cm Mass Sampling T v T \ r \ i Pr*"in "f AV»O c f C .... ... _ . mm 1 i U[J I L U 1 r\Cl n i l U I U J L O MIIII vJII R a i n f o r e s t (mature), n a t u r a l , loamy sand < 2 58 88 130 s o i l , p l a t e a u , Ivory C o a s t , Hu t te l 1975 2-5 53 91 130 R a i n f o r e s t (mature) , n a t u r a l , sandy loam < 2 68 88 130 s o i l , t ha lweg , Ivory C o a s t , Hu t te l 1975 2-5 64 89 130 R a i n f o r e s t (mature) , n a t u r a l , c l a y < 2 81 68 130 loam s o i l , Ivory C o a s t , Hu t te l 1975 2-5 61 91 130 R a i n f o r e s t (mature) , n a t u r a l , Ghana, < 2 92 29 120 Jen i k 1971 2-5 83 39 120 R a i n f o r e s t (mature) , n a t u r a l , G ian t < 3 87 32 90 Podzo l , Cen t ra l Amazon, K l i nge 1976 >3-6 72 38 90 R a i n f o r e s t (mature) , n a t u r a l , < 3 48 56 107 L a t o s o l , Cen t ra l Amazon, K l i nge 1976 >3-6 43 63 107 10 differences could also ari s e from d e f i c i e n c i e s in the data res u l t i n g from: small numbers of samples often not selected randomly; sampling at d i f f e r e n t (often unspecified) times of the year; separation of roots into various d i f f e r e n t morphological and size categories (range of size classes was from < 2 mm to ^ 6.4 mm) using various techniques, and sampling to d i f f e r e n t depths, thus missing an undetermined amount of roots. Only a few studies have been undertaken to evaluate the effects of s i t e , s o i l type, age of stand, and contribution by understory vegetation to root d i s t r i b u t i o n . For example, in Cryptomeria japonica plantations roots became more abundant at greater depths as s o i l moisture increased from dry to moist (Karizumi 1976). In a Picea sp. stand in Germany, there was a smaller biomass of roots at greater depths on the good s i t e s when they were compared to the poor ones ( Z o t t l 1964). A similar trend for < 2 mm roots was observed in a t r o p i c a l rainforest where the s o i l s studied ranged from loamy sand to clay loam, ( i . e . root d i s t r i b u t i o n was shallowest on the clay loam) (Huttel 1975). However, in that study no such trend was evident in the case of 2-5 mm roots. With increasing stand age in a Pinus sp. plantation there was an increase in biomass of roots at greater depths (Hausdorfer 1957). 11 In a mature Picea glauca and Abies lasiocarpa stand, the depth d i s t r i b u t i o n s of the roots of the overstory and understory vegetation components were similar (Kimmins and Hawkes 1978); in a Pinus sp. stand, grass roots were more shallow than tree roots (HausdQrfer 1957). Considering the available information, i t i s clear that unqualified generalizations about root d i s t r i b u t i o n are impossible at this time. 2.2 Biomass of Fine Roots and Ratios to Shoots Numerous studies have been c a r r i e d out to ascertain fine root biomass; however, comparison between them i s complicated by l i m i t a t i o n s similar to those discussed in Section 2.1. The difference in the upper size l i m i t of roots that were studied varied between 0.6 mm and 10 mm (Table 2.2). Maximum biomass reported was 18.4 t ha" 1 for < 5 mm roots in a Pseudotsuga menziesii stand in Montana (Weaver 1977); the lowest was 0.01 t h a - 1 for < 2 mm roots in Betula sp. stands in Japan (Karizumi 1976). Root-to-shoot r a t i o s , i . e . the r a t i o of fine root biomass to above-ground tree biomass, also varied, r i s i n g from a low of 0.001 for < 2 mm roots in a Pinus densiflora plantation in Japan (Karizumi 1976) to 0.079 for < 5 mm roots in a Liriodendron t u l i p i f e r a stand in Tennessee (Harris and others 1973). Table 2.2 Biomass and root-to-shoot rat ios for roots <10 mm in diameter in forest ecosystems Root-to-Species (age), Si te and/or Soi l Root Shoot Descr ipt ion , Locat ion, Source Size Biomass Ratio Coniferous Forests -mm- t ha~l Abies amabil is (23), na tura l , 1150 m, <2 3.6-10.6. Washington State, Vogt and others 1981 (sampling over 1 yr) Abies amabili s (180), natura l , 1150 m < 2 8.7-17.7 Washi ngton State , Vogt and others 1981 (sampling over 1 year) Abies balsamea (40-50), natura l , 1700 < 1.6 4.3 0.032 trees per ha, New Brunswick, Baskerv i l l e 1966 Abies balsamea (mature), sand s o i l , < 10 11.2 Newfoundland, Damman 1971 Abies f i rm a (20), moderately moist < 2 0.4 0.004 s o i l , Japan, Karizumi 1976 2-5 1.0 0.011 < 5 1.4 0.015 Abies las iocarpa (mature), na tura l , < 5 7.1 1810 m, Montana, Weaver 1977 Abies las iocarpa (mature), na tura l , < 5 15.1 2360 m, Montana, Weaver 1977 Chamaecyparis obtusa (38), p lanta t ion , <2 1.1 0.014 moderately moi st s o i l , Japan, 2-5 3. 5 0.045 Karizumi 1976 < 5 4.6 0.059 Chamaecyparis p i s i f e r a (38), moder- < 2 1.1 0.005 ately moist s o i l , Japan, Karizumi 2-5 1.9 0.008 1976 < 5 3. 0 0.01 3 Cryptomeria japonica (34), p lantat ion , < 2 1 . 0 0.006 dry s o i l , Japan, Karizumi 1976 2-5 1.7 0.009 < 5 2.7 0.015 Table 2.2 continued Rootr to-Species (age), Si te and/or Soi l Root Shoot Descr ipt ion , Locat ion, Source Size Biomass Ratio Coniferous Forests Cryptomeria japonica (34), p lantat ion, moderately moist s o i l , Japan, Karizumi 1976 Crypt s o i l , omeria japonica (29), moist Japan, Karizumi 1976 Larix l ep to l ep i s (45), p lanta t ion , moderately moist s o i l , Japan, Karizumi 1976 Picea abies (55), p lanta t ion , southern Sweden, Nihlgard 1972 Picea glauca (39), p lanta t ion , s i l t 1oam s o i l , Maine, Safford and Bel l 1972 Picea glauca and Abi es 1 a si ocarpa (230), na tura l , sandy-nutrient-poor s o i l , B . C . , central Kimmins and Hawkes 1978, Kimmins Picea mari ana (65), sand s o i l , Newfoundland, Damman 1971 Picea sp. (number of stands) , USSR, Orlov 1969 Picea sp. (67), poor s i t e , Germany, Zott l 1964 Picea sp. (68), medium s i t e , Germany, Zot t l 1964 •mm- t ha" 1 < 2 1.8 0.01 4 2-5 3.3 0.027 < 5 5.1 0.041 < 2 1. 5 0.008 2-5 2.4 0.01 3 < 5 3.9 0.021 < 2 0.3 0.002 2-5 0.6 0.004 < 5 0.9 0.006 < 5 2.0 0.005 < 3 7.0 -6.4 1.9 0.009 6.4 7.9 0.037 < 10 8.8 -0.6 1.5-3.1 -< 2 8.4 -< 2 6.2 Table 2.2 continued . . . Root-to-Species (age), S i te and/or Soi l Root Shoot Descr ipt ion , Locat ion, Source Size Biomass Ratio Coniferous Forests -mm- t ha~l Picea sp. (57), good s i t e , Germany, Zot t l 1964 Picea sp. (200), na tura l , sand s o i l , northern Taiga , USSR, Marchenko and Karlov 1962 Picea sp. and Betula sp. (200), sand so i1 , fores ted Tundra, USSR, Marchenko and Karlov 1962 Pinus dens i f lora (35), p lanta t ion , moderately moi st s o i l , Japan, Karizumi 1976 Pinus radiata (10), p lanta t ion , f ine textured s o i l , 1350 trees per ha, I A u s t r a l i a , Moir and Bachelard 1969 Pi nus radiata (20), p lantat ion , f ine textured soi 1 , 700 trees per ha, I A u s t r a l i a , Moir and Bachelard 1969 Pinus radiata (34), p lan ta t ion , f ine textured s o i l , 700 trees per ha, I A u s t r a l i a , Moir and Bachelard 1969 Pinus radiata (27-39), p lanta t ion , stony sandy s o i l , South A f r i c a , Heth and Donald 1978 Pinus s y l v e s t r i s (18), p lanta t ion , sand s o i l , central Sweden, Persson 1979. Overstory component Understory component (Call una vulgari s and Vaccinium vi t i s - i daea) < 2 4.8 — < 1 1 -5 < 5 0.9 5.4 6.3 0.003 0.016 0.01 9 < 1 1 -5 < 5 3.0 4.7 7.7 0.022 0.035 0.057 < 2 2-5 < 5 0.1 1 .1 1 . 2 0.001 0.007 0.008 4-3 3.5 -4-3 3.1 -4-3 2.1 -< 5 13.7 -< 2 0.3 < 2 1.0 Table 2.2 continued . . . Root-to-Species (age), S i te and/or Soi l Root Shoot Descr ipt ion , Location, Source Size Biomass Ratio Coniferous Forests -mm- t ha"1 Pinus s y l v e s t r i s (120) p lantat ion , sand s o i l , central Sweden, Persson 1979. Overstory component < 2 1.2 Understory component (Calluna vulgaris < 2 2.2 and Vacci nni urn vi t i s- i daea) Pi nus sp. (65-70), p lanta t ion , sand s o i l , Hausdorfer 1957. Overstory < 2 1.5 component Understory component (grass) < 2 0.7 Pi nus sp. (125), p lanta t ion , sand so i l Hausdorfer 1957. Overstory component < 2 1.4 Understory component (grass) < 2 1.6 Pi nus sp. (number of stands), < 6 2.7-3.6 USSR, Orlov 1969 Pi nus strobus (42) moderately moist < 2 0.1 0.001 s o i l , Japan, Karizumi 1976 2-5 0.5 0.007 < 5 0.6 0.008 Pinus s y l v e s t r i s (33), p lanta t ion , heavy clay s o i l , southern Scotland, <. 5 3.4 0.01 8 Ovington and Madgwick 1959 Pinus s y l v e s t r i s (28), p lanta t ion , <10 3.5 F in land , Malkonen 1974 Pinus s y l v e s t r i s (47), p lanta t ion , <10 2.7 F in land , Malkonen 1974 Pinus s y l v e s t r i s (45), p lanta t ion , <10 4.1 F in land , Malkonen 1974 Pinus taeda (15), p lanta t ion , <10 4.3 0.047 Tennessee, Harris and others 1977 Tab!e 2.2 conti nued Root-to-Species (age), Si te and/or Soi l Root Shoot Descr ipt ion , Locat ion, Source Size Biomass Ratio Coniferous Forests -mm - t ha -1 Pseudotsuga menziesii (40), na tura l , 320 m, gravel ly loamy sand, low product iv i ty stand, Washington State, Keyes and Grier 1981 < 2-2 5 5 8. 2. 1 0. 3 2 5 0. 0. 0. 033 009 042 Psuedotsuqa menziesii (40), natural , 320 m, s i l t loam, high product iv i ty stand, Washington State, Keyes and Grier 1981 < 2-2 5 5 2. 1. 4. 7 8 5 0. 0. 0. 006 004 01 0 Pseudotsuga menziesii (36), p lanta t ion , sand to sandy loam s o i l , B r i t a i n , Reynolds 1970 < (most 6 <1) 13. 7 Pseudotsuga menziesii (70), dry s i t e , Oregon, Santantonio 1978 < 5 7. 1 0. 01 7 Pseudotsuga menziesii (170), moderate s i t e , Oregon, Santantonio 1978 < 5 8. 5 0. 01 6 Pseudotsuga menziesii (120), wet s i t e , Oregon, Santantonio 1978 < 5 9. 0 0. 01 7 Pseudotsuga menziesii (480), na tura l , Oregon, Santantonio 1974 < 5 9. 7 0. 016 Pseudotsuga menziesii (mature), na tura l , 1830 m, Montana, Weaver 19 77 < 5 18. 4 — Pseudotsuga menziesii (mature), na tura l , 1650 m, Montana, Weaver 1977 < 5 6. 0 Tsuga canadensis (38), moderately moist s o i l , Japan, Karizumi 1976 < 2-< 2 •5 5 0. 3. 4. 8 6 4 0. 0. 0. 006 029 035 Table 2.2 continued . . . Root-to-Species (age), S i te and/or Soi l Root Shoot Descr ipt ion , Locat ion, Source Size Biomass Ratio Deciduous Forests -mm- t ha"1 Betula dayurica (32-40), moderately moi st soi1 , Japan, Karizumi 1976 < 2 2-5 < 5 0.01 0.1 0.1 0. 002 0.019 0.021 Betula piaty phy 11 a v. japonica (19-39), moderately moist s o i l , Japan, Karizumi, 1976 Fagus grandi f1ora, Acer rubrum, and Betula a l l eghaniens i s , na tura l , coarse 1oam soi1 , New Hampshire, Safford 1974 Fagus s y l v a t i c a , na tura l , Germany, Meyer and Gottsche 1971 < 2 2-5 < 5 < 3 < 2 2-5 < 5 0. 01 0.1 0.1 12.5 2.6 3.9 6.5 0.002 0.01 5 0.017 Fagus sy lva t i ca (90), natura l , southern Sv/eden, Ni hi gird 19 72 Liriodendron t u l i p f e r a (48), p 1 antati on, si It loam s o i l . Tennessee, Harris and others 1973 Quercu s mongolica v. grossesrrata (36-46), moderately moist s o i l , Japan, Karizumi 1976 Quercus robur (149), natura l , heavy clay so i1 , southern Sweden, Andersson 1970 Overstory and shrub component. Non-woody plant component Quercus spp. (100), natura l , a l l u v i a l s i t e , Czechoslovakia, Miroslav 1976 < 5 6.0 0.016 < 5 9.0 0.079 < 2 0.03 0.002 2-5 0.2 0.01 8 < 5 0.02 0.020 < 5 3.4 0.01 7 < 5 2.6 0.01 3 O 1.4 0.004 18 Table 2.2 continued . . . Root-to-Species (age), Si te and/or Soi l Root Shoot Descr ipt ion , Locat ion, Source Size Biomass Ratio Tropical Rainforests -mm i - t ha-1 Rainforest (mature), na tura l , loamy < 2 7. 3 sand s o i l , plateau, Ivory Coast, 2- 5 3. 6 Huttel 1975 _< 5 10. 9 Rainforest (mature), na tura l , sandy < 2 5. 3 loam soi l , t h a l w e q , Ivory Coast, 2- 5 3. 3 Huttel 1975 _< 5 8. 6 Rainforest (mature), natura l , clay < 2 4. 8 loam s o i l , Ivory Coast, Huttel 2- 5 2. 8 1 975 <_ 5 7. 6 Rainforest (mature), na tura l , < 2 1. 8 Ghana, Jemk 1971 2- 5 4. 4 _< 5 6. 2 Rainforest (mature), na tura l , Giant < 3 1 1. 4 Podzol, central Amazon, Klinge 1976 3- 6 3. 1 < 6 14. 5 Rainforest (mature), na tura l , < 3 9. 8 Latoso l , central Amazon, Klinge 1976 3- 6 4. 5 6 14. 3 19 It would have been preferable to relate fine root mass to foliage mass of trees. However, data on foliage mass have rarely been reported in studies dealing with fine roots. The disadvantage of the root-to-shoot r a t i o i s the large amount of dead mass in the stem which may overshadow the re l a t i o n s h i p between roots and fol i a g e . In a Cryptomeria japonica plantation, biomass and the root-to-shoot r a t i o increased on s i t e s with dry s o i l in comparison to those with moderately moist s o i l , then dropped on those with moist s o i l (Karizumi 1976). Pseudotsuga menziesii stands in Oregon had 7.1 t ha" 1 of < 5 mm roots on dry s i t e s , increasing to 9.0 t ha" 1 on wet s i t e s while root-to-shoot r a t i o s remained similar (Santantonio 1978). In Washington State, Keyes and Grier (1981) compared low-productivity with high-productivity Pseudotsuga menziesii stands. They found 10.5 t ha" 1 versus 4.5 t ha" 1 of roots ^ 5 mm in diameter in the low-productivity and high-productivity stands, respectively. Root-to-shoot ratios were over 4 times greater on the low productivity stand as compared to the high productivity stand. Understory roots made up 7.9 of the t o t a l 9.8 t ha" 1 of roots ^ 6.4 mm in diameter in a rather open, mature Picea  glauca and Abies lasiocarpa stand in central B r i t i s h Columbia (Kimmins and Hawkes 1978). In a 125-year-old Pinus sp. plantation, grass roots < 2 mm in diameter constituted 1.6 of the t o t a l 3.0 t ha" 1 of roots < 2 mm in diameter; in a 65 to 70-year-old plantation they constituted 0.7 of the t o t a l 2.2 t ha" 1 (Hausdorfer 1957). Non-woody plant roots < 5 mm in 20 diameter made up 2.6 of the t o t a l 6.0 t ha" 1 in a 149-year-old Quercus robur stand in southern Sweden. Time of sampling has a d i s t i n c t influence on biomass. For example, in a 120-year-old Pinus s y l v e s t r i s plantation with a mean biomass of 1.2 t ha" 1 for overstory < 2 mm roots, the annual range was from 0.8 to 1.5 t ha" 1 (Persson 1979). Similar fluctuations were also evident in 18-year-old plantations in the same study. In Pseudotsuga menziesii stands on dry s i t e s the mean biomass of roots < 5 mm in diameter was 7.1 t ha" 1 with an annual range from 4.7 to 9.0 t ha" 1 (Santantonio 1978). On moderate s i t e s in the same study the range was 5.2 to 10.6 t ha" 1 while on wet s i t e s i t was 5.5 to 11.4 t ha" 1 . No seasonal pattern to the fluctuations was apparent in the above cases. On the other hand, in Washington State, Vogt and others (1981) have found evidence of a peak in September in biomass of < 2 mm roots in an Abies amabilis stand. However, the peak in September was d i f f e r e n t s t a t i s t i c a l l y from values determined at other times of the year only in a 180-year-old stand, and not in a 23-year-old stand. Keyes and Grier (1981), studying a low-productivity Pseudotsuga  menziesii stand, found a maximum biomass of roots < 2 mm in diameter of 8.3 t ha" 1 in June and a minimum of 2.1 t ha" 1 in December. In contrast, they found v i r t u a l l y no seasonal change in the biomass of t h i s size category of roots (2.7 t ha" 1) in the high-productivity stand. There may be seasonal fluctuations in root biomass in other studies which remain undetected due to large sampling errors. In any event, regardless of the s t a t i s t i c a l significance of seasonal fluctuations, a single sample from one time should not be extrapolated throughout the year, yet this has been done many of the studies reported in Table 2.2. 22 2.3 Length and Surface Area of Fine Roots In terms of function of the roots in the absorption of water and nutrients, length i s a more appropriate unit of measure than biomass. When d i f f e r e n t diameter classes of roots are to be compared, surface area i s probably a more appropriate unit; since, for a given length, the larger roots w i l l have a greater surface area exposed to the s o i l . Estimates of the length of roots per unit area reported in the l i t e r a t u r e varied from 0.075 to 48 km m ~ 2 (Table 2.3). Dry s i t e s in 58-year-old Picea sp. plantations had a greater root length than wet s i t e s (Kern and others 1961). Understory root length was 0.022 km m"2 of the t o t a l root length of 0.147 km m - 2 on one of the dry s i t e s and 0.013 of 0.0883 km m"2 on the wet s i t e . Kalela (1957) looked at annual va r i a t i o n in root length of a 75-year-old Pinus sp. plantation and found a peak in < 1 mm roots of 0.93 km n r 2 in July and a minimum of 0.38 km n r 2 in October. There was no such peak in roots > 1 mm in diameter. Sampling errors were not reported in the above studies. Surface area has not been reported often in the l i t e r a t u r e . Karizumi (1976) found values to a maximum of 2.1 m2 m"2 for £ 5 mm roots of a Chamaecyparis obtusa plantation in Japan. He reported lower values of 0.31 m2 n r 2 for Pinus d e n s i f l o r a . In an 18-year-old Pinus s y l v e s t r i s plantation, Persson (1978) presented information indicating that he found surface areas of < 1 mm roots ranging from 0.1 to 4.5 m2 nr 2 during a growing Table 2.3 Lengths of roots <6 mm in diameter in forest ecosystems Species (age), S i te and/or Soi l Root Descr ipt ion , Locat ion, Source Size Length Coniferous Forests -mm- km m~2 Picea s i tchens i s (11), p lanta t ion , peaty < 0.5 6.75 gley, Scot land, Ford and Deans 1977 Picea sp. (58), p lantat ion , dry s i t e , Germany, Kern and others 1961. Overstory <. 2 0.12 Understory component i 2 0.022 Picea sp. (58), p lanta t ion , dry s i t e , Germany, Kern and others 1961. Overstory <. 2 0.16 Understory component 2 0.0071 Pi cea sp. (58), p lanta t ion , moist s i t e , Germany, Kern and others 1961. Overstory <. 2 0.11 Understory component <. 2 0.0 Picea sp. (58), p lan ta t ion , wet s i t e , Germany, Kern and others 1961. Overstory <Z 0.075 Understory component ± 2 0.013 Picea sp. (40), p lanta t ion , sand s o i l , F in land , Kalela 1950 < 1 0.18 Pi cea sp. (80), p lantat ion , sand soi 1, F in land , Kalela 1 950 < 1 0.25 Picea sp. (130), p lanta t ion , sand s o i l , F i n l a n d , Kalela 1950 < 1 0.24 Pi nus sp. (40), p lantat ion , sand soi1 , F in land , Kalela 1950 < 1 0.22 Pi nus sp. (80), p lanta t ion , sand s o i l , F in land , Kale la 1 950 < 1 0.21 Table 2.3 continued . . . 24 Species (age), S i te and/or Soi l Descr ipt ion , Locat ion, Source Size Root Length Coniferous Forests Pi nus sp. (110), p lanta t ion , sand so i1 , F in land , Kale la 1950 -mm-< 1 km m""2 0.10 Pi nus sp. (110), p lanta t ion , sand, Kalela 1957 Pi nus sp. (150), peat bog, Fi nland, Heikurainen 1 957a Pseudotsuga menziesi i (20), p lantat ion gravel ly sandy loam, southern B r i t i s h Columbia, Nnyamah and others 1978 < 1 < 2 2-5 < 5 < 5 Pseudotsuga menziesii (36), p lanta t ion , sand to sandy loam s o i l , B r i t a i n , < 6 Reynolds 1970 (most <1) 0. 53 0.62 0.027 0.65 4.5 7.7 Deciduous Forests Eucalyptus marginata, natura l , various landforms, western A u s t r a l i a , Carbon 0.4 and others 1980 (avg.) 48 Tropical Rainforests Rainforest (mature), na tura l , Giant Podzol, Central Amazon, Klinge 1973 Rainforest (mature), na tura l , Latoso l , Central Amazon, Klinge 1973 < 3 3-6 < 6 < 3 3-6 5.4 0.47 5.9 1.9 0.7 25 season. He found no indication of a d i s t i n c t seasonal pattern in root surface area but did comment upon large sampling errors. 2.4 S o i l Physical and Chemical Parameters Root d i s t r i b u t i o n near the surface was determined by factors such as oxygen a v a i l a b i l i t y , p r e c i p i t a t i o n , and rate of nutrient c y c l i n g in the stands investigated by Karizumi (1976). In these stands the physical and chemical properties of the s o i l determined the root d i s t r i b u t i o n at greater depths. Simple correlations were determined by McGinty (1976) between biomass of roots ^ 25 mm in diameter and s o i l properties of a hardwood forest in Montana. The strongest relationship was with bulk density with a co r r e l a t i o n c o e f f i c i e n t of 0.40. Kern and others (1961) studied the relationship between root length and s o i l properties in Picea sp. plantations. They found an inverse r e l a t i o n s h i p between root length and the sum of available calcium, potassium, and phosphorus, while other parameters such as available water, bulk density, and pH were of minor importance. As was the case for the other information reported in thi s review, each ecosystem appears to be unique in the properties determining root d i s t r i b u t i o n . Perhaps when more studies are c a r r i e d out, some generalizations w i l l be possible, but t h i s i s not the case at t h i s time. 26 3. THE STUDY AREA 3.1 Location and Description The study area i s on the west-facing slopes of the Coast Mountains of southwestern B r i t i s h Columbia. It i s situated within Garibaldi Provincial Park at 49°45'N l a t i t u d e , 123°03'W longitude, approximately 10 km NE of Squamish (Fig. 3.1). Three plant associations were chosen in the study area. These associations are representative of the major types present in old-growth forests commonly found on the slopes of the mountains in the area. Three plots were established within each association. Brief descriptions of the associations follow: 1. The Oplopanaco (horridi) - Thujetum plicatae Association (abietetosum amabilis subassociation) (OPHO THPL) (Brooke and others 1970) was located on the concave lower slopes of the elevational transect at 1250 m (Fig. 3.2, 3.3). The moss layer was poorly developed. In contrast, there was a well developed herb layer of Rubus pedatus J.E. Smith, Veratrum v i r i d e A i t . , Athyrium f i l i x - f e m i n a (L.) Roth., Streptopus amplexifolius (L.) DC., Dryopteris austriaca (Jacq.) Woynar, and T i a r e l l a t r i f o l i a t a var. t r i f o l i a t a L. In the shrub layer Vaccinium membranaceum Dougl., V. alaskaense Howell, and V. ovalifolium Smith dominated, but were not abundant. Oplopanax horridum (Smith) Miq. was common in two of the three r e p l i c a t e s . (The 27 Figure 3 . 1 . Loca t ion of the sampl ing s i t e at 49° 45 ' N and 123° 03* W. 28 F igure 3 . 2 . Loca t ion of r e p l i c a t e s 1, 2 , and 3 o f the Oplopanaco ( h o r r i d i ) - Thujetum p l i c a t a e (9) A s s o c i a t i o n , Ab ie to ( amab i l i s ) - Tsugetum mertensianae (A) A s s o c i a t i o n , and V a c c i n i o (membranacei) - Tsugetum mertensianae (V) A s s o c i a t i o n . Figure 3.3. A view of the Oplopanaco (horridi) - Thujetum plicatae Association (abietetosum amabilis subassociation). The box houses a recording thermograph. 30 above species of the herb layer are referred to as 'understory vegetation' in the following sections.) Abies  amabilis (Dougl.) Forbes dominated the tree layer of the plots, with 72 to 78 % of the volume; the remainder of the tree layer was occupied by Tsuga mertensiana (Bong.) Carr., with sporadic T. heterophylla (Raf.) Sarg., except for one Thuja p l i c a t a Donn. on rep l i c a t e 1 of the pl o t s . Mean height of a l l tree species on the plots ranged from 26.4 to 36.2 m; mean diameter at breast height (DBH) ranged from 45.9 to 66.1 cm; stand density from 180 to 319 trees per ha; volume was between 888 and 1157 m3 ha" 1; basal area ranged from 64.5 to 70.1 m2 ha" 1; and biomass (determined by Krumlik (1979) using allometric equations) was between 419 and 482 t ha" 1 (Table 3.1). Tree d i s t r i b u t i o n was irregular and crown closure was incomplete. As a result there was a heavy cover of understory vegetation. 2. The Abieto (amabilis) - Tsugetum mertensianae Association (variant abieto-tsugetum mertensianae) (ABAM TSME) (Brooke and others 1970) was found at 1350 m, on gently sloping benches on the middle of the elevational transect (Fig. 3.2, 3.4). The moss layer was dominated by Dicranum  scoparium Hedw. with scattered Rhytidiopsis robusta (Hook.) Broth. Vaccinium membranaceum followed c l o s e l y by V. alaskaense and V. ovalifolium (the shrub species constituted the 'understory vegetation' of these p l o t s ) . The herb layer was i n s i g n i f i c a n t . Tsuga mertensiana accounted for 12 to 69 % of the number of trees and 17 to Table 3 . 1 . C h a r a c t e r i s t i c s o f the ove rs to ry component o f the sample p l o t s ; mod i f ied a f t e r Kruml ik (1979). A s s o c i a t i o n Oplopanaco ( h o r r i d i ) - Ab ie to (amab i l i s ) - Vacc i n i o (membranacei) -Thujetum p l i c a t a e Tsugetum mertensianae Tsugetum mertensianae R e p l i c a t e 1 2 3 1 2 3 1 2 3 P l o t area (ha) 0.141 0.139 0.179 0.095 0.105 0.106 0.186 0.091 0.100 Dens i ty ( t rees per ha) 319 180 190 673 457 311 1018 1275 520 Mean t ree ages 295 424 346 340 294 434 358 344 367 Mean t r e e DBH (cm) 45.9 66.1 62.7 44.9 45.9 55.9 28.1 27.9 43.5 Mean t r ee he igh t (m) 26.4 36.2 33.2 24.3 25.6 29.6 11.9 13.2 17.0 Basa l area (m 2 h a - 1 ) 64.5 70.1 65.3 119.2 83.2 87.5 82.5 93.1 102.4 Volume (m 3 h a - 1 ) 888 1157 912 1272 930 1206 523 609 826 C o n t r i b u t i o n by spec ies (% by volume) Tsuga mertensiana 22* 28* 26* 67 54 17 83 75 58 Abies amab i l i s 78 72 74 33 46 83 12 22 42 Chamaecyparis noo tka tens i s 0 0 0 0 0 0 5 3 0 Mass (met r i c t h a - 1 ) 419 482 435; 731 511 550 389 458 510 * A m inor , but u n t a l l i e d amount o f Tsuga h e t e r o p h y l l a was p resen t . Figure 3.4. A view of the Abieto (amabilis) - Tsugetum mertensianae Association (variant abieto-tsugetum mertensianae). The Stenenson screen houses a recording hygrothermograph and maximum and minimum thermometers. 33 67 % of the tree stem volume; Abies amabilis constituted the remainder. Mean height of both tree species on the plots ranged from 24.3 to 29.6 m, mean DBH from 44.9 to 55.9 cm, stand density from 311 to 673 trees per ha, volume from 930 to 1272 m3 ha" 1, basal area from 83.2 to 119.2 m2 ha" 1; and biomass was between 511 and 731 t ha" 1 (Table 3.1). Tree d i s t r i b u t i o n was f a i r l y regular and crown closure v i r t u a l l y complete. In contrast to the OPHO THPL (above) there was l i t t l e understory vegetation in thi s 'assoc i a t i o n . 3. Uppermost on the elevational transect on gentle knolls at 1450 m was the Vaccinio (membranacei) - Tsugetum mertensianae Association (Brooke and others 1970) (VAME TSME) (Fig. 3.2, 3.5). The moss layer in t h i s association consisted' of Dicranum scoparium with lesser amounts of Rhytidiopsis robusta. The shrub layer was dominated by Rhododendron albiflorum Hook, followed by Vaccinium  membranaceum (the shrub species constituted the 'understory vegetation' of these p l o t s ) . The herb layer was i n s i g n i f i c a n t . Tsuga mertensiana accounted for the majority of the volume (between 58 and 83 %) but Abies  amabilis was present in almost equal numbers (between 36 and 64 % of the trees ) . A minor component of Chamaecyparis  nootkatensis (D. Don) Spach. was present in two of the three p l o t s . Mean height of the trees ranged from 11.9 to 17.2 m, mean DBH from 27.9 to 43.5 cm, stand density from 34 Figure 3.5. A v iew of the V a c c i n i o (membranacei) - Tsugetum mertensianae A s s o c i a t i o n . The box houses a reco rd ing thermograph. 35 520 to 1275 trees per ha, stem volume from 523 to 826 m3 ha" 1, basal area from 82.5 to 102.4 m2 h a - 1 ; and biomass from 389 to 510 t ha" 1 (Table 3.1). Tree d i s t r i b u t i o n was clumpy and crown closure incomplete. As a result there was a heavy cover of understory vegetation. For purposes of brevity, the associations at 1250, 1350, and 1450 m w i l l be abbreviated as OPHO THPL, ABAM TSME, and VAME TSME, respectively. A summary of selected c h a r a c t e r i s t i c s of the associations appears in Table 3.2. Descriptions of the overstory component of the plots are presented in Table 3.1 and by Krumlik (1979). 3.2 Climate and Microclimate A climate station was established in a clearcut (1060 m) 1 km SE of the OPHO THPL plots in late 1975 and maintained u n t i l the spring of 1978. Data on the microclimate of each of the three plant associations were also c o l l e c t e d from the f a l l of 1975 u n t i l the f a l l of 1977. At each station maximum, minimum, and continuous temperatures were measured. Comparison of the temperature data showed that the clearcut had a milder climate (annual mean in 1976 of 5.7°C) than the stations on the study s i t e s . Of the three study s i t e s , the ABAM TSME Association was the warmest with an annual mean in 1976 of 4.1°C. It was followed by the OPHO THPL Association (3.8°C), while the coldest conditions occurred on the VAME TSME Association (3.5°C). Frost-free periods in 1976 ranged from Table 3 . 2 . Summary of c h a r a c t e r i s t i c s of the three p l an t a s s o c i a t i o n s . P lan t a s s o c i a t i o n : E l e v a t i o n (m): S i t e p o s i t i o n : R e l i e f shape: Slope ( ° ) : Aspec t : Drainage c l a s s : Seepage: Parent m a t e r i a l : Oplopanaco ( h o r r i d i ) -Thujetum p l i c a t a e 1250 lower s lope concave 5 to 12 west i m p e r f e c t l y t o moderately we l l d ra ined present c o l l u v i a l veneer / moraine b lanke t Ab ie to (amab i l i s ) -Tsugetum mertensianae 1350 mid-s lope t e r r a c e s t r a i g h t 0 to 20 none moderately we l l d ra ined absent moraine b lanket V a c c i n i o (membranacei) Tsugetum mertensianae 1450 upper s lope and c r e s t convex 0 to 40 v a r i a b l e we l l to moderately we l l dra ined absent s a p r o l i t e / b e d r o c k CO 37 119 days on the VAME TSME to 142 days on the clearcut. In 1977 the values were 109 and 135 days respectively. The temperature data are summarized in Appendix 1 and frost free periods in Appendix 2. Pr e c i p i t a t i o n was recorded on the clearcut in a rain gauge. It was greatest during the f a l l and winter with the maximum of 430 mm f a l l i n g in October 1975. Drier conditions prevailed during the spring and summer. The lowest monthly value recorded was 10 mm in September 1975. In 1976 t o t a l p r e c i p i t a t i o n was 2093 mm and in the following year i t was 1841 mm. During the f a l l and winter of 1975-76 p r e c i p i t a t i o n was 1991 mm,* predominantly as snow. An unusually deep snowpack resulted, remaining on the VAME TSME u n t i l August of 1976. In the following f a l l and winter, p r e c i p i t a t i o n was only 1047 mm. Consequently, the snowpack had disappeared from the VAME TSME by July in 1977. Generally, the OPHO THPL was free of snow f i r s t with the ABAM TSME a close second. The VAME TSME s t i l l had snow as much as one month afte r i t had melted off the ABAM TSME. Pr e c i p i t a t i o n data are summarized in Appendix 3. Krajina (1969) characterized the Mountain Hemlock Biogeoclimatic Zone in southwestern B r i t i s h Columbia as occurring between 900 and 1800 m (elevation l i m i t s vary from windward to leeward sides of mountains). He summarized the zonal climate employing Koppen's c l a s s i f i c a t i o n (Appendix in Trewartha 1968) as predominantly Dfc. That means i t i s cold and snowy, the coldest monthly mean temperature i s below -3°C, 38 the warmest month mean temperature i s over 10°C, winters are humid, summers are cool and short, and fewer than 4 months have a mean temperature over 10°C. The data from the forested ecosystems generally f a l l within the l i m i t s he gave for t h i s zone. 3.3 Geology and S o i l s Rock fragments taken from s o i l p i t s in the plots were analyzed by Dr. W.B. Danner of the Department of Geological Sciences at the University of B r i t i s h Columbia, Vancouver, B r i t i s h Columbia. Most of the rocks were pink and blue-grey dacite originating from Garibaldi Group volcanic flows (Mathews 1958a, b). Second in abundance were the f o l i a t e d and partly micaceous hornblende-quartz d i o r i t e s of the batholiths. Next were dense and heavy porphyritic rocks, possibly also from Garibaldi Group volcanic flows. Least abundant were dark green schistose rocks which may have come from metamorphosed older volcanic flows. This order of importance was maintained over the entire study area. Dr. Danner's report i s included in Krumlik's (1979) d i s s e r t a t i o n . In the OPHO THPL a c o l l u v i a l veneer covered a moraine blanket (Table 3.2). The ABAM TSME was covered by a moraine blanket, while sapro l i t e covered bedrock in the VAME TSME. 39 The s o i l s of the OPHO THPL were sometimes gleyed and in one case showed o r t s t e i n development (Table 3.3). Groundwater has a s i g n i f i c a n t influence on s o i l development in these ecosystems. Not a l l the s o i l s in the OPHO THPL were gleyed, probably due to the absence of reducing conditions in them either because of oxygen d i f f u s i o n through the water or the high dissolved oxygen content of the groundwater. On the ABAM TSME and VAME TSME the s o i l s were Orthic Humo-Ferric and Ferro-Humic Podzols with one Ortstein Humo-Ferric Podzol. A l l of the s o i l s of the three plant associations were in the s i l t loam (CSSC 1978) texture range. There appeared to be l i t t l e difference in the measured humus parameters between the associations (Table 3.3). The range in depths was from 4 to 23 cm. Bulk density ranged from 0.13 to 0.23 g cm"3. No data for pH of humus from modal s o i l p i t s were ava i l a b l e . Coarse fragment content increased with depth and from the OPHO THPL to the ABAM TSME and to the VAME TSME. In the B horizon the content was between 9.8% (ABAM TSME) and 16.4% (VAME TSME) of the volume. The highest measured value was in the C horizon of the OPHO THPL (44.0%) (Table 3.3). Bulk density of the s o i l increased with depth. There was l i t t l e difference between the associations in the values for bulk density when individual horizons were compared. Table 3 . 3 . Summary of c h a r a c t e r i s t i c s o f modal s o i l s based on two p i t s i n each of three r e p l i c a t e s of the OPHO THPL and ABAM TSME a s s o c i a t i o n s , and two p i t s i n two r e p l i c a t e s and th ree p i t s i n a t h i r d r e p l i c a t e o f the VAME TSME A s s o c i a t i o n . Data cour tesy o f Dr. J . P . Kimmins, F a c u l t y o f F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver, B .C. Oplopanaco ( h o r r i d i ) -Thujetum p l i c a t a e A s s o c i a t i o n Ab ie to (amab i l i s ) -Tsugetum mertensianae A s s o c i a t i o n Vacc in i o (membranacei) Tsugetum mertensianae A s s o c i a t i o n S o i l Subgroup* (CSSC 1978) GLOT.FHP(l) GL.HFP(2) 0.FHP(2) O.HFP( l ) O.FHP(3) O.HFP(2) OT.HFP(l ) TY .FO( l ) O.HFP(3) O.FHP(2) TY .FO( l ) Texture s i l t loam s i l t loam s i l t loam mean (n) range mean (n) range mean (n) range Th ickness of Hor izons (cm) LFH 12.8 (6) 8-22 9 .3 (6) 8-15 10.7 (7) 4-23 A 3.6 (5) 1-9 7.00 (6) 3-17 B 18.5 (6) 12-31 31.5 (6) 26-40 30.0 (7) 15-70 BC 25.0 (3) 17-33 18.0 (3) 10-33 16.9 (4) 6-26 Table 3.3 cont inued Oplopanaco ( h o r r i d i ) - Ab ie to (amab i l i s ) - Vacc in i o (membranacei) -Thujetum p l i c a t a e ' Tsugetum mertensianae Tsugetum mertensianae A s s o c i a t i o n A s s o c i a t i o n A s s o c i a t i o n mean (n) range mean (n) range mean (n) range Coarse Fragments (> 2 mm) (% volume by h o r i z o n ) * * A not present 9.8 (2) 6-14 16.4 (6) 6-55 B 12.1 (4) 1-24 13.4 (6) 6-24 24.5 (7) 6-51 BC 29.6 (3) 1-43 19.8 (5) 9-35 41.4 (5) 19-85 C 44.0 (1) not sampled 32.7 (3) 19-53 Bulk Densi ty (g cm )** LFH 0.18 (4) 0 .13 -0 . ,20 0.18 (6) 0 .14-0 .23 0.18 (6) 0 .15-0.21 A not present 0.4 (2) 0 . 4 - 0 . 5 0.6 (6) 0 . 5 - 0 . 8 B 0.9 (4) 0 . 5 - 1 , ,2 0.7 (6) 0 . 5 - 1 . 2 0.8 (7) 0 . 5 - 1 . 4 BC 1.3 (3) 0 . 8 - 1 , .7 1.0 (5) 0 . 7 - 1 . 2 1.3 (5) 1 .2-1 .4 C 2 .5 (1) not sampled 1.4 (3) 1 .1-1 .6 Table 3.3 cont inued Oplopanaco ( h o r r i d i ) -Thujetum p l i c a t a e A s s o c i a t i o n Ab ie to (amab i l i s ) -Tsugetum mertensianae A s s o c i a t i o n Vacc in i o (membranacei) -Tsugetum mertensianae A s s o c i a t i o n mean (n) range mean (n) range mean (n) range pH i n C a C l 2 LFH not a v a i l a b l e not a v a i l a b l e not a v a i l a b l e A not present 3.0 (2) 2 .8-3.1 2.9 (6) 2 .6 -3 .1 B 4 .6 (6) 4 . 0 - 5 . 0 4.6 (6) 3 .8 -5 .2 4 .3 (7) 3 . 0 - 5 . 3 BC 4 .9 (6) 4 . 6 - 5 . 2 5.3 (5) 4 . 8 - 5 . 4 5.0 (5) 4 . 5 - 5 . 3 C 5.0 (3) 4 . 9 - 5 . 0 not sampled 4.7 (3) 4 . 5 - 5 . 0 Methods are as f o l l o w s : Bulk d e n s i t y - Sample o f known volume (water i s poured i n t o p l a s t i c l i n e d excava t i on and measured) i s d r i e d to 105°C and weighed ( repor ted va lue does not i n c l u d e coarse f ragments ' mass and volume). Coarse fragments - Volume o f minera l ma te r i a l > 2 mm. pH i n C a C l 2 - 1:2 (w/w) s l u r r y , o rgan ic mat te r / 0.01M C a C l 2 o r 1:1 (w/w) s l u r r y , minera l s o i l / 0 . 0 1 M C a C l 2 , de te rmina t ion by pH e l e c t r o d e . * Not v e r i f i e d by chemical a n a l y s i s . GL = G leyed , 0T = O r t s t e i n , FHP = Ferro-Humic P o d z o l , 0 = O r t h i c , HFP = Humo-Ferr ic P o d z o l , TY = T y p i c , F0 = F o l i s o l . Number of p i t s so c l a s s i f i e d are i n paren theses . * * Means are not weighted by depth o f hor i zon i n the s o i l p i t s . Rep l i ca te 2 of the f i r s t a s s o c i a t i o n was not sampled f o r bulk d e n s i t y a n d , i n the case o f minera l s o i l , f o r coarse f ragments. S o i l pH in 0.01 M CaCl 2 showed an increase with depth in a l l p l o t s . Generally the s o i l s were a l l a c i d i c , r e f l e c t i n g the accumulation of a c i d i c decomposition products in the humus and the abundant leaching. 44 4. METHODS 4.1 F i e l d Numerous treatises exist on methods for sampling root systems (Kinman 1932, Lott and others 1950, Hough and others 1965, Mi t c h e l l and Woods 1966, USSR Academy of Sciences 1968, Yamakura and others 1972, Overton and others 1973, Karizumi 1974a, Safford 1976, Brown and Thilenius 1977, Bohm 1979, and others mentioned in the l i t e r a t u r e review). The methods used in t h i s study were chosen for th e i r s i m p l i c i t y and e f f i c a c y in achieving the objectives. Random sampling was chosen since a number of studies showed that fine roots tended to be randomly d i s t r i b u t e d in stands. (Moir and Bachelard 1969, Safford and B e l l 1972, McQueen 1973, Roberts 1976, Santantonio and others 1977). On the other hand, Nnyamah and others (1978), found that the density of roots of Pseudotsuga menziesii decreased with increasing distance from the boles of trees. In t h i s thesis, the sampling methods made i t impossible to determine i f there was any relationship between biomass of fine roots of trees in a sample and the distance of that sample from a tree. 45 4.1.1 Roots In each of the nine plots, four random points were marked. Root samples were taken 1 m N of these points between 2 and 9 June, 1 m to the SE between 18 and 21 July, and 1 m to the SW between 14 and 17 September 1977. Coring tubes (120.7 mm inside diameter with a 6 mm wall) were used for sampling. They consisted of one short cylinder (305 mm) for sampling the forest f l o o r , and a second, longer cylinder (711 mm) for sampling the mineral s o i l . One end of each cylinder was sharpened, while the-other was reinforced to accommodate a steel cap of high t e n s i l e strength. The units were driven into the ground with a sledge-hammer and withdrawn with a short extractor bar which was inserted through holes in the reinforced top of the c y l i n d e r . A large corer was chosen because i t was possible to reach inside to remove the sample and also because fewer roots would be pushed aside as the corer was hammered downward. In the event that a stone was encountered in sampling, i t was excavated (unless i t was over 50 cm in diameter) and the coring continued. Large stones were not encountered frequently. Sometimes bedrock prevented the f u l l 60-cm depth from being sampled. 46 The cores were separated into the following depth increments: 0-5, 5-10, 10-20, 20-30, 30-40, 40-50, and 50-60 cm. The f i r s t 10 cm were divided because of the rapid changes in root abundance near the surface. F i r s t l y , the short core was driven to a depth of 5 cm in the forest f l o o r , and then the sample was removed with a modified trowel. The corer was then driven further into the forest floor and the next sample was removed. When mineral s o i l was reached, or the short corer was inserted to i t s l i m i t of 20 cm, the large corer was used. Only one depth increment was taken at a time. This was done to reduce the problem of compaction of samples. Individual depth increments from the four cores taken within a plot were bulked to form one sample for each depth increment. This eliminated the p o s s i b i l i t y of measuring within-plot variation in the s t a t i s t i c a l treatment, but was necessary to reduce the number of samples to be processed. Within 48 hr, the samples were moved to the laboratory where they were stored at 2°C u n t i l processed. 4.1.2 S o i l Water After the cores were removed as described above, samples of organic matter or mineral s o i l from 5, 10, 20, and 40 cm depths were placed into 136 ml aluminum containers with a i r t i g h t l i d s for volumetric water content determination. As in the root sampling, bedrock or stones occasionally prevented a f u l l complement of samples from being taken. 47 The seasonal variation in s o i l water potential was measured at one location in each plot using a Wescor HR-33T dew point microvoltmeter and PT51-10 hygrometers. The hygrometers were calibra t e d in 0.1, 0.2, 0.5, and 0.9 mol (kg H 20)" 1 NaCl solutions. These corresponded to osmotic potentials of -462, -915, -2281, and -4158 kPa. Several 250 ml bottles of the solutions, each containing 8 hygrometers, were placed into an insulated Haake FK-2 constant temperature c i r c u l a t o r housed inside a constant temperature (21.1°C) room. Four hours were required for equilibrium to be established and then readings were taken. Calibration was c a r r i e d out at 10°C and 25°C. The above methods followed those of Nnyamah and Black (1977b) and Wiebe and others (1971). I n s t a l l a t i o n of the hygrometers involved excavating a 40 x 60 cm trench to a depth of 100 cm or u n t i l compacted t i l l or bedrock were encountered. The s o i l horizons were separated in the excavation. A steel rod, of a diameter equal to that of the hygrometer, was inserted 30 cm into the 40-cm-wide wall at one end of the p i t . The hygrometer was then pushed into t h i s 9 hole and sealed into place with s o i l or organic matter, depending on the horizons involved. Instruments were placed at 5, 10, 20, and 40 cm, and above any compacted t i l l or bedrock i f these were encountered before reaching 100 cm in depth in the p i t . The reason for placing the instruments over the compacted t i l l or bedrock was to sample a l l potential sources of water available to the vegetation in the event that s u r f i c i a l s o i l horizons dried out. The same s o i l that had been 48 removed from the p i t was c a r e f u l l y replaced, so that at least 60 cm of the wires buried at the same depth and were in the same materials as the sensing unit. This procedure was c a r r i e d out to minimize thermal gradients in the wire near the hygrometer. After the hole had been completely f i l l e d i n , a l l leads were staked in place to prevent w i l d l i f e such as bears from tearing probes out of the ground. Readings were taken from the units every two weeks from 14 June to 2 October — corresponding to the time when the ground was free of snow. The i n s t a l l a t i o n procedures and performance of instruments such as these are described in d e t a i l by Nnyamah and Black (1977b). 4.1.3 S o i l Temperature S o i l temperature was recorded in the p i t s described above with Kahlsico three-probe remote thermographs. Sensors were inserted perpendicular to the faces of the s o i l p i t s at least 30 cm from the hydrometers at depths of 5, 10, and 20 cm. I n i t i a l c a l i b r a t i o n was c a r r i e d out on the s i t e , in ice water. Also, readings were cross-checked whenever the s o i l water potential was measured, since the hygrometers also contained thermocouples. 49 4.2 Laboratory t 4.2.1 Root Washing and Separation In order to avoid a systematic error due to possible decay of roots in cold storage, the order of plot analysis was random. A l l samples from a plot were processed at one time. To reach the measurement stage t h i s took at least 12 hours for each set of 7 samples from a p l o t . The f i r s t set of samples was processed by 5 August, the second by 27 August, and the t h i r d by 9 October 1977. Each sample was treated by emptying i t onto a table and removing a l l roots > 2 mm in diameter. A l l of the roots < 2 mm in diameter that were attached to the larger roots were removed. If the sample contained a considerable biomass of < 2 mm roots then the < 2 mm roots that had been removed were cut into lengths of 5 cm or l e s s , and added back to the sample. This sample of s o i l and roots was mixed thoroughly and subsampled by quartering, i . e . the p i l e of sample material was halved, one half was removed, and the remaining half was remixed and again bisected. The process was continued u n t i l the mass of roots remaining was s l i g h t l y over 6 g. This amount was s u f f i c i e n t to permit the measurement of root physical parameters and also was s u f f i c i e n t for the chemical analyses which constituted the second part of t h i s study. (Courtin, P. 1978. Concentrations of macronutrients in fine roots of a coastal subalpine forest in B r i t i s h Columbia. University of B r i t i s h Columbia. 70 p. 50 BSF Thesis.) Ty p i c a l l y , the forest f l o o r required considerable subsampling. Where the sample contained less than 6 g of < 2 mm roots the entire sample was processed. Often, the 50-60 cm sample did not require subsampling and, in some cases, was i n s u f f i c i e n t for routine chemical analysis. The physical parameters for the entire root sample were obtained by multiplying the measured subsample parameters by the r a t i o of the mass of t o t a l sample divided by the mass of the subsample. The sample or subsample was placed onto a 2-mm-mesh screen and washed with a water spray to separate the organic debris or mineral s o i l from the roots. The clumps of roots and dead organic matter remaining after washing were teased apart with tweezers in a p l a s t i c dish f i l l e d with water. Fungal mycelia attached to mycorrhizal roots were removed as much as possible at t h i s time. The material passing through the 2-mm screen was mostly dead and decaying segments of roots. However, there was also a component of I k e root fragments. Many of these were unsuberized roots. For t h i s reason, the unsuberized root data underestimate the true values. The d i s t i n c t i o n between l i v e and dead roots (retained on the screen) was based on root appearance since no s a t i s f a c t o r y technique for ide n t i f y i n g functionally a l i v e roots has yet been developed for large-scale studies. Thus, a l l roots that were sound with no indication of separation of the cortex from the endodermis and no signs of internal decay (based on microscopic examination of sections of representative samples) were 51 considered a l i v e . It i s possible to use nuclear staining (Knievel 1973, Holden 1975) to determine the number of l i v e and dead c e l l s in a sample of roots that are considered to be a l i v e . This would have provided a check on the method based on morphological c r i t e r i a . Nuclear staining was not done due to the limited time available and the necessity for carrying out the analysis immediately after sampling. A l l roots were separated•into overstory vegetation and understory vegetation components based on previous excavation of the root systems of individuals of the species present in the three plant associations. They were also separated into size categories: those < 2 mm and those from 2 to 5 mm (Fig. 4.1, 4.2, 4.3). Overstory roots < 2 mm were divided further into unsuberized and suberized components during the estimation of root length and surface area. Unsuberized roots lacked any sign of development of secondary thickening of the cortex, were translucent, and a creamy color. Suberized roots had a corky cortex, were opaque, and brown to black in color. Root t i p s were counted. The treatment of these data was not an objective of t h i s thesis but, due to the importance of this information, root t i p data are presented in Appendix 4. To test the significance of mineral p a r t i c l e contamination of roots, f i r s t the ash content remaining after heating overstory < 2 mm and 2-5 mm roots in a muffle furnace at 475°C for 4 hours was calculated. Then, the content of roots not growing in mineral s o i l ( i . e . only in organic horizons) was subtracted from the ash content of the roots from their 52 Figure 4 . 1 . Selection of < 2 mm overstory roots. Translucent t i p s are part of the unsuberized category. Grid l i n e s are 1 cm apart. Figure 4.2. Understory < 2 mm roots. Grid l i n e s are 1 cm apart. Figure 4.3. Example of roots a f t e r s o r t i n g . Upper l e f t a re unders tory < 2 mm r o o t s , upper r i g h t are ove rs to ry > 2 mm r o o t s , lower l e f t are the overs to ry < 2 mm r o o t s , lower r i g h t are a l s o ove rs to r y < 2 mm roo ts which c o n s t i t u t e the subsample t ha t has been used f o r es t imates o f l e n g t h , su r face a r e a , and number of t i p s . The smal l con ta i ne r i n s i d e the l a rge one a t the lower l e f t has the unsuber ized t i p s in i t . Th is sample was from a depth o f 40 t o 50 cm on the ABAM VAME a s s o c i a t i o n . G r i d l i n e s are 1 cm apa r t . respective depth increments. The difference was considered to be due to contamination of roots by adhering mineral p a r t i c l e s and was to be applied in correcting the root data i f necessary. 4.2.2 Root Length and Surface Area The < 2 mm roots were cut into 1 cm segments and spread randomly onto a 20 x 20 cm glass plate. The lengths and diameters of unsuberized overstory roots were measured with a micrometer eyepiece i n s t a l l e d into a dissecting microscope; then they were removed for drying and weighing. The lengths and diameters of the remaining suberized roots < 2 mm and a l l understory roots < 2 mm were estimated according to methods outlined by Newman (1966) and applied by Ambler and Young (1977). The intersections of roots with 20 p a r a l l e l l i n e s scribed onto the glass plate at 0.9 cm intervals were counted. Applying the formula given by Newman (1966) gave an estimate of length: J I N A L = (1) 2H L = length of roots in cm N = number of intersections A = area of plate = 400 cm2 H = length of l i n e = 400 cm 30 of these intersections were measured with the These were averaged to y i e l d a mean root diameter. where Diameters at microscope. 55 Whenever possible, a sample s u f f i c i e n t to give between 100 and 200 intersections was used. When th i s meant subsampling, the data (including those for unsuberized roots) were subsequently corrected by the r a t i o of dry mass of t o t a l sample divided by the dry mass of the subsample. Sometimes, when only a small sample was present, fewer than 100 intersections were counted. This only occurred for depths at which few roots were found and was not thought to influence interpretations. As a test of t h i s method of c a l c u l a t i n g root length, a cord 1.0 mm in diameter and 150 cm long was cut into 1 cm segments and placed onto the plates. Ten counts, each followed by a rearrangement of the segments of s t r i n g on the plate, gave the following r e s u l t s : X = 142 cm SE = 8.14 n = 10 where X is the mean, SE i s the standard error, and n i s the size of sample This suggests the method yielded results 5.1 % too low. This is not a large error considering the other l i m i t a t i o n s in root sampling. Non-random placement of roots on the gri d would have created a bias in the r e s u l t s . To test for t h i s , the g r i d was turned 90° to the d i r e c t i o n used for the f i r s t count and a second count was made. The counts were v i r t u a l l y i d e n t i c a l : 1. X, =111 SE = 13.9, 2. X 2 = 108 SE = 13.7 n,, 2 = 11 where X, is mean of f i r s t counts 56 X 2 is mean of perpendicular counts The length of roots £ 2 mm was measured with a ruler, and their diameter with vernier c a l i p e r s . Several measurements were taken along irregular pieces and a mean diameter was determined. For pieces longer than 20 cm, diameters were measured at 5 cm intervals and a mean diameter was calculated. Surface area of roots was calculated by assuming a c y l i n d r i c a l shape: irD x L = SA (2) where D = diameter of roots in cm L = length of roots in cm SA = surface area of roots in cm2 4.2.3 Root Biomass Following measurements for ca l c u l a t i n g length and surface area, the < 2 mm roots were dried at 70°C for 24 hr in a forced draught oven. Roots 2-5 mm in diameter were dried u n t i l no further weight loss was detected (usually t h i s required less than 48 hr). After drying, a l l roots were placed into a desiccator u n t i l they were cool; then they were weighed. A l l results reported in thi s thesis are based on dry biomass. 57 4.2.4 S o i l Physical and Chemical Parameters Gravimetric s o i l moisture was determined by drying the samples at 105°C for 12 hr (no further moisture loss occurred afte r t h i s time) followed by cooling in a desiccator, and weighing. The mean weekly s o i l temperatures were calculated from the recorder charts by integration of the area under the trace of the pen. Maximum and minimum weekly values were read d i r e c t l y from the traces on the charts. A l l of the other physical and chemical properties described below were determined for the samples from the 14-17 September c o l l e c t i o n . Bulk density of a l l samples was determined by oven-drying a subsample and applying a correction for moisture content to the whole sample. Values are reported as bulk density exclusive of coarse fragments (mineral material > 2 mm) and roots of a l l si z e s . Coarse fragments between 2 mm and 10 cm in diameter were separated from organic materials by hand picking and from mineral s o i l by sieving. Their volume was determined by measuring the amount of water they displaced in a graduated cylin d e r . Mean diameters of coarse fragments averaging over 10 cm in diameter were determined in the f i e l d . Their volumes were calculated from these measurements using the equation for the volume of a sphere. 58 S o i l organic matter was calculated by multiplying the carbon content of samples by 1.724 (the conversion factor used in s o i l studies). Carbon was determined by dry combustion in a LECO furnace ( A l l i s o n and others 1965). For determining t o t a l nitrogen, a semimicro Kjeldahl digestion was used in conjunction with a Technicon autoanalyzer. Ammonium saturation (Chapman 1965) was used to determine cation exchange capacity. Hydrogen ion a c t i v i t y was measured with a combination electrode in 1:1 w:v water and 1:2 w:v 0.01 M CaCl 2 (Black 1965). 59 5. RESULTS AND DISCUSSION I n i t i a l examination of the data revealed a c o r r e l a t i o n between dry root mass, length, and surface, area. Regression analysis revealed a strong linear r elationship for the majority of parameters (Table 5.1). The table does not include understory 2-5 mm roots due to l i m i t a t i o n s in the data. For the other root categories the poorest f i t to a straight l i n e was for understory root length and mass (R 2 = 0.68). In these cases the data did not exhibit c u r v i l i n e a r relationships, nor did transformation result in any improvement. Therefore, li n e a r regression was employed in a l l cases. P a r t i c u l a r l y good correspondence to a straight l i n e occurred for overstory root length and mass, and surface area and mass data with R2 equal to or exceeding 0.92. A similar r e l a t i o n s h i p between root mass, length, and surface area has been reported by Persson (1978). As a result of the good co r r e l a t i o n between the parameters measured, the results of more detailed analysis were usually i d e n t i c a l , regardless of the parameter examined. Therefore, only mass data were considered in detailed analysis. A l l of the parameters measured for roots were summarized by plant association in Section 5.3. 60 Table 5 . 1 . R e l a t i o n s h i p s between l e n g t h , su r face a r e a , and mass of r o o t s . Root Category R e l a t i o n s h i p Standard E r r o r of Y R 2 Unsuber ized Length"1" = 2810 Mass* 4.52 0.92 Sur face a r e a * * = 346 Mass 0.455 0.95 Overs tory Length = 802 Mass 461 0.92 < 2 mm Sur face area = 126 Mass 61.2 0.93 Overs tory Length = 28.0 Mass 4.94 0.92 2-5 mm Sur face area = 26.0 Mass 3.24 0.96 Understory Length = 2838 Mass 378 0.68 < 2 mm Sur face area = 2 1 . 6 Mass 2.47 0.85 Length i n cm * Mass i n g * * Sur face area i n cm2 61 Analysis of variance (ANOVA) was to be employed to detect s i g n i f i c a n t differences in fine root mass-volume (o 0.05) between the plant associations, plots, depths, and times and to detect s i g n i f i c a n t interactions between these factors. Two major assumptions of ANOVA are that the data are randomly di s t r i b u t e d and that the variances are homogeneous (Sokal and Rohlf 1969). Neither of these conditions were s a t i s f i e d as ascertained by a X 2 test for goodness of f i t to a normal frequency d i s t r i b u t i o n and B a r t l e t t ' s test for homogeneity of variance. Therefore, the data (Appendix 5) were transformed by taking the logarithm base 10 of each datum to which 1.0 was f i r s t added. Analysis of variance was then c a r r i e d out with the following s t a t i s t i c a l model: ? i j k l " 1 1 + v i + P / V ( i ) j + D k + VDik + P/VD(i)jk + T l + VTil + P / V ^ D j l + D T k l + VDTfki + e ( i ) j k l where * i j k l " variate v = mean e f f e c t Vi = e.ffect of i l e v e l of fixed plant association i = 1,2,3 p / V ( i ) j = e f f e c t of j l e v e l of random plot nested within i l e v e l of plant association j = 1,2,3 Dk = e f f e c t of k l e v e l of fixed depth k = 1, ...,.7 T l = e f f e c t of 1 l e v e l of fixed time 1 = 1,2,3 e(i)jkl s error = p/VDT ( n-)j k l 62 When depth increments could not be included in the case of understory 2-5 mm roots due to many samples without roots in them the model became: Y i j l = y + V i + p / V ( i ) j + T, + V T ^ + £ where the symbols are as explained above except for the missing depth factor and e ( i ) j l = P / V T ( i ) j l Interaction between plant association, depth and time was only s i g n i f i c a n t in understory < 2 mm roots. In the other analyses i t was included in the error term (Table 5.2). The results of the ANOVA w i l l be applied to the appropriate sections to follow. As the f i n a l step in analysis, correlations were sought between the root mass data from the 14-17 September c o l l e c t i o n period and s o i l physical and chemical parameters. These s o i l data were c o l l e c t e d from the cores that were used for sampling the roots. 5.1 Variation of Roots with Depth 5.1.1 Overstory Roots In the ANOVA there was almost a s i g n i f i c a n t interaction of plot within plant association and depth (p = 0 . 0 6 ) for < 2 mm roots and a d e f i n i t e one (/> = 0 . 0 0 3 ) for 2-5 mm roots (Table 5.2). This made i t necessary to consider the depth d i s t r i b u t i o n for each p l o t . In the case of unsuberized roots, the interaction was not s i g n i f i c a n t . This was probably due to Table 5 .2 . P r o b a b i l i t i e s o f F -va lues from ANOVA of log™ root mass-volume. S i t e r e f e r s to p lan t a s s o c i a t i o n and p l o t r e f e r s to r e p l i c a t e s of the p lan t a s s o c i a t i o n s . P r o b a b i l i t i e s not s i g n i f i c a n t a t p = 0.05 are u n d e r l i n e d . Source o f V a r i a t i o n Degrees o f Freedom Tes t i ng Term Overstory Understory Unsuber ized < 2 mm 2-5 mm < 2 mm 2-5 mr S i t e 2 P l o t / S i t e 1.0 0.1 0.1 0.06 0.003 P l o t / S i t e 6 E r r o r 0.02 0.000 0.000 0.000 0.5 Depth 6 P/S x D , 0.000 0.000 0.000 0.000 _* S x D 12 P/S x D 0 .3 0.02 0.05 0.000 -P/S x D 36 E r r o r 0.5 0.06 0.003 0.7 -Time 2 P/S x T 0.003 0.6 0.3 0.9 0.6 S x T 4 P/S x T 0.4 0.7 0.2 0 .8 0.4 P/S x T 12 E r r o r 0 .3 0.006 0.01 0.07 ** D x T 12 E r r o r 0.000 0.6 0.02 0.05 -Error"1" 96 + E r r o r i n c l u d e s SDT except f o r understory < 2 mm roots where SDT was s i g n i f i c a n t (P = 0 . 0 4 ) . * Numerous zero va lues made i t imposs ib le to ca r r y out the ANOVA f o r the depth f a c t o r . * * E r r o r term used i n t h i s ANOVA. the large v a r i a b i l i t y between the three sampling times which was the only 'r e p l i c a t i o n ' of any observation for the samples from a plo t ; (there was no such large v a r i a b i l i t y between sampling times for the remaining categories of overstory roots). The mass-volume of < 2 mm roots f e l l off quickly as depth increased (Fig. 5.1). This trend was weakest for the plots of the OPHO THPL and greatest for those of the ABAM TSME and VAME TSME. The peaks in the 0-5 cm depth increment for the ABAM TSME were markedly d i f f e r e n t , ranging from 3.2 to 7.1 kg n r 3 . Otherwise, variation was much less within depth increments of the r e p l i c a t e plots of the plant association. Roots 2-5 mm in diameter had peaks in mass-volume just below the surface, but occasionally there were peaks at greater depths (Fig. 5.2). Generally, the peaks occurred in the 5-10 cm increment in the VAME TSME plots, between 5 and 20 cm in the ABAM TSME plots, and from 5 to 30 cm in the OPHO THPL plot s . Only the ABAM TSME 1 and VAME TSME 1 curves were similar to one another. As in the case of the roots < 2 mm, the trend was for a more gradual decrease in mass-volume from lower i n i t i a l s u r f i c i a l values on the OPHO THPL plots as compared to those from the other associations. The greatest value occurred in the VAME TSME 2 plot, second highest were both VAME TSME 1 and ABAM TSME 1, while VAME TSME 3 was t h i r d followed by ABAM TSME 65 O V E R S T O R Y < 2 m m ROOT M A S S - V O L U M E (kg m 3 ) 1 2 3 4 5 6 T 3 0 ~ P L O T 1 V A M E T S M E A S S O C I A T I O N 6 0 Figure 5.1. Variation in overstory <2 mm root mass-volume f o r r e p l i c a t e s of the plant associations. Depth begins at the surface of the forest f l o o r . O V E R S T O R Y 2 TO 5 m m ROOT M A S S - V O L U M E (kg m 3 ) 0 . 5 1.0 1.5 2 . 0 2 . 5 V A M E T S M E A S S O C I A T I O N i 1 .5 —I— 2 . 0 2 . 5 PLOT 3-^ J9* A B A M T S M E A S S O C I A T I O N I 0 . 5 1 .0 1 . 5 2 . 0 2 . 5 O P H O T H P L A S S O C I A T I O N l gure 5.2. Variation in overstory 2-5 mm root mass-volume with depth f or r e p l i c a t e s of the plant associations. Depth begins at surface of fo r e s t f l o o r . 67 The values reported in F i g . 5.1 and 5.2 had standard errors that were, sometimes, as large as the mean. These large standard errors were probably due to the var i a t i o n in root d i s t r i b u t i o n from one sample to the next. It might be argued that the large standard errors were because the samples cons t i t u t i n g the replicates for a plot were taken at d i f f e r e n t times. This i s not the case for < 2 mm suberized roots and 2-5 mm roots, since there was no s t a t i s t i c a l l y s i g n i f i c a n t difference between samples taken at d i f f e r e n t times (Table 5.2). There was a s t a t i s t i c a l difference between times in the mass of unsuberized roots per unit volume of s o i l . Therefore, samples of t h i s root category from the three times could not be combined. A similar large v a r i a t i o n between samples in close proximity to one another was noted in a Pinus radiata stand by Bowen (1964). Means for the < 2 mm root mass-volume in the OPHO THPL were lower than those for the corresponding depth in the ABAM TSME and VAME TSME (Table 5.3). Below 10 cm the difference was no longer s i g n i f i c a n t as determined by Duncan's multiple range test (DMRT) on transformed data at a 0.05. Only the means of the 5-10 cm increment were s i g n i f i c a n t l y d i f f e r e n t from the others in the ABAM TSME and VAME TSME with values of 2.9 and 4.2 kg m"3 respectively. Generally, the trend was for the OPHO THPL to have the least rapid decrease in the < 2 mm root mass-volume with increasing depth. The ABAM TSME and VAME TSME associations both had larger i n i t i a l density at the surface with a more rapid decrease with depth than in the OPHO THPL. Table 5 . 3 . V a r i a t i o n in o v e r s t o r y , suber i zed < 2 mm root massr volume w i th depth f o r the p lan t a s s o c i a t i o n s . In a l l cases the mean i s from 9 samples. The standard e r r o r i s i n parentheses . Depth (cm) A s s o c i a t i o n OPHO ABAM VAME THPL TSME TSME kg m - 3 0-5 2.77 + 4 .98 5.14 (0 .97) (2 .98) (1.31 ) 5-10 1.77 + 2.92 + 4.19 (1.35) (1.22) (1.89) 10-20 0.88 1.76 2.31 (0.54) (0.60) (1.16) 20-30 0.65 1.09 1.06 (0.50 ) (0.43 ) ( 0 . 6 3 ) 30-40 0.45 0.69 0.62 (0 .24 ) (0 .39 ) (0 .40 ) 40-50 0.36 0.37 0.40 (0.26 ) (0 .29 ) (0.44) 50-60 0.30 0 .28 0.30 (0 .29 ) ( 0 .26 ) (0 .29) + Transformed va lues are s i g n i f i c a n t l y d i f f e r e n t (Duncan's m u l t i p l e range t e s t , a 0.05) from others w i t h i n t h i s depth. 69 Standard errors were based on 9 samples (three samples taken at di f f e r e n t times from each of the three r e p l i c a t e plots for a plant association). A l l the standard errors were large, tending to increase in re l a t i o n to the mean as depth increased. In the 2-5 mm roots of the OPHO THPL the peak in mass-volume was at 20-30 cm (0.65 kg m"3) gradually decreasing towards the surface and at greater depths (Table 5.4). Maximum density in the ABAM TSME was between 5 and 20 cm (1.3 kg m~3), while i t was between 5 and 10 cm in the VAME TSME (1.8 kg n r 3 ) . S u r f i c i a l values for the roots of the OPHO THPL to a depth of 20 cm were s i g n i f i c a n t l y lower than those of corresponding depths from the other associations based on a DMRT of transformed data at a 0.05. From the trends in the decline of mass-volume with depth shown in Table 5.3 i t i s evident that some roots remained unsampled at the 60 cm depth. As root size increased to 2-5 mm, there was a slower decline in mass-volume with depth (Table 5.4), suggesting also an underestimate in biomass of t h i s size of root. Ash contents of the roots increased considerably from the surface downward. This is probably due to mineral contamination. Actual values are presented in Appendix 6. Differences between percent values for roots < 2 mm growing only in organic matter and those growing in mineral s o i l ranged from 0.05 % in the surface 5 cm (mostly forest floor) to 4.40 % at 50-60 cm in the ABAM TSME (Table 5.5). In contrast, the actual mass-volume values before correction for ash content Table 5 .4 . V a r i a t i o n in ove rs to r y 2-5 mm root mass-volume w i th depth f o r the p lan t a s s o c i a t i o n s . In a l l cases the mean i s from 9 samples. The standard e r r o r i s i n parentheses . Depth A s s o c i a t i o n (cm) OPHO ABAM VAME THPL TSME TSME *g m 0 - 5 0.32 + 0.88 1.06 (0.35) (0.48) (0 .68) 5-10 0.57 + 1.25 1.83 (0.42) (0 .72 ) ( 0 . 6 1 ) 10-20 0.47 + 1.28 1.18 (0.33) ( 0 . 5 8 ) (0.52) 20-30 0.65 0.68 0.64 - ( 0 . 7 0 ) (0 .48 ) (0.61 ) 30-40 0.39 0.40 0.48 (0 .31) (0.32) (0.51) 40-50 0.27 0.15 0.22 (0 .22) (0.15) (0.33) 50-60 0.26 0.32 0.29 (0.27) (0.43) (0.36) Transformed va lues are s i g n i f i c a n t l y d i f f e r e n t (Duncan's m u l t i p l e range t e s t , a 0.05) from others w i t h i n t h i s depth. Table 5 . 5 . E f f e c t o f minera l contaminat ion of overs to ry roots < 2 mm i n diameter on mass-volume and biomass es t imates f o r the three p lan t a s s o c i a t i o n s . . Oplopanaco ( ho r r i d i ) - Ab ie td (amab i l i s) - Vaccin ium (membranacei) -Depth Thujetum p l i c a t a e Tsugetum mertensianae Tsugetum mertensianae (cm) M inera l C o r r e c t i o n * M ine ra l M inera l Co r rec t i on Minera l Minera l Co r rec t i on M inera l Inc luded % Excluded Included % Excluded Included % Excluded 0-5 2 .77* * 0.05 2.77 4. 98 0.05 4.98 5.14 0.08 5.13 5-10 1.77 0.26 1.77 2.92 0.93 2.89 4.19 0.55 4.17 10-20 0.88 0.53 0.88 1.76 2.74 1.74 2.31 1.59 2.27 20-30 0.65 1.33 0.64 1.09 3.76 1.05 1.06 2.12 1.04 30-40 0.45 1.68 0.44 0.69 4.98 0.66 0.62 3.29 0.60 40-50 0.36 2.38 0.35 0.37 4.56 0.35 0.40 1.97 0.39 50-60 0.30 2.10 0.30 0.28 4.40 0.27 0.30 2.26 0.29 To ta l g dm" 3 7.19 7.14 12.09 11.94 14.02 13.89 g dm" 2 492 488 814 801 935 925 C o r r e c t i o n i s d i f f e r e n c e between mean ash content of samples and mean ash content of roots not growing i n minera l s o i l . Percentages are based on ash content o f sample. Unless s p e c i f i e d o the rw i se , a l l va lues are g d m - 3 . 72 were almost the same as the values after correction. If there had been more roots growing in mineral s o i l the impact of contamination would have been greater and a correction l i k e that used by Meyer and Gottsche (1971) for beech roots would have been required. Roots 2-5 mm in diameter had less contamination by mineral p a r t i c l e s than the smaller roots. Therefore, the effect on mass-volume would have been even less than noted for the < 2 mm roots. Thus, i t was unnecessary to correct any roots in t h i s study (including understory) for mineral contamination. Some of the residue remaining after ashing i s due to opaline s i l i c a within the c e l l s . In leaves of some species of the Pinaceae the amount can be as high as 0.18 % by mass (Klein and Geis 1978). No studies on the amount in the c e l l s of roots of the species in t h i s study have been conducted. However, Marchenko and Karlov (1962) commented on the s i l i c a content of fine roots of other species, suggesting that i t may have been due to i n t e r c e l l u l a r deposition, but not discounting s u r f i c i a l mineral contamination. Considering the importance of t h i s mineral contamination to estimates of biomass and nutrient content of roots (Kimmins and Hawkes 1978), more examination of the contribution of opaline s i l i c a to ash content i s needed. 73 Table 5.6 shows the percentage of root biomass for each depth increment based on the biomass in g m - 2 to 60 cm. In a l l plant associations, over 90 % of the unsuberized root biomass was above 30 cm. For suberized < 2 mm roots on ABAM TSME and VAME TSME, over 90 % of the biomass was above 40 cm; on the OPHO THPL the value decreased to 86 % above th i s depth. The difference between unsuberized and suberized roots i s not necessarily s i g n i f i c a n t in terms of water absorption or plant nutrient absorption since these occur in both (Kramer 1946, Kramer and Bullock 1966, Carson 1974, Russel 1977, Wilson and Atkinson 1978). In the 2-5 mm roots, over 90 % of the biomass was above the 50-cm depth in a l l plant associations. In comparison with other coniferous forests, the values found in t h i s study are well above the low of 24 % of the < 6 mm roots above the 30-cm depth in a stand of Pseudotsuga  menziesi i (referred to as P. taxi f o l i a by Reynolds (1970). Generally, the values from t h i s study are 10 to 20 % lower than the upper extreme where 96 % of the < 3 mm roots were above 30 cm in a Picea glauca plantation (Table 5.10 and 2.1). Also, 90 % of the root mass was above 36 to 45 cm in t h i s study while the range for < 2 mm roots in the l i t e r a t u r e was from 23 cm in a Picea glauca and Abies lasiocarpa stand to 125 cm for < 2 mm roots in a Pinus densiflora plantation (Table 2.1). Overall, the values found in t h i s study are intermediate between those reported in the l i t e r a t u r e for coniferous stands. With the additional information from t h i s thesis i t becomes even more Table 5 .6 . Cumulat ive percentage of roo t biomass i n the s o i l f o r the th ree ove rs to ry root ca tego r i es i n the th ree p lan t a s s o c i a t i o n s . Depth (cm) Unsuber ized < 2 mm 2-5 mm OPHO THPL ABAM TSME VAME TSME OPHO THPL ABAM TSME VAME TSME OPHO THPL ABAM TSME VAME TSME 0-5 58 39 68 28 31 28 6 11 12 5-10 69 63 79 46 48 50 18 27 34 10-20 87 87 88 64 70 75 37 60 62 20-30 92 96 94 77 84 86 63 78 77 30-40 94 97 97 86 92 92 79 88 88 40-50 97 99 99 94 97 97 90 92 93 50-60 100 100 100 100 100 100 100 100 100 75 evident that the d i s t r i b u t i o n of roots cannot be extrapolated from one area to another with any degree of confidence. 5 . 1 . 2 Understory Roots The OPHO THPL had the greatest mass-volume ( 0 . 4 9 kg m"3) of < 2 mm roots at the surface, the VAME TSME had less mass-volume of root ( 0 . 3 8 kg m" 3), and the ABAM TSME had the least mass-volume of roots ( 0 . 1 5 kg m~3) (Table 5 . 7 ) . This trend changed in the 5 - 1 0 cm depth increment, with the VAME TSME peaking at 0 . 4 8 kg n r 3 while the OPHO THPL and ABAM TSME associations had declined from the s u r f i c i a l values to 0 . 2 0 and 0 . 0 4 5 kg m"3, respectively. This ranking of associations was maintained in a l l deeper increments, as was the trend for a decline in mass-volume with increasing depth. Due to large standard errors (derived using the method applied for overstory roots), only the s u r f i c i a l values for the OPHO THPL and ABAM TSME associations were s t a t i s t i c a l l y d i f f e r e n t when a DMRT was performed on the transformed data ( a 0 . 0 5 ) . The values from the VAME TSME were s i g n i f i c a n t l y d i f f e r e n t from those of the other associations to a depth of 40 cm and from those of the OPHO THPL to 50 cm. Definite differences between the associations were found as shown by the s i g n i f i c a n t plant association by depth interaction (Table 5 . 2 ) . There was also a s i g n i f i c a n t plant association by depth by time interaction. Table 5 .7 . V a r i a t i o n in unders tory < 2 mm root mass-volume w i t h depth f o r the p l an t a s s o c i a t i o n s . In a l l cases the mean i s from 9 samples. The s tandard e r r o r i s i n parentheses . A s s o c i a t i o n ABAM VAME TSME TSME kg m" 3 0-5 0.487 a * 0.149 b 0.380 c (0 .259) (0 .175) (0 .316 ) 5-10 0.203 a 0.045 b 0.483 c (0 .114) (0 .044) ( 0 .358 ) 10-20 0.118 a 0.041 a 0.267 •b (0.073 ) (0.050) (0.158) 20-30 0.078 a 0.039 a 0.268 b • (0 .087) (0.045) (0.226) 30-40 0.061 a 0.015 a 0.169 b (0 .112) (0.019) (0 .166) 40-50 0.051 a 0.013 ab 0.127 b (0 .099) (0 .017) (0.108) 50-60 0.027 a 0.010 a 0.100 a 0.042 0.019 0.086 * Numbers w i th d i f f e r e n t l e t t e r s f o l l o w i n g them are s i g n i f i c a n t l y d i f f e r e n t (Duncan's m u l t i p l e range t e s t , a 0.05 on t ransformed va lues ) from o thers w i t h i n t h i s depth . No t e s t s were done to compare the d i f f e r e n t depth increments . Depth (cm) OPHO THPL 77 Understory roots 2-5 mm in diameter in the 0-5 cm depth increment were lowest in mass-volume in the ABAM TSME (0.045 kg m"3) which was s i g n i f i c a n t l y d i f f e r e n t (DMRT, o 0.05) from the OPHO THPL (0.31 kg n r 3 ) and VAME TSME (0.50 kg n r 3 ) associations (Table 5.8). At the 5-10 and 10-20 cm depths, only the VAME TSME was s i g n i f i c a n t l y d i f f e r e n t from the others. Below th i s there were no s i g n i f i c a n t l y d i f f e r e n t values. The general trend for 2-5 mm roots in the OPHO THPL was for a decline in mass-volume from the surface downward with mean values intermediate between those of the other associations (Table 5.8). The ABAM TSME mass-volume peaked at 0.11 kg n r 3 in the 5-10 cm increment, then dropped as depth increased. Within any depth increment, the means from the ABAM TSME were always the lowest. In contrast, the greatest means were evident in the mass densities of the VAME TSME (the maximum value was 0.75 kg n r 3 in the 5-10 cm depth increment). Summarizing the data as cumulative percentages of the t o t a l biomass showed that the majority of understory < 2 mm roots were above 40 cm with 89 % in the OPHO THPL and ABAM TSME associations and 83 % in the VAME TSME (Table 5.9). Understory 2-5 mm roots had 89 % of the biomass above 40 cm in the OPHO THPL and 99 % and 95 % above 30 cm in the ABAM TSME and VAME TSME, respectively. Table 5 .8 . V a r i a t i o n i n unders tory 2-5 mm root mass-volume w i th depth f o r the p lan t a s s o c i a t i o n s . In a l l cases the mean i s from 9 samples. The s tandard e r r o r i s i n parentheses . A s s o c i a t i o n ABAM VAME TSME TSME kg m - 3 0-5 0.308 0.045 b 0.502 (0.206) (0.062) (0.375) 5-10 0.154 0.113 0.752 b (0.202) (0.229) (0.645) 10-20 0.085 0.080 0.374 b (0.118) (0.191) (0.320) 20-30 0.087 0.037 0.109 (0.135) (0.071) ( 0 .148 ) 30-40 0.039 0.001 0.040 (0.071) (0 .004 ) (0 .062) 40-50 0.055 0.0 0.015 (0.095) - (0.030) 50-60 0.001 0.0 0.001 0.003 - 0.002 * Transformed va lues fo l l owed by ' b ' are s i g n i f i c a n t l y d i f f e r e n t (Duncan's m u l t i p l e range t e s t , a 0.05) from o thers w i t h i n t h i s depth . No t e s t s were done t o compare the d i f f e r e n t depth increments . Depth (cm) OPHO THPL Table 5 .9 . Cumulat ive percentage of root biomass i n the s o i l f o r the two unders tory root s i z e s i n the th ree p l an t a s s o c i a t i o n s . Depth (cm) < 2 mm 2-5 mm OPHO THPL ABAM TSME VAME TSME OPHO THPL ABAM ' TSME VAME TSME 0-5 36 34 14 31 11 22 5-10 51 45 32 46 40 54 10-20 68 64 52 64 81 86 20-30 80 82 71 81 99 95 30-40 89 89 83 89 100 99 40-50 96 95 93 100 100 50-60 100 100 100 80 5.1.3 Overstory Compared With Understory In the VAME TSME there was a 15 % difference in mass at 30 cm between overstory and understory < 2 mm roots with the understory roots growing proportionally deeper than overstory roots (Table 5.10). There are three possible explanations for t h i s : 1. ' That understory roots are able to exploit a greater volume of s o i l and thus gain an advantage over trees; 2. That understory roots are unable to compete with the tree roots near the surface and are forced to grow deeper to meet their requirements; and 3. That the difference i s due to sampling error. McQueen (1968), in a study of succession from Pinus  s y l v e s t r i s to Fagus s y l v a t i c a , noted that the l a t t e r had smaller diameter roots which, he stated, made i t able to grow deeper, exploit the s o i l more f u l l y and thus become dominant. The shrubs of the VAME TSME also have smaller diameter roots than the trees (0.20 mm on the average as compared to 0.54 mm, respectively), and may thus have an advantage. This tends to support hypothesis 1 above. However, the other hypotheses cannot be discounted. In the ABAM TSME and OPHO THPL there was l i t t l e * d i f f e r e n c e in mass at 30 cm between overstory and understory < 2 mm roots (Table 5.10). In the ABAM TSME there i s a very sparse understory component due to the dense tree canopy. The shrubs present may not be able to r e a l i z e a 81 Table 5 .10 . Changes i n the percentage of roo t mass above 30 cm and depth of 90% of the root mass f o r the root ca tego r i es and p lan t a s s o c i a t i o n s . Sampling depth was 60 cm i n a l l c a s e s . Percentage of Depth o f p l a n t R o o t Mass Above 90% of A s s o c i a t i o n Category 3 0 c m M a s s — cm — VAME TSME ove rs to r y < 2 86 36 2-5 77 45 unders tory < 2 71 47 2-5 95 24 ABAM TSME ove rs to r y < 2 84 38 2-5 78 45 unders tory < 2 82 42 2-5 99 25 OPHO THPL ove rs to r y < 2 77 45 ,. 2-5 63 50 unders tory < 2 80 42 2-5 81 41 o 82 competitive advantage (see hypothesis 1 above), or may not have the required energy to grown deeper (hypothesis 2 above). In the OPHO THPL a high groundwater table probably l i m i t s the growth of both categories of root. Understory 2-5 mm roots were always proportionally more shallow than overstory 2-5 mm roots. This probably r e f l e c t s the size of the plants more than any functional aspects. Instances where the d i s t r i b u t i o n of understory and overstory roots has been j o i n t l y studied are rare. Kimmins and Hawkes (1978) found the d i s t r i b u t i o n of the two categories of roots to be similar in a Picea glauca and Abies lasiocarpa stand in north central B r i t i s h Columbia (Table 2.1). Hausd5rfer (1957) found grasses to be more shallowly rooted than trees (Table 2.1). 5.2 Changes in V e r t i c a l D i s t r i b u t i o n and Root Biomass With  Time 5.2.1 Overstory Root V e r t i c a l D i s t r i b u t i o n As was mentioned e a r l i e r in Section 5.1, unsuberized roots varied s i g n i f i c a n t l y with time. In addition, there was a s i g n i f i c a n t depth by time interaction (Table 5.2). Examination of the data from individual plant associations showed that unsuberized roots grew primarily near the surface with v i r t u a l l y no change below 40 cm during the growing season (Fig. 5.3). Changes above 5 cm were the greatest, r e f l e c t i n g the importance for roots of the surface s o i l layers (mostly U N S U B E R I Z E D ROOT M A S S - V O L U M E (kg m " J ) 0.01 0.02 0.03 30 18-21 J U L Y 2-9 J U N E f V A M E T S M E A S S O C I A T I O N 60 0.01 0.02 —I 0.03 l 60 A B A M T S M E A S S O C I A T I O N I 0.01 0.02 0.03 30 •Pi > f 18-21 JULY vj-14-17 S E P T . 2-9 J U N E 60 O P H O T H P L A S S O C I A T I O N l Figure 5.3. V a r i a t i o n i n unsuber ized root mass-volume w i th t ime f o r the th ree p lan t a s s o c i a t i o n s . Depth begins at the su r face of the f o r e s t f l o o r . 84 forest floor) in these plant associations. The differences between the plant associations that are evident in Figure 5.3 w i l l be discussed in the following section (5.2.3) on overstory root biomass Other categories of roots were not considered here. Suberized < 2 mm root biomass did not vary s t a t i s t i c a l l y between the sample times. Also, there was no s t a t i s t i c a l l y s i g n i f i c a n t interaction between the time factor and the other factors. The s i g n i f i c a n t interaction between depth and time for the 2-5 mm roots i s probably due to random var i a t i o n between plots rather than to a real change since one would not expect the biomass of these larger roots to change s i g n i f i c a n t l y during one growing season. 5.2.2 Understory Root V e r t i c a l D i s t r i b u t i o n In view of the marginally s i g n i f i c a n t interaction between plant association, depth, and time in the case of < 2 mm understory roots i t was no surprise that some unusual trends were evident in the data. Among these were the peak in the 2-9 June sample from 20-30 cm in the VAME TSME, the high s u r f i c i a l values from the 2-9 June sample as opposed to the other times in the ABAM TSME (Fig. 5.4). These unusual trends may be due to random variation between plots since sampling errors were almost always as large as the mean. On the other hand, the high values noted in the 2-9 June sample may also be the result of root growth before 2-9 June or in the previous f a l l or U N D E R S T O R Y < 2 m m ROOT M A S S - V O L U M E (kg m ) 0.1 0.2 0 .3 0 . 4 0.5 0.6 3 0 6 0 2 - 9 J U N E ^ \ 14-17 S E P T . . . • • / " 18-21 JULY V" V A M E T S M E A S S O C I A T I O N I 0.1 0.2 0 .3 0.4 0 .5 0 . 6 E o CL UJ Q 3 0 6 0 / ; J 2 - 9 J U N E - v • • • -j^/s 1 4 - 1 7 S E P T . J M Ii a'. 1 8 - 2 1 JULY A B A M T S M E A S S O C I A T I O N 0.1 0.2 3 0 6 0 18 -21 J U L Y I 2 - 9 J U N E 1 4 - 1 7 S E P T O P H O T H P L A S S O C I A T I O N I F igu re 5.4. V a r i a t i o n i n understory < 2 mm roo t mass-volume w i th t ime f o r the th ree p lan t a s s o c i a t i o n s . Depth begins at the su r face of the f o r e s t f l o o r . 86 winter. The lack of any d i s t i n c t decline in values from the 18-21 July sample to the 14-17 September sample in the VAME TSME and ABAM TSME tends to support t h i s hypothesis. Also, in the surface of the OPHO THPL the 14-17 September sample has the highest value of the three sampling times (Fig. 5.4). It i s also unusual that the curves from 14-17 September do not show the high values evident at greater depths in the 2-9 June sample in the OPHO THPL (Fig. 5.4). Assuming root growth in the previous f a l l or winter t h i s probably i s in response to the greater temperatures deeper in the s o i l after 14-17 September as compared to those at the surface (Table 5.21). The lack of any records prior to the f i r s t sample and after the time of the la s t sample makes the above interpretations regarding root growth before 2-9 June or in the previous f a l l or winter hypothetical. Limitations in the data similar to those for the overstory 2-5 mm roots also apply to the understory roots of th i s s i z e . Thus, their depth d i s t r i b u t i o n was not examined. 5.2.3 Overstory Root Biomass In the previous section i t was necessary to use root mass-volume to compare the uneven depth increments. Since depth increments are no longer being compared units of biomass per unit area (the unit commonly used in ecosystem studies) w i l l be used. 87 Unsuberized roots grew d i f f e r e n t l y in the VAME TSME and ABAM TSME as compared to the OPHO THPL (Fig. 5.5). In the case of the former two associations, there was a s t a t i s t i c a l l y s i g n i f i c a n t peak (DMRT a 0.05 on transformed data) in the 18-21 July sample while there was none in the l a t t e r . Between the 18-21 July sample and 14-17 September sample, there was an increase in the OPHO THPL while there were decreases in the other associations. This difference may be a response to a reduction in root growth in the presence of high levels of groundwater in the OPHO THPL during the f i r s t two sampling periods. Figure 5.5 also shows that unsuberized roots were present in the 2-9 June sample. There are three possible explanations for t h i s : 1. Sampling was not early enough to precede root growth; 2. Root growth i s continuous throughout the winter; 3; Unsuberized roots growing in the f a l l are dormant during the winter but remain white. In r e l a t i o n to the f i r s t p o s s i b i l i t y , sampling began at least two weeks prior to shoot growth. Perhaps t h i s was s t i l l not early enough since studies in Germany of several coniferous species (not those of the present study, however) have shown signs of root growth considerably before shoot growth (Hoffmann 1972). In re l a t i o n to the second p o s s i b i l i t y , other investigators (Reed 1939, Heikurainen 1957a, Vogt and others 88 6 0 0 CM 10 10 < O CD o O cc 3 0 0 2 0 0 1 0 0 O V E R S T O R Y 2- 5 m m • • **. V A M E T S M E ^ A B A M T S M E * %x O P H O T H P L . - -2 - 9 J U N E 18-21 J U L Y 14-17 S E P T . F igu re 5.5. V a r i a t i o n i n ove rs to ry roo t biomass w i th t ime f o r the th ree p l an t a s s o c i a t i o n s . 89 1980) have noted limited root growth throughout the winter during periods when the temperature was low but the ground was unfrozen. In re l a t i o n to the t h i r d p o s s i b i l i t y , white roots have been observed by others (Head 1966, Wilcox 1968, Kozlowski 1979), but upon examination these were inactive, yet had remained white due to the slowness of browning. It i s impossible to ascertain which of the above possible explanations applies in the present study. In spite of the obvious large differences between the plant associations in terms of temporal changes in unsuberized root biomass (Fig. 5.5), yet there was s t i l l a s i g n i f i c a n t time e f f e c t . There was no s i g n i f i c a n t interaction between plant association and time (probably the interaction shown for the VAME TSME and the ABAM TSME was masked by the lack of an interaction in the OPHO THPL). Also, there was no s i g n i f i c a n t interaction between plot-within-plant-association and time (Table 5.2). Increases in the mean biomass of < 2 mm roots of the overstory vegetation were very s l i g h t (Fig. 5.5). It i s unlikely that these arose from suberization of new roots, since the biomass of the unsuberized roots was p r a c t i c a l l y inconsequential in r e l a t i o n to the biomass of the suberized < 2 mm roots. It i s more l i k e l y that the minor increases between 18-21 July and 14-17 September in the OPHO THPL and between 2-9 June and 18-21 July in the VAME TSME (Fig. 5.5) were due to random v a r i a t i o n . 90 The declines between 2-9 June and 18-21 July in the OPHO THPL and between 18-21 July and 14-17 September in the ABAM TSME could also be ascribed to random v a r i a t i o n . However, the declines could have been due to growth of < 2 mm roots into the 2-5 mm size c l a s s . The decrease in root biomass between 2-9 June and 18-21 July in the OPHO THPL may have been a result of root death due to lack of oxygen caused by a high water table. The decrease in the ABAM TSME may have been due to decreased nutrient a v a i l a b i l i t y or some other environmental stress. With the limited r e p l i c a t i o n for a p a r t i c u l a r sample in thi s study i t i s impossible to make any d e f i n i t e statements regarding the above hypothesis about decreases in root biomass. In the 2-5 mm root category i t i s more l i k e l y that the fluctuations were largely due to random variation between plots; however, the arguments presented for the < 2 mm roots also apply to th i s category. In the < 2 mm and 2-5 mm root categories there was no si g n i f i c a n t plant association by time interaction for either root category (Table 5.2). On the other hand, the plot-within-plant-association by time interaction was s i g n i f i c a n t , emphasizing the s i g n i f i c a n t difference between plots (Table 5.2). As mentioned previously, the time effect was not si g n i f i c a n t for either of these root categories. 91 Recently, root productivity has been estimated by summing the increments in fine root biomass from numerous samples taken over the course of a year (Harris and others 1977). This should not be done with unsuberized root biomass data from t h i s study for the following reasons: 1. Sampling was not frequent enough and not over the course of a f u l l year, making i t unlikely that maximum and minimum values have been determined; 2. New growth may already have been p a r t i a l l y suberized in the 18-21 July sample; and 3. Death and predation may have occurred (recently Santantonio (1978) and Persson (1979) have shown that death i s an important f a c t o r ) . 5.2.4 Understory Root Biomass Biomass of roots < 2 mm in diameter changed l i t t l e from one sampling time to the next except for one minor decrease from 2-9 June to 18-21 July in the OPHO THPL (Fig. 5.6). Thus, i t was not surprising to find no s i g n i f i c a n t plant association by time interaction nor any time effect (Table 5.2). Simonovic (1973) measured r e l a t i v e l y small changes in biomass during the year in an oak-hornbeam forest. In A p r i l the underground herb-layer biomass was 22.9 g m"2, i t was 17.6 g n r 2 in June, and 23.1 g n r 2 in September. The most intensive root growth followed flowering or the ripening of seeds. It i s interesting to note the s i m i l a r i t y between his work and the 92 ' E 1 2 0 cn to to < 8 0 g CO 8 4 0 U N D E R S T O R Y <2 m m V A M E T S M E J / - O P H O T H P L A B A M T S M E 2 - 9 J U N E 1 18 -21 J U L Y 1 6 0 r 120 UNDERSTORY 2"5 m m - V A M E T S M E - / - O P H O T H P L ^ - ' • - — - A B A M T S M E . • 14-17 2 - 9 S E P T . J U N E 18-21 JULY 14-17 S E P T . Figure 5 .6 . Variation in understory root biomass with time for the three plant associations. 93 trend in the OPHO THPL (Fig. 5.6). It was only in t h i s association that flowering and ripening of seed occurred at the end of the growing season in most of the plants. In contrast, the shrubs of the other associations flowered early in the spring and seeds ripened throughout the growing season. The absolute amounts of biomass were much higher in the present study than in the oak-hornbeam forest. No other l i t e r a t u r e was discovered wherein the seasonal changes in understory root biomass were measured. Fluctuations from one time to another in the mean biomass of 2-5 mm roots (Fig. 5.6) were probably due to var i a t i o n between cores since they were 1 m apart in the plot s . With such va r i a t i o n there was no s i g n i f i c a n t plant association by time interaction and no time e f f e c t in the ANOVA. Considering the lack of any s i g n i f i c a n t interactions, i t was possible to combine the biomass from the three sampling times together to form a single estimate for each plant association. The results thus obtained w i l l be discussed in Section 5.4. It was mentioned for the overstory roots that the res u l t s should not be used to estimate productivity. This applies also to the understory roots. This i s especially so since i t was impossible to separate the current annual growth from that of previous years. 94 5.3 Variation in Root Biomass Between Plots Comparison (based on transformed data) of individual plots by DMRT revealed that in several instances the biomass of roots from plots within a plant association was not s i g n i f i c a n t l y d i f f e r e n t , e.g. overstory < 2 mm roots of the ABAM TSME plots (Table 5.11). In other cases there was a difference e.g. plot 2 of the VAME TSME in the overstory < 2 mm root category. Considering the results of the DMRT, i t appeared that the magnitude of the differences between plots within the plant associations were not as large as the prob a b i l i t y recorded in the ANOVA (Table 5.2) suggested. On the other hand, there i s some j u s t i f i c a t i o n for a separate analysis for each p l o t . Considering that only a few of the plots were s i g n i f i c a n t l y d i f f e r e n t within a plant association and that the objective of this thesis was to compare plant associations, the decision was made to deal with plant associations rather than individual plots in the following sections. A DMRT was not car r i e d out for the mass-volume of unsuberized roots since the error term would have been i n f l a t e d due to the s i g n i f i c a n t temporal v a r i a t i o n . Table 5 .11 . Resu l t s o f Duncan's M u l t i p l e Range Test on t ransformed ( l o g i o ( x + 1)) root mass-volume ( k g m " 3 ) . Bars connect po in t s not s i g n i f i c a n t l y d i f f e r e n t at a 0 .05 . For r e f e r e n c e , the untransformed means on a u n i t area bas i s are repo r ted . Due t o the presence o f o c c a s i o n a l 0 va lues i n the d a t a , the order o f means do not always agree w i th the t ransformed order i . e . they do not always decrease to the bottom o f the page l i k e the t ransformed va lues d i d . Standard e r r o r s o f the means are in paren theses . The 0 , A , and V symbols represent the OPHO THPL, ABAM TSME, and VAME TSME a s s o c i a t i o n s r e s p e c t i v e l y . Overs tory Understory Unsuber ized < 2 mm 2-5 mm < 2 mm 2-5 mm _2 g.m (60 cm depth) S3. 0 . 2 5 ( 0 . 0 6 ) ££ 0 . 6 9 ( 0 . 5 8 ) A3 0 . 7 4 ( 0 . 5 2 ) VJL 0 . 7 9 ( 0 . 5 4 ) V2 0 . 8 2 IK0.99) A l 0 . 9 9 to.51 ) A2 11.38 t l . 2 4 ) V3 1 .20 : i . 2 i ) 1 .88 :i .05) 03 280 (97) 82_ 490 (200) V2_ 660 (91) 8JL 710 (100) A3 740 (160) El 840 (270) A2 850 (220) 13 990 (250) vi 11200 (180) S3 90 (40) 02 300 (140) A3 310 (67) QJ_ 360 (150) A l 380 (60) V2 390 (56) VI 430 K 1 4 0 ) A2 480 (76) V3 1470 1340) A2 9 (13) AJ_ 19 (2) A3 37 (19) 02 36 (12) 01 59 (21) V2 66 (17) 03 n o (53) 11 120 (25) V.3 270 (35) A2 3.6 (6) A3 28 (27) A± 27 (24) 31 29 (18) 02 48 (29) _G3 72 (8) 12 92 (67) 13 1118 1(57) VI I TOO (43) 96 5.4 Variation in Root Biomass Between Plant Associations There was a consistent increase in biomass of the overstory < 2 mm and 2-5 mm roots from the OPHO THPL to the ABAM TSME and then to the VAME TSME (Fig. 5.7). In both the length and surface area measurements, the OPHO THPL had the lowest values while the ABAM TSME and VAME TSME had very similar values. In contrast, the unsuberized roots had similar biomass values in a l l associations (Table 5.12). For lengths and surface areas of t h i s root category, the OPHO THPL again had the lowest values while the remaining associations had similar values (Tables 5.13 and 5.14). As was mentioned e a r l i e r , there i s a strong relationship between biomass, length and surface area of roots within a pa r t i c u l a r size category. Thus, any of the above may be applied for comparative purposes. However, i f a pa r t i c u l a r size category of roots i s to be compared to roots of other size categories, the choice of unit of measurement i s c r i t i c a l . For example, in this study the r a t i o of roots 2-5 mm in diameter to those < 2 mm in the ABAM TSME was 0.48, 0.19, and 0.10 for biomass, length, and surface area, respectively (Tables 5.12, 5.13, 5.14). These tables also show the insig n i f i c a n c e of the unsuberized root category, e.g. ratios in the ABAM TSME of unsuberized roots to those < 2 mm were 0.0012, 0.0054, and 0.0035 for biomass, length, and surface area, respectively. I 6 0 0 r 1400r-1 2 0 0 h-CU 2 - 5 1 0 0 0 h - . U 2 - 5 800-6 0 0 -4 0 0 200 0 U<2 O 2-5 0<2 [ LU <2 0 2 - 5 0<2 U 2-5 U<2 0 2-5 0<2 OPHO ABAM VAME THPL TSME TSME BIOMASS ( g m " 2 ) 12 10 8 t—' U <2 2-5 U<2 U<2 02-5) rO 2-5 r 0 2-5 0<2 O <2 0<2 OPHO ABAM VAME THPL TSME TSME LENGTH (km m"2) 16r-1 4 h -r U 2 - 5 12 10 8 h ^-U2-5 U <2 r U 2 - 5 2 h -U<2 O 2-5 0<2 0 2 - 5 O <2 U 2-5 U<2 O 2 -5 0 < 2 OPHO ABAM VAME THPL TSME TSME SURFACE AREA ( m 2 m - 2 ) F i g u r e 5.7. Summary by p l a n t a s s o c i a t i o n of b i o m a s s , l e n g t h , a n d s u r f a c e a r e a f o r t h e v a r i o u s s i z e s and c a t e g o r i e s of o v e r s t o r y (0) and u n d e r s t o r y (U) r o o t s . Table 5 .12 . Comparison between ove rs to ry and unders tory root biomass f o r the th ree p lan t a s s o c i a t i o n s . Means are based on 3 p l o t s . The standard e r r o r s are i n paren theses . Unsuber ized (uns) roo ts are separated from the < 2 mm r o o t s . A s s o c i a t i o n S i z e Overs tory Understory U/0 -mm- g m - 2 VAME TSME uns 0.94 (0.23) < 2 940 (256) 136 (77) 0.145 2-5 430 (40) 117 (24) 0.272 ABAM TSME uns 1.0 (0 .3) < 2 810 (61) 22 (14) 0.0272 2-5 390 (87) 20 (14) 0.0513 OPHO THPL uns 0.94 (0.85) < 2 490 (217) 68 (37) 0.139 2-5 250 (140) 50 0.200 Table 5 .13 . Comparison between ove rs to ry and unders tory root leng ths f o r the th ree p lan t a s s o c i a t i o n s . Means are based on 3 p l o t s . The standard e r r o r s are i n paren theses . Unsuber ized (uns) roo ts are separated from the < 2 mm r o o t s . A s s o c i a t i o n S i z e Overs tory Understory U/0 -mm- km m VAME TSME uns 0.030 (0.006) < 2 6 .4 (1 .8) 4 .7 (2 .3 ) 0.734 2-5 0.12 (0.02) 0.027 (0.008) 0.225 ABAM TSME uns 0.034 (0.011) < 2 6.3 (0 .6) 0.67 (0.45) 0.106 2-5 0.12 (0.02) ' 0.0042 (0.0028) 0.0350 OPHO THPL uns 0.030 (0.006) < 2 3.7 (1 .3 ) 2.0 (1 .0 ) 0.541 2-5 0.080 (0.040) 0.019 (0.009) 0.238 100 Table 5 .14. Comparison between ove rs to r y and unders tory root su r face areas f o r the th ree p lan t a s s o c i a t i o n s . Means are based on 3 p l o t s . The standard e r r o r s are i n parentheses . Unsuber ized (uns) roo ts are separated from the < 2 mm r o o t s . A s s o c i a t i o n S i z e Overs tory Understory U/0 -mm- m 2 m" 2-VAME TSME uns 0.036 (0.005) < 2 10.8 (2 .9) 2.9 (1 .4) 0. 269 2-5 1.1 (0 .2) 0.26 (0.06) 0. 236" .ABAM TSME uns 0.037 (0.012) < 2 10.6 (0 .9 ) 0.44 (0.30) 0. 0415 2-5 1.09 (0.21) 0.044 (0.029) 0. 0404 OPHO THPL uns 0.028 (0.024) < 2 6.6 (2 .3) 2.2 (1 .0 ) 0. 333 2-5 0.74 (0.38) 0.18 (0.08) 0. ,243 101 In contrast to the overstory roots, understory roots had the greatest values of root biomass, length, and surface area in the VAME TSME, intermediate values in the OPHO THPL, and the lowest values in the ABAM TSME (Fig. 5.7). The rel a t i o n s h i p between the < 2 mm and 2-5 mm roots for biomass, length, and surface area was similar to that noted above for the overstory roots. Standard errors of the means of biomass for the three replicates sampled within an association were least in the < 2 mm overstory roots of the ABAM TSME and greatest in the 2-5 mm understory roots of the OPHO THPL (Table 5.12). Using the general relationship for a normally d i s t r i b u t e d population, the number of samples necessary for obtaining a standard error that i s within 10 % of the mean (Cochran and Cox 1957) i s 3 in the case of overstory < 2 mm roots from the ABAM TSME and almost 150 in the case of the 2-5 mm roots from the same association. In most instances, three samples were only adequate to give a general idea of the magnitude of sampling that would be required. Understory roots contributed r e l a t i v e l y l i t t l e to the combined biomass of overstory and understory categories. In contrast, the length and surface area values shown in F i g . 5.7 and Tables 5.12, 5.13, 5.14 indicate a greater importance of understory roots. On the VAME TSME, ratios of understory to overstory for roots < 2 mm, for example, were 0.145, 0.734, and 0.269 for biomass, length, and surface area, respectively. On the ABAM TSME Association, the rati o s were the lea s t , namely: 102 0.0272, 0.106, and 0.0415 for biomass, length, and surface area, respectively. Assuming that the surface area of fine roots r e f l e c t s t h e i r a b i l i t y to absorb moisture and nutrients and that d i f f e r e n t sizes of roots have equal absorptive capacity, the < 2 mm roots are r e l a t i v e l y much more important than those that are 2-5 mm in diameter (Fig. 5.7). Shea (1973) reported similar findings in jack pine in Ontario with < 1 mm roots constituting 80-90 % of the t o t a l horizontal root surface area. Also, assuming understory roots have the same absorptive capacity as overstory roots given similar diameter classes and surface areas, the understory component of the VAME TSME and OPHO THPL i s a strong competitor with the trees for moisture and nutrients in spite of the apparent dominance of the trees above the ground. Sohlenius and others (1977) have noted lessened rooting of Pinus s y l v e s t r i s under clones of Calluna vulgaris due to allelopathy of substances released from heather plants and l i t t e r . Allelopathy between understory species and tree species in these associations has not been studied but could be a factor in addition to direc t competition. Values of root biomass reported in the l i t e r a t u r e for coniferous forests range from 1.0 (< 2 mm roots) to 1810 (< 5 mm roots) g m~2 for overstory roots (Table 2.2). In t h i s study, the values for roots £ 5 mm were between 740 and 1320 g n r 2 . Lengths reported in the l i t e r a t u r e range from 0.075 (<, 2 mm) to 7,.7 (< 6 mm) km n r 2 (Table 2.3) and for roots ^ 5 mm in thi s study they were from 3.8 to 6.5 km m~2. Surface area was 103 only reported in two studies and the values ranged from 0.1 to 4.5 m2 nr 2 for < 1 mm roots (Persson 1978), somewhat lower than in this study (7.3 to 11.9 m2 n r 2 ) . The number of root t i p s was only found in one study, which reported them to vary between 4.4 and 112.2 x 10 3 n r 2 (over one year) (Kalela 1950). In t h i s study there were between 4.2 and 8.0 x 105 t i p s n r 2 . Understory roots have also received l i t t l e study. In the l i t e r a t u r e their biomass i s reported to vary between 70 and 790 g n r 2 ; in comparison the range here was 42 to 253 g n r 2 for roots ^ 5 mm. In one study done in a plantation, the length of roots < 2 mm ranged from 0.0 to 0.0071 km n r 2 , considerably below the values found in t h i s study (0.67 to 4.7 km n r 2 for < 2 mm roots). No comparisons for surface area are possible due to the absence of available data. 5.5 Ratio of Overstory Root Biomass to Overstory Above-Ground  Biomass In previous sections, the differences in overstory root biomass could have been a result of edaphic differences between the plant associations; however, they could also have arisen because of differences in above-ground biomass of the overstory vegetation. By c a l c u l a t i n g r a t i o s of fine roots of trees to above-ground tree (shoot) biomass (see Table 3.2 for data) the effect of d i f f e r e n t above-ground tree biomass can be eliminated. Thus, there were almost twice as many < 2 mm roots per unit of shoot mass on the VAME TSME (0.021) as on the OPHO 104 THPL (0.011), with the ABAM TSME being intermediate (0.014) (Table 5.15). Very l i t t l e difference was noted in the ra t i o s for unsuberized roots (range from 1.8 x 10"S to 2.2 x 10~ 5), the 2-5 mm roots showed almost the same trend as the < 2 mm roots but had considerably reduced rat i o s o v e r a l l (0.0095, 0.0067, and 0.0056 forVAME TSME, ABAM TSME, and OPHO THPL associations, respectively). Results of t h i s study f a l l within the range of values reported in the l i t e r a t u r e (Karizumi 1974b and summarized in Table 2.2). However, in the l i t e r a t u r e the r a t i o of root biomass to shoot biomass for < 2 mm roots was generally lower than for 2-5 mm roots. This i s the opposite of what was found in the present study. Since the vast majority of studies reported in the l i t e r a t u r e were done in young stands the difference may be due to stand age. Tree biomass above the ground includes a large amount of dead material in the stem. Foliage biomass was thought to be more clos e l y related to the roots studied. Greater foliage mass would require proportionally more root mass, other factors remaining unchanged. If t h i s r elationship did not hold, then extraneous factors such as climate or edaphic features or internal factors such as genetics would have to be considered as influencing root mass. 105 Table 5 .15 . Ra t i os of biomass of unsube r i zed , < 2 mm, 2-5 mm, and < 5 mm overs to ry roo ts to who le - t ree biomass f o r the th ree p l an t a s s o c i a t i o n s . Means are based on th ree p l o t s , Standard e r r o r s are i n parentheses . P lan t . Overs tory Root Category A s s o c i a t i o n Unsuber ized < 2 mm 2-5 mm To ta l VAME TSME 2.1 x 1 0 " 5 0.021 0.0095 0.031 (0.3 x 1 0 " 5 ) (0.008) (0.0013) (0.009) ABAM TSME 1.8 x 1 0 " 5 0,014 0.0067 0.021 (0.8 x 1 0 " 5 ) (0.003) (0.0023) (0.005) OPHO THPL 2.2 x 1 0 " 5 0.011 0.0056 0.017 (2.1 x 1 0 " 5 ) (0.005) (0.0033) (0.009) 106 Data on foliage biomass were not available, since t h i s component was lumped with twigs (Krumlik 1979). These combined data plotted against mass of < 2 mm roots (including the unsuberized category in t h i s case) show a weak trend for more roots per unit of foliage plus twig in the VAME TSME than in the other associations (Fig. 5.8). The trend for a high r a t i o of < 2 mm and 2-5 mm root biomass to foliage biomass in the VAME TSME was more apparent when the mean of the three plots in t h i s association was compared to the means from the plots in the ABAM TSME and the OPHO THPL (Table 5.16). The ratios for these root categories from the ABAM TSME and the OPHO THPL were s i m i l a r . In contrast, the ratios involving unsuberized roots increased from the ABAM TSME, to the VAME TSME, and to the OPHO THPL. A l l of the above results are comparable to those for root-to-shoot ratios described above. The large standard errors r e l a t i v e to the mean values for the associations makes any s t a t i s t i c a l l y based conclusions impossible. 107 200 400 600 800 1000 BIOMASS O F . OVERSTORY <2 mm R O O T S ( g m " 2 ) F igure 5 .8 . R e l a t i o n s h i p between f o l i a g e p lus twig biomass and ove rs to ry < 2 mm suber i zed p lus unsuber ized root biomass. L e t t e r s represen t the f o l l o w i n g : 0 = OPHO THPL, A = ABAM TSME, V = VAME TSME. Numbers represent p l o t s w i t h i n the a s s o c i a t i o n s . 108 Table 5 .16 . Ra t ios of r o o t - t o - f o l i a g e p lus tw ig biomass f o r the th ree p l an t a s s o c i a t i o n s . In a l l cases the mean i s from 3 samples. The standard e r r o r i s i n parentheses . Root Category < 2 mm 2-5 mm To ta l 0.37 (0.14) 0.17 (0.02) 0.54 (0.16) 0.24 (0.05) 0.11 (0.04) 0.35 (0 .08) 0.22 (0.09) 0.11 (0.06) 0.33 (0.15) P l a n t A s s o c i a t i o n Unsuber ized VAME TSME 3.6 x 10" (0.2 x 10" 1 1 ) ABAM TSME 3.1 x 10"* (1.2 x 10" " ) OPHO THPL 4.1 x 10" (3 .4 x 10 ) 109 5.6 S o i l Physical and Chemical Properties 5.6.1 Results of Analysis Bulk density tended to increase with increasing depth in a l l associations; the lowest value recorded was 0.12 g cm"3 in the OPHO THPL at 0-5 cm while the highest was 0.68 g cm"3 in the ABAM TSME at 50-60 cm (Table 5.17). The values at greater depths were intermediate between those that might be found for ty p i c a l organic matter and for mineral s o i l because at least one of the four cores making up a sample usually consisted e n t i r e l y of decaying wood (Appendix 7 shows diagrams of the cores taken). This affected most of the properties discussed below as well. Also, v a r i a b i l i t y tended to increase with increasing depth, although t h i s trend was not consistent. Coarse fragment volume as a percentage of sample volume was greatest at 20-30 cm in the OPHO THPL. It increased with increasing depth in the ABAM TSME and VAME TSME, and also from the OPHO THPL to the ABAM TSME and to the VAME TSME (Table 5.17). Values ranged from zero near the surface of the OPHO THPL to 53 for the VAME TSME at 50-60 cm. Considerable v a r i a b i l i t y was evident in the values due to var i a t i o n between plots within the associations. Table 5.17. Bulk d e n s i t y and coarse fragment content of the s o i l s taken from 14-17 September 1977 separated by p l a n t a s s o c i a t i o n and depth . In a l l cases the mean i s from 3 samples. The standard e r r o r i s i n parentheses . _ i i r — Bulk Dens i ty Coarse Fragments Depth From 3 Sur face of 0 p H 0 /\BAM VAME OPHO ABAM VAME Fores t F l o o r T H p | _ j$ME TSME THPL TSME TSME „ „ _3 — % of volume cm — g cm 0-5 5-10 20-30 30-40 40-50 50-60 0.12 0.18 0.13 0 3 5 (0 .01) (0 .01 ) (0 .02 ) (0) (4) (9) 0.15 0.24 0.24 0 2 8 (0 .01) (0 .09 ) (0 .17 ) (0) (1) (13) 0.19 0.25 0.27 2 3 10 1 0 " 2 0 (0.06) (0 .04 ) (0 .06 ) (3) (1) (7) 0.21 0.42 0.38 12 6 25 (0.04) (0 .09) (0 .16) (16) (4) (25) 0.33 0.60 0.39 9 17 32 (0.04) (0 .14) (0 .26) (12) (12) (24) 0.37 0.52 0.37 5 32 42 (0.11) (0 .16) (0 .30) (4) (22) (31) 0.49 0.68 0.41 4 43 53 (0.10) (0 .26 ) (0 .20 ) (0) (25) (40) 111 The reader may have noted that the above results (Table 5 . 1 7 ) are di f f e r e n t from those determined using modal s o i l p i t s (Table 3 . 3 ) . The reason for these differences i s probably the decaying wood content referred to e a r l i e r ; however, since d i f f e r e n t sample sizes are involved in core and p i t sampling, a parameter by parameter comparison i s not j u s t i f i e d . Organic matter content varied l i t t l e with depth in the OPHO THPL (Table 5 . 1 8 ) . In the top 5 cm i t was 88 kg n r 3 , at 5 0 - 6 0 cm i t was 107 kg m"3. In the ABAM TSME the surface 0 - 5 cm value was 126 kg m"2 while at 5 0 - 6 0 cm i t was 51 kg n r 2 . The lowest 0 - 5 cm value of 64 kg n r 2 occurred in the VAME TSME with only 19 kg n r 2 at 5 0 - 6 0 cm. Standard errors were large, almost equalling the mean in magnitude at the 5 0 - 6 0 cm depths. Nitrogen followed the same trend as organic matter. It was 1 .54 g m - 3 in the surface 5 cm in the OPHO THPL and 2 . 3 6 g n r 3 at 5 0 - 6 0 cm (Table 5 . 1 8 ) . In the ABAM TSME corresponding values were 2 . 1 8 and 1 .07 g m"3, respectively. The values were 1 .36 g n r 3 at 0 - 5 cm and 0 . 3 2 g n r 3 at 5 0 - 6 0 cm in the VAME TSME. Again, the standard errors were large in r e l a t i o n to the means. Carbon-to-nitrogen (C/N) rati o s were between 40 and 25 in the OPHO THPL with the lower values below 40 cm (Table 5 . 1 8 ) . The values in the ABAM TSME varied between 47 and 2 6 . The highest value was in the 1 0 - 2 0 cm depth increment; the lower value was at 5 0 - 6 0 cm. In the VAME TSME the greatest r a t i o ( 4 6 ) was in the 5 - 1 0 cm depth increment. The r a t i o at the surface ( 0 - 5 cm) was 27 while at 5 0 - 6 0 cm i t was 2 5 . Standard 112 Tab le 5 .18 . Organic ma t te r , n i t r ogen and c a r b o n - t o - n i t r o g e n r a t i o s o f the s o i l s taken from 14-17 September 1977 separated by p l a n t a s s o c i a t i o n and depth . In a l l cases the mean i s from 3 samples. The s tandard e r r o r i s i n paren theses . o o . u r - Organic Mat te r Depth From a  Sur face o f Q p H 0 A B A M V A M E Fores t F l o o r T H p | _ T $ M E T $ M E Ni t rogen OPHO THPL ABAM TSME VAME TSME C/N Ra t i o OPHO THPL ABAM TSME VAME TSME cm kg m" 0-5 88 126 64 1.54 2.18 1.36 40 34 27 (10) (6) (20) 0.31 (0.36) (0 .29) (5) (4) (6) 5-10 98 90 86 1.82 1.42 1.12 33 37 46 (2) (26) (12) (0.46) (0.50) (0.20) (10) (7) (15) 10-20 103 66 85 1.82 0.86 1.20 ' 33 47 42 (21) (23) (6) (0.15) (0.27) (0.25) (9) (20) (9) 20-30 79 83 55 1.53 1.28 0.91 41 38 36 (22) (13) (15) (0.93) (0.24) (0.20) (32) (4) (13) 30-40 106 114 33 2.23 1.76 0.69 28 38 28 (30) (51) (15) (0.17) (0.81) (0.29) (10) (8) . (6) 40-50 102 53 19 2.26 1.11 0.42 26 28 27 (31) (24) 13 (0.53) (0.46) (0.22) (2) (2) (14) 50-60 107 51 19 2.36 1.07 0.32 25 26 25 (62) (35) 15 (0.35) (0.53) (0.19) (11) (5) (21) 113 errors were frequently large in r e l a t i o n to the means. The pH was 3.4 at 0-10 cm in the VAME TSME, with a maximum pH of 4.4 occurring at d i f f e r e n t depths in a l l three plant associations (Table 5.19). The surface values in the OPHO THPL ere somewhat higher than in the other plant associations. There also appeared to be a general increase in pH with increased depth in a l l plant associations. The only difference between the above observations and those where the solution used for pH measurements was 0.01 M calcium chloride was the o v e r a l l drop in pH values. The minimum value was pH 2.8 and the maximum was pH 4.2. Standard errors were not so large in comparison to the means as those for some of the previous s o i l properties. Cation exchange capacity decreased with increasing depth in a l l associations (Table 5.19). The highest value was .376 mol charge n r 3 at 50-60 cm in the OPHO THPL. The lowest value was 130 mol charge m?? at 0-5 cm in the VAME TSME. Standard errors were often large in r e l a t i o n to the means. Volumetric water contents decreased from one time of sampling to the next in a l l plots when data were summarized from 0 to 50 cm (Table 5.20). The maximum surface value of the 2-9 June samples was 0.806 cm3cm"3 in the ABAM TSME while the minimum value of 0.272 cm3 cm"3 was in the VAME TSME in the 14-17 September samples. The l a t t e r association had the lowest recorded s o i l moisture value: 0.245 cm3 cm*3 at the 30-50 cm depth. Standard errors varied considerably in r e l a t i o n to means between plant associations, times of sampling, and Table 5 .19 . Hydrogen- ion a c t i v i t i e s and c a t i o n exchange c a p a c i t i e s o f s o i l s taken from 14-17 September 1977 separated by p lan t a s s o c i a t i o n and depth. In a l l cases the mean i s from 3 samples. The s tandard e r r o r i s i n parentheses. Depth begins from the su r face of the f o r e s t f l o o r . Hydrogen- ion A c t i v i t y i n Water Hydrogen- ion A c t i v i t y i n C a C l 2 Ca t ion Exchange Capac i t y Depth Q p H 0 ABAM VAME OPHO ABAM VAME OPHO ABAM VAME THPL TSME TSME THPL TSME TSME THPL TSME TSME cm 0-5 5-10 10-20 20-30 30-40 40-50 50-60 pH mol charge n r 3 4.0 3.5 3.4 3.7 3.0 2.9 144 193 130 (0 .4 ) (0 .1) (0 .3) (0 .5) (0.1) (0.3) (12) (35) (8) 4.1 3.4 3.6 3.7 2.8 3.1 189 195 171 (0 .4) (0 .1 ) (0 .8) (0 .5) (0.1) (0 .5) (20) (36) (37) 4 .0 3.7 3.7 3.5 3.3 3.1 203 145 202 (0 .5 ) (0 .1) (0 .7) (0 .5) (0.1) (0.6) (39) (8) (14) 4 .3 4.1 4 .0 3.8 3.7 3.6 173 268 247 (0 .4 ) (0 .1) (0 .8) (0 .4) (0.1) (0.8) (47) (78) (29) 4 .4 4.1 4 .4 3.8 3.8 3.9 277 370 173 (0 .4 ) (0 .1 ) (1 .2) (0 .5 ) (0.1) (1.0) (15) (90) (67) 4 .3 4 .3 4 .3 3.7 3.9 3.9 283 269 131 (0 .4) (0 .1) (0 .9) (0 .7) (0.2) (1 .2) (53) (55) (47) 4 .3 4 .4 4 .4 4 .0 4.2 3.9 376 346 *" 145 (0 .3 ) (0 .2 ) (1 .0 ) (0 .4) (0.3) (1.1) (53) (130) (61) Table 5 .20. Vo lumet r i c water contents of s o i l s separated by p lan t a s s o c i a t i o n , t ime of samp l i ng , and depth . In a l l cases the mean i s from 3 samples. The standard e r r o r s are i n paren theses . Depth begins at the su r face of the f o r e s t f l o o r . Depth OPHO THPL 2-9 June Time 18-21 J u l y 14-17 Sept . 2-9 June ABAM TSME Time 18-21 J u l y 14-17 Sept . VAME TSME 2-9 June Time 18-21 J u l y 14-17 Sept . —cm — 0-10 0.617 (0.131) 0.542 (0.008) 0.418 (0.050) 0.806 (0.228) cm 3 cm" 3 0.671 (0.113) 0.361 (0.124) 0.571 (0.156) 0.418 (0.125) 0.272 (0.123) 10-30 0.727 (0.104) 0.644 (0.007) 0.510 (0.111) 0.776 (0.459) 0.530 (0.233) 0.402 (0.140) 0.584 (0.117) 0.487 (0.063) 0.318 (0.049) 30-50 0.898 (0.292) 0.900 (0.258) 0.579 (0.110) 0.548 (0.288) 0.784 (0.574) 0.424 (0.071) 0.306 (0.179) 0.417 (0.204) 0.245 (0.126) 0-50 0.748 0.695 0.502 0.710 0.661 0.400 0.487 0.440 0.278 (0.208) (0.205) (0.108) (0.318) (0.333) (0.104) (0.190) (0.218) (0.097) 1 16 depths. S o i l water potential, as measured by the hygrometers, never dropped low enough to be measurable (-100 kPa i s the upper l i m i t of the unit) in any plant association or at any depth. This i s what might be expected considering the high volumetric water contents reported above. Temperatures in the s o i l were highest in the 18-21 July samples from a l l plant associations. The maximum value was 12.2°C while the minimum was 5.6°C, with both occurring in the surface of the VAME TSME (Table 5.21). The maximum mean temperature in this association at 10-30 cm was 8.4°C, while the minimum was 3.8°C. In a l l associations there was a general decrease in temperature from the 0-10 cm to the 10-30 cm increment in the 2-9 June and 18-21 July sample. In the 14-17 September sample from a l l associations the 10-30 cm increment had values equal to or higher than those in the 0-10 cm increment. Standard errors r e l a t i v e to the means were frequently large. 5.6.2 Correlations With Root Mass-Volume 5.6.2.1 Overstory Roots 117 Table 5 .21 . Temperatures i n the s o i l s a t 0-10 cm and 10-30 cm depths separated by p lan t a s s o c i a t i o n and t ime of sampl ing . In a l l cases the mean i s from 3 samples. The s tandard e r r o r s are i n parentheses . Depth begins at the su r face of the f o r e s t f l o o r . Depth OPHO THPL ABAM TSME VAME TSME Time Time Time 2-9 June 18-21 J u l y 14-17 Sept . 2-9 June 18-21 J u l y 14-17 Sept . 2-9 June 18-21 J u l y 14-17 Sept . cm 0-10 6.4 9.7 7.5 (1 .0) (0 .4 ) (0 .2) 7.8 10.5 6.7 (2 .4) (1 .2) (0 .6 ) 5.6 12.2 6 .8 (2 .6) (1 .3 ) (0 .1) 10-30 4 .6 7.8 7.5 4 .8 8.1 7.3 (0 .4) (0 .1 ) (0 .3) (1 .6 ) (0 .4 ) (0 .2) 3.8 8 .4 7.5 (1 .3) (0 .3 ) (0 .5) 5.5 8 .8 7.5 6 .3 9 .3 7.0 4 .7 10.3 7.2 0-30 (1.2) (1 .1 ) (0 .2) (2 .4 ) (1 .6) (0 .5 ) (2 .0) (2 .4 ) (0 .5) 118 Simple correlations were made for the three plant associations between the s o i l properties discussed in the previous section (excluding s o i l moisture and temperature) and root mass-volume (Table 5.22). The measured s o i l properties were from the 14-17 September sample, therefore the c o r r e l a t i o n was made with the root data from that time. For moisture and temperature correlations the root data were summed so that the depth increments for the parameters being correlated corresponded. Normally, with the lack of s t a t i s t i c a l l y s i g n i f i c a n t differences between the associations, the data from them would be combined. However, there was evidence to show differences in the means from the three associations in terms of below-ground biomass and in r a t i o s of below-ground to above-ground components (Table 5.11, 5.15, 5.16). Thus, separate correlations were carried out for individual plant assoc iat ions. When bulk density approaches 1.4 g cm'3, root growth ceases (Fayle 1975), although some tree species can grow in more dense s o i l s (Minore and others 1969). Also, i t i s easier for roots to grow in s o i l s with lower bulk densities (Harley and Russell 1979). With more coarse fragments the fine f r a c t i o n of the s o i l decreases, making i t more d i f f i c u l t for roots to obtain moisture and nutrients. Thus, to support a given amount of above-ground tree biomass there w i l l be less rooting where bulk density and coarse fragment content are high, than where they are low, provided that moisture and nutrients are adequate for growth to occur. This explains the negative correlations Table 5 .22 . Simple c o r r e l a t i o n s between s o i l p rope r t i es and the OPHO THPL, ABAM TSME, and VAME TSME f o r the th ree c a t e g o r i e s of ove rs to ry root biomass. Unsuber ized < 2 mm 2-5 mm Proper ty — OPHO ABAM VAME OPHO ABAM VAME OPHO ABAM VAME THPL TSME TSME THPL TSME TSME THPL TSME TSME r Bulk d e n s i t y * - 0 . 40 - 0 . 6 3 -0 .42 - 0 . 54 - 0 . 63 - 0 . 55 -0 .04 - 0 . 50 -0 .22 Coarse fragments - 0 . 22 -0 .48 -0 .38 - 0 . 37 - 0 . 48 - 0 . 51 -0 .30 - 0 . 42 - 0 . 2 5 Organic mat ter - 0 . 14 0.34 0.38 - 0 . 00 0. 42 0. 72 0.43 - 0 . 08 0.60 N i t rogen - 0 . 32 0.35 0.52 - 0 . 21 0. 36 0. 67 -0 .19 - 0 . 16 0.31 C/N r a t i o 0 . 11 0.05 0.01 0. 10 0. 18 •0. 41 0.57 0. 19 0.63 PH - 0 . 01 - 0 . 8 0 -0 .32 - 0 . 16 - 0 . 73 - 0 . 53 -0 .68 - 0 . 43 -0 .27 (0.01 M CaCl ) Cat ion exchange c a p a c i t y - 0 . 33 - 0 . 4 3 -0 .29 - 0 . 41 - 0 . 40 - 0 . 16 0.04 - 0 . 36 0.06 M o i s t u r e + - 0 . 30 -0 .11 -0 .02 - 0 . 32 0. 11 0. 10 -0 .07 - 0 . 06 0.46 Temperature^ 0. 35 0.66 0.71 0. 04 0. ,39 0. 44 -0 .39 0. 23 0.04 * n = 21 f o r 7 depths w i th samples from 14-17 September; + n = 27 f o r 3 depths ( 0 - 1 0 , 10 -30 , 30-50 cm) w i th samples from 2-9 June , 18-21 J u l y , and 14-17 September; fi n = 18 (OPHO THPL, ABAM TSME) 14 (VAME TSME) f o r 2 depths (0 -10 , 10-30 cm) wi th samples from 2-9 June , 18-21 J u l y , and 14-17 September. 120 observed in Table 5.22. Within any association the co r r e l a t i o n was lowest for the 2-5 mm roots. This may r e f l e c t the lesser importance of this category of root in moisture and nutrient absorption. The highest co r r e l a t i o n between root mass-volume and bulk density was for the unsuberized roots and the < 2 mm roots of the ABAM TSME (-0.63). Correlations for the unsuberized and < 2 mm roots within the associations were lower for coarse fragments than they were for bulk density. The c o e f f i c i e n t s were always negative. Correlations of root biomass with organic matter concentration varied from -0.00 in the < 2 mm roots of the OPHO THPL to 0.72 in the < 2 mm roots of the same VAME TSME (Table 5.22). There was v i r t u a l l y no corr e l a t i o n between unsuberized and < 2 mm roots and organic matter in the OPHO THPL. This i s probably due to the high organic matter content at a l l depths in this plant association. In contrast, there was a pos i t i v e c o r r e l a t i o n with the unsuberized and < 2 mm roots of the ABAM TSME and VAME TSME associations. Nitrogen also showed both positive and negative correlations with the greatest posi t i v e value of 0.67 with the < 2 mm roots of the VAME TSME and the greatest negative value (-0.32) with the unsuberized roots of the OPHO THPL. The trends for organic matter discussed above are also evident in the correlations of root mass-volume with nitrogen concentration. 121 Carbon-to-nitrogen r a t i o s were always p o s i t i v e l y correlated with fine root biomass. Correlations with the unsuberized roots were weak in a l l plant associations. The best c o r r e l a t i o n with < 2 mm root mass-volume was in the VAME TSME (0.41) while the poorest was in the OPHO THPL. In the case of 2-5 mm roots the best c o r r e l a t i o n was in the VAME TSME (0.63) with the OPHO THPL close behind (0.57). There was only a weak cor r e l a t i o n in the ABAM TSME (0.19). A high C/N r a t i o usually suggests that nitrogen i s less available for plant growth than where the r a t i o i s low. The positive (albeit sometimes weak) co r r e l a t i o n of C/N ratios with a l l categories of fine root in a l l plant associations suggests either that the fine roots of the overstory vegetation are not concentrated where there i s more nitrogen a v a i l a b l e , or that the C/N rat i o s are not an indicator of nitrogen a v a i l a b i l i t y to the overstory vegetation of these plant associations. Hydrogen ion a c t i v i t y was lower near the surface (Table 5.19). Thus, there was a negative c o r r e l a t i o n with root biomass. If t h i s were an a g r i c u l t u r a l s o i l , the low pH would have meant that phosphorus was less available than where the pH was higher (Tisdale and Nelson 1975). However, these s o i l s are not l i k e a g r i c u l t u r a l s o i l s and mycorrhizae probably counteract any effect the low pH at the surface would have on phosphorus a v a i l a b i l i t y to the plants in these associations. 122 Cation exchange s i t e s as measured by CEC r e f l e c t the potential of the s o i l to supply cations, provided that these s i t e s are not predominantly occupied by hydrogen ions. (Base saturation data were not a v a i l a b l e ) . Cation exchange capacity does not give information about cation a v a i l a b i l i t y although i t can give a general idea about th e i r potential a v a i l a b i l i t y . Cation exchange capacity increased with depth in the OPHO THPL and ABAM TSME. This i s r e f l e c t e d in the negative • correlations shown in Table 5.22 (the maximum cor r e l a t i o n was -0.43, with unsuberized roots of the ABAM TSME.) There was a peak in CEC in the 20-30 cm depth of the VAME TSME; therefore, the correlations with root mass-volume for t h i s association were weaker. There was no c o r r e l a t i o n of CEC with the 2-5 mm root category in the case of the OPHO THPL and the VAME TSME. The c o r r e l a t i o n was negative in the ABAM TSME (-0.36). Volumetric water content was either not correlated, or correlated negatively (-0.32 in the case of < 2 mm roots of the OPHO THPL) with root mass-volume (Table 5.22). Considering the high moisture content of these s o i l s in general, i t i s unlikely that any d e f i c i t was experienced by the vegetation. The high positive c o r r e l a t i o n (0.46) in the 2-5 mm roots of the VAME TSME i s a t y p i c a l , considering the other correlations with CEC (Table 5.22) 123 Unsuberized and < 2 mm roots were p o s i t i v e l y correlated with s o i l temperature in a l l associations (Table 5.22). Only for the 2-5 mm roots of the OPHO THPL was there a negative c o r r e l a t i o n . Correlations within a plant association were stronger for unsuberized roots than for < 2 mm and 2-5 mm roots. This suggests that the unsuberized roots are more sensitive to seasonal temperature changes than the suberized roots. 5.6.2.2 Understory Roots Understory roots were similar to overstory roots in many cases, with negative correlations for bulk density and coarse fragment contents (except for no corr e l a t i o n in the 2-5 mm root category of the OPHO THPL) (Table 5.23). Organic matter and nitrogen concentrations were variably, and usually weakly correlated with understory root biomass. If these parameters had ref l e c t e d areas in the s o i l that were important for plant uptake, the cor r e l a t i o n would have been strongly p o s i t i v e . It was weakly so in the VAME TSME for both root categories. It i s not clear why the co r r e l a t i o n of organic matter with 2-5 mm understory roots was higher (0.73) than with the < 2 mm roots (0.54) since the smaller roots should be more sensitive to organic matter and nitrogen concentrations. Table 5 .23 . Simple c o r r e l a t i o n s between s o i l p r o p e r t i e s and the OPHO THPL, ABAM TSME, and VAME TSME p l a n t a s s o c i a -t i o n s f o r the two s i z e ca tego r i es o f unders tory root biomass. < 2 mm 2-5 mm Proper ty OPHO ABAM VAME OPHO ABAM VAME THPL TSME TSME THPL TSME TSME r Bulk d e n s i t y * - 0 . 50 - 0 . 35 - 0 . 50 - 0 . 43 - 0 . 34 - 0 . 42 Coarse fragments - 0 . 23 - 0 . 25 - 0 . 55 0. 05 - 0 . 20 - 0 . 53 Organic mat ter - 0 . 16 0. 21 0. 54 - 0 . 20 - 0 . 04 0 . 73 N i t rogen - 0 . 33 0. 33 0. 67 - 0 . 41 - 0 . ,06 0. 61 C/N r a t i o 0. 07 - 0 . 08 0. .10 0. 06 0. ,04 0. 42 pH 0. 15 - 0 . 27 - 0 . ,55 - 0 . 02 - 0 . ,43 - 0 . 43 (0.01 M C a C l 2 ) Ca t ion exchange c a p a c i t y - 0 . 49 - 0 . 18 - 0 . ,07 - 0 . 48 - 0 , .20 - 0 . 04 M o i s t u r e + - 0 . 17 - 0 . 15 0. ,30 - 0 . 34 0. .16 0. 34 Temperature^ - 0 . 17 0. 23 0. ,67 0. ,53 - 0 , .23 - 0 . 15 * n = 21 f o r 7 depths w i th samples from 14-17 September; + n = 27 f o r 3 depths 0 -10 , 10 -30 , 30-50 cm w i th samples from 2-9 June , 18-21 J u l y , and 14-17 September; n n = 18 , 18, 14 (sometimes 2 r e p l i c a t e s were a v a i l a b l e ) f o r 2 depths ( 0 - 1 0 , 10-30 cm) w i th samples from 2-9 June , 18-21 J u l y , and 14-17 September. 125 Carbon-to-nitrogen ratios were weakly correlated with understory root biomass in a l l associations. The best co r r e l a t i o n (0.42) was with the 2-5 mm root mass-volume. As mentioned in the l a s t section, either the fine roots of the understory vegetation are not related to available nitrogen, or the C/N ratios are not an indicator of nitrogen a v a i l a b i l i t y to the understory species of these plant associations. The strongest negative c o r r e l a t i o n (-0.55) with pH a c t i v i t y was with the < 2 mm roots of the VAME TSME. With the exception of the < 2 mm roots of the OPHO THPL, correlations with root mass-volume were weak and negative. Cation exchange capacity was negatively correlated with root .mass-volume. The strongest c o r r e l a t i o n was -0.49 for < 2 mm roots in the OPHO THPL. The weakest c o r r e l a t i o n was with the 2-5 mm roots in the OPHO THPL (-0.04). Moisture was weakly correlated with understory root mass-volume. With high moisture contents at a l l depths and the peak in < 2 mm and 2-5 mm roots near the surface t h i s was expected. Temperature correlations with < 2 mm understory roots were negative in the OPHO THPL and weakly positive (0.67) in the VAME TSME. This trend was reversed for the 2-5 mm roots with the greatest c o r r e l a t i o n in the OPHO THPL (0.53) (Table 5.23). 126 5.6.3 Results Summarized by Plant Association Mean bulk density was highest in the ABAM TSME and lowest in the OPHO THPL (Table 5.24). Mean coarse fragment content increased from the OPHO THPL, to the ABAM TSME, and to the VAME TSME. Total organic matter and nitrogen contents had the opposite trend. The C/N r a t i o was similar for a l l plant associations. Mean pH in both water and calcium chloride did not d i f f e r between associations. Mean cation exchange capacity was highest in the ABAM TSME, intermediate in the OPHO THPL, and least in the VAME TSME. It did not d i f f e r between the ABAM TSME and VAME TSME. Mean volumetric water content was sampled at a l l three times. It was highest in the OPHO THPL, second highest in the ABAM TSME, and least in the VAME TSME. Mean temperature in the s o i l was also sampled at a l l three times. It was only s l i g h t l y higher in the ABAM TSME with the remaining associations being equal. Large, standard errors r e l a t i v e to the means are evident for the values reported above. In r e l a t i o n to the above results i t should be s l i g h t l y easier for roots to grow in the OPHO THPL than in the other plant associations because of lower mean bulk density and lower mean coarse fragment contents there. Also, nitrogen should be more available to plants in the OPHO THPL than in the other plant associations since i t has a greater mean concentration and the C/N ra t i o s are similar in a l l plant associations (this assumes that for a similar C/N r a t i o , there w i l l be more 127 Table 5 .24 . Summary of s o i l p r o p e r t i e s f o r the OPHO THPL, ABAM TSME, and VAME TSME. Samples were taken on 14-17 September except i n the case of s o i l mo is ture and temperature which were sampled on 2-9 June , 18-21 J u l y , and 14-17 September. In a l l cases the mean i s from 3 samples. The s tandard e r r o r i s i n paren theses . Proper ty OPHO THPL ABAM TSME VAME TSME Mean bulk dens i t y (g c m - 3 ) 0.27 (0.14) 0.41 (0.22) 0.31 (0.19) Mean coarse fragments (7o o f volume) 5 (8) 15 (19) 25 (27) To ta l o rgan ic mat ter (kg m" 3) 97.8 (27.8) 83.4 (36.6) 51.5 (29.6) To ta l n i t r ogen (kg m - 3 ) 1.94 (0.52) 1.38 (0.59) 0.86 (0.43) C/N r a t i o 31 (13) 35 (10) 34 (12) Mean pH i n water 4 .2 (0 .4) 3.9 (0 .4) 4 .0 (0 .8) Mean pH i n C a C l 2 3.7 (0 .4 ) 3.5 (0 .5) 3.5 (0 .8) Mean c a t i o n exchange c a p a c i t y (mol charge m - 3 ) 235 (83) 255 (99) 171 (54) Mean Mo is tu re (cm 3 c m - 3 ) 0.648 (0.203) 0.589 (0.297) 0.402 (0.165) Mean Temperature ( ° C ; 0-30 cm) 7.3 7.6 7.4 (1 .6 ) (2 .1 ) (2 .7) 128 nitrogen available for plant growth where the amount of t o t a l nitrogen i s greater). If base saturation and base composition are similar in the three plant associations, the higher CEC in the OPHO THPL may indicate that cations are more available for root uptake in this association than in the other plant associations. Also, there i s groundwater flowing through the OPHO HPL during most of the growing season. This would have increased the amount of bases available for vegetation uptake in t h i s plant association r e l a t i v e to the other plant associations. (The presence of thi s groundwater i s re f l e c t e d in the higher moisture contents as well.) Applying the above arguments and assumptions to the remaining plant associations, root growth would be s l i g h t l y easier in the VAME TSME due to the lower mean bulk density. However, t h i s would be counterbalanced by the higher mean coarse fragment content in the VAME TSME than in the ABAM TSME. N u t r i t i o n a l l y , there should be more nitrogen and more bases available for vegetation growth in the ABAM TSME than the VAME TSME. In terms of moisture r e l a t i o n s , the ABAM TSME had a greater mean volumetric water content than the VAME TSME. Perhaps t h i s difference was due to the input of groundwater (and bases) to the ABAM TSME. There was no such input to the VAME TSME. On the other hand, as shown in Table 5.20, there was never any moisture d e f i c i t in either the VAME TSME or the ABAM TSME during the growing season. 129 The arguments above assume that the plants growing on the plant association studied have equal a b i l i t i e s to grow and take up nutrients. If t h i s i s true, then the e f f o r t required by the plants to grow roots and obtain nutrients i s less in the OPHO THPL than in the other plant associations. Also, the e f f o r t required for plants to grow roots i s similar in the ABAM TSME and VAME TSME, but nutrients are easier for plants to obtain in the ABAM TSME. Where i t is more d i f f i c u l t for roots to grow and where fewer nutrients are available there should be more e f f o r t expended by a plant to grow and obtain nutrients and this should be reflected in the properties of fine roots and the ratios of these to fol i a g e . The following results that have been presented e a r l i e r tend to support t h i s hypothesis. 1. On the OPHO THPL, the < 2 mm roots of the overstory vegetation had the least biomass, length, and surface area on an area basis when compared to the ABAM TSME and VAME TSME; 2. The ABAM TSME when compared to the other plant associations was intermediate in terms of biomass, length, and surface area of fine roots of the overstory vegetation; 3. On the VAME TSME the < 2 mm roots of the overstory vegetation had the greatest biomass, length, and surface area on an areal basis as well as r e l a t i v e to above-ground components when compared to the OPHO THPL and the ABAM TSME. 130 Other investigators have discovered greater root mass, density, length or volume on i n f e r t i l e and dry s i t e s (Scholtes 1953, Heikurainen 1957b, Rogers and Booth 1959, Kern and others 1961, Bray 1963, Smith 1964, Mikola and others 1966, Meyer 1967, T611e 1967, Lorio and others 1972, Kochendorfer 1973, Karizumi 1976, Santantonio 1978, Keyes and Grier 1981). Garelkov (1973) found similar trends as the above, and he also reported that active roots were concentrated at shallower depths on poorer s i t e s than on richer ones. Preston (1942) found an increase in mycorrhizal roots of juvenile lodgepole pine on the poorest of two s i t e s . Not a l l the results of this study agree with the above hypothesis: 1. The biomass, length, and surface area of unsuberized roots remained roughly the same on a l l associations. This does not mean that uptake of nutrients was the same, since suberized roots and the unsampled mycorrhizae were probably more important in nutrient uptake than unsuberized roots in these associations. 2. The standard errors for fine root data were always large in r e l a t i o n to the means, so that there were no s t a t i s t i c a l l y s i g n i f i c a n t differences between the associations. 3. The biomass, length, and surface area of fine roots of the understory vegetation were greatest in the OPHO THPL and the VAME TSME and least in the ABAM TSME. It i s l i k e l y that the biomass of understory vegetation i s governed more by the amount of l i g h t available through the overstory canopy than by the physical or chemical properties of the s o i l s . (Both the OPHO THPL and the VAME TSME had a r e l a t i v e l y sparse canopy coverage while coverage was r e l a t i v e l y dense in the ABAM TSME.) 132 6. SUMMARY AND CONCLUSIONS A l l of the objectives set out for t h i s study have been met. Detailed analysis was carr i e d out only on biomass due to the close c o r r e l a t i o n between biomass, length, and surface area within a pa r t i c u l a r size category of root. At various stages in the section dealing with r e s u l t s , summaries of length and surface area were presented. In the case of < 2 mm overstory roots there was a rapid decline with increasing depth. Overstory roots 2-5 mm in diameter peaked above 30 cm and then declined rapidly. The least rapid decrease with increasing depth was. in the < 2 mm roots of the OPHO THPL. The ABAM TSME and VAME TSME were similar in terms of the v e r t i c a l d i s t r i b u t i o n of roots. In both there was a more rapid decrease with increasing depth than there was in the OPHO THPL. Mass of roots 2-5 mm in diameter was lower near the surface in the OPHO THPL than in the other associations. Unsuberized roots were separated from the < 2 mm category. Their d i s t r i b u t i o n was the shallowest of a l l the categories of root with 90 % of the biomass above 30 cm in a l l associations. Over 90 % of the < 2 mm roots were above 40 cm in the ABAM TSME and VAME TSME, while 86 % were above t h i s depth in the OPHO THPL. Ninety percent of the biomass of 2-5 mm roots was above 50 cm in the three associations. 133 Near the surface, < 2 mm roots of the understory vegetation did not decrease as consistently with depth as the < 2 mm roots of the overstory vegetation. They did follow similar trends at greater depths. In the OPHO THPL and ABAM TSME, 89 % of the biomass was above 40 cm while 83 % was above t h i s depth in the VAME TSME. In the 2-5 mm category, 89 % of the biomass was above 40 cm in the OPHO THPL, 99 % was above 30 cm in the ABAM TSME, and 95 % was above 30 cm in the VAME TSME. In the VAME TSME, the < 2 mm roots of the understory vegetation grew deeper than the < 2 mm roots of the overstory vegetation. The d i s t r i b u t i o n s of < 2 mm roots of the understory and overstory vegetation were similar on the other associations. Different v e r t i c a l d i s t r i b u t i o n s in 2-5 mm roots were l i k e l y due to differences in plant morphology rather than to functional relationships. Only the v e r t i c a l d i s t r i b u t i o n of unsuberized roots varied s t a t i s t i c a l l y with time. Growth during the season was primarily in the upper 5 cm. Other root categories of the overstory vegetation did not vary s t a t i s t i c a l l y from one sampling time to the next. The v e r t i c a l d i s t r i b u t i o n of < 2 mm roots of the understory vegetation varied during the growing season. There was an indication that root growth had occurred prior to the f i r s t sampling time, but random va r i a t i o n between repli c a t e s could not be ruled out. Changes in overstory and understory 2-5 mm root biomass with time was thought to be due to sampling v a r i a b i l i t y . 134 When data from a l l depths were combined i t was evident that unsuberized roots peaked in the 18-21 July sample in the VAME TSME and ABAM TSME, while they increased continuously throughout the sampling period in the OPHO THPL. Unsuberized roots found in the f i r s t sampling suggested that some previous growth had occurred or that dormant unsuberized roots were present from the previous growing season. Temporal variation in other categories of root of the overstory vegetation and in the 2-5 mm roots of the understory vegetation was probably due to sampling error but could also have been due to growth into the next size category. Biomass of a l l root categories of the overstory vegetation together 5 mm roots) varied between 740 and 1320 g nr 2 with the largest va l ues in the VAME TSME, intermediate values in the ABAM TSME, and the smallest in the OPHO THPL. Lengths were 3.8 to 6.5 km nr 2 and surface areas were from 7.3 to 11.9 m2 nr 2. In both length and surface area the VAME TSME and ABAM TSME were similar and the OPHO THPL was considerably lower. Biomass of a l l root categories of the understory vegetation together 5 mm roots) varied between 44 and 253 g nr 2, length between 0.67 and 4.70 km nr 2, and surface area between 0.48 and 3.16 m2 nr 2. The highest values were always in the VAME TSME, the OPHO THPL had intermediate values, and the ABAM TSME had the lowest. 135 The root biomass of the overstory vegetation was greater than the root biomass of the understory vegetation; however, in terms of length and surface area, the understory was a si g n i f i c a n t contributor to the t o t a l amounts. S t a t i s t i c a l analysis showed s i g n i f i c a n t differences between repl i c a t e plots of the associations. Other differences were found, but generally the v a r i a b i l i t y was high enough to preclude detection of s t a t i s t i c a l l y s i g n i f i c a n t differences between the associations. When the ratios of biomass of overstory $ 5 mm roots to the above-ground biomass of the overstory vegetation were considered, the highest value for the VAME TSME was nearly twice the lowest in the OPHO THPL, while the ABAM TSME was intermediate. This trend was the same as the one for biomass of fine roots alone, indicating that these associations indeed had d i f f e r e n t strategies in apportioning biomass between above-and below-ground parts. There was an indication of a greater r a t i o of biomass of < 2 mm roots of overstory vegetation to biomass of foliage plus twig on the VAME TSME than on the ABAM TSME and the OPHO THPL. This supported the hypothesis that there was more biomass of fine roots of overstory where there was more fo l i a g e . When the ratio s of root biomass to foliage plus twig biomass were summarized by association, the trends were similar to those reported for the ratios of root biomass to above-ground overstory biomass. 136 Correlations between s o i l physical and chemical parameters and root mass-volume were generally weak. Examinations of the mean concentrations of nitrogen and the mean CEC in the associations suggested that nutrient a v a i l a b i l i t y was greatest in the OPHO THPL, intermediate in the ABAM TSME, and least in the VAME TSME. The input into the OPHO THPL as groundwater was not accounted for in the mean values. Since there was no moisture d e f i c i t in any of the associations, i t was hypothesized that nutrient a v a i l a b i l i t y was the factor most strongly a f f e c t i n g the r e l a t i v e amounts of fine roots of the overstory vegetation in the associations. The results of thi s study indicated that the greatest amount of fine roots of the overstory vegetation was in the VAME TSME which was the poorest n u t r i t i o n a l l y ; the ABAM TSME had an intermediate amount of fine roots and was intermediate n u t r i t i o n a l l y ; while the OPHO THPL had the least amount of fine roots and was the riche s t . Due to the large v a r i a b i l i t y between replicates of the plant associations none of the -above conclusions could be based on s t a t i s t i c a l l y s i g n i f i c a n t differences between the associations in terms of the fine root biomass of the overstory vegetation. 137 LITERATURE CITED A l l i s o n , L.E., W.B. BoLlen, and CD. Moodie. 1965. Total carbon. Pp. 1346-1366 in C.A. Black, ed. 1965 Methods of s o i l analysis: Part 2, Chemical and microbiological properties. American Society of Agronomy, Inc. Madison, Wisconsin, USA. Agronomy No. 9. 1572 p. Ambler, J.R. and J.L. Young. 1977. Techniques for determining root length infected by vesicular-arbuscular mycorrhizae. S o i l S c i . Amer. Journal 41: 551-556. Andersson, F. 1970. Ecological studies in a scanian woodland and meadow area, southern Sweden. II. Plant biomass, primary production and turnover of organic matter. Bot. Notiser 123: 8-51, Lund. Baskerville, G.L. 1966. Dry-matter production in immature balsam f i r stands: roots, lesser vegetation, and t o t a l stand. For. S c i . 12: 49-53. Black, C.A., ed. Methods of s o i l analysis: Part 2, Chemical and microbiological properties. American Society of Agronomy, Inc. Madison, Wisconsin, USA. Agronomy No. 9. 1572 p. Bohm, W. 1979. Methods of studying root systems. Springer-Verlag, B e r l i n . 188 p. Bowen, G.D. 1964. Root d i s t r i b u t i o n of Pinus radiata . C.S.I.R.O. Div. of S o i l s D i v i s i o n a l Report 1/64. f4 p. Bray, J.R. 1963. Root production and the estimation of net productivity. Can. J . Bot. 41: 65-72. C.A. Black, ed. Methods of s o i l analysis: Part 2, Chemical and microbiological properties. American Society of Agronomy, Inc. Madison, Wisconsin, USA. Agronomy No. 9. 1572 p. Brooke, C.R., E.B. Peterson, and V.J. Krajina. 1970. The subalpine Mountain Hemlock Zone. Ecol. of W. North America 2(2): 147-349. Brown, G.R. and J.F. Thilenius. 1977. A tool and method for extracting p l a n t - r o o t - s o i l cores on remote s i t e s . J . Range Manage. 30(1): 72-74. Carbon, B.A., G.A. Bartle, A.M. Murray, and D.K. Macpherson. 1980. The d i s t r i b u t i o n of root length, and the l i m i t s to flow of s o i l water to roots in a dry scl e r o p h y l l forest. For. S c i . 26: 656-664. 138 Carson, E.W., ed. 1974. The plant root and i t s environment. University Press of V i r g i n i a . C h a r l o t t e s v i l l e . 691 p. Chapman, H.D. 1965. Cation-exchange capacity. Pp. 891-901 jln C.A. Black, ed. Methods of s o i l analysis: Part 2, Chemical and microbiological properties. American Society of Agronomy, Inc. Madison, Wisconsin, USA. Agronomy No. 9. 1572 p. Cochran, W.G. and G.M. Cox. 1957- Experimental designs. Wiley, New York. CSSC (Canada S o i l Survey Committee, Subcommittee on S o i l C l a s s i f i c a t i o n ) . 1978. The Canadian system of s o i l c l a s s i f i c a t i o n . Can. Dept. Agric. Publ. 1646. Supply and Services Canada, Ottawa, Ont. 164 p. Damman, A.W.H. 1971. Effect of vegetation changes on the f e r t i l i t y of a Newfoundland forest s i t e . Ecol. Monog. 41: 253-270. Fayle, D.C.F. 1975. Extension and longitudinal growth during the development of red pine root systems. Can. J . For. Res. 5: 109-121. Ford, E.D. and J.D. Deans. 1977. Growth of a Sitka spruce plantation: s p a t i a l d i s t r i b u t i o n and seasonal fluctuation of lengths, weights, and carbohydrate concentrations of fine roots. Plant and S o i l 47: 463-485. Garelkov, D. 1973. B i o l o g i c a l productivity of some beech forest types in Bulgaria. Pp. 307-314 i_n IUFRO Biomass Studies; Mensuration, Growth and Y i e l d . College of L i f e Sciences and Agriculture, University of Maine at Orno. Harley, J.L. and R.S. Russell (eds.). 1979. The s o i l - r o o t interface. Academic Press, New York and London. 448 p. Harris, W.F., R.A. Goldstein and G.S. Henderson. 1973. Analysis of forest biomass pools, annual primary production and turnover of biomass for a mixed deciduous forest watershed. Pp. 43-64 in IUFRO Biomass Studies, Mensuration, Growth and Y i e l d . College of L i f e Sciences and Agriculture, Univ. of Maine at Orno. Harris, W.F., R.S. Kinerson, J r . , and N.T. Edwards. 1977. Comparison of belowground biomass of natural deciduous forests and l o b l o l l y pine plantations. Pedobiologia 17: 369-381. Hausdorfer, Von H.D. 1957. Die Durchwurzelung unter Kiefer auf zwei Standorten des Choriner Sanders. Archiv. Forstwes. 6(11/12): 811-827. 139 Head, G.C. 1966. Estimating seasonal changes in the quantity of white unsuberized root on f r u i t trees. Journal of Hort. S c i . 41: 197-206. Hermann, R.K. 1977. Growth and production of tree roots: A review. Pp. 7-28 in J.K. Marshall, ed. The belowground ecosystem: A synthesis of plant-associated processes. Range Science Dept. Series No. 26. Colorado State University, Fort C o l l i n s . Heikurainen, L. 1957a. Uber Veranderungen in den Wurzelverhaltnissen der Kiefernbestande auf Moorboden im laufe des Jahres. Acta. For. Fenn. 65(2): 1-70. Heikurainen, L. 1957b. Ramemannikon juuriston rakenne ja kuivatuksen vaikutus s i i h e n . Acta. For. Fenn. 65(3): 1-83. (German summary). Heth, D. and D.G.M. Donald. 1978. Root biomass of Pinus  radiata D. Don. S. African For. J . 107: 60-76. Hoffmann, Von G. 1972. Wachstumsrhythmik der Wurzeln und Sprossachsen von Forstgeholzen. Flora, Bd. 161: 303-319. Holden, J . 1975. Use of nuclear staining to assess rates of c e l l death in cortices of e e r i a l roots. S o i l B i o l . Biochem. 7: 333-334. Hough, W.A. , F.W.. Woods, and M.L. McCormack. 1965. Root extension of individual trees in surface s o i l s of a natural longleaf pine - turkey oak stand. For. S c i . 11: 223-243. Huttel, C. 1975. Root d i s t r i b u t i o n and biomass in three Ivory Coast rain forest p l o t s . Pp. 123-130 iri Golley, F.B. and E. Medina, eds. Ecological Studies 11. Tropical ecological systems: trends in t e r r e s t r i a l and aquatic research. 398 p. Jenik, J . 1971. Root structure and underground biomass in equatorial forests. Pp. 323-331 in P. Duvigneaud, ed. Productivity of forest ecosystems. UNESCO, Paris. Kalela, E.K. 1950. Mannikoiden ja kuusikoiden juurisuhteista I. Acta. For. Fenn. 57(2): 1-79. (English summary.) (On the horizontal roots in pine and spruce stand I. Kalela, E.K. 1957. Uber Veranderungen in den Wurzelverhaltnissen der Kiefernbestande im Laufe der Vegetationsperiode. Acta F o r e s t a l i a Fennica 65: 5-42. Karizumi, N. 1974a. The mechanism and function of tree root in the process of forest production. I. Method of investigation and estimation of the root biomass. B u l l , of the Government Forest Experiment Station No. 259. Meguro, 140 Tokyo, Japan. 99 p. Karizumi, N. 1974b. The mechanism and function of tree root in the process of forest production. I I . Root biomass and d i s t r i b u t i o n in stands. B u l l , of the Government Forest Experiment Station No. 267. Meguro, Tokyo, Japan. 88 p. Karizumi, N. 1976. The mechanism and function of tree root in the process of forest production. I I I . Root density and absorptive structure. B u l l , of the Government Forest Experiment Station No. 285. Meguro, Tokyo, Japan. 149 p. Kern, K.G. von, W. Moll, and H.J". Braun. 1961. Wurzeluntersuchungen in Rein und Mischbestanden des Hochschwarzwaldes ( V f l . Todtmoos 2/I-IV). Allgemeine Forst und Jagdzeitung. 132: 241-260. Keyes, M.R. and C.C. Grier. 1981. Above- and below-ground net production in 40-year-old Douglas-fir stands on low and high productivity s i t e s . Can. J . For. Res. 11: 599-605. Kimmins, J.P. 1974. Nutrient removal associated with whole-tree logging on two d i f f e r e n t s i t e s in the Prince George Forest D i s t r i c t . Report to B r i t i s h Columbia Forest Service Productivity Committee. 100 p. Kimmins, J.P. and B.C. Hawkes. 1978. Di s t r i b u t i o n and chemistry of fine roots in a white spruce-subalpine f i r stand in B r i t i s h Columbia: Implications for management. Can. J . For. Res. 8: 265-279. Kinman, C.F. 1932. A preliminary report on root growth studies with some orchard trees. Proc. Amer. Soc. Hort. S c i . 29: 220-224. Klein, R.L. and J.W. Geis. 1978. Biogenic s i l i c a in the pinaceae. S o i l S c i . 126: 145-156. Klinge, H. 1973. Root mass estimation in lowland t r o p i c a l rain forests of Central Amazonia, B r a z i l . I. Fine root masses of a pale yellow l a t o s o l and a giant humus podzol. Trop. Ecol. 14(1): 29-38. 141 Klinge, H. 1976. NShrstoffe, Wasser und Durchwurzelung von Podsolen und Latosolen unter tropischem Regenwald bei Manaus/Amazonien. Biogeographica 7: 45-58. Knievel, D.P. 1973. Procedure for estimating r a t i o of l i v e to dead root dry matter in root core samples. Crop S c i . 13: 124-126. Kochendorfer, J.N. 1973. Root d i s t r i b u t i o n under some forest types native to West V i r g i n i a . Ecology 54: 445-448. Kostler, J.N., E. Bruckner, and H. B i b e l r i e t h e r . 1968. Die Wurzeln der Waldbaume. Verlag Paul Parey. Hamburg und B e r l i n . 284 p. Kozlowski, T.T. 1979. Growth and development of trees. Volume II. Cambial growth, root growth, and reproductive growth. Academic Press, New York and London. 514 p. Krajina, V.J. 1969. Ecology of forest trees in B r i t i s h Columbia. Ecology of Western North America 2(1): 1-152. Kramer, P.J. 1946. Absorption of water through suberized roots of trees. Plant Phys. 21: 37-41. Kramer, P.J. and H.C. Bullock. 1966. Seasonal variations in the proportions of suberized and unsuberized roots of trees in r e l a t i o n to the absorption of water. Amer. J . Bot. 53: 200-204. Krumlik, G.J. 1979. Comparative study of nutrient c y c l i n g in the subalpine Mountain Hemlock Zone of B r i t i s h Columbia. University of B r i t i s h Columbia Faculty of Forestry, Vancouver, B r i t i s h Columbia. 196 p. Dissertation. Lorio, P.L., J r . , V.K. Howe, and C.N. Martin. 1972. L o b l o l l y pine rooting varies with microrelief on wet s i t e s . Ecology 53: 1134-1140. Lott, W.L., D.P. Satchell, and N.S. H a l l . 1950. A tracer-element technique in the study of root extension. Proc. Amer. Soc. Hort. S c i . 55: 27-34. Lyr, H. and G. Hoffmann. 1967. Growth rates and growth p e r i o d i c i t y of tree roots. Pp. 181-236 in. J' A« Romberger and P. Mikola, eds. Int. Rev. For. Res. Vol. 2. Academic Press, New York. MalkSnen, E. 1974. Annual primary production and nutrient cycle in some Scots pine stands. Comm. Inst. For. Fenniae 84: 1-87. 142 Marchenko, A.N. and Ye.M. Karlov. 1962. Mineral exchange in spruce forests of the northern taiga and forest tundra in Arkhangelsk Oblast. Soviet S o i l S c i . 7: 722-734. Mathews, W.H. 1958a. Geology of the Mount Garibaldi map-area, southwestern B r i t i s h Columbia, Canada. Part I. Igneous and metamorphic rocks. B u l l . Geol. Soc. Am. 69: 161-178. Mathews, W.H. 1958b. Geology of the Mount Garibaldi map-area, southwestern B r i t i s h Columbia, Canada. Part I I . Geomorphology and Quaternary volcanics. B u l l . Geol. Soc. Am. 69: 179-198. McGinty, D.T. 1976. Comparative root and s o i l dynamics on a white pine watershed and in the hardwood forest in the Coweeta Basin. University of Georgia, Athens, Georgia. 110 p. Dissertation. McQueen, D.R. 1968. The quantitative d i s t r i b u t i o n of absorbing roots of Pinus s y l v e s t r i s and Fagus sylv a t i c a in a forest succession. Oecol. Plant. 3: 83-99. McQueen, D.R. 1973. Changes in understory vegetation and fine root quantity following thining of 30-year Pinus radiata in central North Island, New Zealand. J. Appl. Ecol. 10: 13-Meyer, F.H. 1967. Feinwurzelverteilung bei Waldbaumen in Abhangigkeit vom Substrat. Forstarchiv 38: 286-290. Meyer, F.H. and D. GSttsche. 1971. D i s t r i b u t i o n of root t i p s and tender roots of beech. Pp. 48-52 ijn H. Ellenberg, ed. Integrated experimental ecology. Springer-Verlag, N.Y. Mikola, P., J. Hahl, and E. Tornianen. 1966. V e r t i c a l d i s t r i b u t i o n of mycorrhizae in pine forests with spruce undergrowth. Ann. Bot. Fenn. 3: 406-409. Minore, D., C.E. Smith, and R.F. Woollard. 1969. Effects of high s o i l density on seedling root growth of seven northwestern tree species. U.S.D.A. Forest Service Research Note. PNW-112. 6 p. Miroslav, I. 1976. Floodplain forest in biomass. Pp. 205-229 in IUFRO Biomass Studies, Mensuration, Growth and Y i e l d . College of L i f e Sciences and Agriculture, Univ. of Maine at Orno. M i t c h e l l , D.F. and F.W. Woods. 1966. Root extension in a longleaf pine plantation. Ecology 47: 97-102. Moir, W.H. and E.P. Bachelard. 1969. D i s t r i b u t i o n of fine roots in three Pinus radiata plantations near Canberra, 143 A u s t r a l i a . Ecology 50: 658-662. Newman, E.I. 1966. A method of estimating the t o t a l length of root in a sample. J . Appl. Ecol. 3: 139-145. Nihlgard, B. 1972. Plant biomass, primary production and d i s t r i b u t i o n of chemical elements in a beech and a planted spruce forest in south Sweden. OIKOS 23: 69-81. Nnyamah, J.V. and T.A. Black. 1977a. Rates and patterns of water uptake in a Douglas-fir forest. S o i l S c i . Soc. Am. J . 41: 972-979. Nnyamah, J.V. and T.A. Black. 1977b. F i e l d performance of the dew-point hydrometer in studies of s o i l - r o o t water r e l a t i o n s . Can. J. S o i l S c i . 57: 437-444. Nnyamah, J.V., T.A. Black, and C S . Tan. 1978. Resistance to water uptake in a Douglas-fir forest. S o i l S c i . 126: 63-76. Orlov, A. Ya. 1969. [A method of determining the root biomass of trees in the forest and the p o s s i b i l i t y of c a l c u l a t i n g the annual increment of the organic biomass in the mass of forest s o i l . ] Translated from Russian by Can. Dept. of Fisheries and Forestry Library, Ottawa. Source from Lesovedenie No. 1. 1967. Pp. 64-70, Moscow. c Overton, W.S., D.P. Lavender, and R.K. Herman. 1973. Estimation of biomass and nutrient c a p i t a l in stands of old-growth Douglas-fir. Pp. 91-103 JJT. IUFRO Biomass Studies, Mensuration, Growth and Y i e l d . College of L i f e Sciences and Agriculture, Univ. of Maine at Orno. Ovington, J t D . and H.A.I. Madgwick. 1959. D i s t r i b u t i o n of organic matter and plant nutrients in a plantation of Scots pine. For. S c i . 5: 344-355. Persson, H. 1978. Root dynamics in a young Scots pine stand in Central Sweden. Oikos 30: 508-519. Persson, H. 1979. Fine-root production, mortality, and decomposition in forest ecosystems. Vegetatio 41(2): 101— 109. Preston, R.J., J r . 1942. The growth and development of the root systems of juvenile lodgepole pine. Ecol. Monog. 12: 449-468. Reed, J.F. 1939. Root and shoot growth of shortleaf and l o b l o l l y pines in r e l a t i o n to certain environmental conditions. Duke University School of Forestry B u l l . 4. 52 P. 144 Reynolds, E.R.C. 1970. Root d i s t r i b u t i o n and the cause of i t s s p a t i a l v a r i a b i l i t y in Pseudotsuga taxi f o l i a (Poir.) B r i t t . Plant and S o i l 32: 501-517. Roberts, J . 1976. A study of root d i s t r i b u t i o n and growth in a Pinus s y l v e s t r i s L. (scots pine) plantation in East Anglia. Plant and S o i l 44: 607-621. Rogers, W.S. and G.A. Booth. 1959. The roots of f r u i t trees. S c i . Hort. 14: 27-34. Russell, R.S. 1977. Plant root systems: Their function and interaction with the s o i l . McGraw-Hill Book Co. (U.K.) Ltd. London 298 p. Safford, L.O. 1974. E f f e c t of f e r t i l i z a t i o n on biomass and nutrient cont of fine roots in a beech-birch-maple stand. Plant and S o i l 40:349-363. Safford, L.O. 1976. Seasonal v a r i a t i o n in the growth and nutrient content of yellow-birch replacement roots. Plant and S o i l 44: 439-444. Safford, L.O. and S. B e l l . 1972. Biomass of fine roots in a white spruce plantation. Can. J . For. Res. 2: 169-172. Santantonio, D. 1974. Root biomass studies of old-growth Douglas-fir. Oregon State University. 60 p. M.Sc. Thesis. Santantonio," D. 1978. Seasonal dynamics of fine roots in mature stands of Douglas-fir of d i f f e r e n t water regimes — A preliminary report. Pp. 190-203 in Riedacker, A. and J . Gagnaire-Michard, eds. Symposium: Root physiology and symbiosis, Nancy, September 1978. CNRF Champenoux - 54280 Seichamps, France. 502 p. Santantonio, D., R.K. Hermann, and W.S. Overton. 1977. Root biomass studies in forest ecosystems. Pedobiologia 17: 1-31 . Scholtes, W.H. 1953. The concentration of forest tree roots in the surface zone of some Piedmont s o i l s . Proceedings of the Iowa Academy of Science 60: 243-259. Shea, S.R. 1973. Growth and development of jack pine (Pinus  banksiana Lamb) in r e l a t i o n to edaphic factors in northeastern Ontario. University of Toronto, Dissertation. Simonovic, V. 1973. Study of the root biomass in the herb layer of an oak-hornbeam forest. Biologia (Bratislava) 28: 1:11-22. 145 Smith, J.H.G. 1964. Root spread can be estimated from crown width of Douglas-fir, lodgepole pine, and other B r i t i s h Columbia tree species. For. Chron. 40: 456-473. Sohlenius, B., H. Persson and C. Magnusson. 1977. Root-weight and nematode numbers in a young scots pine stand at Ivantjarnsheden, Central Sweden. Swedish Coniferous Forest Project, Barrskogslandskapets Ekologi, Technical Report 4. 22 p. Sokal, R.R. and F.J. Rohlf. 1969. Biometry: The p r i n c i p l e s and practice of s t a t i s t i c s in b i o l o g i c a l research. W.H. Freeman and Co. San Francisco. 776 p. Sutton, R.F. 1969. Form and development of conifer root systems. Tech. Commun. 7. Commonwealth For. B u l l . , Oxford, England. Tappeiner, J.C. I I . and H.H. John. 1973. Biomass and nutrient content of hazel undergrowth. Ecology 54: 1342-1348. Tisdale, S.L. and W.L. Nelson. 1975. S o i l f e r t i l i t y and f e r t i l i z e r s . MacMillan Publishing Co. Ltd. New York. 694 p. Torrey, J.G. and D.T. Clarkson, eds. 1975. The development and function of roots. Academic Press, New York. 618 p. T S l l e , H. 1967. Durchwurzelungsverhaltnisse m i t t e l a l t e r Kiefernbestande. (Root d i s t r i b u t i o n in middle aged pine stands.) Arch. Forstwes. 16(6/9): 775-779. Trewartha, G.T. 1968. An introduction to climate. 4 e d i t i o n . McGraw-Hill Book Co., New York. 693 p. USSR Academy of Sciences. 1968. Methods of productivity studies in root systems and rhizosphere organisms. Distributed by IBP Central O f f i c e c/o Linnean Soc. Burlington House, P i c c a d i l l y , London, W1V OLQ. Published by Nauka, Leningrad. 240 p. Vogt, K.A., R.L. Edmonds, C.C. Grier, and S.R. Piper. 1980. Seasonal changes in mycorrhizal and fibrous-textured root biomass in 23- and 180-year-old P a c i f i c s i l v e r f i r stands in western Washington. Can. J . For. Res. 10: 523-529. Vogt, K.A., R.L. Edmonds, and C.C. Grier. 1981. Seasonal changes in biomass and v e r t i c a l d i s t r i b u t i o n of mycorrhizal and fibrous-textured conifer fine roots in 23- and 180-year-old subalpine Abies amabilis stands. Can. J. For. Res. 11: 223-229. Weaver, T. 1977. Root d i s t r i b u t i o n and s o i l water regimes in nine habitat types of the northern Rocky Mountains. Pp. 146 239-244 in J.K. Marshall, ed. The belowground ecosystem: A synthesis of plant-associated processes. Range Science Dept. Series No. 26. Colorado State University, Fort C o l l i n s . Wiebe, H.H., G.S. Campbell, W.H. Gardner, S.L. Rawlins, J.W. Cary, and R.W. Brown. 1971. Measurement of plant and s o i l water status. Utah A g r i c u l t u r a l Experiment Station B u l l e t i n 484. 71 p. Wilcox, H.E. 1968. Morphological studies of the root of red pine, Pinus resinosa I. growth c h a r a c t e r i s t i c s and patterns of branching. American J . of Bot. 55(2): 247-254. Wilson, S.A. and D. Atkinson. 1978. Water and mineral uptake by f r u i t tree roots. Pp. 372-382 ijn Riedacker, A., J . Gagnaire-Michard (eds.). Symposium: root physiology and symbiosis. Nancy. France. 502 p. Yamakura, T., H. Saito, and T. Shidei. 1972. Production and structure of underground part of hinoki (Chamaecyparis  obtusa) stand. 1. Estimation of root production by means of root analysis. J. Jpn. For. Soc. 54: 118-125. Yarie,*J. 1978. The role of understory vegetation in the nutrient cycle of forested ecosystems in the Mountain Hemlock Biogeoclimatic Zone. Faculty of Forestry, University of B r i t i s h Columbia, Vancouver, B r i t i s h Columbia. 176 p. Dissertation. Z S t t l , H. 1964. [ F e r t i l i z a t i o n and fine-root d i s t r i b u t i o n in spruce stands.] Staatsforestverwaltung; Bayerns Mitteilungen 34: 333-342. 147 Appendix 1. Summary o f temperature data f o r the Oplopanaco ( h o r r i d i ) -Thujetum p l i c a t a e ( 0 ) , Ab ie to ( a m a b i l i s ) - Tsugetum mertensianae ( A ) , and V a c c i n i o (membranacei) - Tsugetum mertensianae (V) a s s o c i a t i o n s and a nearby c l e a r c u t (C) which was 1 km SE of the H a s s o c i a t i o n , 190 m lower i n e l e v a t i o n on a s l o p e o f 30%, and had an e a s t e r l y aspec t . F = October , November, December; W = Janua ry , February , March; Sp = A p r i l , May, June ; Su = J u l y , August , September. S t a t i o n 1975 1976 1977 Loca t ion F W Sp Su F W Sp Su F o r Abso lu te C -3 -17 -6 2 -5 -8 -4 2 -16 q u a r t e r l y 0 -17 -16 -9 4 -8 -11 -6 0 minimum A -16 -17 -8 4 - 7 * -9 -5 4 _ V -16 -17 -9 3 -8 -11 -7 -1 -Mean C -1 -3 3 9 ' 2 -2 4 9 - 3 * q u a r t e r l y 0 -2 -4 1 7 1 -3 3 8 _ minimum A -3 -4 1 8 2* -2 4 9 _ V -3 -5 1 8 1 -3 2 8 -Mean • C 1 -1 6 13 5 1 8 13 0* q u a r t e r l y 0 1 -2 4 10 3 -1 6 11 A - 1 * -2 4 11 3* 0 6 12 _ V - 1 * -2 3 10 3 -1 5 10 -Mean C 4 1 10 16 8 3 11 17 2* q u a r t e r l y 0 1 0 6 13 5 1 9 14 -maximum A 1* -1 6 14 5* 2 9 15 -V 0 -1 6 13 5 1 8 13 -Abso lu te C 10 7 18 23 14 9 20 23 3* q u a r t e r l y 0 5 5 13 20 9 4 15 17. -maximum A 4 4 15 21 11 4 16 19 -V 7 4 15 20 10 2 15 18 -Mean C _ 5. 7 5 .4 annual 0 - 3. 8 A - 4. 1 V - 3. 5 * Par t o f the data f o r one month i s m i s s i n g . Appendix 2. F ros t f r e e per iods f o r the p lan t a s s o c i a t i o n s and the c l e a r c u t . Oplopanaco ( h o r r i d i ) - Ab ie to (amab i l i s ) - V a c c i n i o (membranacei) CI P3 K T U t Thujetum p l i c a t a e Tsugetum mertensianae Tsugetum mertensianae 2 June to 22 Oct . 1976 = 142 days 2 June to 16 Oct . 1976 136 days 25 June to 22 Oct . 1976 =119 days 25 June to 22 Oct . 1976 119 days 26 May t o 8 Oct . 1977 = 135 days 2 June to 1 Oct . 1977 = 121 days 2 June to 1 Oct . 1977 = 121 days 2 June to 19 Sept . 1977 = 109 days Appendix 3. Precipitation for the clearcut at 1060 m which was 1 km SE of the Oplopanaco (horridi) - Thujetum plicatae Association, 190 m lower in elevation on a slope of 30% and had an easterly aspect. W = January, February, March; Sp = April, May, June; Su = July, August, September; F = October, November, December. 1975 1976 1977 1978 mm W __* 854 485 514 Sp 331 322 Su 346 278 F 1137 562 755 Total 2093 1841 Not available. 150 Appendix 4. Resu l t s o f a n a l y s i s of root t i p counts . The number of root t i p s was counted wh i l e the ove rs to r y < 2 mm roo ts were spread onto the g l ass p l a t e f o r the measurements r e l a t e d to leng th and su r face area (Sec t i on 4 . 2 . 2 ) . Unsuber ized t i p s were counted as they were removed from the s u b e r i z e d r o o t s . Suber i zed roo t t i p s ( u s u a l l y b lack and covered in fungal myce l ia ) were counted on the root from which they were growing. A f t e r count ing the t i p s , t h e i r number was c a l c u l a t e d on the bas i s o f a u n i t volume of s o i l . These data appear i n Appendix 5. Some root t i p s were l o s t dur ing the washing p r o c e s s , t h e r e f o r e the va lues repor ted are underes t imates . Regress ion a n a l y s i s w i th the < 2 mm roo t mass-volume as the indepen-dent v a r i a b l e and sube r i zed roo t t i p s as the dependent v a r i a b l e produced the f o l l o w i n g r e l a t i o n s h i p : # Suber ized T ips = 1133 Mass The s tandard e r r o r o f Y was 1210 and the r 2 va lue was 0 . 8 1 . With such a good c o r r e l a t i o n the t rend i n v e r t i c a l d i s t r i b u t i o n was not ana l yzed . I t would probably have been s i m i l a r t o the r e s u l t s repor ted in t h i s t h e s i s f o r v e r t i c a l d i s t r i b u t i o n of < 2 mm r o o t s . When summarized by p lan t a s s o c i a t i o n , t i p s of ove rs to ry roo ts demon-s t r a t e d tendenc ies s i m i l a r to those of leng th and su r f ace area d i scussed i n t h i s t h e s i s (see f i g u r e be low) . There was a marked d i f f e r e n c e between the OPHO THPL w i th 3.8 x 10 5 sube r i zed (o ld ) t i p s m 2 and the ABAM TSME and VAME TSME a s s o c i a t i o n s w i t h 7.4 x 10 5 sube r i zed t i p s m 2 . Unsuber ized (new) t i p s comprised a smal l amount of the t o t a l number of t i p s ( r a t i o o f unsuber ized t o sube r i zed t i p s was 0 .0725) . There were 104, 118, and 116 t i p s per meter o f ove rs to r y < 2 mm roo ts f o r t o t a l t i p s and 8 . 6 , 8 . 6 , and 6.6 t i p s per m f o r unsuber ized t i p s i n the OPHO THPL, ABAM TSME, AND VAME TSME a s s o c i a t i o n s r e s p e c t i v e l y . Appendix 4 c o n ' t 6 0 0 7 0 0 6 0 0 5 0 0 — 4 0 0 3 0 0 O L D O L D O L D 2 0 0 1 0 0 — — N E W N E W NEW 0 OPHO THPL A B A M T S M E VAME T S M E T IPS x 1 0 3 -2 m I n t e r p r e t a t i o n s s i m i l a r to those d i scussed e a r l i e r i n r e l a t i o n to n u t r i e n t supp ly and roo t growth cou ld be a p p l i e d to the root t i p data a l s o . That i s : there would be fewer root t i p s requ i red on the OPHO THPL than on the o ther a s s o c i a t i o n s because i t i s e a s i e r f o r the p l an t s to ob ta in n u t r i e n t s . The d i f f e r e n c e i n t h i s case i s tha t the ABAM TSME and VAME TSME are q u i t e s i m i l a r . The r e l a t i v e l y constant number of root t i p s per u n i t length o f roo t may be under gene t i c c o n t r o l by the t r e e s , o r i t may be e n v i r o n -men ta l l y induced . Appendix 5. Root mass-volume, su r face a r e a , number of t i p s , l e n g t h , and diameter i n r e l a t i o n to p lan t asso -c i a t i o n , p l o t , dep th , and t ime . U N S U B E R I Z E D OVERSTORY ROOTS B I O M A S S S U R F A C E T I P S LENGTH D I A M E T E R AREA I T E PLOT T I M E D E P T H 3 G / D M 2 3 CM / D M 3 / D M 3 C M / D M MM ^ *' 1 1 0 . 0 0 6 1 9 1 . 107 61 . 2 1 0 . 5 5 0 . 3 3 1 1 2 1 0 . 0 0 6 3 8 0 . 8 7 7 48 . 7 8 . 5 2 0 . 3 3 1 1 3 1 0 . 0 0 0 5 0 0 . 133 17 . 4 1 . 5 4 0 . 2 7 1 1 4 1 0 . 0 0 1 0 9 0 . 162 6 . 3 1 . 4 3 0 . 3 6 1 1 5 1 0 . 0 0 0 4 3 0 . 0 7 1 7 . 3 0 . 8 9 0 . 2 5 1 1 6 1 0 . 0 0 0 6 1 0 . 1 1 4 7 . 7 1 . 3 0 0 . 2 8 1 1 7 1 0 . 0 0 0 4 6 0 . 154 3 .4 1 . 0 5 0 . 4 7 1 2 1 1 0 . 0 0 3 1 1 0 . 6 4 1 146 . 9 8 . 3 9 0 . 2 4 1 2 2 1 0 . 0 0 1 7 3 O . 3 2 8 147 . 9 4 . 8 8 0 . 2 1 1 2 3 1 0 . 0 0 0 7 9 0 . 148 27 . 5 1 . 4 5 0 . 3 3 1 2 4 1 0 . 0 0 0 5 9 0 . 136 2 2 . 2 1 . 3 3 0 . 3 2 1 2 5 1 0 . 0 0 0 1 2 0 . 0 2 9 8 . 9 0 . 4 1 O . 2 3 1 2 e 1 0 . 0 0 0 2 0 0 . 0 5 8 2 . 6 0 . 3 6 0 . 5 1 1 2 7 1 0 . 0 0 0 0 0 0 . 0 0 1 0 .4 0 . 0 2 0 . 2 0 1 3 1 1 0 . 0 0 1 2 0 0 . 4 4 6 8 6 . 5 4 . 7 0 0 . 3 0 1 3 2 1 0 . 0 0 0 3 4 0 . 154 32 . 6 1 . 4 7 0 . 3 3 1 3 3 1 0 . 0 0 1 6 2 0 . 3 3 8 3 3 . 6 2 . 0 7 0 . 5 2 1 3 4 1 0 . 0 0 . 0 0 0 0 . 0 0 . 0 1 3 5 1 0 . 0 0 . 0 0 0 0 . 0 0 . 0 1 3 6 1 O . 0 O 0 2 8 0 . 0 5 9 9 3 0 . 4 8 0 . 3 9 1 3 7 1 0 . 0 0 0 1 5 0 . 0 3 1 4 2 0 . 2 6 0 . 3 9 2 1 1 1 0 . 0 0 5 1 7 1 . 5 6 2 2 4 2 4 1 2 . 7 9 0 . 3 9 2 1 2 1 0 . 0 0 5 0 6 1 . 4 2 2 135 0 8 . 3 0 0 . 5 5 2 1 3 1 0 . 0 0 0 4 7 0 . 168 5 3 0 2 . 1 0 0 . 2 5 2 1 4 1 0 . 0 0 0 0 9 0 . 0 3 3 13 0 0 . 4 5 0 . 2 3 2. 1 5 1 0 . 0 0 . 0 0 0 0 . 0 0 . 0 2 1 6 1 0 . 0 0 . 0 0 0 0 . 0 0 . 0 2 1 7 1 0 . 0 0 . 0 0 0 0 . 0 0 . 0 2 2 1 1 0 . 0 0 4 9 4 1 . 5 5 4 4 9 4 5 1 8 . 5 4 0 . 2 7 2 2 2 1 0 . 0 0 1 0 0 0 . 165 7 9 0 . 2 . 6 3 0 . 2 0 2 2 3 1 0 . 0 0 0 3 7 0 . 128 3 0 3 1 . 4 4 0 . 2 8 2 2 4 1 0 . 0 0 0 5 8 0 . 136 3 6 5 1 . 7 0 0 . 2 5 2 2 5 1 0 . 0 0 0 1 0 0 . 0 4 6 10 0 0 . 3 8 0 . 3 8 2 2 6 1 0 . 0 0 0 1 1 0 . 0 1 3 3 6 0 . 13 0 . 3 1 2 2 7 1 0 . 0 0 0 0 8 0 . 0 7 7 2 2 4 0 . 9 0 0 . 2 7 2 3 1 1 0 . 0 0 0 3 2 0 . 7 4 8 2 2 7 5 8 . 7 5 0 . 2 7 2 3 2 1 0 . 0 0 1 1 5 0 . 2 3 9 103 7 3 . 8 0 0 . 2 0 2 3 3 1 0 . 0 0 0 5 7 0 . 169 14 1 1 . 2 0 0 . 4 5 2 3 4 1 0 . 0 0 0 1 2 0 . 0 6 9 1 1 . 8 0 . 6 3 0 . 3 5 2 3 5 1 0 . 0 0 0 0 1 0 . 0 2 1 2 . 8 0 . 1 5 0 . 4 5 2 3 6 1 0 . 0 0 . 0 0 . 0 0 . 0 0 . 0 2 3 7 1 0 . 0 0 0 0 0 0 . 0 0 4 1 . 5 0 . 0 6 0 . 2 0 3 1 1 1 0 . 0 0 1 4 0 0 . 2 8 1 2 7 . 3 2 . 0 3 0 . 4 4 3 1 2 1 0 . 0 0 0 5 3 0 . 186 4 2 . 3 1 . 9 6 0 . 3 0 3 1 3 1 0 . 0 0 0 3 4 0 . 0 5 4 2 2 . 7 0 . 72 0 . 2 4 S i t e : 1 = OPHO THPL 2 = ABAM TSME 3 = VAME TSME P l o t : as marked Depth (cm): 1 = 0 - 5 , 2 = 5 -10 , 3 = 10-20 , 4 = 20-30 5 = 30 -40 , 6 = 40 -50 , 7-= 50-60 Time: 1 = 2-9 June , 2 = 18-21 J u l y , 3 = 14-17 September Appendix 5 c o n ' t . . . U N S U B E R I Z E D OVERSTORY ROOTS B I O M A S S S U R F A C E T I P S L E N G T H D I A M E T E R AREA P L O T T I M E 3 2 3 3 3 S I T E D E P T H G / D M CM / D M / D M C M / D M MM 3 1 4 1 0 . 0 0 0 2 9 0 . 0 5 4 2 2 . 4 0 . 8 5 0 . 2 0 3 1 5 1 0 . 0 0 . 0 0 . 0 0 . 0 0 . 0 3 1 6 1 0 . 0 0 0 0 2 0 . 0 1 0 0 . 5 0 . 0 6 0 5 0 3 1 7 1 0 .OOOO1 0 . 0 0 1 0 . 3 0 . 0 1 0 2 0 3 2 1 1 0 . 0 0 0 9 4 0 109 15 .8 1 0 4 0 3 3 3 2 2 1 0 0 0 0 5 5 0 187 74 4 2 9 8 0 2 0 3 2 3 1 0 0 0 0 2 0 0 0 4 0 14 5 0 5 5 0 2 3 3 2 4 1 0 0 0 0 0 1 0 0 0 3 1 8 0 0 4 0 2 0 3 2 5 1 0 0 0 0 0 0 0 0 0 1 0 7 0 0 2 0 2 0 3 2 6 1 0 0 0 0 0 3 0 0 0 1 0 6 0 0 2 0 2 0 3 2 7 1 0 0 0 0 0 0 0 0 0 0 3 3 1 1 0 0 0 4 8 0 1 4 4 3 2 8 8 5 13 61 0 3 4 3 3 2 1 0 0 0 1 7 S 0 6 0 4 2 2 9 4 8 2 2 0 2 3 3 3 3 1 0 0 0 0 6 0 0 174 5 7 2 2 3 0 O 2 4 3 3 4 1 0 0 0 0 5 6 0 107 3 0 3 1 44 0 . 24 3 3 5 1 0 0 0 0 1 7 0 0 8 0 13 0 0 6 2 0 . 4 2 3 3 6 1 0 0 0 0 1 2 0 0 5 1 10 1 0 46 0 . 3 6 3 3 7 1 0 0 0 0 0 0 0 0 0 2 0 9 0 0 3 0 2 0 1 1 1 2 0 0 1 8 6 0 2 6 7 7 3 0 4 4 2 3 34 0 37 1 1 2 2 0 0 0 3 5 8 1 3 6 4 2 0 7 8 13 3 9 0 . 3 2 1 1 3 2 0 0 0 1 7 2 0 5 1 9 7 0 9 4 3 6 0 3 8 1 1 4 2 0 0 0 1 6 2 0 5 0 4 8 3 1 5 3 5 0 3 0 1 1 5 2 0 0 0 0 4 7 0 164 5 1 1 0 9 0 4 8 1 1 6 2 0 0 0 0 5 1 0 2 0 0 17 0 1 4 3 0 . 4 5 1 1 7 2 0 0 0 1 6 6 0 4 4 7 4 7 0 3 5 4 0 4 0 1 2 1 2 0 0 2 2 4 0 6 9 8 2 5 8 9 2 4 9 7 5 0 4 5 1 2 2 2 0 0 0 2 2 3 0 8 5 5 127 4 7 16 0 3 8 1 2 3 2 0 0 0 0 3 2 0 108 28 1 1 2 3 0 28 1 2 4 2 0 0 0 0 1 1 0 0 2 8 4 4 0 24 0 3 6 1 2 5 2 0 0 0 0 3 2 0 121 16 9 1 0 3 0 3 7 3 3 5 1 0 0 0 0 1 7 0 0 8 0 13 0 0 6 2 0 4 2 3 3 6 1 0 0 0 0 1 2 0 0 5 1 1 0 1 0 4 6 0 3 6 3 3 7 1 0 0 0 0 0 0 0 0 0 2 0 9 0 0 3 0 2 0 1 1 1 2 0 0 1 8 6 0 2 6 7 7 3 0 4 4 2 3 34 0 3 7 1 1 2 2 0 0 0 3 5 8 1 3 6 4 2 0 7 8 13 3 9 0 3 2 1 1 3 2 0 0 0 1 7 2 0 5 1 9 7 0 9 4 3 6 0 3 8 1 1 4 2 0 0 0 1 6 2 0 5 0 4 8 3 1 5 3 5 0 3 0 1 1 5 2 0 0 0 0 4 7 0 164 5 1 1 0 9 0 48 1 1 6 2 0 0 0 0 5 1 0 2 0 0 17 0 1 4 3 0 4 5 1 1 7 2 0 0 0 1 6 6 0 4 4 7 4 7 0 3 5 4 0 4 0 1 2 1 2 0 0 2 2 4 0 6 9 8 2 5 8 9 2 4 9 7 5 0 4 5 1 2 2 2 0 0 0 2 2 3 0 8 5 5 127 4 7 16 0 3 8 1 2 3 2 0 0 0 0 3 2 0 108 28 1 1 2 3 0 2 8 1 2 4 2 0 0 0 0 1 1 0 0 2 8 4 4 0 24 0 3 6 1 2 5 2 0 0 0 0 3 2 0 121 16 9 1 0 3 0 3 7 1 2 6 2 0 0 0 0 0 6 0 0 5 8 13 3 0 6 5 0 2 9 •| 2 7 2 0 0 0 0 3 7 0 160 2 7 3 1 5 9 0 3 2 1 3 1 2 0 0 0 3 0 4 1 0 6 8 138 2 11 4 7 0 3 0 1 3 2 2 0 0 0 0 5 0 0 196 2 8 4 2 13 0 2 9 1 3 3 2 0 0 0 0 0 0 0 0 0 0 1 3 4 2 0 O 0 0 0 0 0 0 0 0 1 3 5 2 0 0 0 0 0 0 0 0 0 0 1 3 6 2 0 0 0 0 0 1 0 0 0 2 0 8 0 0 3 0 2 0 1 3 7 2 0 0 0 0 0 1 0 0 1 0 2 0 0 12 0 2 6 2 1 1 2 0 0 1 6 8 8 5 2 5 4 1849 0 8 2 0 0 0 2 0 2 1 2 2 0 0 0 5 1 6 2 3 1 3 3 5 3 1 2 2 0 0 • 0 3 3 2 1 3 2 0 0 0 2 7 4 1 178 129 8 10 9 6 0 34 2 1 4 2 0 0 0 1 14 0 3 6 8 21 5 3 0 0 0 3 9 2 1 5 2 0 0 0 0 1 4 0 1 10 4 7 3 1 7 4 0 2 0 2 1 6 2 0 0 0 0 5 4 0 163 14 2 1 31 0 3 9 2 1 7 2 0 0 0 0 0 7 0 0 1 4 3 6 0 2 0 0 2 2 2 2 1 2 0 0 1 7 1 3 6 2 6 2 6 8 2 8 5 5 9 4 o 3 6 Appendix 5 con't 154 UNSUBERIZED OVERSTORY ROOTS BIOMASS SURFACE TIPS LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 3 SITE DEPTH G/DM CM /DM /DM CM/DM MM 2 2 2 2 0 .00762 2 959 214 9 23 06 0 41 ' 2 2 3 2 0 01259 4 056 328 2 28 68 0 45 2 2 4 2 0 00254 0 883 71 5 6 43 0 44 2 2 5 2 0 0 0 0 O 0 O 0 0 0 2 2 6 2 0 00002 0 007 3 6 0 1 1 0 20 2 2 7 2 0 00023 0 057 5 8 0 48 0 38 2 3 1 2 0 00940 2 485 358 2 23 13 0 34 2 3 2 2 0 00534 1 953 178 0 12 34 •o 50 2 3 3 2 0 001 14 0 408 68 6 3 71 0 35 2 3 4 2 0 00026 0 146 21 2 1 27 0 37 2 3 5 2 0 00060 0 200 23 1 " 1 60 0 40 2 3 6 2 0 00023 0 071 14 1 0 72 0 31 2 3 7 2 0 00009 0 017 3 0 0 18 0 31 3 1 1 2 0 01290 4 470 332 1 33 91 0 42 3 1 2 2 0 00210 1 249 241 0 10 84 0 37 3 1 3 2 0 00202 0 857 124 0 7 44 0 37 3 . 1 4 2 0 00102 0 277 30 0 2 77 0 32 3 1 5 2 0 00041 0 125 5 5 1 14 0 35 3 1 6 2 0 00040 0 080 3 6 0 55 0 46 3 1 7 2 0 0001 1 O 01 1 0 7 0 07 0 50 3 2 1 2 0 03320 12 500 1121 0 91 38 0 44 3 2 2 2 0 00229 0 973 152 6 9 99 " 0 31 3 2 3 2 0 00139 0 554 46 4 3 71 0 48 3 2 4 2 0 00006 0 051 7 4 0 48 0 34 3 2 5 2 0 00020 0 089 17 9 0 85 0 33 3 2 6 2 0 00002 0 010 2 4 0 13 0 26 3 2 7 2 0 00003 0 013 2 1 0 12 0 36 3 3 1 2 0 03762 13 790 912 9 115 90 0 38 3 3 2 2 0 00322 0 931 193 2 10 95 0 27 3 3 3 2 0 00164 0 457 84 6 4 74 0 31 3 3 4 2 0 00242 0 920 84 5 6 98 0 42 3 3 5 2 0 001 12 0 203 14 7 0 99 0 65 3 • 3 6 2 0 00036 0 127 1 1 6 0 88 0 46 3 3 7 2 0 00003 0 004 1 1 0 04 0 30 1 1 1 3 0 03446 1 1 990 997 1 87 82 0 44 1 1 2 3 0 00322 2 549 472 3 25 76 0 31 1 1 3 3 0 00969 2 898 207 7 24 98 0 37 1 1 4 3 0 00038 0 154 18 9 1 25 0 39 1 1 5 3 0 00055 0 127 17 9 1 13 0 36 1 1 6 3 0 00054 0 250 18 7 1 75 0 46 1 1 7 3 0 00006 0 023 2 6 0 18 0 41 1 2 1 3 0 00369 1 682 237 0 14 84 0 36 1 2 2 3 0 00045 0 179 44 5 2 09 0 27 1 2 3 3 0 00054 0 170 18 8 1 64 0 33 1 2 4 3 0 00027 0 070 16 5 0 76 0 29 1 2 5 3 0 00001 0 005 1 9 0 06 0 30 1 2 6 3 0 00003 0 019 4 6 0 18 0 32 1 2 7 3 0 00005 0 009 3 8 0 14 0 20 1 3 1 . 3 0 00519 1 898 415 0 18 91 0 32 1 3 2 3 0 00021 0 128 28 5 1 47 0 28 1 3 3 3 0 00005 0 030 1 1 0 0 39 0 24 1 3 4 3 0 00001 0 006 1 4 0 09 0 20 1 3 5 3 0 00004 0 020 3 9 0 25 0 25 1 3 6 3 0 00006 0 018 4 2 0 20 0 29 1 3 7 3 0 oooo i 0 003 " 1 0 6 05 0 23 2 1 1 3 0 00734 2 974 577 0 28 59 0 33 2 1 2 3 . 0 00826 1 920 165 1 12 66 0 48 2 1 3 3 0 00046 0 171 31 6 1 51 0 36 2 1 4 3 0 00005 0 037 18 0 0 59 0 20 2 1 5 3 0 O0015 0 198 58 3 2 57 0 25 2 1 6 3 0 00003 0 021 3 7 0 20 0 33 2 1 7 3 0 00000 0 001 0 2 0 01 0 20 Appendix 5 con't UNSUBERIZED OVERSTORY ROOTS BIOMASS SURFACE TIPS LENGTH DIAMETER AREA ITE PLOT DEPTH TIME 3 G/DM CM 2 3 /DM 3 /DM 3 CM/DM MM 2 2 1~ 3 0 .00192 0 .636 162 .5 7 .46 0 .27 2 2 2 3 0 .00681 3 .533 374 .0 30 .05 0 . 37 2 2 3 3 0 .00241 1 . 154. 137 .8 10 .48 0 . 35 2 2 4 3 0 .00127 0 .568 97 . 1 6 . 34 0 .29 2 2 5 3 0 .00041 0 . 137 23 .3 1 .25 0 .35 2 2 6 3 0 .00063 0 . 161 23 8 1 .70 0 30 2 2 7 3 0 .00037 0 131 29 5 1 55 0 27 2 3 1 3 0 .00943 2 837 309 5 22 79 0 40 2 3 2 3 0 00431 1 714 204 7 17 48 0 31 2 3 3 3 0 00221 1 044 33 4 5 34 0 62 2 3 4 3 0 00191 0 943 46 6 6 83 0 44 2 3 5 3 0 00002 0 029 7 6 0 38 0 25 2 3 6 3 0 00006 0 020 • 1 9 0 15 0 41 2 3 7 3 0 00000 0 000 0 2 0 01 0 20 3 1 1 3 0 01440 8 1 10 617 5 47 63 0 54 3 1 2 3 0 00230 1 524 263 1 14 64 0 33 3 1 3 3 0 00088 0 386 153 5 4 39 0 28 3 1 4 3 0 00042 0 135 26 8 1 38 0 31 3 1 5 3 0 00006 0 086 10 6 0 75 0 37 3 1 6 3 0 00041 0 143 24 5 1 39 0 33 3 1 7 3 0 00036 0 181 33 6 1 96 0 29 3 2 1 3 0 00335 1 825 177 2 14 27 0 41 3 2 2 3 0 00215 1 059 214 9 10 74 0 31 3 2 3 3 0 00088 0 343 71 7 4 15 0 26 3 2 4 3 0 00037 0 187 29 6 1 85 0 32 3 2 5 3 0 00018 0 073 13 1 0 78 0 30 3 2 6 3 0 00004 0 024 4 5 0 22 0 34 3 2 7 3 0 0OOO2 0 016 1 4 0 12 0 41 3 3 1 3 0 00687 2 157 389 1 19 69 0 35 3 3 2 3 0 00240 1 177 156 0 1 1 40 0 33 3 3 3 3 0 00013 0 080 25 4 1 10 0 23 3 3 4 3 0 00005 0 048 19 1 0 76 0 20 3 3 5 3 0 0OOO4 0 025 9 2 0 41 0. 20 3 3 6 3 0 00022 0 087 18. 4 0. 90 0. 31 3 3 7 3 o 00023 o 079 1 1 . 0 0. 68 o. 37 OVERSTORY < 2 MM ROOTS BIOMASS SURFACE TIPS LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 3 SITE DEPTH G/DM CM /DM /DM CM/DM MM 1 1 1 1 2 1622 355 2364 2350 0 48 1 1 2 1 1 7970 297 1663 1630 0 58 1 1 3 1 1 3240 144 411 750 0 61 1 1 4 1 1 5050 145 248 640 0 73 1 1 5 1 0 9147 95 168 410 0 75 1 1 6 1 0 3495 39 123 170 0 73 1 1 7 1 0 3685 37 40 1 10 1 03 1 2 1 1 3 8332 587 2798 2780 0 67 1 2 2 1 2 5200 394 1576 2590 0 48 1 2 3 1 1 9870 226 553 1090 0 66 1 2 4 1 0 9138 121 351 570 0 67 1 2 5 1 0 4750 65 160 290 0 71 1 2 6 1 0 2597 36 57 140 0 80 1 2 7 1 0 0589 7 8 20 0 94 1 3 1 1 1 8347 280 632 1550 0 57 1 3 i 2 1 0 9002 234 1773 1220 0 61 Appendix 5 con't OVERSTORY < 2 MM ROOTS BIOMASS SURFACE TIPS LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 3 SITE DEPTH G/DM CM /DM /DM CM/DM MM 1 3 3 1 0 .8082 152 826 820 0.59 1 3 4 1 0 .4053 63 652 500 0.40 1 3 5 1 0 .6350 1 10 653 760 0.46 1 3 6 1 0 .4531 64 210 370 0.54 1 3 7 1 0 . 1738 29 70 140 0.65 2 1 1 1 3 .7957 512 4937 3340 0.49 2 1 2 1 3 6812 449 3920 2810 0.51 2 1 3 1 2 0873 268 1362 1440 0.59 2 1 4 1 0 6973 77 369 410 0.60 2 1 5 1 0 0482 5 10 30 0.68 2 1 e 1 0 0063 1 3 10 0.67 2 1 7 1 0 0031 1 0 00 0.77 2 2 1 1 6 1201 1204 12670 8330 0.46 2 2 2 1 5 7103 526 6634 3720 0.45 2 2 3 1 2 2820 309 1455 1280 0.77 2 2 4 1 1 0012 134 • 359 650 0.65 2 2 5 1 0 7319 92 356 490 0.60 2 2 6 1 0 4853 93 354 470 0.64 2 2 7 1 0 6443 79 289 450 0.56 2 3 1 1 5 9684 962 7946 6120 0.50 2 3 2 1 2 5776 464 3962 3040 0.49 2 3 3 1 2 9641 412 2363 2520 0.52 2 3 4 1 1 1221 113 608 630 0. 57 2 3 5 1 0 3514 45 181 260 0.55 2 3 6 1 0 0225 4 9 20 0.73 2 3 7 1 0 2043 28 69 140 0.65 3 1 1 1 3 2057 439 3303 2410 0.58 3 1 2 1 3 4379 479 3527 2510 0.61 3 1 3 1 2 8943 374 2659 2190 0.54 3 1 4 1 2 1270 178 730 890 0.64 3 1 5 1 0 9392 78 44 1 440 0.56 3 . 1 6 1 0 1608 17 33 80 0.67 3 1 7 1 0 0542 6 14 30 0.59 3 2 1 1 5 1763 539 2271 2340 0.73 3 2 2 1 2 0194 235 1426 1420 0. 53 3 , 2 3 1 1 3570 124 757 670 0. 59 3 ' 2 4 1 0 2874 32 200 200 0.52 3 2 5 1 0 3135 39 119 150 0.82 3 2 6 1 0 1655 17 48 60 0.85 3 2 7 1 0 0 0 0 00 0.0 3 3 1 1 4 8026 565 8575 4320 0.42 3 3 2 1 4 8778 558 9019 4200 0.42 3 .3 3 1 1 9964 256 2583 1580 0.51 3 3 4 1 1 8343 207 1731 1250 0.53 3 3 5 1 1 5042 158 685 790 0.64 3 3 6 1 1 3916 136 464 610 0.71 3 3 7 1 0 7743 77 323 330 0.74 1 1 1 2 4 4454 422 8962 3800 0.35 1 1 2 2 2 9125 258 1986 1470 0.56 1 1 3 2 1 1291 1 18 649 620 0.60 1 1 4 2 1 . 3152 213 858 1200 0.57 1 1 5 2 0. 4220 52 127 280 0.59 1 1 6 2 0. 9375 1 18 387 600 0.63 1 1 7 2 0. 7833 143 369 750 0.61 1 2 1 2 1 . 8953 373 2946 2130 0.56 1 2 2 2 1 . 2130 169 963 980 0.55 1 2 3 2 0. 3784 72 503 480 0.48 1 2 4 2 0. 2227 44 241 260 0.54 1 2 5 2 0. 3294 51 205 310 0.53 1 2 6 2 0. 2753 46 185 250 0.59 Appendix 5 con't OVERSTORY < 2 MM ROOTS BIOMASS SURFACE TIPS LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 3 SITE DEPTH G/DM CM /DM /DM CM/DM MM 1 2 7 2 0. 1422 22 90 130 0.57 1 3 1 2 1.8939 286 3760 2240 0.41 1 3 2 2 0.4829 62 767 420 0.46 1 3 3 2 0.2321 38 1 18 160 0.77 1 3 4 2 0.2297 30 35 1 10 0.84 1 3 5 2 0.0642 9 21 50 0.63 1 3 6 2 0.0942 8 17 40 0.62 1 3 7 2 0.1457 22 34 90 0.79 2 1 1 2 11.5130 1561 21 140 10350 0.48 2 1 2 2 3.0423 324 3966 2330 0.44 2 1 3 2 1.3644 192 1233 1350 0.45 2 1 4 2 1.7808 178 887 1230 0.46 2 1 5 2 0.6519 106 429 580 0. 58 2 1 e 2 0.3651 31 98 160 0.61 2 1 7 2 0.1462 24 23 1 10 0.72 2 2 1 2 2.1862 340 5151 3030 0.36 2 2 2 2 1.4850 189 2892 1720 0. 35 2 2 3 2 1.7899 256 2195 1630 0.50 2 2 4 2 1.6324 157 508 660 0.76 2 2 5 2 1.2659 113 268 500 0.71 2 2 6 2 0.6298 74 236 320 0.74 2 2 7- 2 0.4867 59 185 320 0.59 2 3 1 2 4.7039 533 4298 3630 0.47 2 3 2 2 2.7381 282 1661 1620 0.55 2 3 3 2 1.2133 138 660 950 0.46 2 3 4 2 0.6774 70 339 420 0.53 2 3 5 2 0.6892 74 243 330 0.72 2 3 6 2 0.8019 83 282 360 0.73 2 3 7 2 0.4795 41 86 150 0.89 3 1 1 2 5.3851 749 4999 5070 0.47 3. 1 2 2 8.1328 731 8253 4820 0.48 3 1 3 2 3.9989 463 3814 3090 0.48 3 1 4 2 0.9352 98 273 570 0.55 3 1 5 2 0.2123 21 101 170 0.40 3 1 6 2 0.0964 16 65 100 0.50 3 1 7 2 0.2512 25 65 120 0.66 3 2 1 2 6.7539 848 6937 5740 0.47 3 2 2 2 1.9280 218 1495 1520 0.46 3 2 3 2 1.2998 125 1 100 830 0.48 3 2 4 2 0.3236 48 266 310 0.49 3 2 5 2 . 0.3990 50 108 200 0.79 3 2 6 2 0.0344 4 22 20 0.68 3 2 7 2 0.0352 6 12 30 0.72 3 3 1 2 7.2476 805 • 6805 5650 0.45 3 3 2 2 4.8517 586 3897 3160 0.59 3 3 3 2 1.6357 149 880 850 0.56 3 3 4 2 1.1492 146 924 830 0.56 3 3 5 2 0.4956 67 1 17 340 0.62 3 3 6 2 0.3575 51 171 260 0.62 3 3 7 2 0.3276 34 136 120 0.91 1 1 1 3 3.6368 387 6053 3240 0.38 1 1 2 3 4.5724 546 6376 3410 0.51 1 1 3 3 0.9155 139 987 810 0.55 1 1 4 3 0.5805 58 223 320 0.58 1 1 5 3 0.4612 61 190 340 0.57 1 1 6 3 0.351 1 56 185 310 0.58 1 1 7 3 0. 1813 26 23 90 0.92 1 2 1 3 2.6804 410 1431 2080 0.63 1 2 2 3 1.0103 168 436 790 0.68 1 2 3 3 0.6943 88 424 540 0.51 Appendix 5 con't OVERSTORY < 2 MM ROOTS PLOT TIME SITE DEPTH BIOMASS 3 G/DM SURFACE AREA 2 3 CM /DM TIPS 3 /DM LENGTH 3 CM/DM DIAMETER MM 1 2 4 3 0.5863 83 200 430 0.61 1 2 5 3 0.4749 59 114 290 0.65 1 2 6 3 0.4715 57 59 230 0.80 1 2 7 3 0.7908 81 83 270 0.96 1 3 1 3 2.5363 275 1994 1850 0.47 1 3 2 3 0.5515 59 245 320 0.59 1 3 3 3 0.4579 49 135 210 0.77 1 3 4 3 0.0882 13 19 50 0.86 1 3 5 3 0.2614 27 65 1 10 0. 77 1 3 6 3 0.061 1 1 1 31 50 0.64 1 3 7 3 0.0927 12 28 50 0. 72 2 1 1 3 5.8249 777 , 5744 4820 0.51 2 1 2 3 2.1716 246 1018 1280 0.61 2 1 3 3 1.1862 1 14 253 650 0.56 2 1 4 3 0.5138 83 172 430 0.61 2 1 5 3 1.1759 146 568 780 0.60 2 1 6 3 0.2853 38 98 160 0. 74 2 1 7 3 0.0217 3 8 20 0.67 2 2 1 3 1.3079 233 1691 1670 0.44 2 2 2 3 2.7903 505 2578 2430 0.66 2 2 3 3 1.6697 215 965 980 0.70 2 2 4 3 1.1404 128 672 660 0.61 2 2 5 3 0.8772 107 262 370 0.93 2 2 6 3 0.6273 70 231 340 0.64 2 2 7 3 0.5414 41 180 2.10 0.62 2 3 1 3 3.3792 426 2068 2030 0.67 2 3 2 3 2.1014 271 899 1370 0.63 2 3 3 3 1.2526 165 322 730 0.72 2 3 4 3 1.2426 187 921 940 0.63 2 3 5 3 0.4424 50 262 290 0.54 2 3 6 3 0.1129 16 57 100 0.53 2 3 7 3 0.0072 1 4 10 0.54 3 1 1 3 5.1160 777 6410 4670 0.53 3 1 2 3 4.7828 625 3174 3930 0.51 3 1 3 3 4.2755 374 1414 1960 0.61 3 1 4 3 1.3763 185 475 820 0.72 3 1 5 3 0.8212 73 309 520 0.45 3 1 6 3 0.8702 102 269 500 0.65 3 1 7 3 0.7177 70 379 500 0.45 3 2 1 3 3.4532 356 1634 2200 0.52 3 2 2 3 4.5674 612 3323 3950 0.49 3 2 3 3 2.1196 273 686 1400 0.62 3 2 . 4 3 0.6413 71 193 370 0.62 3 2 5 3 0.3901 46 173 310 0.48 3 2 6 3 0.2218 25 86 160 0.52 3 2 7 3 0.1488 14 13 50 0.82 3 3 1 3 5.1046 606 6867 4320 0.45 3 3 2 3 3.1316 326 2233 1980 0.52 3 3 3 3 1.2139 134 589 760 0.56 3 3 4 3 0.8630 157 554 680 0.73 3 3 5 3 0.5040 69 197 350 0.63 3 3 6 3 0.3005 28 181 260 0.35 3 3 7 3 0.3895 45 166 230 0.61 Appendix 5 con't OVERSTORY 2-5 MM ROOTS BIOMASS SURFACE LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 SITE DEPTH G/DM CM /DM CM/DM MM ! ! 1 1 0.3037 12 1 16 0 2 4 1 1 2 1 0.2831 9 0 10 8 2 6 1 1 3 1 1.0460 29, 9 32 6 2 9 1 1 4 1 2.1294 62 9 65 9 3 0 1 1 5 1 0.9164 27 4 28 8 3 0 1 1 6 1 0.4405 15 4 17 4 2 8 1 1 7 1 0.4027 14 1 16 9 2 7 1 2 1 1 1.0331 33 4 32 8 3 2 1 2 2 1 0.9339 32 0 37 3 2 7 1 2 3 . 1 0.7878 19 5 18 8 3 3 1 2 4 1 0.8728 27 1 27 9 3 1 1 . 2 5 1 0.3803 15 6 17 7 2 8 1 2 6 1 0.3733 9 0 8 4 3 4 1 2 7 1 0.0408 1 6 1 3 3 8 1 3 1 1 0.6100 16 7 15 8 3 4 1 3 2 1 0.0843 2 8 3 4 2 6 1 3 3 1 0.1462 5 5 4 7 3 7 1 3 4 1 0.0441 2 7 4 2 2 0 1 3 5 1 0.6358 21 0 18 8 3 6 1 3 6 1 0.0617 2 5 2 9 2 8 1 3 7 1 0.0303 1 4 2 1 2 2 2 1 1 1 1.2846 37 9 37 7 3 2 2 1 2 1- 0.9087 30 9 33 8 2 9 2 1 3 1 1.7426 '58 1 66 4 2 8 2 1 4 1 0.4350 13 1 14 3 2 9 2 1 5 1 0.0311 1 4 1 1 4 1 2 1 6 1 0.0139 0 5 0 5 2 9 . 2 1 7 1 0.0 0 0 0 0 0 0 2 2 1 1 0.3882 10 1 9 7 3 3 2 2 2 1 1.0523 21 1 17 8 3 8 2 2 3 1 1.3609 32 8 30 5 3 4 • 2 2 4 1 0.6237 1 1 7 10 1 3 7 2 2 5 1 0.3849 10 9 10 9 3 2 2 2 6 1 0.0738 2 4 3 3 2 3 2 2 7 1 1.0477 25 8 26 1 3 1 2 3 1 1 1.2071 36 5 39 7 2 9 2 3 2 1 0.3280 9 0 8 4 3 4 2 3 3 1 1.9030 52 9 57 5 2 9 2 3 4 1 0.5821 15 3 13 0 3 8 2 3 5 1 0.0851 2 7 3 3 2 6 2 3 6 1 0.0 0 0 0 0 0 0 2 3 7 1 0.0 0 0 0 0 0 0 3 1 1 1 1.7089 34 0 29 1 3 7 3 1 2 1 1.1112 22 8 22 4 3 2 3 1 3 1 • 1.1024 27 3 29 3 3 0 3 1 4 1 0.9757 24 5 26 0 3 0 3 1 5 1 1.0893 24 7 25 3 3 1 3 1 6 1 0.1367 3 4 3 4 3 1 3 1 7 1 0.0652 1 5 1 5 3 1 3 2 1 1 1.8518 42 3 43 5 3 1 3 2 2 1 2.0648 52 8 54 6 3 1 3 2 3 1 1.4984 35 9 37 4 3 1 3 2 4 1 0.0 0 0 0 0 0 0 3 2 5 1 0.0904 3 3 4 7 2 3 3 2 6 1 0.1237 3 0 2 4 3 9 3 2 7 1 0.0 0 .0 0 .0 0 .0 3 3 1 1 1 .2988 26 .5 26 5 3 .2 3 3 2 1 1 .6546 34 1 33 4 3 2 3 3 3 1 1.5879 46 8 51 3 2 9 3 3 4 1 2.0821 56 3 61 0 2 9 3 3 5 1 1.5284 45 5 48 7 3 0 Appendix 5 con't OVERSTORY 2-5 MM ROOTS BIOMASS PLOT TIME 3 SITE DEPTH G/DM 3 3 6 1 1.0936 3 3 7 1 0.6796 1 1 1 2 0.0 1 1 2 2 0.9040 1 1 3 2 0.3720 1 1 4 2 1.1399 1 1 5 2 0.3858 1 1 6 2 0.5000 1 1 7 2 0.3620 1 2 1 2 0.0324 1 2 2 2 1.3493 1 2 3 2 0.2067 1 2 4 2 0.2104 1 2 5 2 0.1474 1 2 6 2 0.0191 1 2 7 2 0.1203 1 3 1 2 O.1707 1 3 2 2 0.4098 1 3 3 2 0.0 1 3 4 2 0.0 1 3 5 2 0.0345 1 3 6 2 0.0396 1 3 7 2 0.0851 2 1 1 2 1.0632 2 1 2 2 2.0062 2 1 3 2 1.6172 2 1 4 2 0.6663 2 1 5 2 0.3699 2 1 6 2 0.3136 2 1 7 2 0.0 2 2 1 2 0.4334 2 2 2 2 1.8282 2 2 3 2 2.0819 2 2 4 2 1.1697 2 2 5 2 0.7986 2 2 6 2 0.1532 2 2 7 2 0.3173 2 3 1 2 1.6970 2 3 2 2 1.2308 2 3 3 2 0.6588 2 3 4 2 0.0153 2 3 5 2 0.3772 2 3 6 2 0.0870 2 3 7 2 0.9349 3 1 1 2 0.1463 3 1 2 2 2.0193 3 1 3 2 1.0219 3 1 4 2 0.2290 3 1 5 2 0.0 3 1 6 2 0.0278 3 1 7 2 0.3370 3 2 1 2 1.8326 3 2 2 2 1.4858 3 2 3 2 1.0158 3 2 4 2 0.4643 3 2 5 2 0.2383 3 2 6 2 0.0513 3 2 7 2 0.0 3 3 1 2 0.5478 3 3 2 2 1.8112 SURFACE LENGTH DIAMETEF AREA 2 3 3 CM /DM CM/DM MM 32 .0 35 . 2 2.9 19.8 22 .8 2.8 0.0 0.0 0.0 21.7 20.7 3.3 7.6 6.9 3.5 28 . 2 25.6 3 . 5 7.5 6.6 3.6 10.6 8.8 3.8 9.0 9.4 3 . 1 1 .7 2.7 2.0 38.5 38.9 3. 1 5.9 5.8 3.3 5.2 4. 1 4.0 6 . 2 8.3 2.4 1 .0 1 .4 2.3 4.3 5.3 2.6 6.6 9.2 2 . 3 12.6 15.4 '2.6 0.0 0.0 0.0 0.0 0.0 0.0 1 .2 1 . 1 3 . 3 1 . 2 1 .8 2. 1 3.8 5.8 2 . 1 28.2 29 . 3 3. 1 56 . 1 60.4 3.0 43.4 43.4 3.2 17.9 18.0 3.2 12.1 15.3 2.5 9.2 1 1 .0 2.6 0.0 0.0 0.0 10.7 11.2 3.0 52.7 56 .9 2.9 54.9 56.6 3 . 1 32.5 34 . 1 3.0 22.9 26.5 2.8 4.7 5.7 2.6 8. 1 9.9 2.6 43.0 46.5 2.9 30.9 28.9 3.4 18.0 18.0 3.2 0.8 1 .2 2.0 10.0 10.3' 3. 1 2.4 2.6 2.9 24.9 26.9 2.9 4.7 5.6 2.7 39.9 38.6 3.3 20.9 19.6 3.4 5.8 6 .8 2.7 0.0 0.0 0.0 0.9 1 .4 2.0 9.3 8.8 3.4 44. 1 48.0 •2.9 38.2 42.5 2.9 27.7 32.6 2.7 11.1 10.8 3.3 5.8 5.7 3.3 1 .3 1 .2 3.4 0.0 0.0 0.0 15.5 19.0 2.6 40.8 41.5 3. 1 Appendix 5 con't OVERSTORY 2-5 MM ROOTS BIOMASS SURFACE LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 SITE DEPTH G/DM CM /DM CM/DM MM 3 3 3 2 0.7886 3 3 4 2 0.6162 3 3 5 2 0.2626 3 3 6 2 0.1529 3 3 7 2 0.4810 1 1 1 3 0.1527 1 2 3 0.2789 1 1 3 3 0.5533 1 4 3 0.3562 1 5 3 0.2501 6 3 0.5183 7 3 0.3987 2 1 3 0.5578 2 2 3 0.6334 2 3 3 0.4455 . 2 4 3 0.9785 2 5 3 0.7446 2 6 3 0.4592 2 7 3 0.8394 3 1 3 0.0 3 2 3 O. 2402 1 3 3 3 0.6464 3 4 3 0.0949 3 5 3 0.0415 3 6 3 0.0492 ^ 3 7 3 0.0284 2 1 1 3 0.9879 2 1 2 3 2.4812 2 1 3 3 0.6969 2 ^ 4 3 0.2102 2 1 5 3 0.7271 2 1 6 3 0.3678 2 1 7 3 0.0 2 2 1 3 0.4684 2 2 2 3 0.4970 2 2 3 3 0.7370 2 2 4 3 1.6028 2 2 5 3 0.8124 2 2 6 3 0.3293 2 2 7 3 0.5426 2 3 1 3 0.3700 2 3 2 3 0.9290 2 3 3 3 0.7444 2 3 4 3 0.8586 2 3 5 3 0.0029 2 3 6 3 0.0385 2 3 7 3 0.0 3 1 1 3 0.1750 3 •( 2 3 2.2459 3 i 3 3 1.9741 3 1 4 3 0.2696 • 3 5 3 0.6379 3 1 6 3 0.2207 3 1 7 3 0.9773 17.7 18.7 3.0 17.4 17.9 3. 1 7.6 9 . 3 2.6 5 . 3 6 . 2 2.7 10.9 10.5 3.3 6 . 3 8.7 2.3 11.3 15.8 2 . 3 13.6 16.8 2.6 9.8 11.0 2.8 7.5 8.2 2.9 15. 1 17.2 2.8 13.0 14.7 2.8 14 . 3 19. 1 2.4 21.0 26.4 2.5 13.8 14.8 3.0 28.4 31 .5 2.9 23 . 3 27 .7 2.7 13.7 15.4 2.8 25.0 27 .4 2.9 0.0 0.0 0.0 9.4 12.9 2.3 19.0 21.0 2.9 2.3 1 .8 4.0 1 .9 2. 1 2.8 2. 1 2.9 2 . 3 1 .5 2.2 2. 1 24.8 29.4 2.7 68 . 3 71.7 3.0 22.6 27.7 2.6 fl .6 12.3 2 . 2 20. 1 21.6 3.0 9.9 11.5 2.7 0.0 0.0 0.0 14.2 20.4 2.2 16. 1 20. 2 2.5 21 .2 21.7 3. 1 40.4 43.4 3.0 24.2 29.9 2.6 10.9 14.3 2.4 15.3 18.4 2.6 11.8 15.9 2.4 33 .6 47.5 2.3 23.4 28 .7 2.6 23.4 25.5 2.9 0.2 0.4 2.0 2.4 3.5 2.2 0.0 0.0 0.0 3.6 2.6 4.4 54.3 64.9 2.7 44.0 48.0 2.9 6.8 8.3 2.6 13.9 15.4 2.9 5.4 5.9 2.9 19.8 20.4 3. 1 Appendix 5 con't ... OVERSTORY 2-5 MM ROOTS BIOMASS SURFACE LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 SITE DEPTH G/DM CM /DM CM/DM MM 3 2 1 3 1 1540 3 2 2 3 3 0100 3 2 3 3 1 4450 3 2 4 3 0 4907 3 2 5 3 0 3506 3 2 6 3 0 0755 3 2 7 3 0 0513 3 3 1 3 0 8259 3 ' 3 2 3 1 0326 3 3 3 3 0 2091 3 3 4 3 0 6106 3 3 5 3 0 1686 3 3 6 3 0 1 175 3 3 7 3 0 0102 31 7 37 1 2 7 74 5 78 6 3 0 38 4 43 7 2 8 1 1 5 12 2 3 0 7 9 9 1 2 8 1 7 2 2 2 5 2 4 3 6 2 1 29 3 36 2 2 6 26 4 27 0 3 1 6 1 7 9 2 5 18 8 22 7 2 6 5 0 6 1 2 6 3 2 3 5 2 9 0 5 0 9 2 0 UNDERSTORY < 2 MM ROOTS BIOMASS SURFACE LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 SITE DEPTH G/DM CM /DM CM/DM MM 1 1 1 0 44260 147 3 960 0 49 1 2 1 0 16177 55 2 1131 0 15 1 3 1 0 08325 30 7 393 0 25 1 4 1 0 03974 10 1 151 0 21 1 5 1 0 04030 14 8 198 0 24 1 6 1 0 00389 3 0 48 0 20 1 7 1 0 03397 5 3 37 0 45 2 1 1 0 25696 81 4 877 0 30 2 2 1 0 19752 1 15 9 1 145 0 32 2 3 1 0 05710 12 8 179 0 23 2 4 1 0 05754 12 4 126 0 31 2 5 1 0 00539 1 5 21 0 23 2 6 1 0 02246 3 8 24 0 52 2 7 1 0 00639 1 1 10 0 35 3 1 1 0 46698 132 9 1529 0 28 3 2 1 0 19363 70 2 843 0 26 3 3 1 0 26631 1 16 7 1044 0 36 3 4 1 0 30439 69 8 629 0 35 3 5 1 O 35668 78 4 555 0 45 3 6 1 0 30929 84 9 513 0 53 3 7 1 0 13143 49 1 409 0 38 2 1 1 1 0 1 1447 19 8 220 0 29 2 1 2 1 0 1 1969 29 6 320 0 30 2 1 3 1 0 05557 15 3 251 0 19 2 1 4 1 0 01485 4 4 75 0 19 2 1 5 1 0 00383 1 4 28 0 17 2 1 6 1 0 00333 1 4 26 0 17 2 1 7 1 0 00186 0 9 17 0 17 2 2 1 1 0 0 0 0 0 0 0 2 2 2 1 0 0 0 0 0 0 0 2 2 3 1 0 00031 0 1 4 0 07 2 2 4 1 0 00072 0 3 12 0 09 2 2 5 1 0 00018 0 1 8 0 04 2 2 6 1 0 00022 0 1 9 0 04 2 2 7 1 0 00017 0 1 4 0 08 Appendix 5 con't 163 UNDERSTORY < 2 MM ROOTS BIOMASS SURFACE LENGTH DIAMETER AREA SITE PLOT DEPTH TIME 3 G/DM 2 3 CM /DM 3 CM/DM MM 2 3 1 ! 0.52563 107 .6 1115 0.31 2 3 2 1 0.10068 15 .3 246 0.20 2 3 3 1 0 . 16324 18 .6 543 0.11 2 3 4 1 0 .05799 19 . 2 414 0.15 2 3 5 1 0 .05205 6 .5 149 0.14 2 3 6 1 0 .00293 1 .2 1 1 0.35 2 3 7 1 0 00099 0 .9 19 0. 14 3 1 1 1 0 31604 49 . 3 936 0. 17 3 1 2 1 0 29375 37 6 620 0. 19 3 1 3 1 0 35044 100 4 1968 0. 16 3 1 4 1 0 42636 58 .9 1057 0. 18 3 1 5 1 0 10613 27 2 491 0. 18 3 1 6 1 0 04949 9 9 85 0. 37 3 1 7 1 0 03668 4 8 42 0. 37 3 2 1 1 0 18455 48 7 990 0. 16 3 2 2 1 0 09682 28 6 442 0.21 3 2 3 1 0 14176 • 32 5 432 0.24 3 2 4 1 0 28827 31 7 267 0.38 3 2 5 1 0 13125 20 O 204 0.31 3 2 6 1 0 07934 7 3 81 0.28 3 2 7 1 0 0 0 0 0 0.0 3 3 1 1 0 20362 33 1 535 0.20 3 3 2 1 0 20426 25 8 321 0.26 3 3 3 1 0 23916 40 5 546 0.24 3 3 4 1 0 68643 94 5 1 156 0.26 3 3 5 1 0 57762 99 4 1211 0.26 3 3 6 1 0 22818 42 4 627 0.21 3 3 7 1 0 14169 24 4 399 0. 19 1 1 1 2 0 22329 101 0 1322 0. 24 1 1 2 2 0 15550 51 9 675 0.25 1 1 3 2 0 1 1595 46 7 328 0.45 1 1 4 2 0 07440 9 8 82 0.38 r 1 5 2 0 02413 5 7 72 '0.25 1 1 6 2 0 01746 6 4 75 0.27 1 1 7 2 0 01263 3 6 33 0. 34 1 2 1 2 0 44830 172 2 1334 0.41 1 2 2 2 0 18898 86 6 642 0.43 1 2 3 2 0 08720 25 8 322 0.25 1 2 4 2 0 03047 20 6 177 0.37 1 2 5 2 0 01642 6 9 90 0.25 1 2 6 2 0 01405 4 2 72 0. 18 1 2 7 2 0 00105 0 7 16 0. 14 1 3 1 2 0 47399 116 8 1206 0.31 1 3 2 2 0 38957 143 3 2108 0.22 1 3 3 2 0 15800 35 7 362 0.31 1 3 4 2 0 06001 16 4 129 0.41 1 3 5 2 0 03987 12 7 148 0.27 1 3 6 2 0 01 181 4 8 58 0.26 1 3 7 2 0 00566 1 2 24 0. 16 2 1 1 2 0 24707 20 9 251 0.26 2 1 2 2 0 04573 9 3 170 0. 17 2 1 3 2 0 02750 7 5 156 0. 15 2 1 4 2 0 02023 6 8 91 0.24 2 1 5 2 0 00103 1 1 20 0. 18 2 1 6 2 0 00330 1 4 24 0. 18 2 1 7 2 0 00113 0 7 13 0. 18 2 2 1 2 0 00149 1 0 16 0. 19 2 2 2 2 0 00080 0 6 32 0.06 2 2 3 2 0 00315 2 0 58 0. 1 1 2 2 4 2 0 00599 2 2 47 0. 15 Appendix 5 con't UNDERSTORY < 2 MM ROOTS BIOMASS SITE PLOT DEPTH TIME ' 3 G/DM 2 2 5 2 0.00274 2 2 6 2 0.00026 2 2 7 2 0.00016 2 3 1 2 0.12842 2 3 2 2 0.04279 2 3 3 2 0.01101 2 3 4 2 0.10024 2 3 5 2 0.01385 2 3 6 2 0.03149 3 5 2 3 7 2 0.05768 3 1 1 2 0.49036 3 1 2 2 1.07820 3 1 3 2 0.23366 3 1 4 2 0.15106 3 1 5 2 0.03643 3 1 6 2 0.11708 3 1 7 2 0.14003 3 2 1 2 0.64986 3 2 2 2 0.27281 3 2 3 2 0.14605 3 2 4 2 0.02207 3 2 5 2 0.07466 3 2 6 2 0.01336 3 2 7 2 0.01156 3 3 1 2 0.24619 3 3 2 2 0.77092 3 3 3 2 0.27664 3 3 4 2 0.44230 3 3 5 2 0.21238 3 3 6 2 0.34022 3 3 7 2 0.11091 1 1 1 3 1.11040 1 1 2 3 0.15951 1 1 3 3 0.13093 1 1 4 3 0.04627 1 1 5 3 0.01274 1.' 1 6 3 0.00791 1 1 7 3 0.00087 1 2 1 3 0.38106 1 2 2 3 0.01366 1 2 3 3 0.00981 1 2 4 3 0.01735 1 2 5 3 0.00578 1 2 6 3 0.00204 1 2 7 3 0.00232 1 3 1 3 0.58182 1 3 2 3 0.36904 1 3 3 3 0.15324 1 3 4 3 0.07237 1 3 5 3 0.04507 1 3 6 3 0.07003 1 3 7 3 0.04571 2 1 1 3 0.04958 2 1 2 3 0.07049 2 1 3 3 0.04176 2 1 4 3 0.00566 2 1 5 3 0.02999 2 1 6 3 0.01981 2 1 7 3 0.01490 2 2 1 3 0.00007 SURFACE LENGTH DIAMET AREA 2 3 3 CM /DM CM/DM MM 0.9 12 0. 23 0. 1 3 0.15 0. 1 3 0.13 32 .9 259 0.40 13.0 183 0.23 5.4 218 0.08 12.6 231 0.17 6. 1 122 0. 16 9.8 106 0.29 6 . 2 36 0.54 113.1 1096 0.33 221 .0 3297 0.21 90.0 1699 0. 17 36 . 7 709 0.17 4.5 71 0.20 16.7 353 0. 15 37 .0 839 0.14 63 . 5 1046 0. 19 76 . 2 1423 0.17 49.6 848 0. 19 7.9 146 0.17 43 . 2 602 0.23 8 . 1 138 0. 19 4.2 114 0. 12 153.2 3372 0. 14 175 . 1 2701 0.21 115.0 1636 0. 22 82 .6 1397 0. 19 24 .6 440 0. 18 99. 1 1316 0.24 35.3 516 0.22 406.6 2692 0.48 87.7 454 0.62 30.0 205 0.46 15.2 136 - 0. 36 4.4 48 0.29 3.2 57 0. 18 0.6 13 0. 16 81.2 428 0.60 7.9 120 0.21 5.5 122 0. 14 6 . 3 119 0.17 1.8 34 0. 17 0.8 16 0. 15 0.7 1 1 0.21 86.2 1798 0. 15 131.6 867 0.48 68.0 670 0.32 20. 1 143 0.45 11.5 121 0.30 27 . 1 280 0.31 9.6 101 Or30 16.4 277 0. 19 15.5 304 0. 16 7.3 152 0. 15 10.8 169 0.20 5.7 73 0.25 2.6 33 0.26 2.7 27 0.32 0.2 4 0. 13 . 165 Appendix 5 c o n ' t . . . UNDERSTORY < 2 MM ROOTS 5 BIOMASS SURFACE LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 SITE DEPTH G/DM CM /DM CM/DM MM . 2 2 2 3 0. 00596 2. 4 51 0. 15 2 2 3 3 0. 01445 4. 5 82 0. 18 2 2 4 3 0. 12604 15. 4 141 0. 35 2 2 5 3 0. 03186 5 . 9 71 0. 27 2 2 6 3 0. 04929 6. 2 81 0. 24 2 2 7 3 0. 01614 4 . 6 89 0. 17 2 3 1 3 0. 27451 44 . 2 326 0. 43 2 3 2 3 0. 02291 7 . 6 130 0. 19 2 3 3 3 0. 05195 15. 3 282 0. 17 2 3 4 3 0. 02325 8. 7 109 0. 25 2 3 5 3 0. 00088 O. 8 17 0. 15 2 3 6 3 0. 00256 0. 9 9 0. 31 2 3 7 3 0. 00009 0. 0 1 0. 13 3 1 1 3 0. 1 1749 66 . 2 2134 0. 10 3 1 2 3 0. 47712 72. 6 1664 0. 14 3 1 3 3 0. 23453 86. 6 1603 0. 17 3 1 4 3 0. 04310 25. 7 588 0. 14 3 1 5 3 0. 08356 35. 9 619 0. 19 3 1 6 3 0. 15826 57 . 8 1310 0. 14 3 1 7 3 0. 14388 89. 6 999 0. 29 3 2 1 3 0. 13360 39. 5 611 0. 21 3 2 2 3 0. 21620 71 . 4 1068 0. 21 3 2 3 3 0. 13800 23. 5 389 0. 19 3 2 4 3 0. 04435 1 1 . 1 139 0. 26 3 2 5 3 0 06194 8 . 8 195 0. 14 3 2 6 3 0. 00551 2 . 4 59 0. 13 3 2 7 3 0. 04578 5, 2 37 0. 44 3 3 1 3 1 08140 181 . 1 2982 0. 19 3 3 2 3 0 93973 145 . 8 2090 0. 22 3 3 3 3 0 64426 42 . 8 626 0. 22 3 3 4 3 0 30930 40. 4 560 0. 23 - 3 3 5 3 0 18137 38 . 9 710 0. 18 3 3 6 3 0 15158 20. 2 436 0. 15 3 3 7 3 0 27214 71 . 6 1290 0. 18 UNDERSTORY 2-5 MM ROOTS BIOMASS SURFACE LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 SITE DEPTH G/DM CM /DM CM/DM MM 1 1 1 1 0 .2060 19.79 21 .87 2 .9 1 1 2 1 0 .0 0.0 0.0 0 .0 1 1 3 1 0 .0026 0. 34 0.55 2 .0 1 1 4 1 0 .0 0.0 0.0 0 .0 1 1 5 1 0 .0 0.0 0.0 0 .0 1 6 1 0 .0025 0.30 0.44 2 .2 1 1 7 1 0 .0 0.0 0.0 0 .0 1 2 1 1 0 .0367 3.93 4.81 2 .6 1 2 2 1 0 .0 0.0 0.0 0 .0 1 2 3 1 0 . 1797 7.77 9.73 2 .5 1 2 4 1 0 .0 0.0 0.0 0 .0 1 2 5 1 0 .0 0.0 0.0 0 .0 1 2 6 1 0 .0 0.0 0.0 0 .0 1 2 7 1 0 .0 0.0 0.0 0 .0 Appendix 5 con't ... UNDERSTORY 2-5 MM ROOTS BIOMASS SURFACE AREA LENGTH DIAMETER PLOT TIME 3 2 3 3 SITE DEPTH G/DM CM /DM CM/DM MM 1 3 1 1 0.127G 1 3 2 1 0.5117 1 3 3 1 0.0578 1 3 4 1 0.2272 1 3 5 1 0.0182 1 3 6 1 0.1592 1 3 7 1. 0.0 2 1 1 1 0.098G 2 1 2 1 0.4138 2 1 3 1 0.1525 2 1 4 1 0.0 2 1 5 1 0.0 2 1 6 1 0.0 2 1 7 1 0.0 2 2 1 1 0.0 2 2 2 1 0.0 2 2 3 1 0.0 2 2 4 1 0.0 2 2 5 1 0.0 2 2 6 1 0.0 2 2 7 1 0.0 2 3 1 1 0.0 2 3 2 1 0.0 2 3 3 1 0.5712 2 3 4 1 0.0 2 3 5 1 0.0106 2 3 6 1 0.0 2 3 7 1 0.0 3 1 1 1 0.7490 3 1 2 1 0.7939 3 1 3 1 0.4536 3 1 4 1 0.0 3 1 5 1 0.0 3 1 6 1 0.0 3 1 7 1 0.0 3 2 1 1 0.7763 3 2 2 1 2.2931 3 2 3 1 0.0381 3 2 4 1 0.0761 3 2 5 1 0.0106 3 2 6 1 0.0151 3 2 7 1 0.0 3 3 1 1 0.3196 3 3 2 1 0.4467 3 3 3 1 0.6941 3 3 4 1 0.4197 3 3 5 1 0.0763 3 3 6 1 0.0151 3 3 7 1 0.0 1 1 2 0.2967 1 2 • 2 0. 1618 1 3 2 0.1148 1 4 2 0.0 1 5 2 0.1303 1 6 2 0.0 1 7 2 0.0 2 1 2 0.2990 2 2 2 0.0 2 3 2 0.3556 2 4 2 - 0.0 2 5 2 0.0 2 6 2 0.2684 2 7 2 0.0 5. 19 5.86 2.8 12 .67 10.41 3.9 4. 15 5.23 2.5 8.98 1 1 .76 2.4 2.13 3. 13 2.2 4.76 4.00 3.8 0.0 0.0 0.0 2.97 3 . 78 2.5 9.75 9.32 3.3 5. 18 6.11 2 . 7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0' 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 10.01 8.48 3.8 0.0 0.0 0.0 0.41 0.61 2. 1 0.0 0.0 0.0 0.0 0.0 0.0 16.73 18.26 2.9 17.51 19.37 2.9 12.11 13.93 2.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 15.11 13.43 3.6 38.98 28.82 4.3 1 .07 1 .68 2.0 2.59 2.38 3.5 0.48 0.72 2. 1 0.54 0. 79 2.2 0.0 0.0 0.0 9.20 10.42 2.8 12.07 12.99 3.0 17.86 19.62 2.9 9.54 10. 13 3.0 2.92 4.66 2.0 0.73 0.63 3.7 0.0 0.0 0.0 9. 12 7 .65 3.8 5.56 6.39 2.8 2.87 2.41 3.8 0.0 0.0 0.0 3.99 3.43 3.7 0.0 0.0 0.0 0.0 0.0 0.0 17 .07 18. 11 3.0 0.0 0.0 0.0 5.85 4.70 4.0 0.0 0.0 0.0 0.0 0.0 0.0 5.70 5.80 3. 1 0.0 0.0 0.0 Appendix 5 con't ... UNDERSTORY 2-5 MM ROOTS 167 BIOMASS SURFACE AREA LENGTH DIAMETER PLOT TIME 3 2 3 3 SITE DEPTH G/DM CM /DM CM/DM MM 1 3 1 2 0.5397 1 3 2 2 0.3634 1 3 3 2 0.0441 1 3 4 2 0.2147 1 3 5 2 0.0 1 3 6 2 0.041 1 1 3 7 2 0.0 2 1 1 2 0.0 2 1 2 2 0.0 2 1 3 2 0.0 2 1 4 2 0.0 2 1 5 2 0.0 2 1 6 2 0.0 2 1 7 2 0.0 2 2 1 2 0.0 2 2 2 2 0.0 2 2 3 2 0.0 2 2 4 2 0.0 2 2 5 2 0.0 2 2 6 2 0.0 2 2 7 2 0.0 2 3 1 2 0.0446 2 3 2 2 0.0 2 3 3 2 0.0 2 3 4 2 0.2020 2 3 5 2 0.0 2 3 6 2 0.0 2 3 7 2 0.0 3 1 1 2 1.1109 3 1 2 2 0.1682 3 1 3 2 1.0306 3 1 4 2 0.1849 3 1 5 2 0.0240 3 1 6 2 0.0096 3 1 7 2 0.0 3 2 1 2 0.1268 3 2 2 2 0.5174 3 2 3 2 0.3433 3 2 4 2 0.0 3 2 5 2 0.0 3 2 6 2 0.0 3 2 7 2 0.0 3 3 1 2 0.1429 3 3 2 2 0.1289 3 3 3 2 0.2471 3 3 4 2 0.0429 3 3 5 2 0.0 3 3 6 2 0.0917 3 3 7 2 0.0073 1 1 1 3 0.5651 1 1 2 3 0.0 1 1 3 3 0.0 1 1 4 3 0.0 1 1 5 3 0.0 1 1 6 3 0.0 1 1 7 3 0.0 1 2 1 3 0.5744 1 2 2 3 0.0 1 2 3 3 0.0 1 2 4 3 0.0 1 2 5 3 0. 1918 1 2 e 3 0.0 1 2 7 3 0.0 17 51 20 86 2 7 16 59 17 19 3 1 2 30 2 89 2 5 6 86 6 73 3 2 0 0 0 0 0 0 2 20 3 06 2 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 54 2 19 2 2 0 0 0 0 0 0 0 0 0 0 0 0 3 72 2 69 4 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 24 60 24 41 3 2 3 75 2 84 4 2 22 19 17 54 4 0 5 06 4 35 3 7 0 72 0 83 2 8 0 37 0 42 2 8 0 0 0 0 0 0 3 04 2 84 3 4 11 20 9 58 3 7 8 55 10 13 2 7 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5 22 6 65 2 5 3 78 5 60 2 1 5 77 5 34 3 4 1 56 2 41 2 1 0 0 0 0 0 0 3 92 4 46 2 8 0 41 0 63 2 0 29 82 38 49 2 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 9 69 7 83 3 9 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 17 3 76 3 5 0 0 0 0 0 0 0 0 0 0 0 0 168 Appendix 5 c o n ' t . . . UNDERSTORY 2-5 MM ROOTS BIOMASS SURFACE LENGTH DIAMETER AREA PLOT TIME 3 2 3 3 SITE DEPTH G/DM CM /DM CM/DM MM 1 3 1 3 0 1274 8 35 10 02 2.7 1 3 2 3 0 3442 10 70 1 1 90 2.9 1 3 3 3 0 01 12 2 06 1 64 4.0 1 3 4 3 0 3398 11 14 9 01 3.9 1 3 5 3 0 0075 0 45 0 57 2.5 1 3 6 3 0 0269 1 66 2 34 2.3 1 3 7 3 0 0076 0 38 0 33 3.7 2 1 1 3 0 1720 4 37 5 34 2.6 2 1 2 3 0 6036 9 36 6 34 4 . 7 2 1 3 3 0 0 0 0 0 0 0.0 2 1 4 3 O 0236 1 13 1 71 2 . 1 2 1 5 3 0 0 0 0 0 0 0.0 2 1 6 3 0 0 0 0 0 0 O.O 2 1 7 3 0 0 0 0 0 0 0.0 2 2 1 3 0 0 0 0 0 0 O.O 2 2 2 3 0 0 0 0 0 0 0.0 2 2 3 3 0 0 0 0 0 0 0.0 2 2 4 3 0 1081 2 98 3 45 2.7 2 2 5 3 0 0 0 0 0 0 0.0 2 2 6 3 0 0 0 O 0 0 0.0 2 2 7 3 0 0 0 0 0 0 0.0 2 3 1 3 0 0858 1 48 1. 57 3.0 2 3 2 3 0 0 0 0 0 0 0.0 2 3 3 3 0 0 0 0 0 0 0.0 2 3 4 3 0 0 •o 0 0 0 0.0 2 3 5 3 0 0 0 0 0 0 0.0 2 3 6 3 0 0 0 0 0 0 O.O 2 3 7 3 0 0 0 0 0 0 0.0 3 1 1 3 0 7538 13 30 14 43 2.9 3 1 2 3 0 6458 12 47 13 30 3.0 3 1 3 3 0 3196 6 47 6 85 3.0 3 1 4 3 0 0 0 0 0 0 0.0 3 1 5 3 0 0652 1 49 1 90 2.5 3 1 6 3 0 0 0 0 0 0 0.0 3 1 7 3 0 0 0 0 0 0 0.0 3 2 1 3 0 0 0 0 0 0 0.0 3 2 2 3 0 7843 18 23 18 81 3. 1 3 2 3 3 0 0258 0 91 1 07 2.7 3 2 4 3 0 0 0 0 0 0 0.0 3 2 5 3 0 0 0 0 0 0 0.0 3 2 6 3 0 0 0 0 0 0 0.0 3 2 7 3 0 0 0 0 0 0 0.0 3 3 1 3 0 5373 7 52 10 85 2.2 3 3 2 3 0 9937 17 85 15 35 3.7 3 3 3 3 0 2146 4 40 4 33 3.2 3 3 4 3 0 2543 6 55 7 1 1 2.9 3 3 5 3 0 18'49 4 41 5 10 2.8 3 3 6 3 0 0 0 0 0 0 0.0 3 3 7 3 0 0 0 0 0 0 0.0 Appendix 6 . Ash content o f roo ts (data cour tesy o f P. Cour t in who c a r r i e d out the chemical a n a l y s i s o f these roots) summarized by p lan t a s s o c i a t i o n , s i z e ca tego ry , and dep th , and of roo ts not growing in minera l s o i l . Mean i s of 9 samples un less otherwise noted a f t e r the mean i n parentheses . The standard e r r o r i s i n parentheses below the mean. Depth From Sur face of Oplopanaco ( h o r r i d i ) -Thujetum p l i c a t a e Ab ie to (amab i l i s ) -Tsugetum mertensianae Vaccin ium (membranacei) -Tsugetum mertensianae Fores t F l o o r Suber i zed 0 c m m _ 0 c-o mm < 2 mm Suber ized 0 c , c-o mm < c mm Suber ized n c < 2 mm 2 " 5 m cm — — — — — - — - — — — percent 0-5 2.11 (0.488) 1.89 (4) (0.456) 1.99 (0.239) 1.70 (0.300) 1.80 (0.288) 1.60 (7) (0.314) 5-10 2.22 (6) (0.671 ) 1.94 (6) (0.533) 2.88 (0.625) 2.07 (0.495) 2.27 (1.198) 1.58 (0.350) 10-20 2.49 (7) (1.050) 2.08 (8) (0.614) 4.69 (2.119) 2.52 (0.709) 3.31 (1.568) 2.13 (0.787) 20-30 3.29 (6) (0.738) 2.06 (6) (0.473) 5.71 (1.852) 2.87 (8) (0.649) 3.84 (7) (1.621) 2.28 (8) (0.939) 30-40 3.64 (8) (1.662) 2.58 (7) (1.150) 6.93 (8) (2.455) 3.28 (7) (1.125) 5.01 (2.712) 2.47 (7) (0.890) 40-50 4.34 (6) (2.701 ) 2.64 (5) (0.888) 6.51 (7) (2.292) 2.52 (4) (0.598) 3.69 (6) (2.257) 2.65 (5) (1.358) 50-60 4.06 (6) (1.087) 2.78 (3) (0.331 ) 6.35 (6) (1.460) 3.44 (4) (0.379) 3.98 (5) (3.027) 2.91 (5) (1.220) Roots not growing in minera l s o i l 1.96 (10) (0.220) 1.94 (6) (0.528) 1.95 (3) (0.074) 1.43 (3) (0.087) 1.72 (10) (0.231) 1.42 (8) (0.277) 170 Appendix 7. Diagrams of s o i l cores sampled f o r root v e r t i c a l d i s t r i b u t i o n , b iomass, l e n g t h , and su r face area de te rm ina t i on . OPHO THPL TIME CORE* 2-13 JUNE 0 10 3 2 0 z ~ 30 I t-Si 4 0 D 50 60 CORE* o — 10 — g 20 30 — 40 — 50 — 60 CORE* 0 10 5 2 0 z ~ 30 I l-°- 40 UJ Q 50 60 1 2 3 4 F H Ae H H H Bf Bg A he Ae 5 6 7 e F i i • i H H H H Ae Ae Ae eg G Bf Bg 10 11 12 F DW H DW Bf H H DW Bf Ahe B1 Ahf Bt Ahe Bf 18-21JULY 1 2 3 F 14-1 7 SEPT 1 2 3 4 H Ahe H Ae DW H Ah? Bf eg 5 6 7 F e 1 i 1 H AA H G M DW G . Ahe c H Ae T Bf e Ae Bg Bf 9 10 11 12 F H H DW Ae Bf Ahe DW A^ e Bf G Bf A flf? sg Bf A e Bf G Bg G DW Ahe Bf _A£_ Bf eg Cg 10 11 12 F DW H H Ahe H IJUV H Ae Bhf Bf G G Bg G G Bhf Bhf BC G c Bhf b H DW A e — Ahe H DW — Bf Bhf Bf eg — BC Ahe-* Co res : 1-4 = r e p l i c a t e 1, 5-8 = r e p l i c a t e 2 , 9-12 = r e p l i c a t e 3. Hor izon no ta t i ons f o l l o w CSSC (1978) , excep t : G = Rock, Dw = decaying wood. Appendix 7 c o n ' t ABAM TSME TIME CORE * 0 — 10 — 5 30 — I -Q — 50 — 601— CORE * ° E 1 o\-5"20r-a. Q . I -2-13 JUNE 4 0 — 5 0 — 6 0 — CORE * ° E I 0 r -8 ' t 2 3 0 t 4 0 | -UJ o 5 0 I -60 13 14 15 F . 16 i i i H DW H H AC Ae R f Bf s Bf Bf R R 17 18 F 19 20 I i i Ae H DW H Ae Ae H Bf G Bf 6 Bf Bf C Bf 21 22 23 24 F i i " ' I H Bf Ae H Ah H Ahe Ae Ae Ae Bf Ae Bf Bf G Bf G R Bf 18-21 JULY 13 14 15 16 14-17 S E P T =BZ Bf Ae Bf DW Ae DW Arte Bf Bg eg 17 18 19 20 F H H Ae DW H T Ae Bf Ae Bf Bf G Bf Bf T Bf R 21 22 23 F 24 i i i i H H H H DW Ae Ahe Bf Bf DW Bf Bf Bg Bf A? G Bf 13 14 15 F 16 r I i Ac H H H DW DW DW A h r Bhf Ae Bf BC Ae R Bf G G Bf 17 ie 19 F 20 I 1 I H Bf H H Ae H C Bf Ahe Bf BC H Bf G Bf Bf 21 22 23 F 24 r1" 1 I I H H H H Ae Ae Ahe Ahe Bf G Bf Bhf Ahe Bf eg G BC BC * Cores : 13-16 = r e p l i c a t e 1, 17-20 = r e p l i c a t e 2 , 21-24 = r e p l i c a t e 3. Appendix 7 c o n ' t 172 VAME TSME TIME C O R E * °E ioh-2 20 u z - 30t— X E 4 0 Q 5 0 l -60 CORE 0 10 5 20 z - 30 I i-°- 4 0 ui o . SOf-60 1 — CORE O p 10 — 2 20 O 5 30 I c~ 4 0 u o 50 601— 2-13 JUNE 25 26 27 26 F 16-21 JULY 14-17 S E P T H H DW H DW Bf Bf H R Ae Ae G G Bf Bf G R 29 30 31 32 F H Ae H -A£L Bf Bf Afi. Bf eg 33 34 35 36 F H H DW H H Ae Bf Ae Bhf Bf Bf Bf Ae Bf eg C C5 25 26 27 F 26 i 1 H DW H H G Ae Bf At Bf H G Bf R R G 29 30 31 32 F H F H H Ae Ae Bf Ahe Bf A P Bf Bf G G G G eg eg 33 34 35 F 36 1 1 1 H Ap H H Ae Ae Ahe Bf DW DW Bg Bf eg Ae Bf G G 25 26 27 28 F i i 1 H H F H A P A<-Bhf H R Bf R Ae Bf G 29 30 31 32 F H F H H H Bf Bf Bf Bf G BC Bf Bhf R R R Bf R 33 34 35 36 F H H H Ae Bf DW DW H BC H eg Ahe * Co res : 25-28 = r e p l i c a t e 1, 29-32 = r e p l i c a t e 2 , 33-36 = r e p l i c a t e 3. 

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