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Ecology of the alpine and timberline vegetation of Big White Mountain, British Columbia Eady, Karen 1971

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ECOLOGY OP THE ALPINE AND TIMBERLINE VEGETATION OP BIG WHITE MOUNTAIN, BRITISH COLUMBIA by KAREN EADY B.Sc., McGill University, 1965 M.Sc, University of Calgary, 1967 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OP DOCTOR OF PHILOSOPHY in the Department of Botany We accept this thesis as conforming to the required standard THE UNIVERSITY OP BRITISH COLUMBIA January, 1971 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of Botany  The University of British Columbia Vancouver 8, Canada Date Feb. 1. 1971 ii Abstract During the summers of 1968 and 1969, a plant ecological i study was carried out on Big White Mountain, in the Okanagan Highland of southern British Columbia. The main objectives of the research were to produce an ecosystematic classification of the vegetation, and to determine the environmental factors important in differentiating the plant communities. The vegetation was studied by the phytosociological methods of Braun-Blanquet, as modified by Krajina. A number of environ mental features were noted for each plot, and soil samples were collected by horizon. Physical and chemical analyses of the soils were done in the laboratory. Fourteen plant associations, with nine variations, were distinguished in the study area. These communities were compared with one another, using an index of floristic similarity. In general, there is a very low degree of similarity among the communities, thus supporting the initial classification system. The communities were compared with those described in other alpine and subalpine areas. Trees occurring in the subalpine parkland were found to be much older than the krummholz forms found in the alpine area. It was suggested that there has been a recent migration of tree species into the alpine area. The occurrence^ of conifer seed lings in alpine and timberline communities was presented. No conifer seedlings were found in the tree island communities. The soils were classified according to the Canadian system of soil classification. Four orders are represented in the re search area: Brunisolic, Regosolic, Podzolic and Gleysolic. • iii The soils are generally shallow, with weak horizon development. Important chemical properties are the acidic pH, narrow carbon: nitrogen ratios, low cation exchange capacities, and very low amounts of exchangeable cations. In an analysis of environmental variables, the communities were grouped according to hygrotope. The environmental data were summarized for each group. Prom an analysis of variance, all the factors were significant either at the 1% or 5% level, except relief. Based on Duncan's New Multiple Range Test, each community was discussed, mentioning the environmental factors which were found to be significant in differentiating it. It was concluded that general environmental factors (with hygrotope the most important) are more significant in distinguishing the communities than the physical and chemical soil properties. Detailed soil moisture data were presented for a number of alpine and subalpine communities. Several of the communities were found to undergo soil moisture stress. In the zonation of the research area, the subalpine park land area was placed in the Engelmann Spruce - Subalpine Fir Zone. The alpine and low alpine areas constitute the Alpine Zone. The timberline vegetation is composed of the' subalpine parkland and parts of the low alpine area. The alpine zone of Big White Mountain is not as well developed as it is in the coastal area or the Rocky Mountains. It was concluded that much further work needs to be done in order to properly characterize the alpine zone in British Columbia. iv TABLE OF CONTENTS Page 1. Introduction 1 2. Description of Study Area 5 A. Geographical Location and Physiography 5 B. Climate 5 C. Vegetation 10 D. Soils 1 3. Methods 4 A. Vegetation Analysis 1B. Soil Analysis 6 C. Vegetation Synthesis 19 D. Computational Methods 20 i. Floristic Similarity Index 2ii. Environment Analysis 21 4. Alpine and Timberline Communities 23 Juniperus communis Association 4 Antennaria lanata - Sibbaldia procumbens Association... 31 Juncus parryi Association 43 Antennaria lanata Association 51 Phyllodoce empetriformis - Antennaria lanata Association 58 Picea engelmannii Association 69 Abies lasiocarpa Association 73 Abies lasiocarpa - Picea engelmannii - Vacclnium scoparium Association 78 Abies lasiocarpa - Valeriana sitchensis Association.... 86 Page Carex spectabilis Association 92 Valeriana sltchensis - Castilleja elmeri Association... 98 Carex nigricans Association 109 Polytrichum norveglcum Association 121 Drepanocladus exannulatus Association 127 5. Distribution of Tree Species 133 6. Vegetation Relationships 138 7. Vegetation-Environment Relationships 151 A. Analysis of Environmental Variables 152 B. Soil Moisture l6i C. Topographic-Altitudinal Relationships 166 D. Soil Types and Plant Communities 169 8. Vegetation Zonation 173 9. Summary and Conclusions 6 10. Literature Cited 182 Appendix 1. Checklist of Vascular Plants, Bryophytes and Lichens 194 Appendix 2. Soil Types of Big White Mountain classif ied according to the American, German and World FA0/UNESC0 Classifications 202 Appendix 3. Iron and Aluminum Determinations 205 Appendix 4. Class Limits for Environmental Variables. 211 Appendix 5. Statistical Analysis 214 vi LIST OF TABLES Table Page 1 Some Climatic Data for the Big White Area 9 2 Mean Monthly Temperature (°F) for Big White Mountain (elev. 6050 ft.) 9 3 Precipitation (in.) for Big White Mountain (elev. 6050 ft.) 9 4 Species Significance Scale 15 5 Sociability Scale 16 General Environment, Juniperus communis Association. 25 7 Juniperus communis Association 26 8 Floristic Similarity Indices for the Juniperus  communis Association 27 9 Soil Texture, Juniperus communis Association 28 10 Soil Chemical Analysis, Juniperus communis Association 29 11 General Environment, Antennaria lanata - Sibbaldia  procumbens Association 32 12 Antennaria lanata - Sibbaldia procumbens Association 33 13 Floristic Similarity Indices for the Antennaria lanata - Sibbaldia procumbens Association 35 14 Soil Texture, Antennaria lanata - Sibbaldia  procumbens Association 37 15 Soil Chemical Analysis, Antennaria lanata -Sibbaldia procumbens Association 38 16 General Environment, Juneus parryi Association 45 17 Juncus parryi Association 46 vii Table Page 18 Floristic Similarity Indices for the Juncus parryi Association 47 19 Soil Texture, Juncus parryi Association 48 20 Soil Chemical Analysis, Juncus parryi Association... 49 21 General Environment, Antennaria lanata Association.. 52 22 Antennaria lanata Association 53 23 Floristic Similarity Indices for the Antennaria  lanata Association 54 24 Soil Texture, Antennaria lanata Association 55 25 Soil Chemical Analysis, Antennaria lanata Association 56 26 General Environment, Phyllodoce empetriformis -Antennaria lanata Association 59 27 Phyllodoce empetriformis - Antennaria lanata Association 60 28 Floristic Similarity Indices for the Phyllodoce  empetriformis - Antennaria lanata Association 62 29 Soil Texture, Phyllodoce empetriformis - Antennaria  lanata Association 64 30 Soil Chemical Analysis, Phyllodoce empetriformis -Antennaria lanata Association ; 65 31 General Environment, Picea engelmannii Association.. 70 32 Picea engelmannii Association 71 33 Soil Texture, Picea engelmannii Association 72 34 Soil Chemical Analysis, Picea engelmannii Association 72 35 General Environment, Abies laslocarpa Association... 74 36 Abies laslocarpa Association 75 viii Page Soil Texture, Abies lasiocarpa Association 76 Soil Chemical Analysis, Abies lasiocarpa Association 77 General Environment, Abies lasiocarpa - Picea engelmannii - Vaccinium scoparium Association 79 Abies lasiocarpa - Picea engelmannii - Vaccinium scoparium Association 80 Floristic Similarity Indices for the Abies lasiocarpa - Picea engelmannii - Vaccinium scoparium Association 82 Soil Texture, Abies lasiocarpa - Picea engelmannii -Vacci'nium scoparium Association 83 Soil Chemical Analysis, Abies lasiocarpa - Picea engelmannii - Vaccinium scoparium Association 84 General Environment, Abies lasiocarpa - Valeriana sitchensis Association 87 Abies lasiocarpa - Valeriana sitchensis Association. 88 Floristic Similarity Indices for the Abies lasiocarpa - Valeriana sitchensis Association 89 Soil Texture, Abies lasiocarpa - Valeriana sitchensis Association 90 Soil Chemical Analysis, Abies lasiocarpa - Valeriana sitchensis Association 91 General Environment, Carex spectabilis Association.. 93 Carex spectabilis Association 94 Floristic Similarity Indices for the Carex spectabilis Association 95 Soil Texture, Carex spectabilis Association 96 ix Table Page 53 Soil Chemical Analysis, Carex spectabilis Association 97 54 General Environment, Valeriana sltchensis -Castilleja elmerl Association 100 55 Valeriana sltchensis - Castilleja elmeri Association 101 56 Floristic Similarity Indices for the Valeriana  sltchensis - Castille ja elmeri Association 102 57 Soil Texture, Valeriana sltchensis - Castilleja  elmeri Association . 105 58 Soil Chemical Analysis, Valeriana sitchensis -Castille ja elmeri Association 106 59 General Environment, Carex nigricans Association.... 110 60 Carex nigricans Association Ill 61 Floristic Similarity Indices for the Carex nigricans Association 113 62 Soil Texture, Carex nigricans Association 115 63 Soil Chemical Analysis, Carex nigricans Association.116 64 General Environment, Polytrichum norvegicum Associat- : ion 122 65 . Polytrichum norvegicum Association 123 66 Soil Texture, Polytrichum norvegicum Association....124 67 Soil Chemical Analysis, Polytrichum norvegicum Association 125 68 General Environment, Drepanocladus exannulatus Association 128 69 Drepanocladus exannulatus Association 129 X Table Page 70 Soil Texture, Drepanocladus exannulatus Association 131 71 Soil Chemical Analysis, Drepanocladus exannulatus Association 1372 Diameter, Height and Age Measurements of Abies  lasiocarpa and Picea enge lmannii 135 73 Age of Tree Species in Alpine and Subalpine Parkland Areas 136 74 Occurrence of Conifer Seedlings and Shrubs in Alpine and Timberline Communities 13? 75 Synthesis Table for all Associations 139 76 Floristic Similarity Indices for all Communities.... 141 77 Summary of General Environmental Variables for all Communities 153 78 Summary of Physical and Chemical Soil Data for all Communities 15^ 79 Soil Moisture Percentages for Selected Alpine Communities 163 80 Soil Moisture Percentages for Selected Subalpine Communities 164 81 F Values for Environmental Variables 215 82 Significant Environmental Variables between Communities 216 83 Key to Environmental Variables 217 xi LIST OF FIGURES Figure Page 1 Location of Big White Mountain 6 2 Topographic Map of Big White Mountain and surround ing Area 7 3 Juniperus communis Association 30 4 Antennaria - Sibbaldla Association, Antennaria -Sibbaldia - Salix Variation 39 5 Soil Profile of Antennaria - Sibbaldia Association, Antennaria - Sibbaldia - Salix Variation 39 6 Antennaria - Sibbaldia Association, Carex  phaeocephala Variation 42 7 Antennaria - Sibbaldia Association, Carex brewer! Variation 48 Juncus parryi Association 50 9 Antennaria lanata Association 7 10 Soil profile of Antennaria lanata Association 57 11 Phyllodoce - Antennaria Association, Phyllodoce -Antennaria Variation 67 12 Soil profile of Phyllodoce - Antennaria Association, Phyllodoce - Antennaria Variation 67 13 Phyllodoce - Antennaria Association, Antennaria -Vaccinium Variation r 68 14 Soil profile of Phyllodoce - Antennaria Association, Antennaria - Vaccinium Variation 68 15 Abies - Picea - Vaccinium Association 85 xii Figure Page 16 Soil profile of Abies - Picea - Vaccinium Association 85 17 Valeriana - Castilleja Association, Valeriana -Castille.1a Variation 104 18 Soil profile of Valeriana - Castilleja Association, Valeriana - Castille ,1a Variation 104 19 Valeriana - Castille.ja Association, Trollius laxus Variation 108 20 Carex nigricans Association, Carex - Polytrichadel-phus Variation 117 21 Soil profile of Carex nigricans Association, Carex-Polytrichadelphus Variation, Plot 71 118 22 Soil profile of Carex nigricans Association, Carex -Polytrichadelphus Variation, Plot 77 118 23 Carex nigricans Association, Juncus - Carex -Drepanocladus Variation 120 24 Soil profile of Carex nigricans Association, J uncus - Carex - Drepanocladus Variation 120 25 Polytrichum norvegicum Association 126 26 Drepanocladus exannulatus Association 132 27 Soil profile of Pre pan o cladus exannulatus Association 13xiii Acknowledgements I would like to express my thanks to my supervisor, Dr. V.J. Krajina, for his advice and Criticism, for the verification of the vascular plants, and identification of certain bryophytes. I would also like to thank Dr. W.B. Schofield for the verific ation and identification of the bryophytes, Mr. George Otto for the verification and identification of the lichens, and Dr. C.D. Bird of the University of Calgary for the identification of certain lichens. My thanks are also extended to Dr. L.M. Lavkulich of the Department of Soil Science for his^advice on soils and environ mental analyses; to Mr. B. von Spindler of the Department of Soil Science for performing many of the soil chemical analyses; to Mr. Stephen Borden of the Biology Data Centre for his help with computational and statistical analyses; and to Dr. Jack Maze for his helpful discussions on several aspects of the research. I would like to acknowledge the cooperation of the Kelowna Ski Club and the Big White Ski Development Limited in providing accommodation and access to the research area; and to my field assistants Miss Terry Odium and Miss Linda Majeski. This research was supported through a National Research Council Scholarship and Bursary, a National Research Council grant to Dr. V.J. Krajina, and a grant from the Arctic and Alpine Committee of the University of British Columbia. Finally, I would like to thank my husband, Frank, without whose encouragement and support this research would not have been completed. I 1 1. Introduction y The ecology of the alpine region has been little studied in Canada, essentially due to the inaccessibility and the harsh environmental conditions of such areas. In British Columbia, Archer (1963) contributed a synecological study in Garibaldi Park. Krajina (1959* 1965* 1969) has described some general characteristics for the alpine zone. Praser (1970) studied successional trends on recently deglaciated terrain in Garibaldi Park. The timberline area has been more fully documented in the work of Peterson (1964) and Brooke (1966), who described the vegetation and environment in the parkland subzone of the coast al subalpine zone. Brink (1959) discussed the subalpine forest-heath ecotone in Garibaldi Park. A later work by Brink (1964) dealt with plant establishment in alpine and subalpine regions. Brief mention of the alpine zone in the coastal area has been made by Calder and Taylor (1968) in the Queen Charlotte Islands, and Carl (19^4) and Hardy (1955) in the Forbidden Plateau area of Vancouver Island. No detailed ecological work has previously been done in the interior of British Columbia. Cooper (1916) studied successional trends in the subalpine zone of the Mount Robson area, and mentioned the occurrence of an alpine zone. Munro and Cowan (1944) in Kootenay National Park, and Carl and Hardy (1945) in the Columbia Valley briefly discussed alpine vegetation. Raup (193^* 19^5) made notes of alpine and timber-line vegetation in northern British Columbia. In Alberta, a general description of alpine vegetation was given by Moss (1955). A detailed study was carried out by the author (Beder, 1967). Recently, Bryant and Scheinberg (1970) studied the Interaction of vegetation and frost activity in an alpine fellfield. A great amount of work has been done in the United States, much of the emphasis being on autecological studies. Important works in this field are Billings and Bliss (1959), Billings and Mooney (.1968), Bliss (1956, 1962), Mooney and Billings (1961), Mooney (1963), Spomer (1964), and Spomer and Salisbury (1968). Many valuable synecological studies have also been done. Bliss (1963) worked on alpine communities in the Presidential Range of New Hampshire. Marr (1961) has described the various eco systems in the Front Range in Colorado. Bamberg and Major (1968) worked in several alpine regions in Montana. Daubenmire has dis cussed alpine timberlines (1955) and vegetational zonation in the Rocky Mountains (19^3). Other discussions of timberline are given in Griggs (1938, 1946). Wardle (1965) compared timberlines in North America with those in New Zealand. A number of recent studies have been done in Washington and Oregon. Franklin and Trappe (1963) and Franklin and Dyrness (1969) described alpine and subalpine meadow communities. Franklin et al. (1966) dis cussed invasion of subalpine meadows by trees in Mount Rainier National Park. Douglas (1969) worked on subalpine tree groups in the North Cascade Mountains. The upper subalpine zone in the Olympic Mountains was studied by Kuramoto (1968) and by Fonda and Bliss (1969). The mountain communities of Scotland have been studied by Poore and McVean (1957), and McVean and Ratcliffe (1962). Other British ecologists who have dealt with alpine vegetation are Tansley (1949), Pearsall (1950), and Watt and Jones (1948). In 3 Australia, Costin (1957) and McVean (1969) have described the alpine vegetation. Billings and Mark (1961), and Mark and Burrell (1966) worked in alpine areas of New Zealand. In central 11 Europe, the Zurich-Montpelller school has studied the classific ation and ecological relations of communities in the alpine reg ion (Braun-Blanquet and Jenny, 1926; Braun-Blanquet, 1948). Physiological ecology studies include those of Tranquillini (1963, 1964). Krajina (1933) and Hadac (1969), working in Czechoslovakia, and Szafer, Pawlowski and Kulczynski (1923) and Pawlowski (1935)* working in Poland, have studied the high moun tain vegetation of the Tatra Mountains. In Scandinavia, Nord-hagen (1936) studied the subalpine-alpine vegetation of Norway. Dahl (1956) studied the vegetation of Rondane, in southern Norway. Gjaerevoll (1956) has worked on the Scandinavian alpine snowbeds. Detailed alpine ecological work has been done in the U.S.S.R. by many botanists (Sukachev, 1965). Many of the ecological studies mentioned above also deal with alpine and subalpine soils. For British Columbia, Farstad and Rowles (i960) briefly mentioned several alpine soils. Recently, detailed work has been done by Sneddon (1969) and van Ryswyk (1969). Baptie (1968) studied the soils of an alpine valley in Alberta. The Canadian system of soil classification (Canada Soil Survey Committee, 1970) discussed the distribution of alpine soils. The major segment of information for North America is derived from the work of Retzer (1956, 1962, 1965) ln the Rocky Mountains, Nimlos and McConnell (1962, 1965) in Montana, and Johnson and Cline (1965) in Colorado. Kubiena (1953) is the basic reference work for European soils. More recently, Romans et al. (1966) worked on alpine soils in Scot land. In Australia, alpine soils have been described by Costin (1955). As can be seen from this brief literature review, much work is being done throughout the world, both from synecological and autecological approaches. The original aim of the present study was to provide detailed synecological information on alpine ecosystems in the interior of British Columbia. Although Big White Mountain has only a very limited area of alpine veget ation, it was selected because of its accessibility. The project was then expanded to include the timberline area of the mountain. This ecotone area of timberline, while interesting in itself, is important in an understanding of the alpine zone. The research was carried out during the summers of 1968 and 1969, with the following objectives: l) to provide data on vegetation and envir onment in an alpine-timberline area, 2) to produce an ecosystem-atic classification of the alpine and timberline vegetation, and 3) to elucidate the environmental factors responsible in the formation of different plant communities. 5 2. Description of Study Area A. Geographical Location and Physiography Big White Mountain, with an elevation of 7603 feet, is located approximately thirty miles southeast of Kelowna, in the Okanagan Highland, a subdivision of the Interior Plateau (Pig. 1 and 2). Access is afforded by a gravel road from Highway 33 to the local ski area at 6050 feet. The following summary is based on Holland (1964). The Okanagan Highland lies between the Monashee Mountains to the east and the Thompson Plateau on the west. It consists of rounded mountains and ridges, and gentle slopes. During the Pleistocene, ice covered the highland, but erosion was not great. There was some rounding of surfaces, but the main effect was the deposition of drift. A large part of the area is underlain by Shuswap gneisses. On Big White Mountain, the main rock types are granite and porphyritic granite, which comprise the Valhalla Intrusions, dated to the Lower Cret aceous (Little, 1957). The highland is drained and dissected by the Okanagan and Kettle Rivers and their tributaries. As seen in Pig.l, Big White Mountain is situated between the Kettle and West Kettle Rivers. The valley of the Kettle River actually forms the eastern boundary of the Okanagan Highland. B. -Climate There are no climatic stations in the alpine and timberline region of the research area. None were set up during the course of.study due to the logistic difficulties involved. Table 1 summarizes some approximate climatic data for the general area, based on a number of A.R.D.A. maps (British Columbia, Canada 1 Land Inventory). A few differences between these data and those 120° U.S.A. Scale : I inch = 30 miles Based on ac. Dept. of Mines & Petroleu Resources Map No. UPS fig. 1. Location of Big White Mountain. 7 Fig 2. Topographic map of Big White Mountain and surrounding area. Scale: 1 inch = 2 miles. Taken from B.C. Department of Lands and Forests, National Topographic Series, Sheets 82 E/NW and 82 E/NE. 8 presented by Krajina (1959, 1965, 1969) for the Alpine Tundra Zone are the higher mean July temperature (60°P compared to 44-52°F), higher absolute maximum temperature (95°F compared to 70-83°P) and longer frost-free period (60 days compared to less than 25) at Big White. Temperature and precipitation have been recorded sporadic ally at an elevation of 6050 feet on Big White. This altitude corresponds to the Engelmann Spruce - Subalpine Fir Zone of Krajina (1965). Tables 2 and 3 present the available information (British Columbia Department of Agriculture, 1965-1968). From these data, it appears that Big White has a maritime precipit ation pattern with a winter maximum and summer minimum. Snow is a very important factor in alpine and timberline areas. Snow may fall during any summer month. In 1969, there were snowfalls on June 28, July 4 and July 6. In 1968, it snowed on August 18. Impassable road conditions during the spring thaw of 1969 prevented any attempt at/obtaining snow depth measurements. When the summer field season began during the last week of June, most of the snow had disappeared, with the exception of late-lying snowbanks. The British Columbia Department of Lands, Forests and Water Resources operates a number of snow courses throughout the province. One of these is located on Big White at an elevation of 55OO feet. Measure ments have been made since 1966 (British Columbia Department of Lands, Forests and Water Resources, 1966-1969). In 1966, the maximum snow depth of 48.1 inches occurred at the end of February. By the end of May, the snow depth was 7.8 inches. The corres ponding figures for 1967 are 71.6 inches at the end of March Table 1 9 Some Climatic Data for the Big White Area Period of observations Mean January temperature • 15°F 1950-1964 Mean July temperature 60°F 1950-196Absolute minimum temperature -40°F 1930-1964 Absolute maximum temperature 95°F 1930-196Average frost-free period 60 days 1950-1964 Mean annual precipitation 30-40 in. 1930-1964 Average precipitation May through September 10 in. 1930-1963 Mean annual snowfall 150-200 in. 1963-1964 Table 2 Mean Monthly Temperature (°F) for Big White Mountain (elev. 6050 ft.) Jan. Feb. Mar. April May June July Aug. Sept. Oct. Nov. Dec. 1965 - 19 - 48 55 54 42 41 - 19 1966 17 21 22 33 44 45 - 50 35 25 22 1967 20 23 22 29 39 - 55 62 54 33 25 11 1968 16 29 28 28 41 43 -Table 3 Precipitation (in.) for Big White Mountain (elev. 6050 ft.) Jan. Feb. Mar. Apri1 May June July Aug. Sept. Oct. Nov. Dec. Annual 1965 - - 3.80 - - 0.40 1.92 5.65 2.55 0.86 2.57 5.83 -1966 4.58 5.43 2.65 3.45 2.51 3.63 2.91 - 1.58 2.90 7.38 8.26 -1967 11.03 6.39 8.07 6.54 2.41 1.65 1.25 0.53 1.40 9.91 3.39 10.16 62.73 1968 7.05 3.35 4.89 9.32 4.05 4.88 - . - - 1 -and 35.0 inches at the end of May; for 1968, 59.5 inches air~fehe end of March and 26.0 inches at the end of May; for 1969, 64.0 inches at the end of February and 9.8 inches at the end of May. It must be remembered that the snow depths in the research area, which is approximately 2000 feet higher than the snow course, would be substantially greater. Winter snowfall at the 605O ft. station on Big White has been reported as being 270.9 inches for the 1965-1966 winter and 479.9 inches during the 1967-1968 winter (British Columbia Department of Agriculture, 1966, 1968). The latter figure seems unusually high, since Hollyburn Ridge in the coastal subalpine mountain hemlock zone averages only 301.5 inches (British Columbia Department of Agriculture, 1966). C. Vegetation The portion of Big White which was studied ranged in alti tude from ca.7100 to 7600 feet. Three altitudinally-delimited areas have been distinguished: alpine (ca.7500-7600 feet), low alpine (ca.7400-7500 feet), and subalpine parkland (ca.7100-7400 feet). The subalpine parkland and parts of the low alpine areas constitute the timberline vegetation, which is an ecotone between the closed subalpine forest (occurring below 7100 feet on Big White) and the alpine region. A more detailed discussion of zonation is presented in section 8. Tree species present in the area are Abies lasiocarpa^  Picea engelmannii, Pinus contorta var. latifolia and Pinus  albicaulis. The two species of pine are rare on the mountain. 1 Authorities of botanical names are listed in Appendix 1. There seems to be no evidence of recent fire in the study area. However, examination of a core from a subalpine fir in the parkland area revealed that the tree had been burned on one side. This was possibly due to a lightning strike. No evidence of fire was noted from another fir in the same tree island. Fourteen plant associations have been distinguished in the present study. Their habitats vary from dry ridge tops and rock outcrops to snowpatches and seepage slopes. Although there are few species of bryophytes and lichens present in the area, they constitute an important part of some communities. D. Soils Four orders of the Canadian system of soil classification (Canada Soil Survey Committee, 1970) are represented in the study area - Brunisolic, Regosolic, Podzolic and Gleysolic. None of the soil great groups or subgroups are restricted to a particular community. The Brunisolic Order is represented by the Alpine Dystrie Brunisol subgroup, which occurs over a wide range of habitat conditions, from ridges and heather communities to snowpatches. and seepage slopes. Orthic Regosols are also in the above-mentioned habitats. The Podzolic Order is dominated by Sombric Humo-Ferric Podzols and Sombric Ferro-Humic Podzols. Sombric Humo-Ferric Podzols are mainly in heather communities and on rocky slopes, while Sombric Ferro-Humic Podzols predominatevin tree islands, as well as on rocky slopes. Of infrequent pccur-rence are Mini Ferro-Humic Podzols and an Orthic Humic Podzol. All the podzolic soils lack an Ae horizon. The Gleysolic Order 12 is represented mainly by Rego Humic Gleysols, and less frequently by Fera Humic Gleysols, Orthic Humic Gleysols and Rego Gleysols. These soils occur only on seepage slopes, in snowpatches and wet moss communities, and usually have an Ah-Cg horizon sequence. The soils of Big White are generally shallow (less than one foot in depth), with weak horizon development (excluding the podzols). Soil development is proceeding slowly, and has not yet reached the point at which plant communities can be diff erentiated by their soil types. The soils are all acidic, with a pH range from 4.1 to 6.3. This is to be expected, as the parent material is predominantly granite, which is an acidic rock. In some of the very shallow soils, organic matter moves downward and accumulates in the G horizon overlying a lithic contact. This occasionally creates a higher organic matter content in the C horizon than in the Ah horizon. The carbon:nitrogen ratios of the soils are generally narrow, ranging from 10 to 20; the value for cultivated soils is between 8 and 15. A narrow carbon:nitrogen ratio indicates that nitrogen should be available for higher plants. There are a number of cases in which the percentage of nitrogen is very low, thus creating a very wide carbon:nitrogen ratio. This can be due either to a loss of nitrogen during the air-drying of the soil sample, or to the presence of undecomposed organic matter in the case of an L-H or Ah horizon. Phosphorus ranges from a low of 2 ppm to a high of 28 ppm. Cation exchange capacities are generally low, ranging from 4.6 to 163.0 me/100 g. (the latter in an L-H horizon). Exchangeable cations are present in very low quantities. The ranges are as follows: calcium 0.02-7.40 me/100 g., magnesium 0.01-1.58 me/100 g., sodium 0.02-0.84 me/100 g., and potassium 0.00-3.26 me/100 g. l 14 3. Methods A. Vegetation Analysis The general approach used accepts the fact that while veg etation can be considered to be continuous, it is still pp.ssible to distinguish discrete communities (Daubenmire, 1966, 1968). The communities were selected subjectively for homogeneity in vegetation and habitat. Transitional communities were not sampled, with one exception. A single rectangular plot was used to sample each community. The plot sizes varied considerably, as a single plot size would not have adequately sampled many of the communities. The major ity of the plots ranged from 10 to 30 m2 in area. A total of 82 sample plots were analyzed. The vegetation analysis was based on the Braun-Blanquet method as modified by Krajina (1933). Subjective estimations of species significance (coverage, combined with abundance, of a species in the plot) and sociability (amount of aggregation or spacing of the individuals of a species) were made separately for each species in the different layers. The percentage cover age of each layer was also estimated. The layers distinguished we re: B layer - shrubs, 20 cm.-10 m in height C layer - herbs and dwarf shrubs, less than 20 cm. in height D layer - bryophytes and lichens E layer - epiphytes The scales used for estimating species significance and 15 sociability (after Krajina, 1933) are shown in Tables 4 and 5. Table 4 Species Significance Scale Class Description Midpoint W) +• Quite solitary, very low dominance (0-1$) 0.5 1 Seldom, very low dominance (1-2$) 1.2 Very scattered, low dominance (2-3$) 2.5 3 Scattered, low dominance (3-5$) 4.0 4 Covering 5-10$ of the plot 7.5 Covering 10-20$ of the plot 15.6 Covering 20-33$ of the plot 26.5 7 Covering 33-50$ of the plot 4l.8 Covering 50-75$ of the plot 62.9 Covering 75-less than 100$ of the plot 87.0 10 Covering 100$ of the plot 100.Table 5 Sociability Scale Class Description + Sociability 0 (individual plants) 1 Groups, up to 4x4 cm2 2 Groups, up to 25x25 cm2 3 Groups, up to 50x50 cm2 4 Groups, up to 1/3 - 3/4 m2 5 Groups, up to 1-2 m2 6 Groups, up to 5 m2 7 Groups, up to 25-50 m2 8 Groups, up to 100 m2 9 Groups, up to 200-250 m2 10 Groups, up to 500 m2 16 Collections were made of all vascular plants, bryophytes and lichens. These were later identified in the laboratory. A number of environmental features were noted for each plot. These included elevation, exposure, slope, land form, wind influence, relief, erosion, and percentage cover of humus, mineral soil and rock. Relief describes the surface shape of the plot (convex, concave, hummocky or straight). Wind Influence and erosion were assessed subjectively, on four-point scales (slight, moderate, strong, very strong for wind; none, slight, moderate, strong for erosion). B. Soil Analysis One soil pit was dug in each plot, and the horizons des cribed for depth and colour. Soil depth, rockiness, root dis tribution, and the presence of ground water were noted for each profile. A total of 230 soil samples were collected for physical and chemical analyses. The soils were then classified according to the Canadian system of soil classification (Canada Soil Survey Committee, 1970), based primarily on soil morphology observed in the field. Correlations of the Canadian system with the American, German and World classifications are pres ented in Appendix 2. The soil samples were screened through a 2 mm. screen. All analyses, both physical and chemical, were done on the less than 2 mm. size fraction. The physical properties determined were texture, moisture percentage, and water retention. Texture was done by the revised hydrometer method (Bouyoucos, 1962), using a reciprocal shaker 17 to agitate the soil suspension. The textural classification followed that of the United States Department of Agriculture (sand —2.00 to 0.05 mm., silt = 0.05 to 0.002 mm., clay = less than 0.002 mm.). The following abbreviations were used in the soil texture tables: S = sand, LS = loamy sand, SL = sandy loam, SiL = silt loam, L = loam. Moisture percentage was determined directly by collecting soil samples, weighing in the field, and oven-drying to constant weight at 105°C (moisture percentage thus calculated on a dry weight basis). A total of 122 samples were collected at three depths (0-3 in., 9-12 in., 15-18 in.), where possible, from 12 sites. Collections were made four times during the summer of 1969. Water retention was studied on the original samples collected by horizon, using a pressure plate and pressure membrane apparatus at 1/3 and 15 atmospheres to approximate field capacity and permanent wilting percentage (Richards and Weaver, 1943; Richards, 1965). The chemical properties measured were pH, total carbon, total nitrogen, available phosphorus, exchangeable cations, cation exchange capacity, and oxalate-extractable iron and alum inum. The determinations of carbon, nitrogen, phosphorus, cation exchange capacity, iron and aluminum were done by Mr. B. von Spindler of the Department of Soil Science, University of British Columbia. The determinations of pH and exchangeable cations were done in the Department of Botany. Analysis of pH was done using a Beckman model N pH meter and a Radiometer pH meter, number 24 on soil samples mixed to a paste consistency (Wilde and Voigt, 1955). A Leco total carbon 18 analyzer was used to determine percent total carbon (Allison, Bollen and Moodie, 1965). These figures, when multiplied by the factor 1.72, were used to express percentage of organic matter. Total nitrogen, expressed as a percentage, was measured by the semimicro-Kjeldahl method (Bremner, i960). Available phosphorus, expressed -in.parts per million, was determined color-imetrically by the dilute acid-fluoride extraction method of Bray and Kurtz (19^5). Exchangeable cations (calcium, magnesium, sodium and potassium) were extracted by leaching soil samples with IN ammonium acetate (pH adjusted to 7) and filtering gravi-metrically (Peech et al., 1947). The concentrations of the cat ions were determined on a Perkin-Elmer, model 303* atomic absorp tion spectrophotometer. The results were expressed in milli-equivalents per hundred grams of soil (me/100 g.). Cation ex change capacity (CEC), expressed in me/100 g., was analyzed by the KC1 saturation method (Jackson, 1958). The methods for the determination of nitrogen, phosphorus, exchangeable cations and cation exchange capacity were those used by the Department of Soil Science, University of British Columbia. Percentages of iron and aluminum were determined only on a selected number of samples, in order to classify the sample as a Bf, Bfh, Bhf or Bm horizon. The samples were ground to 100 mesh and extracted following the acid ammonium oxalate procedure of McKeague and Day (1966). The concentrations of iron and aluminum in the extracts were determined by atomic absorption spectrophotometry. The iron and aluminum determinations are presented in Appendix 3. 19 C. Vegetation Synthesis Association tables were made up for each association, show ing variations separately, where present. The synthetic values of presence and average species significance were determined for each species. Presence is defined as the percentage of plots of a particular association in which-.a species occurs. The presence percentages were converted to classes as follows: Presence Class $ of Plots I 1-20 II 21-4III 41-60 IV 61-8V 81-100 In the cases of communities having fewer than four plots, the fraction of plots in which a species occurred was used instead of presence (eg. 2/3). In calculating average species significance, the numbers of the species significance class were converted to percentages using the midpoint value (see Table 4). The average percentage was then converted back to a species significance class number. In the association tables, the species are arranged by layers, by decreasing presence value within each layer, and by decreasing average species significance value within each presence class. Average species significance and sociability are represented by two numbers (eg. 4.2). Sporadic species (those occurring in only one plot) are listed separately by layers. The altitudinal area is indicated as A (alpine), LA (low alpine) or SB 20 (subalpine parkland). The degree of presence together with the average species significance indicates the importance of a species in the community. The species selected as the Characteristic Combinat ion of Species are those with the highest values of both presence and average species significance, or species which are more or less restricted to the particular community. A summary of presence and average species significance for the major species together with all the associations is provided in a synthesis table (Table 75). Only species occurring in at least one association with a presence of IV or V (or correspond ing fraction) were used. In addition, a few species with a presence less than IV are listed, as they are characteristic species for some associations. D. Computational Methods i. Floristic Similarity Index The S/^rensen Index of Floristic Similarity was used to compare the different plots representing a particular associat ion. The index calculated was that based on dominance, Kg(j (Dahl, 1956), the formula being Kgd = 2c. x 100 a + b where a—sum of species significance values of all species in one plot b = sum of species significance values of all species in second plot c= sum of the lesser of the two species significance values of each species in common to both plots 21 The index ranges from 0 to 100, the highest value occurring when a = b = c. This index was also used to compare the different communit ies with each other. The indices were calculated using a computer program devel oped by Ream (1965) and modified by Mr. Stephen Borden of the Biology Data Centre, University of British Columbia, ii. Environment Analysis Among the twenty-three environmental variables dealt with in this study, twelve had values taken at several depths in the soil profile. Since all plots did not have the same horizons, comparisons on a horizon basis would not have been possible. It was thus considered that comparisons between plots and communities would be facilitated by having only one value per plot for each environmental factor. In order to do this, the variables were weighted by depth in the following manner (method by Mr. Stephen Borden, Biology Data Centre, University of British Columbia). For every variable which had been meas ured at several depths, the value for each horizon was multi plied by the depth of that horizon. The sum of these values for all horizons was divided by the total depth of the soil to give an average value for the plot. A one-way analysis of variance between communities was done for each environmental variable, using the weighted values, where necessary. In the analysis of environmental variables (section 7A), the terms "low", "medium" and "high" are based on the average 22 (weighted or,unweighted) value for each community. 23 4. Alpine and Tlmberline Communities A total of fourteen plant associations, with nine variations, are distinguished in this study. They are described below, arranged along a general gradient of increasing moisture, from the xeric Juniperus communis Association to the subhydric drepanocladus exannulatus Association. An alternative, more conventional, method of arranging the communities would be to group them by altitudinal zone (alpine or subalpine). Within each of these groups, the communities could then be ordered by hygrotope. This method, however, is not well suited to the present study, since the communities do not segregate easily according to altitude. Much of the study area is actually an ecotone between the alpine and subalpine zones; thus, many of the communities are represented in both areas. To present the description of the communities in such a manner would be more confusing than instructive, since it would be repetitious. Hygrotope was found to be the most important factor in delimiting the communities (see section 7A), and is therefore used as the basis for the present arrangement. The altitudinal area (s) for each community is mentioned in the description. Despite using a moisture gradient in the present section, the altitudinal approach is still considered to be a useful one. The topographic and altitudinal relationships of the communities are summarized in section 7C In the description of each association, the floristic simi larity indices of the plots comprising the association are given. This provides an objective check on the subjective classification, since plots within an association should have their highest similarities to each other rather than to a plot ln another association. If the similarity values among all the plots of an association are high, the association is considered to be homogeneous. An association with a very large number of species is usually less homogeneous than one with few species. Few studies have utilized this technique, and thus the evaluat ion of the indices is strictly empirical. Nonetheless, the method is believed to be of considerable value. Juniperus communis Association (Ref. Tables 6, 1, 8, 9, 10; Fig. 3) Characteristic Combination of Species Juniperus communis Carex phaeocephala Arenaria capillaris Festuca brachyphylla Polytrichum piliferum Tortula ruralis Peltigera malacea This association occurs over rock outcrops on ridges and slopes in the low alpine and alpine areas. The relief shape is generally straight, and the exposure is usually southeast. Slope gradients range from 26 to 70$. Rock comprises 85-95$ of the ground surface and humus 4-10$, with very little mineral soil exposed. No erosion was observed. The hygrotope is rated as xeric. There is a we11-developed B layer, ranging from 85-100$ of the coverage, poorly developed C and D layers, and an occasional occurrence of an E layer. The C layer covers 10-35$ of the Table 6 General Environment Juniperus: communis Association. Plot No.. 50 47 48 52 Elevation (ft.) 7550 7450 7450 _ 7425 Phy s-io graphy Landfcrm ledge slope ridge ridge Relief shape straight convex straight straight Exposure SW SE SE SE Slope gradient (%) 70 29 35 26 Layer Coverage (%) B layer C layer D layer E layer 100 10 20 1 98 35 .10 0 98 30 7 1 85 25 35 0 Plot Coverage (%) Humus Mineral soil Rock 5 0 95 4 1 95 10 0 90 10 5 85 Soil Hygrotope Erosion Horizon depth (in.) Ah C R Classification 0-5 5-10 10+ xerxc - none 0-6 0-12 . 0-15 6-12 12+ 12+ 15+ Lithic Orthic Regosol Table 7 Juniperus communis Assoc!ation 26 Plot No. 50 47 48 52 Plot Size (m2) 12 26 14 14 Extent of type (m2) 12 26 14 14 Elevation (ft.) 7550 7450 7450 7425 Altitudinal area A LA LA LA B layer Presence Aver.Sp Signifi 1 Juniperus communis 10.7 9.7 9.7 9.6 V 9 C layer 2 Carex phaeocephala 4.2 3.2 5.2 3.2 V 4 3 Arenaria capillaris 1.2 2.2 3.2 3.1 V 2 4 Festuca brachyphylla 2.2 2.2 3.2 1.2 V 2 5 Vaccinium scoparium 1.2 5.2 - 4.2 IV 4 6 Antennaria lanata 1.2 2.2 - 4.2 IV 2 7 Selaginella densa - 3.1 1.1 2.1 IV 2 8 Sibbaldia procumbens - 2.1 1.1 1.2 IV 1 9 Luzula spicata 1.1 - +.+ 2.1 IV 1 10 June us parryi - 4.2 - 4.2 III 3 11 Potent!11 a di vers!fol!a - 2.2 2.2 - III 1 12 Erigeron peregrinus - 2.2 - 1.2 III 1 13 Trisetum spicatum 1.2 - 2.1 - III 1 14 Arenaria obtusiloba - - 1.1 1.2 III + D layer Bryophytes 15 Polytrichum piliferum Dh 2.2 4.2 3.1 6.2 V 5 16 Tortula rural is Dh 2.2 2.2 3.2 3.1 V 3 17 Barb!1ophozi a hatcheri Dh 3.2 2.1 +.1 - IV 1 18 Dicranum scoparium Dh 1.1 1.2 - 2.1 IV 1 19 Bryum capillare Dh 1.1 1.1 - - III + Lichens 20 Cetraria ericetorum Dh 3.2 - 2.1 4.2 IV 3 21 Peltigera malacea Dh - 3.2 +.+ 2.2 IV 1 22 Cladonia ecmocyna Dh - 1.1 - 4.2 III 2 23 Lecidea granulosa Dh - 1.1 - 1.1 111 + 24 Sol ori na crocea Dh 1.1 1.1 . - - III + E layer 25 Cetraria pinastri EB 1.1 - +.+ - III + Total Species (incl.sporadics ) 21 25 21 23 26 27 28 29 30 31 32 33 Sporadic Species C layer Carex spectabilis Haplopappus lyallii Hieracium gracile Saxifraga bronchial is Vaccinium caespitosum D layer Bryophytes Barb! 1 ophozi a lycopodi oides Drepanocladus uncinatus Hypnum revolutum 47(4.2) 50(1.2) 52(2.1) 48(1.2) 48(1.2) 52(1.1) 50(4.2) 48(2.2) 34 35 36 37 38 39 40 41 42 43 44 Polytrichadelphus lyallii Rhacomitri um canescens ? Tetraplodon mnioides Lichens Cetraria islandica Cladonia carneola Corn! cul ar! a aculeata Peltigera canina Stereocaulon alpinum E layer Alectoria minu'scula Parmeliopsis hyperopta Psoroma hypnorum 47(1.1) 47(1.1) 52(3.1) 47(2.1) 47(1.1) 50(1.2) 50(1.1) 52(4.1) 48(+.+) 48(+.+) 50(1.1) 27 plot, while the D layer occupies 7-35$. The E layer only occurs in two plots, with a cover of one percent. Juniperus communis, which is the dominant species in the community, is the only species in the B layer, with a presence of V and average species significance of 9. There are only three constant species in the C layer, all with low cover values. These are Carex phaeocephala, Arenaria capillaris and Festuca  brachyphylla. Polytrichum pillferum and Tortula ruralis com prise the constant bryophytes, while there are no lichens with a presence of class V. Peltigera malacea, with a presence of IV, and Tortula ruralis show a high preference for this association. Table 8 gives the floristic similarity indices for the four plots comprising the association. The values are all very high, indicating that the association is homogeneous. Table 8 Floristic Similarity Indices for the Juniperus communis Assoc iation 50 47 48 52 ' 50 69 76 63 47 70 74 48 67 52 The soils are all classed as Lithic Orthic Regosols, having an Ah-C-R horizon sequence. Both the A and C horizons are coarse textured. All the samples are classified as loamy sands. Plot No. Ah Horizon Textural class Sand (%) Silt (%) Clay (%) C Horizon Textural class Sand (%) Silt (%) Clay (%) 28 Table 9 Soil Texture Juniperus communis Association 50 47 ,48 52 LS LS LS LS 77.4 82.8 84.8 79.4 22.6 17.2. 15.2 19.0 0 0 0 1.6 LS LS 73.9 79.2 26.1 20.4 0 0.4 29 Table 10 Soil Chemical Analysis Juniperus communis Association .ot No. 50 47 48 52 i Horizon pH 4.9 4.9 4.8 4.5 C (%) 12.1 7.8 9.2 12.7 OM (% ) 20.8 13. 4 15.8 21.8 N (%) 0.1 0.5 0.6 0.9 C/N 201. 17. 15. 15. P (ppm) 7. 9. 17. 18. Ca (me/lOOg) 0. 32 0.08 1. 39 0.13 Mg (me/lOOg) 0.12 0.08 0.14 0.09 Na (me/lOOg) 0.04 0.03 0.02 0.12 K (me/lOOg) 0.23 0. 07 0. 25 0.14 CEC (me/lOOg) 36.6 27.3 74.8 121.0 Horizon pH 5.0 5.2 - -C (%) 8.9 5.7 - -OM- (-%•) 15. 3 9.9 - -N (%) 0.8 0.3 - -C/N 11. 18. - -P (ppm) 8. 6. - -Ca .(me/100g) 0.05 0.27 - -Mg (me/lOOg) 0.07 0.05 - -Na (me/lOOg) 0.04 0.04 - -K (me/lOOg) 0.07 0.02 - -CEC (me/100g) 21.5 23.5 - -30 Pig. 3. Juniperus communis Association, Plot 47. Table 10 presents the soil chemical data for this associat ion. The pH values are slightly lower in the Ah than in the C horizon, but all values are strongly acidic. Organic matter decreases somewhat with depth, but due to the very shallow nature of the soils, there is still a considerable amount of organic matter in the C horizon. Carbon-.nitrogen ratios are generally narrow. There is an extremely low percentage of total nitrogen in the Ah horizon of plot 50, thus making the C/N ratio very wide. The amount of phosphorus and sodium is approximately the same in the A and C horizons. Magnesium, potassium and cation exchange capacity decrease with depth; calcium decreases in one case and increases in another. In general, phosphorus, cation exchange capacity and available cations are quite variable among the plots of the association; carbon:nitrogen ratios are similar. Antennaria lanata - Sibbaldia procumbens Association (Ref. Tables 11, 12, 13, 14, 15; Pig. 4, 5, 6, 7) Characteristic Combination of Species Antennaria lanata Sibbaldia procumbens Polytrichum piliferum Umbilicaria hyperborea Alectoria minuscula Rhizocarpon geographicum This association occurs on ridge tops, predominantly in the alpine area. The relief shape is generally straight. Expos ure is variable, being south, southwest, north, northwest or neutral. Slope gradients range from 0 to 28$. The ground surface is covered by 5-85$ rock, 5-70$ humus and 0-25$ mineral Table 11 General Environment Antennaria lanata - Sibbaldia procumbens Association Antennaria - Sibbaldia - Salix Van* ati on Carex phaeocephala Vari ati on 32 Carex breweri Variati on Plot No. 10 Elevation (ft.) 7600 Physi ography Landform 22 7600 13 7500 28 7500 9 7600 ridge D T x u J. • ux straight , . , , convex to Relief shape straight . straight to convex concave convex to straight 17 7500 ridge 67 7325 straight straight 45 7500 46 7490 ridge cliff face straight convex , to convex Exposure N S neutral SW SW S neutral neutral NW Slope gradient 17 6 0 15 13 12 0 0 28 Layer Coverage {%) B layer 0 5 2 0 0 0 0 0 0 C layer 35 45 30 35 40 30 50 75 60 D layer 50 85 65 90 60 80 75 15 40 Plot Coverage {%) Humus 36 10 38 5 27 20 55 70 58 Mineral Soil 4 10 2 10 3 5 0 25 2 Rock 60 80 60 85 70 75 45 5 40 Soil Hygrotope Erosi on Horizon depth (in.) L-H Ah B C R AM 0-2 Ah2 2-4 Bm 4-12 12+ . xeric strong 0-2 0-3 Bf 3-6 2-12 Cgj 6-14 12+ 14+ 0-6 6-14 14+ 1 2J-0 0-3^ xeric moderate strong none 0-4^ 3^-12 4J--13 12+ 13+ 0-7 7+ xeric xeric strong moderate 0-6 Bf 6-15 Cg 15+ AM 0-7 Ah2 7-10 10+ Classification Alpine Lithic Dystric Orthic Brunisol Regosol Lithic Gleyed Sombric Humo-Ferric Podzol Lithic Orthic Regosol Lithic Orthic Regosol Gleyed Sombric 0rthic Humo-Ferric Regosol Podzol Table 12 Antennaria lanata - Sibbaldia croc—tens As^oci all on Antennaria lanata - Sibbaldia Carex phaeoqeshala  _procu:bens - Sallx cascaden^fs Variation Variation Carex breserl .Variation Plot No. 10 22 13 23 9 •17 67 45 46 Plot Size Cn2) 70 70 70 50 120 105 8 6 .5 Extent of type (n?) 210 104 500 96 360 105 16 6 5 Elevation (ft.) 7600 7600 7500 75 00 7600 7500 7325 75 00 7*90 Altitudinal area A A LA A A. LA • SP. A A B layer Aver.Species Aver.Spec'les Sign!flcance Significance Picea engelnannii - +.+ +.+ + '• _ ' _ Association Aver,Species presence Aver.Species Significance Sigr.i f lcar.ce C layer . 2 Carex phaeocephala 3 Antennaria lanata Sibbaldia procumbens _ Antennaria untJri nel 1 a Festuca'brachyphylla Arenaria capillaris Arenaria obtusiloba Sallx cascadensis Haplopappus lyallii Agrostis variabilis Luzula splcata Trlsetun splcatun Potentilla diversifolia Seduilanceolatun Juniperus consunls Juncus parryl Selaginella densa Carex-breweri Oryas octopetala Carex spectabilis" Vacclnlua scopariun Luzula arcuata Saxifraga bronchial is Pinus albicaulis Q layer Bryophytes Polytrichum piliferun Barbllophozla hatcher! Orthocaulis floerkii Bryun capillare Ceratodon purpureus Gricicla alpestris tophozla alpestris Lichens 33 Solorina crocea 34 Uablllcaria hyperborea 35 Alectoria ninuscula 36 Lectdea granulosa 37 Rhlzocarpon geographicuni 38 Cladonia carneola 39 Peltigera canlna 40 Cetraria erfcetorua 41 Cetraria Island 1ca 42 Stereocaulon alpinun 43 Cornicularia aculeata 44 Cladonia eccocyna 45 Cladonia chiorophaea 3.2 3.2 4.2 3.2 4.2 5.2 .4.2 5.2 3.2 4.2 5.2 5.2 3.2 4.2 5.1 2.1 4.2 3.1 4.1 4.2 1.1 2.2 1.2 1.1 3.1 3.2 3.2 2.1 4.2' 5.2 6.2 3.2 4.2 4.2 4.2 4.2 - 3.1 4.1 • 4.2 2.) 2.1 3.1 3.1 2.1 2.1 3.1 3.2 2.2 2.2 3.2 2.2 1.1 1.1 1.1 1.1 +.+ 1.+ +.+ _ -3.2 . 1.2 - 1.1 1.1 1.1 6.3 5.3 _ ' _ - 2.2 - 1.2 2.2 2.2 2.2 : 1.* Oh 5.2 5.2 6.2 7.2 Oh - 2.1 - 1.1 Oh 2.1 - 2.1 _ Oh 1.1 - _ _ Oh . 1.1 - _ Or - - 1.1 _ Oh - - - 1.1 Dh 3.1 3.2 4.2 4.2 Dr 5.3 6.2 5.3 7.3 Dr 5.3 6.2 5.3 7.3 Dh 2.1 4.2 . 2.1 Or 3.1 4.1 3.1 4.2 Dh 3.1 4.2 2.1 . Oh 2.1 +.+ _ 1.1 Oh - 4.2 4.2 _ Oh 2.1 - _ 3.1 Oh - - 1.1 Oh 3.1 - 4.2 Oh - - 2.1 Dh - - . 3.2 6.3 6.3 7.2 6 4.2 . 3 V 5 4.2 3.2 3.2 .4 ' 7.2 4.2 6 V - 5 4.2 4.2 5.2 . . 4" 4.2 5.2 5 V 5 2J 2.1 5.2 4 . : 3.2 2.1 3 V . 4 4.2 2.2 4.2 4 4.2' . 3.2. 4 v ••• ^ 3.2 3.2 6.2 5 . 2.1 - •• 1' "'• V 3 3.2 3.2 2.1 .3 2.2 1.1 . 1 . .V 3 - 2.2 - . .'• • 5.2 1.2 4 IV . 3.3 2.2 2 4.2 3 IV 3 - 3.2 4.2 . 3 3.1 - ' 1 IV 3 3.2 3.2 4.2 • . 4 - IV 3 3.2 1.1 - 1 2.2 - 1 IV 2 2.2 2.1 - 1 - - - IV 1 - 2.1 - + 1.1 - + IV . i 1.+ +.+ • - . - IV + 2.2 +.2 - + - III + - 1.1 • - • - III • - - - - 8.3 7.2 II. 5 - - - • - - II 3 - 2.2 - • , II + - - 1.2 1.1 II - - - .2.1 2 II + - : - - - • •••'-. II. + ' II + 4.2 5.2 6.2 5 • 4.2 6.2 5 V ' - 5 - 1.1 - • 3.1 - 1 III 1 1.1 - - • - - -' II + 1.1 - - • - - II - - - . - 1.1 - + II + - 1.1 - • - - II • ' ~ - - 1.1 . + II + 2.2 3.1 2 2.2 4.2 3 V 3 5.3 6.3 5.1 5 _ _ IV 5 5.3 6.3 . 5 -m IV 5 - - 8.1 6 1.1 2.1 1 IV 4 3.1 3.1 5.1 4 - _ _ IV 1, - 1.2 3.1 1 2.1 3.1 3 IV 2 1.1 •.2 - • 2.1 1 IV - 3.2 4.1 3 - ZA • 1 . III 3 3.2 2.2 - 2 2.2 - 1 III 1 1.1 1.1 - * 1.1 - + III •- - - . _ II - - • 3.1 1 3.2 1 II 1 - 1.2 - • - - - II + 27 33 20 24 20 Total Species (incl.sporadics) 32 34 29 32 Sporadic Species C layer Ranunculus eschscholtzli Saxifraga ferrugtnea 9(1.1) 46 Abies lasiocarpa 22(1.*) 55 Solldago rultiradiata 17(1.1) 60 47 Carex nardlna 46(6.2) 56 Stellarla laeta 10(1.1) 48' Carex pyrenalca 22(2.2) 49 Erigeron peregrinus 28(1.2) 0 layer 61 SO Hieraclun gracile 67(3.1) Bryophytes 62 51 Luzula lahlenbergil 46(4.2) 63 52 Picea-engelnannii 101+.+) 57 Barbilophozla lycopodioidss 3(1.!) 64 Olcranun-scopariun Pohlia gracilis Pohlia nutans Lichens 13(2.1) 28(1.1) 46(1.1) 46(2.1) 9(2.1) 9(1.2) . 10(3.2) 34 soil. Erosion is moderate to strong, wind being the important factor. The hygrotope is rated as xeric. A poorly-developed shrub layer occurs in only two plots, covering 2-5$ of the area. The moderately-developed herb layer covers 30-75$, while the we11-developed bryophyte and lichen layer occupies 15-90$. Antennaria lanata and Sibbaldia procumbens are the dominant species in the C layer. Other constant sub-dominants are Carex  phaeocephala, Antennaria umbrinella, Festuca brachyphylla, Arenaria capillaris and Arenaria obtusiloba. Polytrichum plll-ferum is the only constant bryophyte. Solorina crocea is the only constant lichen, but due to its low cover and low prefer ence for this community, it is not considered as a characteris tic species. However, Umbilicaria hyperborea, Alectoria minus-cula and Rhlzocarpon geographicum are considered to be charact eristic species because of their high preference for this assoc iation . The association is subdivided into three variations, based mainly on floristic composition. The variations are: a. Antennaria lanata - Sibbaldia procumbens - Salix  cascadensis Variation b. Carex phaeocephala Variation c. Carex breweri Variation Table 13 shows the floristic similarity indices for the nine plots of the association. The blocked-in areas represent the three variations. It can be seen that the values are generally higher within the variations than among them. The indices are still fairly high between the Antennaria - Sibbaldia - Salix and the Carex phaeocephala variations, but are much lower with the Carex brewer! Variation. Table 13 Floristic Similarity Indices for the Antennaria lanata -Sibbaldia procumbens Association 10 22 13 28 9 17 67 45 46 10 68 64 53 22 64 65 13 62 28 61 51 57 51 56 69 59 62 33 40 51 41 9 17 67 45 46 38 42 40 36 35 32 24 28 28 38 35 22 26 30 The variations are described below, by general habitat, floristics and detailed soil data. a. Antennaria lanata - Sibbaldia procumbens - Salix cascadensls  Variation This variation, which is the type for the association, occurs on ridges in the alpine and low alpine areas, with a relief shape varying from straight through convex to concave. Exposure varies from north to south and southwest. Slope gradients range from 0 to 17$. The ground surface is 60-85$ rock, 5-38$ humus and 2-10$ mineral soil. Erosion is strong, and the hygrotope is xeric. The bryophyte and lichen layer is the most prominent, covering 50-90$ of the area. The herb layer covers 30-45$, while the shrub layer covers 0-5$. In addition to the dominant species listed for the assoc iation, the following species are important in differentiating this variation from the others: Salix cascadensis, Haplopappus  lyallii, Sedum lanceolatum (which all show a high preference for this variation), and Dryas octopetala, which is exclusive to this variation. Soil types associated with this variation are Lithic Orthic Regosol (2), Alpine Dystric Brunisol (l) and Lithic Gleyed Sombric Humo-Ferric Podzol (l). Generally, the C horizon has a coarser texture than the Ah or B horizons. The textures of the Ah horizon range from sandy loam to loamy sand and sand. The B horizon, where present, is classed as sandy loam or loamy sand. The C horizon is either sand, sandy loam or loamy sand. In one case, fineness of texture increases with depth (clay content increases from 0 to 7$). The soil chemical data are given in Table 15. The pH increases slightly with depth, but all values are strongly acidic. Organic matter and nitrogen decrease steadily with depth, and the carbon:nitrogen ratios are narrow. Phosphorus, magnesium, potassium and cation exchange capacity decrease with depth. In general, calcium decreases from the A to the B horizon, then increases somewhat from the B to the C horizon. & Table 14 Soil Texture Antennaria lanata - Sibbaldia procumbens Association 37 Vari ati on Plot No. 10 22 13 28 Ah Horizon Textural cl ass LS SL S LS Sand {%) 73.4 67.4 89.4 78.6 Silt (?) 26.5 31.8 10.6 21.4 Clay {D 0.1 0.8 0 0 B Horizon Textural class SL - LS -Sand (%) 63.6 - 84.2 -Silt {%) 35.6 - 15.8 -Clay (I) 0.8 - 0 -C Horizon Textural class S LS SL S Sand (?) 90.8 • 82.8 74.0 92.2 Silt (%) 9.2 17.2 19.0 7.8 Clay {%) 0 0 7.0 0 LS 73.8 24.8 1.4 SL 73.8 21.8 4.4 Carex phaeocephal a Carex breweri Variation Variation 9 17 67 45 46 LS LS SiL LS SL 75.8 78.4 36.4 71 .4 59.1 24.2 21.2 56.8 28.6 38.2 0 0.4 6.8 0 2.7 LS 81.6 18.4 0 LS 84.0 15.8 0.2 LS 76.4 23.2 0.4 Table 15 'Plot No.. L-H Horizon • pH . C (I) OH (1) N (I) C/N • P (pps) Ca (oe/100g) Bg (w/IOOg) Na (ne/lOOg) X (oe/IOOg) CEC (oe/IOOg) Ah Horizon pH C (I) oa (J) N (j) . C/N p (pp») Ca (oe/IOOg) Hg (oe/IOOg) Na (oe/IOOg) K (ce/IOOg) CEC (oe/IOOg) B Horizon pH" C (J) OK (!) N (J) C/N P (ppn) Ca (oe/IOOg) Kg (oe/IOOg) Na (oe/IOOg) K (oe/100g) CEC (oe/IOOg) C Horizon pH C (I) OM (I) N (I) C/N P (ppe) Ca (oe/IOOg) Bg WlOOg) Na (m/tOOg) K (n/IOOg) CEC (w/IOOg) Antennaria - Sibbaldia - Salix Vari ati on Sol) Choral cal Analysts Antennaria lanata • Sibbaldia procurabene Association Carex phaeocephala Variation 4.7 9.3 16.0 ' 0.6 16. 14. U6 0.17 ' O.H 0.18 33.7 28 17 67 4.7 16.8 29.0 0.7 24. 17. . 0.9* 0.20 0.17 0.08 61.8 Carex brenerl ' Variation 4.9 . 4.9 5.1 5.2 4.9 4.9 5.1 5.4 8.* 7.0 • 7.6 ... 10.1 *.o 8.9 10.0 6.8 14.5 12.1 13.1 17.3 .6.9 15.4 17.2 11.7 0.4 . 0.* 0.5 .0.6 0.2- 0.5 0.7 0.5 20. 18. 15. 1B. . • . 18. 20. 14. 14. 10. 8. • 11. 21. 5. 17. 4. 7. 1.9* . 1.06 • 0.47 0.2* 0.27 1.38 ' 0.78 0.42 0.2B 0.11 . 0.09 0.03 0.03 0.17 • 0.09 0.02 0.23 0.16 0.12 0.10 0.15 0.14 . 0.05 0.07 0.3* 0.20 0.13 0.02 0.08 0.13 0.16 0.00 2*.6 25.9 37.9 *8.3 H.O 28.5 14.9 . 32.3 5.5 5.1 _ _ 5.3 3.7 - - 2.8 - - 2.1 6.4 - 4.8 • - _ 3.5 0.3 - 0.2 - - _ 0.1 _ 12. - 16. - _ 17. m 3. - 7. - - ... 6. 0.23 - 0.47 - 0.27 m 0.03 - 0.02 -• 0.01 m 0.13 - • 0.18 - _ _ 0.05 m 0.03 - 0.04 - - 0.00 m 8.3 19.9 ." - • • - - 22.6 . -5.6 • 5.3 5.8 . 5.2 5.3 5.5 ' 5.8 5.7 1.1 2.0' 0.5 2.1 5.5 1.0 0.5 2.9 1.9 3.4 0.9 3.6 9.5 1.7 0.8 5.0 0.1 0.2 0.0 0.1 0.4 0.1 0.0 0.2 18. 11. 13. 21. 16. 12. - 15. 19. 5. ' 7. 4. 8. 11. 4. 6. 5. 0.33 0.42 0.80 2.23 0.30 0.40 0.47 0.80 0.02 0.03 . 0.01 0.02 0.02 0.01 . • • - 0.02 0.02 0.14 0.15 0.32 . 0.55 0.10 0.16 0.08 0.09 0.02 0.04 0.02 0.00 0.04 0.03 0.00 0.08 16.5 17.9 . 16.3 12.8 11.7 22.0 28.5 31.5 39 Fig. 4. Antennaria - Sibbaldia Association, Antennaria -Sibbaldia - Salix Variation, Plot 28. Note high coverage of rock lichens. Krummholz belongs to Abies laslocarpa Association and Picea engelmannii Association. Fig. 5. Soil profile of Antennaria - Sibbaldia Association, Antennaria - Sibbaldia - Salix Variation, Plot 22. This soil is classified as a Lithic Orthic Regosol, with an Ah-C-R horizon sequence. ko In one case, it increases considerably from the A to the C horizon. Sodium increases with depth, b. Carex phaeocephala Variation This variation also occurs on ridges, with a straight relief shape. Exposures are south and southwest. Slope grad ients range from 0-13$. The ground surface is 45-75$ rock, 20-55$ humus, and 0-5$ mineral soil. Erosion varies from none to strong. The hygrotope is xeric. This variation occurs main ly in the alpine area, with one occurrence in the subalpine park land. As in the previous variation, the D layer is the most impor tant, covering 60-80$ of the area. The herb layer coverage is approximately the same, occupying 30-50$. There is no shrub layer. The species important in the differentiation of this varia tion are Carex phaeocephala and Arenaria capillarls, both with a much higher average species significance than in the other two variations. The soils are all classed as Lithic Orthic Regosols, with an Ah-C horizon sequence. The Ah horizon in the alpine sites is a loamy sand, while it is a silt loam in the subalpine plot. The C horizon, where present, is either a loamy sand or sandy loam. The chemical data on pH, organic matter, nitrogen and carbon:nitrogen ratios are as described for the Antennaria -Sibbaldia - Sallx Variation. In plot 9> organic matter is still relatively high in the C horizon because of the shallowness of the soil. Phosphorus, magnesium, potassium and cation exchange capacity decrease with depth, and sodium increases, as in the previous variation. Calcium increases from the A to the C horizon. The available cations are present in smaller quantit ies in the A horizon of the Carex phaeocephala Variation as compared to the Antennaria - Sibbaldia - Salix Variation. In the C horizon, there is less calcium and sodium, and similar amounts of magnesium and potassium, c. Carex breweri Variation This variation occurs in the alpine area on ridges and cliff faces. The relief shape is convex to straight. Exposure is northwest for one plot and neutral for the other. The slope gradient is 28$ in one £lot and 0 in the other. The ground surface is only 5-40$ rock, which is much less than In the other two variations. There is a much higher cover of humus, occupying 58-70$ of the area. Mineral soil ranges from 2-25$ of the plot coverage. Erosion is moderate to strong. The hygrotope is xeric. As in the Carex phaeocephala Variation, there is no shrub layer. The herb layer is the most prominent, covering 60-75$ of the area. The bryophyte and lichen layer is reduced to 15-40$ coverage, this being mainly due to the lack of the rock lichens Umbillcaria hyperborea, Alectorla minuscula and Rhlzo-carpon geographicum. This community is a closed one, whereas the other two are open. The important species differentiating this variation are Carex breweri, which is exclusive to it, and Luzula arcuata, 42 Fig. 6. Antennaria - Sibbaldia Association, Carex phaeocephala Variation, Plot 9. Fig. 7. Antennaria - Sibbaldia Association, Carex breweri Variation, Plot 45. Note high coverage of Antennaria  lanata (light green leaves). which has a high preference for it. The soils vary from Orthic Regosol to Gleyed Sombric Humo-Ferric Podzol. Texture becomes coarser with depth in plot 46. All samples, except for one Ah horizon, are classed as loamy sands. The Ah horizon in plot 46 is a sandy loam. The pH values for the Ah horizon are slightly higher than in the other two variations, but they increase in the C horizon to similar values. Organic matter, nitrogen, carbon:nitrogen ratio, magnesium and sodium are as described for the other variations. Potassium and calcium decrease with depth in one case, and increase in another. Cation exchange capacity increases with depth or remains the same. In the A horizon, the exchangeable cations are all present in smaller quantities than in the Antennaria - Sibbaldia - Salix Variation. There is more calcium and less sodium than in the Carex phaeocephala Variation. In the B horizon, there is less magnesium, sodium and potassium than in the Antennaria - Sibbaldia - Salix community. In the C horizon, there is less sodium and potas sium than in the type variation, and less potassium than in the Carex phaeocephala Variation. Juncus parryi Association (Ref. Tables 16, 17, 18, 19, 20; Fig. 8) Characteristic Combination of Species Juncus parryi Antennaria lanata Polytrichum piliferum Lecidea granulosa This association occurs on south-facing slopes in the alpine and low alpine areas. It is less well developed in the subalpine parkland, occurring there on slopes and ridges having a southern exposure. Slope gradients range from 10 to 35$ > the steeper slopes being In the alpine zone. Relief shape varies from straight to convex to concave. The slopes are fair ly rocky, rocks covering 15 to 60$ of the ground surface. Humus covers 35-82$, while exposed mineral soil occupies only 0-20$. Erosion varies from none to moderate and, in one case, strong. The hygrotope is rated as subxeric. The herb layer is very well developed, occupying 40-85$ of the area. The bryophyte and lichen layer Isiless well developed, coverage being 15-60$. In the C layer, Juncus parryi, with an average species significance of 8, and Antennaria lanata, with an average species significance of 5, are the dominant plants. Other constant species are Arenaria capillaris, Hieracium gracile and Sibbal dia procumbens. Vaccinium scoparium and Erigeron peregrinus are prominent in a few plots. In the D layer, Polytrichum piliferum is the only constant bryophyte, with an average species signif icance of 6. Among the lichens, Lecidea granulosa Is the only constant, with an average species significance of 5. Table 18 gives the floristic similarity indices for the eight plots comprising the association. The majority of the Table 16 General Environment Juncus parryi Associ ation 45 Plot No. 12 Elevation (ft.) 7575 Physi ography Landform Relief shape straight Exposure SE Slope gradient {%) 24 7550 21 7550 40 7500 30 7420 sl ope' straight straight to convex to convex S 34 S 25 convex to concave 35 S 24 59 7350 7325 ridge SE 10 14 63 7275 base of ridge convex straight concave concave SE 12 Layer Coverage {%) C layer 85 D layer 15 70 40 60 45 70 40 65 40 75 45 40 60 75 25 Plot Coverage {%) Humus Mineral Soil Rock 10 65 0 35 35 5 60 65 5 30 65 5 30 55 5 40 35 0 65 70 20 10 Soil Hygrotope Erosi on Horizon depth (in.) L-H 1-0 Ah 0-5 .1 moderate 0-5 Bfh 5-15j Bhf 5-16 Cg 15y + 16+ 0-6 6-13 13+ — subxeric none strong 0-5 Bhf 5-8 Bm 8-20 20+ none none moderate 0-6 Bm 6-12 12+ 0-9 9-13 13+ 0-7 Bfh 7-10 10+ 0-4 4+ Classification 61 eyed Sombric Humo-Ferric Podzol Sombric Lithic Ferro-Humic Orthic Podzol Regosol Sombric Alpine Lithic Sombric Ferro-Humic Dystric Orthic Humo-Ferric Podzol Brunisol Regosol Podzol Orthic Regosol Table 17 Juncus parry! Association Plot Ho. 12 8 21 . 40 30 59 68 63 Plot Size (n?) 10 10 10 . 10 10 8 •6 10 Extent of type (B-) 20 21 27 ' 16 27 15 11 18. Elevation (ft.) 7575 7550 7550 75 00 7420 7350 7325 7275 Altitudinal area A A A LA LA SP SP SP C layer ' Presence. Aver.Sp' Signifii 1 Juncus parry! 8.2 8.2 8.2 8.2 7.2 8.2 7.2 8.2 V 8 2 Antennaria lanata 4.2 4.2 5.2 5.2 4.2 7.2 4.2 7.2 . V. 5 3 Arenaria capillaris '4.2 3.2 3.1 2.2 3.2 . 3.1 . 6.2 . V 4 * Hieraclua graclle 6.2 +.2 . 4.2 3.2 6.2 2.1 2.1 V 4 5 Sibbaldia procur.bens 4.2 3.2 4.2 5.2 . 5.2 3.2 4.2 - V . 4 6 Vacclnluiii scoparius - 3.1 . 1.2 6.2 5.2 _ 3.1 IV. 4 7 Carex phaeocephala 2.2 5.2 4.2 2.2 3.2 3.2 IV 3 8 Antennaria unbrinella 2.1 - 2.2 2.1 2.1 - 3.2 IV . 1 .9 Luzula splcata 1.1 1.1 1.1 - 2.1 3.2 - IV 1 10 Erigeron peregrin'us •- -' -.- 6.2 3.2 5.2 • - 2.2 III 4 11 Agrostls variabilis 1.1 +.1 4.2 - 2.1 - - - III 1 12 Carex pyrenalca 1.2 - - - 4.2 • - 3.2 - II 1 13 Carex spectabilis - - 2.2 - 4.2 - 2.2 - II 1 14 Claytonla lanceolata - - - - - 4.1 4.1 II 1 15 festuca brachyphylla - 4.2 1.1 - 3.1 - - - ll 1 16 Luplnus 1ati foli us - - - 1.2 4.2 II 1 17 Poa cusicki1 +.+ - - +.+ - - 5.2 . II 1 18 Selaglnella densa 1.1 2.1 4.1 '- - - - II 1 19 Trisetua spicatun . +.1 1.1 _ - 4.2 - ' - II 1 20 luzula wahlenbergti 2.2 . 3.2 1.2 - - - II + 21 Arenaria obtusiloba - - 1.1 - 1.1 - _ - II + 22 Carex nigricans 2.3 - ' - . - - 1.2 - II + 23 Luzula sp. - - - - - - 3.2 1.2 II + 0 layer Bryophytes 24 Polytrlchun piliferum 0h4.2 7.2 7.3 .6.2 5.2 6.1 7.2 5.2 V- . 6 25 Ceratodon purpureus Oh . 2.1 - 1.1 1.1 2.1 - Ill + 26 Polytrichadelphus lyallii Dh2.2 - - 4.2 - - - 3.1 II 1 27 Polytrlchun fornosun Dh 3.1 - - 2.2 1.1 - - - II + Lichens 28 Lecidea granulosa Dh - 1.1 3.2 1.1 3.1 4.1 8.2 3.1 V 5 29 Cladonia carneola Oh - 3.2 2.1 4.2 1.1 1.1 IV 2 30 Solorina crocea 0h+.+ 2.1 3.+ 3.1 . 3.1 IV 1 31 Cladonia ecnocyna Dh - 3.2 2.1 3.2 4.2 - 111 2 32 Cetraria tslandica Dh - 3.1 3.2 2.2 - - - II 1 33 Cetraria ericetorun Dh - - - 4.2 - 3.1 II 1 . 34 Peltigera canina Dh +.+ 1.1 - 1.1 - - II + 35 Cladonia sp. Oh - 2.1 - - . 2.1 II + 36 Cladonia pyxidata Oh - 2.1 1.1 - II + Total Species (Incl.sporadics) 23 22 25 21 26 18 15 18 Sporadic Species . C layer 0 layer 37 Anenone occidentals 63(2.2) Bryophytes 38 Arnica 1atifolia. 63(1.1) .39 Carex brevlpes 63(2.1) .49 Barbllophozla hatched 21(3.1) 40 DeschazipsU atropurpurea 59(3.2) . 50 Desr.atodon latifolius 59(3.1) 41 Juniperus co.-nunls 2K+.+) Lichens 42 Luzula glabrata 12(2.2) 43 Phleun alpir.un 12(1.2) 51 Alectorla ninuscula 8(2.1) 44 Potentllla diversifolia 21(1.2) 52 Peltigera canina var. rufescens 40U.+) 45 Sedun lanceolatun 2K+.+) 53 Peltigera lepidcphora 21(+.+) 46 Vacclnlu.i caespi tosur, 8(1.1) 54 Rhizocarpo.i geographicu: 8(4.2) 47 Valeriana sitchensis 63(1.2) 55 Stereocaulon alplnuT 12(1.1) 48 Veronica sornskjoldil 12(3.2) 56 Ur.bi Tlcarla hyperborea 8(4.2) 47 values are quite high. Table 18 Floristic Similarity Indices for the Juncus parryi Association 12 8 21 40 30 59 68 63 12 56 58 64 51 58 45 52 8 74 60 50 54 54 50 21 68 57 57 60 51 40 59 66 53 55 30 50 55 42 59 42 65 68 36 62 The soils associated with this community are Sombric Humo-Ferric Podzol (2), Sombric Ferro-Humic Podzol (2), Orthic Regosol (3), and Alpine Dystric Brunisol (l). Podzols predomin ate in the alpine sites, while regosols are the major soil class in the subalpine plots. The soil texture results are shown In Table 19. Texture becomes coarser with depth. The A horizon samples are mainly loamy sands or sandy loams. The B horizon ranges from sandy loam to sand. Sands predominate in the C horizon. Plot 68, which occurs in the subalpine parkland, is a finer textured soil, all horizons being sandy loams. Table 20 presents the soil chemical data for the associat ion. The values for pH increase slightly with depth, but all 48 Table 19 Soil Texture Juncus parryi Association .ot No. 12 8 21 40 30 59 68 63 L Horizon Textural class LS LS LS SL LS SL SL LS Sand (•%-)• 8-3.2 71. 8 8-2-. 0 5 5.4 79.6 68.8 59. 8 78.0 Silt (%) 16.8 28.2 18.0 41.4 20.4 28.0 36. 4 21.6 Clay (%)• 0 0 0 3.2 0 3.2 3.8 0.4 Horizon Textural class S LS - LS LS - SL -Sand (%) 89. 4 73.4 - 74.4 84.6 - 54.8 • -Silt (%) 10. 6 26.6 - 25.6 15.4 - 42.4 -Clay (%) 0 0 - 0 0 - 2.8 -Horizon Textural class S S S LS S S SL S Sand (%) 94. 8 93.4 85.6 83.2 90.2 88.4 57.8 87.2 Silt (%) 5.2 6.6 14.4 16.8 9.8 9.8 33.8 12.4 Clay (%) 0 0 0 0 0 1.8 8.4 0.4 Table 20 Soil Chemical Analysis Juncus parryi Association 49 Plot No. 12 8 21 Ah Horizon pH 4.7 4.7 5.0 C (%) 8.4 15.1 7.9 OM {%) 14.4 26.0 13.6 N (?) 0.5 0.8 0.8 C/N 16. 18. 10. P (ppm) 6. 13. 18. Ca (me/1OOg) 0.29 1.03 0.62 Mg (me/1OOg) 0.05 0.20 0.07 Na (me/1OOg) 0.14 0.12 0.14 K (me/1OOg) 0.14 0.10 0.13 CEC (me/1OOg) 27.5 31.6 38.4 Horizon pH 4.9 5.2 -C {%) 4.5 8.1 -OM {%) 7.7 14.0 -N {%) 0.3 0.5 -C/N 16. 15. -P (ppm) 5. 6. -Ca (me/1OOg) 0.05 0.38 -Mg (me/1OOg) 0.04 0.04 -Na (me/1OOg) 0.14 0.12 -K (me/1OOg) 0.08 0.03 -CEC (me/1OOg) 8.9 12.0 -Horizon pH 5.1 5.1 5.2 C (%) 1.7 2.9 2.1 OM {%) 2.9 4.9 3.6 N il) 0.1 0.2 0.1 C/N 17. 15. 16. P (ppm) 3. 5. 6. Ca (me/1OOg) 0.32 0.08 0.91 Mg (me/1OOg) 0.01 0.04 0.01 Na (me/1OOg) 0.13 0.13 0.31 K (me/1 OOg) 0.09 0.03 0.01 CEC (me/1OOg) 4.6 6.3 7.3 40 30 59 68 63 4.8 4.8 4.6 4.6 4.7 11.4 6.1 10.5 15.0 7.2 19.7 10.5 18.1 25.9 12.3 0.7 0.1 0.6 0.7 0.3 17. 122. 18. 20. 21. 9. 10. 17. 14. 7. 0.11 0.25 0.95 0.12 0.37 0.07 0.03 0.25 0.05 0.05 0.07 0.12 0.13 0.14 0.14 0.09 0.01 0.37 0.10 0.17 26.4 34.1 28.4 41.2 26.9 5.0 4.9 5.2 7.7 5.7 - 4.0 -13.3 9.7 - 6.9 -0.5 0.3 - 0.2 -16. 20. - 19. -10. 15. - 6. -0.05 0.22 - 0.13 -0.03 0.02 - 0.01 _ 0.05 0.13 - 0.14 -0.01 0.00 - 0.04 _ 22.4 13.2 - 7.3 -5.1 5.1 4.7 5.1 5.0 4.1 3.3 5.1 2.2 2.2 7.1 5.6 8.7 3.8 3.8 0.3 0.2 0.3 0.1 0.2 15. 19. 16. 18. 14. 6. 5. 11. 5. 2. 0.23 0.26 0.29 0.27 0.44 0.02 0.01 0.05 0.01 0.03 0.06 0.13 0.10 0.14 0.11 0.00 0.00 0.13 0.03 0.18 24.9 20.0 17.0 8.9 7.0 Pig. 8. Juncus parryi Association, Plot 8. 51 values are strongly acidic. Organic matter and nitrogen dec rease steadily with depth. Carbon:nitrogen ratios are generally narrow. There is a very low amount of nitrogen In the Ah horizon of plot 30, thus making the carbon:nitrogen ratio very wide. Phosphorus, magnesium, potassium and cation exchange capacity all decrease in quantity with depth; calcium and sodium are variable, decreasing in some plots and Increasing in others. Carbon:nitrogen ratios, phosphorus and cation exchange capacity are generally similar among the various plots; the exchangeable cations vary widely. Antennaria lanata Association (Ref. Tables 21, 22, 23, 24, 25; Fig. 9, 10) Characteristic Combination of Species Antennaria lanata Salix cascadensis Gentiana glauca Polytrichum pillferum This association occurs at the base of slopes, on ridges and on slopes ln the alpine and low alpine areas. The relief shape is hummocky. Exposure is variable, and the slopes are very gentle, ranging from 0-9$. Most of the ground surface is covered by humus (64-90$), with very few rocks (0-12$). There is usually some mineral soil exposed (0-35$). Erosion varies from none to strong. The hygrotope is mesic. The herb layer is predominant, covering 70-85$ of the area. The D layer is fairly we 11-developed, although there are few species. Coverage is 40-60$. Antennaria lanata Is the dominant species, with an average 52 Table 21 General Environment Antennaria lanata Association Plot No. 3 4 16 32 Elevation (ft.) 7600 7575 7475 7450 Physiography Landform ridge base of slope • base of slope slope Relief shape Exposure neutral NE neutral S Slope gradient (%) 0 1 0 9 Layer Coverage (%) C layer 70 70 85 85 D layer 50 40 40 60 Plot Coverage ( %) Humus 64 73 90 88 Mineral soil 35 25 10 0 Rock 1 2 0 12 Soil Hygrotope Erosion strong moderate none none Horizon depth (in. ) L-H - - 2 1/2-0 -Ah 0-6 0-3 0-2 0-2 B Bm 6-10 Bfh 3-15 Bm 2-13 1/2 Bhf 2 1 Cgj 10-19 Cg 19 + C 15-21 Cg 21+ 13 1/2+ 8 1/2 Cl 8 1/2-16 1/2 C2 16 1/2+ Classification Gleyed Gleyed Alpine Sombric Dystric Humo-Ferric Brunisol Podzol Alpine Dystric Brunisol Mini Ferro-Humic Podzol Table 22 Antennaria lanata Associ ati on 53 Plot No. • 3 4 16 32 Plot Size (m2) 10 10 10 10 Extent of type (m ) 40 126 196 24 Elevation (ft.) 7600 7575 7475 7450 Altitudinal area A A LA LA Aver.Species C layer Presence Significance 1 Antennaria lanata 8.3 8.3. 7.3 7.2 V 8 2 Salix cascadensis 5.2 5.2 7.2 5.2 6 3 Carex pyrenai ca 4.2 3.2 5.2 3.2 V 4 4 Gentiana glauca 3.1 2.1 4.2 4.2 4 5 Phyllodoce empetriformis 1.2 4.2 1.2 4.2 V 3 6 Sibbaldia procumbens 2.1 3.2 3.2 3.2 3 7 Juncus parryi 1.1 2.2 2.2 2.2 V 2 8 Luzula spicata 2.1 1.1 2.1 1.1 1 9 Luzula wahlenbergii ' +.+ 1.1 1.2 2.2 V 1 10 Carex spectabilis 1.2 5.3 - 6.2 IV 5 11 Agrostis variabilis 3.1 - 4.2 2.2 IV 3 12 Arenaria obtusiloba 2.1 1.1 2.1 IV 1 13 Festuca brachyphylla - 3.1 4.2 111 2 14 Erigeron peregrinus - 1.1 - 3.2 III 1 15 Vacci ni urn scopari urn - - +.+ 3.2 III 1 16 Carex phaeocephala 2.2 - 1.2 111 + 17 Arenaria capillaris - 1.1 +.1 III + D layer Bryophytes 18 Polytrichum pi 1 iferum Dh 7.3 7.3 6.2 5.2 V 6 19 Lophozia alpestris Dh - +.+ +.+ 2.1 IV + 20 Barbi 1 ophozia hatcheri Dh - +.+ - 2.1 Ml + 21 Ceratodon purpureus Dh - +.+ - 2.1 III + 22 Pohlia nutans Dh - 1.1 1.1 III + Lichens 23 Lecidea granulosa Dh 3.1 3.1 5.2 5.2 V 4 24 Cladonia carneola Dh 2.1 3.1 1.2 4.2 3 25 Sol ori na crocea Dh 2.1 2.1 3.2 3.2 V 3 26 Cetraria islandica Dh 5.2 2.1 2.1 IV 3 27 Cladonia ecmocyna Dh 2.1 - 1.2 4.2 IV 2 28 Cetraria ericetorum Dh - - 4.2 3.2 III 2 Total Species (incl.sporadics) 21 26 31 31 Sporadic Species 38 Bryum sp. 16(1.1) C layer 39 Cephaloziella subdentata 16(+.+) 29 Deschampsia atropurpurea 32(1.1) 40 Dicranum scoparium 4(1.1) 30 Hieraci urn gracile 32(2.2) 41 Kiaeria blyttii 32(4.2) 31 Juncus drummondii 16(2.2) 42 Orthocaulis floerkii 16(1.1) 32 Juniperus communis 32(+.+) 43 Paraleucobryum enerve 32(3.2) 33 Luzula arcuata 3(+.1) 44 Polytrichadelphus lyallii 3(1.+) 34 Picea engelmannii 32(+.+) 45 Polytrichum formosum 16(3.1) 35 Poa cusickii 4(3.1) 46 Polytrichum ? juniperinum 32(5.2) 36 Vaccinium caespitosum 32(6.2) 47 Polytrichum norvegicum 4(3.1) D layer Lichens Bryophytes 16(1.1) 48 Pel ti ge ra canina 32(1.1) 37 Bryum capillare 49 Stereocaulon alpinum 16(1.2) 54 species significance of 8. The other species of high cover in the C layer is Salix cascadensis, with an average species sig nificance of 6. Other constant species are Carex pyrenaica, Gentiana glauca, Phyllodoce empetriformis, Sibbaldia procumbens, Juncus parryi, Luzula spicata and Luzula wahlenbergii. Gentiana  glauca is considered as a characteristic species because of its exclusiveness for this association. In the D layer, Polytrichum  pillferum is the constant bryophyte, with an average species significance of 6. Constant lichens include Lecldea granulosa, Cladonla cameola and Solorlna crocea, all with low cover values. Table 23 shows the floristic similarity indices for the four plots making up the association. Plots 3 and 4, In partic ular, have a very high similarity. Table 23 Floristic Similarity Indices for the Antennaria lanata Associat ion 3 4 16 32 3 80 63 49 4 59 57 16 4 22 The soils of this community are classed as Alpine Dystric Brunisols, Gleyed Sombric Humo-Ferric Podzol;,and Mini Ferro-Humic Podzol. Table 24 Soil Texture Antennaria lanata Association Plot No. Ah Horizon Textural class Sand (%)• Silt (%) Clay (%) B Horizon Textural class Sand (%) Silt (%) Clay (%) C Horizon Textural class Sand (%) Silt (%) Clay (%) 3 4 16 SL SiL SL 5-5.6 47. 2 60 .4 H4.4 50.0 37.2 0 2.8 2.4 SL SiL SL 55.2 46.4 57.2 43.2 51.0 40.4 1.6 2.6 2.4 LS 8 3.3 16.5 0.2 S 94.7 5.3 0 . S 94.0 5.6 0.4 Plot No. L-H Horizon pH C (?) OM (?) N (?) C/N P (ppm) Ca (me/lOOg) Mg (me/lOOg) Na (me/lOOg) K (me/lOOg) CEC (me/lOOg) Ah Horizon pH C (?) OM (?) N (?) C/N P (ppm) Ca (me/lOOg) Mg (me/lOOg) Na (me/lOOg) K (me/lOOg) CEC (me/lOOg) B Horizon pH C (?) OM (?) N (?) C/N P (ppm) Ca (me/lOOg) Mg (me/lOOg) Na (me/lOOg) K (me/lOOg) CEC (me/lOOg) C Horizon pH C (?) OM (?) N (?) C/N P (ppm) Ca (me/lOOg) Mg (me/lOOg) Na (me/lOOg) K (me/lOOg) CEC (me/lOOg) Table 25 Soil Chemical Analysis Antennaria lanata Association 3 4 16 32 .4.2 I9;3 33.3 -1.5 13. 6. : 1.66 0.31 0.34 0.78 29.9 4.9 5.0 4.5 4.5 13.0 10.9 13.0 14.0 22.3 18.7 22.3 24.1 0.8 0.7 0.1 0.9 16. 16. 118. 15. 15. 20. 9. 180.14 0.16 0.19 0.76 0.08 0.06 0.08 0.24 .0.16 . 0.13 0.17 0.13 0.16 0.12 0.17 0.30 32.5 ' 71.3 18.3 78.7 5.8 5.4 5.0 5.1 2.8 5.2 6.6 7.6 4.8 8.9 11.4 13.1 0.2 0.4 0.5 0.5 13. 13. 14. 17. 8. 7. 3. 11. 0.13 0.19 0.10 0.03 0.02 0.02 0.02 0.02 0.16 0.18 0.15 0.11 0.04 0.04 0.06 0.00 21.8 22.8 9.9 31.5 5.9 5.5 5.1 5.2 0.8 0.9 1.3 1.9 1.4 1.6 2.2 3.2 0.1 0.1 0.1 0.1 16. 18. 16. 19. 7. 4. 3. 9. 0.43 0.33 0.03 0.22 0.02 0.01 0.01 0.01 0.12 0.10 0.13 0.10 0.02 0.02 0.07 0.00 21.5 6.8 4.6 15.2 57 Fig. 9. Antennaria lanata Association, Plot 3. Fig. 10. Soil profile of Antennaria lanata Association, Plot 3. This soil is classified as a Gleyed Alpine Dystric Brunisol with Ah, Bm, Cgj and Cg horizons. Soil texture is coarser in the G horizon than in the A or B horizons. The A horizon samples range from silt loam to loamy sand. The B horizon is predominantly loamy sand. The C horizon soils are classed as sands and loamy sands. The soil chemical data are given in Table 25. The pH values increase with depth and are all acidic. Organic matter and nitrogen decrease with depth. Carbon:nitrogen ratios are narrow. The percentage of nitrogen in the Ah horizon of plot 16 is very low; thus, the carbon:nitrogen ratio is extremely wide. Phosphorus, cation exchange capacity and exchangeable cations decrease in amount with depth, with the exception of calcium, which increases in half the plots. Carbon:nitrogen ratios, phosphorus, sodium and magnesium (the latter In the B and C horizons) are similar among the four plots, while cation exchange capacity, calcium and potassium are variable. Phyllodoce empetrlformis - Antennaria lanata Association (Ref. Tables 26, 27, 28, 29, 30; Fig. 11, 12, 13, 14) Characteristic Combination of Species Phyllodoce empetrlformis Antennaria lanata Vaccinium scoparium Polytrichum piliferum Dicranum scoparium Lecidea granulosa This association occurs mainly on slopes, in the alpine, low alpine and subalpine parkland areas. Relief shape varies from hummocky to straight. Exposure is variable, and slope gradients range from 5 to 28$. Humus covers most of the ground surface, from 58-98$. There is very little mineral soil Table 26 General Envlronaent Phyllodoce ontpotrlf orals - Antennaria lanata Association Phyllodoce - Antennaria Variation Plot Ko. 5 6 19 27 31 37 55 5 7 6 66 72 Elevation (ft.) 7575 7500 7500 7460 7450 745 0 7425 7400 7375 7300 7300 Physiography Landfon —^————— slope of'ridge 5'0t" sl opa ridge Relief shape humnocky hunoocky straight hunaocky ^^nyex ^uanocky concave_ straight hunaocky Exposure E HE SE S S £ SI NI NI NI 1 Slope gradient (J) 15 B 28 12 15 11 . 27 5 6 14 . 9 Layer Coverage (1) C layer 80 80 90 80 85 60 75 . 85 5 95 . D layer 40 45 50 50 65 50 45 60 • 60 75 50 Plot Coverage (I) KUQUS 98 94 80 70 90 95 95 95 95 95 98 Klneral soil 2 3 5 20 0 0 0 0 0 0 0 Rock 0 3 15 10 10 5 5 5 '. 5 5 2 Soil . ' • Hygrotop* • . • ... • • Basic . Erosion • —• •• •• •• . • •••• none . Horizon depth (in.) L-H 2-0 - ty.0 - - - -Ah . 0-7 0-9 0-2 H-Ah 0-4 0-7 0-3 0-12 B Bf 7-14 6i 9-21 B> 2-14 Bg 4-12 Bfh 7-17 Bf 3-11 -C Cg 14* 21+ Cg 14+ 12+ 17+ 11-16 Cg 16+ 12-18 0-3J- 0-4 0-8 En 4.10 Bn 8-13 *• 10+ 13+ 0-3j-10+ Cl.s.m»t1.n Gleyed A|p)ne Gleyed ^ ^ Gleyed ^ M( RuIc-ferrlc °'5,,r,c1 DystHc J"''", "7-f«"-'< HumolVerrle D0rtMcI Alpine Dystric Brunisol Ferro-Hu.lc Podzol Bn"""' Brunisol 6™""' Podzol R"os'', PM Antennaria - Vacclntui Variati on 20 7 23 41 29 7580 . 7550 7550 7475 7400 " slope • straight straight .... . . straight . . • straight straight , to convex to concave .  to convex 1 S . E . SI ' SE 19 23 25 13 15 90 •' 65 • • 75 60 90 50 ' 50 • 50 50 . 60 90 83 5B 92 . 94 2 " 2 2 0 0 8 .15 '•' 40 8 6 meslc • I... submeslc • Male ' nnn* slight - 4-0 -w 0-3 0-3 4-12 Bn 3-9. Bi 3-11 12+ 9+ 11+ 0-3T 0-6 ..... Cg12j-« Gleyed ,, , ,. Alpine .Alpine Dystric Brunisol, * p , Oystrlc Dystric ' , . n . t Brunisol Brunisol _Phy11cwioco t--cetrlfar;Ii - A-ttmnrla ..mat a i-.-.octaHon Antannirla l.ii Variation Plot «». Plot Sill lo') s 6 19. 27 11 37 55 57 76 ec 72 20 7 2] (1 29 24 24 2( 21 24 21 2( 24 2( 2( 2( 2a 2a 29 2a • 28 E.tint ol lipa iV) 56 UO 75 189 60 320 32 375 • 150 ICS 155 66 % (6 75 50 El..,lion III.) 757b 75 00 75 OO 7(50 7150 7(50 7(25 7(00 .7375 7J00 7300 7510 75S0 .7550 7(75 7(00 Iltltudlnal ar.a > U I 4 U LA SP SP SP SP SP » < 4 Ll u J>ver.Sp«ctn Slo.nl 1 Ph,tloJoc. ("Ict.'l ferali !.l B.l !.3 8.3 a. 3 1.3 8.2 9.3 !.! i.j 9.3 6.2 5.2 5.2 2.2 2.2 5 V 7 2 Antannarla 1 .fiat i 7.3 6.2 7.2 7.2 7.2 6.2 7.2 8.2 6.2 !.2 6.? 7 8.2 1.2 7.2 8.2 7.2 a • V 7 ] Vacclnlm scopjrlm 6.3 2.1 7.2. 6.2 6.2 5.2 7.2 6.2 7.2 6.2 5.2 7.2 7.2 7.2 8.2 7.2 7 V - 6 4 Crlgeron pcrgjrln'js 5.2 (.2 4.2 (.2 (.2 (.2 6.2 (.2 2.1 6.2 (.2 3.2 3.2 • 1.2 (.2 4.2 4 V 4 5 Juncus pirry 1 2.1 2.1 3.? 3.2 2.1 3.Z 3.2 . (.2 . 5.2 3.2 4.2 2.2 6.2 5 . V ' 4 6 Slb.'aldla procu.-ians I.I 3.1 1.2 2.2 2.1 2.2 3.1 3.1 1.1 2.1 3.2 1.1 1.2 1.1 2.2 2 V 2 7 C.rai srectal'11] ri - 3.2 (.2 3.2 (.2 2.2 5.2 . . . 1.1 5.2 (.2 . 7.2 6.2 . 5 IV 1 S mer.:lun orjclle 2.1 3.1 3.2 3.2 (.2 . 3.2 1.1 3.1 2.1 3.2 3.1 3.1 2 IV 2 9 Carai nigricans 3.1 - 2.1 3.2 2.1 3.2 1.1 2.1 (.2 3.2 3.2 • . IV 2 10 irr-.arla papillaris 1.1 - 3.2 . 2.2 2.2 1.2 - l.t 3.2 3.1 3.2 (.2 2.2 3 IV 2 11 Dtscl-a-psla atrgpurpurea 1.1 2.1 - 3.2 2.1 3.2 . 3.1 . 1.1 . 1.2 1.1 1.1 • IV • 1 12 Irr.lca latlfolla - - - 1.2 - 1.2 6.2 3.2 2.1 3.2 5.2 . III 3 13 Luljla fihlent.rsl 1 (.1 (.2 - 5.2 3.2 _ 3.2 1.2 3.2 3.2 2 III 2 14 Lulula glabraba - - 5.2 3.2 (.2 - (.2 . II 2 15 lsl.1l IP. 1.1 - II 1 16 Cla/tcnta lancealata - 1.1 - - - - 4.1 1.1 3.1 3.1 3.1 1 17 Poa cuslcUl 1.1 - - - 4.2 - - - 3.2 II ' 18 Stti.fr.el.ii dema (9 lupin.,,, I.Mfallu-. 20 Carn p,resileJ 21 Vicclrlu-i caeipltoiun 22 Casttlleja ilreri 23 U-iul» parvlftora 24 Valeria.-) tltchensls ' 25 Ciru phiescaghil a 26 ft.luca brachyprflli 27 luiula splcata . 20 Sallx canadensis 29 Irlietui jplcatun Alienation Awtr.Spflcfoi Presence *'.•«'.';•« Us Sign. Meance- Sit;.-, fierce 4.2 1.1 3.2 2,7 1.2 2.1 1.1 •' 6r,o?k.ftm 30 Pol.trlcfc'ji pltlfem 31 Okr-ici jeaearl un 12 U^ciU alee. trU 33 PBlytrlchidelphut lyillll 34 Polftrlchua forsoiua 35 Pol|trkhj^ nar.e;lcun 36 Dts-atodan lattfallus 37 Ciratotjon purpureus 38 Klaerla Myttll 39 Polytrich.ua ]unlperlnus 40 PoMla nut Jin 41 BarbllcB^oMa lycopodloliiti •42 Barbllophiuta hatcherl " Dl> 6.3 (.2 (.2 5.2 5.2 5.2 7.2 7.2 4.2 5.2 Oh (.2 3.3 1.1 3.1. 3.2 1.2 3.1 5.2 3.1 3.2 6.2 Oti 2.1 3.2 1.1 3.1 - 3.1 6.2 5.1 (.1 _ Oh - - - _ - 2.2 3.1 5.2 Oh - - 1.1 - 3.2 (.2 - -Oh 3.1 (.1 3.2 3.1 Oh - 2.1 1.1 -Oh - . 3.1 - 3.1 S.2 -Oh - 3.1 _ 5.2 - 2.1 Oh - 1.1 1.1 Oh - - 1.1 - • - - - - - • 6.2 7.3 3.2 2.1 2.1. 3.1 6.2 3.2 3.1 1] Llctdaa granulosa Oh 7.2 6.2 6.2 6.2 6.2- 7.2 3.1 7.2 7.2 5.1 3.1 E 6.2 4.2 6.2 7.2 6.2 6 V 6 44 Cladonia fC.nocyna Oh 3.2 2.1 2.2 3.2 4.2 3.2 3.1 3.2 3.2 - 3 3.2 3.1 - 2.1 3.2 2 V ] 45 Cladonia earreola Oh • _ . 3.2 3.2 3.2 _ 2.1 2.1 3.1 1 3.2 . 2.1 2.1 •3.2 2 IV 2 46 C.trarli arte.torui Oh 2.1 2.2 3.2 - - 3.1 - - 1 ' _ 2.1 2.1 2.2 2.2 1 III 1 47 Sglorlna cfOMa Oh . - - ' t.l 2.2 . - - - • • 1.1. 1.1 . 2.1 2.1 1 . III . 48 Cttrarla subalplna Oh . 3.2 - 2.2 3.1 - - - t.2 - 1 ; 11 1 49 Cttrarla lllandlcl- : Dh - . - I.I - 2.2 3.2 • 3.2 - - . II SO Cl .Ionia cMoropha.a Dh - - - - 3.1 3.1 1 51 Peltigera canina Oh - - - •.* - - - - - - • - 1.1 3.2 1 1 52 Peltigera oalacea Oh - - - - - - - - - - 1.. - 2.1 • - * 1 * fetal Sp.cltl (Incl.sporadic:) 19 21 27 26 24 2S 2( 20 21 21 24 31 23 20 20 19 Ssoradlc Species 57 Castllleja rhealloiW T21S.2I 64 Potefitllle dHarslfolla 7(1.1) 67 OrlhocauJIl Moor.ll 31(3.1) C laur 53 tplloblun alplnun 27(1.1) 65 Veronica aar.nskjoldll SSI2.1) 6! Rnaco-.l trlun carescens 7(1.1) 59 Juncus drunondll 72(2.2) 63 lortula norvtglca 13(1.1) S3 Igrostls variabilis 20(1.2) 60 Luruta glabrata-tihlcnbergll 31(4.2) 0 la.er Lichens 54' artraria obtuslloba 7(..l) 61 r-dlcularls bructeosa 72(2.2) . Bryopnytos 55 Irnlca sollls 72(1.1) 62 Fhlaun alplnun 20I..I) 70 Ic'idophl la trlcetoru.i 27(1.2) 56 Carta p,renjlca-nlgrlcans 27(2.2) 63 Plcia tngel.'anall 57(»..) 66- Bryun sp. 19(2.1) - 71 Paltlqara canina var. rufasct ! 1.(2.1) exposed (0-20$). There is usually some rock coverage (0-40$). There is no evidence of erosion. The hygrotope is placed as submesic to mesic. Two vegetation layers are present - the herb layer and the bryophyte-lichen layer. The C layer has a high coverage, from 75 to 95$. The D layer is fairly well developed, covering 40-75$ of the area. The dominant plants in the C layer are Phyllodoce empetri formis , Antennaria lanata and Vaccinium scoparium. Other con stant species of lower coverage are Erigeron peregrinus, Juncus  parryi and Sibbaldia procumbens. In the D layer, Polytrichum  plliferum and Dicranum scoparium are the constant bryophytes, while Lophozla alpestris, with a presence of IV, is also impor tant. Among the lichens, Lecidea granulosa and Cladonia ecmo-cyna are constant. The association is divided into two variations: a« Phyllodoce empetriformis - Antennaria lanata Variation b. Antennaria lanata - Vaccinium scoparium Variation Table 28 gives the floristic similarity indices for the sixteen plots comprising the association. The two variations are shown separately. The values are all very high. The values are higher within each variation than between them. 62 Table 28 Floristic Similarity Indices for the Phyllodoce empetriformis -Antennaria lanata Association 5 6 19 27 31 37 55 57 76 66 72 20 7 23 41 22 5 6 19 27 31 37 55 57 76 66 72 71 79 82 83 78 68 76 75 74 52 73 75 72 76 52 59 68 60 50 83 79 71 66 63 74 63 52 83 79 65 71 74 72 55 79 66 71 68 67 50 54 67 76 68 53 60 59 61 52 68 72 56 54 46 20 7 23 41 29 65 58 67 63 61 55 42 47 41 44 67 62 70 57 64 65 57 63 55 61 71 63 69 60 68 59 47 55 52 54 59 60 59 50 54 61 60 59 59 51 59 49 57 53 52 56 50 50 54 45 24 28 30 24 2S 76 72 72 76 78 69 66 66 76 74 The variations are described below, by general habitat, floristics and soil data. a. Phyllodoce empetriformis - Antennaria lanata Variation This is the type variation for the association. It occurs on slopes in the alpine, low alpine and subalpine parkland areas. Relief shape is predominantly hummocky, but can also be straight or concave. Exposure is variable, and slope gradients range from 5 to 28$. The ground surface is covered by 70-98$ humus, 0-20$ mineral soil and 0-15$ rock. No erosion was observed. The hygrotope is rated as mesic. The herb layer is very well developed, covering 75-95$ of the area. The bryophyte and lichen layer has a coverage of 40-75$. 63 In addition to the dominant species listed for the associa tion, the following species are important in the differentiation of this variation: Carex nigricans, Arnica latifolia and Clay-tonia lanceolata in the C layer (these are lacking in the Anten naria - Vaccinium Variation); Dicranum scoparium, Lophozia  alpestris (both with a higher cover) and Cetraria subalplna (which has a high preference for this community) in the D layer. The predominant soils associated with this variation are Alpine Dystric Brunisols (6). Other soils are Sombric Humo-Ferric Podzols (3)* Sombric Ferro-Humic Podzol (l) and Lithic Orthic Regosol (l). Soil texture (Table 29) becomes coarser with depth. Samples of the A and B horizons are classed as loamy sands or sandy loams. The C horizon ranges from sandy loam to sand. Table 30 presents the soil chemical data. The values for pH are strongly acidic, and increase slightly with depth. Organic matter and nitrogen decrease steadily with depth. Carbon: nitrogen ratios are generally narrow. The percentage of nitrog en in the B horizon of plot 19 is very low. Phosphorus, cation exchange capacity, magnesium and potassium decrease in amounts with depth; calcium and sodium increase in some cases and dec rease in. others, b. Antennaria lanata - Vaccinium scoparium Variation This variation also occurs on slopes, but only in the alpine and low alpine areas. Relief shape is mainly straight, in contrast to the hummocky terrain of the Phyllodoce -Antennaria Variation. Exposure varies, but is never: northerly. Tab!* 29 Soil Texture Phyllodoca enpotrlforuls - Antennaria lanata Association Phyllodoce - Antennaria Variation Antennaria - Vacclnlm Variation 'lot No. 5 6 19 27 31 37 55 57 76 66 72 20 7 23 41 29 ,h Horizon Textural class IS LS LS LS SL SL SL SL SL SL SL SL SL LS LS S Sand (J) 73.6 75.6 75.2 72.4 64.0 56.4 66.8 73.0 66.6 56.8 72.2 66.8 56.4 80.8 78.0 S6.0 Slit (I) 26.4 24.4 24.8 27.6 36.0 43.6 32.4 23.2 33.4 39.4 22.6 32.6 43.6 19.2 22.0 14.0 Clay (I) 0 ' 0 0 0 0 0 0.8 3.8 • 0 1.8 5.2 0.6 0 0 0 0 l Horizon Textural class LS LS SL SL LS LS - LS SL LS SL S LS S LS LS Sand (J) 79.6 74.4 66.2 56.4 75.6 80.8 - 75.0 50.8 84.8 58.4 85.2 81.0 88.4 70.0 77.6 Silt (1) 20.4 25.6 . 33.8 41.8 24.4 19.2 - 22.4 45.2 10.4 36.4 14.8 19.0 11.6 30.0 22.4 Clay (J) 0 . 0 0 1.8 0 0 - 2.6 4.0 4.8 5.2 0 0 0 0 0 Horizon Textural class S '•• LS •. S . LS S LS . LS SL 5 LS SL S S S LS S Sand (i) 86.4 82.4 91.2 86.4 • 66.6 76.5 78.0 69.8 67.8 87.4 66.6 92.6 93.8 88.4 78.4 65.6 Silt (I) 13.6 17.6 8.8 11.B 13.4 23.6 18.4 24.0 9.2. 5.8 26.6 7.0 6.2 11.6 21.2 14.4 Clay (I) 0 0 0 1.6 0 0.5 3.6 .6.2 3.0 6.B 6.8 0.2 0 0 0.4 0 1 t I 1 1 .1 > 1 1 1 1 33S2--355S2 -3253553 3 4 t • 1 • 1 I 1 1 • • 332S--33353 2-22--53 55-. 23S5--55532 • 1 t 1 1 1 1 1 " 1 1 « 3333risJ3§333 ---3--23353 -3-5--3I332 2S32--35533 2-S2--2535-i i i i i i < i «. i i 3 32 3-.•35335 3532--33553 5.0 3.1 5.3 0.1 21. t. 0.33 0.01 0.13 0.05 18.3 3223^35535 -335^-33533 S332--2553-i i i i i t i i i * • 332----S5S33 -235e-,-25353 - 3233^-53335 - 3-33--53S35 33S5--S5333 t i t i • « t • • • i 3 333,,-333 5 3 S-22--53353 -232--55533 • «•••««*•«) •3333^35353 • • '• • - 3333--3353g • • i • t i i • • i i 3^3=--55S23 - 3 3 2--3153;- 2-22--35533 i « j III « '» t i > 3335=-c-33553 2323--5355S -3-5--55533 * • t i • i i i < i i .3 333 3--35 53 s. -332--I5333 -335--55512 3555,5-33333 3-S3--33552 2-2S--55553 SS33--5553S • 1 1 • 1 1 > 1 1 1 1 3323--53S33 -333--33532 -3-5--53-3 3 1 I t 1 1 1 t • 1. 1 t 3333e--53s53 -325---SSS3 ggggg sss 112133 ggggg 1 sss 333233 ggggg ! ===.323333 ggggg i sss 333233 66 Slope gradients are greater than in the type variation, ranging from 13 to 25$. The ground is covered by 58-94$ humus, 0-2$ mineral soil and 6-40$ rock. There is no erosion. The hygro-tope ranges from submesic to mesic. The herb layer is still of high cover, being 75-90$ of the area. There is also a we 11-developed D layer, covering 50-60$. There is a shift in the relative dominance of the major species, with Phyllodoce empetriformis being reduced, and Antennaria lanata assuming the primary role, along with Vaccin ium scoparium. Juncus parryi is also more important in this community. Carex spectabilis and Arenaria capillarls have higher values of average species significance in this variation than in the type variation. Although this community has been placed in the Phyllodoce empetriformis - Antennaria lanata Association, it is probably best regarded as a transition bet ween this association and the Antennaria lanata Association. The soils are all classed as Alpine Dystric Brunisols. Soil texture appears to be coarser than in the type varia tion. The A horizon samples are sandy loams, loamy sands or sand. The B horizon consists of loamy sands or sands, while the C horizon is predominantly sand. The soil chemical data all appear to be as described for the type variation, except that calcium is present in smaller quantities in the B and C horizons of this variation. 67 Fig. 11. Phyllodoce - Antennaria Association, Phyllodoce -Antennaria Variation, Plot 31. Fig. 12. Soil profile of Phyllodoce - Antennaria Association, Phyllodoce - Antennaria Variation, Plot 57. This soil is classified as an Alpine Dystric Brunisol, with an Ah-Bm-C horizon sequence. 68 Fig. 13. Phyllodoce - Antennaria Association. Antennaria -Vaccinium Variation, Plot 7. Fig. 14. Soil profile of Phyllodoce - Antennaria Association, Antennaria - Vaccinium Variation, Plot 7. This soil is classified as an Alpine Dystric Brunisol with L-H, Ah, Bm and C horizons. 69 Picea engelmannii Association (Ref. Tables 31, 32, 33, 3*0 This association is represented by only one plot in the study area. It occurs on a ridge in the alpine region. The relief shape is straight. Exposure is southeast, with a slope gradient of 29$. Humus and rock each cover 50$ of the ground surface. There is no mineral soil exposed. There is no evid ence of erosion. The hygrotope is rated as submesic. There are four vegetation layers in the community. The shrub layer covers 100$, the herb and bryophyte-lichen layers each 15$, and the epiphyte layer 5$. The plot is composed of one Picea engelmannii individual in the B layer, which has a species significance value of 9. There are few species in the other layers. The most important species in the C layer are Carex phaeocephala and Antennaria  lanata. Barbilophozla hatcheri and Polytrichum piliferum are the most important bryophytes, while Cetraria ericetorum and Solorina crocea are the dominant lichens. There are just two epiphytes: Parmeliopsis ambigua-Vand Parmeliopsis hyperopta. The soil is classified as a Lithic Orthic Regosol, with an Ah-C-R horizon sequence. Texturally, the A horizon is a sandy loam, while the C horizon is a loamy sand. The pH increases from the A to the C horizon, but Is strongly acidic in both cases. Organic matter and nitrogen decrease in quantity with depth. The carbon:nitrogen ratios are narrow. Phosphorus, cation exchange capacity and the available cations decrease from the A to the C horizon, with the exception of sodium, Table 31 General Environment Picea engelmannii Association Plot No. 49 Elevation (ft.) 7600 Physiography Landform ridge Relief shape straight Exposure SE Slope gradient (%) 2 9 Layer Coverage (%) B layer 100 C layer 15 D layerE layer 5 Plot Coverage (%) Humus 50 Mineral soil 0 Rock 50 Decaying wood 0 Soil Hygrotope submesic Erosion none Horizon depth (in.) . Ah 0-3 C 3-12 R 12 + Classification Lithic Orthic Regosol 71 Table 32 Picea engelmannii Association Plot No. 2 49 Plot size (m ) 2 & Extent of type (m ) 6 Elevation (ft.) 7600 Altitudinal area A B layer 1. Picea engelmannii 9.7 C layer 2. Carex phaeocephala 4.2 3. Antennaria lanata 3.4. Festuca brachyphylla 2.5. Vaccinium caespitosum 2 iff •** 6. Vaccinium scoparium 2.7. Agrostis variabilis 1.1 8. Antennaria umbrinella9. Arenaria capillaris 2 10. Arenaria obtusiloba 1.11. Luzula spicata 1.1 12. Selaginella densa13. Sibbaldia procumbens 1.14. Trisetum spicatum 1 D layer Bryophytes 15. Barbilophozia hatcheri Dh 4.2 16. Polytrichum piliferum h 4.2 17. Bryum capillare Dh 1.1 18. Pohlia nutans h 1.1 Lichens 19. Cetraria ericetorum Dh 3.2 20. Solorina crocea h 3.2 21.. Lecidea granulosa h 1.1 22. Cladonia carneola Dh +.+ 23. Peltigera canina h +.+ E layer 24. Parmeliopsis ambigua ER 2.1 25. Parmeliopsis hyperopta R 2.1 Table 3 3 Soil Texture Picea engelmannii Association Plot No, 49 Horizon Textural class Sand (%) Silt (%) Clay (%) Table 34 Soil Chemical Analysis Picea engelmannii Association Plot No. 49 Horizon Ah C p.H 4.8 5.3 C (%) 10.2 2.6 OM (%) 17.6 4.4 N . (%) 0.7 0.2 C/N 14. 13. P (ppm) 13. 4Ca (me/lOOg) 1.90 0.60 Mg (me/lOOg) 0.26 0.03 Na (me/lOOg) 0.02 0.03 K (me/lOOg) 0.42 0.08 CEC (me/lOOg) 58.0 14.5 Ah SL 59.2 38.0 2.8 C LS. 83.4 16.6 0 73 which increases slightly. Abies lasiocarpa Association (Ref. Tables 35, 36, 37, 38) Characteristic Combination of Species Abies lasiocarpa Parmeliopsis hyperopta This association, represented by only two plots, occurs on ridges in the alpine area. Relief shape Is either straight or convex. The exposure is southwest, with a slope gradient of 27-30$. Humus covers 55-65$ of the ground surface, rock 35-40$ and mineral soil 0-5$. Erosion is slight or none. The hygro tope varies from submesic to mesic. The shrub layer occupies 95$ of the area. There is a very sparse herb layer, coverage being 5-7$. The bryophyte and lich en layer is somewhat better developed, covering 5-30$. The epiphyte layer is very poorly developed in one plot (only 3$ cover), but well developed In the other plot (30$ cover). The only species ln the B layer is Abies lasiocarpa, with an average species significance of 9. There are only two con stant species in the C layer, both with very low average species significance values: Carex spectabilis and Sibbaldia procumbens. No bryophyte species are constant. Cetraria ericetorum and Lecidea granulosa are the constant lichens, both with very low coverage. Among the epiphytes, Parmeliopsis hyperopta, with an average species significance of 5, and Parmeliopsis ambigua, with a value of 3, are constant. The floristic similarity index for the two plots of the Table 35 General Environment Abies lasiocarpa Association Plot Mo. Elevation (ft.) Physiography Landform Relief shape Exposure Slope gradient (%) Layer Coverage (%) B layer C layer D layer E layer Plot Coverage (%) Humus Mineral soil Rock Decaying wood Soil Hygrotope Erosion Horizon depth (in.) H Ah B 53 7550 44 7535 C R Classification straight SW 27 95 5 5 30 55 5 40 0 submesic slight 0-6 6-13 13 + Lithic Orthic Regosol -ridge-convex SW 30 95 7 30 3 65 0 35 0 mesic none 2-0 0-2 Bh 2-4 Bhf 4-12 BC 12-17 17 + Lithic Mini Ferro-Humic Podzol Table 36 75 Abies lasiocarpa Association Plot No. 53 Plot Size (m2) 30 Extent of type (m2) 96 Elevation (ft.) 7550 Altitudinal area A B layer 1 Abies lasiocarpa 9.7 C layer 2 Carex spectabilis 1.2 3 Sibbaldia procumbens +.+ 4 Vaccinium scoparium 5 Antennaria lanata 6 Carex phaeocephala 7 Vaccinium caespitosum 8 Agrostis variabilis 9 Arenaria capillaris 10 Erigeron peregrinus 11 Festuca brachyphylla 12 Luzula spicata D layer Bryophytes 13 Tortula ruralis Dh -14 Barbilophozia lycopodioides Dh -15 Bryum capillare Dh -16 Barbilophozia hatcheri Dh -17 Polytrichum piliferum Dh -18 Bryum bimum Dh 2.2 19 Dicranum scoparium Dh 2.2 20 Orthocaulis floerkii h 1.1 21 Brachythecium starkei Dh -22 Lescuraea baileyi Dh -23 Lescuraea radicosa h -24 Paraleucobryum enerve Dh -Lichens 25 Cetraria ericetorum Dh 2.1 26 Lecidea granulosa h 2.1 27 Cladonia macrophyl 1 odes Dh -E layer 28 Parmeliopsis hyperopta Eg 6.1 29 Parmeliopsis ambigua g 3.1 30 Cetraria pinastri Eg -44 25 25 7535 A Aver.Species Presence Significance 9.7 2/2 9 2.2 2/2 1 1.1 2/2 + 3.2 1/2 2 2.2 1/2 1 1.2 1/2 + 1.2 1/2 + +.+ 1/2 + +.1 1/2 + +..+ 1/2 + +.+ 1/2 + +.+ 1/2 + 5.2 1/2 4 4.2 , 1/2 3 4.2 1/2 3 3.1 1/2 2 3.2 1/2 2 1/2 1 1/2 1 1/2 + +.2 1/2 + +.2 1/2 + +.1 1/2 + +.1 1/2 + +.2 2/2 1 +.2 2/2 1 1.1 1/2 + 2.1 2/2 5 2.1 2/2 3 +.1 1/2 + Total Species 10 27 76 Table 3 7 Soil Texture Abies lasiocarpa Association Plot No. 5 3 44 Ah Horizon Textural class LS LS Sand (%) 80.8 73.8 Silt (%) 18.0 26.2 Clay (%) 1.2 0 B Horizon Textural class - LS Sand (%) - 72.7 Silt (%)  27.3 Clay (%) - 0 C Horizon Textural class LS LS Sand (%) 77.2 78.0 Silt (%) 22.0 22.0 Clay (%) 0.8 0 Table 38 Soil Chemical Analysis Abies lasiocarpa Associ ati on Plot No. 53 44 H Horizon pH - 4.6 C (?)  27.8 OM (?)  47.N (?) - 1.7 C/N  16. P (ppm) ' - 25Ca (me/1 OOg) - 7.40_ Mg (me/1OOg)  1.04 • Na (me/1OOg)  0.02 K (me/1OOg) - 0.76 CEC (me/1OOg)  163.0 Ah Horizon pH 4.7 4.2 C (?) 9.8 17.5 OM (?) 16.8 30.1 N (?) 0.4 1.C/N 23. 16. P (ppm) 15.Ca (me/lOOg) 0.22 1.63 Mg (me/1OOg) 0.07 0.37 Na (me/1OOg) 0.12 0.03 K (me/lOOg) 3 0.46 CEC (me/1OOg) 108.0 61.5 B Horizon pH - 4.4 C (?)  10.7 OM (?)  18.4 N (?) - 0.6 C/N  18. P (ppm)  10Ca (me/1OOg) - 0.16 Mg (me/1OOg)  0.07 Na((me/I00g)  0.04 K (me/lOOg) - 0.10 CEC (me/1OOg)  50.9 C Horizon pH 4.6 4.8 C (?) 12.1 6.1 OM (?) 20.9 10.5 N (?) 0.8 0.3 C/N 15. 18. P (ppm) 13. 11Ca (me/1OOg) 0.08 0.19 Mg (me/100g) 6 0.03 Na (me/1OOg) 0.12 0.04 K (me/1OOg) 1 0.03 CEC (me/IOOg) 103.0 33.1 78 association is 67, which is relatively high. The soils are classed as Lithic Orthic Regosol and Lithic Mini Ferro-Humic Podzol. Texturally, all the samples from the A, B and C horizons are loamy sands. Among the chemical data, pH generally increases slightly with depth, all values being strongly acidic. In one plot, organic matter and nitrogen dec rease in amount with depth; in the other, they increase. The increase in organic matter is probably due to its downward move ment and accumulation in the very shallow soil. There should be an ample supply of nitrogen available for higher plants, as indicated by the narrow carbon:nitrogen ratio. Phosphorus, cation exchange capacity, calcium, magnesium and potassium dec rease in quantity with depth; sodium increases slightly. In all the chemical data, there is a wide variability between the two plots. Abies lasiocarpa - Picea engelmannii - Vaccinium scoparium  Association (Ref. Tables 39, 40, 41, 42, 43; Fig. 15, 16) Characteristic Combination of Species Abies lasiocarpa Picea engelmannii Vaccinium scoparium Dicranum scoparium Parmellopsis hyperopta Cetraria pinastri This association occurs mainly on ridges in the alpine, low alpine and subalpine parkland areas. Relief shape varies from straight to convex to concave. Exposure is usually southwest, with slope gradients of 2-25$. Humus covers 40-70$ of the Table 39 General Environment Abies lasiocarpa - Picea engelmannii - Vaccinium scoparium Association Plot No. Elevation (ft.) Physiography Landform Relief shape 51 7540 ledge convex' to concave Exposure SW Slope gradient (%) 2 5 54 7500 61 7350 -ridge-SW 15 SW 2 70 7300 straight straight concave NW 24 Layer Coverage (%) B layer C layer D layer E layer 95 7 10 5 85 60 60 30 95 45 50 10 95 60 25 20 Plot Coverage (%) Humus Mineral soil Rock Decaying wood 70 0 30 0 65 10 25 0 40 0 40 20 60 0 30 10 Soil Hygrotope mesic Erosion none Horizon depth (in.) Ah 0-4 B Bh 4-12 C R 12-19 19 + mesic slight submesic none 0-4 1/2 0-3 Bhf 4 1/2- Bhf3-11 14 1/2 14 1/2+ Cgll + mesic none 0-3 Bhf 3-12 12 + Classification Lithic Orthic Humic Podzol Sombric Ferro-Humic Podzol Gleyed Sombric Ferro-Humic Podzol Sombric Ferro-Humic Podzol Table 40 Abies laslocarm 5cea ennelnannll - Vacclnlui scoparlun Association Plot No. .. 5' . ' 54 • 61 70 Plot Size (V) •27 .; 30 . 30 105 Extent of type (u^) 27 ' 30 30 .105 Elevation (ft.) 75i0 . .75 00 7350 7300 Altitudinal area A : LA SP SP Presence Aver.Specks ' B layer Significance 1 Abies lasiocarpa 7.7 7.6 8.6 8.8 V 8 2 •Picea engelnannil C layer 5.7 • . 7-6 '. 7.6 ' 6.6 V. .7 • 3 Vacciniun scopariun •4.2 • 6.2 7.2 5.2 V .6 4 Antennaria lanata 1.2 4.2 3.2 2.2 V 3 5 Phyllodocs-ecpctri forais - 2.2 4.2 7.3 IV . 5 6 Arenaria capiMaris . 2.2 3.1 +.+ IV 1 7 Luzula sp. - . - 2.1 6.3 III 5" 8 Carex phaeocephala 3.2 3.2 111 1 9 Festuca brachyphylla 2.2 3.2 - , - m: 1 "• • 10 Sibbaldia prpcunbens 1.1 ' 3.1 - HI 1 11 Potenti1 la diversifolia 1.1 2.1 . - - in + : 12 Hieraciira gracile - 1.1 1.1 - in 13 Erigeron peregrinus, 0 layer Bryophytes 1.1 in + 14 Dicranuij scopariun Qh 1.2 2.1 5.1 5.2 V 4 15 Polytrichua piliferun C- 2.2 4.2 3.1 - IV 3 16 Lophozia alpestris Or. 1.1 4.1 3.1 - IV 3 17 Pohlia nutans Gh 1.1 - 7.2 - III 5 18 Orthocaulis floerkii Lichens Dh 4.2 2.1 ~ — III 2 19 Cladonia ecnocyna :-. 2.2 2.1 3.1 3.1 V 3 . 20 Lecidea granulosa 6.2 3.1 - IV 4 21 Cetraria ericetorun L- 3.2 1.1 . 1.1 - IV 1 •-22 Sblorina crocea E layer :-. i.i 2.1 +.• IV 1 23 Parceliopsis hyperopta E; 3.2 6.1 4.1 4.1 V 5 24 Cetraria pinastri £:-3.2 4.1 - 5:1 IV 4 25 Parmeliopsis anbigua E: 3.2 - 4.1 4-1 IV 3 Total Species (incl.sporadic :) 31 32 22 14 Sporadic Species C layer 26 Agrostis variabilis 27 Arenaria obtusiloba 28 Carex spectabilis 29 Oeschanpsia atropurpurea 30 Haplopappus lyallii 31 Juncus parryi 32 Luzula spicata 33 Salix cascadensis. 34 Trisetup spicatun D layer 8ryophytes 35 Barbilophozia barbata 36 Barbilophozia hatched 37 Barbilophozia lycopodioide 51(2.2) 54(1.2) 54(1.2) 70(2.1) 51(+.2) 54(2.2) 54(1.1) 54(2.2) 51(1.2) 54(1.1) 5K+.1) 70(4.1) 38 Brachytheciun starke1 61(3.1) 39 Drepanocladus uncinatus 51(1.1) 40 Kiaeria blyttii 70(5.1) 41 Lescuraea bailey! 61(1.1) 42 Lophozia ? kunzeana 61(4.1) 43 Pohlia elongata 54(3.1) 44 Polytrichadelphus lyallii 61(3.1) Lichens 45 Cladonia carneola 54(3.1) 46 Cladonia chlorophaea 54(2.1) 47 Cladonia coccifera 51(1.1) 48 Cladonia pleurota 51(1.1) 49 Peltigera oalacea 54(1.1) 50 Stereocaulon al pi nun 510.1) E laver 51 Alectoria anerlcana 51(1.1) 81 ground surface and rock 25-40$. Mineral soil is exposed In only one plot. Decaying wood occurs In two plots (10-20$). There is generally no observable erosion. The hygrotope ranges from submesic to mesic, most plots being mesic. Since this community is a tree island, the B layer is pre dominant, coverage being 85-95$ of the area. The C layer is mainly well developed, covering 7-60$. The D layer is also well developed, with a cover of 10-60$. The E layer covers 5-30$. The B layer Is composed of two species, which are both dominant in the community - Abies lasiocarpa, with an average species significance of 8, and Picea engelmannii, with a value of 7. Vaccinium scoparium dominates the C layer, with an aver age species significance of 6. The only other constant species in the herb layer is Antennaria lanata. Phyllodoce empetrlformis, with a presence of IV and average species significance of 5, is important in most plots. Dicranum scoparium is the dominant bryophyte, while Cladonia ecmocyna is the dominant lichen. Parmeliopsis hyperopta is the only constant epiphyte. Cetraria  plnastri, though not constant, is considered a characteristic species, because it reaches its highest cover value in this association. Table 41 gives the floristic similarity indices for the four plots of the associon. The highest values are obtained between the two alpine plots (51, 5^) and between the two subalpine plots (6l, 70). 82 Table 41 Floristic Similarity Indices for the Abies lasiocarpa - Picea  engelmannll - Vaccinium scoparium Association 51 54 61 70 51 60 51 44 54 56 43 61 8 1° Most of the soils are Sombric Ferro-Humic Podzols. One soil is classified as a Lithic Orthic Humic Podzol. The subalpine soils are finer-textured than the alpine soils. All the samples from the A, B and C horizons of the subalpine soils are sandy loams. The alpine A horizons are loamy sands; the B horizon Is a sandy loam or loamy sand; the C horizon is a loamy sand or sand. Table 43 presents the soil chemical data. The pH values are all strongly acidic, and increase slightly with depth. Organic matter and nitrogen decrease steadily with depth. The carbon:nitrogen ratios are relatively narrow. Phosphorus, cation exchange capacity, magnesium and potassium decrease in quantity with depth; sodium increases, while calcium decreases from the A to the B horizon, then increases from the B to the C horizon. Organic matter and nitrogen vary widely among the four plots. Phosphorus and cation exchange capacity are similar in the A and C horizons. Calcium and sodium are similar in the B horizon Table 42 Soil Texture Abies lasiocarpa - Picea engelmannii - Vaccinium scoparium Association Plot No. 51 54 61 70 Ah Horizon Textural class LS LS SL SL Sand (%) 79.6 73.6 71.6 69.6 Silt (%) 20.4 25. 0 25.0 2 8.4 Clay (%) 0 1.4 3.4 2.0 B Horizon Textural class SL LS SL SL Sand (%) 58.6 79 .6 56.6 71.4 Silt (%) 38. 8 19. 0 41. 0 24. 2 Clay (%) 2.6 1.4 2.4 4.4 C Horizon Textural class LS S SL SL Sand (%) 7 4-.6 89. 0 58.0 70.4 Silt (%) 24.4 11.0 32.0 20.0 Clay (%) 1.0 0 10.0 9.6 Abies lasiocarpa Table Soil Chemical - Picea engelmannii 43 Analysis - Vaccinium scoparium Associ ati on Plot No. 51 54 61 70 Ah Horizon pH C (?) OM (?) N (?) C/N P (ppm) Ca (me/lOOg) Mg (me/1OOg) Na (me/1OOg) K (me/1OOg) CEC (me/1OOg) 4.2 13.2 22.6 0.7 20.. 10. 0..40 0..17 0.04 0.28 62.8 4.1 6.8 11.6 0.5 14. 7. 1.99 0.31 0.11 0.26 24.6 4.4 12.8 22.0 0.8 16. 23. 0.15 0.16 0.29 0.20 29.9 4.1 9.7 16.7 0.5 18. 9. 0.34 0.21 0.13 0.33 22.0 Horizon PH C (?) OM (?) N (?) C/N P .. (ppm) Ca (me/1OOg) Mg (me/1OOg) Na (me/1OOg) K (me/1OOg) CEC (me/1OOg) 4.6 7.7 13.3 0.4 20. 6. 0.13 0.04 0.04 0.01 48.4 4.8 8.2-14.1 0.4 20. 3. 0.05 0.03 0.12 0.04 17.1 4.8 6.2 10.7 0.4 16. 3. , 0.09 0.03 0.13 0.04 7.5 4.6 6.2 10.6 0.3 23. 3. 0.07 0.03 0.13 0.05 6.1 C Horizon pH C (?) OM (?) N (?) C/N P (ppm) Ca (me/1OOg) Mg (me/1OOg) Na (me/1OOg) K (me/1OOg) CEC (me/1OOg) 4.7 3.8 6.6 0.3 14. 6. 0.25 0.03 0.10 0.02 37.3 5.0 2.6 4.5 0.1 26. 2. 0.45 0.02 0.15 0.02 11.6 4.8 3.2 5.5 0.2 18. 5. 0.42 0.01 0.27 0.03 12.8 4.7 3.8 6.5 0.1 27. 3. 0.27 0.02 0.14 0.04 11.6 85 Fig. 15. Abies - Picea - Vaccinium Association, Plot 51. Taller tree species is Picea engelmannii. In left foreground is Antennaria - Sibbaldia Association. Fig. 16. Soil profile of Abies - Picea - Vaccinium Association, Plot 54. This soil Is classified as a Sombric Ferro-Humic Podzol, with an Ah-Bhf-C horizon sequence. 86 only. Magnesium and potassium are fairly similar in all horizons. Abies lasiocarpa - Valeriana sitchensis Association (Ref. Tables 44, 45, 46, 47, 48) Characteristic Combination of Species Abies lasiocarpa Valeriana sitchensis Arnica latifolia Polytrichadelphus lyallii Parmeliopsis hyperopta This association occurs on seepage slopes in the subalpine parkland. Relief shape is concave to straight. The exposure is southwest, with a slope gradient ranging from 21 to 39$. The amount of rock covering the ground surface is variable, from 5 to 65$. Humus covers 35-75$, while there is no mineral soil exposed. Decaying wood is present in two plots, with a cover of 10-20$. No evidence of erosion was observed. The hygrotope is rated as subhygric to hygric. The shrub layer occupies 75-95$ of the area. The herb layer coverage varies from 20 to 80$. The bryophyte and lichen layer is sparsely developed, covering only 10$ of the plot. The epiphytic layer is better developed, coverage being 5-20$. Abies lasiocarpa is the only species in the B layer, with an average species significance of 9. Valeriana sitchensis and Arnica latifolia dominate the C layer, with average species significance values of 6 and 5, respectively. Other constant species, of lower coverage, are Vaccinium scoparium, Senecio  triangularis, Ve rat rum viride t Claytonla lanceolata, Castille ,1a Abies Table 44 General Environment 1asiocarpa - Valeriana sitchensis Association Plot .No. Elevation (ft.) Physiography Landform Relief shape Exposure Slope gradient (%) Layer Coverage (%) B layer C layer D layer E layer Plot Coverage (%) Humus Mineral soil Rock Decaying wood Soil Hygrotope Erosion Horizon depth (in.) Ah Bin C 56 62 74 7400 7300 7200 —seepage slope — concave concave straight SW SW SW 39 21 22 85 95 75 35 80 20 10 10 10 5 20 20 35 70 75 0 0 0 65 20 5 0 10 20 hygric subhygric subhygric .—none 0-6 0-3 0-7 1 6-12 3-13 7 1/2-15 12+ 13+ 15 1/2+ Classification •Alpine Dystric Brunisol Table 45 Abies lasiocarpa - Valeriana sitchensis Associ ati on 88 Plot No. 56 62 74 Plot Size (m2) 41 126 35 Extent of type (m2) 41 126 35 Elevation (ft.) 7400 7300 7200 Altitudinal area SP SP SP B layer Presence Aver.Species Significance 1 Abies lasiocarpa C layer 9.7 9.8 8.6 3/3 9 2 Valeriana sitchensis 6.3 8.3 4.2 3/3 6 3 Arnica latifolia 5.2 6.2 4.2 3/3 5 4 Vacci ni um scopari um 3.2 5.2 2.2 3/3 4 5 Senecio triangularis 4.2 2.2 3.2 3/3 3 6 Veratrum viride 2.2 4.3 3.2 3/3 3 7 Claytonia lanceolata 3.2 3.1 2.1 3/3 3 .8 Castilleja elmeri 3.2 3.1 1.2 3/3 3 9 Mi tell a breweri 3.2 3.2 1.2 3/3 3 10 Luzula sp. - 7.3 2.2 2/3 5 11 Carex spectabilis 4.2 5.2 - 2/3 4 12 Phyllodoce empetriformis - 4.2 3.2 2/3 3 13 Pedicularis bracteosa - 3.2 3.2 2/3 2 14 Erigeron peregrinus - 3.1 1.2 2/3 1 15 Lupinus 1 ati fol i us - 2.1 2.2 2/3 1 16 Arnica mollis D layer Bryophytes 2.2 1.1 2/3 1 17 Polytrichadelphus lyallii Dh 2.1 4.1 4.2 3/3 4 18 Lophozia alpestris E layer Dh 3.1 3.2 2/3 2 19 Parmeliopsis hyperopta EB 2.1 4.1 4.1 3/3 4 20 Parmeliopsis ambigua EB 3.1 - 4.1 2/3 3 Total Species (incl.sporadics) 26 26 25 Sporadic Species C layer 21 Anemone occidentalis 62(2.1) 22 Antennaria lanata 62(3.223 Arenaria capi 11 aris 62(2.1) 24 Deschampsia atropurpurea 74(1,1) 25 Juncus drummondii 56(1.126 Luzula glabrata 56(3.2) 27 Ranunculus eschscholtzii 62(2.1) 28 Saxifraga ferruginea 56(1.1) 29 Sibbaldia procumbens 56(1.1) 30 Trollius laxus 56(2.2) D layer Bryophytes 31 Barbilophozia hatcheri 62(+.+) 32 Barbilophozia lycopodioides 56(3.1) 33 Brachytheci um curtum 56(2.1) 34 Brachythecium starkei 62(2.2) 35 Bryum 1 pseudotriquetrum 56(1.1) 36 Cephaloziella sp. 74(1.1) 37 Ceratodon purpureus 74(1.1) 38 Dicranum scopari um 62(2.1) 39 Kiaeria blyttii 56(2.1) 40 Lescuraea incurvata 62(1.1) 41 Pohlia gracilis 74(1.1) 42 Pohlia nutans 74(1.1) 43 Pohlia wahlenbergii 56(4.1) 44 Polytrichum piliferum 74(2.2) 45 Rhacomitrium sudeticum 56(2.1) 46 Scapania undulata 56(+.+) E layer 47 Cetraria pinastri 62(4.1) 89 elmeri and Mitella brewer!. In the D layer, Polytr1chadelphus  lyallii Is the only constant bryophyte. Due to the wet condit ions, there are no ground lichens in this community. Among the epiphytes, Parmeliopsis hyperopta is a constant dominant. Table 46 presents the floristic similarity indices for the three plots comprising the association. The values are reason ably high. Table 46 Floristic Similarity Indices for the Abies lasiocarpa -Valeriana sitchensis Association 56 62 74 56 62 61 62 51 14 The soils are all classed as Alpine Dystric Brunisols, with an Ah-Bm-C horizon sequence. Soil texture generally becomes coarser with depth. The A horizon samples are classed as sandy loams or loamy sand. The B horizon is a loamy sand. The C horizon ranges from sandy loam to sand. The soil chemical data are shown in Table 48. The pH is strongly acidic in all horizons, and increases slightly with depth. Organic matter and nitrogen decrease with depth. In plot 56, there is still a considerable amount of organic matter Abies lasiocarpa -Table 4 7 Soil Texture Valeriana sitchensis Association Plot No. 56 62 74 Ah Horizon Textural class LS SL SL Sand (%) 76.0 63.0 68.0 Silt (%) 23.4 33.6 31.2 Clay (%) 0.6 3.4 0.8 Bm Horizon Textural class LS LS LS Sand *(%) 79.6 7 3.6 7 3.4 Silt (%) 18.6 24.0 25.2 Clay (%) 1.8 2.4 1.4 C Horizon Textural class S SL LS Sand (%) 87.0 64.0 77.2 Silt (%) 12.2 31.0 16.6 Clay (%) 0.8 5.0 6.2 Table Soil Chemical Abies lasiocarpa - Valeriana 48 Analysis sitchensis Association Plot 56 62 74 Ah Horizon pH C (?) OM (?) N (%) C/N P (ppm) Ca (me/lOOg) Mg (me/lOOg) Na (me/lOOg) K (me/1OOg) CEC (me/1OOg) 4.7 13.8 23.7 1.0 14. 6. 0.47 0.25 0.15 0.34 44.4 4.2 11.0 18.9 0.7 15. 8. 1.06 0.26 0.27 0.30 23.5 4.7 12.9 22.1 0.9 14. 12. 0.30 0.19 0.27 0.34 26.0 Horizon pH C (?) OM {%) N (I) C/N P (ppm) Ca (me/1OOg) Mg (me/1OOg) Na (me/1OOg) K (me/1OOg) CEC (me/1OOg) 4.7 7.4 12.8 0.5 14. 11. 0.39 0.13 0.14 0.18 36.8 4.7 6.5 11.3 0.4 17. 11. 0.38 0.07 0.12 0.13 13.8 4.8 7.0 12.1 0.5 15. 8. 0.20 0.06 0.15 0.08 24.3 C Horizon pH C {%) OM {%) N (?) C/N P (ppm) Ca (me/1OOg) Mg (me/1OOg) Na (me/1OOg) K (me/1OOg) CEC (me/1OOg) 4.8 6.7 11.5 0.4 16. 8. 0.69 0.13 0.26 0.16 19.3 4.9 2.7 4.6 0.2 16. 4. 0.42 0.02 0.15 0.04 8.3 5.1 1.7 3.0 0.1 14. 7. 0.36 0.02 0.15 0.04 12.0 92 in the C horizon due to the shallowness of the soil. Carbon: nitrogen ratios are narrow. Cation exchange capacity, magnesium and potassium decrease in quantity with depth. Calcium and sodium decrease from the A to the B horizon, then increase somewhat from the B to the C horizon. Phosphorus decreases down the profile in one plot, but in the others it increases from the A to the B horizon, and then decreases from the B to the C horizon. Organic matter, nitrogen and phosphorus appear to be fairly similar among the three plots, whereas cation exchange capacity is variable. The values for calcium are similar only In the B horizon. Magnesium is similar in the A horizon; in the B and C horizons, plots 62 and 70 are close in value. Sodium has similar values in the B horizon; plots 62 and 70 are similar in the A and C horizons. The three plots are similar in potass ium in the A horizon; only plots 62 and 70 are similar in the C horizon. Carex spectabilis Association (Ref. Tables 49, 50, 51, 52, 53) Characteristic Combination of Species Carex spectabilis Antennaria lanata Polytrichum piliferum This association occurs on slopes with some seepage, mainly in the alpine and low alpine areas. The relief shape is predom inantly straight. Exposure is south, west or southwest, with slope gradients ranging from 9 to 50$. Humus covers most of the ground surface, from 70 to 95$. Exposed mineral soil is mostly Table 49 General Environment Carex spectabilis Associ ati on 93 Plot No. Elevation (ft.) Physiography Landform Exposure 11 7575 depression channel Relief shape straight SW Slope gradient (?) 9 14 7500 straight to concave S 50 26 7500 straight 11 slope 39 7475 straight to convex 26 7475 straight S 28 Layer Coverage (?) C layer 65 D layer 20 96 35 85 5 90 40 Plot Coverage (?) Humus 70 Mineral Soil 25 Rock 5 2 10 95 0 5 75 0 25 90 5 5 hygric Soil Hygrotope Erosion Horizon depth (in.) L-H 1-0 Ah B C R 0-3 3-7 7+ subhygric 0-12 Bhf 12-18 18+ subhydric — none — 3-0 0-8 8+ subhydric 0-6 6-13 13+ submesic H-Ahi 0-tj Ah2 1y-12 Bm 12-18 18+ Classification Alpine Dystric Brunisol Sombric Ferro-Humic Podzol Orthic Regosol Lithic Orthic Regosol Alpine Dystric Brunisol Table 50 Carox spectabilis Association Plot No. 11 14 26 39 18 Plot Size (V) 30 27 20 24 30 Extent of type (m^) 110 30 35 36 76 Elevation (ft.) 7575 7500 7500 7475 7475 Altitudinal area A LA A - LA' SP C layer Presence Aver.Spec les Significance 1 Carex spectabilis 8.4 . 9.5 •• 8.4 9.4' 8.4 V '8 . 2 Antennaria lanata 4.2 5.2 5.2 2.2 5.2 V . 5 . 3 Erigeron. peregrinus 2.2 5.2 3.2 , 4.2 3.2 V • 4 4 Sibbaldia procumbens 1.1 1.1 4.2 .1.1 ' 4.2 V 3 5 Vaccinium scopari um 1.2 4.2 3.2 - 4.2 IV 3 6 Arenaria capillaris 3.1 2.2 3.2 3.2 IV 2 7 Juncus parryi • -• 4.2 5.2 III 4 • .8 Phi eun alpinum 4.2 - . 4.2 2.2 '• - III 3 9 Carex pyrenaica 4.2 4.2 - - - 1.2 III . 3 10 Veronica wormskjoldii 1.1 - 3.2 2.2 111 1.' .11 Hieraciun gracile - . 1.1 1.2 . 3.2 III 1 12 'Luzula spicata 1.1 - - +.+ 2.1 III + 13 Phyllodoce e^petriforals 1.2 - 1.2 1.2 - 111 + 14 Juncus drummondi i 2.2 5.2 - - II 3 15 Arnica noil is - . - 5.3 1.2 . - 1! 3 16 Carex nigricans 4.2 - 3.2 - . - II 2 17 Potentilla diversifolia - - 4.2 - 1.1 ' II 1 18 Claytonia lanceolata +.+ 4.2 - - - II 1 19 Agrostis variabilis • - - - 1.2 3.2 II .1 20 Antennaria fries!ana 3.2 - - - 1.1 ll • 1 21 Trisetura spicatun - - 1.1 3.2 - II 1 22 Poa cuslcki i 1.1 - 1.1 - • - II + 0 layer Bryophytes 23 Polytrichum piliferum Oh 4.2 3.2 1.1 3.2 3.2 V 3 24 Polytrichum junlperinum Dh - - 3.2 2.2 5.2 III 3 25 Polytrichum formosum Oh 1.1 1.1 3.2 - ' - III 1 26 Oicranun scoparium Oh 2.1 - 2.1 - - II + 27 Oesnatodon 1 ati fol i us Oh - 1.1 - 2ll - II + 28 Drepanocladus uncinatus Oh 2.1 - 1.1 - - II + 29 Pohlia gracilis Oh - 1.1 2.1 - ' - II + Lichens 30 Cladonia carneola Oh 3.1 - 2.1 - 4.2 III 2 31 Cladonia ecnocyna Dh - - 2.1 - 4.2 II 1 32 Lecldea granulosa Dh 1.1 - - - 3.2 II 1 33 Cetraria ericetorum Dh 1.1 - - - 2.2 II + 34 Stereocaulon alpinum Dh 2.1 - - '.' - • 1.1 II + Total Species (incl.sporad •cs) 28 33 21 28 Sporadic Species C layer 0 layer 35 Arenaria obtusiloba 18C+.1) Bryophytes 36 Arnica latifolia 39(1.2) 51 Barbilophozla hatcheri 26(2.1) 37 Carex phaeocephala 26(1.1) 52 Cephaloziella subdentata 14(+.+) 38 Oeschanpsia atropurpurea 11(1.1) 53 Ceratodon purpureus 26(2.1) 39 Festuca brachyphylla 18(4.2) 54 Klaeria blyttil 11(4.2) 40 Juncus mertensianus 26(1.2) 55 Lophozia alpestris 26(3.1) 41 Juniperus communis 18(1.+) 56 Orthocaulis floerkil 18(1.1) 42 Lupinus 1 a t i foli us 39(2.2) 57 Pohlia nutans 18(1.1) 43 Luzula glabrata 11(3.2) Lichens 44 Luzula nahlenbergii 11(3.2) 45 Luzula sp. 26(3.2) 58 Cetraria subalplna 26(1.2) 46 Salix cascadensis .11(6.2) 59 Cladonia chlorophaea 26(2.1) 47 Seneclo triangularis 26(1.2) 60 Cladonia deformls 26(1.1) 48 Silene parryi 39(3.2) 61 Cladonia sp. . 14(+.+) 49 Sol 1 dago cul11radi 18(4.2) 62 Peltigera canlna . 18(2.1) 50 Vaccinium caespitosum 39(3.2) 63 Solori na crocea 18(1.1) 95 very sparse. Rock covers 5-25$ of the area. There is no evi dence of erosion. . The hygrotope ranges from submesic to sub-hydric. The herb layer is predominant in this community, coverage being 65-96$. The bryophyte and lichen layer is moderately well developed, covering 5-40$. Carex spectabilis, the dominant species, has an average species significance of 8. In the C layer, Antennaria lanata is the subdominant, with an average species significance of 5. Other constants are Erigeron peregrinus and Sibbaldia procumbens. The only important bryophyte is Polytrichum piliferum. None of the lichens is of importance in this wet habitat. The floristic similarity indices for the five plots of the association are shown in Table 51. The values are lower than was the case with many of the previously-described communities. However, these plots all have their highest values of similarity with each other rather than with any other association. Table 51 Floristic Similarity Indices for the Carex spectabilis Associat ion 11 14 26 39 18 11 54 57 49 50 14 52 75 60 26 51 55 39 54 18 Table 5 2 Soil Texture Carex spectabilis Association Plot No. Ah Horizon Textural class Sand (%) Silt (%) Clay (%) B Horizon Textural class Sand (%) Silt (%) Clay (%) C Horizon Textural class Sand (%) Silt (%) Clay (%) 11 14 SL SL 66.8 50.2 33.2 49.4 0 0.4 26 39 SL SL 58.0 . 65.0 39.8 32.8 2.2 2.2 18 H-Ahl Ah2 SL LS 54.8 78.4 45.2 21.0 0 0.6 LS 82.2 17. 8 0 SL 70. 8 27.8 1.4 S 90.4 9.6 0 S 92.4 7.6 0 S 99.0 1.0 0 S 9 0.2 9 . 8 0 S 85.4 14.6 0 LS 76.0 23.6 0.4 Table' 53 Soil Chemical Analysis Carex spectabilis Association lot No. 11 14 26 39 18 -H Horizon H-Ahi pH - _ 4.8 _ 4.5 C (?) - - 30.1 15.1 OM (?) _ _ 51.8 25.9 N (?) _ _ 0.2 a* 1.0 C/N _ _ 167. _ 15. P (ppm) - - 16. 17. Ca (me/1OOg) - _ 4.50 0.10 Mg (me/1OOg) - 1.20 0.21 Na (me/1OOg) - _ 0.25 - 0.32 K (me/1OOg) - _ 1.06 0.44 CEC (me/1OOg) - - 163.0 - 96.5 ) Horizon pH 4.8 4.6 4.4 4.6- 4.9 C (?) 16.1 16.3 12.1 12.1 5.6 OM (?) 27.7 28.1 20.8 20.7 9.7 N (?) 0.1 0.8 0.6 0.5 0.3 C/N 134. 20. 21. 25. 18. P (ppm) 7. 20. 15. 13. 12. Ca (me/1OOg) 0.09 1.28 0.53 1.72 0.15 Mg (me/1OOg) 0.06 0.23 0.16 0.33 0.02 Na (me/1OOg) 0.13 0.27 0.21 0.03 0.14 K (me/1OOg) 0.02 0.54 0.10 0.44 0.04 CEC (me/lOOg) 51.6 62.8 36.6 61.4 48.6 Horizon pH 5.1 4.7 - - 5.0 C (?) 7.5 7.6 - - 3.2 OM (?) 12.9 13.1 - - 5.5 N (?) 0.4 0.5 - - 0.1 C/N 18. 17. - - 23. P (ppm) 4. 21. - - 6. Ca (me/1OOg) 0.29 0.04 - - 0.22 Mg (me/1OOg) 0.02 0.08 - - 0.01 Na (me/1OOg) 0.15 0.31 - - 0.17 K (me/1OOg) 0.03 0.25 - 0.02 CEC (me/1OOg) 14.8 43.9 - - 13.6 Horizon pH 5.0 5.1 4.7 4.8 5.1 C (?) 2.3 1.6 5.6 7.4 2.1 OM (?) 4.0 2.7 9.6 12.8 3.7 N (?) 0.2 0.1 0.4 0.5 0.1 C/N 15. 26. 16. 14. 16. P (ppm) 4. 5. 6. 18. 4. Ca (me/lOOg) 2.26 0.44 0.38 0.69 0.43 Mg (me/1OOg) 0.02 0.01 0.06 0.13 0.01 Na (me/1OOg) 0.49 0.17 0.12 0.04 0.16 K (me/1OOg) 0.02 0.03 0.04 0.19 0.01 CEC (me/1OOg) 6.8 7.0 18.3 79.3 19.5 98 The soils associated with this community are Alpine Dystric Brunisols, Orthic Regosols and a Sombric Ferro-Humic Podzol. The soils are finer-textured in the surface Ah horizon than in the B or C horizons. Surface horizons are classed mainly as sandy loams. The B horizon, where present, varies from sandy loam to sand. The C horizons are mainly coarse-textured sands. Table 53 presents the soil chemical data. The p"H increases slightly with depth, all values being strongly acidic. Organic matter and nitrogen decrease with depth, and carbon:nitrogen ratios are generally narrow. The values for nitrogen in the L-H horizon of plot 26 and in the Ah horizon of plot 11 are very low. Cation exchange capacity decreases with depth. It is very high in the L-H horizon of plot 26; this is due to the high organic matter content. Phosphorus, magnesium and potass ium decrease in quantity with depth. Calcium and sodium are both variable, in some cases increasing with depth and in others decreasing. Most of the chemical data are variable among the plots of the association. Phosphorus is similar in the L-H, Ah and C horizons. Valeriana sitchensis - Castille,1a elmeri Association (Ref. Tables 54, 55, 56, 57, 58; Fig. 17, 18, 19) 99 Characteristic Combination of Species Valeriana sitchensis Castilleja elmeri Carex spectabilis Arnica mollis Erigeron peregrinus Senecio triangularis Ranunculus eschscholtzii Aulacomnium palustre Philonotis americana Brachythecium asperrimum This association occurs on seepage slopes in the subalpine parkland and, less frequently, in the alpine and low alpine areas. The relief shape is mainly concave or straight. Expos ure is variable, with slope gradients ranging from 10 to 28$. The ground surface is predominantly covered by humus (92-100$), with practically no mineral soil exposed. The sites are rarely rocky (rockiness 0-10$). There is no observable erosion. The hygrotope varies from subhygric to subhydric. Both the herb layer and the bryophyte layer are well devel oped. The herb layer coverage is 90-100$, while that of the bryophyte layer is 15-95$. This meadow community is very rich in species. The domin ant species in the C layer are Valeriana sitchensis, Castille,1a  elmeri, Carex spectabilis (all with an average species signific ance of 6), Arnica mollis, Erigeron peregrinus and Senecio triangularis (all with an average species significance of 5). Other constant species include Antennaria lanata, Vaccinium  scoparium, Carex nigricans, Juncus drummondii, Veronica wormsk.1 -oldii and Sibbaldia procumbens. Ranunculus eschscholtzii is considered a characteristic species because of its high Tibia 54 General Envlronaent Valeriana sltchensla - CastlTleJa alaorl Association Valeriana - CaatlHela Vari ati on Troll1u« 1a»ua Vari ati on Plot %. Elevation (ft.) Physiography Landfora Relief shape 38 7475 42 7475 34 7450 43 7375 .seepage slope. 60 7300 65 7300 75 7200 ' 25 7500 36 7400 24 7375 .64 7300 ' 69 7300 . seepage alope -concave straight straight convex to concave to straight straight concave straight to concave 73 7275 straight ' concave convex Exposure .SI ' S SI II SS SI SI 1 S S« SI NI I Slope gradient {%) 25 16 15 14 10 17 '9 26 18 28 17 22 13. Layer Coverage (X) ' • C layer 100 90 95 95 100 95 98 . 92 100 97 95 98 98 0 layer 40 15 45 60 65 85 60 . 60 90 85 95 • 60 95 Plot Coverage (J) Huaua 100 90 95 95 100 96 95 89 100 100 100 .: 100 98 Mneral Soil 0 0 5 0 0 2 0 1 0 0 0 0 0 Rock 0 10 0 •'. 5 . 0 2 5 . 10 0 . : 0 0 0 2 Soil Hygrotope • hygrlc hygrlc subhydrlc subhydrlc hygrlc subhygric hygrlc hygrlr Eroslon nonn nnnn Horlion depth (tn.) L-H - - - - - - - 2-0 . H 8-0 - - - -Ah 0-12 0-8 0-12 0-6 0-10 0-8 0-6 - 0-4 0-12 .0-12 0-12 B - - Bhfg JJ-TJ- - - • - - Bg 4-8 - - - . -C Cg 12-15 Cg] 8-12 *• Cg 12.16 Cg 6+ 10+ 6+ Cg 6+ Cg 0-11 Cg 8+ Cg 12+ . Cg 12+'. -15+ 12+ 16+ _ - 11+ _ _ 12+ Classification Lithic Rego Kunlc Gleysol Gleyed Lithic Orthic Regosol Fera Hunlc Gleysol Lithic Rego Huralc Gleysol Rego Huntc . Gleysol Orthic Regosol' Orthic • Regosol Rego Lithic •Humlc Rego Gleysol Gleysol . Orthic Hunt C Gleysol Rego Huralc Gleysol Rego. • Lithic.-Hunlc Orthic Gleysol Regosol. Table 55 Val er: £-a sitchensis - Castllleja elrc-rl Association 101 Valerl ana sitchensis - CastiT!;.-: sl^ert Varlatlo Trol 11 us laj<us Variation Plot Do. 36 42 34 43 50 65 75 25 36 24 64' 69 73 PUl Size (o2) 15 15 15 15 15 "15 15 10 10 . 4 10 10 10 Extent of type (n2) 60 45 33 35 • C4 40 40 12 28 e 40 105 21 Elevation (ft.) 7475 7475 7450 7375 7:C0 7300 7200 75 00 7400 7375 7300 7300 7275 Altitudinal area LA LA LA SP S? SP S.P A SP SP . SP SP SP 1 Aver.Specles Aver.Specles Presence 5111. Significance Significance 1 Valeriana sitchensis 2.2 7.3 5.2 5.3 7.3 3.3 7.2 6. 3.2 4.2 6.2 6.2 6.2 6.2 5 V 2 Castllleja elrerl 5.2 5.2 7.2 6.2 -.- 6.2 7.2 6 4.2 2.2 5.2 6.2 4.2 4 V 3 Carex spectabilis • 9.4 8.3 7.3 6.2 3.2 3.1 1.2 6 5.2 5.2 2.2 4.1 2.2 *.+ 4 V 4 Arnica nollls 7.2 3.2 7.2 6.2 4.2 4.2 2.1 5 6.2 6.2 6.2 4.2 3.2 2.2 5 V 5 Erigeron peregrinus 4.2 6.2 3.2 5.2 C / 6.2 5.2 .. 5 4.2 1.2 3.2 5.2 6.2 5.2 5 V 6 Seneclo-triangularis 6.3 3.2 6.3. 7.3 5.2 5.3 5.2 5 +.+ 2.2 -2.2 5.2 - 4.2 - .3 V 7 Antennaria Janata 2.2 3.2 4.2 1.2 4.2 6.2 5:2 4 6.2. . 1.2 4.2 4.2 3.2 4 V 8 Vacclnlun scoparium 2.2 1.2 4.2 2.2 4.2 5.2 5.2 4 5.2 1.2 3.2 3.2 . 4.2 4 V 9 Carex nigricans 3.2 +.+ 4.2 2.2 2.1 - 2.1 2 2.2 6.2 4.2 4.1 3.2 3.2 4 V 10 Juncus drunnondi! 5.2 4.2 3.2 5.2 4.2 3.1 2.1 4 3.2 - _ 2.2 2.2 2.2 1 V 11 Veronica worcskjoldi1 2.2 2.1 3.2 2.2 2.1 1.1 3.1 2 3.1 2.1 1.1 1.1 1.1 1 V 12 Sibbaldia procur.tens 2.2 1.1 2.2 1.1 -3.' 2.1 1.1 2 2.1 1.1 2.2 - • . ••.1 1 V 13 Trolllus laxus - - 1.1 - 5:2 5.2 3 8.4 8.4 8.2 9.3 8.2 7.2 8 IV 14 CaltHa leptosepala - - ' - 2.2 1.2 - 2.2 1 . . 7.2 5.2 2.1 4.2 2.2 5 iV 15 Oeschar.psla atropurpurea 4.2 2.2 3.2 3.2 3.2 - - 3 2.1 • - 4.2 - 2.1 4.2 . IV 16 Phyllodoce e^pe'rlforais - 1.2 1.2 2.2 1.2 .4.2 2 5.2 - 2.2 3.2 2.2 3 IV 17 Ranunculus eschscholtzii 4.2 1.1 1.1 1.1 3.1 3.1 1.1 3 . - _ _ 2.1 2.1 IV 18 Phleun alplnun 3.2 1.1 3.2 3.2 - :- . . 1 2.1 1.2 1.1 _ . + IV 19 Arnica latifolla - 3.2 2.2 - 3.2 5.2 7.2 . 4 - - . 4.2 .4.2 2 III 20 Castllleja rhexifolia - 5.2 - 4.2 - 4.2 6.2 4 - ' 1.1 - 4.2 4.2 _ 2 III 21 Veratrun vlrltfa . - - - 5.3 3.2 4.2 3 +.+ +.2 _ 2.2 • + III 22 Juncus rertenslanus - 4.2 4.2 3.2 - 2 4.2 _ 1.1 _ _ 1 II 23 Ctaytonla lanceolata - - - - 2.1 4.1 1 . 4.1 3.1 3.1 2 II 2* Luzula sp. . - . - - 3.1 4.2 • 3.1 2 - 2.2 + II 25 Agrostls thurberiana - - - 3.2 - . - + - •3.1 - 4.2 1 II 26 Wltella breserl - 2.2 4.2 2.1 1.1 1 - - - +.+ - . + II 27 Poa cuslckll - - 4.2 3.2 1 1.1 - - - _ _ + II 28 Luzula glabrata 2.2 - 2.2 3.2 - - - . 1 - 1.2 - + - II 29 Potentllla druMondll - 1.2 - 1.2 2.2. - 1 - - - - - - II 30 Vacclnlufl caespltosurc 2.2 1.2 2.2 - - - - + - - - - - - - II 31 Anemone occidental Is - - 1.2 - - + - - - - - +.+ + II 32 Lupinus latifollus - - - - 4.2 3.2 1 - - - . - - - 1 33 Hieraclun graclle - - 1.1 - - - - > 3.1 - • - - 1 34 Trlsetun splcatun 2.1 - - - - - - + 1.1 - - - - - + 1 D layer Bryophytes .35 Aulaconnlun palustre Dh 2.1 2.1 6.2 4.r 7.2 8.2 5 _ 7.5 9.2 8.3 _ .9.3 7 IV 36 Philonotis anerlcana Dh - - 3.2 2.1 5.1 7.2 4 7.5 1.2 1.1 IV 37 Brachytheciun asperriraun Dh 3.2 - 2.1 - 5.1 3 2.1 2.1 _ 2 III 38 Scapanla subalplna' Oh 2.1 - 2.1 2.1 4.1 2 t.1 + IM 39 Polytrichadelphus lyallii Oh - - 5.2 1.1 4.1 5.2 4 6.2 3 II 40 Polytrlchun junlperlnun Dh - 3.2 - - - 4.1 1 7.2 _ 1.1 4 II 41 Polytrlchun forraosun Dh 6.2 3.1 1.1 - 3 _ 1.1 _ + II 42 Pohlia nutans Oh - - - 5.2 5.1 3 1.1 t.i II 43 Bryum pseudotrlquetrun Dh - - - - 5.1 - ' 2 1.1 + II 44 Brachythectun starkel Dh - - - - . - _ _ - 1.1 4.1 3.1 3.2 2 .11 45 Bryun nuehlenbeckll Dh - - - - . *.1 - + +.+ _ _ 5.2 .... _ 2 II 46 Olcranun scopariun Dh - - 3.2 - 3.1 3.1 1 2.1' _ + II 47 Lophozla alpestris Oh - - 1.1 - 2.1 3.1 .-- • 1 2.1 * II 48 Oescatodon latifollus Oh - - - 3.1 - _ _• _ 6.2 3 49 Ceratodon purpureus Oh - - 3.1 - t.1 m _ _ _ 1 50 Tortula rural Is 'Dh - - - - - - _ 1.1 _ 3.1 m 1 51 Barbilophozia hatcher] Oh - - - _ m + 1 52 Dicranella sp. Oh - - - - - +.t - - - - + 1 Lichens 53 Cladonia sp. Dh - 1.1 - - 1.1 - •_ _ + 1 Association Aver.Specles Significance 6 6 6 Total Species (Incl.sporadlcs) 22 25 29 3 ; 11 26 27 44 23 24 26 24 26 Sporadic Species 62 Picea engelcannll 69K-) 70 Drepanocladus unclnatus 25(1.1) C layer 63 Potentllla dlvBrslfolla 25(2.2) 71 Lophozla obtusa 36(2.1) 0 layer 72 Snlun blyttil 64(5.2) 54 Antennaria frieslana 25(1.1) 73 Pohlia lahlenbergll 25(3.1) 55 Arenaria capf11 arts 25(2.1) Bryophytes 74 Polytrlchun norveglcua 43(1.1) 56 Festuca saxlnontana 25(1.2) 54 Barbilophozia lycopodloldes 69(2.1) 75 Polytrlchun plllferun 42(3.2) 57 Gaultheria hunlfusa 24(4.2) 55 crachytheclun curtun 34(1.1) Lichens 58 Juncus parry] 25(2.2) 55 Bryun blflum 25(1.1) 59 Kalnia polifolla ' 36N-.1) 57 Bryul sp. 64(6.3) 76 Cladonia chlorophaea 43(1.1) 60 Luzula lahlenbergll 25(4.2) 69 Cephalozlella subdentata 25(+.+) 77 Leprarla neglecta 42(1.1) 61 Pedlcularls bracteosa 75(1.1) E3 Drepanocladus exannulatus 25(1.1) 78 Peltigera canina wr. rufescens 25(1.+) 102 preference for this association. The dominant bryophytes in the D layer are Aulacomnium palustre and Philonotis amerlcana, which both have a very high preference for this community. Brachythec-ium asperrimum is exclusive to the association, and thus is regarded as a characteristic species. Very few lichens are found, due to the wetness of the habitat. The association is divided into two variations: a. Valeriana sitchensis - Castille,1a elmeri Variation b. Trollius laxus Variation Table 56 gives the floristic similarity indices for the thirteen plots making up the association. The two variations are shown separately. The values within each variation are generally higher than between them. Floristic Similarity Indices for the Valeriana sitchensis -Table 56 Castille,1a elmeri Association 38 42 34 43 60 65 75 25 36 24 64 69 73 38 42 34 43 60 65 75 25 36 24 64 69 II 50 64 53 30 23 21 45 44 46 43 40 60 52 37 42 50 42 44 55 52 65 31 24 24 24 18 22 25 16 22 32 42 33 37 32 30 31 35 28 31 38 36 39 33 38 25 23 28 37 44 38 36 40 35 48 43 49 29 27 42 51 45 58 713 113 37 46 33 61 47 3^ 42 60 45 73 51 63 51 103 The variations are described below, by general habitat, floristics and soil data. a. Valeriana sitchensis - Castille,1a elmeri Variation This is the type variation for the association. It occurs on seepage slopes in the subalpine parkland and the low alpine area. Relief shape varies from concave to straight. Exposure is mainly southwest, and slope gradients range from 10-25$. Humus covers 95-100$ of the ground surface, mineral soil 0-5$, and rock 0-10$. There is no evidence of erosion. The hygrotope is rated as subhygric to subhydric. The herb layer has a very high coverage of 90-100$. The bryophyte layer is less well developed, but still very signific ant, with a coverage of 15-85$. In addition to the dominant species listed for the associa tion, the following species are important in the differentiation of this variation: Potentilla drummondii (which is exclusive to this variation) and Vaccinium caespitosum, which are both miss ing in the Trolllus laxus Variation; Mitella breweri, Ranunculus  eschscholtzii and Polytrichadelphus lyallli, which are all present in greater quantities in this variation. The soils associated with this variation are Rego Humlc Gleysols (3)» Orthic Regosols (3) and Fera Humic Gleysol (l). Texture generally is coarser at greater depths. The A horizon ranges from sandy loams to loamy sands. The C horizon is mostly sands or loamy sands. Table 58 presents the soil chemical data. The pH increases slightly with depth, all values being strongly acidic. Organic 104-Pig. 17. Valeriana - Castille,1a Association, Valeriana -Castille,1a Variation, Plot 38. Yellow flowers are Senecio triangularis and Arnica mollis, red flowers are Castille,1a elmeri, and light-coloured flowers are Erigeron peregrinus. Fig. 18. Soil profile of Valeriana - Castille,1a Association, Valeriana - Castille,1a Variation, Plot 38. This soil is classified as a Lithic Rego Humic Gleysol, with an Ah-Cg-R horizon sequence. Plot No. Ah Horizon Textural class Sand (?) Silt (?) Clay (?) B Horizon Textural class Sand' (?) Silt (?) Clay (?) C Horizon Textural class Sand (?) Silt (?) Clay (?) Table 57 Soil Texture Valeriana sitchensis - Castilleja elmeri Association Valeriana - Castilleja Variation Trollius laxus Variation 38 42 34 43 60 65 75 25 36 24 64 69 73 SL LS LS SL SL ' LS LS SL LS-S SL SL SL 61.4 77.6 76.0 50.2 61.2 77.2 72.0 62.4 - 85.0 60.2 68.6 69.6 38.4 22.0 22.8 48.0 : 36-0 21.4 28.0 36.8 - 15.0 38.0 30.6 29.0 0.2 0.4 1.2 1.8 2.8 1.4 0 0.8 0 1.8 0.8 1.4 LS 80.8 19.2 0 s S S LS LS , LS SL S S LS SL 92.8 93.2 98.0 72.0 82.2 85.8 69.2 90.4 86.8 80.4 ' 65.2 7.2 6.8 2.0 28.0 17.8 11.4 28.4 9.6 13.2 19.6 27.4 0 0 0 0 0 2.8 2.4 0 0 o 7.4 s 85.6 14.4 0 Plot Ko. 3B 141 Horl ion pH C (J) OS (I) N (J) C/N P (ppm) Ca (™/100j) Kg (ne/IOOg) Na (ne/IOOg) K (ne/lOOg) CEC (i>o/100g) Ah Horizon 4.8 C (J) 13.2 OH (J) 22.7 N (t) 0.9 C/N 15. P (ppi) 19. Ca WlOOg) 0.72 Hg (ne/100g) 0.30 Na (ne/IOOg) . 0.10 K (ms/IOOg) 0.57 CEC (ne/IOOg) . , . 59.7 8 Horizon pH C (I) OM (J) N (J) C/N P (pp>) Ca (ra/IOOg) Kg (m/IOOg) Na (m/100j) K (ne/IOOg) CEC (w/IOOg) . C Horizon A.8 C (t) 6.1 ON «) 10.5 N (J) 0.3 C/N . 21. P (ppn) • • 16. Ca (ne/IOOg) . 0.33 Kg (n/lOOg)' 0.09 Na (oe/lOOg) 0.06 K (oe/100g) 0.24 CEC («/100g) 33.8 Tabla 58 Soil Chanlcal Analysis Valeriana sitchensis - Cast!He la elmeri Association Valeriana . Castlllele Variation Trot 11 us laxus Variation 34 43 60 65 75 . 25 36 24 64 - -- --4.2 4.8 5.3 4.7 4.7 4.8 13.3 16.9 11.5 16.8 8.1 13.7 22.8 29.1 19.7 26.9 13.9 23.5 1.0 1.3 1.0 1.3 0.4 1.0 14. 13. 12. 13. 19. 13. 28. 17. 10. 20. 21. 5. 0.56 0.95 0.51 0.91 0.72 0.59 0.31 0.37 0.20 0.42 0.19 0.26 0.08 0.02 0.14 0.84 0.14 0.28 0.37 0.56 0.29 0.68 0.41 0.60 68.3 53.8 33.6 38.6 28.3 23.4 4.6 5.1 33.2 20.9 - -57.0 35.9 - - ' - • 1.9 . 1.6 - - -17. 13. 21. 15. 3.88 2.22 -1.20 0.80 ' 0.60 ' 0.60 - -2.84 0.22 - -145.0 29.4 - -4.5 4.7 4.8 5.0 4.9 11.2 - 18.5 11.8 8.8 12.3 19.2 • 31.9 20.3 15.2 21.2 0.7 - 1.0 0.1 0.7 0.7 16. - 18. 118. 14. 18. 17. - 6. 8. 8. 11. 0.57 - 1.01 0.73 0.73 1.00 0.19 0.40 0.34 0.18 0.28 0.23 - 0.34 0.48 0.27 0.28 0.21 0.71 0.72 0.30 0.66 36.5 - 55.1 19.5 11.0 34.0 7.5 12.8 -0.5 - 16. - - •. _ 13. _ -_ 0.37 - - _ ' _ 0.08 -0.14 . - - -- 0.10 - -- 29.1 - -• -4.9 ' 5.1 5.2 5.0 5.1 5.3 6.7 3.6 '•7.4 3.7 5.3 6.1 11.4 6.2 12.7 6.3 9.2 10.4 0.3 0.2 0.5 0.3 0.3 0.3 27. 24. . 14. 11. 17. 19. 13. 8. 12. 8. 12. . • 6. 0.21 0.32 0.23 0.35 0.44 0.33 0.05 0.04 0.09 0.05 0.08 0.05 0.05 0.13 0.08 0.15 0.15 0.16 0.05 0.01 0.09 0.09 0.16 0.12 49.6 9.6 , 36.8 6.9 12.4 6.9 6.2 _ - 10.7 - -- - 0.4 -- 16. _ _ -- 5. 0.22 . -_ 0.08 - -- - 0.11 _ 0.07 - - _ - - 31.4 - - -4.7 5.7 5.2 5.2 4.2 4.6 1.1 3.2 -7.2 7.8 1.9 5.5 0.3 0.3 0.1 0.2 16. 17. 16. . - 14. 19. 7. 7. 9. ' -.0.50 1.12 0.40 0.72 -0.09 0.13 0.02 0.06 . -0.11 0.16 0.12 0.19 0.08 0.01 0.00 0.09 -32.7 22.0 25.6 14.8 -107 matter and nitrogen decrease with depth, and carbon:nitrogen ratios are narrow. Organic matter is fairly high in some of the C horizons, because the soils are very shallow. Phosphorus, cation exchange capacity and the available cations all decrease In amount with depth. Potassium is present in particularly large quantities in the A horizon of this community, b. Trollius laxus Variation This variation also occurs on seepage slopes, but almost exclusively in the subalpine parkland. Relief shape varies from concave to straight to convex. Exposure is variable, with slope gradients ranging from 13 to 28$. Humus covers 89-100$ of the ground surface. Exposed mineral soil is only present in one plot (1$), and rocks occur in only two plots (2-10$). There is no discernible erosion. The hygrotope is hygric. The herb layer in this variation is also very high in coverage, being 92-100$. The bryophyte layer is much better developed than in the type variation, with a coverage of 60-95$. The dominant species in this variation is Trollius laxus, with an average species significance of 8. The dominant species listed for the association are present in somewhat lesser amounts. Other species important in differentiating this variation from the type variation are Caltha leptosepala, which is present with much higher cover, and Brachythecium starkei, which is lacking in the Valeriana - Castilleja Variation. The soils of this variation are classed as Humie Gleysols (4), Rego Gleysol (l) and Lithic Orthic Regosol (l). The Ah horizon is finer-textured than in the type variation, Pig. 19. Valeriana - Castilleja Association, Trollius laxus Variation, Plot 25. White flowers belong to Trollius. 109 being classed mainly as sandy loam. The C horizon is similar to that of the Valeriana - Castille .ja Variation, with a range from sandy loam to sand. The soil chemical data are as described for the type variation, except that phosphorus increases slightly with depth instead of decreasing. The quantity of nitrogen in the Ah horizon of plot 64 is very low. The amounts of calcium and sodium differ between the two variations. There is more calcium in the C horizon and more sodium in the A horizon of this variation than in the Valeriana - Castilleja Variation. Carex nigricans Association (Ref. Tables 59, 60, 61, 62, 63; Fig. 20, 21, 22, 23, 24) Characteristic Combination of Species Carex nigricans Deschampsia atropurpurea Claytonia lanceolata Epilobium alpinum Polytrichum norvegicum Polytrichadelphus lyallii This association occurs in snow basins, depressions and temporary ponds In the alpine, low alpine and subalpine parkland areas. Relief shape varies from hummocky to straight to concave. Exposure is mainly southeast or southwest, with slope gradients ranging from 0 to 21$. Humus covers most of the ground surface (89-100$). Mineral soil occurs only in two plots, with a cover of 5-10$. There are usually no rocks present. No evidence of erosion was observed. The hygrotope is rated as mesic to hydric. The herb layer and bryophyte layer are both very well developed. Coverage of the herb layer is 45-100$, while the 110 Table 59 . General Environnent Carex nigricans Association Carex - Polytrlchadelphus Variatio Juncus - Carex - Drepanocladus Variation Plot lio. 1 2 15 33 ; 35 .78 58 77 71 81 Elevation (ft.) 7600 .7575 ' 7475 7460 7450 7400 7380 7350 7300 7450 Physiography Landforn depression depression sno» channel snos . channel slope , snoa channel snos basin . Snos basin sno« . channel temporary pond . Relief shape hucinocky hunocky straight straight straight to concave concave straight , concave concave straight Exposure . SE SE SE SB \ S SB • Sill neutral neutral W Slope gradient (?) 1 3 11 9 8 • 10 . 0 0 2 Layer coverage (?) C layer 95 90 97 100 100 . 99 100 ' 99 100 D layer 60 50 5 80 80 85 . 70 85 85 50 Plot Coverage (?) Hunus 99- 89 95 100 . 100 100 100 100 100 96 Mineral Soil 0 10 5 0 .0 0 0 0 0 0 Rock 1 1 0 0 0 o 0 0 0 4 Soil Hygrotope hygric hygrlc subhydrlc . hygrlc neslc subhygric hygric hydrlc - hygric Erosion none none Horizon depth (in .) tj-0 L-H 1-0 - . - .' - - H 6-0 • -Ah 0-3 •4 Ah 0-2 Ahf 2-4. 0-5 0-12 .0-9 . 0-18 0-6 0-5 0-4 B C Bra 3-11 Bn 3J-HJ Cg 11+ Cgj Dj- 18 Cg 18+ Bhf 4-6 Bra 6-15 15+ - Bhfg 12-30 Cg 5+ Cg 30+ Cg 9-18 18+ 18+ Bn1 6-9 Bm2 9-15 15+ Cg 5+ Classification Gleyed Alpine Dystric Brunisol Gleyed Alpine Dystric Brunisol Sonbric Rego Ferro-h'unlc Huaic Podzol Gleysol Fera Hunlc Gleysol Lithic Rego Hunlc Gleysol Orthic Regosol Alpine Dystric Brunisol Rego . Hunlc Gleysol Rego Hunlc Gleysol Ill . Table 60 Carex nigricans Association •' Juncus wtensianus - Carex Larex nigricans - Polytrichadelphus lyallii Vari at! on • nigricans - .Verianoclndus exannul atus Variation Plot lio. 1 2 15 33 35 .78 : 58 77 71 81 Plot Size (n2) 10 10 10 10 10 10 10 10.. 10 103 Extent of type (a2) 44 70 120 175 28 162 36 .119 . 120 108 Elevation (ft.) 7600 7575 7475 7460 7450 7400 7360 7350 73 00 7450 Altitudlnal area A A LA LA LA SP .SP . SP . SP LA " Aver.Specles Assoclation C layer Presence Aver.Species Slgni ficance bigni ficance • 1 Carex nigricans 9.4 9.4 9.3 9.6 9.6 9.6 ' 9.6 9.5 9.6 9 6.4' V / g 2 Deschanpsia -atropurpurca 1.+ 2-1 +.1 4.2 4.2. •4.2 3.1 4.1 .3 - IV 3 3 Clayton)a lanceolata - +.+ 7.2 +.+ 2.1 6.1 _ 2.1 4 III 4 4 Antennaria lanata 3.2 3.1 6.2 - - 2.2 5.2 1.1 4: III III 4 5 Veronica wornskjoldi i 1.+ 2.1 -• - 6.2 5.2 +.+ . _ • 4 3 6-• Erigeron-peregrinus 1.1 3,1 3.2 2.2. . 4.2 - ,2 III 1 '7 Juncus drunaondii - 3.1 4.2 3.2 1.2 • - :. 1.2 _ 2 1 1 8 Hieraciun gracile - - 3.2 - 2.1. 2.2 - 3.1 • 1.1 1 1.1 III. . 9 Sibbaldia procunbens 2.1 2.1 2.1 - 2.1 _ _ 1 1 10 Phyllodoce enpetrifornis 1.1 +.1 - - 1.1 2.2 + Ill + 11 Arnica latifolia - - - - +.1 1.1 6.2 _ 1.1 .3 II 2 12 Juncus parryi 3.2 - 5.2 - • - 4.2 - . • - 2 - II 2 13 Arnica nol1is '- - 2.2 4.2 • _ • 1 1.2 II. 1 14 Luzula sp. - - - - - 2.1 - 3.2 1.1 + II + 15 Luzula sahlenbargii - 2.2 2.2 1.2 - _ _ + .11 + 16 EpiI obi un alpinua - . - 1.1 - 2.1 - 1.1 + 2.1 II + 17 Carex spectabilis - - - +.3 -• - 1.2 + 1.3 II + 18 Juncus certensianus - - - - - - 2.1 - _ + 7.3. 1 3 19 Phleun alpi nun - - - 5.2 1.1 - 1 - 1 . 1 20 Luzula glabrata 3.2 ' 4.2 - • - - - _ 1 - 1 1 21 Arenaria capillaris +.1 - •• 1.1 - • - •' ".'+ - 1 + D layer Bryophytes 22 Polytrichua norvegicun Dh 8.2 1.1 - - 1.1 2.1 +.+ 4.2 3.1 4 2.1 IV 4 23 Polytrichadelphus lyallii Dh - 1.1 - 9.2 - 8.2 8.2 8.2 2.2 6 . III 6 24 Lophozia alpestris Dh 1.1 1.1 . - - - 2.1 - 3.1 ; 5.1 2 III 2 25 Pohlia nutans Dh - - - - - 2.1 - 1.1 + 3.1 II + 26 Polytrichua piliferun Dh - +.1 3.2 - - - - 1.1 - - II + 27 Ceratodon purpureus Dh 1.1 - 1.1 - - 2.1 - - - • + II + '28 Pohlia gracilis Dh 1.1 2.1 1.1 - - - - - + II + 29 Polytrichua foraosun Oh - 7.2 - - 9.3 - . - - - • 5 - .1 5 30 Orepanocladus exannulatus Dh - - - - - - - - - - 6.2 1 2 31 Brachytheciua colli nun Dh 5.1 2.1 - - - - - - - 1 1 1 32 Orepanocladus aduncus Dh - - - - - - - - . - 5.2 1 1 33 Barbilophozia hatcheri Dh 2.1 - - - - - - 1.1 . - + 1 + 34 Cephaloziella rubella Oh - - . - - - - - . - - - 3.2 1 + 35 Cephaloziella subdentata Dh - - - - . - +.+ - + 1 + Lichens 36 Lecidea granulosa Dh 3.2 1.1 +.1 - - - - 1.1 - + II + 37 Cladonia sp. Dh +.1 +.+ +.+ - - - - . +.+ + .11 + Total Species (incl.sporadics) 23 20 19 8 16 18 12 14 15 11 Sporadic Species 42 Brachytheciua starkei 58(3.1) C laver 43 Oicranun scopariua 71(4.1) 44 Klaeria blyttii 2(1.1) 38 Antennaria friesiana 1(1.1) 45 Lophozia ? ventrlcosa 1(1.1) 39 Antennaria unbrinella 15(1.1) 46 Orthocaulis floerkii 71(7.1) 40 Vacclnlun scopariun 1(1.1) 47 Philonotis anerlcana 1(1.1) 0 layer 48 Pohlia druKiondii 35(1.1) 49 Scapanla subalpina 35(1.1) Bryophytes Lichens 41 Brachytheciua curtun 71(5.1) 50 Cetraria Islandica K+.+) 112 bryophyte layer covers 5-85$. The dominant species is Carex nigricans, which has an average species significance of 9. It is the only constant species in the association. Other important species in the C layer, all of much lower cover, are Deschampsia atropurpurea, Claytonia lan ceo lata, Antennaria lanata and Veronica wormsk.joldil. Epilobium alpinum is considered as a characteristic species because of its preference for this association. The D layer is dominated by bryophytes,as .lichens are very sparse in this wet community. The important species are Polytrichum norvegicum and Polytrlchadelphus lyallii. Polytrichum formosum has a very high cover ln two plots. The association is divided into two variations: a. Carex nigricans - Polytrichadelphus lyallii Variation D• Juncus mertensianus - Carex nigricans - Drepanocladus  exannulatus Variation The floristic similarity indices for the ten plots of the association are shown in Table 6l. Some of the values are not too high, but these plots all have their highest affinity with each other. It can be seen that the Juncus - Carex - Drepano cladus Variation (plot 8l) is not very similar to the Carex -Polytrlchadelphus Variation. However, it is included within the Carex nigricans Association mainly because of the high coverage of Carex nigricans. Table 61 Floristic Similarity Indices for the Carex nigricans Association 1 2 15 33 78 58 77 71 81 1 60 52 47 42 52 43 55 50 18 2 58 55 68 56 48 57 55 19 15 48 44 52 59 51 49 18 33 45 78 73 83 52 17 35 53 40 44 48 18 78 71 81 53 20 58 73 47 17 77 54 21 71 20 81 The variations are described below, by general habitat, floristics and soils data. a. Carex nigricans - Polytrichadelphus lyallii Variation This is the type variation for the association. It occurs In snow basins and depressions in the alpine, low alpine and subalpine parkland areas. Relief shape varies from hummocky to straight to concave. Exposure is generally southeast or south west, and slope gradients range from 0 to 21$. Humus covers 89-100$ of the ground surface, mineral soil 0-10$, and rock 0-1$. There is no erosion. The hygrotope ranges from mesic to sub-hydric. The herb layer covers 90-100$ of the area. The bryophyte 114 layer also has a fairly high coverage, of 5-85$. The dominant species listed for the association also char acterize this snowpatch variation. Deschampsia atropurpurea, Claytonla lanceolata, Antennaria lanata, Veronica wormskjoldii and Polytrichadelphus lyallii, which are important in this variation, are lacking in the Juncus - Carex - Drepanocladus Variation. The soils associated with this variation are Rego Humic Gleysol (3), Fera Humic Gleysol (l), Alpine Dystric Brunisol (3), Sombric Ferro-Humic Podzol (l), and Orthic Regosol (l). Texture is generally coarser at greater depths. The A horizon varies from sandy loam to silt loam, sandy loam being predominant. The B horizon samples are all sandy loams. The C horizon is mainly a loamy sand or a sandy loam. The soil chemical data are shown in Table 63. The pH values increase with depth, but all the values are strongly acidic. Organic matter and nitrogen generally decrease with increasing depth. There is still a considerable amount of organic matter in some of the C horizons due to the very shallow nature of the soils. The amount of nitrogen in the C horizon of plot 2 appears to be very high, whereas the quantity ln the C horizon of plot 78 is very low. The carbon:nitrogen ratios are narrow. Phosphorus, cation exchange capacity, magnesium and potassium decrease in quantity with depth. Calcium and sodium decrease in some cases; in others, they decrease from the A to the B horizon, then increase from the B to the C horizon. Plot No. 1 Ah Horizon, Textural class SL Sand (?) 65.6 Silt (?) 34.4 Clay (?) 0 B Horizon Textural class SL Sand (?) 61.2 Silt (?) 38.8 Clay (?) 0 C Horizon Textural class LS Sand (?) 84.8 Silt (?) 15.2 Clay (?) 0 Table 62 Soil Texture Carex nigricans Association Carex - Polytrichadelphus Variati 2 15 33 35 78 SL SL SL SiL SL 66.8 66.2 60.0 46.4 48.0 32.8 33.3 38.6 50.4 48.6 0.4 0.5 1.4 3.2 3.4 SL SL SL 52.2 65.2 - 52.4 -47.4 33.3 - 44.4 -0.4 1.5 - 3.2 -LS LS LS LS SL 80.4 83.0 80.6 84.0 68.0 17.8 16.6 19.4 16.0 31.4 1.8 0.4 0 0 0.6 Juncus - Carex -Drepanocl adus Variation 58 77 71 81 SL SL L SL 63.0 55.2 51.2 51.0 31.8 40.4 37.6 42.2 5.2 4.4 11.2 6.8 SL - 48.0 - -- 49.5 - -- 2.5 - -SL SL SiL SiL 70.8 62.6 32.2 36.0 22.4 33.8 63.6 51.2 6.8 3.6 4.2 12.8 Plot No. L-H Horizon pH C (J) 0,1 (J) . N (J) C/N P (ppn) Ca (ne/IOOg) »g (ne/IOOg) Na (ne/IOOg) K (ne/IOOg) CEC (WlOOg) Ah Horizon pH C (J) oa «) N (J) c/« P (ppn) Ca (ite/IOOg) Bg <«/100g) Na (ne/IOOg) K (m/100g) CEC WlOOg) 6 Horizon pH C (I) 0» (J) N (J) C/N P (ppn) Ca (ne/IOOg) Bg (ne/IOOg) Na (ne/IOOg) K (ne/IOOg) CEC (ne/IOOg) C Horizon pH C (J) OK (J) N (J) C/N P (pp>) Ca (ne/IOOg) Bg (ne/100g) Na (m/IOOg) K (ne/IOOg) CEC (•e/IOOg) Table 63 •,,'••' Soil Chemical Analysis Carex nigricans Association Juncus - Carex -Carex - Polytrichadelphus Variation Drepanocladus — n 1 2 15 33 35 78 ,. 58 77 71 ' 81 '.5 4.3 _ 4.6 _ 23.9 _ 19.0 - - • 22.1 • -41.1 32.6 _ - - 38.0 • 1.3 1.3 - - - • - 1.6 -18. - 15. - - - '. - . 14. -16. 16. - - • - 21. -3.20 • 0.56 - _ - - - 0.12 -1.58 0.35 - 0.25 -0.23 0.26 - 0.43 -3.26 0.98 - - . - • '- 0.35 -138.0 - 53.6 - - • 58.4 4.8 4.6 • 4.6 4.9 5.1 4.8 5.0 4.6 5.0 5.6 14.1 . 14.6 12.1 12.5 6.6 13.3 5.6 15.9 14.0 2.5 24.3 25.2 20.9 21.6 11.7 22.9 9.7 27.4 24.0 . 4.3 0.9 1.0 • 0.7 0.9 0.5 1.0 0.4 0.9 1.1 0.3 15.' 15. ' 17. 14. 15. 14. 15. 18. 13. 8. 8. 8. 10. . 21. 6. 12. 9. 6. 4. 7. O.08 0.25 . 0.04 0.27 0.61 0.44 0.42 0.03 0.03 0.97 0.10 0.28 0.05 0.13 0.06 0.15 0.05 0.14 ' 0.03 0.19 0.16 0.13 0.15 0.15 . 0.44 0.27 0.26 . 0.27 0.19 . 0.16 0.35 0.59 0.20 0.07 0.00 0.22 0.09 0.27 0.05 0.13 36.4 41.9 35.1 . 41.3 24.3 17.0 . 16.2 31.6 24.1 18.9 5.4 5.3 4.9 5.2 5.6 4.5 5.8 . 8.2 - 7.5 - 3.0 - -7.7 10.0 14.1 12.8 - 5.2 - -0.4 0.4 ' 0.5 - 0.5 - . - 0.2 - -12. 14. 16. - 15. - - 15. - • -3. 6. 10. - 8. - 6. - -0.03 0.06 0.02 ' - 0.44 _ 0.08 • -0.02 0.01 0.02 0.04 - _ 0.02 • 0.13 0.10 • 0.13 0.24 - - 0.14 -0.05 0.03 0.11 - . 0.00 - 0.04 - -12.3 . 18.3 12.7 - • 27.6 - - 13.9 - -5.4 5.4 • 4.9 5.1 5.5 5.0 5.4 5.2 5.5 6.3 1.1 0.9 , 4.4 4.1 2.5 9.1 4.1 1.7 3.5 1.8 1.8 1.5 7.5 .7.1 4.3 15.7 7.1 2.9 6.0 3.0 0.1 0.6 0.2 0.2 0.2 0.1 0.3 0.1 0.3 0.2 13. 2. 18. 18. 15. 76. 16. 14. 11. 9. 3. 4. 4. 16. 8. 11. 2. 5. 2. 5. 0.29 0.37 0.19 0.44 0.38 0.09 0.32 0.23 0.11 1.17 0.01 0.01 0.01 . 0.03 0.02 0.02 0.02 ' 0.02 . 0.01 0.23 0.14 0.14 0.12 0.14 0.13 0.13 0.14 0.15 0.14 0.15 0.04 0.03 0.10 - 0.07 0.00 0.05 • 0.06 0.04 0.03 0.15 16.1 15.8 8.3 • 21.6 16.1 21.5 11.6 18.4 22.0 25.1 Fig. 20. Carex nigricans Association, Carex -Polytrlchadelphus Variation, Plot Jl, Note late-lying snow. Photo taken July 5,1969. 118 Fig. 21. Soil profile of Carex nigricans Association, Carex -Polytrichadelphus Variation, Plot 71. This soil is classified as a Rego Humic Gleysol with H, Ah and Cg horizons. Fig. 22. Soil profile of Carex nigricans Association, Carex -Polytrichadelphus Variation, Plot 77. This soil is classified as an Alpine Dystric Brunisol, with an Ah-Bml-Bm2-C horizon sequence. 119 b. Juncus mertensianus - Carex nigricans - Drepanocladus  exannulatus Variation This community is represented by only one plot (8l). It occurs In a temporary pond in the low alpine area. The relief shape is straight. Exposure is northwest, with a slope gradient of 2$. Humus covers 96$ of the ground surface, and rock 4$. There is no exposed mineral soil. No evidence of erosion is present. The hygrotope ranges from hydric to hygric. The herb layer and the bryophyte layer have similar cover age values. The C layer covers 45$ of the area, and the D layer 50$. There Is thus a much lower coverage of plants than in the type variation. The herbs grow mainly In large clumps, which stand up above the water level while the pond is still in existence. The dominant species in this variation is Juncus mertensia nus, with a species significance of 1. Of the important species in the C layer listed for the association, the only one present here is Carex nigricans, with a lower cover than in the type variation. In the D layer, several species differentiate this variation: Drepanocladus exannulatus, which Is lacking in the type variation, Drepanocladus aduncus and Cephaloziella rubella, both of which are exclusive to this variation. The soil is classed as a Rego Humic Gleysol. The soil texture becomes finer with depth, unlike the Carex - Folytricha-delphus Variation. The A horizon is a sandy loam, while the C horizon is a silt loam. The pH increases with depth; the values are higher than for the type variation. Organic matter and 120 Pig. 23. Carex nigricans Association, Juncus - Carex -Drepanocladus Variation, Plot 8l. Large clumps are Carex nigricans, smaller clumps are Juncus  mertensianus. Pig. 24. Soil profile of Carex nigricans Association, Juncus -Carex - Drepanocladus Variation, Plot 8l. This soil is classified as a Rego Humic Gleysol, with Ah and Cg horizons. 121 nitrogen decrease with depth, and carbon:nitrogen ratios are narrow. The amount of organic matter present in the A horizon is very low. Phosphorus and sodium decrease with depth, as described for the type variation. However, cation exchange capacity, calcium, magnesium and potassium all increase in amounts with depth. In the A horizon, calcium and magnesium are present in greater quantities than in the type variation; in the C horizon, there is more calcium, magnesium and potassium. Polytrichum norvegicum Association (Ref. Tables 64, 65, 66, 67; Fig. 25) Characteristic Combination of Species Polytrichum norvegicum Juncus mertensianus This association, which is represented by only two plots, occurs in temporary ponds in the alpine area. The relief shape is straight. The exposure is neutral, and thus there is no slope gradient. Humus covers 15-75$ of the ground surface, mineral soil 0-15$ and rock 25-70$. There is no evidence of erosion. The hygrotope ranges from hydric to hygric. The D layer is the conspicuous one, covering 60-80$ of the area. The C layer is very sparsely developed, coverage being only 10$. Polytrichum norvegicum is the dominant species, with an average species significance of 8. It is also the only constant bryophyte. Lichens are practically lacking, due to the wetness of the site. The only two constant species in the C layer are Juncus mertensianus and Carex spectabilis, both with lower Table 64 General Environment Polytrichum norvegicum Association Plot No. Elevation (ft.) Physiography Landform Relief shape Exposure Slope gradient (%) 79 7575 80 7575 -temporary pond straight neutral 0 Layer Coverage (%) C layer D layer 10 80 10 60 Plot Coverage (%) Humus Mineral soil Rock 75 0 25 15 15 70 Soil Hygrotope Erosion Horizon depth (in.) Ah Bf Cg -hydric-hygric-none 0-5 5-14 14 + 0-5 5 + Classification Gleyed Sombric Humo-Ferric Podzol Rego Humic Gleysol 123 Table 6 5 Polytrichum norvegicum Association Plot No. ? 79 80 Plot size (m ) 9 10 40 Extent of type (m ) 10 40 Elevation (ft.) 7575 7575 Altitudinal area A A C layer Pres- Aver. ence Species  Signifi cance 1. Juncus mertensianus 3.1 4.2 2/2 4 2. Carex spectabilis 4.2 2.2 2/2 3 3. Juncus drummondii - 2.2 1/2 1 4. Phleum alpinum 2.1 - 1/2 1 5. Poa cusickii 2.1 - 1/2 1 6. Car^ex nigricans - 1.3 1/2 + D layer Bryophytes 7. Polytrichum norvegicum Dh 9.6 7.3 2/2 8 8. Anthelia juratzkana Dh - 4.2 1/2 3 9. Pohlia gracilis Dh - 4.2 1/2 3 10. Kiaeria blyttii Dh 2.1 - 1/2 1 11. Aulacomnium palustre Dh 1.1 - 1/2 + 12. Bryum sp. h - 1.1 1/2 + Lichens 13. Lecidea granulosa Dh 1.1 - 1/2 + Total Species 8 8 Table 6 6 Soil Texture Polytrichum norvegicum Association Plot No. 79 80 Ah Horizon Textural class SL LS Sand (%) 70.8 81.2 Silt (%) 22.8 14.4 Clay (%) 6.4 4.Bf Horizon Textural class SL Sand (%) 76.4 Silt (%) 16.8 Clay (%) 6.8 Cg Horizon Textural class SL SL Sand (%) 72.0 72.4 Silt (%) 21.2 15.2 Clay (%) 6.8 12.4 Table 67 Soil Chemical Analysis Polytrichum norvegicum Associ ati on 125 Plot No. 79 80 Ah Horizon pH 5.0 5.3 C (?) 7.3 1.8 OM (?) 12.6 3.1 N (?) 0.7 0.2 C/N 10. 11. P (ppm). 8. 2Ca (me/1OOg) 0.29 0.39 Mg (me/1OOg) 0.05 0.02 Na (me/1OOg) ' 0.12 0.14 K (me/1OOg) 0.13 0.07 CEC (me/1OOg) 13.6 8.Bf Horizon pH 4.8 C (?) 2.6 OM (?) 5 N (?) 0.2 C/N 16. P (ppm) 4. • Ca (me/1OOg) 0.46 Mg (me/1OOg) 0.01 Na (me/lOOg) 0.12 K (me/1 OOg) 0.07 CEC (me/1OOg) 11.0 Cg Horizon pH 5.0 5.5 C (?) 1.0 0.9 OM (?) 8 1.5 N (?) 0.1 0.1 C/N 20. 14. P (ppm) 2. 3. . Ca (me/1OOg) 0.39 0.52 Mg (me/1OOg) 0.01 0.0Na (me/lOOg) 0.13 0.16 K (rae/lOOg) 0.04 0.09 CEC (me/1OOg) 6.3 14.0 126 Fig. 25. Polytrichum norvegicum Association, Plot 79. In foreground is transition to Carex  nigricans Association. In background are clumps of Carex spectabilis. 127 cover values. The floristic similarity index for the two plots is 53. The soils are classified as a Gleyed Sombric Humo-Ferric Podzol and a Rego Humic Gleysol. Texturally, the soils are fairly fine-textured. The A horizon is either a sandy loam or loamy sand. The B horizon, where present, is a sandy loam. The C horizon is a sandy loam. In plot 80, the texture becomes finer with depth. Table 67 presents the soil chemical data. The pH increases with depth, and all the values are strongly acidic. Organic matter and nitrogen decrease with depth. There is very little organic matter in the Ah horizon of plot 80. Carbon:nitrogen ratios are generally narrow. Amounts of phosphorus, cation ex change capacity and exchangeable cations have no particular relationship with depth that can be judged from the two repres entative plots. The quantities of the cations are also variable between the two plots. Drepanocladus exannulatus Association (Ref. Tables 68, 69, 70, 71; Fig. 26, 27) This association, represented by only one plot, occurs as a narrow band around the edge of a temporary pond in the sub alpine parkland. The relief shape is concave. Exposure is neutral and therefore there is no slope gradient. The entire surface is covered by humus, with no mineral soil or rock. There is no discernible erosion. The hygrotope varies from hydric to hygric. 128 Table 6 8 General Environment Drepanocladus exannulatus Association Plot No. Elevation (ft.) Physiography Landform Relief shape Exposure 82 7350 edge of temporary pond concave neutral Slope gradient (%) Layer Coverage (%) C layer D layer 8 95 Plot Coverage (%) Humus 100 Soil Hygrotope Erosion Horizon depth (in.) Ah Cg hydric-hygric none 0-12 12+ Classification Rego Humic Gleysol Drepanocladus Table 69 exannulatus Association Plot No. 82 2 Plot size (m ) 5 2 Extent of type (m )Elevation (ft.) 7350 Attitudinal area SP C layer Calamagrostis canadensis 3.1 Juncus mertensiarius 2.2 D layer Bryophytes Drepanocladus exannulatus Dh 9.6 Dicranella sp. Dh 4.1 Sphagnum nemoreum h 3.1 130 The D layer is predominant, covering 95$ of the area. The C layer is very sparsely developed, with a coverage of 8$. Drepanocladus exannulatus is the dominant species, with an average species significance of 9. Only two other bryophytes are found here: Dicranella sp. and Sphagnum nemoreum. This community is the only locality for Sphagnum in the research area. The only two species In the C layer are Calamagrostis canadensis and Juncus mertensianus. Calamagrostis is not found in any other community. The soil is classed as a Rego Humic Gleysol. Texture becomes finer with depth. The A horizon is a sandy loam, while the C horizon is a silt loam. The pH increases slightly with depth, but still remains strongly acidic. Organic matter and nitrogen decrease with depth, and the carbon:nitrogen ratios are narrow. Phosphorus, cation exchange capacity, calcium, magnesium and potassium decrease in amounts with depth, while sodium increases slightly. Table 70 Soil Texture Drepanocladus exannulatus Association Plot No. 82 Horizon Ah Cg Textural class SL SiL Sand (%) 61.4 38.0 Silt (%) 34.8 59.2 Clay (%) 3.8 2.8 Table 71 Soil Chemical Analysis Drepanocladus exannulatus Association Plot No. 8 2 Horizon Ah Cg pH 5.5 5.6 C (%) 6.6 2.5 OM (%) 11.4 4.3 N (%) 0.6 0.3 C/N 11- 9. P (ppm) 8. 6. Ca (me/lOOg) 0.22 0.16 Mg (me/lOOg) 0.03 0.01 Na (me/lOOg) 0.12 0.12 K (me/lOOg) 0.08 0.06 CEC (me/lOOg) 24.6 21.7 132 Fig. 26. Drepanocladus exannulatus Association, Plot 82, is represented by the narrow, dark band around the margin of the pond. In foreground is part of Valeriana - Castilleja Association, with Senecio  triangularis in flower. Photo taken Aug. 3, 1969. Fig. 27. Soil profile of Drepanocladus exannulatus Association, Plot 82. This soil is classified as a Rego Humic Gleysol, with an Ah-Cg horizon sequence. Photo taken Aug. 29, 1969, after pond has dried out. t 133 5. Distribution of Tree Species This section presents information on growth habits, distrib ution patterns, measurements of diameter, height and age, and occurrence of seedlings and shrubs of tree species. These data are somewhat scanty, since they were not the major aim of the research. However, it is believed that they are of value, particularly in judging the future vegetation development of the area; that is, forest versus open vegetation. More detailed autecological work should be done on this aspect in order to draw valid conclusions. As mentioned previously, there are four tree species present in the study area: Abies lasiocarpa, Picea engelmannii, Pinus albicaulis and Pinus contorta. Plnus contorta was observed only in one locality (a Juncus parryi community), growing as a single very low shrub one and a half feet in height (see Table 7*0. Plnus albicaulis is slightly more frequent, but is still rare. Only two very low shrubs were noted, the remainder of the occurrences being as seedlings. Abies lasiocarpa and Picea engelmannii are thus the predominant tree species. In the alpine area1, Abies occurs mostly in low krummholz colonies, while Picea occurs as solitary or a few clumped speci mens, usually taller than Abies and not growing in krummholz form. In the subalpine parkland, Abies, in particular, grows much taller, and in some cases is as tall as Picea. 1 Throughout this section, "alpine area" includes the low alpine area. 134 In the alpine area, the tree species occur on ridges, which have less snow cover than the surrounding terrain, and are thus free of snow earlier in the growing season. This is the situation in a region of high snowfall, as in the coastal alpine zone (Krajina, 1965). In the subalpine parkland the trees occur not only on ridges but also on seepage slopes. Table 72 presents the diameter, height and age measurements made on the oldest specimens in each tree island community which was sampled. Table 73 summarizes the ages. In considering either the alpine area or the subalpine parkland, Abies laslo carpa is older than Picea engelmannii. The more interesting comparison is that the subalpine trees are much older than those in the alpine area. These ages, combined with the fact that no dead wood was observed, suggest that there has been a recent migration of tree species into the alpine area. It was hoped that'these migrations could be correlated with a climatic change, but there is Insufficient tree mensuration and climatic data to do this. Such a task would be a separate project in itself. Franklin et al. (1966) have correlated tree invasions into subalpine meadows in the Pacific Northwest with a warming trend in the early part of this century. It is possible that this may be the case for the Big White area also. Table 72 Diameter, Height and Age Measurements^ of Abies lasiocarpa and Picea enoelnannll Community Plot Species" Olaneter Height ( , Height Range (In.) Range (ft.) (ftj Age ^ Height Estimated (yrs.) (ft) Age (yrs.) Picea engelmannii Association Abies lasiocarpa Association sitchensis Assoclati on engelmannii - Vacci ni um  scoparium Assoc! ati on 49 (A) Picea engel. - - 4.0 8 36 2.0 41 39 0.7 44 (A) Abies laslo. 1-3 2.5-8 3.0 8 52 2.5 109 86 1.0 > 2.0 7 47 2.5 . 60 52 1.5 53 (A) Abies laslo. . 3-8 2.5 8 36 3.0 51 48 0.5 1.5 6.5 72 2.0 98 85 1.0 56(SP) Abies lasio. 3-5 7-11 6.0 10 43 2.5 98 65 1.5 62 (SP) Abies lasio. 1.5-8.5 5-15 8.5 15 162 2.0 162+ 7.0 12 105 2.0 229 167 1.0 74 (SP) Abies lasio. 0.8-8.0 5-18 . 8.0 18 98 3.0 116 105 1.0 6.0 17 134 2.0 274 204 1.0 51 (A) Abies lasio. 1-2.5 5-6 2.5 6 55 2.0 55+ 1.3 5 31 2.0 31+ Picea engel. 5-6 - 6.0 12 56 2.5 • 80 75 0.5 6.0 10 52 2.5 60 57 1.0 54 (LA) Abies lasio. 1.5 5.5 23 2.0 53 38 1.0 2.0 5 35 2.0 70 58 0.7 Picea engel. - - 7.0 11 49 2.0 67 58 1.0 3.0 11 37 2.0 48 44 0.7 61 (SP) Abies laslo. 3-7 2.5 7 58 2.0 58+ 2.0 . 7 72 2.0 106 89 1.0 Picea engel.. - - 9.0 12 57 2.0 91 74 1.0 10.0 13 74 2.0 83 80 0.7 70 (SP) Abies laslo. 0.5-7 "3.5-18 " 7.0 15 139 2.0 267 203 1.0 6.0 18 126 3.0 315 189 2.0 Picea engel. 0.8-9.5 5.5-18 9.5. 18 121 2.0 121+ 4.8 • 12 55 2.0 105 80 1.0 Age of oldest trees Table 73 Age1of Tree Species in Alpine and Subalpine Parkland Areas Species Area Average Range Abies laslocarpa Alpine2 74 (6) 51-109 Subalpine 201 (7) 98-315 Picea engelmannii Alpine 59 (5) 41-80 Subalpine 100 (4) 83-121+ 1 Age of oldest individuals 2 Includes low alpine area Table 74 shows the occurrence of conifer seedlings and shrubs in alpine and timberline communities, exclusive of the sampled tree islands. No conifer seedlings were observed in the tree island communities. Picea engelmannii seedlings are the most common, and they are scattered among four communities. More seedlings were found in the Antennaria - Sibbaldia - Salix community than in any other. This corresponds to the statement made earlier that the tree species occur mainly on ridges in the alpine area, since this community occurs on ridges. Table 74 Occurrence of Conifer Seedlings and Shrubs in Alpine and Timberline Communities1 No. of seedlings Community Carex phaeocephala Variation Juncus parryi Association Antennaria lanata Association Phyllodoce empetriformis - Antennaria lanata Association Phyllodoce - Antennaria Variation Valeriana sitchensis - Castille,ja elmeri Association Trollius laxus Variation 1 Exclusive of sampled tree islands Plot No. Abies Picea Pinus a. Shrubs Antennaria lanata - Sibbaldia procumbens Association Antennaria - Sibbaldia - Salix Variation 10 (A 22 (A 13 L 17 (LA Outside 68 (SP) 32 (LA) Outside 27 (A) Outside 31 (LA) 57 (SP) 69 (SP) -11 2 — — l Picea 1 Picea 1 Plnus  contorta dt ft.) Abies, Picea C6~ ft7T~ Several Picea (8 ftT) ->3 138 6. Vegetation Relationships . Relationships between the communities on a vegetational basis are presented in this section. In order to compare all the associations with one another, a synthesis table was constructed (Table 75) in which the major species are listed with their presence and average species significance values shown for all associations. The associations are arranged along a gradient of increasing moisture (as described in section 4). In this manner, it is possible to show the floris tic distinctness of some associations and the overlap of species ln other associations. The first half of the table (down to Juncus mertensianus) presents species which are characteristic of at least one assoc iation. The species which are boxed show the characteristic species for that particular association. The second part of the table (beginning with Selaglnella densaK lists species which occur in at least one association with a presence of IV or V (or corresponding fraction). The species enclosed by dotted lines are the high-presence species for that association. It can be seen that there is some overlap of species In the mesic associations, as expected, but in general, both the characteristic species and the high-presence species follow the moisture trend. That is, there is a different group of species which is most important for each community, from the dry associations through to the wet ones. Species occurring in associations for which they are not characteristic are usually much less important in those associations. Table 75 Syntheslt Table for all Association! Juniperus conuunis Carex phaeocephala Arenaria caplllarls Festuca brachyphylla Tortula ruralls Peltigera raalacea Rhizocarpon geographicura Sibbaldia procumbens Unbillcarla hyperborea Alectorla mlnuscula Polytrlchun piliferum Antennaria lanata Juncus parryI Lecidea granulosa Sallx cascadensls Gentlana glauca Phyllodoce empetriforrais Vaccinium scoparium Dicranum scoparium Picea engelmannii Cetraria plnastri Abies lasiocarpa Parmeliopsis hyperopta Arnica latlfolta Valeriana sitchensis Polytrichadelphus lyallii Carex spectabilis Castllleja elmeri Arnica noil Is Erfgeron peregrinus Senecio triangularis Ranunculus eschscholtzii Aulacoranlum palustre Philonotis ame He ana Brachytheclui asperriraun Carex nigricans Oeschampsfa atropurpurea Clayton!a lanceolata Epi1 obiua alpi nun Polytrichum norveglcua Juncus mertensianus Selaglnella densa Barbilophozia hatcherl luzula spicata Peltigera canina Trlsetum splcatum Sedun lanceolatum Potentllla dlverslfolia Haplopappus lyalllt Antennaria umbrinella Cladonia cameo]a Solorina crocea Hieracium graclle Arenaria obtusiloba Agrostis variabilis Carex pyrenalca Luzula lahlenbergll Cetraria Islandica Cladonia ecmocyna Lophozla alpestris Cetraria erlcetorum Parmeliopsis ambigua Luplnus latifollus Pedicularis bracteosa Luzula sp. Veratrun virlde Ml tell a breierl Caltha leptosepala Juncus drunnondll Phleuo alplnua Troll I us laxus Veronica •ormskjoldll V.5 V.* V.2 V.2 V.3 IV.1 IV.1 ED IV.2 III.3 IV.* IV.1 III. IV. * V. 5 V.3 V.* IV. 3 V. * 11.1 IV.* -V.5 v.* IV.5 -IV.5 -V.5 i .0 V.5 V.5 III.* V.8 IV.* v.5 IV.* -III.* _ III.2 V.3 •8 I.* IV.2 I.* I. V.2 II.* V.2 V.* H V.3 111.1 U. v.* rnrt i.+ II.I in.* n.* II.I IV.5 111.1 III.3 I.+ 11.1 IV. * !.• V. * IV.2 IV.1 11.1 11.1 o innll | 1 | 3 Ablei - Plcti -.* 1/2.* 111.1 -.i 1/2.* IV.1 -.2 Ml.* 111.1 - 1/2.* --.1 2/2.+ 111.1 -.* 1/2.2 IV.3 -.3 1/2.1 V.3 -.1 2/2.1 IV.* - - IV.5 -.2 1/2.2 U - 1/2.1 v.* -.9 - V.T 1/2.+ IV.* - 2/2.9 V.8 -.2 V,1 2/2.1 1/2.* III.* 8 •s IV.2 V.3 I.* I.. V.1 111.1 v.* 2/3.3 3/3.* 37579' 3/3.* 3/3.5 3/3.6 MM 2/3.* 3/3.3 2/3.1 2/3.1 3/3.3 3/3.3 II.*' _ II.I _ II.* IV.2 IV.3 v.* II.+ 11.1 \ III.* cm - II.3 rvTel V.b - V.6 11.3 V.5 v.* V.5 - V.5 - IV.2 - IV.6 - IV.* _ III.? 11.2 V.4 _ IV.3 11.1 11.1 III.* 11.2 1/2.-rrrm 11.1 in.i V.9 IV.3 III.* EA 2/2.3 .1/2.+ 1/2.+ 11.2 1.3 17275 JTvlFJ in.* 11.1 _ 11.+ -.1 _ - - -nv.1i _ III.* 1.+ -.« 1/2.2 - - I.+ 1.+ - -1 . 1 sn-i.+ -.1 1/2.+ - - III.* - • - - -;iv.*, 11.* - 1.+ -.+ • - - - -' - - -111.1 ' IV.2 | 11.1 - 1.+ -.1 - - 11.1 - - -JlV.1 ' _ - _ - - - - - • - - -111.1 1iv.1 ! - - - - - in.* - 11.1 - - - -- •IV.3 , ' v.*, - --.1 - - - - - --! IV.2 1 • V..2 1 -,+ - - . - 111.2 . - - - -111.+ jv.i: ivjj III^* -.3 mz - - - - - -- [tVj • - - in.* - 111.1 I.* 111.1 - • -111.+ 11.* ;rv:n -.1 - - - - - - --111.1 11.1 11.* IIVJ! |v.*' »V.1J I.* III.2 -.1 1/2.+ - -11.1 III.3 -II.* - -111.1 11.1 llv.3l II.* . • _ _ - - - - -111.2 11.1 III.2 <IV.2j TO! _ fT3T - 11.1 - - -11.+ in.3 11.1 tv..+j 111.2 111.1 -.3 s/m liv.3; .iv.i; 27372] II.* II.* in.2 • - -_ _ -.2 ! 2/3.15 - - - - -_ _ 11.1 _ 1.1 _ - I.+ - -_ _ .. _ _ i2/3.2; - - - - -II.+ 11.1 _ _ 111.5 12/3.5- - 11.1 II.* - ----• - --- 1' '3/3.3! L3/3JJ 11.3 III.3 III.2 11.1 ! v.*i Jiv.1! jlV.6| 111.1 1.1 1/2.1 1/2.1 -- - - - • •1.1 111.3 - -140 Floristic similarity Indices between all the associations and variations are presented in Table 76. The rationale for using this technique has been discussed previously (section 4). By this method, it can be shown that some communities have higher similarities than others. The evaluation of the indices is subjective, since there are no tests of significance of the differences for indices based on dominance (Mark and Burrell, 1966). Despite this limitation, however, general conclusions can still be drawn from the indices. In Table 76, the communities are arranged in the order in which they were described in section 4. Each community, with its highest affinities, is discussed separately. The Juniperus communis Association has very low affinities with all the other communities. Its highest similarities are with the Juncus parryi Association and the Carex phaeocephala Variation, but even these values (27$ and 26$ respectively) are not high enough to draw any conclusions. The Antennaria - Sibbaldia - Salix, Carex phaeocephala and Carex breweri Variations, which all belong to the Antennaria-Sibbaldla Association, are arranged separately to show their affinities. The Antennaria - Sibbaldia - Salix Variation has its highest similarity with the other two variations, which is what would be expected. The values are 63$ and 45$. It has some affinities with the Antennaria lanata Association, having a similarity of 42$; this would be primarily through the pres ence of Antennaria lanata. The Carex phaeocephala Variation has a similarity value of 63$ with the Antennaria - Sibbaldia - Salix Juniperus communis Associ ati on Antennaria - Sibbaldia Associ ati on Antennaria - Sibbaldia - Salix Vari ati Carex phaeocephala Vari ati on Carex breweri Vari ati on Juncus parryi Association Antennaria lanata Associ ati on Phyllodoce - Antennaria Association Phyllodoce - Antennaria Vari ati on Antennaria - Vaccinium Vari ati on Picea engelmannii Associ ati on Abies lasiocarpa Associ ati on Abies - Picea - Vaccinium Associ ati on Abies - Valeriana Association Carex spectabilis Associ ati on Valeriana - Castilleja Associ ati on Valeriana - Castilleja Vari ati on Trollius laxus Vari ati on Carex nigricans Associ ati on Carex - Polytrichadelphus Vari ati on Juncus - Carex - Drepanocladus Vari ati Polytrichum norvegicum Associ ati on Drepanocladus exannulatus Associ ati on Table 76 Floristic Similarity Indices for all Communities 23 26 63 19 45 36 27 38 39 37 20 42 31 42 16 20 24 26 38 44 22 27 30 30 50 61 62 24 24 25 23 21 20 10 15 13 8 7 7 8 8 7 8 14 16 14 16 14 19 19 34 31 38 44 7 3 3 5 10 10 19 13 6 53 21 23 19 23 34 36 27 36 14 9 7 6 4 8 16 15 23 21 5 4 11 36 34 7 5 4 6 13 13 20 14 4 4 9 26 22 52 6 7 6 7 13 11 17 12 4 4 10 13 18 18 18 1 1 1 0 2 2 4 2 1 1 3 2 5 6 7 20 2 1 1 1 3 7 5 4 1 2 1 4 9 7 6 9 10 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 25 3 142 Variation, 36$ with the Carex breweri Variation, and 39$ with the Juncus parryi Association. This variation is still consid ered to be most similar to the association in which it has been placed, since the Antennaria - Sibbaldia - Salix Variation is the type for the association. It is, however, not closely allied with the Carex breweri Variation. It shows some similar ity to the Juncus parryi Association, but not enough to be grouped with it. The Carex breweri Variation shows approximate ly the same similarities with the Juncus parryi Association (37$) and the Antennaria lanata Association (42$) as it does with the Antennaria - Sibbaldia - Salix Variation and the Carex  phaeocephala Variation (45$ and 36$ respectively). However, the values are not high enough to warrant placing this variation in another association. Tne Juncus parryi Association has its highest similarity with the Antennaria - Vaccinium Variation of the Phyllodoce -Antennaria Association, with a similarity index of 50$. Other, lower, affinities are with the Antennaria lanata Association (44$), the Phyllodoce- Antennaria Variation (38$), and the Antennaria - Sibbaldia Association (38$, 39$, 37$). The Antennaria lanata Association has its highest affinities with the Antennaria - Vaccinium Variation (6l$). This will be discussed under the Antennaria - Vaccinium similarities. This association has lower similarities with the Phyllodoce -Antennaria Variation (44$), the Juncus parryi Association (44$), the Carex breweri and Antennaria - Sibbaldia - Salix Variations (both 42$). The Phyllodoce - Antennaria and Antennaria - Vaccinium Variations are classed into the Phyllodoce - Antennaria Assoc iation. The Phyllodoce - Antennaria Variation is most similar to the Antennaria - Vaccinium Variation, with a value of 62$. It has much lower similarities with the Antennaria lanata Assoc iation (44$) and the Juncus parryi Association (38$). The Antennaria - Vaccinium Variation has a higher similarity to the Juncus parryi Association than the previous variation, with an Indkx of 50$. This variation has its highest affinities with both the Phyllodoce - Antennaria Variation (62$),which is the type for the association, and the Antennaria lanata Association (6l$). This fact corroborates the statement made in the descrip tion of this association that the Antennaria - Vaccinium Variat ion, while classified in the Phyllodoce - Antennaria Association is best regarded as a transition community between this associa tion and the Antennaria lanata Association. The Picea engelmannii Association has low indices of simil arity with all other associations. It has its highest affinity with the Abies - Picea - Vaccinium Association (38$), mainly due to the common presence of Picea engelmannii. The Abies laslocarpa Association has affinities with the Abies - Valeriana Association (53$) and the Abies - Picea -Vaccinium Association (44$). In this case, the similarity is due to the common presence of Abies lasiocarpa. The Abies - Picea - Vaccinium Association is not too? closely allied to the other tree island communities. It does, however, show its highest similarities with these communities: 144 44$ with the Abies lasiocarpa Association, 4l$ with the Abies -Valeriana Association, and 38$ with the>. Picea enge lmannii Assoc iation. These affinities are due mainly to the tree species. This association also shows similarities with the Phyllodoce -Antennaria Variation, but at a lower level (34$). The Abies - Valeriana Association has its highest affinity for the Abies lasiocarpa Association, with a similarity value of 53$. It has a lower similarity to the Abies - Picea -Vaccinium Association (4l$). It shows some affinities with the Valeriana - Castilleja Variation (36$), due to the common pres ence of several meadow species, such as Valeriana sitchensis, Castilleja elmeri, Senecio triangularis and Veratrum viride. The Carex spectabilis Association has low indices of simi larity with the other communities. Its highest similarities are with the Antennaria lanata Association (36$), the Antennaria-Vaccinium Variation (36$), the Juncus parryi Association (34$) and the Valeriana - Castilleja Variation (34$). The Valeriana - Castilleja and the Trollius laxus Variat ions are grouped into the Valeriana - Castilleja Association. The Valeriana - Castilleja Variation does have its highest affinity with the Trollius laxus Variation, the similarity index being 52$. It also has affinities with the Abies -Valeriana Association and the Carex spectabilis Association, but at much lower levels of similarity (36$ and 34$, respect ively). The Trollius laxus Variation is most similar to the Valeriana - Castilleja Variation (52$). It has quite low affinities for any other community: 26$ for the Abies -145 Valeriana Association, 22$ for the Carex spectabilis Association, and 20$ for the Phyllodoce - Antennaria Variation. Thus, the initial classification is supported. The Carex - Polytrlchadelphus and the Juncus - Carex -Drepanocladus Variations are classified as the Carex nigricans Association. It can be seen that the Carex - Polytrlchadelphus Variation has no high similarity to any other community. It only has a 20$ similarity value for the Juncus - Carex -Drepanocladus Variation. The Juncus - Carex - Drepanocladus Variation actually has a slightly higher similarity for the Drepanocladus exannulatus Association (25$), but is included in the Carex nigricans Association because of the high cover of Carex nigricans. This association is thus very homogeneous. The Polytrichum norvegicum Association has no high affin ities for any other community. The similarity values are all extremely low, the highest being for the Carex nigricans Assoc iation (9$ and 10$) and for the Carex spectabilis Association (9$). This community is also very homogeneous. The Drepanocladus exannulatus Association has only one similarity value which is greater than 3$J this is a value of 25$ for the Juncus - Carex - Drepanocladus Variation. This value is due to the common presence of Drepanocladus exannulatus. However, as mentioned previously, the Juncus - Carex - Drepano cladus Variation is retained in the Carex nigricans Association. Dahl (1956) gives some general rules for distinguishing different associations, alliances and orders. This can be ex tended to variations of associations, in which case the variations of an association should have higher indices of simi larity with each other than with any other association. Table 76 indicates that all the variations do fulfil this rule. In general, there is a very low degree of similarity among the communities. This substantiates the fact that they are distinct groupings of species, which are recognisable in the field. Thus, the indices of similarity support the classif ication system. Similar conclusions were reached by Dahl (1956) and Bliss (1963). Many of the plant communities recognized on Big White Mountain are ecologically comparable to ones described in the mountains of Scandinavia, Scotland, the United States, the U.S.S.R., Europe, Alberta and other parts of British Columbia. Juniperus communis communities on exposed rock outcrops have been described from the alpine zone of Garibaldi Park (Archer, 1963), where Penstemon menzlesii, which is absent on Big White, is a co-dominant. This community is also found in Washington and Oregon in the subalpine parkland (Franklin and Dyrness, 1969). McVean and Ratcliffe (1962) describe a compar able community in the low alpine zone of the Scottish Highlands, in which Juniperus communis ssp. nana is the dominant. The Antennaria lanata - Sibbaldia procumbens Association does not seem to have a counterpart elsewhere. Dryas octopetala has a very limited occurrence in this association. Dryas  octopetala communities have been described in Alberta, Colorado, Montana, Scandinavia, Scotland, the U.S.S.R. and the Pyrenees of Spain and France (Beder, 1967; Marr, 1961; Bamberg and Major, 147 1968; Johnson and Billings, 1962; Dahl, 1956; McVean and Ratcliffe, 1962; Sukachev, 1965; Braun-Blanquet, 1948). This Dryas octopetala community is more widespread on calcareous parent material since the species is calcicolous. The presence of acidic parent material on Big White thus explains its reduced distribution in the area. Sibbaldia procumbens associations have been described in Scandinavia (Gjaerevoll, 1956) and British Columbia (Archer, 1963) as snowpatch communities. On Big White, this community occurs on exposed ridge tops. Either these ridges do have a heavy accumulation of snow, or else a different ecotype of the species has developed here which does not require as much moisture as provided in snowbeds. The Juncus parryi Association on Big White may be ecologic ally equivalent to the dwarf shrub heath-rush community, with Juncus trifldus, found in the Presidential Range of New Hamp shire (Bliss, 1963). It Is also comparable to the Juncus tri fldus association ln Czechoslovakia, which is placed In the order Caricetalia curvulae (Krajina, 1933). Since it is found on warm dry slopes, it may be the community most comparable to the alpine grasslands of Alberta and eastern Oregon and Wash ington. Alpine grasslands are typical of the east slopes of mountains in western North America because of the rain shadow effect produced there. Therefore, with regard to amount of precipitation, Big White shows greater similarity to the coastal mountains than to the Rockies. The Antennaria lanata Association seems to be unique to the study area. An Antennaria lanata variant of the Carex 148 nigricans Association has been described in Alberta (Beder, 1967), but the ecology of this community appears to be different, since it occurs in snowpatches. Marr (1961) describes a Willow-Sedge Hummock Stand-Type in Colorado, which is ecologically similar to the Antennaria lanata Association in that the habitat consists of hummocks elevated by ice beneath them. The European counterpart of this association Is found in the Caricetalia curvulae of Braun-Blanquet and Jenny (1926). The Phyllodoce empetriformis - Antennaria lanata Associat ion corresponds to the Phyllodoce glanduliflora - Vaccinium  scoparium Association in Alberta (Beder, 1967) and the Phyllodoce  coerulea - Vaccinium myrtlllus community in Scandinavia (Dahl, 1956). A similar heath community is described in British Columbia and Washington (Archer, 1963; Brooke, 1966; Peterson, 1964; Franklin and Dyrness, 1969; Kuramoto, 1968), which con sists of Phyllodoce and Casslope mertenslana. It is interesting that Cassiope is completely absent from the study area. The European equivalent of this community is found in the Rhodoreto-Vaccinion alliance of Braun-Blanquet and Jenny (1926) and Krajina (1933). The occurrence of krummholz and tree islands at timberline is a we11-documented phenomenon. The tree species present dep end on the geographical area. Archer (1963) describes an Abies  lasiocarpa - Chamaecyparis nootkatensis association in Garibaldi Park. This area was considered as transitional between the coastal and interior alpine zones. Four tree island communities are., distinguished on Big White, all indicative of the interior zone: the Picea engelmannii Association, the Abies lasiocarpa Association, the Abies laslocarpa - Valeriana sitchensis Assoc iation, and the Abies lasiocarpa - Picea engelmannii - Vaccinium  scoparium Association. The latter association has been describ ed in Alberta by Ogilvie (1961). The Carex spectabilis Association has been described in the alpine zone of Garibaldi Park (Archer, 1963) and in the subalpine parkland in Washington and Oregon (Franklin and Dyrness, 1969). This community seems to have coastal affinities, with no reported occurrences in the Rockies. The Valeriana sitchensis - Castilleja elmeri Association is similar to communities described elsewhere in British Columbia, in Washington and the U.S.S.R. Archer (1963) includes such a community in an alpine meadow group. Fraser (1970) describes a Valeriana - Lupinus - Epilobium angustifolium community on seepage slopes in Garibaldi Park. A Valeriana  sitchensis community occurs in the subalpine parkland and in the lower part of the alpine zone in Washington (Franklin and Dyrness, 1969). Subalpine moist meadows with Valeriana, Troll ius and Caltha are described in the U.S.S.R. (Sukachev, 1965). The Carex nigricans snowpatch community is common in North America, in both alpine and subalpine parkland areas. It has been documented in Alberta (Beder, 1967), British Columbia (Archer, 1963; Peterson, 1964; Brooke, 1966) and Washington (Kuramoto, 1968; Franklin and Dyrness, 1969). Comparable communities are the Polytrichum alpinum - Carex bigelowli snowbeds in Scotland (McVean and Ratcliffe, 1962) and the Carex 150 bigelowil association in Scandinavia (Gjaerevoll, 1956). It is well known that bryophytes form the main cover in habitats with an extremely long snow duration. The Polytrichum  norvegicum Association on Big White has previously been des cribed in British Columbia by Archer (1963) in the alpine zone, Peterson (1964) and Brooke (1966) in the subalpine parkland. Archer's association includes Gymnomitrlum varians as a co-dominant. This community is also known from Scotland (McVean and Ratcliffe, 1962), where Dicranum starkei is a co-dominant, Scandinavia (Gjaerevoll, 1956), Czechoslovakia (Krajina, 1933), central Europe (Braun-Blanquet and Jenny, 1926) and the Pyrenees (Braun-Blanquet, 1948). Reference to the Drepanocladus exannulatus Association was found in a study by Dahl (1956) in Scandinavia and another in the Tatra Mountains of Czechoslovakia by Krajina (1933). This community does not appear to be very common, and has not been previously described from North America. It can be seen from the above discussion that although floras differ considerably among geographical areas, similarit ies in environment produce comparable communities. In addition, many of the communities distinguished on Big White are not only ecologically but also floristically similar to those of other alpine and subalpine areas. 151 7. Vegetation-Environment Relationships This section presents relationships between communities on an environmental basis. No attempt is made to show causal relations, as this is impossible to prove without detailed autecological studies. The section is subdivided into four parts: A. An analysis of all environmental variables measured, with the basic aim of determining which factors are important in differentiating the communities B. A detailed study of soil moisture for a number of alpine and subalpine communities C. A summary of topographic-altitudinal relationships among the communities D. A discussion of the communities associated with each soil type Successional relationships among the communities are not dealt with in the present study for several reasons. One is the fact that the research was not organized for the purpose of studying succession, as only distinct homogeneous stands were chosen for analysis. If transitional areas had also been studied, more could have been said about changes taking place in the communities. Secondly, the vegetation of the study area is developed on parent material of uniform age. Thus, it is not possible to produce a scheme showing changes in vegetation devel opment with different times since deglaciation, as was done by Praser (1970). Finally, It is believed that the plant communit ies in such an alpine area are relatively stable and undisturbed, 152 and the rates of development are slow. Therefore, In this environment, the successional approach Is of very limited use (Dahl, 1956). On a larger scale, it can be speculated whether forests will take over the alpine area, or whether the alpine area will maintain itself. In section 5* it was suggested that there had been a recent migration of tree species from the sub alpine to the alpine area. There are insufficient data to deter mine precisely the direction of change in tree establishment. It appears that in this ecotone area discontinuous changes in climate may cause changes in forest development in a constantly shifting pattern. However, accidental factors such as seed production and dispersal are also Important. A. Analysis of Environmental Variables General environmental variables and physical and chemical soil data are summarized for all the communities in Tables 77 and 78. Instead of presenting actual values, the terms "high", "medium" and "low" are used. The limits for these terms (given in Appendix 4) were chosen in reference to the present data only. These general terms are believed to be more useful for comparis on of the communities. The communities are grouped on the basis of hygrotope, because it is considered to be the most important factor in delimiting the various communities. The communities are discussed in four groups: xeric, mesic, hygric and subhydric. The xeric group contains five communities: the Juniperus  communis Association ,the three variations of the Antennaria -Sibbaldia Association, and the Juncus parryi Association (the Table 77 Community Summary of General Environmental Variabl Altit. Expos. Slope .Relief 1 es for all Communities Wind Erosion Hygrotope Mineral Humus Soil Rock Soi Dep Juniperus communis Association M S hi straight strong none xeric L L H L Antennaria - Sibbaldia Association • Antennaria - Sibbaldia - Salix Variation H SSW L straight-convex very strong strong xeric L M H M Carex phaeocephala Variation M SE L straight very strong moderate xeric M L M L Carex breweri Variation 111 SSE M convex very strong strong xeric M H L M Juncus parryi Association M S M straight-convex strong slight subxeric M M M M Antennaria lanata Association H ENE L hummocky moderate slight mesic H M L H Phyllodoce - Antennaria Association Phyllodoce - Antennaria Variation M SSW HI hummocky moderate none mesic H L L H Antennaria - Vaccinium Variation H S M straight moderate none submesic H L L M Picea enqelmannii Association H ESE H straight strong none submesic M L M L Abies lasiocarpa Association H WSW H straight-convex strong slight mesic M L M L Abies - Picea - Vaccinium Association M SW H straight strong sl ight mesic M L L M Abies - Valeriana Association L WSW H concave moderate none subhygric M L L Id Carex spectabilis Association H SSW M straight moderate none hygric H G H Valeriana - Castilleja Association Valeriana - Castilleja Variation L WSW M concave slight none hygric H L L i Trollius laxus Variation L WSW III straight-concave slight none hygri c H L L M Carex niqricans Association Carex - Polytrichadelphus Variation M SE L straight-concave slight none hygric H L L H Juncus - Carex - Drepanocladus Variation IH NNUI L straight slight none subhydric H L L M Polytrichum norveqicum Association H L straight slight none subhydric M M M L Drepanocladus exannulatus Association L L concave slight none subhydric H L L M L = low, M = medium, H = high See Appendix 4 for class limits Table 78 Summary of Physical and Chemical Soil Data1 for all Communities C"'ty Sand Silt Clay pH Ca Mg Na Juniperus communis Association H L L L L L L Antennaria - Sibbaldia Association Antennaria - Sibbaldia - Salix Variation If L L ' M H L H Carex phaeocephala Variation M M M M M -L M Carex breweri Variation H L L M M "L L Juncus parrvi Association  L L L L L M Antennaria lanata Association H L L M L L M Phyllodoce - Antennaria Associ ati on Phyllodoce - Antennaria Variation H L L M L L M Antennaria - Vaccinium Variation H L L M M L M Picea engelmannii Associ ati nn H L L M H L L Abies lasiocarpa Associ ati nn H L L L M M L Abies - Picea - Vaccinium Association hi L M L L L M Abies - Valeriana Association H L L L L M ft| Carex spectabilis Association H L L L H M H Valeriana - Castllleja Associ ati on Valeriana - Castilleia Variation H L L L M M M Trollius laxus Variation M L L M H H H Carex nigricans Associ ati on Carex - Polytrichadelphus Variation M M L M L L H Juncus - Carex - Drepanocladus Variation L H H H H H M Polytrichum norvegicum Association H L H M L L ft] Drepanocladus exannulatus AssnMatinn (_ H L H L i M L = low, III = medium, H = high See Appendix 4 for class limits 155 latter is included even though its hygrotope is rated as sub-xeric). Among the general environmental factors, exposure, slope, erosion, mineral soil and rock vary among the communities. Altitude is generally medium (except for the Antennaria - Sibbal-j-dia - Salix Variation), relief varies from straight to convex, wind is strong to very strong (in the Antennaria - Sibbaldia Association), humus is medium to low, and soil depth is medium to low. Among the physical and chemical soil factors, calcium, sodium, cation exchange capacity, organic matter, nitrogen and phosphorus are variable among the communities. Sand is generally high, while silt and clay are low (the exception in all cases being the Carex phaeocephala Variation). The pH ranges from medium in the Antennaria - Sibbaldia Association to low in the other communities. Magnesium is low, as is potassium (with the exception of the Juniperus communis Association). The mesic group consists of six communities: the Antennaria  lanata Association, both variations of the Phyllodoce - Antenn aria Association (one of which is submesic), the Picea engel mannii Association (rated as submesic), the Abies lasiocarpa Association and the Abies - Picea - Vaccinium Association. Among the general environmental factors, exposure, slope and soil depth are variable. Altitude ranges from high to medium, and relief is generally straight. Wind is strong in the tree island communities and moderate in the others. Erosion varies from none to slight. Humus is medium in the tree island commun ities and high in the other two. Mineral soil is low, except for the Antennaria lanata Association, and rock is low to 156 medium (in the Picea and Abies Associations). Among the physical and chemical soil factors, calcium, cation exchange capacity, organic matter, nitrogen and phosphorus vary among the communit ies. Sand is high, while silt and clay are low (medium clay in the Abies - Picea - Vaccinium Association). The pH ranges from low in the Abies lasiocarpa and Abies - Picea - Vaccinium Assoc iations to medium in the other communities. Magnesium is generally low (the exception being the Abies lasiocarpa Associa tion). Sodium is medium and potassium is low (exceptions for both being the Picea and Abies Associations). The hygric group contains five communities: the Abies -Valeriana Association (rated as subhygric), the Carex spectabilis Association, both variations of the Valeriana - Castilleja Assoc iation, and the Carex - Polytrichadelphus Variation of the Carex  nigricans Association. Altitude, exposure, slope and soil depth are the general environmental factors which vary among the communities. Relief ranges from straight to concave. Wind is moderate in the Abies - Valeriana and Carex spectabilis Assoc iations, and slight in the others. There is no evidence of erosion in any of these communities. Humus is high (except for the Abies - Valeriana Association), mineral soil is low (the exception being the Carex spectabilis Association), and rock is low. Calcium, magnesium, potassium and cation exchange capacity arewthe soil factors which are variable among the communities. Sand ranges from high to. medium, while silt and clay are low (with the exception of silt in the Carex - Polytrichadelphus Variation). The pH ranges from low to medium. Sodium ranges 157 from high to medium. Organic matter is high in the Valeriana -Castille,1a Association, and medium ln the other communities. Nitrogen is generally high (except for the Carex spectabilis Association), and phosphorus is medium (the exception being the Valeriana - Castilleja Variation). Three communities make up the subhydric group: the Juncus -Carex - Drepanocladus Variation of the Carex nigricans Associat ion, the Polytrichum norvegicum Association and the Drepanocladus  exannulatus Association. Among the general environmental factors, altitude is variable. Exposure is generally neutral (except for the Juncus - Carex - Drepanocladus Variation), slope is low, relief is straight to concave, wind is slight, and there is no erosion. Humus is high, mineral soil and rock are low, and soil depth is medium (the exception in all cases being the Polytrichum  norvegicum Association). Among the physical and chemical soil factors, sand is low, silt is high and pH is high (the exception in all cases being the Polytrichum norvegicum Association). Clay is high, while organic matter and nitrogen are low (with the exception of the Drepanocladus exannulatus Association). Calcium, magnesium and potassium are all low (except for the Juncus - Carex - Drepanocladus Variation). Sodium is medium, while cation exchange capacity and phosphorus are low. The two variations of the Carex nigricans Association were placed in different hygrotope groups in the above discussion. Prom Tables 77 and 78, it can be seen that these communities do differ from one another mainly on the basis of physical and chemical soil factors. This does not mean that the classification of these communities should be altered, but merely indicates environmental differences between them (which is helpful in sub dividing an association). It is interesting to note that the variations of the Valeriana - Castilleja Association also differ mainly due to soil factors. However, the variations of the Phyllodoce - Antennaria Association and the Antennaria - Sibbal dia Association differ equally in respect to both general envir onmental factors and physical and chemical soil factors. A one-way analysis of variance was done for each environ mental variable to determine which factors are significantly different between communities. The P values are presented in Table 8l, Appendix 5. Altitude, exposure, slope, wind, erosion, hygrotope, humus, rock, soil depth, sand, silt, clay, pH, magnesium, sodium, potassium, cation exchange capacity, organic matter and nitrogen are all significant at the 1% level. Mineral soil, calcium and phosphorus are significant at the 5$ level. Relief is not significant. In order to distinguish the commun ities which are significantly different on the basis of each environmental variable, Duncan's New Multiple Range Test was done, a test which has proved to be a very useful tool. The results are shown in Table 82, Appendix 5. Based on this table, as well as Tables 77 and 78, the communities are discussed below, mentioning (in order of importance) the group of envir onmental factors which are significant in differentiating each community from all others. The Juniperus communis Association is best differentiated by its low humus, high rock, xeric hygrotope and high slope. It 159 is somewhat less differentiated by its strong wind, high cation exchange capacity and low pH. In the Antennaria - Sibbaldia Association, the Antennaria -Sibbaldia - Sallx Variation is differentiated by its high rock, strong erosion, xeric hygrotope, low humus and very strong wind. Other less Important factors are its medium pH, low nitrogen, low organic matter and high sand. The Carex phaeocephala Varia tion is best separated by its medium rock, medium humus, xeric hygrotope and very strong wind. Another less important factor is its medium sand. The Carex brewer! Variation is distinguished by its xeric hygrotope, strong erosion and very strong wind, and, to a lesser degree, by its medium pH and medium humus. The Juncus parryi Association is differentiated by its sub-xeric hygrotope, strong wind and medium humus, and, to a lesser degree, by its medium rock and low pH. The Antennaria lanata Association is best separated by its medium mineral soil and mesic hygrotope, and less by its high soil depth, medium pH, low rock and moderate wind. In the Phyllodoce - Antennaria Association, the Phyllodoce-Antennaria Variation is best distinguished by its mesic hygro tope and, to a lesser degree, by Its high humus, moderate wind, lack of erosion and low rock. The Antennaria - Vaccinium Variation is best differentiated by its submesic hygrotope, and less by its high humus, low rock and moderate wind. The Picea engelmannii Association is separated by its sub mesic hygrotope and, to a lower degree, by Its medium humus, medium rock and strong wind. The Abies lasiocarpa Association is distinguished by its high cation exchange capacity, mesic hygrotope, medium humus and strong wind, and less by its medium rock, low pH and high organic matter. The Abies - Picea - Vaccinium Association is best differ entiated by its mesic hygrotope and medium humus and, to a less er degree, by its strong wind and low pH. The Abies - Valeriana Association is separated by its sub hygric hygrotope and medium humus, and less by its moderate wind, low altitude and low pH. The Carex spectabilis Association is best distinguished by its hygric hygrotope, and, to a lower degree, by its high humus, low rock, moderate wind and low pH. In the Valeriana - Castille,ja Association, the Valeriana -Castilleja Variation is differentiated by its hygric hygrotope, and less by its high humus, slight wind, low rock and low pH. The Trollius laxus Variation is separated by its high magnesium, high potassium, hygric hygrotope and high humus, and, to a lesser degree, by its slight wind and low rock. In the Carex nigricans Association, the Carex - Polytrich ade lphus Variation is distinguished by its hygric hygrotope, slight wind and high humus, and less by its low rock, medium silt and medium sand. The Juncus - Carex - Drepanocladus Variation is best differentiated by its high pH, low sand, high clay and subhydric hygrotope. It is less well differentiated by its high humus, slight wind, high silt, low organic matter and low rock. 161 The Polytrichum norvegicum Association is best separated by its high clay, subhydric hygrotope, medium rock, neutral exposure and medium humus. Other less important factors are its low slope, slight wind and low organic matter. The Drepanocladus exannulatus Association is best distin guished by its subhydric hygrotope and high humus, and less well differentiated by its high pH, slight wind, low rock, high silt and low sand. Prom the above discussion, it appears that general environ mental factors are more significant in distinguishing the commun ities than are physical and chemical soil properties. Except ions to this are the Trollius laxus Variation, the Juncus - Carex-Drepanocladus Variation, the Abies lasiocarpa Association and, to some extent, the Polytrichum norvegicum Association. Fonda and Bliss (1969) found that soil fertility levels had no effect on community type distribution. Among the general environmental factors, hygrotope is the only one which constantly contributes highly to the differentiation of the communities. This fact substantiates the statement made earlier that hygrotope is the most important factor in delimiting the various communities. B. Soil Moisture Soil moisture was studied in detail because of the prev iously-mentioned fact that hygrotope is the most important factor in distinguishing the communities. Two main aspects are discussed below - available water and actual field moisture values. Available water is considered to be important because it is the only portion of the water supply which is actually 162 available for plant growth. The primary importance in discuss ing the actual field moisture values is to see whether this value is less than the permanent wilting percentage, which would cause soil moisture stress to the plants. Tables 79 and 80 present soil moisture percentages for a selected number of communities in the alpine and subalpine park land areas. As expected, available water generally decreases with depth (Fonda and Bliss, 1969) in both the alpine and subalpine commun ities. However, available water in the Valeriana - Castilleja Variation in the subalpine parkland is approximately the same in the A and C horizons. In the Phyllodoce - Antennaria Variat ion, available water increases with depth in the subalpine park land. Available water also increases with depth in the Carex -Polytrichadelphus Variation, in both the alpine and subalpine areas. The amount of available water in the surface and bottom horizons is similar in both alpine and subalpine communities of the Juncus parryi Association. For the Phyllodoce - Antennaria Variation, the surface horizon has somewhat more available water in the alpine site, whereas the bottom horizon has higher available water in the subalpine site. In the Abies - Picea -Vaccinium Association, the surface horizon has similar quantit ies of available water in both the alpine and subalpine areas. However, both the B and C horizons have more available water in the subalpine site. Since trees have deep rooting systems, the greater amount of water available at depth may be an explanation Table 79 Soil Moisture Percentages for Selected Alpine Communities (Aver of 2) Community Horizon 1/3 atm. ' 15 atm. Available Horizon July 6/69 July 18/69 Aug.3/69 Aug.29/69 Antennaria - Sibbaldia Associ ati on Antennaria - Sibbaldia - Salix Variation A 39.1 24.1 15.0 A 55.7 37.4 12.1 6.6 (Plot 22 - A) 26.7 C 11.1 15.6 C 60.6 38.5 24.3 16.5 Juncus parryi Association A 35.6 20.6 15.0 A 55.2 33.3 19.2 18.8 (Plot 12 - A) 29.3 15.2 B 14.1 B 44.6 75.9 31.8 23.2 C 16.7 10.5 6-2 B + C 36.5 46.2 17.8 25.4 Antennaria lanata Association A 67.9 31.2 36.7 . A 169.6 115.9 99.1 61.1 (Plot 4 - A) 65.2 22.2 43.0 B 8 67.9 40.8 60.7 33.1 C 13.9 6.3 7.6 C 90.1 101.1 44.6 39.0 Phyllodoce - Antennaria Association Phyllodoce - Antennaria Variation A 49.2 21.2 28.0 A 95.1 75.2 58.9 41.3 (Plot 5 - A) 41.4 12.B 28.6 B B 78.8 67.1 56.4 42.2 C 23.1 14.9 8 + C. 74.9 56.8 48.8 32.2 Abies - Picea - Vaccinium Association A 59.4 30.9 28.5 A 53.1 89.6 12.3 14.4 (Plot 54 - LA) 46.5 26.0 20.5 B B 49.8 43.4 25.4 27.7 C 17.3 10.0 7.3 C 58.2 61.1 25.1 27.1 Valeriana - Castilleia Association Valeriana - Castilleia Variation A 54.1 45.5 8.6 A1 245.5 182.3 27.9 47.8 (Plot 38 - LA) A2 178.0 115.0 63.0 31.7 Carex niqricans Association Carex - Polytrichadelphus Variation A 70.2 54.3 15.9 A 215.7 277.5 110.6 52.5 (Plot 2 - A) 62.5 23.0 -B 39.5 B 25.2 120.3 33.1 32.7 ON Table 80 Soil Moisture Percentages for Selected Subalpine Communities Community Horizon 1/3 tef -of; \l atm. Available Horizon July 6/69 July 18/69 Aug. 3/69 Auq.29/ Juncus parrvi Association (Plot 59) A 48.6 32.0 16.6 A 70.5 25.5 9.6 10.7 C 21.7 15.7 6.0 C 13.7 16.8 - 8.4 Phyllodoce - Antennaria Association Phyllodoce - Antennaria Variation (Plot 57) A 48.1 29.0 19.1 A 73.2 79*3 53^ 47.0 C 39.0 13.2 25.8 C1 C2 90.1 78.7 74.8 48.0 63.3 32.2 48.7 Abies - Picea - Vaccinium Association (Plot 61) A 66.4 37.6 28.8 A 110.6 86.3 26.6 21.2 B 58.3 21.8 36.5 J3.+ C 79.9 97.0 52.7 37.0 C 34.2 14.8 19.4 C 39 .4 65.9 - 26.3 Valeriana - Castilleia Association Valeriana - Castilleia Variation (Plot 60) A 75.1 64.6 10.5 A 200.0 303.9 257.9 223.1 C 23.9 13.3 10.6 C 138.4 54.8 34.5 108.4 Carex niqricans Association Carex - Polytrichadelphus Variation (Plot 58) A1 36.8 20.4 16.4 A1 60.5 59.0 17.6 14.1 A2 34.2 14.3 19.9 A2 49.7 51.1 42.6 30.1 C 32.9 14.1 18.8 A3 48.7 64.6 43.3 38.7 ON 165 for the better growth of trees in the subalpine parkland. In a study by Patten (1963), availability of water was believed to be associated with the most common limiting factors for tree growth and establishment in the particular study area in Montana. It was further stated that if the availability of water were increased, the forested area would expand at the expense of the non-forested areas. Available water in the surface horizon of the Valeriana - Castilleja Variation is similar in the alpine and subalpine parkland areas. For the Carex - Polytrichadelphus Variation, the surface horizon has similar amounts of available water in the alpine and subalpine sites; however, there is much more available water in the subsurface horizon in the alpine site. The actual field moisture values for each community are dis cussed below. In general, moisture decreased throughout the summer. For the Antennaria - Sibbaldia - Salix Variation, the surface horizon fell below permanent wilting percentage (PWP) from the beginning of August onward, while the subsurface horizon remained above wilting percentage. In the Juncus parryi Association at the alpine site, the surface horizon fell below PWP at the beginning of August, while the subsurface horizons always remained above field capacity. At the subalpine site, the surface horizon dried out below PWP from the middle of July, and the subsurface horizon remained at or below wilting percent age for most of the summer. The Antennaria lanata Association and the Phyllodoce - Antennaria Variation (in both the alpine and subalpine areas) generally remained above field capacity throughout the summer in the surface and subsurface horizons. In the Abies - Picea - Vaccinium Association, the surface horizon fell below PWP from the beginning of August onward, in both the alpine and subalpine sites. The C horizon at both sites always remained above PWP. However, the B horizon in the alpine area fell below wilting percentage for a period in August. Both surface and subsurface horizons remained above field capacity throughout the summer ln the subalpine Valeriana - Castille,1a Variation, due to the constant supply of seepage water. In the alpine area, the surface horizon dried out below PWP during a period in August, there being less seepage on an alpine slope than on a subalpine one. In the Carex - Polytrichadelphus Variation, the subsurface horizons remained above PWP through out the summer, at both the alpine and subalpine sites. At the subalpine site, the surface horizon fell below wilting percent age at the beginning of August, whereas at the alpine site this did not occur until the end of August. Thus, certain communities in the study area undergo soil moisture stress. This has been reported in a number of alpine areas (Klikoff, 1965; Johnson and Billings, 1962; Billings and Bliss, 1959). In contrast, Bliss (1966) on Mt. Washington, New Hampshire, and Bamberg and Major (1968), working in Montana, found soil moisture not to be critical. C. Topographic-Altitudinal Relationships In this section, the communities are discussed according to decreasing altitude within the topographic categories of ridges, slopes and depressions. This arrangement is essentially 167 a reordering of the sequence presented in section 4, in order to show the relationships of the communities from a different aspect. In general, snow cover would increase from ridges, which are wind-blown, to depressions with the greatest accumu lation of snow. In the alpine area, the Antennaria - Sibbaldia Association occurs on the highest ridges. According to Krajina (personal communication), the presence of Sibbaldia procumbens indicates that there is heavy snow cover on these ridges (perhaps as cornices), contrary to the general snow-free condition existing on ridge tops. This community is prabably subhygric at the beginning of the vegetative season, becoming xeric at the end of the vegetative season. The Picea engelmannii and Abies  lasiocarpa Associations also occur on ridges in the alpine area, but in mesic habitats. These three associations are not present in the subalpine parkland. The Abies - Picea - Vaccinium Assoc iation, which is a mesic community, occurs on lower ridges in the alpine and low alpine areas, and on high ridges in the sub alpine parkland. This association is, of course, much more widespread in the subalpine parkland. Seven communities occur on slopes. The Antennaria lanata Association occurs on very gentle slopes ln the alpine area, but is lacking in the subalpine parkland. It is a mesic community. The hummocky terrain found in this association may be the result of frost activity. The Phyllodoce - Antennaria Association, also mesic, is found on medium slopes in both the alpine and subalpine parkland areas. The Juncus parryi Association occurs on somewhat steeper slopes than the Phyllodoce - Antennaria Association. It differs considerably from the latter in being subxeric in hygrotope, and would have shorter snow duration. It occurs in both the alpine and subalpine parkland areas, but is more prevalent in the alpine area. The Juniperus communis Assoc iation, a xeric community, is found on very steep slopes in the low alpine and alpine areas only. The Carex spectabilis Assoc iation is best represented in the low alpine area on slopes with temporary seepage; it is practically lacking in the subalpine parkland. It is a hygric community. The Valeriana - Castille,1a Association, also hygric, occurs on seepage slopes rich in nutrients in the low alpine and subalpine parkland areas. This association is most frequent in the subalpine parkland. The subhygric Abies - Valeriana Association occurs also on seepage slopes in the subalpine parkland. This community is present on the same slopes as, and adjacent to, the Valeriana - Castllleja Association, but is much less frequent. The Carex nigricans Association, rated as hygric to sub hydric, is found in depressions with a long duration of snow. This community is common in the alpine, low alpine and subalpine parkland areas. The Polytrichum norvegicum Association, which is subhydric, occurs in temporary ponds in the alpine area. It has the longest snow duration of all the communities. There is a sorting of rocks in one plot, which may indicate the existence of frost action. A similar habitat in the subalpine parkland is occupied by the subhydric Drepanocladus exannulatus Association. In this case, the pond dries up at a later date than in the 169 Polytrichum norvegicum Association. D. Soil Types and Plant Communities In the community descriptions (section 4), the soils assoc iated with each community were mentioned. Very few communities are confined to one soil type. In this section, the alternative approach is followed: that is, the communities associated with each soil type are presented. None of the soil great groups and only three subgroups are restricted to a particular commun ity. The use of both these approaches is helpful in providing more information about the interrelationships of soils and veget ation . Alpine Dystric Brunisols are found in eight communities, mainly in the Phyllodoce - Antennaria Association (both variat ions) and in the Abies - Valeriana Association. Other commun ities with this soil type are the Carex - Polytrichadelphus Variation of the Carex nigricans Association, the Carex spect abilis Association, the Antennaria lanata Association, the Juncus parryi Association and the Antennaria - Sibbaldia - Sallx Variation of the Antennaria - Sibbaldia Association. Orthic Regosols occur in twelve communities, predominantly the Juniperus communis Association, the Antennaria - Sibbaldia Association (all three variations), the Valeriana - Castilleja Variation of the Valeriana - Castllleja Association, and the Juncus parryi Association. Less frequent occurrences are with the Carex spectabilis Association, the Trollius laxus Variation of the Valeriana - Castllleja Association, the Carex - Poly trichadelphus Variation of the Carex nigricans Association, the Phyllodoce - Antennaria Variation of the Phyllodoce - Antennaria Association, the Abies lasiocarpa Association and the Picea  engelmannii Association. In the Podzolic Order, Sombric Humo-Ferric Podzols are associated with six communities, mainly the Phyllodoce -Antennaria Variation of the Phyllodoce - Antennaria Association, and the Juncus parryi Association. The other communities are the Antennaria - Sibbaldia Association (Antennaria - Sibbaldia -Salix Variation and Carex breweri Variation), the Antennaria  lanata Association, and the Polytrichum norvegicum Association. Sombric Ferro-Humic Podzols occur mainly In the Abies - Picea -Vaccinium Association and the Juncus parryi Association. Other occurrences are in the Carex spectabilis Association, the Phyllodoce - Antennaria Variation of the Phyllodoce - Antennaria Association, and the Carex - Polytrichadelphus Variation of the Carex nigricans Association. Of infrequent occurrence are Mini Ferro-Humic Podzols, associated with the Antennaria lanata Assoc iation and the Abies laslocarpa Association, and an Orthic Humic Podzol found in the Abies - Picea - Vaccinium Association. In the Gleysolic Order, the predominant soil type is the Rego Humic Gleysol, which occurs mainly in the Valeriana -Castille,ja Association (both variations) and the Carex nigricans Association (both variations). It is also found with the Drep anocladus exannulatus Association and the Polytrichum norvegicum Association. The other soil types in this order occur very infrequently. The, Fera Humic Gleysol is associated with the Valeriana - Castilleja Variation of the Valeriana - Castilleja 171 Association, and the Carex - Polytrichadelphus Variation of the Carex nigricans Association. The Orthic Humic Gleysol and Rego Gleysol are both found in the Trollius laxus Variation of the Valeriana - Castilleja Association. Soils and vegetation have been considered as dependent variables, both depending on the same group of ecosystem factors, according to the following equation (Jenny, 1941; Major, 1951; Crocker, 1952): V and S = f (cl, o, r, p, t) where V is veget ation, S is soil, and the factors are climate, organisms, relief, parent material and time. Thus soils and vegetation are not causally related to each other. A given set of environmental factors produces a certain vegetation type and a soil type. It is interesting to note that the vegetation types and soil types found in the present study are not very closely related. The Canadian soil classification scheme for alpine soils, as used here, is essentially tentative, as very little work has previously been done In alpine regions. It is the name of the soil, as such, which is Important, but the processes which are acting to produce a given set of horizons which is of signif icance. Unfortunately, very little is yet known about soil genesis in alpine environments. One reason for the lack of correlation between vegetation and soils may be the relatively young age of the area, not enough time having elapsed since glaciation for the maximum development of the soils. Another explanation may be the fact that physical environmental factors (such as low temperature and frost activity) can reduce the effect of the time element, thus slowing the development of the 172 soils. It is also important to realize that the vegetation and soils were compared at different levels in their respective classification systems. The vegetation unit used is the assoc iation (or variation), which is very specific, while the soil subgroup is a more generalized abstract category. A much closer correlation between soil types and vegetation types would be produced if the plant communities were compared with the soil series, since the series is a specific unit of the landscape. However, such a comparison was not possible, as no detailed soil map was available for the study area. 173 '8. Vegetation Zonation Throughout this thesis, reference has been made to alpine, low alpine and subalpine parkland areas. It is necessary now to explain how these areas fit into an altitudinal zonation scheme. The lowest area Is the subalpine parkland, ranging ln altitude from ca. 7100 to 7^00 feet. This area is considered as the upper part of the interior Engelmann Spruce - Subalpine Fir Zone. This zone has been divided into three geographic subzones, within which parkland areas occur at the higher alti tudes (Krajina, 1969). For the coastal subalpine zone, the parkland area has been described as a subzone of the Mountain Hemlock Zone (Krajina, 1965). The low alpine (ca. 7^00-7500 feet) and the alpine (ca. 7500-7600 feet) areas comprise the Alpine Zone (after Krajina, 1965). The timberline vegetation is composed of the subalpine parkland and parts of the low alpine area, and is essentially a transition area or ecotone between the closed subalpine forest and the alpine zone. Since there is such a small elevational difference between the subalpine parkland and the summit of the mountain, it may be suggested that the entire study area belongs to the Engel mann Spruce - Subalpine Fir Zone. According to Krajina (1959), the Alpine Zone in the southeast of British Columbia extends above 7500 feet. This would place the summit of Big White in the alpine zone. An examination of species listed as characteristic for the alpine and/or interior subalpine zones by Krajina (1959) revealed 174 the following information about the flora of Big White: 56/73 vascular plants and 33/87 bryophytes and lichens on Big White are characteristic of the alpine zone; 26 vascular plants and 10 bryophytes and lichens are characteristic of the interior subalpine zone. Upon subtracting the number of species listed as characteristic for both the alpine and subalpine zones (18 vascular plants, and 8 bryophytes and lichens), the revised figures are as follows: 38 vascular plants and 25 bryophytes and lichens are characteristic of the alpine zone; 8 vascular plants and 2 bryophytes and lichens are characteristic of the subalpine zone. Therefore, floristically, the study area belongs to the alpine zone. This conclusion can be considered valid, since it is based on a list of plants characteristic of the alpine zone as found throughout British Columbia. However, a comparison can not be made from one geographic area, such as the Rocky Mountains or the coast, to another. As Krajina (1959) stated, "there are possibly several alpine zones which could be separated on the basis of their phytogeographic and macroclimatic characteristics." The alpine zone has been generally defined as the area above timberline. However, timberline itself is variously interpreted as being the elevation of the forest line (the upper edge of continuous forest), the tree line (altitude of the highest stunted tree), or a point midway across the transition zone between forest and alpine tundra (Daubenmlre, 1955). The definitive empirical climatic data which are used by Krajina (1965) to determine the alpine zone are the monthly mean 175 temperatures being below 50°P throughout the year (after Koppen, 1936). Unfortunately, there are no detailed climatic measure ments from the study area. However, climatic data from Old Glory Mountain, to the south of Big White, indicate that its summit (7700 feet) belongs to the alpine zone. This suggests that the summit of Big White will also climatically belong to the alpine zone. Much of the area studied is located in the transition area between the forest and alpine regions, and is called timberline vegetation. However, it is concluded that the upper part of Big White Mountain constitutes the Alpine Zone, although it is certainly not as well developed as it is in the coastal area or in the Rocky Mountains. As there are no other detailed veget ation studies in alpine areas of interior British Columbia for comparison, no generalizations can be drawn from the present study as to the exact characterization of the interior alpine zone. Much further work thus needs to be done in this neglected area of plant ecology in British Columbia. 176 9. Summary and Conclusions The purposes of this research were to obtain data on veget ation and environment in an alpine-timberllne area, to produce an ecosystematic classification of the vegetation, and to deter mine the environmental factors important in the differentiation of the plant communities. The main results of this study are summarized below: (l) Fourteen plant associations, with nine variations, are distinguished and described along a general gradient of Increas ing moisture. The communities are compared with those described in other alpine and subalpine areas. The Juniperus communis Association occurs over rock outcrops on ridges and slopes in the alpine and low alpine areas. The Antennaria lanata - Sibbaldia procumbens Association occurs on ridge tops, primarily in the alpine area. The assoc iation is subdivided intoothree variations: Antennaria lanata -Sibbaldia procumbens - Salix cascadensis Variation, Carex phaeo cephala Variation, and Carex breweri Variation. The Juncus parryi Association occurs on south-facing slopes In the alpine and low alpine areas. It is less well developed in the subalpine parkland, occurring there on slopes and ridges with a southern exposure. The Antennaria lanata Association occurs at the base of slopes, on ridges and on slopes in the alpine and low alpine areas. The Phyllodoce empetriformis - Antennaria lanata Association occurs mainly on slopes In the alpine, low alpine and subalpine 177 parkland areas. The association is divided into two variations: Phyllodoce empetriformis - Antennaria lanata Variation, and Antennaria lanata - Vaccinium scoparium Variation. The Picea engelmannii Association, represented only by one plot, occurs on a ridge in the alpine area. The Abies laslocarpa Association also occurs on ridges in the alpine area. The Abies laslocarpa - Picea engelmannii - Vaccinium scop arium Association occurs mainly on ridges in the alpine, low alpine and subalpine parkland areas. The Abies lasiocarpa - Valeriana sitchensis Association occurs on seepage slopes in the subalpine parkland. The Carex spectabilis Association occurs on slopes with temporary seepage, mainly in the alpine and low alpine areas. The Valeriana sitchensis - Castilleja elmeri Association occurs on seepage slopes in the subalpine parkland and, less frequently, in the alpine and low alpine areas. The association is divided into two variations: Valeriana sitchensis - Castill eja elmeri Variation, and Trollius laxus Variation. The Carex nigricans Association occurs in snow basins, depressions and temporary ponds in the alpine, low alpine and subalpine parkland areas. The association is divided into two variations: Carex nigricans - Polytrichadelphus lyallii Variat ion, and Juncus mertensianus - Carex nigricans - Drepanocladus  exannulatus Variation. The Polytrichum norvegicum Association occurs in temporary ponds in the alpine area. 178 The Drepanocladus exannulatus Association, represented by only one plot, occurs as a narrow band around the edge of a temporary pond in the subalpine parkland. (2) The soils are classified according to the Canadian system of soil classification (Canada Soil Survey Committee, 1970). The orders and soil types represented in the study area are: Brunisolic - Alpine Dystric Brunisol; Regosolic - Orthic Regosol; Podzolic - Sombric Humo-Ferric Podzol, Sombric Ferro-Humic Podzol, Mini Ferro-Humic Podzol, and Orthic Humic Podzol; Gley-solic - Rego Humic Gleysol, Fera Humic Gleysol, Orthic Humic Gleysol, and Rego Gleysol. The communities associated with each soil type are presented in detail, with a discussion on the lack of close correlation between soil types and vegetation types. (3) The soils are generally shallow, with weak horizon develop ment (excluding the podzols). Soil development appears to be proceeding slowly. Important chemical properties are the acidic pH, narrow carbon:nitrogen ratios, low cation exchange capacities, and very low amounts of exchangeable cations. (4) The distribution of the tree species in the area, together with selected diameter, height and age measurements, is discussed. The krummholz growth form of trees occurs on ridges in the alpine area, while trees occur on ridges and seepage slopes In the subalpine parkland. The subalpine trees are much older than those in the alpine area. It is suggested that there has been a recent migration of tree species into the alpine area. There are insufficient data to correlate the migrations with a 179 climatic change. (5) The occurrence of conifer seedlings and shrubs in alpine and timberline communities (exclusive of the sampled tree islands) is presented. No conifer seedlings were observed in the tree island communities. More seedlings were found in the Antennaria-Sibbaldia - Salix community than in any other. (6) A synthesis table including characteristic species and high-presence species for all associations is discussed. In general, both the characteristic species and the high-presence species follow the moisture trend. Species occurring in associations for which they are not characteristic are usually much less Important in those associations. (7) Floristic similarity indices were calculated a) between all plots and b) between all associations and variations. Simi larity matrices are included in the description of the communit ies to show the values of plots within an association. Plots within an association generally have their highest similarities to each other rather than to a plot in another association. Variations of an association show up clearly in the similarity matrix. In comparing the associations and variations with each other, the highest affinities of each community are presented. In general, there is a very low degree of similarity among the communities. It is concluded that the indices of similarity support the classification system. (8) The topographic-altitudinal relationships of the alpine and subalpine communities are presented. (9) The environmental data are summarized for each community as being low, medium or high (in relation only to the present 180 data). The communities are grouped according to hygrotope, and the environmental factors are discussed for each group. (10) A one-way analysis of variance was done for each environ mental variable. All factors are significant at the 1% level except for mineral soil, calcium and phosphorus, which are significant at the 5$ level, and relief, which is not signific ant . (11) Based on Duncan's New Multiple Range Test, the environment al factors which are significant in differentiating each commun ity are outlined. It is concluded that the general environmental factors are more significant in distinguishing the communities than the physical and chemical soil properties. Hygrotope is the most important of the general environmental factors. (12) Soil moisture was studied for a number of communities in the alpine and subalpine parkland areas. Available water gener ally decreases with depth. The amount of available water in the surface and subsurface horizons is compared for corresponding alpine and subalpine communities. A greater amount of available water at depth is proposed as an explanation for the better growth of trees in the subalpine parkland. The actual field moisture values are discussed for each community. A number of the communities fall below permanent wilting percentage for part of the summer, and thus undergo soil moisture stress. (13) A detailed discussion of vegetation zonation is presented. It is concluded that the subalpine parkland area belongs to the Engelmann Spruce - Subalpine Fir Zone, and the alpine and low alpine areas comprise the Alpine Zone. The subalpine parkland and parts of the low alpine area constitute the timberline veget ation. The alpine zone in the study area is not as well devel oped as on the coast or in the Rocky Mountains. 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Dept. of Botany, Univ. of B.C. 31 PP. Tranquillini, W. 1963. Climate and water relations of plants in the sub-alpine region. In_: The Water Relations of Plants. Brit. Ecol. Soc. Symp. 3: 153-167. Tranquillini, W. 1964. The physiology of plants at high altitudes. Ann. Rev. Pl. Physiol. 15: 345-362. United States Department of Agriculture (U.S.D.A.). i960. Soil classification, a comprehensive system - the seventh approx imation. 265 pp. van Ryswyk, A.L. 1969. Forest and alpine soils of southcentral British Columbia. Ph.D. Thesis, Dept. of Agronomy, Wash. State Univ., Pullman, Wash. Diss. Abstr. 30: 2993-B. Wardle, P. 1965. A comparison of alpine timberlines in New Zealand and North America. New Zeal. Jour. Bot. 3: 113-135. Watt, A.S. and E.W. Jones. 1948. The ecology of the Cairngorms I. The environment and the altitudinal zonation of the vegetation. J. Ecol. 36: 283-304. Wilde, S.A. and G.K. Voigt. 1955. Analysis of soils and plants for foresters and horticulturists. J.W. Edwards, Publ. Inc Ann Arbor, Mich. 117 PP. Appendix 1. Checklist of Vascular Plants, Bryophytes and Lichen 195 The checklist of plants is arranged alphabetically. The nomenclature of the vascular plants is according to Taylor (1966), Hitchcock et al. (1955-1969), Hulten (1968) and Moss (1959). The nomenclature of the bryophytes follows Schofield (1968a, 1968b), Crum et al. (1965), Schuster (1969) and Nyholm (1958). The nomenclature of the lichens is after Otto and Ahti (1967), Hale and Culberson (1966) and Bird (1966). Voucher specimens of all plants are deposited in the herbarium of the Department of Botany, University of British Columbia. Vascular Plants Abies lasiocarpa (Hook.) Nutt. Agrostis thurberiana Hitchc. Agrostis variabilis Rydb. Anemone occidentalis S. Wats. Antennaria friesiana (Trautv.) Ekman Antennaria lanata (Hook.) Greene Antennaria umbrinella Rydb. Arenaria capillaris Poir. var. americana (Maguire) Davis Arenaria obtusiloba (Rydb.) Fern. Arnica latifolia Bong. Arnica mollis Hook. Calamagrostis canadensis (Michx.) Beauv. Caltha leptosepala DC. Carex brevipes Boott. Carex'brewer! Boott. Carex nardina Fries 196 Carex nigricans C.A. Meyer Carex phaeocephala Piper Carex pyrenaica Wahl. Carex pyrenaica Wahl. x Carex nigricans C.A. Meyer: intermediate Carex spectabilis Dewey Castilleja elmeri Fernald Castilleja rhexifolia Rydb. Claytonia lanceolata Pursh Deschampsia atropurpurea (Wahlenb.) Schule Dryas octopetala L. var. hookeriana (Juz.) Breit. Epilobium alpinum L. var. clavatum (Trel.) C.L. HItchc. Erigeron peregrinus (Pursh) Greene ssp. callianthemus (Greene) Cronq. Pestuca brachyphylla Schultes Pestuca saximontana Rydb. Gaultheria humifusa (Grah.) Rydb. Gentiana glauca Pall. Habenaria dilatata (Pursh) Hook. Haplopappus lyallii Gray Hieracium gracile Hook. Juncus drummbndii E. Meyer Juncus mertensianus Bong. Juncus parryi Engelm. Juniperus communis L. var. montana Ait. Kalmia polifolia Wang. var. microphylla (Hook.) Rehd. Lupinus latifolius Agardh var. subalpinus (Piper & Robins.) CP. Smith 197 Luzula arcuata (Wahlenb.) Wahlenb. Luzula glabrata (Hoppe) Desv. Luzula glabrata (Hoppe) Desv. x Luzula wahlenbergii Rupr. : intermediate Luzula parviflora (Ehrh.) Desv. Luzula spicata (L.) DC. Luzula wahlenbergii Rupr. Mite 11a breweri A. Gray Pedicularis bracteosa Benth. Phleum alpinum L. Phyllodoce empetrlformis (Smith) Don Picea engelmannii Parry Pinus albicaulis Engelm. Pinus contorta Loudon var. latifolia Engelm. Poa cusickii Vasey var. purpurascens (Beal) C;L. Hitchc. Potentilla diversifolia Lehm. Potentllla drummondii Lehm. Ranunculus eschscholtzii Schlecht. Salix cascadensis Cook. Saxifraga bronchialis L. var. austromontana (Wieg.) G.N. Jones Saxifraga ferruginea Grah. Sedum lanceolatum Torr. Selaginella densa Rydb. var. scopulorum (Maxon) Tryon Senecio triangularis Hook. Sibbaldia procumbens L. Silene parry! (S. Wats.) C.L. Hitchc. & Maguire Solldago multiradiata Ait. 198 Stellaria laeta Richards. Trisetum spicatum (L.) Richter Trollius laxus Salisb. Vaccinium caespitosum Michx. Vaccinium scoparium Leib. Valeriana sitchensis Bong. Veratrum viride Ait. Veronica wormskjoldii R. & S. Bryophytes Anthelia juratzkana (Limpr.) Trevis Aulacomnium palustre (Hedw.) Schwaegr. Barbilophozia barbata (Schmid) Loeske Barbilophozia hatcheri (Evans) Loeske Barbilophozia lycopodioides (Wallr.) Loeske Brachythecium asperrimum (Mitt.) Sull. Brachythecium collinum (Schleich.) B.S.G. Brachythecium curtum Lindb. Brachythecium starkei (Brid.) B.S.G. Bryum bimum (Brid.) Turn. Bryum capillare Hedw. Bryum muehlenbeckii B.S.G. Bryum pseudotriquetrum (Hedw.) Gaertn., Meyer & Scherb. Bryum sp. Cephaloziella rubella (Nees) Douln Cephaloziella subdentata Warnst. Cephaloziella sp. Ceratodon purpureus (Hedw.) Brid. Desmatodon latifolius (Hedw.) Brid. DIcranella sp. Dicranum scoparium Hedw. sens. lat. The specimens closely resemble D. muehlenbeckii B.S.G., but are not identical to it. They are thus retained within D. scoparium Hedw. in the broad sense. Drepanocladus aduncus (Hedw.) Warnst. Drepanocladus exannulatus (B.S.G.) Warnst. Drepanocladus uncinatus (Hedw.) Warnst. Grimmia alpestris (Web. & Mohr.) Nees, Hornsch. & Sturm Hypnum revolutum (Mitt.) Lindb. Kiaeria blyttii (Schlmp.) Broth. Lescuraea baileyi (Best & Grout) Lawt. Lescuraea incurvata (Hedw.) Lawt. Lescuraea radicosa (Mitt.) Monk. Lophozia alpestris (Schleich.) Evans .ti . Lophozia ? kunzeana (Huben.) Evans Lophozia obtusa (Lindb.) Evans Lophozia ? ventricosa (Dicks.) Dumort. Mnium blyttii B.S.G. Orthocaulis floerkii (Web. & Mohr.) Buch Paraleucobryum enerve (Thed.) Loeske Philonotis americana Dism. Pohlia drummondii (C. Mull.) Andr. Pohlia elongata Hedw. Pohlia gracilis (B.S.G.) Lindb. Pohlia nutans (Hedw.) Lindb. 200 Pohlia wahlenbergii (Web. & Mohr.) Andr. Polytrichadelphus lyallii Mitt. Polytrichum formosum Hedw. Polytrichum juniperinum Hedw. Polytrichum norvegicum Hedw. Polytrichum piliferum Hedw. Rhacomitrium canescens (Hedw.) Brid. Rhacomitrium sudeticum (Punck) B.S.G. Scapania subalpina (Nees) Dumort. Scapania undulata (L.) Dumort. Sphagnum nemoreum Scop. ? Tetraplodon mnioldes (Hedw.) B.S.G. Tortula norvegica (Web.) Wahlenb. Tortula ruralis (Hedw.) Gaertn., Meyer & Scherb. Lichens Alectorla americana Mot. Alectoria minuscula Nyl. Cetraria ericetorum Opiz Cetraria islandica (L.) Ach. Cetraria pinastri (Scop.) S. Gray Cetraria subalpina Imsh. Cladonia carneola (Fr.) Pr. .11 . Cladonia chlorophaea (Florke) Spreng. Cladonia coccifera (L.) Willd. Cladonia deformis (L.) Hoffm. Cladonia ecmocyna (Ach.) Nyl. Cladonia macrophyllodes Nyl. 201 Cladonia pleurota (Plbrke) Schaer. Cladonia pyxidata (L.) Hoffm. Cladonia sp. Cornicularia aculeata (Schreb.) Aeh. Icmadadophlla ericetorum (L.) Zahlbr. Lecidea granulosa (Hoffm.) Ach. Lepraria neglecta (Nyl.) Lett. Omphalodiscus virginis (Schaer.) Schol. Parmeliopsis ambigua (Wulf.) Nyl. Parmeliopsis hyperopta (Ach.) Arn. Peltigera canina (L.) Willd. Peltigera canina (L.) Willd. var. rufescens (Weiss) Mudd Peltigera lepidophora (Nyl.) Vain. Peltigera malacea (Ach.) Funck Psoroma hypnorum (Vahl.) S. Gray Rhizocarpon geographicum (L.) DC. Solorina crocea (L.) Ach. Stereocaulon alpinum Laur. Umbilicaria hyperborea (Ach.) Ach. Appendix 2. Soil Types of Big White Mountain classified according to the American, German and World FAO/UNESCO Classifications Canadian American German World FAO/UNESCO Order: Brum sol Ic Order: Inceptisol Suborder: Ochrept Class: Braunerden Order: Cambisol Great Group: Dystric Brunisol Great Group: Dystrochrept Type: Braunerde Soil Unit: Dystric Cambisol Subgroup: Alpine Dystric Brunisol Order: Regosol ic Order: Entisol Class: A-C-BBden Order: Rhegosol Great Group: Regosol Suborder: Orthent Subgroup: Orthic Regosol Great Group: Udorthent or Cryorthent Type: Ranker Order: Gleysolic Class: Gleye Order: Gleysol Great Group: Humic Gleysol Order: Inceptisol Suborder: Aquept Soil Unit: Humic Gleysol Subgroup: Orthic Humic Gleysol Great Group: Humaquept Type: Anmoorgley : Rego Humic Gleysol : Haplaquept : " : Fera Humic Gleysol : Humaquept Order: Entisol : " Great Group: Gleysol Suborder: Aquent Soil Unit: Haplic Gleysol Subgroup: Rego Gleysol Great Group: Hapl aquent Type: Gley o Canadian^ Order: Podzolic Great Group: Humic Podzol Subgroup: Orthic Humic Podzol Great Group: Ferro-Humic Podzol Subgroup: Mini Ferro-Humic Podzol Subgroup: Sombric Ferro-Humic Podzol Great Group: Humo-Ferric Podzol Subgroup: Sombric Humo-Ferric Podzol American Order: Spodosol Suborder: Humod Great Group: Haplohumod Suborder: Orthod Great Group: Haplorthod Suborder: Orthod German^ Class: Podsolc Type: Podsol Type: Podsol Type: Podsol World FAQ/UNESCO^ Order: Podzol Soil Unit: Humic Podzol Soil Unit: Humo-Ferric Podzol Soil Unit: Humo-Ferric Podzol 1 The Canadian system follows Canada Soil Survey Committee (1970). 2 The American system follows U.S.D.A. (1960). . 3 The German system follows Mtickenhausen (1965), after Kubiena (1953). 4 The World FA0/UNESC0 system is taken from Canada Soil Survey Committee (1970). O 205 Appendix 3. Iron and Aluminum Determinations Community Plot No. Antennaria - Sibbaldia Association Antennaria - Sibbaldia -Salix Variation 10 13 Carex brewer! Variation 45 Juncus parryi Association 8 12 30 40 68 Antennaria lanata Association 3 4 Horizon % Fe % Al %> Fe*Al AFe+Al O.M./Fe Bm 0.49 0.83 1.32 0.31 13.02 C 0.26 0.75 1.01 — 7.27 Bf 1.13 0.95 2.08 1.01 4.27 Cgj 0.69 0.38 1.07 — 1.32 Bf 0.74 0.98 1.72 0.89 4.78 Cg 0.32 0.51 0.83 — — 2.41 Bhf 1.06 0.98 2.04 1.22 13.21 C 0.43 0.39 0.82 — 11.49 Bfh 0.87 1.46 2.33 1.13 8.84 Cg 0.39 o.8l 1.20 — 7.36 Bm 0.84 0.98 1.82 0.57 11.60 C 0.63 0.62 1.25 — 8.92 Bhf 1.02 1.06 2.08 0.83 15.93 Bm 0.63 1.32 1.95 0.70 16.43 C 0.41 0.84 1.25 — 17.29 Bfh 0.64 1.38 2.02 0.79 10.83 C 0.51 0.72 1.23 — — 7.4l Bm 0.63 1.06 1.69 O.69 7.67 Cgj 0.41 0.59 1.00 — 5.29 Bfh 0.49 0.96 1.45 O.87 18.22 C 0.21 0.37 O.58 -- 10.33 Community Plot No, Antennaria lanata Association (continued) 16 32 Phyllodoce - Antennaria Association Phyllodoce - Antennaria Variation 5 6 19 27 31 37 57 66 Horizon % Fe % Al % Fe+Al AFe+Al O.M./Fe Bm 0.49 0.68 1.17 0.35 23.24 C 0.43 0.39 0.82 — 5.16 Bhf 1.12 1.46 2.58 1.14 11.71 Cl 0.61 0.83 1.44 6.54 Bf O.85 1.63 2.48 1.88 5.65 Cg 0.21 0.39 0.60 -- 10.48 Bm 0.53 0.71 1.24 O.25 14.02 C 0.49 0.50 0.99 — 6.33 Bm 0.6l 0.92 1.53 — 10.21 Cg 0.84 1.33 2.17 - - 1.20 Bm 0.73 O.89 1.62 0.74 16.63 C 0.42 0.46 0.88 — 5.90 Bfh 0.83 1.37 2.20 1.35 9.29 C 0.26 0.59 0.85 — 9.35 Bf 1.32 1.28 2.60 1.39 3.17 C 0.48 0.73 1.21 — 18.77 Bm 0.6l 0.97 1.58 0.55 18.38 C 0.40 O.63 1.03 — 11.48 Bm 0.38 0.74 1.12 O.65 19.29 C 0.15 0.32 0.47 — 25.13 Community Plot No, Phyllodoce - Antennaria Variation (continued) 72 76 Antennaria - Vaccinium Variation 7 20 23 29 41 Abies lasiocarpa Association 44 Abies - Picea - Vaccinium Association 51 54 jo Al % Fe+Al AFe +A1 O.M./Fe Horizon % Fe Bhf 1.22 C 0.75 Bm 0.48 C 0.31 Bm 0.68 C 0.49 Bm 0.76 C 0.53 Bm 0.81 C 0.47 Bm 0.35 C 0.20 Bm 0.37 Cg 0.41 Bhf 0.74 BC 0.28 1.63 2.85 1.20 1.95 0.84 1.32 0.46 0.77 1.23 1.91 1.46 1.95 0.84 1.60 0.52 1.05 1.31 2U2 1.09 1.56 0.73 1.08 0.41 0.61 1.06 1.43 0.73 1.14 0.93 I.67 0.52 0.80 O.90 10.59 6.51 0.55 18.27 5.45 17.78 12.22 0.55 17-42 9.96 O.56 11.17 5.19 0.47 18.00 IO.85 0.29 26.14 6.37 O.87 15.88 37.36 Bh 0.32 C 0.28 Bhf 1.26 C 0.74 0.48 0.80 0.37 O.65 1.37 2.63 1.06 1.80 0.15 41.56 23.57 O.83 11.21 6.04 Community Plot No. Horizon % Pe % Al • % Fe+Al AFe+Al O.M./Fi Abies - Picea - Vaccinium Association (continued) 61 Bhf Cg 0.72 0.27 0.84 0.45 1.56 0.72 0.84 14.86 20.26 70 Bhf C 0.58 0.32 1.05 0.47 1.63 0.79 0.84 18.28 20.25 Abies - Valeriana Association 56 Bm C 0.47 0.53 1.02 0.74 1.49 1.27 0.22 . 27.19 21.64 62 Bm C 0.63 0.28 0.94 0.69 1.57 0.97 0.60 17.86 16.54 74 Bm C 0.61 O.51 0.53 0.74 1.14 1.25 — 19.85 5.80 Carex spectabilis Association 11 Bm C 0.64 0.73 0.59 0.48 1.23 1.21 0.02 20.19 5.47 14 Bhf C 0.92 0.32 0.78 0.43 1.70 0.75 0.95 14.28 8.44 18 Bm C 0.57 0.31 0.83 0.47 1.40 0.78 0.62 9.63 11.81 Valeriana - Castilleja Association Valeriana - Castilleja Variation 34 Bhfg 0.71 1.38 2.09 0.87 18.04 C 0.39 O.83 1.22 — 15.92 Trollius laxus Variation 24 Bg 0.27 0.63 0.90 0.03 39.70 Cg 0.16 0.71 0.87 II.69 Community Plot No. Carex nigricans Association Carex - Polytrichadelphus Variation 1 2 15 35 71 77 Polytrichum noryeglcum Association 79 Horizon % Fe % Al % Fe+Al AP&*-A1 O.M./Fe Bm 0.55 0.89 1.44 0.08 14.02 Cg 0.43 0.93 1.36 -- 4.21 Bm 0.71 1.26 1.97 0.23 14.10 Cgj 0.64 1.10 1.74 -- 2.61 Ahf 0.70 1.28 1.98 0.95 24.54 Bhf 0.8o 1.70 2.50 1.47 19.85 Bm 0.74 0.99 1.73 0.70 16.55 C 0.39 0.64 1.03 -- 19.23 Bhfg 0.86 1.50 2.36 0.84 14.90 Cg 0.54 0.98 1.52 -- 7.87 Ah 0.43 0.86 1.29 0.41 55.79 Cg 0.57 0.31 0.88 — 10.53 Bml 0.74 1.32 2.06 0.49 10.23 Bm2 0.83 0.94 1.77 0.20 3.37 C 0.51 1.06 1.57 5.73 Bf 0.74 1.32 2.06 1.10 6.07 Cg 0.32 0.64 0.96 -- 5.47 Appendix 4. Class Limits for Environmental Variables 212 The limits chosen are based on the range of values present for each variable. Variable Altitude Slope Humus Mineral Soil Rock Soil Depth Sand Limits H 75OO-76OO ft M 7400-7500 ft L<7400 ft. H 25 -40$ M 10' -25$ L 0--10$ H 66 -100% M 33 -66% L 0--33% H 12' -18$ M 6 -12% L 0--6% H 66 -100% M 33 -66% L 0--33% H 22--26 in. M 17' -21 in. L 12--16 in. H 68--80$ M 54. -67$ L 40 • -53$ Variable Silt Clay PH Ca (me/lOOg.) Mg (me/lOOg.) Na (me/lOOg.) K (me/lOOg.) Limits H 39-^9$ M 28-38$ L 17-27$ H 8-11$ M 4-7$ L 0-3$ H>5.5 M 5.0-5.5 L 4.5-5.0 H>0.8 M 0.5-0.8 L 0.2-0.5 H>0.18 M 0.10-0.18 L 0.02-0.10 H>0.18 M 0.10-0.18 L 0.02-0.10 H>0.25 M 0.14-0.25 L 0.03-0.14 Variable Limits CEC (me/lOOg.) O.M. N P (ppm) H>40 M 25-40 L<25 H 15-20$ M 9-14$ L 3-8$ H>0.5$ M 0.4-0.5$ L<0.4$ H 12-14 M 8-11 Appendix 5. Statistical Analysis 215 Table 8l P Values for Environmental Variables1 Variable F Value ** Altitude 2.42 Exposure 2.63** Slope 4.26** Wind ** 15.70** Relief 0.80 Erosion 8.61** Hygrotope 44.78** Humus 20.44** Mineral soil 1.99* Rock 20.11 ^** Depth of soil 2.58 Sand . ** 3.24 #* Silt 2.59 Clay 4.53** PH 5.94** Ca 1.84* Mg ** 5.33 Na 2.41** K ** 3.99 CEC 3.63 OM 3.86** N 2.98 P 1.84* 1 Community degrees of freedom = 18; Error degrees of freedom b3 * Significant at 5$ level, ** Significant at 1$ level 3 5 „ 5 3 < 3 - - 5 ...... ?. hi j| «. ? 2 3 9 5 .ii . 5 1 3 «. §§5 3" < 3 H 5 5?, rr «, » § 1 5 r- • s *§, 3 5 s ~ 1 2 *!>=-- „ -- „- 2 s *- r- * j. , 3" 5 =-. ! ** Z- . 2 2 2 2! 2&S =-i & J = J§3 - 2 s 3-3-3 3 2. s-= 2 3?-3 3 . I , = = l-l- 1; 3 33 • ' 2 1 ? 5 . I. • 2 .1 I . - c * ^ S !J! h > 2 2 5 !Siin:i Mil «J«. I ll IIII 11 ill! Table 83 Key to Environmental Variables Variable Assigned Number Altitude 1 Exposure 2 Slope 3 Wind 4 Erosion 5 Hygrotope 6 Humus 7 Mineral soil 8 Rock 9 Depth of soil 10 Sand 11 Silt 12 Clay 13 pH 14 Ca 15 Mg 16 Na 17 K 18 CEC 19 OM 20 N 21 P 22 

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