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Biogeoclimatic ecosystem classification of subalpine and alpine plant communities in the Cariboo Mountains. Osorio, Federico G. 2014

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Biogeoclimatic ecosystem classification of subalpine and alpine plant communities in the Cariboo Mountains. by  Federico G. Osorio BSF, The University of British Columbia Vancouver, 2007     A thesis submitted in partial fulfillment of the requirements for the degree of  Master of Science in The faculty of graduate and Postdoctoral studies (Forestry)        The University of British Columbia (Vancouver)  May 2014 © Federico G. Osorio, 2014 ii  Abstract  Terrestrial ecosystems in British Columbia are catalogued using the Biogeoclimatic Ecosystem Classification (BEC). My research is a continuation of the BEC program, specifically aimed at classifying high-elevation ecosystems (alpine and subalpine) of the western Cariboo Mountains of British Columbia. I‟ve included a literature review describing the factors that contribute to the formation of high-elevation plant communities. My results include: five new site series for the Interior Mountain Heather wet cold alpine subzone; b) seven new site series for the Engelmann Spruce Subalpine Fir wet cold parkland subzone; c) two plant orders (one not previously described); d) 5 alliances (2 not previously described), and 19 plant associations (17 not previously described). These results contribute to the description of Site Associations, Site Orders and Site Alliances for the provincial Alpine classification. To develop the classification, I explored the interaction of vegetation with topographic and edaphic variables, and followed phytosociology methods to develop the nomenclature for the plant communities.      iii  Preface  The design, collection and analysis of the data in this thesis, are primarily my own, except for a few graphs, descriptions, and nomenclature, which were created by Will Mackenzie and are given due credit in the text. Also, a substantial number of relevés (credited in text) were collected by various researchers and provided by Will Mackenzie. Ray Coupé, Will Mackenzie and Curtis Bjork, were very helpful in the identification of plant species, but any errors are my own. No peer reviewed publications have arisen from this thesis.  The data collection took place in the late spring, summer, and fall of 2007-2011; however, the results presented in this thesis are primarily from 2007 through 2009.  There is substantial data collected in 2008 through 2011, it is available upon request and I hope it can be used to further the scientific understanding of alpine ecosystems in BC. The thesis is comprised of the main text (divided into five chapters, Introduction through Epilogue) and has two appendices.               iv  Table of Contents Abstract ............................................................................................................................... ii Preface................................................................................................................................ iii Table of contents………………………………………………………………………….iv List of tables ....................................................................................................................... vi List of figures ...................................................................................................................... x Acknowledgements ........................................................................................................... xv Prologue ........................................................................................................................... xvi Chapter 1: Introduction ....................................................................................................... 1 Setting of the alpine and subalpine ecosystems ........................................................... 2 Snow ........................................................................................................................... 5 Soils............................................................................................................................. 8 Nitrogen (N), litter decomposition, and soil microbes ............................................. 11 Mycorrhizae .............................................................................................................. 18 Plant interactions ....................................................................................................... 19 Succession ................................................................................................................. 20 Overall objective, research question and scope of thesis........................................... 23 Objective ................................................................................................................... 23 Research question ..................................................................................................... 24 Scope of thesis .......................................................................................................... 24  v  Chapter 2 Vegetation Classification ................................................................................. 25 Introduction ................................................................................................................ 25 Conceptual delineations of the AT and the Parkland................................................ 33 Objectives and hypotheses ......................................................................................... 35 Methods ..................................................................................................................... 35 Study area.................................................................................................................. 35 Data collection .......................................................................................................... 37 Data analysis ............................................................................................................. 38 Chapter 3 Results .............................................................................................................. 49 Classification ............................................................................................................. 49 Correlations with soil properties ................................................................................ 64 Chapter 4 Discussion ........................................................................................................ 67 Survey of the vegetation above 1900m.a.s.l. in the western Cariboo Mountains ...... 67 Comparison to previously identified plant communities .......................................... 68 Relevance of phytosociology in modern studies ....................................................... 81 Chapter 5 Epilogue ........................................................................................................... 85 References ......................................................................................................................... 87 Appendix A ....................................................................................................................... 97 Appendix B ..................................................................................................................... 154  vi    List of tables  Table 1 Differentiating criteria for BGC units.  a DCS = diagnostic combination of species. May include character, differential and/or companion species. .......................... 29 Table 2 Methods used to answer the questions presented by De Cáceres and Wiser (2012) to help define vegetation types ......................................................................................... 39 Table 3 Activities related to the determination of membership to vegetation types (from (De Cáceres and Wiser 2012) ........................................................................................... 41 Table 4 List of Results available after each step of the data analysis methodology ......... 49 Table 5 Hierarchical relationship between alliances, site associations, site series and TWINSPAN groups in the ESSFwcp. .............................................................................. 61 Table 6 Hierarchical relationship between alliances, site associations, site series and TWINSPAN groups in the IMAwc................................................................................... 62 Table 7 Synopsis of the vegetation units in the classification. ......................................... 63 Table 8 Comparison of plant communities  from Kootenay National Park identified by Achuff (1984a, left column) to those described in this thesis (right column). ................. 69 Table 9 Comparison of plant communities from Mt. Revelstoke and Glacier National parks identified by Achuff (1984b, left column) to those described in this thesis (right column). ............................................................................................................................ 70 Table 10 Comparison of plant communities from Wells Gray Provincial Park (Battle Mountain) identified by Hämet-Ahti (1979, left column) to those described in this thesis (right column). .................................................................................................................. 74  vii  Table 11 Comparison of plant communities from Mt. Revelstoke National Park identified by Landas and Scotter (1974, left column) to those described in this thesis (right column)............................................................................................................................................ 76 Table 12 Pojar et al. (1987) Definitions of diagnostic values of plant species ................. 98 Table 13 Mackenzie 2012 Ecosystem attributes used to differentiate between units of the same biogeoclimatic hierarchical level. ............................................................................ 98 Table 14 Shows the interim table processed from steps 6 through 9 in for the ESSFwcp. ......................................................................................................................................... 102 Table 15 Shows the tabulation of TWINSPAN groups for the IMAwc after step 6.. .... 106 Table 16 Tabulation resulting from step 8 for the IMAwc. . .......................................... 109 Table 17 Tabulation of the alliances represent the final (highest hierarchical level) grouping for the relevés of the IMAwc........................................................................... 121 Table 18 Tabulation of the alliances represent the final (highest hierarchical level) grouping for the relevés of the ESSFwcp. ...................................................................... 122 Table 19 Key to Site Units of the IMAwc. ..................................................................... 130 Table 20 Site Units of the IMAwc (by Will Mackenzie, revised by Federico Osorio). . 131 Table 21 Key to Site Units of the ESSFwcp ................................................................... 139 Table 22 Site Units of the ESSFwcp3............................................................................. 141 Table 23 Principal Component Analysis for the measured edaphic and environmental variables. ......................................................................................................................... 144 Table 24 Eigenvectors for each principal component resulting from edaphic and environmental variables. ................................................................................................. 144  viii  Table 25 Final Communality Estimates (variance accounted for) for the Varimax Factor rotation on 2 the first two Principal Components. .......................................................... 145 Table 26 Factor Structure shows correlations between variables and components.  ...... 145 Table 27 Variance explained by the Principal Components for three selected variables: Org Matter, NH4-N and NO3-N. .................................................................................... 146 Table 28 Correlations in the new Factor Analysis with reduced variables. .................... 146 Table 29 Communality estimates (variance accounted for). .......................................... 146 Table 30 Results from different classifications methods (step 1 from  Classification Methodology) for all relevés (300). ................................................................................ 154 Table 31 Results from different classifications methods (step 1 from  Classification Methodology) for ESSFwcp relevés (190). .................................................................... 156 Table 32 Results from different classifications methods (step 1 from Classification Methodology) for the IMAwc relevés (110). Coloured groups indicate the method selected for further analysis. ........................................................................................... 158 Table 33 Isopam method (criterion Percent Frequency ≥III). ........................................ 160 Table 34 All Plots (300) TWINSPAN (Sørensen‟s  index, pseudospecies cut levels 0 5 10 20 50). ............................................................................................................................. 166 Table 35 TWINSPAN (Whittaker‟s div., pseudospecies cut levels = 0 5 10 20 50). ..... 176 Table 36 TWINSPAN Sørensen‟s Pseudospecies cut levels = 0 5 10 20 50. ................. 178 Table 37 ESSFwcp Pseudospecies cut levels = 0 5 10 20 50 Presence/Absence ........... 181 Table 38 ESSFwcp Isopam. ............................................................................................ 183 Table 39 All plots (190) TWINSPAN Sørensen‟s  0 5 10 30 50. ................................... 187 Table 40 TWINSPAN fidelity (>18) .............................................................................. 189  ix  Table 41 TWINSPAN (Sørensen‟s index, pseudospecies cut levels 0 5 10 20 50). ....... 192 Table 42 TWINSPAN pseudospecies cut levels 0 5 10 20 50 Presence/Absence. ......... 195 Table 43 ISOPAM. ......................................................................................................... 198 Table 44 My relevés (130) TWINSPAN w/merged groups. .................................... 200 Table 45 Raw soil analysis........................................................................................... 203 Table 46 IMAwc site series species list………………………………………………...209 Table 47 ESSFwcp site series species list……………………………………………....213     x  List of figures Figure 1 Relationships of climate variables to vegetation characteristics for alpine ecosystems (Kikvidze et al. 2005). ..................................................................................... 2 Figure 2 Ordination of 128 stands in the alpine zone of the North Cascades  (Douglas and Bliss 1977). ......................................................................................................................... 7 Figure 3 Climatic-edaphic grid showing relationships of the site groups and associations in the parkland Mountain Hemlock (MH) subzones to gradients of snow duration and soil moisture (Klinka and Chourmouzis 2001). ......................................................................... 8 Figure 4 Fluxes of N for the Niwot Ridge, Colorado alpine ecosystem Bowman and Seastedt (2001).................................................................................................................. 11 Figure 5 Conceptual diagram of the effects of alleviation of stress (e.g. low fertility, low pH) on alpine plant communities and plant diversity (2002). .......................................... 13 Figure 6 Conceptual model showing the generalized pattern of plant species self-replacement with low or high N availability (Bowman 2000). ........................................ 14 Figure 7 Plant and microbial 15N tracer uptake, expressed per square meter (Jaeger et al. 1999). ................................................................................................................................ 16 Figure 8 Pie chart showing proportions of major phylogenetic groups represented in a bacterial clone library from spring alpine tundra wet meadow bulk soil (0–20 cm) (Costello and Schmidt 2006). ........................................................................................... 17 Figure 9 Development of arbuscules of the coarse endophyte and shoot phosphorus concentrations in R. adoncus at different dates Mullen and Schmidt (1993). .................. 19 Figure 10 Diagrammatic representation of the main lines of alpine plant succession in xero- and hydroseres (Cox 1933). ..................................................................................... 20  xi  Figure 11 Distribution of the Alpine Tundra in British Columbia (Mackenzie 2006). .... 32 Figure 12 The study area boundary; the purple line delineates the Cariboo Mountains. . 36 Figure 13 Mt. Elsey Pilot Project Area.. ........................................................................... 37 Figure 14 Southern interior Forest Region (in green) ....................................................... 37 Figure 15 The Biogeoclimatic Ecosystem Classification framework. New upper-level site units are highlighted in grey from Mackenzie (2012). ..................................................... 43 Figure 16 Example of the Modified TWINSPAN parameter selection. ........................... 45 Figure 17 Example of the options selection for ISOPAM. ............................................... 46 Figure 18 Detrended Correspondence Analysis (DCA) for 130 relevés. ......................... 52 Figure 19 Canonical Correspondence Analysis for 130 relevés with the aspect overlaid. 53 Figure 20 NMDS for all 300 relevés and 18 TWINSPAN divisions. ............................... 54 Figure 21 Revised BEC classification for part of the Horsefly Watershed, Horsefly Forest District............................................................................................................................... 97 Figure 22 Previous BEC classification for a section of the Horsefly Watershed, Horsefly Forest District.................................................................................................................... 97 Figure 23 Non-metric multidimensional Scaling (NMDS) Ordination of 130 relevés with 18 TWINSPAN groups.  ................................................................................................... 99 Figure 24 Detrended Correspondence Analysis (DCA) for 130 relevés.. ...................... 100 Figure 25 Canonical Correspondence Analysis for 130 relevés with the aspect overlaid........................................................................................................................................... 101 Figure 26 NMDS for all 300 relevés and 18 TWINSPAN divisions. ............................. 102 Figure 27 Principal Component Analysis (PCA) for 130 relevés with TWINSPAN groups reduced from 18 to 7. ...................................................................................................... 111  xii  Figure 28 Nonmetric multidimensional scaling for 130 relevés with TWINSPAN groups reduced from 18 to 7. ...................................................................................................... 111 Figure 29 Detrended Correspondence Analysis for 130 relevés with TWINSPAN groups reduced from 18 to 7. ...................................................................................................... 112 Figure 30 Detrended Correspondence Analysis (Axis 3 and 1) for 130 relevés with TWINSPAN groups reduced from 18 to 7, marked by convex hulls. ............................ 113 Figure 31 Canonical Correspondence Analysis for 130 relevés with TWINSPAN groups reduced from 18 to 7. ...................................................................................................... 114 Figure 323D representation for the nonmetric multidimensional scaling for 130 relevés with TWINSPAN groups reduced from 18 to 7. ............................................................ 114 Figure 33 Detrended Correspondence Analysis for the IMAwc (110 relevés) with 5 TWINSPAN divisions. ................................................................................................... 115 Figure 34 PCA (using the variance/covariance matrix) for the IMAwc (110 relevés) with 5 TWINSPAN divisions. ................................................................................................ 115 Figure 35 NMDS for the  IMAwc (110 relevés) with 5 TWINSPAN divisions. ........... 116 Figure 36 Canonical correspondence analysis for the  IMAwc (110 relevés) with 5 TWINSPAN divisions. ................................................................................................... 117 Figure 373D representation of Canonical Correspondence Analysis for the IMAwc (110 relevés) with 5 TWINSPAN divisions. ........................................................................... 118 Figure 38 Principal Component Analysis for the ESSFwcp (190 relevés) with 5 TWINSPAN divisions. ................................................................................................... 119 Figure 393D representation of the Detrended Correspondence Analysis for the ESSFwcp (190 relevés) with 5 TWINSPAN divisions. .................................................................. 119  xiii  Figure 40 NMDS for the ESSFwcp (190 relevés) with 5 TWINSPAN divisions. ......... 120 Figure 41 Canonical Correspondence Analysis for the ESSFwcp (190 relevés) with 5 TWINSPAN divisions .................................................................................................... 120 Figure 42 NMDS for the 3 alliances of the IMAwc. . .................................................... 123 Figure 43 NMDS for the three alliances of the IMAwc with only dominant species used in the analysis. ................................................................................................................. 123 Figure 44 DCA for the three alliances of the IMAwc. ................................................... 124 Figure 45 NMDS for the 4 alliances of the ESSFwcp. ................................................... 125 Figure 46 CCA for the four alliances of the ESSFwcp. .................................................. 126 Figure 47 Hierarchical Cluster Analysis for 130 relevés based on Sørensen‟s dissimilarity index as distance measure and group average as linkage method. ................................. 127 Figure 48 Two-way Cluster Analysis  for the IMAwc based on Sørensen‟s dissimilarity index as distance measure and group average as linkage method. ................................. 128 Figure 49 Two-way Cluster Analysis for the ESSFwcp based on Sørensen‟s dissimilarity index as distance measure and group average as linkage method. ................................. 129 Figure 50 Edatopic Grid for the IMAwc with Site Series. ............................................. 134 Figure 51 Landscape profile for IMAwc Site Series . .................................................... 135 Figure 52 Edatopic grid for the IMAwc with site associations (site units).. .................. 135 Figure 53 Landscape diagram for the IMAwc with site associations, created by Will Mackenzie. ...................................................................................................................... 136 Figure 54 Edatopic grid for the ESSFwcp with Site Series. ........................................... 136 Figure 55 Landscape diagram for the ESSFwcp Site Series........................................... 137 Figure 56 Edatopic Grid for the ESSFwcp with site associations (site units). ............... 138  xiv  Figure 57 Landscape diagram for the ESSFwcp with site association nomenclature. ... 139 Figure 58 Loading plot for Principal Components 1 and 2 for measured edaphic and environmental variables . ................................................................................................ 144 Figure 59 Oblique Quartimax Factor Rotation for the first 2 Principal Components .... 146 Figure 60 Rotated Factor Pattern (Oblique Quartimax) for the first 2 Principal Components.. .................................................................................................................. 147 Figure 61 CCA for all edaphic variables. ....................................................................... 148 Figure 62 CCA for the four selected variables and their corresponding species. ........... 149 Figure 63 PCA for the three selected variables and their corresponding species. .......... 150 Figure 64 PCA for all edaphic variables axis 2vs1. ........................................................ 151 Figure 65 PCA for all edaphic variables Axis 3 and 1. .................................................. 152 Figure 66 NMDS for all edaphic variables. .................................................................... 153   xv   Acknowledgements   I express deep gratitude to my friends, who supported me throughout my endeavour into Science. The following thesis would not have been possible without the guidance of Ray Coupé, Will Mackenzie, Suzanne Simard, Michael Feller and Gary Bradfield. The research was enabled through the funding provided by the BC Forest Service Southern Interior Research Branch. I owe the inspiration for this project to, my cat (who‟s passing away could only be remedied by a distraction of this magnitude), and the mountains. The most important lessons learnt throughout this project were provided by, Suzanne Simard, Chip and Joanna McKay, Rene Cornu, Bruce Larson, Thane Isert, the Xeni Gwet‟in, Natasha Bush, Austin Spry, Ifor Thomas (R.I.P.), Agathe Bernard, Frederick G. Hill, Aaron Chance, Mark Hartley, Mike Shipp, Andrew Morrell, Chic Sharp, and my parents; all of who - by doleful example -   highlighted the value and fragility of life, which taught me the meaning of strength and resilience.        xvi   Prologue Alpine environments have played a decisive role in my life since my teenage years when, on the rare smog-free days in Mexico City, I was unexplainably drawn to the distant snow-capped mountains that soared above the city. Eventually this fascination grew so strong that I would wake up at pre-dawn to go for bike ride fuelled by the desire to spend a few minutes close to the mountain tops knowing that once there the day would be ok no matter what. This passion grew stronger during my early days at UBC when the unbearable rain and darkness of the Vancouver winters, became worth it, knowing that I could easily reach the alpine during my time-off. Yet it was during my first summer in BC, at Wells Gray Provincial Park, when I realized that the alpine was so important to me that I would guide my life choices based on how I could maximize my time in those environments.  A series of fortunate events led to a proposition by Ray Coupé (former ecologist for the BC Forest Service Research Branch) to spend a few summers in the subalpine and alpine of the Cariboo Mountains to develop a botanical classification of these ecosystems. This classification was to complement the ongoing provincial Biogeoclimatic Ecosystem Classification for the former Cariboo Forest Region and for the Alpine-Tundra ecosystems (led by Will H. Mackenzie, provincial ecologist for the BC Forest Service). Luckily, Suzanne Simard of the Department of Forest and Conservation Sciences, and the Faculty of Graduate Studies of the University of British Columbia, accepted me as their student.     1  Chapter 1: Introduction   Globally, approximately 3% of the terrestrial land area and 4% of all known higher plant species are within the alpine life zone (Körner 2003) and since “nearly half of mankind depends on water supplies originating in mountain catchments, the integrity and functional significance of the upland biota is a key to human welfare and will receive even more attention as water becomes an increasingly limited resource” (Körner 2003).  The Intergovernmental Panel on Climate Change (2007) has also identified alpine and arctic ecosystems as the most susceptible to change as a result of human induced climate change.  The compounding effects of traditional human activities (e.g. recreation and mining) and climate change create a unique situation in which British Columbia‟s high elevation ecosystems are susceptible to unprecedented pressures; yet, it is unclear how such pressures might affect the subalpine and alpine areas of the province. For example, it is commonly understood that the subalpine and alpine will experience an increase in abundance and distribution of vegetation as growing conditions become more favourable. Yet, one analysis showed a 20% increase in peak snow-water equivalents, which would indicate a shorter growing season and thereby a reduction in the abundance of high-elevation vegetation (Pike et al. 2010).  Therefore, classifying the current botanical character of British Columbia‟s high-elevation vegetation will allow for better monitoring, assessment and remediation of these ecosystems.  Understanding the composition of the main primary producers at high elevations might help us understand some of the key components of biological diversity, ecosystem dynamics and our impacts upon them. The literature reviewed was chosen to highlight how, although scientists have been intrigued by high elevation environments for over a century, we are still (especially in North America) at a very early stage in the extent and intensity of our research. Unlike other places (e.g. Europe, Asia, and South America), in which high elevation environments play a direct and key role in the well-being of their societies, in North America they remain primarily places for recreation and exploration. In British Columbia, a strong interest in mineral exploration and in water availability has recently placed greater emphasis on high elevation environments, which combined with ever increasing accessibility and developments in technology, and the imminent threat of  2  climate change, have recently sparked a new wave of scientific interest in the alpine and subalpine regions of the province.  This thesis summarizes the research conducted over 3 years which aimed at describing the composition of plant communities.  The following introduction is a general review of the current understanding of some of the basic biotic and abiotic factors which might influence plant communities. A specific introduction focused on vegetation classification is included in the second chapter.  Setting of the alpine and subalpine ecosystems Price (1981), Körner (2003), and Martin (2001) considered elevation, aspect, relief, but most importantly climate as the main factors that create alpine ecosystems. Martin attributed the alpine character to: high winds, low temperatures, low effective moisture, and short growing seasons. Billing‟s (1973) research added mesotopographic gradients, nearness to glaciers, glacial history, and susceptibility to disturbance as other important conditions that determine the distribution and occurrence of vegetation in alpine ecosystems. Kikvidze et al. (2005) related climatic components (maximum, minimum temperatures and precipitation), in a global study, to the presence and abundance of plants in alpine environments (Figure 1).  However, this generalized model does not include some of the components that play a major role in the characteristic of vegetation in temperate climates, like those in British Columbia.          Figure 1 Relationships of climate variables to vegetation characteristics for alpine ecosystems. R2 values indicate strength of correlations between components (Kikvidze et al. 2005).  At temperate latitudes, distance from the ocean (oceanity vs. continentality) was determined as  a major driving force behind the climate at high elevations (Nagy and Grabherr 2009). The complexities in composition (biotic and abiotic) and in the Figure 1 has been removed due to copyright restrictions. It showed the relationships of climate variables to vegetation characteristics for alpine ecosystems. Original source:  Kikvidze, Z., Pugnaire, F.I., Brooker, R.W., Choler, P., Lortie, C.J., Michalet, R., and Callway, R.M. 2005. Linking patterns and processes in alpine plant communities: A Global Study. Ecology 80[6]:1395-400.   3  distribution of high elevation soils further explained the spatial patterns in high elevation plant communities.  However, most authors agree that characteristics of snow are the strongest limiting factors for the establishment of plants, their growth, reproduction, diversity, and abundance. Since the strength of snow‟s influence on plant communities is one of the most defining characteristics of high elevation environments, I have included a section on snow in this introduction. In the Columbia Mountains the vertical range for trees is limited by late frosts and short growing seasons combined with the “desiccating effect of winds when soil water is deficient” (Cox 1933). However, temperature and growing season can be of minor consideration when soil moisture imposed restrictions in distribution (Cox 1933). Habeck (1987) observed that the distribution of alpine vegetation in the Northern Rocky Mountains was controlled primarily by length of growing season, temperature, soil moisture, and to a lesser degree by wind, evaporation, precipitation, humidity, light, and soil. He considered the dwarfed state of the vegetation indicative of a brief growing season and water deficits caused by frequent low soil temperatures, dry spells, high evaporation rates, intense light, and poor soil conditions. In addition to these factors, the spatial patterns of the treeline seemed to be greatly influenced by microtopography.  Resler (2006) proposed that surface geomorphic features, such as boulders and terraces, created the necessary conditions - primarily by providing wind shelters - for tree seedling establishment and development.  Wind was considered a controlling factor when it accelerated evapotranspiration rates, thereby decreasing soil water (Resler 2006). Wind also played an integral part in the redistribution of snow, making it difficult to determine whether wind-drifted snow or wind-desiccation (Cox 1933) was the primary effect of wind on the alpine and subalpine plant ecosystems. Wind was also indirectly important when it influenced pollination and seed dispersal, and caused mechanical stress. Valentine and Green (1976) suggested that wind erosion might be a key factor in the creation of alpine soils, which in turn may help explain the variability in plant patterns.  An interesting assertion, by Douglas and Bliss (1977), is that “when steep, abrupt changes in species composition occur[ed], [they] creat[ed] a mosaic of plant communities  4  on the landscape. If these gradients [were] gentle, more gradual changes create[d] a continuum” (Douglas and Bliss 1977). The discussion of plant communities vs. continuum is beyond the scope of this thesis, but more insight into this subject can be found in Daubenmire (1966), McIntosh (1967), Whittaker (1967), Austin and Smith (1990), and Callaway (1997).  To further explain the abrupt changes and continuums in the alpine landscape, the biotic conditions, soils, and plant interactions will be discussed below. But first, it must be acknowledged that geology deserves a category of its own as an influence on vegetation patterns in the alpine. This subject will not be discussed in exclusivity, but it will be alluded to in the discussion of soils.   Bach (2010) proposed that the total mass of a mountain and the range of which it is part, might effectively change the surrounding atmospheric temperature especially when compared to an isolated mountain at a similar latitude. Bach (2010) also analyzed the diverse pathways of frontal systems upon encountering mountain ranges, in relation to the position of the ranges with respect to the prevailing winds. Ultra violet radiation and its elevation gradient, as well as its responses to steepness (as determined by slope angle and position to the sun), is also discussed in Bach‟s report. For example, concerning solar radiation, Scott and Billings (1964) observed that “root and shoot standing crop” in mesic meadows reacted to solar radiation only when combined with other climatic factors. In xeric sites, however, variations in growth were exclusively attributed to changes in solar radiation. All of the above factors create considerable difficulties when analyzing the independent effect of each limiting factor on plant distribution at high elevations. In British Columbia the mean annual temperature in the alpine averages -4 to 0 oC, the mean temperature of the warmest month is less than 10 oC, and the average temperature remains below zero for 7-11 months (Pojar and Stewart 1991). The literature varies considerably on the extent of the effect of temperature on high elevation plant communities. For example, Rae et al. (2006) considered that woody species are primarily influenced by mean and maximum temperatures, and secondly by wind speed and soil temperature; in contrast, the presence of herbaceous species was driven by soil temperature and average air temperature. Conversely, Billings and Mooney (1968), and  5  Martin (2001) argued that since alpine species are cold tolerant, ambient temperature is not a primary factor; instead, “steep local gradients in snow cover and soil moisture often govern productivity and distribution of alpine plants” (Billings 1973). Interestingly, while climate changed gradually with elevation, vegetation appeared to occur in distinct „belts‟ in their life form (functional type), composition, and character (physiognomy) of dominant species; however, species richness did not seem to reflect such distinct zonation (Nagy and Grabherr 2009). The abovementioned abiotic factors, with emphasis in British Columbia, are briefly discussed below.  Snow Accumulation, distribution, and disappearance of snow are frequently considered to be primary factors determining plant community composition. It has been consistently reported that differential snowmelt largely determines the variations in growing season thereby creating the „patchy‟ distribution of vegetation in alpine areas (Brink 1964). The importance of drought caused by early snowmelt, soil properties, or wind, was also analyzed by Brink (1964), who concluded that surfaces with only slightly different water table levels display significant differences in seedling establishment and growth. The distribution of snow might also affect the migration patterns of wildlife, which in turn play an important role in high elevation plant ecosystems.  Several different specific-effects of snow on vegetation have been reported (Heegaard and Vandvik 2004, Huelber et al. 2006) and correlated with the effects of topography on snowmelt. Topography controls snowpack accumulation, length of the growing season, soil water availability and the distribution of plant communities (Fisk et al. 1998). Chen et al.  (2008) found no evident relationship between plant diversity and soil water content but it was reported that the highest species richness and species diversity occurred with intermediate snow depths. Species composition (based on Sørensen‟s similarity index) was also found to be affected by snow depth (Chen et al. 2008).  Snowmelt was considered to be the single most important factor “in determining the distribution of the alpine meadow and alpine desert vegetation” (Shaw 1916), in the  6  Selkirk mountain range of British Columbia. A simple but insightful categorization was suggested by Shaw (1916): Other things being equal, the snow disappears first from the east-facing and south-facing slopes; these slopes therefore usually bear more luxuriant and more varied vegetation than those facing the west and north.  Kudo (1992) described that in fellfield habitats (e.g. scree slopes), the lack of protection from snow resulted in additional susceptibility to cold and drought while allowing for a long growing season. On the other hand, the vegetation under snowbed habitats overwintered under relatively warm and moist conditions but suffered from a short growing period caused by the late snow melt. Other effects of snow observed in British Columbia were described by Brink (1964). These were snow creep (movement inside the snowpack); snow glide (movement of the snowpack); interfacial frost (freezing of the soil/snow transition zone); and, needle ice (micro-frost of bare soil during the growing season).  Douglas and Bliss (1977) attributed the mosaic pattern of alpine vegetation in the North Cascades to snowfall (combined with complex topography), which created variable snow depths and snowmelt timings. In turn, differential snowmelt timings created a wide range of soil moisture regimes.  Walker et al. (1995) considered that rather than length of the growing season – as suggested by Billings and Mooney (1968) and Selby (1980) - snow was the primary limiting factor for alpine species. This hypothesis was supported with Walker et al.‟s (1995) results which showed that “initiation of growth, date of maximum leaf length, leaf number, and flower numbers” (Walker et. al. 1995) for the alpine forbs Bistorta bistortoides and Acomastylis rossii, were related to snowfall patterns. Specifically, they found that the average number of leaves nearly doubled and leaf length increased by ~ 10% in a single year (within the six year study) for B. bistortoides, when higher than normal snowfall amounts occurred as a result of a major ENSO (El Nino) event.  Walker et al. (1995) observed that it was not only timing of snowmelt that influences the phenology of alpine vegetation, but it was also the combination of snowmelt, nutrient availability, soil moisture and air temperature that determined the responses of individual species to varying conditions. However, Kudo (1992) considered  7  time of flowering and fruiting a direct result of snowmelt‟s timing gradient. The snow-release gradient created a diversity of flowering patterns even between adjacent populations of the same species. From this information, Kudo (1992) inferred that reproductive success was closely linked to the variation of snowmelt timing. The effects of differential snowmelt were also used to explain the two-way ordinations presented by Douglas and Bliss (1977) in Figure 2; all plant ordinations are shown with respect to timing of snowmelt for the central, east, and west North-Cascades.     Figure 2 Ordination of 128 stands in the alpine zone of the North Cascades. Stands are included in three ordinations according to region (West, Central, and East).  Most recently, Mackenzie (2006), supported by Klinka and Chourmouzis (2001) - as presented in Figure 3 - proposed „date of snow disappearance‟ as one of the axes for the edatopic grid for the Alpine-Tundra (AT1). Most authors catalogued plant communities into similar „habitat types‟ (i.e. snowbeds, late snowbeds,) which captured differences in moisture gradients and in the influence of snow in alpine environments.  Déry (2010) observed how transportation of snow by wind varies drastically in the Cariboo Mountains, which further helps explain the complexity of the interactions between topography, climate, snow distribution, and the response of plant communities. The effect of snow as a factor with an important influence on other variables (soil conditions and plant interactions) will be further discussed below.                                                      1 Prior to the current revision of the Biogeoclimatic Ecosystem Classification (BEC), both subalpine and alpine ecosystems were classified as „Alpine Tundra‟.  Figure 2 has been removed due to copyright restrictions. It showed the ordination of 128 stands in the alpine zone of the North Cascades. Stands are included in three ordinations according to region (West, Central, and East). Original source:   Douglas, G.W., and Bliss, L.C. 1977. Alpine and high subalpine plant communities of the North Cascades range, Washington and British Columbia. Ecological Monographs 47[2]: 113-150.    8            Figure 3 Climatic-edaphic grid showing relationships of the site groups and associations in the parkland Mountain Hemlock2 (MH) subzones to gradients of snow duration and soil moisture (Klinka and Chourmouzis 2001).   Soils  The importance of soils in alpine and subalpine ecosystems is most evident at treeline. Körner (2003) correlated the upper-most boundaries of trees with soils whose temperatures exceeded 5oC for 128 days at a soil-depth of 5cm. Similarly, Bliss (1971) found that dry matter production was significantly correlated with soil temperature and that shoot growth responded to a combination of air and soil temperature.  Previously, Cox (1933) considered amount of humus, and the resulting differences in water holding capacity, as the most immediate effect of alpine soils on the vegetation. Cox (1933) also suggested that high elevation soils were consistently nutrient-poor and immature in development. Today, contrary to Cox‟s observations, it is recognized that there is a much wider range in soil types and that vegetation responds differently to their diverse structures and distribution. Also, it has been frequently observed that the wide range of high elevation soil typologies support rich and poor, and dry and wet conditions. Hence, alpine soils can no longer be generalized as nutrient poor. For example, in the European Alps, poorly developed soils with sparse vegetation exhibited carbon to nitrogen (C/N) ratios as low as 5 – 8. In contrast, acid raw humus soils under ericaceous shrubs showed C/N ratios of 40 or higher (Körner 2003).   Prior to Körner‟s findings, Brink (1964) found that in areas of Garibaldi Provincial Park a combination of cold, dry soil, with                                                  2 The Mountain Hemlock zone is the subalpine ecosystem in the coastal ranges of BC, analogous to the subalpine ecosystems (Engelmann Spruce – Subalpine Fir parkland (ESSFp)) described in this thesis.  Figure 3has been removed due to copyright restrictions. It showed the climatic-edaphic grid showing relationships of the site groups and associations in the parkland Mountain Hemlock (MH) subzones to gradients of snow duration and soil moisture Original source:  Klinka, K. and Chourmouzis, C. 2001. The Mountain Hemlock Zone of British Columbia. Scientia Silvica Forest Science Department, The University of British Columbia, Vancouver, BC.     9  fluctuations in soil insulation (from snow), and fluctuations in soil temperatures, might determine the critical conditions for „ecesis‟ on subalpine barrens. Brink concluded that rather than cryogenic influences, lack of summer moisture and high summer surface temperatures (consistent with Körner) were the main soil conditions influencing the establishment of vegetation. While analyzing a series of soil textures, Brink (1964) found that plant abundance varied from high on silts, to low to none on coarse gravel. Plant density was correlated with relative moisture-holding capacity. It is also important to understand how fine mineral substrate is accumulated at high elevations so that the interactions of soils with plant communities can be better understood. Körner (2003) considered that:  a) site erosion of bed rock, b) gravity, c) sedimentation by water (liquid or solid) and, d) sedimentation by wind, were the main abiotic factors in the creation of alpine soils. The importance of fine-textured materials in alpine ecosystems was observed by Litaor et al. (2002) when they noticed that cryoturbation aided the intrusion of fine-textured mineral soils into the rhizosphere; consequently, higher clay and sesquioxide content was recorded. The increase in clay and sesquioxide affected the availability of nutrients to plants, particularly phosphorus (P), since specific-surface area of soil particles was increased (Litaor et al. 2002). To further explain the „patchy‟ character of the alpine, Brink (1976) suggested that “soil perturbation, steep slopes, high rates of erosion, frost riving, heaving, solifluction, sorting and other features associated with patterned substrata”(Brink 1976) contributed to the seemingly erratic distribution of alpine soils and plant communities. Valentine (1976) indicated that the compounded complexities of topography, meso- and microclimate, alongside geomorphologic conditions, contributed to the disruption of addition, removals, transferrals, and transformations during soil formation. Other features unique to alpine and subalpine soil formation processes are: “shattering of particles by frost, the disturbance of profiles and horizons by cryoturbation and solifluction and the slowing down of chemical and biological reactions as a result of low temperatures” (Valentine 1976). The difficulties in classifying high elevation soils are accentuated by their youthfulness. This implies such soils: “may not yet have reached a stable equilibrium position even in the short term” (Valentine 1976). Faced with these  10  difficulties, Valentine recognized that there was insufficient knowledge of the rate of soil processes and the role they played on the different high elevation soil types known in 1976. Unfortunately, research on these soils in BC has not been significantly advanced since the initial classifications.  Chemical and physical weathering in alpine soils was evidenced by iron and aluminium accumulation in Podzolic B horizons and clay build up in the B horizons (Sneddon et. al 1972).  More specifically, Sneddon et al. (1972) and van Ryswyk and Okazaki (1979), reported montmorillonite being weathered from vermiculite and micas in the topsoils, and kaolinite weathered from montmorillonite. Considering that there are discrepancies in the recognition of chemical and physical weathering in alpine soils, an alternative explanation is that volcanic ash might be responsible for the availability of weatherable Fe and Al, which explained their high contents in the B horizons (Sneddon et al. 1992). Volcanic ash might also provide high water retention capacities (Valentine 1976).  After an extensive review of the original soils classification, Valentine suggested the following trends for the Central Interior, and Northern Coast Mountains: 1. Humo-Ferric Podzols in the Krummholz zone; 2. Various forms of Brunisols in the treeless alpine (properties varying by bedrock type) and; 3. Turbic and Cryic Regosols in the high alpine where frost disrupts the soil and low temperatures limit chemical weathering.       More recently, Wittneben and Lacelle (2006) described the following trends for the alpine soils in the Columbia Mountains: 1. Regosols with disrupted horizons created by colluviation, cryoturbation and solifluction; 2. Gleysolic soils  in areas with imperfect drainage;  3. Melanic and Sombric Brunisols with turfy Ah horizons under alpine graminoid vegetation;  11  4. Lithic soils at the highest elevations (2300-3600 m.a.s.l.) in the Columbia and Southern Rocky Mountains  Nitrogen (N), litter decomposition, and soil microbes The subalpine and alpine ecological dynamics that influence plant community composition are heavily determined by the interaction between plant individuals, between communities, and most importantly between plants and soil organisms.  The importance of bacteria, fungi and other biotic decomposing organisms is illustrated by the reliance of the flora on the nutrients released by decomposer organisms. N is considered the most limiting nutrient to alpine plant productivity and it is the most common parameter used to assess impacts from human activities in the AT. Nitrogen availability is critical for plant growth at high elevations, and it is predominantly dependant on biological activity and atmospheric inputs.  To date, most research on soil nutrients at high elevations has focused on N availability, N cycling (Figure 4) and the associated plant responses. Understanding the role that N plays in plant community composition is essential for understanding the diversity in AT plant communities and their response to changes in the environment and in succession. In general, alpine plant communities are considered N deficient; however, the capacities and requirements for N vary by species (Miller and Bowman 2003). It is important to consider that while individual plants might have nutrient deficits, a plant community normally shifts in composition to compensate for a certain nutrient deficit (Körner 2003) so that plant communities are not nutrient limited, but their variability reflects the abundance or lack of certain nutrients.  It is widely accepted that the availability and distribution of N has an ample spatial variability mainly driven by soil moisture regimes; this variability is often captured by comparing N dynamics by site type (eg. wet, mesic, xeric), as discussed below.      Figure 4 Fluxes of N for the Niwot Ridge, Colorado alpine ecosystem Bowman and Seastedt (2001). Figure 4 has been removed due to copyright restrictions. It was a diagram of the fluxes of N for the Niwot Ridge, Colorado alpine ecosystem. Original source: Bowman, W.D., and Seastedt, W.D. eds. 2001. Structure and function of an alpine ecosystem – Niwot Ridge, Colorado. Oxford University Press, UK.     12   Fisk et al. (1998) attributed N turnover among alpine plant communities to soil moisture gradients, but others (Steltzer and Bowman 1998, Miller and Bowman 2003, and Klanderud 2005) have suggested that soil N transformations are controlled by plant species (with their corresponding microbial communities) thereby influencing their communities,  structures, and functions. Another example of  N-moisture interactions was presented by Miller and Bowman (2003) who found that soil moisture accounted for 60% of the variation in exchangeable NH4-N, and 30% of the variation in water extractable amino acid N concentrations. The authors also found that in wet meadows, NH4-N and amino acid N concentrations were positively correlated with soil moisture, while in dry meadows, amino acid N concentrations were inversely correlated to soil moisture. For exchangeable NO3-N, a positive correlation with soil temperature but not with soil moisture, was observed. The variation of snow accumulation was also shown to affect decay rates and plant productivity, organic matter accumulation (Liator et. al 2002, Bryant et al. 1998), and microbial processes with their cumulative effects on N mineralization and immobilization rates (Liator et. al 2002).  Researchers in Sweden observed that the nutritional requirements for growth influenced the alpine plant species‟ response to increases in N (Nilsson et al. 2002). They observed that the faster growing species, Deschampsia flexuosa, responded favourably to increases in N when compared to the adjacent slower growing species, Empetrum hermaphroditum (Nilsson et al. 2002). Favourable growth responses and reduced competing vegetation (thereby, possibly reducing secondary metabolites), strengthened the competitive advantage of the faster growing species when N deposition increased (Nilsson et al. 2002). For example, stress tolerant plants such as Ericaceous plants, create litters low in N but high in secondary metabolites (e.g. phenolic allelochemicals) that suppress soil microorganisms and their processes. As faster growing species out-compete ericaceous plants, in response to increases in N, the soil microorganisms are also favoured by the reduction of secondary metabolites. An increase in microbial biomass, allows for faster cycling of nutrients which ultimately keeps a positive feedback  13  mechanism in place. Figure 5 illustrates changes in dominance as the plant communities respond to increases in N.          Figure 5 Conceptual diagram of the effects of alleviation of stress (e.g. low fertility, low pH) on alpine plant communities and plant diversity. Subordinate species include vascular plants, mosses and lichens Nilsson et al. (2002).    Although high elevation ecosystems in the Cariboo Mountains are less predisposed to suffer changes from N deposition (due to the lack of anthropogenic influences), the research discussed below provides valuable insight into some of the possible changes that plant communities might display in response to potential future changes in soil N content. Bowman (2000) observed that, in a Colorado Rocky Mountain alpine ecosystem, Acomastylis rossi3 was at a competitive disadvantage to Deschampsia caespitosa4 in response to increased N deposition. Steltzer and Bowman (1998) also noted that increased N deposition could induce positive feedbacks in the N cycle. The addition of N and its effects are strengthened by changes in plant communities; primarily, the increased abundance of fast growing species triggered positive feedback responses of accelerated N fluxes in the ecosystem.  A generalized description of these interactions is given in Figure 6.                                                        3  Species with lower N cycling characteristics. 4 A higher nutrient requiring species which enhances soil N cycling.  Figure 5 has been removed due to copyright restrictions. It was a conceptual diagram of the effects of alleviation of stress (e.g. low fertility, low pH) on alpine plant communities and plant diversity. Subordinate species include vascular plants, mosses and lichen. Original source: Nilsson, M.C., Wardle, D.A., Zackrisson, O., Jäderlund, A. 2002. Effects of alleviation of ecological stresses on an alpine tundra community over an eight-year period. Oikos 97[1]: 3-17   14          Figure 6 Conceptual model showing the generalized pattern of plant species self-replacement with low or high N availability. In the face of higher N deposition, N conserving species of infertile environments would be replaced by more-competitive species, with biotic characteristics promoting higher N cycling in soils. The new state would require some disturbance or exhaustion of soil N reserves, allowing species adapted to low N conditions to re-establish (Bowman 2000).   In alpine ecosystems in the Colorado Rocky Mountains, Miller and Bowman (2003), found that all species studied can potentially take up all forms, organic and inorganic, of N (NH4-N, NO3-N, and glycine which was the most abundant N-containing amino acid in the soils of the area of study) but different species had different uptake capacities for each form of N. These capacities were similar within species of a given plant community but varied significantly between different plant communities. However, in Sweden, Nilsson et al. (2002) found that addition of organic N had no significant effect on any vascular plant; only Pleurozium schreberi (a moss) responded favourably to its additions. Nilsson et al. (2002) did, however, observe that mineral N additions had a negative impact on non-vascular plant communities, especially for the lichens Dicranum spp., and P. schreberi.  In the Colorado Rockies, NH4-N was the most abundant form of exchangeable N for all sites, yet only half of the species had a high capacity to take up NH4-N; furthermore, only a third of the species preferred NH4-N over the other forms of N (Miller and Bowman 2003). NH4-N and amino acid N concentrations were low across all communities and exchangeable NO3-N concentrations were higher in soils from the dry meadows (Kobresia myosuroides - Carex rupestris). Within the dry meadow communities, three out of five species displayed high capacities for NO3-N uptake, which led the authors to conclude that when/if this form of N is available, it could be widely used by plants. Soils from moist ( Deschampsia caespitosa - Acomastylis rosii) and wet Figure 6 has been removed due to copyright restrictions. It was a conceptual model showing the generalized pattern of plant species self-replacement with low or high N availability. Original source: Bowman WD. 2000.  Biotic controls over ecosystem Response to Environmental Change in Alpine Tundra of the Rocky Mountains. Ambio 29[7]:396–400     15  (Carex Scopulorum) meadows had higher amino acid N concentrations than NO3-N  concentrations, but only species from dry meadow communities exhibited a higher capacity to take up amino acid N than inorganic (NO3-N and NH4-N) forms of N.  This was previously observed by Raab et al. (1996) who concluded that Kobresia myosuroides -a sedge that commonly dominates alpine dry meadow communities- can take up amino acid glycine at higher rates than NO3-N  or NH4-N, an adaptation which presumably allowed the sedge to obtain its N in environments with slow N mineralization rates.  Having observed that amino acid N concentrations had a negative correlation with soil moisture in the dry meadow community, Miller and Bowman (2003) concluded that amino acid N is also a critical form of N for plants during periods of drought. Another interesting finding was that while all alpine plant species take up amino acid N at equal or greater rates than NO3-N and NH4-N, most species have a greater uptake capacity for either NH4-N or NO3-N   in comparison to at least one of the other forms of N (e.g. glycine). That is, plants had a greater capacity to take up either NH4-N or NO3-N, compared to their capacity to take up glycine, yet actual uptake rates were generally higher for glycine. The differential uptake capacities for different forms of N might help explain why Nilsson et al. (2002) observed that although additions of different forms of N and other nutrients had different consequences for humus properties and the decomposing subsystem, all additions had similar above-ground consequences. In general, they observed a decline in Empetrum hermaphroditum, and an enhancement of Deschampsia flexuosa. However, differential uptake capacities by themselves cannot satisfactorily explain Nilsson et al.‟s (2002) abovementioned observations, so we must look at the role of soil microbes in nutrient cycling and ultimately on determining plant communities.  Soil microbial activity is key in the N cycle, but unlike other ecosystems, those at high elevations have a strong alternating pattern of N immobilization and release in response to seasonal changes (i.e. freeze-thaw cycles). Snowmelt and its variability in microclimates created variable litter decay rates at high elevations (Fisk and Schmidt 1995, Bryant et al. 1998, Lipson et al. 1999, Jaeger et al. 1999 and Costello and Schmidt 2006), and it controlled the protein release from winter microbial biomass (Lipson et al.,  16  1999) and ultimately N availability (Jaeger et al. 1999). Soil moisture influenced non-symbiotic N fixation (Bowman et al. 1999, and Costello and Schmidt 2006), yet it did not constrain rates of symbiotic N fixation (as is the case for Trifolium dasyphyllum) (Wojciechowski and Heimbrook, 1984). Lipson et al. (1999) concluded that the key event controlling N availability to alpine plants occurs after snow melt, when protein is released from the winter microbial biomass. Jaeger et al. (1999) found that plant N uptake was much higher during the first half of the growing season than in the rest of the growing season (Figure 7). The differential uptake of N was inversely correlated with microbial N pools, which were measured to be low in the first half of the growing season and increased as the season progressed; the highest microbial N immobilization occurred after plant senescence (Jaeger et al. 1999).  Microbial biomass peaked when soils were cold in autumn, winter, and in early spring, coinciding with the immobilization of N; but, a burst in available N occurred in late spring/early summer coinciding with the perceived increase in protein, enzymatic activity, and amino acids (Lipson et al. 1999).      Figure 7 Plant and microbial 15N tracer uptake, expressed per square meter. Vertical bars on microbial plots represent +1 SEM for n= 8. Vertical bars on the plant plots represent +1 SEM for n = 6-8. Data are from harvests conducted 20 June and 13 August (Jaeger et al. 1999).  Bowman et al. (1999) studied three species of Trifolium, representing dry alpine meadows, wet alpine meadows, and fellfields; they observed that these communities obtained 70-100% of their N requirements from atmospheric N2 fixation.  Wojciechowsi and Heimbrook (1984) estimated very low seasonal inputs (5mg N m-2) by cyanobacteria, making their contribution most likely significant only in the long term build-up of N in the developing soils. Bowman et al. (1999) noticed no difference between dry meadows, Figure 7 has been removed due to copyright restrictions. It was a graphic showing plant and microbial 15N tracer uptake, expressed per square meter. Original source: Jaeger, C.H., Monson, R.K, and Fisk M.C. 1999. Seasonal partitioning of nitrogen by plants and soil Microorganisms in an alpine ecosystem. Ecology 80[6 ]:1883-1891.    17  wet meadows and fellfield communities, in terms of proportion of N obtained from fixation. On the other hand, the different contributions to percent-cover by the Trifolium plants resulted in a range of annual inputs of N from fixation, from 127 mg m-2 per year in wet meadows to 810 mg m -2  per year in fellfields. High cover, combined with high fixation rates in the fellfield communities, resulted in the highest overall N inputs from Trifolium. These results indicate that availability of water does not constrain the plants‟ distribution or their rates of symbiotic N fixation (Bowman et al. 1999), yet non-symbiotic N fixation was found to be constrained by water availability (Wojciechowski and Heimbrook 1984). Considering that N mineralization is constrained by water availability, symbiotic N fixation in fellfield communities (and possibly in other dry communities) might be of critical importance as the primary source of N for plants and microbes within these communities.  Also, freezing of soils, rather than drying, was identified as the main factor controlling water availability; however, freeze-thaw cycles, combined with soil anoxia from saturated conditions, were associated with high dynamism and diversity in bacterial community structure (Figure 8).        Figure 8 Pie chart showing proportions of major phylogenetic groups represented in a bacterial clone library from spring alpine tundra wet meadow bulk soil (0–20 cm). The bacterial groups are primarily related to the ones involved in iron cycling, nitrogen cycling (fixation, nitrification, and denitrification), hydrogen oxidation and carbon breakdown (Costello and Schmidt 2006).      Figure 8 has been removed due to copyright restrictions. It was a Pie chart showing proportions of major phylogenetic groups represented in a bacterial clone library from spring alpine tundra wet meadow bulk soil (0–20 cm).  Original source: Costello, E. K., & Schmidt, S. K. 2006. Microbial diversity in alpine tundra wet meadow soil: novel Chloroflexi from a cold, water‐saturated environment. Environmental microbiology 8(8), 1471-1486.   18  Mycorrhizae Spatial variability and soil moisture are also important in determining mycorrhizal infection rates. The degree of mycorrhizae infection was presumed to be highest in the upper 5 cm of alpine soils and in well drained soils (Körner 2003). Barnola and Montilla (1997) noticed half as many mycorrhizae infections in the upper 5cm as within the 10-30cm soil depth, as well as lower infection rates in poorly drained soils.  Körner (2003) compiled the following list of mycorrhizae and their typical associates in alpine vegetation: ectomycorrhizae (Salix, Dryas, Polygonyum, Kobresia), ericoid mycorrhizae (Ericaceae), vesicular-arbuscular mycorrhizae (VAM) (forbs, grasses, and sedges) and orchid mycorrhizae. Körner (2003) also mentions the presence of mutualistic „dark-septate hyphae‟ associations, which enhanced growth and P uptake in Carex firma within the high-alpine (Haselwandter and Read 1982). The relative proportion of mycorrhizal plants was found to decrease with increasing altitude (Ruotsalainen et al. 1997) which was also observed by Haselwandter and Read (1980) for Vaccinium myrtillus and V. vitis-idaea in the Austrian Alps. Routsalainen et al. (1997) speculated that the inverse relation between altitude and mycorrhizae abundance might exist because plants at higher elevations can afford to share less photosynthesized carbon with the fungi, thereby decreasing the rates of mycorrhizal associations; thus, since photosynthetic nutrient use efficiency (PNUE) decreased along an altitudinal gradient, the rates of mycorrhizal infection followed the trend (Routsalainen et al., 1997).  Barnola and Montilla (1997) found that in poorly drained fertile soils of the higher alpine the rate of mycorrhizal infection for a species of Carex was less than in the lower alpine‟s well drained infertile soils, which further supports the above observations in which wetter soils have less mycorrhizae infections than drier soils. The relationship between mycorrhizae and other nutrients, at high-elevations, is still poorly understood. Körner (2003) suggested that there was no correlation between a carbon surplus and degree of mycorrhizal infection for Carex curvula. Mullen and Schmidt (1993) showed the relation (Figure 9) between phosphorus and rates of VAM infections in the Colorado Rocky Mountains; the presence of arbuscules decreased with increased P accumulation in the roots and shoots of Ranunculus adoneus.  Further  19  research is needed to confidently assess how nutrients, soil conditions, and other environmental factors might influence rates of mycorrhizae infections at high elevations.         Figure 9 Development of arbuscules of the coarse endophyte and shoot phosphorus concentrations in R. adoncus at different dates Mullen and Schmidt (1993).     Plant interactions  In alpine ecosystems, facilitation was found to be more important than competition between plant individuals and communities (Callaway et al. 2002, Nilsson et al. 2002, and Körner 2003). Amongst the positive effects that plants can have on each other, Callaway et al. (2002) mentioned: accumulation of nutrients, provision of shade, amelioration of disturbance and protection from herbivores. However, the role of competition or facilitation for soil moisture is poorly documented. Bowman and Seastedt (2001) alluded to interspecific competition between the dry meadow dominant Kobresia myosuroides and the moist meadow co-dominant Deschampsia caespitosa, as the main factor excluding the latter from dry meadows in which it is out-competed by the former; yet, it is unclear whether they were competing for nutrients, water, or both. Nonetheless, it has been documented that a strong relationship exists between time of snowmelt and nutrient availability. Therefore, it is probable that facilitation plays an important role in reducing snow-driven stresses, such as reduction of growing seasons and anoxic soil conditions.  Kikvidze et al. (2005) suggested that more species can coexist (higher community richness) when they were spatially arranged by niche differentiation as opposed to Figure 9 has been removed due to copyright restrictions. It was chart showing  development of arbuscules of the coarse endophyte and shoot phosphorus concentrations in R. adoncus at different dates. Original source: Mullen, R.B., and Schmidt, S.K. 1993. Mycorrhizal infection, phosphorus uptake, and phenology in Ranunculus adoneus: implications for the functioning of mycorrhizae in alpine systems. Oecologia 94:229-234.    20  random, unstructured spatial distributions. This was evidenced by the climatic patterns observed in plant interactions in three different alpine climates: a) mild-climates had greater plant biomass and stronger competition; the greater plant biomass reduced intraspecific patchiness and increased local richness; b) cold-climates had little plant biomass, so a high proportion of species benefited from strong facilitative effects of neighbouring plants; c) in intermediate climates (relative to the abovementioned) there were intermediate amounts of plant biomass and weak interactions - presumably from counterbalancing of competition by facilitation - as well as a random distribution of plants (Kikvidze et al. 2005). Niche differentiation in relation to soil moisture and nutrient uptake capacities is likely to explain the arrangements of plant communities as well as the type and intensity of interactions amongst plants at high elevations.  Also, the facilitation provided by pioneering species, which increased shade and therefore decreased losses in soil moisture, was found to be important for plant establishment in what would have been uninhabitable soil conditions (Körner 2003).  Succession  At high elevations there is widespread disagreement on what plant communities, if any, represent early succession stages of development and which are at climax stages. Due to the large amount of spatial and temporal variability in high-elevation plant communities, it is commonly assumed that all plant communities are at a climax stage (Mackenzie person. comm.).  However, as early as 1933, this view was challenged; Cox (1933) concluded that the development of the vegetation cycle in areas of the Rocky Mountains followed different pathways (Figure 10).       Figure 10 Diagrammatic representation of the main lines of alpine plant succession in xero- and hydroseres. Especially long enduring locally distributed edaphic associations can be recognized by the broken lines which continue beyond them (Cox 1933).  Figure 10 has been removed due to copyright restrictions. It was flow chart showing the diagrammatic representation of the main lines of alpine plant succession in xero- and hydroseres. Original source:  Cox, C.F. 1933. Alpine plant succession on James Peak, Colorado. Ecological Monographs 3[3]: 299-372.    21     Martin (2001) suggested that herbivory, trampling, and occasional fires (specifically in subalpine grasslands) were the most recurrent forms of disturbance at high elevations. Short growing seasons and long life cycles added to the difficulty of understanding succession at high elevations (Körner 2003); nonetheless, researchers in New Zealand and Europe have recently begun to determine the importance and the extent of succession in alpine ecosystems.  In British Columbia there is very scarce literature suggesting succession stages in high elevation non-vegetated communities. Brett et al. (1998) identified the Luzula wahlenbergii – Saxifraga tolmiei association as a “pioneer community dominated by Saxifraga tolmiei, Luzula wahlenbergii and Marsupella brevissima [which] occurs where soil instability and cool temperatures restrict vegetation and soil development” (Brett et al. 1998). Furthermore a transition to the Cassiope mertensiana – Phyllodoce empetriformis order was signalled by the presence of Phyllodoce empetriformis and dwarf Tsuga mertensiana as a result of soils being protected from sheet-wash, which enabled accumulation of organic matter (Brett et al. 1998). They also found that the Luetkea pectinata association “likely represents the initial stage of succession on alpine sites and, as organic material accumulates and the soil is stabilized by vegetation, [it] is succeeded by zonal heath communities” (Brett et al. 1998). Recently formed fluvial terraces, moraines, and boulder fields were observed to support the Anaphalis margaritacea – Lupinus arcticus association in the initial successional stages in an avalanche bowl in Church Mountain, BC (Brett et al. 1998).  Sequences in alpine plant communities can easily be observed within a small geographic scale; although they are not specifically considered succession patterns, such sequences might provide some insight into the developmental stages that alpine plant communities might experience at a broader geographic and time scale. For example Brett et al. (1998) identified the bog -> heath -> tree island community sequence, which was related to transitioning soils: Fibrisols -> Orthic or Eluviated Dystric Brunisols -> Podzols, respectively. The physical transition in this sequence ranged from standing water to 5 cm, 1 – 1.5 m, and 2 m above standing water, respectively. Once again, it is apparent that moisture (which is primarily determined by snow) and its influences are the  22  driving factors for the community sequences found at high elevations. Moreover, the influences that currently determine soil moisture and plant community sequences might be similar to those that shape the alpine over longer time scales. Tscherko et al. (2005) analyzed sixteen plants species, occurring in ecosystems ranging from 4 to 135 years after an Austrian‟s glacial retreat, and noticed that the abundance of annuals decreased as the abundance of perennials increased during the time period. The pioneering vegetation colonized the de-glaciated ground 14-20 years after glacial retreat. The specific ecological adaptations of the first colonizers allowed for the subsequent colonizers to become established after sufficient plant biomass had accumulated. N-fixing legumes did not appear until relatively late in succession, and woody species, e.g. Salix spp., briefly appeared during years 43-48, after which they were replaced by Poacea species. In Alaska, researchers concluded that no single factor or mechanism accounted for primary succession, but rather the changes in the competitive balance at different successional changes, provided the pathways for change in species dominance (Chapin et al. 1994).  Kikvidze (1993) confirmed some of Cox‟s (1933) findings, by concluding that during alpine succession, patches of vegetation facilitated the invasion of more specialized alpine species which resulted in an increased frequency of typical alpine species. Kikvidze (1993) mentioned that Campanula saxifraga - a pioneer petrophyte - is replaced by Campanula biebersteiniana which is a „typical‟ species of alpine carpets and meadows. Therefore, Kikvidze (1993) confirmed that a change in ecological conditions, mainly soil structure, was associated with the formation of specialized and highly diverse plant communities. The opposite response was observed, in New Zealand, during a simulated secondary succession event (i.e. road construction) in which an alpine cushionfield‟s cover was reduced from 59% to 34% (Roxburgh et al. 1988).  Despite very short growing seasons and slow growth rates, there is some evidence of succession in the high elevation plant communities. The difficulty in assessing succession stages lies in comparing sites with equivalent physical characteristics at different time periods since major disturbance events. While landslides, avalanches, major rockfall events, and trampling, commonly occur at high elevations, the successional stages of the  23  corresponding plant communities are likely to manifest over a time scale beyond our life-span and perhaps beyond recordings in literature. As knowledge on the subject develops and its relevance to operational and recreational activities at high elevations grows, it might become more apparent which, if any, plant communities indicate disturbance and provide the best path (e.g. for mining reclamation) for site restoration and for classification.      Overall objective, research question and scope of thesis The goal of this thesis is to identify and classify the plant communities of the sub-alpine and alpine ecosystems - within the western edge of the Cariboo Mountains of British Columba - for the purpose of furthering our basic knowledge regarding the botanical description, the distribution of species and the site conditions in which plants become established. Since a comprehensive system for the classification of plant ecosystems is widely used and well recognized in B.C., this system was followed for the classification of the vegetation studied in this project. Furthermore, the system used to derive the classification explores the relationship between environmental factors and the vegetation‟s response to them. The introduction to chapter 2 provides an overview to the origin of the classification system, its objectives, the principles behind it, and its relevance to high elevation ecosystems.  Objective  To produce a Biogeoclimatic Ecosystem Classification for a subsection of ESSF wet cold parkland and the Interior Mountain Heather Alpine wet cold subzones  (which are currently being developed by Mackenzie, W.H.) to the site series level in the Cariboo Mountains  24  Research question What is the spatial arrangement of the vegetation at high elevations within the western Cariboo Mountains of British Columbia?  Scope of thesis The scope of my thesis is to describe, analyse, and tabulate approximately 130 sites (vegetation, topographic, and edaphic conditions), combined with ~170 relevés (provided by the BC Forest Service). I hypothesised that the topographic and edaphic variables could be used to explain the spatial arrangement of the vegetation in the ESSFwcp and IMAwc subzones.     25    Chapter 2 Vegetation Classification  Introduction Although there is significant variability within non-forested high elevation ecosystems in British Columbia the provincial government is yet to adopt an official classification for these ecosystems5.  As population increases and land resource availabilities decrease, these remote and often pristine ecosystems are experiencing increasing pressures towards use and development.  To have sustainable development, we should develop at least a basic understanding of the structure and basic function of these ecosystems. A comprehensive and standardized classification of all ecosystems in British Columbia, including those at high elevations, is essential for sound ecosystem management. This thesis reinforces such efforts (led by the BC Forest Service) in which the principles behind the biogeoclimatic ecosystem classification (BEC) system are being applied to high elevation forested and non-forested ecosystems. High-elevation plant communities were the first ecosystems to be described by many plant ecologists (Braun-Blanquet 1926, Poore 1957). However, the complexity of their spatial arrangement led the ecologists to establish an approach whereby field observations and the intimacy of the ecologist with the area, were the main strengths of the descriptions (Shimwell 1972). Presumably, this was a consequence of the high degree of spatial variability of high-elevation ecosystems and of the limited capacity of statistical computations of the time. Also, the main purpose for the development of the classification techniques of the Zurich-Montpellier school of phytosociologists (which is the basis for the BEC classification) was to provide „academic‟ accounts of the arrangement of high-elevation ecosystems. Conversely, the study and classification of lower elevation ecosystems - primarily forested ecosystems and their plant communities - relied more on „objective‟ quantifications of site conditions (e.g. soil nutrient capacities,                                                  5 While several comprehensive classifications exist (e.g. Brett et al. 1998, Hämet-Ahti 1979), as of the completion of this thesis, the Ministry of Forests, Research Branch, has not „officially‟ adopted a province wide classification.   26  soil moisture deficits, leaf area index, site index and plant communities) in order to attain quantitative descriptions of the ecosystems. These descriptions and classifications of forested ecosystems were initially driven by „management‟ purposes, such as the optimization of volume and quality of lumber, and so it became imperative to understand how to select soil (edaphic) and site (topographic) conditions to obtain the desired stand characteristics. This was initially achieved by a significant effort in measuring (quantifying) as many variables as possible, which is in strong contrast to the descriptive approach in field ecology. At the heart of this juxtaposition (mensurative vs. descriptive research) lies the question of the validity of „objectively‟ measuring ecosystem parameters and the validity of „subjectively‟ describing the ecosystems‟ character. When scientists in British Columbia began establishing ecological classification systems for the province, rather than favouring one approach over another they sought to integrate the strengths of both the descriptive and the quantitative approaches.  From this philosophy, the biogeoclimatic ecosystem classification (BEC) system was developed.  The BEC system has enabled scientists, land managers and field technicians to understand and convey a large amount of information about terrestrial ecosystems in simple and concise terms. The BEC system for the description of terrestrial ecosystems is used on a daily basis by foresters developing site prescriptions, for reclamation purposes, for Predictive Ecosystem Mapping (Kopp and Cleland 2014), climate change modelling (Hamann and Wang 2006), and multi-million dollar conservation/land management projects like the ecosystem-based management plans in the Great Bear Rainforest (Kopp and Cleland 2014).  At its core, BEC subscribes to the phytosociology tradition of the Zurich-Montpellier approach to vegetation classification (Pojar et al. 1987), which in turn adheres to the traditions of central European phytosociology (Thorn and Hall 1980). Beyond its various uses, the BEC system is widely accepted in the peer-reviewed academic system which is evidenced by the number of citations (298 for Pojar et al. (1987) and 82 for the Field Guide to forest site identification and interpretation of the Cariboo Region (Steen and  27  Coupe 1997)). Yet, there is some resistance from the Anglo-American Plant Ecology6 (AAPE) approach to using descriptive methods for the study of plant communities (Ewald 2003). The BEC approach for classification seeks to reconcile some of the resistance from the AAPE to the Central European Phytosociology (CEPS) tradition by incorporating abiotic factors and repeatable modes of analysis to the traditional phytosociology descriptions.  The classification of British Columbia into „biogeoclimatic ecosystem‟ units began when “Krajina came to the University of B.C. in 1949 and began formulating the principles of Biogeoclimatic Ecosystem Classification (BEC). Krajina and his students, from the Department of Botany at UBC, developed BEC during the 1960's and early 1970's” (BEC WEB 2006).  In the 1970‟s, Ray Coupé and Ordel Steen, under the coordination of W.J. Watt, began to develop the BEC for the Cariboo Region. This project was formalized with the publication of „A Field Guide to Forest Site Identification and Interpretation for the Cariboo Forest Region Land Management Handbook number 39‟ in 1990.  Within this guide, most of the forested ecosystems of the Cariboo have an in-depth description, which is easy to follow and very well rationalized. The guide has provided land use planners and managers a sound basis for sustainable resource management. Similar BEC guides were developed for the other forest regions of British Columbia. The objectives of the classification (in the former Cariboo Region) were: (1) to provide a framework for organizing ecological information and management experience about ecosystems; (2) to promote a better understanding of forest ecosystems and their interrelationships; (3) to provide resource managers with a common “language” to describe sites; and (4) to improve the user‟s ability to prescribe and monitor site-specific treatments (Steen and Coupé 1997). To achieve these objectives, “the system groups ecosystems at three levels of integration: regional, local, and chronological. At the regional level, vegetation, soils, and topography are used to infer the regional climate and to identify geographic areas that have relatively uniform climate. These geographic areas are termed biogeoclimatic units.                                                  6 Which “strives to base its very applicability on repeatable sampling designs and modes of analysis.”(Ewald 2003)  28  At the local level, segments of the landscape are classified into site units that have relatively uniform vegetation, soils, and topography. Several site units are distributed within each biogeoclimatic unit, according to differences in topography, soils, and vegetation.  In order to arrange ecosystems at the three levels of integration, the BEC system combines four classifications: vegetation, climatic (zonal), site, and seral. Vegetation classification is most important to developing the ecosystem classification. However, the climatic and site classifications are the principal classifications used in the application of the BEC system” (BEC WEB 2006). Table 1 clarifies how some of the BEC terminology relates to climate and vegetation (Trowbridge et al. 2002).   29  Table 1 Differentiating criteria for BGC units.  a DCS = diagnostic combination of species. May include character, differential and/or companion species.  Zonal hierarchy Climate Vegetation DCS a Other Zone broadly homogeneous climate climatic climax ecosystems are members of the same zonal plant order exclusive DCS Grouping of subzones with affinities in climatic characteristics and zonal ecosystems Subzone one type of regional climate climatic climax ecosystems are members of the same zonal plant association exclusive DCS basic unit of BEC Variant regional variation, e.g. drier, wetter, snowier, warmer, colder distinct climax plant subassociation; or changes in proportion and vigour of certain plant species; or variations in successional development, or overall pattern of vegetation over the landscape exclusive or non-exclusive DCS difference in ecosystem productivity from subzone zonal site; intensity of soil-forming processes phase local, not regional, variation in climate due to local relief, e.g. topographic or topoedaphic  zonal ecosystems distinct from that of the subzone/variant surrounding the phase  not specified can be mapped for management or descriptive purposes; small in area relative to surrounding subzone/variant  The purpose of using the BEC procedure for the classification of vegetation is to develop ecologically-meaningful classes of plant communities that: a) can be floristically distinguished; b) represent a group of communities that have affinities in floristic composition and appearance; and, c) occupy a floristically-defined segment of edaphic and local climatic gradients (Brett et al. 1998). Jennings et al. (2009) described the characteristics of plant associations as units that have: (1) uniform physiognomy and  30  structure, (2) uniform habitat, (3) definite floristic composition, and (4) recurring distribution across a landscape or region.  The plant association is the basic category in the vegetation classification; alliances, orders, and classes are groups of associations, and subassociations are divisions of an association (Pojar et al. 1987). Plant associations with similarities in floristic composition, life form, and structure are grouped into alliances which represent major segments of the geomorphic gradient that occur in one or several related climates (Pojar et al. 1987); therefore, alliances are more compositionally and structurally variable, more geographically widespread, and occupy a broader range of habitat conditions than associations (Jennings et al. 2009). A major goal of ecology is to explain why communities change in a systematic fashion across space (McGill et al. 2006). The BEC system provides a way of explaining how communities change across space, which is a necessary first step in understanding what factors lead to such changes. Observing the full spectrum of change in plant distribution in alpine environments (from subalpine to high alpine) can provide useful information in analyzing stress gradients and the response of the vegetation to a variety of biotic and abiotic conditions. Also, in classifying high elevation plant communities, in the context of succession stages7, two approaches might be followed: a) to infer, based on the site‟s physical condition, whether it has been subject to disturbance at a different point in time than the surrounding sites, or b) to ignore whether disturbance has occurred and to recognize that for practical purposes the time scale for succession at high elevation is too long, and hence to consider all current plant communities to be at a mature to climatic seral stage. In British Columbia the second approach might be the best when contextualizing high elevation plant communities in a biogeoclimatic ecosystem classification approach The BEC research program aims at understanding and describing all land-based ecosystems within British Columbia; since non-forested ecosystems have been the last to be described, this thesis will complement the provincial classification for high elevation ecosystems, which is the last step in achieving a complete description of all terrestrial plant ecosystems of the province. The exact extent of high elevation ecosystems in                                                  7 As in the case in the BEC system  31  British Columbia is not well defined.  Martin (2001) determined that 17% of British Columbia‟s land mass was classified as the Alpine Tundra (AT) biogeoclimatic zone. Even if 17% is an overestimation of the extent of the AT, it is most likely an accurate estimate of high elevation ecosystems, forested and non-forested, in British Columbia. A substantial effort from the Ministry of Forests is underway to classify the high elevation ecosystems of British Columbia from a BEC approach. Such an approach, when applied to a botanical classification of high elevation non-forested ecosystems, presents various difficulties. The influence of snowpack, its variability in size, distribution, and timing of release (snowmelt) appear to determine the occurrence and composition of plant communities at high elevations. Hence, the former AT zone of the original BEC classification has been divided into three different zones (Figure 11): the Costal Mountain Heather Alpine (CMA), the Interior Mountain Heather Alpine (IMA), and the Boreal Altai Fescue Alpine (BAFA), following Mackenzie (2006). The new zonation was based on the observation that the differences in climatic components (snowpack depth, precipitation, temperature and continentality) were sufficient to merit a more thorough description and classification. For further information on the character of the different alpine areas of BC and the northwest North America, please consult Pojar and MacKinnon (2013). Some broad differences between these zones are; the CMA has a much lower treeline (at equivalent latitudes and aspects) than its IMA counterpart. In general, the CMA has the snowiest climate, followed by the IMA, and the BAFA has the driest climate. Nonetheless, latitudinality is a significant factor in differentiating between the IMA and the BAFA; whereas, continentality separates the CMA from the BAFA and from the IMA (Mackenzie, 2006). Plant communities in the CMA have heather associations with a higher percent cover of the land (generally speaking) than the IMA. The BAFA is dominated by grass, dwarf willow and birch (Betula glandulosa) communities. To account for these differences, recent classifications (Brett et al. 1998, Klinka and Chourmouzis 2001) and the proposed classification of Mackenzie (2006) place greater importance on snowpack depth than the classification for the other biogeoclimatic zones.   32        Figure 11 Distribution of the Alpine Tundra in British Columbia. BAFA = Boreal Altai Fescue Alpine CMA = Coastal Mountain Heather Alpine and IMA = Interior Mountain Heather Alpine (Mackenzie 2006).  Alpine regions and numerous remote areas of British Columbia were not thoroughly described in the initial BEC descriptions due to  limited access and the lack of tangible „interests‟ in these places (except for mining and grazing). Furthermore, the definitions for alpine and subalpine ecosystems have only recently been formalized.   In order to interpret and map alpine regions, the transitional zone between continuous forests and treeless alpine had to be defined, identified, and differentiated. Hitherto, Trowbridge et al. (2002) wrote, “parkland and alpine were mapped together during our small scale legacy mapping and this mapping was adequate for most users.” As ecology research advances and the BEC project matures, upper elevation forests, the subalpine, and the alpine need to be differentiated following biogeoclimatic principles (Trowbridge et. al. 2002). Consequently, over the past few years (after 2002), the transitional band between the continuous forest and the subalpine has been defined as a „woodland‟; the subalpine forests above the „woodlands‟ have been classified as „parkland‟ and the alpine has been recognized as the zone above the „parkland‟ (Appendix A  Figures 21 and 22). For example, in the Southern and Central Interior, the ecosystems above the wet cold Engelmann Spruce Subalpine Fir (ESSFwc) subzones are classified as ESSFwet-cold-woodland and ESSFwet-cold-parkland for the woodlands and parklands above the ESSFwet-cold subzone. It has also been determined that the ESSFwet-cold-parkland‟s upper limit (in other words, the subalpine-alpine boundary) is found when ecosystems have no trees as a result of severely limited growing seasons, although krummholz forms of trees may be present. The latest clarification and recognition of the highest elevation BEC ecosystems was the first step toward a thorough description of subalpine and alpine zones.  It is possible that the results from my research could be considered a „variant‟ Figure 11 has been removed due to copyright restrictions. It showed a map with the Distribution of the Alpine Tundra in British Columbia. Original source: Mackenzie, W.H. 2006. A Biogeoclimatic ecosystem classification of alpine and subalpine zones in British Columbia Ministry of Forests, Research Branch. Working Draft    33  within the abovementioned subzones, or that they will simply be used to develop a classification of a wider scope and scale.  Conceptual delineations of the AT and the Parkland As previously mentioned, the „subalpine‟ ecosystems were categorized under the „parkland‟ subzone during the High Elevation Summit in Vancouver, BC (Trowbridge et. al 2002) for BEC classification purposes. While the following descriptions are for „subzones‟, the scope of my research is limited to a small specific geographic area within the subzones, so it pertains to a yet undefined variant within the subzones. The parkland is described as the high elevation ecosystem above the continuous forest in which islands of trees are found occurring with krummholz, heath, herb meadows and grasslands. Hämet-Ahti (1965) divides the subalpine into „orohemiarctic and upper oroboreal‟ zones, which he bases on the differences between un-canopied plant communities and those occurring with trees.  Shaw (1916) recognized that “alpine meadow and forest formations interlock, the former frequently extending far down below the timber belt in the depressions, while the latter reaches upward on the exposed flanks and ridges.” Although he did not explicitly mention the area as a „subalpine‟ ecosystem, he distinguished it from what he describes as „Alpine Meadows and Alpine Desert‟. Bliss (1971) recognized the region in which no vascular plants can grow, but yet support insects, spiders, Collembola, enchytraeid worms and other heterotrophic organisms as the Aealian Zone. Currently, Mackenzie (2006) refers to this zone as the upper AT. Although it has little or no plant life, the recognition of this zone8 is necessary to develop an inclusive classification for the province.  Vestal (1915) described the alpine zone as the area below perpetual snow and above the forest line; moreover, he included “the transitional area in which subalpine krummholz and scattered trees alternate with alpine grasslands” (Vestal 1915) in the alpine zone, solely for practical purposes. McLean (1970) recognized the ecosystem of „limited‟ tree stands intermingling with large meadows as the „Abies lasiocarpa zone‟. He                                                  8 If actually present in BC.   34  acknowledged that it corresponded to Krajina‟s (1959) Engelmann Spruce Subalpine Fir BEC zone of British Columbia. McLean (1970) also defined the zone above the upper limit of krummholz development as “truly alpine in nature and tundra-like in appearance.” The literature is consistent in the general distinctions between subalpine and alpine ecosystems; however, there is disagreement on whether krummholz should be included in the alpine or in the subalpine zones.  Bliss (1971) observed that the herb fields and low shrub communities above tree line differed floristically and structurally from the alpine; also, they displayed stronger affinities with forest meadows. Douglas and Bliss (1977) supported this zonation but recognized that Dansereau (1957) and Löve (1970) included the krummholz belt with the subalpine zone. However, Bliss (1971), Krajina (1969), and Franklin and Dyrness (1973) included the krummholz in the alpine zone.  The vegetation and floristic data acquired by Douglas and Bliss (1977), from various samples in the mountains of the Pacific Northwest and BC, further support the inclusion of krummholz with the AT zone. Most recently, in coastal British Columbia, Brett et al. (1998) defined “non-forested plant communities as those communities where trees are permanently absent, or where they can survive only as prostrate or stunted shrubs < 3 m tall (krummholz).” Brett et al. (1998) also specified that this interpretation for the “non-forested plant communities” is commonly used to map the lower alpine limit. Ray Coupé (former Regional Ecologist for the Ministry of Forests, Williams Lake, BC) has included trees > 3 m and < 16 m tall in the „parkland‟ subzone for the former Cariboo Region, thereby integrating the krummholz into the AT zone. It has proven a useful approach since it allows for accurate, simple, and inexpensive mapping of the parkland-AT boundary. Furthermore, since the areas studied in this research represent only a small section of the ESSFwcp and IMAwc subzones, I tried to follow the definitions and delineations used for the specific areas of interest. For example, I included krummholz in the AT to be consistent with the provincial classification approach, for mapping purposes (it‟s very difficult to make out individual krummholz from aerial photography), and for functional purposes since krummholz have more similarities with shrubs than with trees.    35  Objectives and hypotheses To address the overall objective, the following objectives and hypotheses were used: 1. To conduct an intensive survey of the vegetation above 1900m.a.s.l. in the western Cariboo Mountains.  2. To detect and describe patterns in the distribution of the vegetation H0 = The vegetation is randomly distributed. H1= The vegetation is organized in discernible units.  3. If H0 is rejected, to explain some of abiotic factors that contribute to the organization of the vegetation.  H0 = None of the measured edaphic and topographic variables explain the discernible units of vegetation.  H1 = Some or all of the measured variables contribute to the explanation of the discernible units of vegetation. 4.  To describe the composition (species inventory) and distribution (occurrence of species in the landscape) of the observed vegetation units.  Methods Study area  The Cariboo Mountain range is “separated from the Rockies by the Fraser River and [is] bounded on the southeast by the McLennan River, Albreda Pass and the valley of the Albreda and North Thompson rivers. The northern limit is less well defined. Although it extends into the bend of the Fraser River, in the north, the alpine portion breaks down in the vicinity of Goat River” (Whipple 1992).   36  The research was conducted in the westernmost boundary of the Cariboo Mountains (Figure 12). The boundary of the study area (Figure 12) follows the distribution of high-elevation ecosystems within the Quesnel, 100-Mile House and Central Cariboo Forest Districts, of the Southern Interior Forest Region (Figure 14). The area that lies above 1900 m.a.s.l. (which is the lower limit of the study area) is 71,224 ha, but over half of this area is either permanently glaciated, or too steep to access. The pilot project area (Figure 13) was the area in which preliminary surveys took place to assess the feasibility of successfully following the experimental design. This area was more intensively studied than any other area within the project‟s boundary (yellow line Figure 12). It was selected for its accessibility, consists of 646.2 (ha), and is representative of the bedrock geology and climatic influences of the study area.  Sampling for the relevés (the standardized sampling unit/methodology in BEC) and snow-depth measurements was more intensive in the pilot project area than the rest of the study area.    Figure 12 The yellow line marks the study area boundary; the purple line delineates the Cariboo Mountains. All new relevés were conducted with the yellow boundary but were combined with relevés previously conducted with the purple boundary.  37   Figure 13 Mt. Elsey Pilot Project Area. The red line marks the 1900m contour, and the green asterisks show the location for relevé plots and snowpack measurements from 2007.       Figure 14 Southern interior Forest Region (in green)   Data collection Following the procedures described by Pojar et al. (1987), 130 relevés were sampled in 2007 and 2008. The relevés (for which approximate radiuses varied from 1m to 10m) were chosen, in situ, to represent sites of homogeneous vegetation structure and composition. At each site, vegetation (all vascular plants, and the main lichens and Figure 14 has been removed due to copyright restrictions. It showed the Southern Interior Forest Region. Original Source: MFLNO - Ministry of Forests Lands and Natural Resource Operations. 2012. Forest Analysis and Inventory Branch Biogeoclimatic subzone variant mapping. Victoria BC. Available online at: <http://www.for.gov.bc.ca/HRE/becweb/resources/maps/WallMaps.html>   38  mosses) percent cover, soil physical characteristics (including parent material), and topographic conditions were recorded. Snow depth (using a 320cm probe) was also recorded at a few sites throughout the study area (based on accessibility) and more intensively within the pilot project area (Mt. Elsey). The snow depth data were used to broaden my insight into the distribution of snow across the landscape rather than as explanatory variables.  The snow depth data was also used to identify areas that represented atypically high or low snow depths, and to compare inter-annual variation in snow depth in order to get a sense of the length of the growing season. Data from an additional 170 relevés were provided by the Ministry of Forests; these relevés were primarily from within the study area, although some were from adjacent forest regions and subzones with similar climate. Hence, the analysis was performed using a) only the relevés that I sampled (130), and then b) using the combined dataset (300 relevés). From the first 130 relevés 45 soil samples, from 37 relevés - chosen to represent a wide range of sites - were collected (seven of these samples were from a different horizon from the same soil pit so they were excluded from the statistical analysis) and sent to the BC Forest Service lab (Research Branch Analytical Chemistry Laboratory, Victoria, BC; supervised by Clive R. Dawson) for analysis of total N and total C (by elemental analyzer), pH (calcium chloride), NO3-N and NH4-N (see footnote)9, organic matter content (loss of ignition) and % fine particles (<2mm oven dried, milled and sieved) in the soil (full results are shown in Appendix B Table 45).  Data analysis  The relevés were transcribed from the field forms to electronic versions using VPRO (Vegetation and Environment Nexus Professional) software available through the Ministry of Forests (BEC WEB 2006). The dataset was imported into JUICE (Tichý 2002) software to produce tabulation tables, to conduct modified TWINSPAN (Roleček et al. 2009) and ISOPAM (Schmidtlein et al. 2010) analysis, as well as for basic data                                                  9 determined by a 2N KCl extraction (1 hour shake, 2.5 g soil to 25 ml solution) followed by colorimetric analysis of ammonium-N and nitrate-N using an OI-Analytical “Alpkem Flow System IV” segmented flow automated chemistry analyzer.  39  sorting. PC-ORD V6 (McCune and Mefford 1999) was used for the ordinations (see footnote 11) and cluster analysis. Generally speaking, ordination methods order plot data in terms of their similarity in floristic composition while clustering methods aggregate plot data into discrete groups based on floristic composition. TWINSPAN (two-way indicator species analysis) uses a hybrid approach with reciprocal averaging ordination methods and dendrograms; thus it is a combined approach of classificatory and ordination methods (Henderson 2003). Tabulation following Pojar et al. (1987) and Klinka and Chourmouzis (2001) was used for organizing, sorting, and identifying diagnostic combinations of species (DCS) based on the initial TWINSPAN output.  The associations and site series were determined by using the ordinations10 to verify that clusters reoccurred in distinct patterns along environmental gradients. This information was combined with my field observations to locate each site series and site associations in edatopic grids and to produce landscape diagrams. Identification keys were produced to facilitate the identification of these units in the field.  The procedure I followed addresses the four activities and criteria for definition of vegetation types identified by(De Cáceres and Wiser 2012) and is summarized in Table 2.     Table 2 Methods used to answer the questions presented by De Cáceres and Wiser (2012) to help define vegetation types Definition of vegetation types (De Cáceres and Wiser 2012) Approach used   Membership determination. How do I group my vegetation observations? or Which type does this particular observation belong to?  Clustering relevés based on TWINSPAN results combined with tabling procedures (Appendix A Tables 14, 15 and 16 and Appendix B Tables 30 to 44). The resulting vegetation groups were verified using Hierarchical Cluster Analysis (Appendix A Figures 47 to 49)                                                  10 Preference was given to Non-metric dimensional scaling (NMDS), but Principal Component Analysis (PCA), Detrended Correspondence Analysis (DCA), and Canonical Correspondence Analysis (CCA) were also used to look for the best fit of vegetation clusters and to find correlations between vegetation and environmental factors.   40  Definition of vegetation types (De Cáceres and Wiser 2012) Approach used   Characterization. What are the attributes of my grouping?  Using indirect gradient analysis of site environmental attributes through ordinations (Appendix A Figures 23 to 46), factor analysis of soil variables (Appendix A Figures 58 to 60 and Tables 23->29) and diagnostic character species determination (Appendix A Tables 17 and 18)  Validation. Is this grouping acceptable for my purpose? I followed the required combination of diagnostic character species for associations according to Pojar et al. (1987), Klinka and Chourmouzis (2001) and Mackenzie (2012) with slight modifications (e.g. the use of absolute frequency for cut-off limits).  I also conducted field validation of the units (i.e. do the assigned units occur in the landscape at a scale readily identifiable in the field?)  Naming. How do we refer to this grouping?  Associations were  named according to the naming conventions of Klinka and Chourmouzis (2001), Mackenzie (2012), and (Jennings et al. 2009).  The first part of the process, determining membership for vegetation units, would ideally always produce the exact same results, but one of the limitations of using TWINSPAN is that it does not always reproduce the original membership statements (De Cáceres and Wiser 2012). Also, the different options provided in the modified TWINSPAN (Roleček et al. 2009) and the different choices of pseudospecies cut levels, modified the resulting membership assignments (Appendix B Table 1 to 16  ). Therefore, following the definitions of Table 3, my procedure can be considered as „indicative assignments‟ that serve as precursors for „consistent assignments‟, which are obtained from the tabling procedures. Following the suggestion by (De Cáceres and Wiser 2012) that TWINSPAN should be modified to provide membership rules based on the location of plots on ordination axes, every step of the TWINSPAN results was overlaid in various ordination bi-plots (Appendix A Figures 23 – 46).    41  Table 4 provides a list of the resulting figures and tables after each step of the data analysis methodology. This procedure (explained below) was used to reduce the original dataset (300 relevés with several environmental variables) into the final classification units presented in Tables 5 and 6. These units were used as the basis for the descriptions presented in landscape diagrams, edatopic grids and site unit keys (Appendix A Figures 50-57 and Tables 19-22). The soil analysis (Appendix A Figures 58-66 and Tables 23-29) was used to further explore the factors leading to the formation of the plant associations observed in the classification.     Table 3 Activities related to the determination of membership to vegetation types (from (De Cáceres and Wiser 2012)       Similarities and differences within and among vegetation clusters were identified using presence classes (categorical frequency) and species significance (median % cover for each group).  Each group was then classified as an association or a subassociation depending on their relationship to the observed hierarchy (Brett et al. 1998), which was determined by the amount of overlap of the dominant species between the original partitions and the groups under analysis. After associations and subassociations were identified, diagnostic tables were produced following the criteria for differential and dominant-differential species (see Appendix A Table 12 for definitions and criteria). The previous procedure was repeated until unique diagnostic combinations of species were obtained.  Table 3 has been removed due to copyright restrictions. It showed the activities related to the determination of membership to vegetation types. Original Source: De Cáceres, M. and S. K. Wiser. 2012. Towards consistency in vegetation classification. Journal of Vegetation Science 23:387-393.   42   Synthesis of the vegetation is the most important step in ecosystem classification (Pojar et al. 1987); BEC closely follows the Braun-Blanquet approach to vegetation synthesis and the descriptive techniques of the Zurich-Montpellier School (see Shinwell (1972) for a thorough description of the ZM approach). Some procedural and conceptual modifications11 proposed by MacKenzie (2012) were taken into consideration in my approach. Likewise, the variants on the procedures described by Brett et al. (1998) and Klinka and Chourmouzis (2001) were used in the synthesis of the vegetation. Tables 12 and 13 (Appendix A) describe some of the criteria and definitions used.  I followed Jennings et al.‟s (2009) definition for alliances which is: a vegetation classification unit containing one or more associations, and defined by a characteristic range of species composition, habitat conditions, physiognomy, and diagnostic species, typically at least one of which is found in the uppermost or dominant stratum of the vegetation. If associations had species in common in the dominant layer and those same species were absent or infrequent in other nearby associations, the associations which shared dominant species were joined to form an alliance (Jennings et al. 2009). Plant alliances with physiognomically similar vegetation and general affinities in a dominant stratum are grouped into orders which represent broad segments of the edaphic gradient that usually occur in many different climates and each order is represented by one or two prominent climax species (Pojar et al. 1987). Following the approach of Klinka and Chourmouzis (2001), my classification assumes that the alliances are part of the Engelmann-Spruce Sub Alpine Fir and the Mountain Heather orders; therefore, I am equating the Plant Order hierarchical level to the BEC zone level. Figure 15 further explains how the vegetation classification (class, order, alliance, association and subassociation) relate to the BEC zonal classification (zone, subzone, variant).                                                        11 For example the use of an edatopic grid for site associations in addition to the one for site series; and the use of moderately steep slopes (since excessive snow accumulation and soil moisture on flat sites do not properly reflect the alpine climate) for zonal sites.  Figure 15 has been removed due to copyright restrictions. It showed the Biogeoclimatic Ecosystem Classification framework. Original Source:  MacKenzie, W.H. 2012. Biogeoclimatic ecosystem classification of non-forested ecosystems in British Columbia. Prov. B.C., Victoria, B.C. Tech. Rep. 068  43    Figure 15 The Biogeoclimatic Ecosystem Classification framework. New upper-level site units are highlighted in grey from Mackenzie (2012).    During the tabulation process, I followed Pojar et al.‟s (1987) criteria for dividing and assigning vegetation units.  This considers for the Diagnostic Combination of Species (DCS) that: (1) character-species (i.e., the species that differentiate in the absolute sense) are not required in the DCS for any vegetation unit at present; (2) units are recognized by an exclusive DCS that must include at least one differential or dominant-differential species; (3) units that represent the central concept of a higher circumscribing unit are also recognized without a DCS, providing they are differentiated by the absence or low occurrence of species that characterize other units of the same category and circumscription; and (4) plant subassociations are also recognized by non-exclusive DCS's that include at least two differential or dominant-differential species.  Klinka and Chourmouzis (2001) further considered that: a diagnostic species must be either a differential species, which has a much higher presence (proportion of plots of a group in which it occurs) than in other groups, or a dominant differential species, which has higher species significance (percent cover) than in other groups. The exact criteria identified by Klinka and Chourmouzis (2001) for the character-defining species are: Differential species: a species that may be associated with more than one vegetation unit in a hierarchy; has a than in other units of the same hierarchical level within the same higher level unit. Dominant-differential species: a species that may be associated with  more species significance classes greater than in other units of the same hierarchical level within the same higher level unit.  Three different levels of classification were produced in this thesis: a) Site series, b) Site associations, and c) Site Alliances (see Figure 15). The site series and alliances were derived from the procedures described above, while the site associations (for the IMAwc) were mostly developed by W.H.  Mackenzie (unpublished) with some input from me. Also, based on Mackenzie‟s (2012) approach and greatly influenced by the  44  results of my synthesis, we developed the site associations for the ESSFwcp. The information obtained from the soil analysis was used to search for gradients in the distribution of plant communities through principal component and canonical correspondence analysis.    I used the following procedure using my relevés (130), the combined dataset (300 relevés), and also individual zones (IMAwc =110 relevés and ESSFwcp= 190 relevés)12 to produce the classification: 1. Using JUICE Vegetation Software (Tichý 2002), I did the first amalgamation of relevés using TWINSPAN and ISOPAM analysis. For each method, the dataset was used with Braun-Blanquet values or with presence/absence values.  Since this was the first exploration of the dataset, different methods and dataset value-transformations were used to find the best fit.   a. For the modified TWINSPAN (Roleček et al. 2009) analysis I used combinations of the options below: i. Whitaker‟s beta-diversity or  Sørensen‟s dissimilarity indices ii. No. of clusters (range for different iterations): 4-18 iii. Pseudospecies cut levels 0 5 10 20 50 or 0 5 10 30 50                                                  12 IMAwc relevés chosen by elevation  m.a.s.l..> 2100m and for ESSFwcp relevés elevation  m.a.s.l. >1900m and < 2100m (Trowbridge et al. 2002)  45   Figure 16 Example of the Modified TWINSPAN parameter selection.  b. And for the ISOPAM analysis: (Schmidtlein et al. 2010) i. Hierarchical clustering-maximum depth: 10  ii. Optimize cluster no. partition with max. no. of clusters = 6  iii. Threshold for species to be considered diagnostic: 3.5 iv. Vector with stopping rules for hierarchical clustering: c(1,7) v. Distance measures: Bray-Curtis   46   Figure 17 Example of the options selection for ISOPAM.  2. From the above procedures, synoptic tables were produced for each step (for example: step „i‟= modified TWINSPAN with pseudospecies cut levels 0 5 10 20 50 selecting Whitaker‟s diversity and maximum cluster no. = 9) and groups were tabulated selecting species with presence class greater than III (frequency > 41%) and sorted by absolute frequency. From the resulting tables, the species with the highest presence class was selected to name the association and when there were more than one species within the same presence class, the species with highest absolute frequency was chosen.  3. The tables obtained from (2.) were compared for redundancy of associations and visual affinity to field observations.  4. The most accurate clustering method (based on least redundancies for dominant and dominant-differential species and on my perceived fidelity to field observations) was selected and used for further analysis.   47  5. The groups from (4.) were used in ordinations (PCA, DCA, CCA and NMDS) to probe the data structure and develop hypotheses regarding species distribution and vegetation-environment interactions (Pojar et al. 1987). The four abovementioned ordination methods were used in every step of the divisions. Different methods were chosen depending on whether the preference was for strength of correlation to environmental variables or whether tight clustering in vegetation was the main interest. When this step resulted in different associations, the most consistent associations (i.e. those that appeared in more than one ordination method) were chosen.  6. When the same differential species was present in more than one group, associations from step (4.) were regrouped based on the common differential species; in other words, the regrouping resulted as a merger between associations based on the overlapping differential species.  7. The groups resulting from (6.) were tabulated and sorted by species‟ absolute frequency. All species with absolute frequency < 2 were deleted individually (the relevés were kept) and the modified TWINSPAN was performed on this subset, specifying the reduced number of clusters (e.g. for the IMAwc from 9 clusters in the analysis at step (2.) to 5, based on the regrouping from step 6).  8. Groups from (7.) were tabulated and inspected for redundancy in differential species. If dominant-differential species were found in more than one group, these groups were merged.  9. The environmental factors for groups from step (8.) were reviewed for consistency, and outlier relevés were deleted if their elevation was < 1700m.a.s.l.13.  10. The groups from step (9.) were used in the same ordinations as in (5.) to identify correlations between environmental gradients and plant associations.  11. The groups resulting from step (9.) were tabulated and all species with presence class < III were deleted. When differential species were present in more than one                                                  13 Although 1700m.a.s.l. is well below the 1900m. criterion for parkland in the ESSFwcp, it was chosen as a cut off since several of the plots provided by the MoFR (classified as „parkland‟) were within that elevation range.   48  group they were associated only with the group in which they had the highest combination of presence class, species significance and absolute frequency. The resulting groups were labelled „Alliances‟.  12. The alliances from step (11.) were used for DCA and NDMS ordinations to search for groupings of patterns along environmental gradients.     49   Chapter 3 Results Classification Table 4 provides a list of the resulting figures and tables after each step of the data analysis. This procedure (explained below) was used to reduce the original dataset (300 plots with several environmental variables) into the final classification units presented in Tables 5 and 6. These units were used as the basis for the descriptions presented in landscape diagrams, edatopic grids and site unit keys (Appendix A Figures 50-57 and Tables 19-21). The soil analysis (Appendix A Figures 58-66 and Tables 23-29) was used to further explore the factors leading to the formation of the plant associations observed in the classification.  Table 4 List of Results available after each step of the data analysis methodology   Step in the classification data analysis   Corresponding Results  (Appendix) 1.      Using JUICE Vegetation Software (Tichý 2002) I did the first amalgamation of relevés using TWINSPAN and ISOPAM analysis.   Table 1-3 (Appendix B)   2.     From the above procedures synoptic tables were produced for each step and groups were tabulated selecting species with presence class greater than III (frequency > 41%) and sorted by absolute frequency. From the resulting tables, the species with the highest presence class was selected to name the association and when there were more than one species within the same presence class, the species with highest absolute frequency was chosen.   Table 4-16 (Appendix B) 3.      The tables obtained from (2.) were compared for  N/A  50       redundancy of associations and visual affinity to field observations.   4.      The most useful clustering method was selected and used for further analysis.  Modified TWINSPAN using Whitaker‟s beta-diversity index, pseudospecies cut levels 0 5 10 20 50 and max. no. of clusters = 9 for individual zones –IMAwc or ESSFwcp- or max. no. of clusters = 18 for combined zones -–IMAwc + ESSFwcp-)  5.      The groups from (4.) were used in ordinations (PCA, DCA, CCA and NMDS) to probe the data structure and develop hypotheses regarding species distribution and vegetation-environment interactions (Pojar et al. 1987). Different methods were chosen depending on whether the preference was for strength of correlation to environmental variables or whether tight clustering in vegetation was the main interest.    Figures  23-26 (Appendix A) 6.      When the same differential species was present in more than one group, associations from step (4.) were regrouped based on the common differential species.  Tables 14-17 (Appendix A) 7.      The groups resulting from (6.) were tabulated and sorted by species‟ absolute frequency. All species with absolute frequency < 2 were deleted and the modified TWINSPAN was performed on this subset, specifying the reduced number of clusters (e.g. for the IMAwc from 9 clusters in the analysis at step (2.) to 5, based on the regrouping from step (6).   N/A 8.      Groups from (7.) were tabulated and inspected for redundancy in differential species. If dominant-differential species were found in more than one group, these groups were merged.   N/A Step in the classification data analysis     Corresponding Results  51       9.      The environmental factors for groups from step (8.) were reviewed for consistency, and outlier relevés were deleted if their elevation m.a.s.l. was < than 1700m.        N/A     10.      The groups from step  (9.) were used in the same ordinations as in (5.) to identify correlations between environmental gradients and plant associations.  Figures 27– 41 (Appendix A) 11 The groups resulting from step (9.) were tabulated and all species with presence class < III were deleted. When differential species were present in more than one group they were associated only with the group in which they had the highest combination of presence class, species significance and absolute frequency. The resulting groups were labelled „Alliances‟.   Tables 17 and 18 (Appendix A) 12.  The alliances from step (11.) were used for DCA and NMDS ordinations to search for groupings of patterns along environmental gradients.  Figures 42-46 (Appendix A)  Figures 18, 19 and  20 show three different ordination methods (NMDS, DCA and CCA) for the 130 relevés from 2007 and 2008, overlaid with the first TWINSPAN resulting in 18 groups. NMDS and DCA produced similar results in that % rock cover, altitude and organic matter were the environmental variables that met the minimum correlation criterion (r2≥ .2, default in PC-ORD), and the TWINSPAN groups are arranged similarly along the axis. The canonical correspondence analysis (Appendix A Figure 25) differed from NMDS and DCA, in that aspect, % slope, and altitude were the environmental variables that met the minimum correlation criterion (same as above). To explore the arrangement of the vegetation across different aspects, I overlaid aspect as a grouping variable. Figure 19 shows how species are arranged in different aspects; there are few notable patterns with the exception of the wet and snowbed species (e.g. Caltha Step in the classification data analysis     Corresponding Results  52  leptosepala and Carex nigricans) which aligned with the „flat‟ aspect (red triangles in the top half of Figure 19). For the three ordinations, one of the consistent patterns is the segregation of alpine species (e.g. Lepraria neglecta, Cassiope tetragona, Carex phaeocephala, Silene acaulis) from subalpine species (e.g. Heracleum maximum, Senecio triangularis, Viola glabella, Mitella spp., Abies lasiocarpa) which reaffirms the division between the IMAwc and the ESSFwcp.   Figure 18 Detrended Correspondence Analysis (DCA) for 130 relevés. Green dots are species. „orgmatter‟ = Percent surface cover of organic matter. „rocks = pct. surface cov. of rocks     53   Figure 19 Canonical Correspondence Analysis for 130 relevés with the aspect overlaid.  Figure 20 shows the NMDS ordination for all 300 relevés, and it shows that at this scale only altitude met the minimum correlation criterion. From Figure 20 a few plant communities are clearly defined such as the wet Sphagnum-Eriophorum communities (bottom left side of the ordination). As with the previous ordinations, the IMAwc communities (e.g. TWINSPAN groups 7 and 2) are well segregated from the ESSFwcp communities (e.g. TWINSPAN groups 3 and 5).   54   Figure 20 NMDS for all 300 relevés and 18 TWINSPAN divisions.  Table 23 (Appendix A)  shows the tabulation for the ESSFwcp using 190 relevés and 9 TWINSPAN groups; from this table, a few useful patterns begin to emerge, for example the Cassiope mertensiana- Luetkea pectinata (group 2) and the Carex spectabilis – Valeriana sitchensis (group 9) associations. However, at this level (9 groups) several redundancies are present, most notably Carex nigricans as the dominant or co-dominant species for groups 5, 6 and 7, so this overlap provides a good basis for merging groups. Group 3 Salix barrattiana is comprised of only 1 relevé which was located at 1515m.a.s.l, therefore it is considered as an outlier and discarded for further analysis.  Table 24 (Appendix A) shows the tabulation for the IMAwc using 110 relevés and 9 TWINSPAN groups; in this tabulation, Cassiope mertensiana is the dominant or co-dominant species for groups 1 through 4 which were merged for further analysis. Likewise, Artemisia norvegica is the dominant or co-dominant species for groups 8 and 9, and is also strongly present in groups 5 through 7; therefore, these groups were merged  55  for further analysis. The merging of groups in Table 23 and 24 (Appendix A) (4 groups merged and one deleted) was used only to define the new number of clusters to be selected in the next TWINSPAN. The species list was reduced based on the absolute frequency of species (absolute frequency ≥ 3) and a new TWINSPAN was performed for the IMAwc. The results produced in Table 23 (Appendix A) for the ESSFwcp was satisfactory enough (the redundancies in dominant species were eliminated by merging groups) that simply merging groups and deleting species below the absolute frequency criterion was sufficient to produce the final tabulation.  For exploratory purposes the same process of TWINSPAN, tabulation and group merging was applied to the first 130 unsegregated (IMAwc and ESSFwcp plots were not divided) relevés (Appendix B Table 16) and various ordinations were performed; a total of 7 groups and a species reduction based on absolute frequency ≥ 2 was deemed appropriate due to the smaller size of this dataset. Figure 27 (Appendix A) shows the PCA, for the 130 relevé dataset, in which altitude, percent rock cover, and percent organic matter met the minimum correlation criterion (r2 = .2); group 1 (Artemisia norvegica – Arnica latifolia) and group 5 (Cassiope mertensiana-Phyllodoce empetriformis) are clustered to the right hand side of axis 1 and are inversely correlated to altitude and positively correlated with increasing percent cover of organic matter, while group 2 (Artemisia norvegica – Lepraria neglecta) is clustered to the left of axis 1 and positively correlated with increasing altitude and percent rock cover. This pattern of clustering reflects the division between ESSFwcp and IMAwc along an altitudinal gradient but it is important to note that the resulting associations differ in species composition from the associations developed from the analysis performed on the larger dataset, in which relevés were segregated by zone before the analysis.  Figures 28, 29 and 30 (Appendix A) confirm the segregation described above, but the degree of clustering varies slightly depending on the ordination method. Figure 21 (Appendix A) shows the CCA for the same dataset and it differs from the previous ordinations in that in addition to the three variables (altitude, percent rock cover, and percent organic matter), Soil Moisture Regime (SMR) and percent cover of water met the minimum correlation criterion(r2= .2). These additional variables help explain the location of group 7 (Abies  56  lasiocarpa – Caltha leptosepala) which is positively correlated to increasing SMR and percent cover of water (mid-bottom left side of the ordination‟s centre). Figure 22 (Appendix A), the 3D representation of the NMDS analysis, shows the clustering of group 5 around the 3rd and 1st axis while group 1 is primarily clustered along the 1st and 2nd axis; in 3D, the correlations with the environmental variables are slightly better represented for group 5 by showing its intermediate position between increasing percent rock cover and increasing altitude which helps explain the wide range of habitats that is occupied by the  Cassiope mertensiana-Phyllodoce empetriformis association.  The following sets of ordinations (Appendix A Figure 23 to 41) were done using the intermediate stage of tabulations (step 10) for the IMAwc and ESSFwcp, which were used for the final classification. The main purpose of these ordinations is to illustrate how as groups are merged the clusters become tighter and better ordered around environmental variables. Figure 33 (Appendix A) the DCA for the 110 relevés from the IMAwc resulted in well-defined clusters of the TWINSPAN groups but only percent cover of rocks met the correlation criterion (r2=.2) for environmental variables. Groups 1, 2 and 3 are inversely correlated to percent rock cover and they represent the wetter and lusher (snowbed, meadow and mesic) communities, while groups 4 and 5 are positively correlated to percent rock cover and they represent the drier and sparser (e.g. exposed ridges and fellfields) communities. Figure 34 (Appendix A), shows that in the PCA for the same dataset, group 5 is inversely correlated to percent cover of organic matter while the other groups show a weak correlation to percent organic matter. Figure 35 (Appendix A), the NMDS analysis shows weakly defined clusters for each group yet there is a clearly defined trend in which groups 1 and 2 are positively correlated to percent organic matter while groups 4 and 5 are positively correlated to percent rock cover; this further confirms the trends observed in Figures 33 and 34 (Appendix A). In the CCA for the same dataset (Appendix A Figure 36) in addition to percent rock cover and organic matter, elevation (m.a.s.l.), rooting depth, percent bedrock and percent cover of mineral soil met the minimum correlation criterion (r2= .2). These additional environmental variables further highlight the affinity of group 5 to increasing elevation and the positive correlation of group 2 with increasing rooting depth which strengthens the trends  57  observed in Figures 33, 34, and 35 (Appendix A). The 3D representation of the CCA (Appendix A Figure 37) for the IMAwc dataset illustrates that group 5 is inversely correlated to rooting depth, percent organic matter, and percent cover of water, that it is intermediate in percent cover of bedrock and mineral soil and positively correlated with elevation and percent rock cover. Figure 37 (Appendix A) also serves to highlight that group 1 is primarily correlated with increasing organic matter, and that group 2 is best explained by a positive correlation with rooting depth.  From the above ordinations it became apparent that the variability in distribution of the plant communities in the IMAwc could be better explained by further reducing the amount of associations by merging the groups that responded similarly to environmental gradients; therefore groups were merged based on the arrangement of patterns across the environmental gradients and on co-occurrences of differential and dominant-differential species.  The following sets of ordinations (Appendix A Figures 38 to 41) represent the exploration of patterns across environmental gradients for an interim reduction of 9 TWINSPAN groups to 5 groups for the ESSFwcp. Unlike the above analysis for the IMAwc, tabulation for this step was not deemed necessary since the patterns were sufficiently well defined by the ordinations, so that it became apparent that the variability explained would not vary considerably between 5 and 4 groups. Nonetheless, the ordinations are presented since they were used to confirm that enough variability would be explained by merging nine groups into four.  Figure 38 (Appendix A) shows the PCA for the 190 relevés for the ESSFwcp and 5 TWINSPAN groups; most notably the wet community (group 4) is positively correlated with the first Axis and it is orthogonal to the other four groups. The 3D representation of the DCA for the same dataset (Appendix A Figure 39), shows a tight clustering of groups 2, 3, 5 and 6 which are adjacent to each other and primarily correlated to Axis 3, while group 4 is scattered. It is interesting to note that in the PCA group 4 was the only one to show a defined response to an axis (presumably moisture) yet in the DCA it was the only group to be poorly explained by the ordination; from this it can be understood that groups 2, 3, 5 and 6 are best explained by a combination of environmental factors other than moisture. The NMDS analysis for the ESSFwcp (Appendix A Figure 40) shows well  58  defined clusters for each group, yet no environmental variables met the minimum correlation criterion (r2= .2). Regardless, it is interesting to observe that groups 4 and 2 lie in opposite sides of the plot as do groups 5 and 3 while group 6 is intermediate to the above groups. Yet, the lack of correlation with any single environmental gradient limits the generalization of this observation and all that can be confirmed is that the corresponding plant communities are well segregated in the landscape and are responding to different combinations of environmental factors. The CCA for the ESSFwcp (Appendix A Figure 41) was the only analysis in which some environmental factors  (seepage depth, percent cover of water and altitude) met the minimum correlation criterion (r2= .2). From Figure 21 it becomes apparent that group 4 is positively correlated to moisture (seepage depth and percent cover of water), that group 5 is inversely correlated to altitude while group 2 is positively correlated to altitude; the relation of plant communities to these observations becomes better defined in the last set of ordinations and through the final tabulation.  Table 17, (Appendix A) the final tabulation for the IMAwc shows the final grouping of the previously defined associations into alliances. Three alliances were identified: Cassiope mertensiana, Carex spectabilis and Artemisia norvegica; broadly, these could be defined as wet, mesic and dry communities or late, intermediate and early snowmelt communities, respectively. Table 18, (Appendix A) shows the final tabulation for the ESSFwcp which arranges the associations into the: 1) Carex nigricans, 2) Philonotis fontana, 3) Cassiope mertensiana-Luetkea pectinata and 4) Abies lasiocarpa-Valeriana sitchensis alliances; roughly the gradient explained by these alliances is: 1) water saturated (snowbeds), 2) wet (late snowmelt), 3) mesic (intermediate snowmelt), and 4) dry (early snowmelt) communities. Figure 42 and 43 (Appendix A) shows the NMDS analysis for the alliances of the IMAwc and their relation to percent rock cover which was the only environmental variable to meet the minimum correlation criterion. In Figure 42 (Appendix A) it can be observed that Alliance 3 (Artemisia norvegica) has a minimal overlap with Alliance 1 (Cassiope mertensiana) and 2 (Carex spectabilis), while Alliances 1 and 2 show a considerable overlap and are inversely correlated to percent rock cover. By further  59  reducing the dataset so that only dominant and dominant-differential species are considered, the alliances become better segregated with no overlap between Alliances 1 and 3, while Alliance 2 occupies an intermediate position along Axis 1 (Appendix A Figure 43); this confirms the position of these communities from a wet to dry gradient. Figure 44 (Appendix A), the DCA for the alliances of the IMAwc, confirms the separation between Alliances 3 and 1 along the first axis and further segregates Alliance 2 along the second axis. Furthermore, in Figure 24 it can be noted that only the second alliance shows a negative correlation with percent rock cover which would help explain the lush character of the mesic plant communities.  The ordinations for the alliances of the ESSFwcp resulted in well-defined clusters for each alliance in which only the second and third alliance overlap. Figure 45 (Appendix A) shows the NMDS analysis for these alliances and it clearly segregates Alliance 4 (Abies lasiocarpa – Valeriana sitchensis) on the left side of the ordination with an inverse correlation to seepage depth while Alliance 2 (Philonotis fontana) is positively correlated to seepage depth on the right side of the first axis. Alliance 1 (Cassiope mertensiana – Luetkea pectinata) is positioned below Alliance 4 along the second axis and is moderately inversely correlated to seepage depth. Alliance 3 (Carex nigricans) has a considerable overlap with Alliance 2 and it is positive correlated with seepage depth. Figure 46 (Appendix A), the CCA for the alliances of the ESSFwcp, shows the location of the alliances in relation to altitude, percent surface cover of water, mineral soils and rocks, percent slope gradient and seepage depth; although the clusters for each alliance are weaker than in Figure 25, their position along environmental factors is more explicit than in the NMDS. With the CCA, it is confirmed that Alliance 2 is positively correlated to seepage depth and percent surface cover of water and that Alliances 4 and 1 are positively correlated with altitude, percent slope gradient, percent cover of mineral soils and rocks, which confirms their position in the landscape as exposed, water shedding sites.   To independently confirm the grouping patterns from the alliances derived from TWINSPAN and tabulation procedures, hierarchical cluster analyses were applied to  60  each vegetation dataset (my ESSFwcp and IMAwc 130 relevés, the total IMAwc‟s 110 relevés and the total ESSFwcp‟s 190 relevés). Figures 47, 48 and 49 (Appendix A) confirm the grouping patterns through the cluster analysis and they show the distribution of species per alliance. Although it is not a perfect fit, most relevés were placed in the same group as they were in the TWINSPAN/tabulation procedure, which further indicates that the associations and alliances are real patterns that explain the distribution of plant communities throughout the landscape.  Tables 5, 6, and 7 show the synopsis for the alliances and their relation to site units and site series. The nomenclature (e.g. 103 At71 = Angled mountain heather-Crowberry) is described in Tables 20  and  22 (Appendix A). These are further explained in Figures 50 -57 (Appendix A) which show the theoretical arrangement of the associations along soil moisture and snowmelt gradients as well as their expected position in the landscape. Tables 19 and 21 (Appendix A) show the key to identifying site units in the field (following the format by Steen and Coupe 1997), based on a combination of topography and plant community characteristics; Tables 20 and 22 (Appendix A) provide a brief description of these site associations. Although they are gross simplifications of the results, their primary purpose is to facilitate the identification of plant communities in the field based on observations that do not require a high degree of expertise; for further information on how to use these keys, diagrams and descriptions, refer to Steen and Coupe (1997).  61  Table 5 Hierarchical relationship between alliances, site associations, site series and TWINSPAN groups in the ESSFwcp that were found in the present study.. Presence classes (PC) as percent frequency are shown in roman numerals (I = 1-20, II = 21-40, III= 41-60, IV = 61-80, V = 81-100) and species significance (SS) is shown in percent cover. Refer to Table 21 Appendix A for the site unit nomenclature    62  Table 6 Hierarchical relationship between alliances, site associations, site series and TWINSPAN groups in the IMAwc that were found in the present study. Presence classes (PC) as percent frequency are shown in roman numerals (I = 1-20, II = 21-40, III= 41-60, IV = 61-80, V = 81-100) and species significance (SS) is shown in percent cover.  TWINSPAN No. 1 2 3 4 5 SiteNo. of Releves 16 21 21 21 31 SeriesPrescence SpeciesAlliance Cassiope mertensiana Class Significance Cassiope mertensiana Phyllodoce empetriformis Luetkea pectinata Site Association 111 Ah03Cassiope mertensiana V 10 V 15 III 5 II 1 I 2 03Phyllodoce empetriformis III 3 II 4 III 7 I 5 I 2 03Luetkea pectinata III 5 IV 3 II 3 I 2 I 4Carex nigricans Site Association 112 As01Carex nigricans V 35 II 1 II 2 . . . .Carex pyrenaica Luzula piperi Site Association 113 As10Carex pyrenaica . . III 3 . . II 1 I 2Luzula piperi II 5 IV 1 I 2 I 1 I 2 03Allience Carex spectabilisCarex spectabilis Antennaria lanata Arnica latifolia Site Association 101 Am03Carex spectabilis II 2 III 4 V 5 I 1 II 2 05Antennaria lanata II 1 IV 7 IV 15 II 1 II 2Arnica latifolia I 4 II 1 III 3 . . I .Allience Artemisia norvegicaArtemisia norvegica Carex phaeocephala Site Association          105 At29Artemisia norvegica . . II 2 III 7 IV 2 V 1 01Carex phaeocephala . . . . I 2 I 3 I 1Artemisia norvegica Antennaria lanata Site Association                    106 At22Antennaria lanata II 1 IV 7 IV 15 II 1 II 2 01Silene acaulis Salix nivalis Potentilladiversifolia Site Association 102 Af01Silene acaulis . . I 1 . . IV 1 IV 1Potentilla villosa . . . . . . I 1 I 2 04Salix nivalis . . I 1 I 1 IV 5 II . 04Sibbaldia procumbens . . III 1 II 1 III 2 IV 3 04Cassiope tetragona Empetrum nigrum Site Association                       103 At71Cassiope tetragona . . . . I 1 II 1 III 20Empetrum nigrum . . . . I 1 I 1 III 02Festuca brachyphylla I 1 I 1 I 2 III 1 III 3 02Lepraria neglecta . . I 5 II 10 . . V 30 02Salix nivalis Dryas octopetala Site Association              103 At29Dryas octopetala . . I 1 . . III 6 . 63     Table 7 Synopsis of all plant alliances and associations found to date, occurring within the ESSFwcp and IMAwc. Citations are included for previously described vegetation units and n.n. (nomen novum) is added if the name, hierarchy change, or vegetation unit is not previously published. Only Orders follow phytosociology nomenclature, the other levels are named with their Latin name, following Brett et al. (1998).     Order     Alliance      Association                          Phyllodoco-Cassiopion  (Brooke 1965)  Cassiope mertensiana ( Brett et al. 1998)   Anemone occidentalis- Cassiope mertensiana n.n.     Luetkea pectinata- Cassiope mertensiana n.n.    Phyllodoce empetriformis- Luetkea pectinata- Cassiope mertensiana n.n.; (Brooke et al. 1970)   Carex pyrenaica- Luzula piperi n.n.    Carex nigricans n.n.; (Archer 1963)  Artemisia norvegica n.n.    Carex phaeocephala - Artemisia norvegica n.n    Antennaria lanata - Artemisia norvegica n.n.    Silene acaulis- Salix nivalis - Potentilla villosa n.n.    Cassiope tetragona- Empetrum nigrum n.n.    Salix nivalis - Dryas octopetala n.n.   Carex nigricans (Brett et al. 1998)   Carex nigricans (Archer 1963)   Aulacomnium palustre- Caltha leptosepala- Carex nigricans n.n.    Eriophorum angustifolium- Carex nigricans n.n.   Carex spectabilis (Brett et al. 1998)   Antennaria lanata - Carex spectabilis n.n.      Abietion lasiocarpae n.n.    Abies lasiocarpa n.n.    Cassiope mertensiana- Barbilophozia floerkei- Abies lasiocarpa n.n.   Juniperus communis- Empetrum nigrum- Abies lasiocarpa n.n.    Carex spectabilis- Valeriana sitchensis - Abies lasiocarpa n.n.   Valeriana sitchensis- Anemone occidentalis    Carex spectabilis- Valeriana sitchensis- Erigeron peregrinus n.n.          64   Correlations with soil properties  I chose to do a PCA with all variables (Appendix A Table 23) to search for correlation patterns between the vegetation and the edaphic variables. Table 23 (Appendix A) shows the result from the PCA and Figure 58 (Appendix A) shows the loading plot for principal components 1 and 2 (cumulative explained variation = 60%). In Figure 58 (Appendix A) it is interesting that slope gradient is inversely correlated with aspect and elevation and that pH, NO3-N and NH4-N are orthogonal to aspect, elevation and slope; as such, we can determine that the first principal component represents the edaphic variables while the second component represents topographic variables. This correlation pattern is confirmed in Table 24 (Appendix A) which shows the eigenvector loading for each principal component. A Varimax rotation was chosen for the factor analysis and the final communality estimates are shown in Table  25 (Appendix A) which indicates that only Organic Matter, NH4-N and NO3-N account for a significant proportion of the variance explained (.89, .59 and .76 respectively). An Oblique Quartimax rotation was applied for the first two principal components (Appendix A Figure 60) which resulted in similar patterns as the Varimax rotation, yet the Factor Structure (Appendix A Table 27) shows that the abovementioned variables are primarily loaded in the first principal component, while the second principal component was heavily weighted by aspect. Table 27 (Appendix A) also shows that only Organic Matter, NH4-N and NO3-N met the pre-established minimum correlation criterion of r2 ≥.7, so these are the variables that were selected for a new factor analysis with reduced variables. The value of reducing the amount of variables is evident when we compare Table 24 (Appendix A), in which it requires 8 principal components to account for all the variation in the data, Table 27 (Appendix A), in which the same amount of variation is explained by less than half of the number of principal components. By selecting the variables that met the r2 ≥ .7 correlation criterion (Appendix A Table 27), the number of variables was  65  reduced to three from eight. Therefore, the value of the Factor Analysis is that we can explain most of the variation (94%) using only two principal components (Appendix A Table 27) and three variables (Appendix A Table 28 and 29). Figure 60 (Appendix A)  shows the new rotated factor pattern with the three abovementioned variables, and it shows that the nitrogen variables load on the second principal component and are orthogonal to the first principal component which accounts for Organic Matter. With this in mind, the same dataset (Appendix A Table 28) was used in ordinations to search for the links between vegetation and edaphic/topographic variables. The CCA for the dataset (Appendix A Figure 61) chosen after the Factor Analysis, displays NH4-N and NO3-N in different planes which are correlated with different plants. From Figure 62 (Appendix A) it is interesting to note that Caltha leptosepala is strongly influenced by NO3-N, that Erigeron peregrinus, Salix nivalis and Dryas octopetala are closely related to NO3-N while Abies lasiocarpa, Mitella spp., Carex spectabilis and Carex nigricans are positively correlated to NH4-N. However, Arnica latifolia, Cassiope mertensiana, Phyllodoce glanduliflora, Cassiope tetragona and Empetrum nigrum are inversely correlated to NH4-N and NO3-N. These patterns can further be explained by the findings of  Miller and Bowman (2003) in which soil moisture accounted for 60% of the variation in exchangeable NH4-N in wet meadows and that exchangeable NO3-N showed a positive correlation with soil temperature, but not with soil moisture. Surprisingly, none of the abovementioned species are strongly correlated with organic matter which is different from the observations made in the ordinations (e.g. Appendix A Figure 36) from the relevé dataset.  Figure 63 (Appendix A), the PCA for the reduced dataset (using only the three variables chosen from the Factor Analysis) shows that Organic Matter met the minimum correlation criterion (r2= .2) with the vegetation matrix. Figure 63 (Appendix A) was useful to observe the segregation of ESSFwcp species (right hand side) from the IMAwc species (left hand side) and the positive correlation of the species from lush communities (wet and mesic) with organic matter, while species from the dry and exposed sites (e.g.  Lepraria neglecta (Leprnnegl), Cassiope tetragona (Casstet) and Phyllodoce glanduliflora (Phylglan)) are inversely correlated with organic matter. By ignoring the results from the factor analysis and using the full dataset of edaphic,  66  topographic and vegetation variables, a few important patterns are revealed. For example, in the CCA with all variables (Appendix A Figure 61) in addition to the three abovementioned variables, pH and elevation met the minimum correlation criterion (r2= .2). Elevation is helpful in distinguishing ESSFwcp from IMAwc species and pH shows a correlation with Dryas octopetala, Saxifraga bronchialis, Carex nardina, and Draba spp. A PCA for the full dataset (Appendix A Figure 64 and 65) shows tighter clusters which segregate ESSFwcp and IMAwc species with an orthogonal relation and it indicates that species from wet and mesic communities show a positive correlation to the nitrogen variables. When analysing the third and first principal components (Appendix A Figure 66), pH and elevation meet the minimum correlation criterion which helps illustrate the plant-environment correlations. An NMDS analysis for the full vegetation-environment-edaphic dataset (Appendix A Figure 66), results in pH and elevation as the only variables that meet the minimum correlation criterion (r2= .2) and strengthens the observation of positive correlations between Draba spp., Dryas octopetala, Pedicularis lanata, Potentilla diversifolia, Carex nardina (amongst others) with pH; as with all the previous ordinations, elevation simply emphasizes the division between ESSFwcp and IMAwc species.   67  Chapter 4 Discussion  The main goal was to describe the vegetation patterns above 1900m.a.s.l. through discernible units - recurrent across the landscape - which hold sufficient homogeneity and affinity to abiotic factors so that they can be recognized as ordered patterns rather than random arrangements. The protocol by Pojar et al. (1987) for describing plant ecosystems, combined with the approach of Klinka and Chourmouzis (2001) to developing the biogeoclimatic ecosystem classification, proved robust enough that it held its validity when reproduced through independent statistical procedures (highlighted by the similarity in the results of two-way cluster analysis (Appendix A Figures 48 and 49) to those of the tabulation/ordination procedure). The results of my classification process, confirm that certain repeatable and objective arrangements in the vegetation do exist and that both statistical packages and empirical evidence organize vegetation units in very similar ways. My results are consistent with the results by Willner (2011), who found that the assignments of the vegetation into units using TWINSPAN and DCA matched the subjective assignments for 64% to 99% of the relevés used.  Survey of the vegetation above 1900m.a.s.l. in the western Cariboo Mountains The four objectives of the study were all met in that the vegetation at high-elevations in the western mountains of the Cariboo region was thoroughly observed and recorded, vegetation patterns were detected and described, vegetation units were delineated, and their composition and distribution across the landscape were described. This resulted in the rejection of H0, that vegetation is randomly distributed, since the data supported H1, showing the vegetation was arranged in discernable units identified through statistical methods (TWINSPAN, ordinations and cluster analysis) and these units correspond to empirical observations that were readily identifiable in the field. This information contributes to a legacy that began over 60 years ago and has proven essential to the management of ecosystems in British Columbia.  For the third objective, H0 was rejected and H1 was upheld, with the results depending on the ordination method and the number  68  of relevés used in the analysis. The fourth objective of this chapter was met by producing a thorough inventory and classification of the subalpine and alpine vegetation in the research area; this is the first time a classification has been proposed for the high-elevation vegetation of the western Cariboo Mountains.  The plant associations I identified are mostly unique (as indicated by the „nomen novum’ designation in Table 7) but share similarities with some of the communities identified in past studies. The studies used for the comparisons (Tables 8 – 11)did not specifically create a nomenclature for the identified plant associations, so the name in parenthesis that I have provided is my own interpretation based on the dominant species that they identified.  Comparison to previously identified plant communities  Several authors have observed and described the subalpine and alpine vegetation of British Columbia (Shaw 1916, Eady 1971, Service et al. 1974, Buttrick 1977, Douglas and Bliss 1977, Hämet Ahti 1979, Achuff 1984a, Achuff 1984b, Brett et al. 1998, Klinka and Chourmouzis 2001); however, there are no specific studies in the Western Cariboo Mountains. While the differences in composition (species arrangement and richness) and positions in the landscape are notable between my sites and those of coastal (Douglas and Bliss 1977, Brett et al. 1998, Klinka and Chourmouzis 2001), northern (Buttrick 1977) and southern (Eady 1971) locations, they are more subtle when compared with other sites in the Columbia Mountains (Hämet-Ahti 1979, Achuff 1984a, Achuff 1984b, Service et al. 1974). Because of the similarity in location and classification, Tables 8-11 compare some of the plant communities identified in nearby areas of the Columbia Mountains with those identified in my research.    69   Table 8 Comparison of plant communities  from Kootenay National Park identified by Achuff (1984a, left column) to those described in this thesis (right column). Kootenay National Park (Achuff 1984a) Osorio, FG’s (unpublished)  equivalent sites (See Appendix A Figures 51 – 54 for the IMAwc and Figures 55-58 for the ESSFwcp, Appendix A Table 20 and Table 22 provide descriptions for the IMAwc and the ESSFwcp respectively).   20-50% Phyllodoce glanduliflora 15-35% Cassiope mertensiana 2-20% Antennaria lanata on mesic, gentle to moderate slopes, on warm to neutral aspects with well drained soils often subject to solifluction.  Intermediate between the IMAwc Site Series 03- Site Unit 111/Ah03 (Phyllodoce spp. – Cassiope mertensiana – Luetkea pectinata) and the Site Series 01- Site Unit 101/Am03 (Carex spectabilis-Antennaria lanata- Arnica latifolia). Although the species mix is similar, the observed position on the landscape of these communities differs from Achuff‟s (1984) primarily in that mine occupy wetter segments of the landscape. The equivalent unit, considering physical characteristics only (landscape position), would be the IMA wc Site Series 04 – Site Unit 102/Af01 (Silene acaulis- Salix nivalis- Potentilla villosa). Arctic willow-cinquefoil (10-35% Salix arctica – 1-3% Potentilla diversifolia, 20% Antennaria lanata). Occurs in Upper subalpine to alpine, mesic to subhygric, areas of deep snow accumulation  Silene acaulis- Salix nivalis – Potentilla villosa. IMAwc Site Series 04 Site unit 102 Af01. Submesic to subhygric with shallow snow pack and/or exposed sites.   70  Kootenay National Park (Achuff 1984a) Osorio, FG’s (unpublished)  equivalent sites (See Appendix A Figures 51 – 54 for the IMAwc and Figures 55-58 for the ESSFwcp, Appendix A Table 20 and Table 22 provide descriptions for the IMAwc and the ESSFwcp respectively).  White mountain avens – snow willow- moss campion  (10-50% Dryas octopetala – 3-20% Salix nivalis – Silene acaulis). Occurs in Alpine, mesic to Subxeric, south to west aspects No Site Series identified. Site Unit 105 At29 has the same topographic position as the IMAwc Site Series 04, but restricted to base parent materials (eg. Limestone).  Black alpine sedge- woolly pussytoes (10-75% Carex nigricans – 1-35% Antennaria lanata). Occurs in Alpine, mesic to subhygric, wide range of slopes and aspects.  Carex nigricans Site Units 113 As01 and 112 As01.  Carex nigricans – Aulacomnium palustris- Caltha leptosepala ESSFwcp Site Series 07, Site Unit 115 Wa13. Carex spectabilis IMAwc Site Series 05, Site Unit 101 Am03. Restricted to snowbeds and water accumulating areas (depressions).    Table 9 Comparison of plant communities from Mt. Revelstoke and Glacier National parks identified by Achuff (1984b, left column) to those described in this thesis (right column).  Mt. Revelstoke and Glacier National Parks (Achuff 1984b) Osorio, FG’s (unpublished) 5-20% Picea engelmannii 5-10% Abies lasiocarpa 5-15% Valeriana sitchensis 5-10% Erigeron peregrinus. Occurs on mesic to subhygric Upper subalpine sites (1950-2000 m.a.s.l.) on Abies lasiocarpa- Carex spectabilis – Valeriana sitchensis ESSFwcp Site Series 04 Site Unit Sk14Bl. They occupy a similar position in the landscape as identified by Achuff (1984b).  Table 8 Continu d from previous page  71  Mt. Revelstoke and Glacier National Parks (Achuff 1984b) Osorio, FG’s (unpublished) mainly Brunisolic soils often with gleying  1-6 % Picea engelmannii 3-10% Abies lasiocarpa/ 5-30% Phyllodoce glanduliflora 10-55% Cassiope mertensiana. Occurs on mesic Upper Subalpine (2040-2250m) on moderate slopes with southerly to easterly aspects, with well drained Dystric Brunisols.  Abies lasiocarpa – Cassiope mertensiana- Barbilophozia floerkei. ESSFwcp Site Series 03 and 05, Site Unit 103 Sk01. These sites occur on very xeric to subhygric sites, with average to late snowmelt.  5-30% Abies lasiocarpa  1-3% Tsuga mertensiana/ 20-40% Cassiope mertensiana- 5-10% Phyllodoce  Same as above. However there is no Tsuga mertensiana in my research area.  empetriformis  10-20% Luetkea pectinata. Occurs on mesic, Upper Subalpine (1900-2100m) with well drained Dystric Brunisols and Humo-Ferric Podzols.  <1- 5%Picea engelmannii 3-10% Abies lasiocarpa 35-60% Rhododendron albiflorum 20-55% Vaccinium membranaceum. Occurs in the lower Subalpine (1800-2000 m.a.s.l.) on southerly, moderate to steep slopes. Soils are well drained Orthic Ferro-Humic Podzols and Dystric Brunisols.  Abies lasiocarpa- Vaccinium membranaceum. ESSFwcp Site Series 03, Site Unit 103 Sk01. Occur on very xeric to subxeric sites with normal to latest snowmelt timing.  2-10% Abies lasiocarpa 2-5% Pinus albicaulis 1-5% Picea engelmannii / 20-40% Vaccinium membranaceum – 5-35% Cassiope mertensiana. Occurs on mesic, Upper Subalpine (1950-2150 m.a.s.l.) on Same as above.   72  Mt. Revelstoke and Glacier National Parks (Achuff 1984b) Osorio, FG’s (unpublished) moderate westerly and southerly slopes with well drained Dystric Brunisols.  10-50% Abies lasiocarpa  5-10%  Salix spp. 2-15% Valeriana sitchensis. Occurs on mesic lower to upper Subalpine (1580 to 1960 m.a.s.l.) with steep and mostly easterly aspects on Dystric Brunisols and Regosols.  Abies lasiocarpa- Carex spectabilis – Valeriana sitchensis ESSFwcp Site Series 04 Site Unit Sk14Bl. They occupy a similar position in the landscape as identified by Achuff (1984b). 8-35% Salix ssp. 5-20% Tsuga mertensiana  2-10% Abies lasiocarpa/ 10-20% Vaccinium membranaceum. It occurs  Empetrum nigrum – Juniperus communis ESSFwcp Site Series 02, Site Unit 102 Sk62. They occur on very xeric  on mesic sites in the lower and upper Subalpine (1700-2170 m.a.s.l) on steep to very steep slopes with various aspects, on well drained Dystric Brunisols. to xeric sites which are permanently exposed or have early snowmelt timing. Trees (Abies lasiocarpa) are scattered and have low abundance but are typically present in these sites. There is no Tsuga mertensiana in my research area. 10-50% Phyllodoce glanduliflora  15-50% Cassiope mertensiana 2-20% Antennaria lanata. Occurs on mesic upper Subalpine to Alpine sites (2040 – 2500 m.a.s.l.) with gentle to steep slopes on southerly and westerly aspects with well drained Dystric Brunisols and Orthic Humo-Ferric Podzols. Cassiope mertensiana – Phyllodoce empetriformis – Luetkea pectinata. IMAwc Site Series 03, Site Unit 111 Ah03. They occur on gentle to moderate slopes on cool to neutral aspects.  10-50% Dryas octopetala – Salix nivalis – Silene acaulis. It occupies a restricted range on mesic to subxeric No Site Series identified, Site Unit 105 At29. Equivalent in topographic position as the IMAwc Site Series 04, but restricted to  73  Mt. Revelstoke and Glacier National Parks (Achuff 1984b) Osorio, FG’s (unpublished) Alpine sties (2300 – 2650 m.a.s.l.) on southerly aspects with well drained Regosols and Brunisols in which solifluction is common.  base parent materials (eg. Limestone). 10-75% Carex nigricans – 1-5% Antennaria lanata. It occurs on mesic to subhygric upper Subalpine to Alpine sites (2100 – 2400 m.a.s.l.) with a wide range of slopes and aspects on moderately well drained Dystric Brunisols and Regosols.  Carex nigricans Site Unit 112 As01, Carex spectabilis Site Series 05. They occur on subhygric to subhydric sites above 2100 m.a.s.l.  3-30% Erigeron peregrinus – 10-35% Valeriana sitchensis. Occurs on mesic to subhygric upper Subalpine sites (1970-2200 m.a.s.l.) on moderate to steep, southerly and westerly aspects. Soils are well to moderately drained Dystric Brunisols and Orthic Humo-Ferric Podzols.  Carex spectabilis – Valeriana sitchensis – Erigeron peregrinus. ESSFwcp Site Series 01, Site Unit 112 Am02. They occupy submesic to mesic sites in the subalpine, and they seldom extend above 2100 m.a.s.l., except on steep slopes on warm aspects.  40-55% Antennaria lanata 5-15% Cassiope mertensiana – 10-15% Phyllodoce empetriformis. It occurs on mesic Upper Subalpine to Alpine (2220 – 2460 m.a.s.l.) sites with southerly and westerly aspects on well drained Dystric Brunisols and Podzols  Artemisia norvegica – Antennaria lanata. IMAwc Site Series 01, Site Unit 106 At22. It occurs in submesic to mesic sites on moderately steep slopes with neutral to warm aspects, usually between 2100 and 2250 m.a.s.l. The mountains in my research areas don‟t support much vegetation above 2300 m.a.s.l. since such elevations are usually on high ridges, rock outcrops or peaks.   74  Mt. Revelstoke and Glacier National Parks (Achuff 1984b) Osorio, FG’s (unpublished) 8-40% Carex spp. Occurs on moderate to steep snow avalanche slopes in southerly aspects in the lower Subalpine and upper subalpine ( 1390 – 2380 m.a.s.l.) on well drained Regosols and Brunisols.  Carex spectabilis. Site Series 05, Site Unit 101 Am03. They occur on subhygric to subhydric sites, on gentle to very steep slopes on all aspects with average to late snowmelt timing.    Table 10 Comparison of plant communities from Wells Gray Provincial Park (Battle Mountain) identified by Hämet-Ahti (1979, left column) to those described in this thesis (right column).  (Hämet-Ahti 1979) Osorio, FG (unpublished) Heaths (Phyllodoce empetriformis – Cassiope mertensiana – Luetkea pectinata). Occur as narrow belts  Cassiope mertensiana – Phyllodoce empetriformis – Luetkea pectinata. IMAwc Site Series 03, Site Unit 111.   surrounding tree stands, on gently sloping southern aspects. Ah03. They occur on gentle to moderate slopes on cool to neutral aspects. They are not restricted to being adjacent to tree stands, and they often form extensive communities on well drained plateaus with late to latest snowmelt timing Dry Meadows (Antennaria lanata – Sibbaldia procumbens – Erigeron peregrinus). Occur on low ridges and dry gentle slopes. These communities appear to be restricted to the eastern part of the North Cascade range (Douglas and Bliss 1977) and to the Interior Wet belt, where Artemisia norvegica – Antennaria lanata. IMAwc Site Series 01, Site Unit 106 At22. It occurs in submesic to mesic sites on moderately steep slopes with neutral to warm aspects.  75  (Hämet-Ahti 1979) Osorio, FG (unpublished) precipitation is not as high as in coastal areas.  Mesic Meadows (Valeriana sitchensis – Senecio triangularis). Occur on mesic sites on gently sloping cool aspects in the vicinity of tree stands.  These communities appear to be common in many mountains of western North America.  Carex spectabilis – Valeriana sitchensis – Erigeron peregrinus. ESSFwcp Site Series 01, Site Unit 112 Am02. They occupy submesic to mesic sites in the subalpine, and they seldom extend above 2100 m.a.s.l., except on steep  slopes on warm aspects. Mesic mesotrophic meadows (Erigeron peregrinus – Caltha leptosepala- Carex nigricans).  They are irrigated by seepages around springs or brooks and they are not as dense or tall as mesic meadows. Most authors include such communities in the Carex nigricans meadows.  Carex nigricans – Aulacomnium palustris- Caltha leptosepala ESSFwcp Site Series 07, Site Unit 115 Wa13. At the Site Series level, this community is included under the Carex nigricans plant community, but at the Site Unit level it is differentiated by the presence of other herbaceous species (Caltha leptosepala and Ranunculus eschscholtzii). They occupy hygric to subhygric sites with late to latest snowmelt timing.  Moist oligotrophic meadows (Carex nigricans – Aulacomnium palustris) Intermediate between the dry meadows and the wet meadows, they lie on flat or near flat surfaces that are seasonally underwater. They are often associated with snow-bed habitats and therefore have a very short growing season. They occasionally occur along streams and in Same as above   76  (Hämet-Ahti 1979) Osorio, FG (unpublished) other permanently wet places.  Wet meadows (Carex spectabilis – Carex illota – Calamagrostis canadensis). These communities occur on wetter sites than the above, but drier than the fens and marshes.  Carex spectabilis. Site Series 05, Site Unit 101 Am03. Marshes ( Calamagrostis canadensis) Common along brooks and in depressions, covered by water for long periods (usually until the end of July)   Not separated from the above plant community. Although such plant communities were observed, their limited extent did not merit them being segregated as a particular plant community.  Fens (Carex physocarpa – Eriophorum angustifolium). Wetter than shallow marshes and covered by water for most of the summer. These communities are rarely mentioned in the literature and are normally included in the „sedge meadows‟, but they can be expected to be rather common in timberline areas.  Carex nigricans – Eriophorum angustifolium. Site Series ESSFwcp 07, Site Unit 114 Wf12. At the Site Unit level, the communities characterized by Eriophorum angustifolium were segregated, since they occur on the lower limits of the ESSFwcp, but are prevalent in the ESSFwcw and ESSFwc.    Table 11 Comparison of plant communities from Mt. Revelstoke National Park identified by Landas and Scotter (1974, left column) to those described in this thesis (right column). Canadian Wildlife Service, Landas and Scotter (Service et al. 1974) Osorio, FG (unpublished) Abies lasiocarpa - Vaccinium membranaceum- Rhododendron albiflorum  Site Series 03 Abies lasiocarpa – Vaccinium membranaceum. Site unit 103 Sk01 (Bl – Mountain heather – Barbilophozia). The site series and site unit cover the closed canopy forest communities.   77  Canadian Wildlife Service, Landas and Scotter (Service et al. 1974) Osorio, FG (unpublished) Abies lasiocarpa – Rhododendron albiflorum (Open) Site Series 02 Empetrum nigrum – Juniperus communis. Site Unit 102 Ro Sk62 (Bl – Juniper-Crowberry). The equivalent sites, for ridges and rocky sites, presents scattered trees with a scarce vascular plant community in which juniper and crowberry are commonly found. Rhododendron albiflorum is occasionally present but not in as high abundance. Although the species composition differs considerably between Landas and Scotter‟s and my identified plant communities, they occupy the same topographic position, which is why I consider them equivalents. Presumably, the difference in species composition reflects the drier climate of my research area compared to Mt. Revelstoke National Park.   Abies lasiocarpa – Cassiope mertensiana North facing ridgelines Site Series 05 Abies lasiocarpa – Mitella spp. – Cassiope mertensiana. Site Unit 103 Sk01 (Bl – Mountain heather – Barbilophozia). This site series and these site units share similar characteristics as those described by Landas and Scotter.  Abies lasiocarpa- Luzula hitchcockii  Broad meadows and forest fringing Same as above. I included the Luzula hitchcockii understory communities, with  78  Canadian Wildlife Service, Landas and Scotter (Service et al. 1974) Osorio, FG (unpublished) meadows those above.  Eriophorum angustifolium-Sphagnum  Site Series 07 (ESSFwcp) Carex nigricans - Ranunculus eschscholtzii. Site unit 114Wf 12 Narrow-leaved cottongrass  Carex seepage community Same Site Series as above, Site Unit 115 Wa13 Marsh marigold – Black alpine sedge – Glowmoss. Carex nigricans basin community Same site series, Site Unit 113 As01 Alpine sedge  Carex nigricans north slope community Same as above  Luetkea pectinata  Site Series 03 (IMAwc) Phyllodoce spp. – Cassiope mertensiana – Luzula spp. Site Series 06 (ESSFwcp) Cassiope mertensiana – Luetkea pectinata Site unit 111 Ah03 White and Pink mountain-heathers –Partridgefoot.  Cassiope mertensiana Same as above  Valeriana sitchensis dominant meadow -Senecio triangularis – Carex spectabilis sub-type -Lupinus latifolius- Carex spectabilis sub-type  - Veratrum viride sub-type  -Epilobium angustifolium sub-type  Site Series 05 (IMAwc) Carex spectabilis, Site Series 01 (ESSFwcp) Erigeron peregrinus – Lupinus arcticus – Valeriana sitchensis Site Unit 101 Am03 Valerian Pasqueflower, 112 Am02 Showy sedge- Valerian – Fleabane (equivalent to the „Lupinus latifolius’ sub-type).  While the physical characteristics of the sites are similar, my sites do not have Ligusticum canbyi, which is absent from  79  Canadian Wildlife Service, Landas and Scotter (Service et al. 1974) Osorio, FG (unpublished) the entire research area, and Anemone occidentalis seems to be much more prevalent in my sites.   Antennaria lanata  Site Series 01 (IMAwc) Antennaria lanata – Artemisia norvegica. Site Unit 106 At22 Wolly Pussytoes – Mountain sagewort.  The abundance of this community in my research area is a marked contrast to the lack thereof in Landas and Scotter‟s study. Also, Artemisia norvegica is widely distributed and abundant throughout my sites, while it appears to be absent from the sites in Mt. Revelstoke.       As noted in Tables 8-11, several of the plant communities identified in the literature have a similar nomenclature amongst them and with mine, which reflects similar species arrangements and abundance, yet the position in the landscape varies considerably depending on the geographic location. These findings reinforce the need for intensive site-specific studies in which plant communities are identified and recorded based on observed boundaries rather than following predefined quadrats. For example, a recent study of subalpine meadows (Wagner et al. 2013) found very few differences in plant community composition over a 1000Km transect in subalpine sites in the interior ranges of NW North America. Contrary to my results, the researchers found that even across such a large geographic area, the identified plant communities virtually matched those described by Hämet-Ahti (1979). However, it‟s possible that the results of Wagner et al.  80  (2013) highlight the geographic homogeneity of subalpine meadows, which is in great contrast to the heterogeneity of alpine vegetation. Their results might also indicate that a 4m2 quadrat might not be sufficient to capture the true character of the subalpine vegetation and that difference in position on the landscape (topographic location) might be a key differentiating factor when comparing plant communities of similar composition. The BEC methodology uses relevés as the basis for plot sampling, which unlike systematic sampling (as used by Wagner et al. 2013), allows the researcher to capture plant communities which often occur at different scales. Allowing for different scales during sampling, effectively enables the researcher to take into account the variability in composition and distribution unique to the topography and edaphic conditions which host the plant communities. For example, snowbed communities could easily be missed by a transect with fixed quadrat size, since their occurrence in the landscape is confined to localized depressions, which might be missed by a random sample.  The need to carefully examine bedrock type and its influence on plant communities needs to be better understood. On sites underlain with calcareous bedrock (limestone) the relatively high presence and extent of Antennaria racemosa, in the ESSFwcp, and Dryas octopetala, in the IMAwc, is a well-defined pattern that strongly differentiates these sites from those described in my research and in the literature. Similarly, the presence of Aconitum delphiniifolium in the north-most section of my research area (Two Sisters mountain, near the Bowron Lakes) was assumed to be a consequence of the unique bedrock (volcanic and mudstone) of the area; however, upon reading Wagner et. al. (2013), I am uncertain as to whether it is bedrock or latitude which explains the presence of this species which is otherwise absent from my research area.  Contrary to the original assumption that the subalpine vegetation falls within the Picea Engelmannii – Abies Lasiocarpa Order, my results - as well as those of Landas and Scotter (Service et al. 1974) - indicate that it would best be described by a new Order: Abies Lasiocarpa (Abieton lasiocarpae). Likewise, the prevalence of Artemisia norvegica in my study area differs significantly from the more southern sites (Wagner et al. 2013), which could merit rethinking the nomenclature for the Order. However, re-naming a  81  well-established Order (as implied in the BEC system) falls beyond the scope of this thesis.  Relevance of phytosociology in modern studies As a subdiscipline of plant ecology, phytosociology describes the relations between plant species in communities using gradient analysis and classification as its complementary tool (Ewald 2003). Classification is effective in searching for patterns in discontinuous data and ordination is appropriate when continuity in the data exists (Ewald 2003). The tabulation procedure in Central European Phytosociology (CEPS) tradition represents a bottom (locally defined units) to top (synthetic units) approach which is considered „inductive‟. One of the main reasons for tensions between CEPS and the Anglo-American Plant Ecology (AAPE) tradition is that the former tries to describe unique structures (sometimes considered a realm of Natural History) while the latter is dedicated to finding „universal laws‟ (Ewald 2003). Science, as opposed to Natural History, tries to reduce complexity to simple equations and testable predictions. Classifications, such as the ones produced through the BEC approach are largely devoid of equations, and the testability of its predictions is restricted by elements of subjectivity inherent in the identification of plant communities in the field. The methodological sequence I presented should help address the issue of predictability, since it can be applied to any dataset and the resulting groups can be found solely through an automated process in which empirical evidence can be ignored (yet I would personally never rely only on the results from an automated procedure without field verification). Similar methodologies have recently been used to successfully describe and identify plant communities in Iran and the Himalaya (Khan et al. 2013, Noroozi et al. 2014). Considering that in traditional phytosociology, table sorting required „great experience‟ and its mystic nature required „guidance by a master‟ (Mucina 1997), I have attempted to make every step in my procedure explicit so that it can be reproduced by anyone and light can be shed on what could otherwise be seen as a mystical procedure.  Although I tried to create a procedure devoid of subjectivity, the choice of the number of  82  associations or site series is subjective in that I chose the initial number of TWINSPAN divisions and also decided how far I could reduce the groups without loss of information. However, I feel that since we are trying to describe communities which can be visually recognized in the field, the final number of units is decided by the scale of interest. Furthermore, this is an important aspect of the BEC methodology, which describes it as “subjective sampling without preconceived bias” (Pojar et al. 1987), since the output of the tabulation procedure could be full of redundant associations if empirical observations and knowledge is ignored. However, this can be understood if we acknowledge that vegetation patterns are naturally multi-faceted and the units chosen for description will invariably change depending on the scale of observation; also, as noted by Jennings et al. (2009), results vary continuously due to historical and environmental stochasticity. This is one of the fundamental criticisms of phytosociology, since data collection is preceded by subjective pattern recognition and a certain degree of subjectivity is required for cleaning the diagonal tabulations from plots that are considered outliers or redundant (Ewald 2003). Also, there is a degree of tautology in the selection of diagnostic character species since they define the associations which define their status (Ewald 2003). Notwithstanding these limitations, the BEC approach to vegetation classification backed by multivariate procedures (ordinations and cluster analysis) does contribute to the predictive capacity of ecology. Unfortunately, the results of my analysis show that most of the soils, topography, and vegetation variables showed weak correlations. It is important to keep this in mind when developing predictive ecosystem models which could be supplemented by the information I provided, yet one should not rely exclusively on the vegetation-environment correlations that I have described. More intensive studies would be required to better understand the complex interactions that determine the patterns of species composition in high-elevation plant communities.  Some of the shortcomings of my sampling methodology, which should be improved on, include: a low sample number (considering the extent of the area and the variability of the plant communities), a pre-defined elevation cut-off for relevé locations, lack of soil variables used (such as in situ incubations for mineralizable N), lack of repeat measurements for the same areas over a longer time period, and a limited number of  83  measured variables. As such, my research serves as a general indication of what patterns might exist in the vegetation. Further research is necessary to better understand the complexity of the interactions between climate, topography, geology, snow distribution, and soils, and their influence on the physical arrangement of plant communities.  The classification is intended to be used by anyone with basic plant identification skills, a basic understanding of topographic elements, and a general sense of snow distribution (e.g. prevailing wind direction, cold and warm aspects). For example, upon arriving at a new location, after identifying the presence and measuring the abundance of the plants on a site, and the position of that site in the landscape, anyone can determine what site unit or site series any given plant ecosystem belongs to, by comparing the new site to the descriptions in the classification. The classification at a site series level (Appendix A Figures 50, 51, 54, 55 Appendix B Tables 46, 47) places greater emphasis on user-friendliness. That is, in addition to the tabling procedure (where the most frequent and most abundant species define the vegetation unit), when I labelled the site series, I used species that were sometimes very frequent but not always very abundant. The purpose for this was to maximize the chances that if a site is described using the site series nomenclature, the description accurately conveys the site‟s qualities. Also, if there is only limited or no vegetation data available for a site, but there is good topographic and snow distribution data, the classification provides an informed „best-guess‟ as to what plant communities might occupy the site.  The procedure I followed for naming plant communities follows the guidelines for a biogeoclimatic ecosystem classification, but it can be used by anyone trying to hierarchically label plant associations.  Naming plant communities can be useful when describing habitat quality (e.g. bird and animal habitat) since having a standardized nomenclature ensures that equivalency between sites can be established. Understanding how vegetation occupies certain positions in the landscape is useful for reclamation and restoration plans. Also, understanding the extent of the present distribution of the vegetation at high-elevations can provide good baseline information to monitor the effects of climate change on vegetation.   84  The results of my classification include: five new site series for the Interior Mountain Heather wet cold alpine subzone; b) seven new site series for the Engelmann Spruce Subalpine Fir wet cold parkland subzone; c) two plant orders (one not previously described); d) 5 alliances (2 not previously described), and 19 plant associations (17 not previously described). In addition to the abovementioned uses, these results contribute to the description of Site Associations, Site Orders and Site Alliances for the BEC high-elevation classification.       85  Chapter 5 Epilogue  At high-elevations the patterns in the arrangement of the vegetation are easily observed and well defined. At first I thought that describing these patterns and uncovering some of the explanatory variables behind the unique arrangements would be a fairly straightforward endeavour. But during the initial stages of the research it became clear that the same plant communities overlaid very different soils and bedrock, and that they occupied different topographic positions. Although I believe my research is a good starting point in understanding the factors behind the patterns in the vegetation, it falls short of truly understanding how different climatic and physical variables create the observed vegetation patterns, since I did not find strong enough statistical correlations to unequivocally identify how edaphic and topographic variables determine arrangements in the vegetation. Also, since the BEC system emphasizes the effects of climate on plant communities, further mountain climate research (and/or data availability) is required; specifically, better understanding on snow distribution, depth, and date of disappearance is necessary to define concepts like „zonal‟ sites and to explain atypical variations. Furthermore, if various subzones and variants are included for the AT, their boundaries should be reflected in climatic differences and confirmed with climatic evidence.   While understanding the causes behind the vegetation patterns is an important part of a classification, verifying that such patterns are actually real patterns and not just expressions of the researchers interpretations, is just as important. The majority of the effort in this thesis was in using analytical methods to describe the vegetation, rather than simply relying on my empirical observations. I believe that the use of classification statistical packages, combined with tabulation, and verified through ordinations, resulted in a clear corroboration that the empirical observations could be quantitatively and objectively derived. Also, a major task in this thesis was to balance the number of objectively identified plant communities with what made sense to describe at a meaningful scale. For example, the results from TWINSPAN suggest that I should have described many more plant communities than I actually did. Following the tabulation guidelines allowed me to reduce the amount of plant communities to a more representative number. Yet, as it can be observed when comparing the site units to the  86  site series, (and to those in the literature) there is no exact number of plant communities that capture the entire variation in distribution and abundance of the vegetation. Rather, it reflects a choice made by the researcher that involves balancing the existing delineations in the vegetation, with those that are useful for descriptive and mapping purposes. Likewise, the nomenclature used in describing the plant communities can be obtained from the tabulation process (as was the case for the site units), but it might be useful to include some of the researcher‟s bias to highlight species that are readily identifiable and diagnostic of vegetation types, yet quantitatively elusive. This is why the names chosen for the site series do not strictly follow the results from the tabulation but rather they reflect my own interpretation on what plants can be most useful when identifying plant communities.   The classification should provide scientists and land managers a nomenclature that can be used to convey a large amount of information about specific sites and their vegetation, through simple terminology. Also, it provides a starting point from which future comparisons can be made spatially (different geographic locations) and temporally (future studies). Moreover, this thesis is a continuation of the classification tradition that has been used to describe the terrestrial ecosystems of British Columbia for over half a century; it was a great honour to have had an opportunity to contribute to the BEC legacy.     87  References Achuff, P. 1984a. Ecological Land Classification of Kootenay National Park, British Columbia. Vol. I. Integrated Resource Description. Environment Canada. Achuff, P. L. 1984b. Ecological land classification of Mount Revelstoke and Glacier national parks, British Columbia. Alberta Institute of Pedology, University of Alberta. Austin, M. and T. Smith. 1990. A new model for the continuum concept. Pages 35-47  Progress in theoretical vegetation science. Springer. Bach, J. D. 2010. Mountain climate. Draft in mountains and people. 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Tuexenia 31: 271–282 Wittneben, U. and Lacelle, L. 2006. Soils: The Columbia Mountains and the Southern Rockies. Ministry of Sustainable Resource Management, Soils Landscape of BC. Online at < http://srmwww.gov.bc.ca/soils/landscape/3.5columbia.html>  Wojciechowski, M.F., and  Jeimbrook, M.E. 1984.  Dinitrogen fixation in alpine tundra, Niwot Ridge, Arctic and Alpine Research 16:1-10    97  Appendix A   Figure 21 Revised BEC classification for part of the Horsefly Watershed, Horsefly Forest District.   Figure 22 Previous BEC classification for a section of the Horsefly Watershed, Horsefly Forest District.   98   Table 12 Pojar et al. (1987) Definitions of diagnostic values of plant species             Table 13 Mackenzie 2012 Ecosystem attributes used to differentiate between units of the same biogeoclimatic hierarchical level.  Table 12 has been removed due to copyright restrictions. It showed the definitions of diagnostic values of plant species. Original Source:  Pojar, J., K. Klinka, and D. V. Meidinger. 1987. Biogeoclimatic ecosystem classification in British Columbia. Forest Ecology and Management 22:119-154.  Table 13 has been removed due to copyright restrictions. It showed ecosystem attributes used to differentiate between units of the same biogeoclimatic hierarchical level. Original Source:  MacKenzie, W.H. 2012. Biogeoclimatic ecosystem classification of non-forested ecosystems in British Columbia. Prov. B.C., Victoria, B.C. Tech. Rep. 068    99   Figure 23 Non-metric multidimensional Scaling (NMDS) Ordination of 130 relevés with 18 TWINSPAN groups. „orgmatter‟ = Percent surface cover of organic matter.  100   Figure 24 Detrended Correspondence Analysis (DCA) for 130 relevés. Green dots are species. „orgmatter‟ = Percent surface cover of organic matter. „rocks = pct. surface cov. of rocks .  101   Figure 25 Canonical Correspondence Analysis for 130 relevés with the aspect overlaid. Altitude measured in m.a.s.l. and Slope % indicates percent slope (steepness).   102   Figure 26 NMDS for all 300 relevés and 18 TWINSPAN divisions.  Table 14 Shows the interim table processed from steps 6 through 9 in for the ESSFwcp. Note that –for the next analysis- groups 7, 6, 5 were merged into a new group as well as groups 1 and 9, due to the shared differentials Carex nigricans and Carex spectabilis respectively. Group 3 was deleted since it only had 1 plot and its elevation was 1515m.a.s.l. Presence classes (PC) as percent frequency are shown in roman numerals (I = 1-20, II = 21-40, III= 41-60, IV = 61-80, V = 81-100) and species significance (SS)  is shown in percent cover. Group No.   Abs. 1  2  3  4  5  6  7  8  9  No. of relevés  Freq. 37  40  1  9  4  2  11  15  71  Group 1   PC SS   ESSFwcp             Carex spectabilis 107 IV 7 II 4 . . III 1 II 3 . . IV 11 II 6 IV 16 Cassiope mertensiana 89 III 6 V 26 . . III 1 II 2 III 1 III 3 II 2 II 12 Arnica latifolia 80 III 3 III 3 . . I 2 II 10 . . I 3 I 1 III 5 Phyllodoce empetriformis 76 IV 3 IV 6 . . I 1 . . . . I 2 I 1 II 5 Picea engelmannii 23 III 4 I 2 V 1 . . . . . . . . I 3 I 1 Abies lasiocarpa 108 . 29 IV 9 . . II 1 . . III 1 I 2 II 1 II 7 Polytrichum juniperinum 17 I 20 I 2 . . I 1 III 30 . . I 1 . . I 8 Lepraria neglecta 12 I 15 I 4 . . . . . . . . . . . . I 9 Polytrichum commune 8 I 18 . . . . . . . . . . I 1 I 21 I 3 2                     Abies lasiocarpa 108 . 29 IV 9 . . II 1 . . III 1 I 2 II 1 II 7  103  Group No.   Abs. 1  2  3  4  5  6  7  8  9  No. of relevés  Freq. 37  40  1  9  4  2  11  15  71  Valeriana sitchensis 98 II 7 III 5 . . II 1 II 1 . . II 4 I 5 IV 15 Vahlodea atropurpurea 93 II 2 III 4 . . III 2 . . V 2 IV 5 II 5 III 4 Cassiope mertensiana 89 III 6 V 26 . . III 1 II 2 III 1 III 3 II 2 II 12 Arnica latifolia 80 III 3 III 3 . . I 2 II 10 . . I 3 I 1 III 5 Phyllodoce empetriformis 76 IV 3 IV 6 . . I 1 . . . . I 2 I 1 II 5 Anemone occidentalis 69 I 3 III 6 . . . . . . . . III 7 I 3 III 11 Luetkea pectinata 65 I 2 V 11 . . II 1 III 8 V 7 III 5 II 4 I 5 Vaccinium membranaceum 37 II 8 III 5 . . . . . . . . . . I 1 I 9 Barbilophozia floerkei 31 I 8 III 13 . . . . III 2 . . I 11 I 3 I 11 3                     Salix barrattiana 3 . . . . V 19 . . . . . . . . I 15 I 20 4                     Carex spectabilis 107 IV 7 II 4 . . III 1 II 3 . . IV 11 II 6 IV 16 Vahlodea atropurpurea 93 II 2 III 4 . . III 2 . . V 2 IV 5 II 5 III 4 Cassiope mertensiana 89 III 6 V 26 . . III 1 II 2 III 1 III 3 II 2 II 12 Senecio triangularis 85 I 1 I 2 . . IV 6 . . . . IV 10 IV 2 IV 11 Juncus drummondii 24 . . I 2 . . IV 3 III 5 . . II 21 II 2 I 2 Leptarrhena pyrolifolia 23 . . I 1 . . IV 8 III 1 III 15 I 1 III 9 I 1 Juncus mertensianus 17 . . I 1 . . IV 4 II 1 . . I 2 II 2 I 1 Parnassia fimbriata 15 . . . . . . III 3 . . . . I 4 II 2 I 3 Philonotis fontana 14 . . . . . . V 19 II 5 . . I 5 I 2 I 2 Equisetum arvense 14 . . I 1 . . IV 7 . . . . . . I 2 I 4 Petasites frigidus 7 . . I 1 . . III 9 . . . . . . I 5 . . Carex pyrenaica 8 I 1 I 7 . . II 32 . . . . . . . . I 1 Sanionia uncinata 6 . . . . . . II 27 . . III 1 I 2 I 1 . . 5                     Luetkea pectinata 65 I 2 V 11 . . II 1 III 8 V 7 III 5 II 4 I 5 Carex nigricans 52 I 1 I 4 . . II 8 V 36 V 61 V 48 IV 22 I 2 Hieracium gracile 43 II 2 II 2 . . I 1 III 8 . . . . . . II 5 Barbilophozia floerkei 31 I 8 III 13 . . . . III 2 . . I 11 I 3 I 11 Juncus drummondii 24 . . I 2 . . IV 3 III 5 . . II 21 II 2 I 2  104  Group No.   Abs. 1  2  3  4  5  6  7  8  9  No. of relevés  Freq. 37  40  1  9  4  2  11  15  71  Leptarrhena pyrolifolia 23 . . I 1 . . IV 8 III 1 III 15 I 1 III 9 I 1 Polytrichastrum alpinum 16 I 1 I 2 . . . . III 4 . . I 1 . . I 2 Dicranum scoparium 14 I 2 I 2 . . . . III 3 . . . . I 2 I 1 Warnstorfia exannulata 4 . . . . . . . . III 1 . . . . I 2 . . Poa alpina 20 I 1 I 2 . . I 1 III 16 . . I 1 . . I 2                      Polytrichum juniperinum 17 I 20 I 2 . . I 1 III 30 . . I 1 . . I 8 Eriophorum angustifolium 11 . . . . . . II 2 II 18 III 1 . . II 20 I 10 6                     Abies lasiocarpa 108 . 29 IV 9 . . II 1 . . III 1 I 2 II 1 II 7 Vahlodea atropurpurea 93 II 2 III 4 . . III 2 . . V 2 IV 5 II 5 III 4 Cassiope mertensiana 89 III 6 V 26 . . III 1 II 2 III 1 III 3 II 2 II 12 Veratrum viride 77 I 1 II 2 . . II 13 . . III 1 I 3 II 2 IV 11 Luetkea pectinata 65 I 2 V 11 . . II 1 III 8 V 7 III 5 II 4 I 5 Carex nigricans 52 I 1 I 4 . . II 8 V 36 V 61 V 48 IV 22 I 2 Leptarrhena pyrolifolia 23 . . I 1 . . IV 8 III 1 III 15 I 1 III 9 I 1 Aulacomnium palustre 20 . . . . . . II 6 . . III 5 I 1 III 19 I 18 Phyllodoce glanduliflora 19 II 6 I 3 . . . . . . III 1 I 1 . . I 6 Eriophorum angustifolium 11 . . . . . . II 2 II 18 III 1 . . II 20 I 10 Rhytidiadelphus squarrosus 9 . . . . . . II 8 . . III 1 I 7 I 7 I 5 Racomitrium canescens 8 . . I 1 . . . . . . III 20 . . I 2 I 6 Agrostis humilis 8 . . . . . . II 1 . . III 4 . . I 1 I 4 Polytrichum strictum 6 . . . . . . . . . . V 6 . . I 1 I 5 Sanionia uncinata 6 . . . . . . II 27 . . III 1 I 2 I 1 . . Calliergon stramineum 4 I 5 . . . . . . . . III 2 . . I 11 . . 7                     Carex spectabilis 107 IV 7 II 4 . . III 1 II 3 . . IV 11 II 6 IV 16 Vahlodea atropurpurea 93 II 2 III 4 . . III 2 . . V 2 IV 5 II 5 III 4 Cassiope mertensiana 89 III 6 V 26 . . III 1 II 2 III 1 III 3 II 2 II 12 Senecio triangularis 85 I 1 I 2 . . IV 6 . . . . IV 10 IV 2 IV 11 Anemone occidentalis 69 I 3 III 6 . . . . . . . . III 7 I 3 III 11 Luetkea pectinata 65 I 2 V 1 . . II 1 II 8 V 7 III 5 II 4 I 5  105  Group No.   Abs. 1  2  3  4  5  6  7  8  9  No. of relevés  Freq. 37  40  1  9  4  2  11  15  71  1 I Carex nigricans 52 I 1 I 4 . . II 8 V 36 V 61 V 48 IV 22 I 2 Juncus drummondii 24 . . I 2 . . IV 3 III 5 . . II 21 II 2 I 2 8                     Senecio triangularis 85 I 1 I 2 . . IV 6 . . . . IV 10 IV 2 IV 11 Carex nigricans 52 I 1 I 4 . . II 8 V 36 V 61 V 48 IV 22 I 2 Caltha leptosepala 23 . . I 1 . . II 1 . . . . II 8 V 8 I 5 Leptarrhena pyrolifolia 23 . . I 1 . . IV 8 III 1 III 15 I 1 III 9 I 1 Aulacomnium palustre 20 . . . . . . II 6 . . III 5 I 1 III 19 I 18 Eriophorum angustifolium 11 . . . . . . II 2 II 18 III 1 . . II 20 I 10 Polytrichum commune 8 I 18 . . . . . . . . . . I 1 I 21 I 3 Sphagnum russowii 4 . . . . . . . . . . . . . . II 22 . . Pohlia wahlenbergii 3 . . . . V 2 . . . . . . . . I 33 . . Salix barrattiana 3 . . . . V 19 . . . . . . . . I 15 I 20 Carex nardina 3 I 1 . . . . . . . . . . . . I 30 . . 9                     Carex spectabilis 107 IV 7 II 4 . . III 1 II 3 . . IV 11 II 6 IV 16 Valeriana sitchensis 98 II 7 III 5 . . II 1 II 1 . . II 4 I 5 IV 15 Vahlodea atropurpurea 93 II 2 III 4 . . III 2 . . V 2 IV 5 II 5 III 4 Senecio triangularis 85 I 1 I 2 . . IV 6 . . . . IV 10 IV 2 IV 11 Arnica latifolia 80 III 3 III 3 . . I 2 II 10 . . I 3 I 1 III 5 Veratrum viride 77 I 1 II 2 . . II 13 . . III 1 I 3 II 2 IV 11 Anemone occidentalis 69 I 3 III 6 . . . . . . . . III 7 I 3 III 11 Erigeron peregrinus 69 I 1 I 3 . . I 1 . . . . I 2 II 5 IV 7 Phleum alpinum 38 I 1 I 1 . . II 1 . . . . II 2 I 1 III 3 Aulacomnium palustre 20 . . . . . . II 6 . . III 5 I 1 III 19 I 18 Pseudoleskea radicosa 4 . . I 1 . . . . . . . . . . . . I 22 Carex scirpoidea 4 . . . . . . . . . . . . . . . . I 20 Salix barrattiana 3 . . . . V 19 . . . . . . . . I 15 I 20 Rhizomnium glabrescens 3 . . . . . . . . . . . . . . . . I 24   106  Table 15 Shows the tabulation of TWINSPAN groups for the IMAwc after step 6. Presence classes as percent frequency are shown in roman numerals (I = 1-20, II = 21-40, III= 41-60, IV = 61-80, V = 81-100). Group No. Abs. 1 2 3 4 5 6 7 8 9 No. of relevés Freq. 4 12 19 7 14 2 14 15 23 Cassiope mertensiana 55 IV V IV V III . II III I Kiaeria blyttii 2 III . . . . . . . . Luetkea pectinata 32 IV III IV IV II . . I . Carex spectabilis 45 III II III III V III I IV I Hieracium gracile 39 III III IV . IV . I III I Antennaria alpina 31 III . I II I . III III II Carex nigricans 26 III V II II II . . I . Stereocaulon alpinum 15 III . . . . . I I II Dicranum undulatum 2 III . . . . . . . . 2            Cassiope mertensiana 55 IV V IV V III . II III I Luetkea pectinata 32 IV III IV IV II . . I . Antennaria lanata 51 . III IV II V . II III II Hieracium gracile 39 III III IV . IV . I III I Phyllodoce empetriformis 33 . III II V III . I II I Carex nigricans 26 III V II II II . . I . Juncus drummondii 11 . III II . I . . . . 3            Cassiope mertensiana 55 IV V IV V III . II III I Luetkea pectinata 32 IV III IV IV II . . I . Sibbaldia procumbens 52 . . III . III . II V IV Antennaria lanata 51 . III IV II V . II III II Carex spectabilis 45 III II III III V III I IV I Hieracium gracile 39 III III IV . IV . I III I Phyllodoce glanduliflora 36 . I IV . I . II I III Polytrichum piliferum 32 . I III . II . III II II Erigeron peregrinus 28 II . III II IV III I II I Luzula piperi 26 . II IV I II . I II . Polytrichastrum alpinum 18 II I III . II . I I . Carex pyrenaica 17 . . III . . . I II I Cladonia species 16 . I III . I . I I I 4            Cassiope mertensiana 55 IV V IV V III . II III I Luetkea pectinata 32 IV III IV IV II . . I . Carex spectabilis 45 III II III III V III I IV I Phyllodoce empetriformis 33 . III II V III . I II I Abies lasiocarpa 21 . . . III II V II I I Anemone occidentalis 13 . II . III III . . . .  107  Group No. Abs. 1 2 3 4 5 6 7 8 9 No. of relevés Freq. 4 12 19 7 14 2 14 15 23 Senecio triangularis 10 . I . III III . . . . Caltha leptosepala 7 . I . III II . . . . 5            Cassiope mertensiana 55 IV V IV V III . II III I Artemisia norvegica 62 . . II I IV III IV V V Sibbaldia procumbens 52 . . III . III . II V IV  Antennaria lanata 51 . III IV II V . II III II Carex spectabilis 45 III II III III V III I IV I Hieracium gracile 39 III III IV . IV . I III I Phyllodoce empetriformis 33 . III II V III . I II I Erigeron peregrinus 28 II . III II IV III I II I Vahlodea atropurpurea 24 . II II . IV . I I I Phleum alpinum 21 II . II II IV . I I I Arnica latifolia 19 II I II . IV . . I I Claytonia lanceolata 9 . . I . III . . . I Anemone occidentalis 13 . II . III III . . . . Valeriana sitchensis 11 . I I II III . . . I Senecio triangularis 10 . I . III III . . . . 6            Artemisia norvegica 62 . . II I IV III IV V V Carex spectabilis 45 III II III III V III I IV I Silene acaulis 37 . . . . I III IV III IV Erigeron peregrinus 28 II . III II IV III I II I Abies lasiocarpa 21 . . . III II V II I I Saxifraga bronchialis 10 . I . . . III II . I Potentilla villosa 10 . . . . . III II . II Brachythecium species 7 . I I . I III I I . Poa arctica 5 . . . . II III . I . Picea species 4 . . . . I III I . I Oxyria digyna 3 . . I . . III . I . Lupinus arcticus 3 . . . . I III . . . Salix species 2 . . . I . III . . . Salix arctica 2 . . . . . III I . . Epilobium latifolium 2 . . . . . V . . . Pinus albicaulis 2 . . . . . III I . . Rhinanthus minor 1 . . . . . III . . . Arnica cordifolia 1 . . . . . III . . . Antennaria racemosa 1 . . . . . III . . . Anemone parviflora 1 . . . . . III . . .  108  Group No. Abs. 1 2 3 4 5 6 7 8 9 No. of relevés Freq. 4 12 19 7 14 2 14 15 23 Silene parryi 1 . . . . . III . . . Achillea millefolium 1 . . . . . III . . . 7            Artemisia norvegica 62 . . II I IV III IV V V Silene acaulis 37 . . . . I III IV III IV Polytrichum piliferum 32 . I III . II . III II II Antennaria alpina 31 III . I II I . III III II Festuca brachyphylla 30 . I I I I . III III III Luzula spicata 29 . I . . II . III III III Salix nivalis 29 . . I I I . IV III II Cassiope tetragona 24 . . . . I . III II III Flavocetraria nivalis 16 . . . . . . IV . II Dryas octopetala 13 . . . I . . V . . Thamnolia vermicularis 6 . . . . . . III . . 8            Cassiope mertensiana 55 IV V IV V III . II III I Artemisia norvegica 62 . . II I IV III IV V V Sibbaldia procumbens 52 . . III . III . II V IV Antennaria lanata 51 . III IV II V . II III II Carex spectabilis 45 III II III III V III I IV I Hieracium gracile 39 III III IV . IV . I III I Silene acaulis 37 . . . . I III IV III IV Lepraria neglecta 32 . . I . II . . III V Antennaria alpina 31 III . I II I . III III II Festuca brachyphylla 30 . I I I I . III III III Luzula spicata 29 . I . . II . III III III Salix nivalis 29 . . I I I . IV III II Solorina crocea 25 . I II I . . II III II Trisetum spicatum 20 . I I . II . II III I 9            Artemisia norvegica 62 . . II I IV III IV V V Sibbaldia procumbens 52 . . III . III . II V IV Silene acaulis 37 . . . . I III IV III IV Phyllodoce glanduliflora 36 . I IV . I . II I III Lepraria neglecta 32 . . I . II . . III V Festuca brachyphylla 30 . I I I I . III III III Luzula spicata 29 . I . . II . III III III Polytrichum juniperinum 28 . II II . II . I I III Cassiope tetragona 24 . . . . I . III II III  109  Group No. Abs. 1 2 3 4 5 6 7 8 9 No. of relevés Freq. 4 12 19 7 14 2 14 15 23 Empetrum nigrum 21 . . . . I . II II III Cornicularia aculeata 17 . . . . I . II I III  Table 16 Tabulation resulting from step 8 for the IMAwc. Note that the associations from the previous tables were regrouped when differential species were shared, yet the new analysis still produced redundant differentials (groups 4 and 5 below). Presence classes (PC) as percent frequency are shown in roman numerals (I = 1-20, II = 21-40, III= 41-60, IV = 61-80, V = 81-100) and species significance (SS) is shown in percent cover.           IMAwc  Group No.  1  2  3  4  5  No. of  16  21  21  21  31  1  Abs.Freq. PC SS         Cassiope mertensiana 55 V 10 V 15 III 5 II 1 I 2 Phyllodoce empetriformis 33 III 3 II 4 III 7 I 5 I 2 Luetkea pectinata 32 III 5 IV 3 II 3 I 2 I 4 Carex nigricans 26 V 35 II 1 II 2 . . . . Stereocaulon alpinum 15 I 15 . . . . I 4 II 5              2             Cassiope mertensiana 55 V 10 V 15 III 5 II 1 I 2 Sibbaldia procumbens 52 . . III 1 II 1 III 2 IV 3 Antennaria lanata 51 II 1 IV 7 IV 15 II 1 II 5 Carex spectabilis 45 II 2 III 4 V 5 I 1 II 5 Hieracium gracile 39 II 1 IV 1 III 1 I 1 I 2 Phyllodoce glanduliflora 36 I 1 IV 4 I 2 II 1 III 5 Luetkea pectinata 32 III 5 IV 3 II 3 I 2 I 4 Erigeron peregrinus 28 I 6 III 1 III 7 I 3 I 3 Luzula piperi 26 II 5 IV 1 I 2 I 1 I 2 Carex pyrenaica 17 . . III 3 . . II 1 I 2              3             Artemisia norvegica 62 . . II 2 III 7 IV 2 V 7 Cassiope mertensiana 55 V 10 V 15 III 5 II 1 I 2 Antennaria lanata 51 II 1 IV 7 IV 15 II 1 II 5 Carex spectabilis 45 II 2 III 4 V 5 I 1 II 5 Hieracium gracile 39 II 1 IV 1 III 1 I 1 I 2 Phyllodoce empetriformis 33 III 3 II 4 III 7 I 5 I 2 Erigeron peregrinus 28 I 6 III 1 III 7 I 3 I 3 Arnica latifolia 19 I 4 II 1 III 3 . . I 1 Antennaria alpina 31 I 13 I 1 I 20 III 1 III 1  110            IMAwc  Group No.  1  2  3  4  5  No. of  16  21  21  21  31  Arnica angustifolia 3 . . . . I 20 I 1 . . Aulacomnium palustre 3 . . . . I 20 I 1 . .              4             Artemisia norvegica 62 . . II 2 III 7 IV 2 V 7 Sibbaldia procumbens 52 . . III 1 II 1 III 2 IV 3 Silene acaulis 37 . . I 1 . . IV 1 IV 1 Polytrichum piliferum 32 I 2 II 10 I 5 III 3 II 7 Antennaria alpina 31 I 13 I 1 I 20 III 1 III 1 Festuca brachyphylla 30 I 1 I 1 I 2 III 1 III 3 Salix nivalis 29 . . I 1 I 1 IV 5 II 5 Luzula spicata 29 I 5 . . I 1 III 1 III 2 Dryas octopetala 13 . . I 1 . . III 6 . .              5             Artemisia norvegica 62 . . II 2 III 7 IV 2 V 7 Sibbaldia procumbens 52 . . III 1 II 1 III 2 IV 3 Silene acaulis 37 . . I 1 . . IV 1 IV 1 Phyllodoce glanduliflora 36 I 1 IV 4 I 2 II 1 III 5 Lepraria neglecta 32 . . I 5 II 10 . . V 30 Antennaria alpina 31 I 13 I 1 I 20 III 1 III 1 Festuca brachyphylla 30 I 1 I 1 I 2 III 1 III 3 Luzula spicata 29 I 5 . . I 1 III 1 III 2 Polytrichum juniperinum 28 I 2 II 1 I 4 I 1 III 15 Cassiope tetragona 24 . . . . I 1 II 1 III 3 Empetrum nigrum 21 . . . . I 1 I 1 III 2 Rhizocarpon geographicum 4 . . I 1 . . I 1 I 20   111   Figure 27 Principal Component Analysis (PCA) for 130 relevés with TWINSPAN groups reduced from 18 to 7.  Figure 28 Nonmetric multidimensional scaling for 130 relevés with TWINSPAN groups reduced from 18 to 7.  112   Figure 29 Detrended Correspondence Analysis for 130 relevés with TWINSPAN groups reduced from 18 to 7.  113   Figure 30 Detrended Correspondence Analysis (Axis 3 and 1) for 130 relevés with TWINSPAN groups reduced from 18 to 7, marked by convex hulls.  114   Figure 31 Canonical Correspondence Analysis for 130 relevés with TWINSPAN groups reduced from 18 to 7.  Figure 323D representation for the nonmetric multidimensional scaling for 130 relevés with TWINSPAN groups reduced from 18 to 7.  115   Figure 33 Detrended Correspondence Analysis for the IMAwc (110 relevés) with 5 TWINSPAN divisions.  Figure 34 PCA (using the variance/covariance matrix) for the IMAwc (110 relevés) with 5 TWINSPAN divisions orgmatter‟ = Percent surface cover of organic matter.    116   Figure 35 NMDS for the  IMAwc (110 relevés) with 5 TWINSPAN divisions, „orgmatter‟ = Percent surface cover of organic matter.  117   Figure 36 Canonical correspondence analysis for the  IMAwc (110 relevés) with 5 TWINSPAN divisions.    118   Figure 373D representation of Canonical Correspondence Analysis for the IMAwc (110 relevés) with 5 TWINSPAN divisions.  119   Figure 38 Principal Component Analysis for the ESSFwcp (190 relevés) with 5 TWINSPAN divisions.  Figure 393D representation of the Detrended Correspondence Analysis for the ESSFwcp (190 relevés) with 5 TWINSPAN divisions.  120   Figure 40 NMDS for the ESSFwcp (190 relevés) with 5 TWINSPAN divisions.  Figure 41 Canonical Correspondence Analysis for the ESSFwcp (190 relevés) with 5 TWINSPAN divisions „seepage‟= seepage depth, pcntwate= percent surface cover of water.   121  Table 17 Tabulation of the alliances represent the final (highest hierarchical level) grouping for the relevés of the IMAwc. Alliance No.  1  2  3  No. Of relevés  37  21  52    Diagnostic Values    Cassiope mertensiana d V 12 III 5 II 1 Luetkea pectinata d IV 5 II 3 I 2 Carex nigricans d III 14 II 2 . . Hieracium gracile d III 1 III 1 I 1 Luzula piperi d III 1 I 2 I 1 Phyllodoce glanduliflora d III 2 I 2 II 2 Stereocaulon alpinum cd I 15 . . II 5          Carex spectabilis d III 4 V 5 II 3 Antennaria lanata dd III 4 IV 15 II 2 Antennaria alpina cd I 5 I 20 III 1 Arnica latifolia d I 2 III 3 I 1 Arnica angustifolia cd . . I 20 I 1 Aulacomnium palustre cd . . I 20 I 1 Erigeron peregrinus d II 1 III 7 I 3 Phyllodoce empetriformis d II 3 III 7 I 3          Artemisia norvegica d I 2 III 7 V 5 Sibbaldia procumbens d II 1 II 1 IV 2 Silene acaulis d I 1 . . IV 1 Cassiope tetragona d . . I 1 III 3 Festuca brachyphylla d I 1 I 2 III 2 Lepraria neglecta d, cd I 5 II 10 III 30 Luzula spicata d I 5 I 1 III 1 Salix nivalis d I 1 I 1 III 5             122  Table 18 Tabulation of the alliances represent the final (highest hierarchical level) grouping for the relevés of the ESSFwcp. Alliance No.  1  2  3  4  No. of relevés  40  9  32  108    Diagnostic Values          Barbilophozia floerkei d III 8 . . I 3 I 3 Cassiope mertensiana dd,cd V 20 III 1 II 2 II 4 Luetkea pectinata d V 8 II 1 III 3 I 1 Phyllodoce empetriformis d IV 5 I 1 I 1 II 3 Vaccinium membranaceum d III 1 . . I 1 I 4   Carex pyrenaica cd I 1 II 31 . . I 1 Equisetum arvense d I 1 IV 4 I 1 I 1 Juncus drummondii d I 1 IV 1 II 2 I 1 Juncus mertensianus d I 1 IV 1 II 1 I 1 Leptarrhena pyrolifolia d I 1 IV 3 III 2 I 1 Parnassia fimbriata d . . III 1 I 1 I 1 Petasites frigidus d I 1 III 6 I 5 . . Philonotis fontana dd, cd . . V 15 I 4 I 2 Sanionia uncinata dd . . II 30 I 1 . . Senecio triangularis d I 1 IV 2 III 2 III 5          Caltha leptosepala d I 1 II 1 III 3 I 4 Carex nigricans dd I 1 II 3 V 36 I 1 Carex nardina cd . . . . I 30 I 1 Pohlia wahlenbergii c . . . . I 33 . . Sphagnum russowii c . . . . I 22 . . Vahlodea atropurpurea d III 2 III 1 III 3 III 2          Abies lasiocarpa dd IV 3 II 1 II 1 IV 21 Anemone occidentalis d III 4 . . II 3 III 6 Arnica latifolia d III 2 I 2 I 2 III 3 Carex spectabilis d II 2 III 1 III 5 IV 7 Dicranum muehlenbeckii c I 9 . . . . I 40 Erigeron peregrinus d I 1 I 1 II 1 III 3 Pleurozium schreberi c . . . . I 3 I 35 Salix barrattiana c . . . . I 15 I 20 Valeriana sitchensis d III 3 II 1 II 2 IV 10 Veratrum viride d II 1 II 13 II 1 III 3  123   Figure 42 NMDS for the 3 alliances of the IMAwc. „rock‟= percent surface cover of rocks.   Figure 43 NMDS for the three alliances of the IMAwc with only dominant species used in the analysis.  124   Figure 44 DCA for the three alliances of the IMAwc.  125    Figure 45 NMDS for the 4 alliances of the ESSFwcp.  126   Figure 46 CCA for the four alliances of the ESSFwcp.  127   Figure 47 Hierarchical Cluster Analysis for 130 relevés based on Sørensen‟s dissimilarity index as distance measure and group average as linkage method. myplotsnewgroupsshortsppcllusterDistance (Objective Function)Information Remaining (%)3.2E-021006.8E+00751.4E+01502E+01252.7E+010291912311393844124120116887847833 2 1258561775 28914 1312230104511811562575812865645063601175121814979801611954461411155485282531037697102723673563537426 113130129101716968104107961101 33175989112108131247475106100661269586841590123123261147 9 939298272041183299224012710510967708743251218 3494newgroup1234567 128   Figure 48 Two-way Cluster Analysis  for the IMAwc based on Sørensen‟s dissimilarity index as distance measure and group average as linkage method. IMA Allience Two-Way Cluster AnalysisInformation Remaining (%)0 25 50 75 1001007550250108946766816186172645158812110651499310641091051041019010310100605910210730572276243631213529342827262033322370632562956968319809737498371737285485840424152475145544474773843465655969989131192539198508281618439186877879775ArtemisiFestucaLuzula sLeprariaPolytricPolytricSolorinaSalix niPolygonuPotentilStellariCampanulSibbaldiTrisetumSilene aMinuartiStereocaCladinaAntennarPhyllodoCassiopeEmpetrumPotentilRhytidioRhizocarGentianeCarex naCarex alPolytricVeronicaSedum laAbies laCladinaCladoniaSolidagoAgrostisGentianaCladoniaSelagineCetrariaPeltigerPicea enDryas ocSaxifragThamnoliAlectoriUmbilicaCetrariaFestucaCarex phCassiopePhyllodoDiphasiaSenecioAntennarPhleum aHieraciuErigeronArnica lVahlodeaCarex spPoa cusiPediculaAnemoneMyosotisArnica aValerianVeratrumPolytricCarex pyPeltigerDiphasiaLuetkeaDicranumLuzula pRacomitrPoa alpiJuncus dAnemoneCaltha lJuncus mLuzula pPolytricClaytoniLupinusPoa arctPolytricCarex niBarbilopTrichophLuzula aEpilobiuPohlia nPleuroziAulacomnSaxifragRacomitrPolytricPolytricArtemisiOxyria dAllience1 2 3Matrix CodingPresence Absence 129   Figure 49 Two-way Cluster Analysis for the ESSFwcp based on Sørensen‟s dissimilarity index as distance measure and group average as linkage method.   ESSF Allience two-way cluster analysisInformation Remaining (%)0 25 50 75 100100755025018615411421182372443696854184149501835215751185181451601594749984144158481781611881125338121177791131271347313139317055206783851521651626156151916366282377172518602162225965141657642164866381168167365888Abies laPicea enLuzula pMitellaEpilobiuSelagineThalictrOrthiliaCladoniaPhyllodoCarex spVahlodeaValerianArnica lAntennarCassiopePhyllodoLuetkeaAnemoneSibbaldiTrisetumLuzula pBarbilopMitellaPseudoleHieraciuPolytricPoa alpiAnemonePolytricDicranumAntennarRacomitrPoa cusiDiphasiaCladoniaCladinaCladinaSalix niCetrariaArtemisiViola glCampyliuPediculaErythronViola orDiphasiaVacciniuPolytricJuniperuGaultherArtemisiRhododenPinus alRibes laDicranumBarbilopDicranumCetrariaCladoniaDicranelCladoniaCladoniaSenecioJuncus mPhleum aSalix arPeltigerParnassiPhilonotEquisetuSaxifragMimulusBrachythJuncus dRhytidiaTiarellaSanioniaVeratrumCastilleBryum psPetasiteErigeronTrolliusCarex naLeptarrhVeronicaCaltha lCastillePoa arctCalamagrWarnstorEpilobiuEpilobiuLeprariaPotentilPicea glEriophorCalliergSolorinaCarex pyFestucaLuzula sCarex niRanunculPleuroziLuzula aAulacomnAnemoneDryopterSalix baParnassiAntennarPeltigerRhytidioRubus peEmpetrumStreptopArnica cKalmia mRacomitrAgrostisPolytricPolytricSphagnumMitellaArnica mVacciniuSalix baPohlia wAllience1 2 3 4Matrix CodingPresence Absence 130  Table 19 Key to Site Units14 of the IMAwc. 1a. Very shallow soils with very high (60%+) coarse fragment content; permanently exposed to earliest snowmelt; ridge-crests, summits and adjacent site types.  2a. Very xeric to xeric substrate; lichens are the dominant life-form; high exposure to wind; vascular plant cover restricted to few species mostly woody and cushion-like (Moss-campion, alpine azalea) creating micro-habitats for other vascular plants.     103 At71 Angled mountain-heather- Crowberry 2b. Xeric to subxeric sites; sparse to abundant vascular plant cover with higher species diversity than 2a; flat to steep slopes; site unit extends from crests to upper slopes.   102 Af01 Moss-campion – Snow willow - Potentilla  2c. Submesic to mesic, rapidly drained soils; sheltered crests and warm aspects.    105 At29 Dunhead sedge – Mountain sagewort 1b. Slightly developed to moderately well-developed and well drained soils; average snowmelt, moderately steep to gentle slopes  3a. Solifluction lobes common; water receiving and shedding sites; middle and lower slopes; extensive Mountain sagewort and Wolly pussytoes with abundant vascular plant cover (plant cover > bare ground) restricted in height.   106 At22 Woolly pussytoes – Mountain sagewort 3b. Water receiving, concave sites; moderately well-developed, subhygric to hygric soils; warm to neutral aspects.   101 Am  Woolly pussytoes – Showy sedge – Mountain arnica 1c. Late to latest snowmelt; imperfectly drained soils; gentle to very steep slopes; upper                                                  14 Nomenclature of Site Units developed by Will Mackenzie.  131  to lower slope position. 4a. Subxeric to hygric soils; abundant moisture supply; cool to neutral aspects; extensive heathers frequently > 50% cover; low species diversity.   111 Ah03 White & Pink mountain-heathers- Patridgefoot  4b. Snowbeds (depressions); commonly saturated, sandy soils; flat to gentle slopes; low species diversity frequently dominated by Alpine sedge, Tufted bulrush common, occasional Sphagnum and Bluejoint grass.  112 As01 Alpine Sedge  1d. Poorly developed or no soil development with good drainage and highly unstable; moderately steep to very steep sites; cool aspects; barren appearance (fellfields) with very sparse vegetation; latest snowmelt; sparse but common Pyrenees‟s Sedge, Piper‟s woodrush and Mountain sorrel.   113 As10 Pyrenees Sedge – Piper’s woodrush.      Table 20 Site Units of the IMAwc (by Will Mackenzie, revised by Federico Osorio). 101 Am Woolly pussytoes – Showy sedge - Mountain arnica ecosystems are common in the IMAwc where it typically dominates gentle warm and neutral aspects. The site distribution extends to drier sites with relative unstable parent materials and warm aspect sites below snowbeds with improved moisture regime. The snow pack of these sites is continuous through the winter and soils do not freeze; but, the snow pack can be thinned and compacted during periods of high winds or mid-winter thaw. Release from snow occurs in the last week of June to the 2nd week of July. The 101 Am is characterized by a higher cover of woolly pussytoes, mountain sagewort, and showy sedge but better differentiated from other units by a high cover and diverse composition of “meadow” forbs such as subalpine fleabane, mountain arnica, Sitka valerian, western springbeauty, and western pasqueflower. Other species Table 19 Continued from previous page   132  may also occur such as mountain hairgrass, mountain-heathers, alpine timothy, slender hawkweed, Cusick‟s bluegrass, and bracted lousewort.  102 Af  Moss campion - Snow willow ecosystems occur on higher elevation ridges and slopes and on lower elevation neutral aspect flat ridges and gradual upper slopes. They have a thin discontinuous snow pack which melts by the last week of May in years with normal snowpack. These ecosystems commonly have sparse vegetation cover because of high rock cover or extensive cryoturbation. Soils are generally thin with a high percent of bedrock and rapid drainage. The effects of winter wind exposure, cryoturbation, and rapid drainage result in plant communities with relatively low percent vegetation covers which are dominated by rock cover and lichens such as Lepraria neglecta. At a distance, these ecosystems may appear to be barren rocky habitats, however there is often high species diversity resulting from an abundance of specialist species adapted to such extreme conditions. Vegetation cover is typically low but species diversity is high in these ecosystems. Complex microsite variation results in a variety of fellfield and tundra species on many sites. The cushion plant moss-campion occurs in scattered patches on most sites and dwarf snow willow may occur with low to high cover. In microsites with better winter snow protection mountain sagewort, sibbaldia, angled mountain-heather, mountain harebell and others may occur. On more wind-exposed microsites species such as alpine fescue, spiked wood-rush, or alpine pussytoes may occur.  103 At Mountain-avens - Snow willow ecosystems are very uncommon in the IMAwc occurring on wind-swept and rapidly drained ridge crests primarily in areas of base-rich parent materials and lighter snow fall. White mountain-avens occurs with moderate to high cover and is diagnostic for this ecosystem. Other fellfield species are common with dwarf snow willow, alpine fescue, and ragged paperdoll lichen occurring on most sites.  103 At71 Four-angled mountain-heather - Crowberry ecosystems occur on rocky windswept upper slopes and summits which have a thin, discontinuous winter snowpack and experience early snowmelt. Soils are poorly developed and deflated, with a very high percent of surface cobbles. Because of high surface roughness, snow may be captured in small pockets which increases habitat variability and hence higher species diversity. Vegetation cover is variable. Four-angled mountain-heather occurs consistently across all sites but crowberry, or alpine-azalea may be dominant in some localities. Yellow mountain-heather, mountain sagewort, moss campion, sibbaldia, and alpine fescue are common associates. The vascular plants in this site unit are often the first ones to bloom in the spring, with alpine-azalea and moss campion often blooming up to 1 month before species of other site units.  105 At Dunhead sedge - Mountain sagewort ecosystems are uncommon in the IMAwc occurring on wind-swept middle and upper slopes with relatively deep soils. Snow cover is thin and discontinuous but high vegetation cover limits cryoturbation features on these sites. These ecosystems commonly have high vegetation cover and Table 20 Continued from previous page   133  appear as grassland in many cases. Dunhead sedge is the site dominant but a high cover of mountain sagewort and moss campion is typical. Tundra species such as spiked wood-rush, diverse-leaved cinquefoil, showy sedge, and long-leaved starwart occur commonly. The moss layer may have higher cover of heron‟s-bill and haircap mosses.  106 At Woolly pussytoes - Mountain sagewort ecosystems occur on gradual upper slopes and moderately steep insolated slopes on neutral to warm aspects upper slopes. Soils are moderately well developed Sombric or Melanic Brunisols. A moderate snow cover regime largely protects site from soil freezing in the winter but ecosystems become snow-free relatively early in the growing season (mid-July) due to insolation. These ecosystems are generally adequately watered but may experience late summer moisture deficits in drier years. The low statured vascular vegetation generally has a  high cover dominated by woolly pussytoes. Mountain sagewort is also prominent and with high cover on most sites. Secondary species occurring with low cover include a range of snowbed and meadow species: yellow mountain-heather, sibbaldia, subalpine fleabane, slender hawkweed, alpine timothy, and others. Moss layer cover is commonly low but Lepraria neglecta or Polytrichum piliferum may occur where cryoturbation leads to exposed mineral soil. Species diversity is often low as most of the favourable growing sites are well occupied by a few dominant species.  110 Ah White and Pink mountain-heathers - Partridgefoot ecosystems primarily occupy cool and neutral aspect, and snowbed sites that have well drained stable soils. A relatively deep snowpack protects vegetation from soil freezing and winter desiccation. Sites become snow free from mid to late July. Soils are imperfectly drained, shallow to moderately deep and often have a gleyed horizon.  In general, vegetation cover is very high to continuous and dominated by mountain-heathers (Cassiope mertensiana, Phyllodoce empetriformis). Partridgefoot and slender hawkweed are common associates; other species of snow beds such as woolly pussytoes, sibbaldia, showy sedge, and others will occur on many sites but usually with low cover. Snowbed sites with soil movement or higher moisture favour alpine forbs so these sites might be embedded with both the Am## or Am## ecosystems. Later snowbeds and sites directly below late snowbeds are too extreme for the Ah## and usually occupied by the As.  111 Am Alpine sedge ecosystems occur in late snow lay areas on all aspects. These ecosystems are protected from winter conditions by deep snow deposits and soils do not freeze. Alpine sedge ecosystems commonly experience prolonged soil saturation during and following snow melt but they also occur on snow beds sites that are well-drained. These ecosystems are one of the last to become snow free in the spring with exposure occurring from mid-July to mid-August. The vegetation is usually well-developed and always dominated by black alpine sedge where sometimes it can be the only species present. This site unit is often adjacent to mountain-heather or woolly pussy-toes ecosystems and a scattering of species from these ecosystems can occur such as partridgefoot, white mountain-heather, woolly puusytoes, and slender hawkweed.  112 As Pyrenean sedge - Piper’s wood-rush ecosystems occur on the latest snow Table 20 Continued from previous page   134  lay areas of the subzone on cool aspect lee slopes where cornices and deep snow accumulates. Extended snow duration and snow melt saturation lead to frequent soil creep and nivation. Exposed wet fine-textured soils and low vegetation cover results in the frequent occurrence of needle ice cryoturbation features in the shoulder season when snow cover is absent. Sites become snow free as late as early August in extreme years but snow may disappear as early as first week of July in years with lower snowfall. The periodic extreme late snow release years likely create the conditions for this ecosystem. The sparse vegetation of this ecosystem is characterized by species tolerant of high soil instability and short growing season such as small-flowered or curved woodrush and Pyrenean sedge. Other species of snow beds may occur infrequently such as partridge-foot, woolly pussytoes, white mountain-heather, and chocolate chip lichen, and mountain sorrel.   Figure 50 Edatopic Grid for the IMAwc with Site Series.  Table 20 Continued from previous page   135   Figure 51 Landscape profile for IMAwc Site Series (e.g. 01 = Antennaria lanata – Artemisia norvegica).  Figure 52 Edatopic grid for the IMAwc with site associations (site units). Soil Moisture classes are labelled 0-7 ranging from very xeric to subhydric, and Snow Cover Regime classes are labelled A –F ranging from earliest exposed (most shallow snow cover) to last exposed (deepest snow cover).  136   Figure 53 Landscape diagram for the IMAwc with site associations, created by Will Mackenzie.  Figure 54 Edatopic grid for the ESSFwcp with Site Series.   137   Figure 55 Landscape diagram for the ESSFwcp Site Series.   138   Figure 56 Edatopic Grid for the ESSFwcp with site associations (site units).   139   Figure 57 Landscape diagram for the ESSFwcp with site association nomenclature (eg. 110 SK14 = Subalpine Fir – Showy Sedge – Valerian).  Table 21 Key to Site Units of the ESSFwcp  1a. Tree cover > 40%, soils deeper than 20cm  2a. Moderate to well-drained, submesic to subhygric soils; average to moderately-late snowmelt; warm to neutral aspect; Showy-sedge, arnica and sitka valerian are sparse but common in the understory and highly abundant in the adjacent openings.  ESSFwcp 110 Sk14   Bl – Showy Sedge – Valerian      2b. Very rapid to well drained, xeric to submesic soils; early to average snowmelt; upper slope or broad ridge crests, frequently on but not restricted to cool aspects. Heathers and huckleberry are common in the understory with occasional white rhododendron, violets and mitreworts.    140   ESSFwcp 103 SK01 Bl – Mountain-heather – Barbilophozia   1b. Tree cover 5-40%; shallow, blocky soils.  2c. Very rapid to rapid drainage, very xeric to xeric soils; earliest to early snowmelt; crest or ridge-top slope position. Trees are predominantly scattered rather than clumped and intermixed with crowberry, juniper, mountain sagewort and heathers; yellow glacier lilies and spring-beauty are found in moisture-collecting areas within the site unit.     ESSFwcp 102Ro SK62 Bl- Juniper - Crowberry 1c. Tree cover less than 10% 3a. Submesic to mesic, well drained to mod. well drained soils with very weak or no humus layer.  4a. Neutral to warm aspect,  mod. steep to  gentle slopes, early to average snowmelt.   ESSFwcp 101 Am03 Valerian - Pasqueflower 4b. Slightly cool to cold aspect, very steep to gentle slopes, late snowmelt. Abundant or patchy heathers with frequent herbaceous species   ESSFwcp 111 Ah03b White mountain-heather - Pasqueflower 3b. Mesic to subhygric, mod. well drained to imperfectly drained soils with weak to moderate (.5 to 2cm) humus layer. Claytonia?  5a.  Warm to neutral aspect, early to average snowmelt, lush herbaceous layer.    ESSFwcp 112 Am02 Showy sedge – Valerian - Fleabane 5b. Cool aspect, late snowmelt, abundant moisture and a thick heather layer with partridgefoot dominating areas of excessive moisture and/or weak soil development.    ESSFwcp 111 Ah03 White mountain-heather - Partridgefoot Table 21 Continued from previous page   141  3c. Subhydric to Hygric soil moisture with weak or no humus layer, late to very late snowmelt, depressions or bottom of slopes. 6a. Weak soil development with high sand content, snow-bed areas found in any slope position, late snowmelt sometimes 1 or 2 months later than adjacent sites; very gentle or flat slopes; alpine sedge is strongly dominant and sometimes the only species, tufted bulrush is common on these sites.   ESSFwcp 113 As01 Alpine Sedge    6b. Deep poorly drained soils with occasional thick (10cm+) organic layer, saturated < 30cm from surface; at or near water source.   ESSFwcp 115 Wa13 Marsh marigold – Alpine Sedge       Table 22 Site Units of the ESSFwcp3. 110 ESSFwcp  Sk14   Bl – Showy Sedge – Valerian ecosystems are typically raised microsites in which earlier snowmelt –than in surrounding openings- allows for the establishment and development of tree islands. Trees are, on average, less than 16m tall and rarely exceed 200yrs of age due to stressful growing conditions. Due to low-light availability the herb layer under the canopy is usually sparse, but locally abundant in the exposed (warm) side of the tree-clumps, and in canopy openings. These tree-islands usually expand through layering (low branches become new stems) since conditions for cone development and seed germination are usually unfavourable. The herb layer is composed of species found in the adjacent openings and it is characterized by the presence of showy sedge, Sitka valerian, mountain sagewort and arrow-leaved groundsel.    103 ESSFwcp  SK01 Bl – Mountain-heather – Barbilophozia site units are commonly found on cool aspects with later snowmelt. Tree islands are similar in character and composition to the Sk14 although trees might also be present as scattered individuals and less-define „clumps‟ on steeper sites. The combination of long-lasting deep snowpacks Table 21 Continued from previous page   142  and well drained sites results in ecosystems well suited for tree species and a vigorous shrub layer. The understory is typically dominated by heathers and occasionally by black huckleberry. Under closed canopy conditions the understory is very sparse mostly restricted to mosses like Barbilophozia and shade tolerant herbs such as mountain hairgrass, partridgefoot and mountain arnica.   102Ro ESSFwcp SK62 Bl- Juniper – Crowberry ecosystems occur on ridges, summits and crests below 2100m.a.s.l. They are characterized by early snowmelt, high wind and sun exposure, and poor soil development with a high percent of coarse fragments. They are well drained sites in which tree growth is limited by moisture deficits and high wind exposure. Trees occur both as krummholz and as scattered individuals. While the sites are characterized by the presence of crowberry, a wide variety of shrubs and herbs can be found. On the warmer sites, common juniper can be found and it occasionally forms extensive mats. On moister holding sites yellow glacier lily can be prolific alongside showy sedge, fleabane, mountain sagewort and spring beauty. When this site unit occurs at its lower elevation boundaries with reduced wind exposure it results in highly fragmented forested sites with a diverse understory characterized by bracted lousewort. Heathers are commonly found in this site unit but are rarely dominant.   101 ESSFwcp  Am03 Valerian – Pasqueflower sites form small to extensive non-forested communities embedded in predominantly forested landscapes. Good soil drainage with a limited moisture supply (either from snowmelt or rainfall, rather than from sub irrigation or water-body proximity) creates ecosystems in which soil moisture is the main limiting factor to growth. Soils can range from poorly developed to moderately well developed. Sitka valerian and pasqueflower are commonly found on these sites and can be locally dominant yet these sites are characterized by a wide variety of species including small-flowered wood-rush, bracted lousewort, alpine timothy, mountain hairgrass, mountain sagewort, fleabane, bluegrass and hawkweed.   112 ESSFwcp  Am02 Showy sedge – Valerian – Fleabane ecosystems represent the typical subalpine 'meadow' communities. They occur on gentle to moderately steep slopes with good soil development. They represent the non-forested component of the subalpine matrix which is perpetually kept as non-forested sites since the length of the growing season is too short, and the competition from the herbaceous layer is too high to sustain tree species. This site unit has a high degree of species richness with a well-developed herb layer with vigorous growth. While showy sedge, fleabane, arctic lupine and Sitka valerian are typically dominant or co-dominant, Indian hellebore can be locally dominant on sub-irrigated sites with rapid drainage. These sites can also occur along stream banks and around lakes, in which case they can be adjacent to different non-forested site units.   111 ESSFwcp  Ah03b White mountain-heather – Pasqueflower site units are commonly found on neutral to cool aspects with deep snowpacks, forming extensive non-Table 22 Continued from previous page   143  forested ecosystems. Well drained soils prevent saturation and result in sites with higher species diversity than the Ah03 (below). White and pink mountain-heathers can form extensive mats but are usually intermixed with pasqueflower, mountain hairgrass, showy sedge and wood-rushes. This site unit is very similar to the Ah03 (below) albeit slightly drier (better drained soils with lower moisture input) and can occur on warmer sites.    111 ESSFwcp  Ah03 White mountain-heather – Partridgefoot ecosystems form the predominant non-forested sites on cool aspects with late snowmelt. Extensive mats of white mountain-heather result in sites with a very high percent vegetation cover with low diversity levels. Rocky areas and outcrops within these ecosystems are characterized by partridgefoot and Alaska saxifrage.  Soils are typically saturated with no moisture deficit, often subirrigated or in close proximity to ephemeral streams and rivulets. Raised microsites within this site unit are typically dominated by trees with low vigour and high stress from snowcreep and frequent avalanches. Frequent snow-sluffs (and avalanches) can extend this site unit well into the ESSFwcw3.    113 ESSFwcp  As01 Alpine Sedge ecosystems occur in late snow lay areas on warm and neutral aspects. These ecosystems are protected from winter conditions by deep snow deposits and soils do not freeze. Alpine sedge ecosystems commonly experience prolonged soil saturation during and following snow melt but they also occur on  snow beds sites that are well-drained. These ecosystems are one of the last to become snow free in the spring with exposure occurring from early to mid-July. The vegetation is usually well-developed and always dominated by black alpine sedge where it can form a near monoculture in some cases. This site unit is often adjacent to mountain-heather or woolly pussy-toes ecosystems and a scattering of species  from these ecosystems can occur such as partridgefoot, white mountain-heather,  woolly puusytoes, and slender hawkweed.  ESSFwcp 115 Wa13 Marsh marigold – Alpine Sedge site units are characterized by excessive moisture which limits the establishment of tree species. The communities within this site unit can be dominated by alpine sedge, sphagnum, white mountain marsh-marigold, bluejoint reedgrass.  Seepage is commonly present within the top 30 cm of soil, and deep organic soils are common. Rein orchids and globeflower can be locally co-dominant on raised microsites. This site unit can occur in a wide range of topographic conditions and different snowmelt timing, as long as there is an abundant water supply on imperfectly drained soils.      Table 22 Continued from previous page   144  Table 23 Principal Component Analysis for the measured edaphic and environmental variables. Number Eigenvalue Percent Cum Percent  1 3.2437 40.546 40.546  2 1.5789 19.736 60.282  3 1.2333 15.416 75.697  4 0.8924 11.156 86.853  5 0.5242 6.553 93.406  6 0.3622 4.528 97.934  7 0.1610 2.012 99.946  8 0.0043 0.054 100.000    Figure 58 Loading plot for Principal Components 1 and 2 for measured edaphic and environmental variables .  Table 24 Eigenvectors for each principal component resulting from edaphic and environmental variables.      1 2 3 4 5 6 7 8 Slope Gradient 0.13766 -0.55247 0.41597 -0.32146 0.20175 0.59779 0.03690 0.00564 Aspect -0.10851 0.63334 -0.29254 -0.21914 0.31750 0.59180 -0.04876 0.00986 Elevation -0.06209 0.23643 0.57408 0.68688 0.36395 0.07919 0.01007 0.00607 Org Matr 0.49384 -0.05807 -0.24853 0.32384 -0.15874 0.22504 0.02432 0.71392 NH4-N 0.45717 0.09337 -0.00801 -0.24415 0.55515 -0.36496 0.52993 0.01974 NO3-N 0.47933 0.17298 0.20523 -0.20909 0.10439 -0.19260 -0.77770 -0.04080 pH 0.21301 0.43790 0.48743 -0.23319 -0.60191 0.07446 0.32101 -0.00451             NO3-N NH4-N pH Slope Gradient Elevation Aspect Organic Matter  145         Table 26 Factor Structure shows correlations between variables and components. The underlined correlations indicate ≥ .70, the pre-established cut-off value. Factor Structure Slope Gradient Prin comp 1 0.0997854 Prin comp 2  -0.433246 Aspect -0.03507 0.5698424 Elevation -0.044323 0.2076307 Org Matter 0.8599166 -0.471357 NH4-N 0.770289 -0.061525 NO3-N 0.8634563 0.0648154 pH 0.4202639 0.4035163    Table 25 Final Communality Estimates (variance accounted for) for the Varimax Factor rotation on 2 the first two Principal Components (Table 12). Final Communality Estimates Slope Gradient    0.191  Aspect 0.324 Elevation 0.043  Org Matr 0.896 NH4-N 0.593 NO3-N 0.765 pH 0.372     146     Figure 59 Oblique Quartimax Factor Rotation for the first 2 Principal Components  Table 27 Variance explained by the Principal Components for three selected variables: Org Matter, NH4-N and NO3-N. Number Eigenvalue Percent Cum Percent  1 3.0548 76.371 76.371  2 0.7097 17.743 94.114  3 0.2355 5.886 100.00     Table 28 Correlations in the new Factor Analysis with reduced variables. Factor Structure     NH4-N 0.5499684 0.8304846 Organic Matter 0.9946802 0.6653981 NO3-N 0.5585119 0.8529756  Table 29 Communality estimates (variance accounted for). Final Communality Estimates    NH4-N 0.69180 Org Matr 0.99340 NO3-N 0.72908    NO3-N Organic Mater NH4-N pH H Slope Gradient Aspect Elevation  147   Figure 60 Rotated Factor Pattern (Oblique Quartimax) for the first 2 Principal Components. Although biologically this pattern is obvious, it‟s important to notice that the statistical soil analysis correspond to the biological conditions.                Organic Matter NH4-N NO3-N  148   Figure 61 CCA for all edaphic variables.     149     Figure 62 CCA for the four selected variables and their corresponding species.  150    Figure 63 PCA for the three selected variables and their corresponding species.   151   Figure 64 PCA for all edaphic variables axis 2vs1.  152   Figure 65 PCA for all edaphic variables Axis 3 and 1.       153   Figure 66 NMDS for all edaphic variables.        154  A Appendix B   Table 30 Results from different classifications methods (step 1 from  Classification Methodology) for all relevés (300). Coloured groups indicate the method selected for further analysis.                 Twinspan w/Sørensens pseuspp. cut levels 0 5 10 20 50 Six groups               1  2  3  4  5  6         Artemisia  norveigica Carex  spectabilis Juncus  drummundii Cassiope mertensiana Carex nigricans Cassiope mertensiana       Sibbaldia  procumbens Senecio triangularis Cassiope mertensiana Juncus  drummundii               Isopam                   1   2     3 4   5   6   7   8   9   Artemisia norvegica Lepraria neglecta Hieracium gracile Festuca brachyphylla Abies lasiocarpa Luetkea pectinata Luetkea pectinata Cassiope mertensiana Cassiope mertensiana Salix nivalis Artemisia norvegica Antennaria lanata Silene acaulis   Cassiope mertensiana Cassiope mertensiana  Phyllodoce empetriformis Cassiope tetragona   Artemisia norvegica           Abies lasiocarpa 10  11  12  13  14  15  16       Arnica latifolia Carex nigricans Senecio triangularis Erigeron peregrinus Valeriana sitchensis Senecio triangularis Senecio triangularis      Rubus pedatus       Carex spectabilis Valeriana sitchensis Valeriana sitchensis                                          Twinspan w/Sørensens pseuspp. cut levels 0 5 10 20 50 18 groups                     1  2   3  4  5  6  7  8  9 Silene acaulis Dryas octopetala Artemisia norvegica Picea engelmannii Senecio triangularis Senecio triangularis Valeriana sitchensis Carex spectabilis Cassiope mertensiana  spic      Lept pyr Car nigri  Phy emp Phy emp 155    Trisetum atum arrhena olifolia ex cans llodoce etriformis llodoce etriformis   10  11  12  13  14  15  16  17  18 Juncus drummondii Cassiope mertensiana Carex nigricans Cassiope mertensiana Carex nigricans Carex nigricans Abies lasiocarpa Cassiope mertensiana Carex spectabilis Carex pyrenaica Abies lasiocarpa   Carex nigricans Salix barclayi Caltha leptosepala   Luetkea pectinata Cassiope mertensiana Epilobium species Carex nigricans   Leptarrhena pyrolifolia         Vahlodea atropurpurea                                 Phleum alpinum    Table 30 Continued from previous page  156   Table 31 Results from different classifications methods (step 1 from  Classification Methodology) for ESSFwcp relevés (190). Coloured groups indicate the method selected for further analysis. 1  2  3  4  5  6  7  8  9  TWINSPAN (Whittaker’s div., pseudospecies cut levels = 0 5 10 20 50)                          Picea engelmannii Senecio triangularis Valeriana sitchensis Carex spectabilis Carex nigricans Cassiope mertensiana Arnica latifolia Abies lasiocarpa Vahlodea atropurpurea    Carex nigricans Senecio triangularis Valeriana sitchensis   Luetkea pectinata     Cassiope mertensiana      Veratrum viride Senecio triangularis            TWINSPAN  Sørensen’s   Pseusdospp. cut levels = 0 5 10 20 50                             Picea engelmannii Senecio triangularis Juncus drummondii Caltha leptosepala Carex spectabilis Valeriana sitchensis Cassiope mertensiana Arnica latifolia Carex spectabilis    Carex nigricans Epilobium species   Vahlodea atropurpurea Senecio triangularis Luetkea pectinata   Cassiope mertensiana                 Senecio triangularis                  TWINSPAN   Pseusdospp. cut levels = 0 5 10 20 50 Presence/Absence                              Picea engelmannii Carex nigricans Cassiope mertensiana Vahlodea atropurpurea Caltha leptosepala Valeriana sitchensis Carex spectabilis Vahlodea atropurpurea Cassiope mertensiana       Senecio triangularis     Senecio triangularis Abies lasiocarpa                              Artemisia norvegica         Isopam                                   Cassiope mertensiana Abies lasiocarpa Cassiope mertensiana Phyllodoce empetriformis Carex spectabilis Carex nigricans Leptarrhena pyrolifolia Vahlodea atropurpurea Erigeron peregrinus     Carex spectabilis   Valeriana sitchensis Arnica latifolia   Senecio triangularis Cassiope mertensiana    157     10       Cassiope mertensiana Artemisia norvegica     Philonotis fontana Luetkea pectinata   Valeriana sitchensis                 Senecio triangularis                 Veratrum viride                 Table 31 Continued from previous page  158  Table 32 Results from different classifications methods (step 1 from Classification Methodology) for the IMAwc relevés (110). Coloured groups indicate the method selected for further analysis. IMAwc All plots (110)                             Group No.                      1 2  3  4  5  6  7  8  9  TWINSPAN Sorensen 0 5 10 30 50                                Luetkea pectinata Cassiope mertensiana Luetkea pectinata Carex nigricans Cassiope mertensiana Antennaria lanata Silene acaulis Salix nivalis Artemisia norvegica Cassiope mertensiana Carex nigricans Carex nigricans   Luetkea pectinata Carex spectabilis Abies lasiocarpa              Dicranum scoparium     Antennaria lanata Hieracium gracile Saxifraga bronchialis         TWINSPAN fidelity (>18) Duf.-Leg.                                 Cassiope mertensiana Cassiope mertensiana Carex nigricans Carex nigricans Phyllodoce glanduliflora Erigeron peregrinus Salix arctica Artemisia norvegica Artemisia norvegica Luetkea pectinata Carex nigricans Dicranum scoparium Vahlodea atropurpurea Luzula piperi Arnica latifolia Epilobium latifolium Salix nivalis Sibbaldia procumbens         Marsupella species Trichophorum cespitosum Carex pyrenaica Valeriana sitchensis Silene acaulis Flavocetraria nivalis Lepraria neglecta TWINSPAN (Sørensen’s  index, pseudospecies cut levels 0 5 10 20 50)                           Cassiope mertensiana Cassiope mertensiana Luetkea pectinata Carex spectabilis Abies lasiocarpa Trisetum spicatum Salix nivalis Artemisia norvegica Artemisia norvegica Luetkea pectinata Carex nigricans Hieracium gracile     Epilobium latifolium Silene acaulis Dryas octopetala Sibbaldia procumbens Lepraria neglecta         Cassio merten         Poa alpina              159                        TWINSPAN (Whittaker's div. pseudospecies cut levels 0 5 10 20 50)                                    Cassiope mertensiana Cassiope mertensiana Cassiope mertensiana Antennaria lanata Erigeron peregrinus Silene acaulis Salix nivalis Artemisia norvegica Artemisia norvegica Carex nigricans Luetkea pectinata Caltha leptosepala Carex spectabilis Vahlodea atropurpurea Abies lasiocarpa   Carex spectabilis Phyllodoce glanduliflora   Antennaria lanata     Arnica latifolia Saxifraga bronchialis     Lepraria neglecta                   TWINSPAN Sørensen’s  index pseudospecies cut levels 0 5 10 20 50 Presence/Absence                       Luetkea pectinata Cassiope mertensiana Cassiope mertensiana Carex spectabilis Abies lasiocarpa Silene acaulis Salix nivalis Artemisia norvegica Artemisia norvegica Cassiope mertensiana Carex nigricans Carex nigricans     Epilobium latifolium Poa alpina Dryas octopetala Sibbaldia procumbens Lepraria neglecta         Luetkea pectinata         Trisetum spicatum             Isopam                          Salix nivalis Artemisia norvegica Artemisia norvegica Cassiope mertensiana Antennaria lanata             Silene acaulis Lepraria neglecta     Hieracium gracile         Table 32 Continued from previous page  160  Results from step 2 Objective 1 All Plots (300)  Table 33 Isopam method (criterion Percent Frequency ≥III). Group No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 No. of relevés 6 22 20 18 13 29 22 15 12 4 34 19 32 27 18 9              Phyllodoce empetriformis III I I I II IV IV II V IV II I III II I . Abies lasiocarpa III I I II V II II III V V I II II III I . Dryas octopetala III . I II I I . . . . . . . I . . Polytrichum piliferum III III IV II II I II . I . I . I . . . Solorina crocea III II III I . II II . . . I . I . . . Cladina mitis III I I I I . . I I . . . . . . . Dicranum scoparium III . I I . I II . I . I I I I I . Empetrum nigrum III III I II II . . . . . I . I . . . Solidago multiradiata III I I I . . I . . . . . . . . . Thamnolia vermicularis IV . . I . . . . . . . . . I . . Antennaria alpina IV II I IV I I I I . . I I I I . . Cetraria ericetorum IV . . I . I . I . . . . I I . . Cladonia uncialis IV . . . . . . . . . . . . . . . Sibbaldia procumbens IV IV IV III II I III . I . I I III I . . Gentiana glauca IV I I I . . . . . . . . I . . . Artemisia norvegica V V V IV III I II . . . I I III II I II Salix nivalis V II II IV . I I I . . . I I . . I Cassiope tetragona V III I I I . . . . . . . . I . .                   Lepraria neglecta . V II II II I I . I . . . I I . . Phyllodoce glanduliflora I IV III I II II I I II II I . I I . . Silene acaulis II III II IV II . I . . . . . . I . . Polytrichum juniperinum II III I I I I I . I . I I I I . .  161  Polytrichum piliferum III III IV II II I II . I . I . I . . . Empetrum nigrum III III I II II . . . . . I . I . . . Sibbaldia procumbens IV IV IV III II I III . I . I I III I . . Artemisia norvegica V V V IV III I II . . . I I III II I II Cassiope tetragona V III I I I . . . . . . . . I . .                   Arnica latifolia . . III I II I III II II V I I IV III II I Erigeron peregrinus . . III I II II III . . . I II V II IV II Phleum alpinum . . III I . I II . . . I II III I II II Hieracium gracile . I V I I III III I II . II I III I . . Antennaria lanata I II V II . III III I . . II II III I . . Carex spectabilis I I III II III II IV I III V III III IV IV IV III Vahlodea atropurpurea I I III I I II IV III II III II III IV III III II Phyllodoce glanduliflora I IV III I II II I I II II I . I I . . Cassiope mertensiana II I III I I V V V V V III III III I I . Luzula spicata II II III III I . . . . . I . . I . . Polytrichum piliferum III III IV II II I II . I . I . I . . . Solorina crocea III II III I . II II . . . I . I . . . Sibbaldia procumbens IV IV IV III II I III . I . I I III I . . Artemisia norvegica V V V IV III I II . . . I I III II I II                   Poa alpina . I II II . I I . I . I I II I . . Carex nardina . I I II I I . . . . . I . I . . Campanula lasiocarpa . II . II . . . . . . . . . . . . Lepraria neglecta . V II II II I I . I . . . I I . . Potentilla diversifolia . I I III I . I . . . . . I I . . Stellaria longifolia . I I II . . . . . . . . I . . . Saxifraga bronchialis . I . II I . . . . . I . I . . . Antennaria lanata I II V II . III III I . . II II III I . . Carex spectabilis I I III II III II IV I III V III III IV IV IV III Trisetum spicatum I I II II I I I . . . I I II I . I Selaginella densa I I I II . I I . . . . . I I . . Stereocaulon alpinum I II . II II I . . . . I . . . . . Carex phaeocephala I I I II . . . . . . . . . . . . Minuartia obtusiloba I I . II . . . . . . . . . . . . Festuca brachyphylla I II II IV I I I . . . I . I I . . Silene acaulis II III II IV II . I . . . . . . I . . Group No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 No. of relevés 6 22 20 18 13 29 22 15 12 4 34 19 32 27 18 9   162  Carex albonigra II I . II . . I . . . . . . . . . Luzula spicata II II III III I . . . . . I . . I . . Abies lasiocarpa III I I II V II II III V V I II II III I . Dryas octopetala III . I II I I . . . . . . . I . . Polytrichum piliferum III III IV II II I II . I . I . I . . . Empetrum nigrum III III I II II . . . . . I . I . . . Antennaria alpina IV II I IV I I I I . . I I I I . . Sibbaldia procumbens IV IV IV III II I III . I . I I III I . . Artemisia norvegica V V V IV III I II . . . I I III II I II Salix nivalis V II II IV . I I I . . . I I . . I                   Carex spectabilis I I III II III II IV I III V III III IV IV IV III Abies lasiocarpa III I I II V II II III V V I II II III I . Artemisia norvegica V V V IV III I II . . . I I III II I II                   Hieracium gracile . I V I I III III I II . II I III I . . Antennaria lanata I II V II . III III I . . II II III I . . Luetkea pectinata I I I I . V V IV II . III III II I I . Cassiope mertensiana II I III I I V V V V V III III III I I . Phyllodoce empetriformis III I I I II IV IV II V IV II I III II I .                   Arnica latifolia . . III I II I III II II V I I IV III II I Valeriana sitchensis . I I . I II III II I II I II IV IV V V Veratrum viride . . I . . I III II I . I II III III V V Erigeron peregrinus . . III I II II III . . . I II V II IV II Anemone occidentalis . . I . II II IV II I . II I IV II III . Dicranum species . . I I . I III I I IV I . I I I . Hieracium gracile . I V I I III III I II . II I III I . . Antennaria lanata I II V II . III III I . . II II III I . . Carex spectabilis I I III II III II IV I III V III III IV IV IV III Vahlodea atropurpurea I I III I I II IV III II III II III IV III III II Luetkea pectinata I I I I . V V IV II . III III II I I . Cassiope mertensiana II I III I I V V V V V III III III I I . Luzula piperi II I II . . II III . . . I I II I I . Phyllodoce empetriformis III I I I II IV IV II V IV II I III II I . Sibbaldia procumbens I IV IV III II I III . I . I I III I . . Group No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 No. of relevés 6 22 20 18 13 29 22 15 12 4 34 19 32 27 18 9   163  V                   Vaccinium membranaceum . . . . I I II III IV . . . I I I . Barbilophozia floerkei . . I . I I II III I II I I I I . . Vahlodea atropurpurea I I III I I II IV III II III II III IV III III II Luetkea pectinata I I I I . V V IV II . III III II I I . Cassiope mertensiana II I III I I V V V V V III III III I I . Abies lasiocarpa III I I II V II II III V V I II II III I .                   Barbilophozia lycopodioides . . . . . I I I III III I I I I I . Vaccinium membranaceum . . . . I I II III IV . . . I I I . Barbilophozia species . . . . I I I . III II . . I I . . Carex spectabilis I I III II III II IV I III V III III IV IV IV III Cassiope mertensiana II I III I I V V V V V III III III I I . Phyllodoce empetriformis III I I I II IV IV II V IV II I III II I . Abies lasiocarpa III I I II V II II III V V I II II III I .                   Arnica latifolia . . III I II I III II II V I I IV III II I Rubus pedatus . . . . . I . I II V I . . I I . Luzula parviflora . I I . I II II I I V I I I II II II Mitella breweri . . . . . I I . . V . . . II II I Brachythecium species . . I . . I II I I III I I II I II II Barbilophozia lycopodioides . . . . . I I I III III I I I I I . Dicranum species . . I I . I III I I IV I . I I I . Carex spectabilis I I III II III II IV I III V III III IV IV IV III Vahlodea atropurpurea I I III I I II IV III II III II III IV III III II Cassiope mertensiana II I III I I V V V V V III III III I I . Picea engelmannii II I I I II I I I II III I . I II . . Phyllodoce empetriformis III I I I II IV IV II V IV II I III II I . Abies lasiocarpa III I I II V II II III V V I II II III I .                   Carex nigricans . . I . I II II II I . V III II I . . Carex spectabilis I I III II III II IV I III V III III IV IV IV III Luetkea pectinata I I I I . V V IV II . III III II I I . Group No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 No. of relevés 6 22 20 18 13 29 22 15 12 4 34 19 32 27 18 9   164   mertensiana II I III I I V V V V V III III III I I .                   Carex nigricans . . I . I II II II I . V III II I . . Senecio triangularis . . . . . I II I . . II V III III V V Equisetum arvense . . . . . . . I . . . III I . . II Leptarrhena pyrolifolia . . . . . . I I . . I IV I I . I Aulacomnium palustre . . . I . . I I . . I III I I I I Juncus mertensianus . . . . . I I . . . I III I . . I Caltha leptosepala . . . . . I . I . . I IV I I . I Philonotis fontana . . . . . . . . . . I III I I . . Parnassia fimbriata . . . . . . I . . . . III I . I . Epilobium species . . . . . . I . . . I III I I I . Juncus drummondii . . I . . I I I . . II III I I I . Carex spectabilis I I III II III II IV I III V III III IV IV IV III Vahlodea atropurpurea I I III I I II IV III II III II III IV III III II Luetkea pectinata I I I I . V V IV II . III III II I I . Cassiope mertensiana II I III I I V V V V V III III III I I .                   Arnica latifolia . . III I II I III II II V I I IV III II I Senecio triangularis . . . . . I II I . . II V III III V V Valeriana sitchensis . I I . I II III II I II I II IV IV V V Veratrum viride . . I . . I III II I . I II III III V V Erigeron peregrinus . . III I II II III . . . I II V II IV II Pedicularis bracteosa . . I . I . I I . . . I III I III II Phleum alpinum . . III I . I II . . . I II III I II II Anemone occidentalis . . I . II II IV II I . II I IV II III . Hieracium gracile . I V I I III III I II . II I III I . . Antennaria lanata I II V II . III III I . . II II III I . . Carex spectabilis I I III II III II IV I III V III III IV IV IV III Vahlodea atropurpurea I I III I I II IV III II III II III IV III III II Cassiope mertensiana II I III I I V V V V V III III III I I . Phyllodoce empetriformis III I I I II IV IV II V IV II I III II I . Sibbaldia procumbens IV IV IV III II I III . I . I I III I . . Artemisia norvegica V V V IV III I II . . . I I III II I II                   Arnica latifolia . . III I II I III II II V I I IV III II I Senecio triangularis . . . . . I II I . . II V III III V V Valeriana sitchensis . I I . I II III II I II I II IV IV V V Veratrum viride . . I . . I III II I . I II III III V V Carex spectabilis I I III II III II IV I III V III III IV IV IV III Group No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 No. of relevés 6 22 20 18 13 29 22 15 12 4 34 19 32 27 18 9   165                                     Vahlodea atropurpurea I I III I I II IV III II III II III IV III III II Abies lasiocarpa III I I II V II II III V V I II II III I .                   Senecio triangularis . . . . . I II I . . II V III III V V Valeriana sitchensis . I I . I II III II I II I II IV IV V V Veratrum viride . . I . . I III II I . I II III III V V Epilobium angustifolium . . . . I I I . . . . . I I III V Erigeron peregrinus . . III I II II III . . . I II V II IV II Pedicularis bracteosa . . I . I . I I . . . I III I III II Anemone occidentalis . . I . II II IV II I . II I IV II III . Carex spectabilis I I III II III II IV I III V III III IV IV IV III Vahlodea atropurpurea I I III I I II IV III II III II III IV III III II                   Senecio triangularis . . . . . I II I . . II V III III V V Valeriana sitchensis . I I . I II III II I II I II IV IV V V Thalictrum occidentale . . . . . . I . . . . . I I II V Heracleum maximum . . . . . . . . . . . . . I II V Veratrum viride . . I . . I III II I . I II III III V V Epilobium angustifolium . . . . I I I . . . . . I I III V Mitella pentandra . . . . . . . I . . . . I . I III Trollius albiflorus . . . . . . . I . . . I I I I III Viola glabella . . . . . I I . . . . . I I I III Carex spectabilis I I III II III II IV I III V III III IV IV IV III Calamagrostis canadensis I . . . . . . . . . I I . I I III Group No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 No. of relevés 6 22 20 18 13 29 22 15 12 4 34 19 32 27 18 9   166   Table 34 All Plots (300) TWINSPAN (Sørensen’s  index, pseudospecies cut levels 0 5 10 20 50). Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50 Artemisia norvegica 108 III IV V . . II II . . . . . . . . I I IV Poa alpina 41 III I I . . . I . . III . . I . IV I I II Silene acaulis 40 V IV IV . . . I . . . . . . . . I . I Trisetum spicatum 37 V I I . . . I . V . . . . II . . I II Festuca brachyphylla 37 III III III . . . I . . . . . . I . I I I Luzula spicata 35 III III III . . . I . . . . . . I . I . I Cladonia ecmocyna 23 III I I . . . . . . . . . . . . I I I Epilobium anagallidifolium 22 III . . . . . I . . . . . II I . I I I Potentilla diversifolia 15 III I I . . . I . . . . . . . . . . I Dryas octopetala 14 III V I . . . . . . . . . . . . I I . Cetraria ericetorum 13 III I I . . . I . . . . . . . . . I I Saxifraga bronchialis 12 III II I . . . . . . . . . . I . I . I Carex phaeocephala 12 III I II . . . . . . . . . . . . . . . Salix arctica 11 III . . . . . I . V . . . . . . . I I Minuartia obtusiloba 8 V I I . . . . . . . . . . . . . . . Peltigera malacea 8 III . I . . . . . . . . . . . . . . I Cladonia chlorophaea 5 III . . . . . I . . . . . . . . . I I Cladonia fimbriata 2 III . . . . . . . . . . . . . . . . I Stellaria longipes 1 III . . . . . . . . . . . . . . . . . Antennaria monocephala 1 III . . . . . . . . . . . . . . . . . Bromus vulgaris 1 III . . . . . . . . . . . . . . . . . Peltigera ponojensis 1 III . . . . . . . . . . . . . . . . .                      Artemisia norvegica 108 III IV V . . II II . . . . . . . . I I IV Antennaria alpina 47 . III III . . . I . V . . . . . I I I I  167  Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50 Silene acaulis 40 V IV IV . . . I . . . . . . . . I . I Festuca brachyphylla 37 III III III . . . I . . . . . . I . I I I Salix nivalis 36 . IV II . . . I . V . . . . . . . I I Luzula spicata 35 III III III . . . I . . . . . . I . I . I Dryas octopetala 14 III V I . . . . . . . . . . . . I I . Carex nardina 13 . III I . . I . . . . . I . . . I I .                      Artemisia norvegica 108 III IV V . . II II . . . . . . . . I I IV Sibbaldia procumbens 88 . II IV . . . I . . . . . I I . I I III Antennaria lanata 88 . I III . . . I . . III . . III II III . II III Phyllodoce glanduliflora 55 . II III . . . I . . . . . I . II II I II Polytrichum piliferum 51 . II III . . . I . . . . . . I . I I II Antennaria alpina 47 . III III . . . I . V . . . . . I I I I Lepraria neglecta 44 . . IV . . . . . . . . . . . . I I I Silene acaulis 40 V IV IV . . . I . . . . . . . . I . I Festuca brachyphylla 37 III III III . . . I . . . . . . I . I I I Luzula spicata 35 III III III . . . I . . . . . . I . I . I Empetrum nigrum 27 . I III . . . . . . . V . . . . I . I Cassiope tetragona 25 . II III . . . . . . . . . . . . I . I                      Picea engelmannii 32 . I I V . II I . . . V . . . . II I I Salix barrattiana 3 . . . V . I I . . . . . . . . . . . Pohlia wahlenbergii 3 . . . V V . . . . . . I . . . . . . Carex aquatilis 2 . . . V V . . . . . . . . . . . . .                      Senecio triangular 95 . . . . V V I . . I . I I . I I I I 168  Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50 is V II V II I I Carex nigricans 78 . . I . V III I . . . V V V V V I II II Salix barclayi 8 . . . . V . I . . . . . . . . . I I Pohlia wahlenbergii 3 . . . V V . . . . . . I . . . . . . Carex aquatilis 2 . . . V V . . . . . . . . . . . . . Carex lenticularis 1 . . . . V . . . . . . . . . . . . . Viola lanceolata 1 . . . . V . . . . . . . . . . . . . Aster species 1 . . . . V . . . . . . . . . . . . . Carex canescens 1 . . . . V . . . . . . . . . . . . .                      Carex spectabilis 152 . . II . . IV IV V . III . II III IV I III II IV Valeriana sitchensis 109 . . I . . III V . . III . I III . I II II II Erigeron peregrinus 97 . . I . . IV IV . . . . II I . . I I III Abies lasiocarpa 96 . II II . . III II . . . V II . . . V II I Senecio triangularis 95 . . . . V V IV . . III . IV III . II I I II Carex nigricans 78 . . I . V III I . . . V V V V V I II II Juncus drummondii 35 . . . . . IV I V V V . II III I IV . I I Caltha leptosepala 30 . . . . . V I . . . V IV III I . . I I Leptarrhena pyrolifolia 24 . . . . . V I . V III V II V . I . I I Aulacomnium palustre 23 . . I . . III I . V . . III III I . . I I Trollius albiflorus 20 . . . . . III II . . . . . . . . . I I Parnassia fimbriata 15 . . . . . IV I . V . . II II . . . I I Equisetum arvense 14 . . . . . IV I . V III . . I . . . I . Eriophorum angustifolium 11 . . . . . III I . . . V III . . . . . . Rhytidiadelphus squarrosus 10 . . . . . III I . . . . . III . . . . .  169  Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50                      Carex spectabilis 152 . . II . . IV IV V . III . II III IV I III II IV Vahlodea atropurpurea 117 . I I . . II III . . III . II IV IV I II III IV Valeriana sitchensis 109 . . I . . III V . . III . I III . I II II II Arnica latifolia 99 . . I . . II III . . . . . II . II III II IV Erigeron peregrinus 97 . . I . . IV IV . . . . II I . . I I III Senecio triangularis 95 . . . . V V IV . . III . IV III . II I I II Anemone occidentalis 82 . . . . . I III . . . . . III II I I II III Veratrum viride 80 . . . . . II IV . . . . III I . I I I II                      Carex spectabilis 152 . . II . . IV IV V . III . II III IV I III II IV Phyllodoce empetriformis 109 . . I . . II I V V . . . . III I IV IV III Luetkea pectinata 97 . . I . . I I V . III . III IV II IV II V II Solorina crocea 35 . I II . . . . V . . . . . . I . I I Juncus drummondii 35 . . . . . IV I V V V . II III I IV . I I Carex pyrenaica 25 . I I . . . . V . V . . . . . I I I Polytrichum sexangulare 6 . I . . . . . V . . . . . . II . I .                      Cassiope mertensiana 145 . II II . . . I . V III V II V III IV IV V IV Phyllodoce empetriformis 109 . . I . . II I V V . . . . III I IV IV III Phleum alpinum 59 . . I . . II II . V . . . . I . . I III Antennaria alpina 47 . III III . . . I . V . . . . . I I I I Trisetum spicatum 37 V I I . . . I . V . . . . II . . I II Salix nivalis 36 . IV II . . . I . V . . . . . . . I I  170  Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50 Juncus drummondii 35 . . . . . IV I V V V . II III I IV . I I Leptarrhena pyrolifolia 24 . . . . . V I . V III V II V . I . I I Aulacomnium palustre 23 . . I . . III I . V . . III III I . . I I Juncus mertensianus 20 . . . . . II I . V III . II III I I . I I Epilobium latifolium 15 . . . . . I I . V . . . . . . I I I Parnassia fimbriata 15 . . . . . IV I . V . . II II . . . I I Philonotis fontana 15 . . . . . II I . V V . II III I I . . . Equisetum arvense 14 . . . . . IV I . V III . . I . . . I . Salix arctica 11 III . . . . . I . V . . . . . . . I I Saxifraga lyallii 8 . . I . . I I . V III . I . . . . I I Sanionia uncinata 7 . . I . . I . . V . . I III . . . . . Myosotis asiatica 5 . . I . . . . . V . . I . . . . . I Marchantia species 5 . . . . . . I . V . . I . . I . . . Carex lachenalii 3 . . . . . . . . V . . . . I . . . I Hamatocaulis vernicosus 1 . . . . . . . . V . . . . . . . . .                      Carex spectabilis 152 . . II . . IV IV V . III . II III IV I III II IV Cassiope mertensiana 145 . II II . . . I . V III V II V III IV IV V IV Vahlodea atropurpurea 117 . I I . . II III . . III . II IV IV I II III IV Valeriana sitchensis 109 . . I . . III V . . III . I III . I II II II Luetkea pectinata 97 . . I . . I I V . III . III IV II IV II V II Senecio triangularis 95 . . . . V V IV . . III . IV III . II I I II Antennaria lanata 88 . I III . . . I . . III . . III II III . II III Luzula piperi 48 . I I . . . I . . III . . . I II . I III Poa alpina 41 III I I . . . I . . III . . I . IV I I II Brachythecium species 39 . . . . . . II . . III . . . . . I I I  171  Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50 Juncus drummondii 35 . . . . . IV I V V V . II III I IV . I I Carex pyrenaica 25 . I I . . . . V . V . . . . . I I I Veronica wormskjoldii 25 . . I . . I I . . III . . I . . . I II Leptarrhena pyrolifolia 24 . . . . . V I . V III V II V . I . I I Epilobium species 23 . . . . . II I . . V . I II . . . I I Juncus mertensianus 20 . . . . . II I . V III . II III I I . I I Philonotis fontana 15 . . . . . II I . V V . II III I I . . . Equisetum arvense 14 . . . . . IV I . V III . . I . . . I . Anemone multifida 10 . . . . . . I . . III . I I . II . . I Saxifraga lyallii 8 . . I . . I I . V III . I . . . . I I Campylium stellatum 8 . . I . . . . . . V . . . . . I I I Scapania species 5 . . . . . . . . . III . II . . I . I . Carex lenticularis 2 . . . . . . . . . III . . I . . . . . Bryum weigelii 2 . . . . . . I . . III . . . . . . . . Drepanocladus aduncus 1 . . . . . . . . . III . . . . . . . . Pellia species 1 . . . . . . . . . III . . . . . . . . Phyllodoce species 1 . . . . . . . . . III . . . . . . . . Paludella species 1 . . . . . . . . . III . . . . . . . . Cephalozia species 1 . . . . . . . . . III . . . . . . . .                      Cassiope mertensiana 145 . II II . . . I . V III V II V III IV IV V IV Abies lasiocarpa 96 . II II . . III II . . . V II . . . V II I Carex nigricans 78 . . I . V III I . . . V V V V V I II II Picea engelmannii 32 . I I V . II I . . . V . . . . II I I  172  Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50 Caltha leptosepala 30 . . . . . V I . . . V IV III I . . I I Empetrum nigrum 27 . I III . . . . . . . V . . . . I . I Leptarrhena pyrolifolia 24 . . . . . V I . V III V II V . I . I I Rubus pedatus 14 . . . . . . I . . . V . . . . II I . Eriophorum angustifolium 11 . . . . . III I . . . V III . . . . . . Polytrichum strictum 11 . . I . . . I . . . V I I II . I . I Kalmia microphylla 7 . . . . . . . . . . V II . . . I I I Rhododendron albiflorum 7 . . . . . I I . . . V . . . . I I . Arnica cordifolia 5 . . . . . . . . . . V . . . . I . I Streptopus lanceolatus 5 . . . . . . I . . . V . . . . . . . Ptilidium species 3 . . . . . . . . . . V . . . I . I . Vaccinium ovalifolium 2 . . . . . I . . . . V . . . . . . . Sphagnum squarrosum 2 . . . . . . . . . . V . I . . . . . Veronica beccabunga 2 . . . . . . . . . . V . . I . . . . Viola nephrophylla 2 . . . . . I . . . . V . . . . . . . Carex leptalea 1 . . . . . . . . . . V . . . . . . . Carex prasina 1 . . . . . . . . . . V . . . . . . . Carex pauciflora 1 . . . . . . . . . . V . . . . . . . Tsuga heterophylla 1 . . . . . . . . . . V . . . . . . . Carex lasiocarpa 1 . . . . . . . . . . V . . . . . . . Sphagnum capillifolium 1 . . . . . . . . . . V . . . . . . .                      Luetkea pectinata 97 . . I . . I I V . III . III IV II IV II V II Senecio triangularis 95 . . . . V V IV . . III . IV III . II I I II Veratrum viride 80 . . . . . II IV . . . . III I . I I I II Carex nigricans 78 . . I . V III I . . . V V V V V I II II Caltha leptosepala 30 . . . . . V I . . . V IV III I . . I I  173  Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50 Aulacomnium palustre 23 . . I . . III I . V . . III III I . . I I Eriophorum angustifolium 11 . . . . . III I . . . V III . . . . . . Sphagnum russowii 5 . . . . . . . . . . . III . . . . . I                      Carex spectabilis 152 . . II . . IV IV V . III . II III IV I III II IV Cassiope mertensiana 145 . II II . . . I . V III V II V III IV IV V IV Vahlodea atropurpurea 117 . I I . . II III . . III . II IV IV I II III IV Valeriana sitchensis 109 . . I . . III V . . III . I III . I II II II Luetkea pectinata 97 . . I . . I I V . III . III IV II IV II V II Senecio triangularis 95 . . . . V V IV . . III . IV III . II I I II Antennaria lanata 88 . I III . . . I . . III . . III II III . II III Anemone occidentalis 82 . . . . . I III . . . . . III II I I II III Carex nigricans 78 . . I . V III I . . . V V V V V I II II Juncus drummondii 35 . . . . . IV I V V V . II III I IV . I I Caltha leptosepala 30 . . . . . V I . . . V IV III I . . I I Leptarrhena pyrolifolia 24 . . . . . V I . V III V II V . I . I I Aulacomnium palustre 23 . . I . . III I . V . . III III I . . I I Juncus mertensianus 20 . . . . . II I . V III . II III I I . I I Philonotis fontana 15 . . . . . II I . V V . II III I I . . . Rhytidiadelphus squarrosus 10 . . . . . III I . . . . . III . . . . . Sanionia uncinata 7 . . I . . I . . V . . I III . . . . . Petasites frigidus 7 . . . . . II I . . . . . III . . . I .                      Carex spectabilis 152 . . II . . IV IV V . III . II III IV I III II IV  174  Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50 Cassiope mertensiana 145 . II II . . . I . V III V II V III IV IV V IV Vahlodea atropurpurea 117 . I I . . II III . . III . II IV IV I II III IV Phyllodoce empetriformis 109 . . I . . II I V V . . . . III I IV IV III Carex nigricans 78 . . I . V III I . . . V V V V V I II II                      Cassiope mertensiana 145 . II II . . . I . V III V II V III IV IV V IV Luetkea pectinata 97 . . I . . I I V . III . III IV II IV II V II Antennaria lanata 88 . I III . . . I . . III . . III II III . II III Hieracium gracile 82 . . II . . I I . . . . . I I III I II IV Carex nigricans 78 . . I . V III I . . . V V V V V I II II Polytrichum juniperinum 45 . I II . . . I . . . . . I II III I I II Poa alpina 41 III I I . . . I . . III . . I . IV I I II Juncus drummondii 35 . . . . . IV I V V V . II III I IV . I I Dicranum scoparium 29 . I I . . I I . . . . . I . III I I I                      Carex spectabilis 152 . . II . . IV IV V . III . II III IV I III II IV Cassiope mertensiana 145 . II II . . . I . V III V II V III IV IV V IV Phyllodoce empetriformis 109 . . I . . II I V V . . . . III I IV IV III Arnica latifolia 99 . . I . . II III . . . . . II . II III II IV Abies lasiocarpa 96 . II II . . III II . . . V II . . . V II I                      Cassiope mertensiana 145 . II II . . . I . V III V II V III IV IV V IV Vahlodea atropurpurea 117 . I I . . II III . . III . II IV IV I II III IV Phyllodoce empetriformis 109 . . I . . II I V V . . . . III I IV IV III  175  Group No.  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 No. of Rel. 2 10 44 1 1 8 63 1 1 2 1 9 7 9 8 34 49 50 Luetkea pectinata 97 . . I . . I I V . III . III IV II IV II V II                      Carex spectabilis 152 . . II . . IV IV V . III . II III IV I III II IV Cassiope mertensiana 145 . II II . . . I . V III V II V III IV IV V IV Vahlodea atropurpurea 117 . I I . . II III . . III . II IV IV I II III IV Phyllodoce empetriformis 109 . . I . . II I V V . . . . III I IV IV III Artemisia norvegica 108 III IV V . . II II . . . . . . . . I I IV Arnica latifolia 99 . . I . . II III . . . . . II . II III II IV Erigeron peregrinus 97 . . I . . IV IV . . . . II I . . I I III Sibbaldia procumbens 88 . II IV . . . I . . . . . I I . I I III Antennaria lanata 88 . I III . . . I . . III . . III II III . II III Hieracium gracile 82 . . II . . I I . . . . . I I III I II IV Anemone occidentalis 82 . . . . . I III . . . . . III II I I II III Phleum alpinum 59 . . I . . II II . V . . . . I . . I III Luzula piperi 48 . I I . . . I . . III . . . I II . I III       176  ESSF wcp (190 plots)   Table 35 TWINSPAN (Whittaker’s div., pseudospecies cut levels = 0 5 10 20 50). Group No.  1 2 3 4 5 6 7 8 9 No. of Relevés Freq. 1 31 21 32 4 42 7 31 21 Picea engelmannii 24 V . . I II I . III I Salix barrattiana 3 V I I . . . . . . Pohlia wahlenbergii 3 V I . . . . . . . Carex aquatilis 2 V I . . . . . . .             Carex spectabilis 107 . III III V . II IV IV IV Vahlodea atropurpurea 93 . III II III III III III II V Senecio triangularis 85 . IV V V . I . I II Carex nigricans 52 . IV . II V II I I I Juncus drummondii 24 . III I I II I . . . Leptarrhena pyrolifolia 23 . III I I IV I . . I Caltha leptosepala 23 . III I I II I . . .             Carex spectabilis 107 . III III V . II IV IV IV Valeriana sitchensis 98 . II V V . II II II IV Senecio triangularis 85 . IV V V . I . I II Veratrum viride 77 . II V IV II II I I II Erigeron peregrinus 69 . II III V . I IV I III Epilobium angustifolium 28 . . IV II . . I I II Thalictrum occidentale 18 . . III I . . . . I Heracleum maximum 15 . . III I . . . . .             Carex spectabilis 107 . III III V . II IV IV IV Valeriana sitchensis 98 . II V V . II II II IV Vahlodea atropurpurea 93 . III II III III III III II V Senecio triangularis 85 . IV V V . I . I II Arnica latifolia 80 . I I IV . III V III III Veratrum viride 77 . II V IV II II I I II Anemone occidentalis 69 . II I IV . II II I III Erigeron peregrinus 69 . II III V . I IV I III Phleum alpinum 38 . I I III . . III . III Pedicularis bracteosa 32 . I II III . I . . II Claytonia lanceolata 21 . . I III . . I I I             Vahlodea atropurpurea 93 . III II III III III III II V Cassiope mertensiana 90 . II . I IV V II III V  177  Group No.  1 2 3 4 5 6 7 8 9 No. of Relevés Freq. 1 31 21 32 4 42 7 31 21 Abies lasiocarpa 75 . I I II III III . V III Luetkea pectinata 65 . II . I IV V . I II Carex nigricans 52 . IV . II V II I I I Leptarrhena pyrolifolia 23 . III I I IV I . . I Eriophorum angustifolium 11 . II I . III . . . . Polytrichum strictum 6 . . . I IV . . I I Sphagnum squarrosum 2 . . . . III . . . .             Vahlodea atropurpurea 93 . III II III III III III II V Cassiope mertensiana 90 . II . I IV V II III V Arnica latifolia 80 . I I IV . III V III III Phyllodoce empetriformis 76 . I I II . IV I IV IV Abies lasiocarpa 75 . I I II III III . V III Luetkea pectinata 65 . II . I IV V . I II Barbilophozia floerkei 31 . I . I . III I I I             Carex spectabilis 107 . III III V . II IV IV IV Vahlodea atropurpurea 93 . III II III III III III II V Arnica latifolia 80 . I I IV . III V III III Erigeron peregrinus 69 . II III V . I IV I III Artemisia norvegica 46 . I I II . I IV I IV Hieracium gracile 43 . I . II . II III I III Phleum alpinum 38 . I I III . . III . III Antennaria lanata 37 . I . I . II III . II Dicranum species 28 . . . I II II III I I Polytrichum piliferum 19 . . . I . I III I II Polytrichum juniperinum 17 . I . I II I IV . I Coelocaulon species 8 . . . . . I III I I             Carex spectabilis 107 . III III V . II IV IV IV Cassiope mertensiana 90 . II . I IV V II III V Arnica latifolia 80 . I I IV . III V III III Phyllodoce empetriformis 76 . I I II . IV I IV IV Abies lasiocarpa 75 . I I II III III . V III Picea engelmannii 24 V . . I II I . III I Barbilophozia species 17 . . . . . I . III I             Carex spectabilis 107 . III III V . II IV IV IV Valeriana sitchensis 98 . II V V . II II II IV  178  Group No.  1 2 3 4 5 6 7 8 9 No. of Relevés Freq. 1 31 21 32 4 42 7 31 21 Vahlodea atropurpurea 93 . III II III III III III II V Cassiope mertensiana 90 . II . I IV V II III V Arnica latifolia 80 . I I IV . III V III III Phyllodoce empetriformis 76 . I I II . IV I IV IV Abies lasiocarpa 75 . I I II III III . V III Anemone occidentalis 69 . II I IV . II II I III Erigeron peregrinus 69 . II III V . I IV I III Artemisia norvegica 46 . I I II . I IV I IV Hieracium gracile 43 . I . II . II III I III Phleum alpinum 38 . I I III . . III . III Vaccinium membranaceum 37 . . I I . II . I III Sibbaldia procumbens 36 . I . II . I I II III      Table 36 TWINSPAN Sørensen’s Pseudospecies cut levels = 0 5 10 20 50. Group No.  1 2 3 4 5 6 7 8 9 No. of  1 1 2 9 19 53 46 7 52            Picea engelmannii 24 V . . . . I I . II Salix barrattiana 3 V . . I . I . . . Pohlia wahlenbergii 3 V V . . I . . . . Carex aquatilis 2 V V . . . . . . .             Senecio triangularis 85 . V III IV IV V I . I Carex nigricans 52 . V . IV IV I II I I Salix barclayi 8 . V . . . I I . I Pohlia wahlenbergii 3 V V . . I . . . . Carex aquatilis 2 V V . . . . . . . Carex lenticularis 1 . V . . . . . . . Aster species 1 . V . . . . . . . Carex canescens 1 . V . . . . . . . Viola lanceolata 1 . V . . . . . . .             Valeriana sitchensis 98 . . III I II V II II III Senecio triangularis 85 . V III IV IV V I . I Phleum alpinum 38 . . III . II II . III II Juncus drummondii 24 . . V II III I I . .  179  Group No.  1 2 3 4 5 6 7 8 9 No. of  1 1 2 9 19 53 46 7 52 Epilobium species 23 . . V . III I I . I Poa alpina 20 . . III . . I I I I Parnassia fimbriata 15 . . III I III I . . I Philonotis fontana 14 . . V . III I I . . Equisetum arvense 14 . . III I II I I . . Carex pyrenaica 8 . . III . I . I . I Agrostis humilis 8 . . III I I I I . . Campylium stellatum 6 . . III . I . I . I Salix species 6 . . III . I . . . I Saxifraga lyallii 6 . . III . I I I . . Mimulus lewisii 6 . . III I I I . . . Brachythecium frigidum 4 . . III . I I . . I Arnica angustifolia 2 . . III . . . I . . Carex lenticularis 2 . . III . I . . . . Petasites frigidus 1 . . III . . . . . . Drepanocladus aduncus 1 . . III . . . . . . Palustriella species 1 . . III . . . . . . Agrostis exarata 1 . . III . . . . . .             Senecio triangularis 85 . V III IV IV V I . I Veratrum viride 77 . . . III II IV II I II Erigeron peregrinus 69 . . . III II IV I IV II Carex nigricans 52 . V . IV IV I II I I Leptarrhena pyrolifolia 23 . . . III III I I . I Caltha leptosepala 23 . . . V III I I . . Aulacomnium palustre 20 . . . III II I I . I Eriophorum angustifolium 11 . . . IV I I I . .             Carex spectabilis 107 . . . II IV IV II IV IV Vahlodea atropurpurea 93 . . . . IV III III III III Cassiope mertensiana 90 . . . I III I V II IV Senecio triangularis 85 . V III IV IV V I . I Luetkea pectinata 65 . . . II III I V . II Carex nigricans 52 . V . IV IV I II I I Juncus drummondii 24 . . V II III I I . . Epilobium species 23 . . V . III I I . I Leptarrhena pyrolifolia 23 . . . III III I I . I Caltha leptosepala 23 . . . V III I I . . Juncus mertensianus 17 . . . I III I I . .  180  Group No.  1 2 3 4 5 6 7 8 9 No. of  1 1 2 9 19 53 46 7 52 Parnassia fimbriata 15 . . III I III I . . I Philonotis fontana 14 . . V . III I I . .             Carex spectabilis 107 . . . II IV IV II IV IV Valeriana sitchensis 98 . . III I II V II II III Vahlodea atropurpurea 93 . . . . IV III III III III Senecio triangularis 85 . V III IV IV V I . I Arnica latifolia 80 . . . . I III III V III Veratrum viride 77 . . . III II IV II I II Anemone occidentalis 69 . . . I II III II II II Erigeron peregrinus 69 . . . III II IV I IV II             Vahlodea atropurpurea 93 . . . . IV III III III III Cassiope mertensiana 90 . . . I III I V II IV Arnica latifolia 80 . . . . I III III V III Phyllodoce empetriformis 76 . . . I I I IV I IV Abies lasiocarpa 75 . . . II I I III . IV Luetkea pectinata 65 . . . II III I V . II Barbilophozia floerkei 31 . . . I I I III I I             Carex spectabilis 107 . . . II IV IV II IV IV Vahlodea atropurpurea 93 . . . . IV III III III III Arnica latifolia 80 . . . . I III III V III Erigeron peregrinus 69 . . . III II IV I IV II Artemisia norvegica 46 . . . II I II I IV III Hieracium gracile 43 . . . . I I II III II Phleum alpinum 38 . . III . II II . III II Antennaria lanata 37 . . . . II I II III I Dicranum species 28 . . . . . I II III I Polytrichum piliferum 19 . . . . . I I III I Polytrichum juniperinum 17 . . . . I I I IV I Coelocaulon species 8 . . . . . . I III I             Carex spectabilis 107 . . . II IV IV II IV IV Valeriana sitchensis 98 . . III I II V II II III Vahlodea atropurpurea 93 . . . . IV III III III III Cassiope mertensiana 90 . . . I III I V II IV Arnica latifolia 80 . . . . I III III V III Phyllodoce empetriformis 76 . . . I I I IV I IV  181  Group No.  1 2 3 4 5 6 7 8 9 No. of  1 1 2 9 19 53 46 7 52 Abies lasiocarpa 75 . . . II I I III . IV Artemisia norvegica 46 . . . II I II I IV III   Table 37 ESSFwcp Pseudospecies cut levels = 0 5 10 20 50 Presence/Absence          Group No.  1 2 3 4 5 6 7 8 9        No. of  1 2 9 7 10 56 11 27 67 Picea engelmannii 24 V . . . . I II I II Pohlia wahlenbergii 3 V III . . I . . . . Salix barrattiana 3 V . . . I I . . . Carex aquatilis 2 V III . . . . . . .             Senecio triangularis 85 . III IV IV IV V . II I Carex nigricans 52 . V III III IV I I II II Caltha leptosepala 23 . III I IV V I . . I Eriophorum angustifolium 11 . III . II III I . . I Polytrichum commune 8 . III . . I I I . I Salix barclayi 8 . III . . . I . . I Pohlia wahlenbergii 3 V III . . I . . . . Carex aquatilis 2 V III . . . . . . . Aster species 1 . III . . . . . . . Carex canescens 1 . III . . . . . . . Eriophorum species 1 . III . . . . . . . Viola lanceolata 1 . III . . . . . . . Carex lenticularis 1 . III . . . . . . .             Vahlodea atropurpurea 93 . . III V I III . V III Cassiope mertensiana 90 . . IV I I I I IV V Senecio triangularis 85 . III IV IV IV V . II I Luetkea pectinata 65 . . III II II I . II IV Carex nigricans 52 . V III III IV I I II II Juncus drummondii 24 . . IV II II I . I I Leptarrhena pyrolifolia 23 . . IV III III I . I I  182  Epilobium species 23 . . III IV . I . I I Juncus mertensianus 17 . . IV III I I . I . Parnassia fimbriata 15 . . III IV I I . I . Equisetum arvense 14 . . III III I I . . I Philonotis fontana 14 . . IV IV I . . I .             Carex spectabilis 107 . . II III II IV IV III III Vahlodea atropurpurea 93 . . III V I III . V III Senecio triangularis 85 . III IV IV IV V . II I Veratrum viride 77 . . . III III IV . II II Carex nigricans 52 . V III III IV I I II II Leptarrhena pyrolifolia 23 . . IV III III I . I I Epilobium species 23 . . III IV . I . I I Caltha leptosepala 23 . III I IV V I . . I Mitella species 18 . . I III . I . I I Juncus mertensianus 17 . . IV III I I . I . Parnassia fimbriata 15 . . III IV I I . I . Equisetum arvense 14 . . III III I I . . I Philonotis fontana 14 . . IV IV I . . I . Rhytidiadelphus squarrosus 9 . . I III I I . . I Petasites frigidus 7 . . II III . . . . I             Senecio triangularis 85 . III IV IV IV V . II I Veratrum viride 77 . . . III III IV . II II Carex nigricans 52 . V III III IV I I II II Leptarrhena pyrolifolia 23 . . IV III III I . I I Caltha leptosepala 23 . III I IV V I . . I Aulacomnium palustre 20 . . II II III I . . I Eriophorum angustifolium 11 . III . II III I . . I             Carex spectabilis 107 . . II III II IV IV III III Valeriana sitchensis 98 . . II I I V I III II Vahlodea atropurpurea 93 . . III V I III . V III Senecio triangularis 85 . III IV IV IV V . II I Arnica latifolia 80 . . . II . III III IV III Veratrum viride 77 . . . III III IV . II II Anemone occidentalis 69 . . II II . III II III II Erigeron peregrinus 69 . . . I II IV I III I Pedicularis bracteosa 32 . . . I . III . II I             Carex spectabilis 107 . . II III II IV IV III III Group No.  1 2 3 4 5 6 7 8 9 No. of  1 2 9 7 10 56 11 27 67  183  Arnica latifolia 80 . . . II . III III IV III Phyllodoce empetriformis 76 . . II . . I III III IV Abies lasiocarpa 75 . . I II I I IV II IV Artemisia norvegica 46 . . . . I II IV IV I Sibbaldia procumbens 36 . . I . . I III III I Bryum species 13 . . . I . . IV I I             Carex spectabilis 107 . . II III II IV IV III III Valeriana sitchensis 98 . . II I I V I III II Vahlodea atropurpurea 93 . . III V I III . V III Cassiope mertensiana 90 . . IV I I I I IV V Arnica latifolia 80 . . . II . III III IV III Phyllodoce empetriformis 76 . . II . . I III III IV Anemone occidentalis 69 . . II II . III II III II Erigeron peregrinus 69 . . . I II IV I III I Artemisia norvegica 46 . . . . I II IV IV I Hieracium gracile 43 . . . I . I . IV II Phleum alpinum 38 . . II I . II . III I Antennaria lanata 37 . . II I . I . III I Sibbaldia procumbens 36 . . I . . I III III I Veronica wormskjoldii 22 . . II . . I . III I Poa cusickii 15 . . . . . I I III .             Carex spectabilis 107 . . II III II IV IV III III Vahlodea atropurpurea 93 . . III V I III . V III Cassiope mertensiana 90 . . IV I I I I IV V Arnica latifolia 80 . . . II . III III IV III Phyllodoce empetriformis 76 . . II . . I III III IV Abies lasiocarpa 75 . . I II I I IV II IV Luetkea pectinata 65 . . III II II I . II IV  Table 38 ESSFwcp Isopam. Group No. 1 2 3 4 5 6 7 8 9 10 No. of 29 22 20 8 7 28 8 9 33 26             Abies lasiocarpa IV V III II II I II III II I Phyllodoce empetriformis IV III IV V I I I III II I Cassiope mertensiana V III V V I II IV V I I Vaccinium membranaceum III I II I I . . II I I Luetkea pectinata IV I IV IV . II III V I I             Group No.  1 2 3 4 5 6 7 8 9 No. of  1 2 9 7 10 56 11 27 67  184  Group No. 1 2 3 4 5 6 7 8 9 10 No. of 29 22 20 8 7 28 8 9 33 26 Abies lasiocarpa IV V III II II I II III II I Phyllodoce empetriformis IV III IV V I I I III II I Valeriana sitchensis II III IV V . I II II IV V Vahlodea atropurpurea II III IV II II II IV V IV II Cassiope mertensiana V III V V I II IV V I I Carex spectabilis I V IV III IV III III III IV IV Arnica latifolia I III IV IV IV I I IV IV II Barbilophozia species I III . II . . . . I . Picea engelmannii I III I . . I . I I .             Abies lasiocarpa IV V III II II I II III II I Phyllodoce empetriformis IV III IV V I I I III II I Valeriana sitchensis II III IV V . I II II IV V Vahlodea atropurpurea II III IV II II II IV V IV II Cassiope mertensiana V III V V I II IV V I I Carex spectabilis I V IV III IV III III III IV IV Arnica latifolia I III IV IV IV I I IV IV II Luzula parviflora I II III II . I . III II II Anemone occidentalis I I IV V II II . IV IV I Luetkea pectinata IV I IV IV . II III V I I Polytrichastrum alpinum . . III . . I . I I . Antennaria lanata I I III . I I II III II . Hieracium gracile I II III III II I I III II . Sibbaldia procumbens I II III III III I . II II . Barbilophozia floerkei II I III . . I . III . .             Phyllodoce empetriformis IV III IV V I I I III II I Valeriana sitchensis II III IV V . I II II IV V Cassiope mertensiana V III V V I II IV V I I Senecio triangularis I I II IV . III V III IV V Veratrum viride I I II III . II II III III V Carex spectabilis I V IV III IV III III III IV IV Erigeron peregrinus I I II IV III I II II V III Arnica latifolia I III IV IV IV I I IV IV II Anemone occidentalis I I IV V II II . IV IV I Artemisia norvegica I I II IV IV I I . III I Claytonia lanceolata I I . III . . . . II I Dicranum species I I II III II I . II I I Luetkea pectinata IV I IV IV . II III V I I  185  Group No. 1 2 3 4 5 6 7 8 9 10 No. of 29 22 20 8 7 28 8 9 33 26 Hieracium gracile I II III III II I I III II . Veronica wormskjoldii . . I III . I II I II . Sibbaldia procumbens I II III III III I . II II . Lycopodium species . I . IV II . . . I .             Carex spectabilis I V IV III IV III III III IV IV Erigeron peregrinus I I II IV III I II II V III Arnica latifolia I III IV IV IV I I IV IV II Epilobium anagallidifolium . I I . III I I . II I Artemisia norvegica I I II IV IV I I . III I Flavocetraria nivalis . . . . III . . . . . Coelocaulon species I . . II III . . . . . Stereocaulon alpinum I . . . III . . . . . Polytrichum juniperinum I . I I III I I . I . Sibbaldia procumbens I II III III III I . II II .             Senecio triangularis I I II IV . III V III IV V Carex spectabilis I V IV III IV III III III IV IV Carex nigricans I I I . . V IV IV II .             Leptarrhena pyrolifolia . . . I . II V II I I Vahlodea atropurpurea II III IV II II II IV V IV II Cassiope mertensiana V III V V I II IV V I I Senecio triangularis I I II IV . III V III IV V Carex spectabilis I V IV III IV III III III IV IV Equisetum arvense . . . . . . IV I I I Aulacomnium palustre I . I . . II IV . I I Juncus mertensianus . . I . . I IV II I I Juncus drummondii I . I . . II IV II I I Parnassia fimbriata . . . II . I IV II I I Caltha leptosepala I . . . . II III II I I Petasites frigidus I . . . . . III I . I Luetkea pectinata IV I IV IV . II III V I I Philonotis fontana . . . . . I V . I I Carex nigricans I I I . . V IV IV II .             Abies lasiocarpa IV V III II II I II III II I Phyllodoce empetriformis IV III IV V I I I III II I Vahlodea atropurpurea II III IV II II II IV V IV II  186  Group No. 1 2 3 4 5 6 7 8 9 10 No. of 29 22 20 8 7 28 8 9 33 26 Cassiope mertensiana V III V V I II IV V I I Senecio triangularis I I II IV . III V III IV V Veratrum viride I I II III . II II III III V Carex spectabilis I V IV III IV III III III IV IV Arnica latifolia I III IV IV IV I I IV IV II Luzula parviflora I II III II . I . III II II Anemone occidentalis I I IV V II II . IV IV I Luetkea pectinata IV I IV IV . II III V I I Antennaria lanata I I III . I I II III II . Carex nigricans I I I . . V IV IV II . Hieracium gracile I II III III II I I III II . Barbilophozia floerkei II I III . . I . III . .             Valeriana sitchensis II III IV V . I II II IV V Vahlodea atropurpurea II III IV II II II IV V IV II Senecio triangularis I I II IV . III V III IV V Veratrum viride I I II III . II II III III V Carex spectabilis I V IV III IV III III III IV IV Erigeron peregrinus I I II IV III I II II V III Pedicularis bracteosa I . I II I . I . III II Arnica latifolia I III IV IV IV I I IV IV II Anemone occidentalis I I IV V II II . IV IV I Phleum alpinum . I II II I I I II III I Artemisia norvegica I I II IV IV I I . III I             Valeriana sitchensis II III IV V . I II II IV V Senecio triangularis I I II IV . III V III IV V Veratrum viride I I II III . II II III III V Carex spectabilis I V IV III IV III III III IV IV Heracleum maximum . . . . . . . . I III Epilobium angustifolium I . I II I . . . I III Thalictrum occidentale . . I . . . . . I III Erigeron peregrinus I I II IV III I II II V III    187  IMAwc Table 39 All plots (190) TWINSPAN Sørensen’s  0 5 10 30 50. Group No. absfreq 1 2 3 4 5 6 7 8 9 Cassiope mertensiana 55 V V IV IV IV III . II II Kiaeria blyttii 2 V . . . . . . . . Luetkea pectinata 32 V III V I IV II . . I Diphasiastrum alpinum 8 III II . . I I . I . Antennaria species 3 III II . . . I . . .             Cassiope mertensiana 55 V V IV IV IV III . II II Luetkea pectinata 32 V III V I IV II . . I Carex spectabilis 45 . III . II III IV II I II Phyllodoce empetriformis 33 . IV . III II III . I II Carex nigricans 26 . V V V II II . . I             Cassiope mertensiana 55 V V IV IV IV III . II II Luetkea pectinata 32 V III V I IV II . . I Hieracium gracile 39 . II IV II III IV . I I Polytrichum juniperinum 28 . . IV I II I . I II Carex nigricans 26 . V V V II II . . I Poa alpina 21 . . IV . II I II I II Dicranum scoparium 15 . . V . II I . I I Luzula parviflora 10 . II IV I I . . I I Marsupella species 5 . II IV . . I . . I Racomitrium sudeticum 3 . . IV . I . . . . Juncus drummondii 11 . . IV I II I . . .             Cassiope mertensiana 55 V V IV IV IV III . II II Phyllodoce empetriformis 33 . IV . III II III . I II Carex nigricans 26 . V V V II II . . I Vahlodea atropurpurea 24 . II . III I III . I I             Cassiope mertensiana 55 V V IV IV IV III . II II Luetkea pectinata 32 V III V I IV II . . I Sibbaldia procumbens 52 . . . . III II . III IV Antennaria lanata 51 . II II II IV IV . II II Carex spectabilis 45 . III . II III IV II I II Hieracium gracile 39 . II IV II III IV . I I Phyllodoce glanduliflora 36 . . II I III I . II III Luzula piperi 26 . . II II IV II . I I  188  Group No. absfreq 1 2 3 4 5 6 7 8 9             Cassiope mertensiana 55 V V IV IV IV III . II II Artemisia norvegica 62 . . . . II III II IV V Antennaria lanata 51 . II II II IV IV . II II Carex spectabilis 45 . III . II III IV II I II Hieracium gracile 39 . II IV II III IV . I I Phyllodoce empetriformis 33 . IV . III II III . I II Erigeron peregrinus 28 . . . . II IV II I I Vahlodea atropurpurea 24 . II . III I III . I I Phleum alpinum 21 . II . . II III . I I Arnica latifolia 19 . . . . I IV . . I Anemone occidentalis 13 . II II II . III . . . Valeriana sitchensis 11 . II . . . III . . I             Silene acaulis 37 . . . . . I IV IV IV Abies lasiocarpa 21 . II . . I II IV III I Saxifraga bronchialis 10 . . . I . . IV I I Salix arctica 2 . . . . . . IV . . Epilobium latifolium 2 . . . . . . IV . .                                       Artemisia norvegica 62 . . . . II III II IV V Sibbaldia procumbens 52 . . . . III II . III IV Silene acaulis 37 . . . . . I IV IV IV Polytrichum piliferum 32 . . II . II I . III II Antennaria alpina 31 . II . . I I . III III Festuca brachyphylla 30 . . . I I I . III III Luzula spicata 29 . II . . . I . III III Salix nivalis 29 . . . . I I . V II Abies lasiocarpa 21 . II . . I II IV III I Flavocetraria nivalis 16 . . . . . . . IV I Dryas octopetala 13 . . . . I . II IV I Umbilicaria species 9 . . . . I . . III I                          Artemisia norvegica 62 . . . . II III II IV V Sibbaldia procumbens 52 . . . . III II . III IV Silene acaulis 37 . . . . . I IV IV IV Phyllodoce glanduliflora 36 . . II I III I . II III  189  Group No. absfreq 1 2 3 4 5 6 7 8 9 Lepraria neglecta 32 . . . . I I . . IV Antennaria alpina 31 . II . . I I . III III Festuca brachyphylla 30 . . . I I I . III III Luzula spicata 29 . II . . . I . III III Cassiope tetragona 24 . . . . . I . II III Empetrum nigrum 21 . . . . . I . II III    Table 40 TWINSPAN fidelity (>18)     Group No.  1 2 3 4 5 6 7 8 9 No. of Freq. 2 4 3 5 20 22 3 16 35                         Cassiope mertensiana 55 18.3 18.3 8.1 11.7 11.7 6.4 --- 2.6 1 Kiaeria blyttii 2 100 --- --- --- --- --- --- --- --- Luetkea pectinata 32 26.3 6.6 26.3 1.1 12.9 2.7 --- --- 0.2 Antennaria species 3 31.4 7.9 --- --- --- 0.3 --- --- --- Diphasiastrum alpinum 8 23.6 5.9 --- --- 3.8 0.2 --- 0.4 ---                                     Cassiope mertensiana 55 18.3 18.3 8.1 11.7 11.7 6.4 --- 2.6 1 Phyllodoce empetriformis 33 --- 22.4 --- 14.4 4.9 8.2 --- 0.6 2.1 Carex nigricans 26 --- 27.4 27.4 27.4 2.5 2.8 --- --- --- Dicranum undulatum 2 --- 21.2 --- --- --- 0.7 --- --- --- Kobresia myosuroides 2 --- 21.2 --- --- --- 0.7 --- --- --- Luzula species 2 --- 22.4 --- --- --- --- --- --- 0.3 Dicranum fuscescens 1 --- 25 --- --- --- --- --- --- ---             Luetkea pectinata 32 26.3 6.6 26.3 1.1 12.9 2.7 --- --- 0.2 Polytrichum juniperinum 28 --- --- 24.4 2.2 3.4 1.8 --- 0.9 8.8 Carex nigricans 26 --- 27.4 27.4 27.4 2.5 2.8 --- --- --- Poa alpina 21 --- --- 26.1 --- 3.7 1.1 6.5 0.2 3.9 Dicranum scoparium 15 --- --- 63.8 --- 4 1.2 --- 1 0.2 Luzula parviflora 10 --- 4.4 31.6 2.8 2.8 --- --- 0.3 0.1  190  Diphasiastrum sitchense 6 --- --- 18.4 --- 1.7 0.3 --- 2.6 --- Marsupella species 5 --- 6.3 44.9 --- --- 0.2 --- --- 0.1 Anemone multifida 5 --- --- 21.2 --- 1.9 1.6 --- --- --- Racomitrium canescens 3 --- --- 26.2 --- --- --- --- 0.9 0.2 Racomitrium sudeticum 3 --- --- 62 --- 0.3 --- --- --- --- Juncus drummondii 11 --- --- 35.5 3.2 5 1.5 --- --- --- Schistidium apocarpum 2 --- --- 30.7 --- --- --- --- --- 0.2             Carex nigricans 26 --- 27.4 27.4 27.4 2.5 2.8 --- --- --- Vahlodea atropurpurea 24 --- 3.5 --- 20.1 2.2 13.9 --- 2 0.2 Luzula arcuata 14 --- --- --- 20.2 --- 2.3 --- --- 8.3 Trichophorum cespitosum 3 --- --- --- 37.3 --- --- --- --- 0.2 Dicranella palustris 1 --- --- --- 20 --- --- --- --- --- Carex praegracilis 1 --- --- --- 20 --- --- --- --- --- Barbilophozia lycopodioides 1 --- --- --- 20 --- --- --- --- --- Cladonia deformis 1 --- --- --- 20 --- --- --- --- ---             Phyllodoce glanduliflora 36 --- --- 5.7 2.1 18.5 1 --- 3.2 9.4 Luzula piperi 26 --- --- 6.1 8.8 23.2 2.8 --- 0.9 0.4 Carex pyrenaica 17 --- --- --- --- 19.8 0.3 --- 7.7 1.6 Kiaeria species 7 --- --- --- --- 27.4 --- --- --- 0.2             Antennaria lanata 51 --- 2 3.6 5.2 13.8 19.5 --- 3.2 3.8 Carex spectabilis 45 --- 8.5 --- 5.5 10.3 20.4 3.8 0.1 3.4 Erigeron peregrinus 28 --- --- --- --- 7.9 26 7.1 1 0.8 Phleum alpinum 21 --- 5.4 --- --- 5.4 21.8 --- 0.3 0.6 Arnica latifolia 19 --- --- --- --- 2.7 48 --- --- 0.4 Claytonia lanceolata 9 --- --- --- --- --- 27 --- --- 0.9 Valeriana sitchensis 11 --- 9.1 --- --- --- 24.3 --- --- 0.1 Senecio triangularis 10 --- 9.4 --- --- 0.4 19.9 --- --- --- Caltha leptosepala 7 --- --- --- --- --- 31.8 --- --- ---             Silene acaulis 37 --- --- --- --- --- 0.1 22 19.3 23.2 Abies lasiocarpa 21 --- 3.5 --- --- 0.1 2.9 25 10.8 1.2 Saxifraga bronchialis 10 --- --- --- 3.4 --- --- 38 3 1.1 Epilobium anagallidifolium 3 --- --- --- 6.9 --- 0.4 19.2 --- --- Minuartia obtusiloba 8 --- --- --- --- --- --- 18.5 2.6 3.4 Brachythecium species 7 --- --- --- --- 0.4 3.2 19.3 --- 0.6 Poa arctica 5 --- --- --- --- --- 3.5 20.9 0.7 --- Selaginella species 5 --- --- --- --- --- --- 24.8 --- 2.9 Picea species 4 --- --- --- --- --- 0.5 25.5 --- 0.7 Group No.  1 2 3 4 5 6 7 8 9 No. of Freq. 2 4 3 5 20 22 3 16 35  191  Oxyria digyna 3 --- --- --- --- 0.6 --- 27 --- 0.2 Lupinus arcticus 3 --- --- --- --- --- 1.9 26.2 --- --- Salix species 2 --- 10.7 --- --- --- --- 19 --- --- Salix arctica 2 --- --- --- --- --- --- 66.7 --- --- Epilobium latifolium 2 --- --- --- --- --- --- 66.7 --- --- Pinus albicaulis 2 --- --- --- --- --- --- 28.1 1 --- Bromus vulgaris 1 --- --- --- --- --- --- 33.3 --- --- Stellaria longipes 1 --- --- --- --- --- --- 33.3 --- --- Elymus species 1 --- --- --- --- --- --- 33.3 --- --- Rhinanthus minor 1 --- --- --- --- --- --- 33.3 --- --- Arnica cordifolia 1 --- --- --- --- --- --- 33.3 --- --- Antennaria racemosa 1 --- --- --- --- --- --- 33.3 --- --- Anemone parviflora 1 --- --- --- --- --- --- 33.3 --- --- Silene parryi 1 --- --- --- --- --- --- 33.3 --- --- Achillea millefolium 1 --- --- --- --- --- --- 33.3 --- ---             Artemisia norvegica 62 --- --- --- --- 3.2 7.4 4 20.2 32 Silene acaulis 37 --- --- --- --- --- 0.1 22 19.3 23.2 Polytrichum piliferum 32 --- --- 6.4 --- 7 1.9 --- 18.2 5.7 Antennaria alpina 31 --- 4.2 --- --- 0.7 1.2 --- 21 13.9 Festuca brachyphylla 30 --- --- --- 2.9 0.2 1.4 --- 18.2 17.2 Luzula spicata 29 --- 4.6 --- --- --- 1.4 --- 18.2 17.2 Salix nivalis 29 --- --- --- --- 0.8 0.2 --- 49.7 10.4 Flavocetraria nivalis 16 --- --- --- --- --- --- --- 49 3.7 Dryas octopetala 13 --- --- --- --- 0.2 --- 10.7 37.7 0.1 Umbilicaria species 9 --- --- --- --- 0.5 --- --- 37.1 0.2 Solidago multiradiata 10 --- --- --- --- --- --- --- 28.7 2.7 Thamnolia vermicularis 6 --- --- --- --- --- --- --- 37.5 --- Alectoria ochroleuca 4 --- --- --- --- --- --- --- 25 ---             Artemisia norvegica 62 --- --- --- --- 3.2 7.4 4 20.2 32 Sibbaldia procumbens 52 --- --- --- --- 12.1 6.4 --- 9.2 28.7 Silene acaulis 37 --- --- --- --- --- 0.1 22 19.3 23.2 Lepraria neglecta 32 --- --- --- --- 0.2 3.3 --- --- 59.3 Cassiope tetragona 24 --- --- --- --- --- 0.2 --- 15.5 26 Empetrum nigrum 21 --- --- --- --- --- 0.3 --- 8.3 27.8 Potentilla diversifolia 11 --- --- --- --- --- --- --- 1.1 23.4 Campanula lasiocarpa 11 --- --- --- --- --- --- --- 1.1 23.4 Cornicularia aculeate 17 --- --- --- --- --- 1.4 --- 2.7 23.5 Stellaria longifolia 8 --- --- --- --- --- --- --- --- 22.9  Group No.  1 2 3 4 5 6 7 8 9 No. of Freq. 2 4 3 5 20 22 3 16 35  192  Table 41 TWINSPAN (Sørensen’s index, pseudospecies cut levels 0 5 10 20 50). No. of  4 12 19 21 2 1 13 15 23 Group No.  1 2 3 4 5 6 7 8 9 Cassiope mertensiana 55 IV V IV IV . . II III I Kiaeria blyttii 2 III . . . . . . . . Luetkea pectinata 32 IV III IV II . . . I . Carex spectabilis 45 III II III V III . I IV I Hieracium gracile 39 III III IV III . . I III I Antennaria alpina 31 III . I I . . IV III II Carex nigricans 26 III V II II . . . I . Stereocaulon alpinum 15 III . . . . . I I II Dicranum undulatum 2 III . . . . . . . .                         Cassiope mertensiana 55 IV V IV IV . . II III I Luetkea pectinata 32 IV III IV II . . . I . Antennaria lanata 51 . III IV IV . . II III II Hieracium gracile 39 III III IV III . . I III I Phyllodoce empetriformis 33 . III II III . . I II I Carex nigricans 26 III V II II . . . I . Juncus drummondii 11 . III II I . . . . .                         Cassiope mertensiana 55 IV V IV IV . . II III I Luetkea pectinata 32 IV III IV II . . . I . Sibbaldia procumbens 52 . . III II . . II V IV Antennaria lanata 51 . III IV IV . . II III II Carex spectabilis 45 III II III V III . I IV I Hieracium gracile 39 III III IV III . . I III I Phyllodoce glanduliflora 36 . I IV I . . II I III Polytrichum piliferum 32 . I III I . . III II II Erigeron peregrinus 28 II . III III III . I II I Luzula piperi 26 . II IV I . . I II . Polytrichastrum alpinum 18 II I III I . . I I . Carex pyrenaica 17 . . III . . . I II I Cladonia species 16 . I III I . . I I I                         Cassiope mertensiana 55 IV V IV IV . . II III I Artemisia norvegica 62 . . II III III . IV V V Antennaria lanata 51 . III IV IV . . II III II Carex spectabilis 45 III II III V III . I IV I  193  No. of  4 12 19 21 2 1 13 15 23 Hieracium gracile 39 III III IV III . . I III I Phyllodoce empetriformis 33 . III II III . . I II I Erigeron peregrinus 28 II . III III III . I II I Vahlodea atropurpurea 24 . II II III . . I I I Phleum alpinum 21 II . II III . . I I I Arnica latifolia 19 II I II III . . . I I Anemone occidentalis 13 . II . III . . . . . Senecio triangularis 10 . I . III . . . . .                         Artemisia norvegica 62 . . II III III . IV V V Carex spectabilis 45 III II III V III . I IV I Silene acaulis 37 . . . I III V IV III IV Erigeron peregrinus 28 II . III III III . I II I Abies lasiocarpa 21 . . . II V . II I I Saxifraga bronchialis 10 . I . . III V II . I Potentilla villosa 10 . . . . III . II . II Brachythecium species 7 . I I I III . I I . Poa arctica 5 . . . I III . . I . Picea species 4 . . . I III . I . I Oxyria digyna 3 . . I . III . . I . Lupinus arcticus 3 . . . I III . . . . Salix species 2 . . . I III . . . . Salix arctica 2 . . . . III V . . . Epilobium latifolium 2 . . . . V . . . . Pinus albicaulis 2 . . . . III . I . . Rhinanthus minor 1 . . . . III . . . . Arnica cordifolia 1 . . . . III . . . . Antennaria racemosa 1 . . . . III . . . . Anemone parviflora 1 . . . . III . . . . Silene parryi 1 . . . . III . . . . Achillea millefolium 1 . . . . III . . . .                         Silene acaulis 37 . . . I III V IV III IV Poa alpina 21 . II II I . V I I II Trisetum spicatum 20 . I I I . V I III I Dryas octopetala 13 . . . I . V V . . Saxifraga bronchialis 10 . I . . III V II . I Epilobium anagallidifolium 3 . I . I . V . . .  194  No. of  4 12 19 21 2 1 13 15 23 Minuartia obtusiloba 8 . . . . . V I I I Selaginella species 5 . . . . . V . I I Salix arctica 2 . . . . III V . . . Bromus vulgaris 1 . . . . . V . . . Stellaria longipes 1 . . . . . V . . . Elymus species 1 . . . . . V . . .                         Artemisia norvegica 62 . . II III III . IV V V Silene acaulis 37 . . . I III V IV III IV Polytrichum piliferum 32 . I III I . . III II II Antennaria alpina 31 III . I I . . IV III II Festuca brachyphylla 30 . I I I . . IV III III Luzula spicata 29 . I . I . . III III III Salix nivalis 29 . . I I . . V III II Cassiope tetragona 24 . . . I . . III II III Flavocetraria nivalis 16 . . . . . . IV . II Dryas octopetala 13 . . . I . V V . . Thamnolia vermicularis 6 . . . . . . III . .                         Cassiope mertensiana 55 IV V IV IV . . II III I Artemisia norvegica 62 . . II III III . IV V V Sibbaldia procumbens 52 . . III II . . II V IV Antennaria lanata 51 . III IV IV . . II III II Carex spectabilis 45 III II III V III . I IV I Hieracium gracile 39 III III IV III . . I III I Silene acaulis 37 . . . I III V IV III IV Lepraria neglecta 32 . . I I . . . III V Antennaria alpina 31 III . I I . . IV III II Festuca brachyphylla 30 . I I I . . IV III III Luzula spicata 29 . I . I . . III III III Salix nivalis 29 . . I I . . V III II Solorina crocea 25 . I II I . . II III II Trisetum spicatum 20 . I I I . V I III I                         Artemisia norvegica 62 . . II III III . IV V V Sibbaldia procumbens 52 . . III II . . II V IV Silene acaulis 37 . . . I III V IV III IV  195  No. of  4 12 19 21 2 1 13 15 23 Phyllodoce glanduliflora 36 . I IV I . . II I III Lepraria neglecta 32 . . I I . . . III V Festuca brachyphylla 30 . I I I . . IV III III Luzula spicata 29 . I . I . . III III III Polytrichum juniperinum 28 . II II I . . I I III Cassiope tetragona 24 . . . I . . III II III Empetrum nigrum 21 . . . I . . II II III Cornicularia aculeate 17 . . . I . . II I III             Group No.  1 2 3 4 5 6 7 8 9 No. of Freq. 14 20 5 17 3 16 9 13 13                       Table 42 TWINSPAN pseudospecies cut levels 0 5 10 20 50 Presence/Absence. Group No.  1 2 3 4 5 6 7 8 9 No. of relevé 4 12 19 21 2 1 13 15 23                      Cassiope mertensiana 55 IV V IV IV . . II III I Kiaeria blyttii 2 III . . . . . . . . Luetkea pectinata 32 IV III IV II . . . I . Carex spectabilis 45 III II III V III . I IV I Hieracium gracile 39 III III IV III . . I III I Antennaria alpina 31 III . I I . . IV III II Carex nigricans 26 III V II II . . . I . Stereocaulon alpinum 15 III . . . . . I I II Dicranum undulatum 2 III . . . . . . . .             Cassiope mertensiana 55 IV V IV IV . . II III I Luetkea pectinata 32 IV III IV II . . . I . Antennaria lanata 51 . III IV IV . . II III II Hieracium gracile 39 III III IV III . . I III I Phyllodoce empetriformis 33 . III II III . . I II I Carex nigricans 26 III V II II . . . I . Juncus drummondii 11 . III II I . . . . .             Cassiope mertensiana 55 IV V IV IV . . II III I Luetkea pectinata 32 IV III IV II . . . I . Sibbaldia procumbens 52 . . III II . . II V IV Antennaria lanata 51 . III IV IV . . II III II  196  Group No.  1 2 3 4 5 6 7 8 9 No. of relevé 4 12 19 21 2 1 13 15 23 Carex spectabilis 45 III II III V III . I IV I Hieracium gracile 39 III III IV III . . I III I Phyllodoce glanduliflora 36 . I IV I . . II I III Polytrichum piliferum 32 . I III I . . III II II Erigeron peregrinus 28 II . III III III . I II I Luzula piperi 26 . II IV I . . I II . Polytrichastrum alpinum 18 II I III I . . I I . Carex pyrenaica 17 . . III . . . I II I Cladonia species 16 . I III I . . I I I             Cassiope mertensiana 55 IV V IV IV . . II III I Artemisia norvegica 62 . . II III III . IV V V Antennaria lanata 51 . III IV IV . . II III II Carex spectabilis 45 III II III V III . I IV I Hieracium gracile 39 III III IV III . . I III I Phyllodoce empetriformis 33 . III II III . . I II I Erigeron peregrinus 28 II . III III III . I II I Vahlodea atropurpurea 24 . II II III . . I I I Phleum alpinum 21 II . II III . . I I I Arnica latifolia 19 II I II III . . . I I Anemone occidentalis 13 . II . III . . . . . Senecio triangularis 10 . I . III . . . . .             Artemisia norvegica 62 . . II III III . IV V V Carex spectabilis 45 III II III V III . I IV I Silene acaulis 37 . . . I III V IV III IV Erigeron peregrinus 28 II . III III III . I II I Abies lasiocarpa 21 . . . II V . II I I Saxifraga bronchialis 10 . I . . III V II . I Potentilla villosa 10 . . . . III . II . II Brachythecium species 7 . I I I III . I I . Poa arctica 5 . . . I III . . I . Picea species 4 . . . I III . I . I Oxyria digyna 3 . . I . III . . I . Lupinus arcticus 3 . . . I III . . . . Salix species 2 . . . I III . . . . Salix arctica 2 . . . . III V . . . Epilobium latifolium 2 . . . . V . . . . Pinus albicaulis 2 . . . . III . I . .  197  Group No.  1 2 3 4 5 6 7 8 9 No. of relevé 4 12 19 21 2 1 13 15 23 Rhinanthus minor 1 . . . . III . . . . Arnica cordifolia 1 . . . . III . . . . Antennaria racemosa 1 . . . . III . . . . Anemone parviflora 1 . . . . III . . . . Silene parryi 1 . . . . III . . . . Achillea millefolium 1 . . . . III . . . .             Silene acaulis 37 . . . I III V IV III IV Poa alpina 21 . II II I . V I I II Trisetum spicatum 20 . I I I . V I III I Dryas octopetala 13 . . . I . V V . . Saxifraga bronchialis 10 . I . . III V II . I Epilobium anagallidifolium 3 . I . I . V . . . Minuartia obtusiloba 8 . . . . . V I I I Selaginella species 5 . . . . . V . I I Salix arctica 2 . . . . III V . . . Bromus vulgaris 1 . . . . . V . . . Stellaria longipes 1 . . . . . V . . . Elymus species 1 . . . . . V . . .             Artemisia norvegica 62 . . II III III . IV V V Silene acaulis 37 . . . I III V IV III IV Polytrichum piliferum 32 . I III I . . III II II Antennaria alpina 31 III . I I . . IV III II Festuca brachyphylla 30 . I I I . . IV III III Luzula spicata 29 . I . I . . III III III Salix nivalis 29 . . I I . . V III II Cassiope tetragona 24 . . . I . . III II III Flavocetraria nivalis 16 . . . . . . IV . II Dryas octopetala 13 . . . I . V V . . Thamnolia vermicularis 6 . . . . . . III . .             Cassiope mertensiana 55 IV V IV IV . . II III I Artemisia norvegica 62 . . II III III . IV V V Sibbaldia procumbens 52 . . III II . . II V IV Antennaria lanata 51 . III IV IV . . II III II Carex spectabilis 45 III II III V III . I IV I Hieracium gracile 39 III III IV III . . I III I Silene acaulis 37 . . . I III V IV III IV  198  Group No.  1 2 3 4 5 6 7 8 9 No. of relevé 4 12 19 21 2 1 13 15 23 Lepraria neglecta 32 . . I I . . . III V Antennaria alpina 31 III . I I . . IV III II Festuca brachyphylla 30 . I I I . . IV III III Luzula spicata 29 . I . I . . III III III Salix nivalis 29 . . I I . . V III II Solorina crocea 25 . I II I . . II III II Trisetum spicatum 20 . I I I . V I III I             Artemisia norvegica 62 . . II III III . IV V V Sibbaldia procumbens 52 . . III II . . II V IV Silene acaulis 37 . . . I III V IV III IV Phyllodoce glanduliflora 36 . I IV I . . II I III Lepraria neglecta 32 . . I I . . . III V Festuca brachyphylla 30 . I I I . . IV III III Luzula spicata 29 . I . I . . III III III Polytrichum juniperinum 28 . II II I . . I I III Cassiope tetragona 24 . . . I . . III II III Empetrum nigrum 21 . . . I . . II II III Cornicularia aculeate 17 . . . I . . II I III  Table 43 ISOPAM. Group No.  1 2 3 4 5 No. of  17 13 16 32 32 Artemisia norvegica 62 IV V V I IV Sibbaldia procumbens 52 III IV III I IV Silene acaulis 37 III V III . II Polytrichum piliferum 32 III II II I III Antennaria alpina 31 III IV I I I Festuca brachyphylla 30 III IV I I II Salix nivalis 29 V III I I I Luzula spicata 29 III III II I II Cassiope tetragona 24 III II IV . I Flavocetraria nivalis 16 IV I II . . Dryas octopetala 13 III I . I I Umbilicaria species 9 III . . . I         Artemisia norvegica 62 IV V V I IV Sibbaldia procumbens 52 III IV III I IV Silene acaulis 37 III V III . II  199  Group No.  1 2 3 4 5 No. of  17 13 16 32 32 Lepraria neglecta 32 . IV V I II Antennaria alpina 31 III IV I I I Festuca brachyphylla 30 III IV I I II Salix nivalis 29 V III I I I Luzula spicata 29 III III II I II Potentilla diversifolia 11 I III . . I         Artemisia norvegica 62 IV V V I IV Sibbaldia procumbens 52 III IV III I IV Silene acaulis 37 III V III . II Phyllodoce glanduliflora 36 I I IV II III Lepraria neglecta 32 . IV V I II Polytrichum juniperinum 28 II II III II I Cassiope tetragona 24 III II IV . I Empetrum nigrum 21 II II III . I         Cassiope mertensiana 55 II . I V III Antennaria lanata 51 I II I III V Carex spectabilis 45 I II I III IV Phyllodoce empetriformis 33 II II I III II Luetkea pectinata 32 I . I IV II Carex nigricans 26 . . . IV I         Artemisia norvegica 62 IV V V I IV Cassiope mertensiana 55 II . I V III Sibbaldia procumbens 52 III IV III I IV Antennaria lanata 51 I II I III V Carex spectabilis 45 I II I III IV Hieracium gracile 39 . I I II V Phyllodoce glanduliflora 36 I I IV II III Polytrichum piliferum 32 III II II I III Erigeron peregrinus 28 . I I I III Luzula piperi 26 I . . II III Vahlodea atropurpurea 24 I . I II III Phleum alpinum 21 . I . I III Arnica latifolia 19 . . . I III                           200   Table 44 My relevés (130) TWINSPAN w/merged groups. Group No.  2 4 3 7 1 6 5 No. of relevés 33 4 20 4 7 37 26           Artemisia norvegica 59 V . I . IV III I Coelocaulon species 19 III . . . I I I Empetrum nigrum 18 III . . . . I I Festuca brachyphylla 25 III . . . I I I Lepraria neglecta 39 V . I . . I I Sibbaldia procumbens 49 IV . I . III II I Silene acaulis 22 III . I . . I I           Arnica latifolia 57 II III III . IV III II Barbilophozia floerkei 11 . III I . . I II Cassiope mertensiana 59 I V III II III II V Cladonia ecmocyna 2 . III . . . . . Dicranum brevifolium 2 . III . . . . . Dicranum muehlenbeckii 8 . III . . . I I Luetkea pectinata 26 I IV I II . I III Phyllodoce empetriformis 60 II IV IV . III II V Vaccinium membranaceum 15 . V I . . I I           Arnica latifolia 57 II III III . IV III II Barbilophozia species 16 I II III . II I I Carex spectabilis 75 II . V II . V II Cassiope mertensiana 59 I V III II III II V Phyllodoce empetriformis 60 II IV IV . III II V Picea engelmannii 18 I II III . I I I Valeriana sitchensis 46 I II III III III III II           Abies lasiocarpa 79 II . . V III II III Aulacomnium palustre 9 . . . IV . I I Calamagrostis canadensis 4 . . . III . I . Caltha leptosepala 7 . . . V . I . Carex scirpoidea 5 I . . III . . I Coeloglossum viride 2 . . . III . . . Erigeron peregrinus 49 II . I III IV IV II Eriophorum angustifolium 2 . . . III . . . Juncus drummondii 4 I . . III . . I Leptarrhena pyrolifolia 3 . . . III . . I  201  Group No.  2 4 3 7 1 6 5 No. of relevés 33 4 20 4 7 37 26 Marsupella species 4 . . . III . I I Ranunculus eschscholtzii 5 . . . III I I . Salix barclayi 4 . . . III . . I Senecio triangularis 35 . . I V III III II Sphagnum species 5 . . . III . I . Trollius albiflorus 11 . . . IV I I I Valeriana sitchensis 46 I II III III III III II Veratrum viride 29 . . I V I III I           Abies lasiocarpa 79 II 200 180 V III II III Anemone occidentalis 44 I . II . III III III Anemone parviflora 6 . . . . IV I . Antennaria racemosa 5 . . . . IV . . Arnica latifolia 57 II III III . IV III II Artemisia norvegica 59 V . I . IV III I Brachythecium species 17 I . II . III I I Cassiope mertensiana 59 I V III II III II V Claytonia lanceolata 21 I . I II III II I Erigeron peregrinus 49 II . I III IV IV II Hieracium gracile 40 II . II . III III II Lycopodium species 18 II . I . III I I Pedicularis bracteosa 16 . . . . III II I Peltigera species 6 I . I . III I . Phyllodoce empetriformis 60 II IV IV . III II V Poa alpina 15 I . . . III I I Pogonatum species 15 I . I . III I I Salix arctica 6 . . . . III I I Saxifraga bronchialis 9 I . . . III . I Selaginella densa 9 I . I . III I . Senecio triangularis 35 . . I V III III II Sibbaldia procumbens 49 IV . I . III II I Vahlodea atropurpurea 49 I II II II III III IV Valeriana sitchensis 46 I II III III III III II Veronica wormskjoldii 12 I . . . III I I           Anemone occidentalis 44 I . II . III III III Arnica latifolia 57 II III III . IV III II Artemisia norvegica 59 V . I . IV III I Carex spectabilis 75 II . V II . V II  202  Group No.  2 4 3 7 1 6 5 No. of relevés 33 4 20 4 7 37 26 Erigeron peregrinus 49 II . I III IV IV II Hieracium gracile 40 II . II . III III II Phleum alpinum 31 I . . . II III I Senecio triangularis 35 . . I V III III II Vahlodea atropurpurea 49 I II II II III III IV Valeriana sitchensis 46 I II III III III III II Veratrum viride 29 . . I V I III I           Abies lasiocarpa 79 II 200 180 V III II III Anemone occidentalis 44 I . II . III III III Carex nigricans 24 I . I . . II III Cassiope mertensiana 59 I V III II III II V Luetkea pectinata 26 I IV I II . I III Luzula parviflora 34 I . II II II I III Phyllodoce empetriformis 60 II IV IV . III II V Vahlodea atropurpurea 49 I II II II III III IV       203  Table 45 Raw soil analysis.  Org Matr C:NTotal CTotal N LOI NH4-NNO3-N < 2mm pH% % % ppm ppm %BEC Site Site ElevationAspect SlopeSubzone Series Unit  m.a.s.l.deg. %IMAwc 23.9 1.45 44.3 16.6 0.1 92 17 3.9 o1 110/Am03 2105 227 5IMAwc 17.6 1.08 33.3 33.9 8.4 76 16 4.3 o4 104 2120 260 25IMAwc 8.4 0.59 17.4 9.9 2.3 63 14 3.8 o2/o4 105/At29 2155 221 40IMAwc 5.5 0.35 12.3 5.3 0.2 58 16 4.2 o4 106/At22 2150 115 25IMAwc 1.1 0.08 3.6 1.8 0.6 73 13 5.6 o2 104 2158 0 0IMAwc 0.8 0.05 2.7 1.5 0.4 85 16 4.7 o2 102/At71 2215 120 40IMAwc 1.1 0.08 3.6 2.9 0.3 79 14 4.3 o2 102/At71 2295 121 42IMAwc 2.6 0.15 6.2 4.2 0.4 70 17 3.9 o3 111/Ah03 2198 221 13IMAwc 2.5 0.16 5.7 3.5 0.3 74 15 4.3 o2 102/At71 2295 121 42IMAwc 3.1 0.20 6.8 4.3 0.2 70 16 3.6 o1-03 106/At22 2200 225 35IMAwc 3.2 0.22 6.9 4.0 0.4 83 15 3.9 o2 102/At71 2264 162 10IMAwc 18.0 1.11 34.0 17.9 0.7 72 16 4.1 o1 110/Am03 2105 227 5IMAwc 9.9 0.71 19.8 6.3 4.7 59 14 3.9 o4 103/Af01 2163 0 0IMAwc 9.0 0.53 19.6 9.8 0.8 69 17 4.6 o4 104 2120 260 25IMAwc 10.8 0.70 23.1 14.3 0.9 77 16 3.9 o4 106/At22 2150 115 25ESSFwcp 6.1 0.47 12.1 22.1 0.4 67 13 3.8 o1 112/Am02 1920 300 10ESSFwcp 24.1 2.10 50.8 70.5 37.0 60 11 5.3 o7 116/Aw15 2005 223 55ESSFwcp 6.2 0.55 13.2 44.8 0.1 61 11 3.6 o1 111/ Ah02.2 1968 0 0ESSFwcp 3.0 0.18 6.7 4.4 0.2 83 16 3.8 o1 Sileaca-Artenor1960 0 0ESSFwcjp 2.4 0.13 6.2 3.6 0.0 84 18 4.1 o1-o2 Sileaca-Artenor1960 0 0ESSFwcp 4.4 0.30 9.4 7.7 0.0 84 15 3.5 o1 112/Am02 1960 0 0ESSFwcp 2.6 0.15 6.2 5.3 0.0 75 18 3.9 o1 112/Am02 1960 0 0IMAwc 5.8 0.38 13.4 7.0 0.0 77 15 3.8 o4/o1 106/At22 2248 152 16IMAwc 2.6 0.18 6.8 7.8 0.0 76 15 4.1 o2 103/Af01 2292 0 0IMAwc 5.3 0.36 12.5 20.9 0.0 90 15 3.8 o5 112/As01 2122 127 7IMAwc 7.4 0.43 14.9 17.2 0.0 90 17 3.6 o5 112/As01 2122 127 7IMAwc 4.2 0.29 9.7 12.7 0.0 74 15 3.8 o3-o1 106/At22 2180 192 13ESSFwcp 6.1 0.26 12.2 11.1 0.3 78 23 3.7 o5 110/Sk14 2097 227 80IMAwc 4.6 0.38 10.1 12.4 1.4 72 12 3.5 o3-o1 106/At22 2180 192 13ESSFwcp 5.6 0.39 11.8 21.2 0.3 81 15 3.8 o4 112/Am02 2015 197 70ESSFwcp 8.2 0.52 16.5 14.6 1.4 84 16 3.5 o4 110/Sk14 2017 195 70ESSFwcp 4.5 0.26 9.3 14.7 0.8 78 17 3.7 o5 110/Sk14 2024 65 50ESSFwcp 8.4 0.66 17.9 2.7 0.0 91 13 3.9 o4-o1 103/Sk01 1896 329 25ESSFwcp 4.2 0.30 8.5 3.0 0.0 89 14 4.1 o5-o4 101/Am03 1906 137 30ESSFwcp 3.6 0.27 8.3 4.1 0.0 84 13 4.0 o3 Sk60 1889 39 22ESSFwcp 2.1 0.11 5.8 3.7 0.0 62 20 3.8 o4-o1 101/Am03 1893 62 30ESSFwcp 2.1 0.17 5.0 3.8 0.0 87 12 4.3 o5-o1 101/Am03 1895 0 0ESSFwcp 6.7 0.45 13.9 9.6 0.0 37 15 3.7 o2-o3 102/Af01 1916 227 76ESSFwcp 8.0 0.59 15.6 13.9 0.0 41 13 3.7 o2 102/Af01 1960 205 10IMAwc 20.0 1.59 43.2 7.1 2.7 50 13 3.9 o4 106/At22 2077 154 8ESSFwcp 11.3 0.84 24.9 8.6 0.0 57 13 3.7 o4 110/Sk14 1935 103 65ESSFwcp 8.3 0.66 17.7 8.2 0.4 46 13 3.7 o1 101/Am03 1995 0 0ESSFwcp 11.4 0.83 24.6 16.1 0.4 39 14 4.1 o4-o5 110/Sk14 1990 161 35ESSFwcp 7.0 0.46 16.0 4.9 0.0 50 15 3.7 o1 101/Am03 1956 116 5ESSFwcp 8.4 0.54 17.3 5.1 0.0 56 16 3.4 o4-o5 110/Sk14 1921 126 12Total C and N Available NBy Elemental Analyzer 204  Table 46 IMAwc site series species list. Bars indicate the frequency and/or abundance of common species (rows) expected in each site series columns) IMAwc Site Series 02 04 01 03 05       Shrub Layer      Vaccinium scoparium l lll l   Juniperus communis  l    Cassiope mertensiana  ll lll llllll ll Phyllodoce spp. l ll l llllll  Empetrum nigrum llllll lll  l  Salix nivalis lll llllll l ll lll Cassiope tetragona llllll l  l  Loiseleuria procumbens llll l          Herb Layer     Luzula spicata llll lll l ll l Luzula spp.   l lll ll llll Arnica latifolia l l lll  lll Erigeron peregrinus     ll Lupinus arcticus    l ll Valeriana sitchensis  l l ll lll Veratrum viride  lll lll ll ll  205  IMAwc Site Series 02 04 01 03 05 Anemone occidentalis    lll lll Senecio triangularis  ll ll l llll Claytonia lanceolata     ll Erythronium glanduliflora  lll ll  l Silene parryi    l ll Ranunculus eschscholtzii    I III Sibbaldia procumbens llll llllll ll l l Castilleja miniata    ll ll Calamagrostis canadensis   ll lll Carex nigricans l ll ll lll llllll Carex spectabilis   l lll lll Luetkea pectinata l l ll lll  Vahlodea atropurpurea  lllll lll l   Festuca brachyphylla lll ll l ll  Poa alpina ll ll    Festuca altaica ll ll llll ll ll Artemisia norvegica     ll lll  206  IMAwc Site Series 02 04 01 03 05 Trollius albiflorus lll llll ll   Potentilla villosa    ll lll Saxifraga ferruginea ll lll lllll ll l Antennaria lanata llll lll llII l  Epilobium anagallidifolium ll ll  lll  Saxifraga bronchialis lll lll ll l  Minuartia obtusiloba    l lll llll Phleum alpinum  l ll ll  Silene acaulis lllll lll    Stellaria longifolia ll lll ll l  Epilobium latifolium   l l ll Oxyria digyna  lll lll l l  Leptarrhena pyrolifolia    ll ll Potentilla diversifolia l ll ll l        Moss/Lichen Layer       Barbilophozia spp.    l ll ll Lescuraea radicosa      lll  207  IMAwc Site Series 02 04 01 03 05 Brachythecium  l l l ll ll Polytrichum spp.  lll lll l ll lll Dicranum spp.  ll ll l l  Lepraria neglecta llllll lll ll   Cetraria nivalis llll lll l   Coelocaulon aculeatum  lll lll ll   Campylium stellatum    l ll Marsupella spp.     l ll    208  Table 47 ESSFwcp site series species list. Bars indicate the frequency and/or abundance of common species (rows) expected in each site series columns)  ESSF wcp Site Series 01  03 04 05 06 07 02         Tree Layer        Abies lasiocarpa ll llllll lllll llll l l lll Picea spp.  ll llll lll ll l l  Pinus albicaulis l lll ll    llll         Shrub Layer        Rhododendron albiflorum  lll l     Vaccinium membranaceum  llllll lll l l  l Ribes lacustre   lll l l  ll l Juniperus communis       lllll Cassiope mertensiana l ll l llll llllll ll l Phyllodoce empetriformis l ll l lll lll ll ll Empetrum nigrum   l   l lllll  209  ESSF wcp Site Series 01  03 04 05 06 07 02 Salix barclayi ll   lll  ll          Herb Layer        Luzula spp.  ll ll l ll l l llll Arnica latifolia lll l llll ll l  ll Erigeron peregrinus lllll  lll ll  ll ll Lupinus arcticus lllll ll lll l   l Valeriana sitchensis lllll lll lllll l lll l ll Veratrum viride lll  ll l ll  l Anemone occidentalis lll l ll l lll ll llll Senecio triangularis llll l ll l ll l l Pedicularis bracteosa lll ll lll l   llll Mitella spp.  ll ll l llll lll   Claytonia lanceolata lllll  ll   lll lll Erythronium glanduliflora ll  ll    lllll  210  ESSF wcp Site Series 01  03 04 05 06 07 02 Silene parryi lll  ll    llll Ranunculus eschscholtzii    l ll lllll  Sibbaldia procumbens l  l  ll  lll Thalictrum occidentalis ll  l     Castilleja miniata lllll  ll   l ll Calamagrostis canadensis l    l lll  Caltha leptosepala      llll  Carex nigricans    l ll lllll  Carex spectabilis llll ll llll l ll ll ll Luetkea pectinata  ll ll lll llll lll  Leptarrhena pyrolifolia     l  llll lll Heracleum lanatum  lllll  l     Vahlodea atropurpurea  ll  l ll llll ll                       211  ESSF wcp Site Series 01  03 04 05 06 07 02   Moss Layer  Barbilophozia spp.   ll ll llll ll ll  Lescuraea radicosa  ll l ll l lll lll  Brachythecium  ll l lll ll l  lll Polytrichum spp.    lll lll  ll llll Sphagnum spp.      l ll   

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