Interspecific Associations, Phenology, and Environment of Some Alpine Plant Communities on Lakeview Mountain, Southern British Columbia

INTERSPECIFIC ASSOCIATIONS, PHENOLOGY, AND ENVIRONMENT OF SOME ALPINE PLANT COMMUNITIES ON LAKEVIEW MOUNTAIN, SOUTHERN BRITISH COLUMBIA

MARILYN JEAN RATCLIFFE

B.Sc, The University. Of Victoria, 1979

A THESIS SUBMITTED IN PARTIAL FULFILMENT OF

THE REQUIREMENTS FOR THE DEGREE OF

MASTER OF SCIENCE

THE FACULTY OF GRADUATE STUDIES

Botany Department

We accept this thesis as conforming

to the required standard

THE UNIVERSITY OF BRITISH COLUMBIA

July 1983

In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission.

Department of BOTANY

The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3

Date JULY 29, 1983

DE-6 (3/81) i i

Abstract

Three major alpine plant communities were identified on

Lakeview Mountain, Cathedral Provincial Park, using multivariate analysis of percentage cover data. Communities were dominated by Kobresia myosuroides, Carex scirpoidea (with one transitional area dominated by both Kobresia myosuroides and Carex scirpoidea), or by Carex scirpoidea and Carex capitata (with

Salix nivalis as an additional dominant at one site). Community composition and distribution had little relationship with aspect or with the soils and microclimatic factors measured.

Phenology was recorded for vascular species during the summer of 1980. Later flowering times were observed for a number of species in Kobresia myosuroides or Carex scirpoidea/Carex capitata dominated vegetation, and plants generally flowered earlier on southern aspects.

Small scale patterns in the form of significant associations between species-pairs were detected in all communities, using a plotless point-line sampling technique.

Patterns were abundant at this scale, with a total of 182 significant positive associations and 103 significant negative associations recorded between different species pairs. These interspecific associations varied considerably between sampled sites in the study area, with many occurring only once.

Possible association-generating mechanisms have been discussed, and characteristics of the genotype, rather than the taxonomic species, have been suggested as critical in the formation of associations. A competitive hierarchy of dominant species has also been proposed, based on interspecific association and phenological data.

Soils within the study area are classified as Alpine

Dystric Brunisols, and are coarse textured, strongly acidic, low in available nutrients, and high in organic matter.

Climate was relatively uniform over the study area during the 1980 growing season, as were microclimatic air and soil temperature profiles and air humidity profiles. Lower soil temperatures, however, occurred beneath Kobresia myosuroides dominated vegetation. iv

Table of Contents

Abstract ii List of Tables vi List of Figures vii Acknowledgements viii I. INTRODUCTION 1 THE ALPINE ZONE 2 LITERATURE REVIEW 4 OBJECTIVES 10 11 . STUDY AREA 11 LOCATION 11 LAND USE 13 GEOLOGY AND GEOMORPHOLOGY 15 GEOLOGY 15 GEOLOGICAL HISTORY 15 SOILS 16 CLIMATE 17 VEGETATION 18 ANIMALS 23 III. METHODS 24 VEGETATION 24 TRANSECT PLACEMENT 24 QUADRAT SAMPLING DESIGN 26 ANALYSES OF QUADRAT DATA 28 POINT-LINE SAMPLING 33 ANALYSIS OF POINT-LINE DATA 34 PHENOLOGY 38 NOMENCLATURE 39 SOILS 39 CLIMATE 41 IV. RESULTS 43 VEGETATION 43 COMMUNITY TYPES 43 1 . Transect A: 43 2. Transect B: 46 3. Transect C: 48 4. Transect D: 51 5. Transect E: ...... 54 6. Transect F: 57 7. All Transects: ..60 PHENOLOGY 67 1. Constant Aspect: ...74 2. Constant Community Type: 75 INTERSPECIFIC ASSOCIATIONS 77 1 . Transect A: 78 2. Transect B: ..' 81 3. Transect C: 81 4. Transect D: .84 5. Transect E: 90 6. Transect F: 93 7. All Transects: 97 POSITIVE ASSOCIATIONS: 97 V

NEGATIVE ASSOCIATIONS: 115 CONSTANCY OF INTERSPECIFIC ASSOCIATIONS 129 SPECIES ORDINATIONS 132 SOILS 135 MORPHOLOGY 135 PHYSICAL AND CHEMICAL PROPERTIES 135 CLIMATE 138 MESOCLIMATE 138 MICROCLIMATE 138 V. DISCUSSION 146 COMMUNITIES 146 PHENOLOGY 150 INTERSPECIFIC ASSOCIATIONS 152 POSSIBLE MECHANISMS GENERATING POSITIVE ASSOCIATION 154 1. Niche Differentiation: 154 2. Balanced Competitive Abilities: 158 3. Additional Mechanisms: 159 POSSIBLE MECHANISMS GENERATING NEGATIVE ASSOCIATION164 1. Morphology: 1 64 2. Abiotic Effects: 164 3. Competitive Exclusion: 165 GENOTYPIC RESPONSE 168 DOMINANT SPECIES 170 SOILS 175 CLIMATE 176 CONCLUSIONS 178 VI . SUMMARY 181 VEGETATION 181 SOILS 184 CLIMATE 184 VII. LITERATURE CITED 185 APPENDIX A - GEOLOGICAL HISTORY 213 APPENDIX B - SOILS 215 APPENDIX C - PERCENTAGE COVER DATA FOR SIX TRANSECTS ....217 APPENDIX D - PRINCIPAL COMPONENTS ANALYSIS OF SOIL DATA .224 vi

List of Tables

I. Selected Climatic Data for Weather Stations in Southern British Columbia (within the 1941-1970 period) . 19 II. Transect Characteristics 24 III. Methodology for Physical and Chemical Soil Analyses 40 IV. Associated Species 77 V. Positive Associations for 11 Sample Groups 98 VI. Negative Associations for 11 Sample Groups 116 VII. Constancy of Positive Associations 130 VIII. Constancy of Negative Associations 131 IX. Species Pairs Associated Both Positively and Negatively in Different Vegetation Groups 133 X. Physical and Chemical Properties from Soil Profiles within 11 Vegetation Groups 136 XI. Mesoclimatic Data For Three Weather Stations, 1: S84°W, 2481 m; 2: S2°E, 2402 m; and 3: N29°E, 2475 m. 139 vii

List of Figures

1. Location of Study Site in Cathedral Provincial Park, British Columbia ..12 2. Position of Transects (A-F) in the Study Site 25 3. Quadrat Sampling Design 27 4. Transect A Multivariate Analyses 44 5. Transect B Multivariate Analyses 47 6. Transect C Multivariate Analyses 50 7. Transect D Multivariate Analyses 53 8. Transect E Multivariate Analyses 56 9. Transect F Multivariate Analyses 58 10. Centered PCA - All Data Sets 62 11. Centered and Standardized PCA - All Data Sets 63 12. RA - All Data Sets 64 13. Cluster Analysis - All Data Sets 66 14. Transect A Phenology 69 15. Transect B Phenology 69 16. Transect C Phenology 70 17. Transect D Phenology 71 18. Transect E Phenology 72 19. Transect F Phenology 73 20. Group A Positive and Negative Associations 80 21. Group B Positive and Negative Associations 82 22. Group C1 Positive and Negative Associations 83 23. Group C2 Positive and Negative Associations 85 24. Group D1 Positive and Negative Associations 87 25. Group D2 Positive and Negative Associations 88 26. Group D3 Positive and Negative Associations 89 27. Group E1 Positive and Negative Associations 91 28". Group E2 Positive and Negative Associations ....92 29. Group F1 Positive and Negative Associations 94 30. Group F2 Positive Associations 95 31. Group F2 Negative Associations 96 32. Microclimatic data for 11 vegetation groups during summer 1980 140 vi i i

Acknowledgement

I wish to thank my supervisor, Dr. Roy

Turkington, for his assistance, enthusiasm, and

guidance throughout the course of this study. Thanks

are also extended to my commitee members, Drs. G.

Bradfield and J. Maze, for providing valuable

suggestions and critically reviewing the manuscript.

I am grateful to Drs. Piet de Jong and Malcom

Greig for statistical consultation, and to Dr. T.

Ballard for advice regarding soil sampling and

analysis and for providing laboratory facilities. The

computer program for analysis of interspecific

associations was written by Dave Zitton. I also thank

Barry Wong and John Emanuael for computing

consultation. Aid in the identification and

verification of plant species was provided by Dr. G.W.

Douglas and Terry Mcintosh.

Immeasurable appreciation is extended to Dr. G.W.

Douglas for providing advice, inspiration, and

untiring interest during this research, particularly

in the first field season. I also thank George A.

Douglas, Mr. A.G. Ratcliffe, Melanie Madill, and

Angela Chen for field and data manipulation assistance, and Tom Fleet and Karl Gehringer of

Cathedral Lakes Resort for their interest in the study ix

and for their friendship.

Finally, very special thanks are due to Dr. K.

Wilf Nicholls. His assistance with final figures and, most importantly, his unfailing patience, support, and

understanding, made the completion of this thesis possible.

Financial assistance through a National Science and Engineering Research Council of Canada postgraduate scholarship is gratefully acknowledged. 1

I . INTRODUCTION

Alpine vegetation ecology has been essentially limited to the consideration of community patterns in relation to abiotic factors, such as time of snow release, microclimate, and physical and chemical properties of the soil. Savile (1960) has stated that in severe environments such as the alpine, competition is unimportant compared to physical factors,

"allowing essentially random occurrence of plants without distinct associations and frequent coexistence of related species that have extremely similar requirements". This opinion is supported by Bliss (1962). Whittaker (1975) argued along similar lines and considered evolution to be affected more strongly by selection for survival in relation to problems of the physical environment and less strongly by selection involving interaction and competition with other species. Even

Darwin (1859), who emphatically stressed the importance of biotic factors such as competition, considered the "struggle for life" to be almost exclusively with the elements when one reached the "arctic regions or snow-capped summits". As well, studies of pattern in the alpine zone have mainly been done on a large community scale, with small-scale pattern within communities and biotic interactions between neighbouring species rarely being investigated.

In terrestrial ecosystems other than the alpine, however, the study of small-scale pattern or non-random species distributions within plant communities has received much attention. This pattern has been attributed to both abiotic 2

(Blaser and Brady 1950, Harper and Sagar 1953, Snaydon 1962) and biotic factors (Watt 1947, Harper e_t al. 1961, Mack and Harper

1976, Turkington and Harper 1979). In view of the lack of this type of approach in the alpine zone, and the untested assumptions of many workers, the study of small-scale pattern within alpine communities merits attention.

THE ALPINE ZONE

The term "alpine" is defined here as the area above the elevational limit of upright trees, where tree species, if present, assume a dwarfed or krummholz appearance (Douglas and

Bliss 1977). This definition has wide application in the northern hemisphere and is similar to an earlier recommendation made by Love (1970). The alpine zone may begin at elevations as low as 300 m in the extreme north, or as high as 3500-4000 m in tropical areas, where altitude compensates for latitude in lowering the average temperature (Billings 1974a). The alpine zone in the southern interior of British Columbia begins at approximately 2200 m.

The alpine climate is characterized by cool temperatures, both in winter and summer. With increasing altitude, the temperature of the ambient air decreases at an average rate of

5-6° C per 1000 m, with a corresponding decrease in the diurnal temperature range (Billings 1973, Barry and Van Wie 1974,

Bennett 1976). Mean annual temperatures, for example, can be as low as -2° C, with daily maximums of 10-11° C for June through

August, in south-central British Columbia 3

(B.C. Dept. Agri. 1974). The low density of the alpine atmosphere lessens its ability to store heat (Bennett 1976), and the velocity of wind increases with increasing altitude, due to thermal discontinuity (Tranquillini 1964, Flohn 1974). The intensity of solar radiation, particularly in the ultra violet

(U.V.) spectrum, also increases with altitude (i.e., on average, a 50% increase in clear-sky U.V. from sea level to 3650 m in elevation) (Gates and Janke 1966, Caldwell 1968) and is primarily due to a decrease in atmospheric scattering and absorption capacity (Terjung et al. 1969, Barry and Van Wie

1974). Precipitation generally tends to increase with altitude in mid-latitude mountains due to air mass lifting and cooling

(Flohn 1974), and results in a high degree of snow accumulation compared to lower elevations (Barry and Van Wie 1974). Due to snow accumulation, the growing season for alpine plants rarely exceeds 90 days in mid-latitude areas (Billings 1973).

Alpine areas are characterized by a relatively small number of perennial vascular plant species, which include both evergreen and herbaceous growth forms. A total of 170 alpine vascular species have been reported from the Rocky Mountains of

Colorado (Holm 1927), and 165 have been recorded for the eastern

North Cascade Range in Washington and British Columbia (Douglas and Bliss 1977). Rosette species, sedges, grasses, cushion plants, lichens, and mosses are common, with most species assuming a prostrate or dwarf growth form (Bliss 1969, Billings

1974a). The heterogeneity of alpine microtopography often results in marked changes in physical factors such as soil 4

temperature and moisture, depth of thaw, wind speed, and

snowpack over distances of only a few centimeters (Marr 1961,

Billings 1973, Billings 1974a). Steep environmental gradients

such as this result in abrupt changes between plant communities,

while areas with more gradual environmental changes tend to have

transition zones between communities. (Bliss 1969). This mixture

of abrupt and gradual environmental changes is reflected in a mosaic of relatively small and often discrete communities over

the landscape.

LITERATURE REVIEW

Plant ecologists have been investigating various aspects of

the vegetation of alpine zones in North America since the early

1900's, e.g., Cooper (1908), Taylor (1922), Cox (1933), and

Whitfield (1933). The majority of these studies emphasize

community patterns and associated environmental parameters

(Bliss 1956, Billings and Bliss 1959, Johnson and Billings 1962,

Mooney et a_l. 1962, Archer 1963, Bliss 1963, Scott and Billings

1964, Klikoff 1965, Beder 1967, Marr 1967, Bamberg and Major

1968, Eady 1971, Knapik et al. 1973, Crack 1977, Douglas and

Bliss 1977, Hrapko and La Roi 1978, Helm 1982), and have been

recently reviewed by Bliss (1969). A few alpine studies document cyclic vegetation dynamics in relation to environmental

factors such as freeze/thaw phenomena (Billings and Mooney 1959,

Johnson and Billings 1962, Mark and Bliss 1970). This emphasis

on community-orientated studies appears to be due to at least

two factors. First, the visually obvious vegetation patterns 5

invite study, and their description is a first step in ecological investigations. Second, the emphasis on abiotic

factors may be partially due to alpine climatic extremes which are often severe from a human standpoint. This leads to the

natural assumption that these conditions must be "harsh" or

"stressful" for the plants, and therefore important determinants of distribution.

Alpine studies that have shifted emphasis away from community description fall into two camps. First, there are

those that have considered the role of competition in vegetation

that forms closed swards (Rochow 1970, Billings 1974a, Callaghan

1976), but work in this area has been extremely limited. Shaver et al. (1979) have studied intergenotypic competition in Carex aquatilis Wahl. populations in arctic vegetation. Grime (1979), however, uses arctic and alpine vegetation as an example of

stress-tolerant (S-selection) plants and suggests that competition (refering exclusively to the capture of resources)

is relatively unimportant in such high stress environments.

Second, studies concerned with various functional or physiological aspects of the vegetation are common. For example, relatively low photosynthetic temperature optima of 15°

C - 20° C have been reported for alpine species (Hadley and

Bliss 1964, Mooney et a_l. 1964, Scott and Billings 1964, Mark

1975), with vascular plant photosynthesis recorded at temperatures as low as 3° C (Anderson and McNaughton 1973) and -

5° C (Mark 1975). Increased light intensities in the alpine likely contribute to the rapid summer growth rates (Tranquillini 6

1964, Tieszen and Bonde 1967, Billings 1974a). In Colorado, for

example, the average growth rate of Saxifraga rhomboidea Greene

early in the growing season- was 3.5 cm/week, while maximum peduncle elongation rates of 14 cm/week have been recorded for

Polygonum bistortoides Pursh (Holway and Ward 1965). Due to

this rapid growth, daily productivity rates for alpine herbaceous perennials are quite similar to those found for more

temperate ecosystems (Bliss 1966), although average annual productivity of alpine vegetation as a whole is comparatively

low (Lieth 1975). For example, in the Wyoming alpine, values

range from 1.2 g/m2/day on xeric sites to 11.1 g/m2/day on mesic

sites (Scott and Billings 1964), which is comparable to values

reported for temperate field situations (Odum 1 960, Ovington e_t al. 1963). As well, alpine plants generally exhibit higher root and shoot respiration rates than do species from lower elevations (Mooney 1963, Klikoff 1968, Higgins and Spomer 1976), which has been interpreted as a metabolic adaptation to survival

in cold temperatures (Mooney e_t al. 1964). In view of the physiological specializations found in alpine species, it does not seem prudent to rule out the possibility of competitive

interactions, particularly since resources, such as available

soil nutrients, may be in short supply (Bliss 1963, Nimlos and

McConnell 1965, van Ryswk 1969, Bockheim 1972, Sneddon et al. 1972, Knapik et al. 1973).

Recently, there has been a shift in the nature of general plant ecological research from the early study of communities and continua (e.g., Clements 1904, Gleason 1926, Braun-Blanquet 7

1932, Tansley 1939), to the study of small-scale vegetation pattern involving populations of individuals within communities.

A very limited number of early population and experimental

studies were done (Tansley 1917, Clements and Weaver 1924,

Sukatshew 1928), followed by approximately thirty years in which

this approach was virtually disregarded by plant ecologists, although population studies continued in systematics and the applied sciences of forestry and agriculture. The problem of

identifying individuals and the lack of communication between

investigators of natural vegetation and those of managed systems have been postulated by Harper (1977a) as possible reasons for

this gap within plant ecology research. The population and

individual approach has gained increasing popularity since the

investigation by Watt (1947) of processes at this scale within plant communities (see Newman 1982a).

The methods used to describe and detect pattern have changed as the detail of enquiry by investigators has changed.

Visually obvious community patterns were initially recorded

subjectively. This was followed by more detailed scrutiny with species presence/absence or percentage cover recorded in quadrats or along a transect of points (Greig-Smith 1964).

Patterns within grassland communities based on random or contiguous quadrat data have been documented by early ecologists

(Blackman 1935, Clapham 1936, Archibald 1948). Later studies attributed small-scale patterns to such factors as soil nutrients (Blaser and Brady 1950, Kershaw 1958, 1959), soil pH

(Snaydon 1962), microtopography (Harper and Sagar 1953, Kershaw 8

1963), species morphology (Kershaw 1963), and cyclic

regeneration of species (Watt 1947, Goodall 1952, Greig-Smith

1952). The concept of plant species themselves influencing

pattern has also been considered (Harper et al. 1961, Harper

1964, 1967). This conventional, quadrat approach has been

critisized by Greig-Smith (1961) who stated that the scale of

study (i.e., the quadrat) is imposed by the investigator and has

little relevance to the organisms living within it. Recently, a

finer scale of discrimination considering above-ground physical contacts between different species has been employed. This method detects patterns between species at the scale of the

individual and has been used in bryophyte/1ichen communities

(Yarranton 1966), and in temperate grasslands (Turkington et al. 1977, Aarssen et al. 1979, Turkington and Harper 1979a,b,

Aarssen 1983). This methodology indicates a particular type of

small-scale structure, which suggests, by its very nature, possible causalities. Biotic interactions between species

(e.g., competition) and differential use of resources have been

suggested as important factors influencing pattern in these

studies. Interspecific associations do not prove the existence of biotic interaction, but it is unlikely, for example, that two

individuals growing in physical contact are not influencing each other to some degree. For example, Ross and Harper (1972) found the growth of individual Dactylis glomerata L. seedlings was greatly influenced by the distance and morphology of neighbouring plants, while Mack and Harper (1977) found proximity and placement of neighbouring species accounted for 9

69% of the growth and reproductive variation in a sand dune

annual.

Attitudes of plant ecologists are seemingly dependent on

the scale of investigation. Harper and his students have been

largely responsible for redirecting emphasis to the ecology of plant populations rather than communities and to the consideration of the role of natural selection in maintaining and changing these populations. As natural selection affects

the individual, and evolution occurs within populations, the community is determined by processes operating at the individual and populational levels. It is necessary to determine the processes that generate patterns in order to understand the

structure of a community (Ricklefs 1979). Description alone will not accomplish this. The community as a "whole" must also be considered, however, as the evolution and variation of each species is related to the community (Mcintosh 1970), and studies of species in isolation may have little relevance to the behavior of the species in a community context. Greig-Smith

(1979) has stressed the importance of bridging the gap between community and population ecology, stating that vegetation pattern is a continuum of both scale and intensity, begining with the growth form of the individual. 10

OBJECTIVES

The primary objective of this thesis is to determine the

nature and extent of patterning at a small scale within alpine

plant communities, and to compare this, as well as large-scale

community patterns, to measured abiotic and phenological data.

It is hypothesized that vegetation patterns may, to some extent,

be influenced by the plant species themselves. Specifically,

the study objectives are to: (1) detect small-scale patterns in alpine vegetation, using a modified method of interspecific association analysis for point sampling data; (2) relate

significant (p<0.05) interspecific associations to community composition and phenology; (3) relate interspecific associations and community composition to aspect, soil physical and chemical properties, and microclimate; and (4) interpret the results in terms of various abiotic or biotic processes potentially controlling species patterns in alpine vegetation. 11

II. STUDY AREA

LOCATION

The alpine zone of Lakeview Mountain in Cathedral

Provincial Park, British Columbia, was chosen for study.

Lakeview Mountain is part of the Okanogan Range in the north• eastern portion of the Cascade Mountains, and is located in the southern interior of British Columbia, at approximately

49°03'N, 120°09'W, just north of the Washington-British Columbia border (Fig. 1). The Cascade Mountains are separated from the

Coast Mountains to the north by the Fraser River and merge into the Kamloops plateau to the east; they extend from this point through to southern Oregon.

The alpine zone of Lakeview Mountain ranges from approximately 2200 m to 2600 m in elevation, and has continuous vegetation interspersed with boulder fields, rock rivers, stone polygons, and other forms of patterned ground. The study site lies to the north-east of Lakeview peak (Fig. 1), and has an elevation range of 2402 - 2500 m. This area was chosen for its relatively diverse and continous dry sedge vegetation, present on virtually all aspects, and for the comparatively long snowfree or growing season. More western alpine areas are characterized by greater snow accumulation, later snowmelt, and relatively cold, wet summers. In addition, species composition often changes more gradually in the eastern Cascades as environmental gradients are not as steep compared to more western areas (Douglas and Bliss 1977). 1 2

Figure 1 - Location of Study Site in Cathedral Provincial Par British Columbia

Br^TISH_CC^MBIA, CANADA . WASHINGTON~USA~ i _ 1 13

LAND USE

On May 2, 1968, 16,480 ha of land were established as

Cathedral Provincial Park, 30 years after it was first proposed.

On May 6, 1969 an additional 703 ha were included, which previously contained mineral claims (Cartwright 1970). On

September 11, 1975, approximately 25,911 ha were added to the park and the present area now comprises 33,468 ha (R.R. Howie pers. comm.). Cathedral Provincial Park is managed as a class

'A' wilderness park by the British Columbia Parks Branch,

Dept. of Recreation and Conservation, with recreation as the primary use (Cartwright 1970). The park completely encloses a small recreational reserve in the Haystack Lakes area (Fig. 1), where management does not exclude resource extraction should this later become feasible (Travers 1975).

The Cathedral Lakes area has been used for recreation

(e.g., hunting) since the late 1930's. In 1967, the main cabins of Cathedral Lakes Resort were built, and non-hunting, paying guests began to visit the area (Bill Fleet pers. comm.). There is no motorized public access, although hiking trails are available and transport may be obtained through Cathedral Lakes

Resort, which still retains two small lots within the park boundaries. An estimated 749 visitors use the park annually

(Dalziel 1971), but this was before the main lodge was completed, and is a modest estimate for present day use. Major activities for park visitors include camping, fishing, and hiking (Cartwright 1970).

The alpine and subalpine meadows within the park have had a 1 4

history of grazing by domestic sheep and cattle. In the late

19'th and early 20'th century, sheep were grazed in the alpine zone (Cartwright 1970), with subsequent sheep grazing occurring for approximately 10 years in the 1940's, in both alpine and subalpine areas (Tom Fleet pers. comm.). Sheep were then replaced by cattle, and grazing was generally limited to montane and subalpine meadows, with a few strays sometimes reaching the alpine zone (Bill Fleet pers. comm.). The class 'A' status of

Cathedral Park excludes the grazing of domestic animals, although two leases in the park currently permit the grazing of

157 head of cattle, the number maintained in the area before park establishment (R.R. Howie pers. comm.). The permit held by the Terbasket family includes the alpine zone on the east side of Lakeview Mountain, although this area is rarely used.

Cathedral Park is not considered key rangeland for any major wildlife species, but it is adjacent to range for a remnant herd of California Bighorn Sheep (350-400 individuals) and hunting is allowed in the park between August 31 and September 13 (Travers

1975).

Recent disturbance within Cathedral Provincial Park appears limited to hiking, grazing, and frequent small fires, caused primarily by lightning. Open, eroding soil pits carelessly left by a previous researcher are an additional disturbance in the alpine zone of Lakeview Mountain. 15

GEOLOGY AND GEOMORPHOLOGY

Geology

The Cascade Mountains are characterized by folded, metamorphosed sedimentary \ and volcanic rocks of Paleozoic and

Mesozoic origin, with intrusions of granitic and granodiorite batholiths (Holland 1964, McTaggart : 1970). Paleozoic sedimentary and volcanic -rocks include argillite, cherty argillite, limestone, quartzite, andesite, and volcanic breccia.

Mesozoic intrusive rock includes granodiorite, quartz monzonite, quartz diorite, granite, and syenite (Geol. Map Can., 1955).

Most of the Princeton Map area (which includes Cathedral Park) is underlain by intrusive and extrusive igneous rocks (Rice

1960). Bodies of "red" and "white" granodiorite have been found, in addition to "grey", in the Cathedral Park area and are partially derived from granitization of volcanic rocks (Rice

1960). Four geologic group members are exposed in Cathedral

Park (1) Permian sediments, (2) Triassic lava, (3) Jurassic granitic plutons, and (4) Eocene (Tertiary) sediments and volcanic material (Melcon 1975).

Geological History

Prior to the Pleistocene epoch, the Cascade Mountains experienced periods of marine sedimentation, vulcanism, compression, folding, erosion, and uplift (Daly 1912, Rice 1960,

Rudkin 1964, McTaggart 1970). It is likely that the Pleistocene ice sheets did not attain elevations over 2134 m in southern 16

B.C. (Nasmith 1962, Holland 1964), although Melcon (1975) gives evidence for a 2286-2377 upper limit. Alpine glaciation, however, was extensive. Surficial ash deposits from the late

Pliocene to Recent times have been reported within and to the south of the study area (e.g., Powers and Wilcox 1964, Fryxell

1965, Wilcox 1965, van Ryswyk 1969, Bockheim 1972). A more detailed account of the geological history of Cathedral

Provincial Park is given in Appendix A.

Lesser landforms or microtopographical features within the study area on Lakeview Mountain include tors in areas of quartz monzonite bedrock (Melcon 1975) and patterned ground such as sorted stone polygons and stone stripes. Talus areas and felsenmeer of broken balsalt also occur. The majority of these features originated in periglacial conditions accompanying the

Fraser glaciation (Melcon 1975) and continue today as a result of frost action. Frost hummocks and solufluction lobes occur on lower alpine slopes where soil moisture is adequate and result from freeze-thaw cycles in conjunction with gravity.

SOILS

Soils within the alpine zone of Lakeview Mountain have been examined in detail by van Ryswk (1969), who characterized the most extensive soil type as Alpine Brown with discontinuous ash layers - comparable to the Alpine Dystric Brunisol of the

Canadian System.

Structure soils such as sorted stone patterns and stone rivers are also found on Lakeview Mountain, as well as broken 17

rock, rock headwall, and talus units. All soils have been influenced by volcanic ash, burying and mixing of horizons, frost heaving, solufluction, surface erosion, and colluvial activity (van Ryswk 1969). Slow rates of chemical weathering due to cold temperatures and the erosive action of physical processes contribute to the general immaturity of soil profiles found in alpine areas (Retzer 1974).

Buried charcoal fragments (van Ryswyk 1971) may indicate that the treeline was at higher elevations in the past (van

Ryswyk and Okazaki 1979). A more detailed account of soil types within the study area is presented in Appendix B.

CLIMATE

The study area lies within the south interior climatic region of British Columbia described by Kendrew and Kerr (1955), which extends from the crest of the Coast Mountains east to the

Rockies, and from the 49'th parallel north to Prince George.

The climate is classified as mild continental with warm summers, cold winters, and low precipitation due to the rainshadow effect of the Coast and Cascade Mountains. Precipitation is well distributed over the year with some snow during winter and thunderstorms with rain and hail common in summer. Diurnal and seasonal temperature ranges are more extreme than those found in coastal areas to the west (Chilton 1981). Altitude and site exposure can cause considerable variation within this climatic regime.

The closest weather stations to the study area are at 18

Hedley Nickle Plate (49°20'N, 119°59'W) and Hedley (49°21'N,

120°05'W), both approximately 50 km from the study site.

Selected climatic data for these stations is shown in Table I., which includes data from the Old Glory weather station

(49°09'N, 117°55'W), 160 km to the east. The Old Glory station is located above the treeline and is within the south interior climatic region. It is the closest approximation to the alpine climate on Lakeview Mountain.

VEGETATION

The alpine zone of Lakeview Mountain is characterized by relatively diverse dry sedge vegetation, with an average of 30-

35 vascular plant species and 20-30 lichen and moss species occurring with each dominant. The treeline is at approximately

2200 m, with vegetation below this point dominated' by Abies lasiocarpa (Hook.) Nutt., Picea engelmani i Parry, and Larix lyallii Pari., with Vaccinium scoparium Leiberg and Phyllodoce empetriformis (Sw.) D. Don common in the understory. Krummholz forms of these tree species occur at the alpine-subalpine transition. Carex scirpoidea Michx. var. pseudosc irpoidea

(Rydb.) Cronq., is the most ubiquitous dominant species on dry, well-drained Lakeview Mountain sites, often co-dominanting with

Carex capitata L. Vascular species occurring with these dominants include Potentilla diversifolia Lehm., Arenaria obtusiloba (Rydb.) Fern., and Festuca ovina L. Lichen species such as Cetraria islandica (L.) Ach., C. cucullata (Bell.) Ach.,

C. nivalis (L.) Ach., and Thamnolina vermicular is (Sw.) Schaer. TABLE I - SELECTED CLIMATIC DATA FOR WEATHER STATIONS IN SOUTHERN BRITISH COLUMBIA (WITHIN THE 1941-1970 PERIOD)* J-A DENOTES JUNE TO AUGUST.

STATION TEMPERATURE ( C) AVERAGE FROST PRECIPITATION (CM) YEARS OF MEAN DAILY MIN. MEAN DAILY MEAN DAILY MAX. FREE PERIOD MEAN TOTAL MEAN RAIN RTAN SNOW RECORD ANN. J-A ANN. J-A ANN. J-A (DAYS) ANN. J-A ANN. J-A ANN. J-A

HEDLEY

49°21 'N 120°05'W 2 10 8 18 14 26 148 29 10 22 10 75 0 524 m

HEDLEY

NICKEL PLATE 49°20'N 119°59'W -3 k 2 11 8 18 48 54 14 21 14 330 10 1769 m

OLD GLORY

49°g9'N 117 55'W -5 4 -2 8 111 20 73 16 18 12 558 44 23 2347 m

* B.C. Dept. of Agriculture, 1974 20

are also prevalent. Communities dominated by Carex scirpoidea have been reported for the North Cascades (Douglas and Bliss

1977), the Snowy Mountains of central Montana (Bamberg and Major

1968), and for the Sierra Nevada Mountains of California (Major and Bamberg 1963). The existence of Carex capitata as a dominant is apparently limited to the North Cascades (Douglas and Bliss 1977), and the St. Elias Mounatins, Yukon (Douglas, pe r s. c omm.).

Vegetation dominated by Kobresia myosuroides (Vill.) Fiori is relatively frequent on similar dry well-drained sites with early snowmelt, and includes species of lesser abundance such as

Carex scirpoidea, Arenaria obtusiloba, Cetraria nivalis, and

C. cucullata. Kobresia myosuroides communities have been reported from the North Cascades (Douglas and Bliss 1977), the

Rocky Mountains (Cox 1933, Marr 1967, Bamberg and Major 1968,

Knapik et al. 1973, Hrapko and La Roi 1978, Komarkova and Webber

1978, Bell and Bliss 1979), the Sierra Nevada Mountains (Major and Bamberg 1963), and as far north as the St. Elias Mountains of the Yukon and Alaska (Hanson 1951, Douglas 1980).

Dry, well-drained habitats are also dominated by Dryas octopetala L., Salix nivalis Hook., or Salix cascadensi s

Cockerell. Species occurring with Dryas octopetala include

Lupinus lyallii A. Gray, Arenaria obtusiloba, and Potentilla diversifolia. Dryas octopetala occurs as a dominant species over a wide geographical area, and has been reported from the

North Cascades (Douglas and Bliss 1977), the Rocky Mountains of

Alberta (Beder 1967, Bryant and Scheinberg 1970, Hrapko and La 21

Roi 1978, Knapik et a_l. 1973), more southern Rocky Mountain regions (Johnson and Billings 1962, Holway and Ward 1965, Marr

1967, Bamberg and Major 1968), and northern areas in Alaska and the Yukon (Hanson 1951, Price 1971). Salix nivalis occurs with less abundant species such as Cerastium beeringianum Cham. &

Schlecht., Potentilla diversifolia, and Lupinus lyallii . Plant communities dominated by Salix nivalis have been reported from alpine areas in the North Cascades (Douglas and Bliss 1977),

Montana (Bamberg and Major 1968), and Alberta (Knapik et al. 1973). Potentilla diversi folia is a major species in vegetation dominated by Salix cascadensis, with Silene acaulis,

Carex sc i rpoidea, Dryas octopetala, Cetraria nivalis,

C. cuculllata, and C. islandica also common. This community has been reported from only two areas, the North Cascades of

Washington and British Columbia (Douglas and Bliss 1977), and the Medicine Bow Mountains of Wyoming (Billings and Bliss 1959).

Snowbed sites on Lakeview Mountain are dominated by Carex breweri Boott var. paddoensis (Suksd.) Cronq., Carex nigricans

Retz., or Antennaria lanata (Hook.) Greene vegetation types.

Carex breweri dominates sites which are snowfree by late July and occurs with Sibbaldia procumbens L., Arenaria obtusiloba,

Erigeron aureus Greene, and Polytrichum piliferum Hedw. This community appears to be restricted to the North Cascades

(Douglas and Bliss 1977). Vegetation dominated by Antennaria lanata also becomes snowfree by July, with dry conditions occurring by late summer. Other common species include Carex scirpoidea, Carex breweri, Carex nigricans, and Polytrichum 22

juniperinum Hedw. This community also occurs in the subalpine zone of Lakeview Mountain and has been reported from the alpine zone of the North Cascades (Douglas and Bliss 1977), the

Olympics (Bliss 1969), and the Rocky Mountains (Beder 1967,

Hrapko and La Roi 1978, Knapik et al. 1973). Carex nigricans dominates poorly-drained sites where snow persists until early to late August. Greater than 90% cover of C. nigricans is common, with relatively few species occurring with this dominant, e.g., Salix cascadensis and Ranunculus eschscholtzii

Schlecht. This community is widespread and has been reported from the alpine and subalpine zones of the Cascade Mountains

(Douglas and Bliss 1977, Meredith 1972), the Olympic Mountains

(Bliss 1969), Kuramoto and Bliss 1970), and the Rocky Mountains

(Beder 1967, Knapik et al. 1973, Hrapko and La Roi 1978, Helm

1982).

A brief account of vegetation within the study area has been included in a description of Similkameen Valley plant communities (McLean 1970). Nearby Red Mountain, also within

Cathedral Park, has been sampled by Douglas and Bliss (1977), contributing data to their alpine and subalpine zone plant community study of the North Cascade Mountains. The western

North Cascades of Washington have been included in a general survey of Oregon and Washington vegetation (Franklin and Dyrness

1973), and alpine vegetation within the interior plateau region of south-central British Columbia has been studied by Eady

(1971). The ecology of Larix lyallii (Alpine Larch) within the eastern North Cascades has been considered by Arno and Habeck 23

(1972) .

ANIMALS

Hoary Marmot (Marmota caligata [Eschscholtz], Columbian

Ground Squirrel (Spermophilus columbianus columbianus [Ord]),

Mule Deer (Odocoileus hemionus hemionus [Rafinesque3), Pika (Ochontona princeps [Richardson]), California Bighorn Sheep

(Ovis canadensis californiana Douglas), White-tailed Ptarmigan

(Lagopus leucurus), and Mountain Goat (Oreamnus americanus

[Blainvilie]) have been observed grazing alpine vegetation within the study area. Other herbivorous mammals living within or ranging into the alpine zone of Cathedral Park include

Northwestern Chipmunk (Eutamias amoenus [Allen]), Cascade

Mantled Groundsquirrel (Spermophilus saturatus [Rhoads]), Deer

Mouse (Peromyscus maniculatus [Wagner]), Vole (Microtus sp.), and Snowshow Hare (Lepus amer icanus Erxleben). Carnivores such as Lynx (Lynx canadensis Kerr), Black Bear (Ursus americanus

Pallas), Coyote (Canis latrans Say), and Weasel (Mustela sp.) range in to the alpine zone (Chess Lyons unpubl.) 24

III. METHODS

VEGETATION

Transect Placement

The study site is located in the high alpine zone of

Lakeview Mountain (Fig. 1). Six 2 m X 30 m belt transects were established at this site - chosen to include the most common dominant species present, as well as a range of aspects, for comparative purposes (Table II, Fig. 2). Transects were positioned immediately following snow release in June, 1980, using the remains of the previous years vegetation as the primary placement criterion. Each transect was placed in an

Table II - Transect Characteristics

TRAN. ASPECT SLOPE ELEV. DOMINANT SPECIES

A N58°W 9% 2450 _ Carex scirpoidea & C. capitata 2447 m

B S80°W 1 3% 2438 C. scirpoidea & C. capitata 2434 m

C N29°E 16% 2444 C. scirpoidea & C. capitata 2439 m

D S2°E 14% 2405 C. scirpoidea & Kobresia myosuroides 2401 m

E N1 6°W 1 0% 2426 C. scirpoidea & K. myosuroides 2423 m

F S56°E 7% 2475 C. scirpoidea & K. myosuroides 2473 m 25

Figure 2 - Position of Transects (A-F) in the Study Site. 26

attempt to include two areas dominated by different species and separated by a transition zone. Transects were 2 m wide as vegetation homogeneity tended to decrease with greater width and only one direction of major variation (vertical or downslope) was desired. Downslope variation was required to allow for future ease in identification of community types and transition zones, as well as for subsequent grouping of horizontal point- line data corresponding to these communities. Point-line sampling and association analysis are described in later methods sections. Transects were limited to 30 m in length so that a variety of aspects and vegetation types could be sampled within prevailing time constraints.

Quadrat Sampling Design

At 1 m intervals along the length of each transect, a 2m sampling row was established across the transect. Along each row, three 20 X 50 cm quadrats were randomly selected (using random number tables) from a possible 10 and placed perpendicular to the slope contours (Fig. 3). Each quadrat was divided into 20 sections, each 5X10 cm, aiding in the visual estimation of crown cover to the nearest 5% (less than 5% = T

[trace]) for each species. Crown cover is the percentage of ground covered by a vertical projection of the shoots of a plant species; intraspecific overlap is ignored. A total of 90 quadrats were sampled per transect, giving a sampling intensity of 15% for percentage cover within each belt transect. Sample sizes of 6.2% (Bliss 1963) and 4% (Douglas and Bliss 1977) have 27

Figure 3 - Quadrat Sampling Desi gn

SOIL PIT 1 m

0 ©

TEMPERATURE DIODES 30 m

ROWS OF EQUIDISTANT POINTS (EVERY 2cm) 1 m

QUADRATS (20 X 50cm) 28

been found to adequately describe similar vegetation and have satisfied the minimal area criterion of Cain (1938).

The 20 X 50 cm quadrat is well suited to the small stature of herbaceous alpine species (Bliss 1963) and allows relatively accurate visual cover estimates (Daubenmire 1968). Elongate or rectangular quadrats have been found to be the most efficient

(fewer quadrats needed to obtain a representative sample) when orientated parallel to the direction of greatest change or variance in the sampling area (Clapham 1932, Bormann 1953). A greater number of different species are likely to be included in each quadrat with this shape and placement (Kershaw 1973).

Analyses Of Quadrat Data

The multivariate techniques of ordination and cluster analysis were used with percentage cover data to aid in the definition of community types within each transect. This was necessary to assess the variation of communities with aspect.

As well, relationships between climate, soils, and phenological data, and different communities or different stands of the same community could be determined. Division of transects was also needed so that small scale vegetation data (point-lines) recorded within transects could be grouped and analyzed separately, according to the community type in which they were collected. This was done for comparative purposes. Also, a degree of vegetation homogeneity is necessary to satisfy statistical assumptions of the point-line or association analysis. The identification of community types must be made 29

with practical considerations in mind, however, as too many units have limited comparative and general information value.

In addition to separate multivariate analyses for each transect, data from all transects were combined for further analyses. This was done to aid floristic and abundance data comparisons between different stands of each community, as well as for an overall comparison of sampled communities within the study area.

Cluster analysis is a classification or data reduction technique used to separate data into homogenous groups (Everitt

1974), although groupings can often be arbitrary where communities intergrade continuously (Goodall 1978b, Whittaker

1978). Ordination is also a tool for data reduction, but is generally used to depict the range of variation in a data set rather than discontinuities. The two approaches of classification and ordination are, in theory, very, different, but may show similar trends in practice (Greig-Smith 1964) and can act to supplement and evaluate each other (Pritchard and

Anderson 1971, Whittaker and Gauch 1978). Discontinuities in a data set will be revealed as clusters in an ordination, often more clearly than with conventional clustering procedures

(Williams et al. 1969, Goodall 1978b).

Quadrats with 50% or more rock were deleted from each data matrix before any multivariate techniques or data transformations were applied. This was done to minimize the distortion produced by extreme sample outliers, particularly evident with ordination methods (Gauch et al. 1977). In 30

addition, species present in less than 5 quadrats were removed before ordination and clustering. Such species tend to encode little ecological information (Gauch 1977) and often distort results, particularly since centering and standardization techniques give equal weight to all species (Goodall 1978a,

Whittaker and Gauch 1978, Pimentel 1979). Transect quadrat data were grouped and averaged for each sampling row (Fig. 2), producing composite samples (from three original quadrats where rock was less than 50%). This resulted in thirty composite samples or strata per transect, which were then used for ordination and clustering. Grouping of samples can often clarify results by reducing the effects of sample error and chance differences in species abundances (Gauch 1973a, Gauch and

Whittaker 1981). In addition, a large data set is unwieldy when using two-dimensional scatter plots and cluster dendrograms.

Multivariate ordination techniques used were centered principal components analysis (PCA), centered and standardized

(PCA) (Orloci 1966, Gittins 1969), and reciprocal averaging

(RA) - a term first used by Hill (1973), although Hirschfeld

(1935) proposed the original algorithm. Data were analyzed by these ordination methods using the Ordiflex computer program package (Gauch 1977). This was used in conjunction with plotting programs (Wong unpubl.) which provided scatter diagrams showing sample numbers.

The indirect ordination methods used have been previously reviewed by Gauch and Whittaker (1972), Beals (1973), and Gauch et al. (1977). Principal components analysis (PCA) is limited 31

to relatively homogenous samples as it assumes a linear relationship amoung variables (Austin and Noy-Meir 1971, Orloci

1978) . Centered PCA subtracts the mean value for each species from the original values before eigenanalysis. The contribution of each species is therefore proportional to its variance and, as this tends to increase with mean, abundant species are often stressed (Noy-Meir e_t al. 1975, Pimentel 1979). Centered and standardized PCA standardizes species to unit variance, equalizing species contributions. This tends to emphasize rare and absent common species, rather than abundant ones (Goodall

1978a, Pimentel 1979). The variance accounted for by each axis is typically reduced by centering and standardizing (Austin and

Greig-Smith 1968). Involution of axes is more common with non- standardized data (Austin and Noy-Meir 1971), as are distortion due to sample clusters, and point scatter due to noise (Gauch et' al. 1977). When species abundances are important criteria, however, data standardization often gives poor results, as the single presence of a species receives a large score (Pimentel

1979) . Reciprocal averaging (RA) is useful when relatively high sample heterogeneity is present, with the resulting sample clusters producing less distortion than with PCA. A curvature of sample positions is produced by the second axis, however, and the ordination of outlying samples near the periphery of the scatter plot may cause compression of remaining samples in the opposite direction (Gauch et_ al. 1 977). In addition, the simultaneous double standardization of species and samples employed by RA tends to emphasize rare species and unique sites 32

(Pimentel 1979). RA is more suited to the non-linearity of ecological data than is PCA, however, with the first axis producing more accurate sample placement with- less distortion

(Whittaker and Gauch 1978). The application of a variety of multivariate techniques to each data set is advisable for comparative purposes.

An agglomerative hierarchial clustering method was employed using the MIDAS statistical package (Fox and Guire 1976).

Ward's method of minimum variance was used with a Euclidean distance measure. Cophenetic correlation was calculated for each cluster analysis. This measure of "best fit" was developed by Sokal and Rohlf (1962) and relates original distances (from the secondary distance matrix) to distances (cophenetic values) indicated by the cluster dendrogram. This product-moment correlation coefficient tends to vary between 0.6 and 0.95 depending on the clustering technique and data structure, with high values implying that the dendrogram is a reasonable illustration of sample affinities based on species distributions and abundances (Sneath and Sokal 1973).

Clustering techniques have been reviewed by Sneath and

Sokal (1973) and Everitt (1974), and the application of these methods occurs frequently within the literature. Clustering methods will tend to find groupings within a data set, regardless of the structure (Orloci 1975, Gauch and Whittaker

1981) and the comparison of clustering results with those from ordination is advised as groupings are not so rigidly defined in the latter. Recognition of sample groups can be facilitated if 33

the dendrogram is examined for large changes between fusions

(Everitt 1974).

Ward's method of minimum variance maximizes the variance between classes and minimizes that between them at each computational step, an optimal feature in a classification technique (Goodall 1978). Although groups or differences tend to be greatly emphasized with this method (Everitt 1974), consistently useful and interpretable results are found

(Pritchard and Anderson 1971). This algorithm gives results that are consistent with an analysis of Euclidean distances between samples in a data set (Orloci 1975).

Point-line Sampling

A plotless line sampling system described by Pielou (1967) and Stowe and Wade (1979) was used to detect the presence or absence of small scale associations between plant species.

Quadrats are generally not suitable to test for this scale of association, as results will depend on the size of quadrat used.

With relatively large quadrats, for example, spatially separated species appear positively associated, and, as quadrats approach the size of individual plants, associations become negative due to species exclusions (Greig-Smith 1964). Plotless techniques, using points rather than quadrats, therefore are more appropriate.

Three 2 m long lines of equidistant points (every 2 cm) were placed 15 cm apart at each 1 m interval along each transect

(Fig. 3). A fourth line was placed within this area if >25% of 34

any of the three lines contacted rock. The species occurring nearest to each sampling point, but within a radius of 1 cm from this point, was recorded. Any portion of the shoot constituted an occurrence and mosses.and lichens were also recorded. If no species occurred within 1 cm, then a blank was recorded and and treated as a species in the subsequent analysis. Vegetation of low stature, such as grassland, alpine, and bryophyte communities, are best suited to this sampling method, especially if there are extensive bare patches within the vegetation.

Approximately 9000 points were sampled per transect. The

2 cm interval was chosen because the average diameter of most of the vascular plant species present was estimated to be slightly greater than this. Each successive point at this scale was found to contact either the same individual plant contacted previously or the closest different individual. There are no statistical limits on the choice of a distance between points, however, if the distance is too small, inefficient sampling may result, with many blanks recorded or the same individual plant recorded many times. If the distance is too large, the species recorded will not necessarily be adjacent.

Analysis Of Point-line Data

Lines of point data were grouped within each 2 m X 30 m transect according to the vegetation types.and transition zones suggested by multivariate analysis of quadrat data. Homogeneity of grouped lines is required to satisfy the statistical assumptions of this analysis. The transition from the last 35

species of one line to the first species of the next is necessarily ignored during analysis. The chains of species occurrences were collapsed so that sequential occurrences of the same species became a single record. This process limits the results to interspecific associations only. At the present time, no adequate method exists to sample for intraspecific associations, primarily because of the extreme difficulty in determining individuals, particularly in species with extensive vegetative growth.

The algorithm for analysis of point-line data was originally proposed by Pielou (1967), and was subsequently used by Stowe and Wade (1979), where it was termed the species juxtapositions method. A species juxtaposition is, simply, the side by side occurrence of two different species, or a transition between them. This algorithm has since been restructured by de Jong and Greig (1983), because Pielou's

(1967) method is not correct for all possible cases. As well, the model of randomness is difficult to interpret, and the tests for randomness are less than rigorous and do not allow for the essentially directionless nature of point-line data. The new algorithm (de Jong and Greig 1983) is based on a first order

Markov chain model, as is Pielou's (1967) method. The sequence of species in a Markov chain is random - the transition from one species (e.g., 'A') to any different species has the same probability, provided the species are present in equal proportions. Successive observations are dependent only in that the transition from 'A' must be to a species other than 'A' - no 36

two adjacent species are the same in a Markov sequence. The

Markov sequence becomes the expected values for the analysis, with probabilities dependent on the relative abundance of each species in the data matrix. For example, two common species are expected to follow each other more often than would two rare species. The actual calculation of expected values begins with the number of times each species is involved in a transition and uses an iterative procedure (de Jong and Greig 1983) to arrive at the expected number of transitions between each species-pair.

As the analysis is less accurate when very low observed values are present, rare species (recorded less than 6 times in the data set) were grouped into a miscellaneous category before the calculation of expected values. This cut-off point was suggested by Pielou (1967). The matrix of expected values is then compared to the observed transition frequencies in a goodness of fit or chi-square test. Significantly large chi- square values (p<0.05) indicate departures from the Markov model, and, therefore, significant non-random structure within the data set. This analysis differs from that proposed by

Pielou (1967) in that symmetrically opposed elements in both matrices (observed and expected) are combined (the matrix is folded) before the chi-square analysis. This removes the directional aspect of the data. For example, transitions from species 'A' to species 'B' are grouped with transitions from 'B' to 'A', as no biological distinction can be made between the two.

The analysis proposed by Pielou (1967) determined non- 37

randomness of the entire data set only. Stowe and Wade (1979) extended the analysis of significantly non-random data sets with the application of a matrix-residual, to test for deviations from randomness for each species-pair. The appropriate standardized residual for folded matrices is given by de Jong and Greig (1983). This statistic is normally distributed and may be compared to the standard normal table to determine if significant (p<0.05) differences exist between observed and expected values for each species-pair. A positive residual value indicates positive association between two species, and a negative residual value indicates negative association.

Positive and negative associations mean that two species tend to occur together more often, or less often, respectively, than would be expected if their distribution were random. Species with both observed and expected values of less than 5 interspecific transitions were not considered in the presentation and interpretation of results, so as to eliminate significant associations with no biological or practical meaning. For example, a spurious, though highly significant, positive association would be detected between two species with an observed value of 1 and and expected value of 0.0001.

This plotless point-line or species juxtapositions method is allied to the contact sampling method developed by Yarranton

(1966) for bryophyte/lichen communities and subsequently used in pastures (Turkington et al. 1977, Aarssen et al. 1979,

Turkington and Harper I979a,b, Aarssen 1983). The method of calculating the chi-square statistic is these studies was 38

modified by de Jong et al. (1983). Coincidentally, the analysis and methods of calculation used in this thesis (de Jong and

Greig 1983) are very similar to the method proposed by de Jong et al. (1983), although both the method of data collection, models for randomness, and subsequent interpretation are different. Yarranton's (1966) method was not used in this study because it depends on a relatively continuous ground cover - a criterion not often met in alpine areas, due to extensive patterned ground (e.g., stone circles, rock rivers, frost boils, etc.)

Phenology

Phenological data were recorded during the 1980 growing season for vascular plant species at all transect sites. Plots

(2 m X 2 m) were established in each subjectively determined vegetation type in June, 1980. When all transects were completely snowfree (June 18, 1980), the times of vegetative growth, flowering, fruiting, seed dispersal, and dormancy stages were recorded at weekly intervals until September 4, 1980. At this point, all species were in late fruiting, seed dispersal, or dormancy stages. Stages were recorded when at least 50% of a species population (estimated visually) had reached that point. 39

Nomenclature

Nomenclature and taxonomy follows, with some exceptions,

Hitchcock and Cronquist (1973) for vascular plants, Lawton

(1971) for mosses, and Hale and Culberson (1970) for lichens.

The use of Lupinus lyalli i A. Gray follows Dunn and Gillett

(1966), Oxytropis monticola Gray ssp. monticola follows Elisens and Packer (1980), and Draba cana Rydberg follows Mulligan

(1971). Thamnolina vermicular is (Sw.) Schaer. has been included with T. subuliformis (Ehrh.) W. Culb. as they are not distinguishable in the field, requiring chemical tests for positive identification. In the text, only the binomial is used, omitting the variety or subspecies, if the species has only one variant in the study area. Voucher specimens have been deposited in the herbarium of the University of British

Columbia.

SOILS

Soil pits were established in June 1980 within each subjectively determined vegetation type in each transect. A and

B horizons were described in each profile to minimum depths of

30 cm. Composite samples were collected from both A and B horizons and laboratory analysis conducted on the fine (<2 mm) fraction (Table III). Soil color was described using Munsell color charts with moist and dry soil in natural light. 40

Table III - Methodology for Physical and Chemical Soil Analyses

ANALYSIS METHOD OR TECHNIQUE REFERENCE

Texture Hydrometer Method Bouyoucous (1951)

pH pH Meter-Glass Electrode Type 29b Bates (1954)

Total Carbon Leco Induction Furnace (Model 521) Described in & % Organic and Leco Carbon Analyzer (572-200) Black (1965) Matter

Total Cation Ammonium Acetate Method (pH 7.0)- Schollenberger Exchange Technicon Autoanalyzer II and Simon (NH4+-N) (1945)

Exchangeable Ammonium Acetate Method (pH 7.0)- Schollenberger Cations Perkin-Elmer Atomic Absorption and Simon (K, Ca, Mg) Spectrophotometer (KC1 extraction) (1945)

Total Macrokjeldahl Method -Acid Kjeldahl Nitrogen Digestion. Colorimetric Analysis - (1883), Technicon Autoanalyzer II Bremner (1960)

Available NH4F in HC1 Extraction Bray and Kurtz Phosphorus Colorimetric Determination with (1945) Spectrophotometer 41

CLIMATE

The climate of Lakeview Mountain in summer (1980) was

monitored at three stations with aspects and elevations

corresponding to Transects B, C, and D. Temperature and

atmospheric moisture were recorded with Schreibstreifen R. Fuess

hygrothermographs placed in white Steveson screens (louvered

boxes) with double tops, allowing air to circulate, and

minimizing surface heating. Sensors were between 18 and 25 cm

above the ground. Precipitation was collected with a Tru-check

raingauge set 60 cm above the ground surface. Windspeed was

measured with Belfort 3-cup totalizing anemometers set 60 cm

above the ground. Station 1 was located S84°W at 2481 m, above

Transect B, with a hygrothermograph, anemometer, and raingauge.

Station 2 was located S2°E at 2402 m, below Transect E, with a

hygrothermograph and anemometer. Station 3 was situated above

Transect C at an aspect of N29°E and an elevation of 2475 m,

with a hygrothermograph only.

Microclimate was recorded at each soil pit site in each

transect (a total of 14 sites) at weekly intervals during the

summer of 1980. Soil temperatures were measured with laboratory

calibrated Varah FD 300 diodes sealed with silicon. Diodes were

placed at depths of 2, 10, and 20 cm below the surface in

relatively undisturbed soil approximately 10 cm from the pit edge (Fig. 2). Resistances were measured with a battery powered

6.75 V bridge-meter. Air temperature and humidity were monitored with a battery powered fan psychrometer at heights of

2, 10, and 20 cm above ground. Soil moisture was determined for 42

June 25 - July 1, 1980, and September 6-7, 1980 by gravimetric samples taken at depths of 5-10 cm and 20-30 cm at each pit site. Percent water was obtained after drying soils at 110° C for 24 hours. 43

IV. RESULTS

VEGETATION

Community Types

1. Transect A;

All data from the 90 quadrats in transect A were used to form thirty composite samples or strata (Appendix C). The 13 species having less than 5 occurrences in these composite samples were removed, leaving 43 species (variables) for analysis. The data matrix is characterized by fairly consistent high cover values for Carex sc irpoidea (mean 28%), Carex capitata (mean 28%), and Cetraria islandica (mean 24%), with

Potentilla diversifolia and Cetraria nivalis both averaging 15%

(Appendix C). Total species cover in samples ranged from 170-

293% (mean 220%), with the highest total cover found in samples

20-30 (262%).

Multivariate Analyses:

Analysis by centered principal components analysis (PCA), centered and standardized PCA, and reciprocal averaging (RA)

(Fig. 4), accounted for 47%, 29%, and 35% of sample variation in the first two axes, respectively. Axis 3 accounts for only a small amount of additional variance. All three analyses and axes combinations indicate little structure within this data set. Samples 20-30, however, illustrate affinity (possibly due to the presence of Carex nardina), as do anomalous samples 1 and

17-19 which contain less than 12% Carex capitata. Samples 6 and 44

Figure 4 - Transect A Multivariate Analyses

CENTERED PRINCIPAL COMPONENTS ANALYSIS CENTERED AND STANDARDIZED PRINCIPAL COMPONENTS ANALYSIS

.17

. 13 U 15 .16

• 21

23* .18 •20 26*

.11 ,5 *9 .19 •27

.1*

RECIPROCAL AVERAGING MINIMUM VARIANCE CLUSTER ANALYSIS USING EUCLIDEAN DISTANCE

2. . 5 *

28%. -22 29

f*\ \C S >J\ \D O — rsj u** t£ _^r*» CT* CO O 45

10 occur at a large distance from other sample points in the RA scatter plot - both have a high (30%) cover of Polytrichum piliferum, which averages 2% in the remaining samples. •

A cluster analysis using minimum variance or Ward's Method in conjunction with Euclidean distance is illustrated in Fig. 3.

Distances drawn between fusions in this and subsequent cluster dendrograms are proportional to the actual Euclidean distances.

The cluster of samples 20-30 occurs at a relatively large distance from the final fusion of all samples in Fig. 3.

Clusters within the remaining samples are less distinct.

Cophenetic correlation relates original Euclidean distances to those resulting from dendrogram construction, where distances are recalculated at each step. The value for this cluster analysis is relatively low at .4966. High values (e.g., >.6) imply that the dendrogram is a reasonable illustration of sample affinities.

Vegetation "Groups:

Little structure is apparent in this data set, although the cover of dominant species varies considerably in some samples.

Analyses indicate, however, that this variability does not warrent data set division. The entire data set is classed as one community type, dominated by Carex capitata and Carex scirpoidea. Thus, point-line data for this transect will not be sub-divided before association analysis. 46

2. Transect B:

Thirty composite samples, formed from a total of 90 original quadrats, (Appendix C) were used for multivariate analysis. Thirty-nine species were used after the 6 species with less than 5 occurrences in these samples were removed. The data matrix is characterized by uniformly high cover values for both Carex scirpoidea (mean 26%) and Carex capitata (mean 40%)

(Appendix C). Cetraria islandica averages 22% cover, and

Cetraria nivalis, 10%.

Multivariate Analyses:

Analysis by centered PCA, centered and standardized PCA, and RA (Fig. 5), accounted for 48%, 34%, and 37% of sample variation in the first two axes, respectively. Sample points form no distinct groups with the first three axes of these ordinations. Sample 1 occupies an outlying position, possibly due to the absence of Cetraria cucullata, a low value for

Potentilla diversifolia (9%), and relatively high cover of

Cladonia sp. (18%). RA produced a number of outliers, which include samples 6, 8, and 23, perhaps due to their relatively high cover values for Silene acaulis.

The dendrogram resulting from cluster analysis (Fig. 5) shows a small cluster of samples 14 and 22-27 a relatively large distance from the final fusion. The remaining samples fuse close to this final linkage. The cophenetic correlation is low at .5010, implying that some distortion of sample affinities has occurred during dendrogram construction. 47

Figure 5 - Transect B Multivariate Analyses

CENTERED PRINCIPAL COMPONENTS ANALYSIS CENTERED AND STANDARDIZED PRINCIPAL COMPONENTS ANALYSIS

29*

17 *27

21 26 *20 18 '1 .16 • 2,« • If 3 28 30 13 •18 14. "2

'9* 2}

•25

RECIPROCAL AVERAGING MINIMUM VARIANCE CLUSTER ANALYSIS USING EUCLIDEAN DISTANCE

29.

30. II 19

* * » 15 21 «26 • • • 28 13 20 27 • •« 12

• 3

:mnrnIr^n[iii^^ tt — c^< rn-yr* — O — CO O "\ ••D U~\ o J\ CTNtnrv. \o

Vegetation Groups:

The majority of samples within Transect B appear very similar, but irregularities, such as low Cetraria cucullata in samples 1-4, high C. cucullata in sample 17, and high Thamnolina vermicular is and Polytrichum piliferum in sample 12, occur throughout the :data set (Appendix C). These outlying samples have little affinity with each other and do not constitute a separate vegetation group. The entire transect is proposed as one community type, dominated by Carex capitata and Carex scirpoidea. Point-line data for this transect will not be sub• divided before association analysis.

3. Transect C:

The composite data matrix of 30 samples (Appendix C) was formed from an original 88 quadrats; two quadrats with >49% rock were omitted. Forty-four species were used for multivariate analysis, after the removal of 4 infrequent species (<5 occurrences). High cover values for both Carex scirpoidea

(range 8-45%; mean 26%) and Carex capitata (range 5-60%; mean

30%) dominate this data matrix. A distinguishing feature is the presence of Salix nivalis in 12 composite samples (2-8, 13-15, and 21-22), with percentage cover ranging from 8-48%, with a mean of 25%. On average, these samples are also characterized by low cover values for both Carex scirpoidea and Carex capitata, which range at higher values of 22-60% and 20-53%, respectively, in the remaining samples. 49

Multivariate Analyses:

The application of a centered PCA, a centered and standardized PCA, and RA (Fig. 6), resulted in 64%, 29%, and 46% of sample variation explained by the first two axes, respectively. The affinity of samples 2-8, 13, 14, and 21-22

(containing Salix nivalis), is illustrated with axes 1 and 2.

Sample 14 (a lower amount of Salix nivalis) is often positioned midway between this group and the remaining samples. Sample 15

(containing 13% Salix nivalis) occurs closest to the samples dominated solely by Carex capitata and Carex sc i rpoidea, possibly due to its 47% cover of Carex capitata. The presence of Penstemon procerus in samples 11, 16, 20, and 25 (25% in sample 25), likely accounts for their outlying positions in the

RA scatter plot. Scatter plots of the first and third axes were very similar to this, although sample affinities were not illustrated with axes 2 and 3 - possibly due to the small amount of variation accounted for.

Cluster analysis produced two major clusters fusing a large distance from the final cluster of all samples (Fig. 6).

Samples containing Salix nivalis (2-8, 13-14, and 21-22) form one of these distinct groups, with sample 15 included with the remaining data points. The cophenetic correlation is .6330, implying that the dendrogram is a reasonable illustration of sample affinities.

Vegetation Groups:

On the basis of these analyses, it is proposed that transect C be grouped as follows: 50

Figure 6 - Transect C Multivariate Analyses

CENTERED PRINCIPAL COMPONENTS ANALYSIS CENTERED AND STANDARDIZED PRINCIPAL COMPONENTS ANALYSIS

.29

.13 . '9 23.

22 *. 18 ^ 1•8 i

19 .25 • 27 3 * 16 . 27« .23 .20

. 2

26 *10

.26

RECIPROCAL AVERAGING MINIMUM VARIANCE CLUSTER ANALYSIS USING EUCLIDEAN DISTANCE

2315 29".V.30

27 io* •

3 6*-7 •»

• 20 51

1) Samples 2-8, 13, 14, and 21-22, characterized by medium to high cover (mean 26%) of Salix nivalis, with correspondingly lower values for Carex scirpoidea and Carex capitata.

2) Samples 1, 9-12, 15-20, and 23-30, characterized by relatively high cover (>17%) of both Carex scirpoidea and Carex capitata.

Point-line data will therefore be analyzed for associations separately within these groups.

4. Transect D:

A composite data matrix of 30 samples (Appendix C) was formed from 74 of the original quadrats; as sixteen quadrats with >50% rock were removed. Forty-three species were used for analysis, after the 6 species having less than 5 occurrences were deleted. The transect is characterized by a high proportion of both rock (3-45%; mean 12%), and bare or disturbed ground (5-30%; mean 8%), in the quadrats used for analysis.

Cover values' for vascular plants and cryptogams are relatively low as a result, with total cover of species in composite samples ranging from 110-130%, compared to an average total plant cover of 220% in Transect A samples. Carex scirpoidea occurred in all samples with a range of 5-25% cover (mean 17%), as did Arenaria obtusiloba (mean 12%), and Potentilla diversifolia (mean 8%). The lichens, Cetraria nivalis and

Cetraria islandica, occurred in all samples with average cover of 5% and 4%, respectively. 52

Multivariate Analyses:

Analysis by centered PCA, centered and standardized PCA,

and RA, all illustrate two major groups in this data set with

axes 1 and 2 (Fig. 7) and with axes 1 and 3, with a smaller,

intermediate group apparent with centered PCA and RA. Axes 2

and 3 do not illustrate these groups, however, possibly because

of the small amount of total variation they explain. The first

two axes of each ordination account for 65%, 26%, and 36% of

sample variation, respectively. Samples 9, 20-21, 23, and 26-29

form a group characterized by high values for Kobresia

myosuroides (25-38%) and correspondingly low values for Carex

scirpoidea (7-12%; mean 8%). Samples 1-7, 11-18, and 25 form a

second group dominated by Carex scirpoidea (13-30%; mean 21%),

with virtually no Kobresia myosuroides present. The remaining

samples contain intermediate values for both Carex scirpoidea

(mean 15%), and Kobresia myosuroides (mean 13%), a group which

includes sample 23 ("high Kobresia") in the centered and

standardized PCA and RA scatter plots. Sample 7 is outlying in

the RA scatter diagram, possibly due to the relatively high

cover of Carex nardina•

The cluster analysis dendrogram groups "high Kobresia

myosuroides" samples in a distinct cluster a large distance from

the final fusion. The cluster of remaining samples has two

major divisions which separate samples with intermediate values

for Carex scirpoidea and Kobresia myosuroides (8, 10, 19, 22,

24, and 30) from samples dominated solely by Carex scirpoidea.

The cophenetic correlation is .8144, implying that the 53

Figure 7 - Transect D Multivariate Analyses

CENTERED PRINCIPAL COMPONENTS ANALYSIS CENTERED AND STANDARDIZED PRINCIPAL COMPONENTS ANALYSIS

*23

• 16

•17 •17

.12

27* •,„ 2/

L5 .13

5 . •*

"28

. . 10

RECIPROCAL AVERAGING MINIMUM VARIANCE CLUSTER ANALYSIS USING EUCLIDEAN DISTANCE

•9

•29

•2*

•15 •23

•19

18 co j- p-j o o en o • 54

dendrogram is a good illustration of sample affinites.

Vegetation Groups:

Sample groupings for Transect D are as follows:

1) Samples 9, 20, 21, 23, and 26-29, dominated by Kobresia

myosuroides.

2) Samples 8, 10, 19, 22, 24, and 30, characterized by

intermediate values for both Kobresia myosuroides and Carex

scirpoidea, and appearing to form a "transition zone" to either

side of samples containing extremely high cover values for

Kobresia myosuroides (Appendix C).

3) Samples 1-7, 11-18, and 25, dominated by Carex scirpoidea.

Point-line data will be sub-divided into these three groups

before association analysis.

5. Transect E:

Eighty original quadrats were used to form a composite data

matrix of 30 samples (Appendix C), as 10 quadrats had 50% or

more rock. Multivariate analyses were applied to 47 species out

of a total of 54, as 7 species occurred in less than 5 composite

samples. The transect is characterized by high cover values for

Kobresia myosuroides in samples 1-11 (range 13-58%; mean 40%),

with this species virtually absent from the remaining samples.

Carex scirpoidea occurs in all samples with cover values ranging

from 6-48% (mean 28%). Cetraria nivalis (6-30% cover), Cetraria

cucullata (5-20%), Cornicularia aculeata (5-20%), and Arenaria

obtusiloba (5-20%) are similarly ubiquitous.

Multivariate analyses: 55

A centered PCA was applied to the data set, and the resulting first two component axes accounted for 77% of sample variation. The first two axes of a centered and standardized

PCA, and a RA accounted for 32% and 52% of sample variation, respectively (Fig. 8). Each of these analyses (including plots of axes 1 and 3) indicated two large, distinct groups of samples

1-11 (dominated by Kobresia myosuroides) and samples 12-30

(dominated by Carex scirpoidea [mean 36% cover]). These groups were not clear with axes 2 and 3. Low values for Carex scirpoidea (mean 16%) occur in the 1-11 group, as well as consistently high values for Cetraria cucullata (mean 14%), compared to the remaining samples (mean 5%). Samples 19 and 26 are outlying in the centered PCA and RA scatter plots (30% and

22% cover of Silene acaulis, respectively), as is sample 1 (12%

Carex capitata, a species otherwise absent or present only in trace amounts).

Cluster analysis resulted in two groups of samples 1-11 and samples 12-30, both a relatively large distance from the final fusion of all samples. Within the 12-30 sample cluster, a subset of samples 13, 16, 19, 21, 23, and 26 appears, possibly due to their relatively high Silene acaulis component. The cophenetic correlation for this dendrogram is .7646, implying a fairly good illustration of sample affinities.

Vegetation Groups:

Sample groups for Transect E are as follows:

1) Samples 1-11, characterized by high cover of Kobresia myosuroides and correspondingly low cover of Carex scirpoidea. 56

Figure 8 - Transect E Multivariate Analyses

CENTERED PRINCIPAL COMPONENTS ANALYSIS CENTERED AND STANDARDIZED PRINCIPAL COMPONENTS ANALYSIS

.23

23. .18

*27

*3 *9

' 12

2* '3 •16 18. 25. 27. 2f to 15 30

, 29 .22

RECIPROCAL AVERAGING MINIMUM VARIANCE CLUSTER ANALYSIS USING EUCLIDEAN DISTANCE

. 16 .9

24 .18

27 15 . 22

28* 10 ' 17 29. -47 57

2) Samples 12-30, characterized by high cover of Carex scirpoidea, with little or no Kobresia myosuroides.

Corresponding point-line data will be sub-divided into these two groups before association analysis.

6. Transect F;

Thirty composite samples were formed from the complete original data matrix of 90 quadrats (Appendix C). Forty-three species were used for multivariate analyses after the 3 species with less than 5 occurrences in the composite samples were deleted. The data set is characterized by high cover values for

Kobresia myosuroides (mean 49%) in samples 1-4, and 6-7, with lower amounts in samples 5, 8, and 9 (mean 18%). The remaining samples average 32% Carex capitata, with no Kobresia myosuroides. Carex scirpoidea is a variable dominant throughout the data set, ranging from 5-43% cover and averaging 24%.

Multivariate Analyses:

Analysis by centered PCA, centered and standardized PCA, and RA

(Fig. 9) resulted in the first two axes accounting for 68%, 31%, and 47% of sample variation, respectively. The variablility of samples is indicated, particularly in the standardized PCA scatter diagram. Samples 1-4, 6, and 7 show affinity in all three analyses. Samples 5, 8, and 9 (averaging 20% Carex capitata, 24% Carex scirpoidea, and 18% Kobresia myosuroides) occur in a somewhat intermediate position between this group and the remaining sample points in the centered PCA and RA scatter plots, with samples 25 and 26 included with these intermediates 58

Figure 9 - Transect F Multivariate Analyses

CENTERED PRINCIPAL COMPONENTS ANALYSIS CENTERED AND STANDARDIZED PRINCIPAL COMPONENTS ANALYSIS

. 30

"28

23* »9

29. ,, 22 • . *2' . 18

. 26

.8 *15

• 9

, 16

*30

•24

' 7

• 26 .2 5 .6

MINIMUM VARIANCE CLUSTER ANALYSIS USING EUCLIDEAN DISTANCE RECIPROCAL AVERAGING

*30

28* 27 \

• III

. '5 59

in the centered PCA scatter plot of axes 1 and 3. Samples 25

and 26 are outlying in most ordinations, and have an average 42%

cover of Carex scirpoidea with no Carex capitata or Kobresia

myosuroides present. Sample 15 has high Carex capitata (57%)

and low Carex scirpoidea (6%). The above affinities were not

illustrated with axes 2 and 3 in any of the analyses.

Cluster analysis produced a distinct group of samples 1-4,

and 6-7, a large distance from the final fusion of all samples.

Samples 5, 8, and 9 group with the remaining sample points,

which also fuse a large distance from the final linkage. The

cophenetic correlation is .8163, indicating that the dendrogram

is a good illustration of sample similarities and differences.

Vegetation Groups:

Sample groups within Transect F:

1) Samples 1-4, 6, and 7, dominated by Kobresia myosuroides, and

averaging 16% Carex scirpoidea, with little or no Carex capitata.

2) Samples 5, and 8-30, with an average of 30% Carex capitata and 27% Carex scirpoidea. Kobresia myosuroides is absent in all

samples except 5, 8, and 9.

Point-line data for these two vegetation groups will therefore be analyzed for associations separately. 60

7. All Transects:

Composite samples from all six transects were compiled into one data set of 180 cases for additional multivariate analyses, to compare different stands of each community and to determine overall relationships between sampled areas. Sixty-three species were used after deletion of 5 species which occurred in less than 5 of the 180 composite samples. Correspondence of sample points to original data sets in Figures 10-13 is indicated by symbols.

Multivariate Analyses:

Analysis with centered PCA resulted in 56% of sample variation accounted for by the first two axes (Fig. 10), while centered and standardized PCA and RA accounted for only 22% and

30%, respectively (Figs. 11, 12). The affinity of samples dominated by Carex capitata and Carex scirpoidea (data sets A,

B, C2, and F2) is apparent with the first three axes of all three ordinations, with no samples grouped according to their original data matrix. Group C1, with the additional dominant,

Salix nivalis, has the majority of its samples clustered together in the two PCA scatter plots, but a distinct group of all these points occurs only with RA. Samples with high values for Kobresia myosuroides (data sets D1, E1, and F1) appear together in all ordinations. Samples co-dominated by Carex scirpoidea and Kobresia myosuroides (data set D2) occurred in an intermediate position between this group and samples dominated solely by Carex scirpoidea (data sets D3 and E2) in the centered

PCA and RA scatter plots. Separation of Carex scirpoidea 61

Figures 10-13 - Ordination and Cluster Analysis of 11 Data Sets. Data sets are indicated by the following symbols in these figures: Data set A: • ; B: o ; C1: 0 ; C2: • ; D1: v ; D2: 4 • D3: • ; El: + ; E2: t ; F1: * ; and F2: • . 62

Figure 10 - Centered PCA - All Data Sets

AXIS 1 63

Figure 11 - Centered and Standardized PCA - All Data Sets

• • • • • • • * + • •

• v V o° • R?l • • V V ,4! ' o *o * o ° + + ° o • • m T o •• mmo 0 v • *»• • o • 00

AXIS 1 64

Figure 12 - RA - All Data Sets

AXIS 1 65

dominated sample points from data sets D3 and E2 is apparent with all analyses (particularly RA), possibly because of the appreciable Cornicularia aculeata and Silene acaulis components and higher total cover found in E2 (Appendices 1D and 1E).

Cluster analysis (Fig. 13), corroborates sample affinities illustrated by ordination, with four major groups apparent. The two groups to the left comprise Carex capitata/Carex scirpoidea dominated samples with the majority of samples containing Salix nivalis forming a sub-cluster. Samples with high values for

Kobresia myosuroides cluster to the right, a very large distance from the final fusion of all samples, with sub-groups correlating to the original data sets. The remaining samples are contained within a cluster a large distance from the cluster of samples high in Kobresia myosuroides, with further division separating different Carex scirpoidea dominated data sets (D3 and E2). Intermediate Kobresia myosuroides/Carex scirpoidea dominated samples (with 1 "high Kobresia myosuroides" sample) form a small cluster within the larger grouping containing data set D3. The cophenetic correlation is relatively low at .5736, implying that some distortion of sample affinites has likely occurred during dendrogram construction.

Vegetation Groups:

Analysis of compiled transect data sets supports the validity of vegetation groups determined by individual transect analysis. Similarity between Carex capitata/Carex scirpoidea dominated data sets is illustrated, as well as between site differences in Carex scirpoidea dominated and Kobresia FIGURE 13 - CLUSTER ANALYSIS - ALL DATA SETS

H39 67

myosuroides dominated vegetation.

Phenology

Phenological stages for a total of 45 vascular plant species were recorded during the summer of 1980, within permanent 2 m square plots in each sampled transect. Data corresponding to the 11 vegetation groups previously outlined are presented in Figs. 14-19. As phenology was, on average, recorded weekly, differences of less than 10 days in these figures will not be discussed. The timing of phenological phases was quite different between species within each vegetation group. Although intraspecific differences were observed in vegetation sampled at different aspects, the duration of each phenological phase was comparatively consistent within a species. Snow release for all transects took place between June 16 and 18 in 1980. Most species flowered 15-40 days after snow release - all except Carex capitata had assumed at least vegetative growth at this time. In general, flowering occurred for 18-40 days and fruiting for 10-30 days before seed dispersal. Consistent exceptions to this general trend include the late development of Sedum lanceolatum, Potentilla fruticosa, and Solidago multiradiata (mid to late July), and the early floral maturation of Draba incerta, Draba paysoni i, and Draba lonchocarpa, which flowered immediately after snow release. In addition, Taraxacum ceratophorum has a relatively short flowering period ranging from 8-14 days and Dryas octopetala and

Agoseris glauca flower for 10 days only. The fruiting period 68

Figures 14-19 - Phenology of vascular species within 11 vegetation groups from June 20 to Sept. 10, 1980. Solid bars indicate the flowering period, and open bars containing D, V, F, and S indicate dormant, vegetative, fruiting, and seed dispersal stages, respectively. Vegetation was completely snowfree by JUne 20, 1980. 69

Figure 14 - Transect A Phenology

ANDROSACE SEPTENTRI ONAI IS ANTENNARIA ALP INA ARENARIA OBTUSILOBA CAREX CAPITATA CAREX NARDINA CAREX SCIRPOIDEA CERASTIUM BEER INGIANUM DRABA INCERTA DRABA PAYSONII ERIGERON AUREUS FESTUCA OVINA HAPLOPAPPUS LYALLII LUPINUS LYALLII LUZULA CAMPESTRIS OXYTROPIS MONT I COLA PENSTEMON PROCERUS POA RUPESTRIS POLEMONIUM PULCHERRIMUM POLYGONUM VIVIPARUM POTENTILLA DIVERS IFOLIA POTENTILLA NIVEA SALIX NIVALIS SEDUM LANCEOLATUM SENECIO LUGENS SILENE ACAULIS SOL I DAGO MULT I RAD I ATA STELLARIA LONGIPES TRISETUM SPICATUM

Figure 15 - Transect B Phenology

10/8 20/8 30/8 10/S 70

Figure 16 - Transect C Phenology

GROUP C1

GROUP C2

I Ll _ F IS s s F s i r •Pi S 20/6 30/6 10/7 20/7 30/7 10/8 20/8 30/8 10/9 71

Figure 17 - Transect D Phenology GROUP D1

GROUP D2 ANTENNARIA ALP INA ARENARIA OBTUSILOBA CALAMAGROSTIS PURPURASCENS CAREX SCIRPOIPEA CERASTIUM BEER INGIANUM DRABA CANA DRABA INCERTA DRABA PAYSONI I ERIGERON COMPOSITUS FESTUCA OVINA HAPLOPAPPUS LYALLI I KOBRESIA MYOSUROIDES OXYTROPIS MONT I COLA POTENTILLA DI VERS IFOL tA POTENTILLA NIVEA SEDUM LANCEOLATUM SILENE ACAULIS SOL I DAGO MULT I RAD I ATA TARAXACUM CERATOPHORUM TRISETUM SPICATUM

GROUP D3

20/6 30/6 10/7 20/7 30/7 30/8 10/9 72

Figure 18 - Transect E Phenology

GROUP

20/6 3o"/6 \U/7 20/7 30/7 10/8 20/8 30/8 10/9 73

Figure 19 - Transect F Phenology

GROUP F1

GROUP F2 ANDROSACE SEPTENTRIONALIS ANTENNARIA UMBRINELLA ARENARIA OBTUSILOBA CAREX CAPITATA CAREX NARDINA I CAREX PHAEOCEPHALA CAREX SCIRPOIDEA I

CERASTIUM BEER INGIANUM I DRABA INCERTA J DRABA PAYSONII I ERIGERON AUREUS FESTUCA OVINA HAPLOPAPPUS LYALLII LUPINUS LYALLII LUZULA CAMPESTRIS OXYTROPIS MONT I COLA PENSTEMON PROCERUS POA SP. POTENTILLA DIVERSIFOLIA POTENTILLA NIVEA SEDUh LANCEOLATUM SENECIO LUGENS SILENE ACAULIS SOL I DAGO MULTIRAD I ATA STELLARIA LONGIPES TRISETUM SPICATUM —i ' r 20/6 30/6 10/7 20/7 30/7 10/8 20/8 30/8 10/9 74

for Taraxacum ceratophorum is 6-10 days, and the complete fruiting period for Sedum lanceolatum lasted less than 6 days when observed. Major phenological differences between vegetation groups at a constant aspect and between the same community types at different aspects will be outlined in the following sections.

1. Constant Aspect;

Transect C (N29°E):

The phenology of vascular species in groups C1 and C2 is essentially the same, with only two species differing by 10 days or more. Poa alpina flowers 16 days earlier, and Carex nardina

10 days later, in C1 than in C2. Flowering periods for these two species terminate at the same time in both groups.

Transect D (S2°E):

Three species have major phenological differences within this transect. Arenaria obtusiloba and Oxytropis monticola flower an average of 15 days earlier in group D3. Kobresia myosuroides flowers 14 days longer in group D2 than in group D1, with fruiting beginning 10 days later as well.

Transect E (N16°W):

The majority of species within this transect have similar phenological timing. A few species, however, tend to flower earlier in group E2 than in group E1. Senecio lugens,

Haplopappus lyallii, • and Carex scirpoidea flower 10 days 75

earlier, although the duration of flowering is the same in both

groups. Trisetum spicatum flowers 16 days earlier and Cerastium

beeringianum 20 days earlier in E2.

Transect F (S56°E): • '

Many species have similar phenological timing within this

transect, although several species flower later in group F1 than

in group F2. Carex nardina, Carex scirpoidea, Cerastium

beeringianum, and Potentilla diversifolia flower 10-16 days

later in F1 .

2. Constant Community Type:

Carex capitata/Carex scirpoidea Community Type (Groups A, B, C2,

F2):

Species generally begin periods of flowering and fruiting

sooner in group F2 (S56°E) than in groups A (N58°W), B (S80°W),

and C2 (N29°E). For example, Potentilla diversifolia,

Potentilla nivea, Oxytropis monticola, Haplopappus lyallii,

Erigeron aureus, and Androsace septentrional is flower and/or

fruit 10-16 days earlier in F2 than in the other groups. A few

species flower 12-18 days later in C2 than in the remaining

groups (e.g., Cerastium beeringianum, Carex scirpoidea, and

Carex nardina). The duration of flowering is 16-20 days longer

for Stellaria longipes in this group.

Some exceptions to this general trend are evident.

Flowering occurs latest in group B (14-20 days) for Solidago multiradiata, Polemonium pulcherrimum, and Trisetum spicatum, 76

and latest in group A (10-12 days) for Penstemon procerus. Seed dispersal of Draba paysonii occurs 10 days earlier in north- facing groups (A and C2) than in south-facing groups (B and F2), while Silene acaulis begins seed dispersal 10 days earlier in F2 than in C2.

Carex scirpoidea Community Type (Groups D3 and E2):

In general, species begin phenological phases earlier in group D3 (S2°E) than in group E2 (N16°W), or at similar times in both groups. A number of species flower 10-20 days sooner in group D3: Festuca ovina, Draba paysoni i, Polemonium pulcherrimum, Oxytropis monticola, Silene acaulis, Taraxacum ceratophorum, Potentilla diversifolia, and Potentilla nivea. In addition, Taraxacum ceratophorum and Potentilla nivea fruit 10 days earlier and Silene acaulis and Erigeron compositus begin seed dispersal 10 days earlier in D3.

Kobresia myosuroides Community Type (Groups D1, E1, and F1):

Phenological stages tend to be initiated earliest in group

D1 (S2°E) and latest in group E1 (N16°W). For example, Trisetum spicatum flowers 10 days sooner in D1 than in F1 (S56°E), and 18 days earlier in D1 than in E1. Silene acaulis, Potentilla diversifolia, Potentilla nivea, and Kobresia myosuroides all flower 10-16 days earlier in D1 than in the remaining groups and

Sedum lanceolatum flowers 24 days later in E1 than in D1 and F1.

Cerastium beeringianum flowers 12 days sooner in Fl than E1 (it does not occur in D1). Carex scirpoidea is an exception to this 77

trend, flowering 12-15 days later in F1 than in E1 and D1.

Interspecific Associations

Positive and negative associations for 11 vegetation groups are given in Figs. 20-31. Species are indicated by three-letter codes in these figures and are listed in Table IV.

Table IV - Associated Species

AGL - Agoseris glauca (Pursh) Raf. AAL - Antennaria alpina (L.) Gaertn. AOB - Arenaria obtusiloba (Rydb.) Fern. CPU - Calamagrostis purpurascens R. Br. *CAL - Caloplaca sp. Th. Fr. *CAN - Candelariella sp. Mull. Arg. CCA - Carex capitata L. CNA - Carex nardina Fries CPH - Carex phaeocephala Piper CSC - Carex scirpoidea Michx. CBE - Cerastium beeringianum Cham. & Schlecht *CCU - Cetraria cucullata (Bell.) Ach. *CIS - Cetraria islandica (L.) Ach. *CNI - Cetraria niva1is~Tl.) Ach. *CMI - Cladina mitis (Sandst.) Hale & Culb. *CCH - Cladonia chlorophaea (Florke ex Somm.) Spreng. *CLA - Cladonia sp. Wigg. *CAC - Cornicularia aculeata (Schreb.) Ach. nDES - Desmatodon sp Br id. DIN - Draba incerta Pays. DLO - Draba lonchocarpa Rydb. DPA - Draba paysoni i Macbr. DOC - Dryas octopetala L. EAU - Erigeron aureus Greene ECO - Erigeron compositus Pursh FOV - Festuca ovina L. HLY - Haplopappus lyalli i Gray KMY - Kobresia myosuroides (Vill.) Fiori *LVU - Letharia vulpina (L.) Hue LLY - Lupinus lyallii A. Gray LCA - Luzula campestris (L.) DC. *OUP - Ochrolechia upsaliensis (L.) Mass. OMO - Oxytropis monticola Gray *PCA - Peltigera canina (L.) Willd. PPR - Penstemon procerus Dougl. PRU - Poa rupestris Vasey POA - Poa sp. L. 78

PPU - Polemonium pulcherrimum Hook. nPJU - Polytrichum juniperinum Hedw. nPPI - Polytrichum piliferum Hedw. PDI - Potentilla diversifolia Lehm. PFR - Potentilla fruticosa L. PNI - Potentilla nivea L. SNI - Salix nivalis Hook. SLA - Sedum lanceolatum Torr. SDE - Selaqinella densa Rydb. SLU - Senecio lugens Rich. SAC - Silene acaulis L. SMU - Solidaqo multiradiata Ait. SLO - Stellaria lonqipes Goldie TCE - Taraxacum ceratophorum (Ledeb.) DC. *TVE - Thamnolina vermicular is (Sw.) Ach. exSchaer , TSP - Trisetum spicatum (L.) Richter BLA - Bare ground

Lichen species in this list are identifiable by a *, and moss

species by a n. Of the species sampled, 88% had at least one

association in at least one vegetation group.

1 . Transect A:

Point-line data from the entire transect were used in

association analysis. This data set of 9285 points comprised 41

vascular, lichen, and moss species (species which had less than

5 occurrences were grouped into a miscellaneous category). A

total of 38 significant (p<0.05) positive associations and 31

significant (p<0.05) negative associations (Fig. 20) were detected, out of a possible 861 different species-pair

combinations. Carex capitata, one of two dominant vascular

species, is significantly positively associated with 3 lichens

and 2 vascular species, while co-dominant, Carex scirpoidea, is positively associated with 5 vascular plants. Carex capitata

and Carex scirpoidea are negatively (p<0.05) associated with 7 and 3 species, respectively. The lichen, Cetraria islandica, is 79

Figures 20-31 - Positive and negative associations between species pairs in 11 vegetation groups. Species are indicated by three-letter codes and are connected by , , or , indicating p<0.05, p<0.0l, and p<0.00l levels of significance, respectively. 80

Figure 20 - Group A Positive and Negative Associations

POSITIVE

SDE CCH

NEGATIVE L LY PD

T VE_"_'_ _ ecu CNI

P P

S LO

S N I CCH

OUP • AOB

OMO 81

negatively associated with 7 species (including 2 lichens) as well as bare ground.

2. Transect B:

The point-line data set of 9298 points was not subdivided before analysis of the 40 plant species. Forty-four significant positive and 27 significant negative associations (Fig. 21) occurred out of the 820 different species-pair combinations possible. Carex scirpoidea was positively associated with 5 vascular species and negatively associated with 2 lichens. Co- dominant, Carex capitata, was involved in 4 positive associations (3 with lichen species) and 7 negative associations

(6 vascular species and 1 moss). Other major species include

Arenaria obtusiloba (6 positive associations), Cetraria cucullata (5 negative associations), and Selaginella densa (5 negative associations).

3. Transect C:

Based on results of quadrat data analysis, point-line data were sub-divided into two groups before analysis. Group C1 was dominated by Salix nivalis, Carex scirpoidea, and Carex capitata. Group C2 was dominated by Carex scirpoidea and Carex capitata but had no Salix nivalis.

Group C1:

Data involving 3897 points and 44 species were used in association analysis. A total of 40 significant (p^0.05) positive and 18 significant negative associations (Fig. 22) were 82

Figure 21 - Group B Positive and Negative Associations

POSITIVE

••• AOB CNA . • •' t '• /

DES CBE CSC TSP ' PPR / \ / _ _ S AC T VE SMU / ^PDI \ A

ecu .. E AU PP I PCA C I S SDE

CCH 0 U P

CCA _____'C AC. CMI - CN I

H LY

NEGATIVE

CN I 83

Figure 22 - Group Cl Positive and Negative Associations

POSITIVE pD,

C N I DOC

CCA C I S E AU P PR C NA

ecu- TVE CSC LC A C AC OUP S LU SOE

A OB PP I "SLO B LA C LA OMO X^ L LY

PCA AAL. .': S N I D P A FOV_ _ - - SAC P RU TS P

NEGATIVE SOE D PA .PPI "CIS--' CSC S LO TVE

S AC B L A

CCA L LY / CNI

C LA

ecu 84

detected out of 990 possible species-pair combinations. Salix nivalis has the greatest number of significant associations: 11 positive with vascular species and bare ground and 5 negative -

4 with lichens and 1 with Carex capitata. Carex capitata is positively associated with 4 lichens and 3 vascular species and negatively associated with 2 lichens and 3 vascular plants.

Carex scirpoidea has 3 positive associations with vascular species and 1 negative association with bare ground. Cetraria

islandica has a total of 7 negative associations.

Group C2:

Point-line data comprising 5470 points and 43 species were analyzed. Thirty-four significant positive associations and 17

significant negative associations (Fig. 23) occurred out of 946 possible species pair combinations. The co-dominant vascular

species, Carex capitata and Carex scirpoidea, were positively associated with vascular species only; 3 and 2, respectively.

Carex capitata has 5 negative associations, including 1 lichen and Carex scirpoidea is negatively associated with 2 vascular

species. Other major species include Arenaria obtusiloba (4 positive associations), and Cetraria islandica (4 negative associations).

4. Transect D:

Point-line data were sub-divided into three groups based on quadrat data analysis. Kobresia myosuroides is the vascular dominant of group D1, and Carex scirpoidea of group D3. Group

D2 is a transition zone with Kobresia myosuroides and Carex 85

Figure 23 - Group C2 Positive and Negative Associations

POSITIVE CNI C N A I I I KMY I C AC

ecu LLY OUP S LO

C I S SDE CLA

T VE / SLU / *~ P P I P PR / I OMO BLA I I I • FOV TSP C PH CSC / A OB' I I CCA .. SAC E AU LCA PDI " CBE

NEGATIVE

S LO CLA CNI

OMO CSC CCU FOV TSP

PDI C I S C PH SDE CCA TVE

SAC K MY CAC

PPI A 0 B PPR 86

scirpoidea as co-dominants.

Group D1:

This data set comprises 25 species (2122 points) and has

comparatively few significant associations - 13 positive and 4

negative (Fig. 24), with 325 species-pair combinations possible.

Kobresia myosuroides was positively associated with 2 lichens

and 1 vascular species and negatively associated with only 1

lichen species.

Group D2:

The transitional grouping of 1303 points includes data for

25 species. A total of 11 significant positive and 3

significant negative associations were detected (Fig. 25), with

325 species-pair combinations possible. Kobresia myosuroides

has 3 positive associations (with 2 lichens and 1 vascular

species) and 2 negative associations, with 1 lichen and with

Carex scirpoidea. Carex scirpoidea is positively associated

with 2 vascular species.

Group D3:

This larger data set (4362 points) contains data for 38

species. Twenty-five significant positive and 6 significant

negative associations (Fig. 26) occurred out of 741 possible

species-pair combinations. The dominant vascular species, Carex

scirpoidea, was positively associated with 2 vascular species and negatively associated with bare ground. Other major species

include Arenaria obtusiloba (4 positive associations),

Selaginella densa (5 positive, 3 negative), and Cetraria

islandica (3 positive). 87

Figure 24 - Group D1 Positive and Negative Associations

POSITIVE

ecu

CAC LVU CNI KMY

AA L PDI

AOB CSC SMU / BLA PJ U

T CE

OUP SDE CLA CAN

NEGATIVE

A°B CLA KMY

C N I SDE

CSC BLA 88

Figure 25 - Group D2 Positive and Negative Associations

POSITIVE

AOB P N I CLA BL A S LA

TC E CSC SMU

CAN

CAC HLY PDI KMY

C I S C N I ecu

NEGATIVE

CLA KMY CSC

CAC PD I 89

Figure 26 - Group D3 Positive and Negative Associations

POSITIVE

CSC AG L SMU PN I CAN ppi

A A L

OMO CLA BLA- AO B TC E

PCA SOE TS P

CNA OUP PDI CCU PFR

FOV SAC CAC CNI C I S

C PU

NEGATIVE

CSC BLA SMU

CCU

CLA AOB SDE

TC E 90

5. Transect E:

Sub-division of point-line data into two groups occurred

before association analysis. Group E1 was dominated by Kobresia myosuroides and group E2 by Carex scirpoidea, with Kobresia myosuroides virtually absent. No transition zone was

identified.

Group E1:

Data comprising 3447 points and 35 species were analyzed.

A total of 30 significant positive and 14 significant negative

associations were detected (Fig. 27), with 630 different

species-pair combinations possible. Kobresia myosuroides was

positively associated with 4 lichen species and negatively

associated with 3 vascular species, 1 lichen, 1 moss, and bare

ground. Nine species were positively associated and 2 species

negatively associated with bare ground. Other major vascular

species include Selaginella densa (4 positive associations, 1

negative), Arenaria obtusiloba (4 positive, 2 negative), and

Carex scirpoidea (4 positive, 1 negative).

Group E2:

This data set contained 6117 points for 43 species.

Thirty-eight significant positive assocations and 17 significant

negative associations (Fig. 28) occurred out of 946 possible

species-pair combinations. The dominant vascular species, Carex

scirpoidea, has no significant positive associations and only 1

significant negative association with a moss (Polytrichum

piliferum). Several less abundant species had a relatively

large number of associations, for example, Silene acaulis (8 91

Figure 27 - Group E1 Positive and Negative Associations

POSITIVE

LVU OU P

TSP PD1 CMI

NEGATIVE

SDE CSC

CCU CNI T VE \ • • \ .• • N s \

AOB' /PI OUP

/ / / ..KMY BLA CIS

FOV" /

/ CLA

LCA 92

Figure 28 - Group E2 Positive and Negative Associations

POSITIVE

E A U D LO

AOB „ ECO D I N

TCE LL Y OU P \ \ \ CAC ... CAL SDE

CNA CN I CLA

i PP I I C I S BLA I TS P ' S LO i CB E

~ ~ -SAC .. • H LY CCU FOV

PRU LCA TVE SMU OMO

NEGATIVE

P P I CSC

CLA

BLA P D

PN IAOB CNA

CAC

SDE TVE

SAC / FOV CN I CCU L L Y

^OU P- 93

positive associations, 4 negative), Selaginella densa (6 positive, 5 negative), Arenaria obtusiloba (4 positive, 4 negative), and Ochrolechia upsaliensis (5 positive, 4 negative).

6. Transect F;

Based on results of quadrat data analysis, point-line data were sub-divided into two groups. Kobresia myosuroides is the vascular dominant of group F1 and Carex scirpoidea and Carex capitata are vascular co-dominants of group F2.

Group F1:

Data involving 1822 points and 27 species were used in association analysis. Seventeen significant positive and 8 significant negative associations (Fig. 29) were detected from

378 possible species-pair combinations. Kobresia myosuroides was positively associated with 3 vascular species and bare ground and negatively associated with 2 vascular species and 1 lichen. Other major species include Selaginella densa (4 positive associations, 4 negative), and Cetraria islandica (4 positive, 2 negative).

Group F2:

The data set of 8178 points comprised 39 plant species.

The largest number of significant associations occurred in this data set - 54 positive (Fig. 30) and 43 negative (Fig. 31), out of 780 possible species-pair combinations. Carex scirpoidea is positively associated with 4 vascular species and 1 moss and negatively associated with 2 lichens and the vascular co- dominant, Carex capitata. Carex capitata is positively 94

Figure 29 - Group F1 Positive and Negative Associations

POSITIVE

BLA PCA

AOB KMY LLY SDE CLA

CAC OU P POA

CSC CCA CIS CCU S LO

CNI TVE PJ U FOV

NEGATIVE

CAC CIS

I AOB SDE KMY CCA

CCU CLA 95

Figure 30 - Group F2 Positive Associations

L L Y 96

Figure 31 - Group F2 Negative Associations 97

associated with 2 vascular species and 5 lichens and negatively associated with 5 vascular species, one lichen, and bare ground.

Other important species include Arenaria obtusiloba (6 positive associations, 6 negative associations), Polytrichum piliferum (7 positive, 5 negative), Cornicularia aculeata (6 positive, 6 negative), Ochrolechia upsaliensis (7 positive, 3 negative),

Cladonia sp. (5 positive, 7 negative), and Selaginella densa (5 positive, 4 negative).

7. All Transects:

Positive Associations:

Significant positive associations for 11 data- sets consist of 182 different species pair combinations (including bare ground), and are presented alphabetically, by species, in

Table V. These comprise 4.4% of the total number of possible species-pair combinations summed over all data sets. Abundant species, other than dominants, tend to have a greater number of different associations than species with lower counts. The ubiquitous Arenaria obtusiloba had the largest number of different positive associations (22), while some rare species such as Calamagrostis purpurascens, Draba lonchocarpa, and

Potentilla fruticosa were positively associated only once.

Carex scirpoidea has very few positive associations where it is the vascular dominant - 2 in group D3 and zero in group E2.

Many more associations occurred where this species was a co- dominant with Carex capitata. 98

Table V - Positive Associations for 11 Sample Groups.

These are presented alphabetically with respect to species. Numerals refer to the number of times each species was contacted in each data set. Lichen species are indicated by a • and moss species by a n. Species codes are given under each species heading and are formed from the first letter of the generic epithet and the first two letters of the specific. Associations for each species are indicated by codes, with *, **, and *** refering to p<0.05, p^O.01, and p^O.001 levels of significance, respectively. 99

POSITIVE INTERSPECIFIC ASSOCIATIONS

TRANSECT A B Cl C2 01 D2 D3 El E2 Fl F2

DOMINANT CSC CSC SNI CSC KMY CSC CSC KMY CSC KMY CSC VASCULAR CCA CCA CSC CCA KMY CCA SPECIES CCA

Agoserfs ------31 gl auca CSC* (AGL) CLA*

Antennarla 17 - 13 11 10 16 62 49 alplna SNI* (AAJL) AOB*** PNI***

Arenarla 402 410 208 299 244 128 460 192 390 105 501 obtusiloba FOV*** FOV* FOV*** (AOB) CSC** CSC* CSC** CSC** CSC*** PDI* LCA* LCA* LCA** CCU** CCU* CCU** TVE* CNA* CBE* TSP*** TSP* TSP* TSP*** TSP*** CAC* SNI*** SLO** E AU* EAU* SLU* TCE** TCE* TCE* AAL*** PNI* SMJ* SMJ* BLA* BLA*** BLA*** DLO** LLY*** KMY***

Cal an agrostis - - - - 13 9 72 purpurascens CIS*** (CPU) 1 00

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT CI C2 DI D2 D3 El E2 Fl F2

Ca lop I aca 17 18 29 sp. SAC*** (CAL) OUP*** CLA***

Candelarlella 17 17 33 1 7 sp. CLA*** (CAN) KMY* BLA*

Carex 1186 1709 310 784 - - 1 6 - 36 1312 capitata ecu*** ecu*** ecu** ecu** (CCA) CNI*** CNI** CNI* CNI** CBE* CBE*** SL0**» CIS* CIS*** CIS*** CIS* CIS*** HLY* PDI*** PDI** PDI EAU* TVE** DOC* CPH** CMI ** LLY** CSC* PPI CAC*

Carex 254 46 1 8 1 6 44 78 207 nardIna CAC*** CAC*** CAC** CAC*** (CNA) OUP*** OUP*** PPR** AOS*

CNI CNI CN I *** SDE* OMO*** PDI***

Carex 43 1 5 phaeocephal a CCA** -j-yjr*** (CPH) 101

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT B Cl C2 0! D2 D3 El E2 Fl F2

Carex 1088 962 230 677 120 168 812 227 854 106 780 scirpoidea 0M0**OMO* * OMO* OMO*** (CSC) PDI** poi* PDI»* PDI* AOB** AOB* AOB** AOB** AOB*** SAC*** SAC* SAC** PPR*** PPR*** PPR** PPR*** PPU* SMJ* SMU** SMU*** EAU« TCE** AGL* PNI* CCA* DES*** HLY** TSP**

Cerastium 42 1 7 23 26 beer i ng I an un CCA* CCA*** (CBE) CNI* AOB* HLY***

Cetraria 620 768 176 349 54 26 52 261 195 100 589 cucul 1 ata CCA*** CCA*** CCA** CCA** (CCU) PDI * PDI** FOV** FOV* AOB** AOB* AOB** TVE** TVE*** SLU* SLU*** CIS** CIS* ClS*** CIS*** CIS** CIS*** KMY*** KMY* KMY*** KMY*** CMI** SLO***

Cetrar1 a 1261 869 294 504 83 44 134 192 100 1 1 1 687 Is1 and ica CNI*** CN1*** CN1*** CNI** CN1** CNI** CNI*** (CIS) TVE*** jy£*** TVE*** TVE*** CCA* CCA*** CCA*** CCA* CCA*** CMI*** CCU* CCU* ecu*** ecu*** ecu*** ecu*** PFR*** CPU*** LVU* 1 02

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT A B CI C2 DI D2 D3 El E2 F2

Cetrar I a 479 536 189 401 114 50 138 391 541 96 431 nival Is CCA*** CCA** CCA* CCA** (CNI ) CIS*** CIS*** CIS*** CIS** CIS** CIS*** CIS*** CAC*** CAC* CAC* CAC* CBE* FOV* CNA** CNA* CNA*** KMY*** KMY* KMY** LVU** LVU**

PPI PPI1 PDI1

CI adIna 39 71 1 5 1 5 m Itls CIS*** (CMI ) CCA** ecu* CAC***

Cladonia 26 37 28 chiorophaea SDE** (CCH) CLA*** CLA*** CLA** CAC* PPI*** PPI***

CIadon ia 1 1 5 147 89 54 78 43 138 67 1 1 1 60 208 sp. SDE*** SDE*** SDE*** SDE* SDE** SDE*** SDE*** SDE*** SDE*** SDE*** (CLA) CCH*** CCH*** CCH** EAU*** CAC* PPI*** pp|*** PPI*** PPI** PPI*** PPI*** DPA** BLA* BLA* BLA** BLA** PCA*** PCA*** PCA***

CAN*** PNI* AGL* CAL*** OUP** OUP** 103

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT A B CI C2 01 D2 03 El E2 Fl F2

Corntcularla 322 232 54 93 114 82 449 327 682 97 470 aculeata PPI*** (CAC) CNA*** CNA*** CNA*** CNA*** SDE*** SDE*** SDE*** SDE* SDE*** SDE* OUP*** OUP** OUP* OUP** TVE** CNI*** CNI* CNI* CNI* CCH* CLA*

AOS1 LLY*** LLY* KMY*** KMY* LVU** HLY**» PNI CMI *** CCA***

Degnatodon 11 13 1 5 30 9 7 6 sp. TVE* (DES) fSC***

Draba 12 - 7 - - - 1 6 1 5 20 1 ncerta BLA*** (DIN) OUP**

Draba - 6--- - 6 - 20 I one hocarpa AOB** (DLO)

Draba 40 1 8 33 1 1 24 39 payson 11 SNI*** (DPA) CLA** 3LA*

Dryas 47 octopetal a PDI*** (DOC) CCA*

Erlgeron 80 123 85 71 11 25 6 73 aureus CLA*** (EAU) FOV* FOV** PPI* PPI CCA* CSC* AOB* AOB* LLY* 1 04

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT B Cl C2 01 02 D3 El E2 Fl F2

ErIgeron 75 68 con posltus LLY** (ECO) SDE*** OUP***

Festuca 427 344 88 191 41 39 1 12 73 201 57 255 ov ina SAC*** SAC*** SAC** SAC*** SAC*** SAC*** SAC*** SAC*** (FOV) AOB*** AOB*** PPR*** OMO** OMO* OMO* EAU* EAU** SNI*** CNI* CCU** CCU* SMU** SLU* PJU*

Haplopappus 16 11 14 20 33 29 152 lyal I i i CCA* (HLY) CAC* CBE*** LCA* SAC* CSC*

Kobresia 28 497 1 70 558 476 83 myosuroides CCU*** CCU' CCU*** ecu*** (KMY) CAC*** CAC* CNI*** CNI* CNI** PDI*** PDI** CAN* TVE* LLY* AOB*** BLA* POA* OUP**

Letherta 14 27 13 52 34 55 28 vul pina CAC** (LVU) CNI** CNI** CIS* 1 05

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT A B CI C2 DI D2 03 El E2

LupInus 58 139 42 31 113 45 54 lyal I I I SNI* SNI*** (LLY) pp,»*»

CAC*** CAC* SLO*** SDE*** SDE*** SDE* SDE* BLA*** AOB*** ECO** KMY* CCA** EAU**

Luzula 253 179 19 65 12 30 156 20 112 can pestr 1 s SAC** (LCA) AOB* AOB* AOB** HLY* OUP*

Ochrolechla 42 45 13 20 69 31 99 76 238 40 127 upsallensls CAC*** CAC** CAC* CAC** (OUP) CNA*** CNA*** PPI*** PPI** SDE*** SDE** SDE*** SDE** SDE*** SDE*** SDE*** CLA** CLA** CAL*** DIN*** ECO*** LCA* KMY***

s OxytropI 65 105 48 74 74 34 12 76 monticola CSC*** CSC* CSC*** (OMO) FOV** FOV* FOV*** SMJ*** SMJ« SAC** SAC* SAC*** SNI* SLO* TSP* BLA*** CNA***

Pel tiger a - 13 16 12 8 9 30 14 8 11 28 canIna SDE*** SDE*** (PCA) SNI*** CLA*** CLA*** CLA*** PPI*** 106

POSITIVE INTERSPECIFIC ASSXIATIONS (continued)

TRANSECT A B Cl C2 Dl D2 D3 E1 E2 Fl . F2

Penstemon 44 31 17 91 - - - - 40 163 procerus FOV*** (PPR) CSC*** CSC*** CSC** CSC*** CNA** SLO* PJU** SDE**

Poa - 9 22 18 - - - - 50 rupestrIs SNI* (PRU) SAC***

Poa 6 ------1014 sp. KMY* (POA)

Polemon inn 21 23 pulcherr in un CSC* (PPU)

Polytr Ichum 13 22 34 46 54 j un I peri nun BLA*** (PJU) FOV* PFR*

Polytrichun 149 257 172 181 24 73 268 1 1 192 p i I i ferun SNI*** (PPI ) CAC*** SDE* SDE*** SDE** SDE*** SDE* SDE*** CLA*** CLA*** CLA*** CLA** CLA*** CCH*** CCH*** EAU* EAU** OUP*** OUP** LLY*** TVE** TVE** BLA*** BLA*** CNI*** CNI* CCA*** PCA* 1 07

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT A B Cl C2 Dl D2 D3 El E2

Potent 1 I la 551 641 128 341 137 65 207 37 144 diverslfol la CSC** CSC* CSC** CSC* (PDI) AOB* CCU* CCU** CCA*** CCA** DOC*** SAC* SAC** KMY*** KMY** CNI**

Potent I I la - - - - 17 - 22 fruticosa CIS*** (PFR)

Potent I I la - - - - 25 40 165 25 96 n ivea AOB* (PNI) CLA* AAL*** TCE** SMJ* BLA*** CAC* CSC*

SalIx 73 - 607 nival Is LLY*** LLY*** (SNI) PPI*** FOV*** BLA** AAL* SLO** AOB*** PRU* SAC*** OMO* TSP* PCA*** DPA***

Seduii 16- 7 - - 9 - - 6 I anceol atun BLA*** (SLA) 1 08

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT A B CI C2 01 D2 03 El E2 Fl F2

Selaglnel la 299 205 62 104 148 49 268 98 296 101 269 densa CLA*** CLA*** CLA*** CLA* CLA** CLA*** CLA*** CLA*** CLA*** CLA*** (SDE) CCH** CAC*** CAC*** CAC*** CAC* CAC*** CAC* TVE* TVE* PPI* PPI*** PPI** pp|*** PPI * pp,*** PCA*** PCA*** OUP*** OUP** OUP*** OUP** OUP*** OUP*** OUP*** LLY*** LLY*** LLY* LLY* CNA* BLA* BLA** ECO*** PPR**

Seneclo 27 39 38 33 - - 8 43 - 42 lugens CCU* CCU*** (SLU) AOB* FOV*

S i I ene 117 96 113 131 - 12 27 68 205 10 86 acaul Is LCA** LCA* (SAC) FOV*** FOV*** FOV** FOV*** FOV*** FOV*** FOV*** FOV*** CSC*** CSC* CSC** CAL*** •J-3P*** ygp*** TSP** OMO** OMO* OMO*** SMJ** SNI*** PDI* PDI** PDI * BLA** BLA** PRU*** SLO*** HLY*

Sol tdago 69 192 - 18 13 26 146 - 24 17 125 multiradlata OMO*** OMO** (SMU) SAC** CSC** CSC** CSC*** PNI* AOB* AOB* FOV** 1 09

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT B Cl C2 Dl D2 03 El E2 Fl F2

StelI aria 99 106 38 85 33 198 long I pes CCA*** (SLO) AOB** SNI** LLY*** OMO*** PPR* SAC*" CCU*

Taraxacum - 19 26 77 23 ceratophorun AOB** AOB* AOB* (TCE) CSC** PNI**

Thannol ina 777 796 259 451 1 5 I 46 90 38 212 verm icularis Cl S*** Cl S*** Cl S*** CIS*** CIS*** CIS*** (TVE) CAC** SOE* SDE* AOB* DES* CCU** ecu* CCU* CCA** BLA ** CPH PPI** KMY*

Tr isetun 49 66 51 96 11 10 20 17 51 14 117 spicatuti SAC*** SAC*** SAC** (TSP) AOB*** AOB* AOB* AOB*** AOB*** SNM CSC** OMO* 1 10

POSITIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT A B CI C2 DI D2 D3 El E2 Fl F2

Blank 49 20 138 29 178 133 333 265 477 16 219 (BLA) SNI** TVE** PJU*** CLA* CLA* CLA** CLA** SLA*** CAN* PPI*** pp|*** SDE* SDE** AOB* AOB*** SAC** OMO*** KMY* DIN*** LLY*** DPA** 111

The positive associations detected by association analysis are too numerous to discuss individually. The following categories consider possible trends within these associations, based on their locations in the study area and gross species morphology.

1) Community and Aspect Relationships:

Positive associations varied considerably between sampled sites, with the majority showing no relationship to specific community types or aspects, or even the relative abundances of the species involved. Examples include the positive association of Arenaria obtusiloba with Trisetum spicatum (groups B, C2, D3,

E1, and E2), and Lupinus lyalli i with Selaginella densa (groups

C2, E1, E2, and F1). As well, associations existing each time a species pair occurs together are limited to very rare species

(present in one or two data sets). Examples include the positive associations of Agoseris glauca with Carex scirpoidea and Cladonia sp. in group D3, Dryas octopetala with Arenaria obtusiloba in group C1, and Salix nivalis with Lupinus lyallii in groups A and C1. Only two associations between common species occurred everywhere counts greater than 25 were attained for both - Silene acaulis with Festuca ovina and Cladonia chorophaea with Cladonia sp.

A small number of associations, however, do indicate trends based on certain communities and/or aspects. Nine positive associations are essentially restricted to vegetation dominated by Carex capitata and Carex scirpoidea, although a few also 1 1 2

occur in group Fl , where these species are present, but not dominant. Several of these nine associations are restricted to north aspects (e.g., Carex capitata with Cerastium beeringianum) or to south aspects (e.g., Erigeron aureus with Polytrichum piliferum). No positive associations appear restricted to the remaining vegetation types, although a small number appear excluded from them. For example, the association between

Arenaria obtusiloba and Carex scirpoidea does not occur in Carex scirpoidea dominated vegetation, and Selaginella densa and

Polytrichum piliferum are not associated in Kobresia myosuroides dominated areas where both are present. The occurrence of 7 positive associations appears related to aspect. For example,

Kobresia myosuroides is associated with Cetraria cucullata when they occur together, with the exception of two vegetation groups at S56°E (transect F), and is also associated with Cornicularia aculeata at north aspects only (groups C2 and E1). Silene acaulis is positively associated with bare ground in two vegetation groups at N16°W (transect E) only.

A large number of positive associations occur only once and may be either fortuitous or specifically related to vegetation type or aspect. These include the association of Potentilla nivea with Cornicularia aculeata and with Carex scirpoidea in group E1 (Kobresia myosuroides community type), and the association of Carex scirpoidea with Taraxacum ceratophorum in group D2 (Carex scirpoidea/Kobresia myosuroides dominated). 113

2) Associations Between Vascular Species:

i) Monocot/Monocot

Two percent (4) of the 168 different positive plant/plant associations are between two monocot species, and each of these occurs only once. Carex capitata is positively associated with

Carex phaeocephala and with Carex scirpoidea where they do not co-dominate. As well, Carex sc irpoidea is associated with

Trisetum spicatum, and Kobresia myosuroides is associated with

Poa sp. Only one monocot species (Kobresia myosuroides) was positively associated with bare ground.

ii) Dicot/Dicot

Positive associations between dicot species are much more prevalent than those between monocots, and comprise 20% of the total number. Seventy-nine percent of these, however, occur only once. The most common positive dicot/dicot associations were detected a maximum of three times, e.g., Arenaria obtusiloba with Taraxacum ceratophorum, and Silene acaulis with

Oxytropis monticola and with Potentilla diversifolia. Salix nivalis has the largest number of different dicot associations - all occurring in group C1. In addition, the largest number of plant associations with bare ground (8) involved dicot species.

iii) Monocot/Dicot

Twenty-six percent of the positive plant/plant associations occur between monocot/dicot species pairs, with 63% of these 1 1 4

occurring only once. The most common monocot/dicot association is between Festuca ovina and Silene acaulis, and was detected 8 times (Silene acaulis was rare elsewhere). Other frequent associations include Carex scirpoidea with Arenaria obtusiloba

(5 times), Solidaqo multiradiata (3), Potentilla diversifolia

(4), and Penstemon procerus (4); Arenaria obtusiloba with

Trisetum spicatum (5), and Festuca ovina (3); Carex capitata with Potentilla diversifolia (3); and Oxytropis monticola with

Festuca ovina (3).

3) Associations Between Vascular Species and Lichens, Mosses, or

Clubmoss:

The largest number (30%) of positive species associations fall into this category, with the involved vascular species divided almost evenly between monocots and dicots. Seventy-one percent of these associations occur only once, with the most frequently detected association occurring 5 times (Carex capitata with Cetraria islandica). Other frequent associations include Carex capitata with Cetraria nivalis (4 times) and with

Cetraria cucullata (4); Kobresia myosuroides with Cetraria cucullata (4) and with Cetraria nivalis (3); and Lupinus lyallii with the clubmoss, Selaginella densa (4). Twenty-five percent of the associations in this category involve the two dominant species, Carex capitata and Kobresia myosuroides, while the third major dominant, Carex scirpoidea, has only one.

4) Associations Between Lichens, Mosses, and Clubmoss: 1 15

Associations in this category comprise 21% of the total number of positive species associations, with a relatively small percentage (42) occurring only once. Some of the most frequently occurring positive associations fall into this category - e.g., Selaginella densa and Cladonia sp. are associated in all groups except D2 (although counts over 40 occur for both here). Other frequent associations include

Selaginella densa with Ochrolechia upsaliensis (7 times),

Polytr ichum piliferum (6), and Cornicularia aculeata (5);

Cetraria islandica with Cetraria nivalis (7), Thamholina vermicularis (6), and Cetraria cucullata (6); and Cetraria nivalis with Cornicularia aculeata (4). Six different associations occur between bare ground and lichens, mosses, or clubmoss.

Negative Associations:

Significant negative associations for 11 data sets total

103 different species pair combinations (including bare ground)

- 79 fewer than the significant positive associations detected.

Significant negative associations comprise 2.4% of the total number of possible species-pair combinations summed over all data sets, and are presented in Table VI. Abundant species tend to have the greatest number of different negative associations, as was apparent with positive associations. Cetraria islandica and Carex capitata had 18 and 16 different negative associations, respectively, while many less frequent species were involved in only one each (eg., Draba paysonii, Cladonia 1 16

Table VI - Negative Associations for 11 Sample Groups.

The format is identical to that for Table V. 1 1 7

NEGATIVE INTERSPECIFIC ASSXIATIONS

TRANSECT A B CI C2 DI D2 D3 El E2 Fl F2

DOMINANT CSC CSC SNI CSC KMY CSC CSC KMY CSC KMY CSC VASCULAR CCA CCA CSC CCA KMY CCA SPECIES CCA

ArenarI a 402 410 208 299 244 128 460 192 390 105 501 obtusi loba CAC** CAC* CAC* (AOB) CIS* CIS* SAC* PPI* PPI** PPI* SDE* SDE*** SDE* SDE* SDE** PPR* CLA* CLA* CLA* CLA* TVE** KMY*** PNI* CNA* CCA*** OUP*

Carex 1186 1701 7099 310 784 1 6 36 1312 capitata CSC** CSC* CSC*** (CCA) PPI* PPI** PPI*** CCH* SNI* SNI SDE** SDE** SDE* CLA*** CLA* CLA** FOV** FOV*** FOV* FOV*** SAC* SAC* SAC* CNA* CNA* TSP* LLY* CAC* KMY* KMY*** KMY*** TVE* AOB*** BLA** 1 18

NEGATIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT B Cl C2 Dl D2 03 El E2 Fl F2

Carex 254 46 44 78 207 nard ina CCA* CCA*** (CNA) AOB* FOV* PPI ** CCU* CIS*** TVE*

Carex 43 1 5 phaeocephal a CSC* (CPH) CIS*

Carex 1088 962 230 677 120 168 812 227 854 106 780 scirpoidea CCA** CCA* CCA*** (CSC) PPI** PPI* TVE** TVE* TVE* CCU** BLA* BLA* BLA*** SLO* CPH* KMY* CIS**

Cetraria 620 768 1 76 349 54 26 52 261 195 100 589 cucul 1 ata CNI*** CN1*** CN1*** CNI*** CNI*** (CCU) CLA* CLA** CLA** CLA* CLA* SDE** SDE** SDE* SDE*** SDE* SDE* OUP* OUP** OUP** OUP* CSC**

LLY** LLY* SNI** CNA* OMO* 1 19

NEGATIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT Cl C2 Dl D2 03 El E2 Fl F2

Cetrar i a 1261 869 294 504 83 44 134 192 100 111 687 i si and lea SNI* SNI*** (Cl S) OUP* OUP* OMO* OMO* OUP*** BLA** BLA** AOB* AOB*

CAC*** CAC*** CAC* CAC* CAC** CAC SLO** SLO** #* run*** SAC** SAC* FOV** PDI* PDI PPI**

SDE* SDE** SDE* DPA* LLY* CPH* CLA*** CNA*** CSC** TSP*

Cetraria 479 536 89 401 114 50 138 391 541 96 431 n iv al is CCU*** CCU*** CCU*** CCU*** ecu*** (CNI) CLA* PDI** PDI* CLA** SDE* SDE* SDE* SDE* SLO* SDE** SNI OUP* OUP** SAC*** SAC** PPI*

Cladonia 26 37 1 0 1 5 28 chlorophaea CCA* (CCH) 1 20

NEGATIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT A B CI C2 DI D2 03 El E2 Fl F2

Cladonia 115 147 89 54 73 43 138 67 111 60 208 sp. FOV* FOV* FOV* (CLA) CCA*** CCA* CCA** CNI * CNI** ccu* ecu** ecu** ccu* ecu* KMY* KMY** KMY* KMY* AOB* AOB* AOB* AOB* CIS*** PDI*

CornicuI aria 322 232 54 93 114 82 449 327 682 97 470 aculeata CIS*** CIS*** CIS* CIS* CIS** CIS*** (CAC) AOB** AOB* AOB* CCA* PDI* SAC*** SAC** SLO* PPR* FOV*

Draba 40 18 33 1 1 - 7 24 39 payson i I CIS* (DPA)

Festuca 427 344 88 191 41 39 112 73 201 57 255 ov Ina CCA** CCA*** CCA* CCA*** (FOV) CLA* CLA* CLA* TVE** SDE* SDE** CIS** KMY* OUP* CNA* CAC* 121

NEGATIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT A B CI C2 DI 02 D3 El E2 Fl F2

Kobresia - - - 28 497 170 - 558 - 476 83 myosuroides CCA* CCA*** CCA*** (KMY) CLA* CLA** CLA* CLA* CSC* PPI** BLA*** LCA** FOV* AOB*** SDE*

LupInus 58 139 42 31 113 45 54 lyalI I i TVE* (LLY) CIS* CCA*

CCU** CCU"

Luzula 253 179 19 65 - 12 30 156 20 112 canpestrls KMY** (LCA)

Ochrolechia 42 45 13 20 69 31 99 76 238 40 127 upsallensts CIS* CIS* CIS*** (OUP) ccu* ecu** ecu** ccu* CNI * CNI** SAC* FOV* AOB*

Oxytropis 65 105 48 74 - 74 34 12 76 monticola CIS* CIS*

(OMO) ccu* 1 22

NEGATIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT A B Cl C2 01 D2 D3 El E2

Penstemon 44 31 17 91 procerus AOB* (PPR)

Polytrichim 149 257 172 181 - - 24 73 268 pi I Iferun CCA* CCA**

(PPI) CSC** csc# TVE* AOB* AOB** SMU* CIS**

KMY**

Potenti I la 551 641 128 341 137 65 207 37 144 diversifol ia CN I ** CN I * (PDI) TVE** CIS* CIS*** CAC* BLA*

Potenti I la - - - - 25 40 165 25 96 "'vea AOB* (PNI)

Sal ix 73 - 607 nival is CCA* CCA***

(SNI) TVE* TVE*»* CIS* CIS*** CNI*** CCU** 1 23

NEGATIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT B Cl C2 01 D2 03 El E2 Fl F2

Selaginella 299 205 62 104 1 48 49 268 98 296 101 269 densa CCA** CCA** CCA* (SOE) CCU** CCU** CCU* CCU*** CCU* CCU* SAC* SAC** FOV* FOV** AOB* AOB* AOB* AOB* AOB** CNI* CNI* CNI' CNI' CNI** CIS* CIS** CIS* TCE* TVE* KMY*

S iIene 11 7 96 1 1 3 1 31 27 68 205 10 86 acaul is CIS** CIS* (SAC) SDE* SDE** AOB* CCA* CCA* CCA* OUP* CNI*** CNI* CAC*** CAC**

Solidago 69 192 1 8 1 3 26 146 24 1 7 125 multiradiata PPI*

(SMU) BLA*

StelI aria 99 106 38 85 1 4 33 198 long I pes CNI* (SLO) CIS** CIS** CSC* CAC*

23 1 24

NEGATIVE INTERSPECIFIC ASSOCIATIONS (continued)

TRANSECT CI C2 DI D2 03 El E2 Fl F2

Tharinol Ina 777 796 259 451 1 5 7 146 90 38 212 verm Icul ar Is LLY*

(TVE) SNI * SNI1 PDI*** FOV**

CSC** CSC* CSC* PPI* CCA** TSP* TSP* AOB* SDE** PPR* CNA*

Trisetun 49 66 51 96 11 10 20 17 51 14 117 spicatun CCA* (TSP) TVE* TVE* CIS*

Blank 49 20 133 29 178 133 333 265 477 16 219 (BLA) CIS** CIS** CSC* CSC* CSC*** SMJ* KMY*** PDI* CCA** 125

chorophaea, and Taraxacum ceratophorum. Negative associations have been grouped into the same five categories used for positive associations.

1) Community and Aspect Relationships:

The vast majority of negative associations showed no relationship to specific community type, aspect, or the relative abundances of the species involved. Examples include the associations of Arenaria obtusiloba with Cladonia sp. (groups

D1, D3, E2, and F2), and with Selaginella densa (groups B, D3,

E2, F1, and F2); and the association of Silene acaulis with

Cornicularia aculeata (groups E2 and F2). Associations existing each time a species-pair occurs together are again limited to very rare species. For example, Salix nivalis, present in two data sets only, is associated both times with Carex capitata,

Thamnolina vermicular is, and Cetraria islandica. Low counts (16 or less contacts) for at least one member of the following species-pairs occurred at the only locations where associations between them were not detected: Carex capitata/Kobresia myosuroides, Carex phaeocephala/Carex scirpoidea, and Carex phaeocephala/Cetraria islandica.

A small number of negative associations do indicate trends based on certain communities and/or aspects. Seven negative associations are restricted to vegetation dominated by Carex capitata and Carex scirpoidea, and group F1, where these species are present, but not dominant. Only one of these associations

(Oxytropis monticola with Cetraria islandica) is restricted to 126

north aspects, and three occur at all aspects but N29°E (Festuca ovina/Cladonia sp. and Carex capitata/Polytrichum piliferum) or

N58°W (Arenaria obtusiloba/Polytrichum piliferum). In addition, the association between Kobresia myosuroides and Cladonia sp. occurs only where Kobresia myosuroides is a dominant or co- dominant, while Carex scirpoidea and Kobresia myosuroides are negatively associated only where they both co-dominate (group

D2). A single association (Selaginella densa/Cetraria nivalis) appears excluded from vegetation dominated solely by Carex sc i rpoidea. The occurrence of three negative associations appears related to aspect. Carex scirpoidea and Polytrichum piliferum, and Silene acaulis and Selaginella densa, are associated at north-west aspects only (groups A and E2), while

Cetraria nivalis is associated with Ochrolechia upsaliensis in two community types (dominated by Kobresia myosuroides and by

Carex scirpoidea) at an aspect of N16°W only (transect E).

Many significant negative associations occur only once.

For example, Kobresia myosuroides is associated with bare ground only where it is a dominant.species at a north aspect (group

E1). Other examples include the association of Carex scirpoidea with Cetraria cucullata (group B), Kobresia myosuroides with

Selaginella densa (group C1), and Arenaria obtusiloba with

Potentilla nivea and Carex nardina (group E2).

2) Associations Between Vascular Species:

i) Monocot/Monocot 1 27

Ten percent of the 97 different negative plant/plant associations are between two monocot species, with 60% of these occurring a single time. The most frequent are Carex capitata with Carex scirpoidea (4 times), Festuca ovina (4), and Kobresia myosuroides (4). Two monocot species (Carex scirpoidea and

Kobresia myosuroides) are negatively associated with bare ground.

ii) Dicot/Dicot

Negative association between dicot species is very rare only 3% of the total, with each occurring just once. Arenaria obtusiloba is involved in all three of these associations, with

Silene acaulis, Penstemon procerus, and Potentilla nivea. Only one dicot species (Solidago multiradiata) is negatively associated with bare ground.

iii) Monocot/Dicot

Seven percent of the negative plant/plant associations occur between monocot/dicot species-pairs, with 71% of these occurring just once. The most frequent are Carex capitata with

Silene acaulis (3 times) and with Salix nivalis (2).

3) Associations Between Vascular Species and Lichens, Mosses, or

Clubmoss:

The largest number (64%) of negative species associations fall into this category, with 42% of these involving monocot species, and 58% involving dicots. Sixty-one percent of 1 28

associations in this category occur only once, with the most

common occurring 5 times (Arenaria obtusiloba with Selaginella

densa). Other frequent associations include Kobresia

myosuroides with Cladonia sp. (4 times); Carex scirpoidea with

Thamnolina vermicularis (3); Carex capitata with Selaginella

densa, Polytrichum piliferum, and Cladonia sp.; and Arenaria

obtusiloba with Cladonia sp. (4), Polytrichum piliferum (3),

and Cornicularia aculeata (3). Twenty-one percent of the

associations in this category involve one of the three major

dominant species - Carex capitata, Carex scirpoidea, or Kobresia

myosuroides.

4) Associations Between Lichens, Mosses, and Clubmoss:

Associations in this category comprise 15% of the total

number of negative species associations, with a relatively small

percentage (33) occurring only once. Some of the most

frequently occurring negative associations fall into this

category - as was found with positive associations. Examples

include the associations of Cetraria cucullata with Selaginella

densa (6 times), Cetraria nivalis (5), Cladonia sp. (5), and

Ochrolechia upsaliensi s (4); and Cetraria islandica with

Cornicularia aculeata (6). Only one lichen species (Cetraria

islandica) is negatively associated with bare ground. 1 29

Constancy Of Interspecific Associations

Constancy of association for a species-pair is the number of times the association was detected, expressed as a percentage of the total number of times an association was possible (i.e., the two species must occur in the same data set). The constancy of significant positive (Table VII) and significant negative

(Table VIII) associations are presented as a summary of Tables V and VI, respectively. Unique and common associations are readily identified with constancy data. Constancy values of 50% or more are often limited to rare species present in only 1 or 2 data sets (indicated by *), for example, Salix nivalis, Dryas octopetala, Agoseris glauca, and Eriqeron compositus. These species are indicated by a * in Tables VI and VII. Positive associations between more abundant species with constancy values of 50% or greater include Arenaria obtusiloba/Taraxacum ceratophorum (75%); Cornicularia aculeata with Carex nardina

(57%) and Selaginella densa (54%); Carex nardina/Selaqinella densa (54%); Carex capitata with Cetraria cucullata (57%),

Cetraria islandica (71%), and Cetraria nivalis (57%); Cetraria cucullata with Cetraria islandica (54%) and Kobresia myosuroides

(67%); Cetraria islandica with Cetraria nivalis (64%) and

Thamnolina vermicular is (60%); Cladonia sp. with Polytrichum piliferum (56%) and Selaginella densa (91%); Penstemon procerus/Carex scirpoidea (67%); Silene acaulis/Festuca ovina

(80%); and Selaginella densa with Ochrolechia upsaliensis (64%) and Polytrichum piliferum (67%). Relatively few negative associations have constancy values of 50% or more. These 1 30

Table VII - Constancy of Positive Associations

Percentage of positive associations between species pairs in data sets where both occur together. Species present in <3 data sets are indicated by a *.

* A G L AAL AOB 14 CPU CAL CAN CCA CNA CPH 33 c s c|ioo 45 14 20 40 C B E 27 57 CCU 33 71 C I S 54 C N I 5743 20 64 C M I 20 20 20 CCH C L A [WO 25 17 50 CAC 14 57 452017 9 DES D I N D L O 33 D P A •DOC EAU 25 14 12 • ECO FOV 27 18 25 HLY 17 11 20 11 KMY 33 67 50 33 L V U 14 28 14 LLY 11 22 50 25 LCA 33 12 12 OUP 25 28 18 36 20 50 17 O MO 14 38 38 PCA 30 PPR 20 67 17 P R U POA 50 • P P U 50 PJU 17 33 P P I I 14 22 405611 14 11 22 12 P D I 4314 36 18 9 to 33 • P F R 50 P N I 25 17 17 • SNI 50 50 50 50 100 50C0 tD 50 SLA SDE 1791 54 50 44 64 20 67 SLU 28 14 SAC 25 30 8011 22 38 50 30 50 SMU 22 33 11 28 20 12 SLO 12 14 50 12 TCE 75 25 25 TVE 10 14 33 X60 K>14 20 22 20 TSP 45 12 50 30 BLA 27 17 36 20 12 17 12 '7 22 5025 18 _20_ 10 A C CCCCCCCCCCCCCCDDDDDEEF H K L LLOOPPPPPPPPPPSSSSSSSTTTB O P AACNPSBC INMCLAE I LPOACO L M V LCUMCPROPjPDFNNLDLAMLCVSL B U LNAAHCEUS I IHACSNOACUOV Y Y U YAPOARUAUUIIRl IAEUCUOEEPA * * 131

Table VIII - Constancy of Negative Associations

Percentage of negative associations between species pairs in data sets where both occur together. Species present in <3 data sets are indicated by a *.

* A G L A A AOB CPU CAL CAN CCA 14 CNA 14 40 CPH CSC 43 33 CBE CCU 14 9 C I S 14 33 9 CN I C M I 45 CCH 25 CLA 36 43 CAC 45 9 18 27 14 D E S 54 0 1 N D L O D P A • DOC E A U t EC O FOV H L Y 5714 27 9 KMY LVU 17 75 67 LLY LCA 14 2211 O U 25 O MO| 36 2718 PCA 12 25 P P R P R U 17 17 P O A • P P U P J U P P I P D I 33 4314 22 11 11 PFR 25 17 18 18 P N I 9 9 I S N 17 SLA 100 50C050 SDE S L U 45 43 SAC 542745 18 17 S M U 10 43 SLO 20 20 20 T C E T V El 2512 12 T S P| 25 14 14 30 BLA 10 17 10 14 9 11 10 100 14 27 18 20 17 11 AACCCCCCCCCCCCCCCDDDDDEEFHKLLLOOPPPPPPPPPPSSSSSSSTTTB fnM?MCNP;?BC INMCLAEI L POACOL MV LC UMC PRO PJ PD F N N L DL AMLCV SL LBULNAAHCEUS I IHACSNOACUOVYYUYAPOARUAUUI I R | IAEUCUOEEPA 132

include Cornicularia aculeata/Cetraria islandica (54%); Carex capitata with Festuca ovina (57%) and with Kobresia myosuroides

(75%); Cetraria cucullata/Selaqinella densa (54%); and Kobresia myosuroides/Cladonia sp. (67%).

A number of species-pairs are positively associated in some data sets and negatively associated in others (Table IX). In the majority of cases, only single associations of one or both types occur. The vascular species, Arenaria obtusiloba and

Carex capitata are involved in the largest number of these situations.

Species Ordinations

As all forms of pattern, including interspecific association, are dependent on the scale of investigation, a consideration of species associations at the level of the quadrat (20 X 50 cm) seems useful for comparative purposes. PCA and RA were used to ordinate species at this scale, in an attempt to illustrate similarities between species on the basis of co-occurrence within quadrats. Initially, species were ordinated using percentage cover data, which yielded very little useful information on the associations between them. As these analyses summarize the range of data variation, rare species with little difference in percentage cover values tend to group together, and more abundant species, with greater cover variation, appear outlying. In addition, these abundant species are widely separated in the scatter plots, even when present in all quadrats, due to cover variation differences. This effect was somewhat more evident with PCA, than with RA. Percentage 1 33

Table IX - Species Pairs Associated Both Positively and Negatively in Different Vegetation Groups

SPECIES ASSOCIATION GROUP

Arenaria Carex + B obtusiloba nardina - E2

Kobresia + F1 ;myosuroides - E1

Potentilla + D2 nivea - E2

Thamnolina + B vermicularis - E1

Cornicularia + C1 aculeata - A,F1,F2

Carex Carex + F1 capitata scirpoidea - A,B,F2

Lupinus + F2 lyallii - C1

Polytrichum + E1 piliferum - A,B,F2

Cornicularia + F2 aculeata - Cl

Thamnolina + C1 vermicularis - C2

Cetraria Potentilla + F1 nivalis diversifolia - A,B

Polytrichum + A,E2 piliferum - F2

Thamnolina Polytrichum + C2,F2 vermicularis piliferum - A

Selaginella + A,C1 densa - E2

Kobresia Bare ground + F1 myosuroides - E1 134

cover values were then converted to presence/absence data in an attempt to overcome these cover variation difficulties. PCA and

RA of presence/absence data separated rarer species from more abundant ones, with this effect only slightly minimized when species of <5 occurrences were removed. Although species present in all quadrats appeared close together in the scatter plots, so did rare species with only absences in common. As well, placement criteria for rare and relatively common species depended more on the total number of presences and absences shared, than on co-occurrence in particular quadrats.

Although these analyses were less than satisfactory to illustrate association at the scale of the quadrat, one major point has been emphasized. The grouping of common species due to their presence in most quadrats provides a number of examples of large scale positive associations which are not present at a smaller scale of 2 cm. .- For example, Carex scirpoidea is negatively associated at 2 cm with its co-dominant, Carex capitata in groups A, B, and F2, although both species are present in each sampled quadrat in these groups, and are therefore positively associated at this scale. Other examples include Cornicularia aculeata/Cetraria islandica, Carex capitata/Festuca ovina, and Cetraria cucullata/Cetraria nivalis

- all negatively associated at 2 cm and positively associated in

20 X 50 cm quadrats. It seems that even relatively small quadrats tend to give spurious associations compared to those detected at the scale of the individual plant. 135

SOILS

Morphology

Alpine Dystric Brunisol soils within the study area were described in each vegetation grouping. Profile descriptions were very similar between sites. The following profile of transect A is typical:

Horizon Description

A 0-14 cm; color (dry) dark brown (1OYR 3/3), (wet) very dark brown (1OYR 2/2); loamy sand, gravel 1-2 cm in diameter, <5%; weak, amorphous structure; friable; plentiful fine to medium roots; strongly acid (pH 5.2); abrupt, irregular boundary.

B 14-37+ cm; color (dry) dark yellowish brown (10YR 4/4), (wet) dark yellowish brown (1OYR 3/4); loamy sand, gravel (angular) 2-5 cm in diameter, 25%; weak, amorphous structure; friable; abundant fine and very fine roots; strongly acid (pH 5.2).

The major rooting depth in these soils ranged from 0 to 18-28 cm, with the majority of roots occurring 0-15 cm below the surface.

Physical And Chemical Properties

Physical and chemical soil properties were determined within the 11 vegetation groups previously described (Table

X). All profiles are strongly acid (pH 4.8-5.6), and pH values may increase or decrease with depth. No relationship between pH and vegetation type was observed.

All soils are coarse textured, and range from sandy loam to sand, with loamy sand predominating. Organic matter is highest in the A horizon and decreases with depth for all profiles. TABLE X - PHYSICAL AND CHEMICAL PROPERTIES FROM SOIL PROFILES WITHIN 11 VEGETATION GROUPS. L/S AND S/L INDICATE LOAMY SAND AND SANDY LOAM, RESPECTIVELY.

VEG- TEXTURE-2 mm (%) TEXTURE ORGANIC TOTAL AVAIL. EXCHANGEABLE CATIONS TOTAL CATION GROUP HORIZON pH 1%) (II {%T GRAVEL CLASS MATTER N P (meq/100g) EXCHANGE CAPACITY

SAND SILT CLAY {%) {%) ppm K Mg Ca (meq/lOOg)

A A 0-l4cm 5.2 79 18 3 53 L/S 10.7 • 32 11.1 .19 .63 3.92 20.8 B 5.2 2 14-37+ 85 13 52 L/S 5.8 .21 20.4 .09 .26 1.69 16.5

B A 0-10cm 84 5.1 13 3 58 L/S 13.0 .46 13.6 .44 .98 5.07 25.9 B 10-41+ 88 5.1 9 3 52 SAND 5.2 .22 10.4 .09 • 29 1.88 17.9

C1 A 0-14cm 5.0 80 15 4 54 L/S 6.6 .24 12.4 .16 .35 1.74 16.0 B 14-35+ 5-3 69 26 5 49 S/L 1.7 .06 31.9 .11 • 31 1.36 7.8

C2 A 0-17cm 5.3 84 14 2 31 L/S 8.7 .32 9.4 .22 1.02 4.82 18.0 B 17-30+ 4.9 80 15 4 33 L/S 2.3 .13 10.2 .10 .17 .70 12.0

D1 A 0-l4cm 5.4 70 27 3 28 S/L 11.3 .49 7.4 .20 1.62 7.46 27.4 B 14-30+ 5.0 82 15 3 26 L/S 2.1 .12 7.8 .08 .17 1 .22 10.4

D2 A 0-13cm 5.4 21 78 1 60 L/S 10.3 .54 6.8 .23 .98 6.60 30.9 B 13-35+ 5.6 75 21 4 44 L/S 4.9 .29 10.4 .14 .66 4.80 21.5

D3 A 0-12cm 4.8 86 11 3 32 L/S 8.0 .31 10.9 . 12 • 52 2.70 21 .5 B 12-30+ 5.3 83 15 2 35 L/S 4.7 .14 19.2 • 19 .46 2.30 15.2

E1 A 0-15cm 5.3 82 16 2 43 L/S 9.6 .33 10.1 .24 .69 5.72 23.6 B 5.2 88 10 15-35+ 2 44 SAND 1.5 .08 12.7 .06 .14 1 .26 7.3

E2 A 0-15cm 5.2 81 46 15 4 L/S 7.7 • 32 8.6 .25 .98 5.90 21 .5 B 15-35+ 5.3 84 13 3 40 L/S 3.8 .10 18.2 .06 .20 1.35 10.8

F1 A 0-17cm 4.8 4 83 13 29 L/S 6.2 • 23 10.0 .14 • 30 2.29 20.7 B 17-30+ 5.0 4 65 30 21 S/L 2.1 .08 19-3 .11 .06 .60 8.9

F2 A 0-15cm 5 80 16 4 42 .1 L/S 10.0 .32 17.4 .12 .33 2.02 21.7 B 15-35+ .9 88 11 1 4 60 SAND 5.0 .20 28.0 .09 .20 1.43 14.7 137

Total nitrogen, exchangeable cations, and total cation exchange capacity appear correlated with organic matter and decrease with depth - often substantially. Values for calcium are relatively high - particularly in the A horizon of vegetation groups D1 and

D2 (7.46 and 6.60 meq/100 g, respectively) compared to the value for group D3 (2.70 meq/100 g) at the same aspect. Magnesium is also comparatively high in the A horizon of D1 (1.62 meq/100 g), as is total cation exchange capacity in D1 (27.4 meq/100 g) and

D2 (30.9 meq/100 g). In contrast to other analyzed minerals, phosphorus increases with depth. Particularly high values for phosphorus were detected in the B horizons of group C1 (31.9 ppm), and F2 (28 ppm).

Principal Components Analysis:

Physical and chemical soil properties of the 11 vegetation groups previously outlined were analyzed with centered and standardized PCA (Appendix D). Values for A and B horizons were analyzed separately, with the first two axes of each ordination accounting for 68% and 75% of the total variation, respectively.

Three vegetation groups, B, D1, and D2, appear outlying from the main point cluster after analysis of A horizon values (Appendix

D). This is likely due to the relatively high amounts of exchangeable cations and total cation exchange capacity in these samples (Table X). Group D2 appeared outlying after analysis of

B horizon data as well, although groups B and D1 occurred within the main point cluster.

First axis eigenvectors corresponding to A horizon values for sand, clay, and phosphorus have similar negative values, as 138

do eigenvectors corresponding to B horizon values for silt, clay, and phosphorus (Appendix D). This indicates that these parameters tend to vary together, due to interdependency or external agents.

CLIMATE

Mesoclimate

Mesoclimate of Lakeview Mountain was monitored with three weather stations from June 20 to September 6, 1980 (Table XI).

Mean daily temperatures were very similar between stations and increased until the July 21-31 period (mean daily max. 18°C), and subsequently decreased. Mean daily temperatures of 2-6°C were common, with a mean daily minimum of -3.4°C recorded. Wind speed was slightly higher at station 1 than at station 2, and mean values ranged from 7.3-20.5 km/hr for both sites. Relative humidity was also increased at station 2, compared to 1 and 3.

Precipitation was very low throughout the time period measured, with an accumulation of only 2.84 cm.

Microclimate

Temperature profiles for the 1980 growing season show the same general patterns for all 11 vegetation groups (Fig. 32).

As the actual dates on which readings were taken vary, not all graphs are directly comparable. Vegetation groups within a given transect were measured on the same days, as were transects

E and F (with the exception of the July 22 and 23 readings, respectively). All transects have several reading dates in common. The fluctuations in air temperature maxima over the 139

Table XI - Mesoclimatic Data For Three Weather Stations, 1: S84°W, 2481 m; 2: S2°E, 2402 m; and 3: N29°E, 2475 m.

TEMPERATURE °C WIND km/hr HUMIDITY DATE STATION MEAN MEAN MEAN % PPT cm 1980 DAILY DAILY DAILY MEAN RECORD. MEAN (60 cm) MIN. MAX. MAX. DAILY

20/6- 1 •1 .7 2.6 6.9 16.5 31.3 75 0.70 2 •2.2 2.9 6.6 11.0 13.8 83 30/6 3 •1.0 3.1 7.6 77

1/7- 1 1 .2 5.4 11.4 10.0 14.6 66 0.43 2 0.4 5.1 11.2 11.2 18.4 71 10/7 3 2.0 6.3 11.4 64

1 1/7- 1 0.5 4.3 9.9 13.7 24.7 78 0.63 2 0 4.1 9.4 9.5 16.8 84 20/7 3 1 .7 5.4 10.4 79

21/7- 1 3.7 9, 16, 14.2 19.5 52 0.01 2 4.0 10, 18, 11.0 15.8 64 31/7 3 5.0 1 1 , 17, 54

1/8- 1 0.3 5.2 10, 7.3 13.4 75 0.21 2 0 5. 1 1 1 , 8.5 15.8 78 10/8 3 1.0 6.2 10, 74

11/8- 1 1 .2 0 10 10.1 17.8 79 0.66 2 0.8 0 10 9.5 14.8 82 20/8 3 2.2 9 10, 78

21/8- 1 •1.9 2, 8, 11.8 20.6 66 0.04 2 •3.4 2 8, 8.5 11.3 70 31/8 3 •1.2 3, 8, 65

1/9- 0.2 4.5 10.0 20.5 38.6 70 0.16 0.8 4.8 10.3 19.6 26.6 72 6/9 0.3 4.0 9.2 70 1 40

Figure 32 - Microclimatic data for 11 vegetation groupings during summer 1980. Air temperature readings taken at +2 cm are indicated by , +10 cm by , and +20 cm by . Soil profile temperatures at -2 cm are distinguished by — — , -10 cm by — • — • , and -20 cm by . Readings were taken between 12:00 pm and 4:00 pm and approximate daily maxima. 141

GROUP A 30i

GROUP a

30-1

2CH

GROUP CJ

30-1

20-

10-

GROUP C2

301

30 10 20 30 10 20 30 JUNE JULY AUGUST 1 42

GROUP CM 30i

2CH

GROUP D3

30n

2CH 143 144

1980 growing season are quite dramatic, ranging from 0°C to

24°C, with the lowest temperatures observed near the begining of

July and the highest near the end of August or later.

Soil temperatures are observed to track air temperatures, sometimes with a 1 or 2 day lag period. The influence of air temperature decreased with increasing soil depth and temperatures become generally cooler and more stable. When an extreme drop in air temperatures occurs (e.g., 0°C in transect D on July 5, 1980), soil temperatures remain 3-8°C warmer.

Subsurface soil temperatures at a depth of 2 cm are the most widely fluctuating and often surpass the prevailing air temperatures. Highs of 24°C were reached at one location (group

E2) due to surface heating. It is interesting to note that subsurface temperatures in Kobresia myosuroides dominated communities are generally cooler than air temperatures by at least 3~4°C, while in Carex scirpoidea and Carex scirpoidea/Carex capitata dominated communities at the same aspect, these temperatures are often greater than air temperatures. This is particularly evident in transects E and

F. The most southern transect (D) had subsurface temperatures under Kobresia myosuroides vegetation (group D1) equalling or slightly greater than air temperatures, while transitional and

Carex scirpoidea communities (groups D2 and D3) had values often

2-5°C higher than air temperatures.

Humidity readings were taken concurrently with air temperature readings at 2, 10, and 20 cm above the ground surface. On any given day, percent humidity was relatively 145

constant between transects - reflecting the mesoclimatic humidity status. At any given site, humidity generally

increased with proximity to the vegetation cover - a difference averaging 2-5% between 2 and 20 cm above the surface.

Complete data for soil moisture levels through the 1980 growing season are regrettably not available. Samples collected in June and July were weighed in the field and subsequently stored in plastic bags at room temperature until the end of the field season. Many of the waterproof tags used to identify samples were unreadable by this time due to the intense microbial activity afforded by these warm, moist conditions.

The few results available indicate that the highest moisture levels were present in A horizon soils (likely due to the high amounts of organic matter) and the greatest loss of moisture occurred in the week following snowmelt. No major differences between transects or vegetation groups were discernible. 146

V. DISCUSSION

This study has examined selected alpine plant communities of Lakeview Mountain, as well as the corresponding abiotic factors of soils and microclimate. Interspecific associations in each community and sampled site have been a major focus, with phenological patterns also measured. This discussion will consider various aspects of the vegetation, beginning with communities and associated abiotic parameters, then phenology, and, finally, interspecific associations. Soils and microclimatic data will then briefly be discussed.

COMMUNITIES

The most extensive communities or vegetation types with early snowmelt in the study area were sampled. Subsequent analysis of percentage cover data revealed three major community types. These were dominated by either Kobresia myosuroides,

Carex scirpoidea (with one transitional area dominated by both

Kobresia myosuroides and Carex scirpoidea), or by Carex sc i rpoidea and Carex capitata (with Salix nivalis as an additional dominant at one site). Communities dominated by both

Carex capitata and Carex scirpoidea were the most common type.

Multivariate analyses of composite data from all six transects

(Figs. 10-13) indicates the high degree of compositional similarity between different stands of the same community, although some variability is evident. In other words, no sample groups occur which would correspond to individual sampled transects. Samples dominated solely by Carex scirpoidea are an 147

exception, and some separation of groups D3 and E2 occurs in the ordination scatter plots. This is possibly due to the higher cover of Cornicularia aculeata and Silene acaulis, higher total plant cover, and greater species richness in E2.

None of the dominants in the sampled communities have a conspicuous associated flora, although species abundances and overall species richness do change. Average species richness is lowest in Kobresia myosuroides communities (29 species), compared to the remaining communities, which have similar numbers of species (mean 42). The dense, tussock-forming growth form of Kobresia myosuroides may restrict the ability of other species to colonize a site.

The distribution of these communities has no evident correlation with aspect. This is possibly because of the moderate slopes (7-16%) of sampled transects. As well, community distribution appears to have little relationship with measured soil parameters. Soils from each vegetation group were analyzed for a number of physical and chemical properties. PCA of A horizon data indicates three vegetation groups, B, D1, and

D2, which are outlying from the main point cluster due to relatively high values for exchangeable cations and total cation exchange capacity. Each of these three groups, however, represents a different community type, as do outlying groups in the analysis of B horizon data. High levels of available phosphorus occurred within the B horizons of two sites: one dominated by Carex scirpoidea and Carex capitata alone (group

F2), and one with the additional dominant, Salix nivalis (C1). 1 48

The reason for these differences is not apparent at the present t ime.

Little relationship between community type and climatic factors were detected. Air temperatures at both the meso and microclimate scale varied only slightly between sites, with no aspect distinctions. Subsurface soil temperatures (2 cm depth) are an exception, and were generally 3-4°C cooler than air temperatures in Kobresia myosuroides communities. Temperatures at a depth of 2 cm were often greater than prevailing air temperatures in the remaining sampled communities. The Kobresia myosuroides community at S2°E had subsurface temperatures relatively higher than at other aspects of this community, although still much lower than in the adjacent Carex sc irpoidea community. The high insulative capacity of the dense Kobresia myosuroides growth form is a likely causal factor. Patterns of wind speed and humidity were essentially uniform over the study area.

The distribution of communities within the study site appears to have little relationship to the aspect, soil, and climatic factors measured. This contrasts with previous studies in similar environments. For example, Bell (1973) and Douglas and Bliss (1977) reported that Kobresia myosuroides will grow only on wind swept sites that are essentially snowfree in winter, however, in this study, all communities (including

Kobresia myosuroides) became snowfree at approximately the same time in 1980 (June 18-20), were uniformly covered with snow before this time, and were not particularly wind swept. Douglas 1 49

and Bliss (1977), also reported that Carex capitata dominated communities occur on snowbed sites with low snow accumulation and moist soils in the eastern North Cascades, while Carex

scirpoidea dominates on dry, well-drained slopes and becomes

snowfree earlier. These environmental bases for community distribution were not substantiated in this study. The particular environmental requirements reported by Douglas and

Bliss (1977) are difficult to reconcile, especially since Carex

scirpoidea and Carex capitata are often co-dominants in the same community.

Major (1951) has stated that the vegetation of a region is a function of mesoclimate, parent materials, topography, biota, and time - the same factors Jenny (1941) related to soil properties, and a rather restricted version of Billings (1952).

Evidence presently available indicates that the abiotic variables listed by Major (1951) are essentially constant within the study area, with the exception of higher amounts of certain soil nutrients at a few sites. As different community types occurred at each of these sites, nutrient differences at the scale measured appear to have little influence on the distribution of these plant communities. It is therefore appropriate to consider the remaining biotic factors, which include neighbour relationships (considered in this study), predator/prey interactions, and grazing. Elements of chance

(seed dispersal, germination, establishment, etc.), local disturbance, and micro-scale differences in soil nutrients may also be important, but are outside the scope of the present 150

study.

PHENOLOGY

Alpine plants tend to flower and go through other phenological stages very quickly (Bliss 1971), aided by the almost universal production of overwintering floral primordia

(Mark 1965, 1970; Billings 1974b). Initiation of growth often occurs before the winter snow cover has completely melted, at or near freezing temperatures (Billings and Bliss 1959, Mooney and

Billings 1961). In the present study, Draba incerta, D. paysonii, and D. lonchocarpa begin growth before snowmelt, and this may be responsible for their rapid flowering. Most other species, however, flowered 15-40 days after snow release, slightly later than the average reported by Billings (1974b) for alpine species in general (10-20 days) and by Douglas and Bliss

(1977) for western Cascade alpine species (14-20 days). The period from floral expansion to seed dispersal in this study

(28-70 days) is less restricted than the period previously reported for most alpine species (28-35 days) (Holway and Ward

1965) and also for western Cascade alpine species (18-44 days)

(Douglas and Bliss 1977). The relatively early snowmelt and subsequently longer growing season of the eastern Cascade alpine, compared to the western Cascades and other alpine areas, may have some influence on these discrepancies. The degree of snow accumulation and time of melt is largely responsible for reported variations in alpine phenology (Bliss 1956, Holway and

Ward 1963, 1965; Mark 1970). In this study, however, 151

phenological differences are due to other factors, as all sites became snowfree at approximately the same time.

The phenological progression of a species may be influenced by the community type in which it is resident. For example, a number of species (e.g., Carex sc i rpoidea, Carex nardina,

Potentilla diversifolia) flower later (10-20 days) and have other phenological stages similarly delayed in vegetation dominated by Kobresia myosuroides, when compared to Carex scirpoidea, or Carex scirpoidea/Carex capitata communities at the same aspect. The identity of these species often varies between sites (e.g., Senecio lugens in transect E, Cerastium beeringianum in transect F). A similar phenological delay was observed for some species within communities dominated by Carex scirpoidea and Carex capitata, when compared to 'pure' Carex scirpoidea communities. These two community types were not sampled at the same aspect, although group E2 (N16°W) and C2

(N29°E) are both north facing. Species flowering later in the

Carex sc irpoidea/Carex capitata dominated vegetation of C2 include Cerastium beeringianum and Potentilla diversi folia (10 days later), and Carex nardina and Carex scirpoidea (20 days later). The latter dominant species also flowers comparatively later in the remaining Carex sc irpoidea/Carex capitata communities as well. The phenology of species within this community type remained constant where Salix nivalis was an additional dominant (group C1).

While aspect appears to have little influence on the distribution of communities, it does, in some cases, influence 152

the timing of phenological stages. A number of species tend to

flower earlier in south facing communities. These species often differ between community types, although Potentilla diversifolia and Silene acaulis consistently flower and/or fruit 4-12 days earlier at the most southerly aspect sampled for a particular community. Other species such as Carex capitata show little phenological variation due to aspect.

Relationships between plant phenology and interspecific association will be considered in the following section, as will other potential association-generating factors. The possible role of species interactions in determining community distribution and structure will also be discussed.

INTERSPECIFIC ASSOCIATIONS

Small scale patterns in the form of significant associations between species pairs were detected in all communities sampled. A total of 182 different significant positive associations and 103 different significant negative associations were found. Of the species sampled, 88% had at least one association in one vegetation group. The total number of associations, however, comprises only 6.8% of all possible species-pair combinations (summed over all vegetation groups).

Associations varied considerably between and within communities.

The associations are probably quite stable (i.e., persistent), due to the age of these communities and the slow rate of vegetative spread characteristic of alpine plants (Billings

1974a, Bell and Bliss 1979). 153

The intensity of patterning at this scale is rather

remarkable considering previous views held by ecologists (Savile

1960, Bliss 1962, Whittaker 1975), who suggested that species within arctic and alpine communities are essentially random in

their distribution. Clearly, it is not possible to make conclusive statements about the processes generating these patterns from associations alone - controlled experimental procedures are necessary for definitive interpretations.

Associations do, however, present some preliminary implications.

A positive association indicates that two species are often physical neighbours and potentially in each others sphere of

influence. Positive association has been equated to the term

'cohabitation', by Harper et a_l. (1961), which is defined as a proximity such that interaction between the two species is considered plausible. This suggests a sharing of habitat or resources (Werner 1979). Coexistence is a similar term, but

implies continuous interaction between species (Aarssen 1983).

Co-occurrence, is defined by Werner (1979), as a mutual presence of species only, and will be used in this thesis as no a priori assumptions based on proximity alone are implied. Negative associations indicate a frequent spatial separation of two species with a consequent infrequency of potential interactions.

Negative association has often been interpreted as avoidance of interaction (Aarssen 1983), or as a result of past competition

(MacArthur 1972, May 1974, Schoener 1975). There is little evidence, however, demonstrating that species divergence has resulted from coevolution of competitors. Observed patterns may 1 54

be explained by a variety of alternate processes, such as predation, response to disturbance or climatic fluctuations, preadaptation to differing resource levels, differential

tolerance of environmental extremes, etc. (Harper 1969; Connell

1975, 1980; Wiens 1977).

Certain associations suggest, by their very nature, potential causalities, although clear-cut explanations based solely on sampling data are not possible. The identification of possible processes or mechanisms creating and/or maintaining associations will lead, ultimately, to the erection of new hypotheses. Positive and negative associations at a given site may be collectively due to a wide variety of different factors and each species pair must be considered separately.

Possible Mechanisms Generating Positive Association

1. Niche Differentiation:

Species may have sufficient differences in their ecology

(i.e., separation on one or more niche dimensions) which allow them to co-occur. This is similar to the term "ecological combining ability" used by Harper (1964, 1967, 1977b) in discussions of coexistence. Niche separation functions in reducing competition and increasing the individual productivity of two co-occurring species, although niche overlap does not necessarily confer competition if resources are plentiful

(Pianka 1979). Resource partioning among a wide variety of co- occurring species has been reviewed by MacArthur (1972),

Schoener (1974a,b), and Pianka (1976). Definite ecological 155

differences have been reported for virtually all co-occurring species studied, and include differences in nutrient requirements, use of space, and temporal activity (Pianka 1979).

Niche complementarity often occurs between different species, and may result in a given population suppressing its own growth more than the growth of other species (Cole 1960). Separation can occur on a wide variety of niche dimensions, which may be morphological or physiological in nature. Several possibilities within the alpine zone are considered here, a) Temporal Partioning:

Asynchronous phenologies (e.g., timing of flowering, fruiting, etc.) may allow species to make their greatest resource demands on the environment at different times allowing co-occurrence. Concurrent differences in the seasonality of growth has been reported for positively associated grassland species (Turkington and Harper 1979c,

Fowler and Antonovics 1981). As well, phenological variation may account for the co-occurrence of species in semi-desert

(Bykov 1974), tropical rainforest (Medway 1972, Frankie et al. 1974), deciduous woodland (Grubb 1977), and understory herb communities (Bratton 1976). The characteristic late maturation of polyploids may provide the niche separation necessary for the co-occurrence of intraspecific individuals with differing ploidy levels (Lewis 1976). The present study has provided little evidence that positive associations among alpine species are maintained by temporal partioning. A few isolated possibilities do occur, however. For example, Solidago multiradiata has 156

little or no overlap of flowering period with positive associates. A similar separation of flowering times was observed for the positively associated Arenaria obtusiloba and

Draba lonchocarpa (although other Draba species with congruent phenologies are not positively associated with Arenaria obtusiloba). As the alpine growing season is very short, little opportunity for major phenological differences exists, b) Morphology:

As a large proportion of alpine plant biomass is underground (Bliss 1966, Webber 1974, Webber and May 1977), differences in root morphology may be an important factor allowing niche separation of alpine species. Dissimilarities in size, shape, arrangement, and depth of roots may result in two species exploiting different volumes of soil and, concurrently, different nutrient and moisture pools, while growing together.

This has been postulated for species in old fields (Wieland and

Bazzaz 1975, Parrish and Bazzaz 1976), and pastures (Berendse

1979). In the present study, species with thick, branching taproots such as Arenaria obtusiloba, Oxytropis monticola, and

Erigeron aureus are positively associated with the fibrous- rooted Festuca ovina at some sites. In addition, the rhizomatous Carex scirpoidea produces fibrous-rooted shoots and is a common positive associate of Arenaria obtusiloba, Oxytropis monticola, Potentilla diversifolia, Silene acaulis, Solidago multiradiata, Penstemon procerus, and Aqoseris glauca - all variously taprooted. Many more examples exist. In contrast, very few positive associations occur between similarly fibrous- 157

rooted monocots.

Niche separation may also occur when differences in alpine

floral morphology allow partioning of specific pollinators

(Billings 1974b, Macior 1974). This is a strong possibility in

the alpine zone, where insect-mediated pollination is common

(Petersen 1977) and pollinator diversity and abundance is low

(Holway and Ward 1965, Brinck 1974, Moldenke 1976). Although partioning of pollinators has been reported for plant species in more temperate climates (Reader 1975, Parrish and Bazzaz 1978), no data are available for the contribution of this factor to positive alpine associations, c) Physiology:

It seems very likely that physiological variation between alpine species may provide an opportunity for co-occurrence, as has been suggested for lowland species. For example, co- dominant annuals in a successional field were found to differ in their daily patterns of leaf water potential, light saturation points, and rates of photosynthesis (Wieland and Bazzaz 1975).

In addition, different nutrient levels in co-occurring forest species, including species in the herb layer (Siccama et. al. 1970), have been postulated as expressions of niche differentiation, due to differences in nutrient uptake rates, use of soil space, or in the type of nutrient sources exploited

(Woodwell 1974, Woodwell et al. 1975).

A substantial amount of physiological variation has been reported for alpine species, resulting in differential nutrient requirements, rates of uptake, and tissue nutrient levels 158

(Larcher et a_l. 1975, Rehder 1976), dissimilar storage and mobilization rates for carbohydrates and lipids (Mooney and

Billings 1960, Hadley and Bliss 1964), and differences in photosynthetic and respiration rates and temperature optimums

(Mooney et al. 1964, Bliss and Hadley 1964, Scott and Billings

1964, Mark 1975). Physiological differences between immediate neighbours in the alpine zone have yet to be studied.

2. Balanced Competitive Abilities;

Co-occurrence as the result of niche separation is a commonly held view (MacArthur 1972, Newman 1982b) and is a corollary of Gause's competitive exclusion principle (Gause

1934, Hardin 1960), which states,that two species with identical ecological niches will not coexist. Theoretical conclusions based primarily on competitive exclusion within animal populations in laboratory culture, for example, Paramec ium

(Gause 1934), and Drosophila (Ayala et §_1. 1973), may have little relationship to plants. Plants have the potential for more subtle responses to the considerable niche overlap present between species at a given site. Aarssen (1983) has proposed that selection for a balancing of competitive abilities between species pairs may be common. This would act to reduce intraspecific competition among individuals of a strong competitor, and would also prevent competitive exclusion of a slightly weaker competitor. Although no direct evidence is available to substantiate this mechanism in the present study, it is a possibility, considering the age of these alpine 159

communities and the large number of positive associations detected (79 more positive than negative).

3. Additional Mechanisms:

a) Predation:

Predation, such as selective grazing, may lead to the suppression of a superior competitor and allow co-occurrence with a weaker competitor (Harper 1969, Grime 1979). At this study site, marmots have been observed to selectively graze floral heads of Oxytropis monticola. Only controlled experiments could help determine if this contributes to the subordinate community role or positive associations of this species. Specific grazing habits of other animals were not observed. b) Available Nutrients:

Three vegetation groups in the study area (B, D1, D2) were found to have relatively high amounts of exchangeable cations in the A horizon. This indicates large scale differences, as composite samples, taken from various points in each soil pit, were used for nutrient analysis. Trends within the significant positive associations detected do not appear to reflect these higher nutrient levels, although two exceptions were noted.

Carex sc i rpoidea and Solidago multiradiata are positively associated in these three vegetation groups only, while the positive association between Kobresia myosuroides and Potentilla diversifolia is restricted to two of these groups (D1 and D2).

Differences at a small scale (e.g., centimeters) may have a much 160

greater influence on the formation of positive associations.

Certain species, through the action of symbionts, may

locally increase the levels or availability of nutrients,

inducing positive association with other plant species. This could be particularly important in the alpine zone due to the

low nutrient status of alpine soils, the slow rate of nutrient absorption and assimilation in alpine species, and the reduced microbial activity present under low soil temperatures (Nimlos et al. 1965, Bamberg and Major 1968, McCown 1975). Nitrogen and phosphorus are the major limiting nutrients in alpine species

(Rehder 1976a), with a steady decrease of these nutrients occurring in plant tissues over the growing season (Mooney and

Billings 1961).

It is well known that many legume species have Rhizobium nodules on their roots which fix atmospheric nitrogen (Corby

1981). Nodules were observed on the roots of Lupinus lyallii and Oxytropis monticola seedlings. These species were relatively infrequent in the communities sampled and did not have an exceptionally large number of positive associations. No negative associations occurred between these and other vascular species, however, with the exception of Lupinus lyallii and

Carex capitata in group CI. As Lupinus lyallii is positively, and Carex capitata is negatively, associated with Salix nivalis in this group, this could be a secondary result of the Carex capitata/Salix nivalis negative association.

Peltigera canina is the only lichen containing nitrogen- fixing blue-green algae (Nostoc) present at this study site. 161

This lichen significantly contributes to nitrogen stores in many tundra communities (Forman and Dowden 1977, Alexander et al. 1 978). Peltigera canina is positively associated with only

4 different species (2 vascular and 2 lichen) and has no negative associations - possibly due to the infrequent occurrence of this species. Blue-green algae such as Nostoc often occur with moss in moist tundra sites (Granhall and

Selander 1973, Alexander et a_l. 1978) and significant nitrogen fixation has been found even in relatively dry, meadow sites in the tundra (Schell and Alexander 1973). It does not seem likely that moss species sampled in this study were affiliated with large amounts of blue-green algae, because of the predominance of negative associations between moss and vascular species.

Lichens without blue-green algal symbionts have been reported to function more generally as nutrient traps accummulating minerals such as phosphorus and transforming calcium into an available form (Syers and Iskander 1973).

Minerals are leached out of living lichen thalli and released after lichen decomposition (Milbank and Kershaw 1973). This process may contribute to the frequent positive associations observed between certain vascular species and lichens. As lichens occur at the soil surface only, they occupy an environment very different from the deep rooting vascular plants. Positive vascular/lichen associations may therefore result where competitive exclusion of other vascular species has occurred, and positive associations between different lichen species may occur secondarily within such zones of 'vascular 162

exclusion'.

Mycorrhizal root infections are found in some tundra species (Haselwandter and Read 1980, Read and Haselwandter 1981) and benefits to the host plant are well known, particularly where nutrient levels are low (Harley 1969, Sanders et al.

1975). Mycorrhiza may also enhance the nutrient status of neighbouring, non-infected species, through local increases in soluble nutrients. Mycorrhizal infections (endophytic) have been reported in alpine ericaceous species (Pearson and Read

1973, Haselwandter and Read 1980), in a variety of herbaceous alpine species in Austria (including the genera Silene,

Potentilla, Carex, Dryas, Arenaria, and Cerastium) (Read and

Haselwandter 1981), and in almost all arctic and alpine shrub and dwarf Salix species in Alaska (Miller and Laursen 1978,

Laursen and Chmielewski 1980, Linkins and Antibus 1980).

Mycorrhizal infection of Salix nivalis may be a factor contributing to the large number of positive vascular associations observed for this species. Only lichens and Carex capitata were negatively associated with Salix nivalis. In addition, increased levels of phosphorus occurred in the B horizon soils of vegetation where Salix nivalis was one of the dominant species. Ectomycorrhizal infections have been reported for Kobresia myosuroides in European alpine areas (Fontana 1963,

1977; Haselwandter and Read 1980; Read and Haselwandter 1981), a species with few positive vascular associations in this study.

A highly competitive species may gain the most benifit from local increases in nutrients. As well, differential rates of 1 63

absorption may occur for different types of mycorrhizal fungi,

although these are not known (Haselwandter and Read 1980).

c) Microenvironmental Effects:

Certain species may modify microsite conditions and thereby

enhance the growth, and particularly the germination and

establishment, of other plant species (Harper 1964) - events

which occur infrequently in the alpine (Billings 1974b). This

is another possible explanation for the positive associations of

Salix nivalis. This species has leaves orientated close to the

ground and an open growth form, with 'gaps' of bare soil

(demonstrated by association analysis (Table 4)) between leaves.

This configuration produces a shady micro-environment with

relatively high humidity and moisture levels, warmer

temperatures, and protection from wind. The ubiquitous positive association between Festuca ovina and Silene acaulis may also be due to the protection and warmer temperatures afforded by this cushion species. Seedling and small Festuca ovina individuals were growing within most Silene acaulis plants observed during this study. Seedlings within cushion species such as Silene acaulis and Diapensia lapponica L. have been previously reported

(Griggs 1956, Bliss 1963). The occurrence of moss species in dry sites has also been linked to the microsite conditions produced by the vascular canopy - particularly with respect to reduced water losses (Stoner e_t al. 1978), and may contribute to positive association between mosses and vascular species.

A wide variety of possible factors may contribute to positive associations between species. As extensive testing of 164

these and other hypotheses is required before cause and effect

relationships can be established, the preceding interpretations

are necessarily speculative in nature.

Possible Mechanisms Generating Negative Association

1. Morphology:

Species with a clumped growth form, such as the dense,

tussock-forming Kobresia myosuroides, will have relatively fewer

opportunities to grow next to members of a different species

than would a species with a more open growth form. This may

result in negative associations with species less able to

'combine' with the clumped individual. Species with very

similar morphologies - particularly with respect to roots - may

also be infrequent neighbours. For example, taprooted species

rarely form positive associations with each other and

occasionally form negative associations (e.g., Arenaria

obtusiloba and Silene acaulis in group B). This trend is also apparent with fibrous-rooted species, such as Festuca ovina and

Carex nardina.

2. Abiotic Effects:

Differences in resource requirements and environmental tolerances between species may form the basis for negative association within any given community. Previous workers with low elevation species have reported that microtopography

(Struick and Curtis 1962, Bratton 1976), soil disturbance

(Fitter 1982), soil depth (Kershaw 1958), and heterogeneity of 165

soil pH (Dowries and Beckwith 1951), and soil minerals in

grassland (Snaydon 1962), chalk heath (Grubb et al. 1969), and

limestone heath (Etherington 1981) can affect plant distribution

over distances as short as 60 cm. In addition, differences in

site requirements for seed germination and establishment (Grubb

1977), tolerance of soil chemicals, particularly at germination

(Williams and Harper 1965, Williams 1969), and nutrient

requirements (Harper et al. 1961), can result in the spatial

separation of species at a given site. The soil and climatic parameters sampled in the present study differed little between

sites, with the exception of higher levels of exchangeable cations in the A horizon of groups B, D1, and D2. These

relatively large scale differences were not reflected in the

significant negative associations detected. No negative associations are restricted to these vegetation groups, or, conversely, conspicuously absent from these groups. There is, therefore, no indication that increases in nutrient levels at this scale reduces the number of negative associations.

Investigation at a much finer scale, however, may indicate differentiation of these factors at the level of the individual plant. Negative association may be influenced at this finer scale.

3. Competitive Exclusion:

Species or individuals with high competitive abilities may exclude certain other species from their immediate surroundings.

A negative association between the stronger and the weaker competitors may then result. Past competition cannot be 166

conclusively inferred from negative association alone, however, as other factors may be responsible (Connell 1975, 1980). In extreme cases, species diversity at any particular site may be lowered due to competitive exclusion (Grime 1973a). An endless list of physiological and/or morphological characters can confer an increase in relative competitive ability for plant species in general. These include such factors as more efficient uptake of nutrients, increased growth rate and size (Black 1960), large root systems and/or associated mycorrhiza, deposition of organic material unfavorable to other species (Grime 1973b), and release of allelopathic chemicals (Rice 1974). In extreme environments such as the alpine, relative growth rates, seed production, and/or seedling survival, and subsequent competitive ability, may be increased for species best able to withstand unfavorable conditions (e.g., drought, freezing, intense U.V., etc.) due to a specialized morphology, plasticity, or general resistance.

Kobresia myosuroides is an extreme example, with high tolerance of both drought and cold (Ehle.ringer and Miller 1975) and a readiness to grow even under subfreezing conditions. Leaf elongation in this species occurs during winter periods near 0°C

(Bell and Bliss 1979). Other herbaceous winter-green species have been reported (Bell 1974), although the growth of most alpine species is restricted to temperatures above 0°C (Holway and Ward 1963, Billings 1974b). Experimental data on the relative competitive abilities of alpine species is very limited, although a small number of studies suggest that competition may be a significant factor in this zone. For 167

example, certain rosette species such as Draba cannot grow when heavily shaded (Billings 1974a), Thalspi alpestre L. individuals become larger when growing in isolation (Rochow 1970), and in

the Norwegian alpine tundra, only single tillers of Carex bigelowii Torr. individuals invade each small region of adjacent

soil (with frequent abortion of newly initiated ramets), which

is interpreted as a competition-avoidance strategy (Callaghan

1976). This is in direct contrast to Grime's (1979) categorization of alpine species as S-strategists. Grime (1979.) states that S-strategists are species able to withstand extreme environmental stress (an anthropomorphic viewpoint), and must necessarily be weak or non-competitors. The primarily vegetative regeneration strategy of alpine plants (as S- strategists) does not, however, preclude the existence of competition.

As sampling occurred at a very fine scale (every 2 cm) in this study, it is conceivable that secondary negative associations are generated. For example, species that are negatively associated with Carex capitata often have additional negative associations with various species of lichen. Possibly, these additional negative associations are generated because the lichen species are positively associated with Carex capitata.

Cole (1957) termed this secondary type of association as

"partial" association in a discussion of species co-occurrence within quadrats. 168

Genotypic Response

Positive and negative species associations vary

considerably between sampled sites in the study area. Species

pairs occurring together at a number of sites are rarely

associated in all of them. As well, only a small number of

interspecific associations had any relationship to community or

aspect. These data suggest that population differentiation of

alpine species occurs over very short distances. The genotype

or individual and not the taxonomic species may be the critical

ecological unit in the alpine zone, as has been argued by Harper

(1977a,b, 1982) and Aarssen (1983) for lowland species.

Genotypes may form associations with a variety of different

species, depending more on certain characteristics of the

genotypes initially present, and less on the taxonomic or

specific identity of that genotype.

Fine-scale biotic specialization between genotypes of

temperate species has been reported by Allard and Adams (1969),

Turkington and Harper (1979c), Joy and Laitinen (1980), and

Aarssen (1983). In these studies, increased total yield was obtained from clones of immediate neighbours, compared to non- neighbouring clones of the same two species. Genotypes which have differentiated as a result of evolutionary co-adaptation may be termed "biotic ecotypes" (Aarssen 1983).

The importance of genetic differences within a species,

including the degree of plasticity (Jacob and Monod 1961), and their relationship to environmental factors (abiotic ecotypes) has long been recognized (Turesson 1922, 1923; Heslop-Harrison 169

1964, Langlet 1971). Alpine ecotypes are common in species with

arctic/alpine distributions, and differ in physiological factors

such as photosynthetic and respiration rates, photosynthetic

light saturation points, metabolic acclimation to temperature,

etc. (Mooney and Billings 1961, McCown and Tieszen 1972,

Tranquillini 1964, Tiezen and Wieland 1975, Billings e_t al. 1971, Billings 1974a). Differentiation of alpine plant populations, attributed to snow cover, temperature, and length of growing season, have been reported on a more local scale after transplant studies along elevational gradients (Clausen e_t al. 1948, Ward 1969, Rochow 1970, Pearcy and Ward 1972). On an even finer, microenvironmental scale, ecotypes from a variety of different habitats in a given area have been reported for tundra species, differing in phenology (Holway and Ward 1965, May

1976), seed germination requirements (Amen and Bonde 1964), relative rates of seed germination (Sayers and Ward 1966), and competitive abilities (Shaver e_t a_l. 1979). These genetically based differences are attributed exclusively to environmental factors in these studies.

The site specific responses (in the form of positive and negative associations) of species in the present study, and the lack of major environmental differences between sites, suggests that biotic specialization or interaction between genotypes may also be responsible for population differentiation in the alpine zone. The unique selection pressures (e.g., differences in grazing, disturbance, etc.) operating during the history of a site will, of course, be influencing factors. Fine scale 170

genetic variation between neighbouring alpine species has yet to

be studied experimentally.

Dominant Species

The variability of species associations between different

sampled stands of the same community or vegetation type has been

noted. The similarity of these stands, however, is defined chiefly on the basis of the dominant or most abundant species.

It is inferred by the use of the term 'dominant', that the most

abundant species also acquire the greatest proportion of

resources available to the community (McNaughton and Wolf 1970), and therefore possess the greatest competitive ability within

that community. Observations of low elevation dominants has led

to the suggestion that these species also have broader niches

than species with lower biomass and density (Levins 1968,

McNaughton and Wolf 1970, Parrish and Bazzaz 1976). It is very probable that the narrower realized niche of subordinate species

is at least partially due to the presence of neighbouring dominants. For example, removal of dominant grass species in an old field increased the net productivity of less abundant

species by almost three-fold (Pinder 1975), and a similarly dramatic effect on the growth of sand dune annuals was attributed to neighbouring species (Mack and Harper 1977).

The three major dominant species in the present study were observed to have fewer positive and negative associations than many other less abundant species in the community, although similar numbers of associations were also observed. In 171

addition, most negative associations between vascular plants involve a dominant species. Insight into the relative competitive abilities of these three dominant species may be obtained by examining the interspecific associations and phenology of each under varying conditions. These data suggest a competitive hierarchy of dominant species within the study area, with Kobresia myosuroides as the most competitive, compared to all other species, followed by Carex capitata, and finally by Carex scirpoidea. This argument will be supported by the following examples:

1) Kobresia myosuroides:

No negative associations occur between vascular species in

Kobresia myosuroides communities, other than those of the dominant species with Carex capitata, and with Carex scirpoidea, where they co-dominate in group D2. Positive associations between Kobresia myosuroides and vascular species are rare, with lichen associations more common. As well, a relatively large number (5) of vascular species are positively associated with bare ground in the largest data set that Kobresia myosuroides dominates (E1).

Kobresia myosuroides dominated vegetation has abrupt transitions with other communities, with Kobresia myosuroides as the sole dominant in all but extremely rare and localized instances. The total number of different associations is low in some Kobresia myosuroides dominated stands, but this is likely due to the small number of sampling points in these data sets, as well as the comparatively lower species richness previously 1 72

discussed.

The delay of phenological stages for some species within

Kobresia myosuroides communities has previously been noted. A consistent lack of negative association is present between these and the dominant species. Suppression of Carex scirpoidea phenology has been observed at north facing aspects of this community. The phenology of Carex capitata, however, remained unchanged in Kobresia myosuroides communities.

2) Carex capitata:

This species is present as a co-dominant only at sites also co-dominated by Carex scirpoidea. The vast majority of positive

Carex capitata associations involve lichen species, while negative associations with vascular species predominate. Carex capitata is negatively associated with Carex scirpoidea where they co-dominate (except transect C) and positively associated when the two species occur together in Kobresia myosuroides communities. Carex capitata is also strongly negative with

Salix nivalis where this species in an additional dominant.

The later flowering times of some species in Carex sc i rpoidea/Carex capitata communities, compared to Carex scirpoidea communities, has been noted. Of interest is the delayed flowering of Carex scirpoidea where it co-dominates with

Carex capitata.

3) Carex scirpoidea:

This species has very few associations where it is the sole vascular dominant. It has a relatively larger number in communities co-dominated by Carex scirpoidea and Carex capitata 173

or dominated by Kobresia myosuroides, and associations detected

here may be specifically excluded from Carex scirpoidea

dominated vegetation, even when the former associate is

abundant. On average, less abundant species in Carex scirpoidea

communities are involved in more associations than the dominant

plant. Carex scirpoidea is rarely negatively associated with

vascular • species other than Carex capitata and Kobresia

myosuroides, and has no positive lichen associations, in

contrast to the other dominant species.

These observations suggest that Carex scirpoidea is the

least competitive of the dominant species, as evidenced by the

lack of negative association with, and phenological suppression

of, subordinate vascular species, as well as the lack of

positive lichen and moss associations. Kobresia myosuroides is

indicated as the most competitive, due to its sole dominant

status, as well as the phenological suppression of Carex

scirpoidea and other species, and lower species richness in

communities it dominates. Less strongly competitive subordinate

species appear to have a greater ability to form positive

associations than do the dominant plants.

It is interesting that Carex capitata, and particularly

Carex scirpoidea, are associated with different species, as well as different numbers of species, when they are dominants. This

is further support for the possible existence of biotic ecotypes on a local scale, discussed earlier. Similar negative association between co-dominant species has been reported for old-field annuals and implies niche differentiation (Wieland and 174

Bazzaz 1975), due to preadaptation or possibly competitive exclusion (Connell 1975, 1980).

It is also suggested that the relative competitive abilities of these three major dominants may be the critical

factor influencing the distribution of communities within the study area. Such large scale patterning was previously attributed entirely to abiotic factors in the alpine zone. The degree of competitive dominance is mediated by the environment, but, in an essentially uniform area (as suggested by parameters measured in this study), initial establishment (due to chance, available safe sites, bare soil, etc.) and' subsequent vegetative spread of dominant species may determine community patterning.

Once species are established, little chance for germination or

invasion by other potential dominants' is possible - particularly in dense Kobresia myosuroides communities. This has been termed

'premptive competition' or competition for space, by Werner

(1979). As the likelihood of death is generally greatest for any plant during germination and establishment (Harper 1967,

Sarukhan and Harper 1973), only a dominant surviving past a certain size will be able to suppress or displace other colonizing species (Connell 1975). The spatial pattern of dominant species may then determine the presence and placement of other plants (Werner 1979). A large number of morphological and physiological factors can confer relative competitive dominance in a given site and require experimental study for identification. 175

SOILS

Soils within the study area on Lakeview Mountain have been

characterized by van Ryswk (1969) as Alpine Brown with

discontinuous ash, following the U.S. classification system

(Soil Survey Staff 1975), and by Green and Lord (1979) as Alpine

Dystric Brunisols, following the Soil Survey Committee of Canada

(1968). The coarse texture of soil in this study area, (sandy

loam to sand) is characteristic of alpine soils (Faust and

Nimlos 1968, Sneddon 1969, van Ryswyk 1969, Sneddon et al. 1972,

Luckhurst 1973), as is the large proportion of gravel and stone

(Nimlos et al. 1965).

Soils are strongly acidic, with pH values corresponding closely to those obtained by van Ryswyk (1969). The acidic

nature of other alpine soils has been associated with the presence of volcanic ash (Sneddon 1969) and high levels of organic matter (Bliss 1963). Levels of organic matter were high

in the study area, particularly in A horizons (2 to 5 times as high as B horizons) and were within the range reported by van

Ryswyk (1969) and Douglas and Bliss (1977) for similar soils within the Cascade alpine zone, although higher values were

frequently detected by these authors. Organic matter accumulation is largely due to low temperatures, which restrict oxidation processes (Nimlos and McConnell 1965).

Total nitrogen (%) was, on average, within the range reported by van Ryswyk (1969) for discontinuous ash soils, although some values were slightly higher. Values for nitrogen and other nutrients (except phosphorus) tend to decrease with 176

depth (in correlation with levels of organic matter) in this study area and in other alpine regions (Nimlos and McConnell

1965, Sneddon et al. 1972, Knapik et al. 1973), with overall nutrient levels low compared to more temperate regions (Bliss

1963, Bockheim 1972, Douglas and Bliss 1977). Levels of available phosphorus are within the range reported by Douglas and Bliss (1977) for the western North Cascades, although well below the maximum values published by these authors. In addition, phosphorus increased with depth in the present study, in contrast to the findings of Douglas and Bliss (1977), but in agreement with other alpine areas (Nimlos and McConnell 1965).

Values for exchangeable cations and total cation exchange capacity were also within the range reported by van Ryswyk

(1969) and Douglas and Bliss (1977). Calcium levels, however, were comparatively higher ' than in the latter study. Soil properties were relatively constant between sampled sites on

Lakeview Mountain. The few differences that were detected have been discussed previously.

CLIMATE

As previously outlined, mesoclimate (measured at three weather stations) was relatively constant over the study area during the 1980 growing season (Table 10). Wind speed was comparatively higher at station 1 (above transect C) than at station 2 (below transect D).

Microclimatic air and soil temperature profiles also differed little between sampled sites, with the exception of the 177

cooler subsurface temperatures (2 cm depth) in Kobresia myosuroides dominated vegetation, possibly due to its higher

insulative- capacity. Similar temperature monitoring in the western Cascade alpine (Douglas and Bliss 1977) indicated that

subsurface temperatures were usually higher than air

temperatures in sparsely vegetated areas (fellfields). In addition, increasing vegetation cover was correlated with decreasing temperatures at depths of 2-30 cm by these authors.

Soil temperatures fluctuated with prevailing air temperatures in the present study, with the steepest gradients between soil at depths of 2 cm and 10 cm. This was similarly

reported by Douglas and Bliss (1977). Air and subsurface temperatures reached a recorded maximum of 24°C at one transect

(E) and minimum temperatures near 0°C were noted for most sites

in early July, 1980. Temperatures tended to increase as the growing season progressed. Much higher maximum subsurface temperatures have been reported previously for subalpine and alpine areas in western North America (35-49°C), but vegetative cover at measured sites was often lower than in the present study (Bamberg and Major 1968, Ballard 1972, Douglas and Bliss

1977). Soil temperatures were measured at 50 cm below the surface by van Ryswyk (1969) in Alpine Brown, discontinuous ash soils of the study area during 1965 and 1966. Temperatures ranged from 6-8°C over the growing season, with a recorded maximum of 10°C (August) and minimum of 1°C (October). These temperatures are very similar to those obtained at a depth of 20 cm in the present study. Winter depth of frost penetration was 1 78

estimated to reach 180 cm in these soils, and has an inverse

relation to the degree of snow accumulation (van Ryswyk 1969).

CONCLUSIONS

Microscale patterns in the vegetation were detected.

Preliminary studies suggest that the distribution of communities within the study area are possibly related to some of the measured abiotic factors. It is possible that biotic factors, such as the relative competitive abilities of dominant species, and the resultant vegetative spread of each, may play a role.

The distributional patterns are unrelated to aspect.

The phenology of some alpine species appears to be influenced by the dominant species they occur with, or by the aspect at which they grow. Phenological differences between species had little relationship to interspecific associations.

The large number of interspecific associations detected belie previous views held by alpine ecologists on the lack of patterning at this scale. Patterns within communities were assumed non-existent, and species distributions essentially random, while large-scale community patterns were related to abiotic factors. Although conclusive explanations for the existence of these associations are not possible with sampling data alone, trends within these data indicate that temporal partioning and symbiotic nitrogen fixation in legume and lichen species have little influence. Species morphology, particularly with respect to roots, and microenvironmental effects (e.g., seedlings in cushion plants) appear to be more important 1 79

factors. As a wide variety of possible processes or mechanisms

may contribute to the formation of species associations,

extensive testing of the hypotheses and interpretations

presented in this thesis is required before cause and effect

relationships can be established.

Interspecific associations varied considerably over the

study area, with many associations detected only once.

Associations of particular plant species often changed at each

sampled site. This suggests that population differentiation of

alpine species occurs over very short distances. The formation

of associations may depend more on certain characteristics of

the genotypes initially present, and less on the taxonomic or

specific identity of that genotype. As well, less strongly

competitive subordinate species form more positive associations

than do dominant plants. A high degree of competitive ability

is suspected for dominant species that have frequent positive

associations with lichen species and few positive (and many

negative) associations with vascular species.

The plotless point-line, or species juxtapositions, method

used to detect interspecific association was found to be useful

for low-growing alpine vegetation, interspersed with bare

patches or rock. The distance between points is flexible, and

the recording of species sequences is objective. The necessity

of a plotless technique was apparent after a comparison of

resulting small scale associations, with 'those detected using quadrat data. One problem with this and other existing plotless methods, is that results are limited to interspecific 180

associations only. This is primarily due to the extreme difficulty in determining individual plants in closed vegetation patches. As intraspecific association may provide important preliminary data on the intensity of competition between members of the same species, further research is needed to provide an adequate sampling method for this type of association. 181

VI. SUMMARY

VEGETATION

1. Selected alpine plant communities of Lakeview Mountain,

Cathedral Provincial Park were examined, as well as the

corresponding abiotic factors of soils and microclimate.

Multivariate analysis of percentage cover data for 6, 2 m X 30 m

transects revealed three major community types, dominated by

Kobresia myosuroides, Carex scirpoidea (with one transitional

area dominated by both Kobresia myosuroides and Carex

scirpoidea), or by Carex scirpoidea and Carex capitata (with

Salix nivalis as an additional dominant at one site). The

distribution and composition of these communities had little

relationship with aspect or with the soils and microclimatic

parameters measured. Subsurface soil temperatures, however,

were generally 3-4° C cooler in Kobresia myosuroides

communities, than at other sites.

2. Phenological stages for vascular species were recorded

during the summer of 1980, within each sampled transect. A

number of species have delayed phenology in Kobresia myosuroides

communities, compared to other sampled communities. A similar

delay was observed for some species within Carex

scirpoidea/Carex capitata dominated sites, when compared to areas dominated solely by Carex scirpoidea. In addition, a

number of species tend to flower earlier in south facing sites, although other species show little phenological variation with aspect. 182

3. Small scale patterns in the form of significant

associations between species-pairs were detected in all

communities. A plotless point-line sampling technique was used

to determine species juxtapositions (i.e., the transitions

between different species). Subsequent analysis compared

observed species sequences to expected values generated using a

Markov chain model for random distribution. A normally-

distributed standardized residual statistic was used to

determine significant (p^0.05) positive and negative

interspecific associations. This method is most useful for low-

growing vegetation interspersed with bare patches or rock.

Similar plotless techniques, such as the contact sampling

method, depend on continuous ground cover. The necessity of

plotless techniques to determine small scale associations was

apparent after associations at the scale of a 20 X 50 cm quadrat

were found to be very different from those detected at 2 cm

intervals with point-lines.

4. Patterns were abundant at this small scale, with a total of

182 significant positive associations and 103 significant

negative associations recorded between different species-pairs.

Of the species sampled, 88% had at least one association in one

vegetation group (sampled stand). Although it is not possible

to make conclusive statements about the processes generating

these patterns from associations alone, certain associations

suggest potential causalities. Possible mechanisms generating

positive associations have been discussed. Briefly, these are:

i) Niche differentiation - particularly temporal partitioning, 183

differential morphology, and physiological variation. ii) Balancing of competitive abilities, with subsequent prevention of competitive exclusion. iii) Predation, leading to the suppression of superior competitors, allowing co-occurrence with weaker ones. iv) Local increases in available nutrients by certain species through nitrogen-fixing symbionts and mycorrhizal root infections, and v) Modification of microsite conditions by certain species, which proves beneficial for other species.

Possible mechanisms generating negative association were also discussed: i) Similar morphology - particularly with respect to roots

(niche overlap). ii) Differences in resource requirements and environmental tolerances between species. iii) Competitive exclusion of weaker competitiors by stronger ones.

As positive and negative species associations vary considerably between sampled sites in the study area, and few major environmental differences exist between sites, the genotype or individual and not the taxonomic species may be the important ecological unit in the alpine zone.

5. Insight into the relative competitive abilities of the three major dominant species may be obtained by examining the interspecific associations of each under varying conditions, as well as phenological data for subordinate species. Data suggest 184

a competitive hierarchy of dominant species within the study area, with Kobresia myosuroides as the most competitive, compared to all other species, followed by Carex capitata, and, finally, by Carex sc irpoidea. The relative competitive abilities of these three major dominants may be a critical factor affecting the distribution of communities at the study site. Less strongly competitive subordinate species appear to have a greater ability to form associations than do the dominant plants.

SOILS

Soils within the study area are classified as Alpine

Dystric Brunisols, following the Canadian System, and are coarse textured, strongly acidic, and high in organic matter. Low nutrient levels parallel other alpine studies, and are relatively constant between sampled sites.

CLIMATE

Mesoclimate was relatively uniform over the study area during the 1980 growing season, as were microclimatic air and soil temperature profiles and air humidity profiles. Maximum subsurface temperatures were lower than that previously reported for alpine areas in western North America, with the lowest temperatures occurring beneath Kobresia myosuroides dominated vegetation. 185

VII. LITERATURE CITED

Aarssen, L.W. 1983. Interactions and coexistence of species in pasture community evolution. Ph.D. thesis. Univ. of British Columbia, Vancouver, B.C. 249 p.

Aarssen, L.W., R. Turkington, and P.B. Cavers. 1979. Neighbour relationships in grass-legume communities. II. Temporal stability and community evolution. Can. J. Bot. 57: 2695- 2703.

Alexander, V., M. Billington, and D.M. Schell. 1978. Nitrogen fixation in arctic and alpine tundra, p. 539-550. Iri L.L. Tieszen (ed.) Vegetation and Production Ecology of an Alaskan Arctic Tundra. Ecological Studies 29, Springer- Verlag, New York.

Allard, R.W., and J. Adams. 1969. Population studies in predominantly self-pollinating species. XIII. Intergenotypic competition and population structure in barley and wheat. Amer. Natur. 103: 621-645.

Amen, R.D., and E.K. Bonde. 1964. Dormancy and germination in alpine Carex from the Colorado Front Range. Ecology 45: 881-884.

Anderson, J.E., and S.J. McNaughton. 1973. Effects of low soil temperature on transpiration, photosynthesis, leaf relative water content, and growth among elevationally diverse plant populations. Ecology 54: 1220-1233.

Archer, A.C. 1963. Some synecological problems in the alpine zone in Garabaldi Park. M.Sc. thesis. Univ. of British Columbia, Vancouver, B.C. 129 p.

Archibald, E.E.A. 1948. Plant populations. 1. A new application of Neyman's distribution. Ann. Bot. N.S. 12: 221-235.

Arno, F., and J.R. Habeck. 1972. Ecology of alpine larch (Larix lyallii Pari.) in the Pacific Northwest. Ecol. Monogr. 42: 417-450.

Austin, M.P., and P. Greig-Smith. 1968. The application of quantitative methods to vegetation survey. II. Some methodological problems of data from rain forests. J. Ecol, 186

56: 827-844.

Austin, M.P., and I. Noy-Meir. 1971. The problem of non- linearity in ordination: experiments with two-gradient models. J. Ecol. 59: 763-773.

Ayala, F.J., M.E. Gilpin, and J.G. Ehrenfeld. 1973. Competition between species: theoretical models and experimental tests. Theoret. Pop. Biol. 4: 331-355.

Ballard, T.M. 1972. Subalpine soil temperature regimes in southwestern British Columbia. Arct. Alp. Res. 4: 147-166.

Bamberg, S.A., and J. Major. 1968. Ecology of the vegetation and soils associated with calcareous parent materials in 3 alpine regions of Montana, USA. Ecol. Monogr. 38: 127-167.

Baptie, B. 1968. Ecology of the alpine soils of Snow Creek Valley, Banff National Park, Alberta. M.Sc. thesis. Univ. of Calgary, Calgary, Alberta. 135 p.

Barry, R.G., and CC. Van Wie. 1974. Topo- and microclimatology in alpine areas, p. 73-83. I_n J. Ives and R.G. Barry (eds.) Arctic and Alpine Environments. Methuen, London.

Bates, R.G. 1954. Electrometric pH determinations. John Wiley and Sons, Inc., New York.

Beals, E.W. 1973. Ordination: Mathematical elegance and ecological naivete. J. Ecol. 61: 23-35.

Beder, K. 1967. Ecology of the alpine vegetation of Snow Creek Valley, Banff National Park, Alberta. M.Sc. thesis. Univ. of Calgary, Calgary, Alberta. 243 p.

Bell, K.L. 1974. Autumn, winter, and spring phenology of some Colorado alpine plants. Amer. Midi. Nat. 91: 460-464.

Bell, K.L., and L.C. Bliss. 1979. Autecology of Kobresia bellardi i: why winter snow accumulation limits local distribution. Ecol. Monogr. 49: 377-402.

o 187

Benedict, J.B. 1966. Radiocarbon dates from a stone-banked terrace in the Colorado Rocky Mountains, U.S.A. Geogr. Ann. 48A: 24-31.

Bennett, R.C. 1976. Notes on alpine climate, p. 14-20. In H.A. Luttmerding and J.A. Shields (eds.) Proceedings of the Workshop on Alpine and Subalpine Environments. Res. Anal. Branch, B.C. Min. Environ., Victoria, B.C.

Berendse, F. 1979. Competition between plant populations with different rooting depths. I. Theoretical considerations. Oecologia 43: 19-26.

Billings, W.D. 1952. The environmental complex in relation to plant growth and distribution. Quart. Rev. Biol. 27:251- 265.

Billings, W.D. 1973. Arctic and alpine vegetations: similarities, differences, and susceptibility to disturbance. Bioscience 23: 697-704.

Billings, W.D. 1974a. Adaptations and origins of alpine plants. Arct. Alp. Res. 6: 129-142.

Billings, W.D. 1974b. Arctic and alpine vegetation: plant adaptations to cold summer climates, p. 404-443. I_n J. Ives and R.G. Barry (eds.) Arctic and Alpine Environments. Methuen, London.

Billings, W.D., and L.C. Bliss 1959. An alpine snowbank environment and its effects on vegetation, plant development, and productivity. Ecology 40: 388-397.

Billings, W.D., and H.A. Mooney. 1959. An apparent frost hummock-sorted polygon cycle in the alpine tundra of Wyoming. Ecology 40: 16-19.

Billings, W.D., P.J. Godfrey, B.F. Chabot, and D.P. Bourque. 1971. Metabolic acclimation to temperature in arctic and alpine ecotypes of Oxyria diqyna. Arct. Alp. Res. 3: 277- 289.

Black, CA. (ed.) 1965. Methods of soil analysis, vol. 2. Amer. Soc. Agron., Madison, Wis. 1572 p. 188

Black, J.N. 1960. The significance of petiole length, leaf area, and light interception in competition between strains of subterranean clover (Trifolium subterraneum L.) grown in swards. Aust. J. Agric. Res. 11: 277-291.

Blackman, G.E. 1935. A study by statistical methods of the distribution of species in grassland. Ann. Bot. Lond. 49: 749-777.

Blaser, R.E., and N.C. Brady. 1950. Nutrient competition in plant associations. Agron. J. 42: 128-135.

Bliss, L.C. 1956. A comparison of plant development in microenvironments of arctic and alpine tundras. Ecol. Monogr. 26: 303-337.

Bliss, L.C. 1962. Adaptations of arctic and alpine plants to environmental conditions. Arctic 15: 117-144.

Bliss, L.C. 1963. Alpine plant communities of the Presidential Range, New Hampshire. Ecology 44: 678-697.

Bliss, L.C. 1966. Plant productivity in alpine microenvironments on Mt. Washington, New Hampshire. Ecol. Monogr. 36: 125- 1 55.

Bliss, L.C. 1969. Alpine community patterns in relation to environmental parameters, p. 167-184. In K.N.H. Greenridge (ed.) Essays in Plant Geography and Ecology. Nova Scotia Museum, Halifax, Nova Scotia.

Bliss, L.C. 1971. Arctic and alpine plant life cycles. Annual Rev. Ecol. and Syst. 2: 405-438.

Bliss, L.C, and E.B. Hadley. 1964. Photosynthesis and respiration of alpine lichens. Amer. J. Bot. 51: 870-874.

Bliss, L.C, and CM. Woodwell. 1965. An alpine podzol on Mt. Katahdine, Maine. Soil Sci. 100: 274-279.

Bockheim, J.G. 1972. Effects of alpine and subalpine vegetation on soil development, Mount Baker, Washington. Ph.D. thesis. Univ. of Washington, Seattle, Washington. 171 p. 189

Bormann, F.H. 1953. The statistical efficiency of sample plot size and shape in forest ecology. Ecology 34: 474-487.

Bouyoucos, G.J. 1951. A recalibration of the hydrometer method for making mechanical analysis of soil. Agron. J. 43: 434- 438.

Bratton, S.P. 1976. Resource division in an understory herb community: responses to temporal and microtopographic gradients. Amer. Natur. 110: 679-693.

Braun-Blanquet, J. 1932. Plant sociology; the study of plant communities (Transl. by G.D. Fuller and H.S. Conard) Transl. of 1st ed. of Pflanzensoziologie (1928). McGraw- Hill, New York and London. 438 p.

Bray, R.H., and L.T. Kurtz. 1945. Determination of total, organic, and available forms of phosphorus in soils. Soil Sci. 59: 39-45.

Bremner, J.M. 1960. Determination of nitrogen in soil by the Kjeldahl Method. J. Agr. Sci. 55: 1-23.

Brinck, P. 1974. Strategy and dynamics of high altitude faunas. Arct. Alp. Res. 6: 107-116.

British Columbia Department of Agriculture. 1974. Climate of British Columbia. Climatic Normals 1941-1970. Queens Printer, Victoria. 90 p.

Broad, J. 1973. Ecology of alpine vegetation at Bow Summit, Banff National Park. M.Sc. thesis. Univ. of Calgary, Calgary, Alberta. 93 p.

Bryant, J.P., and E. Scheinberg. 1970. Vegetation and frost activity in an alpine fellfield on the summit of Plateau Mountain, Alberta. Can. J. Bot. 48: 751-771.

Bykov, B.A. 1974. Fluctuations in the semidesert and desert vegetation of the Turanian plain, p. 243-251. In R. Knapp (ed.) Vegetation Dynamics, Handbook of Vegetation Science, Part 8. Junk, The Hague. 190

Cain, S.A. 1938. The species-area curve. Amer. Midi. Nat. 17: 725-740.

Caldwell, M.M. 1968. Solar U.V. radiation as an ecological factor for alpine plants. Ecol. Monogr. 38: 243-268.

Callaghan, T.V. 1976. Growth and population dynamics of Carex biqelowii in an alpine environment. Strategies of growth and population dynamics of tundra plants, part 3. Oikos 27: 402-413.

Cartwright, D. 1970. Cathedral Provincial Park enlargement - socio-economic and administrative problems. M.F. thesis. Univ. of British Columbia, Vancouver, B.C. 137 p.

Chilton, R.H.R. 1981. A summary of climatic regimes of British Columbia. B.C. Min. Environ., Air Studies Branch. Queens Printer, Victoria. 46 p.

Clapham, A.R. 1932. The form of the observational unit in quantitative ecology. J. Ecol. 20: 192-197.

Clapham, A.R. 1936. Overdispersion in grassland. J. Ecol. 24: 232-251.

Clausen, J., D.D. Keck, and W.M. Hiesey. 1948. Experimental studies on the nature of species. III. Environmental responses of climatic races of Achillea. Carnegie Inst. Washington. Pub. No. 581. 129 p.

Clements, F.E. 1904. Development and structure of vegetation. Rep. Bot. Surv. Nebr., 7.

Clements, F.E., and J.E. Weaver. 1924. Experimental vegetation. Carnegie Inst. Wash. Publ. 355: 1-172.

Cole, L.C. 1957. The measurement of partial interspecific association. Ecology 38: 226-233.

Cole, L.C. 1960. Competitive exclusion. Science 132: 348-349.

Connell, J.H. 1975. Some mechanisms producing structure in 191

natural communities: a model and evidence from field experiments, p. 460-490. I_n M.L. Cody and J.M. Diamond (eds.) Ecology and Evolution of Communities. Harvard Univ. Press, Cambridge.

Connell, J.H. 1980. Diversity and the coevolution of competitors, or the ghost of competition past. Oikos 35: 131-138.

Coombs, H.A. 1939. Mt. Baker, a Cascade volcano. Geol. Soc. Am. Bull. 50: 1493-1510.

Cooper, W.S. 1908. Alpine vegetation in the vicinity of Long's Peak. Bot. Gaz. 45: 319-337.

Corby, H.D.L. 1981. The systematic value of leguminous root nodules, p. 657-669. I_n R.M. Polhill and P.H. Raven (eds.) Advances in Legume Systematics, Part 2. Proc. Int. Legume Conf., Roy. Bot. Gard., Kew.

Cox, C.F. 1933. Alpine plant succession on James Peak, Colorado. Ecol. Monogr. 3: 299-372.

Crack, S.N. 1977. Flora and vegetation of Wilcox Pass, Jasper National Park, Alberta. M.Sc. thesis. Univ. of Calgary, Calgary, Alberta. 284 p.

Crandell, D.R. 1965. The glacial history of western Washington and Oregon, p. 341-353. I_n H.E. Wright, Jr. and D.G. Frey (eds.) The Quaternary of the United States. Princeton Univ. Press, Princeton, New Jersey.

Crandell, D.R., D.R. Mullineaux, R.D. Miller, and M. Rubin. 1969. Pyroclastic deposits of recent age at Mount Rainier, Washington. U.S. Geol. Surv. Prof. Pap. 450-D. 64 p.

Dalziel, B.R. 1971. An assessment of the recreation potential of the Ashnola Valley. Prov. Parks Branch, Dept. Rec. and Con., Victoria, B.C.

Daly, R.A. 1912. Geology of the North American Cordillera at the Forty-Ninth Parallel, Parts I, II, and III. Memoir 38, Geol. Surv. Can. 857 p. 1 92

Darwin, C. 1859. The origin of species. Harvard Facsimile 1st ed. 1964.

Daubenmire, R. 1968. Plant communities: A textbook of plant synecology. Harper & Row, N.Y. 300 p.

Douglas, G.W. 1980. Vegetation. I_n Biophysical Inventory Studies of Kluane National Park. Parks Canada, Winnipeg.

Douglas, G.W., and L.C. Bliss. 1977. Alpine and high subalpine plant communities of the north Cascades Range, Washington, USA, and British Columbia, Canada. Ecol. Monogr. 47: 113- 150.

Downes, R.G., and R.S. Beckwith. 1951. Studies in the variation of soil reaction. I. Field variation at Baroga, N.S.W. Australia. J. Agr. Res. 2: 60-72.

Dunn, D.B., and J.M. Gillett. 1966. The lupines of Canada and Alaska. Can. Dept. Agri., Monogr. No. 2. 89 p.

Eady, K. 1971. Ecology of the alpine and timberline vegetation of Big White Mountain, British Columbia. Ph.D. thesis. Univ. of British Columbia, Vancouver, British Columbia. 239 P-

Ehleringer, J.R., and P.C. Miller. 1975. Water relations of selected plant species in the alpine tundra., Colorado, USA. Ecology 56: 370-380.

Elisens, W.J., and J.G. Packer. 1980. A contribution to the taxonomy of the Oxytropis campestris complex in northwestern North America. Can. J. Bot. 58: 1820-1831.

Etherington, J.R. 1981. Limestone heaths in south-west Britain: their soils and the maintenance of their calcicole- calcifuge mixtures. J. Ecol. 69: 277-294.

Everitt, B. 1974. Cluster analysis. Heinemann Ed. Books, London.

Faust, R.A., and T.J. Nimlos. 1968. Soil micro-organisms and soil nitrogen of the Montana alpine. Northwest Sci. 42: 101-107. 193

Fitter, A.H. 1982. Influence of soil heterogeneity on the coexistence of grassland species. J. Ecol. 70: 139-148.

Flohn, H. 1974. Contribution to a comparative meteorology of mountain areas, p. 55-71. I_n J. Ives and R.G. Barry (eds.) Arctic and Alpine Environments. Methuen, London.

Fontana, A. 1963. Micorrize ectotrofiche in una Ciperacea: Kobresia belliardi Degl. Giorn. Bot. Ital. 70: 639-641.

Fbrman, R.T.T., and D.L. Dowden. 1977. Nirtogen fixing lichen roles from desert to alpine in the Sangre . De Cristo Mountains, New Mexico. Bryologist 80: 561-570.

Fowler, N., and J. Antonovics. 1981. Competition and coexistence in a North Carolina grassland. I. Patterns in undisturbed vegetation. J. Ecol. 6.9: 825-841 .

Fox, D.J., and K.E. Guire. 1976. Documentation for MIDAS, 3rd ed. Statistical Research Laboratory, The University of Michigan. 203 p.

Frankie, G.W., H.G. Baker, and P.A. Opler. 1974. Comparative phenological studies of trees in tropical wet and dry forests in the lowlands of Costa Rica. J. Ecol. 62: 881- 913.

Franklin, J.F., and C.T. Dyrness. 1973. Natural vegetation of Oregon and Washington. U.S. For. Serv. Gen. Tech. Rep. PNW- 8. 417 p.

Fryxell, R. 1965. Mazama and Glacier Peak ash layers; relative ages. Science 147: 1288-1290.

Gates, D.M., and R. Janke. 1966. The energy environment of the alpine tundra. Oecol. Plant. 1: 39-62.

Gauch, H.G. 1973. A quantitative evaluation of the Bray-Curtis ordination. Ecology 54: 829-836.

Gauch, H.G. 1977. ORDIFLEX - A flexible computer program for four ordination techniques: weighted averages, polar ordination, principal components analysis, and reciprocal 1 94

averaging. Release B. Ecology and Systematics, Cornell Univ., Ithaca, New York.

Gauch, H.G., and R.H. Whittaker. 1972. Comparison of ordination techniques. Eoclogy 53: 868-875.

Gauch, H.G., and R.H. Whittaker. 1981. Hierarchical classification of community data. J. Ecol. 69: 537-557.

Gauch, H.G., R.H. Whittaker, and T.R. Wentworth. 1977. A comparative study of reciprocal averaging and other ordination techniques. J. Ecol. 65: 157-174.

Gause, G.F. 1934. The struggle for existence. Waverly Press, Baltimore.

Geological Map of Canada. 1955. Dept. Mines and Technical Surveys, Geol. Surv. Can. Map 1045A.

Gittens, R. 1969. The application of ordination techniques, p. 37-66. I_n I.H. Rorison et al. (eds.) Ecological Aspects of the Mineral Nutrition of Plants. Symp. Br. Ecol. Soc. 9, Blackwell Sci. Publ., Oxford.

Gleason, H.A. 1926. The individualistic concept of the plant association. Bull. Torrey Bot. Club 53: 7-26.

Goodall, D.W. 1952. Quantitative aspects of plant distribution. Biol. Rev. 27: 194-245.

Goodall, D.W. 1978a. Numerical classification, p. 247-288. In R.H. Whittaker (ed.) Classification of Plant Communities. Junk, The Hague.

Goodall, D.W. 1978b. Sample similarity and species correlation, p. 99-149. In R.H. Whittaker (ed.) Ordination of Plant Communities. Junk, The Hague.

Granhall, U., and H. Selander. 1973. Nitrogen fixation in a subarctic mire. Oikos 20: 175-178.

Green, A.J., and T.M. Lord. 1979. Soil survey of the Princeton 195

map area, British Columbia. Soil Surv. Rep., 14. Queens Printer, Ottawa.

Greig-Smith, P. 1952. The use of random and contiguous quadrats in the study of the structure of plant communities. Ann. Bot., Lond., N.S. 16: 293-316.

Greig-Smith, P. 1961. Data on pattern within plant communities. I. The analysis of pattern. J. Ecol. 49: 695-702.

Greig-Smith, P. 1964. Quantitative plant ecology, 2nd edition. Butterworths, London. 256 p.

Greig-Smith, P. 1979. Pattern in vegetation. J. Ecol. 67: 755- 779.

Griggs, R.F. 1956. Competition and succession on a Rocky Mountain fellfield. Ecology 37: 8-20.

Grime, J.P. 1973a. Control of species diversity in herbaceous vegetation. J. Environ. Manage. 1: 151-167.

Grime, J.P. 1973b. Competitive exclusion in herbaceous vegetation. Nature 242: 344-347.

Grime, J.P. 1979. Plant strategies and vegetation processes. John Wiley & Sons, New York. 222 p.

Grubb, P.J. 1977. The maintenance of species richness in plant communities: the importance of the regeneration niche. Biol. Rev. 52: 107-145.

Grubb, P.J., H.E. Green, and R.C.J. Merrifield. 1969. The ecology of chalk heath: its relevance to the calcicole- calcifuge and soil acidification problems. J. Ecol. 57: 175-212.

Hadley, E.B., and L.C. 'Bliss. 1964. Energy relationships of alpine plants on Mt. Washington, New Hampshire. Ecol. Monogr. 34: 331-357.

Hale, M.E., Jr., and W.L. Culberson. 1970. A fourth checklist of 196

the lichens of the continental United States and Canada. Bryologist 73: 499-543.

Hanson, H.C. 1951. Characteristics of some grassland, marsh, and other plant communities in western Alaska. Ecol. Monogr. 21: 317-378.

Hardin, G. 1960. The competitive exclusion principal. Science 131: 1292-1297.

Harley, J.L. 1969. The biology of mycorrhiza, 2nd ed. Leonard Hill, London. 247 p.

Harper, J.L. 1964. The individual in the population. J.Ecol. (Suppl.) 52: 149-158.

Harper, J.L. 1967. A Darwinian approach to plant ecology. J. Ecol. 55: 247-270.

Harper, J.L. 1969. The role of predation in vegetational diversity, p. 48-62. Ijn Diversity and Stability in Ecological Systems. Brookhaven Symp. Biol., 22.

Harper, J.L. 1977a. The contributions of terrestrial plant studies to the development of the theory of ecology. Acad. Nat. Sci. Phil., Spec. Publ. Vol. 12: 139-157.

Harper, J.L. 1977b. Population biology of plants. Academic Press, London. 892 p.

Harper, J.L. 1982. After description, p. 11-25. I_n E.I. Newman (ed.) The Plant Community as a Working Mechanism. Spec. Publ. No. 1, Brit. Ecol. Soc, Blackwell Scientific Publ., Oxford.

Harper, J.L., and G.R. Sagar. 1953. Some aspects of the ecology of buttercups in permanent grassland Proc. British Weed Control Conf. p.256-264.

Harper, J.L., J.N. Clatworthy, I.H. McNaughton, and G.R. Sagar. 1961. The evolution and ecology of closely related species living in the same area. Evolution 15: 209-227. 1 97

Haselwandter, K., and D.J. Read. 1980. Fungal associations of roots of dominant and sub-dominant plants in high-alpine vegetation systems with special reference to mycorrhiza. Oecologia 45: 57-62.

Helm, D. 1982. Multivariate analysis of alpine snow-patch vegetation cover near Milner Pass, Rocky Mtn. National Park, Colorado, U.S.A. Arct. Alp. Res. 14: 87-95.

Heslop-Harrison, J. 1964. Forty years of genecology. Adv. Ecol. Res. 2: 159-247.

Higgins, P.D., and G.G. Spomer. 1976. Soil temperature effects o root respiration and the ecology of alpine and subalpine plants. Bot. Gaz. 137: 110-120.

Hill, M.O. 1973. Reciprocal averaging: an eigenvector method of ordination. J. Ecol. 61: 237-249.

Hirschfeld, H.O. 1935. A connection between correlation and contingency. Proc. Camb. Phil. Soc. 31: 520-524.

Hitchcock, C.L., and A. Cronquist. 1973. Flora of the Pacific Northwest. Univ. Washington Press, Seattle, Wash. 730 p.

Holland, S.S. 1964. Landforms of British Columbia - a physiographic outline. B.C. Dept. of Mines and Petrol. Res., Bull. 48. Queens Printer, Victoria. 138 p.

Holm, T. 1927. The vegetation of the alpine region of the Rocky Mountains in Colorado. Nat. Acad. Sci. Mem. 19: 1-45.

Holway, J.G., and R.T. Ward. 1963. Snow and meltwater effects in an area of Colorado alpine. Amer. Midi. Nat. 69: 189-197.

Holway, J.G., and R.T. Ward. 1965. Phenology of alpine plants in northern Colorado. Ecology 46: 73-83.

Hrapko, J.O., and G.H. La Roi. 1978. The alpine tundra vegetation of Signal Mountain, Jasper National Park. Can. J. Bot. 56:309-332. 198

Jacob, F., and J. Monod. 1961. Genetic regulatory mechanisms in the synthesis of proteins. J. Mol. Biol. 3: 318-356.

Jenny, H. 1941. Factors of soil formation. McGraw-Hill, New York. 281 p.

Johnson, P.L., and W.D. Billings. 1962. The alpine vegetation of the Beartooth Plateau in relation to cryopedogenic processes and patterns-. Ecol. Monogr. 32: 105-135. de Jong, P., and M. Greig. 1983. First order Markov chains with a zero diagonal transition matrix. Biometrics (in press) de Jong, P., L.W. Aarssen, and R. Turkington. 1983. The use of contact sampling in studies of association in vegetation. J. Ecol. (in press).

Joy, P., and A. Laitinen. 1980. Breeding for coadaptation between red clover and timothy. Hankkija's Seed Publ. No. 13. Hankkija Plant Breeding Institute, Finland.

Kendrew, W.G., and D. Kerr. 1955. The climate of British Columbia and the Yukon Territory. Can. Dep. Transport, Meteorol. Div., Toronto. Queens Printer, Ottawa. 222 p.

Kershaw, K.A. 1958. An investigation of the structure of a grassland community. I. The pattern of Agrostis tenuis. J. Ecol. 46: 571-592.

Kershaw, K.A. 1959. An investigation of the structure of a grassland community. II. The pattern of Dactylis glomerata, Lolium perenne, and Trifolium repens. III. Discussion and conclusion. J. Ecol. 47: 31-53.

Kershaw, K.A. 1963. Pattern in vegetation and its causality. Ecology 44: 377-388.

Kershaw, K.A. 1973. Quantitative and Dynamic Plant Ecology, 2nd ed. Edward Arnold, London. 308 p.

Kjeldahl, J. 1883. Neue Methode zur Bestimmung des Stickstoffs in organischen Korpern. Z. Anal. Chem. 22: 366-382. 199

Klikoff, L.G. 1965. Microenvironmental influence on vegetation pattern near timberline in the central Sierra Nevada. Ecol. Monogr. 35: 187-211.

Klikoff, L.G. 1968. Temperature dependence of mitochondrial oxidative rates of several plant species in the Sierra Nevada. Bot. Gaz. 129: 227-230.

Knapik, L.J., G.W. Scotter, and W.W. Pettapiece. 1973. Alpine soil and plant community relationships of the Sunshine Area, Banff National Park. Arct. Alp. Res. 5: A161-A170.

Komarkova, V., and P.J. Webber. 1978. An alpine vegetation map of Niwot Ridge, Colorado. Arct. Alp. Res. 10: 1-29.

Kuramoto, R.T., and L.C. Bliss. 1970. Ecology of subalpine meadows in the Olympic Mountains, Washington. Ecol. Monogr. 40: 317-347.

Langlet, 0. 1971. Two hundred years genecology. Taxon 20: 653- 722.

Larcher, W., A. Cernusca, L. Schmidt, G. Grabherr, E. Notzel, and N. Smeets. 1975. Mt. Patscherkofel, Austria, p. 125- 139. I_n T. Rosswall, and O.W. Heal (eds.) Structure and Function of Tundra Ecosystems. Swedish Nat. Sci. Res. Coun. (Stockholm), Ecol. Bull. No. 20.

Laursen, G.A., and M.A. Chmielewski. 1980. The ecological significance of soil fungi in arctic tundra, p. 432-488. I_n G.A. Laursen and J.F. Ammirati (eds.) Arctic and Alpine Mycology. The first international symposium on arcto-alpine mycology. Univ. Wash. Press, Seattle.

Lawton, E. 1971. Moss flora of the Pacific Northwest. Hattori Botanical Laboratory, Nichinan, Miyazaki, Japan. 362 p.

Lieth, H. 1975. Primary production of the major vegetation units of the world, p. 203-215. In H. Lieth and R.H. Whittaker (eds.) Primary Productivity of the Biosphere. Springer- Verlag, New York.

Levins, R. 1968. Evolution in changing environments. Princeton Univ. Press, Princeton, N.J. 120 P. 200

Linkins, A.E., and R.K. Antibus. 1980. Mycorrhizae of Salix rotundifolia in coastal arctic tundra, p. 509-525. I_n G.A. Laursen and J.F. Ammirati (eds.) Arctic and Alpine Mycology. The first international symposium on arcto-alpine mycology. Univ. Wash. Press, Seattle.

Love, D. 1970. Subarctic and subalpine: where and what? Arct. Alp. Res. 2: 63-73.

Luckhurst, A.J. 1973. Stone sheep and their habitat in the northern Rocky Mountain foothills of British Columbia. M.Sc. thesis. Univ. of British Columbia, Vancouver, B.C.

MacArthur, R.H. 1972. Geographical ecology: patterns in the distribution of species. Harper & Row, New York. 269 p.

Macior, L.W. 1974. Pollination ecology of the Front Range of the Colorado Rocky Mountains. Melanderia 15: 1-59.

Mack, R.N., and J.L. Harper. 1977. Interference in dune annuals: spatial pattern and neighbourhood effects. J. Ecol. 65: 345-363.

Major, J. 1951. A functional, factorial approach to plant ecology. Ecology 32: 392-412. .

Major, J., and S.A. Bamberg. 1963. Some Cordilleran plant species new for the Sierra Nevada of California. Madrono 17: 93-109.

Mark, A.F. 1965. Flowering and seedling establishment of narrow- leaved snow tussock, Chionochloa rigida. N.Z. J. Bot. 3: 180-193.

Mark, A.F. 1970. Floral initiation and development in New Zealand alpine plants. N.Z. J. Bot. 8: 67-75.

Mark, A.F. '1975. Photosynthesis and dark respiration in 3 alpine snow tussocks (Chionochloa sp.) under controlled environments. N.Z. J. Bot. 13: 93-122.

Mark, A.F., and L.C. Bliss. 1970. The high-alpine vegetation of central Otago, New Zealand. N.Z. J. Bot. 8: 381-451. 201

Marr, J.W. 1967. Ecosystems of the east slope of the Front Range in Colorado. Univ. Colorado Stud. Ser. Biol. No. 8. 134 p.

May, D.E. 1976. The response of alpine tundra vegetation in Colorado to environmental variation. Ph.D. thesis. Univ. of Colorado, Boulder, Colorado. 164 p.

May, R.M. 1974. On the theory of niche overlap. Theoret. Pop. Biol. 5: 297-332.

McCown, B.H. 1975. Physiological responses of root systems to stress conditions, p. 225-237. I_n F.J. Vernberg (ed.) Physiological Adaptation to the Environment. New York.

McCown, B.H., and L.L. Tieszen. 1972. Comparative periodic trends in carbohydrate and lipid levels in arctic and alpine plants and their physiological, ecological significance. Plant Physiol. 49: (suppl.) 6.

Mcintosh, R.P. 1970. Community, competition, and adaptation. Q. Rev. Biol. 45: 259-280.

McLean, A. 1970. Plant communities of the Similkameen Valley, British Columbia, and their relationships to soils. Ecol. Monogr. 40: 403-424.

McNaughton, S.J., and L.L. Wolf. 1970. Dominance and the niche in ecological systems. Science 167: 131-139.

McTaggart, K.C. 1970. Tectonic history of the northern Cascade Mountains, p. 1-5 and 155-166. I_n J.O. Wheeler (ed.) Structure of the Southern Canadian Cordillera. Geol. Assoc. Can. Spec. Paper No. 8.

Medway, L. 1972. Phenology of a tropical rainforest in Malaya. Biol. J. Linn. Soc. 4: 117-146.

Melcon, P.Z. 1975. Tors and weathering on McKeen Ridge, Cathedral Provincial Park, British Columbia. M.A. thesis. Simon Fraser Univ., Burnaby, B.C. 183 p.

Meredith, D.H. 1972. Subalpine cover associations of Eutamias amoenus and Eutamias townsendii in the Washington Cascades. 202

Amer. Midi. Nat. 88: 348-357.

Milbank, J.W., and K.A. Kershaw. 1973. Nitrogen metabolism, p. 289-309. In V. Ahmadjian and M.E. Hale (eds.) The Lichens. Acad. Press, New York.

Miller, O.K., and G.A. Laursen. 1978. Ecto and endomycorrhizae of arctic plants at Barrow, Alaska, p. 229-237. In L.L. Tieszen (ed.) Vegetation and Production Ecology of an Alaskan Arctic Tundra. Ecological Studies 29, Springer Verlag, New York.

Misch, P. 1966. Tectonic evolution of the northern Cascades of Washington State. Canadian Inst. Mining and Metallurgy Spec. Vol. 8: 101-148.

Moldenke, A.R. 1976. California, USA: Pollination ecology and vegetation types. Phytologia 34: 305-361.

Mooney, H.A. 1963. Physiological ecology of coastal, subalpine, and alpine populations of Polygonum bistortoides. Ecology 44: 812-816.

Mooney, H.A., and W.D. Billings. 1960. The annual carbohydrate cycle of alpine plants as related to growth. Amer. J. Bot. 47: 594-598.

Mooney, H.A., and W.D. Billings. 1961. Comparative physiological ecology of arctic and alpine populations of Oxyria. Ecol. Monogr. 31: 1-29.

Mooney, H.A., G.S. Andre, and R.D. Wright. 1962. Alpine and subalpine vegetative patterns in the White Mountains of California. Amer. Midi. Nat. 68: 257-273.

Mooney, H.A., R.D. Wright, and B.R. Strain. 1964. The gas exchange capacity of plants in relation to vegetation zonation in the White Mountains of California. Amer. Midi. Nat. 72: 281-297.

Mulligan, G.A. 1971. Cytotaxonomic studies of the closely-allied Draba cana, D. cinerea, and D. groenlandica in Canada and Alaska. Can. J. Bot. 49: 89-93. 203

Mullineaux, D.R. 1964. Extensive Recent pumice lapilli and ash layers from Mount St. Helens volcano, southern Washington. Geol. Soc. Amer. Spec. Pap. 76: 285 (abstr.)

Naysmith, H. 1962. Late glacial history and surficial deposits of the Okanagan Valley, British Columbia. B.C. Dept. Mines, Bull. 46.

Naysmith, H., W.H. Mathews, and G.E. Rouse. 1967. Bridge River ash and some other Recent ash beds in British Columbia. Can. J. Earth Sci. 4: 163-170.

Newman, E.I. (ed.) 1982a. The Plant Community as a Working Mechanism. Spec. Publ. No. 1 Brit. Ecol. Soc, Blackwell Scientific Publ., Oxford. 128 p.

Newman, E.I. 1982b. Niche separation and species diversity in terrestrial vegetation, p. 61-77. In E.I. Newman (ed.) The Plant Community as a Working Mechanism. Spec. Publ. No. 1, Brit. Ecol. Soc, Blackwell Scientific Publ., Oxford.

Nimlos, T.J., and R.C. McConnell. 1962. The morphology of alpine soils in Montana. Northwest Sci. 36: 99-112.

Nimlos, T.J., and R.C. McConnell. 1965. Alpine soils in Montana. Soil Sci. 99: 310-321 .

Nimlos, T.J., R.C. McConnell, and D.L. Pattie. 1965. Soil temperature and moisture regimes in Montana alpine soils. Northwest Sci. 39: 129-138.

Noy-Meir, I., D. Walker, and W.T. Williams. 1975. Data transformation in ecological ordination. II. On the meaning of data standardization. J. Ecol. 63: 779-800.

Odum, E.P. 1960. Organic production and turnover in old field succession. Ecology 41: 34-49.

Okazaki, R., H.W. Smith, R.A. Gilkeson, and J. Franklin. 1972. Correlation of West Blacktail with Pyroclastic Layer '7' from the 1800 A.D. erruption of Mount St. Helens. Northwest Sci. 46: 74-89. 204

Orloci, L. 1966. Geometric models in ecology. I. The theory and application of some ordination methods. J. Ecol. 54: 193- 215.

Orloci, L. 1975. Multivariate analysis in vegetation research. Junk, The Hague. 276 p.

Orloci, L. 1978. Ordination by resemblance matrices, p. 239-275. In R.H. Whittaker (ed.) Ordination of Plant Communities. Junk, The Hague.

Ovington, S.D., D. Heitkamp, and D. Lawrence. 1963. Plant biomass and productivity of prairie, savanna, oakwood, and maizefield ecosystems in central Minnesota. Ecology 44: 52- 63.

Parrish, J.A.D., and F.A. Bazzaz. 1976. Underground niche separation in successional plants. Ecology 57: 1281-1288.

Parrish, J.A.D., and F.A. Bazzaz. 1978. Pollination niche separation in a winter annual community. Oecologia 35: 133— 1 40.

Pearcy, R.W., and R.T. Ward. 1972. Phenology and growth of Rocky Mountain populations of Deschampsia caespitosa at three elevations in Colorado. Ecology 53: 1171-1178.

Pearson, V., and D.J. Read. 1973. The biology of mycorrhiza in the Ericaceae. I. The isolation of the endophyte and synthesis of mycorrhizas in aseptic culture. New Phytol. 72: 371-379.

Petersen, B. 1977. Pollination of Thlaspi alpestre by selfing and by insects in the alpine zone of Colorado, USA. Arct. Alp. Res. 9: 211-215.

Pianka, E.R. 1976. Competition and niche theory, p. 114-141. I_n R.M. May (ed.) Theoretical ecology: principles and applications. Blackwell Scientific Publ., Oxford.

Pianka, E.R. 1979. Evolutionary ecology, 2nd ed. Harper & Row, New York. 397 p. 205

Pielou, E.C. 1967. A test for random mingling of the phases of a mosaic. Biometrics 23: 657-670.

Pimentel, R.A. 1979. Morphometries, the multivariate analysis of biological data. Kendall/Hunt, Iowa. 276 p.

Pinder, J.E. 1975. Effects of species removal on an old-field plant community. Ecology 56: 747-751.

Porter, S.C., and G.H. Denton. 1967. Chronology of neoglaciation in the North American Cordillera. Amer. J. Sci. 265: 177- 210.

Powers, H.A., and R.E. Wilcox. 1964. Volcanic ash from Mount Mazama (Crater Lake) and from Glacier Peak. Science 144: 1334-1336.

Prest, V.K. 1957. Pleistocene geology and surficial deposits, p. 443-495. In C.H. Stockwell (ed.) Geology and Economic Minerals of Canada. Geol. Surv. Can. Econ. Geol. Series No. 1 .

Price, L.W. 1971. Vegetation, microtopography, and depth of active layer on different exposures in subarctic alpine tundra. Ecology 52: 638-647.

Pritchard, N.M., and A.J.B. Anderson. 1971. Observations on the use of cluster analysis in botany with an ecological example. J. Ecol. 59: 727-747.

Read, D.J., and K. Haselwandter. 1981. Observations on the mycorrhizal status- of some alpine plant communities. New Phytol. 88: 341-352.

Reader, R.J. 1975. Competitive relationships of some bog ericads for major insect pollinators. Can. J. Bot. 53: 1300-1305.

Rehder, H. 1976. Nutrient turnover studies in alpine ecosystems, part 2: phytomass and nutrient relations in the Caricetum- Firmae. Oecologia 23: 49-62.

Retzer, J.L. 1956. The alpine soils of the Rocky Mountains. J. Soil Sci. 7: 22-32. 206

Retzer, J.L. 1965. Present soil forming factors and processes in arctic and alpine regions. Soil Sci. 99: 38-44.

Retzer, J.L. 1974. Alpine soils, p. 771-802. In J. Ives and R.G. Barry (eds.) Arctic and Alpine Environments. Methuen, London.

Rice, E.L. 1974. Allelopathy. Academic Press, New York.

Rice, H.M.A. 1947. Geology and mineral deposits of the Princeton map-area. Memoir No. 243. Geol. Surv. Can. 136 p.

Ricklefs, R.E. 1979. Ecology, 2nd ed. Chiron Press. New York. 966 p.

Rochow, T.F. 1970. Ecological investigations of Thlaspi alpestre L. along an elevational gradient in the central Rocky Mountains. Ecology 51: 649-656.

Ross, M.S., and J.L. Harper. 1972. The occupation of biological space during seedling establishment. J. Ecol. 60: 77-88.

Rudkin, R.A. 1964. The Lower Cretaceous, p. 156-1 68. I_n R.D. McCrossan and R.P. Glaister (eds.) Geological History of Western Canada. Alberta Soc. Pet. Geol.

Sanders, F.E.T., B. Mosse, and P.H.B. Tinker. 1975. Endomycorrhizas. Academic Press, London.

Sarukhan, J., and J.L. Harper. 1973. Studies on plant demography: Ranunculus repens L., R. bulbosus L., and R. acris L. I. Population flux and survivorship. J. Ecol. 61: 675-716.

Savile, D.B.A. 1960. Limitations of the competitive exclusion principal. Science 132: 1761.

Sayers, R.L., and R.T. Ward. 1966. Germination responses in alpine species. Bot. Gaz. 127: 11-16.

Schell, D.M., and V. Alexander. 1973. Nitrogen fixation in arctic coastal tundra in relation to vegetation and micro- 207

relief. Arctic 26: 130-137.

Schoener, T.W. 1974a. Resource partitioning in ecological communities. Science 185: 27-39.

Schoener, T.W. 1974b. The compression hypothesis and temporal resource partitioning. Proc. Nat. Acad. Sci. USA. 71: 4169- 41 72.

Schollenberger, C.J., and R.H. Simon. 1945. Determination of exchange capacity and exchangeable bases in soil - ammonium acetate method. Soil Sci. 59: 13-25.

Scott, D., and W.D. Billings. 1964. Effects of environmental factors on standing crop and productivity of an alpine tundra. Ecol. Monogr. 34: 243-270.

Shaver, G.R., F.S. Chapin III, and W.D. Billings. 1979. Ecotypic differentiation in Carex aquatilis on ice-wedge polygons in the Alaskan coastal tundra. J. Ecol. 67: 1025-1046.

Siccama, T.G., F.H. Bormann, and G.E. Likens. 1970. The Hubbard Brook Ecosystem Study: Productivity, nutrients, and phytosociology of the herbaceous layer. Ecol. Monogr. 40: 389-402.

Snaydon> R.W. 1962. Micro-distribution of Trifolium repens L. and its relation to soil factors. J. Ecol. 50: 133-143.

Sneath, P.H.A., and R.R. Sokal. 1973. Numerical taxonomy, the principles and practice of numerical classification. W.H. Freeman, San Francisco. 573 p.

Sneddon, J.I. 1969. The genesis of some alpine soils in British Columbia. M.Sc. thesis. Univ. of British Columbia, Vancouver, B.C. 131 p.

Sneddon, J.I., L.M. Lavkulich, and L. Farstad. 1972. The morphology and genesis of some alpine soils in British Columbia, Canada. I. Morphology, classification, and genesis. II. Physical, chemical, and mineralogical determinations and genesis. Soil Sci. Soc. Amer. Proc. 36: 100-110. 208

Soil Survey Committee of Canada. 1968. Proceedings of the 7th National Meeting. Can. Dep. Agr., Soil Res. Inst., Ottawa.

Soil Survey Staff. 1975. Soil Taxonomy: A Basic System of Soil Classification for Making and Interpreting Soil Surveys. Soil Conserv. Serv., U.S. Dep. Agric. Handbook 436. 754 p.

Sokal, R.R., and F.J. Rohlf. 1962. The comparison of dendrograms by objective methods. Taxon 11: 33-40.

Stoner, W.A., P.C. Miller, and W.C. Oechel. 1978. Simulation of the effect of the tundra vascular plant canopy on the productivity of four plant species, p. 371-385. I_n L.L. Tieszen (ed.) Vegetation and Production Ecology of an Alaskan Arctic Tundra. Ecological Studies 29, Springer- Verlag, New York.

Stowe, L.G., and M.J. Wade. 1979. The detection of small-scale patterns in vegetation. J. Ecol. 67: 1047-1064.

Struik, G.J., and J.T. Curtis. 1962. Herb distribution in an Acer saccharum forest. Amer. Midi. Nat. 68: 285-296.

Sukatschew, W. 1928. Einige experimentelle Untersuchungen uber den Kampf urns Dasein zwischen Biotypen derselben Art. Z. indukt. Abstamm. -u. VererbLehre 45: 54-74.

Syers, J.K., and I.K. Iskander. 1973. Pedogenic significance of lichens, p. 225-248. I_n V. Ahmadjian and M.E. Hale (eds.) The Lichens. Acad. Press, New York.

Tansley, A.G. 1917. On competition between Ga1ium saxatile L. (G. hercynicum Weig.) and Galium sylvestre Poll. (G. asperum Schreb.) on different types of soil. J. Ecol. 5: 173-179.

Tansley, A.G. 1939. The British Isles and their vegetation. Cambridge Univ. Press, Cambridge.

Taylor, W.P. 1922. A distributional and ecological study of Mt. Rainer, Washington. Ecology 3: 214-236.

Terjung, W.H., R.N. Kickert, G.L. Potter, and S.W. Swarts. 1969. 209

Energy and moisture balances of an alpine tundra in mid- July. Arct. Alp. Res. 1: 247-266.

Tieszen, L.L., and E.K. Bonde. 1967. The influence of light intensity on growth and chlorophyll in arctic, subarctic, and alpine populations of Deschampsia caespitosa and Trisetum spicatum. Univ. of Colo. Studies, Series in Biol. No. 25. 21 p.

Tieszen, L.L., and N.K. Wieland. 1975. Physiological ecology of arctic and alpine photosynthesis and respiration, p. 157- 200. In F.J. Vernberg (ed.) Physiological Adaptation to the Environment. New York.

Tipper, H.W. 1971. Glacial geomorphology and Pleistocene history of central British Columbia. Can. Geol. Surv. Bull. 196. 87 P.

Tranquillini, W. 1964. The physiology of plants at high altitudes. Ann. Rev. Pi. Physiol. 15-: 345-362.

Travers, O.R. 1975. Cathedral Provincial Park Expansion Proposal Impact Evaluation. Resource Planning Unit, E.L.U.C. Secretariat. 60 p.

Turesson, G. 1922. The genotypical response of the plant species to the habitat. Hereditas 3: 211-350.

Turesson, G. 1923. The scope and import of genecology. Hereditas 4: 171-176.

Turkington, R., and J.L. Harper. 1979a. The growth, distribution and neighbour relationships of Trifolium repens in a permanent pasture. I. Ordination, pattern and contact. J. Ecol. 67: 201-218.

Turkington, R., and J.L. Harper. 1979b. The growth, distribution and neighbour relationships of Trifolium repens in a permanent pasture. II. Inter- and intra-specific contact. J. Ecol. 67: 219-230.

Turkington, R., and J.L. Harper. 1979c. The growth, distribution and neighbour relationships of Trifolium repens in a permanent pasture. IV. Fine-scale biotic differentiation. 210

J. Ecol. 67: 245-254.

Turkington, R., P.B. Cavers, and L.W. Aarssen. 1977. Neighbour relationships in grass-legume communities. I. Interspecific contacts in four grassland communities near London, Ontario. Can. J. Bot. 55: 2701-2711. van Ryswyk, A.L. 1969. Forest and alpine soils of south-central British Columbia. Ph.D. thesis. Washington State Univ., Pullman, Washington. 178 p. van Ryswyk, A.L., and R. Okazaki. 1979. Genesis and classification of modal subalpine and alpine soil pedons of south-central British Columbia. Arct. Alp. Res. 11: 53-68.

Ward, R.T. 1969. Ecotypic variation in Deschampsia caespitosa L. Beauv. from Colorado. Ecology 50: 519-522.

Watt, A.S. 1947. Pattern and process in the plant community. J. Ecol. 35: 1-22.

Webber, P.J. 1974. Tundra primary productivity, p. 445-473. I_n J. Ives and R.G. Barry (eds.) Arctic and Alpine Environments. Methuen, London.

Webber, P.J., and D.E. May. 1977. The magnitude and distribution of below-ground plant structures in the alpine tundra of Niwot Ridge, Colorado. Arct. Alp. Res. 9: 157-174.

Werner, P.A. 1979. Competition and coexistence of similar species, p. 287-310. In O.T. Solbrig, S. Jain, G.B. Johnson, and P.H. Raven (eds.) Topics in Plant Population Biology. Columbia Univ. Press, New York.

Westgate, J.A., D.G.W. Smith, and H. Nichols. 1969. Late Quaternary pyroclastic layers in the Edmonton Area, Alberta, p. 179-186. I_n S. Pawluk (ed.) Pedology and Quaternary Research. Univ. Alberta, Edmonton, Alta. 218 p.

Whitfield, C.J. 1933. The ecology of the vegetation of the Pike's Peak Region. Ecol. Monogr. 3: 75-105.

Whittaker, R.H. 1975. Communities and ecosystems, 2nd ed. 21 1

MacMillan, New York. 385 p.

Whittaker, R.H. 1978. Approaches to classifying vegetation, p. 3-31. In R.H. Whittaker (ed.) Classification of Plant Communities. Junk, The Hague.

Whittaker, R.H., and H.G. Gauch. 1978. Evaluation of ordination techniquies, p. 277-336. In R.H. Whittaker (ed.) Ordination of Plant Communities. Junk, The Hague.

Wieland, N.K., and F.A. Bazzaz. 1975. Physiological ecology of three codominant successional annuals. Ecology 56: 681-688.

Wiens, J.A. 1977. On competition and variable environments. Amer. Sci. 65: 590-597.

Wilcox, R.E. 1965. Volcanic-ash chronology, p. 807-816. I_n H.E. Wright, Jr. and D.G. Frey (eds.) The Quaternary of the United States. Princeton Univ. Press, Princeton, New Jersey.

Williams, J.T. 1969. Biological Flora of the British Isles: Chenopodium rubrum L. J. Ecol. 57: 831-841.

Williams, J.T., and J.L. Harper. 1965. Seed polymorphism and germination. I. The influence of nitrates and low temperatures on the germination of Chenopodium album. Weed Res. 5: 141-150.

Williams, W.T., G.N. Lance, L.J. Webb, J.G. Tracey, and M.B. Dale. 1969. Studies in the numerical analysis of complex rain-forest communities. III. The analysis of successional data. J. Ecol. 57: 515-535.

Woodwell, G.M. 1974. Variation in the nutrient content of leaves of Quercus alba, Quercus coccinea, and Pinus rigida in the Brookhaven forest from bud-break to abscission. Amer. J. Bot. 61: 749-753.

Woodwell, G.M., R.H. Whittaker, and R.A. Houghton. 1975. Nutrient concentrations in plants in the Brookhaven oak- pine forest. Ecology 56: 318-332. 212

Yarranton, G.A. 1966. A plotless method of sampling vegetation. J. Ecol. 54: 229-237. 213

APPENDIX A - GEOLOGICAL HISTORY

During the late Paleozoic and early Mesozoic eras, a large inland sea covered much of western North America, from Alaska to California. Marine sedimentation and extensive vulcanism characterized this period. The sea drained near the end of the Triassic and the sediments and lava were compressed and folded during a period of deformation in the Jurassic (Rice 1960). These metamorphosed rocks were then intruded by magma, which crystallized to form granitic plutons, such as Lakeview granodiorite and and Cathedral quartz monzonite (Daly 1912). Early in the Cretaceous period, erosion exposed earlier intrusive rock (Rice 1960) and the Cathedral Park area was included within a new, narrow marine basin (Rudkin 1964). Sedimentation and vulcanism characterized this time, with subsequent termination by Late Cretaceous orogeny (folding and faulting), followed by erosion and peneplain formation (McTaggart 1970). No rock of the Cretaceous age is presently exposed in Cathedral Park (Melcon 1975). Lavas and local lake sediments were deposited on the leveled land during the Tertiary and are represented in Cathedral Park as Mid-Eocene Princeton basalt and sedimentary rock (Rice 1960). Vulcanism and sedimentation were followed by folding and erosion, and the granites, diorites, and granodiorites of the eastern North Cascades were produced during this time (Misch 1966). Further uplift occurred in the late Pliocene and Pleistocene epochs with subsequent erosion and weathering forming deep, dissected valleys with residual hills and mountains of more resistant rock, termed monadnocks, rising above the elevated Teriary peneplain (R ice 1960, Holland 1964). Lakeview Mountain is an example of such a monadnock.

During the Pleistocene epoch, the Cordilleran ice sheet covered most of British Columbia, with the final advance, the Fraser Glaciation (late Wisconsinan), lasting approximately 15,000 years (Crandell 1965). The Fraser glaciation produced the bulk of glacial features seen in British Columbia (Tipper 1971). Tipper (1971) cites evidence for a pre-Fraser, Fraser, and very limited post-Fraser glaciation in the central interior of B.C. The chronology of a subsequent Hypsithermal Interval and Neoglaciation has been outlined by Porter and Denton (1967). Ice sheet development began in mountainous terrain with the advance of alpine glaciers. Mountain ice sheets gathered around the Coast and Cariboo-Columbia Mountains and subsequently merged. A south-eastern segment of ice formed after this coalescence, which flowed across the Washington-British Columbia border after additions from the Monashee Mountains, south Coast Mountains, and other local centers (Nasmith 1962, Tipper 1971). There is evidence that the continental ice covered mountains as high as 2590-2620 m in the study area, although elevations above 2286 m were not reached in Wisconsin time (Prest 1957, Rice 1960). Holland (1964) and Nasmith (1962) both state, however, that the ice did not attain elevations over 2134 m at its 214

maximum in southern British Columbia. Evidence for a 2286-2377 m upper limit of effective glacial erosion has been given by Melcon (1975) for Cathedral Provincial Park where tor landforms (rock exposed in situ by chemical and physical weathering) occur exclusively above this upper limit. Intense alpine glaciation occurred above the upper limit of the Cordilleran ice sheet and produced cirque basins on north and north-east aspects and serrate peaks and ridges; in contrast to the smooth and rounded topography of areas below the ice sheet (Holland 1964). The effects of alpine glaciation above 2440 m is marked in Cathedral Park.

Vulcanism was extensive during the late Pliocene and early Pleistocene and volcanic cones such as Mt. Baker and Glacier Peak were superimposed on existing Cascade Mountain peaks during this time (Coombs 1939). Surfical ash deposits reported within and to the south of the study area (Bockheim 1972, van Ryswyk 1969) resulted from a number of recent volcanic eruptions. Major sources of ash are likely to be Glacier Peak, Mt. Mazama, and Mt. St. Helens in the Cascade Mountains. The oldest ash source is Glacier Peak, erupting 12,000 years ago and distributing ash to southern British Columbia, Alberta, and Montana (Powers and Wilcox 1964, Fryxell 1965, Wilcox 1965). Extensive ash deposits resulted from the Mt. Mazama eruption at Crater Lake in southern Oregon approximately 6600 years B.P., occurring as far north as southern British Columbia (Powers and Wilcox 1964, Wilcox 1965, Westgate et al. 1969). Mt. St. Helens produced three ash deposits, dated at 160 year B.P. (Mullineaux 1964), 500 years B.P., and 3000 years B.P. (Crandell et al. 1969), although known ash distributions suggest only the 3000 year B.P. ash reached as far north as Cathedral Park (Okazaki et al. 1972, Nasmith et al. 1967). Little or no ash from the most recent Mt. St. Helens eruption (1980) is evident in the study area. It is not likely Mt. Rainier ash (2300 and 2000 years B.P.) (Crandell et al. 1969, Wilcox 1965), or Bridge River ash (2400 years B.P.T" (Nasmith et al. 1967) reached Cathedral Park. 215

APPENDIX B - SOILS

Soils of the Princeton map area, which includes the Cathedral Park study area, have been classified by Green and Lord (1979). Soils at the forest-alpine transition within this map area were determined by these workers to be members of the Alpine Subgroups of the Dystric Great Group of the Brunisolic Order (Soil Survey Committee of Canada, 1968). Soils within the alpine zone of Lakeview Mountain have been examined in detail by van Ryswyk (1969) and van Ryswyk and Okazaki (1979). Three different soil types were characterized by van Ryswyk (1969), the most extensive (over 50% of the area studied) being Alpine Brown with discontinuous ash layers, van Ryswyk (1969) stated that this soil type was comparable to the Alpine Dystric Brunisol of the Canadian System, although a later report (van Ryswyk and Okazaki 1979) referred to this soil as an Orthic Sombric Brunisol. Alpine Brown, Discontinuous Ash soils are included within the Incepticol Order of the U.S. Soil Classification System. Douglas and Bliss (1977), working near the study area in the North Cascade alpine of Washington, reported Inceptisols beneath a wide variety of alpine plant communities from snowbed sites to well-drained dry graminoid and sedge vegetation. Alpine Brown, Discontinuous Ash soils are also comparable to the Alpine Turf Great Soil Group described by Retzer (1956, 1962), Johnson and Billings (1962), and Nimlos and McConnell (1962) for the Rocky Mountains of Colorado, Wyoming, and Montana, and by Bockheim (1972) for Mt. Baker in the North Cascades. Alpine Brown, Discontinuous Ash soils develop on moderately acid parent material of medium to coarse texture. They are characterized by thin organic surface layers (L-H) and moderately thick turfy Ah over Bm horizons (van Ryswyk 1969).

Alpine Brown, Continuous Ash profiles were also characterized on Lakeview Mountain (van Ryswyk 1969). These soils have a B horizon similar to the Bf horizon found in Podzol soils. Sombric Humo-Ferric Podzols have been classified in this alpine zone on the basis of sesquioxides and organic matter accumulation in the illuvial B horizon (van Ryswyk and Okazaki 1979), Alpine Podzols (Spodosols) have also been reported under alpine krummholz and heath vegetation in the North Cascades (Douglas and Bliss 1977) with further reports from several alpine areas in British Columbia (Sneddon 1969, Sneddon et al. 1972, Alberta (Baptie 1968, Broad 1973), and Maine (Bliss and Woodwell 1965).

Structure soils such as sorted stone patterns and stone rivers have also been described on Lakeview Mountain (van Ryswyk 1969) and are correlated with the reduction of snow cover found on exposed ridges. Extensive lichen cover suggests recent stability. In addition, broken rock units occur on promentories, rock headwall and talus units occupy cirque basins, and snowbank units characterize lee positions. 216

Buried horizons are common in Lakeview alpine soils, indicating severe disturbance by cryoturbation as well as slope wash and wind erosion. Charcoal fragments, larger than would be possible from existing woody vegetation, have been found in surface and buried horizons at altitudes up to 2440 m (van Ryswyk 1969). The past existence of a more extensive forest of coniferous trees and ericaceous shrubs has been argued by van Ryswyk and Okazaki (1979). A charcoal fragment found 70 cm below the soil surface at an elevation of 2485 m on Lakeview Mountain has been dated at 9120 years B.P. (van Ryswyk 1971), Ash buried with the charcoal has been identified as Mt. Mazama and Mt. St. Helens ash, deposited between 3000 - 3500 years B.P., indicating burial occurred after 3000 years B.P. A period of intense solufluction at the close of a minor glacial interval 1200 - 1000 years B.P. has been reported by Benedict (1966) and is possibly when horizon burial occurred.

Soil parent materials are considered residual and consist primarily of coarse, partially weathered quartz monzonite crystals (Melcon 1975) and other medium-grained diorite type rocks with fine-grained volcanic materials (van Ryswyk 1969). Fine silt or clay is absent from the parent material, and the small amounts that occur in these soils are formed in situ or deposited by percolating ground water (Melcon 1975). Much of the drainage in these soils occurs as seepage over nearly impermeable C horizons. All soils have been influenced by volcanic ash, burying and mixing of horizons, frost heaving, solufluction, surface erosion, and colluvial activity (van Ryswyk 1969). Slow rates of chemical weathering due to cold temperatures and the erosive action of physical processes contribute to the general immaturity of soil profiles found in alpine areas (Retzer 1974). 217

APPENDIX C ~ PERCENTAGE COVER DATA FOR SIX TRANSECTS

Percentage cover data for composite samples from transects A-F are presented in this appendix. Rare species have less than 5 occurrences in each transect data set or cover values of less than 5% in all transect samples. Trace (T) denotes less than 5% cover. 218

TRANSECT A COMPOSITE SAMPLES

1 2 3 It 5 6 7 8 9 10 11 12 13 15 16 20 21 22 Ik 26 VASCULAR SPECIES 23 25 27 28 29 JO

ARENARIA OBTUSILOBA 15 8 8 8 T T 8 T 12 T T 13 T T 23 T 15 T 7 8 12 18 T 9 10 10 13 13 CAREX CAPITATA 3 25 23 38 27 20 *5 <0 17 28 25 15 37 33 28 12 12 30 37 35 33 13 32 27 33 hi <<3 38 CAREX NAROINA T T T 5 T 17 25 18 23 25 25 18 T 12 7 7 CAREX SCIRPOIDEA 12 17 20 28 28 itO 22 23 15 17 28 17 33 32 33 27

ANDROSACE SEPTENTRIONALIS T T T T T T ANTENNARIA ALP INA T T T T T T ANTENNARIA UMBRINELLA T T T T T T T ARENARIA RUBELLA DRABA CANA T T T DRABA INCERTA T DRABA LONCHOCARPA T T T ORABA PAYSONII TTTT TT T TTTT T T T T HAPLOPAPPUS LYALLII T T T T T T T POA ALPINA T T POA SP. T POLYGONUM VIVIPARUM POTENTILLA NIVEA SALIX NIVALIS 15 SEDUM LANCEOLATUM TTTT T TT TTT SENECIO LUGENS TTTTT T TT TT T T SIBBALDIA PROCUMBENS T TARAXACUM CERATOPHORUM T T

LICHENS 5 BRYOPHYTES

CETRARIA CUCULLATA 8 7 5 T 10 5 5 6 23 23 18 13 7 13 6 T T T 12 15 18 7 10 12 12 13 20 12 25 10 CETRAR1A ISLANDICA 20 22 20 15 16 1(0 25 12 9 16 lit 20 27 37 38 38 33 Ul 33 28 20 22 30 20 15 28 23 23 28 20 CETRARIA NIVALIS 13 13 10 8 9 5 6 13 5 8 15 12 11 T T T T T 6 15 8 T 15 12 17 8 15 15 18 13 CLAD INA MITIS T T 5 T T T T T T T T T T T T T T T T T T T T CLADONIA SP. T T T 6 5 T T T T T T T T T 5 T T 6 5 T T T T T T T T T T CORNICULARIA ACULEATA T 7 7 5 T T T 9 20 20 17 13 8 T T T T T T 12 18 12 17 23 28 23 18 12 17 DESMATOOON SP. T T 10 T T T T T T T T OCHROLECHIA UPSALIENSIS T T T T 5 T T T T T T T T T T T T T T T 5 T T T T POLYTRI CHUM JUNIPERINUM T T T T 10 T T POLYTRICHUM PILIFERUM T T T T 29 5 7 T 30 10 T T T T T T T T T T T T T T T T THAMNOLINA VERMICULARIS 6 13 13 12 17 17 8 8 T 8 10 8 5 5 8 10 13 15 15 7 8 13 15 8 8 12 12 12 18 12 20 RARE LICHENS

CALOPLACA SP. T T T T T T T T T T T T T T T T T T T T T T T T CANOELARIELLA SP. T T T T T T T T T T T T T T T T T T T T T T T CLADONIA CHLOROPHAEA T T T T T T T T T T T T T T T T T T T T T T T DACTYLINA RAMULOSA T T T T T T LETHARIA VULPINA T T PELT IGERA CAN 1NA T T T T T T T T T T T T T

BARE GROUND T T T T T T T 10 8 T T T T 8 T T T T T 8 8 T T T T T T T T ROCK 5 T T T 7 T 20 11 5 8 T T T 6 12 5 T T T T T T T T T 5 219

TRANSECT B COMPOSITE SAMPLES

1 2 3 1« 5 6 7 8 10 11 12 14 16 VASCULAR SPECIES 9 13 15 17 18 19 20 21 22 23

ARENARIA OBTUSILOBA 5 6 12 18 111 5 T 6 T 8 15 8 6 18 T 17 T 13 20 T 15 15 20 8 13 15 T 7 12 CAREX CAPITATA 30 45 48 42 55 50 58 57 48 48 37 35 32

ANDROSACE SEPTENTRIONALIS T T T T T ANTENNARIA ALPINA T ARENARIA RUBELLA

CAREX BREWERI1 T T T • • 1 DRABA LONCHOCARPA TTTT T T T T 5 DRABA PAYSONII T T T T T T LUPINUS LYALLII T PENSTEMON PROCERUS 10 POA RUPESTRIS TTTT T TT TT POLEHONIUM PULCHERRIMUM T 12 8 T POLYGONUM VIVIPARUM T POTENT ILLA NIVEA T TTT T T RANUNCULUS ESCHSCHOLTZI I y T T T SEOUM LANCEOLATUM TTTTTT T T T SENECIO LUGENS T T T T T T T TTTTTT T T T T T T SIBBALDIA PROCUMBENS T jjjfTTTTTTTT STELLARIA LONGIPES T T T T T T T T T T TTTTTTTTTT T T T T T

LICHENS & BRYOPHYTES ? ]? ,y 20 )8 20 20 12 12 20 13 T 5 9 22 20 22 17 23 18 20 15 10 15 27 28 33 22 35 28 20 26 23 22 22 20 28 13 10 8 TT 13 10 20 11 8 12 CETRARIA NIVALIS T7 T T T 1T0 1T3 1T0 1T2 T T T T T T T T T 7 5 CLAD I NA MITIS 7 5 T T T T TI iT T T T T T T CLADONIA CHLOROPHAEA 7 T T T T T T T T T T T T T CLADONIA SP. 1188. 5 7 5 T T T T 6 7 5 6 5 5 5 T T T T 5 T T T T T I T T T T CORNICULARIA ACULEATA T T T 8 8 T 5 13 8 6 8 13 8 12 17 8 13 15 17 27 20 17 28 22 18 10 6 8 OCHROLECHIA UPSALIENSIS TTT T T T T T T T T T T T T T T T T T T 6TTT5TTT5 T T T PELT IGE RA CAN INA 5 T T T T T POLYTRI CHUM PILIFERUM 25 12 13 40 12 17 11 13 10 8 5 30 8 5 T 15 18 10 12 10 15 5 T 12 8 T T T 5 5 - - - T 7 22 13 22 THAMNOLINA VERMICULARIS 11 15 17 13 10 12 15 15 10 10 17 37 12 12 8 10 10 13 15 17 15 12 7 7 12 12 7 12 10 17 RARE LICHENS & BRYOPHYTES

CALOPLACA SP. T TTTTTTTTTTTTT T T T T T T T T TTTTTTTTTTTT CANDELARI ELLA SP. TTTTTTTT T T T T T T T T T TTT DESMATODON SP T T T T T T T T T T T T T LETHARIA VULPINVUI A TT TTT^T^ J T T T

BARE GROUND T T 5 T T T 5 T T T T T T T 5T57TTT57TT57T5 ROCK T T T T T 13 T T TTTT TT10T13 220

TRANSECT C COMPOSITE SAMPLES

1 2 } A 6 8 10 11 12 1A 16 VASCULAR SPECIES 5 7 9 13 15 17 18 19 20 21 22 23 2A 25 26 27 28 29 30

ANTENNARIA ALP 1NA T T T T T T T T T T T 6 ARENARIA OBTUSILOBA 8 8 7 10 10 12 T 6 8 18 8 11 12 8 13 13 13 15 10 T 7 8 7 15 8 15 10 12 20 8 CAREX CAPITATA 32 12 A2 11 22 30 13 20 A3 38 30 32 20 18 A7 35 A7 35 38 1(2 17 5 28 60 33 25 38 22 35 27 CAREX NARDINA T T T 7 T 5 10 CAREX SCIRPOIDEA 25 18 8 15 15 20 13 12 38 A5 AO 32 20 35 17 28 23 25 25 27 9 22 27 32 20 53 35 33 A2 A3 DRABA PAYSONI1 T T T 5 T T T T T T T T T T T T T T T T T T T ERIGERON AUREUS 5 T 6 f 5 T 7 6 5 T T 8 5 6 7 T 6 T T T T 5 T T T 6 T T T FESTUCA OVINA 13 8 7 5 5 T 5 5 T 10 10 7 9 8 7 10 7 10 13 10 T 5 10 10 8 15 12 10 8 10 LUPINUS LYALLI1 T 7 T 5 7 8 5 T T T 5 8 7 T T T T T T T LUZULA CAMPESTRIS T T T 6 T T T 6 T T T T T T T T T 8 T T T T T T 5 T T 5 7 OXYTROPIS MONTI COLA T T T 5 5 T T 5 T 8 T T T 5 9 T T 7 6 T T T 5 T T PENSTEMON PROCERUS T T 13 5 10 25 POA ALPINA 8 T T T T T T T T 5 T T POA RUPESTRIS T 5 T T T T T 5 T T T T 5 T T T T T T POTENTILLA DIVERSIFOLIA 15 12 18 8 6 6 T 7 13 17 18 11 5 12 10 15 18 13 13 12 8 13 8 15 17 17 15 27 23 20 SAL IX NIVALIS A3 17 23 22 25 17 22 20 8 13 A8 AO SELAGINELLA DENSA T 6 T T 5 T T T T T 7 6 T T 8 T T T T 8 T 5 T T T T 5 9 12 SENECIO LUGENS 7 5 6 T T T 7 T T T T T 5 T T T T T T T 7 T SILENE ACAULIS T 9 8 17 6 8 T 15 T 5 8 8 T T T T T T T T T T 7 T 12 7 STELLARIA LONGIPES T T T T T T T T T T T T T T T T T T T T T T T T T T 5 T T TRISETUM SPICATUM T 8 T 7 T T T T 6 12 T 6 T T T T T 6 18 5 T T 5 7 T T T T 6 T RARE VASCULAR SPECIES

ANDROSACE SEPTENTR1ONAL1S T T T T T T T T ARENARIA RUBELLA T T CAREX PHAEOCEPHALA T 13 7 23 1 CERASTIUM BEERING ANUM T T T T T T T T T T T T T T T DRABA INCERTA T T T T T T DRABA LONCHOCARPA T T T T T T T T T T T T DRYAS OCTOPETALA T 38 T 1 HAPLOPAPPUS LYALLI T T T T T T T T T T T T T T T KOBRESIA MYOSUROIDES 17 POA SP. T T POLEMONIUM PULCHERR1 MUM 13 POLYGONUM VIVIPARUM T T T T T T T T POTENTILLA NIVEA T T T SEDUM LANCEOLATUM T T T T T T T T T T T T SOLI DAGO MULTIRADIATA 8 T T T LICHENS S BRYOPHYTES

CETRARIA CUCULLATA 10 T T T T T T T T 8 T 9 22 20 25 8 28 12 17 16 7 8 17 22 13 T 15 8 12 15 CETRARIA ISLANDICA 27 T 18 9 T 22 16 8 28 30 15 18 12 12 25 13 28 22 13 30 18 7 17 15 IA 17 20 8 11 7 CETRARIA NIVALIS 12 7 5 T T 8 T T 5 T 5 13 15 10 13 15 10 17 15 8 9 6 15 8 T 8 15 15 13 8 CLADONIA SP. T T T T 6 8 7 5 9 T T T T T T T T T T T T T T T T 6 T T T T CORNICULARIA ACULEATA T T T T T T T T T T T T 6 8 8 T 8 15 17 T T T 8 8 13 DESMATODON SP. T T T T T T T T T 5 T T T T 7 T T T T T T T T T POLYTRICHUM PILIFERUM 8 T T T T T 7 9 15 20 5 9 8 10 26 13 8 12 6 9 T 5 T T T T T T T THAMNOLINA VERMICULARIS 12 7 18 10 T 8 9 6 13 13 10 12 10 10 8 7 10 10 T 10 T T 7 7 T 8 10 10 12 12 RARE LICHENS 6 BRYOPHYTES

CALOPLACA SP. T T T T T T T T T T T T T T T T T T T T T T T T T T T T T T CANDELARI ELLA SP. T T T T T T T T T T T T T T T T T T T T T T T T T T T T T CLADINA MIT IS T T T T T T T T T T T T T CLADONIA CHLOROPHAEA T T T T T T T T T T T T T T T T T T T T T T T T T T T T DACTYLINA RAMULOSA T T T T OCHROLECHIA UPSALIENSIS T T T T T T T T T T T T T T T T T T T T T T T T T 1 PELTIGERA CAN NA T T T T T T T T T T T T T T T T T T T T T T POLYTRICHUM JUNIPERINUM T

BARE GROUND 12 7 10 7 8 8 15 17 7 T 7 8 12 10 12 12 T 5 8 12 T 7 8 5 8 12 12 15 7 10 ROCK 5 T T 27 17 T 18 T T 5 13 30 17 T T T T T 13 7 5 5 5 T T T 5 221

TRANSECT D COMPOSITE SAMPLES

VASCULAR SPECIES 1 2 3 ** 5 6 7 8 9 10 11 12 13 lA 15 16 17 18 13 20 21 22 23 2A 25 26 27 28 29 30

ANTENNARIA ALP INA TTT6T T TT5TTT7TT T85TT T

ARENARIA OBTUSILOBA 13 13 15 6 13 10 15 9 8 T T 12 8 13 12 20 12 18 9 T 10 15 25 8 18 12 7 17 1"t 15

CALAMAGROSTIS PURPURASCENS T T T 5 15 12 10 1*4 lA 8 8 5 T 8 T T T

CAREX NARDINA 5TT TT12 .TT CAREX SCIRPOIDEA 17 22 23 25 20 20 18 23 12 20 25 22 13 22 22 30 23 15 10 7 9 15 8 8 25 5 7 10 8 17 ER IGERON COMPOSITUS TTTTTT5 TTTT T FESTUCA OVINA T5TTTTT6T5 TTTT5TTTTT8TTTTTT6T HAPLOPAPPUS LYALLII T7T6 10 TT TT KOBRESIA MYOSUROIDES 11 33 15 15 32 38 8 25 lA T 30 33 32 28 17 LUZULA CAMPESTRIS 6T TTTT T T TT OXYTROPIS MONT I COLA T T 11 10 T T T T POTENTILLA DIVERSIFOLIA 77T10 10 10 10 8 15 25 10 868TTT8568 10 86666 12 96 POTENTILLA NIVEA TTTT10 9TTTT10 8 10TT57 10 557T

SELAGINELLA DENSA 17 787T8T10T557TTT TTTTTTTTTT13 15 10T SILENE ACAULIS 6 5 5 5 T T T SOLIDAGO MULTI RADI ATA T6 TT T 15 TTT7588T T T7 T TARAXACUM CERATOPHORUM TT5TT T 55TT8TTTT 6 6 T 5 T T RARE VASCULAR SPECIES

AGOSERIS GLAUCA T T T T T T ANDROSACE SEPTENTRIONAL IS T T T T ARENARIA RUBELLA TTTT TTTT CAREX CAPITATA CAREX PHAEOCEPHALA T CERASTIUM BEERINGIANUM T T T T T DRABA AUREA T T DRABA CANA TTTT T T T T T T T DRABA INCERTA T T T T T TT TTTTT DRABA PAYSONII T T T T T T T T T T T T T ERIGERON AUREUS T T LUPINUS LYALLII TTTT T T POA ALPINA TTTT T POLEMONIUM PULCHERRIMUM TTTT T POTENTILLA FRUTICOSA 18 RANUNCULUS ESCHSCHOLTZI I SEDUM LANCEOLATUM T T T T T T T T T . . . . . , T T T T TRISETUM SPICATUM TTT T TT T TT TTT TTTT TTT LICHENS t, BRYOPHYTES CETRARIA CUCULLATA TTTTTTT66T5TTTTTTT7TTTT9TTT95T CETRARIA ISLANDICA TTTTT TTTT55TTTTTTTT5TTTTTTT68 CETRARIA NIVALIS TTTTTTT55TT5TTTT5TT57T7T8 10 7698 CLADONIA SP. TTTTTTTTT 5TTT5T55TT5TT88T15 10 CORNICULARIA ACULEATA T7TT55575TT8TT5 10 755T7T5T58868 LETHARIA VULPINA TT TTTTTT TT TT10 7T6TTTTT8TTTTT OCHROLECH IA UPSALIENSIS TTTTTTTTTTTTTTTTTTTTTTT6TT9785 POLYTRICHUM JUNIPERINUM TTT TT 10 TT TT RARE LICHENS S BRYOPHYTES

CANDELARI ELLA SP. TTTTTTTTTTTTTTT TTTTTTTTTTT CLAD INA MITIS ' T T CLADONIA CHLOROPHAEA T T T T T T DESMATODON SP. T T T T T PELTIGERA CAN INA T T TTTTT, T TT• . T i T TTTTT THAMNOLINA VERMICULARIS T TTTTTTTTTT TTTT TTTTTTT BARE GROUND 10 12 15 13 7TTT5T5TT5TTTTTT5 10TT85 17 5 8 7 13 18 ROCK 10 8 12 5 22 38 7 T 35 20 T 30 12 13 10 T T 32 T T 5 5 T 25 5 12 8 8 222

TRANSECT E COMPOSITE SAMPLES

1 It 11 IA VASCULAR SPECIES 2 3 5 6 7 8 9 10 12 13 15 16 17 18 19 20 21 22 23 2A 25 26 27 28 29 30

ANTENNARIA ALPINA T T T T T 7 T T T T T T T T T ANTENNARIA MICROPHYLLA T 5 T T 6 T T ARENAR1 A OBTUSILOBA T 8 5 T 6 20 5 9 T 7 12 13 10 5 8 8 8 10 5 5 8 17 12 12 15 10 12 12 20 10 CAREX CAPITATA 12 T T T T CAREX NARDINA T T 5 8 T T T T 5 T T T T T T T T CAREX SCIRPOIDEA 15 12 10 30 22 25 12 11 7 6 22 27 AO 33 37 33 27 33 38 28 A8 A2 28 23 A2 32 AO A7 AO 38 DRABA PAYSONI1 T T T T T 5 T T T T T T T T T T T T T T 5 T T T T ERIGERON COMPOSITUS T T 5 T T 7 T T T T 5 T T T FESTUCA OVINA T T T T 5 5 T 5 5 7 7 T 7 5 10 T 8 8 5 9 8 12 7 5 10 10 5 7 5 HAPLOPAPPUS LYALLI1 T T T 5 T T T T T T T 12 T 8 12 12 8 8 18 5 T T 10 KOBRESIA MYOSUROIDES 13 A8 53 30 32 30 A5 50 58 52 30 7 LUPINUS LYALLI1 T T T T T T T 8 T T 5 8 T T T T T T T T T T T T T T 6 T T LUZULA CAMPESTRIS T T T T T T T T T T T 7 5 6 8 T 8 5 T T T 8 6 8 7 5 8 7 8 T OXYTROPIS MONTI COLA T T T T T T 13 POA RUPESTRIS T T T T T T T T 10 T T 7 T 10 T 5 T T POTENTILLA DIVERSIFOLIA T T T T T T 5 T T T T 9 10 5 5 7 6 T 9 5 7 10 7 T T 5 10 12 5 POTENTILLA NIVEA T T T T T T T T T T T 5 T 6 T 5 T 18 7 T T T SELAGINELLA DENSA T T T 6 10 5 10 8 T 6 T 8 T T 5 6 7 T 5 18 10 8 8 7 7 T 13 8 T 10 SENECIO LUGENS T T T T 5 T T T T T T T T T T 5 T T SILENE ACAULIS T 5 7 5 10 12 7 13 T 30 10 13 18 6 T 22 TARAXACUM CERATOPHORUM T T T T T 6 T 8 T T T T TRISETUM SPICATUM T T 5 T T T T 8 8 6 T T T T T T . T T T 5 T 6 T RARE VASCULAR SPECIES

ANDROSACE SEPTENTRIONALIS T T T ANTENNARIA UMBRINELLA T ARENARIA RUBELLA T T T T T T T T T T T T CERAST1UM BEERING1ANUM T T T T T T T T T T T T T DRABA INCERTA T T T T T T T T T T T T T T T T DRABA LONCHOCARPA T T T T T T T T T T T T T T T T T T ERIGERON AUREUS T T T T T T T T T T T T T T T T T POA SP. T POLEMONIUM PULCHERR1 MUM T T POLYGONUM VIVIPARUM T 8 T SAL IX NIVALIS 5 T SEDUM LANCEOLATUM T T T T T T T SOLI DAGO MULTIRADIATA T 7 T STELLARIA LONGIPES T T T T T

LICHENS 5 BRYOPHYTES

CETRARIA CUCULLATA 12 17 15 15 10 20 10 12 18 12 8 T T 7 6 7 8 7 T T T 8 T T T 5 7 T T 10 CETRARIA ISLANDICA T 7 8 5 6 5 8 5 7 T T T T T T T T T T T T 17 T T T T T 7 8 7 CETRARIA NIVALIS 17 27 30 15 22 20 14 22 22 17 15 7 10 10 18 8 10 22 18 10 10 22 12 8 12 6 8 10 15 10 CLADONIA SP. T T T T T T 5 5 5 T T T T T 5 5 T T T T T T T T T T 5 T T T CORNICULARIA ACULEATA 7 5 8 10 5 10 5 8 15 15 13 8 12 18 13 12 12 15 12 10 18 12 15 13 12 10 13 10 13 20 LETHARIA VULPINA T T T T 10 T T R T T T T T T 6 T T T T T 7 T T T T 8 T T OCHROLECHIA UPSALIENSIS T T T 10 T T T T T T T T 5 T T T T T 5 T T T T T T T 7 T T T POLYTRICHUM PILIFERUM T T T 10 5 T T T 5 8 12 7 T 15 T T T 6 T T 10 8 8 10 15 THAMNOLINA VERMICULARIS 10 8 8 5 5 10 6 5 T T T T T T 5 T T 7 T T T T T T T T T T T RARE LICHENS 6 BRYOPHYTES

CALOPLACA SP. T T T T T T T T T T T T T T T T T T T T T T T T T CANDELARIELLA SP. T T T T T T T T T T T T T T T T T T T T T T T CLAD 1NA MIT 1S T T T T T T . CLADONIA CHLOROPHAEA T T T T T T T T T T T T T T T T T T T T T T DESMATODON SP. T T T T T T T T T T T T T T PELT 1GERA CANINA T T T T T T T T T T T T T T T T T T T T T T T T POLYTRICHUM JUNIPERINUM T T T T T T BARE GROUND T T 5 T T T 13 10 T 12 8 8 10 10 7 10 5 10 22 22 7 8 7 18 13 12 15 7 7 T ROCK T 7 8 15 22 15 5 T T T 22 32 30 10 15 15 35 10 8 28 27 5 25 33 35 28 23 25 13 5 223

TRANSECT F COMPOSITE SAMPLES

1 2 4 3 5 6 7 8 9 10 11 12 13 14 16 18 VASCULAR SPECIES 15 17 19 20 21 22 23 24 25 26 27 28 29 30

ANTENNARIA UMBRINELLA T T T T T T T ARENARIA OBTUSILOBA T 5 8 8 13 T 15 9 7 8 17 7 15 T T 10 27 13 5 T T T 13 14 22 27 5 20 18 T 6 CAREX CAPITATA T 7 22 T 20 17 40 20 23 45 53 57 A3 42 33 A7 32 35 45 37 8 22 23 42 22 CAREX NARDINA T T 12 12 T T 7 17 15 8 7 T 5 18 10 7 CAREX PHAEOCEPHALA 15 T 5 T T T CAREX SCIRPOIDEA 20 17 30 12 17 T 12 27 25 20 20 18 6 27 27 23 20 40 27 27 18 25 25 30 42 43 38 32 22 27 ERIGERON AUREUS T T T T 7 T T T T T T 7 15 T 5 T 8 FESTUCA OVINA 12 T 11 8 T 8 9 13 13 10 12 13 10 7 10 10 12 12 8 8 6 10 7 13 8 12 12 12 KOBRESIA MYOSUROIDES 9 13 43 53 A3 42 25 60 55 17 12 LUPINUS LYALLII T 7 T T T 6 5 T 6 T T T 7 T T T T T T T T T LUZULA CAMPESTRIS T 7 T T T T 5 7 T T 5 7 T T T 5 T T 7 T 7 5 7 T T 7 10 OXYTROPIS MONT I COLA T T T T T 12 T T 10 8 T T T 8 T T 5 T 8 8 5 PENSTEMON PROCERUS T 10 T T 17 T 8 7 T 10 T T T T 23 6 13 11 13 8 8 7 5 POA SP. T T 5 5 T T T POTENTILLA DIVERSIFOLIA 15 15 15 12 15 7 11 13 15 T 10 20 12 5 13 17 13 16 13 25 17 9 7 T 12 11 13 13 13 12 SELAGINELLA DENSA 8 12 18 7 5 8 8 T T 6 8 T T 5 T 12 T T T T T 5 T T T 10 T T 5 SENECIO LUGENS T T T T T T 6 T T T 13 SILENE ACAULIS T T T T T T T T T 15 T T 8 SOL I DAGO MULT I RAD I ATA T T 10 5 T 5 8 16 12 7 T 10 T T T 13 T 17 STELLARIA LONGIPES T T T 5 T T T 8 8 5 T T T T T T T T 5 5 T T 5 T T T T 5 7 TRISETUM SPICATUM T T T T T T 6 T T T T T T T T T T T T T 6 6 8 7 T 7 RARE VASCULAR SPECIES

ANOROSACE SEPTENTRIONALIS T T TTTT T T ARENARIA RUBELLA T CERASTIUM BEER INGIANUM T T T T T T T T T T DRABA CANA T T T DRABA INCERTA DRABA PAYSONII T T TT . . _ _ T T HAPLOPAPPUS LYALLI I TTT POLEMONIUM PULCHERRI MUM 1 - POTENTILLA NIVEA T T. SEDUM LANCEOLATUM TTT T TTTT T T T T T T T T T T T T . TARAXACUM CERATOPHORUM

LICHENS S BRYOPHYTES

8 5 T 7 o8 ij i1z2 i/ io io ^ 13 13 15 12 12 10 10 22 8 in in 1? in CETRARIA CUCULLATA „ 3 , / 115 13 17 18 10 5 13 13 15 12 12 10 1 0 1177 1155 22 I1S5 1177 8 17 CETRARIA ISLANDICA 100 13 9 13 200 7 2S5 188 188 71 11 ^ 270n ,1 22 « ™ c li \o lr 11 11 1 '° '? '? Z '2 23 5 23 25 25 20 15 25 28 25 22 23 T T 5 15 CETRARIA NIVALIS 11 7 6 6 10 T 7 17 17 10 6 13 9 13 13 T 10 6 8 18 8 6 15 17 8 T T 6 20 7 CLADONIA SP. 6 10 15 6 T 10 6 6 7 T 10 7 5 T T 6 5 T 5 T T 5 8 12 6 10 T T T T CORNICULARIA ACULEATA 10 18 12 15 27 17 12 10 8 12 18 15 13 17 25 32 10 25 27 7 6 8 T 8 5 3 6 7 2 2 LETHARIA VULPINA T T T '° " " ^ ' ' 'I " « '° " 'I I ! Z I ! ? Z 'Z ? T T T T T T T T T 6 T T T OCHROLECHIA UPSALIENSIS 10 7 T T T 7 6 T T T 6 T T T T T T 5 5 T T T T 5 6 10 5 6 T 6 POLYTRI CHUM JUNIPERINUM 23 T T T 5 10 8 T T 6 15 7 5 T T 8 T POLYTRI CHUM PILIFERUM T T T 5 T 5 T 10 15 T 6 T T T T T 5 T 10 5 18 11 THAMNOL NA VERMICULARIS 13 I T T T T 6 T 7 8 6 8 T T 6 5 10 8 T T T 13 7 12 8 8 T T T 12 7 RARE LICHENS 6 BRYOPHYTES

CALOPLACA SP. T T T T T T T T T T T T T T T T TTT T CANDELARI ELLA SP. T T T T T T T T T T T T T T TTT T CLAD INA MIT IS T T T T T T T T T T T T T T T T T CLADONIA CHLOROPHAEA T T T T T T T T " T T T T T T T T T T T T T T T T T T T I T T T T DACTYLINA RAMULOSA T T T T T DESMATODON SP. T T TTT T T T T T T T PELT I GERA CAN I NA T T T TTT T T T T T T T T T T T T T T T T T T TTT T BARE GROUND 10 8 13 12 15 17 T T T 8 5 T 5 10 T 12 T 5 5 6 12 7 T 10 5 11 T 8 T 7 ROCK 7 5 T T T 7 10 8 T T T T T T 5 8 T 7 5 7 10 8 17 18 T 8 224

APPENDIX D - PRINCIPAL COMPONENTS ANALYSIS OF SOIL DATA

Physical and chemical soil properties within 11 vegetation groups were analyzed with centered and standardized PCA. Data for this analysis are given in Table X. A scatter plot is provided for A horizon data. Vegetation groups are indicated by letters and numbers, described previously.

PCA Eigenvectors for Soil Variables

A Horizon B Horizon

Axis 1 Axis 2 Axis 1 Axis 2

% Variance (50%) (18%) (45%) (30%)

% sand .27832 .53672 .20359 .50231 % silt .34389 .44853 . 19359 .49998 % clay .29243 .25348 .17153 .43382 % organic matter .32565 .25138 .41011 .079477 % nitrogen .43162 . 13390 .43730 .01 9385 phosphorus (ppm) .23633 .22467 .11216 . 10309 potassium (meq) .20949 .44542 . 19399 .36564 magnesium (meq) .37569 . 1 2566 .36416 .301 1 5 calcium (meq) . 1 7979 .29738 .381 10 .24967 CEC (meq) .39142 . 1 1 081 .45184 .073933 225

PCA Scatter Plot of 11 Vegetation Groups using A Horizon Soil Data

* D3 D2

C2 * El

F2 A # E2 + * Fl C1

AXIS 1