Vegetation and Environment Patterns of Liard River Hot Springs Provincial Park, British Coluwbia \

$Vegetation and Environment Patterns of Liard River Hot Springs Provincial Park, British Coluwbia \$

VEGETATION AND ENVIRONMENT PATTERNS

OF LIARD RIVER HOT SPRINGS PROVINCIAL

PARK, BRITISH COLUMBIA

Terry Charles Reid

B.Ed., University of Calgary, 1970

A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF

THE REQUIREMENTS FOR THE DEGREE OF

MASTER OF SCIENCE

in the ~epdrtment

0 f

Biological Sciences

@ Terry Charles Reid 1978

SIMON FRASER UNIVERSITY

April 1978

Name: Terry Charles Reid

Degree: Master of Science

Title of Thesis: Vegetation and Environment Patterns of Liard River Hot Springs Provincial Park, British Coluwbia \

Examining Committee:

Chairman: L. J. Albright

R. C. Brooke Senior Supervisor

R. W. Mathewes

Date ~~~p/oved:\q dl 197~5 / PARTIAL. COPYRIGHT LICENSE

I hereby grant to Simon Fraser University the right to lend my thesis, project or extended essay (the title of which is shown below) to users of the Simon Fraser University Library, and to make partial or single copies only for such users or in response to a request from the library of any other university, or other educational institution, on its own behalf or for one of its users. I further agree that permission for multiple copying of this work for scholarly purposes may be granted by me or the Dean of Graduate Studies. It is understood that copying or publication of this work for financial gain shall not be allowed without my written permission.

Title of Thesis/Project/Extended Essay

Vegetation and environment patterns of Liard River Hotsprings

Provincial Park, British Columbia

Author: (sfgnature)

Terry Charles Reid

(date) ABSTRACT

Vegetation-environment patterns of Liard River Hot Springs Park in northern British Columbia, including the influences provided by the thermal springs, are described in this study. Specific objectives were to analyze the microclimatic patterns, describe and classify the soils and vegetation, and to determine soil influences on community types and selected species.

Field work was conducted from May 1974 to July 1975.

Seven microclimatic station8 monitored air temperature, soil temperatures, precipitation, and relative humidity. The general temperature regime of the area is that of a microthermal boreal cold continental climate.

Microclimatic differences between thermally influenced and non-thermally influenced sites include air temperature, soil temperature, the estimated frost-free period, and precipitation.

~dilprofile descriptions were made from 29 sample plots representing the ran&,,of plant communities, parent materials, and topographic positions

'\ - - represented in-tke-studyarea. Approximately 125 soil samples were chemically analyzed. Soils were classified as Brunisols (Orthic Melanic, Orthic

Eutric, Degraded Eutric, and Degraded Dystric), Regosols (Orthic), Gleysols (Rego, and Carbonated), Organic (Hydric Mesisol, and Hydric Humisol), and

Sub-aqueous (Lake Chalk Protopedon). Tufa (calcium carbonate) was prominent in many of the soils.

Sixty-three sample plots were established in the study area for vegetation analysis. Five aquatic (Chara - Potamogeton, Chara - Utricularia, Mimulus - Riccia, Erigeron - Mimulus, and Typha - Scirpus), six non-forested (Aster - Muhlenbergia, Viola - Spiranthes, Eleocharis - Triglochin, Oplopanax - Gymnocarpiurn, Matteuccia - Actaea, and Apocynum - Cynoglossum), and four iv forested vegetation types (Picea mariana - Pleurozium, Larix - Potentilla, Populus - Viburnum,and Picea glauca - H~locomium)were described and classified. Several types were found only in thermally influenced areas.

The height of the water table was a frequent environmental difference between vegetation types.

Vegetation-environmental relationships were examined for both vegetation types and selected species. Phenological observations on selected species revealed that development in thermally influenced areas was at least one week earlier than development in areas not under a thermal influence from the hot springs. Potentially limiting soil factors could be documented for ten vegetation types: the Chara - Potamopeton, Chara - Utricularia, Mimulus - Riccia, Erigeron - Mimulus, Aster - Muhlenbergia, Viola - Spiraothes, Eleocharis - Triglochin, Larix - Potentilla, Apocynum - Cynoglossum, and Picea mariana - Pleurozium communities. Nine selected species were statistically evaluated for significant relationships with selected edaphic chemical characteristics by using correlation and, where appropriate, regression methods. Significant results between the cover of

Aralia nudicaulis, Rosa acicularis, Cornus stolonifera, and Triglochin palustris and various soil variables were demonstrated. Significant soil variables were correlated with exchangeable calcium. DEDICATION

This is dedicated to my family, because their numerous sacrifices made this study possible. ACKNOWLEDGEMENTS

I would like to thank Drs. R. C. Brooke, R. M. F. S. Sadleir, and M. McClaren for their assistance during field and/or analysis stages of the study. They also provided helpful comments during the writing of the thesis. Dr. Brooke also provided Figures 4, 8, 24, and 28.

The Environment and Land Use Committee Secretariat provided the loan of some meteorological instruments and provided analytic help by reading the hygrothermographs and summarizing the temperature data. Soil analysis was performed by the Department of Soil Science at the University of

British Columbia. Selected bryophyte determinations were kindly provided by Dr. W. B. Schofield of the University of British Columbia and the late

Dr. C. C. Chuang of the British Columbia Provincial Museum.

Financial support for soil analysis and the field portion of the study was provided by the British Columbia Parks Branch. The remainder of the study was financed by Simon Fraser University (President's

Research Grant and Teaching Assistantships) and by my wife. vii TABLE OF CONTENTS

Page

TITLE PAGE APPROVAL PAGE ...... ii ABSTRACT ...... iii DEDICATION ...... v ACKNOWLEXEMENTS ...... vi TABLE OF CONTENTS ...... vii LIST OF TABLES ...... xi i LIST OF FIGURES ...... xv INTRODUCTION AND OBJECTIVES ...... 1 REGIONAL AND STUDY AREA DESCRIPTIONS ...... 4 Location ...... 4 Previous Work ...... 4 Physiography ...... 6 Geology ...... 8 Thermal Springs and Tufa Deposits ...... 11 Human History ...... 15 Climate ...... 16 Soils ...... 16 Vegetation ...... 17 CLIMATE AND MICROCLNTES ...... 20 Objectives...... 20 Methods ...... 20 Stations and Instrumentation...... 20 Air Temperature ...... 22 viii.

Page Measurements ...... 22 Monthly Temperatures ...... 22 Microclimates ...... 23 Mean Annual Temperatures ...... 23 Frost-free Period ...... 24 Soil Temperature ...... 24 Precipitation ...... 25 Relative Humidity ...... 25 Results and Discussion...... 25 Air Temperature...... 25 Monthly Temperatures ...... 25 Microclimates ...... 27 Mean Annual Temperatures ...... 30 Frost-free Period ...... 30 Soil Temperature...... 32 Precipitation ...... 35 Relative Humidity ...... 39 SOILS ...... 42 Objectives ...... 42 Methods ...... 42 Description and Classification of the Soils ...... 43 Results ...... 43 Brunisolic Order ...... 45 Regosolic Order ...... 49 Gleysolic Order ...... 50 Organic Order ...... 52 ix

Page

Sub-aqueous Soils...... 54 Discussion...... 54 Chemical Characteristics of the Soils...... ,...... 55 Results ...... 5 5 Discussion...... 58 VEGETATION ...... 60 Introduction ...... 60 Methods...-...... 60 Vegetation Units...... 65 Chara - Potamogeton (C - P) Vegetation Type 67

Chara - Utricularia (C - U) Vegetation Type 68 Mimulus - Riccia (M - R) Vegetation Type.. . 69 Erigeron - Mimulus (E - M) Vegetation Type. 72 Typha - Scirpus (T - S) Vegetation Type.... 75 Aster - Muhlenbernia (A - M) Vegetation Type,...... 77 Viola - Spiranthes (V - S) Vegetation Type. 80 Eleocharis - Triglochin (E - T) Vegetation Type ...... 84 Oplopanax - Gymnocarpiurn (0 - G) Vegetation Type...... 88 Matteuccia - Actaea (M - A) Vegetation Type 89 Apocynum - Cynoglossum (A - C) Vegetation Type ...... 93 Picea mariana - Pleurozium (Pm - P) Vegetation Type...... 96 Page

Larix . Potentilla (L . P) Vegetation Type . 99 Populus . Viburnum (P . V) Vegetation Type . 105 Picea glauca . Hylocomium (Pg . H) Vegetation Type ...... 109 VEGETATION . ENVIRONMENT RELATIONSHIPS ...... 113 Relationship of Individual Species to Micrometeorological Differences ...... 113 Relationship of Vegetation Types to the Soil Order ... 115 Relationship of Vegetation Types to Soil Chemical Characteristics ...... 117 Method of Analysis ...... 117

Cation Exchange Capacity ...... 121 Percent Base Saturation ...... 122 Exchangeable Calcium ...... 122 Exchangeable Magnesium ...... 123 Calcium/Magnesium Ratio ...... 123 Exchangeable Sodium ...... 123 Exchangeable Potassium ...... 124 Percent Nitrogen and the Carbon/Nitrogen Ratio ...... 125 Available Phosphorous ...... 127 Percent Sulfur ...... 129 Relationship of Individual Species to Soil Chemical Characteristics ...... 129 General Approach ...... 129 Page Spearman Rank Correlation Coefficients ..... 132 Product-Moment Correlation Coefficients .... 135 Regression Equations ...... 139 Discussion ...... 143 SUMMARY ...... 151 APPENDICES A . Mean daily maximum and minimum air temperatures (F) summarized by months ...... B . Mean weekly maximum and minimum air temperatures (F) summarized by months ...... C . Frost-free period (days) ...... D . Summary of soil temperature data (F) ...... 161 E . Summary of precipitation data (inches) ...... 162 F . Surmnary of relative humidity data (2.)...... 163 G . Soil chemical analysis results classified by vegetation type ...... 164 H . Terminology used in the VEGETATION section and in Appendix I...... 173 I . Site and edaphic characteristics classified by vegetation type ...... 174 J . Sulfate dissolved in thermal springs (mg/l) ...... 185 K . Summary of weighted mean edaphic values to the bottom of the maximum rooting concentrations as

classified by vegetation type and profile segment 186 L . Plant species list ...... 187 BIBLIOGRAPKY ...... 199 LIST OF TABLES

Page

Table I. Water temperatures [C (F)] at Liard River Hot Springs Park...... Table 11. Climatic and microclimatic stations of the

study area...... ,......

Table 111. Comparison of observed temperatures (F) with

long-term averages......

Table IV. Statistically significant (pi 0.05) maximum and

minimum air temperature differences between

pairs of stations......

Table V. Estimates of annual mean air temperatures...... Table VI. Annual mean soil temperatures...... Table VII. Comparison of observed precipitation (in.) with

long-term averages......

Table VIII. Statistically significant (pQ 0.05) differences

in precipitation (mm) between pairs of stations.

Table IX. Station relative humidity data (%) summarized

by microclimatic group...... ,......

Table X. Synopsis of soil categories classified and described...... Table XI. Mean chemical characteristics of soils

classified by order and horizon......

Table XII. Definitions for strata limits...... Table XIII. Species significance scale...... xi i i Page

Table XIV. Categories of species used to characterize and distinguish between vegetation types...... Table XV. Summary of the characteristic combination of species for vegetation types ...... Table XVI. Characteristic combination of species and synthesis table (C - P)...... Table XVII. Characteristic combination of species and synthesis table (C - U) ...... Table XVIII. Characteristic combination of species (M - R) ... Table XIX. Synthesis table (M - R) ...... Table XX. Characteristic combination of species (E - M) ... Table XXI. Synthesis table (E - M) ...... Table XXII. Synthesis table (T - S) ...... Table XXIII. Characteristic combination of species (A - M) ... Table XXIV. Synthesis table (A - M) ...... Table XXV. Characteristic combination of species (V - S). .. Table XXVI. Synthesis table (V - S) ...... Table XXVII. Characteristic combination of species (E - T) ... Table XXVIII. Synthesis table (E - T) ...... Table XXIX. Synthesis table (0 - G) ...... Table=. Characteristic combination of species (M - A). .. Table XXXI. Synthesis table (M - A)...... Table XXXII. Characteristic combination of species (A - C) ... Table XXXIII . Synthesis table (A - C)...... Table XXXIV. Characteristic combination of species (Prn - P),. Table XXXV. Synthesis table (Pm - P) ...... xiv Page Table XXXVL. Characteristic combination of species (L - P). . 101 Table XXXVII. Synthesis table (L - P).' ...... 102 Table XXXVIII. Selected differences between the Eleocharis - Triglochin and Larix - Potentilla vegetation types...... Table XXXIX. Characteristic combination of species (P - V).. Table XL. Synthesis table (P - V) ...... Table XLI. Characteristic combination of species (Pg - H). Table XLII. Synthesis table (Pg - H) ...... Table XLIII. Selected phenological observations for 1975....

Table XLIV. Distance between the soil surface and the water

table classified by soil order......

Table XLV. Weighted mean soil values classified by vegetation type...... Table XLVI. Species - edaphic Spearman correlation coefficients (rs) ...... 134 Table XLVII. Non-significant (p) 0.05) species - edaphic product-moment correlation coefficients (r) .... 13 7 Table XLVIII. Statistically significant (pC0.05) species - edaphic product-moment correlation

coefficients (r) ...... 138 Table XLIX. Species - edaphic regression equations...... 140 xv LIST OF FIGURES

Page

Figure Liard River Hot Springs Park...... 5

Figure 2. Park features, microclimatic station locations, and

vegetation patterns of the area studied...... 7

Figure 3. Hot springs and wetlands are located in a

depression which may have served as a meltwater

channel separating the terrace (left) and plateau hillside (right) ...... 10 Figure 4. A spring in the Epsilon Hot Pool complex ...... 11 Figure 5. Canadian thermal springs ...... 12 Figure 6. The Hanging Gardens are formed from tufa precipitated from the waters of Chi Warm Spring .... 13 Figure 7. Monthly mean, mean maximum and minimum, and extreme

maximum and minimum air temperatures for control station LCP ...... 26 Figure 8. A comparison of daily temperature range patterns... 31

Figure 9. Number of continuous weeks of freezing ((OC or <32F)

soil temperatures...... 34 Figure 10. Total precipitation for the twelve month period July 1974 to June 1975...... 37 Figure 11. An Orthic Melanic Brunisol in an Apocynum - Cynoglossum vegetation type (Plot 14) ...... 4 6 Figure 12. An Orthic Eutric Brunisol in a Matteuccia - Actaea vegetation type (Plot 16) ...... 47 Figure 13. A Degraded Eutric Brunisol in a Populus - Viburnum vegetation type (Plot 05) ...... 48 xv i

Page

Figure 14. An Orthic Regosol in an Apocynum - Cynoglossum vegetation type (Plot 55) ...... 50 Figure 15. A Rego Gleysol (peaty phase) in a Picea mariana - Pleurozium vegetation type (Plot 48) ...... 51 Figure 16. A Hydric Mesisol in an Eleocharis - Triglochin vegetation type (Plot 59) ...... 53 Figure 17. Mimulus guttatus (yellow monkey flower) dominates

the vascular flora. The water is occupied by the

rare Riccia fluitans, tufa-forming Chara vulgaris,

and the calciphyte Cratoneuron filicinum...... 69

Figure 18. Erigeron philadelphicus (Philadelphia fleabane)

and Mimulus gut tatus (yellow monkey flower) characterize the vascular flora...... 72 Figure 19. This vegetation, characterized by Typha latifolia

(common cattail) and Scirpus validus (common great

bulrush), is rare in the study area...... 75

Figure 20. Muhlenber~ia glomerata (bog muhly) and Aster

junciformis (rush aster) are conspicuous on these tufa islands located in a thermal pond ...... 77 Figure 21. Viola - Spiranthes vegetation (Plot 34) was found on small tufaceous islands surrounded by thermal

water...... 81

Figure 22. The continuous vegetation cover consists of such

characteristic species as Tofieldia glutinosa

(sticky tofieldia), Lobelia kalmii (Kalm's

lobelia), and Parnassia palustris (grass-of- xvii

Page

Parnassus) ...... 81

Figure 23. Scirpus caespitosus (tufted clubrush), Eleocharis

pauciflora (beaked spike-rush), Triglochin

palustris (marsh arrow-grass), and Carex spp.

(sedges) are common on hummocks. Moose disturbed

the water-saturated soil ...... 84

Figure 24. Oplopanax horridum (Devil's club) was rare in the

study area...... 88

Figure 25. Matteuccia struthiopteris (ostrich fern) is the most abundant species in the Matteuccia - Actaea vegetation type...... 90

Figure 26. Dogbane (Apocymun androsaemifolium) is the most abundant species in the Apocynum - Cynoglossum vegetation type...... 93 Figure 27. A conspicuous shrub layer is absent from this

community. Picea mariana (black spruce),

Pleurozium schreberi, Ptilium crista-castrensis,

and Hylocomium splendens are all abundant species.. 96

Figure 28. Humus hummocks in a fen-like habitat provided the rooting environment for the Larix - Potentilla vegetation type. The foreground is occupied by

Eleocharis - Triglochin, and the background has a Picea mariana - Pleurozium cover...... 100

Figure 29. Tall Populus tremuloides (trembling aspen) and

Betula papyrifera (paper birch) characterize the

tree layer of this vegetation. Characteristic xviii

Page,

understory species include Viburnum edule (high-

bush cranberry), Cornus stolonifera (red osier dogwood), and Aralia nudicaulis (sarsaparilla) ..... 105 Figure 30. The vegetation, consisting largely of Picea glauca

(white spruce) and Hylocomium splendens was located on a steep hillside...... 110 Figure 31. Relationships between soil orders and vegetation types...... 116 Figure 32. Regression lines for plant species and edaphic variables...... 142 INTRODUCTION AND OBJECTIVES

Aquatic springs, in contrast to most aquatic systems, have a relatively constant chemical composition, water velocity, and temperature

(Odum 1959). Communities may thus be studied under approximately constant environmental conditions. Thermal springs have additional features that make them of interest to an ecologist. These features include the establish- ment of a thermal gradient, and low species diversity within the community

(Brock 1970). Brock (1970) also noted that the usual isolation of thermal springs is analogous to an island in terms of biotic movement.

The uniqueness of thermal springs has prompted a number of investigations. North American studies date back to at least 1897 when studies were conducted on the hot spring vegetation of Yellowstone National Park.

Pioneering work usually consisted of compiling an annotated list of thermal spring species (e.g. Brues 1924, 1928, 1932), whereas more recent investigators are often interested in more specific problems such as the relationship between environmental factors and species distribution and abundance

(e.g. Brock 1969, Bott and Brock 1969, Brock and Darland 1970).

Scant attention has been paid to North American boreal thermal springs.

Brandon (1965) discusses the geology and geochemistry of Canadian thermal areas, while Nava and Morrison (1974) made a brief reconnaissance of several Alaskan thermal areas.

The uniqueness of boreal thermal spring vegetation has often been commented on (Camsell 1954, Nava and Morrison 1974, Porsild and Crum 1961,

Scotter et al. 1971, Scotter and Cody 1974). Luxuriant vegetation and the occurrence of species that do not grow beyond the thermal influence of the springs are two characteristics that have often been noted. Botanical information on North American boreal springs is comparatively

sketchy. Henry (1933) described a pack train trip to collect at Toad River

Hot Springs, British Columbia but few collections were cited in the paper.

Although plants were collected at Hutlinana River Hot Springs, Alaska, the

collection was not listed in the paper (Nava and Morrison 1974). Surveys

in connection with the International Biological Program have resulted in

sketchy botanical information for Coal River Springs, Yukon Territory

(Dennington, undated), Toitye Hot Springs, Northwest Territories (Cody and

Acton 1973), and Lymnaea Springs, Northwest Territories (Cody and Sneddon

1973). More detail is available for collections from the Nahanni National

Park area, Northwest Territories which contains many thermal springs

(Scotter et al. 1971, Scotter and Cody 1974). Published collections exist

for Hole-in-the-Wall Hot Spring, Northwest Territories (Porsild 1961) and for Liard River Hot Springs, British Columbia (Porsild and Crum 1961).

The only work dealing with the plant ecology of a North American boreal thermal spring is a small (34 p.) thesis concerning the plant

communities of Hole-in-the-Wall Hot Spring (Arnold 1961). This work

established five plant communities on the basis of sampling 30 rn2 of vegetation (10 belt transects of 15 cm by 20 m). These communities were

then related to humus thickness and the distance to the water table.

In summary, the knowledge of North American boreal thermal spring vegetation remains in an exploratory state. Ecological information on

the vegetation of these areas is extremely brief.

The Interpretation Division of the Parks Branch, Department of

Recreation and Conservation, co-sponsored the following study with Simon

Fraser University. The field portion of the study was carried out at

Liard River Hot Springs Park, Mile 496.5 Alaska Highway, British Columbia, and lasted from late May of 1974 through July 1975. The major objectives of this study were:

1. to determine the climate of the study area.

2. to determine if microclimatic differences occurred between thermally

influenced and non-thermally influenced areas of the study area.

3. to classify and describe the soils within the study area.

4. to classify and describe the plant community types within the

study area.

5. to establish relationships between environmental variables and the

plant community types and certain selected species. REGIONAL AND STUDY AREA DESCRIPTIONS

Location

The study area was confined to Liard River Hot Springs Park, Mile 496.5,

Alaska Highway, British Columbia. This park is located in the north-central part of the province (59' 26' N latitude and 126' 06' W longitude) approximately 56 kilometers (35 miles) south of the Yukon-British Columbia border.

The park consists of approximately 668 hectares (1,650 acres) and is located to the north and east of the intersection of the Alaska Highway and the

Liard River (Figure 1). Important study area features are shown in Figure

2. Field work was concentrated in this portion of the park since all known thermal pools are located in this area.

Previous Work

The construction of the Alcan Military Road resulted in easy access to areas adjacent to this highway. The road from Fort Nelson, British

Columbia to Watson Lake, Yukon Territory was investigated for soils

(Leahey 1943), geology (Williams 1944), mammals (Rand 1944a, Baker 1951), and birds (Rand 1944b). The study area is given brief coverage in some of these publications. A recent publication (Erskine and Davidson 1976) records bird observations for the Fort Nelson area. The geology of the region which includes the study area has been mapped by Gabrielse (1962).

The study area has been the subject of several Parks Branch reports of which Lyons (1954, 1956) contains the most descriptive detail of the earlier series. The park vegetation was later studied by Porsild and

Crum in their 1961 publication for the Natural Museum of Canada. Several additions to the flora were subsequently included in Brayshaw (1971).

Pavlick (1974) has produced a park interpretive assessment and interpretive plan which also includes a comprehensive section on the physical setting.

Other recent work concerns the study area plant communities (Reid 1974b), birds and mammals (Reid 1975b), historical botany (Reid 1975b, 1976), and microclimate and soils (Reid 1977). Several additions to the flora and fauna were noted in these reports. The known park vascular flora is about

300 species, with over 230 species included in the present study (Appendix L).

An additional thermal springs complex was also discovered (Reid 1975b) and . is referred to in this paper as the "Tau" Warm Springs complex. Park use and demographic patterns have also been recently reported (Reid 1974a,

Physiography

The Liard River separates two different physiographic regions: the

Liard Plateau to the north and the Rocky Mountains to the south of the river (Bostock 1970). Holland (1964) regards the Liard Plateau as the southern end of the Mackenzie Mountains. He describes the plateau as generally' consisting of round or flat summits and timbered ridges below

1,524 m (5,000 feet) of elevation which are incised to elevations below

762 m (2,500 feet) by Liard River tributaries. North trending Devonian to Triassic folded sedimentary rocks underlie the plateau (Holland 1964).

In the study area ridges of the Liard Plateau are present in the form of spurs which extend toward the Liard River on the eastern boundary of

L.f.. . .*::;:(..'.'. ,.

the park and outside the western boundary of the park (Figure 1). Mt. Ole is one of these ridges (Pavlick 1974). The hotsprings and associated wetlands are located at 457 m (1,500 feet) elevation (Souther and Halstead

1973) on the margin of the plateau surface.

Geology

The bedrock geology has been mapped by Gabrielse (1962). The following account is summarized from his investigation.

Folded and faulted sedimentary rock are found around the park. Tight folding of strata is characteristic of the Terminal Ranges of the northern

Rocky Mountains which are located south of the Liard River. Some of these strata have been overturned but this did not occur near the vicinity of

Liard River Hot Springs Park. By contrast, the rocks north of the Liard

River generally occur in relatively simple, open folds.

Middle Devonian calcareous deposits of limestone and dolomite are known to occur within four miles both north and south of the park.

Immediately north of the park is an anticline of this carbonate rock which is overlain by the clastic strata found in the park. Glacial drift covers much of the bedrock, particularly in the Liard Valley and Liard Plateau.

The study area is underlain by upper Devonian and Missippian clastic strata. This map-unit may include shale, slate, argillite, siltstone, and sandstone. The bedrock outcrops in Liard River Hot Springs Park consists of these rock types. Siltstone is prominent at the Liard River and Porsild and Crum (1961) reported Elt. Ole bedrock as argillite and sandstone. A veneer of glacial drift covers much of the study area. Pavlick (1974) has reported finding (within the park) samples of granite and quartzite which he stated were probably deposited by glacial transport.

During the Pleistocene, the Cassiar Mountains were the source of a northeasterly and easterly moving ice sheet (Gabrielse 1962, Prest et al.

1968). Much of this ice flowed easterly across the Liard Plateau

(Holland 1964, Prest et al. 1968), and Gabrielse (1962) has reported glacial erratics near the top of Mount Halkett (located about 18 km

[ll miles] north of the park). Some ice also flowed eastward down the

Liard River Valley (Holland 1964, Prest et al. 1968) from its source in the

Kechika River area of the North Rocky Mountain Trench (Prest et al. 1968). .

No evidence was found of ice covering the higher mountains of the Terminal

Range located south of the park. Ice therefore covered the present park area plus the Liard Plateau to the north and at least the lower elevations of the Rocky Mountains to the south.

Pavlick (1974) has suggested that the two Liard Plateau ridges which are found on either side of the hot springs (Figure 1) would have acted as an embayment area. Glacial meltwaters flowing into this embayment would have their velocity reduced which could have resulted in the deposit of sediments. In this fashion a glacial river terrace would be built up from the present Liard River to near the base of the plateau hillside

(Figure 3). A glacial meltwater channel would separate the terrace and plateau hillside, and a tufa-alluvial fan complex would develop in the region of the thermal springs between the meltwater channel and the plateau hillside. This postulation is consistent with the known information for the study area.

.Postglacial time for northern British Columbia has been estimated as beginning some 10,000 years ago (Heusser 1960). . -

' ,;&j g > , .,'4 ? Figure 3. Hot springs and wetlands are located in a $B@~M~Q@ may have served as a meltwater channel separ&M& kt@ @mace (left) and plateau hillside (right). The foreground pond is the "Large Cold ~ond"and the other is Alpha Warm Ponded Swamp. View is from' Mt, Ole.

- Table I. Water temperatures (C (F$ at Liard River Hot Springs Park Date July 18, 1973a Aug. 3, 1974 May, 1975~

Hot Springs and Swamp :' Alpha Spring: outlet 53.3 (128) >50,0 (7122) m (m) : dam 43.3 (110) 43.2 (110) 40.9~ (106)' ' Epsilon Springs 48.3 (119) 44.4 (112) 42,7 Beta Spring 41.7 41.0 (106) 38.9' {::;Ic 41.1 ti:;] m (m) 39.W (102 Swamp 36.1 rn (m) 34.4 94) 34% $1 31.3' (88)'

.= Warn, Springs and swam^ Alpha Warm Ponded Swamp 31.1 (88) 31.0 (88) 25.6 (78) 24.4 m 22.7 (73) 23.3 m 21.7 (71) "Tau" Springs: western m m 19.7 (67) : eastern m m 17.6 (64) 13.9 (57) m 12.3 (54) a: from Pavlick 1974. f four weekly readings on May 6, 13, 27, and June 3. eks of data are missing.

.4 .4 Thermal Springs and Tufa Deposition

Figure 4. A spring in the Epsilon Hot Pool complex.

The pool inlet is at the bottom ~f the spring. MLcroclimatic station EP was located immediately adjacent to a similar spring in this complex.

Souther and Halstead (1973) defined springs as "discharges of groundwater of sufficient volume to cause flowage at the surface", which may be considered thennal "if their temperatures are more than 5' C or 10' F above the mean annual air temperature of the region". They subdivided them1 springs into warm springs if the water temperature is below 32.2 C (90 F), or hot springs if the water temperature is above 32.2 C (90 F), Souther and

Halstead (1973) defined mineral springs as generally having "more than 1,000 parts per million of total dissolved solids". Liard River springs are mineral

(Pavlick 1974) and are all thermal with both hot and warm springs being present (Table I). These springs may be classified as the submarine type

(Jennings 1971) with the inlet below the water surface (Figure 4). 1 Legend

Figure 5. Canadian thermal springs

The location of Liard River Hotsprings is indicated by the arrow. (Modified from Souther and Halstead 1973.)

The Liard River thermal springs are not a unique phenomenon for

British Columbia. The Cordilleran Region of Canada has many thermal

springs (Figure 5) due to the relatively high precipitation in a mountainous

topography (Souther and Halstead 1973). The high relief would permit the

circulation of water through the spring systems.

Pavlick (1974) mentions several mechanisms by which thermal springs

may be created. These include volcanic activity, rising magmatic gasses,

exothermic chemical reactions such as the oxidation of sulfide minerals, or

the water being heated by the thermal gradient as it travels down a thrust

plane. He suggested that this latter possibility may be the mechanism for 13 the Liard River Hot Springs since faulting is known for the general area.

The recharge area for the springs is believed to be in the hills to the north of the perk randon on 1965). According to this view water would flow through Devonian limestone bedding planes to the discharge area in me

Liard Valley. The height difference of roughly 457 hl (1,900 feet) between the Liard Plateau and the Liard Valley would presumably account for deep

groundwater circulation.

The Hanging Gardens are formed from tufa precipitated from the waters of Chi Warm Spring. (March 1975.) Within the Liard thermal springs complex, tufa deposits are common

and form the tufa dykes surrounding Alpha and Epsilon Ponded Swamps, the

tufa dome surrounding Beta Hot Pool, and the Hanging Gardens (Figure 6).

Tufa (calcium carbonate) accumulation in the study area can be explained

by the following generalized scheme summarized by Jennings (1971): CaC03 (solid) + H20 + C02 (dissolved) caw + 2 HCQ~- f 1 C02 (air)

\ This author states that the amount of dissolved CaC03 varies

directly with the partial pressure of COZ, and inversely with water

temperature. Precipitation of calcium carbonate would occur when a

saturated solution loses C02 (by diffusion to the atmosphere or due to

C02 uptake by tufa-forming vegetation) or H20 (by evaporation). Jennings

(1971) points out that these activities are promoted by turbulence and ,

aeration such as water flowing over an initial surface irregularity.

Tufa deposition has been similarily explained by Sweeting (1973,

p. 108) as being due to:

"either an extensive loss of C02 or the action of algae, mosses, and other lime-secreting plants. In the tropics, in addition to these two factors, evaporation of water is important."

The ability of some wetland species to secrete lime is an important 1 tufa-forming mechanism. Cratoneuron commutatum and Chara vulgaris are

especially noted for this ability in the study area (Porsild and Crum 1961,

Pavlick 1974). Cratoneuron camnutaturn is an important lime-secreting

species in other areas of the world (e.g. the Plitvice lakes, Jugoslavia - see Sweeting 1973). Prescott (1970) discussed Chara vulgaris and noted:

..L- 1 Plant nomenclature authorities are given in Appendix L, p. 187 ". . . in their physiology the plants cause lime to become deposited on stem and 'leave' . . . calcareous deposits may be formed largely by Chara over long periods of time."

The differential lime-secreting ability of different species could be noted in the field. An example occurs in the Mimulus - Riccia vegetation

type in which Chara vulgaris was heavily lime encrusted but Mimulus guttatus was not.

Human History

The following account of the human history of the study area is based

on Camsell (1954) and Tarasoff (1974).

The thermal springs were probably known by trappers, prospectors, and

the Teslin Indians. The first written record of the springs on the Liard

is believed to have been in the diary of Robert Campbell, a Hudson Bay

Company Factor stationed at Fort Liard in 1835 and the now defunct Fort

Halkett after 1839. The first known record is in 1898 when William Ogilvie

and Charles Camsell explored the Liard. They camped at the springs both

that year and the next year. A trapper and his daughter (Tom and Jane

Smith) lived at the thermal springs for two years in the early 1920's. On

their journey to Fort Liard with their fur cache, their craft overturned,

the trapper was drowned, and the daughter was rescued by Indians and sub-

sequently taken to Fort Liard. However, when the two did not appear at

Fort Liard in 1925, friends asked Lt. Col. J. Scott Williams to land on

the Liard River during his mineral reconnaissance flights in that area.

His subsequent landing and exploration of the hot springs area apparently

resulted iri the myth of the "Tropical Valley". Newspaper reports later featured reports of such exotic species as monkeys, bananas, parrots, and dinosaurs in a remote, northern, lush, tropical valley. Camsell fought this myth, and again visited the hot springs in 1935 during a geological aerial reconnaissance, but the myth prevailed until the construction of the Alcan

Military (Alaska) Highway in 1942. The hot springs (known to the Americans as Theresa Hot Springs) received intensive use for the first time as it was used as a bathing facility for a nearby highway construction camp. The

Alaska Highway was opened to the public in 1947.

Climate

The regional climate has been characterized by Chapman (1952) and

Krajina (1965) as Dfc under the Koppen system. This is typically a cold snowy forest climate with no distinct dry season and a cool short summer, i.e. a microthermal boreal cold continental humid climate.

The Koppen symbols are (Trewartha 1968):

D - cold snow-forest climates, average temperature of the coldest month is below 26.6 F (-3 C) with the warmest month averaging over 50 F (10 C) f - no distinct dry season and the driest month has more than 1.2 in. (3 cm) of precipitation c - cool short summer with less than four months averaging over 50 F (10 C)

Additional details are provided later in the CLIMATE AND MICROCLIMATES section.

Soils

There is little published information on regional soils. The British Columbia Lands Service (undated, a) indicated that generally weakly developed Brunisolic soils are most common in the Liard River system.

Leahey (1943) was concerned with agricultural possibilities and described the park region as follows (p. 3):

"...The soil is mostly a deep sandy loam with the surface soil being mostly dark grey, but in some places it is a yellowish to reddish brown in color. Drainage for the most pact is good to excessive. This land is level to undulating and at present is heavily wooded with spruce and poplar. The area while small may be of local significance as it lies adjacent to the hot springs and the so-called "Tropical Valley"."

Edaphic results of this study are provided later in the SOILS section.

Vegetation

The region is considered to be in the Boreal White and Black Spruce

Biogeoclimatic Zone (Krajina 1969, 1973). Zonal plant indicator combinations include Picea glauca, Picea mariana, Pinus contorta, Pinus banksiana, Larix laricina, Ledum groenlandicum, and Betula papyrifera (Krajina 1965, 1969).

Pinus banksiana does not occur within the study area.

There are several distinct plant communities within the park. A rich mixed forest consists of aspen (Populus tremuloides), birch (Betula papyrifera), white spruce (Picea glauca), red osier dogwood (Cornus stolonifera), high-bush cranberry (Viburnum edule), and sarsaparilla

(Aralia nudicaulis), Picea glauca and Hylocomium splendens largely comprise the white spruce - feathermoss forest type. A black spruce (Picea mariana) and moss (Pleurozium schreberi, Hylocomium splendens, and ~tiliumcrista-castrensis) type of forest vegetation is found on poorly drained organic soils. Wetland hummocks are occupied by a larch - yellow rose plant community characterized by Larix laricina 18 and Potentilla fruticosa with Labrador tea (Ledum groenlandicum), bog birch (Betula glandulosa), and bog cranberry (Oxycoccus microcarpus) also present. Conspicuous stands of ostrich fern (Matteuccia struthiopteris) are found in several localized areas, and small patches of dogbane

(Apocynum androsaemifolium) are found on steep calcareous slopes.

Tufaceous islands are occupied by northern bog violet (Viola nephrophylla), great sundew (Drosera anglica), rush aster (Aster junciformis), bog muhly (Muhlenbergia glomerata), and the mosses Campylium stellatum and

Drepanocladus vernicosus (?). Some aquatic communities have yellow monkey-flower (Mimulus guttatus) and Philadelphia fleabane (Erigeron philadelphicus) emerging from the water. Submerged vascular species include sago pondweed (Potamogeton pectinatus) and bladderwort (Utricularia vulgaris) but the alga Chara vulgaris (stonewort) and the mosses Riccia fluitans and Cratoneuron filicinum are also abundant. The vegetation patterns of the study area are presented in Figure 2.

Vegetation results of this study are provided later in some detail in the VEGETATION section.

The park has been previously studied from a floristic and phyto- geographic viewpoint (Porsild and Crum 1961). The unique temperate nature of much of the study area's flora was emphasized in this paper.

These authors noted 82 species of vascular plants which were growing either adjacent to the thermal springs or in areas affected by the springs.

Of these species, 43 were considered as distinctly temperate speciks with several having the study area as their northernmost known limit with the remaining species found farther north only in thermally favoured areas such as hot springs. An additional 39 species were considered temperate or thermophile species that are always restricted to "especially favoured and protected habitats" at latitudes north of the park.

The 82 species that Porsild and Crum (1961) selected from the flora of Liard River Hot Springs Park were considered temperate or thermophile species on the basis of their known collection sites (circa 1961) in northern North America. However, considerable collecting has been done throughout the north since their publication and it was considered that northern post-1961 collections may necessitate a revision in the status of some of these species.

A report (Reid 1976) investigated the northern post-1961 collection sites known for these 82 species. This investigation involved a literature search of these more recent collections from northern British Columbia, northern Alberta, Alaska, Yukon Territory, and western portions of the

North West Territories. Collection sites were then plotted on distribution maps (one map for each species), and habitat comments were noted. These more recent collections result in the removal of 49 species from the

Porsild and Crum list. No conclusions could be reached on the current status of 19 other species, but 14 species must still be considered as existing only inthermally favored areas in the north. These species are all found in the study area and include (Reid 1976) :

Carex laeviculmis Muhlenbergia racemosa

Carex preslii Parietaria pennsylvanica

Carex sitchensis Parnassia parviflora

Cynoglossum boreale Phalaris arundinacea

Glyceria striata Sanicula marilandica

Lactuca biennis Urtica lyallii

Lathyrus ochroleucus Viola rugulosa CLIMATE AND MICROCLIMATES

Objectives

The major objectives of this portion of the study were to:

1. determine the climate of the study area;

2. determine microclimatic patterns between thermally influenced and non-

thermally influenced regions; and

3. determine if there is a difference between the microclimates of these

two thermal regimes.

Methods

Stations and Instrumentation

Seven stations were established on the basis of the above objectives.

One "control" climatic station and six microclimatic stations (three

thermal and three non-thermal) were installed in the study area (Figure 2).

Station characteristics are summarized in Table 11. The bases of the

Stevenson-type screens were placed 20 cm above the ground surface for microclimatic stations, and 1.2 m for the control station. These heights were selected in order to facilitate vegetation (microclimatic stations) and meteorological (control station) comparisons as noted above.

All stations were equipped with:

1. a Fuess or Casella hygrothermograph, a separate maximum thermometer,

and a separate mimimum thermometer all mounted in a Stevenson-type

screen.

2. a Tru-check rain gauge. Table 11. Climatic and microclimatic stations of the study area

Station Station Type Code Remarks

Control LC P greatest similarity to a standard meteorological station; established near the "Large Cold Pond"

Thermal EP established at the Epsilon Hot Pools complex AOS established at the stream draining Alpha Hot Po01 WPS established immediately adjacent to Alpha Warm Ponded Swamp

Non- thermal MA established in the moose-churned fenland adjacent to the Epsilon Hot Pools complex in a Larix - Potentilla community type C PF established in the Picea type forest near the "Large Cold Pond" OF established in the Ostrich Fern Glade in a Matteuccia - Actaea comunity type

3. YSI No. 44004 thermistors at a soil depth of 5 cm and 10 cm.

Casella hygrothermographs were installed throughout the measurement period at stations CPF, WPS, and AOS; a Fuess instrument was at station LCP.

Station OF initially held a Casella hygrothermograph but repeated clockwork malfunction necessitated replacement with a Fuess instrument on March 25,

1975. Stations EP and MA originally held a Fuess and Casella respectively but instrument corrosion at the thermal pool installation became visible by November 13, 1974 and the two instruments were switched at this time.

Stations EP and MA then contained a Casella and Fuess hygrothermograph respectively for the duration of the measurements. Air Temperature

Measurements

Air temperature measurements began June 18, 1974 and ended July 15,

1975. Both terminal months have incomplete monthly data so analysis concentrated on the twelve month period from July 1, 1974 to June 30, 1975.

Air temperature measurements were of two types. A continuous 24-hour thermograph trace was attempted but instrument malfunctions resulted in periodic gaps in the data wllich became more pronounced in the colder months

(Appendix A). The second set of measurements was obtained from the maximum and minimum thermometers during the weekly servicing of the stations.

These readings were also used during the field work to maintain thermograph calibrations. Thermometers were read at approximately the same time each week.

Thermograph charts provided daily maximum and daily minimum temperatures. In accordance with standard meteorological procedures, the maximum temperature is taken for the 24 hour period 8 AM to 8 AM whereas

the minimum temperature is taken for the 24 hour period 4 PM to 4 PM

(M. Rose 1976, personal communication). An alternative time interval is

the 24 hour period commencing at midnight (R. C. Brooke 1976, personal

communication), but the need for comparisons between my data and published data (using standard meteorological procedures) suggested the adoption of meteorological time interval periods for daily maximum and daily minimum

thermograph data.

Monthly Temperatures

Data analysis for determination of the park temperature pattern used

the information from the control station LCP. Daily thermograph data were 2 3

summarized on a monthly basis and calculations were made Eor the monthly mean maximum temperature, monthly mean minimum temperature, and monthly mean temperature. The monthly mean temperature was calculated as the average of the mean maximum temperature and the mean minimum temperature.

Microclima tes

Microclimatic patterns were assessed using data from the three

thermal (EP, AOS, and UPS) and the three non-thermal (MA, CPF, and WPS) microclimatic stations. Control station LCP was excluded from this analysis

since it was mounted much higher than all other stations and would be expected to be in a somewhat different temperature regime (Geiger 1965).

Daily thermographs could not be used due to gaps in their data, so the weekly maximum and minimum temperatures were used for analysis purposes.

It was hypothesized that temperature differences occurred between some microclimatic stations. Some doubt existed as to whether or not temperature data conformed to the normal distribution so the most conservative approach was to select a non-parametric statistical tool to test the hypothesis,

The statistical test chosen was the Wilcoxon Rank-Sum Test (Wilcoxon and

Wilcox 1964). This statistic provides information similar to that obtained

from the parametric F-test plus a Duncan Multiple Range Test. The Wilcoxon

test's advantage is that it can provide a distribution-free test of

significance between any two microclimatic stations without involving

recalculation or 2 priori assumptions about the thermal regime of the

stations involved.

Mean Annua 1 Temperatures

Daily thermograph and weekly maximum and minimum temperatures were

individually used to calculate mean annual temperatures. These values were

calculated by taking the average of the mean maximum and mean minimum temperatures from daily (thermograph) or weekly (maximum-minimum) data.

Frost- free Period

Hygrothermograph data supplied information on the number of frost-free days. This could be observed directly for only three (non-thermal) stations as the remaining four stations had their last spring frost prior to obtain- ing data. The observation period for all stations did include that period of the year at which freezing temperatures could occur (i.e. temperatures equal to or less than 0.0 C or 32.0 F). The observed number of days with frost was then used as a basis to calculate an estimate of the number of frost-free days.

Soil Temperature

Instantaneous soil temperatures at 5 and 10 cm depths were measured on a weekly basis by connecting a YSI model 43 TE telethermometer to permanently installed YSI No. 44004 thermistors. Readings were taken at approximately the same time each week.

Thermistor calibrations were checked prior to installation and again at the end of the measurement period by immersion in an ice-water bath.

Thermistor calibration changed for some instruments over the 1974-1975 measurement period. Changes of two or less fahrenheit degrees were noted for nine thermistors, and the LCP - 10 cm and WPS - 5 cm installations had changed 2.2 and 3.5 degrees respectively. Drastic changes (approximately ten fahrenheit degrees) were observed for the AOS - 5 cm and EP - 10 cm thermistors although the other thermistor installed at these thermal locations showed less than a two degree chaage. It was assumed that calibration changes were a linear function of time and where necessary the data were corrected accordingly. 2 5 Precipitation

Precipitation was measured during the snow-free period by a Tru-check rain gauge mounted at each station. Each installation was made in an area free of obstructions with the gaugeorificeapproximately 60 cm above, and parallel to, the ground surface. Oil was added to restrict evaporation between the weekly readings.

During the months of snowfall, weekly snow depth was measured on the top of the Stevenson-type screen. This snow was removed after the weekly measurement was recorded. Snowfall was converted to the rainfall equivalent by using the commonly accepted conversion factor of 1 unit of fresh snow =

0.1 unit of water.

A statistical evaluation of the data was performed by using the

Wilcoxon Rank-Sum Test (Wilcoxon and Wilcox 1964). This test was selected for reasons similar to those noted in a previous section on Air Temperature.

Relative Humidity

Daily relative humidity data were obtained from hygrographs. Instru- ment calibrations were monitored (in season) by using a sling psychrometer during the weekly servicing of the stations. Data reduction involved the conversion of daily maximum and minimum data into mean monthly maximum and minimum values for each station.

Results and Discussion

Air Temperature Monthly Temperatures

During the measurement period only two months (July and August) were July Aug. Sept. Oct. Nov. Dec. Jan. Feb. March April May June (31)a (31) (30) (19) (30) (31) (17) (22) (31) (30) (30) (30) Figure 7. Monthly mean, mean maximum and minimum, and extreme maximum and minimum air temperatures for control station LCP a: Number of days of data. completely free of freezing temperatures, but all twelve months recorded at least one measurement well above the freezing point (Figure 7). Mean monthly temperatures declined steadily from August to January and then increased until the last complete measurement month in June.

However, northern British Columbia is subject to great climatic variations which may persist for several months (Strachan 1959). Conse- quently, these observations may not have been typical, so data were compared with long term averages for other stations. Long term averages were available for Watson Lake Airport, Yukon Territory (elevation 689 m or 2,262 ft.), and Fort Nelson Airport, British Columbia (382 m or 1,253 it.). Liard

River Hot Springs Park (approximately 457 m or 1,500 ft.) is situated approximately half-way between these two centers.

Data from Fort Nelson and Watson Lake (Table 111) suggests that the study area long term average temperature pattern may then differ from

Figure 7 by a steeper monthly mean temperature slope from July to January and a somewhat decreased slope from February through June.

Temperature and precipitation data (Tables 111 and VII) support the

Dfc (cold snow-forest climate with a cool short sumer and no distinct dry season) climatic classification (see REGIONAL AND STUDY AREA DESCRIPTIONS - Climate for details).

Microclimates

Statistically significant (p6 0.05) monthly and yearly comparisons between thermally influenced and non-thermally influenced regions are given in Table IV (data summarized in Appendix B). Table 1x1. Comparison of observed temperatures (F) with long-term averagesb

Mean Temperatures Dif. Mean Temperatures Dif. Location Max. Min. Daily from Max. Min. Daily from Avg .a Avg .a

July 1974 January 1975 Fort Nelson Airport 70.3 47.6 59.0 -3.1 0.3 -15.3 -7.5 2.3 Liard Hot Springs Watson Lake Airport

August February Fort Nelson Airport 67.2 46.9 57.0 -1.6 13.4 -12.7- 0.4 -0.9 Liard Hot Springs 67.2 46.1 56.6 Watson Lake Airport 66.5 43.5 55.0 -0.4

September March Fort Nelson Airport 57.9 34.8 46.4 -1.5 26.4 3.0 14.7 -0.8 Liard Hot Springs Watson Lake Airport

October Apr i 1 Fort Nelson Airport 40.2 24.7 32.4 -1.6 49.7 24.0 36.8 2.8 Liard Hot Springs Watson Lake ~ir~ort

November May Fort Nelson Airport 22.3 7.0 14.6 4.8 62.5 36.9 49.7 0.3 Liard Hot Springs Watson Lake Airport

December June Fort Nelson Airport 6.3 -8.1 -0.9 4.4 70.1 46.9 58.5 0.4 Liard Hot Springs 13.1 -4.8 4.2 Watson Lake Airport 13.1 -6.5 3.3 11.9

. Difference from Average . Compiled from field data (station LCP for Liard Hot Springs) and Canada - Department of the Environment (Monthly Record). . Missing 12 days of data. . Missing 14 days of data. e. Missing 6 days of data. f. Missing 8 days of data. Table IV. statistically significant (pd 0.05) maximum and minimum air temperature differences between pairs of stationsa

Comparison Maximum Minimum Month Thermal: EP WPS AOS EP WPS AOS AOS EP EP EP WPS AOS Non-thermal: CPFCPFCPFOF OF MA OF MA OF CPFMA CPF

-19 74 July August Sept.-Oct. b Novemb er December -19 75 January February March April May June

Yearly Values -k* * ~r 3~ ~r***.~rk**.k dr

a: Based on weekly readings of separate meteorological maximum and minimum thermometers. Blank areas in the table indicate a non- significant comparison. Station comparisons not appearing on the table are non-significant. b: Combined months since my absence from the study area resulted in insufficient data (n = 6) to evaluate separately. *: Significant difference (p - 0.05) *: Significant difference (p = 0.01) m: Missing data

Stations significantly differing in minimum air temperatures were found in all monthly intervals whereas no significant differences in maximum air temperatures could be detected for May and June (1975) and July

(1974). This is expected as the thermal influences of the hot pools would be masked by the generally warm maximum summer air temperatures. Under all other conditions (e.g. maximum air temperatures from fall through early spring, and minimum air temperatures throughout the year) at least some statistically significant differences in air temperatures were observed between thermally influenced and non-thermally influenced microclimatic stations.

Mean Annual Temperatures

Yearly mean temperature values for the period July 1, 1974 to June 30,

1975 are presented in Appendix A but the incompleteness of the daily thermograph data precludes much reliance being placed on the calculated values.

The relatively complete weekly maximum and weekly minimum data base during this period permits an estimation of station mean yearly temperatures (Table V) .

Table V. Estimates of annual mean air temperatures

Thermally Non- thermally Station Type Control 1nf luenckd Influenced Weeks of Data 50 49 50 50 50 50 48.5 Station LCP EP AOS WPS MA CPF OF

Annual Mean (F) 3 1 37 36 34 32 3 2 32 Annual Mean (C) -0.6 2.8 2.2 1.1 0.0 0.0 0.0

From these data, estimated annual mean temperatures average 2.0 2 0.7 C (35.7 2 1.2 F) for the thermally influenced stations but only 0.0 2 0.0 C

(32.0 --k 0.0 F) for the non-thermally influenced stations. These data suggest annual mean temperature differences between the two thermal regimes, perhaps in the order of two celsius degrees or three or four fahrenheit degrees.

Frost-f ree Period

Data for the frost-free period is given in Figure 8 and Appendix C. Figure 8. A comparison of daily temperature range patterns

A pronounced difference existed between thermally and non-thermally influenced stations. The number of estimated frost-free days ranged from

82 to 85 for the former group and from 108 to 120 for the latter group.

Mean estimated frost-free period for the three thermally influenced microclimatic stations were 116 -+ 5.7 days but only 84 1.4 days for the non-thermally influenced microclimatic stations. This four to five week difference in the estimated frost-free period between the two thermal regimes may ultimately be expected to result in earlier and longer growing seasons in thermally favoured areas.

Soil Temperature

Soil temperature data is summarized in Appendix D. Maximum mean monthly soil temperatures occurred during May for station EP. Weekly field data for this station showed sharply increased soil temperatures began in

April and lasted through much of May before they returned to somewhat lower levels. This pattern may be due to increased temperatures in the water from the hot springs complex during this interval but no data is available that would permit a test of this hypothesis. All other stations appeared to have maximum soil temperatures during July (of 1975) as may be expected from an increased duration and angle of solar radiation.

Four of the five stations with incomplete January data had minimum calculated mean monthly soil temperatures during that month. These stations included the three thermal microclimatic stations (EP, AOS, and WPS) as well as station MA. The remaining station with incomplete (January) data was station OF which showed lowest soil temperatures during February. Station

CPF also showed minimum soil temperatures during February. Little variation 33 was shown in the low soil temperatures from December through March for non- thermal station LCP.

The phenomenon of lowest soil temperatures occurring earlier for

thermally influenced stations than for non-thermally influenced stations may be a sampling artifact due to incomplete January data. An alternative

explanation may be that very heavy snowfall during this month (averaging approximately 1.8 times as much snowfall as had fallen in the two previous months combined - Appendix E) would effectively insulate the ground from the cold air temperatures. Thermal influences from the springs (at thermally

influenced stations) may then have had a chance to gradually modify soil

temperatures to the point where they began increasing in February. Station Ill1 MA may have been in a somewhat intermediate thermal regime for soil tempera- j tures since it was located in somewhat swampy ground and was only about

30 m (100 ft.) from thermal station EP. It could then have lowest soil

temperatures during the same month as the three thermally influenced

stations.

With the exceptions of thermally influenced stations EP and AOS,

there was little variation in weekly soil temperatures from late November

until mid April. During this period, only stations EP and AOS had weekly

spot readings consistently above freezing (0 C or 32 F). This is in

sharp contrast with all other installations which recorded from 21 to 27 weeks of freezing temperatures (Figure 9). Station MA has incomplete

10 cm data due to a thermistor failure. Continuous weekly freezing soil

temperatures usually started in November and ended in May. WPS AOS E P Station

Figure 9. Number of continuous weeks of freezing (( 0 C or ~32F) soil temperatures

Stations AOS and EP are Located at major thermal influences. Open histogram: 5 cm depth. Shaded histogram: 10 cm depth.

Annual mean soil temperatures are presented in Table VI. The station MA appears to have some thermal influence as previously noted.

Stations EP and AOS had a higher thermal influence than station WPS due to the EP and AOS thermistors being installed in saturated soils with much higher water temperatures than recorded for the Warm Ponded Swamp (Table I).

The WPS thermistors were also installed comparatively well above the ground water depth. Table VI. Annual mean soil temperatures

Major Moderate Thermal Thermal Non- thermal Station Type Influences Influences Influences Weeks of Data 46 4 7 47 47 48 48 47 Station EP AOS WPS MA LC~OF CPF

Annual Mean (F): 5 cm 60 58 4 2 43 3 9 38 37 : 10 cm 70 56 39 m 37 38 36 AnnualMean (C): 5 cm 15.6 14.4 5.6 6.1 3.9 3.3 2.8 :10cm 21.1 13.3 3.9 m 2.8 3.3 2.2

a: Not a "control station" since thermistor installation is similar to other stations.

From these data, the average annual mean temperature for stations + under a major thermal influence were 15.0 - 0.6 C and 17.2 + 3.9 C (59.0 ' 1.0 F and 63.0 f 7.0 F) at the 5 and 10 cm depths respectively. Non- thermally influenced stations averaged only 3.3 2 0.4 C and 2.8 0.4 C

(38.0 f 0.8 F and 37.0 f 0.8 F) at 5 and 10 cm depths. Stations under a moderate thermal in•’luence averaged 5.8 f 0.2 C (42.5 * 0.5 F) at 5 cm beneath the soil surface.

These data illustrate the differences in soil temperature regimes

b that exist in the study area. These microclimatic differences may ultimately be partially expressed in the form of vegetation cover, composition, vigor, or phenology.

Precipitation

Precipitation data are given in Appendix E. Annual totals are cons'idered to be a reasonable approximation of the probable long term average for the study area (Table VII) despite unusually heavy rains in July, 1974 and June, 1975 which resulted in closure of the Alaska Highway on both occasions. Table VII. Comparison of observed precipitation (in.) with long-term averagesa

Fort Liard Watson Month Nelson Hot Springs - Lake Airport Stn. LCP Airport

1974 July

August

Sept .-Oct.

November

December

1975 January

February

March

April

May

June

Total

a: compiled from field data and Canada - Department of the Environment (Monthly Record). b: numbers in brackets are differences from long-term averages.

During the peak of the growing season, at least 89 unn (3.5 in.) of

rainfall was recorded in each of June (1975), July (1974), and August (1974).

Comparison with long term averages suggest that the thermal springs area may

receive approximately 5 cm (2 in.) per month during this period.

Snowfall began in mid November and lasted for approximately five months 3 7

in. mm in. 26

.24

AOS WPS LC P OF C PI? NON-FOREST FOREST

Figure 10. Total precipitation for the twelve month period July 1974 to June 19 75. Open histogram: total precipitation Shaded histogram: total snowfall until it melted in mid-April. Snow depth was moderate (approximately 0.5 m or 1.6 ft.). This agrees well with measured values in other northern communities.

Twelve month precipitation totals for the stations ranged from 485 to

622 mm (19.1 to 24.5 in - Figure 10). Largest total precipitation (over 610 mm or 24 in.) was recorded at thermally influenced stations EP and AOS.

Station MA (located approximately 30 m or 100 ft. from the thermally influenced station EP) recorded the third highest total precipitation

(approximately 584 mm or 23 in.). By contrast, the non-thermally influenced station CPF (located in a black spruce forest community type) recorded the minimum precipitation (only 483 mm or 19

The observed between-station variation of 137 mm (5.4 in.) of total precipitation suggested that there may be a statistically significant variation between stations. A statistical evaluation of monthly total precipitation and monthly snowfall is given in Table VIII. Significant (p4 0.05) differences for both precipitation parameters existed for two sets of comparisons (EP - CPF, and AOS - CPF). A third comparison (MA - CPF) resulted in a significant difference for only monthly total precipitation.

Some loss of sensitivity would be expected for snowfall measurement as contrasted with total precipitation since the sample size is smaller

(n = 5 and n = 11 respectively).

Table VIII . Statistically significant (p& 0.05) differences in precipitation (nun) between pairs of stationsa

Precipitation Thermal: EP AOS MA Non- thermal : C PF C PF CPF

Total

Snowfall

a: Based on monthly totals for rainfall or fresh snow increment (data from Appendix E). Station comparisons not given are non-significant, *: Significant difference (p = 0.05) ** Significant difference (p = 0.01) ns: Not significant

Stand density has an influence on precipitation reaching the ground surface (Geiger 1965, Brooke et al. 1970). The influence of stand density can be quite marked as is illustrated by station OF (Figure 10). Dense, tall ostrich fern growth during the summer (reaching higher than the top 39 of the rain gauge) appears to result in a depressed ground-level precipitation value. In the winter when the fern growth is leveled and under snow the observed precipitation as snowfall is higher than would be expected on the basis of the recorded total precipitation (Figure 10). In coniferous forests (as at station CPF which was in a black spruce forest vegetation type), crown interception losses from wetting and evaporation are to be expected (Geiger 1965). These factors suggest that precipitation differences are due to stand density effects.

Howeve.r, the CPF rain gauge was placed in an area as free of obstructions as could be found at the site. This placement would make it more susceptible to drip zone influences (which receive perhaps 10% to 20% more precipitation than open land - Geiger 1965) rather than interception losses. Stand density effects would appear to be minimized in this situation.

Relative Humidity

Relative humidity data are summarized in Appendix F. These data are quite incomplete due to instrument malfunctions. Station LCP has the most complete data base (345 days) but this station can not be used to compare microclimatic patterns since this instrument was mounted at a greater height than other instruments and would be in a different relative humidity

regime (Geiger 1965).

Comparisons between statioy are of dubious value unless complete data

from both stations are available. The daily hygrograph data are considered

to be "probably accurate to within 5 per cent" (M. Rose 1976, personal communication). The comparative inaccuracy of these readings plus the

incompleteness of the data base effectively precludes a detailed analysis Table LX. Station relative humidity data (X) sunsrarized by microclimatic group

19 74 1975 July Aug. Sept. Oct. Nov . Dec . Jan. Feb. March April May June Max.Min. Max.Hin. Max.Hin. Max.Min. Max.Min. Max.Min. Max.Min. Max.Min. Max.Min. Max.Min. Max.Min. Max.Fiin.

Control stationa 86 46 90 46 87 43 85 51 85 69 75 61 67 55 65 11 72 38 73 25 74 26 76 28 (30)~ (31) (30) (27) (30) (31) (20) (24) (31) (30) (311 (30)

Thennally Influenced stat ionsb 88 58 90 65 91 61 89 69 91 82 81 76 84 72 78 64 83 61 74 43 70 36 73 41

Eion-thermally Influenced stationrc 92 59 92 68 91 60 90 70 87 77 81 70 69 58 80 56 82 55 80 37 72 28 81 41

a: station SCP b: rtationr EP. AOS, and WPS c: stations NA, CPF, and OF d: valuer in bracket8 represent the number of days used in the calculation. 41 and interpretation. The available data suggests that winter and spring minimum relative humidities are higher in thermally influenced areas than in non-thermally influenced areas (Table IX). This is consistent with the proximity of thermally influenced areas to hot pool vapors. Frequent showers, evaporation, and transpiration factors would tend to minimize relative humidity differences between the two microclimatic regimes during the warmer months. Objectives

Soils were studied to determine the edaphic environment of the vegetation. Specific major objectives of this portion of the study were to:

1. classify and describe the s~ilsof the study area,

2. determine the edaphic chemical environment of particular community

types and species in the study area, and

3. ultimately, to consider how the l?daphic environmental variables may

influence the distribution and abundance of plant community types

and individual species.

A brief survey (Leahey 1943) explored the region for soils with agricultural potential. The closest published soil descriptions are

from the Fort Nelson area, British Columbia (Valentine 1971), and the

Liard River Valley, Northwest Territories (Day 1966).

The following section presents this descriptive portion of the study. The edaphic-vegetation relationship is presented in the VEGETATION - ENVIRONMENT RELATIONSHIPS section.

Methods

Soil data were obtained from 29 vegetation plots located within

representative plant communities of the area. The soils were developed

on a range of parent materials and topographic positions. Profile

descriptions were made for each soil pit and soil samples were taken for chemical analysis.

Pits were dug either to the unweathered parent material, to impermeable layers, to the water table, or to depths exceeding the maximum root penetration. Pits were not dug nor were profile descriptions made for areas containing aquatic plant communities. In these cases, only the top 2 cm of sediments were collected for chemical analysis.

Profile descriptions conformed to recommendations of The System of

Soil Classification for Canada (Canada Soil Survey Committee 1974) hereafter abbreviated as SSCC (1974). Field profile description emphasized the following characteristics: depth, thickness, color, boundary, stoniness, and root distribution of each horizon. Notes were also made on parent material, drainage, water regime, ground water depth, estimated profile stoniness, and impermeable layers.

Soil samples were collected from most recognizable horizons for subsequent laboratory chemical analysis. Mixing of different horizon samples was minimized by not collecting samples from very thin (<5 cm or 2 in.) horizons. Collected samples were later air dried and sieved (2mm).

Approximately 125 samples of the 42 mm soil fraction were then chemically analyzed by the Department of Soil Science at the University of British

Columbia.

Soils were classified and described on the basis of criteria given in

SSCC (1974) or Kubiena (1953).

Description and Classification of the Soils

Results

Four of the eight Canadian Orders were present in the study area: Brunisols, Regosols, Gleysols, and Organic soils. Subaquatic soil samples were also taken but submerged soils are not recognized by SSCC (1974). A

European classification system (Kubiena 1953) was used for this category and one additional group, approximately corresponding to an Order, was recognized: the Sub-aqueous soils not forming peat.

A synopsis of soil categories described and provisionally classified is given in Table X. Results of the chemical analysis are presented in

Appendix G.

Table X. Synopsis of soil categories classified and described

Order Great Group Subgroup

Brunisolic Melanic Brunisol Orthic Melanic Brunisol Eutric Brunisol Orthic Eutric Brunisol Degraded Eutric Brunisol Dysrric Brunisol Degraded Dystric Brunisol

Regosolic Regosol Orthic Regosol

Gleysolic Gleysol Rego Gleysol Carbonated Gleysol

Organic Mesisol Hydric Mesisol Humisol Hydric Humisol sub-aqueousa protopedona Lake Chalk protopedona

a: Based on Kubiena 1953.

The following section describes the soils found in the study area.

Tax~nomiccriteria applicable to the study area are summarized from SSCC

1974 or Kubiena 1953, diagnostic horizons are underlined, and nondiagnostic horizons that may be present are in parentheses. Brunisolic Order

"This order consists of soils with sola indicative of good to imperfect drainage or of good to moderate oxidizing conditions which have developed under forest, mixed forest and grass, grass and fern, or heath and tundra vegetation associations representative of forest, alpine, or tundra communities." (SSCC 1974 p. 111)

Profile type: (L-H) , (Ah), (Aej or Ae) , $J Three Great Groups were identified.

Melanic Brunisol Great Group

"...generally lack F and H horizons. They have mineral- organic (Ah) surface horizons thicker than 2 inches (5 cm), ... Bm horizons in which the base saturation (NaC1) is loo%.. ." (SSCC 1974 p. 111-112)

Profile type: (L) , 9(> 5 cm) , &I (100% base saturation) One Subgroup was recognized - an Orthic Melanic Brunisol (Figure 11) with diagnostic profile type (L), Ah, &, (Ck), (Cca).

One profile occurred in an Apocynum - Cynoglossum cormnunity type (plot 14). The L horizon was thin (1 cm) and consisted of leaves, the Ahk horizon

was thick (some 25 cm) and contained consolidated tufa chunks as did the

Bmk horizon. This was underlain by a Ck horizon with no visible tufa.

All mineral horizons had 100% base saturation with the pH in these horizons

ranging from 7.0 to 8.1.

This soil pit was dug in the non-forested area above Psi Warm Pool.

This area may have once been a garden site. Eutric Bmriisol Great group

If.. .organic surface horizons (L-N) over Bm harisons in which the base saturation - (NaCl) ia POm and the pH ,(CaCA2) is usually 5,s or higher.. ,my have thin Ah horieonr [42 &aches (5 cm)] under forat communities or moderlitely thick Ab horizons uader alpine vegetation at~dclimate. They my have weakly expressed (Aej) or &$rongly expreased (Ae) eluvial horizons. Tbsy may 'also have horizons of illuviation that fail to meet thg requi'rmmts of the Bt as defined, The parent matetial is usually calcareous." (SsCC 1974 p. 113)

Profile Type : s,(Ah), (Ae j or Ae) , @J (100% ba~esaturation with pH'usually b 5.5) 45

Two Subgroups were recognized - the ~rthicEutrie B5yisi$of etdd-the

Degraded Eutrfc Brunisol, The profile sequence for an Orthic Eutric Bruaisal is s,(Ak) ,

i&a, (Ckf, (Cca) and they etsualEy have calcareous parent raert~rials. %is subgroup was Eowd f.n the Matteuccia - hctaaa (Plots 12, 16, and 39 - Figure 12) and in the (15 and 44) catarnunity type@. These soii ~ikshad frodl 3 to 16 cta of ur$aair materia$ and Ed haultauas with 10a baee ihturation and pH values; from 7,2 to 8.2, Ahk and calcareous C horizons were frequently present. "ph .'-

The profile sequence E, or Ae, l&or a,(Ck) characterizes a Degraded Eutric Brunisol. The main distinguishing characteristic of this soil is the presence of an eluvial horizon (Aej or Ae) over the Bm horizon. This Subgroup was found in the Populus - Viburnum vegetation type (Plots 05 and 43 - Figure 13). These soil pits have thick organic horizons over the mineral soil eluvial horizon. Bmk horizons had a base saturation of 100% with pH values generally 7.3 or higher. One Bmk horizon (located beneath a 6 cm thick, pH 5.2 Ae horizon) had a pH of

Qg~d$M, A DegraW Eutrie $runis01 in a PaJ@fye - Viburnum vegetation type (Plot 05). Dystric Brunisol Great Group It ...organic surface horizons (L-H) over Bm horizons in which the base saturation (NaC1) is usually 65 to 100% and the pH (CaC12) usually 5.5 or lower.. .may have thin Ah Horizons [<2 inches (5 cm)] under forest cornunities ...may also have weakly expressed (Aej) or strongly expressed (Ae) eluvial horizons ..." (SSCC 1974 p. 115)

Profile Type: E, (Ah 5 cm), (Ae or Aej), -Bm (65-100% BS, pH4 5.5) One subgroup was recognized - the Degraded Dystric Brunisol Subgroup which has a profile type of z,Aej or &, & or Bmcc, (C). These soils have an eluvial horizon and a Bm horizon lacking sufficient illuvial material for a podzolic B horizon. The one Brunisolic soil pit with low base saturation and pH was found in a Populus - Viburnum cornunity type

(plot 09) ,

Regosolic Order

"These are well- and imperfectly drained mineral soils with good to moderate oxidizing conditions, having horizon development too weak to meet the requirements of soils in any other order." (SSCC 1974 p. 123)

Regosol Great Group

"Only one great group has been established, therefore its definition is the same as that of the order." (SSCC 1974 p. 123)

The one Subgroup recognized was the Orthic Regosol Subgroup (Figure 14) characterized by profile type (L-H), (Ah), -Ck or C. Two soil pits in the Apocynum - Cynoglossum community type (13 and 55) were classified as Orthic Regosols. Both pits had a thin (1 or 2 cm) organic horizon over an Ahk

/ horizon with tufaceous material occurring below this. Base saturation was

100% for Ahk and C horizons with measured pH values from 7.3 to 7.9. Gleysolic Order

"These soils are saturated with water and are under reducing conditions continuously or durins some period of the year ...Matrix colors of low chroma within 20 inches (50 cm) of the mineral surface. I' (SSCC 1974 p. 131)

One Great Group was oheerved in the study area. , .

. , Gleyqol. Graa t Orouq

", ..a@ Ah Ezarizon or an ,& horizon up to 3 inchee (8 em) thick.. .my have orgcrnic twrface layers, up to 16 iachcs (40 a) of mixed peat,..ar up ra 24 inches (60 cm) af fibric nos8 peat.. .I' (S6CG 1974 p, 134 cind 133)

The Rego Ql~ysoland Carbonated Qleyed Subgroups occur bn the pa& Figure 15. i4 Rego Gleysol (peaty phase) in a, PScaa marigna -.,Pleuroeium vegetation type (Plot 48).

(Ah), 3 or a or Ccak and they therefore lack a B horizon. Two so31 pits belonged to this category; one occurred in an ~leocharis- Trialochin community ty,pe (plot 07) and the other in a Picea mariana - Pleurozium

community type (48). The organic horizon in the Eleocharis -Triglochin

community is very thin (2 cm) but the comparatively thick (25 cm) organic

horizons in the Picea mariana - Pleuroziwm community (Figure 15) would

permit classification as the peaty phase of this Subgroup. It is interesting

to note that the more common soil pit classified for each of these community

types is that of the Organic Order. This may suggest that these Gleysolic

J. Qrder pits occur in areas that are in a comparatively early stage of soil n V aevelopment , k=I The second category recognized was the Carbonated Gleysol Subgroup which is defined as ". . .Gleysols with an effervescent (carbonate) surface horizon" (SSCC 1974 p. 135). All soil pits in this category consisted entirely of one or more tufaceous horizons. This Subgroup comprised soil pits in the following community types: Erigeron - Mimulus (53 and 54),

Aster - Muhlenbergia (31-33, 40, 41), and Viola - Spiranthes (35-38).

Organic Order

"These are soils that have developed dominantly from organic deposits. The majority of them are saturated for most of the year ...but some of them are not usually saturated for more than a few days. They contain 30% or more of organic matter and must meet the following specifications: ...the organic materials must extend to a depth of at least 16 inches (40 cm). ...Mdneral layers thinner than 16 inches (40 cm), beginning within a depth of 16 inches from the surface, may occur within the organic soil. A mineral layer, or layers taken cumulatively, thinner than 16 inches (40 cm) may occur within the upper 32 inches (80 cm)," (SSCC 1974 p. 154)

Profile type (basic): -Of and/or 9 and/or -Oh (,40 cm)

Two Great Groups were recognized within the study area.

Mesisol Great Group

"These organic soils have a dominantly mesic middle tier, or middle and surface tiers if a terric1 , lithic2, hydric3, or cryic4 contact occurs in the middle tier." (SSCC 1974 p. 156) 1: mineral horizon 3: water 2: bedrock 4: permanently frozen

Profile type: dominated by @

Only one subgroup was recognized - the Hydric Mesisol Subgroup

(Figure 16) which has a hydric contact layer beginning at a depth between

40 cm and 130 cm (SSCC 1974) from the surface. This water contact layer was observed in the Eleocharis - Triglochin (58, 59), Larix - Potentilla 53 (IT), and Picea mariana - Pleurozium (04, 46) community types. Mineral horizon Ck was occasionally present in the profiles. The lowest pH values in the Larix - Potentilla and Eleocharis - Triglochin profiles wete only slightly acid (6.2) but the Picea mariana - Pleurozium community profiles had extremely acid (4.4 and 4.2) surface horizons.

Figure 16. A Hydric Mesisol in an Eleocharis - Triglochin vegetation type (Plot 59).

Humisol Great Group

"These are organic soils with a dominantly humic middle tier, or middle and surface tiers if a terric, lithic, hydric, or cryic contact occurs in the middle tier." (SSCC 1974 p. 158)

Profile type: dominated by

A Hydric Humisol Subgroup profile (distinguished by a hydric contact beginning between 40 cm and 130 cm) was recognized in a Larix - Potentilla community type (plot 63). The only difference between this profile and other Organic Order soils was the presence of organic matter in a more decomposed state.

Sub-aqueous Soils

"This division is composed of two sub-divisions. The first consists of primitive soil formations always covered with water and not forming peat, in which primarily lower plants (algae) serve as parent material for humus formation. They are always characterized by a very simple profile structure in which B horizons are usually completely missing." (Kubiena 1953 p. 83)

Profile type: (A)C, or AC, or Ag

One Great Group equivalent was recorded in the study area.

Protopedon "Great Group"

This sub-aqueous raw soil is recognized by lacking both peat formation and a macroscopically disting- uishable humus horizon. (based on Key A, Kubiena 1953 p. 85)

The only "Subgroup" recognized was the Lake Chalk Protopedon "Subgroup".

This Lake Chalk raw soil has been defined as:

"Extremely calcareous sediments low in clay, super- ficially colonized by organisms, however showing still no humus horizon formation: the chalk fraction is of organic origin (...formation by chalk precipi- tating plants) ." (Kubiena 1953 p. 88) Profile type: Cca or (A)-Cca

These tufaceous aquatic soils were identified in the following community types: Chara - Utricularia (plots 18-23), Chara - Potamogeton (24-30), and Mimulus - Riccia (49-52).

Discussion

It is possible to make some comparisons with other northern soils. The Fort Nelson, British Columbia area had 22.5% of the surveyed area belonging to the Luvisolic Order (Valentine 1971). This Order was also recorded in the Northwest Territories but only in small amounts in the

Liard River area (6% of surveyed area - Day 1966) and the Mackenzie River area (2.7% of surveyed area - Day 1968). By contrast, Brunisolic Order soils are rare in the Fort Nelson surveyed area (3.0% - Valentine 1971) but formed a conspicuous component of the Liard River area (36% - Day 1966) and the Mackenzie River area (30.3% - Day 1968). Podzolic Order soils have not been recognized in any of these studies.

The absence of Luvisolic soils in my study area plus the comparatively high number of soil pits in the Brunisolic Order suggest that the soils in my study area had a greater affinity to the Liard River and Mackenzie

River soils than they did to the Fort Nelson area soils. These results are consistent with the general observations that Brunisolic soils are most common in the Liard River system and the northern Rocky Mountain Trench

(B. C. Lands Service, undated-a) whereas in the Fort Nelson area, Luvisolic soils dominate on glacial material (B. C. Lands Servlce,.4 undated-b).

Chemical Characteristics of the Soils

Results

Detailed results of the chemical analyses of 29 soil pits and 119 horizons are presented in Appendix G. Variations between chemical characteristics of soils within the same Order are presumed due to difference in drainage, vegetation, and topography.

These data have been surmnarized and are presented as the mean value for horizon and Order (Table XI). This table is best interpreted as PIC(,-'.-, .,.. -*- YN -tea-'.. . 0000 ddd dd ooo 0 5 7

indicating soil dynamics at a gross level. The following discussion is

based on Table XI.

$he pH values of organic horizons ranged from medium acid (5.8) in the

Gleysolic Order through slightly acid (6.4) in the Brunisolic soils to

neutral (6.8 and 7.0) in the Organic and Regosolic soils. The widespread

presence of tufa in the study area is reflected in the neutral to moderately

alkaline pH values of most mineral horizons.

Organic horizons had extremely high (3130 meq/100 g) cation exchange

capacities (CEC). Base saturation (BS) values ranged from 46 to 91% for

these horizons and varied directly with pH. Brunisolic A horizons had a

moderate (16 meq/100 g) CEC whereas the few samples in the Regosolic and

Organic A horizons had a high CEC (42 and 50 respectively). The mixed

sub-aqueous [(A)-~ca) horizons also had a high (77) CEC but the C master

horizons generally ranged from moderate (19) in the Organic Order, through moderately low (13) in Gleysolic soils, to low ((10) in the Brunisolic

and Regosolic Orders. The high base saturation (from 93 to 200%) of these

mineral horizons reflect the presence of calcareous materials in the

profiles.

Exchangeable cations (caw, M~*, ~a+, K') were recorded in the greatest

amounts in the organic horizons and tended to decrease with depth. The

calcium cation was the dominant ion present which is indicative of the wide-

spread presence of CaC03. Much less common was the magnesium catioa, and

the monovalent cations (~a' and K') were the least common.

Mean percentage nitrogen content paralleled the pattern for organic

matter content. The highest (1.9%) and lowest (0.1%) nitrogen values were

recorded for the Brunisolic organic and Bm horizons respectively. Organic

horizon C/N ratios ranged from 21.4 to 33.2 with mineral horizons tending to have higher ratios. The reported ratio of 53.47 for the Brunisolic Bm,

Bmk horizon was calculated after excluding three exceptionally high values recorded from the Apocynum - Cynoglossum and Picea glauca - Hylocomium community types. These atypical values were ~rimarilydue to very low nitrogen levels and the reported ratio would have increased some 15 times if these data had been included in the calculations.

The mean value of available phosphorous was highest in organic horizons with values ranging from 55 ppm in the Brunisols to 6 ppm in the Organic

soils. Mineral horizons had much less available phosphorous with levels below 3 ppm.

The mean sulfur content showed the highest (0.1%) value in the C horizons of Organic Order soils, and ranged downwards to at or near O"L.

There appeared to be a tendency for lower horizons within the soil profile

to have more sulfur than surface horizons. This may in part be due to the

sulfate carrying thermal waters (Pavlick 1974, Reid 1975b) tending to

evaporate and precipitate the sulfate during tufa formation.

The Ca/Mg ratio has the greatest significance in mineral (rather than

organic) horizons. Within one soil type, the ratios were lowest in the

organic horizons (12 to 22) and highest in the C horizons (16 to 92). This

is also consistent with the distribution of tufaceous material in the soil

pits and with the general tendency for pH and percentage base saturation

in the profiles.

Discussion

The C horizons appear to have unusually high organic matter contents.

This apparent anomaly is largely due to the widespread presence of tufa in

the study area. As outlined in the Thermal Springs and Tufa Deposits discussion, tufa may be formed when C02 is removed from carbonate waters.

Within the study area, the lime-secreting ability of such species as

Chara vulgaris would then result in an accumulation of calcium carbonate around the plant. The result of this process is a mineral horizon that

incorporates much organic matter. In addition, some fraction of the organic matter in the C horizons may be due to the presence of small roots in

analyzed samples.

Comparisons with other northern soils (Day 1966, Day 1968, and

Valentine 1971) show that the soils in the study area differ from other

regions in that the study area soils have a higher percentage of organic matter and nitrogen but a lower amount of available phosphorous. These

results are consistent with the widespread prevalence of tufaceous material

in the study area. The calcium saturation of the exchange complex may

then tie up the phosphorous in the form of calcium phosphate as discussed

in the VEGETATION - ENVIRONMENT RELATIONSHIPS section. This process has been reported for mildly alkaline sola in the Fort Nelson area (Valentine

1971). VEGETATION

Introduction

The major objective of this portion of the study was to describe and classify the v&etation of the study area.

A basic vegetation unit is the plant association. This has been defined as:

"An association is a plant community of definite floristic composition, presenting a uniform physiognomy, and growing in uniform habitat conditions.'' (Flahault and Schroter, 1910, Third International Botanical Congress).

The plant association concept represents a useful level of abstraction in phytosociological studies (Becking 1957, Mueller-Dombois and Ellenberg

The original intent was to describe the vegetation of the study area in terms of plant associations. However some plant communities or phytocoenoses could not be found in a sufficient (perhaps five or more) number of stands to abstract a plant association. The more general term

"vegetation type" or "(plant) community type" rather than "plant association" is therefore used throughout this study.

Methods

The basic techniques used were those based on the Zurich-Montpel

~chdolof phytosociology (Braun-Blanquet method). These phytosociological techniques have had wide use in Europe and Scandinavia as well as in

Washington, Oregon, Quebec, and British Columbia (see Brooke et al. 1970). 6 1 Investigations in British Columbia using these basic techniques with slight modifications following Krajina have included: Beil 1974, Bell 1965,

Brayshaw 1965, Brooke 1965, Brooke et al. 1970, Kojima and Krajina 1975,

McMinn 1965, Mueller-Dombois 1965, Orloci 1965, Peterson 1965, Revel 1972, and Wali and Krajina 1973. Investigators in Kluane National Park, Yukon

Territory have also used this method (Hoefs et al. 1975). Detailed descriptions of these methods are given by Braun-Blanquet (1932), Becking

(1957), Poore (1955, 1956, 1962), Moore (1962), and Mueller-Dombois and

Ellenberg (1974).

The Braun-Blanquet approach employs a subjective selection of sample sites and has been criticized on this basis. However, Moore et al. 1970, found no essential difference in the resulting community pattern when vegetation was sampled both by subjective methods and by strictly random methods. They concluded that for a conscientious, careful researcher

I! ...no advantage could be seen in randomization.. .I1 (p. 10). The Zurich-Montpellier approach was considered appropriate for my study since it has been considered "highly desirable for initial surveys in areas of complex community pattern for which no previous data are available" (Beil 1974 p. 205) and also facilitates comparisons with other vegetation studies in the province.

Reconnaissance surveys were carried out in the study area over a two week period prior to establishing sample plots. These trips resulted in familiarity with the vegetation pattern of the area. Sample plots were located on a subjective basis in areas of relatively homogeneous vegetation structure and composition - species were judged to be relatively evenly distributed on the plot surface. Subjective decisions were necessary for there is no satisfactory objective technique to test for homogeneity of vegetation (Mueller-Dombois and Ellenberg 1974).

Transitional areas between different vegetation patterns were not sampled. In total, 63 sample plots were established in the study area.

It is widely recognized that the size of plot necessary to sample the vegetation is a function of the vegetation character: - forested plots should be larger than herbaceous plots. An attempt was made to standardize plot shape in the form of a square, with a size of 100 m 2 for forest communities, 25 m2 for herbaceous communities, and 1 m2 for some flor- istically simple submerged communities. Variations from these standards occurred due to the unique nature of a sample site e.g. plots on small islands would be of irregular shape and would be smaller than the desired area.

Each vegetation plot was divided into strata based on the height and growth form of the species present (Table XII).

For each species, a visual estimate of cover was made by strata.

Cover may be defined as the vertical projection from the crown area of a species to the ground (Mueller-Dombois and Ellenberg 1974). In this study, cover was expressed as a visually estimated percentage of total plot area. This estimate is of ecological importance because (in conjunction with stratification) cover has implications for light and temperature relations, and rainfall interception (Mueller-Dombois and Ellenberg 1974).

Estimated percent cover for each species was later converted to species significance classes (Krajina 1960) by Table XI11 since phytosociological data are customarily expressed as species significance classes rather than percent cover. Table XII. Definitions for strata limits

Sub - Stratum stratum Definition

A (tree) layer A1 trees over 30.5 m (100 feet) tall A2 trees from 18.3 - 30.5 m (60 - 100 feet) tall A3 trees from 9.1 - 18.3 m (30 - 60 feet) tall B (shrub) layer B 1 woody plants from 1.8 - 9.1 m (6 - 30 feet) tall B 2 woody plants from 0.3 - 1.8 m (1 - 6 feet) tall

C (herb) layer C all herbaceous plants plus woody plants less than 0.3 m (1 foot) tall

D (moss) layera Dh bryophytes and lichens occurring on humus Ddw bryophytes and lichens occurring on decaying wood Dms bryophytes and lichens occurring on mineral soil bryophytes and lichens occurring on rock bryophytes submerged in water

a: Includes current year tree seedlings.

Mensurational data were obtained from forest communities. All trees

with a minimum diameter of 2.5 cm (1 inch) at breast height (dbh) were

measured with a diameter tape. All tree heights were measured by means of

a Haga Altimeter. Increment cores were also taken from selected trees

to determine their age. Gross volumes (uncorrected for decay and defect)

of all tree species were determined from standard tree volume tables

(Browne 1962).

Each vegetation plot was also described in terms of general strata

coverage, general plot coverage, estimated wind exposure, land type, land

form, relief shape, exposure, slope gradient, and slope above plot.

Selected vegetation plots had soil profile descriptions made and soil

samples taken for subsequent analysis. Table XIII. Species significance scalea

Class Description

solitary, with insignificant cover seldom, with insignificant cover very scattered, with small cover covering 2 - 5% of the plot covering 6 - 10% of the plot covering 11 - 25% of the plot covering 26 - 33% of the plot covering 34 - 50% of the plot covering 51 - 75% of the plot covering B 76% of the plot

a: based on the Domin-Krajina Scale. The major differences are that Class 10 (~100%cover) was not employed in my study, and non-overlapping percentage ranges were used to avoid ambiguity.

Standards (definitions, estimation and calculation methods) employed

in vegetation analysis and synthesis are given in Appendix H.

All plot data were initially sorted into broad categories based on

gross appearance (forest, non-forest, and aquatic). These categories were

successively refined by a critical comparison of vegetation and environmental data for similarities and differences, and conformity to a general pattern of relationships. This process closely resembles the successive approximation method (Poore 1962) and tabular comparison techniques given

in detail by Mueller-Dombois and Ellenberg (1974).

The main criteria used for differentiating between vegetation types were dominance, presence or constancy, and relative exclusiveness (Table XIV). This synthesis resulted in a characteristic combination of species (Braun-

~lan~uet1932) that may be used to typify the vegetation type. Plots belonging to the same vegetation type were then placed in a synthesis table on the basis of strata, presence or constancy values, and cover values. Table XIV. Categories of species used to characterize and distinguish between vegetation types.a

Symbol Species Category Definition

cd constant dominant found in 81 - 100% of plots representing the vegetation type; mean species significance 3 5.0 c constant found in 81 - 100% of plots representing the vegetation type; mean species significance < 5.0

e exclusive 1007, of occurrences are within this vegetation type se semi-exclusive 80 - 99% of occurrences are within this vegetation type s selective 51 - 79% of all occurrences are within

this vegetation type ,

P preferential widely found but the best development (as indicated by the highest calculated cover value) occurs within this vegetation type

a: species that occurred in two or more strata were rated on the basis of the stratum with the highest cover value

Sporadic species (those present on only one plot in the vegetation type) were not used for diagnostic purposes. Two vegetation types which were each described on the basis of only one study plot were characterized by the use of exclusive species.

Vegetation Units

Distinguishing floristic characteristics of the vegetation types sampled in the study area are sunrmarized in Table XV. Standardized vegetation summary tables for each vegetation type are provided as a supplement to the text. Environmental summary tables are in Appendix I. Table XV Sunnnary of the characteristic combination of species for vegetation typesa

- Submerged Aquatic bergent Aquatic Non-Forested Forested

Potamgeton pectinatus Utricularie vulg4r~s Char4 vulgaris

Riccta fluitans Cra toneuror. filicinurn Himulus guttatus Erigeron philadelphicus PIagiamium ellipticum

Typha latifolia Scirpua validus Corex lsnuginoaa

Aster junciformis Huhlenbcrgia glomeraca Glyceria striata Drepanocladus vernicosus7 Coarpylllum stellrtum Viola nephrophylla Spiranthes romanzoffians Drorera anglica Au lacorrnium palustre

Triglochin palustria Pinguicula vulgari s E1eoch.ris pauciflorr Clrex ricroglochfn

. Oplopanax horrfdum Cpaocarpius dryopteris Streptopur ampIcxifolius htteuccia struth~opter~s Actaea rubra Herocleum lanatum Viola canadensis

Apocynum androsaemifollum Cynoglossm boreale Erigeron acris Sanicula ~rilandica

Dicu mariana Pleurotium schreberi PtLlium crista-castrensis

Lrix laricina Potentilla fruticosa Juniperus ccmmunis lktula slandulosa Rubue arcticus Oxycoccus microcarpus Arctostaphylos -bra Tomenthypnum nitens

Populus tremuloides Becula papyrifera Viburnum edule Cornus stolonifera Aralia nudicaulis

Picea glauca Hylocomlum splendens

a: Vegetation types are arranged within each major category in order of increaeing dryness. Abbreviations are defined Ln the text. Species found ~n more than one stratum are given as the highest observed cover value among their strata. b: Number of sample plotr for the vegetation type. c: Cover value. d: Ruder of plots in which the rpeciee wa recorded. 67 The significance of edaphic variables is presented in the VEGETATION - ENVIRONMENT RELATIONSHIPS section and is not discussed herein.

Chara - Potamogeton (C - P) Vegetation Type

References: Table XVI; Appendix I. / Potamogeton pectinatus and Chara vulgaris are the only two species present in this non-thermal aquatic vegetation type (Table XVI). Water depth averaged 36 cm. Chara was the dominant species in terms of plant cover and exhibited less lime encrustation than was noted in the Chara - Utricularia vegetation type.

& -

Table XVI. Characteristic combination of species and synthesis table (C-P)

Species Plot Number Significance Total Con- Cover Stratum Symbols Species 30 25 29 24 26 27 28 Mean Range stancy Value / 7

C cd,p Chara 8 8 8 9 8 5 4 7.1 4-9 7 5643 vulgaris

,cd,e Potamogeton 6 5 5 3 4 7 7 5.3 3-7 7 2229 pectinatus

Lake Chalk Protopedon (LCP) soils occurred in all study plots.

Moose were frequently observed feeding in this vegetation type. In his review of moose food habit studies in North America, Peek (1974) noted that pondweeds (Potamogeton spp. ) appear to be a preferred aquatic component of diet. A Yellowstone Park moose feeding study recorded 52 observations of moose feeding on Potamogeton pectinatus (plus four records for P. alpinus), 68 but the lime-encrusted Chara sp. was recorded as a food item on only two

occasions (McMillan 1953).

Chara - Utricularia (C - U) Vegetation Type References: Table XVII; Appendix I.

This aquatic community had an average water depth of 30 cm and was

well developed in the thermal waters of Alpha Warm Ponded Swamp. Chara

vulgaris and Utricularia vulgaris are the only vascular plant species

present (Table XVII). r-

Table XVII. Characteristic combination of species and synthesis table (C-U)

Species Plot Number Significance Total Con- Cover Stratum Symbols Species 21 18 22 19 20 23 Mean Range stancy Value / 6

C cd Chara 9 8 7 7 6 4 6.8 4-9 6 4550 vulgaris

c,e Utricularia 5 5 3 5 4 7 4.8 3-7. 6 1617 vulgaris

The lime-secreting ability of Chara has been frequently noted

(Porsild and Crum 1961, Pavlick 1974, and Prescott 1970). Field observations

indicated that previously encrusted Chara could act as a substrate for new

Chara plants. The small tufaceous islands present in the Warm Ponded Swamp

may be formed by this mechanism. Soils were classified as Lake Chalk

Protopedon (LCP) . < Moose were occasionally observed in this vegetation type. These 69 animals may have an influence on the plants through their feeding activity and by their wading. A Yellowstone Park study resulted in only two observations of moose feeding on lime-encrusted Chara sp., but 57 observations for Utricularia vulgaris (MeMillan 1953). Wading stirs up a lot of sediment and field observations noted that sediment had settled on the plants in the two study plots with the lowest apparent surface velocities. L

Mimulus - Riccia (M - R) Vegetation Type References: Tables XVIII and XIX; Figure 17; Appendix I.

This emergent vegetation was found near the outlet of Psi (Plot 52) 7 0 and in "Tau" (Plots 49-51) Warm Springs. Water covered an average of

96% of the plot surface. The herbaceous layer of this vegetation is characterized by Mimulus guttatus (Figure 17),and Epilobium adenocaulon,

Riccia fluitans and Cratoneuron filicinum were most common in the D layer (Table XVIII) .

Table XVIII. Characteristic combination of species (M-R)

Stratum Constancy Symbols Species / 4

C 4 cd ,P Mimulus guttatus

2 e Epilobium adenocaulon

Dw 4 C,se,P Riccia fluitans; Cratoneuron filicinum

The C layer was dominated by Mimulus guttatus and Chara vulgaris

(Table XIX). Epilobium adenocaulon occurred on two plots and is exclusive to this vegetation type. Other rare vascular species found in this vegetation type were Carex leptalepa, Epilobium palustre, and Veronica americana (Table XIX). Glyceria striata, located on tufaceous islands is a rare northern species restricted to thermally favored areas. Phalaris arundinacea is similarly restricted. The bryophyte layer consisted of Riccia fluitans and Cratoneuron filicinum. Riccia appears to be a rare species in western Canada (Porsild and Crum 1961). Cratoneuron is an obligate calciphyte (Porsild and Crum 1961), and occurred in shallow water associated with Mimulus. Table XIX. Synthesis table (M-R)

Species Plot Number Signif j cance Total Constancy Cover Stratum Species 50 49 52 51 Mean Range /4 Value

C Mimu lu s guttatus Chara vulgaris Cicuta douglasii (Glycer ia striata)a,b (Ga lium triflorum) Epilobium adenocaulon

Dw Riccia flui tans Cra toneuron f ilicinum

Sporadic species:

C 50 (Carex leptalepa)c + 49 (Rubus pubescens) + 50 (Cinna latifolia) + 49 (Vaccinium vitis-idaea) + 50 (Epilobium palustre)c + 49 (Aralia nudicaulis) + 50 (Mitella nuda) + 52 (Veronica americana)' 3 50 (Viola renifolia) + 52 Erigeron philadelphicus 1 49 (Epilobium leptophyllum) + 52 Mentha arvensis 1 49 (Parnassia palustris) + 52 Phalaris arundinaceab +

a: Species in brackets observed on moist ground, decaying wood, or tufa. b: Restricted to thermally favored areas in the north (Reid 1976). c: Not otherwise recorded in the study area.

All soils were classified as Lake Chalk Protopedon (LCP) with tufa noted on plots 49 and 52. Water depth averaged only 12 cm with little or no apparent surface flow. j$~%gerait- UmuZus ?CE-Ml.~V@P%&:- %@$ : Refer~nces: %bles XIZ and XXT; Piggure. 18; Appendix I.

, . , * Figure 18.. Erigeron philadelphicus (Philadelphia fleabane) and Mimulus guttatus (yellow monkey flower) characterize the vascular flora. (Plot 53; July 6, 1975).

This vegetation type was found in the streams from Chi (Plot 08) and Psi (Plots 53, 54) Warm Springs. The constant dominants Erigeron m- adelphicus and Mimulus guttatus (Figure 18) and the constants Chara vulgaris and Planiomnium ellipticum characterize this vegetation type

(Table XX). Table XX. Characteristic combination of species (E-M)

Stratum Constancy Symbols Species /3

3 cd,s,p Erigeron philadelphicus cd Mimulus guttatus c Chara vulgaris

Dw 3 CYP Plagiomnium ellipticum

All plots were subject to nutrient import and export via flowing water. The mean plot coverage was 50% water, 47% mineral soil, and 3%

decaying wood.

Soils were classified as Carbonated Gleysols which were formed from unconsolidated tufa in areas of very poor drainage, The water table was

at or near the surface with flowing water present on the plots.

Erigeron philadelphicus was well developed in this vegetation.

Mimulus guttatus, Plagiomnium ellipticum, and Chara vulgaris were well

represented in all study plots (Table XXI). The sporadic Phalaris

arundinacea is restricted to thermally favored areas in the north.

The C stratum averaged 87% plot cover but the D strata averaged only 24%. The sporadic bryophyte Riccardia sp. is probably R. pinguis which has previously been recorded in the study area and is characteristic of

northern fens (Porsild & Crum 1961).

The presence of Mimulus slowed the surface current. In these

quieter waters Riccardia would form mats beneath Mimulus. Chara would

occasionally grow on the surface of these floating Riccardia mats but

was also found in the water. Wetter portions of the tufa were often 74

Table XXI. Synthesis table (E-M)

Species Plot Number Significance Total Constancy Cover Stratum Species 54 53 08 Mean Range 13 Value

C Erigeron philadelphicus Mimulus guttatus Chara vulgaris Solidago canadensis Viola nephrophylla Aster modestus Lycopus uni f lorus Ihs Plagiomnium ellipticum

Sporadic species:

C 54 Mentha arvensis 4 54 Circaea alpina + Dms 53 Cratoneuron filicinum 3 53 Cicuta douglasii + 08 Philonotis sp 5 1; 08 Phalaris + 08 Riccardia sp. 3 arundinaceaa Dh 53 Plagiomnium ellipticum 5 Ddw 53 Riccia fluitans 6 08 Riccardia speb 6

a: Restricted to thermally favored areas in the north (Reid 1976). b: May be E. pinguis. See text. c: May be g. fontana which has been previously recorded from the study area (Porsild and Crum l96l), and has been identified in the Typha - Scirpus vegetation. (Specific name is the author's suggestion.) colonized by Viola nephrophylla, with Solidago canadensis on the drier areas. Lycopus uniflorus was noted beneath the Solidago. .pF"~F - -- ,I.-* , 4

fareacea: T&le XXII; Figure 19; Appmdix I*

. $,hire vegetation, cheiractogized by $y~& !, la,tifoli@ (colaglon cat tai!) and $cirpus va14W [ra~lnrton reat b~lrush), Is rare %n the study area. (FZot 10; July 13, 1974).

This vegetation type, .represented by one plot (Figure X9), ms ' located in a depremion with only 5% cf the plor~cov~tsdby urkct, The er regime vas judged hydric. Eerbs covered IM of the plot b(rt brgophytes

covered ody 14%. 6011 samples were not taken but sema tufa trig mrpd qr~

This vegetation type is eh~racrerieedby the exclu8iva spreiw ,%*

rea, of the park. Table XXIZ. Synthesis table (T-S)

Plot Plot Stratum Species 10 Stratum Species 10

B2 Salix glauca 2 C Mentha arvensis 'Alnus incana C Typha latifolia 5 Aster junciformis Scirpus validus 5 Epilobium leptophyllum Cicuta douglasii 5 Equisetum arvense Carex lanuginosa 4 Juncus alpinus Carex aquatilis 4 Parnassia palustris Mimulus guttatus 4 Sphenopholis intermedia Aster modestus 3 Viola renifolia Carex interior 3 Lycopus uniflorus 3 Dh Philonotis fontana? Solidago canadensis 3 Plagiomnium el lip ti cum? Galium boreale 2 Brachythecium sp.

The flora was generally indicative of wet or moist areas and/or a calcareous soil. Salix glauca was the only shrub species present. The most conspicuous members in the herb cover were: Typha latifolia, Scirpus validus, Cicuta douglasii, Carex lanuginosa, C. aquatilis, C. interior,

Mimulus guttatus, Aster modestus, Lycopus uniflorus and Solidago canadensis

(Table XXII). The major components of the bryophyte layer were Philonotis fontana (?), Plagiomnium ellipticum (?), and Brachythecium sp.. These bryophytes were all found in wet areas of the plot. Plagiomnium was concentrated at the base of taller vegetation such as Lycopus. The

Brachythecium was similarily concentrated at the bases of Typha and, to a lesser degree, Cicuta.

The small size (15m 2 ) of the area covered by this vegetation type, the intermediate vigor of Typha, the shallowness and limited coverage of standing water, and the presence of Typha with low vigor in the area surrounding the study plot may indicate that this vegetation type has l~eflilikdi'tr~~iod of qms;ll.di@g$in the 5mm~diartefuture, ft Tr polsaible that: rha 3J$a& - @%em% vegetatiasa type meeventuaZly die&ljp.e.sr P$ the

Aster - Muhlenbergia (A-M) Vegetation Type References: Tables XXIII and XXIV; Figure 20; Appeodix I,

Figure 20. Muhlenbergia glomerata (bog muhly) and Aster ~unc~fomSs (rush aster) are conspicuous on these tufa islands located in a thermal pond. (Plot 41; August 19, 1974). 78 averaged 90% tufaceous mineral soil and 10% water. Bryophyte cover was as well developed as that of the herbaceous layer (both averaging about 70%) resulting in a continuous vegetation cover on the islands.

Aster junciformis, Muhlenbergia glomerata, Campylium stellatum, and

Drepanocladus vernicosus (?) were constant dominants characteristic of this vegetation type (Table XXIII). Aster and Muhlenbergia are preferential species and Glyceria striata is a selective and preferential species.

Glyceria is rare in the north and is restricted to thermally favored areas.

Table XXIII, Characteristic combination of species (A-M)

Presence Stratum / 5 Symbols Species

5 cd,p Aster junciformis; Muhlenbergia glomerata 4 S,P Glyceria striata

Dms 5 cd Campylium stellatum; Drepanocladus vernicosus (?)

Carbonated Gleysols (CG) comprised these very poorly drained, tufaceous soils. The water table was only 15 cm or less from the plot surface.

In addition to the characteristic combination of species, the herbaceous layer had species indicative of a calcareous and moist substrate:

Erigeron philadelphicus, Parnassia palustris, Viola nephrophylla, and

Eleocharis &auciflora (Table XXIV). In the bryophyte layer, Drepanocladus vernicosus is considered characteristic of northern fens (Porsild and

Crum 1961) and both 2. sendtneri and Cratoneuron commutatum are obligate calciphytes usually indicating a calcareous substrate (Porsild and Crum Table XXIV. Synthesis table (A-M)

Species Plot Number Significance Total Presence Cover Stratum Species 33 40 41 31 32 Mean Range /5 Value

C Aster junciformis Muhlenbergia glomerata Erigeron philadelphicus Glyceria striataa Parnassia palustris Viola nephrophylla Eleocharis pauciflora Solidago canadensis

Dms Drepanocladus 6 8 7 3 6 vernicosus? Campy lium 53787 stellatum Drepanocladus . 4 . 4 . send tneri Hypnum .4.3. lindbergii

Sporadic species:

C 33 Drosera anglica 5 C 31 Primula mistassinica? + 33 Lobelia kalmii 3 32 Triglochin palustris 1 33 Pinguicula vulgaris 3 41 Sphenopholis interrnedia?l Dms 33 Cratoneuron comutatum 5 41 Viola renifolia 1 31 Pohlia sp.b 2

a: Restricted to thermally favored areas in the north (Reid 1976). b: May be P. nutans previously reported for the study area (Porsild and . Crum 19z1), and identified in the Picea mariana vegetation type in this study. The tufa-forming Cratoneuron commutatum and the lime-encrusted

Drepanocladus sendtneri were located at the edge of the water. Immediately above this were Erigeron philadelphicus, Aster junciformis, and Viola nephrophylla, Parnassia palustris and Solidago canadensis. The region

from mid-slope to the island top was occupied by Muhlenbergia glomerata,

Drepanocladus vernicosus, Campylium stellatum, and Hypnum lindbergii.

Viola - Spiranthes (V-S) Vegetation T~E References: Tables XXV and XXVI; Figures 21 and 22; Appendix I,

Small tufa islands in a thermal swamp comprised the habitat for

this type of vegetation (Figure 21). Island heights above the water sur-

face were from 12 to 16 (mean: 14.4) cm. Bryophyte cover (average of 76%)

exceeded that of herbaceous cover (48%). The Carbonated Gleysols (CG) had very poor drainage with standing water observed on one plot.

Constant dominants for this vegetation type include Viola nephrophylla,

Drepanocladus vernicosus (?), and Campylium stellatum (Table XXV). Constant

species include: Spiranthes romanzoffiana (which is also semi-exclusive),

Drosera anglica, Lobelia kalmii, Parnassia palustris, Aster junciforrni.~,

Muhlenbergia glomerata, and Erigeron philadelphicus (Figure 22). An

exclusive species is Aulacomnium palustre.

The vegetation is generally indicative of a wet and/or calcareous

substrate. In addition to the characteristic combination of species, the

coqunity also includes Carex oederi, Tofieldia glutinosa, Triglochin

palustris, T. maritimum, and Solida~ocanadensis (Table XXVI). Drepanocladus

vernicosus (?) is considered characteristic of northern fens, and the Figure 21. Viola - Spiranthes vegetation (Plot 34) was found on mall tufaceaus islands surrounded by thermal water. (July 31, 1974).

Figure 22. The continuous vegetation cover consists of such characteristic species as Tofieldig glutinosa (sticky tofieldia), Lobelia kalmii (Kalm's lobelia), and Parnassia palustris (grass-of- Parnassus). (Plot 34; July 31, 1974). 82

Table XXV. Characteristic combination of species (V-S)

Stratum Presence Symbols Species / 5

5 cd,~ Viola nephrophylla c,se,P Spiranthes romanzoffiana C,S,P Drosera anglica; Lobelia kalmii

C 3 P Parnassia palustris c Aster junciformis; Muhlenbergia glomerata; Erigeron philadelphicus

Dms 5 cd Drepanocladus vernicosus (?); Campylium stellatum 4 e Aulacomnium palustre

sporadic species g. sendtneri and Crantoneuron commutatum are both con-

sidered as obligate calciphytes usually indicating a calcarous substrate

(Porsild and Crum 1961). Crantoneuron is also considered to be an important

tufa-forming moss (Porsild and Crum 1961, Sweeting 1973, Pavlick 1974).

Vertical zonation was evident on the plots. Immediately above the water line, Aster junciformis, Carex oederi, Erigeron philadelphicus, and

Triglochin spp. were located. The mid-slope position was often occupied by Parnassia palustris and Lobelia kalmii. Viola nephrophylla, Muhlen- bergia glomerata, Tofieldia glutinosa, Spiranthes romanzoffiana, and

Solidago canadensis occurred near the top of the hummocks. Bryophytes

showed a similar tendency with the water-line positions occupied by

Cratoneuron commutatum and Drepanocladus sendtneri. 2. vernicosus was always present immediately above the water-line and could extend to the mid-slope position. Campylium stellatum and Aulacomnium palustre would be found above the mid-slope position, with the latter species tending to be more prominent on the southern side of the island. Table XXVI. Synthesis table (V-S)

Species Plot Number Significance Total Presence Cover Stratum Species 38 35 34 37 36 Mean Range / 5 Value

Viola nephrophy lla Drosera anglica Aster junciformis Parnassia palustris Lobelia kalmii Muhlenbergia glomerata Spiranthes romanzo f f iana Erigeron philadelphicus Carex oederi Tofieldia glut inosa Triglochin palustris Solidago canadensis Triglochin mari timum

Drns Drepanocladus vernicosus (?) Campy1 ium stellatum Aulacomnium palustre

Sporadic species:

C 38 Juncus alpinus + Dms 35 Cratoneuron coagnutatum 4 34 Drepanocladus sendtneri 5 An environmental difference between th2s vegetation type and As,ter -

Muhlenbergia tjas in the mean maximum island height (11.2 cm for A-M and

14.4 cm for V-S). Thix difference is statistically significant at p = 0.05 (t = 2.772, df = 8) and would be indicative of slight variations in the distance to the water table, This difference is also reflected in the average water cover on the plot surface (10% for A-M and 1% for V-5).

Eleocharis - Trigloehin (&T) Vegetation Type References: Tablee WCVIS and XXVIII; Figure 23; Appendix L*

H Figure 23. Scirpus caespitosus (tufted clubrush), Eleoeharis pauciflora (beaked spike-rush), Triglochin palustrig (marsh arrow-grass), and Carex spp, (sedges) are common on hummocks. Moose disturbed the water saturated soil. (Plot 06; July 1, 1974).

Plants were concentrated on small hunrmocks (Figure 23) scattered throughout an area pockmarked by moose tracks. The characteristic combination of species included: Carex microglochin, C. oederi, C. garberi, Salix candida (C stratum), Triglochin palustris, Pinguicula vulgaris, -Tofieldia glutinosa, Aster junciformis, Eleocharis pauciflora, Calama- grostis inexpansa, and Campylium stellatum (Table XXVII). Species found on only one plot and not otherwise recorded for the study area were Campanula aurita, Carex diandra, and Salix alaxensis (Table XXVIII). The tufa-forming

obligate calciphyte (Porsild and Crum 1961) Cratoneuron commutatum was also

recorded.

Table XXVII. Characteristic combination of species (E-T)

Stratum Constancy Symbols Species /5

C 5 C,S Carex oederi; Salix candida

c 9 P Triglochin palustris; Pinguicula vulgaris; Carex garberi c Tofieldia glutinosa; Aster junciformis

4 S 9 P Eleocharis pauciflora s Calamagrostis inexpansa

2 e Carex microglochin

Dh 5 c Campyllium stellaturn

Plots averaged a small (4%) amount of water cover with humus (47%) and mineral soils (48%) comprising the remaining cover. Organic accumula-

tions (averaging 25 cm in thickness) made up the plant-occupied hummocks

whereas the mineral soils generally occupied the depressional sites disturbed

by moose tracks. Plant roots were generally concentrated within the top

17 cm in these very poorly drained soils. Soils were classified as either Table XXVIII. Synthesis table (t-'I)

Species Plot Number- Significance - Total Constancy Cover Stratum Species 07 59 60 58 06 Mean Range I5 Value

B2 Potentilla fruticosa Salix candida Larix laricina

C Carex oederi Trlglo~hin palustris Pin&uicula vulgaria Tofieldia glutinosa Salix candida Carex garheri Aster junciformia Scirpus caespitoaus Eleocharis pauciflora Potentilla f ruticosa Parnass ta palustri~ Triglochin mritimum Selaginella selaginoides Calaroagrostis inexpanse Ca rex capillaris Larix laricina Salix athebascensts Primla rnistassinica Carex mic roglochin Juncur slpinus Ag ropy ?on caninum Muhlenbergia f$loaerata

Dh Campyliua rtellatum Cratoneuron cOnnutatum

Sporadic species. 82 60 Betula glandulosa + C 59 Salix rlaxensil + 60 Betula glandulosa 1 C 07 Clyceria atriataa 1 60 Pyrola chlorant 1 07 Aster modrstus ? + 60 Campanula aurita'3 + 07 Habenaria hyperborea 7 + 60 Drosera angllca + 07 Solidago canadensis + 60 Spiranther raunzofftana + 07 Viola nephrophyla + 58 Lobelia kalmii + 59 Picea glauca + 06 Carex diandrab +

a. Rare species In the north (Retd 1976). b: Not otherwise recorded in the study area Rego Gleysols (RG) or Hydric Mesisols (HM) depending on humus thickness. Many species are common to both the Eleocharis - Triglochin and the Larix - Potentilla vegetation types. Shared elements include: Aster junciformis, Parnassia palustris, Triglochin palustris, Carex oederi, -C. garberi, 5. capillaris, Primula mistassinica, Scirpus caespitosus, Selaginella selaginoides, Pinguicula vulgaris, Salix candida, Agropyron caninum, Potentilla fruticosa, and Larix laricina. The major environmental difference between these^ two vegetation types is the average depth to the water table which was 13 cm for Eleocharis - Triglochin but 48 cm for Larix - Potentilla. The higher water table in the Eleocharis - Triglochin vegetation type was associated with an increased abundance of Eleocharis pauciflora, Triglochin palustris, and Pinguicula vulgaris in the herbaceous layer, a marked decrease of Potentilla --fruticosa and Larix laricina and absence of Juniperus communis in the shrub stratum, These two vegetation types often occurred adjacent to each other. In all instances, Eleocharis - Triglochin occupied the depressional sites whereas Urix - Potentilla was found on slightly higher ground (see Figure 28). --- The major faunal influence on the Eleocharis - Triglochin vegetation type is that caused by the physical movement of moose through the community.

The water-saturated tufaceous substrate was easily disturbed by walking moose (Figure 23).

Field observations suggested that moose do not step on the larger hummocks. These areas could eventually increase in height as organic matter gradually accumulated. Increasing distance from the water table may

then result in a vegetation change to the point where the vegetation could

C. be classified as a Larix - Potentilla type. ' &ference8: Table XXIX; Figure 24; Appea&ix 1.

-. - - I

Figure 24, Oplopanax horridurn (~eviZ'1 #Sub) war rare in the study area.

The exclusive species Oplopanax hoq~$dun,*nocarpiup dryopte.ris,

and Streptopus amplexifoiius chiracterizaz this vegetation type which is rare in the study area (Figure 24).

Only one localized area of Oplopanax occurred in a seasonal stream bed. The community probably received some seepage in this midslope position, Wpulus - Viburnum vegetation type occupied the sides of the creekbed. The*Oplopanax - Gymnocarpicum type of vegetation was restricted to the bottom of the gully where seepage-nutritional effects would be accentuated. The water regime was considered hygric for this plot. The plot was largely (96%) covered with humus. Shrub coverage was

83%, due mainly to Oplopanax. Herbs covered 45% of the plot surface with bryophyte coverage only 2%.

Table XXLX. Synthesis table (0-G)

Plot Plot Stratum Species 03 Stratum Species 03

B1 Alnus incana 3 Streptopus amplexifolius 3 Viola selkirkii 2 B 2 Oplopanax horridum 9 Adoxa moschatellina + Viburnum edule 3 Alnus incana + Populus tremuloides + Cinna latifolia + Galiun triflorum + C Oplopanax horridum 5 Maianthemum canadense + Actaea rubra 4 Mertensia paniculata + Gymnocarpium dryopteris 4 Populus sp. + Aralia nudicaulis 3 Rubus pubescens + Equisetum arvense? 3 Viola canadensisa f Mitella nuda 3

a: Variety rugulosa may be rare in the north (Reid 1976).

Other shrub species included scattered Alnus? incana and Viburnum -edule. Actaea rubra, Gymnocarpium dryopteris, Aralia nudicaulis, Equisetum arvense (?), Mitella nuda, Strepropus amplexifolius, and Viola selkirkii were abundant in the herbaceous layer (Table XXIX).

Matteuccia - Actaea (M-A) Vegetation Type

References: Tables XXX and XXXI; Figure 25; Appendix I, Figure 25. ?Aatte; ccia struthiopteris (0s trich fern) fq the moot abundazit- species in the Matteuccia - Actaea Qegetation type, (Plot 42; August 21, 1974).

Matteuccia struthiopteris, Heracleum lanatum, Ribes triste, Sonchus

sp., and Urtica lyallii are exclusive to this vegetation type. The characteristic combination of species also includes Actaea rubra, Viola selkirkii, and -V. canadensis (Table XXX). This community was generally located in concave sites or at the base of slopes. These factors may indicate a possible seepqge-influenced nutritional effect. The largely (96%) humus covered plot was dominated by herbaceous layer vegetation (average of 94% of the plot cover). Shrub and pryophyte strata were sparsely present. Plant roots were concentrated in the humus portions of the soil profile with humus thickness and the maximum rooting concentration both 91 averaging 15 cm. The imperfectly to moderately well drained soils were classified as Orthic Melanic Brunisols (oMB) or Degraded Eutric Brunisols

(DEB).

Table XXX. Characteristic combination of species (M-A)

Stratum Presence Symbols Species / 5

C 5 cd,e Matteuccia struthiopteris C,S Actaea rubra 3 e Heracleum lanatum P Viola canadensis 2 e Ribes triste; Sonchus sp.; Urtica lyallii S,P Viola selkirkii

Matteuccia struthiopteris has a mean species significance of 8.8 and dominates the community (Figure 25). Heracleum lanatum or Viola canadensis are conspicuous in the moister and drier sites respectively. The remaining conspicuous herbaceous species is Actaea rubra, but several other species are,also found in this vegetation type (Table XXXI). Urtica lyallii and

Sanicula marilandica are rare species in the north. Cornus stolonifera, a preferred item in moose diet (Peek 1974), was extensively browse3 One plot (39) was located in an open area by Psi Warm Pool. The open area surrounding the pool may have been a garden area some 20 years ago. Vegetation adjacent to the Matteuccia - Actaea type were typically on.somewhat drier sites and consisted of the Populus - Viburnum vegetation Table XXXI. Synthesis table (M-A)

Species Plot Number Significance Total Presence Cover Stratum Species 12 39 16 02 42 Mean Range /5 Value

Mat teuccia 89999 s truthiopteris Actaea rubra 534++ Heracleum 84.+. lana tum Viola ,.455 canadensisa Aster modestus 2 1 2 . , Ga 1ium 2++. . triflorum Cornus +I+. . stolonifera Mitella nuda 1. .++ Aralia .+3.. nudicaulis Viola selkirkii 3 3 . . . Ribes triste 3.+.. Botrychium .2+.. virginianum Smilacina .+.2. stellata Ma ian t hemum .I+.. canadense Sonchus sp. +I... Urtica lyalliib + + . . .

Sporadic species:

Betula papyri•’era Botrychium lanceolatum Populus balsamifera Equisetum arvense Cornus stolonifera Habenaria obtusata? Apocynum Circaea alpina androsaemifolium Rosa acicularis Taraxacum ceratophorum Urtica gracilis Vic ia americana Populus balsamifera Sanicula mari landicab Asplenium viride Adoxa moschatellina Aster ciliolatus Osmorhiza depauperata Prunus virginiana

a: Variety rupulosa may be rare in the north (Reid 1976). b: Rare species in the north (Reid 1976). Apocynum - Cynoglossum (A-C) Vegetation Type References: Tables XXXII and WUIfII; Figure 26; Appendix I.

Figure 26. fiagbaxle (A- &&r~sag&C~&&) is the wet alzundant species in the &a.c,yaam - Crn,o&L~@rm vegetation type, (Plot 53; Judy 7, 19?5),

The major characteristic species in this vegetation type is the constant dominant and semi-exclusive Apocynqm androaaemif~I.f& ($igure %), Exclusive species are Cynoglassum boranle , Erigeron atria, 4rcta.tspbgloa uva-ursi, and Rubus pedatqs.. Other herbaceous species characrerfzi- thi~ vegetation are: Smilacina s tellata, Vicia americane, Lathmcl &molpha,

Agyopyron caninurn, Solidago canadensis, Ssnicula wrtlandica, and Cal-grostis canadensis (Table XXXII). 9 4

Table XXXII. Characteristic combination of species (A-C)

Stratum Presence Symbols Species /5

B2 4 P Amelanchier alnifolia 3 SSP Prunus virginiana

C 5 cd,se,p Apocynum androsaemifolium c,e Cynoglossum boreale C, S Smilacina stellata; Vicia americana; Lathyrus ochroleucus c Agropyron caninum; Solidago canadensis

Erigeron acris Sanicula marilandica

3 s Calamagrostis canadensis

2 e Arctostaphylos uva-ursi; Rubus pedatus

Plots averaged a 29' slope gradient. Soils were rapidly or well drained. An impermeable tufa layer occurred in all soil pits with the main rooting concentration being immediately above it. Tufa was rarely exposed on the plot surface. Soils were classified as Orthic Regosols (OR), or

Orthic Melanic Brunisols (OMB). The humus layer averaged only 1 cm in depth.

The flora was generally indicative of dry tufaceous conditions. The presence of Viola nephrophylla in two plots was considered due to the geographic proximity to Gamma Hot Pool. This vegetation type is dominated by Apocynum androsaemifolium (Figure 26), but the diagnostic Cynoglossum boreale, although constant had a comparatively minor cover, Cynoglossum is rare in the north, as are Lathyrus ochroleucus, Sanicula marilandica, and'hctuca biennis. Other rare species occurring in some plots of this community include: Carex eburnea, Epilobium angustifolium, Lactuca biennis, Oryzopsis asperifolia, and Senecio pauperculus (Table XXXIII). Table XXXIII. Synthesis table (A-C)

Plot Number Significance Total Preaence Cover Stratum Species 55 14 13 57 56 Hean Range /5 V. lue

81 Prunus virgir~iana

B2 Shepherdla canadensis Juniperus c-nis Panus virginiana Amelanchier alnifolia

C Apocynum androsaenifolium Solidago canadensis Smilacina stellata Cynoglossum borealsa Vicia uericana Agropyron caninus Lathyrue ochroleucuaa Qlium boreale hnicula p.rilandicaa Erigeron acris Aralia nudicaulis Gal-grostis canadensis Carex garberi Arctostaphylos uva-ursi Viola canadensis Prunus virginiana Amelanchier alnifolia Botrychium vlrginianum Carex concinna Calium triflorum Rubue pedatur Viola nephrophylla

Dh Peltigera aphthosa

Dt Cladonia? sp. 5.5.7 3.4 0-7 3 i440

Sporadic apeciee:

Bl 14 Populua treeuloides 2 C 14 Taraucum ceratophoru + 13 Orytopria aaperifollac + 82 55 Picea glauca 3 57 Rubur idawl + 55 Cornur stolonifera 1 57 Senacio p.uciflorua + 55 Ribes oxyacanthoidea + 57 Senecio p.uparcuiurc + 55 Rosa acicularis + 14 Populus balsamifera + Dh 55 Hylocaiur apldenr 3 13 Cladonia? rp. + Aster ciliolatua 57 Pice. glauca + Rubus pubeacens Carex eburneaC Cornus canadensis Linnrea borealis Epilobiui. angustifolbumc Dt 55 Peltlgera aphthcae 7 trctuca bienniaa*' 55 Hylocaiu aplradeoa 3 Botrychiw lanceolatua 57 Tortslla tortuou 3 Nentha ervensia 56 Tortella frasilil 3 Hitella nuda 56 Ditrichum flexicaulm 3 Populus balaarnlfera

a: Pare aptcies in the north (Reid 1976). b: Variety rugulosa may be rare in the north (Reid 1976). c: Not othervire recotdod in the study area. This community was found on slopes below the Populus - Viburnum vegetation. The &ocynum - Cynoglossum sites were on much steeper slopes (averaging 29' vs. lo0) with less humus thickness (mean of 1 cm 'vs. 11 cm).

Picea mariaaa - Pleurozium (Pm-P) Vegetation Type References: Tables XXXIV and XXXV; Figure 27; Appendix I.

Figure 27. A cc~~picuousshrub layer is absent from this clwmunity. Picea mariana (black spruce), Pleurozim echreberi, _e_Ptilium crista-castrensis, and Hylocomium splendeps _e__Lare all abundant species. (Plot 45; June 25, 1975).

The constant dominant species and semi-exclusive species Picea arariana was characteristic of this type of vegetation. Exclusive species were -Carex vaginata, Smilacina trifolia, and Equisetum scirpoides. The characteristic combination of species also included: Geocaulon lividum, Listera cordata, Pleurozium schreberi, Hylocomium slendens, Ptiliyu crista-castrensis, Cornus canadensis, Equisetum arvense, Vaccinium vitis- idaea and Linnaea borealis (Table XXXIV). P

Table XXXIV. Characteristic combination of species (An-P)

Stratum Constancy Symbols Species / 4

A3 4 cd ,se Picea mariana C 4 c,se Geocaulon Jividum; Lis tera cordata C, Cornus canadensis; Equisetum arvense c Vaccinium vitis-idaea; Linnaea borealis

Carex vagina ta

Smilacina trifolia; Equisetum scirpoides

Dh 4 PA ,P Pleurozium schreberi; Ptilium crista- castrensis cd Hylocomium splendens

Moneses uniflora and Galium trifidum were sporadic species rare in the study area (Table XXXV).

This community occurred on flat relief surrounding the wetlands (see

Figure 28). The tree layer was conspicuous and the bryoflora was very prominent. The herbaceous layer was not well developed (Figure 27). -Picea mariana and Larix laricina had an average maximum age of 110 and 102 years respectively but Betula papyrifera averaged only 78 years.

Mean maximum height was 18 m (59 feet) for Picea, 16 m (51 feet) for Larix, and 10 m (32 feet) for Betula. Mean dbh of the oldest trees were 18.3 cm

(7.2 in.), 15.7 cm (6.2 in.), and 8.1 cm (3.2 in.) respectively for the three species. Gross volume averaged 8.7 cu. m/ha (124 cu. ft./acre) of which about 97% was due to 2. mariana. Table XXXV. Synthesis table (h-P)

Species Plot Number Significance Total Coartancy Cover Stratum Species 48 04 45 46 Mean Range /4 Value

A2 Picea rsriana 2..4

A3 Picea mariana Larix laricina Betula papyrifera

B1 Picea mariana 3433 Betula papyrifera .3.+ B2 Lodum groenlandicum .+.+ C Cornus canadensis Equisetum arvense Vaccinium vltis-idaea Geocaulon lividum Linnaea borealis Listera cordata Ledun groenlandicum Rubus pubescens Catex vaginata Hitella nude Smilacina trifolia Rosa acicularis Galium boreale Equisetum scirpoides

Dh Pleurozium schreberi 7778 Hylocoralum splendens 6665 Ptiliula crista-castrensis 5656 Peltigera aphthosa .+.2

Ww Cladonia rp. 25.4 Ptilium crista-ceatrcnsis 1.23 Pleurozium schreberi +..3 Hylocomium splendens I.+.

Sporadic species: Populus balramifera + nPianthaur calu&nsa Galin trifle Populus baleamifera + Rubur arcticur ? Roaa acicularir + Betula papyrilera Plcea uriana + SAlix umticoLa + Betula papyrifera + Dicranu tauticu Galiur criflorum + Pohlia wtana Ilabenaria orbiculata + C.lypo8.i. rp. Noneres uni f loraa + Ptilidiu ciliate Petasites palmtun + Tetrapblr pellucid. Viola renifolia + hltigera aptlthou Equisetum pratense +

a: Mot othervise recorded in the study area. All sites had poor drainage with an average ground water depth of 39 cm from the surface. Humus had accumulated to a mean thickness of 34 cm. Soils were classified as Rego Gleysols (RG) or Hydric Mesisols (HM) depending on whether sufficient humus had accumulated to permit the Organic Order of soil classification. This vegetation type occupies an intermediate topographic position between the Populus - Viburnum community located on higher topography with better drainage and the Larix - Potentilla community bordering the herb and graminoid dominated wetlands.

Squirrel activity is prominent in this community type as evidenced-Ky midden heaps and pruning of P. mariana. Moose trails are common and well- used in this community. -Rosa acicularis, Salix monticola (?), Cornus canadensis, and Equisetum arvense appeared to have been browsed, Salix spp. are well documented components in moose diets (Peek 1974). -- - Y--

Larix - Potentilla (L-P) Vegetation Type References: Tables XXXVI, XXXVII, and XXXVIII; Figure 28; Appendix I.

Most plants were situated on humus-covered raised hummocks in this fen-like habitat (Figure 28). Species exclusive to this vegetation type include: Oxycoccus microcarpus, Carex dioica, Arctostaphylos rubra, and

Tomenthypnum nitens. Other elements in the characteristic combination of species include: Potentilla fruticosa, Betula glandulosa, Carex interior, -C. aquatilis, C. capillaris, Rubus arcticus, Ledum groenlandicum, Primula mistassinica, Selaginella selaginoides, Viola nephrophylla, Galium boreale, Triglochin palustris, Tofieldia glutinosa, and Hylocomium splendens

(Table MXVI). Empetrum nigrum, Juncus arcticus, and Orchis rotundifolia were also exclusive to this community type. Figure 28. Humus hummocks in a fen-like habitat provided the rooting environment for the Larix - Potentilla vegetation type. The foreground is occupied by Eleocharis - Triglochin, and the background has a Picea mariana - Pleurozium cover,

Tree growth is very slow in this habitat with Larix laricina reaching a mean maximum height of only 4 m (14 feet) in an average of 115 years. -Picea glauca had slightly better growth attaining mean heights of 5 m (16 feet) in 99 years. Mean dbh for these two species were 6.9 cm

(2.7 in.) and 6.6 cm (2.6 in.) respectively. Gross volume is negligible under these growth conditions. The oldest Larix and Picea (245 and 230 years respectively) for the entire study area were found in this vegetation type

Soil drainage ranged from poorly to very poorly drained with ground water at an average depth of only 48 cm. This closely corresponded with the main rooting concentration (to 50 cm). Humus accumulation exceeded + 101 a mean of 60 cm under these conditions, and soils were classified either as Hydric Humisols (HH) or Hydric Mesisols (HPI) depending on the state of humus decomposition.

Table XXXVI. Characteristic combination of species (L-P)

Stratum Presence Symbols Species /5

B2 5 cd,s,g Potentilla fruticosa C,S,P Ledum groenlandicum CJP Larix laricina 4 se Betula glandulosa

S 9 P Juniperus communis 5 c,se Rubus arcticus C,SJP Carex capillaris; Primula mistassinica C ,S Selaginella selaginoides 4 e Oxycoccus microcarpus; Carex dioica se Carex interior; C. aquatilis

3 e Arc tostaphylos rubra

Dh 5 c Hylocomium splendens

4 e Tomenthypnum nitens

The comparatively large flora (e.g. 48 species for plot 63 - Table XXXVII) may be due to the variety of micro-habitats present. The island- like hummocks would provide various exposures and distances from the water table, The four-tiered stratification (B1, B2, C, and D) is indicative of this diversity. - 'Moose heavily browsed Salix candida and Cornus stolonifera in this vegetation type. Salix spp. and Cornuss~oniferaarewell known elements in moose diets (Peek 1974). Some browsing of Smilacina stellata, Juncos Table XXXVII. Synthesis table (L-P)

Species Plot Number Significance Total Presence Cover Stratum Species 63 62 01 61 17 Mean Range I5 Value

B1 Larix laricina 5455. Picea glauca 442.5

B2 Potentilla fruticosa Ledum groenlandicum Juniperus comrnunis Larix laricina Betula glandulosa Shepherdia canadensis Picea glauca Salix candida Lonicera dioica Cornus stolonifera Arnelanchier alnifolia

Viola nephrophylla Galium boreale Potentilla fruticosa Carex capillaris Primula mistassinica Triglochin palustris Selaginella selaginoides Rubus arcticus Tofieldia glutinosa Scirpus caespitosus Vaccinium vitis-idaea Carex interior Ledum groenlandicum Oxycoccus microcarpus Triglochin maritimum Carex aquatilis Pyrola asarifolia Pinguicula vulgaris Rubus pubescens Carex dioica Aster junciformis Carex garberi Linnaea borealis Arctostaphylos rubra Carex concinna Picea glauca Sa 1ix candida Betula glandulosa Table XXXVII. (continued)

Species Plot Number Significance Total Presence Cover Stratum Species 63 62 01 61 17 Mean Range 15 Value

Calamogrostis inexpansa Salix athabascensis Taraxacum ceratophorum Agropyron caninum Larix laricina Smilacina stellata Juncus arcticus Empetrum nigrum Carex oederi Lonicera dioica Orchis rotundifolia Rosa acicularis Parnassia palustris Juncus alpinus Cornus stolonifera Shepherdia canadensis Senecio pauciflorus

IIha Hylocomium splendens 45553 4.4 3-5 5 1360 Tomenthypnum nitens .3445 3.2 0-5 4 760 Pleurozium schreberi 3.41. 1.6 0-4 3 230 Peltigera aphthosa .I++. 0.6 0-1 3 14

Sporadic species:

B2 63 Rosa acicularis Habenaria hype rborea 01 Alnus incana Listera cordata 01 Betula papyrifera Betula papyrifera 01 Populus balsamifera Drosera rotundifolia 61 Salix glauca Viburnum edule Juniperus conanunis C 63 Salix glauca 63 Picea mariana Hypnum lindbergii 01 Solidago canadensis Larix laricina 01 Eleocharis pauciflora Potentilla fruticosa 01 Aster modestus Campylium stellatum 01 Bromus ciliatus Drepanocladus 01 Comus canadensis vernicosus? Picea glauca

a: Bryoflora also includes Rhizomnium punctatum (?), Pohlia nutans, and Planiochila aspleniodes. arcticus, Triglochin maritimum, and Pyrola asarifolia also occurred. _ /- The similarity of some elements in the flora of this community to that of the Eleocharis - Triglochin vegetation has been noted in the discussion of that community. The Larix - Potentilla vegetation had greater hummock relief, greater height above groundwater, and a much denser shrub cover (Figure 28, Table XXXVIII).

Table XXXVIII. Selected differences between the Eleocharis - Triglochin and Larix - Potentilla vegetation types

Eleocharis - Triglochin Larix - Potentilla n Mean Range n Mean Range

Hummock relief (cm) 5 2 7 11-40 4 56 46-70

Height above 3 13 8- 18 2 48 40-57 groundwater (cm)

Shrub cover (B total, %)

The Larix - Potentilla vegetation may represent r ouccessional stage from the Eleocharis - Triglochin vegetation. This passibility is partially reinforced by the observation that such species as Carex oederi, g. garberi,

Primula mistassinica, and Aster junciformis which were common to both vegetation types would be found on the lower-most portions of the Larix -

Potentilla hummocks. It is believed that this portion of the hunrmocks would bear the closest relationship to the water conditions existing on the Eleocharis - Triglochin plots. Conversely, the top-most portion of the Larix - Potentilla hummocks would represent an environment not found on the Eleocharis - Triglochin hummocks. Important Larix - Potentilla diagnostic species such as Larix ,&u&?im, Arctostaphylos rubra, and -Carex dioica were concentrated in these regions.

Populus - Viburnum (P-V) Vegetation Type References: Tables XXXIX and XL; Figure 29; Appendix I.

Figure 29. Tall Populus tremuloides (trembling aspen) and Betula papyrifera (paper birch) characterize the tree layer of this vegetation. Characteristic I understory species include Viburnum edule (hi~h- bush cranberry), Cornus stolonifera (red-osier dogwood), and Aralia nudicaulis (sarsaparilla). (Plot 43; August 22, 1974). Exclusive species were Populus tremuloides and Rubus parviflorus.

Other characteristic species included: Betula papyrifera, Viburnum edule, -Rosa acicularis, Aralia nudicaulis, Cornus canadensis, Maianthemum canadense, Actaea rubra, Mertensia paniculata, and Hylocomium splendens (Figure 29, Table XXXIX) .

Table XXXIX. Characteristic combination of species (P-V)

Stratum Constancy Symbols Species / 3

*1 2 e Populus tremuloides

A2 2 'P Betula papyrifera 2 3 cdYp Viburnum edule C7P Rosa acicularis 2 P Cornus stolonifera

3 cd,P Aralia nudicaulis C,P Cornus canadensis c Maianthemum canadense; Actaea rubra

Rubus parviflorus (?) Mertensia paniculata

D 3 c Hylocomium splendens

Mitella nuda, Pyrola asarifolia, P. secunda, and Ribes oxyacanthoides were additional species in the C layer. Lycopodium annotinum was a

sporadic species rare in the study area (Table XL). Common bryophytes included Brachythecium frigidum, B. salebrosum, and Plagiomnium drumnondii on humus and decaying wood. Ptilium crista-castrensis, Dicranum sp. and

Cladonia sp. were also found on decaying wood.

Both A and B strata were well represented with both layers averaging 107 Table XL. Synthesis table (P-'Via

Species Plot Number Significance Total Constancy Cover Stratum Species 05 09 43 Mean Range 13 Va he

A1 Populus tremuloides

A 2 Betula papyrifera Picea glauca

A3 Betula papyrifera Populus tremuloides Picea glauca

B 1 Picea glauca Cornus stolonifera

B2 Viburnum edule Rosa acicularis Cornus stolonifera

Aralia nudicaulis Viburnum edule Cornus canadensis Maianthemum canadense Rosa acicularis Actaea rubra Mertensia paniculata Rubus parviflorus? Mitella nuda Pyrola asarifolia Pyrola secunda Ribes oxyacanthoides Lathyrus ochroleucusb Viola renifolia Lonicera dioica Cornus stolonifera Rubus pubescens

Dh Hylocomium splendens 3++ Brachythecium frigidum + + . Brachythecium salebrosum + + . Plagiomniumdrummondii + + . Ddw Hylocomium splendens 422 Plagiomniumdrummondii + 5 . Brachythecium frigidum + 4 . Ptilium crista-castrensis 3 3 Dicranum sp. +3. Brachythecium salebrosum + + . Cladonia sp. ++. Table XL. (continued)

Sporadic species:

A2 43 Populus tremuloides 5 ~h~ 05 Ptilium crista-castrensis 2 Peltigera aphthosa 1 B1 09 Viburnumedule 4 Dicranum fuscescens + Populus tremuloides 3 Drepanocladus uncinatus + Alnus incana 2 Eurhynchium pulchellum + Rosa acicularis 1 Plagiomnium cuspidatum + B2 05 Betula papyrifera ~dw~05 Brachythecium Lonicera dioica erythrorrhizon 09 Ribes oxyacanthoides Bryum capillare Rubus idaeus Cetraria pinastri 43 Populus tremuloides Cladonia chlorophaea Prunus virginiana Dicranum fuscescens Picea glauca Drepanocladus uncinatus Mnium marginatum C 05 Linnaea borealis Pylaisella polyantha Bromus inermis Tortula princeps Equisetum arvenee Usnea sp. Lycopodium annot inumc 09 Plagiomnium cuspidatum Picea glauca Campylium chrysophyllum Vaccinium vitis-idaea Leskeela nervosa 09 Equisetum pratense Lophzia longidens Rubus idaeus Peltigera aphthosa 43 Petasites palmatus Habenaria orbiculata Populus tremuloides

a: The bryoflora is more completely known for plots 05 and 09 than for any other plots in the study area. These two plots had most of their non-vascular identifications completed by Dr. C.C. Chuang before his untimely death. The absence of a particular species from plot 43 (or from any other plot) does not necessarily imply that the species is absent from that plot but may indicate that the species was present in such small amounts that it was not included in the comparatively few packages selected for subsequent identification by Dr. W. B. Schofield. b: Rare species in the north (Reid 1976). c: Not otherwise recorded for the study area. d: Ptilidium pulcherrimum was present on plot 05 but the substrate was not recorded. 109 over 40% ground cover. Herbs averaged 35% with bryophytes only 3% of estimated ground cover.

Mean maximum height reached by Populus tremuloides was 37 m (120 feet) with a mean age of 142 years. Similar data for Betula papyrifera were 19 m

(62 feet) in 117 years and Picea glauca reached 24 m (80 feet) in 92 years.

The mean dbh for the three species were 43.2 cm (17.0 in.), 23.9 cm (9.4 in.), and 20.8 cm (8.2 in.) respectively. The gross volume had the highest recorded average 20.6 cu. m/ha (295 cu. ft./acre) of any vegetation type within the study area. Populus tremuloides accounted for the bulk of this gross volume.

The water regime was considered mesic for these well or moderately well drained soils. Soils were classified either as Degraded Eutric

Brunisols (DEB) or Degraded Dystric Brunisols (DDB) depending upon base saturation and pH characteristics.

Fire scars were present on the taller aspen (Populus tremuloides).

This, plus the intolerance of established birch (Betula papyrifera) to fire (Fowells 1965) would suggest that the fire was not of recent origin.

This vegetation type was often found on the same hillside as the

Picea glauca - Hylocomium vegetation but the Populus - Viburnum always occupied the more gentle slope (average of 10' vs. 41•‹ for the Picea glauca community type).

Picea glauca - Hylocomium (Pg-H) Vegetation Type

References: Tables XLI and XLII; Figure 30; Appendix I. Figure 30. The vegetatiea, t tg of $ices glayc,e (white spruce) and kylocomium sphndens was located on a steep hillside. (Plot 44; August 24, 1974).

Picea glauca is the most abundant vascular species in this vegetatioa

(Figure 30). Viburnum edule, Cornus stolonifera, Bromus inermis, Aralici nudicaulis, Pyrola chlorantha, Hylocomium splendens, Ptilium crista-

~astrensis,and Peltigera aphthosa are other elements of the characteristic

combination of species (Table XLI).

Average age and height for Picea glauca was 107 years and 26 m (85

feet) with a 24.9 cm (9.8 in.) dbh. This species was almost entirely

(99%) responsible for the gross volume of 11.3 cu. m/ha (162 cu, ft./acre).

. The sites were located on tufaceous parent material. The Degraded

Eutric Brunisols (Dm)were well drained. Table XLI. Characteristic combination of species; (Pg-H)

Stra tum Constancy Symbols Species /3

A2 3 c Picea glauca

B2 3 c Viburnum edule; Cornus stolonifera

C 3 C, S Bromus inermis c Aralia nudicaulis

2 s Pyrola chlorantha

3 cdYp Hylocomium splendens c Ptilium crista-castrensis; Peltigera aphthosa

This vegetation type consisted largely of Picea glauca and Hylocomium splendens (Figure 30). Rosa acicularis, Pyrola secunda, Vicia americana, and Cladonia sp. were present on more than one plot site. Abies lasiocarpa and Corallorhiza ttifida were sporadic species rare in the study area

(Table XLII) .

No tree seedlings were found on the plots representing this community.

This lack of regeneration may be due to the high density of Hylocomium splendens (Table XLII) which would dry out on these steep (averaging a 41' slope) southern slopes and then present an unfavorable rooting medium

(Fowells 1965). Table XLII. Synthesis table (Pg-H)

Species Plot Number Significance Total Constancy Cover Stratum Species 15 44 47 Mean Range / 3 Value

A2 Picea glauca

A3 Picea glauca

B1 Picea glauca

B2 Viburnum edule Cornus stolonifera Rosa acicularis

C Aralia nudicaulis Viburnum edule Bromus inermis Rosa acicularis Cornus stolonifera Pyrola secunda Vicia americana Pyrola chlorantha

Dh Hylocomium 9 9 9 9.0 - 3 8767 splendens Ptilium 2++ 1.3 +-2 3 40 crista-castrensis Pelt igera 11+ 1.0 +-1 3 3 7 aphthosa Cladonia sp. ++. 0.7 0-+ 2 7

Sporadic species:

A1 44 Picea glauca 3 C 15 Linnaea borealis + 15 Ribes oxyacanthoides + A3 15 Betulapapyrifera 3 47 Juniperus conmwnis 1 47 Geocaulon lividum + C 15 Abies lasiocarpaa + 15 Corallorhiza trifidaa + Dh 44 Viburnumedule + 15 Lathyrus ochroleucusb + seedling

a: Not otherwise recorded for the study area. b: Rare species in the north (Reid 1976). - VEGETATION - ENVIRONMENT RELATIONSHIPS

Previous sections separately discussed vegetation, microclimate, and soils. This section considers interactions at the level of both the individual species and the vegetation type with the environment.

Relationship of Individual Species to Micrometeorological Differences

Field results (see CLIMATE AND MICROCLIMATES section) suggested that marked differences in micrometeorological parameters existed between areas thermally influenced by springs and areas not under this thermal influence.

Consequently, phenological differences should exist between those members of a species growing in thermally influenced and non-thermally influenced areas.

Several species were selected which grew in both thermal regimes.

Phenological observations of flowering and fruiting were made on these species at irregular time intervals 9s opportunity permitted. Data were generally collected during the weekly servicing of microclimatic stations.

Smilacina stellata, Mimulus guttatus, Potentilla fruticosa, Heracleum lanatum, and Taraxacum scanicum were one (or more) week(s) "earlier" in development in thermally influenced areas as contrasted to areas not under a thermal influence (Table XLIII). This was true even for those non-thermal areas with a warm southwestern exposure on a non-forested portion of a hillside (such as the slope above Psi Warm Pool).

A previous section determined the presence of microclimatic differences in air and soil temperatures and the frost-free period between thermally and non-thermally influenced areas. The limited data presented herein Table XLIII. Selected phenological observations for 1975.

Thermally Influenced Areas Non-thermally Influenced Areas

Species Date statusa Location Date Statusa Location

Smilacina stellata May 25 •’1. Alpha Hot Pool June 2 •’1.

Mimulus guttatus May 27 •’1. Alpha Hot Pool Outlet Stream June 3 •’1. Delta Hot Pool June 3 veg . Fern Creek Potentilla fruticosa May 27 fl. Alpha Warm Ponded Swamp June 3 •’1. Epsilon Hot Pool June 10 •’1. fenland adjacent to Epsilon Hot Pool

Heracleum lanatum May 25 •’1. Alpha Hot Pool June 2 fl. slope above Psi Warm Pool June 10 •’1. Epsilon Hot Pool June 10 pre- slope above •’1. Epsilon Hot Pool June 12 fr. Alpha Hot Pool

Taraxacum scanicum Apr. 29 f 1. Beta Hot Pool 6 •’1. slope above Psi Warm Pool May 11 f r. Beta Hot Pool June 2 fr. slope above Psi Warm Pool w

a: veg. = lacking reproductive organs; •’1. = flower fully open; fr. = in fruit. may be detected in the vegetation by phenological observations.

I Relationship of Vegetation Types to the Soil Order

I This edaphic-vegetation relationship may be interpreted in two ways (Figure 31). One relationship is the percent of profiles of a soil Order I in a given plant community type and the second is the percent of a plant I community type in a given soil Order. On this basis, 33 percent of all Brunisolic profiles will be associated with the Matteuccia - Actaea

vegetation type but 100 percent of the soil profiles in this community type I were associated with the Brunisolic Order (Figure 31). I There was a general tendency for soils in an order to be associated with several plant communities. The single exception to this were the Regosolic Order profiles which were confined to the Apocynum - Cynoglossum

vegetation type.

Profiles within a given community type were usually within the same

Order. However, the Apocynum - Cynoglossum community type occurred in Regosolic and Brunisolic soils and both the Eleocharis - TrLglochin and Picea mariana - Pleurozium vegetation types occurred on Gleysolic as well

as Organic Order soils.

The Figure 31 soil sequence may approximate an increasing amount of

water (Table XLIV). Only shallow pits were dug for the Regosolic profiles

due to the presence of consolidated tufa but their topographfc location

suggests that these soils are probably the greatest distance from the

water table. Brunisolic

Organic

Regosolic

Gleysolic

Sub-aqueous C-P C-U R E-M A-M V-S E-T M-A A-C Pm-P L-P P-V Pg-H Aquatic Vegetation Non-Forested Forested Vegetation Types Types Vegetation Types

c.l Figure 31. Relationships between soil orders and vegetation types C-l Q\ Vegetation type abbreviations are defined in the VEGETATION section. Each vertical soil order scale represents 0 - 100%. Open histogram: % of soil pits in a given vegetation type. Closed histogram: % of vegetation plots in a given soil order. Table XLIV. Distance between the soil surface and the water table classified by soil order

Distance (cm) Soil Order n Mean Range

Brunisolic 9 Organic 6 Regosolic 2 Gleysolic 6 Sub-aqueous 13

Relationship of Vegetation Types to Soil Chemical Characteristics

Method of Analysis

Field measurements showed that most plant roots were concentrated in the soil organic and mineral surface horizons well above the depth of the deepest observed roots. The most critical portion of the soil environment may then be considered to be that portion of the soil profile from the surface to the bottom of the maximum root concentration. The soil environment beneath this depth will have a comparatively reduced nutritional significance for the plant community type.

This assumption has been implicit in research from other workers.

Waring and Major (1964) established a nutrient gradient based on the top

30 cm of soil and noted (p. 196):

"The surface 30 cm probably include most of the feeder root system of trees and nearly all the root systems of herbaceous plants."

. Wali and Krajina (1973) evaluated edaphic characteristics using variable profile depths that included horizons with very abundant, abundant, and few roots, but excluded horizons with very few to solitary roots.

The presence of consolidated tufa at various depths in various regions of the park necessitated the use of variable profile depths for analysis purposes. The region of maximum rooting concentration was defined as all horizons from the surface down to and including horizons with four or more roots per unit area (see SSCC 1974, p.235). Horizons used for analysis purpose8 are indicated in Appendix G.

Vegetation was compared to soil variables by summarizing the soil data to maximum rooting concentration for each soil profile within a vegetation type. A weighted mean value based on horizon depth was calculated for most variables by multiplying each horizon edaphic value by the horizon depth, summing these values, and then dividing by total horizon depth. This weighted mean value was considered to be a more accurate reflection of the soil environment of the community type than a calculation based on only a simple mean value without reference to horizon thickness. The three exceptions to this method of calculation are the reported values for percent base saturation, the carbon-nitrogen ratio, and the calcium-magnesium ratio which were all calculated directly from the appropriate weighted mean soil values. These calculations are presented as Table XLV, and this table as well as Appendix G is used to elucidate vegetation-edaphic interactions.

The following sections discuss the relationship between each soil variable akd the community type. Caution must be exercised in interpreting soils under the influence of groundwater or seepage because, as Waring and Major (1964) have pointed out, the latter situation is analogous to hydroponic culture methods. Consequently more nutrients may be available to the plants than would be apparent on the basis of a chemical soil Table XLV. Weighted mean soil values classified by vegetation typea

Vegetation Data (meq/ 100g) B S OM N P S C/N Ca/Mg Typeb ~ase' pH CEC caw ~gtf ~a' K* (%) (%) (%) (ppm) (%) ratio ratio

- C-P 1-1 7.5 86 68 9.6 0.3 0.28 91 35 1.03 (1 0.03 20 7

" C-U 1-1 7.2 4 26 0.9 0.1 0.06 100 20 0.10 41 0.05 119 30

r PI-R 4-4 7.3 112 90 8.6 0.2 0.15 88 39 0.78 3 0.03 29 10 E-M 2-4 7.6 17 36 2.0 0.1 0.08 100 21 0.23 1 0.02 54 18 A-M 1-1 7.6 9 30 1.5 0.1 0.12 100 22 1.67 < 1 0.05 8 21 V- S 1-1 7.6 16 62 3.2 0.3 0.14 100 23 0.14 Ll 0.09 96 19 E-T 3-3 7.4 27 61 3.5 0.2 0.10 100 28 0.47 l(O.01 35 17 M-A 3-8 6.4 179 116 9.8 0.2 2.28 72 66 2.31 29 (0.01 17 12 A-C 3-6 7.4 36 47 1.0 0.1 0.35 100 20 0.50 3 0.04 23 47 Pm- P 3-6 5.9 171 82 6.6 0.2 0.99 53 65 0.88 12 0.05 43 12

a: T-S and 0-G were not edaphically sampled. b: Abbreviations are defined in the VEGETATION section. c: Values represent the number of: (pits) - (horizons) used in the calculations. analysis.

A pH measurement not only measures relative acidity or a ' kalinity

but also has important implications for the probable availability of soil

nutrients Eor plants. Iron and manganese may be so soluble unler acidic

conditions that their concentrations may be toxic to some plan1:s whereas

under basic conditions these nutrients are relatively unavaila 11e and

deficiency symptoms may appear (Buckman and Brady 1967). Phosphorous

is most available under slightly acid to neutral conditions (01). cit.).

Although instances are known of plants thriving well outside their

generally preferred pH conditions (Wilde 1958), some genera1iz:~tionsare

possible. Wilde (1958) has stated that if the pH is between 5.0 to 7.0

(strongly acid to neutral), then the reaction class need not influence the

choice of tree species to be planted. Herbaceous species may prefer less

acid conditions and a range of 5.6 to 7.8 (medium acid to mildly alkaline)

is often given for "most plants" (Comar et al. 1962). This same pH range

is considered desirable for northern agricultural soils (Day 1966).

If these criteria are used to evaluate the soils to the bottom of

the main root concentration, then the mean soil values are favorable for

herbaceous vegetation. These values tend towards the low side for the

Populus - Viburnum (pH 5.8) and Pfcea mariana - Pleurozium (pH 5.9)

community types and are predictably towards the top end of the suggested

range for vegetation types with a tufaceous substrate. The calcareous

substrate of the Picea glauca - ~locomiumcommunity type produced a pH

value in excess of that suggested for trees by Wilde. However it is known that in the interior of Alaska, white spruce may be found to 914 m

(3,000 ft.) on limestone but only 762 m (2,500 ft,) or less if limestone is not the substrate. This species is known to tolerate soils with a pH of 7.5 but show chlorosis symptoms at pH 8.3 (Fowells 1965).

It should be noted that the Picea rnariana - Pleurozium community soil pits have an extremely acid surface layer in which the pH may be approximately 4.3 (Appendix G). This surface layer is considerably more acid than the weighted mean value of 5.9 given in Table XLV.

Cation Exchange Capacity

Cation exchange capacity (CEC) is a measure of the ability of soils to adsorb cations. This CEC is larger with increasing fineness of the soil texture (e.g. clay) and with increased organic content of the soil. Con- sequently, considerable variation can exist in soil horizons and values may range from practically zero to in excess of 200 meq/100 g.

The following scale characterizing the nutrient holding power of soils based upon the CEC has been proposed by Comar et al. (1962):

CEC (meq/100 g)

very low low moderately low moderate moderately high high

Most plant associations in this study must be characterized as high on this basis (Table XLV). Exceptions to this generalization are the

Viola.- Spiranthee, and Erigeron - Mimulus community types which would be rated as moderate, Aster - Muhlenbergia would be low, and Chara - Utricularia would be rated as very low. The influence of water in these cornunities may suggest a nutrient availability greater than that indicated by soil analysis.

Percent Base Saturation

Percent base saturation (%BS) is calculated as the sum of exchangeable basic cations (~a*, M~*, ~a+, K') divided by the Cation Exchange Capacity times 100%. High base saturation values "usually implies that the optimum amounts of calcium, magnesium, and potassium are available for plant growth"

(Day 1966). Comar et al. (1962) have suggested that agricultural land should have a minimum of at least 60% BS for mineral soils and 40% BS for organic soils. All community types are judged to have fertile soils

(Table XLV). In many instances the calculated value exceeds 100% BS, indicating the abundance of calcium carbonate in the horizons,

Exchangeable Calcium

Calcium (~a*) is a major nutrient that is important in plants for root development, control of cell wall permeability, and as calcium pectate of the middle lamella of cell walls. Forest soils may have from 1 meq/100 g to several times that concentration (Wilde 1958). Waring and Major (1964) state values under 2 meq/l00 g are limiting to most plants. Calciphilous forest trees are satisfied with 5 meq/100 g (Wilde 1958). Calcium is probably not limiting vegetation growth on any plot.

Indeed the widespread prevalence of tufaceous material is indicated by the generally high calcium levels. These values ranged from 26 to 143 meq/100 g for the community types sampled in this study (Table XLV).

Exchangeable Magnesium

Magnesium (M~*) is an essential nutrient used in chlorophyll, phosphorous utilization, and as an activator for enzymatic reactions

(Wilde 1958), This nutrient may be in critical supply in concentrations less than 0.5 meq/100 g (Waring and Major 1964).

All community types exceed the potentially critical level. The least

recorded magnesium is in the Chare -, Utricularia community type with

0.9 meq/100 g (Table XLV).

Calcium/Magnesium Ratio

The calcium/magnesium ratio may be used as one method to judge nutrient balance. Forest soils typically have ratio values between 3 and 5 although wide ratios (in excess of 30) do not prove unfavorable to most forest species (Wilde 1958). Very narrow ratios (less than 1) suggest calcium deficiencies (Waring and Major 1964).

Ratios ranged from 7 to 47 (Table XLV) in this study. The latter value occurred in the Apocynum - Cynoglossum community and was primarily due to a comparatively low (1 meq/100 g) magnesium content. This was the only vegetation type that had a large ratio.

Exchangeable Sodium

Sodium (~a~)is not essential for plant growth but has been traditionally determined in order to calculate percentage base saturation

(Waring and Major 1964). If the exchange complex has more than 10 to

15% sodium saturation, plant nutritional disorders may be indicated (Comar et al. 1962). Calculations based on Table XLV show that all community types have less than a 2% sodium saturation. Nutritional disorders are not indicated

on this basis.

Exchangeable Potassium Potassium (Kt) is a major plant nutrient with largely regulatory or catalytic functions in the plant. These include roles in carbon dioxide

assimilation, carbohydrate transformation, protein synthesis, and cell

division (Wilde 1958). Virgin forest soila typically have available

potassium in concentrations from 0.13 to 0.51 meq/100 g, whereas various

plant species may require from 0.08 to 0.38 meq/100 g (Wilde 1958).

The following rating scale is adapted from Comar et al. (1962):

very low 4 0.08 low 0.08 - 0.15 moderate 0.16 - 0.23 moderately high 0.24 - 0.31 high )0.31 The following community types have a high potassium availability (Table XLV): Matteuccia - Actaea (2.28 meq/100 g), Picea mariana - Pleurozium (0.99), Picea ~lauca- Hylocomium (0.82), Populus - Viburnum (0.76), Larix - Potentilla (0.43), and Apocynum - Cynoglossum (0.35). Moderately high potassium concentrations were recorded in the

Chara - Potamogeton community type (0.28 meq/100 g). Low concentrations were recorded in the Mimulus - Riccia (0.15), Viola - Spiranthes (0.14),

Aster -v Muhlenbergia (0.12), and Eleocharis - Triglochin (0.10 meq/100 g) community types. Erigeron - Mimulus and Chara - Utricularia community types were rated as very low for potassium availability (0.08 and 0.06 meq/100 g respectively).

Waring and Major (1964) have considered values below 0.10 meq/l00 g as indicating unfavorable available potassium levels for most plants. If this value is accepted as indicating a low nutrient status,then the Mimulus - Riccia, Viola - Spiranthes, and Aster - Muhlenbergia community types would be deleted from the preceeding list of community types with low potassium levels. The Eleocharis - Triglochin, Erigeron - Mimulus, and Chara - Utricularia community types would still be considered as having an unfavorable available potassium level.

It should be noted that the communities listed as being potentially limiting in potassium are all aquatic communities or have soils subject to a seepage effect. The availability of this nutrient to the vegetation may then not be adequately reflected in the analysis of soils. Water analysis suggests that these communities are under the influence of water containing at least 7 mg/l of dissolved potassium (Pavlick 1974, Reid 1975b).

Percent Nitrogen and the Carbon/Nitrogen Ratio

Nitrogen (N) is an element essential for plant growth since it is required in the amino group of proteins. The nitrogen content of soils may be therefore used as a crude index of soil fertility (Waring and Major 1964).

Nitrogen shortages may result from leaching, or from being in a •’omun- available to plants (such as the unavailable nitrogen in raw humus podzols or peat bogs), Microorganisms are also heavy nitrogen users and competition for this nutrient may result in insufficient quantities being available to higher plants, especially under neutral or alkaline soil reactions (Wilde

1958). High C/N ratios may imply that nitrogen is being used by microorganisms and that little nitrogen is available for higher plants (Waring and Major 1964). if t, r Virgin forest soils have from 0.1 to 0.3% nitrogen in the top six t- I inches and most tree species receive adequate nitrogen when it is present

at the 0.2% level (Wilde 1958). However, Waring and Major (1964) point

out that the presence of mycorrhizae, the ability of many trees to utilize

the ammonium form as well as the nitrate form, plus the generally lower

.L nitrogen requirements of trees compared to annuals may suggest more 9 stringent requirements for herbaceous vegetation. The following general guide has been proposed for agricultural soils (Comar et al. 1962):

very low low moderate high

The use of the above scale should also be made in reference to the

relative availability of nitrogen to higher plants. A C/N ratio of 13

or less generally implies conditions favorable for nitrogen availability

(Comar et al. 1962, Day 1966).

If the above considerations are used to interpret nitrogen levels,

a number of community types had soils to maximum root concentration rating high in nitrogen level. These vegetation types were: Matteuccia - Actaea, Aster - Muhlenbergia, Larix - Potentilla, Chara - Potamogeton, -Picea mariana, Mimulus - Riccia, Apocynum - Cynoglossum, Populus - Viburnum, and Eleocharis - Triglochin. In all instances except the Aster - Muhlenbergia community type, the C/N ratio was in excess of the optimum level for nitrogen availability. Of the remaining associations, the Matteuccia - Actaea community type had the lowest C/N ratio (17) and the highest recorded percent nitrogen (2.31),

leaving little doubt that this community had the greatest amount of available nitrogen. The highest C/N ratio (43) occurred in the Picea mariana community type which suggests a comparatively poor nitrogen availability.

This is to be expected because the previously noted acidic surface portion of the profile would prove a relatively poor medium for decomposition (Russell 1961), resulting in a typically higher C/N ratio. The Aster - Muhlenbergia community had a very narrow C/N ratio (8) in a poorly drained soil and these conditions have been interpreted by Comar et al. (1962) as indicating a lack of nitrogen fixation and therefore a deficiency of nitrogen rather than a high nitrogen availability.

Other vegetation types had less favorable nitrogen regimes. The Picea glauca - Hylocomium community had a moderate amount of nitrogen (0.38%), and low nitrogen levels occurred in the Erigeron - Mimulus, Viola - Spiranthes and Chara - Utricularia communities (0.23%, 0.14% and 0.10% respectively). These four communities had C/N ratios in exceas of

45 which would further support their potential nitrogen deficiencies.

The latter three vegetation types are subject to water flow or seepage effects and these reported values may be less than the actually available nitrogen.

Available Phosphorous

Phosphorous (P) is a nutrient used in cell development, phosphory- lation, and as a catalyst in respiratory reactions. Virgin forest soils have from 10 to 200 ppm available phosphorous with most trees being sufficiently supplied at 50 ppm, although pioneer deciduous species may require only 10 to 15 ppm (Wilde 1958). The following rating scale has been used by Comar et al. (1962): Available P (ppm)

very low low moderate moderately high high

A similar table was used by Day (1966).

No community had a high available phosphorous level (Table XLV).

The Matteuccia - Actaea and Populus - Viburnum vegetation types would

be classed as moderately high, and moderate amounts of available phosphorous

were noted in the Picea glauca and Picea mariana communities. All remaining community types (Larix - Potentilla, Mimulus - Riccia, Apocynum - Cynoglossum, Eleocharis - Triglochin, Erigeron - Mimulus, Viola - Spiranthes, Chara - Potamogeton, Chara - Utricularia, and Aster - Muhlenbergia) were rated as

/' very low. The difficulty of evaluating the nutrient availability to vegetation

in an aquatic environment or under the influence of seepage water has been

noted previously. This problem is compounded for available phosphorous

because previous water analysis (Pavlick 1974, Reid 1975b) did not test

for this nutrient, However the generally low rating8 on the above rating

scale plus the fact that the forest esnanunities generally had considerably

less available phosphorous than the 50 ppm given as sufficient for most

'trees would suggest that phosphorous may be deficient within the study area.

This may be due to calcium tying up phosphorous as calcium phosphate as has

been reported for mildly alkaline sola in the Fort Nelson area (Valentine

1971). This effect is discussed in a later section. Percent Sulfur

Typical surface mineral soil horizons may have from 0.02 to 0.50% sulfur (S) with organic soils containing more sulfur (Buckman and Brady

1960). All analyzed horizons (Appendix G) have less than the maximum typical surface value. Plots closest to thermal pools perhaps may be expected to show a sulfur accumulation (Appendix J) above these values but such was not the case. The plots closest to these sulfur pools

(Plots 07, 39, 58, 59) did however have the highest sulfur content.

However, in most cases the higher sulfur content of some horizons could be correlated with the organic nature of these horizons. The lowest horizon of plot 39 (Matteuccia - Actaea conanunity type) did have a comparatively high sulfur content (0.345%) and was mineral in content, thus suggesting a seepage influence from the adjacent (Psi) warm pool.

It would then appear that soils closest to the springs may have a higher sulfur content, perhaps due to seepage influences, but this is not sufficient to cause atypical sulfur levels.

Relationship of Individual Species

to Soil Chemical Characteristics

General Approach The previous section considered vegetation - environmental relationships at the community type level of complexity. An alternative is to consider these relationships at the species level. A major advantage of using this level of integration is that the comparatively easy quantification of species data will permit a more rigorous statistical analysis. This section addresses itself to the statistical consideration of species - environment relationships.

Species that had a comparatively large amount of edaphic data were used for analysis. These species were: Aralia nudicaulis, Rosa acicularis, Viola nephrophylla, Solidago canadensis, Cornus stolonifera, Trirlochin palustris, Aster junciformis, Picea glauca, and Linnaea borealis.

Additional species were not selected since the preliminary statistical analysis (Table XLVI) suggested that there was little likelihood of obtain- ing statistically significant results with a narrower bivariate data base.

Environmental data used in this portion of the analysis was limited to the edaphic environment due to the absence of a sufficient bivariate data base for other environmental parameters. A11 edaphic variables used for this stage of analysis were mean values weighted by horizon thickness after converting each horizon by a bulk density factor. This conversion results in edaphic variables expressed on a volume basis (a 1 cm thick horizon) rather than a weight basis and is considered to more adequately reflect the nutritional environment of the roots (Waring and Major 1964,

Brooke et al. 1970). Bulk density factors of 0.1 g/cc for L, F, H; 0.2 g/cc for Of, Om; 1.0 g/cc for A horizons and aquatic sediments; and 1.2 g/cc for other mineral horizons were assumed for the study area. These values were based on actual determinations for Mackenzie River (Day 1968) and Fort Nelson (Valentine 1971) area soils. These values are considered generally consistent with values reported for other areas (Buckman and

Brady 1969, Millar et al. 1965, Brooke et al. 1970). A summary of these converted values is presented in Appendix K. Like other workers (e.g.

Brooke et al. 1970, Kojima and Krajina 1975, Wali and Krajina 1973, Waring and Major 1964), no attempt was made to determine the root concentration of individual species. 13 1

A number of edaphic parameters were obtained at each soil pit. A review of the literature (Kojitha and Krajina 1975, Wali and Krajina 1973,

Waring and Major 1964) and previous analysis (see Relationship of Vegetation

Types to Soil Chemical Characteristics) resulted in the selection of seven edaphic variables that were believed to have the greatest potential relationship with individual species. The variables selected were: available phosphorous, % nitrogen, exchangeable calcium, exchangeable magnesium, exchangeable potassium, % organic matter, and pH.

These soil chemical characteristics were evaluated for the total soil pit to maximum rooting concentration and, where appropriate, this was subdivided into the organic portion and mineral portion. Humus (when present) would be the first medium encountered by a germinating plant but more mature plants may have roots in the mineral horizons. Consequently, different portions of the soil profile may be more critical in influencing plant growth at different times in the life cycle of the species.

The decision as to which statistical technique to use resulted in unexpected difficulties. The most obvious technique of multiple regression analysis has been successfully used on a single species with no recognizable drawbacks (Slough 1976). However, when Waring and Major (1964) examined plant yield - sot1 chemical characteristics multiple regression equations derived for two species, they concluded that (p 194):

"as the coefficients came out of the machine tested for significance, the equations were biologically meaningless" and

. "nor can biological conclusions be derived from either the magnitude or signs of individual coefficients in the equations". 13 2 Wali and Krajina (1973) derived plant - soil multiple regression equations for several species and noted (p. 360):

"Elimination of significant variables can also occur from the regression equation and as yet, there is no test really to check it except the judgment and experience of the researcher".

These findings appear to cast serious doubts as to the use of multiple

regression techniques for several species.

The statistical analysis method used was the calculation of a

correlation coefficient for each species and environmental variable. The

significance of these coefficients were then evaluated and the bivariate vegetation - environmental data showing statistically significant results were then further analyzed by calculating regression equations and then

evaluating the significance of the slope. These straight-forward techniques

are time-honored (Snedecor 1956, Greig-Smith 1964, Bishop 1971). i

Spearman Rank Correlation Coefficients

The Spearman rank correlation coefficient (Snedecor 1956) was

! selected for the initial analysis. This statistic (rs) has several f 1 F advantages for preliminary analysis: I i - it is distribution free and will then permit use of untransformed k i L data (thereby reducing the likelihood of making an error in the preliminary b 6 stage) . ", - it is an appropriate statistical tool when one data set is a It measurement (edaphic variables) and the other data set is an estimation

C (species cover). 1 - it does not underestimate the association for curvilinear data as Clelland 1957).

- it is fully 91% as powerful as the product-moment correlation coefficient (Bradley 1968).

In the calculation of this statistic, any tied values that appeared were averaged rather than applying a correction factor since the correction factor is negligible (usually in the order of one or two one-hundredths -

Tate and Clelland 1957). The level of significance chosen for these rs calculations was p--L0.10. This rather high value was chosen in view of the slightly lessened sensitivity of this statistic plus the use of tied values rather than applying a correction factor. The object of using p = 0.10 was thus to reject bivariate data with no apparent correlation but to include bivariate data with borderline significance which could then be tested more rigorously in a later analytical stage.

The calculated rs values were used to test the hypothesis Ho: p = 0

(the population coefficientp is not significantly different from zero) versus H1:p # 0 (the coefficient is significantly different from zero).

Standard tables (Tate and Clelland 1957, Table G, p. 132) were entered at n degrees of freedom and tabulated rs values exceeding the calculated values at the desired level of significance resulted in rejection of No and acceptance of HI, i.e. a statistically significant relationship existed.

This stage of analysis is summarized in Table XLVI. The percent plot cover of Aralia nudicaulis and Rosa acicularis showed significant organic horizon correlations with available phosphorous, exchangeable calcium, percent organic matter, pH, and exchangeable potassium. Exchangeable calcium and one other edaphic characteristic were correlated with the plot cover of Cornus stolonifera (percent organic matter), Triplochin palustris Table XLVI. Species - edaphic Spearman correlation coefficients (rs) a

Soil Aralia Rosa Viola Solidago Characteristic ~orizons~nudicaulis acicularis nephrophylla canadensis

Available Phosphorous

% Nitrogen

Exchangeable Calcium

Exchangeable Magnesium

% Organic Matter

Exchangeable Potassium

a: Sample size for T/O/M is: A.n.: 10/10/7; R.a.: 91914; V.n.: 9/4/6 (pH: 7/3/5;K: 71416) ; S.C.: 91418 (pH: 91317); (2,s.: 7/2/5; T.p.: 71413 (pH: 61413); A. j. : 71413 (pH: 61413); P.g.: 71715; L.b.: 7/7/3 b: T = total soil horizons to maximum rooting concentration 0 = organic portion of horizons M = mineral portion of horizons c: Missing data (due to low sample size) are indicated by "m"

Asterisks indicate significant correlations at the following levels: Cornus Triglochin Aster Picea Linnaea stolonifera palustris junciformis glauca borealis

-0.500 - 1. OOO* m

0.134 -0.800 m

-0.777* - 1 ,ooo* m

-0.491 -0.800 m

0,223 -0.800 m

-0.071 0.350 m

-0.330 -0,850 m (available phosphorous), and Aster junciformis (pH). Picea glauca was correlated with exchangeable magnesium. No significant correlations were found between tested soil characteristics and plot cover of Viola nephrophylla, Solidago canadensis, or Linnaea borealis, nor were any tested species correlated with percent nitrogen.

Product-Moment Correlation Coefficients

The second stage of analysis used only those data considered significant

(p 6 0.10) after evaluating the rs statistic. This second analytic stage involved calculating the more rigorous product-moment correlation coefficient

*-- (r) The non-normally distributed percent data should be converted into more normally distributed data by using an arcsin transformation (Snedecor 1956,

Bishop 1971). This transformation is necessary, for the tails of the normal distribution curve extend to infinity on either side of the mean. With percent data however, the limits imposed by confinement within the range

0% to 100% will result in a decreasing standard deviation as the sample mean moves from the middle area towards either end of the scale. This will result in the tail being abruptlyitruncated at that end of the percent range. This discrepancy from the normal distribution curve can be corrected by using an arcsin transformation in which an angle 8 is selected so that sin (3 = J%cover/100 (Bishop 1971). All estimated percent species cover data as well as percent nitrogen and percent organic matter data were transformed by using standard tables

(Snedecor 1956, Table 11.12.1, p. 318-319).

The transformation of vegetation data is straightforward, well documented, and easily achieved. However a survey of the literature 136 revealed there is no concensus of opinion as to either the need for, or the appropriate transformation to use for edaphic variables. This problem is illustrated in Wali and Krajina (1973) in which multiple regression equations are given for several species using both raw and logarithmically transformed edaphic data - in a few instances entirely different edaphic parameters between raw and transformed data appeared influential on the same species.

Kojima and Krajina (1975) used raw data for most (pH, ~a*, ~g*, ~a+,

K+, OM, N, P, and C/N) of their analysis although various transformations were used for cation exchange capacity, base saturation and soil texture.

In view of the present lack of standardization of appropriate transformations for soil characteristics, the most conservative approach at present may be to use raw or untransformed edaphic data for analysis purposes.

This approach was generally taken in my study with the exceptions of percent nitrogen and percent organic matter which used an arcsin transformation in accordance with previously noted statistical theory.

These transformed vegetation data and mostly untransformed environmental data were then used to calculate the product-moment correlation coefficient (r). These determinations were made on a Commodore SR-9190R electronic calculator by a key-stroke sequence designed to convert the pre- programmed linear regression function into the correlation coefficient.

Each calculated coefficient of correlation was tested for significance with the previous statistical hypotheses Hg: p = 0 versus H1: p # 0. Rejection of Hg and the subsequent acceptance of HI at a given probability level means a significant result, i.e. there is a significant correlation between the vegetation and the environmental parameter.

The minimum probability level at which a result was considered significant in this portion of the study was p = 0.05. Tests of significance were performed by entering standard tables (Bishop 1971, Table 79, p. 197) with n-1 degrees of freedom.

Table XLVII. Non-significant (p > 0.05) species - edaphic product-moment correlation coefficients (r)

Edaphic Species Characteristic Horizona r d f P

Aralia nudicaulis exch. ca* 0 -0.380 9 )0.10 exch. K+ 0 0.280 9 20.10 % aM 0 0.535 9 0.10 Rosa acicularis exch. caw 0 -0.585 8 0.10 % OM 0 0.489 8 >0.10 PH 0 -0.468 8 )0.10 Triglochin palus tris exch. caff' T -0.694 6 0.10 exch. ca* 0 -0.854 3 0.10

Aster junciformis exch. ca* T 0,277 6 )0.10 PH T 0.473 5 ? 0.10 Picea glauca exch. M~~ T 0,470 6 70.10

a: 0 = organic portion; T = total evaluated portion

The twenty bivariate data sets previously considered tentatively

significant by the rs statistic (Table XLVI), were used for this part of

the analysis. The use of this more rigorous statistic (r) and probability

level (p 4 0.05) resulted in eleven instances of being unable to reject Hg. These situations in which no statistically significant species - soil correlation could be detected at this stage of the study are presented in

Table XLVII. Exchangeable calcium, percent organic matter, and one other

soil characteristic were not correlated with Aralia nudicaulis (exchangeable 138 potassium) and Rosa acicularis (pH). Triglochin palustris was not

correlated with exchangeable calcium. Non-significant correlations with

Aster junciformis (exchangeable calcium and pH) and Picea glauca (exchangeable magnesium) resulted in the removal of these two species from further

consideration.

In nine instances I$-,was rejected and a significant correlation was

then shown to exist between the species ground cover and soil characteristics

(Table XLVIII). Available phosphorous was correlated with Aralia nudicaulis, -Rosa acicularis, and Triglochin palustris. Additional correlations were Aralia nudicaulis with pH, Rosa acicularis with exchangeable potassium, and Cornus stolonifera with exchangeable calcium and percent organic matter.

Table XLVIII. Statistically significant (p 5 0.05) species - edaphic product-moment correlation coefficients (r)a

Edaphic Species Characteristic ~orizonb r r2 d f P

Aralia nudicaulis avail. P pH Rosa acicularis avail. P avail. P exch. K+

Cornus stolonifera exch. ~a++ exch. Caw % OM

Triglochin palustris avail. P 0 -0.896 0.803 3 0.050

a: Edaphic variables evaluated only to the bottom of the maximum rooting concentration. r b: 0 = organic portion; M = mineral portion; T = total evaluated port ion. 139

Table XLVIII also gives the square of the coefficient of correlation

(r2). The usefulness of this value is best illustrated by an example.

The r2 value for Triglochin palustris is 0.803 which can be interpreted to indicate that, based on these data, 0.803 or 80.3% of the variability in estimated percent ground cover of this species is due to the available phosphorous supply in the organic portion of the soil. All the data shown to be significant (and therefore summarized in Table XLVIII) were then used

for further analysis.

Regression Equations

This third stage of analysis was the calculation of a least-squares

regression line for those data that had a significant (p = 0.05) correlation.

This analytic stage is desirable for the correlation coefficient merely

shows the existence of a relationship but the regression equation will

indicate the nature of the relationship existing between the vegetation and

the environmental variable (Greig-Smith 1964). The regression line was

calculated by using a pre-programmed Commodore SR-9190R electronic

calculator. The regression equations appear in Table XLIX.

A linear regression equation may be visualized as a technique to place the linear line-of-best-fit onto a scattergram, or may be viewed as a mathematical tool to predict one variable from another variable. A

simple linear regression equation will have the form Y = a + bX in which the dependent variable (Y) is estimated from the independent variable (X).

In my study the species percent ground cover (Y) may be estimated from

the appropriate edaphic variable (X). The Y intercept is given by a, and

the slope of the line or change in Y for each unit change in X is given by b. The slope of the regression line (b) was also tested for significance. Table XLIX. Species - edaphic regression equations

Edaphic Species Characteristic ~orizon~Regression Equation SEE t d f P

Aralia nudicaulis avail. P 0 Y = 1.17 + 0.206X 8.14 2.649 8 0.050

PH 0 Y = 93.31 - 12.629X 8.08 -2.692 8 0.050

Rosa acicularis avail. P T Y = 2.04 + 0.384X 6.06 2.230 7 0.100~ 0 Y = 1.16 + 0.174X 2.45 8.127 7 0.001

Cornus stolonifera exch. ~a* T Y = 17.41 - 0.003X 5.00 -2.681 5 0.050 M Yz25.12-0.003X 3.03 -5.455 3 0.025

Triglochin palustris avail. P 0 Y = 8.53 - 1.150X 1.85 -2.856 2 0.200~

a: 0 = organic portion; M = mineral portion; T = total evaluated portion. b: Not statistically significant. 141

No relationship would exist between the vegetation and the environment if the slope was zero but a relationship would exist if the slope is significantly different from zero. The hypotheses tested were Hg: b = 0 versus

HI: b # 0. A test of significance of b is Student's t distribution in which t = b/SES with n-2 degrees of freedom (Snedecor 1956). Standard tables (Snedecor 1956, Table 2.7.1, p. 46) were entered with n-2 degrees of freedom and the significance of t evaluated (Table XLIX). Two instances were noted in which the slope did not differ significantly from zero. The

Triglochin palustris slope may be due to the small sample size resulting in only two degrees of freedom. The slope for available phosphorous of the total soil pits (to maximum rooting concentration) for Rosa acicularis also showed no significant regression line. Alternative explanations for

the lack of a significant (p = 0.05) slope in these two instances may be that the relationship is not a linear one or that the variability inherent in the statistic b may have caused the lack of significance in these cases

(Snedecor 1956).

Table XLIX also gives the standard error of the estimate (SEE) of

these regression lines. The SEE has properties similar to that of the standard deviation. If a pair of lines were constructed parallel to the calculated regression line at a vertical distance of one SEE from the regression line, then about 68% of the sample points should fall between the constructed lines. Similarly, pairs of lines constructed at vertical distances from the regression line equal to two and three times the SEE

should contain about 95% and 99.7% respectively of all plotted points

(Spiegel 1961).

All analyzed data with significant (p 4 0.05) slopes are presented in graphical form in Figure 32. Each y axis represents the angle 8 (expressed -Rosa acicularis -Rosa acicularis

Available Phosphorous (organic), Exchangeable Potassium (organic), meq/l cm x 1 m2 meqll cm x 1 m2

Cornus stolonifera

Figure 32. Regression lines for plant species and edaphic variables

a: arcsin transformation

10 20 30 % Organic mat tera (mineral) , per 1 cm unit horizon Cornus stolonifera Cornus stolonifera

Exchangeable calcium (total), Exchangeable calcium (mineral), meq/l cm x 1 m2 meq/l cm x 1 m2

Aralia nudicaulis Aralia nudicaulis 1

Available phosphorous (organic), pH (organic), mepll cm x 1 m2 per 1 cm unit horizon in degrees) which is the arcsin transformation of the field-estimated percent ground cover. These may be converted to the raw data by the

following formula:

estimated % ground cover = (sin 0)2 x 100%

Based on my data (Figure 32), increasing amounts of available

phosphorous in the organic horizons were associated with increasing ground

cover for Aralia nudicaulis and Rosa acicularis. A negative relationship

existed for exchangeable calcium (total and mineral portions) and the ground cover of Cornus stolonifera, organic matter of the mineral horizons and Cornus stolonifera, and for pH of the organic horizons and Aralia nudicaulis cover. Rosa acicularis showed a positive relationship with

exchangeable potassium in the organic horizons.

Discussion

The present study used correlation coefficients and regression

equations for analysis purposes. A similar analysis technique has been previously utilized in other British Columbia studies. However, species

that I have analyzed were not reported as correlation coefficients or

regression equations by Wali and Krajina (1973), and Kojima and Krajina

(1975) used the plant association rather than the species as the unit

for analysis, so no direct comparisons are possible.

However, indirect comparisons can be made with Wali and Krajina (1973).

Their work in the McLeod Lake region of central interior British Columbia

resulted in the pbblication of graphs representing the relative abundance

of various species plotted against relative nutritional gradients. One of

the nutrients selected was the relative calcium gradient.

In the present study, significant relationships were shown to exist I between the percent cover of some species and various soil chemical k characteristics. These edaphic parameters include exchangeable calcium,

exchangeable potassium, available phosphorous, percent organic matter, and

pH. Since exchangeable calcium is related with the other edaphic factors,

it is then possible to regard each of these soil factors as a function of

exchangeable calcium. This procedure has the advantage of facilitating

, comparisons with Wali and Krajina (1973) and emphasizes the role of calcium

in the study area.

A disadvantage of this procedure is that it may lead to errors in

interpretation. For example, Aralia nudicaulis and Rosa acicularis both

showed a greater plot cover with larger amounts of available phosphorous

in their organic horizons; i.e. there is a direct relationship between plot

cover and phosphorous. If it can be shown that the amount of available

phosphorous is a function of the amount of exchangeable calcium, then the

plot cover of these species is related only indirectly or secondarily to

the calcium level. -Rosa acicularis, Cornus stolonifera, and Aralia nudicaulis were previously shown to have significant relationshbs between ground cover and

certain soil characteristics. These relationships are considered in the

following discussion. -Rosa acicularis -Rosa acicularis had a percent plot cover that increased with the available phosphorous of the organic horizons (Figure 32). An interpretation

of this relationship necessitates a review of the factors affecting avail-

ability of phosphorous to plants.

Organic phosphates are not available to plants and phosphorous intake

is "almost exclusively as inorganic phosphate ions" (Russell 1961, p. 38). C U 5 Microbial decomposition of organic phosphorous results in phosphoric acid I which reacts with an alkali to form soluble or insoluble inorganic B i phosphates (Koj ima and Kra jina 19 75). Buckman and Brady (1969) note that insoluble calcium salts are formed

at pH values of 6.5 and above. Calcium phosphate will increase in solubility

as the pH drops below 6.5 (Lutz and Chandler 1946). Soluble phosphorous

(H2P04' and HP04--) is then available for plant intake (Russell 1961). At

pH values of 5.0 and below, phosphorous once again becomes increasingly

Y unavailable as it is tied up in iron and aluminum phosphates with low

solubilities (Millar et al. 1965). Phosphorous is believed to be most i available to plants at pH values between 4.5 to 6.5 (Lutz and Chandler b 1946), or pH 5.0 to 7.0 (Millar et al. 1965), or pH 6 to 7 (Buckman and

I Brady 1969). The sample pH values for the Rosa acicularis organic horizons

ranged from 5.5 to 7.5. The presence of a high ~a* concentration depresses phosphorous

solubility (Millar et al. 1965). Russell (1961, p. 481) noted:

"The calcium ions which hold phosphate in a soil may be calcium ions in solution, exchangeable yalcium ions forming calcium phosphate on the surface of clay particles, or calcium ions anchored on the surface of calcium carbonate crystals. This process is of primary importance in inherently neutral or calcareous soils..."

Phosphorous availability to plants in the study area is therefore due

to a complex of factors including organic matter, microorganisms, soil pH,

and calcium ions. The first three Factors are related to the amount of

calcium present in the soil. . From this viewpoint, phosphorous availability may be considered to be

related to the amount of calcium. A significant negative (product-moment)

correlation (r = -0.754, df = 8, p = 0.020) was observed between available phosphorous and exchangeable calcium of the organic horizons, and available

phosphorous (Y) may be estimated from exchangeable calcium (X) (Y = 106.35 -

A large amount of exchangeable calcium would result in phosphate

fixation and a low amount of available phosphorous. Calcium phosphate

formation has been previously reported for mildly alkaline sola in the

Fort Nelson area and it was suggested that:

"theadditionof P as well as N will probably be necessary for successful crop growth" (Valentine 1971, p. 48).

The percent cover of Rosa acicularis was also inversely related to

exchangeable potassium of the organic horizons (Figure 32). The amount of

exchangeable potassium is related to the amount of exchangeable calcium

since calcium will replace potassium on the cation exchange site due to

their relative cation exchange efficiencies (Millar et al. 1961). This F effect would be accentuated (due to the law of mass action) in soils with

large amounts of calcium (Millar et al. 1961). Buckman and Brady (1969,

p. 500) noted: k ! "in certain soils potassium deficiency is apparently due i to the presence of excess calcium carbonate".

This inverse relationship between potassium and calcium has been quantified

L by Russell (1961), and Russell (1961) and Becket (1964) quantified the LI I relationship between potassium and calcium plus magnesium.

A significant (product-moment) correlation was noted between exchange-

able potassium and exchangeable calcium (r = -0.726, df = 8, p = 0.020),

and exchangeable potassium (Y) could be estimated from the exchangeable b caiciwn (X) (Y = 35.53 - 0.011X, t = -2.794, df = 7, p = 0.050). On i this basis, the amount of exchangeable potassium may be viewed as a

secondary effect of the amount of exchangeable calcium. A graph of the relative abundance of Rosa acicularis and the relative amount of replaceable (water soluble plus exchangeable) calcium is given by Wali and Krajina (1973, p. 309). They found a bimodal pattern with relative abundance peaks at 5.9% and 31.8% of maximum calcium levels.

The lower calcium level had the higher abundance peak and the species was not found on soils containing more than approximately 60% of the maximum calcium levels.

No evidence for a Rosa acicularis bimodal pattern was gathered in

the present study. The Wali and Krajina (1973) graph does indicate that

the species occurs in greatest abundance towards the lower end of the

calcium gradient. These results are consistent with the present study when the available phosphorous and exchangeable potassium are considered as expressions of the exchangeable calcium.

Cornus stolonifera

Cornus stolonifera had a percent plot cover in an inverse relationship

with the amount of exchangeable calcium in the total and mineral horizons

(Figure 32). Wali and Krajina (1973) plotted the relative abundance of

this species against the relative amount of repfaceable calcium. Their

graph (p. 309) suggests that Cornus stolonifera is more tolerant of lower

calcium levels. Calcium levels greater than that recorded at the highest

species significance resulted in a very rapid drop in relative abundance.

Data from the present study are consistent with their results.

An inverse relationship between the percent plot cover of Cornus

stolonifera and the percent organic matter of the mineral horizons was

documented (Figure 32). Soil organic matter amounts depend upon the balance

between supply and decomposition (Ovington 1954). Lutz and Chandler note

(1946, p. 164) "Calcium is of outstanding importance in its influence on decomposition" with decomposition aided by neutralizing decomposition

products, improving soil physical properties, and encouraging an active

bacterial population. A significant (product-moment) correlation

(r = 0.970, df = 4, p = 0.010) was observed between mineral horizons

exchangeable calcium and the arcsin transformation of percent organic matter, and the arcsin transformed percent organic matter (Y) may be

estimated from the exchangeable calcium (X) (Y = 2.29 + 0.004X, t = 6.878,

df = 3, p = 0,010). Percent organic matter may then be viewed as a

secondary effect of exchangeable calcium with a similar relationship to

Cornus stolonifera percent plot cover as noted in the previous paragraph.

Aralia nudicaulis

Aralia nudicaulis showed a percent plot cover in a direct relationship

with available phosphorous of the organic horizons (Figure 32). As pre-

viously noted, low available phosphorous levels tend to be associated with

high exchangeable calcium levels. However, the (product-moment) correlation

between phosphorous and calcium was not considered significant (r = -0.489,

df = 9, p > 0.100).

b The equivalent relationship was found to be significant for Rosa acicularis data and that discussion noted the relationship between calcium

and organic matter, pH, and phosphorous. Aralia and Rosa had very similar percent organic matter (sample means f standard deviations of 63.98 f 9.81% and 63.19 f 11.29% respectively)

and pH ranges (5.6 to 7.4 and 5.5 to 7.5). However, exchangeable calcium

data showed far more variability in Aralia nudicaulis sites than in Rosa

acicularis sites (1887 f 2609 and 1345 f 599 meq/l cm x 1 m2 respectively).

Examination af Aralia nudicaulis calcium data showed that sample heterogene-

ity was due to the inclusion of a lake chalk ~roto~edonsoil sam~le PA)- cca] which had a very high amount of exchangeable calcium (9700 meq/l cm x 1 m2) and which must be classified as organic under SSCC 1974 guidelines due to the 38.9% organic matter content of the sample (Appendix G - Plot 49). Calcium data from the remaining nine plots involved one or more of the L, F, and H organic horizons. Data conversion from the original weighted basis to the volume basis that is used in this portion of the analysis used bulk density factors of 0.1 g/cc For the nine L-F-H conversions but 1.0 g/cc for the single protopedon sample. The consequent heterogeneity in calcium values result in lack of a significant product-moment correlation.

A Spearman rank correlation coefficient does not underestimate the association for curvilinear data as does the more conventional product- moment correlation coefficient (Tate and Clelland 1957) and the Spearman coefficient showed a significant correlation between available phosphorous and exchangeable calcium (rs = -0.855, df = 10, p = 0.010).

These results suggested that the single anomalous calcium value

should be deleted from subsequent ca~culations. When the protopedon soil was omitted, there was a significant (product-moment) negative correlation

L between exchangeable calcium and available phosphorous (r = -0.883, df = 8, p = 0.001), and available phosphorous (Y) could be estimated from exchangeable calcium (X) (Y = 222.25 - 0.163X, t = -4.986, df = 7, p = 0.005). Aralia nudicaulis also showed a percent plot cover in an inverse

relationship with the pH of the organic horizons (Figure 32). There is a

tendency for high pH values to be associated with high amounts of exchangeable calcium but the product-moment correlation was not considered signifi-

cant (r = 0.499, df = 9, p > 0.100). However, a Spearman rank correlation coefficient showed a significant relationship between pH and exchangeable

calcium (rs = 0.797, df = 10, p = 0.010). Removal of the anomalous lake chalk protopedon soil pit from the calculations resulted in a significant (product-moment) correlation between exchangeable calcium and pH (r = 0.738, df = 8, p = 0.020), and pH (Y) could be estimated from the exchangeable calcium (X) (Y = 4.11 f

0.002X, t = 2.891, dE = 7, p = 0.025).

Wali and Krajina (1973, p. 309) plotted the relative abundance of

Aralia nudicaulis against the relative amount of replaceable calcium and their graph shows maximum abundance occurred at low (14.9% of the maximum) calcium levels. Their results are consistent with the present study when available phosphorous and pH are viewed as expressions of the exchangeable calcium. SUMMARY

The major objectives of this field study were (from INTRODUCTION

AND OBJECTIVES) :

- to determine the climate of the study area, - to determine if microclimatic differences occurred between thermally

influenced and non-thermally influenced areas of the study area,

- to classify and describe the soils within the study area, - to classify and describe the plant community types within the study

area, and

- to establish relationships between the plant community types and

selected species with soil variables.

The following sections summarize the study conclusions.

Climate

1. Climatic air temperature and precipitation patterns were documented

and found to conform to the "Dfc" designations of the Koppen

classification scheme. .

Microclimatic Differences

1. Weekly maximum and minimum microclimatic air temperature data:

a) showed statistically significant differences between

thermally influenced and non-thermally influenced areas, and

b) suggested that the annual mean temperature for thermally

influenced areas may be two celsius degrees (three or four

fahrenheit degrees) warmer than the 0 C (32 I?) recorded for

non- thermally influenced areas. Based on daily thermograph data, thermally influenced microclimatic

areas received an estimated frost-free period of 116 5.7 days

but non-thermally influenced areas received only 84 2 1.4 days. Weekly soil temperature spot readings:

a) showed that most areas have their maximum soil temperatures

in July ;

b) suggested that thermally influenced areas appear to have

minimum soil temperatures occurring earlier than non-thermally

influenced areas. This phenomenon may be due to the snowpack which

would insulate the thermally influenced soil from the low air

temperatures in winter;

c) showed that soils in the immediate vicinity of hot springs

do not freeze. This is in sharp contrast with the 21 to 27 consec-

utive weeks of frozen soil noted in all other areas.

Weekly precipitation data showed statistically significant differences

between some thermal (and non-forested) and a non-thermal (and forested)

microclimatic station. These differences may be due to stand density

effects. .

Incomplete data suggests that minimum relative humidities during

winter and spring may be higher in thermally influenced areas than

in non-thermally influenced areas.

Soils

1. Study area soils consisted of Brunisols (Orthic Melanic, Orthic

Eutric, Degraded Eutric, and Degraded Dystric), Regosols (Orthic),

Gleysols (Rego, and Carbonated), Organic soils (Hydric Mesisols,

and Hydric ~umisols), and Sub-aqueous soils (Lake Chalk Protopedon). 2. The presence of tufa in the study area is shown in the mineral

horizons by:

a) physical presence;

b) the generally neutral to moderately alkaline reaction

classes;

c) the high base saturation (93 - 100%);

d) the dominance of ~a*on the exchange complex; and

e) the generally high Ca/Mg ratio.

The generally high organic matter content of the mineral horizon

may be indicative of tufa formation around certain lime-secreting

species. The generally lower amount of available phosphorous in

the study area (as contrasted with other northern soils) may also

be interpreted as indicating the widespread prevalence of tufa.

Vegetation

1. Fifteen vegetation types were classified and described along with

important environmental influences:

a) Chara - Potamogeton was fed on by mbose in this

non- thermal, aquatic habitat.

b) Chara - Utricularia was located in a thermal pond and

experienced feeding and siltation effects from moose.

c) Mimulus - Riccia was localized in warm springs.

d) Erigeron - Mimulus had a warm spring stream habitat. e) Typha - Scirpus was rare in the study area and may

eventually disappear due to successional trends.

f) Aster - Muhlenbergia and Viola - Spiranthes occupied small

(averaging 11.2 and 14.4 cm above the water respectively) tufaceous islands in thermal waters. This difference is statistically sig-

nificant (p = 0.05) and suggests that small variations in the

distance to the water table may be reflected in the vegetation

occupying a habitat.

g) Eleocharis - Triglochin was concentrated on small organic

hummocks in water-saturated soil (averaging 13 cm to the water table)

pockmarked by moose tracks.

h) Oplopanax - Gymnocarpium was rare in the study area and was

located in a dry stream bed where it would probably receive some seepage

influences.

i) Matreuccia - Actaea grew from a nitrogen-rich 15 cm organic

accumulation on hygric soils that may have a seepage influence.

j) Apocynum - Cynoglossum contained several species rare for

the north and was found on dry tufaceous soils averaging a 29'

slope gradient on a southwestern exposure.

k) Picea mariana - Pleurozium was growing in organic accumu-

lations averaging 39 cm to the water table.

1) Larix - Potentilla had some floristic similarities to Eleocharis - Triglochin and was localized to humus covered hummocks

averaging 48 cm to the water table.

m) Xulus - Viburnum occupied mesic soils with a gentle (10')

slope gradient.

n) Picea glauca - Hylocomium was localized to steep (41')

southern dry calcareous slopes.

Vegetation - Environment Relationships

1. Phenological observations on selected species showed that plants in a thermally influenced microclimatic regime are at least one week

earlier in development than those same species in a non-thermally

influenced regime. 2. Regosolic soils were limited to the Apocynum - Cynoglossum community type. All other soils harbored more than one vegetation

type 3. Vegetation types were examined for chemical edaphic factors to the

bottom of the maximum rooting concentration. This examination

classified edaphic variables on a relative scale and revealed the

following potentially limiting relationships:

a) Chara - Potamogeton was very low in available phosphorous; b) Chara - Utricularia was very low in cation exchange capacity,

exchangeable potassium, nitrogen, and available phosphorous;

c) Mimulus - Riccia was very low in available phosphorous

and may be low in exchangeable potassium;

d) Erigeron - Mimulus was very low in exchangeable potassium

and available phosphorous, and low in nitrogen;

e) Aster - Muhlenbergia was very low cn available phosphorous, low in cation exchange capacity, and may be low in exchangeable

potassium and in nitrogen availability; f) Viola - Spiranthes was very low in available phosphorous, low in nitrogen, and may be low in exchangeable potassium;

g) Eleocharis - Triglochin was very low in available phosphorous and low in exchangeable potassium; h) Apocynum - Cynoglossum was very low in available phosphorous; i) Picea mariana - Pleurozium had very acid (pH approximately 4.3)

surface horizons ; j) Larix - Potentilla was very low in available phosphorous. The nutritional status of community types subject to seepage may

not be adequately shown by chemical analysis.

4. Selected species were statistically examined for relationships

between species cover and chemical edaphic factors to the bottom of

the maximum rooting concentration. The statistical evaluation was

made on the basis of product-moment correlation coefficients and

regression equations. Tests of significance were also conducted.

Significant results were obtained for:

a) Aralia nudicaulis and available phosphorous of the organic

horizons: r = 0.684 (p = 0.050); Y = 1.17 + 0.206 X (p = 0.050). b) Aralia nudicaulis and pH of the organic horizons:

r = -0.689 (p = 0.020); Y = 93.31 - 12.629 X (p = 0.050).

c) Rosa acicularis and available phosphorous of the total pit:

d) Rosa acicularis and available phosphorous in the organic

horizons: r = 0.951 (p = 0.001); Y = 1.16 + 0.174 X (p = 0.001).

e) Rosa acicularis and exchangeable potassium in the organic

horizons: r = 0.710 (p = 0.050); Y = -2.72 + 0.541 X (p = 0.050). f) Cornus stolonifera and exchangeable calcium in the total

pit: r = -0.786 (p = 0.050); Y = 17.41 - 0.003X (p = 0.050). g) Cornus stolonifera and exchangeable calcium in the mineral

horizons: r = -0.953 (p = 0.010) ; Y = 25.12 - 0.003X (p = 0.025). h) Cornus stolonifera and percent organic matter in the mineral

horizons: r = -0.944 (p = 0.010); Y = 26.59 - 0.910X (p = 0.025).

i) Triglochin palustris and available phosphorous in the

organic horizons: r = -0.896 (p = 0.050); Y = 8.53 - 1.150 X 15 7

(p = 0.200).

5. Edaphic variables that had a significant effect on species cover

were statistically evaluated by correlation coefficients, regression

equations, and tests of significance for their relationship with

exchangeable calcium. Exchangeable calcium was positively correlated

with pH and percent organic matter, and negatively correlated with

available phosphorous and exchangeable potassium. ApRNDXX A: Xccm daily mximm and piaiaw air temperatures (F) 6-rired by months

brimrm Minimum -Control l'herully Influenced Not Themally Influaced 01 Themally fnfluenced Not Thermally Influenced &mth UP IP AOS UPS M CPF OF l&P EP AOS VPS PU CPF OF

1974 June

JYly

August

September

October

November

December

1975 Jmuclry

February

March

llnl

Iml APPENDIX D: Summary of soil temperature data (F)

Mean Depth Mean Annual 19 74 Station (cm) ~nnual~Rangea Jul. Aug. Sept. Oct. Nov. Dec.

Thermally Influenced

AOS 5 58 10 5 6

Semi-thermally InEluenced

WPS 5 42 10 39

MA 5 43 10 m

Non-thermally Influenced

LC P 5 39 10 3 7

C PF 5 3 7 10 36

OF 5 38 10 38

a: Calculated from August 1974 through July 1975. This change in the twelve month analysis period is due to the inclusion of more 10 cm depth data and the desire to minimize year to year variations in determining the month of maximum soil temperature. b: Figures in brackets are the number of weekly instantaneous readings used in calculating the means. c: Missing one weekly reading. d: Missing two weekly readings. m: Missing data 1975 Absolute Jan. Feb. March April May June July Max. Min. Range

APfE13DIX F: Sumtary of relative humidity data CL)

-June Julv &. Slpr. E. -Nov. -Dec . -Jan. -Peb. Aprfl -July Stn. &x.Xin. &x.Pfin. bx.Uin. U8x.Nin. H.x.Nin. Uax.Hin. Hax.Win. Uax.Win. H.x.Xin. *x.Xin. &x.Xin. W.x.Hin. Uax.Hin. Max.Xin.

Coatral Station

Tbarr~llly Influenced Stationa

ED 68 44 87 61 91 69 (12) (30) (28)

AOS 87 53 84 64 83 73 (13) (31) (22)

YPS 83 41 94 48 93 57 (12) (31) (31) lion-therrullr Influenced Stationa

HA 86 50 89 53 90 60 (13) (31) (31)

CE' 68 54 94 58 96 l0 (13) (26) (31) aC 9149 9577 9383 (8) (121 (11)

a: Wers tn btacketa npremt the Wb.rs of deys u8ed for cdculatf~~. APPeLlDIX G: Soil chemical aruly8is cla88ified by vegetation type

&tison (meq/ 1001) - (X) (ratio) De8ig- Text- Depth Avail. nation ure (cm) pR CEC a* ns* N.+ K+ BS CRl N S p (PP) CfR Q/%

Cbra-Potamogeton Vegetation Type

Plots 24-30 (Lake Chalk Protopedon)

(A)-cc~ d ,2

Chra-Utricularia Vepetation Type

(We Chlk Protopedon)

7.2 3.63 26.12 0.88 0.06 0.06 100.00 19.61 0.100 0.050 0.4 119.42 29.53

Wmalu8-Riccia V~etatimType -Flot 49 (We Chalk Protopedon) (A)-Cca m > 2 7.2 130.19 97.00 9.90 0.24 0.13 82.39 38.93 0.853 0.003 8.1 26.47 9.80 -Plot 50 (klc CbUr Rocopedon) (A)-Cca a > 2 7.3 131.17 103.50 9.95 0.1 0.15 66.62 48.19 0.799 0.068 2.4 34.98 10.40

Llot 51 (We Cbaik Protopadon)

(A)-Cu r ) 2 7.3 •’30.33 106.50 10.33 0.32 0.17 90.03 67.36 0.835 0.023 1.6 32.90 10.29 RlM 52 (We CbU. ?rocopdoo)

a: r - 4aca dwing b: Values for this borizoo 0- she lurcl of tw -lee. APPENDIX C: (continued)

Horizon (meqf 100~) - (2) (ratio)

Iksig- Text- Depth Avail. nation urea (cm) pH CBC fl tlg* N.+ K+ BS an N S P (ppm) C/N CalMg

Erfperon - Mmulus Yepetation Type -Plot 53 (Carbumaced Gleyaol) Ccd S-T 10 7.6 17.96 32.25 2.02 0.10 0.08 100.00 21.64 0.316 0.043 0.3 39.72 15.97

-Plot 54 (Carbonated Cleyaol) Gca S-T 4 7.6 21.50 53.00 2.80 0.22 0.1L 100.00 25.33 0.407 (0.001 0.9 36.10 18.93

btcr - I4uhlenbergla Vexetatiar Type- Plot. 31-33, 40.41 (Carbosrted Cley.01)

Cca T s 7.6 8.75 30.50 1.44 0.14 0.12

a: S* wd; T * cmsoli&td tob; Sl * dlt b: m * d.cr d8.W dotted lbe: imdlutes the WCCa of the .uciu rooting c40c.atratioa 3'". q a d'o 0 o APPENDIX G: (continued)

Horizon (meq/lOOg)

Desig- Text- Depth nation urea (cm) pH CEC caff M~* ~a+ K+

Matteuccia - Actaea Vegetation Type

Plot 12 (Orthic Eutric Brunisol)

L 0 1 6.7 163.66 98.75 F 0 10 6.4 186.24 120.00 H...... 0 5 6.4 161.17 102.00 Ahk 1 L 5 7.3 20.31 36.00 Ahk2 SiL 10 7.0 27.31 42.25 Cca T 15 mb m m Bmkb 1 S 10 8.2 8.94 29.00 Bmkb 2 CoS 10 7.7 7.50 27.50 Bmkb 3 S ,lo 7.7 8.88 30.75

Plot 16 (Orthic Eutric Brunisol)

L ?-\I . Ak Hb Ak Bmk 1 ~rnklC B1nk2~ cklC Ck2

Plot 39 (Orthic Eutric Brunisol)

L 0 F 0 H...... 0 Ahk SL Ckl VCoS Ck2 L Bmkb 1 VCoS ~mkb2c S Ck SiL

a: 0 = organic; L = loam; Si = silt; T = consolidated tufa; S = sand; Co = coarse; C1 = clay; V = very b: m = data missing c: Values for this horizon are the mean of two samples dotted line: indicates the bottom of the maximum rooting concenfration (Z) (ratio) APPENDM G: (cmtiaued)

Horizon (WQ I - (X) (ratio) ksig- Text- Depth Avail. artion urea (cm) pH CBC c.++ &* Na+ K+ BS Ql N S p (PP) CIN CclHg

&mcyaum - Cynogloaaua Vepetation Type Blot 13 (Orthic Rego.01)

L-P 0 1 mb 128.33 109.00 6.70 0.07 1.45 89.78 61.71 1.682 < 0.001 23.2 21.28 23.19

Cca T >6 m 0.12 41.00 0.45 0.05 0.06 100.00 19.24 0.049 0.009 1.2 227.76 91.11

Plot 14 (Orthic Melanic Bcmi.01)

L 0 1 6.3 148.46 97.50 7.00 0.02 5.50 74.12 62.78 1.m 0.014 81.6 22.65 13.93 "t ...... L 30 7.6 18.19 34.25 0.38 0.05 0.16 100.00 10.97 0.243 0.015 1.0 26.19 90.13 bkl ClL 25 7.0 4.44 21.25 0.25 0.06 0.03 100.00 16.26 0.008 iO.001 0.2 1178.94 85.00

W2 ClL 33 7.2 8.75 31.75 0.38 0.06 0.04 100.00 15.22 t0.001 0.011 0.1 8828.31 83.55

L-F 0 2 7.0 U1.m 115.00 5.38 0.14 1.58 92.61 56.13 1.510 (0.001 19.2 21.56 21.38

-APmLX G: (continued) Horizon (meq/lOW (X) (ratio)

Derig- Text- Depth Avaf 1. lution urea (caf pA CEC c.* Ngce Na+ K+ BS W H S P (PP) C/N Cams

krix - Poteatflla Vexetation Type -Plot 17 (tiydric Mesimol) (h 0 21 6.5 213.90 163.75 13.12 0.45 1.19 74.10 67.16 1.277 0.003 11.2 30.51 10.96

omk

Q -Plot 63 * ah1 ah2.... obk

@•’ ck

a: 0 - mmic; C1 - clay b: m-d8tarfsriag dotted line: indiutes tbe botta of the u*Lu root$ul; coocmtratim 171a APPENDIX G: (continued)

Horizon (meq/lOOg)

Desig- Text- Depth nation urea (cm) pH CEC ~a* M~* ~a' K+

Populus - Viburnum Vegetation Type

Plot 05 (Degraded Eutric Brunisol)

L-F 0 B 0 A e L Bmk...... 1 SL Bmk 2 FS Bmk3 S Ck S

Plot 09 (Degraded Dystric Brunisol)

F-L 0 10 6.0 138.30 80.00 13.75 0.14 3.25 A e S iL 5 5.5 7.31 4.50 0.90 0.04 0.07 Bm...... Si 3 1 5.3 6.50 3.05 0.58 0.04 0.08 Ck 1 S i 14 6.2 3.75 3.75 0.68 0.06 0.06 Ck2 Si > 40 7.2 2.50 3.25 0.52 0.04 0.05

Plot 43 (Degraded Eutric Brunisol)

F-L 0 5 6.3 170.11 H 0 5 5.9 161.81 Aej m 1 m m Bmk...... S iL 19 7.5 16.12 Cca T 16 m m Ck 1 S iL 12 7.5 4.31 Ck2 S i > 13 7.8 7.19

a: 0 = organic; S = sand; L = loam; F = fine; Si = silt; T = consolidated tufa b: m = data missing dotted line: indicates the bottom of the maximum rooting concentration (% ) (ratio) APPEM)IX C: (continued)

Horizon (rrpf 100g) (X ) (ratio)

Dasig- Text- Depth Avail. rution urea fcm) pli CEC c.* -'+ %+ K+ BS M U S p (PP) CIR Ca/%

Hem &lauu - ltylocorium VapsUtion Type -Plot U (Orthic Eutric Brueisol) F-tl 6.2 167.80 110.00 7.50 0.16 1.88 -1.... 7.6...... 26.12 37.25 1.00 0.07 0.12 Llt2 7.4 14.50 32.50 1.02 0.06 0.08

Bd3 7.5 22.00 40.25 1.32 0.08 0.09

Cu 7.2 9.19 30.75 0.82 0.08 0.04 ck 8.4 17.25 38.75 1.02 0.07 0.06

Cu 3 P m m m 1

Plot 44 (Orthic Eutric Bruaiaol)

I-8 7.4 129.50 117.50 5.90 0. 1.95 7.4 34.75 49.50 1 0.06 0.59 APPENDIX H: Terminology used in the VEGETATION section and in Appendix I

Presence and Constancy Values This value is a simple fraction representing the number of times a given species is found in the plots representing a vegetation type. Presence is used when all plot sizes were not equal and constancy is used for equal sized plots.

Cover Value This value is calculated from all plots of a vegetation type. The calculated cover value reflects the mean cover per plot within the vegetation type. This calculation was made by summing the cover for that species within the vegetation type, dividing by the number of plots within the vegetation type, and then multiplying by 100. For this calculation, species significance + = 0.1% plot cover, significance 1 = 0.5% cover, and significance 2 = 1% cover.

Wind Exposure Wind exposure ratings are estimates trom vegetation cover density and topographic position and are based on a 10 point scale. Separate estimates were made for the A, B, and C vegetation layers and a value reported as 7/5/3 may be considered as having a high wind exposure for trees, moderate for shrubs, and low for herbs.

Soil Classification Soils were classified on the basis of recommended standards (SSCC 1974) as outlined in the SOILS section. Soil classification abbreviations used in Appendix I are defined in the appropriate part of the VEGETATION section.

Drainage Soil drainage is based on recommendations of SSCC 1974.

Moisture Regime Moisture regime refers to the estimated norhal soil moisture condition during the growing season. Five categories (dry, mesic, hygric, hydric, and aquatic) were used. Categories were assigned on the basis of topography, soil texture, drainage, seepage, and depth to the water table or presence and abundance of standing water.

Chemical Analysis (Weighted) Appendix I soil data are reported on a basis weighted by horizon thickness. This was considered desirable because thicker horizons would tend to have a proportionately greater influence on the vegetation than would thinner horizons and a summary based only on the number of horizons would mask the horizon thickness influence. Horizons used for these calculations were those to the bottom of the observed rooting concentration for each soil pit. APPENDIX I: Site and edaphic characteristics classified by vegetation type 'Part i. Site characteristics

Chara - Potamogeton Chara - Utricularia

Plot Data

Plot Number 30 25 29 24 26 27 28 Mean 21 18 22 19 20 23 Mean Plot Size (m2) 1 1 Extent of Type (m 2) Analysis Date (M/D/Y)

Physiography

Land Type lacust rine lacustrine Land Form pond pond------Relief Shape: Contour flat flat : Profile flat - flat Exposure (OT) all all Slope Gradient (O) Slope Above Plot (m)

Vegetation Cover (%)

C total

Plot Cover (%)

Water 100 100 I-' < 0 APPENDIX I: (Part i; continued)

Mimulus Riccia Erigeron - Mimulus Typha - Scirpus

Plot Data Plot Number Mean 54 53 08 Mean 10 Plot Size (m2) 25 10 Extent of Type (m 2 ) 175 150 40 15 Analysis Date (M/D/Y) 7/6/75 7/6/75 7/7/74 7/13/74 Wind Exposure (A/B/C) -/-/I -1-12 -/-/I -1-14 -1-12 -1313

Physiography Land Type alluvial tufaceous- alluvial Land Form - thermal stream 7benchland - depression Relief Shape: Contour ------flat- convex flat : Profile - - - - flat concave Exposure (OT) 195 195 129 192 178 128 147 235 170 a 11 Slope Gradient (O) 2 2 2 1 2 4 2 12 6 0 Slope Above Plot (m) - - - - 30 18 >I50 > 66 0

Vegetation Cover (%) B total C total ~h/~ms~ Ddw Dw D total

Plot Cover (%) HumusIMineral soilb Decaying Wood Water w a: Dh in Mimulus - Riccia and Typha - Scirpus; Dms in Erigeron - Mimulus. Ll b: Humus in Mimulus - Riccia and Typha - Scirpus; Mineral Soil in gigeron - Mimulus.

APPENDIX I: (Part i; continued)

Oplopanax - Eleocharis - Triglochin Gymnoca rpium

Plot Data Plot Number Mean 03 Plot Size (m2) 2 5 Extent of Type (m2) 2 75 Analysis Date (M/D/Y) 6/26/74 Wind Exposure (A/B/C) - 1616 -/3/2

Physiography Land Type lacustrine alluvial Land Form flatland stream bed Relief Shape: Contour flat concave : Profile flat concave Exposure (OT) a 11 190 Slope Gradient (O) 0 0 6 Slope Above Plot (m) 0 0 60

Vegetation Cover (%)

B 2 B total C total Dh Ddw Dms D total

Plot Cover (%) Humus Decaying Wood Mineral Soil Water Rock APPE1IDIX I: (Part 1; continued)

Hacteuccia - Actaea Apocvnum - Cvnoglos8um

?lot hta Plot Number 12 39 16 02 42 -Mean 55 14 13 57 56 Uean PIO~ ~iae(m2) 12 25 20 20 10 20 20 - iixtcnt of ~ype(m2) 25 60 100 8100 8100 35 3 2 25 45 30 Analysis Date (M/nN) 7/15/74 8/15/74 7/22/74 6/24/74 8/21/74 7/7/75 7/19/74 7/18/74 7/9/75 7/1/75 Wind- Exposure (A/B/c) -/-/I -2 -1313 -2 -/-/5 -/3/3 -1516 -1314 -/3/6 -/3/5 , -/4/5 -1415

Physionravhy Lnd Type colluvial - tufa colluvial -tufa dome - dome knd Form -ban of .lope- -bench- mid slope -lover rlope- mid slope Eslief Shape: Contour convex convex flat concave flat flat : Profile CO0V.X flat concave flat Eaposure ("1) ZOO 087 a11 20 2 205 247 2 15 245 255 245 Slope Gradient (0) 5 4 0 4 5 18 17 55 30 27 Slope Above Plot (m) 108 0 91 85 91 21 76 15 21 20 Vegetation Cover (2) 8 1 BZ B coul C toul Dh Wv Dr D total

Plot Cover (X) LBnus Decaying wood Rock APPENDIX I: (Part i; continued)

Picea mariana - Pleurorium brix - Potentilla

Blot DIE. Plot Number 48 W 45 46 -Hean 63 62 01 61 17 Plot Sire (m2) 100 U 7 100 3 1 Extent of Type (d) >1m 2500 >lo00 2200 12000 >10000 4000 m 4000 Amlysir Date (H/D/Y) 7/2/75 6/27/74 6/25/75 6/28/75 7/12/75 7/11/75 6/11/74 7/11/75 7/26/74 Wind Exposure (A/B/C) 7/6/4 6/3/2 6/5/3 7/5/5 61514 -/7/5 -1514 -/7/5 - 15 14 - I5 I6 Rlysiography tad Type lacustrine ulluvium Land Form flatland fenland Relief Shape: Contour f 1st -convex- flat -convex- : Profile f lat - convex- flat -convex- Exposure fOT) all- 205 all Slope Gradient (O) flat- 3 0 Slope Above Plot (m) nil - 15 ni1 - Vegcta tion Cover A2 A3 A total B1 12 B coral C total Dh Ddu D total

Plot Cover (X) Iluslus Decaying Wood Menauration 4cof OIdest Tree (years): Pice& up.' 105 100 100 135 110 230 65 35 65 Betula ppyrifera 65 90 a 78 krix lartcina 110 110 85 102 245 105 40 120 65 %%kt of Oldest Tree (feet): HCWsp.' 62 5 1 59 63 59 23 14 22 7 Betufa pepyrifera 45 43 9 32 - - - krix larteiru 56 - 53 43 51 22 14 20 9 7 Grass Voltme (w.ft./acr~) Pice. sp.. 113 106 134 127 120 2 41 1 < 1 Setula papyrifera

a: 2. mriana in Picea ~rirna- Pleurotium; P. tl.UC. in krix - Potmtilla. APPENDIX I: (Part i; continued)

POUU~US - ~iburarrn - Picu glauu - Ilyloccdum

Plot wu Plot Number plot Sire (J) &tent of Tgpe (m2) kulysia hte (M/D/Y) Wind Expowre (A/B/C)

Physiography knd vpc alluvirl Cufacewe had Form low ridge lw ridge lwer aloye mid- slope Relief Sbpe: Contour convex flat flat complex flat concave : Profile convex CODUVC convex complex Exposure (OT) 275 085 210 220 205 222 Slope Gradient (O) 16 5 9 55 30 3 7 .Slope Above Plot (m) 0 46 96 41 40 45

Vegetation Cover (X) A1 A2 A3 A total B 1 B2 B total C total Dh Ww D total

Plot Cover (X) IlrPua Decaying hod

lksurrtioa &ge of Oldest Tree (years): Populus truloides Betula papyriferr 1rS Pkcea glruu 95

mt@t OE '~.11e*t rree (feet): Populus truloides Betula plpyrlfera 75 Pice. gl-u 75

Crosa Volme (cu. f t. /acre) ?opalus treatloides Ikmla p.pyri.fesa 79 mcea &uu 63 Toul L42 APPENDIX I: (continued) part ii. Edaphic characteristics

Chara - Potornogeton Chara - Utricularia

Plot Data Plot Number 30 25 29 24 26 27 28 Mean 21 18 22 19 20 23 Mean Analysis Date (M/D/Y) 6/23/75 6/23/75 - Classification LCP -- \- LCP Parent Material lakebed lakeb ed - Drainage aquatic aquatic- Moisture Regime aquatic aquatic - Standing Water Depth (cm) 40 28 44 22 36 44 38 36 30 28 30 35 20 30 Impermeable Layer (cm) ni 1 nil Stoniness of Profile (%) 0 0 Thickness of Humus (cm) 0 - 0 Root Distribution: main concn. to (cm) maximum depth to (cm) Chemical Analysis -Mean a pH 7.2 CEC (meq/100 g) 3.6 26.1 0.9 0.1 K+ (meq/100 g) 0.1 Organic Matter (%) 19.6 Nitrogen (%) 0.10 Available Phosphorous (ppm) 0.4 Sulfur (%) 0.05 Base Saturation (%) 100.0 C/N ratio 119.4 Ca/Mg ratio 29.5 a: Samples combined prior to analysis. b: m = missing data. 4PPMDIX I; (Part ii. continued)

Plot Data Plot Number 50 49 5 2 51 41 35-38 Analysis Date (M/D/Y) 7/4/75 6/23/75 Classification UP CG CC Parent Material stream sediments - tufa tuf. Drainage aquatic vpd vpd Moisture Regime aquatic hydric bydric Standing Water Deprh (cm) 10 15 l5 7 0- 15 0-16 Impermeable Layer nil Stoniness of Profile (7.) Thickness of Humus (a) Root Distribution: main concn. to (cm) wximum depth to (cm)

Chemical Analysis (Weighted)* Weighted neona kinhted ilcana md pH 7.3 7.2 7.3 7.3 7.3 7.7 7.5 7.6 7.6 CEC (meq1100 8) ~a*(meq1100 g) ng++ (meq1100 g) ~a+imeq/100 g) K+ (meq/lOO g) Organic Matter (7.) Nitrogen (%) Available Phosphoroi Sulfur (%) Base Saturatzon (4.) C/N ratio Ca/Mg ratio

a: Weighted by horizon thickness. b: Tufa. c: m = missing data. d: Samples combined prior to annlyris APPENDIX I: (Part ii. continued)

Eleocbria - Triglochin Pktteuccia - Ac~u Apoc~rrnu~~- ~~loaaum

Plot Data Plot Number 07 59 58 -Hean 12 39 16 Ik.n 55 14 13 nun Analysis Date (M/D/Y) 9/5/74 7/14/75 7/14/75 9/10/74 9/10/74 9/9/74 7/14/75 9/12/74 9/8/74 Classification RG Lh( W OEB om om 01 (18 OR Parent Material -lake bed - -lake bed - tufa lake bad tufa Drainage -vpd - id mwi wd wd wd r d Moisture Regime -hydric - -hygric - -moderatelv dm- Ground Water Depth (cm) 12 8 18 13 6 7 >135 b130 .I11 254 Impermeable Zayer nil- nil- Stoniness of Profile (Z) -0- 45 Thickness of Humus (cm) 2 35 39 25 15 1 Root Distribution: main concn. to (cm) 17 15 19 maximum depth to (cm) 42 75 5 1

Chemical Analysis Weighted Weighted Weighted (Weighted)a -Heana Wna PH 7.4 % -73 CEC (meqf100 g) 26.5 178.6 36.3 ~az(meq/100 g) 60.9 116.3 46.5 Mg (meq/ZOO g) 3.5 9.8 1.0 Na+ (meq/ 100 g) 0.2 0.2 0.1 K+ (meqllOO g) 0.1 2.3 0.4 Organlc Matter (%) 28.1 66.1 20.0 Nltrogen (%) 0.47 2.31 0.50 Available Phosphorous 0.7 29.3 3.1 (PP~) Sulfur (%) (0.01 c0.01 0.04 Base Saturation (%) 100.0 72.0 100.0 C/N ratio 35.0 16.6 23.5 CaiMg ratio 17.4 11.8 47.5

a: Ueighted by horizon thickness. - b: Unconsolidated tufa. c. Consolidated tufa. d: m = missing data. &-I]( X; (Part ii. coatinued)

Pic- ~fi.lll- PhUt02i~ krix - fotmtill~ POVUlu. - VibrtrmE Pice. X~UU- Ihl~caPium

48 (24 46 -Wua 05 09 43 h1y.i~bte wan) 7/16/75 9/5/74 7/16/75 9/7/74 9/5/74 9/4;74 Clmrifiutioa kG m 111 DEB DDB DEB -0- Panat Ifaterial -I&? bed - lake bed lake bed tufa -tufa- hiluge ps- vd vd ad -ud- Ibiscure Regime -hydric - -reic - -drg - Gmod Water Depth (a) 22 48 47 48 >110 >90 ?90 >97 ,105 >70 ,88 Lsprnuble kyrr oil- -nil- nil 31-70" Stoninem of Profile (X) 40" 60b Thietne.. of almu. fc~) 11 3 6 4 Root Dincribution: rin conen. to (a) mxI.IUP depth to (a)

&iTSRtIX K: S-ry of weighted mean edaphic valusu to the bottom of the mximum rooting concentration as classified by vegetation type and profile scgscnta

, Unit horizon. 1 cm thick Vqetetirn ~rofile ~kta wq/l a x 1 12 BS M B P s C/R cam Typeb kguntC bscd pR CS Caw XO* Ei.- KT (%) (XI (%) (%) (%) ratio ratio

C-P 90.6

C - U 100.0

n-R 88 .O

E-n 100.0

A-n 100.0

v-S 100.0

L-T LOO. 0

W-A 72.0

A-C 100.0 85 .O 100.0

R-P 22.6

L-P 88.9

P-v 100.0 62.8 10.0 rrr-n 100.0 62.4 100.0 APPENDIX L: Plant species list

The following species were collected from sample plots or the surrounding area within the park. Voucher specimens are deposited in the

Simon Fraser University Herbarium (SFUV).

Nomenclature of the vascular species generally follows Hulten (1968).

Exceptions were due to considerations based on Argus (1973)) Hitchcock et al. (1955-1969)) Hubbard (1969)) Moss (1959)) Szczawinski (1959, 1962)) or

Taylor (1963, 1966, 1973). Bryophyte nomenclature was based on Crwn et al.

(1973) and nomenclature for lichens follows Hale (1969).

Vascular Taxa

Abies lasiocarpa (Hook.) Nutt.

Achillea millefolium L. ssp. lanulosa (Nutt.) Piper v. lanulosa

Actaea rubra (Ait.) Willd. ssp. rubra

Adoxa moschatellina L.

Agropyron caninum (L.) Beauv. ssp. majus (~asey)Hitch.

v. andinum (Scribn. and Smith) Hitch.

v. latiglume (Scribn. and Smith) Hitch.

v. majus

Agrostis scabra Willd.

Alnus crispa (Ait.) Pursh ssp. sinuata (Regel) Hult.

Alnus incana (L.) Moench. ssp. tenuifolia (~utt,) Breitung

Amelanchier alnifolia Nutt.

Andromeda polifolia L.

Anemone parviflora Michx. v. grandiflora Ulbr.

Antennaria pulcherrima (Hook.) Greene Apocynum androsaemifolium L.

Aquilegia brevistyla Hook.

Arabis glabra (L.) Bernh.

Arabis hirsuta (L.) Scop. v. pynocarpa (Hopkins) Rollins

Arabis holboellii Hornem. v. retrofracta (Graham) Rydb.

Aralia nudicaulis L.

Arctostaphylos rubra (Rehd. and Wilson) Fern.

Arctostaphylos uva-ursi (L.) Spreng. v. adenotricha Fern. and Macbr.

Arnica chamissonis Less. ssp. foliosa (Nutt.) Maguire

Artemesia campestris L. ssp. borealis (Pall.) Hall and Clem.

v. purshii (Bess.) Cronq.

Asplenium viride Huds.

Aster ciliolatus Lindl.

Aster junciformis Rydb.

Aster modestus Lindl.

Astragalus eucosmus Robins ssp. eucosmus

Barbarea orthoceras Ledeb.

Betula glandulosa Michx.

Betula papyrifera Marsh. v. humilis (Regel) Fern.

v. cornrnutata (Regel) Fern.

Botrychium lanceolatum (Gmel.) Angstr.

Botrychium virginianum (L.) Swartz

Bromus ciliatus L.

Bromus inermis Leyss.

Calamagrostis canadensis (Michx.) Beauv. ssp. canadensis

Calamagrostis inexpansa Gray

Calypso bulbosa (L.) Oakes campanula aurita Greene

Cardamine pennsylvanica Muhl.

Carex aquatilis Wahlenb. ssp. aquatilis

Carex capillaris L.

Carex concinna R. Br.

Carex diandra Schrank

Carex dioica L. ssp. gynocrates (Wormsk.) Hult.

Carex eburnea Boott

Carex garberi Fern. v, bifaria Fern.

Carex interior Bailey

Carex lanuginosa Michx.

Carex leptalepa Wahlenb.

Carex microglochin Wahlenb.

Carex oederi Retz. ssp. viridula (Michx.) Hult.

Carex praegracilis Boott

Carex rostrata Stokes

Carex vagina ta Tausch

Castilleja raupii Pennell

Chenopodium album L.

Chenopodium capitatum (L.) Aschers.

Chenopodium hybridum L. ssp. giganthospermum (Aellen) Hult.

Cicuta douglasii (DC.) Coult. and Rose

Cinna latifolia (Trev.) Griesb.

Circaea alpina L.

Corallorhiza trifida Chat.

Cornus canadensis L.

Cornus stolonifera Michx. Corydalis aurea Willd.

Corydalis sempervirens (L.) Pers.

Cyndg~ossumboreale Fern.

Cypripedium calceolus L. v, parviflorum (Salis.) Fern.

Cypripedium passerinurn Richards.

Delphinium glaucum S. Wats.

Dracocephalum parviflorum Nutt.

Drosera anglica Huds.

Drosera rotundifolia L.

Dryas drununondii Richards

Dryas integrifolia Vahl ssp. integrifolia

Dryopteris austriaca (Jacq.) Woynar

Elaeagnus commutata Bernh.

Eleocharis pauciflora (Lightf.) Link.

Elymus glaucus Buckl. v. glaucus

Elymus innovatus Beal

Empetrum nigrum L.

Epilobium adenocaulon Haussk.

Epilobium angustifolium L.

Epilobium latifolium L.

Epilobium leptophyllum Raf.

Epilobium palustre L.

Equisetum arvense L.

Equisetum hiemale L. v. pseudohiemale (Farw.) Morton

Equisetum pratense Ehrh.

Equisetum scirpoides Michx.

Equisetum variegatum Schleich. ssp. variegatum Erigeron acris L. ssp. politus (E. Fries) Schinz and Keller

Erigeron philadelphicus L.

Erysimum cheiranthoides L. ssp. altum Ahti

Fragaria virginiana Duchesne v. glauca Wats.

Galium boreale L.

Galium trifidum L.

Galium triflorum Michx.

Geocaulon lividum (Richards.) Fern.

Geranium bicknellii Britt.

Geum macrophyllum Willd. ssp. perincisum (Rydb.) Hult.

Glyceria striata (Lam.) Hitch. ssp. stricta (Scribn.) Hult.

Goodyera repens (L.) R. Br. v. ophioides Fern.

Gymnocarpium dryopteris (L.) Newm.

Habenaria dilatata (Pursh.) Hook.

Habenaria hyperborea (L.) R. Br.

Habenaria obtusata (Banks ex Pursh.) Richards.

Habenaria orbiculata (Pursh.) Torr.

Hedysarum mackenzii Richards

Heracleum lanatum Michx.

Hieracium umbellatum L.

Juncus alpinus Vill.

Juncus arcticus Willd. ssp. ater (Rydb.) Hult.

Juncus nodosus L.

Juniperus communis L. ssp. nana (Willd.) Syme

Juniperus horizontalis Moench

Kobresia simpliciuscula (Wahlenb.) Mack.

Lactuca biennis (Moench) Fern. 192 Lappula myosotis Moench

Larix laricina (Du Rot) K. Koch v. alaskensis (Wight) Raup

Lathyrus ochroleucus Hook.

Ledum groenlandicum Oeder

Linnaea borealis L. v. americana (Forbes) Rehd.

Listera borealis Morong

Listera cordata (L.) R. Br.

Lobelia kalmii L.

Lonicera dioica L. v. glaucescens (Rydb.) Butters

Lycopodium annotinum L. ssp. annotinum

Lycopodium complanatum L.

Lycopodium obscurum L. v. dendroideum (Michx.) D. C. Eaton

Lycopus unif lorus Michx.

Maianthemum canadense Desf. v. interius Fern.

Matteuccia struthiopteris (L.) Todaro

v. pensylvanica (Willd.) Morton

Melilotus alba Desr.

Melilotus officinalis (L.) Lam.

Mentha arvensis L.

Mertensia paniculata (Ait.) G. Don. v. paniculata

Mimulus guttatus DC.

Mitella nuda L.

Moneses uniflora (L.) Gray

Muhlenbergia glomerata (Willd.) Trin. v. cinnoides (Link) Hem.

Oplopanax horridum (Smith) Miq.

Orchis rotundifolia Banks

Oryzopsis asperifolia Michx.

Osmorhiza depauperata Phil. Oxycoccus microcarpus Turcz.

Parnassia palustris L. ssp. neogaea (Fern.) Hult. Petasites palmatus (Ait.) Gray

Phalaris arundinacea L.

Picea glauca (Moench) Voss

Picea mariana (Mill.) BSP.

~inguiculavulgaris L.

Pinus contorta Loud. v. latifolia Engelm.

Plantago major L. v. pilgeri Domin

Poa pratensis L.

Poa trivialis L.

Populus balsamifera L. ssp. balsamifera

ssp. trichocarpa (Torr. and Gray) Hult.

Populus tremuloides Michx.

Potamogeton filiformis Pers. ' Potamogeton pectinatus L. Potentilla fruticosa L.

Potentilla norvegica L. ssp. monspeliensis (L.) Aschers and Graebn.

Primula mistassinica Michx.

Prunus pennsylvanica L. f.

Prunus virginiana L.

Pyrola asarifolia Michx.

Pyrola chlorantha Sw. v. occidentalis Gray

Pyrola secunda L.

Ranunculus abortivus L.

Ranunculus macounii Britt. v. mco-mii

Rhododendron lapponicum (L.) Wahlenb. Ribes glandulosum Grauer

Ribes hudsonianum Richards.

Ribes lacustre (Pers.) Poir.

Ribes oxyacanthoides L.

Ribes triste Pall.

Rosa acicularis Lindl.

Rosa woodsii Lindl.

Rubus arcticus L. ssp. acaulis (Michx.) Focke

Rubus idaeus L. ssp. melanolasius (Dieck) Focke

Rubus parviflorus Nutt.

Rubus pedatus Sm.

Rubus pubescens Raf.

Salix alaxensis (Anderss.) Cov. v. longistylis (Rydb.) Schneid.

Salix arbusculoides Anderss, v. puberula Anderss.

Salix athabascensis Raup

I Salix barclayi Anderss. v. barclayi

Salix bebbiana Sarg,

Salix brachycarpa Nutt. ssp. niphoclada (Rydb.) Argus

Salix candida Fluegge ex Willd.

Salix glauca L.

Salix monticola Bebb

Salix novae-angliae Anderss.

Salix reticulata L. ssp. reticulata

Sanicula marilandica L.

Scirpus caespitosus L.

Scirpus validus Vahl.

Selaginella selaginoides (L.) Link Senecio pauciflorus Pursh

Senecio pauperculus Michx.

Shepherdia canadensis (L.) Nutt.

Smilacina stellata (L.) Desf.

Smilacina trifolia (L.) Desf.

Solidago canadensis L. v. salebrosa (Piper) Jones

v. subserrata (DC,) Cronq.

Solidago multiradiata Ait. v. scopulorum Gray

Sonchus uliginosus Bieb.

Sorbus scopulina Greene

Sphenophous intermedia (Rydb.) Rydb.

Spiranthes romanzoffiana Cham.

Stellaria longipes Goldie v. altocaulis (Hulten) Hitch.

Strepropus amplexifolius (L.) DC. / Taraxacum ceratophorum (Ledeb.) DC.

Taraxacum scanicum Dahls.

Tofieldia glutinosa (Michx.) Pers.

Trifolium hybridum L.

Triglochin maritimum L.

Triglochin palustris L.

Typha latifolia L.

Urtica gracilis Ait.

Urtica lyallii S. Wats.

Utricularia intermedia Hayne

Utricularia vulgaris L. ssp. macrorhiza (Le Conte) Clausen

Vaccinium alaskaense Howell

Vaccinium uliginosum L. ssp. alpinum (Bigel.) Hult. 196

Vaccinium vitis-idaea L. ssp. minus (Lodd.) Hult.

Veronica americhna Schwein.

Viburnum edule (Michx.) Raf.

Vicia americana Muhl.

Viola adunca Sm.

Viola canadensis L. v, rugulosa (Grenne) Hitch.

Viola nephrophylla Greene

Viola renifolia Gray v. brainerdii (Greene) Fern.

Viola selkirkii Pursh.

Non-vascular Taxa

Alectoria sp.

Amblystegium serpens (Hedw.) B.S.G.

Aneura pinguis (L.) Dumort.

Aulacomnium ~aiustre(Hedw.) Schwaegr.

Brachythecium erythrorrhizon B.S.G.

Brachythecium frigidum (C. Muell.) Besch.

Brachythecium salebrosum (Web. and Mohr.) B.S.G.

Brachythecium starkeii (Brid.) B.S.G.

Bryum capillare Hedw.

Calypogeia sp.

Campylium chrysophyllum (Brid.) J. Lange

Campylium stellaturn (Hedw.) C. Jens.

Cetraria pinastri (Scop.) S. Gray

Chara vulgaris L.

Cladonia chlorophaea (Flk.) Spreng.

Cratoneuron commutatum (Hedw.) Roth v. falcatum (Brid.) Moenk. Cratoneuron filicinum (Hedw.) Spruce

Dicranum fuscescens Turn.

Dicranum tauricum Sapeh.

Ditrichum flexicaule (Schwaegr.) Hampe

Drepanocladus sendtneri Monk.

Drepanocladus uncinatus (Hedw.) Warnst.

Drepanocladus vernicosus (Lindb.) Warnst.

Eurhynchium pulchellum (Hedw.) Jenn.

Evernia mesomorpha Nyl.

Hylocomium splendens (Hedw.) B.S.G.

Hypnum lindbergii Mitt.

Hypogymnia physodes (L.) Nyl.

Leskeela nervosa (Brid.) Loeske

Lophozia longidens (Lindb.) Macoun

Mnium marginatum (With.) Brid. ex P. Beauv.

Orthotrichum obtusifolium Brid.

Parmelia sulcata Tayl.

Peltigera aphthosa (L.) Willd.

Philonotis fontana (Hedw.) Brid.

Plagiochila aspleniodes (L.) Dumort.

Plagiomnium cuspidatum (Hedw.) Koponen

Plagiomnium drurmnondii (Bruch. and Schimp.) Koponen

Plagiomnium ellipticum

Pleurozium schreberi (Brid.) Mitt.

Pohlia nutans (Hedw.) Lindb.

Ptilidium ciliare (Web.) Hampe

Ptilidium pulcherrimum (Web.) Hampe Ptilium crista-castrensis (Hedw.) De Not.

Pylaisiella polyantha (Hedw.) Grout

Rhizomnium puHctatum (Hedw.) Koponen

Riccardia sp.

Riccia fluitans L.

Sphagnum sp.

Tetraphis pellucida Hedw.

Tomenthypnum nitens (Hedw.) Loeske

Tortella fragilis (Drumrn.) Limpr.

Tortella tortuosa (Hedw.) Limpr.

Tortula princeps De Not.

Usnea comosa (Ach.) Ach. B IBLIOGRAPHY

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