"Science, Faculty of"@en . "Botany, Department of"@en . "DSpace"@en . "UBCV"@en . "Wade, Leslie Keith"@en . "2011-09-20T21:44:31Z"@en . "1965"@en . "Master of Science - MSc"@en . "University of British Columbia"@en . "The Sphagnum bogs of the Tofino-Ucluelet area of the western coast of Vancouver Island were studied from vegetational, edaphic, and historical aspects. An integrated approach to these three aspects was attempted in order to give in a relatively limited time as complete a picture as possible of the bog ecosystem.\nThe bog vegetation was studied on 110 sample plots using analytical and synthetic methods of the Zurich-Montpellier school of phytosociology. Ten different vegetation types were described and characterized, nine belonging to the bog ecosystem and one to the surrounding scrub forest. The nine bog vegetation types consist of five distinct associations and one association composed of five variants. The vegetation types studied are summarized below, in order of increasing floristic complexity. \nLow moor bog associations:\n1. Caricetura pluriflorae\n(Carex plurlflora association)\n2. Scirpeto-Sphagnetum mendoclnl\n(Sclrpus caespitosus - Sphagnum mendocinum association)\n3. Oxycocceto-Sphagnetum papillosi \n(Oxycoccus quadripetalus - Sphagnum papillosum association)\nHigh moor bog association:\n4. Ledeto-Sphagnetum caplllacei\n(Ledum groenlandicum - Sphagnum capillaceum association)\nPeripheral bog associations: (Bog-forest transition) \n5. Pineto-Sphagnetum capillacei\n(Pinus contorta - Sphagnum capillaceum association)\na. Pineto-Sphagnetum capillacei sphagnosum papillosi\n(Pinus contorta hummock variant)\nb. Pineto-Sphagnetum capillacei myricosum galis\n(Myrica gale variant)\nc. Pineto-Sphagnetum capillacei chamaecyparosum nootkatensis\n(Chamaecyparis nootkatensis variant)\nSecondary succession variants established after fire:\nd. Pineto-Sphagnetum capillacei vacciniosum vitis-idaeae\n(Vaccinium vltis-idaea variant)\ne. Pineto-Sphagnetum capillacei vacciniosum parvifolii\n(Vaccinium parvifollum variant) Scrub forest association surrounding bogs:\n6. Pineto-Chamaecypareto-Sphagnetum recurvi\n(Pinus contorta - Chamaecyparis nootkatensis - Sphagnum fecurvum association) (Bog forest)\nEdaphic considerations were limited to the analysis of soils from representative sample plots of each association and variant. Soils were analyzed for available cations, including Ca\u00E2\u0081\u00BA\u00E2\u0081\u00BA, Mg\u00E2\u0081\u00BA\u00E2\u0081\u00BA, Na\u00E2\u0081\u00BA, K\u00E2\u0081\u00BA, adsorbed phosphate, total nitrogen, cation exchange capacity, percent base saturation pH, and soil moisture. The results of the soil analyses were wherever possible correlated with trends in the development of plant associations. Climatic factors were regarded as constant over so limited an area as the one under study.\nHistorical considerations included a pollen analysis from a representative core in the center of the major study bog, and a radiocarbon dating to determine the age of a representative bog. The results of the pollen analysis appeared to confirm previous ideas that the bog did not develop from a lake, but rather it developed from a wet seepage forest habitat. The radiocarbon dating indicated the age of the bog at only 390\u00C2\u00B1 90 years B.P., thus explaining partially the apparent very juvenile phase of the bogs of the area.\nThe general hypothesis is suggested that the distribution of the bog plant associations is primarily dependent upon a complex of environmental factors that are dependent upon topography."@en . "https://circle.library.ubc.ca/rest/handle/2429/37514?expand=metadata"@en . "VEGETATION AND HISTORY OF THE SPHAGNUM BOOS OF THE TOFINO AREA, VANCOUVER ISLAND by LESLIE KEITH WADE B.Sc. The University of B r i t i sh Columbia 1963 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n the Department of BOTANY We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA September 196$ In presenting th i s thes i s in p a r t i a l f u l f i lmen t of the requirements for an advanced degree at the Un ivers i ty of B r i t i s h Columbia, I agree that the L ibrary sha l l make i t f r ee l y ava i l ab l e fo r reference and study. I fur ther agree that per-mission for extensive copying of t h i s thes i s for scho la r l y purposes may be granted by the Head of my Department or by his representatives., It is understood that copying or p u b l i -ca t ion of t h i s thes i s for f i n a n c i a l gain sha l l not be allowed without my wr i t ten permiss ion. Department of The Un ivers i ty of B r i t i s h Columbia Vancouver 8, Canada pate z y SEPT. / i t s Abstract The Sphagnum bogs of the Tofino-Ucluelet area of the western coast of Vancouver Island were studied from vegetational, edaphic, and h i s tor ica l aspects. An integrated approach to these three aspects was attempted i n order to give i n a re lat ive ly l imited time as complete a picture as possible of the bog ecosystem. The bog vegetation was studied on 110 sample plots using analytical and synthetic methods of the Zurich-Montpellier school of phytosociology. Ten different vegetation types were described and characterized, nine belonging to the bog ecosystem and one to the surrounding scrub forest. The nine bog vegetation types consist of f ive dist inct associations and one association composed of f i ve variants. The vegetation types studied are summarized below, i n order of increasing f l o r i s t i c complexity. Low moor bog associations: 1. Caricetura p lur i f lorae (Carex p lu r l f l o ra association) 2. Scirpeto-Sphagnetum mendoclnl (Sclrpus caespitosus - Sphagnum mendocinum association) 3. Oxycocceto-Sphagnetum pap i l los i ' -\u00E2\u0080\u00A2 (Oxycoccus quadripetalus - Sphagnum papillosum association) High moor bog association* U* Ledeto-Sphagnetum capl l lacel (Ledum groenlandlcum - Sphagnum capillaceum association) Peripheral bog associations! (Bog-forest transition) 5>. Pineto-Sphagnetum capi l lacei (Pinus contorta - Sphagnum capillaceum association) a. Pineto-Sphagnetum capi l lace i sphagnosum pap i l lo s i (Pinus contorta hummock variant) b. Pineto-Sphagnetum capi l lacei myricosum galis (Myrica gale variant) c. Pineto-Sphagnetum capi l lace i chamaecyparosum nootkatensis (Chamaecyparis nootkatensis variant) Secondary succession variants established after f i r e * d. Pineto-Sphagnetum capi l lacei vacciniosum vit is- idaeae (Vaccinium v l t i s - idaea variant) e. Pineto-Sphagnetum capi l lacei vacciniosum pa rv i f o l i i (Vaccinium parvifollum variant) Scrub forest association surrounding bogs: 6. Pineto-Chamaecypareto-Sphagnetum recurvi (Pinus contorta - Chamaecyparis nootkatensis - Sphagnum fecurvum association) (Bog forest) Edaphic considerations were l imited to the analysis of so i l s from representative sample plots of each association and variant. Soils were analyzed for available cations, including Ca ' f + , Mg + + , Na + , K*, adsorbed phosphate, tota l nitrogen, cation exchange capacity, percent base saturation pH, and s o i l moisture. The results of the s o i l analyses were wherever possible correlated with trends i n the development of plant associations. Climatic factors were regarded as constant over so l imited an area as the one under study. Histor ica l considerations included a pollen analysis from a 112 J representative core in the center of the major study bog, and a radio-carbon dating to determine the age of a representative bog. The results of the pollen analysis appeared to confirm previous ideas that the bog did not develop from a lake, but rather i t developed from a wet seepage forest habitat. The radiocarbon dating indicated the age of the bog at only 390- 90 years B.P., thus explaining partially the apparent very juvenile phase of the bogs of the area. The general hypothesis is suggested that the distribution of the bog plant associations is primarily dependent upon a complex of environmental factors that are dependent upon topography. IV, TABLE OF CONTENTS Chapter I II III 17 INTRODUCTION DESCRIPTION OF THE AREA . .... A. Topography and vegetation . B. Climate C. Geology . . . . . . . . . . METHODS . . . . A. General . . . B. Reconnaissance C. Plot selection and analysis D. Mensuration E. Synthesis \u00E2\u0080\u00A2 . F. Soil sampling and analysis. 1. Field procedure . . . . 2. Analysis . G. Pollen analysis H. Radiocarbon dating DESCRIPTION OF THE ASSOCIATIONS A. Caricetum pluriflorae (Carex pluriflora association), B. Scirpeto-Sphagnetum mendocini (Scirpus caespitosus - Sphagnum mendocinum association) . . . . . . . C. Qxycocceto-Sphagnetum papillosi (Oxycoccus quadripetalus - Sphagnum papillosum association) . . . . D. Ledeto-Sphagnetum capillacei (Ledum groenlandicum - Sphagnum capillaceum association) . . .. . . . . . . . . . . . . Page 1 .. 1 ;, l 7 9 11 11 11 11 13 13 15 15 15 16 16 18 19 21 23 28 v: Chapter ' Page E. Pineto-Sphagnetum capillacei (Pinus contorta - Sphagnum capillaceum association). .. 31 1. Pineto-Sphagnetum capillacei sphagnosum papillosi (Pinus contort a hum-nock variant) 33 ' 2. Pineto-Sphagnetum capillacei myricosum galis (Pinus contorta - Sphagnum capillaceum - Myrica gale variant) . . . . . . . . 37 3. Pineto-Sphagnetum capillacei chamaecyparosum nootkatensis (Pinus contorta - Sphagnum capillaceum -Chamaecyparis nootkatensis variant) . . . . . . . 39 U . Pineto-Sphagnetum capillacei vacciniosum vitis-idaeae (Pinus contorta - Sphagnum capillaceum -Vaccinium vitis-idaea variant $. Pineto-Sphagnetum capillacei vacciniosum parvifolii (Pinus contorta - Sphagnum capillaceum -Vaccinium parvifolium v a r i a n t ) . 7 U8 F. Pineto-Chamaecypareto-Sphagnetum recurvi (Pinus contorta - Chamaecyparis nootkatensis -. Sphagnum recurvum association) (Bog forest) . . . . . . . . . . . . . . . . . . . . . \u00C2\u00A3l V SOILS 58 A. Soil moisture . $9 B. Total nitrogen 60 C. Available sodium . .. 60 D. Available potassium. . . . . . . . . . . . . . . . . . . 60 E. Available, calcium . 60 F. Available magnesium. . . . . . . . . . . . . . . . . . . 60 G. Adsorbed phosphate . . . . . . . . . . . . . . . .. - 61 H. Cation exchange capacity . . . . . . . . . . . . . . . 6 1 I. Percent base saturation.c;. . . . . . . . . . . . . . . 61 J. pH . . . ' 61 K. Discussion of soils * . . . . . . . 61 Chapter Page VT LIFE FORMS 6h VII HISTORY 67 VIII POLLEN ANALYSIS \u00E2\u0080\u00A2 70 DC SUCCESSIONAL TRENDS. . . . . . . . . 76 A. Regional level 76 B. Association level. 78 X SUMMARY AND CONCLUSIONS 83 1. BIBLIOGRAPHY 87 2. BIBLIOGRAPHY OF PUBLICATIONS USED IN THE IDENTIFICA-TION OF VASCULAR PLANTS 90 3 . BIBLIOGRAPHY OF PUBLICATIONS USED FOR IDENTIFICATION OF BRYOPHYTES AND LICHENS 91 APPENDIX I Check l i s t of plant species 93 APPENDIX II Explanation and Legend for Synthesis tables Synthesis tables I - VI . . . 97 APPENDIX III Soils data . . . . . . . . . . . . . . . ........ . 118 APPENDIX IV Radiocarbon dating 12U v i i LIST OF FIGURES Figure .. Page 1. View of interior of study bog, May 2 2. View of interior of study bog, August 2 3. Profile of terrace U lx. Climatic data 8 3>. Bog profile . . . l8a 6. Sketch map of study bog, showing distribution of associations 18b 7. Carex pluriflora association surrounded by Oxycoccus quadripetalus - Sphagnum papillosum association . . . . . 20 8. Juncus oreganus in deepest water of Carex plurlflora association ^ 20 9. Sclrpus caespitosus - Sphagnum mendocinum 2U 10. Sphagnum capillaceum and S. papillosum invading Scirpus caespitosus - Sphagnum mendocinum association 2k 11. Gentiana douglasiana in low moor bog association 26 12. Oxycoccus quadripetalus in Oxycoccus quadripetalus - -Sphagnum papillosum association . 26 13. Mosaic community of Pinus contorta hummock variant and Oxycoccus quadripetalus - Sphagnum papillosum association . . . . . . 27 l l i . Trientalis arctica, Linnea borealis, and Cornus canadensis growing in Pinus contorta hummock varia n t . . . 27 15. High moor Sphagnum fuscum hummock with associated species 31 16. Apargidium boreale growing on Sphagnum fuscum hummock . . 31 17. Rhacomitrium lanuginosum, Empetrum nigrum, and Kalmia polifolia growing on dry hummock surrounded by Carex pluriflora association. . . . . . . 3\u00C2\u00A3 18. Pinus contorta hummock variant surrounded by low moor bog association . . . . . . . . . . . . . . . 35 19. Myrica gale variant . 38 Figure Page 20* Thuja plicata and Pinus contorta growing in mature high moor community 38 21. Characteristic form of old Tsuga heterophylla in Long Beach area bogs . . I l l 22. Pinus contorta, Tsuga heterophylla and Thuja plicata at edge of bog forest hi 23 . Secondary successional communities on burned area of bog. Open area represents Vaccinium vitis-idaea variant. . . . U3 2iu Boschniakia hookeri parasitic on Gaultheria shallon of Vaccinium vitis-idaea variant h3 25. Ledum groenlandicum, dominant shrub of Ledum groenlandi-cum - Sphagnum capillaceum association hh 26. Kalmia polifolia, co-dominant shrub of Vaccinium v i t i s - idaea variant . . . i . . . hk 27. Coptis trifoliata and Empetrum nigrum growing in Vaccinium vitis-idaea variant i . Ii5 28. Nephrophyllidium crista-galli growing in wet depressions of burned area of bog . U5 29 . View of bog forest, Pinus contorta predominating. . . . . 52 3 0 . View of bog forest, with Chamaecyparis nootkatensis, Thuja plicata, Tsuga heterophylla, and Pinus contorta . . 52 3 1 . Characteristic form of old Pinus contorta in bog forest of Long Beach area (height 75 feet, age 260 years). . . . 53 3 2 . Life form distribution 66 3 3 . Pollen profile 75 3U. Successional trends . 81 IX. Acknowledgement I wish to express sincere thanks to Dr. 7. J . Krajina for providing the i n i t i a l stimulus for this study, and for his constant help and guidance throughout i t s completion. His help i n the ident i f icat ion of many vascular plants, bryophytes and lichens i s also greatly appreciated. Grateful thanks are also extended to Dr. G. E. Rouse, who provided generous assistance with the pollen analysis, geology, and radiocarbon dating aspects of the thesis, and to Dr. W. B. Schofield who read the manuscript and gave valuable advice* I wish also to thank the National Research Council, Ottawa, for financing this study, and Dr. T. M. C. Taylor and Dr. G. H. N. Towers for providing study f a c i l i t i e s i n the Department of Botany at the University of B r i t i sh Columbia. Last ly, I am grateful to my fellow students, Richard T. Kuramoto for assistance i n the f i e l d and for helpful advice, Robert C. Brooke for valuable discussions and advice, and to my fiancee, Adele C. Templeton, for her many hours of work on a l l aspects of the thesis. CHAPTER I INTRODUCTION . . Sphagnum bogs have long been a popular subject for ecological studies, probably because they represent ecosystems that are in many aspects totally different from any otherj ecosystems that are unique among temperate climate plant communities. Because of prevailing extreme edaphic conditions, bogs display entirely different vegetational aspects in comparison to other eco-systems. Perhaps more than many other types of ecosystems bogs appear as sharply defined units, having within their confines many plant and animal species as well as many peculiar edaphic conditions that are otherwise absent from large regions. In addition, the physiogonomy of bogs is usually quite unique for most areas. It has also long been thought that Sphagnum bogs exhibit notable affinities with subarctic areas, particularly with regard to the interesting question of subarctic and boreal relics. Although Sphagnum bogs have been very extensively studied from many aspects in Great Britain and continental Europe, relatively l i t t l e detailed work exists for North America, particularly western North America. The bulk of the many bog studies that have been done in western North America consist of investigations into the stratigraphy of bogs, pollen analyses, and li s t s of observed plant species. Many studies, especially those of Rigg, Hansen, and Heusser, are comparative studies of general bog features over extensive areas. While these studies convey a clear picture of the patterns of bog distribution and the general bog types for coastal western North America, they seldom attempt to investigate bog plant associations or successional sequences between actual associations. As a result relatively l i t t l e is known of the detailed patterns of bog vegetation in this area. A Swedish 2 worker (Osvald, 1933) attempted a br ief vegetational study of the bogs of southwestern mainland Br i t i sh Columbia, and established tentative socia-tions for several major bogs of the area. Subsequently Krajina (195>9) and Orloci (I960) have described several bog associations i n the course of regional vegetation studies. The purpose of this thesis i s primarily to present i n deta i l the vegetation of the Sphagnum bogs of a def inite l imited area, and thereby to establish a basic series of bog associations which can be used as a comparison for future bog studies i n other areas. The Tofino-Ucluelet region was chosen as a study location because of i t s many readily accessible, yet re lat ive ly undisturbed bogs. The problem of the bizarre forest of dwarf conifers that also exists i n the bog areas provided an additional highly intr iguing aspect. In addition to a detailed c lass i f icat ion of the bog associations, an important aspect of the study included the correlation of the plant associations with environmental data, and the integration of an h i s tor ica l approach i n the form.of a pollen analysis and radiocarbon dating. ' . CHAPTER II DESCRIPTION OF THE AREA A. Topography and Vegetation Between the villages of Ucluelet and Tofino on the western coast of Vancouver Island exists a broad undulating terrace that appears to have been formed at least partially as a result of post-glacial land uplift. The terrace reaches its best development in the Long Beach area in the vicinities of WLckanninish and Wreck Bays, where i t averages approximately seventy feet above sea-level. The surface of the terrace is characterized by a gentle topography of very low hills and broad shallow valleys, extensive flat areas, and a total absence of any sharp relief features (Fig. 3). The vegetation of the terrace is of four very distinct types, the distribution of three of which appears to be controlled at least indirectly by topography. These three are the Sphagnum bogs, the scrub forest of the valleys, and the climax western hemlock-amabilis f i r forest of the low h i l l s . The distribution of the fourth type, the Sitka spruce forest, appears to be controlled by the proximity of the open ocean. This latter type consists of a pure Sitka spruce (Picea sitchensis) forest that lies immediately adjacent to the open ocean, and occupies, in most cases, the bank of the terrace as well as a narrow strip along the surface of the terrace. The trees of this strip rise between 100 and 120 feet in height above a dense shrub layer of salal (Gaultheria shallon). Nowhere is this strip of pure Sitka spruce more than a few hundred feet wide, and nowhere is i t found except fronting the open ocean, the inference being that its distribution is in some manner controlled by the presence of the open ocean. 2 Inland from the belt of Sitka spruce the prof i le of the terrace generally slopes s l i ght ly downward, leaving a l i p of s l i ght ly higher land that appears to effect ively block drainage from many areas of the terrace. This portion of the terrace i s occupied by three vegetational types the distr ibution of which i s controlled by drainage, which i n turn i s at least par t i a l l y controlled by topography. The lowest parts of the region are shallow basin- l ike areas f i l l e d with shallow Sphagnum bogs, very similar i n character to the muskegs found along the coastal areas to the north (Heusser I960). These bogs are very frequent on parts of the terrace, part icular ly i n the v i c i n i t y of Wreck Bay and Wickanninish Bay. Most of the area between the Sphagnum bogs i s densely covered by a bizarre scrub forest composed of dwarf conifers of several species. Pre-dominant among these are curiously shaped forms of lodgepole or shore pine (Pinus contorta), yellow cedar (Chamaecyparis nootkatensis), and western hemlock (Tsuga heterophylla). To a lesser extent are also found western red cedar (Thuja p l l cata ) , western yew (Taxus brev i fo l i a ) , and occasionally western white pine (Pinus monticola). Bog species are also usually present, part icular ly Ledum groenlandicum and Sphagnum recurvum, prompting the name \"bog forest\" to be adopted for this vegetational type. Trees of this bog forest are seldom over seventy feet i n height. The f i n a l vegetation type i s the normal climax forest of the region, the western hemlock-amabilis f i r forest typ ica l of the wetter subzone of the Western Hemlock Zone (Krajina, 1959; Or loci , 1961). This vegetation type i s restr icted to the higher parts of the terrace and generally occurs above the scrub or bog forest, probably indicating better drainage as a probable distr ibutional factor. Besides including western hemlock and amabilis f i r a high percentage of western red cedar i s commonly present, while within a half mile of the open coast amabilis f i r i s usually completely lacking. PROFILE OF TERRACE Sitka Spruce of terrace Bog Forest Beach i Sands and gravels Glacial marine til l Western Hemlock Amabilis F i r Bog Upper Triassic volcanics Figure 3. Profile of terrace. Although the climax forest i s rapidly being destroyed by logging, a fen excellent stands remain, with trees up to l \u00C2\u00A3 0 and 200 feet i n height. The climax forest meets and merges with the Sitka spruce forest along many parts of the terrace. The transit ional region, however, i s very narrow. In other areas the bog forest borders the Sitka spruce s t r i p , the transit ion again being narrow and well defined. Apart from these four vegetational types, variations occur as secondary suocessional stages on burned and logged areas, and as alluvium colonists along stream banks, where red alder (Alnus rubra) i s often dominant and Sitka spruce i s common. The bogs of the area served as the main theme of the present study, although, to a lesser extent, the bog forest also received attention. Area of the bogs throughout the area varies from about two acres to approximately eighty acres; aer ia l photographs show the presence of much larger bogs on some of the islands of the v i c i n i t y , notably on Varges Island near Tofino. A l l of the observed bogs appear to have reached only juvenile stages, and evidence of mature stages, such as i s indicated by raised or high moor conditions, i s very l imited. The bulk of the bog surface i s composed of Sphagnum species and herbaceous plants such as Carex, Rhynchospora, Juncus, and Scirpus, many of which are typical of early stages i n bog succession (Rigg, 1925; Hansen, 191*0). The predominant Sphagnum species are Sphagnum papillosum, Sphagnum recurvum, and Sphagnum mendocinum, a l l of which are -characteristic of topogenoua or low moor bogs (Heusser, I960). Portions of the bog are completely submerged fo r a l l except very dry periods i n dry summers, and i n these areas the plant associations closely ref lect the > probable marsh ancestry of the area. Ericaceous shrub development, such as i s usually found i n mature high moor bogs, i s absent except i n l imited areas around the margin of some of 6 the bogs. It exists also as a secondary feature of seme burned portions of the bog. The common bog shrubs of the region, Ledum groenlandicum, Kalmia polifolia, Bmpetrum nigrum, Oxycoccus quadripetalus, and Vaccinium uliginosum, are present on the bog but are of very scattered occurrence and low vigor. In localized areas the initiation of high moor development is indicated by the invasion of Sphagnum capillaceum and Sphagnum fuscum, particularly the latter species. Invasion is followed by the development of a Sphagnum hummock formation. In these areas the bog shrubs, notably Ledum groenlandicum and Oxycoccus quadripetalus, attain better development both in size and quantity than in other parts of the bog. Throughout the entire bog area Pinus contorta is represented by stunted, dwarfed specimens spaced at fairly regular intervals over the bog surface. These trees, seldom over lf> feet in height and often more than 100 years old, support an association of plants on their basal hummocks that is typical of much drier conditions than the bog surface itse l f . For the purposes of this study these stunted trees and their associated hummock plants have been treated as a unit, in many aspects different from the \"inter-hummock\" areas. Chamaecyparis nootkatensis is also present on the bog surface, but attains only the stature of a prostrate spreading shrub, seldom more than 2 to 3 feet high. Towards the bog margin, however, i t grows as a dwarfed tree up to Vy feet in height and often over 100 years in age. Chamaecyparis nootkatensis is the commonest and most characteristic tree \"\". species of the transitional area between bog and bog forest. Juniperus communis var. montana, Thuja plicata, and Tsuga heterophylla are less common on the bog. ' Sections of many of the bogs of the area show definite evidence of past fires, both by the presence of charred logs and stumps, and in some cases by the absence of a nell developed organic layer. The secondary vegetation that \u00E2\u0080\u00A2 . \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 i ' ' ' ' / has developed on these areas shows marked differences in composition and structure from the unbumed parts of the bog. Conditions are much drier and lichens, especially Cladonla species, are well developed on the humus layer. Pinus contorta is the dominant tree species and ranges in its occurrence from dwarfed, isolated individuals to scattered, dense clumps of trees. The bogs are very shallow, the accumulated peat,averaging one meter thick and in exceptional situations extending to two meters in depth. Xn most instances a layer of clay or clay mixed with organic material is evi-dent immediately beneath the peat deposits. Below the clay layer are the sand and gravel outwash deposits that characterize the region. These underlying sands are strongly cemented near their surface and effectively impede drainage of the.bogs. Radiocarbon (C^) dating of a peat sample collected in the basal portion of a typical bog profile indicated the very recent date of 3\u00C2\u00B0oi 90 years B.P. (see Appendix 3). The sample was obtained from a depth of \\ meters, immediately overlying a gray s i l t and sand layer. B. Climate The climate of the area is mild marine humid, designated by the Koppen scale (Koppen and Geiger, 1936) as Cfb (Krajina, 1959). Extremes in temperature are rare, with a mean for the warmest months, July and August, of 58\u00C2\u00B0 F., and a mean for the coolest month, January, of Ul\u00C2\u00B0 F. (Fig. 4). Freezing temperatures are uncommon and as a result, decomposers may be active almost the entire year. The extreme temperatures recorded over an eight year period are 19\u00C2\u00B0 F. and 91\u00C2\u00B0 F. Rainfall is extremely high, with an' annual mean of 120.97\" recorded at the Tofino airport over a nine year period. Of this amount only k*Sn f e l l as snow. 8 Cl imat ic data \u00E2\u0080\u0094 Tofino A i r p o r t op 60 55 50 45 -40 Wean Monthly Temperatures \u00E2\u0080\u00A2 J a n . F e b . March A p r i l May June J u l y A u g . S e p t . O c t . Nov. D e c . Inches p p t . 20 Mean Monthly P r e c i p i t a t i o n J a n . F e b . March A p r i l May June J u l y A u g . S e p t . Annual mean 4 9 \u00C2\u00B0 F Nov. O e c . 15 \u00E2\u0080\u00A2 10 1 1 . 6 9 1 4 . 0 4 1 3 . 2 4 11 .37 4 . 2 4 3 . 3 0 2 . 6 3 3 . 0 9 5 . 0 8 1 3 . 3 9 1 6 . 4 7 1 7 . 4 3 i n c h e s p p t . Annual t o t a l 1 2 0 . 9 7 \" Figure h. Climatic data. Summer, although the driest and warmest period, is relatively cool and \u00E2\u0080\u00A2wet when compared to the summer dry situation that exists in much of British Columbia. Summer fogs are common and often last from daybreak until early afternoon. In summary, the climate is typical of that required to produce temperate rain forest conditions, that well f i t s the coastal Sitka Spruce forests as well as the Western Hemlock-Amabilis Fir climax forests of this region. C. Geology The geology of the Tofino-Long Beach-Ucluelet area has been the subject of very l i t t l e investigation. Dolmage (1920} carried out a study of the area and reported that a formation of considerable extent, composed of well sorted sands, gravels, and clays, formed a uniformly level plain approximately fi f t y feet above sea-level, extending from Ucluelet to^Clayoquot Sound along the coast, and inland ten miles to Kennedy Lake. He speaks of this plain as being of probable marine origin, with no overlying glacial material being present, and of the probability that i t was uplifted following glaciation. According to Dolmage, the gravels, sands, and clays are sorted into thin beds lying horizontally on a rough uneven surface of Upper Triassic volcanic rocks, the Vancouver volcanics. The rocky headlands and islands of the region constitute the only visible parts of this latter formation today, which is prevailingly of dark colored andesite. Observations by Rouse (personal communication, May 1965) and myself suggest the following modifications of this interpretation. Bordering Wreck Bay is a steep, rapidly eroding c l i f f composed of clay, sand, and gravel, which forms the seaward edge of the previously mentioned terrace (Fig. 3). The c l i f f face at the north-western end of Wreck Bay shows two distinct sequences: the lower portion consists of approximately 65 feet of glacial marine t i l l , composed of blue-gray clay with many angular rock fragments, occasional large boulders, and numerous marine shells. This is identical to glacial marine t i l l of the lower Fraser River valley reported by Armstrong (1956j 1957j I960). The upper sequence, which makes up the remaining 10 feet of the terrace at this site, consists of inter-bedded lenses of coarse sand and gravel, the particles of which are a l l smoothly rounded. No marine shells are visible in this latter sequence. This upper sand-gravel sequence thickens noticeably toward the southeast along the beach c l i f f . Distinct cross-bedding is evident, the nature of which indicates a general northerly source direction. The character of the upper sequence, in particular the cross-bedding of the sand lenses, suggests either a glacial outwash or river deposit. In the latter case, emergence of the lower marine sequence above high tide would have occurred before deposition of the outwash or river deposits. Sand and gravel lenses which closely match those of Wreck Bay are to be found immediately below the organic accumulations of the bogs, most of which are located about half a mile inland from Wreck Bay. Bog initiation is very recent, as shown by the Radiocarbon (C^) dating of 390\u00C2\u00B1 90 years B.P., (before present) and probably did not occur until drainage of the area became severely restricted, presumably about 400 years B.P. The formation of a higher l i p of land on the terrace on which the Sitka spruce forest now stands and the formation of a hardpan in the underlying sands were the most likely barriers to drainage (Fig. 3). CHAPTER III METHODS A. General The ecology of the Long Beach area bogs was approached from several viewpoints. The study consisted mainly of an investigation of the plant associations and successional trends occurring in the bogs. In addition, pollen analysis was utilized to partially reconstruct the history of the bog vegetation, in particular that of the i n i t i a l bog stages. Dating of the bog was obtained by means of a radiocarbon (C^) test taken from a basal sample of the bog peat. In this manner an attempt was made to orient the study towards providing as complete a picture as possible of the Long Beach bog ecosystem. The synecological methods used in the study are basically those used ; by Krajina (19^2) and his students in previous ecological studies in British Columbia. B. Reconnaissance Reconnaissance was carried out for short periods in June 1963 and for two weeks in August 1963.L These periods were spent in becoming familiar with the flora of the area and in selecting tentative plant.associations. It was decided after reconnaissance of the many bogs of the area-to-concentrate studies on one representative bog (Fig. 6), and to utilize observations' from surrounding bogs as a basis for comparison. C. Plot selection and analysis One hundred ten plots were analyzed during the spring and summer of 1961*. These plots ranged over ten different tentative plant association types. Each type of vegetation that appeared f l o r i s t i c a l l y homogeneous and that appeared, under similar conditions i n other areas, to repeat i t s e l f was tentatively regarded as an association. Homogeneity was defined on the basis of the regular distr ibution of plant species throughout the area i n question. It was attempted, therefore, to locate a l l plots i n f l o r i s t i c a l l y homogeneous areas. Size of a plot was determined by both the complexity and extent of the association, and ranged from 0.5 m for a f l o r i s t i c a l l y very o simple association without trees or shrubs, to 100 m for the highly complex surrounding bog forest. Analysis of the vegetational aspects of the plots was done using the basic methods of the Zurich-Montpellier school (Braun-Blanquet, 1932} Krajina, 1933} Becking, 1957). For each plot estimates were made of the percent cover occupied by each of the following strata: A. Tree layer A^ - dominant tree species over 40 feet i n height Ag - trees 15 - 40 feet i n height A^ - trees and t a l l shrubs 6 - 1 5 feet i n height B. Shrub layer B^ - shrubs between 6 inches and 6 feet i n height Bg - shrubs under 6 inches i n height . C. Herb layer - a l l herbaceous plants, regardless of height D. Bryophyte and l ichen layer Dh - bryophytes and lichens growing on humus , Ddw- bryophytes and lichens growing on decaying wood E. Epiphyte layer (includes bryophytes, l ichens, and vascular plants) - epiphytes i n the A layer Eg - epiphytes in the B layer EQ - epiphytes in the C layer While this scale varies in its arbitrary limits from that used by other workers (Mueller-Dombois, 1959J Peterson, 196U; Orloci, 1961), i t is felt that this modification enables a better representation to be made of bog conditions. All species in each plot were also rated according to species signifi-cance, sociability, and vigor. The eleven point species significance scale of Krajina and .Domin (1933) as slightly modified by Mueller-Dombois (1959) was used as a measure of abundance and dominance. Sociability, or dispersion, was also rated according to an eleven point scale adopted from Krajina and Domin (1933). Vigor, a rating of the relative vitality of each species, was interpreted on a four point scale. The scales used for species significance, sociability, and vigor are given in Appendix I. D. Mensuration In a l l associations bearing trees, representative specimens of a l l \ tree species were measured for height, diameter at breast height, and age. Height was measured with a Relascope. Age was ascertained by counting the rings of cores collected four feet above ground by an increment borer. Counting was done under a binocular microscope in the laboratory. In many cases the cores or portions of them proved too decayed for accurate age counts. E. Synthesis ~\"; ~ The data from a l l analyzed plots was arranged in groups of tentative communities. Much comparison and correlation was then made to check that sufficient floristic similarity existed between sample plots of a tentative association to ver i fy i t s entity as a dist inct association. Constancy, the frequency with which one species occurs within a group of plots of uniform s ize, was calculated for each species. The degree of constancy was expressed i n the f ive class scale of Braun-Blanquet (1932) as s l i ght ly modified by Brooke ( 1 9 6 5 ) . Average cover value, an expression of dominance, was calculated by converting the species significance values. The conversion scale used i s that of Brooke ( 1 9 6 5 ) , see Appendix I. Vegetation units that were tentatively cal led associations i n the f i e l d were occasionally found after synthesis to be less dist inct than previously thought. These units which differed from each other by only minor, although s igni f icant ly different aspects, were then delegated the rank of association variant, rather than that of association (Domin 1 9 3 6 ) . Each plant association i s defined by i t s characteristic combination of species, a category which includes both constant and characteristic species* Constant species - species present i n more than Q0% of the plots of an association (Peterson, 19610 Constant dominants - constant species of over 10% cover (Peterson, 196U) Constant non-dominants - constant species of less than 10% cover Characteristic species (Braun-Blanquet, 1932) Exclusive species - species completely or almost completely confined to one community Selective species - species found most frequently i n a certain ' community but also, though rarely, i n other communities Preferential species - species present i n several communities \u00E2\u0080\u00A2 1 5 more or less abundantly but predominantly or with better vitality in one certain community. Associations and variants were given the Latinized names common to the proceedings of British and Continental European plant ecologists, a procedure designed to standardize the naming of vegetation units on a world-wise basis. Each association was named using one or more (as neces-sary for sufficient differentiation) names of dominant or otherwise characteristic species (Braun-Blanquet, 1953)* F. Soil Sampling and Analysis 1. Field procedure: Pits were dug in at least two plots of each tentative vegetation type, and soil samples collected for analysis. Although soil profiles were at best poorly developed, effort was made to collect samples from each apparent horizon. For. this reason samples were not always collected at precisely the same depths in pits of the same community type. When horizons were not distinguishable, samples were collected at regular intervals in the pit. -All pits were extended into the underlying sand and a sample collected of the sand. Maximum depth of roots and depth of maximum concentration of roots were noted for each pit. Immediately after collecting, soils were weighed and subsequently dried. 2. Analysis: Soils were oven-dried and weighed. Soil moisture was calculated for each sample using the i n i t i a l field weights and the final oven-dried weights. The pH was measured for each, soil sample collected, using the Beckmann Model N pH meter. Before measuring, each soil sample was mixed with distilled water until a thick paste consistency was reached, after which 16 i t was allowed to stand overnight. Selected soil samples were analyzed for cation-exchange capacity and for exchangeable Na+, K+, Ca + +, and Mg++. The method used was extraction by NH^ OAc using a modified technique developed by the Dept. of Soil Science, University of British Columbia, in which the cations are extracted by centrifugation rather than by buchner funnel. The soils were also analyzed for adsorbed phosphate using the Bray and Kutz method #1 ( 1 9 U 5 ) , and for total nitrogen using the modified Kjeldahl method (Jackson, 1\u00C2\u00B06U). ' Determination of cation exchange capacity, exchangeable cations, and adsorbed phosphate was carried out by the Dept. of Soil Science, University of British Columbia. G. Pollen Analysis - o A core for pollen analysis was collected from the central area of the bog, using a Hiller peat borer. Samples were collected at 10 cm depth intervals from the bog surface to the underlying clay-sand transition area. In the laboratory the samples were boiled in KOH, washed and centri- . fuged, and the residue containing the pollen grains and spores stained with safranin and mounted on slides. Slides from 20 cm intervals through the bog profile were analyzed by counting and identifying 200 pollen grains from each slide. The count of 200 pollen grains per slide has been shown statistically to give a coefficient of reliability of . 9 , an acceptably accurate figure (Barkley, 193U). Identification of representative genera present in the samples was made either by Rouse (Dept. of Botany, University of British Columbia) or by myself using modern pollen reference slides for comparison. H. Radiocarbon dating ' A sample of the basal peat overlying the compact clay was collected for radiocarbon (C^) dating. The sample was dated by Geochron Laboratories Inc., Cambridge, Mass. CHAPTER I? DESCRIPTION OF THE ASSOCIATIONS The area of the study bog is occupied by ten recognizable vegetation types, five of which are much more distinctive than the others. The five distinctive vegetation types differ widely from each other in both composi-tion and structure and are regarded as five distinct plant associations. Those that display minor, although significant differences, are regarded as variants of a sixth association. The ten described units, then, consist of six plant associations, one of which is composed of five association variants. Most plots analyzed were situated in the study bog (Fig. 6), but many plots were analyzed from other bogs in the region. The vegetational types described were found to be typical for almost a l l of the Long Beach area bogs that were visited. The terra plant association refers to a basic homogeneous classification unit defined by Krajina (19!?2) as follows: \"A plant association is a definite uniform plant canmunity . . . . that is in equilibrium with a certain complex of environ-mental factors, . . . .} its floristic structure . . . . lies within limits governed not only by the ecotope . . . . , but also by the historical factors of vegetational development . . . . This interpretation of the plant association, unlike some others, places emphasis on both vegetation and environment, resulting in a more inclusive concept than that which considers vegetation alone. For this reason, ideally, a l l possible environmental factors are considered in a synecological study. BOG PROFILE LEGEND Cp Carex p l u r i f l o r a a s s o c i a t i o n Mg M y r i c a g a l e v a r i a n t O-Sp Oxycoccus - Sphagnum p a p i l l o s u m a s s o c i a t i o n Cn Chamaecyparis n o o t k a t e n s i s v a r i a n t L-Sp Ledum - Sphagnum c a p i 11 aceum a s s o c i a t i o n V v - i V a c c i n i u m v i t i s - i d a e a v a r i a n t Ph P i n u s c o n t o r t a hummock v a r i a n t P-Vp P i n u s - V a c c i n i u m p a r v i f o l i u m v a r i a n t A l l v a r i a n t s belong to the P i n u s - Sphagnum c a p i 11 aceum a s s o c i a t i o n . Figure 5. Bog profile. C O So Figure 6. Sketch map of study bog, showing distribution of associations. B O G FOREST B O G FOREST Carex p l u r i f l o r a a s s o c i a t i o n ! j M y r i c a g a l e v a r i a n t S c i r p u s - Sphagnum mendocinum a s s o c i a t i o n Chamaecyparis n o o t k a t e n s i s v a r i a n t Oxycoccus - Sphagnum p a p i l l o s u m a s s o c i a t i o n V a c c i n i u m v i t i s - i d a e a v a r i a n t Ledum - Sphagnum c a p i l l a c e u m a s s o c i a t i o n P i n u s - V a c c i n i u m p a r v i f o l i u m v a r i a n t A l l v a r i a n t s b e l o n g t o t h e P i n u s - Sphagnum c a p i l l a c e u m a s s o c i a t i o n . 19 A similar concept is implied by the term \"biogeocoenosis\" of Sukachev (1950), and the term \"ecosystem\" of Tansley (1935)\u00E2\u0080\u00A2 The latter term, however, can be applied at any level of organization and does not necessarily refer only to the basic classification unit. Thus one can speak of a \"bog eco-system\" or a \"forest ecosystem\", or even an ecosystem in a drop of pond water. At whatever level i t is applied, however, the all-inclusive aspect is implied. In some cases, associations are described floristically on the basis of constant species alone, no characteristic species being present (Methods, p. Iii). In bogs and other vegetation types, the plant associations often differ from one another more by varying dominance of one or more of the same group of species, than by the presence or absence of particular species. In such areas associations cannot be described on the basis of fidelity (characteristic species) although in other features they do not differ from other associations. The following descriptions of the bog plant associations represent, in part, a summary of the synthesis tables (Appendix II). A. Caricetum pluriflorae (Carex piuriflora association) The Caricetum is floristically the simplest of a l l the bog associations. An aquatic type, i t is covered with standing water in a l l but the driest periods of very dry summers, and in average summers may not dry out at a l l . It occurs in .the lowest areas of the Long Beach bogs and is usually of quite limited extent. Its occurrence possibly reflects relic marsh or fen condi-tions. Often the Caricetum pluriflorae occupies hollows l i t t l e more than 1\u00C2\u00A7- m in diameter, while at most i t covers areas up to 5 m across. In the center of the bog, where i t is best represented, its distribution is often broken up by patches of the surrounding Oxycoccus - Sphagnum papillosum association. 1 20 Figure 7. Carex pluriflora association surrounded by Oxycoccus quadripetalus - Sphagnum papillosum association Figure 8. Juncus oreganus in deepest water of Carex pluriflora association / The association is characterized by 15-25 cm of normally standing water in -which, after heavy rains, a slight current is often noticeable. In the bottom of the pool, a layer of dy (Kubiena, 1953) is found. The soil profile is seldom over UO cm in depth, at the bottom of \u00E2\u0080\u00A2which the under-lying sands are found. The dy is underlain by 20-25 cm of peat, beneath which a gray clay-like material extends down to the sand. Roots are abundant only in the brown organic layer. As in a l l the Long Beach bog associations, the pH in the surface layers is strongly acid and ranges from 3.U to U.l. At the sand level i t averages Floristically, the association is dominated by a usually luxuriant growth of Carex pluriflora, a preferential, as well as the only constant species. An aquatic form of Sphagnum recurvum is common (average species significance I4.O) and S. papillosum is also frequently found. Apart from Carex, the only commonly occurring vascular plant species is Sanguisorba microcephala. Sporadic species (those species occurring in 20$ or less of the plots of an association) include the following: Vaccinium uliginosum Kalmia polifolia Carex obnupta Gejntiana sceptrum Xn a few areas the water is deeper than 30 cmj in such situations Carex pluriflora is sparse or non-existent. Under such conditions Juncus oreganus occurs frequently. The presence of this species possibly indicates the last remnant of an association that preceded bog development, and that ~ flourished in the deeper waters of a marsh or fen environment. B. Scirpeto-Sphagnetum mendocini (Scirpus caespitosus - Sphagnum mendocinum association) The Scirpeto-Sphagnetum mendocini is of rather limited extent in the study bog but covers large areas of some of the surrounding bogs. It consists primarily of areas dominated by clumps of Scirpus caespitosus, among which is either standing water or mats of Sphagnum papillosum and S, mendocinum. Rhynchospora alba is a common preferential constituent, and i t often occurs in the standing water. Where the Scirpeto-Sphagnetum covers large areas of the bog, Pinus contorta hummock and Carex pluriflora communities are often found scattered throughout. Floristically, the Scirpeto-Sphagnetum is more complex than the Carex pluriflora association, as is shown by the following lists Characteristic combination of speciesJ constant dominants: Scirpus caespitosus Sphagnum papillosum constant non-dominants: Kalmia polifolia Oxycoccus quadripetalus Agrostis aequivalvis . Sanguisorba microcephala Trientalis arctica Gentiana douglasiana Prosera rotundifolia Tofieldia occidentalis Sphagnum mendocinum Sphagnum capillaceum characteristic species: selective: Scirpus caespitosus preferential: Rhynchospora alba Sporadic species: Chamaecyparis nootkatensis Pinus contorta Apargidium boreale Coptis asplenifolla Coptjs trifoliata Cladonia pacifica Bazzania ambigua Sphagnum papillosum is the dominant Sphagnum species here, as in most 23 of the Long Beach bog associations. Where tree species such as Pinus contorta and Chamaecyparis nootkatensis occur in the association proper, they do so only as small, twisted shrubs growing in standing water. Floristically, the Scirpeto-Sphagnetum is a typical low moor association, characteristic of an early stage in bog succession. As in the Carex pluriflora association, the depth of organic accumula-tion is relatively shallow. The upper 20-30 cm of the soil profile are a dark brown peaty organic material, underneath which is often a 10 cm layer of sticky gray to gray-brown clay-like material. Underlying the clay at the 40-60 cm level is sand. As in the Carex pluriflora association, l i t t l e organic accumulation appears to have been of Sphagnum. Roots are abundant only in the dark brown organic layer, especially in the top 10-1 j? cm. The pH of the surface layers ranges from 3.4 to 4.1, and at the sand level from U.2 to 5.U, values roughly comparable to those of the other bog associations of the area. C. Oxycocceto-Sphagnetum papillosae . , (Oxycoccus quadripetalus - Sphagnum papillosum association) The Oxycocceto-Sphagnetum is the most extensive plant association in the bogs of the Long Beach region, and covers large areas of the interior of each of the larger bogs. The association occurs throughout its distribu-tion as part of a mosaic community, the other part of the mosaic being composed of the Pinus contorta hummock association variant, to be discussed later. Within the mosaic community, these two associations play very different roles. The Pinus hummocks support plants otherwise found only in drier areas such as the bog periphery, while the Oxycocceto-Sphagnetum is a typical low moor association supporting only species adapted to very wet environments. Because of the great differences between the two habitats, the two parts of the mosaic are treated as separate entities. Figure 9. Scirpus caespitoaua - Sphagnum mendocinum aasociation Figure 10. Sphagnum capillaceum and S. papillosum invading Scirpus caespitosus - Sphagnum mendocinum association The Oxycocceto-Sphagnetum is a low-lying almost flat association, which in early spring is extremely wet, the water level reaching the top of the Sphagnum papillosum mat that dominates the association* Although no single species is characteristic of the association, many of the species present are constants, and Sphagnum papillosum here reaches its greatest dominance and vigor (av. species significance 9.0, av. vigor 2.9). This species is characteristic of low moors and early bog succession stages. Sphagnum capillaceum, a bog species more often associated with high moors, is found (species significance 1.5) on small slightly raised \"micro-huramocks\" that occur throughout the association. Ericaceous shrubs are numerous, but except for Oxycoccus quadripetalus, they are of low density and vigor. Oxycoccus quadripetalus, however, is a constant although not dominant plant (species significance 2.8). Characteristic combination of species: constant dominants: Apargidium boreale Sphagnum papillosum constant non-dominants: Oxycoccus quadripetalus Agrostis aequivaivis Drosera rotundifolia Kalmia polifolia Trientalis arctica Tofieldia occidentalis Sporadic species: Vaccinium uliginosum Empetrum nigrum Thuja plicata Sphagnum tenellum Cephalozia biscuspidata Myrica gale Pinus contorta Scirpus caespitosus . ' Sphagnum fuscum Polytrichum commune The accumulation of peat is much deeper than in the previously described associations, reaching a depth of 50-100 cm. The organic material varies in Figure 12. Oxycoccus quadripetalus in Oxycoccus quadripetalus -Sphagnum papillosum association. 27 Figure 13* Mosaic community of Pinus contorta hummock variant and Oxycoccus quadripetalus - Sphagnum papillosum association. Figure l U . Trientails arctica Linnaea boreal is and Cornus canadensis growing in Pinus contorta hummock variant. color from brown near the surface to gray-brown at deeper levels, and appears to be composed partially of Sphagnum remains. Underlying the organic layers is a clay-like yellow-gray to gray-brown material that extends down to the sands which are encountered at depths between 7$ and l\u00C2\u00A30 cm. Roots are numerous in the top 20-30 cm, but rapidly decrease below this level. The pH varies at the surface from 3.U to 3.7, and at the 50 cm depth from U.6 to U.8. The Oxycocceto-Sphagnetum papillosae, by virtue of its wide extent, reflects the dominant character of the Long Beach area bogs. D. Ledeto-Sphagnetum capillacei (Ledum groenlandicum - Sphagnum capillaceum association) The Ledeto-Sphagnetum capillacei is primarily an association character-istic of high moor conditions (Hansen, 19U7; Rigg, 192\u00C2\u00A3j Osvald, 1933), and as such is of limited occurrence in the Long Beach bogs. In the larger bogs of the region, i t occurs in small areas around the periphery, while in a few. of the smaller bogs i t is slightly better developed. It appears probable \u00E2\u0080\u00A2 that this association has developed recently in this region, as i t is not extensive nor has as much organic accumulation occurred as in areas where i t is the dominant, well-established association (e.g. Lulu Island, Ladner, and Pitt Meadows bogs in the Vancouver area of British Columbia). In the Long Beach bogs the Ledeto-Sphagnetum is characterized by a surface topography of large hummocks composed of Sphagnum capillaceum and S. fuscum, among which are depressions in which Sphagnum papillosum and S. mendocinum predominate. Ledum groenlandicum, the dominant'shrub of this association, grows extensively on the tops of the hummocks, while other ericaceous shrubs, notably Kalmia polifolia and Oxycoccus quadripetalus, are also common. Carex obnupta is abundant (species significance 7.U) in both the depressions and on the. hummocks, although i t reaches its greatest density and vigor on the hummocks. The Long Beach Ledeto-Sphagnetum capillacei differs from that of typical high moor bogs in several respects: by the presence of depressions between the Sphagnum hummocks; the presence of S. papillosum and S. recurvum; the dominance of S. capillaceum over S. fuscum; and in the relatively shallow accumulation of organic material. In typical high moor conditions, the association is dominated by Sphagnum fuscum, contains few or no wet depressions (and thus none of the Sphagnum species typical of depressions), and has usually a very extensive accumulation of organic material (Osvald, 1933; Hansen, 19U7; Wilde, 1 9 5 8 ) . To a lesser degree than the Oxycocceto-Sphagnetum, the Ledeto-Sphagnetum is also part of a mosaic community, the other portion of which is the Pinus contorta hummock association variant. Although sufficiently different to warrant treatment as separate entities, there is less apparent difference between the composition of the Pinus hummock variant and the Ledeto-Sphag-netum, than between the Pinus hummock variant and the Oxycocceto-Sphagnetum. As in the Oxycocceto-Sphagnetum, no exclusive, selective, or preferen-t i a l species are present, although the l i s t of constants is relatively long. Characteristic combination of species: constant dominants: Ledum groenlandicum Carex obnupta \u00E2\u0080\u00A2 Sphagnum papillosum Sphagnum capillaceum constant non-dominants: Kalmia polifolia Qxycoccus quadripetalus > Apargidium boreale Prosera rotundifolia Trientalis arctica ' \ ' Sporadic species: Myrica gale Sanguisorba microcephala Carex pluriflora Scirpus caespitosus Deschampsia caespitosa Maianthemum dilatatum Coptis asplenifolia Polytrichum commune Sphagnum recurvum Bazzania ambigua Dicranum scoparium Riccardia palmata ? The accumulation of organic material is the deepest of any of the bog associations, reaching a depth of 65-120 cm in the soil pits studied. Its color ranges from light brown near the surface to dark brown at the lowest levels. Largely undecompo3ed Sphagnum appears to make up the bulk of the accumulation. Underlying the organic accumulation is an indistinct layer of clay-like material that merges gradually with the organic layer above and the sands below. The sand occurs between 90-170 cm from the surface. Roots, especially of Carex obnupta, are abundant in the top 40-65 cm. The pH at the surface varies from 3.3 to 3.7 and at the 65 cm depth from 3.7 to U.5. A pH of U.6 -was recorded from a depth of 100 cm. E. Pineto-Sphagnetum capillacei (Pinus contorta - Sphagnum capillaceum association) This is the most variable of the- bog associations studied. It occurs primarily on drier parts of the bogs such as the bog margins and hummocks around tree bases, and as a secondary successional stage on burned areas. The dominant species is Pinus contorta, which almost always occurs as a stunted tree of peculiar rounded form (Fig. 18). Sphagnum papillosum is also here the dominant Sphagnum species, although S. capillaceum reaches greater development here than elsewhere in the bogs. Ericaceous shrubs also reach their greatest local development here, but they do not achieve the density and vigor that typifies their occurrence in high moors. The Pineto-Sphagnetum capillacei is composed of five distinct variants. These variants, though relatively similar in habitat and floristic Figure 15. High moor Sphagnum fuscum hummock with associated species. Figure 16. Apargidium boreale growing on Sphagnum fuscum hummock 32 composition, appear to have developed their present similarities secondarily. Floristically, they are similar, although significant minor differences do exist. The variants of the Pineto-Sphagnetum capillacei, in order of increasing complexity, are as follows: (a) sphagnosum papillosi (Pinus contorta hummock variant) (b) myricosum galis (Myrica gale variant) (c) chamaecyparosum nootkatensis (Chamaecyparis nootkatensis variant) (d) vacciniosum vitis-idaeae (Vaccinium vitis-idaea variant) (e) vacciniosum parvifolii (Vaccinium parvifolium variant) The last two variants occur as secondary successional stages after fi r e . , Characteristic combination of species: constant dominants: Pinus contorta Carex obnupta Sphagnum papillosum constant non-dominants: Ledum groenlandicum Kalmia polifolia Empetrum nigrum Oxycoccus quadripetalus Apargidium boreale Trientalis arctica ' Prosera rotundifolia Sphagnum capillaceum Frullania nisquallensis ' . . ' I I \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 ' ' . ' i characteristic species: preferential: Empetrum nigrum Kalmia polifolia Vaccinium vitis-idaea \u00E2\u0080\u00A2> Vaccinium uliginosum Myrica gale Cladonia crispata Cladonia pacifica Cladonia rangiferina In addition to the characteristic combination of species, the association includes many sporadic species, the most notable of which are: Picea sitchensis ' Carex pauciflora Nephrophyllidium crista-galli Habenaria saccata Boschniakia hookeri Pteridium aquilinum Veratrum viride Following are descriptions of the five variants: 1. Pineto-Sphagnetum capillacei sphagnosum papillosi (Pinus contorta hummock variant) The Pinus contorta hummock variant occurs as part of the mosaic community already discussed. It occurs throughout the open central areas of a l l the bogs and most often with the Oxycoccus - Sphagnum papillosum association. Less often i t occurs with the Ledum - Sphagnum fuscum and the Scirpus - Sphagnum mendocinum associations. The variant is never extensive since i t develops from a gradual build-up of organic material to form a hummock around the base of one or several dwarf pines. The hummocks are composed mainly of Sphagnum, especially S. capillaceum. The tops of the hummocks provide a drier habitat than the surrounding association and thus support species of plants otherwise not ,34 found in the wet central bog areas. Notable among these are Empetrum nigrum. Linnaea borealis, Vaccinium vitis-idaea, Cornus canadensis, Maianthemum dilatatum, Lycopodium clavatum, and Rhacomitrium lanuginosum, a l l species adapted to relatively dry habitats. In addition the pine trees support a large number of epiphytes that are otherwise not present in the surrounding association. Typical of these are the lichens Cladonia bellidiflora, Usnea plicata, and Sphaerophorus globosus, but many bryophytes are also present. The vegetation of the hummocks of this variant is somewhat similar to that of the hummocks of the Ledum - Sphagnum capillaceum association, although the hummocks differ widely in structure and origin. The pine trees that form the dominant feature of the variant are a l l dwarfed specimens with peculiarly rounded tops (Fig. 18). They range between 4-10 (16) feet in height, with a trunk diameter at 3 feet above ground of 1-6 inches. Ages of representative 6-10 foot specimens varies from 60-110 years. The Sphagnum hummocks commonly grow up and ultimately engulf the lowest branches of the pines so that low foliage-covered branches are often completely buried in Sphagnum capillaceum and S. fuscum for several feet of their lengths. It is possible that layering occurs in this way, for the vertically emerging branch tips often closely resemble young pines in their form. No rooting from the branches was observed. In addition to hummock development, the Pinus hummock variant usually includes small water f i l l e d depressions in which Lysichitum americanum is often present. Lysichitum affects unfavorably the growth of Sphagnum species in its immediate vicinity because of the shading caused by its large leaves. During bog development the Sphagnum grows up around the Lysichitum, eventually leaving i t confined to a small pit (Turesson, 1916} Osvald, 1933). The roots of the Lysichitum usually extend underneath the Sphagnum accumulation to where the pH is less acidic (see Successional Trends). High moor development usually completely eliminates the Lysichitum (Osvald, 1 9 3 3 ) . Shading from prostrate Pinus contorta and Chamaecyparis nootkatensis also apparently inhibits the growth of Sphagnum, for many water-filled pools are found in the bog below such specimens, particularly in the association under discussion. Characteristic combination of species: constant dominants: Pinus contorta Carex obnupta y Sphagnum papillosum Sphagnum recurvum constant non-dominants: Ledum groenlandicum Kalmia polifolia Apargidium boreale Drosera rotundifolia Sphagnum mendocinum Scapania bolanderi No characteristic species are found in this variant. Vascular plant sporadic species: Chamaecyparis nootkatensis 1 Tsuga heterophylla Carex pauciflora Deschampsia caespitosa Qxycoccus quadripetalus Linnaea borealis Agrostis aequivalvis Sphagnum capillaceum Frullania nisquallensls Dicranum scoparium Vaccinium vitis-idaea Coptis asplenifolia Maianthemum dilatatum Lycopodium clavatum The soil profile is very similar to that of the surrounding Qxycoccus -Sphagnum papillosum association. Approximately 40-60 cm of brown peat composed partly of decomposing Sphagnum overlies a 10-20 cm thick layer of black, sticky material containing few plant roots. Below the black, sticky material is the sand layer at a depth of 75-90 cm. Roots are most abundant in the top I 4 O - 6 O cm and are uncommon below the brown organic layer. Very few roots are in the sand. At the surface the pH varies from 3.U to 3.9, and at the sand level from U.5 to $.h 2. Pineto-Sphagnetum capillacei myricosum galis (Pinus contorta - Sphagnum capillaceum - Myrica gale variant) The Myrica gale variant occurs only in a single large section of the study bog and is absent from most of the nearby bogs. Although i t contains most of the other species of the association, i t is dominated by a dense growth of Myrica gale (species significance 8.6). Chamaecyparis nootkatensis \u00E2\u0080\u00A2 is second in abundance (species significance U.8), occurring most commonly as a prostrate spreading shrub, although some dwarf trees (up to 10 feet in height) are present. Pinus contorta occurs here as a round-topped tree of larger size than those in the Pinus contorta hummock variant, and reaches an average height of 1$ feet. The surface of the Myrica gale variant presents a rough uneven pattern of dry hummocks dominated by shrubs, and wet depressions dominated by Sphagnum papillosum, S. mendocinum, S. recurvum, and Carex obnupta. Sphagnum capillaceum is the dominant Sphagnum of the hummocks. In addition to the hummock-depression pattern, many small water-filled pools occur underneath the dense shrubby Chamaecyparis specimens. The character of these pools is much the same as that of those described for the Pinus contorta hummock variant, although Lysichitum americanum is a much less common inhabitant. Sphagnum recurvum and Carex obnupta appear to be the most common inhabitants of the pools, which are heavily shaded by the overhanging shrubs. Sphagnum recurvum was the only Sphagnum observed growing in shaded locations, such as below dense bog shrubs and in the bog forest. Figure 20. Thuja plicata and Pinus contorta high moor community. growing in mature Characteristic combination of species: constant dominants: Myrica gale Chamaecyparis nootkatensis Sphagnum papillosum ' Sphagnum mendocinum constant non-dominants: Pinus contorta Qxycoccus quadripetalus Ledum groenlandicum Trientalis arctica Apargidium boreale Coptis asplenifolia Sphagnum capillaceum Frullania nisquallensis Dicranum scoparium characteristic species: selective: Myrica gale preferential: Sphagnum mendocinum Carex canescens Vascular plant sporadic species include only Lysichitum americanum. In the soil profile a dark brown peaty layer extends to 75 cm in depth and overlies a narrow layer of clay-like material. Underlying the clay at the 85 cm level is sand. Roots are abundant to a depth of 45-50 cm and are s t i l l fairly common to a depth of 70 cm. Below 70 cm roots were rare. 3\u00C2\u00AB Pineto-Sphagnetum capillacei chamaecyparosum nootkatensis (Pinus contorta - Sphagnum capillaceum - Chamaecyparis nootkatensis variant) The Chamaecyparis nootkatensis variant occurs commonly around the margins of the bogs in a band varying from 10-150 feet in width. It contains both bog and forest elements, and may be considered as a transitional type between bog and bog forest. It resembles the Myrica gale variant, differing from i t mainly in the absence of Myrica gale, the greatly increased dominance of Chamaecyparis nootkatensis, and the decreased dominance of Sphagnum species. Chamaecyparis nootkatensis dominates the B or shrub layer, but is usually subdominant to Pinus contorta in the A layer. In the A layer Chamaecyparis reaches a maximum of 1$ feet in height and 10 inches dbh, and is seldom more than 1J>0 years old. Pinus contorta reaches a maximum of 30 feet in height and 16 inches dbh, with a maximum age of 250 years. Thuja plicata and Tsuga heterophylla are also present, but are of low dominance and poor vigor. The hummock-depression pattern is similar to that of the Myrica gale variant, except that the hummocks comprise a proportionately greater area. Sphagnum species (except for S. capillaceum which occupies the hummocks) are of much lower dominance than in the Myrica gale variant, because of the decreased extent of the depressions which they normally inhabit. As in the Myrica gale variant, many small pools occupied by Carex obnupta and Sphagnum recurvum occur immediately beneath the overhanging Chamaecyparis branches , '.and decumbent trunks. \^ Among the forest species that occur i n this variant are the following! Gaultheria shallon (vig. 1.0) Linnaea borealis Vaccinium ovatum Vaccinium parvifolium Vaccinium ovalifolium Cornus canadensis Maianthemum dilatatum Blechnum spicant The ericaceous shrubs a l l appear to establish only on decaying wood or on the tops of hummocks which are composed solely of organic material. ~ Characteristic combination of species: constant dominants: ^ Chamaecyparis nootkatensis 1 Carex obnupta \u00E2\u0080\u00A2 Figure 21. Characteristic form of old Tsuga heterophylla in Long Beach area bogs. Figure 22. Pinus contorta, Tsuga heterophylla and Thuja plicata at edge of bog forest. Apargldium boreale Sphagnum papillosum constant non-dominants: Pinus contorta Ledum groenlandicum Qxycoccus quadripetalus Drosera rotundifolia Empetrum nigrum Kalmia polifolia Cornus canadensis Maianthemum dilatatum Pleurozium schreberl Isothecium stoloniferum Scapania bolanderl Sphagnum capillaceum epiphytic constant non-dominants: Frullania nisquallensis Dicranum scoparium Polypodium glycyrrhiza characteristic species: ..preferential: Chamaecyparis nootkatensis (only within the bog ecosystem) Vascular plant sporadic species: Picea sitchensis Vaccinium ovalifolium Vaccinium uliginosum Scirpus caespitosus Deschampsia caespitosa Gentiana douglasiana Agrostis aequivalvis Nephrophyllidium crista-galli The soil profile shows a dark brown peat extending to an average depth of 40-60 cm, and overlying a 10-20 cm wide band of black, sticky clay-like material. The sand layers occur at the 80-90 cm level, immediately beneath the clay-like material. Roots are abundant in the top 40 cm and are fairly common down to approximately 60 cm. Below this level few roots exist. The pH in the surface layer i s 3.7 and at the sand level 4.5 to 5 . 2 . Figure 24. Boschniakia hookeri parasitic on Gaultheria shallon of Vaccinium vitis-idaea variant. Figure 25\u00E2\u0080\u00A2 Ledum groenlandicum, dominant shrub of Ledum groenlandicum - Sphagnum capillaceum association* Figure 26. Kalmia polifolia, co-dominant shrub of Vaccinium vitis-idaea variant. Figure 27. Coptis trifoliata and Empetrum nigrum growing in Vaccinium vitis-idaea variant. Figure 28. Nephrophyllidium crista-galli growing in wet depressions of burned area of bog. li. Pineto-Sphagnetum capillacei vacciniosum vitis-idaeae (Pinus contorta - Sphagnum capillaceum - Vaccinium vitis-idaea variant) The Vaccinium vitis-idaea variant occupies large areas of several of the bogs of the Long Beach area, and appears to represent a secondary successional stage after f ire . Charred branches and stumps are evidence of past f ire, and in many areas most of the organic layer also appears to have been burned off. Such areas have an extremely shallow organic layer over-lying sand and some charcoal. The vacciniosum vitis-idaeae is the driest of a l l the bog associations and variants, and supports a dense growth of erica-ceous bog shrubs and lichens. Typically, the topography of the Vaccinium vitis-idaea variant is one of hummocks and depressions. The hummocks are occupied by shrubs such as Empetrum nigrum, Kalmia polifolia, ledum groenlandicum, and Vaccinium v i t i s -idaea, and by high moor mosses such as Sphagnum capillaceum and S . fuscum. The lichens Cladonia pacifica, C. rangiferina, C, uncialis, C. arbuscula, and C. bell idiflora also inhabit the drier hummocks. The moist depressions, by contrast, are occupied by wet habitat species such as Apargidium boreale, Drosera rotundifolia, Nephrophyllidium crista-gall i , and Sanguisorba micro-cephala. The lichen Cladonia crispata is exclusive to the low wet depressions of this variant. The low moor Sphagnum species, S. papillosum, S. mendocinum, and S. recurvum are also present in the wet depressions, but less commonly than in the other bog associations. Pinus contorta is abundant in the vicinity in the Vaccinium parvifolium variant (Fig. 23), but is of only scattered occurrence in the Vaccinium vitis-idaea variant. Small specimens of Thuja plicata occur in the shrub layer, but' Chamaecyparis nootkatensis is missing entirely. Forest species also occur, notably Cornus canadensis, Blechnum spicant, and Maianthemum dilatatum. Characteristic combination of species: constant dominants: Empetrum nigrum Ledum groenlandicum Kalmia polifolia ''7:\"'''\"'' Sphagnum capillaceum . \u00E2\u0080\u00A2 constant non-dominants: - Gaultherla shallon Vaccinium vitis-idaea Pinus contorta Apargidium boreale \ Blechnum spicant Trientalis arctica j Cladonia pacifica Cladonia rangiferina Dicranum scoparium Frullania nlsquallensis characteristic species: exclusive: Cladonia cri3pata preferential: Kalmia polifolia Vaccinium vitis-idaea Vaccinium ullginosum Thuja plicata Carex obnupta Cornus canadensis Sanguisorba mlcrocephala Maianthemum dilatatum Cladonia crispata Pleurozium schreberi Rhytidiadelphus loreus Empetrum nigrum Vaccinium ullginosum Cladonia rangiferina Cladonia pacifica The association also includes many sporadic vascular plants, the most notable of which are: Nephrophyllidium crista-galli ' Habenaria saccata Boschniakia hookeri The soil profile shows a very shallow dark brown organic layer usually not exceeding 10-15 cm in thickness. Below 15 cm the organic layer gradually merges with an ashy gray-brown horizon which frequently contains some charcoal. This horizon merges with the underlying sands at an average depth of U0-60 cm. In certain situations the gray-brown ashy material is exposed at the surface and is colonized mainly by lichens. In such situations i t is apparent that the organic layer has been completely burned off. Roots are most abundant in the top 10 cm, and are rare below hO cm. The pH at the surface varies from 3.h to 3.7 and at the sand level from U.7 to U.8, values no different from those of the wet low moor associations. 5. Pineto-Sphagnetum capillacei vacciniosum parvifolii (Pinus contorta - Sphagnum capillaceum - Vaccinium parvifolium variant) The Vaccinium parvifolium variant, like the Vaccinium vitis-idaea variant, occupies burned areas of some of the bogs and is apparently a secondary successional stage. It i s , however, dominated by dense clumps of Pinus contorta and Thuja plicata rather than by ericaceous shrubs. The pines average between 10 and 15 feet in height, and are not as markedly round-topped as the species is in other bog associations. The physiognomy here is similar to that of Pinus contorta as i t occurs in the bogs of the southwestern B.C. mainland (Ladner, Lulu Island, Pitt Meadows bogs). The specimens of western redcedar are seldom over 7 feet in height and have quite sparse foliage. Western redcedar appears to be regenerating well in the B layer under the pine canopy, whereas the shade intolerant pines show no regeneration at a l l . Many charred stumps and snags indicate past fire. Occasional large pines (av. i|0 foot height, 16 inch dbh, \u00C2\u00B1200 years age) are fire scarred near their bases, and appear to be remnants of a previous forest. These large trees, unlike the younger specimens that dominate the variant, show the characteristic round top that distinguishes most of the lodgepole (shore) pines of the area. ^ Under the tree canopy bog shrubs are less common than forest species such as Gaultheria shallon, Vaccinium parvifolium, and Linnaea borealis. Forest species of herbaceous plants also dominate over bog species, the most characteristic being Blechnum spicant, Cornus canadensis, and Maianthemum dilataturn. Sphagnum species are much less abundant than in the other variants and associations, the two commonest species being S. recurvum, which often grows under shade, and S. capillaceum, which occupies hummocks in open areas of the variant. The number of epiphytic species is much lower than in any of the other tree-bearing vegetation types (perhaps because of the much drier conditions). Characteristic combination of species: constant dominants: '- Pinus contorta Thuja plicata Ledum groenlandicum Gaultheria shallon . . constant non-dominants: Kalmia polifolia Vaccinium vitis-idaea Vaccinium parvifolium ' Oxycoccus quadripetalus Carex obnupta \ \ \u00E2\u0080\u00A2 Blechnum spicant Cornus canadensis Apargidium boreale Maianthemum dilatatum Sphagnum papillosum Eurhynchium oreganum Frullania nisquallensis Vaccinium parvifolium is the only characteristic species. The most notable vascular plant sporadic species are: Menziesia ferruginea Habenaria saccata ' Pteridium aquilinum , Boschniakia hookeri Veratrum viride . ' The soil profile is similar to that of the Vaccinium vitis-idaea variant, except that the peat layer is usually deeper, averaging 20 cm in thickness. Beneath the peat small amounts of charcoal appear scattered through an ashy gray-brown horizon, a further indication of fire. Under-lying the ashy horizon is sand which, due to cementation in the uppermost layers, forms a fairly distinct hardpan. The hardpan occurs at an average depth of 40-45 cm, below which the sand is noticeably bluish in color. Roots are abundant in the organic layer, but are very rare in and below the ashy layer. The pH is similar to that of the other bog associations and variants, averaging from 3.4 to 3.5 at the surface, and 4.4 to 5.4 at the sand level. Of the five variants of the Pinus contorta - Sphagnum capillaceum association, the greatest similarities appear to exist between the Vaccinium vitis-idaea and the Vaccinium parvifolium variants, both of which occur after fire , and between the Myrica gale and the Chamaecyparis nootkatensis variants both of which occur on the margins of the bogs. The Pinus contorta hummock variant, although floristically similar to the other variants, occurs as isolated islands of high moor vegetation surrounded by low moor associations. It has been suggested that the two secondary successional variants, the vacciniosum vitis-idaeae and the vacciniosum parvifolii, together form a vegetation type distinctive enough to be recognized as a separate association (Krajina, personal communication 1965). In this event each of the above variants would be termed variants of the Pineto-Vitis-idaeeto-Sphagnetum capillacei. The presence of Vaccinium vitis-idaea as a relatively abundant species, and the notably increased dominance of Cladonia species over that ' of other variants of the Pineto-Sphagnetum capillaoei, constitute the floristic criteria for such a separation. Edaphically the presence of a charcoal layer constitutes an important differentiating ecological factor. . It is thus suggested that future study in this area might substantiate the recognition of a Pineto-Vitis-idaeeto-Sphagnetum capillacei, a seventh bog association in the bogs of the Long Beach area. F. Pineto-Chamaecypareto-Sphagnetum recurvi (Pinus contorta - Chamaecyparis nootkatensis - Sphagnum recurvum association) (Bog forest) The bog forest occupies extensive lowland areas surrounding the bogs of the Long Beach region, and although not a bog association, is discussed as a vegetational type bordering the bogs and thus exerting considerable influence on them. The forest is composed primarily of several species of dwarfed and peculiarly-shaped conifers, of which the dominant species are Pinus contorta, Chamaecyparis nootkatensis, and Thuja plicata. Dominating the forest aspect are relatively large specimens of Pinus contorta, averaging UO feet but occasionally reaching 70 feet in height. These comparatively large trees are characterized by t a l l , branching trunks which support umbrella-like masses of foliage at their tips (Fig. 31). Their form resembles the deliquescent branching of artgiosperm trees, and is unlike that of any of the other gymnosperms of the region. The apparent mechanism of this phenomenon is early loss of dominance of the apical shoot, with subsequent increased growth of the lateral branches. The actual cause may possibly be due to calcium deficiency (Krajina, personal communication, 1965). Greenhouse experiments carried out by Krajina (1959) indicate that lack of sufficient calcium causes in conifers an increased growth and branching of the lateral branches, resulting in a bushy appearance. Available calcium in the bog associations is much less than in forest associations, a fact that may be the cause or partial cause of the stunted bushy form of a l l the coniferous species in the bogs. Similar growth of Pinus contorta is found in bogs northward to Alaska (Heusser I960, Schofield - personal communication, 52 Figure 30* View of bog forest, with Chamaecyparis nootkatensis. Thuja plicata, Tsuga heterophylla, and Pinus contorta. Figure 3 1 . Characteristic form of old Pinus contorta in bog forest of Long Beach area (height 75 feet, age 260 years). 1965). The largest pines of the bog forest were found to be just under 300 years o ld, while average-sized trees of U0-50 feet varied from 120-180 years old. The other bog forest tree species, with the exception of Thuja p l i ca ta , a l l have dwarfed, bushy forms and seldom grow over UO feet i n height (Fig. 30)\u00E2\u0080\u00A2 Thuja p l i ca ta exhibits quite normal growth except for common stem die-back, and reaches a maximum height of approximately $0 feet at 250 years. Taxus b rev i fo l l a occurs as a common though non-dominant bog forest element, and reaches a maximum height of 25 feet. The following mensurational data were taken from representative bog forest trees: Height ( f t ) Diameter (ins) at U Age (yrs) feet above ground Pinus contorta Uo 11.8 175 U5 10.5 150 U7 12.6 16U U8 1U.0 180 70 ; \u00E2\u0080\u00A2 16.5 '\"':V\" 285 Chamaecyparis nootkatensis 3U; : \u00E2\u0080\u00A2 7.3 ; \ 1 3 6 39 6.2 85 39 7.1 11* Thuja p l icata UO 13.7 : 155 : U9 1U.5 250 Tsuga heterophylla 29.5 10.5 .: 188 32 1U.6 1UU 3U.5 8.9 16U Of these tree species, only Thuja p l icata and Tsuga heterophylla appear to be regenerating under the tree canopy. Many saplings of these two species were observed i n the B layer. No young specimens of either Pinus contorta or Chamaecyparis nootkatensis were observed i n this association. Pinus contorta is completely shade intolerant (Hansen, 19U7} Krajina, personal communication, 196U) and thus does not germinate under a closed canopy forest. The shrub layer is dominated by Gaultheria shallon, but other erica-ceous shrubs are also common. Bog elements are represented by Ledum groen-landicum and Vaccinium vitis-idaea. The herbaceous layer is composed of both bog and forest elements, of which the forest species comprise most of the dominants. Carex obnupta, Apargidium boreale, and Trientalis arctica are examples of apparent invaders from the adjacent bogs. The bryophyte layer is extremely luxuriant and includes Sphagnum recurvum as a constant species. This moss occurs in patches in the wettest depressions of the forest floor. The association is also extremely rich in epiphytes, both in numbers and species. Besides many bryophytes, the l i s t includes many usually terrestrial vascular plants such as Vaccinium parvifolium, Cornus canadensis, Menziesia ferruginea, and Gaultheria shallon. The abundance of epiphytes is a direct reflection of the high precipitation and constant high relative humidity of the region. On a modest scale, the bog forest shows many of the attributes of a temperate rain forest. Characteristic combination of species: constant dominants: ' Pinus contorta \u00E2\u0080\u00A2 . Chamaecyparis nootkatensis Thuja plicata Gaultheria shallon \u00E2\u0080\u00A2\u00E2\u0080\u00A2\u00E2\u0080\u00A2\u00E2\u0080\u00A2\u00E2\u0080\u00A2\u00E2\u0080\u0094. , constant non-dominants: / Tsuga heterophylla Vaccinium parvifolium Vaccinium ovatum Menziesia ferruginea Maianthemum dilatatum Hylocomium splendens Rhytidiadelphus loreus i > Scapanla bolanderi Lepidozia reptans constant non-dominant epiphytes: Isothecium stoloniferum ];!\u00E2\u0080\u00A2 Frullania nisquallensis || ' Antitrichia curtipendula Dicranum scoparium Cornus canadensis No characteristic species are found. Sporadic vascular plant species include: Malus diversifolia Sphagnum recurvum Eurhynchium oreganum Herberta adunca Scapania bolanderi Hypnum circinale Vaccinium parvifolium Pblypodium glycyrrhiza Gentiana sceptrum B cc 0-5 cm 5-20 cm 25-35 cm HO-60 cm below 70 cm Qoodyera oblongifolia The soil profile shows fairly distinctly the typical horizons of a podzol: l i t t e r ; humus whitish-gray leached eluviated layer, gray-brown cemented illuviated layer outwash (or stream deposit) sands and gravels Root distribution is most abundant in the dark brown humus layer, especially the top 30 cm which appear to be dominated by roots of Gaultheria shallon. Roots are few in the gray leached horizon and below. The pH of the soil of this association is higher than that of any other bog association, but is s t i l l highly acidic. In the humus horizon the pH ranges from U.O to li.5 and in the sand level from 5.1 to 5.U. The stunted growth of this forest probably results from the cemented layer lying close to the soil surface, preventing drainage and thus preventing satisfactory aeration. The pedogenic process of podzolization usually results in the formation of a distinct hardpan which is impermeable to water (Wilde, 1958). When such a hardpan develops in low-lying basins with no drainage outlet, the probable result is waterlogging of the soil, preventing adequate aeration for normal forest development. Soil nutrients are unlikely to be the limiting factor, since the quantities of available cations present are relatively high for a region of high precipitation. CHAPTER V SOILS An evaluation of the soil data (Appendix III) results in the basic generalization that no correlation exists between the pattern of distribu-tion of bog plant associations and the relative quantities of available nutrients. The soils of a l l the bog associations contained almost equal amounts of available cations (Na+, IT1\", Ca + +, Mg++), total nitrogen, and adsorbed phosphate.\u00E2\u0080\u00A2 Similarly, cation exchange capacity, pH, and percent base saturation were highly uniform among the various associations. As stated previously, the nutrient content appears relatively rich for an area of high precipitation, and i t thus seems unlikely that nutrients play the role of limiting factors affecting distribution of the associations. . The ratio of magnesium to' calcium (5.0:6.5) is much higher than in most peat soils. Buckman and Brady ( i 9 6 0 ) give an average ratio of magnesium to calcium in peat soils of 1:8. The higher than average value obtained for magnesium may possibly be explained by the proximity of the bogs to the open ocean, and the subsequent effect of salt spray. Salt spray, which contains a very high proportion of magnesium, is, according to local inhabitants, often blown inland short distances by the frequent gales of winter. The effect of such spray may possibly be responsible for the disproportionately high magnesium content encountered in the bog soils. Available calcium, however, is much lower ( 6 . 5 me#) than that of approximately U0 me* which is taken as representative of bog soils (Buckman and Brady, I 9 6 0 ) . The reason for the low calcium figure is due possibly to the rapid leaching that occurs in the region of the Long Beach bogs due to the high precipitation (\u00C2\u00B1120 inches per year) of the area. In such a region, where the soils of the climax forests are well-developed podzols, the effects of leaching are considerable. The results of the soil analysis from the bog forest differ only slightly from those of the bog associations. Available calcium is much higher, averaging 24.0 me$ in the humus layer as against 6.5 me$ for that of the bog associations. Total nitrogen, conversely, is lower, with an average in the surface layers of 0.88$ as against 1.6$ for that of the bog associa-tions. Percent base saturation in the surface layers of the bog forest averages twice as high as that of the bog associations, with an average of 13.0$. Soil moisture is much less than in the bog associations because of the.inferior water-holding capacity of humus in comparison to bog peat. In other aspects measured, however (available Na, K, Mg, adsorbed phosphate, and cation exchange capacity) no difference from the bog associations was evident. Summary of the soil data A. Soil moisture In the surficial layers the soil moisture is highest (1380$) in the Ledum groenlandicum - Sphagnum capillaceum association and lowest. (265$) in the bog forest. The other associations have values somewhere between these two extremes, depending directly on the proportion of Sphagnum in the organic layer. The higher the proportion of Sphagnum, with its unrivalled water holding capacity, the higher the soil moisture. Thus the only true high moor .^association in the bog, the Ledum groenlandicum - .Sphagnum capillaceum association, which has the highest Sphagnum accumulation, has . also the highest percent soil moisture. Correspondingly, the bog forest and the secondary successional variants, which have l i t t l e or no Sphagnum accumulation, have also the lowest percent soil moisture (265$, 500$ resp.). A correlation may also be attempted between tree growth and percent soil moisture. The associations -with high soil moisture support l i t t l e tree growth, probably because of resultant decreased aeration. Topography would thus appear to play the final distributional role, in that i t is responsible for the amount of water accumulation and thus for aeration. B. - Total nitrogen The average value of total nitrogen for the surface layers is 1.6%, with no significant difference between most of the associations. The value (3*9%) for the Ledum groenlandicum - Sphagnum capillaceum association, however, is much higher. The bog forest, with l i t t l e or no peat deposition, shows only 0.88$. Peat generally contains a higher total nitrogen content than other soils (Buckman and Brady, I960). For this reason the Ledum groenlandicum - Sphagnum capillaceum association, which possesses the highest proportion of peat, would.be expected to have the highest total nitrogen content. The underlying sand layers have uniform values averaging 0.08$. C. \u00E2\u0080\u00A2 Available sodium No trends are evident. The surface layers range from 3.9 to 6.3 me# and the underlying sand from 0.5 to 0.9 m.e%. D. Available potassium No trends are evident. The values for the surface layers range from 0.3 to 2.17 me%, in this case the extreme high and the extreme low values being from the same association. The average for the associations is 1.5 rae%. The underlying-sands average 0.03 to 0.09 me%, E. Available calcium No trends are evident except for^the bog forest, which'shows an average figure for the humus of 2U.0 me%- The surface layers of the other /associa-tions average 6.5 me%. The underlying sands range from O.U to 1.1 me%. F. Available magnesium No trends are evident. The surface layers range from 3.6 to 7.3 me%, and average 5.0 me%. The underlying sands range from 0.03 to 0.2 me%. G. Adsorbed phosphate The results appear somewhat chaotic, with differences lacking any-evident pattern. Surface values range from O.U ppm (Scirpeto-Sphagnetum) to 5.3 ppm (Caricetum), while underlying sand values range from 0.9 ppm (Caricetum, Scirpeto-Sphagnetum) to 3.0 ppm (Pineto-Sphagnetum capillacei sphagnosum papillosi). H. Cation exchange capacity No trends are evident. Values in the surface layer, whether of peat in.the bog associations or of humus in the bog forest, range from 2l6 to 278 me$. The highest values are from the bog forest. These values are approximately equivalent to cation exchange capacity figures from representa-tive North American bogs (Buckman and Brady, I960). Values of the underlying sand range from 2? to 5 l me%] with an average of 1;0$. I. Percent base saturation Percent base saturation figures (av. 7.0$) are much lower than the average (33.5$) for representative bogs of other areas (Buckman and Brady,-I960). For the upper layers in the profile the bog forest has the highest figure with an average of 13.0$. Figures for the underlying sand demonstrate no.trends, ranging from 2.5 to 5.6$. J... pH Except for the bog forest, a l l the associations and variants of the bog display remarkable constancy in pH. The uppermost layers of the profiles of a l l the associations have pH values ranging from 3.5 to 3.7. The pH of the underlying sand\" is slightly less acidic, varying from U\".5~tb\" U:9. The values for the bog forest are less acidic, ranging from an average'in the humus layer of 4.5 to an average in the underlying sand layer of 5.3. K. Discussion of soils . In many studies high moors have been found to be considerably more acidic than low moors; this is thought to be due partly to the release of organic acids from the high moor Sphagnum species (Wilde, l\u00C2\u00B058j Buckman and Brady, I960). In the bogs under study, however, the low moor associations such as the Scirpus caespitosus - Sphagnum mendocinum association are just as acidic as the Ledum groenlandicum - Sphagnum capillaceum association, commonly a high moor type. The pH of a soil affects in various ways the availability of the soil's nutrients. Iron, for example, is more available in acidic conditions, calcium in neutral to alkaline conditions. The presence of specific nutrient requirements, therefore, limits an individual species' range of habitats to those of suitable pH. In a broader sense entire associations are limited in their distribution to specific pH limits, simply because the nutrients which the component plants require are available in sufficient quantity only within those limits. Within a given localized . area an association may often be defined at least partly by its pH, and successional sequences are often speculated on the basis of a gradual shift i n pH between associations, usually towards the acidic side. Often the change in pH is the result of the activity of the plants themselves. In the Long Beach area, bog associations can be defined neither on the basis of pH, which is uniform, nor on available soil nutrients, which are also uniform. A correlation between the two conditions probably exists: the uniform pH maintains the availability of the same nutrients in the same relative proportions. The fact that not only the relative proportions of each nutrient are similar, but also that the amounts of each nutrient are very similar for each of the associations, possibly reflects the common origin and composition of the parent material. ~~ ~. The extreme acidity (pH 3.5-3.7) of the bog associations is a direct reflection of the very heavy rainfall of the area. In such areas hydrogen ions displace mineral cations in the upper profile layers, creating an acidic environment and resulting in a very low percent base saturation. In comparison, representative eastern North American bogs, probably because of a much lower rainfall, are less acidic (pH lw0-5.1) and have correspondingly higher percent base saturation values (33.5 - 76.6$ versus 7.0$ for Long Beach bogs) (Buckman and Brady, I960). Such figures probably indicate that the bogs under present study are less productive than bogs as an average. As discussed earlier, profiles from the bog associations show l i t t l e well-defined horizon development. Apart from the bog forest with its podzol formation, the rest of the associations have soil profiles composed primarily of an upper organic horizon, usually of peat, and a lower clay-like horizon. These horizons vary considerably in their respective thicknesses among the various associations. The clay-like horizon is usually partly organic in its highest occurrence in the profile, while towards its base i t merges 1 gradually with the underlying outwash or stream deposit sands. These sands and gravels are well cemented in their upper layers, preventing drainage from the bog associations above them. The cementation in the sands is probably a result of previous podzolization in the soils of a former forest/ which stood on whatsis now bogland. CHAPTER V I L I F E FORMS The distribution of l i f e forms in the associations and variants under study is presented graphically (Fig. 32). The system followed is that of Raunkiaer (193U) as modified'by Braun-Blanquet (1965), as i t is the best known and most widely accepted l i f e form classification in use today. In the present study various trends and correlations are evident. The associations and variants are arranged in order of increasing floristic complexity (Fig. 32). The simplest associations (Caricetum pluriflorae, Scirpeto-Sphagnetum mendocinium) are those with the most extreme edaphic conditions. Conversely, the most complex associations (Bog forest, Chamae-cyparis nootkatensis variant) are those with the least extreme edaphic. conditions (i.e. they are most like the zonal forest in composition). Correlated with this increase in floristic complexity is a distinct change in the proportions of the.life forms. The simplest associations show almost complete dominance of a few l i f e forms. As the associations progress in floristic complexity, however, the dominance of a few l i f e forms is gradually replaced by a more even distribu-tion of l i f e forms within an association. In the more complex associations no one l i f e form is outstandingly dominant. It has often been stated that bogs are \"enclaves of subarctic l i f e \" (Deevey, 1958), and that in many features they resemble subarctic or boreal areas. Many of the plant species found in bogs are thought to be subarctic or boreal relics, and are sometimes numerous enough to impart a very \"northern\" aspect to temperate bogs. While the absence of permafrost in these bogs 65 definitely limits the comparisons to be made, there are, however, some definite similarities, such as the composition of l i f e forms. Apart from bryophytes, which are numerous in a l l the associations, hemicryptophytes constitute the dominant l i f e form, both in number of species and total cover. Next in abundance are the chamaephytes and the nanophanerophytes, although these latter groups do not approach the hemicryptophytes in number. This composition of l i f e forms is very similar to that found in subarctic or alpine regions (Braun-Blanquet, 1932). In the floristically more complex associations which contain many forest elements, the composition of l i f e forms more closely resembles that of warm temperate than of subarctic areas. Specifically, an increase in the proportion of macrophanerophytes, nanophanerophytes, and chamaephytes is evident, correlated with a considerable decrease in the proportion of hemicryptophytes. Geophytes also increase slightly, but never assume a significant position. Therophytes, which are most abundant in warm regions with long growing seasons, are represented only by a single species, Gentlana douglasiana. In conclusion, a distinct correlation may be drawn between the floristically simple bog associations having the most extreme edaphic conditions, and a l i f e form spectrum reflecting boreal or subarctic condi-tions. Similarly, a correlation exists between the more complex bog associa-tions with less extreme edaphic conditions, and a l i f e form spectrum reflecting warm temperate conditions. LIFE FORM DISTRIBUTION Caricetum pluriflorGe Scirpeto-Sphagnetum mendocini Oxycocceto-Sphagnetum paplbsae Pm Pn Ch H G T B Pm Pn Ch H G T B L P m P n C h H G T \u00E2\u0080\u00A2 Number of species Pirwto-Charnaecypareto-Sphagnetum recurvi Pm Pn Ch H G T Figure 32. Life form distribution. CHAPTER VII HISTORY The history is in part a summary.of various previously discussed aspects. The study bog lies directly over apparent glacial outwash or stream deposits and there is l i t t l e evidence of what type of vegetation preceded.bog initiation. It is probable that the seaward flow of water following deglaciation was previously much greater than at present. This is evidenced by the wide extent of the outwash or stream deposits at Wreck Bay. Bog initiation, however, began no earlier than 3\u00C2\u00B00\u00C2\u00B1 90 years B.P. It is thus evident that a considerable period of time must have elapsed between the deposition of the outwash or stream deposits and the beginnings of bog development. It appears most probable that a forest of some type gradually developed through succession on the deposits, only to be replaced in the ; past U00 years by bog vegetation. Bogs can develop only when drainage is seriously impeded, which is the probable reason for the development of bogs on previously forested areas. Drainage of the valley floors where the bogs are located is impeded by two factors: the topography of the landscape which in many areas allows no drainage outlet, and the presence of a cemented layer in the otherwise highly porous sand. This cemented layer forms a distinct hardpan in the uppermost layers of the sand, and effectively prevents drainage from the overlying bog associations. The impervious cemented layer probably had its origin in the soils of the former forest, through the process of podzoliza-tion. Podzolization is the normal soil forming process found in a cool, moist climate where raw humus is accumulated in large amounts. Accumulation is usually due to retarded decomposition. In the Long Beach-Tofino area decomposition is retarded because of the highly acidic environment caused by the heavy precipitation, high accumulation of organic matter, and cool weather. Acids produced by the slowly decomposing humus bring about the solution of sesquioxides of iron and aluminum together with a deflocculation of the colloids. Oxides and colloids are both carried downwards in the soil profile where they are precipitated out in the B horizon (Daubenmire, 1959J Soil Survey Manual, 1 9 5 l ) . One common result of this process is the formation of an impervious hardpan or \"ortstein\", composed of material cemented with precipitated iron and organic matter. Such a process can occur only where large amounts of raw humus are accumulated, thus indicating the probable existence of a previous forest. Once a hardpan becomes well established in a shallow valley with no drainage outlet and a high level of precipitation, the existing forest is probably doomed. Gradually the soil becomes water-logged, aeration is greatly decreased, and conditions become mature for bog invasion. Before conditions become too acidic for its existence a skunk cabbage (Lysichitum americanum) association may thrive in the wet lowland habitat (Krajina, personal communication, 1 9 6 5 ) . Finally Sphagnum species, initially-S. recurvum and S. mendocinum, become established, the forest species gradually die out, and bog species attain supremacy. Thus i t can be said that in such circumstances the forest destroys itself, in that the soil processes initiated because of the presence of the forest eventually are the cause of its destruction. / ,, As time progresses the bog invades more and more of the surrounding scrub forest until large areas of the lowlands are occupied by low moor bogs From this present stage, high moor bogs may in time become established. This hypothetical sequence of events appears to represent the most logical history of the bogs of the area, as i t is substantiated by considerable circumstantial evidence. The results of the pollen analysis also confirm this hypothesis. v A much less probable explanation of the origin of the impervious layer is that of glacial compaction of the sand. This suggestion appears highly improbable, partly because of the known age of the bog. In the event of glacial compaction, the bog would be expected to have originated immediately following deglaciation, approximately 6000-8000 years B.P. Another aspect that appears to refute this possibility is the apparent non-marine origin of the sand (see Geology). CHAPTER VIII POLLEN ANALYSIS The results of the pollen analysis (Fig. 33} reflect both the develop-ment of the present bog vegetation and to a lesser extent, changes that have occurred in the vegetation of the surrounding forest. Evidence of the latter must be interpreted as representing only very local fluctuations, since the time span of the bog (390\u00C2\u00B1 90 years B.P.) is much too short to record changes in the vegetation of the region as a whole. Caution must be taken too, in interpreting vegetational trends on the basis of the analysis of only one sample corej however, basic trends are clearly discernible and are probably reasonably accurate. The organic accumulation of the bog is 1-| meters deep in the area from which the core was taken. The lowest \ meter is a mixture of clay and organic remains and appears to contain very l i t t l e or no Sphagnum. The upper J meter appears to contain mostly remains of undecomposed Sphagnum, while the middle \u00C2\u00A7\u00E2\u0080\u00A2 meter contains a mixture of Sphagnum, plus other plant remains. The core was taken from the center of the study bog in the plant association of widest extent, the Oxycoccus - Sphagnum papillosum association. The pollen profile contains no peat bog species in its lowest depths (Fig. 33). Prevalent in the pollen of this early period of deposition are Pinus, cf. contorta (20$), Alnus, cf. rubra (32$), and Picea, cf. sitchensis 1 {22%). These three species are common invaders on mineral soil, and today often occur as pioneers on recently deglaciated surfaces at low altitudes (Heusser, 1960j Cooper, 1939; Rigg, 1937). That some shallow pools and probable seepage habitats existed at this time is shown by the presence of the following plants: Cyperaceae and Juncaceae (10$), Veratrum (10$), Myrica (1$), and Typha (2$). Of these plants, Typha is restricted to areas of shallow water, while the other plants are a l l restricted to moist habitats. Exceptions occur in the Cyperaceae and Juncaceae, some of which inhabit dry areas. Typha also commonly occurs in the skunk cabbage (Lysichitum americanum) association as a secondary plant after opening of the forest by logging, fire, etc. As i t . i s a completely shade intolerant species, its presence in the bottom of the bog profile probably indicates open seepage habitats, possibly occupied by a skunk cabbage association (Krajina - personal communication, 1965). By this stage the former forest was probably receding in the face of bog development. The pollen record gives no indication of a lake as a stage preceding bog formation, unlike the history of many other bogs in southwestern British Columbia. This observa-tion is to be expected in view of the topography of the area, which includes no depressions deep enough for lake formation. The pollen profile shows a gradual decrease in abundance of Pinus, Picea, and Alnus in upper layers; at the surface pollen of these trees is present only in small quantities (5$, 4$, 1$, respectively). Pinus contorta, however, is s t i l l a very abundant species of the present day bog vegetation. Tsuga, cf. heterophylla, Rhamnus, cf. purshiana, and Salix sp. are a l l present in small numbers in the profile, and reflect no definite changes in the vegetational composition. Cupressaceous pollen, unlike that of other confiers, does not preserve well and is rarely present in the pollen record (Hansen, 1940} Rouse, personal communication, 1965). For this reason the record of the coniferous trees is probably biased, as Thuja plicata, Chamaecyparis nootkatensis, and Juniperus communis var. montana commonly occur in the area today, and almost certainly were extant on the bog during its development. 72 Veratrum, cf. vtrlde is surprisingly abundant (up to 17$) through most of the profile, but disappears completely towards the top. Its disappearance from the pollen record appears to be well correlated with the appearance of pure bog species, in particular Sphagnum species. Lysichitum, cf. americanum is well documented at the 90 cm level, the same point at which Veratrum reaches its greatest abundance. The presence at this period of both Veratrum and Lysichitum in abundance, correlated with the absence of true bog species such as Sphagnum, suggests wet seepage conditions possibly occupied by a skunk cabbage association. Both these species thrive be3t today in wet seepage habitats of close to neutral pH (Krajina - personal communication, 1965). In highly acidic conditions such as are found in Sphagnum bogs, both species exhibit greatly reduced dominance and vigor, while in true high moor bogs they are non-existent (Osvald, 1933J Turesson, 1916). Both species, and especially Lysichitum americanum, may, however, commonly exist as relics in low moor bogs, although they are usually of scattered occurrence and relatively poor vigor. In such situations the roots of the two plants commonly extend beneath the Sphagnum to the less acidic environment below. The ericaceae are first noted at the 100 cm depth, but as the identifi-cation of separate genera of ericaceous pollen is almost impossible, i t cannot be said whether or not the pollen belongs to bog genera such as Ledum and Kalmia, or to woodland genera such as Gaultheria and Menziesia. The ericaceae appear according to the pollen analysis to be reduced in numbers near the top of the profile. Such a reduction may reflect the decrease of Gaultheria shallon, the dominant forest shrub of the area, during bog development. Gaultheria is very uncommon on the bog surface and, along with the major tree species, probably died out in the bog area during bog invasion. As most of the bog is not yet in the bog shrub stage, Kalmia, ledum, and other ericaceous bog shrub pollen probably does not counteract in quantity the loss of salal. Myrica, cf. gale pollen is present in small quantities throughout the profile, an indication of prevailingly wet, but not necessarily bog conditions. Today Myrica gale occurs both in bogs and,, more commonly, around lake edges. Myrica gale grows much better in the latter habitat, where conditions are generally less acidic than in Sphagnum bogs. Throughout the profile the cyperaceous and juncaceous pollen (which . cannot be distinguished from each other) increases steadily from 10$ in the lowest level to a maximum of 53$ at the surface. These plant families, and especially Carex of the Cyperaceae, include the dominant plants of the earlier bog stages and the marsh or fen stages that precede most bog develop-ment (Hansen, 1940, 1950} Rigg, 1 9 2 5 , 19U0; Osvald, 1 9 3 3 ) . The gradual increase of this group of plants throughout the profile reflects the change from a probable wet seepage habitat to the herb-stage topogenous bog that is present today. The Polypodiaceae are present throughout the entire profile, but with greatly increased abundance at the 100 cm level (Fig. 3 3 ) . This point of abundance may simply indicate a locally rich pocket of pollen rather than a general trend. Trends indicated by the rest of the plants identified i n the profile (Composltae, fungal spores) are inconclusive, partly because of the lack of more definitive identification. Identification was carried where possible to the genus level, but in no cases were the pollen grains definitely determined to species. Of the \u00E2\u0080\u00A2 genera identified, however, the following species are the only ones present today in the region studied: Pinus contorta - (much less commonly P. monticola) Picea sitchensis Alnus rubra - (less commonly A. crispa subsp. sinuata) Rhamnus purshiana ' U-Tsuga heterophylla Myrica gale - (M. califomica is now found in the area, but is generally thought to be a recently introduced species) Lysichitum americanum Typha l a t i f o l i a j, Veratrum viride Considering the short time interval involved, these species present today are probably the same as those found throughout the pollen profile. In summary, the pollen analysis appears to reflect a gradual change . in UOO years from a wet seepage forest habitat to a low moor bog in the herb stage. Apparent present day invasion of ericaceous shrubs and high moor Sphagnum species suggests that succession in the area may eventually lead to high moor development, with shrubs instead of herbs as the dominant vascular plants. POLLEN PROFILE Figure 33. Pollen profile. CHAPTER IX SUCCESSIONAL TRENDS A. Regional level At the regional level, as has already been discussed, bog succession appears to have superceded forest succession in the lowest parts of the valleys. At the present time i t appears that the bogs are s t i l l enlarging their areas at the expense of the surrounding bog forest. Several bog elements, notably Ledum groenlandicum, Trientalis arctica, and Sphagnum is especially prevalent, forming solid mats in low wet areas. In some parts of the forest Sphagnum recurvum appears to be spreading at the expense of woodland species such as Mnium glabrescens, Plagiothecium undulatum, and Eurhynchium oreganum. Invasion of the forest by bog elements is a much more plausible, explanation than the opposite sequence of events, namely that the forest has developed from a bog and represents a final stage of succession. This latter sequence appears impossible because no peat accumulation exists in the bog forest soi l profiles. Even under the patches of Sphagnum recurvum almost no accumulation.of peat is present, indicating that the presence of this species is undoubtedly of very recent occurrence. In the transitional area between the bog and the bog forest are found some of the plant species that, according to the pollen analysis, were common during the period of bog initiation. These species, particularly Veratrum viride and Lysichitum americanum, are species that occur primarily in wet seepage habitats such as skunk cabbage associations. Habenaria 77 saccata, although not present in the pollen record, is another species of such habitats which also occurs in the transitional area. Carex obnupta, another species that occurs abundantly in skunk cabbage associations, also occurs with great frequency in the transitional area, especially in the Chamaecyparis nootkatensis variant. A possibility that suggests itself is that present day changes in vegetation, as seen along a transect from forest to bog, reflect the historical change from forest to bog that is recorded in the pollen profile. The probable major criterion governing the changes is in both instances the same* the relative degree of aeration, or other possible environmental factor, which is indirectly controlled by topography. The major changes, greatly simplified, appear to involve the transition from mesic forest to wet seepage habitat to low moor bog. The formation of hardpans in the soils of the valley provide i n i t i a l stimulus by impeding drainage. The excess moisture, because of a lack of sufficient drainage outlets, is probably retained to a large extent within the valleys, and contributes to an ever-rising water table. In such a situation, the slightly higher areas surrounding the bogs would gradually become water-logged, aeration would simultaneously decrease, and bog vegetation would theoretically invade and replace the bog forests. The intermediate stage, the Lysichitum americanum and Veratrum viride inhabited wet seepage habitat, appears to occur in varying degree both in the present day vegetation at the bog periphery, and in the center of the bog as a past stage recorded in the pollen profile. This suggested regional successional sequence, although highly specu-lative, appears to best f i t the observed pattern. Under its implications, the bogs that occupy the lowest parts of the valleys of the region are slowly invading the surrounding bog forests that occupy slightly higher ground. The criterion of invasion appears to be the water-level, which due to underlying impervious hardpans is controlled largely by topography. High precipitation (-120 inches per year) and generally high relative humidity combine to decrease evaporation, -which might otherwise tend to negate the effect of the slowly rising water level. In somewhat similar situations in coastal Alaska and north coastal British Columbia, bogs commonly develop on . gradual slopes, having apparently invaded these sites from the adjacent valley bottoms where they were previously well-established (Heusser, I 9 6 0 ; Rigg, 1925). B. \u00E2\u0080\u00A2 Association level (Fig. 3^) The term \"sphagnum bog\" has been defined as \"that stage in the physio-graphic succession of an area during which its surface is entirely devoid of ordinary \"hard\" soi l and is composed almost entirely of l iving Sphagnum, immediately under which is fibrous brown peat composed mainly or entirely of partially disintegrated Sphagnum, the habitat exercising a distinctly selective influence on its flora\" (Rigg, 1922) . Sphagnum bogs are of two basic types, the low moor bog and the high moor bog. Low moor bogs are formed at or below the level of the water table, and are dependent on ground water for basic moisture requirements. Their surfaces are flat or slightly concave. High moor bogs, by contrast, are not necessarily dependent upon the occurrence of ground water, but receive most of their moisture directly from precipitation. They may occur well above the water table and often occur on slopes. The surface of high moor bogs, or \"raised bogs\", is convex, the center of the bog being sometimes considerably higher than the margins (Daubenmire, 191.7; Wilde, 1958; Rigg, 1925, 19U0, 1938). Low moor bogs may also be designated \"topogenous bogs\", and high moor bogs \"ombrogenous bogs\", the terms referring to the major source of water (Heusser, I960). The bogs under study contain features of both types of bogs, although 79 basically they are typical low moors. The surfaces of a l l the bogs observed , are essentially flat, and coincide approximately with the water table. In early spring large portions of the bog are completely submerged in several inches of water, and even in late summer the surface is s t i l l extremely wet to walk on. The three basic low moor associations occupying the bulk of the bog area are the Carex pluriflora association, the Scirpus caespitosus -Sphagnum mendocinum association, and the Oxycoccus quadripetalus - Sphagnum papillosum association. Small portions of the bogs, however, especially near the periphery, are occupied by a developing high moor community, the Ledum groenlandicum - Sphagnum capillaceum association. Here the surface, in the form of Sphagnum capillaceum and S. fuscum hummocks, is several feet above the water table, although the associations are of too limited an extent to show the typical high moor convex surface. The other associations of the bog appear to represent transitional stages to high moor or to forest development, and often contain both high and low moor elements. On a broad basis, the bog appears to be gradually developing towards a high moor situation, although the change is probably in its i n i t i a l stages. High moor bog species, especially Sphagnum capillaceum and S. fuscum (Wilde, 1958; Osvald, 1933), are common although not dominant species and occupy, besides the high moor association already discussed, the tops of numerous very tiny hummocks throughout the low moor area. In addition, they are very common in the peripheral and secondary successional associa-tions, usually occurring on hummocks around tree bases and on decaying wood. Their general pattern of distribution suggests that they are spreading on the bog surface, and may in time form a continuous high moor bog. Of these two species, Sphagnum capillaceum is by far the most common. S. fuscum, regarded by many workers (Wilde, 1958; Osvald, 1933; and others) as occupying the final stage in high moor development, is restricted to the highest hummocks, and is commonest in the Ledum groenlandicum - Sphagnum capillaceum association. Other peculiarly high moor species such as Careat pauclflora (Krajina - personal communication, 1965) also appear to be gaining in abundance. The Pineto-Sphagnetum capillacei vacciniosum vitis-idaeae, an association variant that represents a secondary successional stage after fire, also displays many high moor characteristics. Floristically, i t consists of hummocks dominated by Sphagnum capillaceum plus a dense growth of ericaceous shrubs. * It differs from normal high moors mainly by its lack of a deep layer of peat accumulation. According to Osvald (1933) who worked on bogs of the southwestern British Columbia mainland, high moor bog species such as Sphagnum fuscum, Ledum groenlandicum, and Kalmia poli-folia, along with many Cladonia species, commonly invade recently burned areas. He termed this community type the Ledum groenlandicum - Sphagnum fuscum sociation (Osvald, 1933). Definite successional sequences are difficult to hypothesizej however, the following tentative sequence appears most logicalt . 81 Carex pluriflora assoc. Scirpus caespitosus -Sphagnum mendocinum assoc. .Oxycoccus quadripetalus -Sphagnum papillosum assoc Ledum groenlandicum -'Sphagnum capillaceum assoc Vaccinium vitis-idaea variant Vaccinium parvifolium variant Pinus contorta \"hummock variant Myrica gale variant Chamaecyparis nootkatensis variant fire Bog Forest Figure_34. Successional trends. This hypothetical pattern suggests the high moor Ledum groenlandicum -Sphagnum capillaceum association as the eventual dominant bog association, towards which many of the other associations and variants are tending. Eventually, Sphagnum fuscum becomes the dominant high moor species (Wilde, 1 9 5 8 ) . It is probable that as the high moor develops, the Myrica gale variant may disappear, since Myrica gale grows poorly in the highly acidic ' conditions of a high moor (Krajina - personal communication, 1 9 6 5 ) . Vege-tation types such as the Pinus hummock and Chamaecyparis nootkatensis variants may retain their tree development but at the same time develop lesser vegetation typical of high moors. The two secondary successional variants appear to be the only vegetation types tending towards the drier bog forest, rather than towards the wetter high moor associations. The reason for this apparent difference in trend may be the fact that the burned areas are situated on higher ground than the low moor bog associations and are thus not affected as much by the rising water table. i The sequence of Caricetum pluriflorae to Ledeto-Sphagnetum capillacei appears very distinctive as a series of serai changes from low moor to high moor. Caricetum is floristically the simplest, as well as the wettest association. In much of the bog, however, i t apparently is being replaced directly by the Oxycocceto-Sphagnetum papillosi rather than by the Scirpeto-Sphagnetum raendocini, which i s of rather limited occurrence. A definite increase in floristic oomplexity is evident as progression is made towards high moor development. Finally, the bog forest apparently is being replaced by peripheral bog associations, i n i t i a l l y by the Chamaecyparis nootkatensis variant. Further evidence of this change is provided by numerous dead snags near the margin of the bog, indicating that forest trees probably once lived in what is now part of the bog, > \" CHAPTER X SUMMARY AND CONCLUSIONS The basic results and conclusions of the study are summarized as follows. (1) The distribution of the bogs, the scrub or \"bog forest\", and the climart forests of the terrace is dependent indirectly on \u00E2\u0080\u00A2 topography. Drainage of the soils of the region is impeded by impervious hardpans developed as a result of podzolization. \u00E2\u0080\u00A2 The development of hardpans, correlated with lack of adequate drainage outlets, produces in the valleys water-logged soils in which aeration is apparently insufficient for forest develop-ment. Bogs, therefore, occupy the lowest parts of the valleys, where drainage, and thus aeration, is probably minimal. Scrub forest (bog forest) occupies the gentle slopes of the valleys where drainage i s better than on the valley floor, while the climax western hemlock-amabilis f i r forest occupies the highest parts of the terrace, where drainage is not a limiting factor. (2) The phytosociological aspect of this study resulted in the description and characterization of ten different vegetation types, nine belonging to the bog and one belonging to the surrounding bog forest. The nine bog vegetation types consist of four distinct associations and a f i f t h association composed , of five variants. Two of these variants occur on burned areas ; and represent secondary successional stages. 8U (3) A soil analysis revealed no correlation between community type ++ ++ and the following soil characteristics: available Ca , Mg , Na+, K+, adsorbed phosphate, cation exchange capacity, and percent base saturation. In a l l these aspects the differences among community types are negligible. (U) Soil moisture is highest in the,bog associations, intermediate in the bog-forest transitional Associations, and lowest in the secondary successional variants and the bog forest. The associa-tion having the greatest dominance of Sphagnum species has also the highest percent soil moisture. The amount of soil moisture . is interpreted as being directly proportional to the degree of ' aeration, thus supporting the hypothesis of ( l ) . (5) Total nitrogen is highest in the bog associations and lowest in ...the bog forest. A correlation appears possible between total nitrogen and amount of accumulation of Sphagnum species, as the . high moor association shows a much higher total nitrogen figure than any of the other associations. (6) Available Mg + + is much higher than in most bog areas, the higher figure possibly being accounted for by salt spray. (7) pH was not demonstrated to play any role in the distribution of bog plant associations within a bog area. The surface peat of , a l l associations and variants is extremely acidic (3.5 to 3.7 at the surface) but no differences in pH are found between them. '.' (8) The bog forest differs from the bog associations by the following soil-analytical aspects: higher available Ca + +, less acidio pH, lower soil moisture and lower total nitrogen. (9) . The age of the bogs, based on radiocarbon dating of a basal peat sample, is approximately400 years. From this and other evidence, r ) a tentative history of the bogs of the area is suggested as follows: . a) A post land-uplift period of glacial outwash or stream deposition. . b) A period of plant succession on the outwash or stream deposits, terminating in the development of a forest. c) A period of increasing lack of soil aeration caused by the development of hardpans, eventually resulting in water-logged soils and subsequent forest decline. d) Initiation of the bogs approximately U00 years ago, the preliminary stages including shallow water areas and a possible marsh vegetation. e) Gradual development during the past 1|00 years of a low moor bog and recently, indications of high moor development. The results of a pollen analysis appear to verify the preceding statement. The pollen profile indicates at its lowest level an abundance of coniferous trees, the presence of seepage habitat species, and the absence of true bog species such as Sphagnum. Higher in the profile the disappearance of seepage habitat species such as Veratrum and Lysichitum is correlated with the appearance of Sphagnum and other bog species. On a regional level the bogs of the area appear to be gradually increasing their areas by invading the surroundingbog forest. The criterion of invasion is suggested to be a gradually rising water table that renders the bog forest soils too water-logged to support even a scrub forest, hence the subsequent development of conditions favorable for bog vegetation. 86 On an association level, the sequence of succession appears to be from floristically simple low moor associations, through more complex low moor associations, to the high moor associa-tion. At the bog margins the bog forest appears to be giving way i n i t i a l l y to the Chamaecyparis nootkatensis variant, and subsequently to the Myrica gale variant or the Ledum groen- landicum - Sphagnum capillaceum association. 1. BIBLIOGRAPHY Armstrong, J.E. 1956. Surficial geology of Vancouver area, B.C. Geo. Survey of Canada, paper 5 5 - 4 0 . ., 1957. Surficial geology of New Westminster Map-Area, B.C. Geo. Survey of Canada, paper 5 7 - 5 . I960. Surficial geology of Sumus Map-Area, B.C. Geo. Survey of Canada, paper 5 9 - 9 . Barkley, F. 1934. The statistical theory of pollen analysis. Ecology 15* 2 8 3 - 2 8 9 . Becking, R.W. 1957. The Zurich-MontpelDier school of phytosociology. Bot. Rev. 23J 411-488. Bird, H. 1 9 2 3 . On the \"boreal\" character of bogs and an a r t i f i c i a l modification. Ecology 4 : 2 9 3 - 2 9 6 . Bowman, P.W. 1 9 3 1 . Study of a peat bog near the Matamek River, Quebec Canada, by the method of pollen analysis. Ecology 1 2 : 6 9 4 - 7 0 8 . Braun-Blanquet, J. 1932. Plant Sociology. The study of plant communities. English trans, of Pflanzensoziologie. Edited by G.D. Fuller and H.S. Conard. 439 pp. Braun-Blanquet, J. 1 9 6 4 . Pflanzensoziologie. Springer-Verlag, Wein. 865 pp. Brooke, R.C. 1 9 6 5 . Ecotopes of plant communities in the ecosystem classification of the coastal Subalpine Zone in southern British Columbia. Ph.D. Thesis, Dept. Bot., Univ. of B.C. Buckman, H.O. and N.C. Brady, i 9 6 0 . The nature and properties of soils. Macmillan 6 t h Ed. 565 pp. Canada, Dept. Transport. 1 9 5 6 - 1 9 6 3 . General summaries of hourly, weather observations. Meteor. Br. Clements, F.E. 1928. Plant succession and indicators. H.W. Wilson Company, N.Y. Cooper, W.S. 1937. The problem of Glacier Bay, Alaska: a study of glacier variations. Geog. Rev. 27: 3 7 - 6 2 . \"\" 1939. A fourth expedition to Glacier Bay, Alaska. Ecology 20: 130-155. Dachnowski-Stokes, A.P. 1930. Peat profiles in the Puget Sound basin of Washington. J. Wash. Acad. Sci. 20: 193-209. Daubenmire, R.F. 1959. Plants and environment. A textbook of plant autecology. 2nd Ed. J. Wiley and Sons, inc., 1*22 pp. Davis, N.F.G., and W.H, Mathews. 19)4.. Four phases of glaciation with illustrations from southwestern British Columbia. J. Geol. 52: U03-U13. Deevey, E.S. 1958. Bogs. Scientific American - Oct. 2-8. Dolmage, V. 1920. West coast of Vancouver Island between Barkley and Quatsino Sounds. Canada Dept. of Mines, Geol. Sur. Sumra. Rpt. Hansen, H.P. 1938. Postglacial forest succession and climate in the Puget Sound Region. Ecology 19J 528-51*2. . 1939. Paleoecology of a central Washington Bog. Ecology 20: 563-568. . 191*0. Paleoecology of two peat bogs in southwestern British Columbia. Amer. J. of Bot. 27: ll*U-ll*9. 191+0. Paleoecology of a montane peat deposit at Bonaparte Lake Washington. Northwest Sci. l l * : 60-69. . 191*1. Paleoecology of a bog in the spruce-hemlock climax of the Olympic Peninsula. Amer. Mid. Nat. 25: 290-297. . 19l*3\u00C2\u00BB A pollen study of two bogs on Orcas Island, of the San Juan Islands, Washington. Bull. Tor. Bot. Club 70: 236-2U3. . 19U7. Postglacial forest succession, climate, and chronology in the Pacific Northwest. Trans, of the Amer. Phil. Soc. Vol. 37, part I. . 1950. Pollen analysis of three bogs on Vancouver Island, \"Canada. J. Ecology 38: 270-276. Heusser, C.J. 1959. Radiocarbon dates of peats from North Pacific North America. Amer. J. Sci. Radiocarbon Supplement 1: 29-3U. ' . I960. Late-Pleistocene environments of North Pacific North America. Am. Geog. Soc. special pub. no. 35. 368\"pp. \" Jackson, M.L. 1961;. Soil chemical analysis. Prentice-Hall, Inc. 1*98 pp. Krajina, V.J. 1933. Die pflanzengesellschaften des Njynica Tales in . den Vysoke Tatry (Hohe Tatra). Beihefte zum Botanischen Centrallblatt. Bd. 50, Abtlg. 11, 77l*-957 (I. Teil): Bd. 51, Abtlg. 11, 1-221; (II Teil). . 1958. Ecological requirements of Douglas-Fir, western hemlock, Sitka spruce and western redcedar. Presented at the meetings of the Royal Soc. of Canada. June 1958. . 1959. Bioclimatic zones in British Columbia. \"Univ. of B.C. Bot. Series 1. 47 pp. . 1965. Biogeoclimatic zones and biogeocoenoses of \"British Columbia. In: Ecol. Western N. Am. 1: 1-17. Lewis, F.J. and E.S. Dowding. 1926. The vegetation and progressive changes of peat areas (muskegs) in central Alberta. J. Ecology 14: 317-341. Metson, A.J. 1961. Methods of chemical analysis for soil survey samples. R.E. Owen, Gov. Printer, Wellington, New Zealand. 208 pp. McMillan, C. 1956. The edaphic restrictions of Cupressus and Pinus i n the coast ranges of central California. Ecol. Mono. 26: 177-212. Mueller-Dcmbois, D. 1959. The Douglas-fir forest associations on Vancouver Island in their i n i t i a l stages of secondary succession. Ph.D. thesis, Dept. Biol, and Bot., Univ. of B.C. Osvald, H. 1933. Vegetation of the pacific coast bogs of North America. Acta Phytogeographica Suecica, Uppsala 5s 1-32. Peterson, E.B. 1964. Plant associations in the subalpine mountain hemlock zone in southern British Columbia. Ph.D. thesis, Dept. Biol, and Bot., Univ. of B.C. Ratcliffe, D.A. 1964. Mires and bogs. In: The vegetation of Scotland. Oliver and Boyd Ltd., Edinburgh. 613 pp. Rigg, G.B. 1914. Notes on the flora of some Alaskan Sphagnum bogs. PI. World. 17t 167-182. :. 1916. The toxicity of bog water. Amer. J. Bot. 8: 436-437. ' 1916. Physical conditions in Sphagnum bogs. Bot. Gaz. 6 l : 159-163. , . 1916. A summary of bog theories. PI. World. 19: 310-325. . 1917. Forest succession and rate of growth i n Sphagnum bogs. J. Forestry 15: 726-739. \" . 1918. Growth of trees in Sphagnum. Bot. Gaz. 65: 359-362. . 1922. The Sphagnum bogs of Mazama Dome. Ecology 3: 321-324. . 1925. Some Sphagnum bogs of the North Pacific coast of America Ecology 6: 260-278. ' \u00E2\u0080\u00A2 . 1933. Notes on a Sphagnum bog at Fort Bragg, California. Science 77: 535-536. . 1937. Some raised bogs of southeastern Alaska \u00E2\u0080\u00A2with notes on flat bogs and muskegs. Amer. J. Bot. 21;; 191+-198. . 19U0. Comparisons of the development of some Sphagnum bogs of the Atlantic Coast, the Interior, and the Pacific Coast. Amer. J. Eot. 27: 1-lU. . 19l;0. The development of Sphagnum bogs in North America. Bot. Rev. 6\u00C2\u00BB 666-693. Rigg, G.B. and E.S. Harrar. 1931. Root systems of trees growing in Sphagnum. Amer. J. Bot. l 8 s 391-397. Rigg, G.B. and CT. Richardson. 193U. The development of Sphagnum bogs in the San Juan Islands. Amer. J. Bot. 21: 610-622. . 1938. Profiles of some Sphagnum bogs of the Pacific Coast of North America. Ecology 19: 1;08-1;34. Rigg, G.B. and T.G. Thompson. 1922. A bog forest. Ecology 3: 207-213. . .1922. Birch succession in Sphagnum bogs. J. Forestry 20:. 1-3. Sears, P.B. and E. Janson. 1933. The rate of peat growth in the Erie Basin. Ecology l l ; : 3U8-355. Transeau, E.N. 1903. On the geographic distribution and ecological relations of the bog plant societies of northern North America. Bot. Gaz. 36: 1;01-1;20. Turesson, G. 1916. Lysichiton camtschatcense (L) Schott, and its behavior in Sphagnum bogs. Amer. J. Bot. 3: 189-209. Wilde, S.A. 1958. Forest soils. Ronald. 537 pp. 2. BIBLIOGRAPHY OF PUBLICATIONS USED IN THE IDENTIFICATION OF VASCULAR PLANTS Gilkey, H. 1961. Handbook of northwest flowering plants. Binfords and Mort. hXk pp. ^ T V - . Hitchcock, A.S. 1950. Manual of the grasses of the United States. U.S. Govt. Printing Office, Wash. 1051 pp. Hitchcock, A.S., A. Cronquist, M. Oweriby, and J.W. Thompson. 1955. Vascular plants of the Pacific Northwest.. Part $t Compositae. ' .Univ. of Wash. Press. 353 pp. \u00E2\u0080\u00A2 . 1 9 5 9 . Vascular plants of the Pacific Northwest. Part Us Ericaceae through Campanulaceae. Univ. of Wash. Press. 510 pp. . 1 9 6 1 . Vascular plants of the Pacific Northwest. Part 3s Saxifragaceae to Ericaceae. Univ. of Wash. Press. 510 pp. . 1964. Vascular plants of the Pacific Northwest.Part 2: Salicaceae to Saxifragaceae. Univ. of Wash. Press. 597 pp. Feck, M.E. 19Ul. A manual of the higher plants of Oregon. Binfords and Mort. 866 pp. Szczawinski, A.F. 1 9 5 9 . The orchids of British Columbia. B.C. Prov. Mus., Dept. of Educ, Handbook No. 1 6 . Victoria, B.C. 12U pp. 1 9 6 2 . The heather family (Ericaceae) of British Columbia. B.C. Prov. Mus., Dept. of Educ, Handbook No. 19. Victoria, B.C. 205 pp. Taylor, T.M.C. 1963. The ferns and fern-allies of British Columbia. B.C. Prov. Mus., Dept. of Educ, Handbook No. 12. Victoria, B.C. 172 pp. 3. BIBLIOGRAPHY OF PUBLICATIONS USED FOR IDENTIFICATION OF BRYOPHYTES AND LICHENS Al t i , T. 1 9 6 1 . Taxonomic studies on reindeer lichens (Cladonia, sub-genus Cladina). Societas Zoologica Botanica Fennica \u00E2\u0080\u00A2Vanamo'i Helsinki. 160 pp. Conard, H.S. 1 9 5 6 . How to know the mosses and liverworts. Wm. C. Brown Co., Iowa. 226 pp. Dixon, H.M. 1 9 5 4 . The students handbook of British mosses. Sumfield and Day, Ltd., London. 582 pp. Frye, T.C. and L. Clark. 1949. Hepaticae of North America. Vol. 6 , nos. 1 - 5 . 1018 pp. Grout, A.J. 1 9 0 3 . Mosses with the hand lens and microscope. A non-technical handbook of the more common mosses of the north-eastern United States. Publ. by the Author. Brooklyn. 416 pp. Grout, A.J. 1 9 2 8 . Moss flora of North America. Publ. by the author. Hale, G.E. 19!?0. Lichens of the State of Washington. Univ. Wash. Press. 191 PP. Hale, M.E. 1961. Lichen Handbook. A guide to the lichens of eastern North America. Smithsonian Institution, Wash. D.C., 178 pp. Lamb, I.M. 1963. Index nominum lichenum. The Ronald Press Co., N.Y. 809 pp. Macvicar, S.M. 1926. The students handbook of British hepatics. Sumfield, London. U6U pp. 93 APPENDIX I Check l i s t of plant species CHECK LIST OF SPECIES Trees Abies amabilis (Dougl.) Forbes Chamaecyparis nootkatensis (Lamb.) Spach. Malus diversifolia Anderson Picea sitchensis (Bong.) Carr. Pinus contorta Dougl. Pinus monticola Dougl, Taxus brevifolia Nutt. Thuja plicata Don. Tsuga heterophylla (Raf.) Sarg. Shrubs Empetrum nigrum L. Gaultheria shallon Pursh. Kalmia polifolia Wang. Ledum groenlandicum Oedr. Linnaea borealis L. Menziesia ferruginea Smith Myrica gale L, Oxycoccus quadripetalus Gilib. Vaccinium ovalifolium Smith Vaccinium ovatum Pursh. Vaccinium ullginosum L. Vaccinium vitis-idaea L. subsp. minus (Lodd.) Hult. Agrostis aequivalvis (Trin.) Trin. Agrostis diegoensia Vasey Apargidium boreale (Bong.) T. & G. Blechnum spicant (L.) Roth Boschniakia hookeri Wlprs, Calamagrostis nutkaensis (Presl.) Steud. CareK canescens L. Carex obnupta Bail Carex pauciflora Lightf. Carex pluriflora Hulten Coptis asplenifolia Salisb. Coptis trifoliata (L.) Salisb, Cornus canadensis L. Deschampsia caespitosa (L,) Beauv. Drosera rotundifolia L. Vaccinium parvifolium Sm, Herbs Gentiana douglasiana Bong, Gentiana sceptrum Griseb. Goodyera oblongifolia Raf. Habenaria saccata Greene Jiincus oreganus Wats. Listera cordata (L.) R, Br, Lycopodium clavatum L, Lysichitum americanam Hulten & St. J. Maianthemum dilatatum (Wood) Nels, & Macb. Nephrophyllidium crista-galli (Menzies) Gilg. Polypodium glycyrrhiza D, C. Eaton Pteridium aquilinum (L.) Kuhn. Rhynchospora alba (L.) Vahl, Sanguisorba microcephala Presl. Scirpus caespitosus L. Tofieldia occidentalis S.Wats. Trientalis arctica Fisch. Veratrum viride Ait. Bryophytes Antitrichia curtipendula (Hedw.) Brid. Aulacomnium palustre Schwaeg, Bartramia pomiformis (L.) Hedw. var, crispa Bazzania ambigua (Lindenb.) Trev. Blepharstoma trichpphyllum (L.) Dum. Calypogeia trichomanis (L.) Corda Cephalozla bicuspidata (L.) Dum. Cephalozla lammersiana Spruce Dicranum bergeri Bland. Dicranum fusce'scens Turn. Dicranum scoparium Hedw, Diplophyllum albicans (L.) Dum. Eurhynchium oreganum (Sull.) Jaeger & Sauerb. Eurhynchium stokesii (Turn.) B. & S. Frullania nisquallensis Sull. Herberta adunca (Dicks.) Gray Hookeria lucens (Hedw.) Sm. Hylocomium splendens B. & S. Hypnum circinale Hook. Isothecium stoloniferum (Hook.) Brid. Lepidozia reptans (L.) Dum. Macrodiplophyllum plicatum (Lindb.) H. Perss, Metzgeria conjugate Lindb. Mnium glabrescens Kindb. Mylia taylori (Hook.) S. F, Gray Neckera douglasii Hook, Pellia epiphylla ( L . ) Corda Plagiochila asplenioides (L.) Dum. Plagiothecium undulatum B. & S, 96 Pleurozlum scbreberi (Brid.) Mitt. Polytrichum commune L. Polytrichum strictum Smith Pore11a navicularis (Lehm. & Lindenb.) Lindb. R'adula bolanderi Gottsche Rhacomitrium lanuginosum Brid. Rhytidiadelphus loreus (L., Hedw.) Warnst. Riccardia sp. S. F, Gray Scapania bolanderi Aust. Sphagnum capillaceum (Weiss.) Schrank Sphagnum compactum De Cand. Sphagnum fuscum Klingg, Sphagnum mendocinum Sull, & Lesq, Sphagnum papillosum Lindb, Sphagnum recurvum Beauv. Sphagnum tenellum Ehrh. Sphenolobus minutus (Crantz) Steph, Tortella tortuosa (L. Turn.) Limpr, Lichens Cladonia arbuscula (Wallr.) Rabenk. subsp. beringiana Ahti Cladonia bellidlflora (Ach,) Schaer. Cladonia crispata (Ach.) Fw. Cladonia pacifica Ahti. Cladonia rangiferina (L.) Web. Cladonia uncialis (L.) Web, f. turgescens Del. Igmadophila ericetorurn (L.) Zahlbr. Parmelia physodes L. Sphaerophorus globosus (Huds.) Vain. Usnea pj.icnta (L.) Wigg, 97 .APPENDIX II Explanation and Legend for Synthesis tables Synthesis tables I - VI I Explanation and Legend for Synthesis tables. Species ratings are given by three figures (e.g. 4.8.3) which represent species significance, sociability and vigor. The following scales were used for floristic evaluation: Species significance Corresponding cover value + solitary, with small dominance 1. seldom, with small dominance 2. very scattered, with small dominance 3. scattered, with small dominance 4. often, with l/20 to l/lO dominance 5. often, with lAO to l/5 dominance 6. any number, with l/5 to l/3 dominance 7. any number, with l/3 to l/2 dominance 8. any number, with l/2 to 3A dominance 9. any number, with dominance more than 3/4, but less than complete 10. any number, with complete dominance Sociability + individual, sociability none 1. up to 4 x 4 cm 2. 25 x 2$ cm2 3. 50 x 50 cm2 4. 1/3 - 2/3 m2 5. 1 - 2 m2 6. 5 m2 7. 25 - 50 m2 8. 100 m2 9. 200 - 250 m2 10. at least 500 m2 .1 .5 1 5 10 20 33 50 75 95 Vigor 0 - none + -1 -2 -3 -poor fair good excellent Species are grouped by sublayer, in decreasing order of presence and total cover degree. Sublayer abbreviations are as described in Methods (Chapter III). Five presence classes are used: V - species which occur in 81 - 100$ of the plots (constants) IV - species which occur in 6 l - Q0% of the plots i n - species which occur in hX - 60% of the plots II - species which occur in 21 - k0% of the plots I - species which occur in 20% or less of the plots (sporadics) Cover value for each species was calculated by the following formula cover value = sum of cover value (%) for a species x 1 Q Q % number of plots in particular unit Addendum: In a l l synthesis tables, Agoseris glauca should read Apargidium boreale. SYNTHESIS TABLE I: CARICETUM PUR I FLORAE Number of p lots P l o t number P lo t s i ze (meters 2 ) Extent of type (meters?) Date Percent cover Vegetation: l a ye r : B2 C Oh Humus decaying eood L i f e form B layer D layer Sphagnum recurvum Sphagnum papillosum Total species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 48 41 39 38 61 52 51 47 46 45 44 37 59 53 50 49 43 42 54 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 \u00E2\u0080\u00A2 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 50 12 100 50 35 15 10 80 20 150 25 50 40 12 50 40 25 12 12 24/7/ 22/7/ 22/7/ 22/7/ 30/7/ 25/7/ 25/7/ 24/7/ 24/7/ 23/7/ 23/7/ 22/7/ 28/7/ 25/7/ 25/7/ 25/7/ 23/7/ 23/7/ 25/7/ 64 64 64 64 64 64 64 64 64 64 64 64 64 64 64 64 64 64 64 0 2 0 0 0 0 0 0 0 0 0 0 0 0 8 0 0 0 0 25 40 30 20 20 30 45 45 20 35 25 25 25 50 50 35 35 25 65 35 75 95 90 5 5 6 70 20 25 60 80 2 5 0 2 3 35 5 100 100 100 100 100 100 98 100 98 100 97 100 100 100 100 100 100 100 100 0 0 0 0 0 0 2 0 2 0 3 0 0 0 0 0 0 0 0 Sub-layer 1 Pn Vaccinium ullginosum 2 2 Pn Kalmia p o l i f o l i a 2 C l aye r 3 H Carex p l u r i f l o r a 6.2.1 4 H Sangulsorba microcephala 5 H Agoseris glauca 1.+.1 6 H Carex obnupta 7 H Gent!ana sceptrum 7.2.2 1.1.1 8.3.2 3.2.2 7.3.2 7.3.2 6.3.2 5.2.1 2.2.1 9.4.3 2.2.1 7.5.2 3.2.1 1.+.1 9.5.3 1.1.1 2.+.1 Species Con- Cover s i gn i f i cance Vigor stancy value H R \u00C2\u00BB R 0.1 0-2 1.0 \u00E2\u0080\u0094 I 5 0.03 0-1 1.0 \u00E2\u0080\u0094 I 0.5 5.2.2 3.2.2 3.2.2 1.1. 7.3.2 1.+.1 2.1.1 2.1.1 7.4.2 7.4.2 6.2.1 7.3.2 6.3.2 7.4.3 6.3.2 8.4.2 8.5.2 7.3.2 7.3.2 6.2.2 9.5.3 6.8 5-9 2.0 1-3 V 4936 2.+.1 2.2.1 5.2.1 3.2.1 3.2.1 2.2.1 1.2.1 4.2.1 2.0 0-5 1.1 1-2 iy 392 2.2.1 4.2.1 0.6 0-4 1.2 1-2 III 92 +.+.2 0.2 0-3 2.0 1 27 2.+.1 0.2 0-2 1.0 1 8 2.1.1 8.4.2 6.2.1 5.2.2 8.3.2 8.4.3 1.+.1 2.1.1 2.1.1 4.0 0-9 1.8 1-3 IV 3013 2.1.1 1.+.1 1.1.1 2.1.1 3.2.1 1.+.1 1.+.1 1.+.1 1.1.1 1.4 0-7 1.0 0.5-2 IV 334 4 4 4 4 4 4 3 3 3 3 3 3 2 L i f e f o ra B H Pn Proportion by: Species: Number 2.0 5.0 2.0 i 22.2 55.5 22.2 Total Cover: cover 3347.0 5455.0 5.5 t 38.0 61.9 0.06 101 SYNTHESIS TABLE I I. SCIRPETO-SPHAGNETUM HEHDOCIHI Number of p lots 1 2 3 4 5 6 7 8 9 10 P lo t number 99 98 94 97 103 101 93 95 102 100 P lot s i ze (meters^) 4 4 4 4 4 4 4 4 4 4 Extent of type (acres) 1 1 1/4 1 1 1 1/3 1/4 1 1 Date 4/9/ 4/9/ 3/9/ 4/9/ 4/9/ 4/9/ 3/9/ 3/9/ 4/9/ 4/9/ 64 64 64 64 64 64 \u00E2\u0080\u00A264 64 64 64 Percent coverage Vegetation: layer: 8^ 0 0 0 0 5 0 0 0 0 0 82 5 5 5 15 5 5 5 5 8 3 Total 8 5 5 5 15 10 5 5 5 8 3 C 55 55 50 50 60 65 40 40 65 50 Oh 80 80 60 80 70 75 60 70 75 65 Humus 100 100 100 100 100 100 100 100 100 100 Oecaylng good 0 0 0 0 0 0 0 0 0 . 0 Species Con- Cover L i f e Sub- S ign i f icance Vigor stancy value form B layer layer HI R 11 R 1 Pn Ka la ia pol l f o l l a 1 3.3.2 0.3 0-3 2 2 2.+.1 2.+.1 1.+.1 4.2.1 4.3.2 2.+.1 2.+.1 1.+.1 3.2.1 1.+.1 2.2 1-4 1.1 1-2 V 295 2 Ch Oxycoccus quadrl petal us 2 3.2.2 2.2.2 2.+.2 1.+.1 2.2.2 1.+.1 2.+.1 2.+.1 2.1.2 1.1.1 1.8 1-3 1.5 1-2 V 130 3 Pn Vaccinium ullglnosum 2 3.2.2 2.2.2 1.1.1 4.2.2 2.1.1 1.1.1 1.2.1 1.4 0-4 1.4 1-2 IV 185 * Ch Empetrum nigrum 2 1.2.1 1.2.1 3.2.1 +.+.+ 0.4 0-3 0.8 +-1 III 66 5 Pn Ledum groenlandicum 2 1.+.1 1.+.1 +.+.1 1.+.1 0.4 0-1 1 II 35 6 Pm Thuja p l l c a t a 2 +.+.1 2.+.1 +.+.+ 0.3 0-2 0.8 +-2 II 22 7 Pn Chamaecyparis nootkatensis 2 1.+.1 1.0 0-1 1 I 5 8 Pm Pinus contorta 2 +.+.1 0.05 0-+ 1 \u00E2\u0080\u0094 I 1 C layer 9 H Sclrpus caespitosus 8.5.2 8.5.2 8.4.2 8.5.2 8.5.3 9.5.3 8.3.3 7.3.2 8.5.3 8.5.2 8.0 7-9 2.4 2-3 V 7450 10 H Agrost is aequlvalv ls 4.2.2 4.2.2 4.2.2 4.2.2 4.2.2 4.3.2 4.2.2 4.2.2 3.2.2 3.2.2 3.8 3-4 2.0 V 900 11 H Sanguisorba mlcrocephala 2.+.1 1.+.1 3.1.1 2.+.2 3.2.2 3.2.3 4.2.2 2.1.1 3.2.2 2.+.1 2.5 1-4 1.6 1-3 V 345 12 6 T r l e n t a l l s a r c t l ca 1.+.1 2.+.1 3.+.1 2.+.1 +.+.1 1.+.2 2.+.2 2.+.1 2.+.2 2.+.1 1.8 +-3 1.3 1-2 V 121 13 T Gentiana douglaslana 2.+.1 1.+.1 1.+.1 +.+.+ 1.+.+ 1.+.1 2.+.1 3.+.2 1.+.1 2.+.1 1.5 1-3 1.0 +-2 V 106 14 H Orosera r o t und l f o l l a 1.+.1 2.1.1 1.+.1 1.+.1 1.+.1 1.+.1 2.+.1 1.+.1 1.1 0-2 1.0 V 55 15 G Tof 1e1d1 a occ iden ta l ! : 1.+.1 +.+.1 1.+.1 +.+.1 2.+.1 2.+.1 1.+.1 +.+.1 1.+.1 +.+.2 1.1 +-2 1.1 \u00E2\u0080\u00A2-? V 48 16 H Rhynchospora alba 4.2.2 2.1.1 4.3.2 4.2.2 4.2.2 1.2.2 4.2.2 3.2.2 2.7 0-4 1.5 1-2 IV 665 17 H Carex canescens 3.2.2 2.2.2 3.2.2 2.2.2 3.2.2 +.2.2 4.2.2 2.2.1 2.0 0-4 1.9 1-2 IV 285 18 H Carex obnupta 2.2.2 +.+.1 +\u00E2\u0080\u00A2.+.+ 2.2.2. 2.2.2 0.7 0-2 1.5 1-2 III 32 19 H Carex p i u r l f l o r a 1.+.1 2.+.1 4.2.2 3.2.2 1.0 0-4 1.5 1-2 II 165 20 H Gentiana sceptrum 2.+.1 3.2.2 +.+.1 0.6 0-3 1.3 1-2 II 61 21 H Agoserls glauca 1.+.2 3.2.1 0.4 0-3 1.5 1-3 I 55 22 H Coptls a s p l e n l f o l l a 2.1.2 0.2 0-2 2.0 I 10 23 H Coptls t r i f o l l a t a +.+.1 0.05 0-+ 1.0 \u00E2\u0080\u0094 1 5 D laver 24 B Sphagnum papillosum h 7.3.2 8.5.2 8.3.2 8.5.2 8.5.3 8.5.2 8.5.3 8.4.3 8.5.2 7.1 0-8 2.3 2-3 V 6500 25 B Sphagnum mendocinum h 3.2.2 4.2.2 +.+,+ 3.2.2 4.2.2 3.2.2 1.1.1 4.3.2 8.5.2 4.2.2 3.5 *-8 1.8 *-2 V 1306 26 B Sphagnum capil laceum h 3.1.2 3.2.2 2.1.1 3.2.2 3.2.2 2.2.1 2.1.1 2.1.1 2.1.1 2.1.2 2.4 2-3 1.5 1-2 v 260 27 L Cladonla rang l fer lna h 2.1.1 1.1.1 1.1.1 1.1.1 0.6 0-2 0.9 +-1 III 30 28 B Rhacomitrium lanuglnosum h 1.1.2 1.2.2 0.2 0-1 2.0 II 10 d i 2.2.1 0.2 0-2 1.0 II 10 29 B Cephalozla blcuspldata h 1.1.2 1.1.2 0.2 0-1 2.0 II 10 d> 1.+.1 0.1 0-1 1.0 11 5 30 L Cladonla padf1ca h 5.3.2 1.1.+ 0.6 0-5 1.3 +-2 1 205 31 B Bazzanla amblgua d> +.+.1 0.05 O-t . 1.0 \u00E2\u0080\u0094 1 1 Total species 22 21 21 20 18 18 18 17 16 15 L i f e form H B Pn L Ch G Pm \u00E2\u0080\u00A2 T Proportion by: Species: Number 12.0 6.0 4.0 2.0 2.0 2.0 2.0 1.0 2 38.8 19.4 13.0 6.5 6.5 6.5 6.5 3.2 Total Cover: cover 6026.0 8107.0 275.0 235.0 196.0 169.0 23.0 106.0 t 63.8 32.2 1.1 0.9 0.8 0.7 0.1 0.4 SYNTHESIS TABLE III. OJYCOCCErO-SPHAGNETUM PflPILLOSAE Number of plots 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Plot number 55 56 50 66 19 18 17 58 57 64 25 20 63 67 65 Plot size (meters?) 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 Extent of type (ecres) * 4 4 4 2 1 1 4 1/100 4 1 3 4 4 4 Date 26/7/ 28/7/ 30/7/ 31/7/ 8/6/ 8/6/ 3/6/ 28/7/ 28/7/ 31/7/ 12/6/ 8/6/ 31/7/ 31/7/ 31/7/ 64 64 64 64 64 64 64 64 64 64 64 64 64 64 64 Percent coverage Vegetetion: layer: Bs 15 15 5 3 2 5 5 8 15 7 5 5 8 2 2 C 30 50 60 55 45 35 40 50 45 60 35 40 60 45 50 Oh \u00E2\u0080\u00A2 98 9B 95 98 98 95 98 98 98 100 100 100 99 98 100 \"d. 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 Total 0 9B 98 95 98 98 97 98 98 98 100 100 100 99 98 100 Humus 100 100 100 100 100 98 100 100 100 100 100 100 100 100 100 Decaying Wood 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Species Con- Cover Life Sub- Significance Vigor stancy value form 6 laver layer \u00E2\u0080\u00A2 II a R 1 Ch Oxycoccus quadripetalus 2 3.2.2 3.2.2 2.2.2 2.1.2 2.1.1 2.1.1 2.2.1 3.2.2 3.1.2 3.1.2 3.2.2 4.2.2 3.1.2 2.1.2 3.2.2 2.8 2-4 1.8 1-2 V 370 2 Pn Kalmia polifolia 2 3.2.1 3.3.1 2.+.1 2.2.1 2.2.2 1.2.1 3.2.1 3.1.1 2.*.2 3.+.1 2.2.1 1.*.1 1.*.+ 1.+.1 2.0 1-3 1.0 <-2 V 217 3 Pn Ledum groenlandicum 2 3.2.1 3.2.2 1.+.1 l . t . t 1.*.1 3.2.1 0.8 0-3 1.1 +-2 III 111 4 Pn Vaccinium ulIglnosum 2 3.2.2 2.2.2 0.3 0-3 2.0 \u00E2\u0080\u0094 1 40 5 Pn Myrica gale 2 3.2.1 3.2.1 0.4 0-3 1.0 \u00E2\u0080\u0094 1 40 6 Ch Empetrum nigrum 2 + + + 0.2 0-1 + \u00E2\u0080\u0094 1 7 7 P> Pinus contorta 2 +.+.\u00C2\u00AB\u00E2\u0080\u00A2 0.03 0-* + I 0. S Pm Thuja plicata 2 0.05 O-t + \u00E2\u0080\u0094 1 0. C laver 9 H Agoserls glauca 3.2.2 3.2.2 4.2.2 4.2.2 7.3.2 7.2.2 6.3.2 5.2.2 4.2.3 5.3.2 2.1.2 7.3.2 4.2.2 4.0 0-7 2.1 2-3 V 1827 10 H Agrostis aequlvalvls 4.2.2 4.2.2 4.2.2 4.2.2 1.2.2 3.2.2 4.2.2 5.2.3 3.2.2 2.*.1 4.2.2- 4.2.2 4.2.2 3.0 0-5 2.0 1-3 y 743 11 s Trlentalls arctlce 3.1.2 5.1.2 4.*.2 3.\u00C2\u00BB.2 2.*.2 1.+.1 2.1.2 3.+.1 1.+.1 Z.+.2 3.1.2 3.+.1 3.1.2 3.+.2 2.+.2 2.7 1-5 1.7 1-2 y 467 12 H Orosera rotundifolia 2.1.2 2.*. 2 3.+.2 3.2.2 3.2.2 2.-..1 3.1.2 3.t.2 3.1.2 2.1.2 2.1.2 2.1.2 3.1.2 1.-\u00C2\u00BB.2 3.1.2 2.5 1-3 1.9 1-2 y 310 13 G Tofleldle occidentals 1.*.1 \u00E2\u0080\u00A21.+.2 3.1-.2 1.+.1 1.*.1 1.*.2 3.2.2 2.->.1 3.*.2 1.1.1 1.3 0-3 1.4 1-2 v 133 14 H Sangulsorba mlcrocephala 4.2.2 4.2.2 \u00E2\u0080\u00A2.+.1 3.2.2 2.-0 2.1.1 3.2.2 4.2.2 2.1.1 7.2.2 4.2.2 3.2.2 2.6 0-7 1.7 1-2 IV 707 15 T Gentiana douglaslana 1.+.1 1.+.1 3.+.2 3.1.2 3...1 4.1.2 1.+.1 3.1.2 3.1.3 1.5 0-4 1.6 1-3 IV 247 16 H Carex pluriflora 4.2.2 4.3.1 4.3.2 6.3.2 5.2.2 6.3.1 5.3.2 9.3.2 7.3.2 3.3 0-9 1.8 1-2 i n 1B70 17 H Carex obnupta 3.2.2 2.2.2 3.+.1 3.2.1 !.*.* 5.2.2 3.2.2 3.2.2 1.5 0-5 1.5 \u00C2\u00AB-2 i n 310 18 H Rhynchospora alba 4.3.2 3.3.2 5.2.2 4.2.2 1.1 0-5 2.0 \u00E2\u0080\u0094 II 300 19 H Scirpus caespitosus 1.2.2 0.07 0-1 2.0 \u00E2\u0080\u0094 i 3. 20 B Sphagnum papillosum h 9.6.3 9.5.3 9.5.3 9.5.2 9.7.3 9.7.3 9.7.3 9.5.3 9.5.3 9.4.3 9.6.3 9.7.3 9.5.3 9.5.2 9.5.3 9.0 9 2.9 2-3 y 9500 21 B Sphagnum capillaceum h 1.1.1 1.1.1 2.2.2 1.2.2 2.2.2 1.1.2 1.1.1 2.2.2 3.2.2 1.1.2 4.2.2 4.1.1 1.5 0-4 1.7 1-2 IV 207 22 B Sphagnum recurvum h 1.1.1 1.1.1 2.3.2 1.1.1 2.2.2 0.5 0-2 1.4 1-2 II 23 23 B Sphagnum tenel lum h 5.3.3 0.3 0-5 3.0 \u00E2\u0080\u0094 I 133 2t B Sphagnum fuscum h 2.2.2 3.3.2 0.3 0-3 2.0 \u00E2\u0080\u0094 i 40 25 S Cephalozia blcuspidata h 2.1.2 0.13 0-2 2.0 \u00E2\u0080\u0094 i 7 26 B Polytrichum comnune h 1.2.2 0.1 0-1 2.0 \u00E2\u0080\u0094 1 3. Total species 18 17 15 14 14 14 14 13 13 12 12 12 11 9 9 Life form H B Pn Ch 6 Pm T Proportion by Species: Number 1 8 30.8 7 26.9 4 15.4 7.7 2 7.7 2 7.7 1 3.8 Total cover: cover % 6070.3 34.4 9913.3 56.3 406 2.3 377 2.1 600 3.4 1.4 247 1.4 103 SYNTHESIS TABLE IV. LEDETO-SPHAGNETUM CAPILLACEI Number of p lots 1 2 3 4 5 6 7 8 9 10 P lot number 85 69 68 91 90 86 84 70 87 26 P lot s i ze (meters^) 4 4 4 4 4 4 4 4 4 4 Extent of type (acres) 1/2 1/5 1/10 1/20 1/100 1/20 1/5 1/10 1/20 1 Date 27/8/ 4/8/ 4/8/ 28/8/ 28/8/ 27/8/ 27/8/ 4/8/ 27/8/ 12/6/ 64 64 64 64 64 64 64 64 64 64 Percent coverage Vegetation: l ayer : S-| 15 0 0 15 0 0 15 0 0 0 B 2 20 25 30 10 15 20 35 20 20 15 Total B 30 25 30 20 15 20 40 20 20 15 C 65 70 65 65 40 80 40 60 70 50 Oh 95 96 98 95 100 100 100 98 100 100 Od. 5 0 0 2 0 0 0 0 0 0 Total D 100 96 98 98 100 100 100 98 100 100 Humus 95 100 100 98 100 100 100 100 100 100 Decaying Wood 5 0 0 2 0 0 0 0 0 0 Species Con- Cover L i f e Sub- S igni f icance Vigor stancy value form B layer layer M R 11 R 1 Pn Ledum groenlandicum 1 5.2.2 5.2.1 4.3.2 2.0 0-5 1.7 1-2 2 5.2.2 5.3.2 5.2.2 4.2.1 3.4.1 5.2.1 6.3.2 3.2.1 4.2.1 4.3.1 4.4 3-6 1.4 1-2 V 1530 2 Pn Kalmia po l l f o l l a 2 3.+.1 3.2.2 3.2.1 3.4.1 3.4.1 4.2.1 4.2.1 2.4.2 3.2.2 2 4.1 3.0 2-4 1.3 1-2 V 520 3 Ch Oxycoccus quadrlpetal us 2 3.2.2 3.2.2 3.2.2 3.2.2 3.2.2 4.2.2 3.2.2 3.2.2 3.2.2 2.2.1 3.0 2-4 1.9 1-2 V 510 4 Ch LInnaea boreal i s 2 3.2.1 1.1.1 3.1.2 2.2.1 3.2.1 1.2.1 3.2.1 1.6 0-3 1.1 1-2 IV 220 5 Ch Empetrum nigrum 2 2.2.1 4.3.2 2.2.1 1.2.1 1.1.1 1.0 0-2 1.2 1-2 111 130 6 Pn Chamaecyparis nootkatensis 1 3.+.1 0.3 0-3 1 2 4.4.1 4.3.1 0.5 0-4 1 \u00E2\u0080\u0094 l l 151 7 Pn Myrica gale 2 2.2.1 2.4.1 0.4 0-2 1 \u00E2\u0080\u0094 I 40 C layer 8 H Carex obnupta 8.4.2 6.3.2 7.4.2 8.5.2 7.5.2 9.6.3 6.3.2 7.3.2 8.5.2 8.4.3 7.4 6-9 2.2 2-3 V 6110 9 H Agoserls glauca 4.2.2 4.2.2 5.2.2 4.2.2 3.2.2 4.3.2 4.2.2 4.3.2 6.2.2 3.8 0-6 2.0 \u00E2\u0080\u0094 V 1180 10 H Drosera ro tund! fo l i a 3.+.2 3.1.2 2.4.2 1.4.1 1.4.1 2.4.2 1.4.1 3.4.2 2.4.1 2.1.2 2.0 1-3 1.6 1-2 V 205 11 G T r i en t a i l s a r c t l c a 2.t.2 1.4.2 1.4.1 2.4.2 2.4.2 1.4.2 2.4.2 3.4.2 3.4.1 1.4.1 1.8 1-3 1.7 1-2 V 144 12 H Agrost is aequlvalv ls 3.2.2 1.2.2 2.4.2 4.3.2 3.2.2 2.4.2 4.2.2 2.2.2 2.1 0-4 2.0 \u00E2\u0080\u0094 IV 335 13 Ch Cornus canadensis 3.2.2 3.1.1 1.4.1 0.7 0-3 1.3 1-2 III 20 14 T Gentiana douglasiana 1.4.2 1.4.2 2.4.2 0.4 0-2 2.0 \u00E2\u0080\u0094 II 20 15 G To f i e l d l a occidental 1s 1.+.1 1.4.1 1.4.1 0.3 0-1 1.0 II 15 16 G Sanguisorba microcephal a 2.2.2 4.2.2 0.6 0-4 2.0 \u00E2\u0080\u0094 1 110 17 H Carex p l u r i f l o r a 3.2.1 3.2.1 0.6 0-3 1.0 \u00E2\u0080\u0094 1 100 18 H Scirpus caespltosus 1.4.1 3.2.2 0.4 0-3 1.5 1-2 1 55 19 H Deschampsia caespltosa 3.2.2 0.3 0-3 2.0 \u00E2\u0080\u0094 1 50 20 G Maianthemum di latatum 1.+.1 4.4 .4 0.2 0-1 0.8 +-1 1 6 21 H Coptis a s p l e n i f o l l a 1.4.1 0.1 0-1 1.0 \u00E2\u0080\u0094 1 1 0 layer 22 8 Sphagnum papillosum h 5.2.1 3.1.2 3.1.1 4.2.2 8.4.2 6.2.2 8.3.2 7.3.2 9.5.3 6.3.2 5.9 3-9 1.9 1-3 V 4010 23 B Sphagnum capil laceum h 3.2.2 3.7.2 1.1.1 7.4.3 5.2.2 7.3.3 7.3.3 3.3.2 6.2.2 6.3.2 4.8 1-7 2.2 1-3 V 2515 24 8 Sphagnum mendocinum h 2.2.2 3.2.2 8.4.3 6.2.2 6.2.2 3.2.2 3.2.2 3.2.2 3.4 0-8 2.1 2-3 IV 1620 25 e Sphagnum fuscum h 9.4.3 7.3.2 8.3.3 7.3.3 4.2.2 3.5 0-9 2.6 2-3 III 2800 26 B Sphagnum tenel 1 um h 5.2.2 3.2.2 4.2.2 3.2.2 8.5.3 2.3 0-8 1.1 2-3 III 1150 27 B Dicranum bergeri h 1.2.2 2.2.2 4.2.3 0.7 0-4 2.3 2-3 II 115 d . 2.1.1 0.2 0-2 1.0 28 8 Pleurozium schreberi h 2.2.2 2.1.2 3.2.2 1.2.2 0.8 0-3 2.0 \u00E2\u0080\u0094 II 75 d> 3.1.2 0.3 0-3 2.0 29 B Aulacomnium palustre h 2.+.2 2.4.2 1.1.1 0.5 0-2 1.6 1-2 II 25 30 B Polytrlchum commune h 4.3.2 5.3.2 0.9 0-5 2.0 \u00E2\u0080\u0094 1 300 31 B Sphagnum recurvum h 2.2.2 4.2.2 0.6 0-4 2.0 \u00E2\u0080\u0094 1 110 32 B Bazzania ambigua d i 1.1.1 0.1 0-1 1.0 \u00E2\u0080\u0094 1 5 33 B Dicranum scoparium da 1.1.1 0.1 0-1 1.0 \u00E2\u0080\u0094 1 5 34 B R iccard la sp. di 1.1.2 0.1 0-1 2.0 \u00E2\u0080\u0094 1 5 Total species 23 21 20 19 > 18 16 15 15 14 12 L i f e form B H Pn Ch G T Proportion by Species: Number 13 9 4 4 3 1 % 38.2 26.5 11.8 11. 3 8.8 2.9 Total cover: cover 12795 8146 2195 880 165 20 , % 52.9 33.6 9.1 3.6 0.7 0.1 SYNTHESIS TABLE V. PINETO-SPHAGNETUM CAPILLACEI Life Sub- Pres- Cover form A layer layer ence value 1 Pm Pinus contorta 2 3 IV 2005 2 Pm Thuja plicata 3 I 167 3 Pm Chamaecyparis nootkatensis 3 I 32 4 Pm Tsuga heterophylla 3 I 1 B layer Pm Pinus contorta 1 V 2166 5 Pn Ledum groenlandicum 1 V 1551 6 Ch Empetrum nigrum 1 2 V 1500 7 Pn Kalmia polifolia 1 V 1073 8 Ch Oxycoccus quadripetal us 2 V 314 9 Ch Vaccinium vitis-idaea 2 IV 176 10 Ch Linnaea borealls 2 IV 135 11 Pn Gaultheria shallon 1 2 III 297 Pn Chamaecyparis nootkatensis 1 II 1128 Pm Thuja plicata 1 II 247 12 Pn Vaccinium uliginosum 2 II 177 Pm Tsuga heterophylla 1 2 II 8 13 Pn Myrica gale 1 I 1146 14 Pn Vaccinium parvifolium 1 I 22 15 Pn Vaccinium ovatum 1 I 15 16 Pm Picea sitchensis 2 I 2 17 Pn Vaccinium oval ifol 1um '1 I 2 18 Pn Iflenziesia ferruginea 1 I 0.2 C layer 19 H Carex obnupta V 2392 20 H Agoserls glauca V 1005 21 G Trientalls arctica V 156 22 H Drosera rotundlfolia V 63 23 Ch Cornus canadensis IV 240 24 H Sanguisorba microcephala IV 215 25 G Maianthemum dilatatum IV 60 26 H Carex pluriflora III 290 27 H Agrostis aequivalvls III 231 28 H Blechnum splcant III 175 29 III Coptls asplenifolia III 78 30 T Gentiana douglasiana II 50 31 G Lysichitum americanum II 31 32 G Tofieldla occidentals II 13 33 ti Carex canescens I 144 34 H Scirpus caespitosus I 56 35 H Coptls trifoliata I 34 36 H Gentiana sceptrum I 32 37 H Deschampsia caespltosa I 21 36 G Nephrophyllldium crista-galU I 14 39 H Calamagrostis nutkaensls I 5 40 H Carex pauciflora I 5 41 Ch Lycopodium clavatum I 5 42 G Habenaria saccata I 3 43 H Rhynchospora alba I 2 44 G Boschniakia hookari I 1 45 G Pteridium aquillnum I 1 46 G Veratrum virlda I 0.2 Life Sub- Pres- Cover form D layer layer ence value 47 B Sphagnum papillosum h V 2718 48 B Sphagnum capillaceum h V 1634 da 49 B Sphagnum recurvum h IV 1257 50 B Sphagnum mendocinum h III 882 51 B Pleurozium schreberi h III 271 dw 52 B Dicranum scoparium h dm III 115 53 B Rhytidladelphus loreus h dw III 98 54 B Aulacomnium palustre h III 56 55 B Sphagnum fuscum h II 821 dt 56 L Cladonia crispata h II 240 57 L Cladonia pacifica h II 157 di 58 B Dicranum bergeri h II 119 dw 59 L Cladonia rangiferina h II 111 dw 60 B Hylocomium splendens h dw II 107 61 B Bazzania ambigua h dw II 13 62 B Rhacomitrium lanuginosa h dw 1 180 63 B Herberta adunca dw 1 55 64 B Sphagnum tenellum h 1 39 65 B Eurhynchium oreganum h dw 1 33 66 B Polytrichum strictum h 1 28 dw 67 B Isothecium stoloniferum dw 1 18 68 B Cephalozla bicuspidata h 1 15 dw 69 B Dicranum fuscescens dw 1 15 70 B Polytrichum commune h 1 14 71 B Plagiothecium undulatum h da 1 8 72 B Mnium glabrescens h da 1 7 73 L Cladonia bellldiflora . da 1 6 74 L Cladonia unci al Is da 1 5 75 B Diplophyllum albicans da 1 3 76 L lemadophila ericetorum da 1 3 77 B Riccardla sp. da 1 3 78 B Eurhynchium stokesil dw 1 - 1 79 B Plagiochila asplenioides da 1 1 80 B Bantramia pomiformis da 1 1 E layer Bryophytes S lichens 81 E Frullania nlsquallensis A D B c V 635 82 E Scapania bolanderl B C IV 150 E Dicranum scoparium A< D D c IV 138 E Herberta adunca A \u00E2\u0080\u00A2 D C II 61 E Isothecium stoloniferum AB II 48 C Life form E layer (continued) 83 E Antitrlchia curtipendula E Bazzanla ambi gua E Olplophyllun albicans E Pleuroziua schreberl E Rhacoraitrlum lanuginosum 84 E Parmella physodes 85 E E 86 E E E 87 E 88 E E Hylocomium splendens E Riccardia sp. E Bartramia poniformis E Mnfum glabrescens Life form Proportion by Species: Number I Sub- Pres- Cover layer ence value A B C II 40 6 C II 19 e C II 16 A C I 44 C I 17 A B I 14 C A I 14 C I 13 C I 11 C I 7 C I 6 B I 6 B I 5 C C I 3 C I 3 C I 3 C I 2 E B H Pn 38 28 16 10 32.0 23.5 13.4 8 Usnea plicata Cephalozia b 1.1.2 3.2.2 3.1.1 3.2.2 2.1.1 3.2.1 1.1.2 1.5 0-3 1.6 1-2 39 8 Pleurozium schreberl h 5.3.3 3.2.2 3.2.2 4.2.2 3.1.1 4.2.2 4.2.1 3.2.2 1.1.1 2.7 0-5 1.8 1-3 V 650 d i 2.2.2 2.1.2 3.2.2 3.1.2 3.1.1 2.1.2 3.2.2 1.6 0-3 1.9 1-2 40 L Cladonia crlspata h 2.1.2 5.2.3 7.3.3 3.2.2 3.1.2 3.3.2 3.3.2 2.4 0-7 2.3 2-3 IV 827 41 B Sphagnum papillosum h 5.3.2 4.2.2 3.2.2 3.2.1 6.2.2 1.2.2 3.3.1 4.3.2 2.6 0-6 1.9 1-2 IV 804 42 8 Rhacomitrium lanuginosum h 4.3.2 6.4.2 3.2.2 1.2.2 4.2.2 3.2.1 3.2.2 2.2 0-6 1.9 1-2 IV 622 d i 3.2.2 4.2.2 1.1.1 2.2.1 0.9 0-4 1.5 1-2 43 B Dicranum scoparium h 3.2.2 1.2.2 2.1.1 3.2.2 4.1.2 3.2.2 4.2.2 2.2.2 2.0 0-4 1.9 1-2 V 440 d i 3.2.2 3.2.1 2.1.1 1.1.1 2.1.1 3.1.2 2.2.2 4.2.2 1.8 0-4 1.5 1-2 44 L Cladonia rangiferina h 3.2.2 4.2.1 4.2.2 3.2.2 3.2.1 1.+.Z 2.2.2 3.3.2 2.1 0-4 1.6 1-2 V 390 d> 2.2.1 2.2.1 3.2.1 1.1.2 0.7 0-3 1.3 1-2 45 B Rhytldiadelphus loreus h 1.1.1 1.2.1 2.2.2 1.1.1 1.1.1 3.1.2 0.8 0-3 1.3 1-2 V 86 d i 1.1.1 2.2.2 1.1.1 1.2.2 0.5 0-2 1.5 1-2 46 B Dicranum bergeri h 3.2.2 3.2.3 2.1.2 3.2.2 4.2.3 4.2.2 2.1.2 1.9 0-4 2.3 2-3 IV 336 47 B Hylocomlum splendens h 2.2.1 3.1.2 0.5 0-3 1.5 1-2 de 2.3.1 2.2.1 1.+.1 3.2.2 3.1.2 2.2.1 1.2 0-3 1.3 1-2 IV 131 48 B Scapania bolanderi d i 1.2.2 2.1.1 2.1.1 1.2.2 1.1.1 2.1.2 1.2.2 1.1.2 1.0 0-2 1.6 1-2 IV 54 49 8 Sphagnum mendocinum h 4.2.2 2.2.2 2.2.2 3.2.2 4.1.1 1.4 0-4 1.8 1-2 III 245 50 8 Aulacomnlum palustre h 2.2.2 1.1.1 2.1.2 1.2.2 3.2.2 0.8 0-3 1.8 1-2 III 72 51 L Cladonia b e l l l d l f l o r a de 1.+.1 1.1.1 1.1.1 1.+.1 1.+.2 0.5 0-1 1.2 1-2 III 22 52 B Eurhynchium oreganum h 1.1.1 1.2.1 0.2 0-1 1.0 \u00E2\u0080\u0094 . d i 2.1.2 2.1.1 0.4 0-2 1.5 1-2 II 27 53 B Cladonia unc 'a l i s h 1.2.3 1.1.3 2.1.3 0.4 0-2 3.0 \u00E2\u0080\u0094 II 18 54 B Plagiothecium undulatum h +.1.+ 0.04 0-4- \u00E2\u0080\u0094 de- +.+.+ 1.+.+ 1.+.1 0.2 0-1 0.7 <-1 II 10 55 8 Sphagnum fuscum li 6.3.2 0.5 0-6 2.0 \u00E2\u0080\u0094 I 300 56 B Sphagnum recurvum h 3.2.2 4.2.2 0.6 0-4 2.0 \u00E2\u0080\u0094 1 136 57 B Polytrichum strictum h 3.2.1 3.2.2 0.5 0-3 1.5 1-2 1 90 d i 2.2.1 0.2 0-2 1.0 \u00E2\u0080\u0094 58 L Icmadophlla erlcetorum d i 1.2.2 2.2.3 0.3 0-2 2.5 2-3 1 13 59 B Polytrichum commune h 2.1.1 1.1.2 0.3 0-2 1.5 1-2 1 13 60 B ttnlum glabrescens d i 2.2.2 0.2 0-2 2.0 \u00E2\u0080\u0094 1 9 61 B Riccardia sp. d i 1.1.2 1.1.1 0.2 0-1 1.5 1-2 1 9 62 B Bazzania ambigua d\u00C2\u00BB 1.1.1 0.09 0-1 1.0 \u00E2\u0080\u0094 1 4 63 B Cephalozia blcuspldata d< 1.1.1 0.09 0-1 1.0 \u00E2\u0080\u0094 1 4 64 B Dicranum fuscescens d i 1.1.2 0.09 0-1 2.0 \u00E2\u0080\u0094 1 4 65 B Eurhynchium stokesll it 1.1.1 0.09 0-1 1.0 \u00E2\u0080\u0094 1 4 66 B Isothecium stolonlferum d i 1.1.1 0.09 0-1 1.0 \u00E2\u0080\u0094 1 4 E layer Bryophytes & lichens 67 E Frul lanla nisquallensis B 2.1.1 2.1.1 3.2.2 4.1.3 3.1.1 3.1.2 1.1.1 4.1.2 3.2.1 3.2.1 3.1.2 2.8 1-4 1.5 1-3 V 477 C 2.1.1 2.1.2 3.2.2 4.1.3 3.1.2 3.1.2 1.1.1 4.1.2 2.2.1 4.2.1 2.5 0-4 1.7 1-3 E Dicranum scoparium A 2.1.1 0.2 0-1 1.0 \u00E2\u0080\u0094 B 3.1.1 2.2.1 0.5 0-3 1.0 \u00E2\u0080\u0094 C 2.2.1 3.1.1 1.1.1 0.5 0-3 1.0 \u00E2\u0080\u0094 II 68 E ' Isothecium stolonlferum C 2.1.1 1.1.1 2.1.1 0.5 0-2 1.0 \u00E2\u0080\u0094 II 22 68 E btacrodiplophyllum plicatum c 2.2.2 1.1.2 2.2.2 0.5 0-2 2.0 \u00E2\u0080\u0094 II 22 E Scapania bolanderi 6 1.1.2 0.09 0-1 2.0 \u00E2\u0080\u0094 C 2.2.2 1.1.2 1.1.1 0.4 0-2 1.7 1-2 II 22 69 E Parmelia physoides B 1.1.2 3.1.2 0.4 0-3 2.0 \u00E2\u0080\u0094 1 50 C 1.1.2 3.1.1 0.4 0-3 1.5 \u00E2\u0080\u0094 E Pleurozium schreber) C 1.1.2 3.1.2 0.4 0-3 2.0 \u00E2\u0080\u0094 1 50 70 E Radula bolanderi C 2.2.2 0.2 0-2 2.0 \u00E2\u0080\u0094 1 9 E Dicranum fuscescens C 1.1.1 0.09 0-1 1.0 \u00E2\u0080\u0094 1 4 E Bazzania ambigua c 1.1.1 0.09 0-1 1.0 \u00E2\u0080\u0094 1 4 E Hylocomlum splendens c 1.+.* 0.09 0-1 + \u00E2\u0080\u0094 1 4 71 E Plagiochl la asplenloldes c 1.+.1 0.09 0-1 1.0 \u00E2\u0080\u0094 1 4 72 E Ant l t r i ch ia curtlpendula c 1.+.+ 0.09 0-1 + \u00E2\u0080\u0094 1 4 73 E Sphaerophorus globosus B 1.1.1 0.09 0-1 1.0 \u00E2\u0080\u0094 1 4 74 E Usnea p l icata A 1.+.+ 0.09 0-1 \u00E2\u0080\u00A2f \u00E2\u0080\u0094 1 4 Vascular plants E Empetrum nigrum c 9.4.2 3.3.2 1.1 0-9 2.0 \u00E2\u0080\u0094 1 1163 E Gaultherla shallon c 5.2.1 2.1.1 0.6 0-5 1.0 \u00E2\u0080\u0094 1 190 E Vaccinium v i t i s - idaea c 4.2.1 2.2.1 0.5 0-4 1.0 \u00E2\u0080\u0094 1 100 E Maianthemum dl latatua c 1.+.1 0.09 0-1 1.0 \u00E2\u0080\u0094 1 4 Total species 43 42 39 38 36 36 36 36 31 29 29 L i fe form B E H G Pn L Ch Pm T Proportion by Species: dumber 24 19 13 7 6 6 5 4 1 1 28.2 22.4 15.3 8.2 7.1 7.1 5.9 4.7 1.2 Total cover: cover 7063 748 3756 328 7971 2642 5593 86 141 { 25.1 2.7 13.4 1.2 28.4 9.4 19.9 0.3 0.5 SYNTHESIS TABLE V. PINETO-SPHAGNETUM CAPILLACEI Var iant : P.-S. c ap i l l a ce i vacciniosum p a r v i f o l l l Number of p lots 1 2 3 4 5 P lot number 105 107 31 32 108 Plot s i ze (meters 2) 25 25 25 25 25 Extent of type (acres) 2 1 1/20 1/20 1 Oate 5/9/ 7/9/ 5/7/ 12/7/ 7/9/ 64 64 64 64 64 Percent coverage Vegetation: layer : A3 30 40 70 65 50 Bl 90 55 70 75 35 B2 15 25 70 45 20 Total 8 95 65 85 75 50 C 25 50 30 25 30 \"h 60 65 20 35 60 \u00C2\u00B0d\u00C2\u00AB 10 10 2 3 15 Total D 70 75 22 38 70 EA 15 15 20 30 15 \u00E2\u0080\u00A2 E B 30 25 25 35 25 E C 35 35 45 40 35 Humus 75 85 95 90 75 Decaying Wood 25 15 5 10 25 Sped es Con- Cover L i f e Sub- S ign i f icance Vigor stancy value form A laver layer M R M R 1 Pm Pinus contorta 3 6.6.2 7.3.2 8.5.2 8.5.2 4.2.1 6.6 4-8 1.8 1-2 V 4860 2 Pm Thuja p l i c a t a 3 2.5.2 4.5.2 ' 2.1.2 7.5.1 3.0 0-7 1.8 1-2 IV 1240 B laver Pm Pinus contorta 1 8.6.1 7.5.2 5.4.2 3.3.2 5.4.1 5.6 3-8 1.6 1-2 V 3400 Pm Thuja p l i c a t a 1 5.5.1 6.5.2 3.+.1 4.3.2 5.3.1 4.6 3-6 1.4 1-2 V 1760 2 1.+.1 3.+.1 3.+.1 1.4 0-3 1 \u00E2\u0080\u0094 3 Pn Ledum groenlandicum 1 5.4.2 5.4.2 4.3.2 5.3.2 4.3.2 4.6 4-5 2 \u00E2\u0080\u0094 V 1600 2 4.2.2 3.2.+ 3.2.2 3.2.2 2.6 0-4 1.3 1-2 4 Pn Gaultheria shal lon 1 4.4.2 4.3.2 4.3.1 5.3.2 5.4.2 4.4 4-5 1.8 1-2 V 1400 2 3.2.+ 4.2.1 4.2.2 2.2 0-4 1.2 +-2 5 Pn Kalmla p o l i f o l l a 1 4.3.2 4.4.2 2.+.1 2.+.2 4.3.2 3.2 2-4 1.8 1-2 V 720 2 4.4.2 3.2.+ 3.+.1 1.+.1 3.2.2 2.8 1-4 1.3 +-2 6 Ch Vaccinium v i t i s - i d a e a 2 2.2.2 1.+.2 2.+.1 3.2.2 3.2.2 2.2 1-3 1.8 1-2 V 250 7 Pn Vaccinium parv i fo l ium 1 1.+.1 2.+.2 2.+.1 3.2.1 +.+.1 1.7 +-3 1.2 1-2 V 152 2 2.1.1 0.4 0-2 1 \u00E2\u0080\u0094 8 Ch Oxycoccus quadripetalus 2 1.2.2 1.+.1 1.2.1 1.2.1 2.+.2 1.2 1-2 1.4 1-2 V 60 9 Ch Empetrum nigrum 1 4.4.2 4.3.2 6.4.2 7.4.2 3.4 0-7 2 \u00E2\u0080\u0094 IV 2060 2 3.2.2 2.2.2 4.2.2 , 2.6 0-4 2 \u00E2\u0080\u0094 10 Ch Linnaea boreal i s 2 3.2.2 3.2.2 1.2.1 3.2.1 2.0 0-3 1.5 1-2 IV 310 11 Pn Vaccinium uliglnosum 2 1.2.2 3.2.2 2.2.1 1.2 0-3 1.7 1-2 III 130 12 Pn Vaccinium ovatum 1 3.+.2 0.6 0-3 2 1 100 13 Pm Tsuga heterophylla 1 2.+.1 0.4 0-2 1 \u00E2\u0080\u0094 1 20 14 Pn Vaccinium oval 1folIum 1 1.+.1 0.2 0-1 1\" \u00E2\u0080\u0094 l i 12 2 +.+.1 0.1 0-+ 1 15 Pn Menzlesla ferruglnea 1 +.+.1 0.1 0-+ 1 \u00E2\u0080\u0094 1 2 C layer 16 H Carex obnupta 4.3.2 5.4.2 1.+.1 3.3.1 4.3.2 3.4 1-5 1.6 1-2 V 910 17 H Blechnum splcant 3.3.1 4.3.2 4.3.2 2.3.1 4.+.2 3.4 2-4 1.6 1-2 V 720 18 Ch Cornus canadensis 3.+.2 3.+.2 4.2.2 4.2.2 3.+.2 3.4 3-4 2 \u00E2\u0080\u0094 V 700 19 H Agoserls glauca 3.2.2 4.2.2 2.+.1 3.2.1 3.2.2 3.0 2-4 1.6 1-2 V 520 20 G Maianthemum di latatum 2.+.1 1.+.+ 2.+.1 1.+.+ 1.+.+ 1.4 1-2 0.7 +-1 V 70 21 H Gentiana sceptrum 1.2.2 1.+.1 1.+.1 2.+.2 1.0 0-2 1.5 1-2 IV 50 22 H Drosera r o tund i f o l i a 1.+.2 1.+.1 1.+.2 1.+.1 0.8 0-1 1.5 1-2 IV 40 23 G T r i en ta i l s a r c t i c a 1.+.1 1.+.1 +.+.+ 1.+.1 0.7 0-1 0.9 +-1 IV 32 24 H Sangulsorba microcephal 3.2.3 3.2.2 3.2.2 1.8 0-3 2.3 2-3 III 300 25 H Coptls a s p l e n l f o l l a 1.+.1 1.+.2 1.+.1 0.6 0-1 1.3 1-2 III 30 26 H Calamagrostis nutkaensl 2.2.2 2.2.2 0.8 0-2 2 \u00E2\u0080\u0094 II 40 27 H Oeschampsia caespltosa 3.2.1 0.6 0-3 1 \u00E2\u0080\u0094 1 100 28 H Carex p l u r i f l o r a 1.2.1 0.2 0-1 1 \u00E2\u0080\u0094 1 20 29 G Lysichitum americanum 2.3.1 0.4 0-2 1 \u00E2\u0080\u0094 1 20 30 G Habenaria saccata 1.+.2 0.2 0-1 2 1 10 31 G Pterldlum apuillnum 1.+.2 0.2 0-1 2 1 10 32 Ch Lycopodlum clavatum 1.2.1 0.2 0-1 1 1 10 33 G Boschniakia hookerl +.+.2 0.1 0-+ 2 1 2 34 G Veratrum vi r ide +:+.2 0.1 0-+ 2 \u00E2\u0080\u0094 1 2 115 Life Sub-form D layer layer 35 B Sphagnum papillosum h 4.2.2 4.2.2 1.1.1 36' B Eurhynchium oreganum h 4.2.2 1.2.1 du 1.1.1 3.2.2 37 B Sphagnum recurvum h 5.3.2 6.3.3 2.2.2 38 B Sphagnum capillaceum h 4.2.2 4.3.2 4.3.2 39 B Pleurozium schreberi h 3.1.2 4.2.2 d\u00C2\u00AB 3.2.2 40 B Scapania bolanderi dw 2.1.1 3.2.3 41 B Dicranum bergerl h 3.2.2 dm 1.2.2 42 B Sphagnum mendocinum h 4.3.2 6.2.3 43 B Hylocomlum splendens h 3.2.1 5.3.2 dw 44 B Rhytidiadelphus loreus h 4.2.2 1.1.1 1.1.1 dw 3.2.2 2.1.1 45 B Cephalozla bicuspidata h 2.1.2 dw 2.1.1 46 B Dicranum scoparium du 3.2.2 47 B Bazzania ambigua h 2.1.2 dw 2.1.2 48 6 Sphagnum fuscum h 3.3.2 49 L Cladonla pacifica h 50 B Plaglothecium undulatum h 1.2.1 51 B Mnlum glabrescens h dw 1.1.1 52 B Sphagnum tenellum h 53 L Cladonla rangiferina h 2.2.2 54 L Cladonia crispata h 55 B Isothecium stoloniferum dw 56 B Polytrichum strictum dw 1.1.1 E layer Bryophytes S lichens 57 E Frullania nisquallensis A 3.2.2 2.1.1 3.2.2 B 3.2.2 3.1.1 4.2.2 C 4.2.2 4.1.2 4.2.2 E Dicranum scoparium B 1.1.1 +.1.1 C 3.1.2 E Scapania bolanderi B 2.1.2 C 3.2.2 3.2.2 E Pleurozl um schreberi C 3.2.2 E Isothecium stoloniferum 8 1.2.1 C 1.2.1 E Plagiothecium undulatum C 1.1.1 1.2.1 58 E Antitrichia curtipendul. C 3.1.2 E Bazzania ambigua C 1.1.2 Species Con- Cover Significance Vigor stancy value III R IH R 2.2.2 2.2.2 2.6 1-4 1.8 1-2 V 450 1.1.1 1.2 0-4 1.3 1-2 V 330 3.2.1 1.4 0-3 1.3 1-2 4.3.2 3.4 0-6 2.3 2-3 IV 1280 5.2.2 3.4 0-5 2 \u00E2\u0080\u0094 IV 1000 2.2.1 4.2.2 2.6 0-4 1.8 1-2 IV 520 2.1.1 1.0 0-3 1.5 1-2 2.2.1 2.2.2 1.8 0-3 1.8 1-2 IV 160 2.2.1 1.2.2 1.2 0-3 1.7 1-2 IV 140 0.2 0-1 2 \u00E2\u0080\u0094 5.3.2 3.0 0-6 2.3 2-3 III 1260 4.3.2 2.4 0-5 1.7 1-2 III 700 3.1.1 0.6 0-3 1 \u00E2\u0080\u0094 1.2 0-4 1.3 1-2 III 230 1.0 0-3 1.5 1r2 3.1.1 1.0 0-3 1.5 1-2 III 140 2.1.1 0.8 0-2 1 \u00E2\u0080\u0094 1.1.1 2.1.2 1.2 0-3 1.7 1-2 III 130 2.1.1 0.8 0-2 1.5 1-2 111 60 0.4 0-2 . 2 \u00E2\u0080\u0094 4.3.2 1.4 0-4 2 \u00E2\u0080\u0094 II 300 3.3.2 3.2.2 1.2 0-3 2 \u00E2\u0080\u0094 11 200 2.1.1 0.6 0-2 1 \u00E2\u0080\u0094 11 30 1.1.1 0.2 0-1 1 \u00E2\u0080\u0094 II 20 0.2 0-1 1 \u00E2\u0080\u0094 3.3.2 0.6 \u00E2\u0080\u00A20-3 2 \u00E2\u0080\u0094 1 100 0.4 0-2 2 \u00E2\u0080\u0094 1 20 1.1.1 0.2 0-1 1 \u00E2\u0080\u0094 1 10 1.1.1 0.2 0-1 1 \u00E2\u0080\u0094 1 10 0.2 0-1 1 ' 10 2.2.1 2.1.1 2.4 2-3 1.4 1-2 4.2.2 2.1.1 3.2 2-4 1.6 1-2 4.2.2 3.2.2 3.8 3-4 2 \u00E2\u0080\u0094 V 900 0.3 0-1 1 \u00E2\u0080\u0094 2.2.2 1.0 0-3 2 \u00E2\u0080\u0094 IV 132 0.4 0-2 2 \u00E2\u0080\u0094 2.1.2 1.6 0-3 2 \u00E2\u0080\u0094 111 220 2.1.2 1.0 0-3 2 \u00E2\u0080\u0094 II 120 0.2 0-1 1 \u00E2\u0080\u0094 1.1.1 0.4 0-1 1 \u00E2\u0080\u0094 II 20 0.4 0-1 1 \u00E2\u0080\u0094 \u00E2\u0080\u00A2 II 20 0.6 0-3 2 \u00E2\u0080\u0094 1 100 0.2 0-1 2 \u00E2\u0080\u0094 1 10 Total species 43 36 35 Life form Pn Ch Pm Proportion by Species: Number 19 10 8 8 7 6 3 3 t 29.7 15.6 12.5 12.5 10.9 9.4 4.7 4.7 Total cover: cover 6870 2730 4116 1522 146 3390 6640 230 % 26.8 10.7 16.1 5.9 0.6 13.2 25.9 0.9 SYNTHESIS TABLE VI. PINETO-CHAMAECYPARETO-SPHAGKETUM RECUP.VII Nunber of plots 1 2 3 4 5 Plot number 72 71 23 27 28 Plot size (neters) 100 100 100 100 100 Date 5/8/ 5/8/ 10/6/ 15/6/ 15/6/ 64 64 64 64 64 Percent coveraga Vegetation: layer: \u00C2\u00BBi 0 15 70 15 30 \u00C2\u00BBz 50 40 30 70 40 A 3 40 35 50 30 25 Total A 75 60 80 80 70 Bl 65 60 40 60 65 B 2 15 25 10 5 8 Total B 65 60 45 60 65 C 15 20 8 20 15 Dh 30 35 25 40 30 \u00C2\u00B0ds 35 15 50 35 40 Total 0 65 50 75 75 70 E\u00E2\u0080\u009E 10 15 25 20 10 EB 45 55 60 60 40 EC 80 85 90 90 60 Kuaus 60 75 35 60 50 Decaying lood 40 25 65 40 50 Species Con- Cover Life Sub- Significance Vigor stancy value fora A laver layer II 8 H R Pa Pinus contorta 1 2.1.2 8.7.2 5.5.2 5 . 0 4.0 0-8 2 V 25ZO 2 4 . 0 2 . 0 2 . 0 3 . 0 2.2 0-4 2 \u00E2\u0080\u0094 2 Pa Chaaaecyparls nootkatensis 1 2.4.1 4 . 0 1.2 0-4 1 \u00E2\u0080\u0094 2 6.6.1 5.6.1 6.5.1 6.6.1 4.6 0-6 1 \u00E2\u0080\u0094 V 2400 3 Pa Thuja plicata 1 5.5.2 3 . 0 1.6 0-5 2 \u00E2\u0080\u0094 2 3 . 0 5.6.2 5.5.2 3.4.2 4.4.2 4.0 3-5 2 \u00E2\u0080\u0094 3 3.5.2 3.4.2 7.6.2 5.4.2 5.4.2 4.6 3-7 2 \u00E2\u0080\u0094 V 2300 4 Pa Tsuga heterophylla 2 3 . 0 2.3.1 6.5.1 7.6.2 3.6 0-7 1.3 1-2 3 3 . 0 2 . 0 1.2.1 6.5.2 3 . 0 3.0 1-6 1.2 1-2 V 1890 5 Pn Vacclnlua parvlfollua 3 3 . 0 0.6 0-3 2 \u00E2\u0080\u0094 1 100 Pa Malus dlvers l fo l la 2 3.5.1 0.6 0-3 1 \u00E2\u0080\u0094 1 100 3 3.5.1 0.6 0-3 1 \u00E2\u0080\u0094 7 Pn Benzlesla ferruglnea 3 3 . 0 0.6 0-3 2 \u00E2\u0080\u0094 1 100 9 Pa Taxus brevlfDlla 2 1.4.2 0.2 0-1 2 \u00E2\u0080\u0094 II 20 3 1.+.2 0.2 0-1 2 \u00E2\u0080\u0094 B layer 9 Pn Gaultheria shallon 1 7.5.2 8.5.2 5.3.2 7.5.2 8.5.2 7.0 5-8 2 \u00E2\u0080\u0094 V 5400 2 5.2.2 5.3.2 3 . 0 3.+.2 3 . 0 3.8 3-5 1.8 1-2 Pn Vacclnlua parvlfollua 1 3 . 0 3.3.1 4.3.2 3.4.2 4.4.2 3.4 3-4 1.8 1-2 V 700 2 3.2.2 2 . 0 Z . O 2 . 0 3 . 0 2.4 2-3 1.6 1-2 10 Pn Vacclnlua ovatua 1 3 . 0 \u00E2\u0080\u00A24.3.2 3.4.2 1.+.2 2.4 1-4 1.8 1-2 V 420 2 3 . 0 0.6 0-3 2 \u00E2\u0080\u0094 Pn Venzlesla ferruglnea 1 3 . 0 I . O 2.4.2 2.3.1 I . O 1.8 1-3 1.4 \u00E2\u0080\u0094 V 160 2 1.+.1 0.2 0-1 1 \u00E2\u0080\u0094 Pa Tsuga heterophylla 1 4.3.2 3 . 0 3 . 0 4.3.2 2.8 0-4 1.4 1-2 IV 600 2 Z.+.1 3 . 0 1.0 0-3 1 \u00E2\u0080\u0094 11 Pn Vacclnlua ovaUfollua \ I . O 3.2.2 2.4.2 2.3.2 1.6 0-3 1.4 1-2 IV 150 Pa Thuja pl icata 1 5.3.2 3.+.Z 2.4.2 2.0 0-5 2 \u00E2\u0080\u0094 III 5Z0 12 Ch Llnnaea boreal Is 2 2.1.2 3 . 0 3.Z.2 1.6 0-3 2 \u00E2\u0080\u0094 III 220 13 Pn Ledua groenlandlcua 1 2 . 0 2 . 0 0.8 0-2 0.6 1-2 II 40 Pa Chamaecyparis nootkatensis 1 3.3.1 0.6 0-3 1 \u00E2\u0080\u0094 1 100 Pa Hal us d lvers l fo l la 1 3.4.1 0.6 0-3 1 \u00E2\u0080\u0094 I 100 2 2 . 0 0.4 0-2 1 \u00E2\u0080\u0094 14 Ch Vacclnlua vltls-ldaea 2 2.1.2 0.4 0-Z 2 \u00E2\u0080\u0094 1 20 C laver 15 6 Halantheaua dlletatua 1 . 0 1.+.+ Z.+.+ 2 . 0 1.4 1-2 0.7 \u00C2\u00AB-1 V 70 16 H Blechnua splcant 5.4.2 4.3.2 5.5.2 5.5.2 3.8 0-5 2 \u00E2\u0080\u0094 IV 1400 17 Ch Cornus canadensis 3.2.2 3.2.1 2 . 0 3.2.1 2.2 0-3 1.3 1-2 IV 320 18 H Care* obnupta 3.3.1 \u00E2\u0080\u00A2 3.3.1 4.5.2 2.0 0-4 0.9 1-2 III 400 19 H Deschaepsla caespltoss 3.3.2 4.3.2 1.2.2 1.6 0-4 2 \u00E2\u0080\u0094 III 310 20 G Veratrua vtrlde 2 . 0 3.3.1 I . O 1.2 0-3 1.3 1-2 III 130 21 H Calaaagrcstis nutkaensls \u00E2\u0080\u00A2 i . o 2.1.3 1 . 0 0.8 0-2 3 \u00E2\u0080\u0094 III 40 22 Ch Lycopodlua clavatua 1.1.1 2.2.1 0.6 0-2 1 \u00E2\u0080\u0094 II 30 23 G Boschnlekla hookerl 1.1.2 1.2.2 0.4 0-1 2 \u00E2\u0080\u0094 II 20 24 G Llstera cordata 1 . 0 0.4 0-1 0.6 1-2 II 20 25 H Gentiana sceptrua 2.1.1 0.4 0-2 1 \u00E2\u0080\u0094 1 20 26 G Lyslchltus aaerlcanua 2.4.2 0.4 0-2 2 \u00E2\u0080\u0094 1 20 27 H Agoserls glauca 1.2.2 0.2 0-1 2 \u00E2\u0080\u0094 1 10 28 G Trlentalls arctlca 1 . 0 0.2 0-1 2 \u00E2\u0080\u0094 1 io 29 H Goodyera oblonglfolle \u00E2\u0080\u00A2.\u00E2\u0080\u00A2.1 0.1 0-.5 1 \u00E2\u0080\u0094 1 2 Life fora E 8 H Pa Pn 6 Ch Proportion by Species: (iuiber 33 24 7 6 6 6 4 1 39.4 28.0 ,8.1 7.0 7.0 7.0 4.7 Total cover: cover 8752 7350 2182 9220 6870 270 590 I 24.9 20.9 6.2 26.2 19.5 0.8 1.7 Life Sub-forn 0 later layer 30 G HylocoBfue tplendens h 4.3.2 5.3.2 de 3.2.2 1.2.1 31 B fihytldladelphus loreus h 4.3.2 4.2.2 de 4.2.2 3.2.2 2.2.2 32 e Sphagnui recurvui h 5.3.2 5.3.2 1.3.1 33 B Scapania bolenderl de 4.2.2 3.2.2 4.2.2 34 B Eurhynchlua oreganua h 1.1.1 de 3.2.2 2.2.1 35 G Lepldozla reptans de 3.2.2 2.1.2 2.1.2 30 B Inlua glebrescens h 1.2.2 3.3.2 6.5.2 de 4.3.2 37 B . Plaglotheclue unduletue h 3.2.1 de 2.1.1 38 B Bazzania anblgua h 2.1.2 de 3.1.2 2.1.2 39 B Cephalozia blcuspldata h 1.1.2 de 2.1.2 2.1.2 40 G Hookerla lucens h 1.1.2 di 1.1.2 41 B Dlplophylluo albicans h 2.2.2 6.4.3 de 3.2.2 4.3.3 42 G Glepharetona trlchophyllun h 1.2.2 di 2.1.2 43 B Olcranua ecoparlui da 44 B Olcranua fuscescens de 1.1.2 45 G Herberts adunca de 3.3.3 45 B Hypnue clrclnale de 47 B Isotheclui stolonlferui de 3.3.3 48 G Pellle eplphylle h 2.4.3 49 G Eurhynchlua stokesll de 50 e Pleglochlla aaplenoldes dl 3.1.1 1.1.2 51 6 Riccardia sp. de 1.1.2 52 6 Cslypogela trlchonanes de 1.1.1 53 B Bartraala poatfornle dl 1.1.1 E liter Bryophytes I lichens E IsothecluB stolonlferui A 7.3.2 B 4.3.2 3.3.3 8.5.3 C 4.3.3 3.2.2 5.3.2 E Herberts adunce A 2.2.2 1.2.1 e 5.2.3 4.3.3 C 6.3.3 5.4.3 54 E Frullenla nlequallensls A 3.2.2 2.2.2 5.2.2 B 4.2.2 4.2.2 5.2.2 C 4.2.3 4.2.3 E Scepanla bolanderi A 2.1.2 3.2.2 B 3.2.2 3.2.3 4.3.2 C 4.2.2 3.2.2 4.3.2 55 E Antltrfchle curtlpendula A 2.3.2 G 2.2.2 C 4.3.2 3.2.2 E Hypnue clrcinele A 4.2.1 e 1.1.1 1.2.1 4.2.2 C 3.2.1 E Olcrenua scoparlun A 2.2.2 2.2.2 . 2.2.1 B 3.2.2 2.1.2 C 2.1.2 2.1.2 E Bazzenla anblgua A 2.1.2 B 3.1.2 2.1.2 C 1.1.2 E Olplophyllun albicans A 2.2.2 C 1.1.1 E Lepldozla reptans B 1.1.2 E Plagfochla aeplenoldee B 2.1.2 C 1.1.1 E Cephalozia blcuspldata B 2.1.2 E Hookerla lucens C 2.2.2 E Inlua glabrescens C 56 E Neckera dougl all 1 G 2.3.3 E Plaglotheelul undulatue B 2.2.1 57 E Cephalozia leaaerilene C 1.1.2 58 E Cladonia bellldlflore C E Olcranua fuscescens C 1.1.2 E Eurhynchlm etokesll c 1.1.2 59 E Netzgerla conjugata c 1.2.2 Vascular el ants E Vacclnlui pervlfollua A 2.1.1 1.\u00C2\u00BB.1 B 2...1 3.4.2 C 3.1.1 2...1 4.4.2 E Cornus canadensle B 2.1.2 l . t . t C 3.+.1 2.+.1 60 E Polypodlua olycyrrhlzi A. 2.Z.2 1.2.1 G 1.2.2 1.\u00C2\u00BB.1 3.2.1 C 1.*.1 E Gaultherla shallon A 3.2.1 2.3.2 2.*.1 G 2.-f.2 3.4.2 C 3.3.1 4.4.2 E Hanzlesla ferruginea A G 2.+.1 C 2.*.1 E Halinthoeus dllatatua B +.+.+ E Vacclnlua ovatua c . 3.3.1 \u00C2\u00A3. Vacclnlua vitis-idaea c 3.+.1 * E EnpetruB nlgrui B 2.+.1 E Llnnsea boreal la S 1.2.2 E thuja plicata A 1...1 E Tauge heterophylle 6 1. Total speclee 46 41 41 Spades Con- Caver Significance Vigor stancy value H R H R 5.4.2 3.2.2 3.8 0-5 2 \u00E2\u0080\u0094 V 1410 4.3.2 3.3.2 2.2 0-4 1.6 1-2 3.2.2 3.2.2 2.6 0-4 2 \u00E2\u0080\u0094 5.3.3 5.4.3 3.8 2-5 2.4 2-3 V 1220 2.4.3 3.4.2 3.2 1-5 2 1-3 V 930 3.2.2 3.2.2 3.4 3-4 2 \u00E2\u0080\u0094 y 700 3.2.1 3.2.1 1.4 0-3 1 \u00E2\u0080\u0094 V 330 2.2.1 1.4 0-3 1.7 1-2 1.2.2 2.1.2 2.0 1-3 2 \u00E2\u0080\u0094 V 170 2.0 0-6 2 \u00E2\u0080\u0094 IV 780 1.1.1 1.0 0-4 1.5 1-2 3.2.1 3.2.1 1.1 0-3 1 \u00E2\u0080\u0094 IV 320 2.2.1 3.1.1 2.0 0-3 1 \u00E2\u0080\u0094 0.4 0-2 2 \u00E2\u0080\u0094 2.1.2 2.1.2 0.8 0-3 2 \u00E2\u0080\u0094 IV 120 0.2 0-1 2 \u00E2\u0080\u0094 M.2 2.1.2 1.6 0-2 2 \u00E2\u0080\u0094 IV 80 1.1.2' 0.4 0-1 2 \u00E2\u0080\u0094 IV 40 1.1.2 1.1.2 0.6 0-1 2 \u00E2\u0080\u0094 1.6 0-6 2.5 2-3 2.1.2 1.8 0-4 2.3 2-3 in 780 0.2 0-1 2 \u00E2\u0080\u0094 in 50 2.1.2 0.8 0-2 2 \u00E2\u0080\u0094 1.1.1 2.1.2 0.6 0-2 1.5 1-2 II 30 1.1.2 0.4 0-1 2 \u00E2\u0080\u0094 II 20 0.6 0-3 3 \u00E2\u0080\u0094 i 100 3.2.2 0.6 0-3 2 l 100 0.6 0-3 3 \u00E2\u0080\u0094 f 100 0.4 0-2 3 I 20 1.1.2 0.2 0-1 2 \u00E2\u0080\u00A2 i 10 0.2 0-1 2 \u00E2\u0080\u0094 \u00E2\u0080\u00A2 t 10 0.2 0-1 2 \u00E2\u0080\u0094 i 10 0.2 0.1 1 \u00E2\u0080\u0094 i 10 0.2 0-1 1 1 10 2.2.2 3.2.1 2.4 0-1 1.7 1-2 4.3.2 4.3.2 4.6 3-e 2.4 2-3 V 2200 4.3.2 3.2 0-5 1.8 2-3 4.3.2 3.3.2 2.0 0-4 1.4 1-2 4.3.3 4.3.2 3.4 0-5 2.2 2-3 4.3.3 3.0 0-5 3 \u00E2\u0080\u0094 y 1470 4.2.2 4.2.2 3.6 2-5 2 \u00E2\u0080\u0094 4.3.3 5.2.2 4.4 4-5 2.2 2-3 V 1400 4.2.3 3.2.2 3.0 0-4 2.8 2-3 2.2.1 1.4 0-3 1.7 1-2 V 700 3.2.2 3.2.2 3.2 3-4 2 \u00E2\u0080\u0094 3.2.2 2.8 0-4 2 \u00E2\u0080\u0094 0.4 0-2 2 \u00E2\u0080\u0094 V 620 2.+.2 3.1.1 1.4 0-3 1.7 1-2 4.3.3 2.2 0-3 1.7 1-2 4.2.2 1.6 0-4 1.5 1-2 2.1.1 3.2.2 2.2 1-4 1.4 1-2 V 440 2.1.2 1.0 0-3 1.5 1-2 2.1.1 1.6 0-2 1.5 1-2 y 200 2.1.1 1.4 0-3 1.7 1-2 0.8 0-2 2 \u00E2\u0080\u0094 0.4 OJ 3 \u00E2\u0080\u0094 IV 150 2.1.1 1.4 0-3 1.7 1-2 0.2 0-1 2 \u00E2\u0080\u0094 0.4 0-2 2 \u00E2\u0080\u0094 in 40 1.1.2 0.2 0-1 1.5 1-2 2.1.2 0.6 0-2 2 n 30 0.4 0-2 2 \u00E2\u0080\u0094 II 30 0.2 0-1 1 \u00E2\u0080\u0094 0.4 0-2 2 \u00E2\u0080\u0094 I 20 0.4 0-2 2 \u00E2\u0080\u0094 i 20 2.2.2 0.4 0-2 2 \u00E2\u0080\u0094 i 20 0.4 0-2 3 \u00E2\u0080\u0094 l 20 0.4 0-2 1 \u00E2\u0080\u0094 i 20 0.2 0-2 2 \u00E2\u0080\u0094 I 10 l.t.1 0.2 0-1 1 \u00E2\u0080\u0094 i 10 0.2 0-1 2 \u00E2\u0080\u0094 i 10 0.2 0-1 2 \u00E2\u0080\u0094 i 10 0.2 0-1 2 \u00E2\u0080\u0094 1 10 0.6 0-2 1 2.*.2 1.4 0-3 1.7 1-2 2.\u00C2\u00BB.1 2.*.2 2.6 2-4 1.4 1-2 V 360 1.2.1 0.8 0-2 1.2 *-2 1.+.1 2..-.1 1.6 0-3 1 \u00E2\u0080\u0094 V 160 1.2.1 1.2.2 1.0 0-2 1.5 1-2 1.2.1 1.2 0-3 1.7 1-2 V 150 0.2 0-1 1 \u00E2\u0080\u0094 1.4 1.3 1-2 IV 330 1.*.2 1.2 0-3 2 \u00E2\u0080\u0094 1.4 0-4 1.5 1-2 1.+.2 0.2 0-1 2 \u00E2\u0080\u0094 2.*.2 0.8 0-2 1.5 1-2 in 50 1.2.1 0.6 0-2 1 \u00E2\u0080\u0094 0.1 0-* + \u00E2\u0080\u0094 2.2.1 0.4 0-2 1 \u00E2\u0080\u0094 II 22 0.6 0-3 1 \u00E2\u0080\u0094 I 100 0.6 0-3 1 l 100 0.4 0-2 1 \u00E2\u0080\u0094 I 20 0.2 0-1 2 \u00E2\u0080\u0094 1 10 0.2 0-1 1 \u00E2\u0080\u0094 I 10 0.2 0-1 + \u00E2\u0080\u0094 i 10 40 36 118 APPENDIX III Soils data i A s s o c i a t i o n : C a r i c e t u m p l u r i f l o r a e P l o t n o . 40 44 50 Date c o l l e c t e d 19/8/64 19/8/64 20/8/64 Root d i s t r i b u t i o n max. d e p t h (cm) 40 30 45 main c o n e , t o (cm) 25 15 10 Chemical A n a l y s i s S o i l T o t a l E x c h a n g e a b l e c a t i o n s Adsorbed Base P l o t Depth M o i s t u r e pH N i t r o g e n C . E . C . Na K Ca Mg Phosphate S a t u r a t i o n n o . (cm) ( ? ) ( ? ) (me?) (me?) (me?) (me?) (me?) (ppm) ( ? ) 40 5-10 1247.5 3 . 4 1.61 2 . 6 5 . 7 0 1 .04 8 . 6 0 4 . 6 5 5 . 3 9 . 3 20-25 7 6 0 . 8 3 . 9 2 . 8 0 85 1 . 4 5 0 . 2 8 1 . 5 0 1 . 2 5 0 . 6 5 . 3 3 0 - 3 5 85.1 4 . 4 0.001 42 0 . 7 0 0 . 0 5 0 . 8 0 0 . 3 7 0 . 9 4 . 6 50 5-10 2 2 3 . 3 4.1 66 0 . 9 0 0 . 0 9 0 . 5 6 0 . 2 0 2 . 7 15-20 1019.4 3 . 9 81 1 . 0 5 0 . 2 7 2 . 2 0 1 . 7 3 1 7 . 6 3 0 - 3 5 9 6 . 2 4 . 4 45 0 . 7 4 0 . 0 5 0 . 4 0 0 . 0 8 2 . 8 40-45 3 5 . 1 4 . 0 44 10-15 7 3 2 . 5 3 . 5 15-20 9 4 . 7 4 . 3 2 5 - 3 0 5 2 . 2 4 . 7 A s s o c i a t i o n : Oxycocceto-Sphagnetum p a p i l l o s a e P l o t n o . 18 Date c o l l e c t e d 15/8/64 Root d i s t r i b u t i o n max. d e p t h (cm) 57 main c o n e , t o (cm) 25 19 16/8/64 50 20 Chemical A n a l y s i s S o i l T o t a l P l o t Depth M o i s t u r e pH N i t r o g e n C . E . C . n o . (cm) ( ? ) ( ? ) (me?) E x c h a n g e a b l e c a t i o n s Na (me?) K (me?) Ca (me?)' Mg (me?) Adsorbed Base Phosphate S a t u r a t i o n (ppm) ( ? ) 10-1.5 1 0 7 3 . 3 3 . 7 247 4 . 5 5 1 . 2 6 4 . 5 8 4 . 8 0 3 . 0 6.1 22-25 7 9 6 . 7 4.1 71 2 . 0 5 0 . 7 6 0 . 9 5 1 . 3 0 1 . 4 7.1 2 8 - 3 2 2 8 1 . 8 3.'9 60 0 . 8 5 0 . 1 6 0.51 0 . 4 9 1 . 2 3 . 4 3 5 - 4 0 1 4 4 . 8 4 . 0 63 0 . 8 2 0 . 1 2 0 . 2 7 0 . 1 9 0 . 9 2 . 2 5 2 - 5 7 5 5 . 5 4 . 8 42 0 . 7 0 0 . 0 4 0 . 2 6 0 . 0 5 1 . 2 2 . 5 19 6-11 1 4 0 8 . 6 3 . 4 1 . 5 5 276 6 . 2 8 1 . 4 9 6 . 4 0 6 . 2 5 7 .4 15-20 1 0 6 5 . 0 3 . 5 2 . 2 4 73 1 .75 0 . 5 7 1.17 1 . 5 5 6 . 9 30-35 8 2 . 7 3 . 8 0 . 1 6 49 0 . 6 9 0 . 0 7 0 . 3 6 0 . 1 4 2 . 6 4 5 - 5 0 1 3 4 . 6 4 . 6 0 . 3 6 61 0 . 8 0 0 . 0 8 0 . 4 4 0 . 2 2 2 . 5 A s s o c i a t i o n : S c i r p e t o - S p h a g n e t u m mendocini P l o t no. 93 Date c o l l e c t e d 6/9/64 Root d i s t r i b u t i o n max. d e p t h . ( c m ) 55 main c o n e . t o (cm) 15 Chemical A n a l y s i s S o i l T o t a l P l o t Depth 1 M o i s t u r e pH N i t r o g e n C . E . C , no. (cm) ( ? ) ( ? ) (me?) 93 5-10 8 2 0 . 6 4 , 1 1 , 7 9 61 20-25 5 5 7 . 0 4 . 2 1 . 8 8 65 2 5 - 3 0 ' 5 P . 7 4 . 7 0 . 1 6 49 5 0 - 5 5 3 6 . 6 5 . 4 34 101 5-10 9 8 5 , 5 3 . 4 244 15-20 2 8 0 . 8 4 . 4 65 2 5 - 3 0 3 . 9 63 60-65 4 . 2 44 101 6/9/64 65 20 E x c h a n g e a b l e c a t i o n s Adsorbed Base \u00E2\u0080\u00A2Na K Ca Mg Phosphate S a t u r a t i (me?) (me?) (me?) (me?) (ppm) ( ? ) 1 . 5 5 0 . 4 7 2 . 2 6 2 . 1 0 0 . 4 11.1 1 . 4 5 0.41 1 . 5 5 1 .45 0 . 5 7 . 5 0 . 6 3 0 . 0 8 0 . 5 0 0.14 0 . 9 2 . 8 0 . 6 8 0 . 0 4 0 . 5 0 0 . 0 6 0 . 9 3 . 8 4 . 2 0 1 . 2 2 5 . 8 0 5 . 2 0 6 . 7 2 . 1 8 0 , 3 6 5 . 4 0 1 . 9 5 1 5 . 2 1 .75 0 . 1 2 5 . 3 5 1 . 6 8 14.1 0 . 9 2 0 . 0 9 1 .75 0.41 7 . 2 A s s o c i a t i o n : Ledeto-Sphagnetum c a p i l l a c e i P l o t n o . 26 68 Date c o l l e c t e d 15/8/64 19/8/64 Root d i s t r i b u t i o n max. d e p t h (cm) 65 95 main c o n e , t o (cm) 40 45 Chemical A n a l y s i s S o i l T o t a l Exchanqeabl e c a t i o n s Adsorbed Base P l o t Depth M o i s t u r e pH N i t r o g e n C . E . C . Na K Ca Mg Phosphate S a t u r a t i n o . (cm) ( ? ) ( ? ) (me?) (me?) (me?) (me?) (me?) (ppm) ( ? ) 26 6-11 1 3 7 6 . 3 3 . 7 3 . 9 3 276 6 . 2 4 1 , 2 7 6.41 6 . 2 0 0 . 6 7 . 3 2 2 - 2 7 1441.2 3 . 4 1 . 5 9 279 6 , 4 0 1 . 2 9 6 . 6 0 5 . 6 5 2 . 5 7.1 30-35 ' 3 4 8 . 2 4 . 1 0 . 0 8 60 1 .15 0 . 1 2 1 . 0 8 0 . 9 4 0 . 9 5 . 5 4 5 - 5 0 62.1 5.1 0 . 0 2 42 0 . 6 0 0 . 0 4 0 . 4 7 0 . 1 5 0 . 8 3 . 0 60-65 4 4 . 9 4 . 5 0 . 0 7 32 0 . 5 9 0 . 0 3 0 . 7 5 0 . 1 0 2 . 2 4 . 6 68 5-10 1 3 9 0 . 9 3 . 3 253 4 . 4 0 1 . 7 6 9 . 4 0 8 . 5 0 9 . 5 30-35 1131.0 3 . 8 82 1 , 6 5 \u00E2\u0080\u00A2 0 . 4 2 3 , 9 5 2 . 0 5 9 . 8 40-45 1 2 4 3 . 0 4 . 4 70 2 , 0 8 0.21 2 . 1 0 1.41 8 , 3 6 0 - 6 5 1 2 3 0 . 0 3 . 7 157 4 , 0 5 0 . 2 8 6 . 4 0 1 .75 7 . 9 95-100 6 5 . 8 4 . 6 . 42 0 . 7 8 0 . 0 5 1 . 3 6 0 . 2 0 5 , 7 A s s o c i a t i o n : P i n e t o - S p h a g n e t u m c a p i l l a c e i sphagnoosum p a p i l l o s i P l o t n o . 75 78 Date c o l l e c t e d 24/8/64 24/8/64 T h i c k n e s s of humus (cm) 20 25 Root d i s t r i b u t i o n max. d e p t h (cm) 105 8 0 - 9 0 main c o n e , t o (cm) 25 25 C h e m i c a l ' . ' A n a l y s i s S o i l T o t a l E x c h a n g e a b l e c a t i o n s Adsorbed Base P l o t Depth M o i s t u r e pH N i t r o g e n C . E . C . Na K Ca Mg Phosphate S a t u r a t i No. (cm) ( ? ) ( ? ) (me?) (me?) (me?) (me?) (me?) (ppm) ( ? ) 75 15-20 1061.5 3 . 9 1.61 300 4 . 0 5 1 .24 6 . 8 0 4 . 4 0 0 . 5 5 . 5 32-37 1 8 2 4 . 8 4 . 2 1 . 2 3 68 1 .75 0 . 2 7 2 . 5 4 1 . 4 3 0 . 7 8 . 8 5 5 - 6 0 6 2 8 . 5 4 . 6 0 . 2 2 66 1 .15 0 . 0 7 2 . 9 5 0 . 8 8 0 . 2 7 . 7 60-75 3 1 8 . 4 4 . 6 0 . 1 6 70 0 . 9 0 0 . 0 8 5 . 0 5 0 . 8 2 0 . 4 9 . 8 105-110 5 1 . 2 5 . 4 0 . 0 4 34 0 . 6 4 0 . 0 4 0 . 7 6 0 . 0 8 3 . 0 4 . 5 78 10-15 1 4 0 0 . 0 3 . 4 256 4 . 9 0 0.91 7 . 8 0 5 . 9 5 7 . 6 20-25 9 7 9 . 3 3 . 9 70 1 . 4 5 0 . 3 8 1.15 1 . 4 3 6 . 3 40-45 7 8 1 . 8 4 . 2 58 2 . 1 8 0.21 0 . 8 5 1 . 0 3 7 . 4 \u00E2\u0080\u00A2 60-65 5 5 7 . 3 3 . 9 61 1 .15 0 . 0 8 0 . 6 2 0 . 4 6 3 . 8 75-80 7 0 . 8 4 . 5 56 0 . 6 3 0 . 0 4 0 . 4 0 0 . 0 7 2 . 0 A s s o c i a t i o n : P i n e t o - S p h a g n e t u m c a p i l l a c e i chamaecyparosum n o o t k a t e n s i s P l o t no. 11 Date c o l l e c t e d 20/8/64 Root d i s t r i b u t i o n max. d e p t h (cm) 95 main c o n e , t o (cm) 30 13 13/8/64 70 23 Chemical A n a l y s i s S o i l T o t a l E x c h a n g e a b l e c a t i ons Adsorbed Base P l o t Depth M o i s t u r e pH N i t r o g e n C . E . C . Na K Ca Mg Phosphate S a t u r a t i n o . (cm) ( ? ) ( ? ) (me?) (me?) (me?) (me?) (me?) (ppm) ( ? ) 13 4-12 4 3 4 . 2 3 . 7 285 4 . 2 4 1 . 8 7 9 . 4 0 8 . 3 0 1.1 8 . 4 19-23 6 1 . 6 4 . 5 43 0 . 5 5 0 . 0 6 0 . 2 2 0 . 0 4 1 . 5 2 . 0 2 5 - 2 9 6 9 . 2 4 . 5 51 0 . 7 5 0 . 0 6 0.15 0 . 0 6 1.1 2 . 0 3 0 - 3 5 7 1 . 0 4 . 5 51 0 . 5 5 0 . 0 4 0 . 1 8 0.11 2 . 4 1 . 7 40-45 4 6 . 0 4 . 5 36 0.51 0 . 0 4 0.11 0 . 0 4 1 . 7 1 . 9 6 5 - 7 0 4 0 . 4 5 . 2 26 0 . 4 9 0 . 0 4 0 . 2 6 0 . 0 3 2 . 4 3 . 2 11 3 - 8 1 0 9 7 . 0 3 . 7 275 5 . 5 0 2.14 4 . 4 0 4 . 3 0 5 . 9 20-25 6 8 7 . 5 3 . 6 219 3 . 4 8 0 . 8 7 3 . 6 0 1 . 9 5 7 . 5 3 5 - 4 0 4 4 2 . 8 3 . 9 216 4 . 0 5 0 . 1 8 2 . 8 2 0 . 2 5 3 . 4 6 5 - 7 0 2 0 8 . 7 4 . 4 36 0 . 7 8 0 . 0 8 2 . 0 5 0.21 8 . 7 90-95 2 7 . 2 4 . 5 36 0 . 7 5 0 . 0 8 1 . 8 2 0 . 2 5 8.1 123 A s s o c i a t i o n : P i n e t o - S p h a g n e t u m c a p i l l a c e i v a c c i n i o s u m v i t i s - i d a e a e P l o t n o . 1 24 30 Date c o l l e c t e d 12/8/64 12/8/64 15/8/64 T h i c k n e s s of humus (cm) 5 9 10 Root d i s t r i b u t i o n max. d e p t h (cm) 50 75 75 main c o n e , t o (cm) 10 10 15 Chemical A n a l y s i s S o i l T o t a l E x c h a n g e a b l e c a t i o n s Adsorbed Base P l o t Depth M o i s t u r e pH N i t r o g e n C . E . C . Na K Ca Mg Phosphate S a t u r a t i o n no. (cm) ( ? ) ( ? ) (me?) (me?) (me?) (me?) (me?) (ppm) ( ? ) 1- 0-5 6 5 3 . 3 3 . 5 1 . 4 0 263 6 . 0 3 2 . 0 2 8 . 0 3 9 . 0 0 5 . 4 9 . 5 15-20 9 4 . 3 3 . 7 0.21 53 0 . 7 5 0 . 0 7 0 . 2 8 0 . 1 0 0 . 9 2 . 3 2 5 - 3 0 6 5 . 3 4 . 9 0 . 0 0 2 43 0 . 5 9 0 . 0 5 0 . 3 3 0.15 0 . 3 2 . 6 4 5 - 5 0 5 0 . 7 4 . 7 0 . 0 0 2 42 0 . 5 9 0 . 0 5 0 . 3 2 0 . 0 4 1 .7 2 . 4 0 - 8 5 5 4 . 9 3 . 4 295 4 . 4 5 1 .77 5 . 6 5 6 . 5 0 6 . 2 30-35 3 7 . 3 4 . 6 49 0 . 6 0 0 . 0 4 0 . 2 7 0 . 0 3 1 . 9 5 5 - 6 0 3 9 . 4 4 . 8 34 0 . 6 0 0 . 0 3 0 . 5 9 0 . 0 5 3 . 7 70-75 2 9 . 2 4 . 8 36 0 . 6 3 0 . 0 4 0 . 7 0 0 . 0 7 4 . 0 3 - 9 4 5 5 . 4 3 . 7 8-12 7 7 . 8 4 . 2 22-27 4 4 . 5 4 . 7 A s s o c i a t i o n : P i n e t o - S p h a g n e t u m c a p i l l a c e i v a c c i n i o s u m p a r v i f o l i i P l o t n o . 31 32 Date c o l l e c t e d 13/8/64 12/8/64 T h i c k n e s s of humus (cm) 15 18 Root d i s t r i b u t i o n max. d e p t h (cm) 75 70 main c o n e , t o (cm) 15 20 Chemical A n a l y s i s S o i l T o t a l E x c h a n g e a b l e c a t i o n s Adsorbed Base P l ot Depth M o i s t u r e pH N i t r o g e n C . E . C . Na K Ca Mg Phosphate S a t u r a t i n o . (cm) ( ? ) ( ? ) (me?) (me?) (me?) (me?) (me?) (ppm) ( ? ) 31 0 - 8 3 . 5 10-15 4 4 6 . 5 4 . 2 0 . 3 3 200 3 . 9 5 0 . 9 8 2 . 8 0 2 . 4 5 2.1 5.1 20-25 5 2 . 2 4.1 0 . 0 7 52 0 . 5 4 0 . 0 5 0 . 0 4 0 . 0 5 1 . 7 1 . 3 70-75 3 0 . 5 5 . 4 0.01 31 0 . 7 4 0 . 0 5 0 . 4 6 0 . 0 3 2 . 6 4.1 32 3 - 8 6 0 3 . 2 3 . 4 222 4 . 0 5 2 . 1 5 3 . 6 4 6 . 4 5 7 . 3 12-18 4 5 1 . 9 4 . 0 70 1 . 2 5 0 . 4 3 0 . 5 0 0 . 9 7 4 . 5 20-25 82.1 4 . 4 38 0 . 4 5 0 . 0 6 0 . 1 8 0 . 0 8 2 . 0 34-37 3 1 . 4 4 . 6 37 0 . 5 3 0 . 0 5 0 . 2 2 0 . 0 3 2 . 2 6 5 - 7 0 2 7 . 3 4 . 4 24 0 . 6 4 0 . 0 5 0 . 6 4 0 . 0 4 5 . 7 A s s o c i a t i o n : P i n e t o - C h a m a e c y p a r e t o - S p h a g n e t u m r e c u r v i P l o t n o . 27 72 Date c o l l e c t e d 13/8/64 19/8/64 T h i c k n e s s of humus (cm) 10 15 Root d i s t r i b u t i o n max. d e p t h (cm) 65 85 main c o n e , t o (cm) 15 30 Chemical A n a l y s i s S o i l T o t a l E x c h a n g e a b l e c a t i o n s Adsorbed Base P l o t Depth M o i s t u r e pH N i t r o g e n C . E . C . Na K Ca Mg Phosphate S a t u r a t i n o . (cm) ( ? ) ( ? ) (me?) (me?) (me?) (me?) (me?) (ppm) ( ? ) 72 5-10 2 7 0 . 5 4 . 5 0 . 8 8 288 4 . 8 0 2 . 1 7 3 7 . 8 0 7 . 2 5 1 . 3 18.1 30-35 4 1 7 . 7 3 . 3 0 . 8 7 185 4 . 1 0 1 . 3 5 9 . 8 0 6 . 4 0 2 . 4 11 .7 4 5 - 5 0 7 5 . 9 4 . 3 0 . 0 3 49 0 . 6 3 0 . 0 6 0 . 1 0 0.11 2.1 1 . 8 5 5 - 6 0 4 6 . 8 4 . 5 0 . 0 6 39 0.71 0 . 0 6 0 . 2 8 0 . 1 0 1 . 4 2 . 9 8 0 - 8 5 4 4 . 0 5 . 4 32 0 , 7 5 0 . 0 5 0 . 4 6 0 . 0 9 2 . 0 4 . 2 27 0 - 5 2 6 2 . 7 4 . 0 295 4 . 4 5 0 . 3 7 1 0 . 2 0 7 . 0 5 7 . 7 6-10 3 5 6 . 6 3 . 9 301 6 . 9 5 1 . 8 7 9 . 6 0 9 . 2 0 9 . 2 2 6 - 3 0 4 0 . 6 4 . 4 41 0 . 7 5 0 . 0 8 0 . 7 2 0 . 8 5 5 . 9 3 5 - 4 0 4 8 . 5 4 . 9 49 0 . 8 4 0 . 0 6 0 . 2 8 0 . 1 0 2 . 6 6 0 - 6 5 3 8 . 8 5.1 42 0 . 6 4 0 . 0 6 0 . 3 3 0 . 0 6 2 . 6 APPENDIX IV Radiocarbon dating 12.$; boratoried, nc. 24 Blackstone Street, Cambridge, Mass. 02139 Telephone TRowbridge 6-3691 2 August 1965 REPORT DF ANALYTICAL WORK RADIOCARBON AGE DETERMINATION Our Sample No.: GXO^S Sample Name: Vancouver I s l a n d peat, Your Reference No, L e t t e r : 18 May 1965 AGE m 3SQ \u00C2\u00B190 c-m y e a r s 8. P. (1560 A. D.) D e s c r i p t i o n : L o c a t i o n : O c c u r r e n c e : . C o l l e c t e d : S u b m i t t e d by: Comments: S i l t y p e a t . West c o a s t o f Vancouver I s l a n d , B r i t i s h Columbia, Canada. B a s a l peat o v e r l y i n g compact clay-- i n t u r n o v e r g l a c i a l outwash g r a v e l s i n t u r n o v e r g l a c i o - m a r i n e t i l l . Depth t o peat i s Vk m e t e r s . E s t . s l i g h t l y l e s s than 10-11,000 y e a r s . Not g i v e n . G. E. Rouse, U n i v e r s i t y o f B r i t i s h C olumbia, Vancouver, Canada. The sample uas t r e a t e d w i t h hot d i l u t e H C l , h ot NaOH, and a g a i n w i t h H C l , to remove a l l s o l u b l e m a t e r i a l s p r i o r to a n a l y s i s . ' j Notes: This date is based upon the Libby half l i f e (5570 years) for C The error stated is \u00C2\u00B1 1\u00C2\u00AB>\" as judged by the analytical data alone. Our modern standard is 95% of the activity of N. B. S. Oxalic Acid. "@en . "Thesis/Dissertation"@en . "10.14288/1.0104856"@en . "eng"@en . "Botany"@en . "Vancouver : University of British Columbia Library"@en . "University of British Columbia"@en . "For non-commercial purposes only, such as research, private study and education. Additional conditions apply, see Terms of Use https://open.library.ubc.ca/terms_of_use."@en . "Graduate"@en . "Vegetation and history of the sphagnum bogs of the Tofino area, Vancouver Island"@en . "Text"@en . "http://hdl.handle.net/2429/37514"@en .