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Vegetation and history of the sphagnum bogs of the Tofino area, Vancouver Island Wade, Leslie Keith 1965

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VEGETATION AND HISTORY OF THE SPHAGNUM BOOS OF THE TOFINO AREA, VANCOUVER ISLAND  by  LESLIE KEITH WADE B.Sc. The U n i v e r s i t y of B r i t i s h 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 t h i s thesis as conforming to the required standard  THE UNIVERSITY OF BRITISH COLUMBIA September 196$  In p r e s e n t i n g  fulfilment  of  the requirements f o r an advanced degree at the U n i v e r s i t y  of  British  Columbia,  available  for  this  thesis  I agree that  in p a r t i a l  the L i b r a r y s h a l l  r e f e r e n c e and s t u d y .  I f u r t h e r agree that  m i s s i o n f o r e x t e n s i v e copying o f t h i s purposes may be granted his  representatives.,  cation of this  thesis  without my w r i t t e n  It  permission.  SEPT.  for  per-  scholarly  i s understood that copying o r p u b l i -  for financial  The U n i v e r s i t y o f B r i t i s h Vancouver 8, Canada  zy  thesis  freely  by the Head o f my Department o r by  Department of  pate  make i t  Columbia  /its  gain s h a l l  not be allowed  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 t o r i c a l aspects.  An integrated approach to these three aspects was attempted i n  order to give i n a r e l a t i v e l y l i m i t e d time as complete a picture as possible of the bog ecosystem. The bog vegetation was studied on 110 sample plots using a n a l y t i c a l and synthetic methods of the Zurich-Montpellier school of phytosociology.  Ten  d i f f e r e n t vegetation types were described and characterized, nine belonging to the bog ecosystem and one to the surrounding scrub f o r e s t .  The nine bog  vegetation types consist of f i v e d i s t i n c t associations and one association composed of f i v e v a r i a n t s .  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 l u r i f l o r a e (Carex p l u r l f l o r a association)  2.  Scirpeto-Sphagnetum mendoclnl (Sclrpus caespitosus - Sphagnum mendocinum association)  3.  Oxycocceto-Sphagnetum p a p i l l o s i  '  (Oxycoccus quadripetalus - Sphagnum papillosum association) High moor bog association* U*  Ledeto-Sphagnetum c a p l l l a c e l (Ledum groenlandlcum - Sphagnum capillaceum association)  -•  Peripheral bog associations! 5>.  (Bog-forest  transition)  Pineto-Sphagnetum c a p i l l a c e i (Pinus contorta - Sphagnum capillaceum association) a. Pineto-Sphagnetum c a p i l l a c e i sphagnosum p a p i l l o s i (Pinus contorta hummock v a r i a n t ) b. Pineto-Sphagnetum c a p i l l a c e i myricosum g a l i s (Myrica gale variant) c. Pineto-Sphagnetum c a p i l l a c e i chamaecyparosum nootkatensis (Chamaecyparis nootkatensis variant)  Secondary succession variants established a f t e r f i r e * d. Pineto-Sphagnetum c a p i l l a c e i vacciniosum v i t i s - i d a e a e (Vaccinium v l t i s - i d a e a variant) e. Pineto-Sphagnetum c a p i l l a c e i vacciniosum p a r v i f o l i i (Vaccinium parvifollum variant) Scrub f o r e s t association surrounding bogs: 6.  Pineto-Chamaecypareto-Sphagnetum r e c u r v i (Pinus contorta - Chamaecyparis nootkatensis - Sphagnum fecurvum association)  (Bog f o r e s t )  Edaphic considerations were l i m i t e d to the analysis of s o i l s from representative sample p l o t s of each association and v a r i a n t .  S o i l s were  analyzed f o r a v a i l a b l e cations, i n c l u d i n g C a ' , M g , N a , K*, adsorbed f+  + +  +  phosphate, t o t a l nitrogen, cation exchange capacity, percent base saturation pH, and s o i l moisture.  The r e s u l t s 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 i m i t e d an area as the one under study. H i s t o r i c a l considerations included a p o l l e n analysis from a  112 J representative core i n 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 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 i s suggested that the distribution of the bog plant associations i s primarily dependent upon a complex of environmental factors that are dependent upon topography.  IV,  TABLE OF CONTENTS Page  Chapter I II  III  1  INTRODUCTION DESCRIPTION OF THE AREA . ....  .. 1  A. Topography and vegetation .  ;, l  B.  Climate  C.  Geology  7 . . . . . . . . . .  METHODS . . .  .  11  . ..  11  A.  General  B.  Reconnaissance  11  C.  Plot selection and analysis  11  13  D. Mensuration E.  Synthesis  •.  13  F.  Soil sampling and analysis.  15  1.  15  Field procedure . . . .  2. Analysis  17  9  .  15  G.  Pollen analysis  16  H.  Radiocarbon dating  16  DESCRIPTION OF THE ASSOCIATIONS A.  18  Caricetum pluriflorae (Carex pluriflora association),  19  Scirpeto-Sphagnetum mendocini (Scirpus caespitosus - Sphagnum mendocinum association) . . . . . . .  21  C. Qxycocceto-Sphagnetum papillosi (Oxycoccus quadripetalus - Sphagnum papillosum association) . . . .  23  D. Ledeto-Sphagnetum capillacei (Ledum groenlandicum - Sphagnum capillaceum association) . . .. . . . . . . . . . . . .  28  B.  v: Chapter  ' E.  Pineto-Sphagnetum capillacei (Pinus contorta - Sphagnum capillaceum association). .. 1.  '  2.  3.  F.  Page  Pineto-Sphagnetum capillacei sphagnosum papillosi (Pinus contort a hum-nock variant)  33  Pineto-Sphagnetum capillacei myricosum galis (Pinus contorta - Sphagnum capillaceum - Myrica gale variant) . . . . . . . .  37  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  Pineto-Chamaecypareto-Sphagnetum recurvi (Pinus contorta - Chamaecyparis nootkatensis . Sphagnum recurvum association) (Bog forest) . . . . . . . . . . . . . . . . . . . . .  V  31  £l  SOILS  58  A.  Soil moisture  .  $9  B.  Total nitrogen  C.  Available sodium  D.  Available potassium. . . . . . . . . . . . . . . . . . .  60  E.  Available, calcium  60  F.  Available magnesium. . . . . . . . . . . . . . . . . . .  G.  Adsorbed phosphate . . . . . . . . . . . . . . . ..  H.  Cation exchange capacity .  I.  Percent base saturation.c . . . . . . . . . . . . . . .  61  J.  pH . . . '  61  K.  Discussion of soils  60 . ..  .  60  60 - 61  . . . . . . . . . . . . . . 6 1  ;  * ......  .  61  Chapter VT VII VIII DC  X  Page 6h  LIFE FORMS  67  HISTORY POLLEN ANALYSIS SUCCESSIONAL TRENDS. . .  • ......  70 76  A. Regional level  76  B.  78  Association level.  SUMMARY AND CONCLUSIONS  83  1.  BIBLIOGRAPHY  87  2.  BIBLIOGRAPHY OF PUBLICATIONS USED IN THE IDENTIFICATION 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 APPENDIX II Explanation and Legend for Synthesis tables Synthesis tables I - VI  93  ...  97  APPENDIX III Soils data . . . . . . . . . . . . . . . . . . . . . . . .  118  APPENDIX IV Radiocarbon dating  12U  v ii 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 i n 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 i n low moor bog association  26  12.  Oxycoccus quadripetalus i n Oxycoccus quadripetalus - Sphagnum papillosum association  13.  lli.  .  Mosaic community of Pinus contorta hummock variant and Oxycoccus quadripetalus - Sphagnum papillosum association . . . . . .  26  27  Trientalis arctica, Linnea borealis, and Cornus canadensis growing i n Pinus contorta hummock v a r i a n t . . .  27  15.  High moor Sphagnum fuscum hummock with associated species  31  16. 17.  Apargidium boreale growing on Sphagnum fuscum hummock . . Rhacomitrium lanuginosum, Empetrum nigrum, and Kalmia p o l i f o l i a growing on dry hummock surrounded by Carex pluriflora association. . . . . . .  31 3£  Pinus contorta hummock variant surrounded by low moor bog association . . . . . . . . . . . . ...  35  Myrica gale variant .  38  18. 19.  Figure  Page  20*  Thuja plicata and Pinus contorta growing i n mature high moor community  38  21.  Characteristic form of old Tsuga heterophylla i n Long Beach area bogs . .  Ill  Pinus contorta, Tsuga heterophylla and Thuja plicata at edge of bog forest  hi  Secondary successional communities on burned area of bog. Open area represents Vaccinium vitis-idaea variant. . . .  U3  Boschniakia hookeri parasitic on Gaultheria shallon of Vaccinium vitis-idaea variant  h3  Ledum groenlandicum, dominant shrub of Ledum groenlandicum - Sphagnum capillaceum association  hh  Kalmia p o l i f o l i a , co-dominant shrub of Vaccinium v i t i s idaea variant . . . i . . .  hk  27.  Coptis t r i f o l i a t a and Empetrum nigrum growing i n Vaccinium vitis-idaea variant i .  Ii5  28.  Nephrophyllidium crista-galli growing i n wet depressions  22. 23. 2iu 25. 26.  of burned area of bog .  U5  29.  View of bog forest, Pinus contorta predominating. . . . .  52  30.  View of bog forest, with Chamaecyparis nootkatensis,  31.  Thuja plicata, Tsuga heterophylla, and Pinus contorta . . Characteristic form of old Pinus contorta i n bog forest  52  of Long Beach area (height 75 feet, age 260 years). . . .  53  32.  Life form distribution  66  33.  Pollen profile  75  3U.  Successional trends .  81  IX.  Acknowledgement  I  wish to express sincere thanks to Dr. 7. J . Krajina f o r providing  the i n i t i a l stimulus f o r t h i s study, and f o r his constant help and guidance throughout i t s completion.  His help i n the i d e n t i f i c a t i o n of many vascular  p l a n t s , bryophytes and lichens i s also greatly appreciated. Grateful thanks are also extended to Dr. G. E. Rouse, who provided generous assistance with the p o l l e n a n a l y s i s , geology, and radiocarbon dating aspects of the t h e s i s , and to Dr. W. B. Schofield who read the manuscript and gave valuable advice* I  wish also to thank the National Research Council, Ottawa, f o r  financing t h i s study, and Dr. T. M. C. Taylor and Dr. G. H. N. Towers f o r 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 s h Columbia. L a s t l y , I am g r a t e f u l to my fellow students, Richard T. Kuramoto f o r assistance i n the f i e l d and f o r h e l p f u l advice, Robert C. Brooke f o r valuable discussions and advice, and to my f i a n c e e , Adele C. Templeton, f o r her many hours of work on a l l aspects of the t h e s i s .  CHAPTER I INTRODUCTION  .  .  Sphagnum bogs have long been a popular subject for ecological studies, probably because they represent ecosystems that are i n 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 i n comparison to other ecosystems.  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. quite unique for most areas.  In addition, the physiogonomy of bogs i s usually 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 r e l i c s . Although Sphagnum bogs have been very extensively studied from many aspects i n 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 i n western North America consist of investigations into the stratigraphy of bogs, pollen analyses, and l i 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 i s  known of the detailed patterns of bog vegetation i n this area.  A Swedish  2 worker (Osvald, 1933) attempted a b r i e f vegetational study of the bogs of southwestern mainland B r i t i s h Columbia, and established tentative s o c i a tions f o r several major bogs of the area.  Subsequently Krajina (195>9) and  O r l o c i (I960) have described several bog associations i n the course of regional vegetation studies. The purpose of t h i s thesis i s primarily t o present i n d e t a i l the vegetation of the Sphagnum bogs of a d e f i n i t e l i m i t e d area, and thereby t o e s t a b l i s h a basic series of bog associations  which can be used as a  comparison f o r future bog studies i n other areas.  The Tofino-Ucluelet  region was chosen as a study l o c a t i o n because of i t s many r e a d i l y a c c e s s i b l e , yet r e l a t i v e l y undisturbed bogs.  The problem of the b i z a r r e forest of  dwarf conifers that also exists i n the bog areas provided an a d d i t i o n a l highly i n t r i g u i n g aspect.  In addition t o a detailed c l a s s i f i c a t i o n of the  bog associations, an important aspect of the study included the c o r r e l a t i o n of the plant associations with environmental data, and the i n t e g r a t i o n of an h i s t o r i c a l approach i n the form.of a p o l l e n 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 u p l i f t .  The  terrace reaches i t s best development i n the Long Beach area i n the vicinities of WLckanninish and Wreck Bays, where i t averages approximately seventy feet above sea-level.  The surface of the terrace i s characterized by a gentle  topography of very low h i l l s and broad shallow valleys, extensive flat areas, and a total absence of any sharp r e l i e f features (Fig. 3). The vegetation of the terrace i s 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 l i e s immediately adjacent to the open ocean, and occupies, i n 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 i n height above a dense shrub layer of salal (Gaultheria shallon).  Nowhere i s this strip of pure Sitka spruce more  than a few hundred feet wide, and nowhere i s i t found except fronting the open ocean, the inference being that i t s distribution i s i n some manner controlled by the presence of the open ocean.  2  Inland from the b e l t of Sitka spruce the p r o f i l e of the terrace generally slopes s l i g h t l y downward, leaving a l i p of s l i g h t l y higher land that appears to e f f e c t i v e l y block drainage from many areas of the t e r r a c e . This portion of the terrace i s occupied by three vegetational types the d i s t r i b u t i o n of which i s controlled by drainage, which i n turn i s at l e a s t p a r t i a l l y c o n t r o l l e d by topography.  The lowest parts of the region are  shallow b a s i n - l i k e areas f i l l e d with shallow Sphagnum bogs, very s i m i l a r 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 t e r r a c e , p a r t i c u l a r l y  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 b i z a r r e 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 l e s s e r extent are a l s o found western red  cedar (Thuja p l l c a t a ) , western yew (Taxus b r e v i f o l i a ) , and occasionally western white pine (Pinus monticola).  Bog species are a l s o usually present,  p a r t i c u l a r l y Ledum groenlandicum and Sphagnum recurvum, prompting the name "bog f o r e s t " to be adopted f o r t h i s vegetational type.  Trees of t h i s bog  forest are seldom over seventy f e e t 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 t y p i c a l of the wetter subzone of the Western Hemlock Zone (Krajina,  1959; O r l o c i , 1961).  This vegetation type i s  r e s t r i c t e d to the higher parts of the terrace and generally occurs above the scrub or bog f o r e s t , probably i n d i c a t i n g better drainage as a probable distributional factor.  Besides i n c l u d i n g western hemlock and amabilis  fir  a high percentage of western red cedar i s commonly present, while within a h a l f mile of the open coast amabilis f i r i s usually completely l a c k i n g .  PROFILE O F  TERRACE  Western Hemlock Sitka  Spruce  Bog Forest  Amabilis F i r Bog  of t e r r a c e  Beach i  Sands  and gravels  Glacial m a r i n e till  Upper Triassic volcanics  Figure 3.  Profile of terrace.  Although the climax forest i s r a p i d l y being destroyed by logging, a fen excellent stands remain, with trees up to l £ 0 and 200 feet i n height.  The  climax forest meets and merges with the S i t k a spruce forest along many parts of the t e r r a c e .  The t r a n s i t i o n a l region, however, i s very narrow.  In  other areas the bog f o r e s t borders the Sitka spruce s t r i p , the t r a n s i t i o n 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 S i t k a spruce i s common. The bogs of the area served as the main theme of the present study, although, to a l e s s e r extent, the bog forest also received a t t e n t i o n .  Area  of the bogs throughout the area varies from about two acres to approximately eighty acres; a e r i a l photographs show the presence of much l a r g e r 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 i m i t e d .  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 t y p i c a l of e a r l y 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 c h a r a c t e r i s t i c of topogenoua or low moor bogs (Heusser, I960).  Portions of  the bog are completely submerged f o r a l l except very dry periods i n dry summers, and i n these areas the plant associations c l o s e l y r e f l e c t 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 i m i t e d areas around the margin of some of  >  6  the bogs. the bog.  It exists also as a secondary feature of seme burned portions of The common bog shrubs of the region, Ledum groenlandicum, Kalmia  p o l i f o l i a , 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 i n i t i a t i o n of high moor development i s indicated by the invasion of Sphagnum capillaceum and Sphagnum fuscum, particularly the latter species.  Invasion i s 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 i n size and quantity than i n other parts of the bog. Throughout the entire bog area Pinus contorta i s represented by stunted, dwarfed specimens spaced at f a i r l y regular intervals over the bog surface.  These trees, seldom over lf> feet i n height and often more  than 100 years old, support an association of plants on their basal hummocks that i s typical of much drier conditions than the bog surface i t s e l f .  For  the purposes of this study these stunted trees and their associated hummock plants have been treated as a unit, i n many aspects different from the "interhummock" areas.  Chamaecyparis nootkatensis i s 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 i n height and often over 100 years i n age. Chamaecyparis nootkatensis i s 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 f i r e s , both by the presence of charred logs and stumps, and i n some cases by the absence of a n e l l developed organic layer. • '  . • '  '  The secondary vegetation that •  '  i /  has developed on these areas shows marked differences i n 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 i s the dominant tree species and ranges i n i t s  occurrence from dwarfed, isolated individuals to scattered, dense clumps of trees. The bogs are very shallow, the accumulated peat,averaging one meter thick and i n exceptional situations extending to two meters i n depth. Xn most instances a layer of clay or clay mixed with organic material i s evident 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 i n the basal portion of a typical bog profile indicated the very recent date of 3°oi 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 i s mild marine humid, designated by the Koppen  scale (Koppen and Geiger, 1936) as Cfb (Krajina, 1959).  Extremes i n  temperature are rare, with a mean for the warmest months, July and August, of 58° F., and a mean for the coolest month, January, of Ul° 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° F. and 91° F. Rainfall i s extremely high, with an' annual mean of 120.97" recorded at the Tofino airport over a nine year period. f e l l as snow.  Of this amount only k*S  n  8  Climatic  data — Tofino A i r p o r t  Wean Monthly Jan.  Feb.  March  April  May  June  Temperatures July  •  Aug.  Sept.  Oct.  Nov.  Dec.  op 60  55  50  45  -  40  Annual Mean Monthly Jan.  Feb.  March  April  May  June  mean  Precipitation July  49°F  Aug.  Sept.  3.09  5.08  Nov.  Oec.  Inches ppt.  20  15  •  10  11.69  14.04  13.24  11.37  4.24  3.30  2.63  13.39  16.47  17.43  inches Annual  ppt. total  120.97"  Figure h.  Climatic data.  Summer, although the driest and warmest period, is relatively cool and •wet when compared to the summer dry situation that exists i n much of British Columbia.  Summer fogs are common and often last from daybreak until early  afternoon. In summary, the climate i s 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 F i r 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 f i 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 i s prevailingly of dark colored andesite.  Observations by Rouse  (personal communication, May 1965) and myself suggest the following modifications of this interpretation. Bordering Wreck Bay i s 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 i s 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 s i t e , 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 i n this latter sequence. This upper sand-gravel sequence thickens noticeably toward the southeast along the beach c l i f f .  Distinct cross-bedding i s evident, the nature of  which indicates a general northerly source direction.  The character of the  upper sequence, i n 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 i n i t i a t i o n  i s very recent, as shown by the Radiocarbon ( C ^ ) dating of 390± 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 i n the underlying sands were the most l i k e l y 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 i n 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 i n the study are basically those used  ;  by Krajina (19^2) and his students i n previous ecological studies i n British Columbia. B.  Reconnaissance Reconnaissance was carried out for short periods i n June 1963 and for  two weeks i n August 1963.  L  These periods were spent i n becoming familiar  with the flora of the area and i n 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 u t i l i z e  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 d i f f e r e n t t e n t a t i v e plant  types.  association  Each type of vegetation that appeared f l o r i s t i c a l l y homogeneous  and that appeared, under s i m i l a r conditions i n other areas, to repeat i t s e l f was t e n t a t i v e l y regarded as an association.  Homogeneity was defined on the  basis of the regular d i s t r i b u t i o n of plant species throughout the area i n question.  It  was attempted, therefore, to l o c a t e 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 a s s o c i a t i o n , and ranged from 0.5 m  f o r 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 f o r the highly complex surrounding bog f o r e s t . Analysis of the vegetational aspects of the plots was done using the basic methods of the Zurich-Montpellier school (Braun-Blanquet, Krajina,  1932}  1933} Becking, 1957). For each plot estimates were made of the  percent cover occupied by each of the following s t r a t a : A.  Tree l a y e r 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  B.  feet i n height  Shrub l a y e r B^ - shrubs between 6 inches and 6 feet i n height Bg - shrubs under 6 inches i n height  .  C.  Herb l a y e r - a l l herbaceous p l a n t s , regardless of height  D.  Bryophyte and l i c h e n l a y e r Dh - bryophytes and lichens growing on humus Ddw- bryophytes and lichens growing on decaying wood  E.  Epiphyte l a y e r (includes bryophytes, l i c h e n s , and vascular plants) - epiphytes i n the A l a y e r  ,  Eg - epiphytes i n the B layer EQ - epiphytes i n the C layer While this scale varies i n i t s 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. A l l species i n each plot were also rated according to species s i g n i f i 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 v i t a l i t y of each species,  was interpreted on a four point scale. The scales used for species significance, sociability, and vigor are given i n 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 i n 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 analy ed plots was arranged i n groups of tentative z  communities. Much comparison and correlation was then made to check that sufficient f l o r i s t i c similarity existed between sample plots of a tentative  association to v e r i f y i t s e n t i t y as a d i s t i n c t association.  Constancy, the  frequency with which one species occurs within a group of plots of uniform s i z e , was calculated f o r each species.  The degree of constancy was expressed  i n the f i v e class scale of Braun-Blanquet (1932) as s l i g h t l y modified by Brooke ( 1 9 6 5 ) .  Average cover value, an expression of dominance, was  calculated by converting the species s i g n i f i c a n c e values. scale used i s that of Brooke ( 1 9 6 5 ) , see Appendix I.  The conversion  Vegetation units that  were t e n t a t i v e l y c a l l e d associations i n the f i e l d were occasionally found a f t e r synthesis to be l e s s d i s t i n c t than previously thought.  These units  which d i f f e r e d from each other by only minor, although s i g n i f i c a n t l y d i f f e r e n t aspects, were then delegated the rank of association v a r i a n t , rather than that of association (Domin 1 9 3 6 ) . Each plant association i s defined by i t s c h a r a c t e r i s t i c combination of species, a category which includes both constant and c h a r a c t e r i s t i c 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 l e s s than 10% cover Characteristic species (Braun-Blanquet, 1 9 3 2 ) Exclusive species - species completely or almost completely confined to one community Selective species - species found most frequently i n a c e r t a i n community but a l s o , though r a r e l y , i n other communities P r e f e r e n t i a l species - species present i n several communities  '  •  1  5  more or less abundantly but predominantly or with better v i t a l i t y i n 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 i n at least two plots of each tentative vegetation type, and s o i l samples collected for analysis.  Although s o i l 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 i n pits of the same community type.  When horizons were not  distinguishable, samples were collected at regular intervals i n 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 f i e l d weights and the f i n a l oven-dried weights. The pH was measured for each, s o i l sample collected, using the Beckmann Model N pH meter. Before measuring, each s o i l sample was mixed with d i s t i l l e d water until a thick paste consistency was reached, after which  -  16 i t was allowed to stand overnight. Selected s o i l samples were analyzed for cation-exchange capacity and for exchangeable Na , K , C a , 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, i n 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°6U). ' 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 i n 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 r e l i a b i l i t y of . 9 , an acceptably accurate figure (Barkley, 193U).  Identification of representative genera  present i n 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  f o r radiocarbon ( C ^ ) Inc.,  Cambridge, Mass.  dating.  The sample was dated by Geochron Laboratories  CHAPTER I? DESCRIPTION OF THE ASSOCIATIONS The area of the study bog i s occupied by ten recognizable types, five of which are much more distinctive than the others.  vegetation The five  distinctive vegetation types differ widely from each other i n both composition 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 i s composed of five association variants. Most plots analyzed were situated i n the study bog (Fig. 6), but many plots were analyzed from other bogs i n 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 i s a definite uniform plant canmunity ....  that i s i n equilibrium with a certain complex of environ-  mental factors, . . . .} i t s f l o r i s t i c structure . . . . l i e s 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 i n a more inclusive concept than that which considers vegetation alone.  For this reason, ideally,  a l l possible environmental factors are considered i n 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  Myrica gale  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  L-Sp  Ledum - Sphagnum capi 11 aceum a s s o c i a t i o n  Vv-i  Vaccinium 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  Pinus - Vaccinium p a r v i f o l i u m v a r i a n t  All  variant variant  v a r i a n t s belong to the Pinus - Sphagnum capi 11 aceum a s s o c i a t i o n .  Figure 5. Bog profile. C O  So  Figure 6.  BOG  Sketch map of study bog, showing distribution of associations.  FOREST  BOG  Carex p l u r i f l o r a a s s o c i a t i o n S c i r p u s - Sphagnum mendocinum  ! association  j  Myrica gale Chamaecyparis  FOREST  variant nootkatensis  variant  Oxycoccus - Sphagnum p a p i l l o s u m a s s o c i a t i o n  Vaccinium v i t i s - i d a e a  Ledum - Sphagnum c a p i l l a c e u m a s s o c i a t i o n  Pinus - Vaccinium p a r v i f o l i u m  All  variants  belong t o the 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 .  variant variant  19 A similar concept i s implied by the term "biogeocoenosis" of Sukachev (1950), and the term "ecosystem" of Tansley (1935)• 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 i n a drop of pond water. At whatever level i t i s applied, however, the all-inclusive aspect i s implied. In some cases, associations are described f l o r i s t i c a l l y 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 f i d e l i t y (characteristic species) although i n 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 I I ) . A.  Caricetum pluriflorae (Carex piuriflora association) The Caricetum is f l o r i s t i c a l l y the simplest of a l l the bog associations.  An aquatic type, i t i s covered with standing water i n a l l but the driest periods of very dry summers, and i n average summers may not dry out at a l l . It occurs i n .the lowest areas of the Long Beach bogs and i s usually of quite limited extent. tions.  Its occurrence possibly reflects r e l i c marsh or fen condi-  Often the Caricetum pluriflorae occupies hollows l i t t l e more than  1§- m i n diameter, while at most i t covers areas up to 5 m across.  In the  center of the bog, where i t i s best represented, i t s distribution i s often broken up by patches of the surrounding Oxycoccus - Sphagnum papillosum association.  1  20  Figure 7.  Figure 8.  Carex pluriflora association surrounded by Oxycoccus quadripetalus - Sphagnum papillosum association  Juncus oreganus i n deepest water of Carex pluriflora association  /  The association i s characterized by 15-25 cm of normally standing water i n -which, after heavy rains, a slight current i s often noticeable. In the bottom of the pool, a layer of dy (Kubiena, 1953) i s found. The s o i l profile i s seldom over UO cm i n depth, at the bottom of •which the underlying sands are found. The dy i s underlain by 20-25 cm of peat, beneath which a gray clay-like material extends down to the sand. abundant only i n the brown organic layer.  Roots are  As i n a l l the Long Beach bog  associations, the pH i n the surface layers i s strongly acid and ranges from 3.U to U.l.  At the sand level i t averages  F l o r i s t i c a l l y , the association i s dominated by a usually luxuriant growth of Carex pluriflora, a preferential, as well as the only constant species.  An aquatic form of Sphagnum recurvum i s common (average species  significance I4.O) and S. papillosum i s also frequently found.  Apart from  Carex, the only commonly occurring vascular plant species i s Sanguisorba microcephala.  Sporadic species (those species occurring i n 20$ or less of  the plots of an association) include the following: Vaccinium uliginosum Kalmia p o l i f o l i a Carex obnupta Gejntiana sceptrum Xn a few areas the water i s deeper than 30 cmj i n such situations Carex pluriflora i s sparse or non-existent. occurs frequently.  Under such conditions Juncus oreganus  The presence of this species possibly indicates the  last remnant of an association that preceded bog development, and that ~ flourished i n the deeper waters of a marsh or fen environment. B.  Scirpeto-Sphagnetum mendocini (Scirpus caespitosus - Sphagnum mendocinum association) The Scirpeto-Sphagnetum mendocini i s of rather limited extent i n the  study bog but covers large areas of some of the surrounding bogs. I t consists primarily of areas dominated by clumps of Scirpus caespitosus, among which i s either standing water or mats of Sphagnum papillosum and S, mendocinum. Rhynchospora alba i s a common preferential constituent, and i t often occurs i n 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.  F l o r i s t i c a l l y , the  Scirpeto-Sphagnetum i s more complex than the Carex pluriflora association, as i s shown by the following l i s t s Characteristic combination of speciesJ constant dominants: Scirpus caespitosus Sphagnum papillosum constant non-dominants: Kalmia p o l i f o l i a  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 t r i f o l i a t a  Cladonia pacifica  Bazzania ambigua Sphagnum papillosum i s the dominant Sphagnum species here, as i n most  23 of the Long Beach bog associations.  Where tree species such as Pinus contorta  and Chamaecyparis nootkatensis occur i n the association proper, they do so only as small, twisted shrubs growing i n standing water.  F l o r i s t i c a l l y , the  Scirpeto-Sphagnetum is a typical low moor association, characteristic of an early stage i n bog  succession.  As i n the Carex pluriflora association, the depth of organic accumulation i s relatively shallow.  The upper 20-30 cm of the s o i l profile are a  dark brown peaty organic material, underneath which i s often a 10 cm layer of sticky gray to gray-brown clay-like material. 40-60 cm level i s sand.  Underlying the clay at the  As i n the Carex pluriflora association, l i t t l e  organic accumulation appears to have been of Sphagnum. Roots are abundant only i n 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 i n the bogs of the Long Beach region, and covers large areas of the interior of each of the larger bogs.  The association occurs throughout i t s 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 i n drier areas such as the bog periphery, while the Oxycocceto-Sphagnetum i s 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 i s a low-lying almost flat association, which i n early spring i s extremely wet, the water level reaching the top of the Sphagnum papillosum mat that dominates the association*  Although  no single species i s characteristic of the association, many of the species present are constants, and Sphagnum papillosum here reaches i t s greatest dominance and vigor (av. species significance 9.0, av. vigor 2.9). This species i s characteristic of low moors and early bog succession stages. Sphagnum capillaceum, a bog species more often associated with high moors, i s 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, i s 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  Kalmia p o l i f o l i a  Agrostis aequivaivis  Trientalis arctica  Drosera rotundifolia  Tofieldia occidentalis  Sporadic species: Vaccinium uliginosum  Myrica gale  Empetrum nigrum  Pinus contorta  Thuja plicata  Scirpus caespitosus  Sphagnum tenellum  Sphagnum fuscum  Cephalozia biscuspidata  Polytrichum commune  .'  The accumulation of peat i s much deeper than i n the previously described associations, reaching a depth of 50-100 cm. The organic material varies i n  Figure 1 2 . Oxycoccus quadripetalus i n 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 i s and Cornus canadensis growing i n 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£0 cm.  Roots are  numerous i n the top 20-30 cm, but rapidly decrease below this l e v e l .  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 i t s 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 i s primarily an association character-  i s t i c of high moor conditions (Hansen, 19U7; Rigg, 192£j Osvald, 1933), and as such i s of limited occurrence i n the Long Beach bogs.  In the larger bogs  of the region, i t occurs i n small areas around the periphery, while i n a few. of the smaller bogs i t i s slightly better developed. It appears probable • that this association has developed recently i n this region, as i t i s not extensive nor has as much organic accumulation occurred as i n areas where i t is the dominant, well-established association (e.g. Lulu Island, Ladner, and Pitt Meadows bogs i n the Vancouver area of British Columbia). In the Long Beach bogs the Ledeto-Sphagnetum i s characterized by a surface topography of large hummocks composed of Sphagnum capillaceum and S. fuscum, among which are depressions i n 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 p o l i f o l i a and Oxycoccus quadripetalus, are also common. Carex obnupta i s abundant (species significance 7.U) i n 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 i n 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 i n the relatively shallow accumulation of organic material. In typical high moor conditions, the association i s 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 i s also part of a mosaic community, the other portion of which i s the Pinus contorta hummock association variant.  Although sufficiently different to  warrant treatment as separate entities, there i s less apparent difference between the composition of the Pinus hummock variant and the Ledeto-Sphagnetum, than between the Pinus hummock variant and the Oxycocceto-Sphagnetum. As i n the Oxycocceto-Sphagnetum, no exclusive, selective, or preferent i a l species are present, although the l i s t of constants i s relatively long. Characteristic combination of species: constant dominants: Ledum groenlandicum Carex obnupta • Sphagnum papillosum Sphagnum capillaceum constant non-dominants: Kalmia p o l i f o l i a  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 i s the deepest of any of the bog associations, reaching a depth of 65-120 cm i n the s o i l 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 i s 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 i n 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 i s Pinus contorta, which almost always occurs as a stunted tree of peculiar rounded form (Fig. 18).  Sphagnum papillosum i s  also here the dominant Sphagnum species, although S. capillaceum reaches greater development here than elsewhere i n the bogs.  Ericaceous shrubs also  reach their greatest local development here, but they do not achieve the density and vigor that typifies their occurrence i n high moors. The Pineto-Sphagnetum capillacei is composed of five distinct variants. These variants, though relatively similar i n habitat and f l o r i s t i c  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. F l o r i s t i c a l l y , they are similar, although significant minor differences do exist. The variants of the Pineto-Sphagnetum capillacei, i n 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 p a r v i f o l i i (Vaccinium parvifolium variant) The last two variants occur as secondary successional stages after f i r e .  ,  Characteristic combination of species: constant dominants: Pinus contorta Carex obnupta Sphagnum papillosum constant non-dominants: Ledum groenlandicum  Kalmia p o l i f o l i a  Empetrum nigrum  Oxycoccus quadripetalus  Apargidium boreale  Trientalis arctica  Prosera rotundifolia  Sphagnum capillaceum  Frullania nisquallensis  '  '  .  • •  '  .  '  .  '  I  ' Ii  characteristic species: preferential:  Empetrum nigrum Kalmia p o l i f o l i a Vaccinium vitis-idaea  • • >  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 i s never extensive since i t develops from a gradual buildup 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 i n 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 i n the surrounding association.  Typical of these are the lichens Cladonia b e l l i d i f l o r a , Usnea  plicata, and Sphaerophorus globosus, but many bryophytes are also present. The vegetation of the hummocks of this variant i s somewhat similar to that of the hummocks of the Ledum - Sphagnum capillaceum association, although the hummocks differ widely i n 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 i n height, with a  trunk diameter at 3 feet above ground of 1-6 inches. 6-10 foot specimens varies from 60-110 years.  Ages of representative  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 i n Sphagnum capillaceum and S. fuscum for several feet of their lengths. It i s possible that layering occurs i n this way, for the vertically emerging branch tips often closely resemble young pines i n 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 i n which Lysichitum americanum i s often present.  Lysichitum affects unfavorably the growth of Sphagnum species  i n i t s immediate vicinity because of the shading caused by i t s 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 i s less acidic (see Successional Trends).  High moor development  usually completely eliminates the Lysichitum (Osvald,  1933).  Shading from  prostrate Pinus contorta and Chamaecyparis nootkatensis also apparently inhibits the growth of Sphagnum, for many water-filled pools are found i n the bog below such specimens, particularly i n the association under discussion. Characteristic combination of species: constant dominants: Pinus contorta Carex obnupta  y  Sphagnum papillosum Sphagnum recurvum constant non-dominants: Ledum groenlandicum  Qxycoccus quadripetalus  Kalmia p o l i f o l i a  Linnaea borealis  Apargidium boreale  Agrostis aequivalvis  Drosera rotundifolia  Sphagnum capillaceum  Sphagnum mendocinum  Frullania nisquallensls  Scapania bolanderi  Dicranum scoparium  No characteristic species are found i n this variant. Vascular plant sporadic species:  1  Chamaecyparis nootkatensis  Vaccinium vitis-idaea  Tsuga heterophylla  Coptis asplenifolia  Carex pauciflora  Maianthemum dilatatum  Deschampsia caespitosa  Lycopodium clavatum  The s o i l profile i s 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 i s the sand layer at a depth of 75-90 cm. i n the top  I4O-6O  Roots are most abundant  cm and are uncommon below the brown organic layer. Very  few roots are i n the sand.  At the surface the pH varies from 3.U to 3.9,  and at the sand level from U.5 to $.h 2.  Pineto-Sphagnetum capillacei myricosum galis (Pinus contorta - Sphagnum capillaceum - Myrica gale variant) The Myrica gale variant occurs only i n a single large section of the  study bog and i s absent from most of the nearby bogs.  Although i t contains  most of the other species of the association, i t i s dominated by a dense growth of Myrica gale (species significance 8.6).  Chamaecyparis nootkatensis •  i s second i n abundance (species significance U.8), occurring most commonly as a prostrate spreading shrub, although some dwarf trees (up to 10 feet i n height) are present.  Pinus contorta occurs here as a round-topped tree of  larger size than those i n the Pinus contorta hummock variant, and reaches an average height of 1$ 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 i s 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 i s  much the same as that of those described for the Pinus contorta hummock variant, although Lysichitum americanum i s 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 i n shaded locations, such as below dense bog shrubs and i n the bog forest.  Figure 20.  Thuja plicata and Pinus contorta growing in mature high moor community.  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 s o i l profile a dark brown peaty layer extends to 75 cm i n depth and overlies a narrow layer of clay-like material. Underlying the clay at the 85 cm level i s sand.  Roots are abundant to a depth of 45-50 cm and are  s t i l l f a i r l y common to a depth of 70 cm. 3«  Below 70 cm roots were rare.  Pineto-Sphagnetum capillacei chamaecyparosum nootkatensis (Pinus contorta - Sphagnum capillaceum - Chamaecyparis nootkatensis variant) The Chamaecyparis nootkatensis variant occurs commonly around the  margins of the bogs i n 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 i n 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 i s  usually subdominant to Pinus contorta i n the A layer.  In the A layer  Chamaecyparis reaches a maximum of 1$ feet i n height and 10 inches dbh, and i s seldom more than 1J>0 years old. Pinus contorta reaches a maximum of 30 feet i n 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 i s 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 i n the Myrica gale variant, because of the decreased extent of the depressions which they normally inhabit.  As i n 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 Carex obnupta  •  1  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  Empetrum nigrum  Ledum groenlandicum  Kalmia p o l i f o l i a  Qxycoccus quadripetalus  Cornus canadensis  Drosera rotundifolia  Maianthemum dilatatum  Sphagnum capillaceum  Pleurozium schreberl  epiphytic constant non-dominants: Frullania nisquallensis  Isothecium stoloniferum  Dicranum scoparium  Scapania bolanderl  Polypodium glycyrrhiza characteristic species: ..preferential:  Chamaecyparis nootkatensis (only within the bog ecosystem)  Vascular plant sporadic species: Picea sitchensis  Vaccinium ovalifolium  Vaccinium uliginosum  Gentiana douglasiana  Scirpus caespitosus  Agrostis aequivalvis  Deschampsia caespitosa  Nephrophyllidium  crista-galli  The s o i l 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 i n the top 40 cm and are f a i r l y  common down to approximately 60 cm. Below this level few roots exist. The pH i n 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• Ledum groenlandicum, dominant shrub of Ledum groenlandicum - Sphagnum capillaceum association*  Figure 26. Kalmia p o l i f o l i a , co-dominant shrub of Vaccinium vitis-idaea variant.  Figure 27.  Coptis t r i f o l i a t a and Empetrum nigrum growing i n Vaccinium vitis-idaea variant.  Figure 28.  Nephrophyllidium crista-galli growing i n 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 i r e .  Charred branches and stumps are evidence of  past f i r e , 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 ericaceous 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 p o l i f o l i a , 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. b e l l i d i f l o r a also inhabit the drier hummocks.  The moist depressions, by  contrast, are occupied by wet habitat species such as Apargidium boreale, Drosera rotundifolia, Nephrophyllidium c r i s t a - g a l l i , and Sanguisorba microcephala.  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 i n the wet depressions, but less commonly than in the other bog associations. Pinus contorta i s abundant i n the vicinity in the Vaccinium parvifolium variant (Fig. 23), but is of only scattered occurrence i n the Vaccinium vitis-idaea variant.  Small specimens of Thuja plicata occur i n 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 p o l i f o l i a  ''7:"'''"''  Sphagnum capillaceum  . •  constant non-dominants: Vaccinium ullginosum  - Gaultherla shallon  \  Vaccinium vitis-idaea  Thuja plicata  Pinus contorta  Carex obnupta  Apargidium boreale  Cornus canadensis  Blechnum spicant  Sanguisorba  Trientalis arctica  j  mlcrocephala  Maianthemum dilatatum  Cladonia pacifica  Cladonia crispata  Cladonia rangiferina  Pleurozium schreberi  Dicranum scoparium  Rhytidiadelphus loreus  Frullania nlsquallensis characteristic species: exclusive:  Cladonia cri3pata  preferential:  Kalmia p o l i f o l i a  Empetrum nigrum  Vaccinium vitis-idaea  Vaccinium ullginosum  Cladonia pacifica  Cladonia rangiferina  The association also includes many sporadic vascular plants, the most notable of which are: Nephrophyllidium crista-galli  '  Habenaria saccata Boschniakia hookeri The s o i l profile shows a very shallow dark brown organic layer usually  not exceeding 10-15 cm i n 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 i s exposed  at the surface and i s colonized mainly by lichens.  In such situations i t i s  apparent that the organic layer has been completely burned off. Roots are most abundant i n 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 i s 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 i n height, and are not as markedly round-topped as the species i s i n other bog associations.  The physiognomy  here i s similar to that of Pinus contorta as i t occurs i n the bogs of the southwestern B.C. mainland (Ladner, Lulu Island, Pitt Meadows bogs). The specimens of western redcedar are seldom over 7 feet i n height and have quite sparse foliage. Western redcedar appears to be regenerating well i n 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 f i r e .  Occasional large pines (av. i|0 foot height, 16 inch dbh, ±200 years age) are f i r e 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 i n 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 i s much lower than i n 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 p o l i f o l i a  Vaccinium vitis-idaea  Vaccinium parvifolium Carex obnupta  ' \\•  Oxycoccus quadripetalus Blechnum spicant  Cornus canadensis  Apargidium boreale  Maianthemum dilatatum  Sphagnum papillosum  Eurhynchium oreganum  Frullania nisquallensis  Vaccinium parvifolium i s the only characteristic species. The most notable vascular plant sporadic species are: Menziesia ferruginea  Habenaria saccata  Pteridium aquilinum ,  Boschniakia hookeri  Veratrum viride  .  '  '  The s o i l profile i s similar to that of the Vaccinium vitis-idaea variant, except that the peat layer i s usually deeper, averaging 20 cm i n thickness.  Beneath the peat small amounts of charcoal appear scattered  through an ashy gray-brown horizon, a further indication of f i r e .  Under-  lying the ashy horizon i s sand which, due to cementation i n the uppermost layers, forms a f a i r l y distinct hardpan.  The hardpan occurs at an average  depth of 40-45 cm, below which the sand i s noticeably bluish i n color. Roots are abundant i n the organic layer, but are very rare i n and below the ashy layer.  The pH i s 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 f i r e , 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 f l o r i s t i c a l l y 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 p a r v i f o l i i , 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 f l o r i s t i c 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 i n this area might substantiate the  recognition of a Pineto-Vitis-idaeeto-Sphagnetum  capillacei, a seventh bog  association i n 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 i s 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 i s unlike that of any of the other gymnosperms of the region.  The apparent mechanism  of this phenomenon i s 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 i n conifers an increased growth and branching of the lateral branches, resulting in a bushy appearance.  Available calcium  in the bog associations i s much less than i n forest associations, a fact that may be the cause or partial cause of the stunted bushy form of a l l the coniferous species i n the bogs.  Similar growth of Pinus contorta i s found  i n bogs northward to Alaska (Heusser I960, Schofield - personal communication,  52  Figure 3 0 * View of bog forest, with Chamaecyparis nootkatensis. Thuja plicata, Tsuga heterophylla, and Pinus contorta.  Figure 3 1 .  Characteristic form of old Pinus contorta i n 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 l d , 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 c a t a ,  a l l have dwarfed, bushy forms and seldom grow over UO feet i n height ( F i g . 30)• Thuja p l i c a t a exhibits quite normal growth except f o r common stem die-back, and reaches a maximum height of approximately $0 feet at 250 years.  Taxus  b r e v i f o l l a occurs as a common though non-dominant bog forest element, and reaches a maximum height of 25 f e e t . The following mensurational data were taken from representative bog forest t r e e s : Height ( f t )  Pinus contorta  Diameter (ins) at U feet above ground  Uo  11.8  175  U5  10.5  150  U7  12.6  16U  U8  1U.0  180  70 Chamaecyparis nootkatensis  Thuja p l i c a t a  3U;  ;  :  •  • 16.5  '"'V" :  7.3 ;  285  \  1  3  6  39  6.2  85  39  7.1  11*  UO  13.7  : U9 Tsuga heterophylla  Age (yrs)  :  155 250  1U.5  188  29.5  10.5  32  1U.6  1UU  8.9  16U  3U.5  .:  Of these tree species, only Thuja p l i c a t a and Tsuga heterophylla appear to be regenerating under the tree canopy. were observed i n the B l a y e r .  Many saplings of these two species  No young specimens of either Pinus contorta  or Chamaecyparis nootkatensis were observed i n t h i s association.  Pinus  contorta i s completely shade intolerant (Hansen, 19U7} communication, 196U)  Krajina, personal  and thus does not germinate under a closed canopy  forest. The shrub layer i s dominated by Gaultheria shallon, but other ericaceous shrubs are also common. Bog elements are represented by Ledum groenlandicum and Vaccinium vitis-idaea.  The herbaceous layer i s 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 i s extremely luxuriant and includes Sphagnum recurvum as a constant species.  This moss occurs i n patches i n the wettest depressions of the  forest floor.  The association i s also extremely rich i n epiphytes, both i n  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 i s 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 .  ' •  Chamaecyparis nootkatensis Thuja plicata Gaultheria shallon  •••••—. ,  constant non-dominants:  /  Tsuga heterophylla  Vaccinium parvifolium  Vaccinium ovatum  Menziesia ferruginea  Maianthemum dilatatum  Hylocomium splendens  Rhytidiadelphus loreus  i>  Sphagnum recurvum Eurhynchium oreganum  Scapanla bolanderi Lepidozia reptans constant non-dominant epiphytes: Isothecium stoloniferum ];!•  Herberta adunca  Frullania nisquallensis  Scapania bolanderi  || '  Antitrichia curtipendula  Hypnum circinale  Dicranum scoparium  Vaccinium parvifolium  Cornus canadensis  Pblypodium glycyrrhiza  No characteristic species are found. Sporadic vascular plant species include: Gentiana sceptrum  Malus diversifolia Qoodyera oblongifolia  The s o i l profile shows f a i r l y distinctly the typical horizons of a podzol:  B  cc  0-5 cm  litter  5-20 cm  humus  ;  25-35 cm  whitish-gray leached eluviated layer,  HO-60 cm  gray-brown cemented illuviated layer  below 70 cm  outwash (or stream deposit) sands and gravels  Root distribution i s most abundant i n the dark brown humus layer, especially the top 30 cm which appear to be dominated by roots of Gaultheria shallon.  Roots are few i n the gray leached horizon and below.  The pH of  the s o i l of this association i s higher than that of any other bog association, but i s s t i l l highly acidic.  In the humus horizon the pH ranges from U.O to  li.5 and i n 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 s o i l surface, preventing drainage and thus preventing  satisfactory aeration.  The pedogenic process of podzolization usually  results i n the formation of a distinct hardpan which i s impermeable to water (Wilde, 1958).  When such a hardpan develops i n low-lying basins with  no drainage outlet, the probable result i s waterlogging of the s o i l , 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 i n the basic generalization that no correlation exists between the pattern of distribution 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 , IT", C a , Mg ), total nitrogen, and +  1  ++  ++  adsorbed phosphate.• 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) i s much higher than i n 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, i s , 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 i n the bog soils. however, i s much lower ( 6 . 5 me#)  Available calcium,  than that of approximately U0 me* which i s  taken as representative of bog soils (Buckman and Brady, I 9 6 0 ) .  The reason  for the low calcium figure i s due possibly to the rapid leaching that occurs in the region of the Long Beach bogs due to the high precipitation (±120 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 s o i l analysis from the bog forest differ slightly from those of the bog associations.  only  Available calcium is much  higher, averaging 24.0 me$ i n the humus layer as against 6.5 me$ for that of the bog associations.  Total nitrogen, conversely, i s lower, with an average  in the surface layers of 0.88$ as against 1.6$ for that of the bog associations.  Percent base saturation i n the surface layers of the bog forest  averages twice as high as that of the bog associations, with an average of 13.0$.  Soil moisture i s much less than i n the bog associations because of  the.inferior water-holding capacity of humus i n comparison to bog peat. In other aspects measured, however (available Na, K, Mg, adsorbed phosphate, and cation exchange capacity) no difference from the bog associations was evident. Summary of the s o i l data A.  Soil moisture In the surficial layers the s o i l moisture i s highest (1380$) i n the  Ledum groenlandicum - Sphagnum capillaceum association and lowest. (265$) i n the bog forest.  The other associations have values somewhere between  these two extremes, depending directly on the proportion of Sphagnum i n the organic layer.  The higher the proportion of Sphagnum, with i t s unrivalled  water holding capacity, the higher the s o i l moisture.  Thus the only true  high moor .^association i n the bog, the Ledum groenlandicum - .Sphagnum capillaceum association, which has the highest Sphagnum accumulation, has . also the highest percent s o i l moisture.  Correspondingly, the bog forest  and the secondary successional variants, which have l i t t l e or no Sphagnum accumulation, have also the lowest percent s o i l moisture (265$, 500$ resp.). A correlation may also be attempted between tree growth and percent s o i l  moisture.  The associations -with high s o i l moisture support l i t t l e tree  growth, probably because of resultant decreased aeration.  Topography would  thus appear to play the f i n a l distributional role, i n 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 i s 1.6%,  with no significant difference between most of the associations. The value (3*9%) for the Ledum groenlandicum - Sphagnum capillaceum association, however, i s much higher. shows only 0.88$.  The bog forest, with l i t t l e or no peat deposition,  Peat generally contains a higher total nitrogen content  than other soils (Buckman and Brady, I960).  For this reason the Ledum  groenlandicum - Sphagnum capillaceum association, which possesses the highest proportion of peat, would.be expected to have the highest total nitrogen content.  The underlying sand layers have uniform values averaging 0.08$.  C. • 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%, i n this case the extreme high and the extreme low values being from the same association. The average for the associations i s 1.5 rae%. E.  The underlying-sands  average 0.03 to 0.09 me%,  Available calcium No trends are evident except for^the bog forest, which'shows an average  figure for the humus of 2U.0 me%tions average 6.5 me%. F.  The surface layers of the other /associa-  The underlying sands range from O.U to 1.1 me%.  Available magnesium No trends are evident.  and average 5.0 me%.  The surface layers range from 3.6 to 7.3 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 i n the surface layer, whether of peat  in.the bog associations or of humus i n the bog forest, range from 2 l 6 to 278 me$. The highest values are from the bog forest.  These values are  approximately equivalent to cation exchange capacity figures from representative North American bogs (Buckman and Brady, I960). Values of the underlying sand range from 2? to 5 l me%] with an average of 1;0$. 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 i n 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 i n 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 i n 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 i s thought to be due partly to the release of  organic acids from the high moor Sphagnum species (Wilde, l°58j 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 s o i l affects i n various ways the  availability of the soil's nutrients. Iron, for example, i s more available i n 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 i n 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 i t s 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 i n pH i s 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 i s uniform, nor on available s o i l nutrients, which are also uniform.  A correlation between the two conditions probably exists:  the uniform pH maintains the availability of the same nutrients i n 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 i s a direct reflection of the very heavy r a i n f a l l of the area.  In such areas hydrogen  ions displace mineral cations i n the upper profile layers, creating an acidic environment and resulting i n a very low percent base saturation. In  comparison, representative eastern North American bogs, probably because of a much lower r a i n f a l l , 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 i t s podzol formation, the rest of the associations have s o i l profiles composed primarily of an upper organic horizon, usually of peat, and a lower clay-like horizon. These horizons vary considerably i n their respective thicknesses among the various associations.  The clay-like horizon i s usually partly organic i n  i t s highest occurrence i n the profile, while towards i t s base i t merges gradually with the underlying outwash or stream deposit sands.  1  These sands  and gravels are well cemented i n their upper layers, preventing drainage from the bog associations above them. The cementation i n the sands i s probably a result of previous podzolization i n the soils of a former forest/ which stood on whatsis now  bogland.  CHAPTER V I  LIFE  FORMS  The distribution of l i f e forms i n the associations and variants under study i s presented graphically (Fig. 32).  The system followed i s that of  Raunkiaer (193U) as modified'by Braun-Blanquet  (1965), as i t i s the best  known and most widely accepted l i f e form classification i n use today. In the present study various trends and correlations are evident.  The  associations and variants are arranged i n order of increasing f l o r i s t i c 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 i n composition). Correlated with this increase i n f l o r i s t i c complexity i s 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 i n f l o r i s t i c complexity, however, the  dominance of a few l i f e forms i s gradually replaced by a more even distribution of l i f e forms within an association.  In the more complex associations  no one l i f e form i s outstandingly dominant. It has often been stated that bogs are "enclaves of subarctic l i f e " (Deevey, 1958), and that i n many features they resemble subarctic or boreal areas. Many of the plant species found i n bogs are thought to be subarctic or boreal r e l i c s , and are sometimes numerous enough to impart a very "northern" aspect to temperate bogs.  While the absence of permafrost i n 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 i n a l l the associations, hemicryptophytes constitute the dominant l i f e form, both i n number of species and total cover. Next i n abundance are the chamaephytes and the nanophanerophytes, although these latter groups do not approach the hemicryptophytes i n number. This composition of l i f e forms i s very similar to that found i n subarctic or alpine regions (Braun-Blanquet, 1932). In the f l o r i s t i c a l l y 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 i n the proportion of macrophanerophytes, nanophanerophytes, and chamaephytes i s evident, correlated with a considerable decrease i n the proportion of hemicryptophytes.  Geophytes also increase slightly, but never assume a  significant position.  Therophytes, which are most abundant i n 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 f l o r i s t i c a l l y simple bog associations having the most extreme edaphic conditions, and a l i f e form spectrum reflecting boreal or subarctic conditions.  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  Pm  Pn  Ch  H  Oxycocceto-Sphagnetum paplbsae  Scirpeto-Sphagnetum mendocini  G  T  B  Pm  Pn  Ch  H  G  T  B  L  P  m  P  n  C  h  H  G  T  Pirwto-Charnaecypareto-Sphagnetum recurvi  •  Number of species  Pm  Figure 32. Life form distribution.  Pn  Ch  H  G  T  CHAPTER VII HISTORY The history i s i n part a summary.of various previously discussed aspects.  The study bog lies directly over apparent glacial outwash or  stream deposits and there i s l i t t l e evidence of what type of vegetation preceded.bog initiation.  It i s 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°0± 90 years B.P.  It  i s 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 i n the ; past U00 years by bog vegetation. Bogs can develop only when drainage i s seriously impeded, which i s the probable reason for the development of bogs on previously forested areas. Drainage of the valley floors where the bogs are located i s impeded by two factors:  the topography of the landscape which i n many areas allows no  drainage outlet, and the presence of a cemented layer i n the otherwise highly porous sand.  This cemented layer forms a distinct hardpan i n the  uppermost layers of the sand, and effectively prevents drainage from the overlying bog associations.  The impervious cemented layer probably had i t s  origin i n the soils of the former forest, through the process of podzolization.  Podzolization i s the normal s o i l forming process found i n a cool,  moist climate where raw humus i s accumulated i n large amounts.  Accumulation  i s usually due to retarded decomposition.  In the Long Beach-Tofino area  decomposition i s 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 i n the s o i l  profile where they are precipitated out i n the B horizon (Daubenmire, 1959J Soil Survey Manual, 1 9 5 l ) .  One common result of this process i s 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 i n a shallow valley with no drainage outlet and a high level of precipitation, the existing forest i s probably doomed.  Gradually the soil becomes water-logged, aeration i s  greatly decreased, and conditions become mature for bog invasion.  Before  conditions become too acidic for i t s existence a skunk cabbage (Lysichitum americanum) association may thrive i n the wet lowland habitat (Krajina, personal communication, 1 9 6 5 ) .  Finally Sphagnum species, i n i t i a l l y - 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 i n such  circumstances the forest destroys i t s e l f , i n that the s o i l processes initiated because of the presence of the forest eventually are the cause of i t s 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 i n time become established. This hypothetical sequence of events appears to represent the most logical  history of the bogs of the area, as i t i s 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 i s 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 i s 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 development of the present bog vegetation and to a lesser extent, changes that have occurred i n 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± 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, i n 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 i s 1-| meters deep i n 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 §• meter contains a mixture of Sphagnum, plus other plant remains. The core was taken from the center of the study bog i n the plant association of widest extent, the Oxycoccus - Sphagnum papillosum association. The pollen profile contains no peat bog species i n i t s lowest depths (Fig. 33).  Prevalent i n 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 s o i l , 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 i s shown by the presence of the following plants: Myrica (1$),  Cyperaceae and Juncaceae (10$), Veratrum (10$),  and Typha (2$).  Of these plants, Typha i s restricted to areas  of shallow water, while the other plants are a l l restricted to moist habitats.  Exceptions occur i n the Cyperaceae and Juncaceae, some of which  inhabit dry areas.  Typha also commonly occurs i n the skunk cabbage  (Lysichitum americanum) association as a secondary plant after opening of the forest by logging, f i r e , etc. As i t . i s a completely shade intolerant species, i t s presence i n 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 i n 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 i n southwestern British Columbia.  This observa-  tion i s to be expected i n view of the topography of the area, which includes no depressions deep enough for lake formation. The pollen profile shows a gradual decrease i n abundance of Pinus, Picea, and Alnus i n upper layers; at the surface pollen of these trees i s present only i n small quantities (5$,  4$, 1$, respectively).  Pinus contorta,  however, i s s t i l l a very abundant species of the present day bog vegetation. Tsuga, cf. heterophylla, Rhamnus, cf. purshiana, and Salix sp. are a l l present i n small numbers i n the profile, and reflect no definite changes i n the vegetational composition. Cupressaceous pollen, unlike that of other confiers, does not preserve well and i s rarely present i n the pollen record (Hansen, 1940} Rouse, personal communication, 1965).  For this reason the record of the coniferous  trees i s probably biased, as Thuja plicata, Chamaecyparis nootkatensis, and Juniperus communis var. montana commonly occur i n the area today, and almost certainly were extant on the bog during i t s development.  72  Veratrum, cf. vtrlde i s surprisingly abundant (up to 17$) 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, i n particular Sphagnum species.  Lysichitum, cf. americanum  i s well documented at the 90 cm l e v e l , the same point at which Veratrum reaches i t s greatest abundance. The presence at this period of both Veratrum and Lysichitum i n 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 i n wet  seepage habitats of close to neutral pH (Krajina - personal communication, 1965).  In highly acidic conditions such as are found i n Sphagnum bogs, both  species exhibit greatly reduced dominance and vigor, while i n true high moor bogs they are non-existent (Osvald, 1933J Turesson, 1916).  Both species, and  especially Lysichitum americanum, may, however, commonly exist as relics i n 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 f i r s t noted at the 100 cm depth, but as the i d e n t i f i cation of separate genera of ericaceous pollen i s 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 i n 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 i s very uncommon on the bog surface and, along with  the major tree species, probably died out i n the bog area during bog invasion.  As most of the bog i s not yet i n the bog shrub stage, Kalmia,  ledum, and other ericaceous bog shrub pollen probably does not counteract  i n quantity the loss of salal.  Myrica, cf. gale pollen i s present i n small  quantities throughout the profile, an indication of prevailingly wet, but not necessarily bog conditions.  Today Myrica gale occurs both i n bogs and,,  more commonly, around lake edges. Myrica gale grows much better i n the latter habitat, where conditions are generally less acidic than i n Sphagnum bogs. Throughout the profile the cyperaceous and juncaceous pollen (which . cannot be distinguished from each other) increases steadily from 10$ i n 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 development (Hansen, 1940, 1950} Rigg, 1 9 2 5 , 1 9 U 0 ; 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 i s 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 i n no cases were the pollen grains definitely determined to species.  Of the •  genera identified, however, the following species are the only ones present today i n 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 i s now found i n the area, but i s 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 . i n UOO years from a wet seepage forest habitat to a low moor bog i n the herb stage.  Apparent present day invasion of ericaceous shrubs and high  moor Sphagnum species suggests that succession i n 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 i s especially prevalent, forming solid mats i n 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 f i n a l stage of succession.  This  latter sequence appears impossible because no peat accumulation exists in the bog forest s o i l profiles.  Even under the patches of Sphagnum recurvum  almost no accumulation.of peat i s present, indicating that the presence of this species i s 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 i n the pollen record, i s another species of such habitats which also occurs i n the transitional area.  Carex obnupta,  another species that occurs abundantly i n skunk cabbage associations, also occurs with great frequency i n the transitional area, especially i n the Chamaecyparis nootkatensis variant. A possibility that suggests i t s e l f i s that present day changes i n vegetation, as seen along a transect from forest to bog, reflect the historical change from forest to bog that i s recorded i n the pollen profile. The probable major criterion governing the changes i s i n both instances the same*  the relative degree of aeration, or other possible environmental  factor, which i s 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 i n 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, i s 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 i n varying degree both i n the present day vegetation at the bog periphery, and i n the center of the bog as a past stage recorded i n the pollen profile. This suggested regional successional sequence, although highly speculative, appears to best f i t the observed pattern. Under i t s 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 i s 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.  • Association level  (Fig. 3^)  The term "sphagnum bog" has been defined as "that stage in the physiographic succession of an area during which i t s surface is entirely devoid of ordinary "hard" s o i l and is composed almost entirely of l i v i n g Sphagnum, immediately under which is fibrous brown peat composed mainly or entirely of partially disintegrated Sphagnum, the habitat exercising a distinctly selective influence on i t s flora" (Rigg, 1 9 2 2 ) . 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. well above the water table and often occur on slopes.  They may occur  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 f l a t , and coincide approximately with the water table. In early spring large portions of the bog are completely submerged i n several inches of water, and even i n late summer the surface i s 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, i n the  form of Sphagnum capillaceum and S. fuscum hummocks, i s 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 i s probably i n i t s 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 i n the peripheral and secondary successional associations, 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 i n time form a continuous high moor bog.  Of these  two species, Sphagnum capillaceum i s by far the most common. S. fuscum, regarded by many workers (Wilde, 1958; Osvald, 1933; and others) as occupying the f i n a l stage i n high moor development, i s restricted to the  highest hummocks, and i s commonest i n the Ledum groenlandicum - Sphagnum capillaceum association.  Other peculiarly high moor species such as  Careat pauclflora (Krajina - personal communication, 1965) also appear to be gaining i n abundance. The Pineto-Sphagnetum capillacei vacciniosum vitis-idaeae, an association variant that represents a secondary successional stage after f i r e , 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 i t s 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 p o l i f o l i a , 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 d i f f i c u l t to hypothesizej however, the following tentative sequence appears most logicalt  .  81  Carex pluriflora assoc.  Scirpus caespitosus Sphagnum mendocinum assoc.  Vaccinium vitis-idaea variant .Oxycoccus quadripetalus Sphagnum papillosum assoc  Vaccinium parvifolium variant Ledum groenlandicum 'Sphagnum capillaceum assoc  Myrica gale variant fire Pinus contorta "hummock variant Chamaecyparis nootkatensis variant  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, 1958).  I t i s probable that as the high moor develops, the Myrica gale  variant may disappear, since Myrica gale grows poorly i n 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 i n trend may be the fact that the burned areas are situated on higher ground than the low moor bog associations and i  are thus not affected as much by the rising water table. 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 i s f l o r i s t i c a l l y the simplest, as well as the wettest association.  In much of the bog, however, i t apparently i s being replaced  directly by the Oxycocceto-Sphagnetum papillosi rather than by the ScirpetoSphagnetum raendocini, which i s of rather limited occurrence.  A definite  increase i n f l o r i s t i c oomplexity i s evident as progression is made towards high moor development. Finally, the bog forest apparently i s 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 i s provided by numerous dead snags near the margin of the bog, indicating that forest trees probably once lived i n what i s 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 i s dependent indirectly on • topography.  Drainage of the soils of the region i s impeded by  impervious hardpans developed as a result of podzolization. • The development of hardpans, correlated with lack of adequate drainage outlets, produces i n the valleys water-logged soils i n which aeration i s apparently insufficient for forest development. Bogs, therefore, occupy the lowest parts of the valleys, where drainage, and thus aeration, i s 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 i s not a limiting factor. (2)  The phytosociological aspect of this study resulted i n 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 s o i l analysis revealed no correlation between community type ++  and the following s o i l characteristics: available Ca Na , +  ++  , Mg  ,  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 i s highest i n the,bog associations, intermediate i n the bog-forest transitional Associations, and lowest i n the secondary successional variants and the bog forest. The association having the greatest dominance of Sphagnum species has also the highest percent s o i l moisture.  The amount of s o i l moisture  . i s interpreted as being directly proportional to the degree of ' aeration, thus supporting the hypothesis of ( l ) . (5)  Total nitrogen i s highest i n the bog associations and lowest i n ...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  ++  i s much higher than i n most bog areas, the higher  figure possibly being accounted for by salt spray. (7)  pH was not demonstrated to play any role i n the distribution of bog plant associations within a bog area.  The surface peat of  , a l l associations and variants i s extremely acidic (3.5 to 3.7 at the surface) but no differences i n pH are found between them. '.' (8)  The bog forest differs from the bog associations by the following soil-analytical aspects:  higher available C a , less acidio pH, ++  lower s o i l moisture and lower total nitrogen. (9) . The age of the bogs, based on radiocarbon dating of a basal peat sample, i s approximately400 years.  From this and other evidence, r  )  a tentative history of the bogs of the area i s 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 i n the development of a forest. c) A period of increasing lack of s o i l aeration caused by the development of hardpans, eventually resulting i n 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 i t s lowest level an abundance of coniferous trees, the presence of seepage habitat species, and the absence of true bog species such as Sphagnum. Higher i n the profile the disappearance of seepage habitat species such as Veratrum and Lysichitum i s 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 i s 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 f l o r i s t i c a l l y simple low moor associations, through more complex low moor associations, to the high moor association.  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 groenlandicum - 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 . B.C.  ., 1957. Surficial geology of New Westminster Map-Area, 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. 1 9 3 4 . 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 : 694-708. Braun-Blanquet, J. 1 9 3 2 . 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. Wein. 865 pp.  Springer-Verlag,  Brooke, R.C. 1 9 6 5 . Ecotopes of plant communities i n the ecosystem classification of the coastal Subalpine Zone i n 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. 1 9 3 7 . The problem of Glacier Bay, Alaska: glacier variations. Geog. Rev. 27: 37-62. ""  1939. A fourth expedition to Glacier Bay, Ecology 20: 130-155.  a study of Alaska.  Dachnowski-Stokes, A.P. 1930. Peat profiles i n 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 i n 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 i n southwestern B r i t i s h 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 i n the spruce-hemlock climax of the Olympic Peninsula. Amer. Mid. Nat. 25: 290-297. . 19l*3» 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 i n 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.  1*98 pp.  Prentice-Hall, Inc.  Krajina, V.J. 1933. Die pflanzengesellschaften des Njynica Tales i n . den Vysoke Tatry (Hohe Tatra). Beihefte zum Botanischen Centrallblatt. Bd. 50, Abtlg. 11, 77l*-957 (I. T e i l ) : Bd. 51, Abtlg. 11, 1-221; (II T e i l ) .  . 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 i n B r i t i s h 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) i n central Alberta. J. Ecology 14: 317-341. Metson, A.J. 1961. Methods of chemical analysis for s o i l 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 i n their i n i t i a l stages of secondary succession. Ph.D. thesis, Dept. B i o l , 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 i n the subalpine mountain hemlock zone i n southern British Columbia. Ph.D. thesis, Dept. B i o l , 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.  :  '  436-437.  1916. Physical conditions i n Sphagnum bogs. Bot. Gaz. 6 l : 159-163.  ,  .  1916. A summary of bog theories.  .  1917. Forest succession and rate of growth i n Sphagnum bogs.  J. Forestry 15: "  Amer. J . Bot. 8:  PI. World.  19: 310-325.  726-739.  . 1918. Growth of trees i n 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. '  • . 1933. Notes on a Sphagnum bog at Fort Bragg, California. Science 77: 535-536. . 1937. Some raised bogs of southeastern Alaska •with notes on f l a t 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 i n North America. Bot. Rev. 6» 666-693. Rigg, G.B. and E.S. Harrar. 1931. Root systems of trees growing i n Sphagnum. Amer. J . Bot. l 8 s 391-397. Rigg, G.B. and CT. Richardson. 193U. The development of Sphagnum bogs i n 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 i n Sphagnum bogs. J . Forestry 20:. 1-3. Sears, P.B. and E. Janson. 1933. The rate of peat growth i n 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 i t s behavior i n Sphagnum bogs. Amer. J. Bot. 3: 189-209. Wilde, S.A. 1958. Forest soils.  2.  Ronald. 537 pp.  BIBLIOGRAPHY OF PUBLICATIONS USED IN THE IDENTIFICATION OF VASCULAR PLANTS  Gilkey, H. 1961. Handbook of northwest flowering plants. Binfords and Mort. hXk pp.  ^TV-.  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. ' . U n i v . of Wash. Press. 353 pp.  • . 1959. Vascular plants of the Pacific Northwest. Part Us Ericaceae through Campanulaceae. Univ. of Wash. Press. 510 pp. . 1961. Vascular plants of the Pacific Northwest. Part 3s Saxifragaceae to Ericaceae. Univ. of Wash. Press. 510 pp. . Vascular plants of the Pacific Northwest.Part 2: Salicaceae to Saxifragaceae. Univ. of Wash. Press. 597 pp. Feck, M.E.  1964.  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 B r i t i s h 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.  A l t i , T.  BIBLIOGRAPHY OF PUBLICATIONS USED FOR IDENTIFICATION OF BRYOPHYTES AND LICHENS 1 9 6 1 . Taxonomic studies on reindeer lichens (Cladonia, sub-genus Cladina). Societas Zoologica Botanica Fennica •Vanamo'i Helsinki. 160 pp.  Conard, H.S. 1 9 5 6 . How to know the mosses and liverworts. Brown Co., Iowa. 226 pp.  Wm.  C.  Dixon, H.M. 1 9 5 4 . The students handbook of British mosses. and Day, Ltd., London. 582 pp.  Sumfield  Frye, T.C. and L. Clark. 1949. nos. 1 - 5 . 1018 pp.  Vol. 6 ,  Hepaticae of North America.  Grout, A.J. 1 9 0 3 . Mosses with the hand lens and microscope. A nontechnical handbook of the more common mosses of the northeastern 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. 809 pp.  The Ronald Press Co., N.Y.  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 G i l i b . Vaccinium ovalifolium Smith Vaccinium ovatum Pursh. Vaccinium parvifolium Sm, Vaccinium ullginosum L. Vaccinium vitis-idaea L. subsp. minus (Lodd.) Hult.  Herbs 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 t r i f o l i a t a (L.) Salisb, Cornus canadensis L. Deschampsia caespitosa (L,) Beauv. Drosera rotundifolia L.  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 A i t .  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 Cladonia b e l l i d l f l o r a (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,  Ahti  97  .APPENDIX II Explanation and Legend for Synthesis tables Synthesis tables I - VI  I  Explanation and Legend f o r 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 f l o r i s t i c evaluation: Species significance  Corresponding cover value  +  solitary, with small dominance  .1  1.  seldom, with small dominance  .5  2.  very scattered, with small dominance  1  3.  scattered, with small dominance  5  4.  often, with l/20 to l/lO dominance  10  5.  often, with l A O to l/5 dominance  20  6.  any number, with l/5 to l/3 dominance  33  7.  any number, with l/3 to l/2 dominance  50  8.  any number, with l/2 to 3 A dominance  75  9.  any number, with dominance more than 3/4, but less than complete  95  10.  +  any number, with complete dominance Sociability  Vigor  individual, sociability none  0 - none  1. up to 4 x 4 cm  +-  2. 25 x 2$ cm  1 - fair  3. 50 x 50 cm  2 - good  2  2  4. 1/3 - 2/3 m  3 - excellent  2  5. 1 - 2 m  2  6. 5 m  2  7. 25 - 50 m  2  8. 100 m  2  9. 200 - 250 m  2  10.  poor  at least 500 m  2  Species are grouped by sublayer, i n decreasing order of presence and total cover degree.  Sublayer abbreviations are as described i n  Methods (Chapter III). Five presence classes are used: V - species which occur i n 81 - 100$ of the plots (constants) IV - species which occur i n 6 l - Q0% of the plots i n - species which occur i n hX - 60% of the plots II - species which occur i n 21 - k0% of the plots I - species which occur i n 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 number of plots i n particular unit  Addendum: In a l l synthesis tables, Agoseris glauca should read Apargidium boreale.  x  1  Q  Q  %  SYNTHESIS  TABLE  I:  CARICETUM PUR I FLORAE  Number of p l o t s P l o t number Plot size  1  2  3  4  5  6  7  8  9  10  11  48  41  39  38  61  52  51  47  46  45  44  0.5  (meters ) 2  0.5  50  Extent of type (meters?) Date  12 22/7/  24/7/ 64  64  0.5 100 22/7/ 64  0.5 50  0.5 35  0.5 15  0.5 10  0.5 80  0.5 20  0.5  0.5  150  25  •  12  13  14  15  16  17  18  19  37  59  53  50  49  43  42  54  0.5 50  0.5 40  0.5  0.5  50  12  0.5 40  0.5 25  0.5 12  0.5 12  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  Percent cover Vegetation:  0  2  0  0  0  0  0  0  0  0  0  0  0  0  8  0  0  0  0  C  25  40  30  20  20  30  45  45  20  35  25  25  25  50  50  35  35  25  65  Oh  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  layer:  B2  Humus decaying eood  Species significance H R  Sub-  Life form  layer  B layer  1  Pn  Vaccinium ullginosum  2  2  Pn  Kalmia p o l i f o l i a  2  2.+.1  Vigor »  Con-  Cover  stancy  value  R  0.1  0-2  1.0  —  I  5  0.03  0-1  1.0  —  I  0.5  C layer 3  H  Carex p l u r i f l o r a  4  H  Sangulsorba  5  H  Agoseris glauca  6  H  Carex obnupta  7  H  Gent!ana sceptrum  6.2.1  8.3.2  microcephala 3.2.2  1.+.1  6.3.2  7.5.2  5.2.2  7.3.2  7.4.2  5.2.1  3.2.1  3.2.2  1.+.1  2.+.1  7.4.2 2.2.1  2.2.1  6.2.1  7.3.2  6.3.2  7.4.3  2.2.1  5.2.1  3.2.1  6.3.2  8.4.2  8.5.2  7.3.2  3.2.1  2.2.1  1.2.1  7.3.2  6.2.2  9.5.3  4.2.1  4.2.1 +.+.2  3.2.2  2.+.1  1.+.1  6.8  5-9  2.0  1-3  V  4936  2.0  0-5  1.1  1-2  392  0.6  0-4  1.2 2.0  1-2  iy III 1  27  1  8  92  0.2  0-3  0.2  0-2  1.0  4.0  0-9  1.8  1-3  IV  3013  1.4  0-7  1.0  0.5-2  IV  334  D layer Sphagnum recurvum  7.2.2  7.3.2  9.4.3  9.5.3  Sphagnum papillosum  1.1.1  7.3.2  2.2.1  1.1.1  Total  B  H  Pn  P r o p o r t i o n by: Species:  Number  i  2.0  5.0  2.0  22.2  55.5  22.2  3347.0 38.0  5455.0  5.5  Total Cover: cover  t  2.1.1  2.1.1  8.4.2  6.2.1  5.2.2  8.3.2  8.4.3  2.1.1  2.1.1  1.+.1  1.1.1  2.1.1  3.2.1  1.+.1  4  species  Life fora  1.1.  61.9  0.06  4  4  4  4  4  1.+.1 1.+.1 3  2.1.1  2.1.1 1.1.1  1.+.1 3  3  3  3  3  2  101  SYNTHESIS TABLE  II.  SCIRPETO-SPHAGNETUM HEHDOCIHI  Number of p l o t s P l o t number P l o t s i z e (meters^) Extent of type (acres) Date  1  2  3  4  5  6  7  8  9  10  99  98  94  97  103  101  93  95  102  100  4  4  4  4  4  4  4  4  4  4  1  1  1  1  1  4/9/  4/9/  4/9/  1/4 3/9/  1  4/9/  1/3 3/9/  1  4/9/  1/4 3/9/  4/9/  4/9/  64  64  64  64  64  64  •64  64  64  64  0  Percent coverage Vegetation:  layer:  8^  0  0  0  0  5  0  0  0  0  82 8 C  5  5  5  15  5  5  5  5  Total  5  5  5  15  10  5  5  5  8 8  55  55  50  50  60  65  40  40  65  3 50  Oh  80  80  60  80  70  75  60  70  75  65  100  100  100  100  100  100  100  100  100  0  0  0  0  0  0  0  0  0  Humus Oecaylng good  3  100 .  0 Species  Life  Sub-  form 1  Pn  B layer  Significance R HI  layer 1  Kalaia poll fol l a  2  3.3.2 2.+.1 2.2.2  1.+.1 2.+.2 1.1.1  4.2.1  4.3.2  2.+.1  2.+.1  1.+.1 4.2.2  2.2.2  1.+.1  2.+.1  1.+.1 2.+.1  2.1.1  1.1.1  Vigor 11  R  0.3  0-3  2  2.2  1-4  1.1  1-2  V  295  2.1.2  1.1.1  1.8  1-3  1.5  1-2  V  130  1.2.1  1.4  0-4  1.4  1-2  IV  185  0.4  0-3  0.8  +-1  0.4  0-1  1  0.3  0-2  0.8  1.0  0-1  1  0.05  0-+  1  —  Ch  Oxycoccus quadrl petal us  2  3  Pn  Vaccinium ullglnosum  2  3.2.2  2.2.2  *  Ch  Empetrum nigrum  2  1.2.1  1.2.1  3.2.1  5  Pn  Ledum groenlandicum  1.+.1  1.+.1  6  Pm  Thuja p l l c a t a  2 2  7  Pn  Chamaecyparis nootkatensis  2  8  Pm  Pinus c o n t o r t a  2  9  Sclrpus caespitosus Agrostis aequlvalvls  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  10  H H  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  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  12  6  Trlentalls arctlca  1.+.1  2.+.1  3.+.1  2.+.1  +.+.1  1.+.2  2.+.2  2.+.2  +-3  T  Gentiana douglaslana  2.+.1  1.+.1  +.+.+  1.+.+  1.5  H  Orosera r o t u n d l f o l l a  1.+.1  1.+.1  1.+.1  1.+.1 1.+.1  2.+.1  14 15  1.+.1 2.1.1  2.+.1 2.+.1  1.8  13  2.+.1 3.+.2  1.+.1  2.+.1  1.+.1  G  Tof 1e1d1 a o c c i d e n t a l ! :  1.+.1 4.2.2  +.+.1 2.1.1  1.+.1 4.3.2  +.+.1  2.+.1  1.+.1 1.2.2  +.+.1  4.2.2  2.+.1 4.2.2  1.+.1 4.2.2  +.+.2 3.2.2  3.2.2  2.2.2  3.2.2  2.2.2  3.2.2  +.2.2  4.2.2  2.2.1  +.+.1  +•.+.+ 2.2.2.  +.+.+ +.+.1  value  1.+.1  2  1.+.1 2.+.1  Cover  3.2.1  2.+.1 3.2.2  +.+.1  Constancy  +.+.+  1.+.1 +.+.1  III  66  II  35  II I  22  I  1  2-3  V  7450  V  900  1.6  1-3  V  345  1.3  1-2  V  121  1-3  1.0  +-2  V  106  1.1  0-2  1.0  V  1.1 2.7  +-2  •-?  V  55 48  0-4  1.1 1.5  1-2  IV  665  2.0  0-4  1.9  1-2  IV  285  0.7  0-2  1.5  1-2  1.0  0-4  1.5  1-2  III II  165  0.6  0-3 0-3  1.3  1-2  1.5  1-3  0.2 0.05  0-2  +-2  5  C layer  16  H  Rhynchospora alba  17  H  Carex canescens  18  H  Carex obnupta  19  H  Carex p i u r l f l o r a  2.2.2  20  H  Gentiana sceptrum  21  H  Agoserls glauca  22  H  Coptls a s p l e n l f o l l a  1.+.1  2.2.2 2.+.1  1.+.1  4.2.2  3.2.2  3.2.2  2.+.1 1.+.2  +.+.1 3.2.1  0.4  2.1.2  32  II I  61  2.0  I  10  0-+  1.0  1  5  55  23  H  Coptls t r i f o l l a t a  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 B  Sphagnum mendocinum Sphagnum capillaceum  h  3.2.2  4.2.2  +.+,+  4.2.2  3.2.2  8.5.2  4.2.2  3.5  *-8  1.8  *-2  V  3.1.2  3.2.2  2.1.1  3.2.2  2.2.1  1.1.1 2.1.1  4.3.2  h  3.2.2 3.2.2  2.1.1  2.1.1  2.1.2  2.4  2-3  1.5  1-2  1306 260  L  Cladonla r a n g l f e r l n a  h  0-2  h  0.2  0-1  0.9 2.0  +-1  Rhacomitrium lanuglnosum  1.1.1 1.2.2  0.6  B  2.1.1 1.1.2  0.2 0.2  0-2  1.0  II  10  0-1  2.0  II  10  0.1  0-1  1.0  11  0.6 0.05  0-5  1.3  +-2  1  5 205  1.0  —  1  +.+.1  —  D laver  26 27 28  di 29  B  Cephalozla blcuspldata  30  L  Cladonla p a d f 1 c a  h  31  B  Bazzanla amblgua  d>  1.1.1  1.1.1  2.2.1  h  1.1.2  1.1.2  d>  Total  species  1.+.1 5.3.2  1.1.+ +.+.1  22  L i f e form  H  21  B  21  20  18  18  18  17  16  Pn  L  Ch  G  Pm  15  •  T  Proportion by: Species:  Number  12.0  6.0  4.0  2.0  2.0  2.0  2.0  2  38.8  19.4  13.0  6.5  6.5  6.5  6.5  3.2  6026.0  8107.0  275.0  235.0  196.0  169.0  23.0  106.0  63.8  32.2  1.1  0.9  0.8  0.7  0.1  0.4  Total Cover:  cover  t  1.0  O-t.  v  III II  30 10  1  SYNTHESIS TABLE III.  OJYCOCCErO-SPHAGNETUM PflPILLOSAE  Number of plots  1  Plot number Plot size (meters?) Extent of type (ecres) Date  55 4  Total  Oh "d. 0  •  Humus Decaying Wood  Life form 1 2 3 4 5 6 7 S  Ch Pn Pn Pn Pn Ch P> Pm  3  4  5  6  7  19 4 2 8/6/ 64  18 4 1 8/6/ 64  17 4 1 3/6/ 64  58 4 4 28/7/ 64  57 4 1/100 28/7/ 64  64 4 4 31/7/ 64  3 55 98 0 98  2 45 98 0 98  5 35 95 2 97  5 40 98 0 98  8 50 98 0 98  15 45 98 0 98  7 60 100 0 100  100 0  100 0  98 0  100 0  100 0  100 0  100 0  26/7/ 64  50 4 4 30/7/ 64  66 4 4 31/7/ 64  15 30 98 0 9B  15 50 9B 0 98  5 60 95 0 95  100 0  100 0  100 0  *  Percent coverage Vegetetion: layer: Bs C  2 56 4 4 28/7/ 64  8  9  10  11  12  13  20 4 3 8/6/ 64  63 4 4 31/7/ 64  67 4 4 31/7/ 64  65 4 4 31/7/ 64  5 35 100 0 100  5 40 100 0 100  8 60 99 0 99  2 45 98 0 98  2 50 100 0 100  100 0  100 0  100 0  100 0  100 0  25 4 1 12/6/ 64  14  15  Species Significance  Sublayer  6 laver Oxycoccus quadripetalus Kalmia polifolia Ledum groenlandicum Vaccinium ulIglnosum Myrica gale Empetrum nigrum Pinus contorta Thuja plicata  2 2 2 2 2 2 2 2  3.2.2 3.2.1  3.2.2 3.3.1 3.2.1  2.2.2 2.+.1 3.2.2  2.1.2  2.1.1 2.2.1  1.+.1  2.1.1 2.2.2 l.t.t  3.2.2  2.2.1 1.2.1 1.*.1 2.2.2  3.2.2 3.2.1  3.1.2 3.1.1 3.2.1  3.1.2 2.*.2  3.2.2 3.+.1  4.2.2 2.2.1  3.2.1  3.1.2 1.*.1  2.1.2 1.*.+  3.2.2 1.+.1  3.2.1  + + +  +.+.«•  Constancy  Vigor  •  II  a  R  2.8 2.0 0.8 0.3 0.4 0.2 0.03 0.05  2-4 1-3 0-3 0-3 0-3 0-1 0-* O-t  1.8 1.0 1.1 2.0 1.0 + + +  1-2 <-2 +-2  4.0 3.0 2.7 2.5 1.3 2.6 1.5 3.3 1.5 1.1 0.07  0-7 0-5 1-5 1-3 0-3 0-7 0-4 0-9 0-5 0-5 0-1  2.1 2.0 1.7 1.9 1.4 1.7 1.6 1.8 1.5 2.0 2.0  2-3 1-3 1-2 1-2 1-2 1-2 1-3 1-2 «-2  9.0 1.5 0.5 0.3 0.3 0.13 0.1  9 0-4 0-2 0-5 0-3 0-2 0-1  2.9 1.7 1.4 3.0 2.0 2.0 2.0  2-3 1-2 1-2  V V  III  1 1 1  — — —  I  1  —  Cover value  370 217 111 40 40 7 0. 0.  C laver 9 10 11 12 13 14 15 16 17 18 19  20 21 22 23 2t 25 26  H H H G H T H H H H  Agoserls glauca Agrostis aequlvalvls Trlentalls arctlce Orosera rotundifolia Tofleldle occidentals Sangulsorba mlcrocephala Gentiana douglaslana Carex pluriflora Carex obnupta Rhynchospora alba Scirpus caespitosus  B B B B B S B  Sphagnum papillosum Sphagnum capillaceum Sphagnum recurvum Sphagnum tenel lum Sphagnum fuscum Cephalozia blcuspidata Polytrichum comnune  s  3.2.2 4.2.2 3.1.2 2.1.2 1.*.1 4.2.2 4.2.2 3.2.2 4.3.2 1.2.2  h h h h h  Proportion by Species:  Number  1  Total cover:  cover %  9.5.3 1.1.1 1.1.1  4.2.2 4.2.2 4.*.2 3.+.2 3.1-.2 •.+.1 1.+.1 4.3.2  4.2.2 4.2.2 3.».2 3.2.2  7.3.2 1.2.2 2.*.2 3.2.2  3.2.2 3.+.2 6.3.2 3.+.1  2.-0 3.1.2 5.2.2 3.2.1  7.2.2  6.3.2  1.+.1 2.-..1  2.1.2 3.1.2 1.+.1 3.2.2 4.1.2 5.3.2  2.1.1 3...1 6.3.1  !.*.*  9.5.2 1.2.2  9.7.3 2.2.2 2.3.2  9.7.3 1.1.2  9.7.3 1.1.1  5.2.2 4.2.2 1.+.1 3.1.2 1.*.2  4.2.3 5.2.3 Z.+.2 2.1.2 3.2.2  5.3.2 3.2.2 3.1.2 2.1.2 2.->.1 2.1.1  2.*.1 3.+.1 2.1.2  3.1.2 9.3.2  9.5.3 2.2.2 1.1.1  2.1.2 4.2.23.1.2 3.1.2  7.3.2 4.2.2 3.+.2 1.-».2  3.*.2 7.2.2 4.2.2  1.+.1 5.2.2 4.2.2  5.2.2  9.5.3 2.2.2  3.2.2 3.+.1 3.t.2 1.*.1 4.2.2  4.2.2 4.2.2 2.+.2 3.1.2 1.1.1 3.2.2  3.1.3  7.3.2  3.2.2  3.2.2  9.5.3 3.2.2  9.4.3 1.1.2  9.6.3 4.2.2  9.7.3 4.1.1 2.2.2  9.5.3  9.5.2  9.5.3  5.3.3 3.3.2  2.2.2 2.1.2  h  h  Total species  Life form  9.6.3 1.1.1 1.1.1  3.2.2 4.2.2 5.1.2 2.*. 2 •1.+.2 4.2.2 1.+.1 4.3.1 2.2.2 3.3.2  1.2.2 18  17  15  14  14  14  14  13  13  12  H  B  8 30.8  7 26.9  4 15.4  7.7  2 7.7  2 7.7  1 3.8  6070.3 34.4  9913.3 56.3  406 2.3  377 2.1  600 3.4  1.4  247 1.4  Pn  Ch  6  Pm  T  12  12  11  9  9  — —  — — — —  V y y y v  IV IV in in  II i  y IV  II I i i  1  1827 743 467 310 133 707 247 1B70 310 300 3.  9500 207 23 133 40 7 3.  103  SYNTHESIS TABLE IV.  LEDETO-SPHAGNETUM CAPILLACEI  Number of p l o t s P l o t number Plot size  (meters^)  Extent of type  (acres)  1  2  3  4  5  6  7  8  9  10  85  69  68  91  90  86  84  70  87  26  4  4  4  4  4  4  4  4  4  4  1/5 4/8/  1/10  1/20  1/100  1/20  1/10  1/20  4/8/  28/8/  28/8/  27/8/  1/5 27/8/  4/8/  27/8/  64  64  64  64  64  1/2 27/8/  Date  64 Percent coverage Vegetation: l a y e r :  64  64  64  S-|  15  0  0  15  0  0  15  0  0  0  B  20  25  30  10  15  20  35  20  20  15  B  30  25  30  20  15  20  40  20  20  15  C  65  70  65  40  60  70  50  95  95  100  100  98  100  100  5  96 0  40 100  80  Oh Od. D  65 98 0  2  0  0  0  0  0  0  100  96  98  98  100  100  100  98  100  100  95  100  100  98  100  100  100  100  100  100  5  0  0  2  0  0  0  0  0  0  Total  Total  64  1 12/6/  2  Humus Decaying Wood  Species Life  Sub-  form 1  Pn  B layer  Significance  layer  Ledum groenlandicum  1  5.2.2  2  5.2.2  5.2.2  4.3.2  11  R 1-2  value  2.0  0-5  1.7  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  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  3.2.2  3.2.2  2.2.1  3.0  2-4  1.9  1-2  V  510  1.6 1.0  0-3  1-2  IV  220  0-2  1.1 1.2  1-2  111  130  0.3  0-3  1  2  Pn  Kalmia p o l l f o l l a  2  3.+.1  3.2.2  3.2.1  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  4  Ch  LInnaea boreal i s  2  3.2.1  3.1.2  2.2.1  3.2.1  1.2.1  3.2.1  5  Ch  Empetrum nigrum  2  2.2.1  1.1.1 4.3.2  6  Pn  Chamaecyparis nootkatensis 1 2  3.+.1  7  Pn  Myrica gale  1.2.1  1.1.1  4.4.1  0.5  0-4  1  —  ll  151  0.4  0-2  1  —  I  40  8.4.3 6.2.2  7.4  6-9  2.2  2-3  V  6110  3.8  0-6  2.0  —  V  1180  2.0 1.8  1.6 1.7  1-2 1-2  V V  205 144  2.1  1-3 1-3 0-4  2.0  —  IV  335  0.7  0-3  1-2  0.4  0-2  1.3 2.0  0.3  0-1  1.0  0.6  0-4  0.6 0.4 0.3  4.3.1 2.2.1  2  R  Cover  3.4.1  4.2.1 3.4.1  2.2.1  M  Constancy  3.4.1  5.2.1 5.3.2  Vigor  2.4.1  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  9  Agoserls glauca Drosera r o t u n d ! f o l i a  4.2.2  4.2.2  5.2.2  3.+.2  Trientails arctlca  2.t.2  2.4.2  4.3.2 3.4.2 3.4.2  2.4.1  G H  2.4.2 1.4.1  4.3.2 2.4.2 1.4.2  4.2.2 1.4.1  11 12  3.1.2 1.4.2  4.2.2 1.4.1 2.4.2  3.2.2  10  H H  3.4.1  2.1.2 1.4.1  Agrostis  3.2.2  1.2.2  2.4.2  4.3.2  3.2.2  2.4.2  4.2.2  2.2.2  13  Ch  Cornus canadensis  3.2.2  3.1.1  14  T  Gentiana douglasiana  15  G  T o f i e l d l a occidental 1s  16  G  17  H  Sanguisorba microcephal a Carex p l u r i f l o r a  18 19  H H  Deschampsia  20  G  Maianthemum d i l a t a t u m  21  H  Coptis a s p l e n i f o l l a  aequlvalvls  Scirpus  1.4.1  2.4.2  8.5.2  1.4.1 1.4.2  2.4.2  1.4.2 1.4.1  1.+.1  1.4.1 2.2.2  3.2.1  4.2.2  3.2.1  caespltosus  1.4.1  caespltosa  3.2.2  3.2.2 1.+.1  4.4.4  1.4.1  —  III  20 20  II II  15  2.0  —  1  110  0-3  1.0  —  1  100  0-3 0-3  1.5  1-2  1  55  2.0  —  50  0.2  0-1  0.8  +-1  1 1  0.1  0-1  1.0  —  1  1  5.9 4.8  3-9 1-7  1.9 2.2  1-3  V V  4010 2515  3.4  0-8 0-9  2.1 2.6  2-3  3.5  2-3  III  2800  2.3 0.7  1.1 2.3  2-3  III II  1150  0-4  0.2  0-2  1.0  0.8  0-3  2.0  —  II  75  0.3  0-3  0.5  0-2  2.0 1.6  1-2  II  0.9  0-5  0.6 0.1 0.1  0-4 0-1 0-1  0.1  0-1  6  0 layer 8 B  Sphagnum papillosum  h  3.1.1  4.2.2  8.3.2  h  3.7.2  1.1.1  7.4.3  8.4.2 5.2.2  6.2.2  Sphagnum capillaceum  5.2.1 3.2.2  3.1.2  23  7.3.3  7.3.3  24  8  Sphagnum mendocinum  h  2.2.2  3.2.2  8.4.3  6.2.2  6.2.2  3.2.2  25  e  Sphagnum fuscum  h  9.4.3  7.3.2  8.3.3  26 27  B  Sphagnum tenel 1 um  h  5.2.2  3.2.2  B  Dicranum bergeri  h  1.2.2  d.  22  28 29  8 B  Pleurozium schreberi  h  2.1.1 2.2.2  d>  3.1.2 2.+.2  Aulacomnium p a l u s t r e  h  30  B  Polytrlchum commune  h  31  B  Sphagnum recurvum  h  32 33  B B  Bazzania ambigua Dicranum scoparium  di da  34  B  R i c c a r d l a sp.  di  1.1.2  species  23  Total  B  L i f e form  3.2.2  3.2.2  4.2.2  3.2.2  8.5.3  3.2.2  1.2.2 1.1.1  5.3.2 2.2.2  4.2.2  1.1.1 1.1.1  21  H  20  19  Pn  > 18  15  16  15  14  Ch  G  T  4  3  1  8.8  2.9  P r o p o r t i o n by Species:  Number  13  %  38.2  Total cover:  cover %  12795 52.9  6.3.2 6.3.2  4.2.3  2.4.2 4.3.2  9.5.3 6.2.2  4.2.2  7.3.3  2.2.2 2.1.2  7.3.2 3.3.2  9  4  26.5  11.8  8146 33.6  2195 9.1  11. 3 880 3.6  165 0.7  20 0.1  ,  12  0-8  1-3  2-3  IV  1620  115  25  2.0 2.0  —  1  300  —  110  1.0 1.0  — —  1 1 1  —  1  5  2.0  5 5  SYNTHESIS TABLE V.  1 2 3 4  PINETO-SPHAGNETUM CAPILLACEI  Life form  A layer  Pm  Pinus contorta  Pm Pm Pm  Thuja plicata Chamaecyparis nootkatensis Tsuga heterophylla  Sublayer 2 3 3 3 3  Presence  IV I I I  Life form  Cover value  2005 167 32 1  D layer  5  Pn  Pinus contorta Ledum groenlandicum  6  Ch  Empetrum nigrum  7  Pn  Kalmia polifolia  1  V  1  V  V  1500  1  V  1073  8 9  Ch Ch  Oxycoccus quadripetal us Vaccinium vitis-idaea  2 2  V IV  314 176  10 11  Ch Pn  Linnaea borealls Gaultheria shallon  2 1  IV  135  h  V  2718  Sphagnum capillaceum  V  1634  49 50 51  B  Sphagnum recurvum Sphagnum mendocinum  h da h  IV  1257  52  B  Dicranum scoparium  53  B  Rhytidladelphus loreus  54 55  B B  Aulacomnium palustre Sphagnum fuscum  56 57  L L  12  III  297  B B  h h dw h dm h  III III  882 271  III  115  III III II  98 56 821  Cladonia crispata  dw h h dt h  II  240  Cladonia pacifica  h  II  157  Dicranum bergeri  di h  II  119  Cladonia rangiferina  dw h  II  111  II  107  II  13  Pleurozium schreberi  B L  dw h dw h dw  Pn  Chamaecyparis nootkatensis  1  II  1128  60  B  Hylocomium splendens  Pm  Thuja plicata  1  II  247  61  B  Bazzania ambigua  Pn Pm  Vaccinium uliginosum Tsuga heterophylla  2 1 2  II  177  62  B  Rhacomitrium lanuginosa  II  8  63 64 65  B B B  Herberta adunca Sphagnum tenellum Eurhynchium oreganum  66  B  Polytrichum strictum  67 68  B B  Isothecium stoloniferum Cephalozla bicuspidata  69 70 71  B B B  Dicranum fuscescens  72  B  Mnium glabrescens  h da h  73 74 75  L L B  Cladonia bellldiflora .  da da  76 77 78 79 80  L B B B B  Cladonia unci al Is Diplophyllum albicans lemadophila ericetorum Riccardla sp. Eurhynchium stokesil Plagiochila asplenioides Bantramia pomiformis  da da da da dw da da  13  Pn  Myrica gale  1  I  1146  14  Pn  Vaccinium parvifolium  1  I  22  15 16 17  Pn Pm Pn  Vaccinium ovatum Picea sitchensis Vaccinium oval ifol 1um  18  Pn  Iflenziesia ferruginea  1 2 '1  I I I  1  I  15 2 2 0.2  C layer 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 36 39 40 41 42 43 44 45 46  58 59  2  H H G H Ch H G H H H III T G G ti H H H H G H H Ch G H G G G  Carex obnupta Agoserls glauca Trientalls arctica Drosera rotundlfolia Cornus canadensis Sanguisorba microcephala Maianthemum dilatatum Carex pluriflora Agrostis aequivalvls Blechnum splcant Coptls asplenifolia Gentiana douglasiana Lysichitum americanum Tofieldla occidentals Carex canescens Scirpus caespitosus Coptls trifoliata Gentiana sceptrum Deschampsia caespltosa Nephrophyllldium crista-galU Calamagrostis nutkaensls Carex pauciflora Lycopodium clavatum Habenaria saccata Rhynchospora alba Boschniakia hookari Pteridium aquillnum Veratrum virlda  V V V V IV IV IV III III III III II II II I I I I I I I I I I I I I I  2392 1005 156 63 240 215 60 290 231 175 78 50 31 13 144 56 34 32 21 14 5 5 5 3 2 1 1 0.2  Cover value  Sphagnum papillosum  1551  2  ence  B B  2166  1  Pres-  47 48  B layer Pm  Sublayer  Polytrichum commune Plagiothecium undulatum  h dw dw h h dw h dw dw h dw dw h  1  180  1 1  55 39  1 1  33 28  1  18 15  1  1  15 14  1  8  1 1 1  7 6 5 3  1  1 1 1 1 1 1  3 3 1 1 1  E layer Bryophytes S lichens 81  E  Frullania nlsquallensis  A  82  E  Scapania bolanderl  cB  E  Dicranum scoparium  C A<  E  Herberta adunca  cA  D B  V  635  IV  150  IV  138  D  II  61  B  II  48  D D •  E  Isothecium stoloniferum  C A  C  Life form 83  84  85 86  87 88  E layer (continued)  E  Antitrlchia curtipendula  E  Bazzanla ambi gua  E  Olplophyllun albicans  E  Pleuroziua schreberl  E  Rhacoraitrlum lanuginosum  E  Parmella physodes  E E E E E E E E E E E  Pres-  Cover  Life  Sub-  Pres-  Cover  ence  value  form  layer  ence  value  A  E  Plagiochila asplenioides  C  I  2  B  E  Plagiothecium undulatum  C  I  2  E E E E E  Radula bolanderi Bazzanla nudicaults Blepharstoma trichophyllum Mylia taylori Sphenolobus ninutus  C C C C C  I I I I I  2 1 1 1 1  C B C  I  302  I  60  C A B C  I  31  I  15  I I  2 2  C 6 C e  Usnea plicata Cephalozia b<cuspidate Macrodi plophy]T um pli catura Cladonia bell Id!flora Olcranum fuscescens Sphaerophorus globosus Hypnum circinale  II  40  II  19  C A C  II  16 I  44  C A B C A C  I  17  E  Empetrum nigrum  I  14  E  Gaultheria shallon  I I  14 13  E  Vaccinium vitis-idaea  I I I I I  11 7 6 6 5  E  Polypodlum glycyrrhiza  E  Cornus canadensis  E E  Linnaea boreal Is Maianthemum dilatatum  E  Tsuga heterophylla  B C C B C B  E  Vaccinium parvifolium  B  C C C B B C C C C C  Hylocomium splendens Riccardia sp. Bartramia poniformis Mnfum glabrescens  Life form  Sublayer  I I I I  89 90 91 92 93  Vascular plants  94  3 3 3 2  E  B  H  Pn  38 32.0  28 23.5  16 13.4  10 8  1678 6.5  8521 32.8  4748 18.3  5411 20  Ch  1  Pm  Proportion by Species:  Number  I  Total cover:  cover  i  9 7.6 279 1.1  6 5.04 522 2.0  6 5.04 2370 9.1  5 4.2  1 0.8  2455 9.5  50 0.2  I  1 0.2  I  0.2  106  SYNTHESIS TABLE V.  PINETO-SPHAGNETUM CAPILLACEI Variant:  P.-S, c a p i l l a c e i sphagnoosun p a p l l l o s l  Number of plots  1  Plot number Plot size (outers ) Extent of type (meters ) Date  33 4 5 13/7/ 64  2  2  Percent coverage Vegetation: l a y e r :  A2 3 A A  Total  Total  B1 B2 8 C  % Total  0°d.  k Humus Decaying Wood  1  Life form  A laver  Pm  Pinus contorta  2 34 4 5 13/7/ 64  10 0 10 65 20 70 30 80 5  0 0 0 40 20 50 35 85 4  85 10 30  89 0 10  65  60  90  95  10  5  3 74 4 16 21/8/ 64  4 82  5  4 12 26/8/ 64  79 4 4 26/8/ 64  0 20 20 65 20 70 25 78 8 85 15  0 15 15 65 25 75 30 75 15 90 10  0 45 45 25 35 55 40 75 10 85 25  15 40  20 65  25 40  90 10  85 15  90 10  6  7  6  9  80  75  81  77  4 4 26/8/ 64  4 4 21/8/ 64  4 4 26/8/ 64  0 20 20 75 15 80 30  0 25 25 20 15 35 40 85 0  70 10 80 20 20  85 25 25  65  30  85 15  100 0  4 8 22/8/ 64  10 76 4 •4 22/8/ 64  11 78 4 8 22/8/ 64  0 0 0 60 15 70 35 85 3 68 0  0 65 65 10 10 20 65 75 0 75 25  0 85 85 40 20 50 50  0 40 40 20 25 40 35  85 2 87 35  85 5 90  15 60  25 50  35 60  95 5  100 0  95 5  12 73 4 6 21/8/ 64  0 30 30 10 15 20 35 75 0 75 40  40 35 45  25 40  95 5  100 0 Species S i g n l f l canoe R III  Sublayer 2 3  1.+.2  1  8.5.2  7.5.1  5.+.1  7.5.1  4.5.1  6.5.1  8.5.2 2.+.2 3.3.2 3.+.2 3.2.2  8.5.1  6.5.1  8.5.1  6.5.1  1.+.1 1.+.1 3.2.2 1.+.2  8.6.2  9.6.1  7.5.1  ?.+.2  4.4.1  7.6.1  6.4.1  1.+.2  3.2.2 3.2.1 4.2.3  3.2.2 3.2.1 3.2.2  4.3.2 4.2.2 4.2.2  4.3.2 4.2.2  3.2.1 3.2.2  1.+.1 1.2.1  3.2.1  3.2.1  3.2.1  3.2.2  Vigor M R  0.08 5.0  0-1 0-9  2.0 1.2  6.4 0.3 2.4 3.4 3.0 0.5 1.7 1.7 0.3 2.3 0.08 0.33 0.5  1-8 0-2 0-4 3-4 1-4 0-2 0-3 0-3 0-2  1.6 1.5 1.5 1.4  0-4 0-1  1.1 2 1 1 1 1 1 2  Constancy  Cover val ue  1-2  V  3612  1-2 1-2 1-2 1-2 1-3 1-2 1-2 1-2  V  4729  V V  708 554  V V  250 170  B layer Pi 2  Pn  Pinus contorta Ledum groenlandicum  1  3 4  Ch Pn  Oxycoccus quadripetalus Kalmia p o l i f o l i a  5 6  Ch Ch  Llnnaea boreal 1s Empetrum nigrum  1  7  Pn  Myrica gale  1 1  6  P«  Thuja p l i c a t a  9  Pn  Chamaecyparis  10 11  Ch  Vaccinium v i t i s - i d a e a Tsuga heterophylla  Pm  1  nootkatensis  6.5.2  4.2.2 3.2.2 3.2.2 2.2.1  3.2.1 3.+.1 3.2.2 2.*.1  3.2.2 3.3.2 3.2.2  2.2.2  2.+.1 2.1.1  3.2.2  2 2 2  1.+.1 2.2.1 4.2.1 3.2.2  2.3.1 3.1.1 3.1.2  2.1.1 3.2.2  2.2.1 4.2.1 1.+.2 2.2.2 3.2.2 2.2.1  1.+.1 2.2.2  1.+.1 2.2.2  4.3.1  3.2.1  3.2.1  3.2.1 3.2.1 2.1.1  1.+.1 1.+.1  2.+.1  1  6.5.1  3.3.1  4.2.1 '3.1.1 3.2.1  3.2.1  1.+.1  3.2.1 1.+.1  0.2 0.5  3.3.1  0.3 0.3 0.04  3.+.1 3.2.2  +.+.+  0-3 0-2 0-2 0-3 0-3 0-3 0-+  1.9 1.3 1.1 1.6 2  +  —  1-2  IV  416  —  II  —  II  50 25  —  —  —  1  83  1  41 0.  1  C laver 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28  H ' Carex obnupta H Agoseris glauca H Agrostis aequlvalvis G Trientails arctica H Orosera r o t u n d l f o l i a H Sangulsorba microcephala Ch Cornus canadensis G Lysichitum americanum H Carex pluri f l o r a H Scirpus caespitosus G T o f i e l d l a occidental 1s T Gentiana douglasiana H Coptis a s p l e n l f o l i a H Carex pauciflora G Maianthemum dilatatum H Deschampsla caespitosa* Ch Lycopodiuffl clavatum  4.3.2 3.2.2 2.+.2  5.3.2 4.2.2 3.2.2 2.+.2  2.2.1  1.+.2  2.*.2 1.2.2  2.1.2 4.2.3 4.+.2  2.+.1  1.+.1 3.2.2  3.2.1  2.2.2 1.*.2  1.+.1 2.4.1  6.3.1 3.2.1 3.2.2 3.*.2 2.+.1 2.1.1 1.+.I 6.3.2  6.3.2 5.2.2 2.2.2 1.+.2 1.+.1 2.+.1 1.*.1  6.3.2 4.2.2 3.2.2 3.*.2 1.+.2 I.+.+ 1.3.1  3.2.2 6.3.2 2.2.2 1.+.2 2.2.2 1.2.1 2.+.1  4.3.2 1.+.1 1.+.2  1.+.1 1.+.1  6.3.2 4.2.2 3.2.2 3.+.2 2.+.1 2.+.1  8.4.2 3.2.2 1.+.2 2.1.2 4.2.2  3.2.1 3.2.2  3.2.2 1.+.1 2.-0  2.2.2 1.+.+  +.+.+ 1.+.3  +.+.1  7.4.2 3.2.1 2.2.2 1.*.2 2.+. 2 3.2.2  2.2.1 3.2.3 2.1.1 3.2.1  1.2.2 1.+.1  1.1.2  2.*.2  8.5.2 3.2.2 5.3.2 2.+.2 2.1.2  5.1 3.3 2.3 1.7 1.8 1.7 1.0 0.5 1.3 0.9 0.3 0.3 0.3 0.2 0.12 0.08 0.04  0-8 0-6 0-5 0-3 1-2 0-4 0-4 0-2 0-6 0-4 0-1 1-1 0-2 0-2 0-1 0-1 0-*  1.8 1.7 2.1 2.0 1.7 1.6 1.1 1.0 1.5 2.0 1.0 1.7 1.5 2.0  + 3 1  —  V V V V V IV III III II II II II 1 1 1 1  —  1  2-3 1-2 2-3 —  1-2 1-3 *-2 —  1-2 — —  1-2 1-2 — —  3066 908 408 170 87 275 116 25 400 170 16 12 16 8 5 4 0.  107  Species Sub-  Life form 29' 30 31 32 33  B  D laver  layer  e e  Sphagnum papillosum Sphagnum recurvum Sphagnum capillaceum  B B  Sphagnum mendocinum Sphagnum fuscum  h h h da h h di h da h  34  B  Pleurozium schreberi  35 36 37  S B B  Aulacomnlum palustre Bazzania ambigua Polytrichum commune  38 39 40 41  B  Cephalozla bicuspidata Rhytidiadelphus loreus Scapania bolanderl Dipiophyllum albicans Dicranum scoparium Polytrichum structura Rhacomitrium lanuginosum  42 43 44  B B B B B B  Significance M R 5.2.2 3.2.2 3.2.2  8.4.3 2.1.1  6.4.2  8.4.3  2.2.3  5.4.3 4.3.3 3.2.2 2.1.2 4.3.2 5.3.3 2.1.2  2.1.1  2.2.2  4.2.2 6.3.2  4.3.2  5.3.3 6.3.3 1.1.1 1.1.1 1.1.1  2.1.2 4.2.2  2.1.2  d« h da da da da da h da  6.3.2 5.3.2 4.2.2  7.3.3 7.3.2 3.2.1 3.2.1 4.2.2  7.3.2 5.3.3 4.2.2  6.3.3 3.2.2 2.1.1 2.2.1 3.2.2  5.3.3  2.2.3 1.2.1  4.2.2  7.3.2 7.3.3 5.2.2  6.3.2 7.3.3 4.2.2  4.2.2  3.2.2  7.3.2 5.3.2 7.3.3 3.1.2 3.2.1  8.3.2 5.2.2 1.2.2 7.2.3  1.2.2 1.1.1  2.2.2 1.1.2 2.2.2  2.1.2  •1.+.+  1.1.2  2.1.2 4.2.2  3.2.2  8.3.2 4.2.1 6.3.2 3.2.2 3.2.1  1.1.1 3.2.1  1.2.3  2.1.2 2.2.2 2.1.1  2.2.2  ConVigor  tt  6.5 4.4 3.8 0.9 2.9 3.6 0.3 0.8 0.6 1.1 0.3 0.5 0.3 0.6 0.3 0.3 0.2 0.2 0.2  4-8 0-7 0-7 0-3 0-5 0-8 0-3 0-2 0-4 0-3 0-2 0-2 0-2 0-4 0-3 0-2 0-2 0-2 0-2  2.7 2.3 1.9 1.8 1.9 2.8 2.0 1.8 1.3 1.7 2.0 1.4 1.7 2.0 1.0 2.5 2.0 1.0 2.0  2.8 3.3 3.6 0.4 2.3 0.08 0.8 1.6 0.08 0.5 0.7 0.2 0.7 0.9 0.08 0.7 0.7 0.3 0.3 0.2 0.3 0.2 0.2 0.08 0.08 0.08 0.08 0.08 0.08 0.08  0-6 1-5 2-5 0-3 1-3 0-1 0-4 0-4 0-1 0-3 0-3 0-1 0-3 0-2 0-1 0-2 0-2 0-3 0-1 0-2 0-2 0-1 0-1 0-1 0-1 0-1 0-1 0-1 0-1 0-1  1.6 1.7 2.0 2.0 1.6 +  0.3  0-3  Cover  stancy value R  2-3 1-3 1-3 1-2 1-3 2-3  V V V  4508 2233 1408  III  712 2200  III  112  —  1-3 1-2 1-2 1-3 +-2 1-2 —  2-3  III II II II I I 1 1 1 1  —  83 54 25 16 125 41 12 8 8 8  E laver Bryophytes & lichens 45  46  47  E  F r u l l a n i a nlsquallensis  A B C B  E  Scapania bolanderl  E  Dicranum scoparium  E  Isothecium stoloniferum  C A B C A  E  A n t i t r l c h i a curtlpendula  B C B  E E  Dipiophyllum albicans Bazzania ambigua  E E  Cephalozla bicuspidata Pleurozium schreberi  E E  Bartramia pomiformis Hypnum c i r c i n a l e  C  48 49  c  1.1.1 4.2.2 4.2.2  3.1.2 3.1.2  3.1.2 3.1.2 4.2.2  3.1.1 3.1.1 3.2.2  3.2.1 3.2.1 3.2.2  3.1.1 3.1.1 4.2.2 3.2.2  4.1.2 4.1.2 4.2.2 2.2.2  1.1.1 2.1.2  4.2.2 3.2.2 4.2.2  4.2.2 4.2.2  3.2.2  3.2.2  3.2.2  3.2.1  1.1.1  2.2.1  3.2.2  2.2.2  2.2.1  3.2.2 1.1.+ 1.1.1 3.2.2  2.2.1  1.+.1 2.1.1  1.1.2  1.1.1 3.2.2 2.2.2  c c c  1.1.1 1.1.2 1.1.2  50 51 52  E E E  Riccardla sp. Sphaeijophorus globosus Herberta adunca  53 54 55 56 57  E E E E E  Cladonla b e l l l d i f l o r a Mylla t a y l o r l Bni un glabrescens Sphenolobus mlnuius Usnea p l i c a t a  C C C C C A  4.2.3 4.2.3  2.1.1 2.2.2  2.1.1  3.2.2 3.2.2 3.2.2  6.1.2 5.1.2 4.1.2  1.1.1  2.2.2  1.1.1  1.1.1 1.1.1  2.2.2  B  C 8 C C B A  1.2.t  3.2.2  5.2.2  2.1.2 1.1.1  2.1.2 2.1.2  3.1.1 3.2.1  2.2.2 2.3.2 1.-1.1 1.1.1 2.2.2  2.2.2 1.1.1 1.1.2  2.1.2 2.1.1 1.1.1  2.2.1 1.1.1 1.1.1  1.1.1 1.1.1  1.1.1  2.1.2 2.1.2  1.+.Z  3.1.1 2.2.2  2.1.2 1.1.1  1.1.2  1.+.1  1.1.1 1.1.1  1.1.1 1.1.2 1.1.1 1.1.1 1.1.1 3.1.1  1.4 1.7 1.0 1.3 1.3 1.0 1.4 1.6 1.0 1.8 1.8 1.0 1.0 2.0 2.0 1.5 'l.O 1.0 1.0 2.0 1.0 1.0 1.0 1.0  1-2 1-2  1-2  V  1033  V  291  V  212  —  1-3 1-3 —  1-2 1-Z  III  '66  III III  62 45  III III 1  37 33 45 41  1 1 1 1  16 6 6 8  —  1 1 1 1  4 4 4 4  —  1  4  —  1  41  —  1-2 1-2 —  1-2 1-2 —  1-2  — —  1  Vascular plants 58  E  Polypodium glycyrrhiza  C  Total species  L i f e form Proportion by Species:  Number  i Total cover:  cover  2  3.2.2 32  37  E  8  H  20 31.8  16 .25.4  10 15.9  1966 7.5  11553 44.2  5342 20.5  31  Ch  5 7.9 1182 4.5  30  30  Pn  4 6.3 1091 4.2  28  6  28  24  Pm  4 6.3  3 4.8  216 0.8  4755 18.2  24  T  1 1.6 12 0.04  24  20  19  2  108  SYNTHESIS TABLE V.  PINETO-SPHAGNETUM CAPILLAC Variant:  P.-S.  c a p i l l a c e i nyricosura  Number of p l o t s  2  3  4  5  110  15  111  106  112  25  25  25  25  25  1  P l o t number P l o t s i z e (meters ) 2  Extent of type (acres)  galls  1  Date  9/9/ 64  27/5/  9/9/  9/9/  64  64  64  64  10/9/  Percent coverage Vegetation:  layer:  A3  20  3  15  15  10  Bl  70  90  60  15  70  B  15  20  15  60  15  B C  75  92  85  15  75  40  40  70  Oh  75  45 55  80  50  55. 85  2  Total  Total  10  20  10  80  °d. D  85  75  90  5  2 87  EA  15  15  10  15  85  EB  15  5  25  15  15  EC  25  60  35  20  20  Humus  85  65  85  90  95  Decaying Wood  15  35  15  10  5 Species  Life form  Sub A layer  1  Pm  Pinus contorta  2  Pm  Chamaecyparis  3  Pn  Myrica gale  Significance M R  layer 3 nootkatensis  5.+.1  3  1.+.1 1.+.+  5.+.2  4.+.2  4.+.1  Vigor H  R  Con-  Cover  stancy  value  3.8  1-5  V  210  0-1  1.4 +  1-2  0.2  —  I  10  1-2  V  8700  V  340  8 layer 1  4  Ch  5  Pn  Oxycoccus quadripetal us Ledum groenlandicum  Pn  Chamaecyparis  Pm  Pinus contorta  9.6.2 6.4.1  9.7.2  9.6.1 5.4.1  8.5.2  8.6.1  8.6  8-9  1.6  2  5.5.2  5.5.1  4.2  0-6  1.3  1-2  2  2.2.2  2.2.1  3.2.1  3.2.1  3.2.2  2.+.1  1.+.1  0-2  1.4 1.0  1-2  1.+.1 1.+.1  2.6 0.8  2-3  2.+.1 5.4.1  2.+.1 6.5.1  —  7.5.1 1.+.1  1.+.1 5.5.2  2.+.1  1 2  6  Ch  1.+.1 6.5.1  nootkatensis 1 2  3.+.1  1  +.+.1  2  1.+.1  Empetrum nigrum  1  1.+.1  3.3.2  1.4  1-2  1.0  4.8  0-7  1.0  V  70  IV  2720  0.8  0-3  1.0  1.7 0.2  0-5  1.3  0-1  1.0  1-2  IV  425  0.6  0-3  2.0  0-2 0-2  1.5 1.0  1-2  IV  160  —  IV  60  IV  60  2  2.2.2  2.2.1  2.2.1  2.2.2  1.+.1 1.2.2  2.2.1  1.+.1  2.1.2  2.+.1 2.2.2  1.6 1.2  1.+.1  1.5  0-2  1.8  1-2  3.2.2  2.+.1  1.0  0-3  1.5  1-2 —  7  Pn  Gaultheria shallon  2  8  Ch  Vaccinium v i t i s - i d a e a  2  9  Pn  Kalmla p o l i f o l l a  1 2  1.+.1  10  Ch  Linnaea boreal Is  2  1.2.1  11  Pm  Tsuga heterophylla  2  +.+.+  2.+.1  1.0  0-2  3.2.1  1.0  1.+.1  +.+.+  0.4  0-3 0-1  1.0 1.0 0.7  +-1  1.+.1  1.1.1  IV  50  III  120  III  14  C layer 12  0  TrlentalIs  13 14  H H  Agoserls glauca Coptls a s p l e n l f o l t a  15  H  16 17 18  H  19 20  H H  Agrostis aequlvalvis Drosera r o t u n d ! f o l l a  21 22  arctica  3.+.3 2.+.2  1.+.1  3.+.2  3.+.1  3.+.2  2.6  1-3  1.8  1-2  V  410  2.1.1  3.+.2  3.2.2  2.2.2  2.4  2-3  1.8  1-2  V  260  3.t.2  1.2.2  2.+.2  2.+.1  2.+.1  2.0  1-3  1.6  1-2  V  170  Carex obnupta  3.3.2  7.4.2  2.3.2  2.2.1  2.8  0-7  1.4  1-2  H  Carex  5.5.3  3.4  0-5  2.5  2-3  IV IV  1140 1100  H  Carex p l u r i f l o r a  2.6  0-4 0-4  1.5 2.3  1-2  IV  500  IV  340  1.6 1.6  0-2  1.3  2-3 1-2  0-3  1.0  1-2  0.6 0.4  0-1  1.0  0-1  0.4  0-1  1.5 +  canescens  3.3.1 3.2.2 2.+.1  Sangulsorba mlcrocephala  2.+.1  Ch  Cornus  1.+.1  T  Gentiana  23  G  Maianthemum d i l a t a t u m  24  G  T o f l e l d i a occidental i s  25  G  Lysichitum americanum  canadensis douglaslana  3.2.2  1.+.1  4.3.2  3.2.2  3.2.1  4.3.2  2.+. 2  2.+.2  2.+.1 3.+.2  2.+.1  5.3.3 4.3.3 2.+.2 3.2.2  1.+.1  1.+.1  1.2.2 1.+.+  1.+.+ 1.+.1  1.+.+  1.+.2  2.2  0.4  0-1  0.2  0-1  1.5 +  1-2 1-2 —  IV  80  III  220  III  30  II  20  II  20  II  20  1  10  109  Species Significance M R  Sub-  Life D laver  form  layer  26 27  B  Sphagnum papillosum  8  Sphagnum capillaceum  28 29 30  B' B B  Sphagnum mendocinum Sphagnum recurvum Rhytidi adel phus loreus  31  B  Pleurozium schreber!  32 33  B 8  Dicranum scoparium Aulacomnium palustre  34 35  B B B  da Hylocomlum splendens dp Herberta adunca Rhacomitrium lanuginosum do  2.3.3  B 8 B  Bazzania ambigua  di di  1.1.2  du du  1.1.2  A B  2.1.1 2.1.1 3.1.2  36 37 38 39 40  B  h h d» h  6.4.2  h h du  3.2.2  h da d« h  Mnium glabrescens Riccardia sp. Scapania bolanderi  5.3.2  8.5.2  8.5.2 4.2.2  1.3.1  3.2.2 1.1.2 7.5.2  3.2.2 7.5.3 6.3.2 2.2.1 2.2.2  1.1.1 1.1.1  1.2.1 1.1.1  1.1.1  1.1.1 2.1.1  4.3.2 1.1.1 1.1.1  7.4.2 3.2.2  9.5.3 3.2.2  7.2  4.2.2  6.2 3.2  2.6 0.4  0.2 1.0 0.4  1.2.1  1.1.1 2.1.2  1.1.1 1.1.1 1.1.1  1.2.1  1.2.1  0.6 0.8 0.6 0.6 0.4 0.4 0.2 0.2 0.2  1.2.2  0.2  1.1.1  5-9 0-4 0-1 0-7 0-6 0-1 0-2 0-2 0-1 0-2 0-1 0-1 0-2 0-2 0-1  Constancy  Vigor D I  R  2.2  2-3  2.0 1.5  —  2.3 2.0 1.0 1.3 2.0 1.0 1.3 1.0  Cover value  V V  5960 510  2-3  IV  3200  —  IV  1060  IV  50  IV III III III  50 40 30  1-2  —  1-2 - — —  1-2 —  1.0 1.0 3.0  —  2.0 2.0 2.0  —  — —  30 20 20  I I I  10  —  I I  10 10  1.0  —  1  10  2-4 2-4 3-4  1.0 1.4 2.0  —  —  V  600  0-2 1-3 0-1 0-3 0-3 0-2 0-2 0-1 0-2 0-1 0-1 0-1 0-1 0-1 0-1 0-1  1.5 1.4 2.0  1-2 1-2  V  170  1.3 1.3 1.0 1.0 1.0 1.0 2.0 1.0 2.0 1.0 1.0 2.0 1.0  1-2 1-2  IV III III  130 120 50  III II 1 1  40 20 10  0-2 0-3 0-2 0-2 0-2 0-1 0-1  1.0 2.0  0-1 0-1 0-1  —  E layer Bryophytes S lichens 41  E  Frullania ntsquallensis  c' E E  42 43  44 45 46  47  E E E E E E E E E E  Olcranum scoparium  B  C B C Herberta adunca C Antltrichia curtlpendula B C Isothecium stolonlferum B C C Bazzania ambigua C Cladonia bellidlflora 01plophyl1um albicans C B Hypnum circinale  4.2.1 4.2.2 3.2.2  2.1.1 3.2.2  2.1.1 1.2.2 1.1.1 3.3.2  Scapania bolanderi  Rhacomitrium lanuginosum C C Riccardia sp. Sphaerophorus globosus B  1.1.1 1.2.1 1.1.1  2.1.1 2.1.2 4.1.2  2.1.1 2.1.1 3.2.2  1.2.1  2.2.2 1.2.2  2.1.1  1.2.1  3.1.2  1.2 1.0 0.6 0.6 0.2  1.2.1 2.2.1 1.2.1  2.2.1 1.+.1 2.1.1 1.1.2  1.1.1 1.1.2  2.4 2.4 3.2 0.8 1.8 0.2  2.1.1 2.1.1 3.1.2  0.4 0.4 0.2  1.1.1  1.+.1 1.1.2  0.2 0.2 0.2 0.2 0.2  1.1.1 1.1.1 1.1.2 1.1.1  1-2  —  — — —  — — — —  —  1 1 1  —  1  — —  10 10 10 10 10  Vascular plants Empetrum nigrum  B  E  Cornus canadensis  C B  E E  Vaccinium vitis-idaea Polypodlum glycyrrhiza  C C B  E E  Tsuga heterophylla Vaccinium parvifolium  C B B  +.+,+ +,+,+  Total species  39  E  48  Life form  Proportion by Species:  Nraber  t Total cover:  cover  I  2.2.1  0.4 0.6 0.4 0.4 0.4 0.2  3.2.2 2.+.+ 2.+.1 2.2.2 1.+.+ 1.+.1  38  33  E  B  H  19 31.7  15 25.0  6 13.3  1394 4.7  11010 37.4  3810 12.9  31  0.2 0.1 0.1  0-+ 0-+  + 1 2  + 1  + +  — — —• — — — — ~  II II  120 40  1 II  20 20  1 1  2 2  23  Ch  5 8.3 710 2.4  Pn  5 8.3 11600 39.4  &  4 6.7 460 1.6  Pm  3 5.0 449 1.5  T  1 1.7 20' 0.06  110 SYNTHESIS TABLE V.  PINETO-SPHAGNETUM CAPILLACEI Variant:  P.-S.  c a p i l l a c e i chamaecyparosum  Number of p l o t s P l o t number Plot size  (meters ) 2  Extent of type  (acres)  Date  nootkatensis  1  2  3  4  11  13  4  3  5 2  100 4  100 2  100 5  100  100  25/5/  26/5/  19/5/  3 19/5/  17/5/  2  64  64  64  64  64  Percent coverage layer:  A  2  5  30  10  10  15  5 10  20  15  8  Total  A3 A  35  20  18  15 20  B1  70  40  65  80  65  45  20  25  50  35  Total  B2 B  75  45  75  85  70  C  20  50  40  45  65  Oh D ,  65 30  65 30  75  55  60  20  20  35  D  85  95  95  80  95  EA  20  15  10  20  25  EB  50  50  45  35  60  EC  50  80  60  60  65  Humus  70  65  80  85  70  Decaying Hood  30  35  20  15  30  Vegetation:  0  Total  Spec! s Life  1  Sub-  form  A laver  Pm  Pinus contorta  Vigor  Constancy  M  R  value  2  2.3.1  5.7.2  3.6.2  3.6.2  3.4.2  3.2  2-5  1.8  1-2  3  1.+.1 2.3.1  4.6.2  3.6.2  3.5.2"  3.3.2  2.8  1-4  1.8  1-2  3.3.1  2.2.1  2.0  2.3.1  1:3.1  0.6  0-3 0-2  1.0  —  0.2  0-1  1.0  —  I  10  7.2 3.0  6-8  1.0  —  V  5660  0-5 1-5  2.5  2-3  2.4  2-3  V  1200  1.8  1-2  V  900  1.3  1-2 V  600  2  Pm  Chamaecyparis  3  Pm  Thuja p l i c a t a  3  4  Pm  Tsuga heterophylla  3  nootkatensis  Cover  Significance M R  laver  3  3.4.1 1.+.1  V  1.0  II  720 240 30  II  B layer Pm  7.4.1  Ch  Chamaecyparis nootkatensis Empetrum nigrum  1  5  1 2  5.4.3  6  Pn  Ledum groenlandicum  1  5.4.3 4.3.2  6.5.1  8.5.1  8.4.1  7.4.1  4.4.2  3.3.2.  3.2.3  5.4.3  1.2.1 3.3.1  1.1.2 3.2.2  1.1.2 3.2.2  2.6  5.3.2  1.1.1  1.2.2  0.6  3.3.2  3.3.2  3.1.2  1.2.1 3.3.2  3-5 0-1  3.2  3-4  2.0  1.1.2 1.2.1  1.2.2  0-1  2.0  1-4  3.1.2  2.2.1 2.1.2  0.6 2.6 2.4  1-2  V V  430 260  3.2.1  1.0  —  IV  320  1.2.1  1.2.1  2.2 0.8  2-3 0-3  1.4 1.8  3.2.1 2.1.2  2.1.2  1.8  IV  160  IV  50  2 7  Pn  Kalmia p o l i f o l i a  1 2  Pm  Pinus contorta  1  4.3.2 3.5.1  3.4.2  1.2.1 4.4.2  8  Ch  2.1.1  2.2.2  3.1.2  Pn  Oxycoccus quadrlpetalus Gaultheria shallon  2  9  1  3.3.1  2  2.2.1 2.2.1  10  Ch  Vaccinium v i t i s - i d a e a  2  2.1.2  3.2.2  11 12  Ch  2  1.1.1  Pm  Linnaea boreal i s Tsuga heterophylla  2.1.2 2.4.1  13  Pm  Thuja p l i c a t a  1  14  Pn  Vaccinium ovatum  1  1.+.1  15  Pn  Vaccinium p a r v i f o l 1 un  1  1.+.1 1.4.1 1.+.1  1 2  1.1.1  1.4.1  16  Pm  Picea s i t c h e n s i s  17  Pn  Vaccinium oval 1folium  2  18  Pn  Vaccinium uliginosum  2  19 20  H H  Carex obnupta Agoseris glauca  4.4.2 4.2.2  5.3.2 5.2.3  21  Ch  Cornus canadensis  2.1.2  22  H  Drosera r o t u n d i f o l i a  1.+.2  2.1.1 2.4.1  23  G  Maianthemum dilatatum  1.4.4  24  H  Carex p l u r i f l o r a  1.2.1  25  H  Coptls a s p l e n l f o l l a  i.+.r  26  G  Lysichitum amerlcanum  27  G  Trlentalls arctica  28  H  Sanguisorba mlcrocephala  29  G  Tofieldia occfdentalis  30  Ch  Lycopodium clavatum  31  H  Blechnum spicant  1.2.2 2.2.1  32  T  Gentiana douglaslana  2.4.2  33  H  Sclrpus  34  H  Agrostis  35  H  Deschampsia  36  G  Nephrophyllldiura c r l s t a - g a l l l  1-2  0-2  1.0  0-3 0-2  2.0  1.3  1.3  1-2  0.4  0-2  1.0  —  0.6  0-2  0.8  4-2  II  30  0-1  1.0  —  II  20  0.4  0-1  1.0  II  20  0.4  0-1  1.0  II  20  0.2 0.2  0-1  1.0  0-1  4  1  10  0.2  0-1  f  10  1.1.2  0.2  0-1  1.0 2.0  —  1  10  6.6  1.4.1  2 2  •  0.4  2.1.1  1.+.+ 1.+.1  3.6  1.4.1 1.2.1 1.+.+  C laver 8.3.2  8.4.2 5.2.3  8.2.2 7.2.3  5.4  4-8 4-7  2.0 2.8  —_  6.3.3  2-3  V V  5100 2660  2.1.1  2.1.1  2.1.2  2.0  2  1.4  1-2  V  100  1.1.2  2.1.2  1.6  1-2  1.8  80  2.1.1  2.1.1  1.6  1-2  0.8  1-2 4-1  V  1.4.4  2.1.2 2.1.1  V  80  4.2.2  3.2.2  2.2  0-4  1.4  1-2  IV  410  3.1.2  3.2.2 1.4.1  2.1.2  1.4  0-3  1.5  1-2  IV  140  1.1.1 1.4.2  3.1.2 1.4.1  1.2.1  1.4  0-3  1.3  1-2  IV  140  0.8  0-1  1.5  1-2  IV  40  1.4.1  2.1.2  0.8  0-2  1.3  1-2  III  40  1-2  III  30  III II 1  30  1  20  1  10  2.2.1 1.+.2  1.4.1 1.+.1 1.+.1  1.4.2  1.2.2  0.6  0-1  1.7  1.1.2  1.1.2  0.6  0-1  2.0  0.6  0-2  1.0  0.4  0-2  2.0  0.4  0-2  2.0  0.2  0-1  +  1.+.1  0.2  0-1  1.0  —  1  10  1.2.2  0.-2  0-1  2.0  —  1  10  1.1.1  caespitosus aequlvalvls caespitosa  2.2.2 1.+.+  —  30 20  Life form  D laver  37 38 39  B B 8  Sphagnum papillosum Sphagnum caplllaceuo Pleurozium schreberi  40 41 42 43 44 45 46  B B 8  Sphagnum recurvum Rhytidiadelphus loreus Plagiotheciun undulatun Herberts adunca Rhacomitriun lanuginosum Scapania bolanderl Hylocomlum splendens  47 48 49 50 51  L L B 8 B  52 53 54 55 56 57 58 59 60 61 62  eB B B  B B B 8 L B B B B B B  Species Significance R »  Subayer h h h di h da  .da da da da li da da Cladonla rangiferina da Cladonla paclfica h Aulacoonlua palustre h Dicranum bergerl h Dicranum 6coparlum da Isothecium stoloniferum da da Dicranum fuscescens da Bazzania ambigua Sphagnum tenellum h da Cladonla b e l l l d l f l o r a Nnlum glabrescens di h Polytrichum commune da Cephalozla bicuspidata da Plagiochila asplenioldes da Bartramla pomi forml s da Eurhynchium oreganum  4.3.2 4.3.2  5.5.2 5.4.2  2.2.2 1.1.2  2.2.2 3.3.2 3.2.2  1.4.4  1.4.4  5.2.2. 4.1.2 2.2.2  6.5.3 4.3.2 2.2.2 2.2.2 6.5.3 3.3.2  4.3.2 2.1.2  4.3.2 3.2.2  2.2.2 5.3.2 2.2.1  2.1.1 5.3.3  1.4.1  4.3.2 2.3.2 3.2.2  3.2.2  1.1.1 2.3.2 1.1.1 1.2.2 2.2.2  2.2.1  2.3.2 3.2.2  3.2.1 1.2.1 3.3.2 3.2.2 2.2.3 1.1.2 2.2.2 2.2.2  1.2.1 1.1.1  1.2.2 1.2.2  3.2.2  1.1.2  3.2.2 3.3.2 1.1.2  2.1.2 1.1.1  4.4.2 2.1.1 2.2.2 2.1.1 1.1.2 1.2.2 1.2.2 4.4.1  Vigor  Constancy  t  8 2-3  0-4  2.2 2.0 2.0 1.8 2.5 1.4 0.5 2.0 2.0 2.0 1.0 1.0 2.0 1.3 2.3 1.6 2.0 2.0 2.0 1.5 2.0 2.0 1.0 2.0 1.0 2.0 2.0 2.0 1.0  3.6 3.6 2.6 2.4 2.7 1.0 1.2 2.4 1.6 2.2 1.0 0.6 2.0 2.2 0.8 0.4 0.4 0.4 1.0 0.8 0.8 0.6 0.4 0.4 0.6 0.4 0.4 0.2 0.2 0.2 0.2 0.2 0.2 0.2  2-4 3-4 2-3 1-4 0-3 0-2 0-2 1-3 0-3 1-3 0-2 0-3 0-4 0-4 0-1 0-2 0-2 0-2 0-3 0-2 0-2 0-2 0-1 0-2 0-3 0-2 0-2 0-1 0-1 0-1 0-1 0-1 0-1 0-1  2.6 2.4 2.0 1.6 1.7 1.7 2.0 1.6 1.5 2.0 1.7 2.0 2.0 2.0 1.0 2.0 2.0 2.0 2.0 1.3 2.0 2.0 1.5 1.0 2.0 1.0 2.0 1.0 1.0 2.0 2.0 1.0 2.0 2.0  2-3 2-3  0.4 1.0 0.2 0.4 1.2 0.4 0.8 0.6 0.4 0.2  0-1 0-2 0-1 0-2 0-3 0-2 0-2 0-2 0-2 0-1  1.5 1.0 1.0 2.0 2.0 1.0 1.0 1.0 2.0 1.0  —  4.6 3.6 0.4 2.0 3.8 1.8 0.8 2.0 1.6 1.6 0.6 0.8 1.2 1.0 0.8 0.8 0.4 1.0 1.0 0.8 0.4 0.8 0.4 0.4 0.2 0.2 0.2 0.2 0.1  4-6 2-5 0-2 2 0-6 0-3 0-1 0-4 0-4 0-3 0-3 0-2 0-3 0-3 0-2 0-2 0-2 0-3 0-3 0-3 0-1 0-4 0-2 0-2 0-2 0-1 0-1 0-1  —  Cover value  V V  1660 920  V IV IV IV III III III  100 1560 230 40 400 240 220  III III III III III  140 130 . 120 40 40  —  1-2 2-3 1-2 4-1  1-2 2-3 1-2 —  II II II II I I I I I I I  — —  1-2 — — — — — — — —  1  120 120 110 20 ZOO 20 ZO 20 10 10 10 Z  V  900  V  350  V  340  V  330  IV IV III  600 100 60  II II II II II  1Z0 4D 40 30 20  —  E laver Bryophytes 4 lichens 63  E  E  E  64  65  Isothecium stoloniferum  Dicranum scoparium  E  Scapania bolanderi  E  Herberta adunca  E E  E E E E E E  66 67 68  Frullanla nlsquallensls  E E E E E E  69  E E  70  E  Cladonla b e l l l d l f l o r a Antitrichia curttpendula  Rhacomitrium lanuginosum Dicranum fuscescens Macrodtplophyllum pi 1 catJB Bazzania ambigua Cephalozla bicuspidata Pleurozium schreberi  A B C A B C A B C B C A 9 C C A B C C C  4.2.3 4.2.2 2.2.2  2.1.1 2.1.1 1.+.1 1.1.1  2.2.1 1.2.1 2.2.2 2.1.2 3.2.2 3.2.2 3.2.2 1.*.1  4.3.2 2.1.2 1.+.1  1.1.1 1.2.1  2.2.2 3.2.2 3.2.2 2.1.1 2.2.2 2.1.2 3.2.2 2.2.2 3.2.2 2.2.2  4.2.3 4.2.3 3.2.2 3.2.2 3.2.2  4.1.3 4.1.3 2.1.2 2.1.2 2.1.2  2.2.2 2.2.2 3.2.2 3.2.2  2.1.2 3.1.2 2.2.3  3.1.2 2.1.2 1.+.2 2.2.2  4.3.2 1.4.1  2.2.2  c c c A C C C  Hylocomlum splendens Bartramla pomlformls 8azzanla nudlcaulis Blepharstoma trfchophyli JD C Hypnum circinale B lilnium glabrescens C MyIIa taylorl Plagiochila asplenioldes B Sphaerophorus globosus  4.2.2 3.2.2 3.2.2 4.2.2 3.1.1 2.2.2  2.2.2 2.2.2  3.2.2 2.+.+ 2.2.2 2.1.2  2.2.2 2.2.2 1.1.2 1.1.2  1.4.1  2.1.1 3.2.2 2.2.1  c  2.1.2 1.1.1 1.1.1 1.2.2 1.1.2  c c  1.4.1  1.1.2 1.2.2  —  1-2 1-2 1-2 —  1-2 1-2 1-3 1-2 — — — — — — —  4-2 — —  1-2 —  —  1 1 1 I 1 I 1 1 1  —  1  100 20 ZD 10 10 10 10 10 10 10  V  60  — — — — — — — —  Vascular plants 71  E  Polypodium glycyrrhfza  E  Eopetrun nigrum  E  Gaultheria shallon  E E E  Vaccinium vitis-idaea Llnnaea boreal Is Kaiantheoun dilatatum  A B C 8 C B C 6 C B  1.1.2 1.4.4  Proportion by Sped es:  Total cover:  Number  J  cover  J  2.2.2  3.2.2  1.1.1 1.1.1 1.1.1 2.3.2  3.2.2  2.2.1 2.1.1  49  45  2.3.1 1.4.1  2.1.1 2.1.2 1.1.1  Total species  Life form  1.4.4  44  43  E  B  H  26 31.8  23 26.1  10 11.4  7 8.0  3440 12.0  6232 Z1.6  8500 29.4  1880 6.5  Pn  4-2 — —  III  —  220  —  III II 1 1  — — — —  60 30 20 10  39  Ch  6 6.8  tsoo 6.Z  Pm  5 5.7 6450 22.3  6  L  T  5 5.7  3 3.4  1 1.1  300 1.03  270 0.9  20 0.06  112  SYNTHESIS TABLE V.  PINETO-SPHAGNETUM CAPILLACEI Variant:  P.-S. c a p i l l a c e i vacciniosum v i t i s - l d a e a e  Number of plots Plot number Plot size (meters ) Extent of type (acres) Date 2  Percent coverage Vegetation: layer: Total  A2  »1 82 B C Oh de 0  EA EB EC Humus Decaying Vood  1  3  4  104 25 1/10 5/9/ 64  16 25  24 25 1/10  7 25 1  11/6/ 64  21/5/ 64  65 75 35 90  n  Total  2  0 0 0 40  »3 A  Total  1  Life form  A laver  layer  Pm  Pinus contorta  2  1/3 27/5/ 64  0 0 0 25 55  5 92 0  65 30 75 15 90 0  5 5  5 10  90 10  80 20  10 0 10 20 50 65 25 95 1 96 2 5  0 0 0 60 15 ' 65 70 65 20 85 0  10  20 30  95 5  80 20  5  6  7  8  9  83 25  5 25  29 25  1/2 26/8/ 64  1/2 20/5/ 64  89 25 1/20 28/8/ 64  88 25 1/10 28/8/ 64  0 0 0 35  0 0 0 25 65 70  0 0 0 25  0 0 0 60  0 0 0 35  65 80  15 75 5 80 0  15 85 2 87 0 10  50 75 45 80 5 85 0  55 65 25 70 5 75 0  15  5 10  5 15  95 5  90 10  90 10  70 75 15 65 3 68 0 5 5  5 5  90 10  90 10  1/2 1/7/ 64  10 30 25 1/20 1/7/ 64  0 10 10 40 50 65 15 75 8 83 5 5  11 1 25 2 16/5/ 64  0 0 0 60 35 70 25 90 5 95 0  15  5 0  90 10  90 10 Species Significance  Sub-  1.+.2  Vigor  0.09  0-1  2.0  2.4 5.3 5.9 2.4 5.6 3.1  0-7 0-9 1-8 0-4 3-7 1-6 0-4 1-5 0-4 1-4 0-2 0-2 1-4 0-3 0-3 0-3 0-1 0-1 0-1 0-1  2.8 2.4 2.0 1.7 2.3 2.0 1.0 1.0 1.7 1.9 1.1 0.9 1.1 1.0 1.8 1.4  1.0  —  1-8 0-5 1-4 0-4 0-4 0-3 +-3 0-2 0-3 0-3 0-3 0-4 0-3 0-1 0-3 0-3 0-2 0-2  1.2 1.9  +-2 1-3  1.1 1.0  1.2 +-1 1-2 1-2 +-1 1-2 1-2  Constancy  Cover value  (  4  V V  4672 3768  V  3063  V V V  577 550 350  V V  68 286  IV III II I I  161 63 10 9 4 4  B layer 2  Ch  Empetrum nigrum  3  Pn  Ledum groenlandicum  *  Pn  Kalmia p o l i f o l i a  5  Pn  Gaultherla shallon  6 7 8  Pn Ch  Vaccinium ullginosum Vaccinium v i t i s - i d a e a Thuja p l i c a t a  Pm Pm  9 10 11 12 13 14  Ch Ch Pm Pn Pm Pn  PI nus contorta Oxycoccus quadripetalus Ltnnaea boreal i s Tsuga heterophylla Vaccinium parvifolium Picea sitchensis Vaccinium ovalifolium  1 2 1 2 1 2 1 2 2 2 1 2 1 2 2 2 2 2 2 1  7.3.3 7.5.2 4.3.2 4.4.2 6.3.2  5.3.2 6.3.2 4.2.2 6.3.2 5.2.2  9.5.2  2.+.1 4.3.2 1.2.2  4.2.1 4.2.2 3.2.2  5.4.1 3.2.1 3.2.1  1.+.1 1.+.1 4.+.1 1.+.1 3.2.2 3.2.2  2.+.1 2.+.1 3.+.1 2.1.1 1.+.2  1.+.+ 1.+.1 1.+.1  1.2.1 1.+.1 7.5.2 4.2.2  1.+.+  8.3.3 1.1.2 3.2.2 1.1.2 4.2.1 1.1.1 3.2.2 2.2.2 2.2.2 1.+.1 3.2.2  8.5.2 6.3.2 4.2.2 7.4.3 4.2.2  8.5.3 6.3.2 4.2.1 6.3.3 4.2.2  4.+.1 4.2.2 2.1.2  4.2.1 3.2.2  2.+.1 2.+.1 2.+.1 2.2.2  2.+.+  1.1.2  1.+.1 3.+.1 2.2.2  6.3.2 5.3.2 4.2.2 5.3.2 6.2.2 2.+.1 4.2.2 2.1.2 +.+.1 1.+.1 2.+.1 3.+.1 3.2.2  1.1.1 1 + + 1.+.1  6.3.3 1.1.3 5.2.2 1.1.2 7.5.2 1.1.2 2.2.1 1.1.1 3.1.2 1.1.2  7.5.3 7.5.3 7.5.2 1.1.2 6.4.2 1.1.2 4.3.1 3.2.1 1.2.1 4.2.2  7.4.2 7.4.2 7.5.2 2.2.2 5.4.2 2.2.2 2.2.1 3.2.1 3.2.1 3.2.2  6.3.3 7.3.2 6.3.2 1.1.2 3.1.1 1.1.1 3.1.2  1.+.+ 2.1.1 2.+.1 2.1.1  1.+.1 1.+.1  1.+.1  1.2.1  4.5.1 1.+.1 2.2.1  1.+.1 3.2.1 2.1.2  1.+.1  1.+.+ 1.2.1  1.4 2.7 2.6 2.4 0.4 1.1 2.3 1.6 1.7 0.6 0.2 0.2 0.09 0.09  2-3 2-3 1-3 1-2" 2-3 — — —  1-2 1-2 0-2 +-1 1-2 —  1-2 1-2 —  1.0  — —  '  C layer 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 30  H H Ch H H G G H H H T H H H H 6 G H 0 G G H  Carex obnupta Agoserls glauca Cornus canadensis Blechnum spicant Sangulsorba mlcrocephala Trientalis arctica Maianthemum dilatatum Orosera rotundlfol!a Agrostis aequivalvls Carex p l u r i f l o r a Gentlana douglasiana Coptls t r i f o l i a t a Coptis a s p l e n i f o l l a Deschampsla casspltosa Gentlana sceptrum Nephrophyllldlum c r l s t a - g a l l l Habenarla saccata Rhynchospora alba Boschniakla hooker! Lysichitum americanum Tofieldia occidentals Scirpus caespitosus  4.2.1 4.2.3 1.+.1 1.+.1 1.+.1  +.+.+ 1.+.1 1.2.2 3.2.1 +,+.+ 4.2.3 3.2.2  3.2.1 4.2.2 3.+.1 4.3.1 2.+.1 2.+.Z 1.+.1 1.+.2 3.2.2 1.+.1 3.2.2  2.2.+ 2.2.1 4.1.1 1.+.+ 2.2.1 3.+.2  8.5.2 2.1.2 3.2.1  2.+.1 4.2.2 2.+.1 2.+.1 2.+.1 3.+.2  +•+.+ 1.+.+  +.+.+  1.1.2  1.+.1 3.+.1 2.+.1 3.2.2  2.1.2 3.1.2  3.1.1 1.+.1  2.1.2 1.2.2  2.+.1 4.2.1 2.+.1 3.+.1 3.+.1 2.+.2 3.+.1 1.+.1 2.+.1  2.1.2 3.1.2  3.2.2 3.2.1 3.1.2  8.6.2 4.2.2 3.2.1 3.2.1 4.2.2 1.+.2 2.1.1 2.+.2  3.3.1 4.2.2 4.2.1 1.2.1 1.+.1 1.+.1 1.+.1  1.+.1 3.2.2  1.2.1 5.2.1 2.2.1 2.2.2 2.+.1 1.+.1  1.+.1  2.+.2 1.+.1  3.+.2  3.2.1 4.2.2 2.1.1 1.+.1 2.+.2 1.+.2 1.+.1  1.+.1 1.+.1 3.2.1  2.+.1 2.2.2 1.2.2  +.+.1 1.1.1  1.+.2 +.1.1  3.2.2 4.1.3 1.+.1 2.+.1 1.+.2 3.+.1 1.+.2  3.5 0.5 2.3 1.9 1.7 1.5 1.3 0.9 1.4 1.1 1.0 0.9 0.7 0.4 0.5 0.3 0.2 0.2 0.13 0.09 0.09 0.04  0-1 0-1 0-1 0-+  1.2 1.7 0.8 1.6 1.6 1.0 1.5 1.8 2.0 1.3 2.0 1.0 1.0 2.0  —  +-2 1-3 —  1-2 —  V V V V V V V IV III III III III II II I 1  1688 918 327 263 181 136 125 45 195 150 141 118  9 9  1.5 1.0 2.0  — —  1 1 1 1 1  1.0  —  1  — —  1-2  100 16 90 45  5 4 4 1  113  Life form  Species Significance M R  Sublayer  OJ.ayer  37 38  B I  Sphagnum i-.apl 11 aceum Cladonia p a c i f i c a  h h  6.3.2  6.3.3 4.3.2  2.4.2 7.4.2  8  Pleurozium schreberl  d> h  1.1.2 5.3.3  3.2.2  39  3.1.1 3.2.2 3.2.2  40 41 42 43 44 45 46 47  L B 8 B L B B B  Cladonia crlspata Sphagnum papillosum Rhacomitrium lanuginosum  di h h h  Dicranum scoparium  di h  Cladonia rangiferina  di h  Rhytldiadelphus loreus Dicranum bergeri Hylocomlum splendens  48 49 50 51  B 8 8 L  52  B  Aulacomnlum palustre Cladonia b e l l l d l f l o r a Eurhynchium oreganum  53 54  B B  Cladonia u n c ' a l i s Plagiothecium undulatum  55 56 57 58 59 60 61 62 63 64 65 66  8 B B L B B B B B B B B  Scapania bolanderi Sphagnum mendocinum  Sphagnum fuscum Sphagnum recurvum Polytrichum strictum Icmadophlla erlcetorum Polytrichum commune ttnlum glabrescens Riccardia sp. Bazzania ambigua Cephalozia blcuspldata Dicranum fuscescens Eurhynchium stokesll Isothecium stolonlferum  d> h di h h de di h h  2.2.2  2.1.2  5.3.2  4.2.2 4.3.2 3.2.2 1.2.2 3.2.2 3.2.2 2.2.1  3.2.2  2.2.1 1.1.1  di di h di di d» d< di  7.3.2 4.3.2 3.2.2 4.2.2 3.1.2 7.3.3 3.2.1 1.2.2  5.2.3 3.2.2  6.4.2 2.1.1 3.2.1  3.2.2 4.2.2 3.2.2 2.1.1  1.1.1 3.2.3  1.1.1 3.2.2  4.2.2  4.2.1 1.2.1  3.2.2  6.3.2 7.3.2 3.2.2  2.1.2  2.2.2 2.2.2 2.1.2  6.2.3 4.2.2  4.2.2 4.2.2  4.3.2 3.2.2  3.1.1 3.1.1 3.2.2 6.2.2 4.2.2  4.2.2  2.1.1 4.2.1  2.1.1 3.2.1 1.1.1  1.1.1 1.1.1 4.2.3  2.2.1 1.2.2  3.2.2  4.2.2 2.2.2  1.+.1 2.1.1  4.2.2  3.2.2 2.1.1  1.2.2 3.2.2  2.2.2  2.2.2  1.1.1 4.1.1  1.1.1 1.+.1  2.1.2 3.1.2 1.2.2 3.2.1 1.1.1 4.1.2 3.1.2 1.+.Z  3.2.2 2.2.2 2.2.2  6.4.2 3.2.1 3.2.1 3.2.2 3.2.2 3.3.2 3.3.1 3.2.2 2.2.1 4.2.2 4.2.2 3.3.2 3.2.1  5.5 3.3 1.5 2.7  8.3.3 1.1.2 1.1.1  1.6 2.4 2.6 2.2 0.9 2.0 1.8  3.3.2 4.3.2  2.2.2  2.1 0.7 0.8 0.5 1.9 0.5  1.1.2  3.1.2 1.2.2  2.2.1 2.3.1  de h . di h h deli h h  5.4.3  2.1.2 3.1.2 3.1.2  2.2.1  2.1.2  1.2.2  2.1.2  1.2.2 1.1.1  1.1.1  1.1.2 3.2.2 1.+.1  1.+.2  1.2.1  1.1.1 1.2.3  2.1.2 1.1.3  +.+.+  1.+.+  2.1.1 2.1.3 +.1.+ 1.+.1  6.3.2 4.2.2 3.2.2  3.2.2 3.2.1 2.2.1 1.2.2  2.2.3 1.1.2  2.1.1 2.2.2 1.1.2  1.1.1  1.1.1 1.1.1 1.1.2  it  1.1.1  di  1.1.1  R  2-8 0-7  2.4 1.9  0-3 0-5 0-3 0-7 0-6 0-6 0-4 0-4 0-4 0-4 0-3 0-3 0-2 0-4 0-3  1.6 1.8  2-3 1-2 1-2 1-3 1-2 2-3 1-2 1-2 1-2 1-2 1-2 1-2 1-2 1-2 1-2  1.9 2.3 1.9 1.9 1.5 1.9 1.5 1.6 1.3 1.3 1.5 2.3 1.5 1.3 1.6 1.8 1.8 1.2  2-3 1-2 1-2 1-2 1-2  1.0 1.5 3.0  —  1.2 1.0 1.4 0.8 0.5 0.2 0.4 0.4 0.04  0-3 0-2 0-4  0.2 0.5 0.6 0.5  0-1 0-6 0-4 0-3 0-2 0-2 0-2 0-2 0-1 0-1 0-1 0-1 0-1 0-1  0.7 2.0 2.0 1.5 1.0 2.5 1.5 2.0 1.5 1.0 1.0 2.0 1.0 1.0  2.8 2.5 0.2 0.5 0.5 0.5 0.5 0.09 0.4 0.4 0.4 0.4 0.2 0.09 0.09 0.09 0.09 0.09 0.09 0.09  1-4 0-4 0-1 0-3 0-3 0-2 0-2 0-1 0-2 0-3 0-3 0-3 0-2 0-1 0-1  1.5 1.7 1.0 1.0 1.0 1.0 2.0 2.0 1.7 2.0 1.5 2.0 2.0 1.0 1.0  0-1 0-1 0-1 0-1 0-1  +  1.1 0.6 0.5 0.09  0-9 0-5 0-4 0-1  0.2 0.3 0.3 0.2 0.2 0.09 0.09 0.09 0.09 0.09  Constancy  Vigor »  0-3 0-1 0-1 0-2 0-2  1-2 1-2 1-2 —  Cover val Lie  V V  3009 1372  V  650  IV IV IV  827 804 622  V  440  V  390  V  86  IV  336  IV IV III III III  131 54 245 72 22  II II  27 18  II I 1 1  10 300 136 90  1 1 1 1 1 1 1 1 1  13 13 9 9 4 4 4 4 4  V  477  —  0-4-  <-1 — —  1-2 —  2-3 1-2 —  1-2 — — — — —  E layer Bryophytes & lichens 67  68  69  70  71 72 73 74  E  F r u l l a n l a nisquallensis  E  Dicranum scoparium  E ' E E  Isothecium stolonlferum btacrodiplophyllum plicatum  2.1.1 2.1.1  Parmelia physoides  E E E E E E  Pleurozium schreber) Radula bolanderi Dicranum fuscescens Bazzania ambigua Hylocomlum splendens Plagiochlla asplenloldes A n t l t r i c h i a curtlpendula Sphaerophorus globosus Usnea p l i c a t a  2.1.1 2.1.2  2.2.1  3.2.2 3.2.2 2.1.1 3.1.1 3.1.1  4.1.3 4.1.3  3.1.1 3.1.2  3.1.2  1.1.1  3.1.2  1.1.1  4.1.2 4.1.2  3.2.1 2.2.1  3.2.1 4.2.1  3.1.2  2.2.1 1.1.1 2.1.1  1.1.1  2.1.1  c  2.2.2  2.2.2  1.1.2  1.1.2  6 C B C  Scapania bolanderi  E  E E E  B C A B C C  2.2.2  1.1.2 1.1.2 1.1.2  C  1.1.2  C C  2.2.2  c c c c  1.1.1 1.+.* 1.+.1 1.+.+  1.1.1 3.1.2 3.1.1 3.1.2 1.1.1  B A  1.1.1 1.+.+  1.0 + 1.0 •f  1-3 1-3 — — — — —  II II  68 22 22  II  —  1-2 —  II 1  22 50  —  —  1 1 1 1 1  —  1  50 9 4 4 4 4 4 4 4  — —  1 1 1  1163 190 100  —  1  4  — — — —  — —  —  1 1 1  Vascular plants E E E E  Empetrum nigrum Gaultherla shallon Vaccinium v i t i s - i d a e a Maianthemum d l l a t a t u a  Total species  L i f e form Proportion by Species:  dumber  1 Total cover:  cover  {  3.3.2  9.4.2 5.2.1 4.2.1  c c c c  2.1.1 2.2.1 1.+.1  43  42  39  38  36  36  36  36  31  29  29  B  E  H  G  Pn  L  Ch  Pm  T  24 28.2  19 22.4  13 15.3  7  6 7.1  6 7.1  5  4 4.7  1  5.9  748 2.7  3756 13.4  328  5593 19.9  86 0.3  141 0.5  7063 25.1  8.2  1.2  7971 28.4  2642 9.4  1.2  2.0 1.0 1.0 1.0  —  SYNTHESIS TABLE  V.  PINETO-SPHAGNETUM CAPILLACEI Variant:  P.-S.  c a p i l l a c e i vacciniosum p a r v i f o l l l  Number of p l o t s P l o t number Plot size  (meters ) 2  Extent of type  (acres)  Oate  1  2  3  4  5  105  107  31  32  108  25  25  25  25  25  2  1  1/20  1/20  5/9/  7/9/  5/7/  12/7/  64  64  64  64  64  50  1 7/9/  Percent coverage Vegetation:  layer:  A3  30  40  70  65  90  55  70  75  35  15  25  70  45  20  Total  Bl B2 8  95  65  85  75  50  C  25  50  30  25  30  "h  60  65  20  35  60  10  10  2  3  15  D°d«  70  75  22  38  70  15  15  20  30  15  30  25  25  35  25 35  Total  EA • E  B  35  35  45  40  Humus  75  85  95  90  75  Decaying Wood  25  15  5  10  25  E  C  S p e d es Life  Sub-  form  A laver  layer  1  Pm  Pinus contorta  3  6.6.2  7.3.2  2  Pm  Thuja p l i c a t a  3  2.5.2  4.5.2 '  8.5.2  Significance M R  Vigor M  R  Con-  Cover  stancy  value  8.5.2  4.2.1  6.6  4-8  1.8  1-2  V  4860  2.1.2  7.5.1  3.0  0-7  1.8  1-2  IV  1240  B laver  3  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 2  5.5.1  6.5.2  3.+.1  4.3.2  5.3.1  4.6  3-6  1-2  V  1760  1.4  0-3  5.4.2  3.+.1 4.3.2  3.+.1  1  1.+.1 5.4.2  1.4 1  4.3.2  4.6  4-5  2  —  V  1600  2  4.2.2  3.2.+  3.2.2  3.2.2  2.6  0-4  1.3  1-2  1  4.4.2  4.3.2  4.3.1  V  1400  V  720  Pn  Ledum groenlandicum  4  Pn  Gaultheria shallon  5  Pn  Kalmla p o l i f o l l a  1  4.3.2  4.4.2  2.+.1  4.4.2  3.2.+  Ch  Vaccinium v i t i s - i d a e a  2 2  2.2.2  Pn  Vaccinium p a r v i f o l i u m  1  1.+.1  1.+.2 2.+.2  3.+.1 2.+.1  2  6 7  3.2.+  2 8  Ch  1.2.2  Ch  Oxycoccus quadripetalus Empetrum nigrum  2  9  1  4.4.2  2  3.2.2  10  Ch  Linnaea boreal i s  2  3.2.2  11  Pn  Vaccinium uliglnosum  2  1.2.2  12  Pn  Vaccinium ovatum  1  13  Pm  Tsuga h e t e r o p h y l l a  1  14  Pn  Vaccinium oval 1folIum  1 2  15  Pn  Menzlesla f e r r u g l n e a  1  1.+.1 4.3.2 3.2.2  2.+.1 2.1.1 1.2.1  5.3.2  —  5.3.2  5.4.2  4.4  4-5  1.8  1-2  4.2.1  4.2.2  2.2  0-4  2.+.2  3.2  2-4  1.2 1.8  +-2  4.3.2  1.+.1  3.2.2  2.8  1-4  +-2  3.2.2 3.2.1  3.2.2  2.2  1-3  1.3 1.8  1-2  V  250  +.+.1  1.7  +-3  1.2  1-2  V  152  0.4  0-2  1  —  1.2 3.4  1-2  1.4  1-2  0-7  2  — —  2.+.2  1-2  6.4.2  1.2.1 7.4.2  2.2.2  4.2.2  , 2.6  0-4  2  1.2.1  3.2.1  2.0  0-3  1.5  1-2  3.2.2  2.2.1  1.2  0-3  1.7  1-2  3.+.2  0.6  0-3  2  0.4  0-2  1  —  0.2  0-1  1"  —  0.1  0-+  1  0.1  0-+  1  2.+.1 1.+.1 +.+.1 +.+.1  V  60  IV  2060  IV III  310 130  1  100  1  20  li  —  12  1  2  910  C layer 16  H  Carex obnupta  4.3.2  5.4.2  4.3.2  3.4  1-5  1.6  1-2  V  H  Blechnum s p l c a n t  4.3.2  2.3.1  4.+.2  3.4  2-4  1.6  1-2  V  720  18  Ch  Cornus canadensis  3.3.1 3.+.2  1.+.1 4.3.2  3.3.1  17  3.+.2  4.2.2  4.2.2  3.+.2  3.4  3-4  2  —  V  19 20 21 22  H  3.2.2  4.2.2  3.2.2  3.0  2-4  V  1.+.+ 1.+.1  1.+.+ 2.+.2  1.+.+  1-2 0-2  Drosera r o t u n d i f o l i a  1.+.2  1.+.1  1.+.1  0-1  1.5  +-1 1-2 1-2  V IV  H  1.4 1.0 0.8  1.6 0.7 1.5  1-2  2.+.1 1.2.2  2.+.1 2.+.1  3.2.1  G H  Agoserls glauca Maianthemum d i l a t a t u m Gentiana sceptrum  700 520  23  G  Trientails arctica  1.+.1  0.7  0-1  0.9  +-1  24  H  Sangulsorba microcephal  1.+.1 3.2.3  1.8  0-3  2.3  2-3  III  300  25  H H  Coptls a s p l e n l f o l l a  0.6  0-1  1.3  1-2  III  30  26  0.8  0-2  2  —  27  H  Oeschampsia  0.6  0-3  1  —  1  100  28  H  Carex p l u r i f l o r a  0.2  0-1  1  —  1  20  29  G  Lysichitum americanum  2.3.1  0.4  0-2  1  —  1  20  30  G  Habenaria saccata  1.+.2  0.2  0-1  2  1  10  31  G  Pterldlum apuillnum  0.2  0-1  2  1  10  32  Ch  Lycopodlum clavatum  0.2  0-1  1  1  10  33  G  Boschniakia hookerl  0.1  0-+  2  1  2  34  G  Veratrum v i r i d e  0.1  0-+  2  1  2  Calamagrostis  nutkaensl  1.+.1 2.2.2  1.+.2  1.+.1 1.+.2 1.+.1  +.+.+  3.2.2  3.2.2  1.+.1 2.2.2  caespltosa  3.2.1 1.2.1  1.+.2 1.2.1 +.+.2 +:+.2  —  IV IV  II  70 50 40 32  40  115  Life form  Species Significance R III  Sublayer  D layer  4.2.2  35 36'  B  Sphagnum papillosum  h  B  Eurhynchium oreganum  37 38 39  B B B  Sphagnum recurvum Sphagnum capillaceum Pleurozium schreberi  h du 1.1.1 h 5.3.2 h 4.2.2 h 3.1.2 d«  40 41  B B  Dicranum bergerl  42 43  B B  Sphagnum mendocinum Hylocomlum splendens  44  B  45  B  dw h dm h h  Scapania bolanderi  4.2.2 4.2.2 3.2.2 6.3.3 4.3.2 4.2.2 3.2.2 3.2.3  2.1.1  1.1.1 1.2.1  6.2.3 5.3.2  Rhytidiadelphus loreus  h 4.2.2 dw 3.2.2  1.1.1 2.1.1  Cephalozla bicuspidata  h 2.1.2 dw  2.2.1 2.1.1 2.2.1 2.2.1  B B  Dicranum scoparium  48 49 50 51  6 L B  Sphagnum fuscum Cladonla pacifica Plaglothecium undulatum  B  Mnlum glabrescens  52 53 54 55 56  B L L B B  du h  Bazzania ambigua  Sphagnum tenellum Cladonla rangiferina Cladonia crispata Isothecium stoloniferum Polytrichum strictum  4.3.2 5.2.2 4.2.2  2.2.2 1.2.2  3.1.1  1.1.1  1.2 1.0  1.0 0.8  3.1.1 2.1.1  2.1.1 3.2.2  1.1.1 2.1.1  2.1.2 2.1.2 3.3.2  dw h h h h  2.6 1.2 1.4 3.4 3.4 2.6 1.0 1.8 1.2 0.2 3.0 2.4 0.6  5.3.2 4.3.2  dw  46 47  2.2.2  3.2.1  2.2.2 4.3.2  3.2.2 1.2.2 4.3.2 3.2.1  2.2.2 1.1.1  1.2.1  3.3.2 2.1.1  2.1.2  1.2  0.8 0.4  4.3.2 3.2.2  1.4  1.2 0.6 0.2 0.2 0.6 0.4 0.2 0.2 0.2  1.1.1  dw 1.1.1 h h h dw dw  3.3.2 2.2.2 1.1.1  1.1.1  1.1.1  Vigor IH  R  1-4 0-4 0-3 0-6 0-5 0-4 0-3 0-3 0-3 0-1 0-6 0-5 0-3 0-4 0-3 0-3 0-2 0-3 0-2 0-2 . 0-4 0-3 0-2 0-1 0-1 •0-3 0-2 0-1 0-1 0-1  1.8  1-2 1-2 1-2 2-3  1.7 1.5 2 2 2 1 1 1 2 2 1 1 1  2-3 2-4 3-4 0-1 0-3 0-2 0-3 0-3 0-1 0-1 0-1 0-3 0-1  1.4 1.6 2  1-2 1-2  1.3  1.3  2.3 2 1.8 1.5 1.8 1.7 2 2.3 1.7 1 1.3 1.5 1.5  Con-  Cover  stancy  value  V V IV IV IV  —  1-2 1-2 1-2 1-2  450 330 1280 1000 520  IV  160 140  III III  1260 700  III  230  III  140  1-2  III  1-2  111  130 60  IV  —  2-3 1-2 —  1-2 1r2 1-2  1  —  —  II  —  —  11 11  —  II  —  300 200 30 20  —  1 1 1 1  — — — —  '  100 20 10 10 10  E layer Bryophytes S lichens  57  58  E  Frullania nisquallensis  E  Dicranum scoparium  E  Scapania bolanderi  E E  Pleurozl um schreberi Isothecium stoloniferum  E E E  A B C B  3.2.2 3.2.2 4.2.2 1.1.1  C B C C  2.1.2 3.2.2 3.2.2  4.2.2 4.2.2  Total species  43  2.1.1  2.4 3.2 3.8 0.3  2.1.1 3.2.2  2.2.2  1.0  0.4 1.6 1.0 0.2 0.4 0.4 0.6 0.2  2.1.2 2.1.2 1.2.1 1.2.1 1.2.1  36  1.1.1  Number  t  Total cover:  cover  %  19  10  29.7  15.6  6870  26.8  2730  10.7  8  12.5 4116  16.1  V  900  IV  132  —  111  —  II  220 120  —  1  —  2 2 2 2 1 1 1 2 2  — —  — — — — —  •  II II  1 1  35  Pn  Life form  Proportion by Species:  2.2.1  3.2.2  8 1.1.1 3.1.2 1.1.2  Bazzania ambigua  3.2.2 4.2.2 4.2.2 +.1.1  3.1.2  C C C C  Plagiothecium undulatum Antitrichia curtipendul.  2.1.1 3.1.1 4.1.2  8  12.5 1522  5.9  7  10.9 146  0.6  Ch  Pm  6  3  3  9.4  4.7  4.7  3390  13.2  6640  25.9  230  0.9  20 20 100 10  SYNTHESIS TABLE VI.  PINETO-CHAMAECYPARETO-SPHAGKETUM RECUP.VII  Nunber of plots Plot number Plot size (neters) Date  Percent coveraga Vegetation: layer:  »i »z 3 A A  Total  Bl B B C  2  Total  Dh Total  °ds 0 E„ EB EC  Kuaus Decaying lood  Life fora Pa  Pinus contorta  Pa  Chaaaecyparls  3  Pa  Thuja plicata  4  Pa  nootkatensis  Tsuga heterophylla  5  Pn Pa  Vacclnlua parvlfollua Malus d l v e r s l f o l l a  7  Pn Pa  Benzlesla ferruglnea Taxus brevlfDlla  9  2  4  72 100 5/8/ 64  71 100 5/8/ 64  23 100 10/6/ 64  0 50 40 75 65 15 65 15 30 35 65 10 45 80  15 40 35 60 60 25 60 20 35 15 50 15 55 85  70 30 50 80 40 10 45 8 25 50 75 25 60 90  15 70 30 80 60 5 60 20 40 35 75 20 60 90  30 40 25 70 65 8 65 15 30 40 70 10 40 60  60 40  75 25  35 65  60 40  50 50  27 100 15/6/ 64  28 100 15/6/ 64  Species Significance 8 II  Sublayer  A laver  2  3  5  1  1 2 1 2 1 2 3 2  4.0  2.1.2 2.0  6.6.1  5.6.1  8.7.2  5.5.2 2.0  2.4.1  3.0 3.5.2  3 3 2 3 3 2 3  3.0 3.0  1  7.5.2 5.2.2 3.0 3.2.2  6.5.1 5.5.2 5.5.2 7.6.2 2.3.1 1.2.1  5.6.2 3.4.2 3.0 2.0  3.4.2 5.4.2 6.5.1 6.5.2  5.0 3.0 4.0 6.6.1 3.0 4.4.2 5.4.2 7.6.2 3.0  3.5.1 3.5.1 3.0 1.4.2 1.+.2  Vigor H  Constancy  Cover value  V  25ZO  V  2400  V  2300  V 1 1  1890 100 100  1 II  100 20  V  5400  V  700  V  420  R  4.0 2.2 1.2 4.6 1.6 4.0 4.6 3.6 3.0 0.6 0.6 0.6 0.6 0.2 0.2  0-8 0-4 0-4 0-6 0-5 3-5 3-7 0-7 1-6 0-3 0-3 0-3 0-3 0-1 0-1  2 2 1 1 2 2 2 1.3 1.2 2 1 1 2 2 2  1-2 1-2  7.0 3.8 3.4 2.4 2.4 0.6 1.8 0.2 2.8 1.0 1.6 2.0 1.6 0.8 0.6 0.6 0.4 0.4  5-8 3-5 3-4 2-3 1-4 0-3 1-3 0-1 0-4 0-3 0-3 0-5 0-3 0-2 0-3 0-3 0-2 0-Z  2 1.8 1.8 1.6 1.8 2 1.4 1 1.4 1 1.4 2 2 0.6 1 1 1 2  —  1.4 3.8 2.2 2.0 1.6 1.2 0.8 0.6 0.4 0.4 0.4 0.4 0.2 0.2 0.1  1-2 0-5 0-3 0-4 0-4 0-3 0-2 0-2 0-1 0-1 0-2 0-2 0-1 0-1 0-.5  0.7 2 1.3 0.9 2 1.3 3 1 2 0.6 1 2 2 2 1  «-1  — — — — — —  — — — — — —  B layer 9  Pn Pn  10  11 12 13  14  Pn  Gaultheria shallon Vacclnlua parvlfollua Vacclnlua ovatua  Pn  Venzlesla ferruglnea  Pa  Tsuga heterophylla  Pn Pa Ch Pn Pa Pa  Vacclnlua ovaUfollua Thuja plicata Llnnaea boreal Is Ledua groenlandlcua Chamaecyparis nootkatensis Hal us d l v e r s l f o l l a  Ch  2 1 2 1 2 1 2 1 2 \ 1 2 1 1 1 2 2  Vacclnlua vltls-ldaea  5.3.2 3.0 4.3.2 Z.O •4.3.2 3.0 2.4.2  8.5.2 5.3.2 3.3.1 2.0 3.0  3.0  I.O  4.3.2  3.0 3.2.2  I.O 5.3.2 2.1.2 2.0  3.0 2.0 3.3.1  7.5.2 3.+.2 3.4.2 2.0 3.4.2 2.3.1 1.+.1 3.0 Z.+.1 2.4.2 3.+.Z 3.Z.2  8.5.2 3.0 4.4.2 3.0 1.+.2 I.O 4.3.2 3.0 2.3.2 2.4.2  3.4.1 2.0 2.1.2  1-2 1-2 1-2 1-2 —  V  160  IV  600  IV III III II 1 I  150 5Z0 220 40 100 100  1  20  — —  V IV IV III III III III II II II 1 1 1 1  —  1  70 1400 320 400 310 130 40 30 20 20 20 20 10 io 2  — —  1-2 —  1-2 — —  1-2 — — — —  C laver 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29  6 H Ch H H G H Ch G G H G H G H  Halantheaua dlletatua Blechnua splcant Cornus canadensis Care* obnupta Deschaepsla caespltoss Veratrua vtrlde Calaaagrcstis nutkaensls Lycopodlua clavatua Boschnlekla hookerl Llstera cordata Gentiana sceptrua Lyslchltus aaerlcanua Agoserls glauca Trlentalls arctlca Goodyera oblonglfolle  Life fora Proportion by Species:  (iuiber  1 Total cover:  cover I  1.0 5.4.2 3.2.2 3.3.1 3.3.2 2.0  1.+.+ 4.3.2 3.2.1 • 3.3.1 4.3.2 3.3.1  2.0 5.5.2 3.2.1  4.5.2 1.2.2 I.O 2.1.3  •i.o 2.2.1  1.1.1 1.1.2  Z.+.+ 5.5.2 2.0  1.0 1.2.2 1.0  2.1.1 2.4.2 1.2.2 1.0 •.•.1  —  1-2 1-2 —  1-2 — — —  1-2 — —  E  8  H  Pa  33 39.4  24 28.0  7 ,8.1  6 7.0  6 7.0  6 7.0  4 4.7  8752 24.9  7350 20.9  2182 6.2  9220 26.2  6870 19.5  270 0.8  590 1.7  Pn  6  Ch  Life forn  30  G  HylocoBfue tplendens  31  B  fihytldladelphus loreus  32 e 33 B 34 B  Sphagnui recurvui Scapania bolenderl Eurhynchlua oreganua  35 30  G B  Lepldozla reptans Inlua glebrescens  37  B . Plaglotheclue unduletue  38  B  Bazzania anblgua  39  B  Cephalozia blcuspldata  40  G  Hookerla lucens  41  B  Dlplophylluo albicans  42  G  Glepharetona trlchophyllun  43 B 44 B 45 G 45 B 47 B 48 G 49 G 50 e 51 6 52 6 53 B  Spades Significance H R  Sublayer  0 later  Olcranua ecoparlui Olcranua fuscescens Herberts adunca Hypnue clrclnale Isotheclui stolonlferui Pellle eplphylle Eurhynchlua stokesll Pleglochlla aaplenoldes Riccardia sp. Cslypogela trlchonanes Bartraala poatfornle  h de h de h de h de de h de h de h de h de h di h de h di da de de de de h de dl de de dl  4.3.2 3.2.2 4.3.2 4.2.2 5.3.2 4.2.2 1.1.1  5.3.2  3.2.2 1.2.2  2.1.2  4.2.2 3.2.2 5.3.2 3.2.2  2.2.2 1.3.1 4.2.2  3.2.2  2.2.1  3.3.2  3.2.1  2.1.2 3.1.2 1.1.2  1.2.1  1.1.2 2.1.2  2.2.2 3.2.2 1.2.2  2.1.2 6.5.2 4.3.2  1.1.2 1.1.1  2.1.2  2.2.1  3.2.1  2.1.2  2.1.2  2.1.2  2.1.2  M.2  1.1.2  1.1.2  2.1.2 1.1.2'  1.1.2  3.1.1  1.1.2 2.1.2  2.1.2  1.1.1 1.1.2  2.1.2  3.2.2 2.4.3 1.1.2  3.1.1  1.2.2  2.4.3  2.1.1  2.1.2  3.3.3  3.2.2 3.2.1  3.2.2 3.3.2 3.2.2 5.4.3 3.4.2 3.2.2 3.2.1  1.1.1 3.2.1 2.2.1  6.4.3 4.3.3  3.3.3  5.4.2 4.3.2 3.2.2 5.3.3  1.1.2  1.1.1  3.8 2.2 2.6 3.8 3.2 3.4 1.4  1.4 2.0 2.0 1.0 1.1  2.0 0.4 0.8 0.2 1.6 0.4 0.6 1.6 1.8  0.2 0.8 0.6 0.4 0.6 0.6 0.6 0.4 0.2 0.2 0.2 0.2 0.2  0-5 0-4 0-4  2-5 1-5  3-4 0-3 0-3 1-3 0-6 0-4 0-3 0-3 0-2 0-3 0-1 0-2  Con- Caver stancy value  Vigor H  2 1.6 2 2.4  2 2  R —  2-3 1-3 —  1  —  1.7 2 2  —  1.5  1-2 —  1-2  1 1 2 2 2 2 2 2 2.5 2.3 2 2  —  —  0-1  2 3 2 3 3 2 2 2 1 1  0-1  1.7  0-1  0-1 0-6 0-4 0-1 0-2 0-2 0-1  0-3 0-3 0-3 0-2 0-1 0-1 0-1 0.1  1.5  V  1410  V V y V  1220 930 700 330  V IV  170 780  IV  320  1-2 —  — — —  IV  120  IV IV  80 40  in in  780 50  II II  30 20 100 100 100 20 10 10 10 10 10  — — — —  2-3 2-3 — —  1-2  i l f  — —  • — — —  I •  i  t  i i 1  E liter Bryophytes I lichens E  IsothecluB stolonlferui  E  Herberts adunce  E  Frullenla nlequallensls  A B C A  e  54  E  55  E  E  Scepanla bolanderi  Antltrfchle curtlpendula  Hypnue clrcinele  C A B C A B C A G C A  e  56 57 58 59  E  Olcrenua scoparlun  E  Bazzenla anblgua  E  Olplophyllun albicans  E E  Lepldozla reptans Plagfochla aeplenoldee  E E E E E E E E E E  Cephalozia blcuspldata Hookerla lucens Inlua glabrescens Neckera dougl all 1 Plaglotheelul undulatue Cephalozia leaaerilene Cladonia bellldlflore Olcranua fuscescens Eurhynchlm etokesll Netzgerla conjugata  C A B C A B C A C B B C B C C G B C C C  c c  4.3.2 4.3.3 2.2.2 5.2.3 6.3.3 3.2.2 4.2.2 4.2.3 2.1.2 3.2.2 4.2.2  3.3.3 3.2.2 4.3.3 5.4.3 2.2.2 4.2.2 4.2.3 3.2.3 3.2.2  4.3.2  3.2.2  1.1.1  1.2.1  2.2.2 3.2.2  2.2.2 . 2.1.2 2.1.2  2.1.2 3.1.2 2.2.2 2.1.2 2.1.2  2.1.2  7.3.2 8.5.3 5.3.2 1.2.1 5.2.2 5.2.2 3.2.2 4.3.2 4.3.2 2.3.2 2.2.2 4.2.1  4.2.2 3.2.1 2.2.1 2.1.2 1.1.2  2.2.2 4.3.2 4.3.2 4.3.3 4.3.3 4.2.2 4.3.3 4.2.3 3.2.2 3.2.2 2.+.2 4.3.3 2.1.1 2.1.2 2.1.1  4.2.2 5.2.2 3.2.2 2.2.1 3.2.2 3.1.1  4.2.2 3.2.2 2.1.1  2.1.2  1.1.1 2.2.2 2.3.3  2.2.2 2.2.1 1.1.2 1.1.2 1.1.2  1.2.2  2.4 4.6 3.2 2.0 3.4 3.0 3.6 4.4 3.0 1.4 3.2 2.8 0.4 1.4  2.2 1.6 2.2 1.0 1.6  1.4 0.8 0.4 1.4  2.1.1  1.1.2  1.1.1 1.1.2  3.2.1 4.3.2 4.3.2 3.3.2 4.3.2  l.t.1  0.2 0.4 0.2 0.6 0.4 0.2 0.4 0.4 0.4 0.4 0.4 0.2 0.2 0.2 0.2 0.2  3-e  0-5 0-4 0-5 0-5 2-5 4-5 0-4 0-3 3-4 0-4  0-2 0-3 0-3 0-4 1-4 0-3 0-2 0-3 0-2 OJ  0-3 0-1 0-2 0-1  0-2 0-2 0-1 0-2 0-2 0-2 0-2 0-2 0-2 0-1 0-1 0-1 0-1  2.4 1.8 1.4 2.2 3 2 2.2 2.8 1.7 2 2 2 1.7 1.7 1.5  1.4 1.5 1.5  1.7  1-2 2-3 2-3  V  2200  1-2  2-3  —  y  1470  V  1400  V  700  V  620  —  2-3 2-3 1-2 — — —  1-2  1-2 1-2 1-2  1-2 1-2  1-2  2 3 1.7 2 2 1.5 2 2  — —  1  —  2 2 2 3  —  1-2  V  440  y  200  IV  150  — —  1-2 —  —  in  40  n  30 30  II I  i  —  i  —  l i  1  —  2 1 2 2 2  — — — — —  I  i i i 1  20 20 20 20 20 10 10 10 10 10  Vascular el ants E E 60  E  E  E  Vacclnlui pervlfollua Cornus canadensle Polypodlua olycyrrhlzi  Gaultherla shallon  Hanzlesla ferruginea  E  Halinthoeus dllatatua  E  Vacclnlua ovatua Vacclnlua vitis-idaea EnpetruB nlgrui Llnnsea boreal la thuja plicata Tauge heterophylle  £.  E E E E  A B C B C A. G C A G C A G C B  c cB  2.1.1 3.1.1 2.1.2 3.+.1 2.Z.2  1.2.2  3.2.1 3.3.1  2...1 2...1 2.+.1 1.».1 1.*.1 2.3.2 2.-f.2  l.t.t  1.2.1 3.2.1 2.*.1 3.4.2 4.4.2 2.+.1  2.*.1  +.+.+  3.+.1  S A  2.».1 1.2.1 1.+.1 1.2.1 1.2.1  2.*.2 2.*.2 2..-.1 1.2.2  1.2.1  1.+.2 2.*.2 2.2.1  1...1 1.<v>  46  41  41  40  1.4 2.6 0.8 1.6 1.0 1.2 0.2 1.2  *  2.+.1 1.2.2  0.6  1.4  1.*.2  . 3.3.1  6  Total speclee  1.».1 3.4.2 4.4.2  36  1.4  0.2 0.8 0.6 0.1 0.4 0.6 0.6 0.4 0.2 0.2 0.2  0-2 0-3 2-4 0-2 0-3 0-2 0-3 0-1 0-3 0-4 0-1 0-2 0-2 0-* 0-2 0-3 0-3 0-2 0-1 0-1 0-1  1 1.7 1.4  1.2 1 1.5 1.7 1 1.3 2 1.5 2 1.5 1 +  1  1-2  1-2 *-2 —  1-2  V  360  V  160  1-2  V  150  1-2  IV  330  in  50  — —  1-2 —  1-2  — — —  1 1 1 2  —  1  —  +  —  — —  II I  l  I  1 I  i  22 100 100 20 10 10 10  118  APPENDIX III Soils data  i  Association:  Caricetum p l u r i f l o r a e  P l o t no.  40  44  19/8/64  19/8/64  max. d e p t h (cm)  40  30  45  main c o n e , t o (cm)  25  15  10  Date c o l l e c t e d  50 20/8/64  Root d i s t r i b u t i o n  Chemical  Analysis  Plot  Depth  no.  (cm)  40  5-10  1247.5  20-25 30-35  Soil  50  44  Moisture  Total pH  Exchangeable  Nitrogen  C.E.C.  Na  (?)  (me?)  (me?)  3.4  1.61  2.6  760.8  3.9  2.80  85.1  4.4  0.001  5-10  223.3  15-20  1019.4  30-35 40-45  Adsorbed  Base  Mg  Phosphate  (me?)  (me?)  (me?)  (ppm)  (?)  5.70  1.04  8.60  4.65  5.3  9.3  85  1.45  0.28  1.50  1.25  0.6  5.3  42  0.70  0.05  0.80  0.37  0.9  4.6  4.1  66  0.90  0.09  0.56  0.20  3.9  81  1.05  0.27  2.20  1.73  96.2  4.4  45  0.74  0.05  0.40  0.08  35.1  4.0  10-15  732.5  3.5  15-20  94.7  4.3  25-30  52.2  4.7  Association:  cations Ca  (?)  Oxycocceto-Sphagnetum  P l o t no.  Saturation  2.7 17.6 2.8  papillosae  18  19  15/8/64  16/8/64  max. depth (cm)  57  50  main c o n e , t o (cm)  25  20  Date c o l l e c t e d  K  Root d i s t r i b u t i o n  Chemical  Analysis  Plot  Depth  no.  (cm)  (?)  10-1.5  1073.3  3.7  247  22-25  796.7  4.1  71  28-32  281.8  3.'9  35-40  144.8  52-57  55.5  6-11  1408.6  3.4  15-20  1065.0  3.5  Soil  19  Moisture  Total pH  Exchangeable  Nitrogen  C.E.C.  Na  (?)  (me?)  (me?)  K  Adsorbed  cations Ca  Mg  Phosphate (ppm)  Base Saturation (?)  (me?)  (me?)'  (me?)  4.55  1.26  4.58  4.80  3.0  2.05  0.76  0.95  1.30  1.4  7.1  60  0.85  0.16  0.51  0.49  1.2  3.4  4.0  63  0.82  0.12  0.27  0.19  0.9  2.2  4.8  42  0.70  0.04  0.26  0.05  1.2  2.5  1.55  276  6.28  1.49  6.40  6.25  7.4  2.24  73  1.75  0.57  1.17  1.55  6.9  6.1  30-35  82.7  3.8  0.16  49  0.69  0.07  0.36  0.14  2.6  45-50  134.6  4.6  0.36  61  0.80  0.08  0.44  0.22  2.5  Association:  Scirpeto-Sphagnetum  P l o t no.  mendocini 93  Date c o l l e c t e d  101  6/9/64  6/9/64  55  65  15  20  Root d i s t r i b u t i o n max.  depth.(cm)  main c o n e . t o (cm) Chemical  Analysis  Plot  Depth  no.  (cm)  Soil  93  101  1M o i s t u r e  Total pH  (?)  Exchangeable K  Nitrogen  C.E.C,  •Na  (?)  (me?)  (me?)  cations  Adsorbed  Base  Ca  Mg  Phosphate  (me?)  (me?)  (me?)  (ppm)  (?) 11.1  Saturati  5-10  820.6  4,1  1,79  61  1.55  0.47  2.26  2.10  0.4  20-25  557.0  4.2  1.88  65  1.45  0.41  1.55  1.45  0.5  7.5  25-30  '5P.7  4.7  0.16  49  0.63  0.08  0.50  0.14  0.9  2.8  50-55  36.6  5.4  34  0.68  0.04  0.50  0.06  0.9  3.8  5-10  985,5  3.4  244  4.20  1.22  5.80  5.20  6.7  15-20  280.8  4.4  65  2.18  0,36  5.40  1.95  15.2  25-30  3.9  63  1.75  0.12  5.35  1.68  14.1  60-65  4.2  44  0.92  0.09  1.75  0.41  7.2  Association:  Ledeto-Sphagnetum  Plot no.  capillacei 26  Date c o l l e c t e d  68  15/8/64  19/8/64  Root d i s t r i b u t i o n max. depth (cm)  65  95  main c o n e , t o (cm)  40  45  Chemical  Analysis  Plot  Depth  no.  (cm)  (?)  Soil  26  68  Moisture  Total pH  Nitrogen (?)  Exchanqeabl e c a t i o n s C.E.C.  Na  (me?)  (me?)  K  Adsorbed  Base  Ca  Mg  Phosphate  (me?)  (me?)  (me?)  (ppm)  (?) 7.3  Saturati  6-11  1376.3  3.7  3.93  276  6.24  1,27  6.41  6.20  0.6  22-27  1441.2  3.4  1.59  279  6,40  1.29  6.60  5.65  2.5  7.1  30-35  ' 348.2  4.1  0.08  60  1.15  0.12  1.08  0.94  0.9  5.5  45-50  62.1  5.1  0.02  42  0.60  0.04  0.47  0.15  0.8  3.0  60-65  44.9  4.5  0.07  32  0.59  0.03  0.75  0.10  2.2  4.6  5-10  1390.9  3.3  253  4.40  1.76  9.40  8.50  9.5  30-35  1131.0  3.8  82  1,65 •  0.42  3,95  2.05  9.8  40-45  1243.0  4.4  70  2,08  0.21  2.10  1.41  8,3  60-65  1230.0  3.7  157  4,05  0.28  6.40  1.75  7.9  65.8  4.6.  42  0.78  0.05  1.36  0.20  5,7  95-100  Association:  Pineto-Sphagnetum  P l o t no.  capillacei  sphagnoosum p a p i l l o s i  75  Date c o l l e c t e d  78  24/8/64  T h i c k n e s s of humus (cm)  24/8/64  20  25  Root d i s t r i b u t i o n max. depth (cm)  105  main c o n e , t o (cm)  80-90  25  25  Chemical'.'Analysis Plot  Depth  No.  (cm)  75  Soil  Total  Moisture  Nitrogen  Na  (?)  (me?)  (me?)  Adsorbed  cations  Base  Ca  Mg  Phosphate  (me?)  (me?)  (me?)  (ppm)  (?)  K  Saturati  15-20  1061.5  3.9  1.61  300  4.05  1.24  6.80  4.40  0.5  5.5  32-37  1824.8  4.2  1.23  68  1.75  0.27  2.54  1.43  0.7  8.8  55-60  628.5  4.6  0.22  66  1.15  0.07  2.95  0.88  0.2  7.7  60-75  318.4  4.6  0.16  70  0.90  0.08  5.05  0.82  0.4  9.8  51.2  5.4  0.04  34  0.64  0.04  0.76  0.08  3.0  4.5  10-15  1400.0  3.4  256  4.90  0.91  7.80  5.95  7.6  20-25  979.3  3.9  70  1.45  0.38  1.15  1.43  6.3  4.2  58  2.18  0.21  0.85  1.03  7.4  61  1.15  0.08  0.62  0.46  3.8  56  0.63  0.04  0.40  0.07  2.0  105-110 78  pH  (?)  Exchangeable C.E.C.  40-45  781.8  • 60-65  557.3  3.9  75-80  70.8  4.5  Association:  Pineto-Sphagnetum c a p i l l a c e i  P l o t no.  chamaecyparosum 13  11  Date c o l l e c t e d  nootkatensis  13/8/64  20/8/64  Root d i s t r i b u t i o n max. depth (cm)  95  70  main c o n e , t o (cm)  30  23  Chemical  Analysis  Plot  Depth  no.  (cm)  (?)  Soil  13  11  Moisture  Total pH  Nitrogen (?)  Exchangeable C.E.C. (me?)  Na (me?)  c a t i ons  Adsorbed  Base  Ca  Mg  Phosphate  (me?)  (me?)  (me?)  (ppm)  (?)  K  Saturati  4-12  434.2  3.7  285  4.24  1.87  9.40  8.30  1.1  8.4  19-23  61.6  4.5  43  0.55  0.06  0.22  0.04  1.5  2.0  25-29  69.2  4.5  51  0.75  0.06  0.15  0.06  1.1  2.0  30-35  71.0  4.5  51  0.55  0.04  0.18  0.11  2.4  1.7  40-45  46.0  4.5  36  0.51  0.04  0.11  0.04  1.7  1.9  65-70  40.4  5.2  26  0.49  0.04  0.26  0.03  2.4  3.2  1097.0  3.7  275  5.50  2.14  4.40  4.30  5.9  3.48  0.87  3.60  1.95  7.5  3-8 20-25 35-40 65-70 90-95  687.5 442.8 208.7 27.2  3.6 3.9 4.4 4.5  219 216  4.05  0.18  2.82  0.25  3.4  36  0.78  0.08  2.05  0.21  8.7  0.08  1.82  0.25  8.1  36  0.75  123  Association:  Pineto-Sphagnetum  capillacei  Plot no.  vacciniosum v i t i s - i d a e a e  1  24  30  12/8/64  12/8/64  15/8/64  5  9  10  max. depth (cm)  50  75  75  main c o n e , t o (cm)  10  10  15  Date c o l l e c t e d T h i c k n e s s of humus (cm) Root d i s t r i b u t i o n  Chemical  Analysis Soil  Plot  Depth  no.  (cm)  1-  0-5  Moisture  Total pH  (?)  Exchangeable  Nitrogen  C.E.C.  (?)  (me?)  Na (me?)  K (me?)  cations Ca  Adsorbed Mg  (me?)  (me?)  Phosphate  Base Saturation  (ppm)  (?)  653.3  3.5  1.40  263  6.03  2.02  8.03  9.00  5.4  9.5  15-20  94.3  3.7  0.21  53  0.75  0.07  0.28  0.10  0.9  2.3  25-30  65.3  4.9  0.002  43  0.59  0.05  0.33  0.15  0.3  2.6  45-50  50.7  4.7  0.002  42  0.59  0.05  0.32  0.04  1.7  2.4  0-8  554.9  3.4  295  4.45  1.77  5.65  6.50  6.2  30-35  37.3  4.6  49  0.60  0.04  0.27  0.03  1.9  55-60  39.4  4.8  34  0.60  0.03  0.59  0.05  3.7  70-75  29.2  4.8  36  0.63  0.04  0.70  0.07  4.0  455.4  3.7  3-9 8-12  77.8  4.2  22-27  44.5  4.7  Association: Plot  Pineto-Sphagnetum  capillacei  vacciniosum p a r v i f o l i i  no.  31  Date c o l l e c t e d  13/8/64  12/8/64  15  18  T h i c k n e s s of humus (cm)  32  Root d i s t r i b u t i o n max.  depth (cm)  main c o n e , t o (cm) Chemical  Analysis  Pl ot  Depth  Moisture  no.  (cm)  (?)  Soil  31  70 20  Total pH  0-8  32  75 15  Exchangeable K  Nitrogen  C.E.C.  Na  (?)  (me?)  (me?)  cations  Adsorbed  Base  Ca  Mg  Phosphate  Saturati  (me?)  (me?)  (me?)  (ppm)  (?)  5.1  3.5  10-15  446.5  4.2  0.33  200  3.95  0.98  2.80  2.45  2.1  20-25  52.2  4.1  0.07  52  0.54  0.05  0.04  0.05  1.7  1.3  70-75  30.5  5.4  0.01  31  0.74  0.05  0.46  0.03  2.6  4.1  3-8  603.2  3.4  222  4.05  2.15  3.64  6.45  7.3  12-18  451.9  4.0  70  1.25  0.43  0.50  0.97  4.5  20-25  82.1  4.4  38  0.45  0.06  0.18  0.08  2.0  34-37  31.4  4.6  37  0.53  0.05  0.22  0.03  2.2  65-70  27.3  4.4  24  0.64  0.05  0.64  0.04  5.7  Association:  Pineto-Chamaecypareto-Sphagnetum  P l o t no.  recurvi  27  Date c o l l e c t e d  72  13/8/64  T h i c k n e s s of humus (cm)  19/8/64  10  15  max. depth (cm)  65  85  main c o n e , t o (cm)  15  30  Root d i s t r i b u t i o n  Chemical  Analysis  Plot  Depth  no.  (cm)  (?)  Soil  72  27  Moisture  Total pH  Exchangeable  Nitrogen  C.E.C.  Na  (?)  (me?)  (me?)  (me?)  K  cations  Adsorbed  Base  Ca  Mg  Phosphate  (me?)  (me?)  (ppm)  (?)  Saturati  5-10  270.5  4.5  0.88  288  4.80  2.17  37.80  7.25  1.3  18.1  30-35  417.7  3.3  0.87  185  4.10  1.35  9.80  6.40  2.4  11.7  45-50  75.9  4.3  0.03  49  0.63  0.06  0.10  0.11  2.1  1.8  55-60  46.8  4.5  0.06  39  0.71  0.06  0.28  0.10  1.4  2.9  80-85  44.0  5.4  32  0,75  0.05  0.46  0.09  2.0  4.2  0-5  262.7  4.0  295  4.45  0.37  10.20  7.05  6-10  356.6  3.9  301  6.95  1.87  9.60  9.20  9.2  26-30  40.6  4.4  41  0.75  0.08  0.72  0.85  5.9  35-40  48.5  4.9  49  0.84  0.06  0.28  0.10  2.6  60-65  38.8  5.1  42  0.64  0.06  0.33  0.06  2.6  7.7  APPENDIX IV Radiocarbon dating  12.$; 24 Blackstone Street, Cambridge, Mass. 02139 Telephone TRowbridge 6-3691 2 August 1965 boratoried, REPORT DF ANALYTICAL WORK RADIOCARBON AGE DETERMINATION  nc.  Your Reference No,  Our Sample No.: GXO^S Sample Name: AGE  Vancouver I s l a n d  m 3SQ ±90 c-m  Letter: 18 May 1 9 6 5  peat,  y e a r s 8. P.  (1560  A. D.)  Description:  Silty  Location:  West c o a s t o f V a n c o u v e r I s l a n d , Canada.  Occurrence:  B a s a l p e a t o v e r l y i n g c o m p a c t 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 over g l a c i o - m a r i n e t i l l . D e p t h t o p e a t 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 .  .Collected:  Not  Submitted by:  G. E. R o u s e , U n i v e r s i t y Canada.  Comments:  The s a m p l e u a s t r e a t e d w i t h h o t d i l u t e H C l , h o t NaOH, and a g a i n w i t h H C l , t o remove a l l s o l u b l e m a t e r i a l s prior to analysis. ' j  Notes:  peat. British  Columbia,  given. of British  Columbia,  Vancouver,  This date i s based upon the Libby half l i f e (5570 years) f o r C The error stated i s ± 1«>" as judged by the a n a l y t i c a l data alone. Our modern standard i s 95% of the a c t i v i t y of N. B. S. Oxalic Acid.  

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