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UBC Theses and Dissertations

Classification and successional relationships of some bog and forest ecosystems near Prince Rupert, British… Banner, Allen 1983

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C L A S S I F I C A T I O N AND SUCCESSIONAL RELATIONSHIPS OF SOME BOG AND FOREST ECOSYSTEMS NEAR PRINCE RUPERT, B R I T I S H COLUMBIA b y A l l e n B a n n e r B . S c , U n i v e r s i t y o f B r i t i s h C o l u m b i a , 1977 A THESIS SUBMITTED I N PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES ( F a c u l t y o f F o r e s t r y ) We a c c e p t t h i s t h e s i s a s c o n f o r m i n g t o t h e r e q u i r e d s t a n d a r d THE UNIVERSITY OF B R I T I S H COLUMBIA A p r i l 1983 © A l l e n B a n n e r , 1983 I n p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r an a d v a n c e d d e g r e e a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e a n d s t u d y . I f u r t h e r a g r e e t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by t h e h e a d o f my d e p a r t m e n t o r by h i s o r h e r r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . D e p a r t m e n t o f The U n i v e r s i t y o f B r i t i s h C o l u m b i a 1956 Main M a l l V a n c o u v e r , C a n a d a V6T 1Y3 D a t e Apul 9-1 J fi£Z (.3/81) ABSTRACT A c l a s s i f i c a t i o n o f t h e bog and f o r e s t e c o s y s t e m s a l o n g f o u r t o p o g r a p h i c t r a n s e c t s n e a r P r i n c e R u p e r t , B r i t i s h C o l u m b i a i s d e v e l o p e d and d e s c r i b e d . The s t u d y a r e a i s c l a s s i f i e d w i t h i n t h e C o a s t a l - C e d a r s - P i n e - Hemlock zone (CCPH) a c c o r d i n g t o t h e B r i t i s h C o l u m b i a M i n i s t r y o f F o r e s t ' s B i o g e o c l i m a t i c c l a s s i f i c a t i o n s y s t e m . D e t a i l e d v e g e t a t i o n , s o i l , s i t e and m e n s u r a t i o n d a t a were c o l l e c t e d f r o m a t o t a l o f 40 r e l e v e s t h a t were c h o s e n a c c o r d i n g t o t h e B r a u n - B l a n q u e t a p p r o a c h o f s a m p l i n g d i s t i n c t , homogeneous e x a m p l e s o f c l i m a x v e g e t a t i o n . V e g e t a t i o n d a t a were a n a l y z e d u s i n g s u b j e c t i v e t a b u l a r c o m p a r i s o n , t o g e t h e r w i t h t h r e e o b j e c t i v e o r d i n a t i o n t e c h n i q u e s ; p o l a r o r d i n a t i o n , p r i n c i p a l c o m p o n e n t s a n a l y s i s and r e c i p r o c a l a v e r a g i n g . T a b u l a r c o m p a r i s o n and o r d i n a t i o n p r o v e d t o be c o m p l e m e n t a r y i n d e v e l o p i n g and e v a l u a t i n g t h e c l a s s i f i c a t i o n , and i n d i s p l a y i n g r e l a t i o n s h i p s w i t h i n and among t h e community t y p e s . I t was r e c o g n i z e d , h o w e v e r , t h a t s u b j e c t i v e r a t h e r t h a n s y s t e m a t i c s a m p l i n g p l a c e s l i m i t s on t h e i n t e r p r e t a t i o n o f t h e o r d i n a t i o n s . The v e g e t a t i o n , s o i l s and l a n d f o r m s o f s i x co m m u n i t y t y p e s m a k i n g up t h e b l a n k e t bog - u p l a n d f o r e s t c o m p l e x o f t h e s t u d y a r e a a r e d e s c r i b e d i n d e t a i l and v e g e t a t i o n - e n v i r o n -ment r e l a t i o n s h i p s a r e d i s c u s s e d . A c o o l , p e r h u m i d , r a i n y , h y p e r o c e a n i c c l i m a t e i s c o n s i d e r e d t h e o v e r r i d i n g f a c t o r influencing vegetation and s o i l development. Important associated c o n t r o l l i n g factors are; s o i l drainage, through i t s influence on s o i l water regimes and nutrient turnover; the two important s o i l forming processes of organic matter accumu-l a t i o n and podzolization; landform and topography, through t h e i r e f f e c t on s o i l drainage; and bedrock geology, through i t s e f f e c t on s o i l nutrient regimes and drainage. Vegetation - environment relationships on the north coastal mainland appear si m i l a r to those described i n South-east Alaska and on the southern and western Queen Charlotte Islands. There are, however, some s i g n i f i c a n t f l o r i s t i c and geologic d i s t i n c t i o n s among these geographic areas. The oceanic blanket bog ecosystems of the north P a c i f i c coast show many relationships with those i n the western B r i t i s h Isles and, to a lesser extent, with those of eastern Newfoundland. The h i s t o r i c a l development of a bog woodland i s recon-structed using pollen analysis, peat stratigraphy and C-14 dating. The succession spans 8700 +/- 210 years i n the following sequence: Pinus contorta - Alnus rubra - ferns/ pioneer a l l u v i a l forest; Picea sitchensis - Alnus rubra -Tsuga heterophylla - (Thu j a pli c a t a) - Lysichiton americanum -ferns/moist productive a l l u v i a l forest on Regosols; Thuja  p l i c a t a - Chamaecyparis nootkatensis - Tsuga heterophylla -Pinus contorta/scrub forest on peaty mineral s o i l s ; Pinus  contorta - Chamaecyparis nootkatensis - ericaceous shrubs -Sphagnum spp. /bog woodland on organic s o i l s . This ecosystem i v sequence i s c o r r e l a t e d with changes i n p a l e o c l i m a t e r e p o r t e d f o r southwestern B r i t i s h Columbia, and may a l s o be a s s o c i a t e d w i t h edaphic f a c t o r s such as changing drainage p a t t e r n s , the formation of cemented s o i l h o r i z o n s and the accumulation of t h i c k o r g a n i c s u r f a c e h o r i z o n s . The p a l y n o l o g i c a l , f l o r i s t i c and edaphic evidence f o r s u c c e s s i o n from f o r e s t t o muskeg i s c o n t r a s t e d with other edaphic evidence f o r s u c c e s s i o n from muskeg to f o r e s t i n north c o a s t a l B r i t i s h Columbia and Southeast A l a s k a . I i n t e r p r e t the r e g i o n a l v e g e t a t i o n and s o i l s as a dynamic complex of ecosystems l i n k e d s u c c e s s i o n a l l y through a c l i m a t i c a l l y s e n s i t i v e pathway. L o c a l s i t e f a c t o r s such as topography, landform, drainage and n u t r i e n t regime are important secondary f a c t o r s c o n t r o l l i n g the d i r e c t i o n of s u c c e s s i o n . The problem of d e f i n i n g the c l i m a t i c climax v e g e t a t i o n along the n o r t h P a c i f i c coast, where s u c c e s s i o n has proceeded i n the absence of major d i s t u r b a n c e s such as f i r e f o r thousands of years, i s addressed. I conclude t h a t the scrub upland f o r e s t s of Thuja p l i c a t a , Chamaecyparis noo t k a t e n s i s and Tsuga h e t e r o p h y l l a t h a t predominate on the moderate slopes throughout the study area, appear to b e s t express the macroclimate and should t h e r e f o r e be c o n s i d e r e d the c l i m a t i c climax. However, t h i s may not apply t o the exposed outer coast where open b l a n k e t bog and bog woodland dominate the landscape, even on s l o p e s . V TABLE OF CONTENTS ABSTRACT i i LIST OF TABLES i x LIST OF FIGURES X LIST OF APPENDICES x v i ACKNOWLEDGEMENTS. x v i i CHAPTER 1 INTRODUCTION 1 1.1 E c o l o g i c a l I n v e s t i g a t i o n s Along North C o a s t a l B r i t i s h Columbia 2 1.2 Th e s i s O b j e c t i v e s 4 1.3 T h e s i s O u t l i n e 5 CHAPTER 2 BOG TERMINOLOGY : A LITERATURE REVIEW 7 2.1 I n t r o d u c t i o n 7 2.2 Bog Genesis and Morphology.... 8 2.3 N u t r i e n t R e l a t i o n s h i p s 12 CHAPTER 3 LOCATION AND DESCRIPTION OF THE STUDY AREA 16 3.1 General D e s c r i p t i o n of the C o a s t a l Cedars - Pine -Hemlock B i o g e o c l i m a t i c Zone 16 3.2 Geographic L o c a t i o n of the Study Transects 17 3.3 Physiography and Geology 21 3.4 C l i m a t e 22 CHAPTER 4 DESCRIPTION AND CLASSIFICATION OF COMMUNITY TYPES...25 4.1 I n t r o d u c t i o n 2 5 4.2 F i e l d Methods 26 4.3 Methods of Data A n a l y s i s 28 4.3.1 Tabular Comparison 29 4.3.2 O r d i n a t i o n Techniques 31 4.3.2.1 Pol a r O r d i n a t i o n (PO) 33 4.3.2.2 P r i n c i p a l Components A n a l y s i s (PCA) 33 4.3.2.3 R e c i p r o c a l Averaging (RA) 3 5 4.3.2.4 Data Adjustments t o Produce the O r d i n a t i o n s . . . . 36 v i 4.4 R e s u l t s 37 4.4.1 T a b u l a r C o m p a r i s o n and O r d i n a t i o n R e s u l t s 37 4.4.2 D e s c r i p t i o n o f Community T y p e s 43 4.4.2.1 E r i c a c e o u s s h r u b - T r i c h o p h o r u m c e s p i t o s u m -Sphagnum s p p . - C l a d o n i a s p p . - S i p h u l a c e r i t i t e s open b o g s 43 M a j o r d i s t i n c t i o n s i n bog v e g e t a t i o n and s u r f a c e f e a t u r e s 45 V e g e t a t i o n and s u r f a c e f e a t u r e s o f t h e f l a t bogs 47 V e g e t a t i o n and s u r f a c e f e a t u r e s o f t h e b l a n k e t b o g s 50 S o i l s and l a n d f o r m s o f t h e open bog c o m m u n i t i e s 6 2 R e l a t i o n s h i p s among t h e r e l e v e s 66 4.4.2.2 Empetrum n i g r u m - V a c c i n i u m u l i g i n o s u m  R h a c o m i t r i u m l a n u g i n o s u m - C l a d o n i a s p p . o u t c r o p c o m m u n i t i e s 68 V e g e t a t i o n 68 S o i l s and l a n d f o r m s 70 4.4.2.3 P i n u s c o n t o r t a - C h a m a e c y p a r i s n o o t k a t e n s i s -( T h u j a p l i c a t a ) - G a u l t h e r i a s h a l l o n - F a u r i a -c r i s t a - g a l l i - P l e u r o z ium s h r e b e r i - Sphagnum s p p . bog w o o d l a n d s 70 V e g e t a t i o n 72 S o i l s and l a n d f o r m s 77 R e l a t i o n s h i p s among t h e r e l e v e s 78 4.4.2.4 T h u j a p l i c a t a - C h a m a e c y p a r i s n o o t k a t e n s i s -T s u g a h e t e r o p h y l l a - V a c c i n i u m ( p a r v i f o l i u m -o v a l i f o l i u m - a l a s k a e n s e ) - G a u l t h e r i a s h a l l o n -C o r n u s u n a l a s c h k e n s i s - R h y t i d i a d e l p h u s l o r e u s -H y l o c o m i u m s p l e n d e n s - Sphagnum g i r g e n s o h n i i c l i m a t i c c l i m a x u p l a n d f o r e s t s 79 V e g e t a t i o n 82 S o i l s and l a n d f o r m s 87 R e l a t i o n s h i p s among t h e r e l e v e s 91 v i i 4.4.2.5 T h u j a p l i c a t a - P i c e a s i t c h e n s i s - T s u g a h e t e r o p h y l l a - A l n u s r u b r a - ( A b i e s  a m a b i l i s ) - L y s i c h i t o n a m e r i c a n u m -G y m n o c a r p i u m d r y o p t e r i s - R h i z o m n i u m  g l a b r e s c e n s - P e l l i a n e e s i a n a a l l u v i a l f o r e s t s 92 V e g e t a t i o n 92 S o i l s and l a n d f o r m s 96 R e l a t i o n s h i p s among t h e r e l e v e s 97 4.4.2.6 A b i e s a m a b i l i s - T s u g a h e t e r o p h y l l a -T h u j a p l i c a t a - V a c c i n i u m ( a l a s k a e n s e -p a r v i f o l i u m ) O p l o p a n a x h o r r i d u s -G y m n o c a r p i u m d r y o p t e r i s - R h y t i d i a d e l p h u s  l o r e u s - R h i z o m n i u m g l a b r e s c e n s P l a g i o t h e c i u m u n d u l a t u m u p l a n d f o r e s t s 98 V e g e t a t i o n 98 S o i l s a n d l a n d f o r m s 105 R e l a t i o n s h i p s among t h e r e l e v e s 107 4.4.3 S e p a r a t e A n a l y s i s o f U n d e r s t o r y and O v e r s t o r y . . . . 1 0 7 4.4.4 S i t e I n d i c e s a n d G r o w t h C l a s s e s I l l 4.5 D i s c u s s i o n 113 4.5.1 O r d i n a t i o n a n d C l a s s i f i c a t i o n 113 4.5.2 V e g e t a t i o n - E n v i r o n m e n t R e l a t i o n s h i p s 115 4.5.3 C o m p a r i s o n w i t h O t h e r S t u d i e s i n B r i t i s h C o l u m b i a and S o u t h e a s t A l a s k a 120 4.5.4 R e l a t i o n s h i p s w i t h O t h e r O c e a n i c R e g i o n s 125 CHAPTER 5 HISTORICAL DEVELOPMENT OF A BOG WOODLAND: A PALYNOLOGICAL STUDY 134 5.1 O b j e c t i v e s o f t h e S t u d y 134 5.2 L o c a t i o n a n d D e s c r i p t i o n o f t h e S t u d y S i t e 134 5.3 M e t h o d s 139 5.3.1 F i e l d M e t h o d s 139 5.3.2 L a b M e t h o d s 139 v i i i 5.3.3 Limitations of the Methodology 140 Pollen p r o f i l e analysis 140 Radiocarbon dating techniques 140B 5.4 Results 140C 5.4.1 Pollen and Peat Stratigraphy 140C Zone HM-1 141 Zone HM-II 141 Zone HM-III 142 Zone HM-IV 143 5.4.2 Absolute Pollen Concentrations 143 5.5 Discussion 144 5.5.1 Pollen and Peat Stratigraphy 144 5.5.2 Absolute Pollen Concentrations 155 5.5.3 Summary of the Bog Woodland Successional Development 156 5.5.4 Comparison with Other Studies of P a c i f i c Coast Bog Development 158 5.5.5 Studies of Forest - Bog Succession in Other Regions 160 5.5.6 Paleoclimatic Interpretations 162 CHAPTER 6 ECOSYSTEM DYNAMICS 165 6.1 Contrasting Evidence for Bog/Forest Succession 165 6.2 The Succession Complex and the Climatic Climax Ecosystem 167 CONCLUSIONS 169 LITERATURE CITED 171 APPENDIX 1 184 APPENDIX II 185 APPENDIX III 192 APPENDIX IV 194 APPENDIX V 201 APPENDIX VI 204 APPENDIX VII 228 APPENDIX VIII 231' LIST OF TABLES Table 1. V e g e t a t i v e f e a t u r e s and c h a r a c t e r s p e c i e s f o r the e r i c a c e o u s shrub - Trichophorum cespitosum - Sphagnum spp. - C l a d o n i a spp. - S i p h u l a c e r a t i t e s open bog community type 46 Table 2. V e g e t a t i v e f e a t u r e s and c h a r a c t e r s p e c i e s f o r the Pinus c o n t o r t a - Chamaecyparis noot- k a t e n s i s - (Thuja p l i c a t a ) - G a u l t h e r i a s h a l l o n -F a u r i a c r i s t a - g a l l i - Pleuroz ium s c h r e b e r i - Sphagnum spp. bog woodland community type 73 Table 3. V e g e t a t i v e f e a t u r e s , stand c h a r a c t e r i s t i c s and c h a r a c t e r s p e c i e s f o r the Thuja p l i c a t a -Chamaecyparis n o o t k a t e n s i s - Tsuga h e t e r o p h y l l a -Vaccinium spp. - G a u l t h e r i a s h a l l o n - Cornus  unalaschkensis - R h y t i d i a d e l p h u s l o r e u s - Hylocomium  splendens - Sphagnum g i r g e n s o h n i i upland f o r e s t community type 83 Table 4. V e g e t a t i v e f e a t u r e s , stand c h a r a c t e r i s t i c s and c h a r a c t e r s p ecies f o r the Thuja p l i c a t a - P i c e a  s i t c h e n s i s - Tsuga h e t e r o p h y l l a - Alnus rubra -(Abies a m a b i l i s ) - L y s i c h i t o n americanum - Gymnocarpium  d r y o p t e r i s - Rhizomnium q l a b r e s c e n s - P e l l i a neesiana a l l u v i a l f o r e s t community type 94 Table 5. V e g e t a t i v e f e a t u r e s , stand c h a r a c t e r i s t i c s and c h a r a c t e r s p e c i e s f o r the Abies a m a b i l i s - Tsuga hetero- p h y l l a - Thuja p l i c a t a - Vaccinium spp. - Oplopanax  h o r r i d u s - Gymnocarpium d r y o p t e r i s - R h y t i d i a d e l p h u s  l o r e u s - Rhizomnium glabre s c e n s - P l a g i o t h e c i u m  undulatum upland f o r e s t community type 101 Table 6. Mensuration data from sample t r e e s of the four f o r e s t and woodland community types 112 X LIST OF FIGURES Figure 1. P r o f i l e of blanket mire complex showing a l l the component peat types. After Moore and Bellamy (1974) 11 Figure 2. Location of the four study transects near Prince Rupert, B r i t i s h Columbia 18 Figure 3. Oblique a i r photo of the study area; looking northwest over Rainbow Lake 19 Figure 4. Climatic diagram for Prince Rupert air p o r t 23 Figure 5. Polar ordination of releves 38 Figure 6. PCA (standardized) ordination of releves, axes l(x) and 2(y) 39 Figure 7. RA ordination of releves, axes l(x) and 2(y) 40 Figure 8. PCA (standardized) species ordination; axes l(x) and 2(y) 42 Figure 9. Blanket bog - upland forest complex near Rainbow Lake 44 Figure 10. Pinus contorta - ericaceous shrub - T r i -chophorum cespitosum - Sphagnum spp. - Cladonia spp.-Siphula cer a t i t e s open blanket bog community; releve 7 44 Figure 11. Ericaceous shrub - Trichophorum cespitosum - Sphagnum spp. open f l a t bog community; releve 6 48 x i F i g u r e 12. E r i c a c e o u s s h r u b - T r i c h o p h o r u m c e s p i t o s u m - Sphagnum s p p . o p e n f l a t bog c o m m u n i t y ; r e l e v e 14 48 F i g u r e 13. P i n u s c o n t o r t a - C h a m a e c y p a r i s n o o k a t e n s i s - e r i c a c e o u s s h r u b - T r i c h o p h o r u m c e s p i t o s u m -- Sphagnum s p p . - C l a d o n i a s p p . - S i p h u l a c e r a t i t e s o pen b l a n k e t bog c o m m u n i t y n e a r R a i n b o w L a k e 51 F i g u r e 14. P i n u s c o n t o r t a - e r i c a c e o u s s h r u b - T r i - c h o phorum c e s p i t o s u m - Sphagnum s p p . - C l a d o n i a s p p . -S i p h u l a c e r a t i t e s o p e n b l a n k e t bog c o m m u n i t y ; r e l e v e 11 51 F i g u r e 15. L a r g e r e d d i s h Sphagnum r u b e l l u m mound a r o u n d t h e b a s e o f P i n u s c o n t o r t a i n a b l a n k e t bog n e a r R a i n b o w L a k e 53 F i g u r e 16. S e r i e s o f t e r r a c e d p o n d s w i t h T r i c h o p h o r u m  c e s p i t o s u m "dams" i n a s l o p i n g b l a n k e t bog w e s t o f D i a n a C r e e k , t r a n s e c t 3 53 F i g u r e 17. S l o p i n g bog w i t h s e r i e s o f t e r r a c e d ponds n e a r Work C h a n n e l 54 F i g u r e 18. Bog - p o o l c o m p l e x n e a r O l i v e r L a k e 54 F i g u r e 19. A M y r i c a g a l e - d o m i n a t e d b l a n k e t bog co m m u n i t y e a s t o f R a i n b o w L a k e ; r e l e v e 28 58 F i g u r e 20. Sphagnum l i n d b e r g i i e n c r o a c h i n g on a pond i n a s l o p i n g b l a n k e t b o g ; r e l e v e ' 30 58 F i g u r e 2 1 . A Sphagnum p a p i l l o s u m hummock d e v e l o p i n g i n a pond 59 F i g u r e 22. A l a t e r s t a g e i n pond s u c c e s s i o n i n a b l a n k e t bog 59 F i g u r e 23. E r i o p h o r u m a n g u s t i f o l i u m ( m a i n l y ) i n a l a t e r s t a g e o f pond s u c c e s s i o n i n a b l a n k e t bog 60 F i g u r e 24. The l i c h e n S i p h u l a c e r a t i t e s ( w h i t e ) and t h e l i v e r w o r t G y m n o c o l e a i n f l a t a ( b l a c k ) ; common a s s o c i a t e s i n d e p r e s s i o n s s u b j e c t t o i n t e r m i t t e n t s u b m e r g e n c e i n open b l a n k e t bog 60 F i g u r e 25. A m i n e r a l f l u s h i n a s h a l l o w , s l o p i n g b l a n k e t b o g ; r e l e v e 30 63 F i g u r e 26. Om h o r i z o n o f an O r t h i h i s t o m o d e r f r o m an open b l a n k e t bog c o m m u n i t y ; r e l e v e 20 63 F i g u r e 27. S p h a g n i c Of h o r i z o n o f an O r t h i h i s t o m o d e r f r o m an open b l a n k e t b o g ; r e l e v e 1 14 64 F i g u r e 28. S h r i n k i n g s e d i m e n t a r y p e a t i n a d r i e d - u p bog p o n d 64 F i g u r e 29. RA o r d i n a t i o n o f e r i c a c e o u s s h r u b - T j r i c h o -phorum c e s p i t o s u m - Sphagnum open bog c o m m u n i t i e s and P i n e - c y p r e s s - c e d a r - G a u l t h e r i a - F a u r i a bog w o o d l a n d c o m m u n i t i e s , a x e s 1 ( X ) and 2 ( y ) 67 F i g u r e 30. Empetrum n i g r u m - V a c c i n i u m u l i g i n o s u m -R h a c o m i t r i u m l a n u g i n o s u m - C l a d o n i a s p p . o u t c r o p c o m m u n i t y ; r e l e v e " 1 69 F i g u r e 3 1 . An O r t h i c R e g o s o l s u p p o r t i n g t h e above o u t c r o p c o mmunity ( F i g . 3 0 ) ; r e l e v e 1 69 x i i i F i g u r e 32. C o m p l e x o f open b l a n k e t bog and bog w o o d l a n d on t h e l e v e l t o g e n t l e s l o p e s and u p l a n d f o r e s t on t h e s t e e p e r s l o p e s ; T s i m p s e a n P e n i n s u l a n e a r P o r t S i m p s o n 71 F i g u r e 3 3 . The p i n e - c y p r e s s - c e d a r - G a u l t h e r i a -F a u r i a - P l e u r o z i u m - Sphagnum bog w o o d l a n d community t y p e : t r a n s e c t 4 71 F i g u r e 34. L o o k i n g w e s t a l o n g t r a n s e c t 2, e a s t o f R a i n b o w L a k e 80 F i g u r e 35. The c e d a r - c y p r e s s - h e m l o c k - z o n a l f o r e s t o f t h e m i d s l o p e c o n t r a s t s w i t h t h e S i t k a s p r u c e a l l u v i a l f o r e s t a t t h e t o e o f t h e s l o p e 80 F i g u r e 36. The c e d a r - c y p r e s s - h e m l o c k - V a c c i n i u m -C o r n u s - R h y t i d i a d e l p h u s z o n a l f o r e s t community t y p e ; r e l e v g 3 3 81 F i g u r e 37. The c e d a r - c y p r e s s - h e m l o c k - V a c c i n i u m -C o r n u s - R h y t i d i a d e l p h u s z o n a l f o r e s t c o m m u n i t y t y p e ; r e l e v S 2 81 F i g u r e 38. An o r t h i c F e r r o - Humic P o d z o l ( f o l i s o l i c p h a s e ) s u p p o r t i n g a c e d a r - c y p r e s s - h e m l o c k z o n a l f o r e s t c o m m u n i t y ; r e l e v e 1 4 88 F i g u r e 3 9. An O r t h i c Humic P o d z o l s u p p o r t i n g t h e c e d a r -c y p r e s s - h e m l o c k z o n a l f o r e s t c o mmunity o f F i g . 37; r e l e v e 2 88 F i g u r e 40. An O r g a n o F o l i s o l c o n s i s t i n g o f 60 cm o f f o r e s t humus ( m o s t l y H) o v e r a 75 cm t h i c k Oh h o r i z o n ; r e l e v e 16 90 x i v Figure 41. A cedar - spruce - hemlock - Lysichiton -Rhizomnium a l l u v i a l forest community; transect 4 90 Figure 42. A cedar - spruce - hemlock - Lysichiton -Rhizomnium community; releve 10 93 Figure 43. A Rego Humic Gleysol supporting the a l l u v i a l forest of Fig. 42.; releve 10 93 Figure 44. The amabilis f i r - hemlock - cedar - Gymno-carpium - Rhizomnium community type. This stand i s r e l a t i v e l y young and spruce - dominated; transect 3 99 Figure 45. The amabilis f i r - hemlock - cedar - Gymno- carpium - Rhizomnium community type. This stand i s dominated by western hemlock and amabilis f i r ; transect 3 99 Figure 46. The Polystichum munitum association of the amabilis f i r - hemlock - cedar - Gymnocarpiurn -Rhizomnium community type; releve 18 100 Figure 47. The Polystichum munitum association of the amabilis f i r - hemlock - cedar - Gymnocarpium -Rhizomnium community type; transect 3 100 Figure 48. A Typic F o l i s o l supporting an amabilis f i r -hemlock - cedar - Gymnocarpium - Rhizomnium upland forest community; releve 9 106 Figure 49. An Orthic Ferro - Humic Podzol ( f o l i s o l i c phase) supporting an amabilis f i r - hemlock - cedar - Gymnocarpium - Rhizomnium upland forest community; releve 27 106 X V F i g u r e 50. P r o f i l e s o f r e l e v e s c o r e s on t h e RA f i r s t a x i s o r d i n a t i o n by s t r a t u m o f a l l r e l e v e s 108 a) A l l s p e c i e s 108 b) Low s h r u b s 108 c ) H e r b s 108 d) B r y o p h y t e s , L i c h e n s and S e e d l i n g s 108 F i g u r e 5 1 . P r o f i l e s o f r e l e v 6 s c o r e s on RA f i r s t a x i s o r d i n a t i o n by s t r a t u m o f f o r e s t and w o o d l a n d r e l e v e s 109 a) A l l s p e c i e s 109 b) T r e e s 109 c ) S h r u b s 109 d) H e r b s , B r y o p h y t e s a n d L i c h e n s 109 F i g u r e 52. L o c a t i o n o f p o l l e n p r o f i l e s a m p l i n g s i t e n e a r Mt. Hayes 135 F i g u r e 53. The p i n e - c y p r e s s - c e d a r - G a u l t h e r i a -F a u r i a - P l e u r o z i u m - Sphagnum bog w o o d l a n d community s u r r o u n d i n g t h e p o l l e n c o r e s a m p l i n g s i t e 137 F i g u r e 54. Mt. Hayes p o l l e n / s p o r e p r o f i l e d i a g r a m 138 F i g u r e 55. L y s i c h i t o n a m e r i c a n u m , an a b u n d a n t s p e c i e s i n a l l u v i a l f o r e s t c o m m u n i t i e s i n t h e s t u d y a r e a 150 x v i LIST OF APPENDICES Appendix I. Edatopic g r i d o f K r a j i n a (1969) 184 Appendix I I . F i e l d forms used i n the r e l e v e sampling 185 Appendix I I I . Cover-abundance s c a l e , v i g o u r s c a l e and s t r a t a d e s i g n a t i o n s used i n the r e l e v e sampling 192 Appendix IV. L i s t of s p e c i e s recorded i n the study area 194 Appendix V. Species groups o u t l i n e d on the PCA sp e c i e s o r d i n a t i o n s ( F i g . 8.) 201 Appendix VI. Complete v e g e t a t i o n t a b l e s and vege-t a t i o n summary t a b l e s f o r each community type 204 Appendix V I I . Data matrix f o r the f i r s t a x i s RA r e l e v e o r d i n a t i o n 228 Appendix V I I I . Environment summary t a b l e f o r the f o r t y r e l e v e s 231 x v i i ACKNOWLEDEMENTS I am g r a t e f u l t o a g r e a t many people who have helped and encouraged me throughout the completion of t h i s t h e s i s . My a d v i s o r , Dr. Hamish Kimmins s t i m u l a t e d my i n t e r e s t i n f o r e s t ecology and c o u n s e l l e d me throughout t h i s study. Dr. Jim Pojar ( t h e s i s committee) i n t r o d u c e d me t o the f a s c i n a t i n g l a n d of "muskeg, rocks and r a i n " and helped me with many aspects of the study. Dr. Glenn Rouse provided l a b f a c i l i t i e s , advice and enthusiasm f o r the palynology study. Dr. Richard Hebda and Dr. Rolf Mathewes a l s o provided advice on the palynology s e c t i o n . Dr. Gary B r a d f i e l d ( t h e s i s committee) o f f e r e d much advice on the o r d i n a t i o n s of the v e g e t a t i o n data. Dr. Les L a v k u l i c h ( t h e s i s committee) and Mr. Rick Trowbridge advised me on s o i l s matters. Ms. Anne Smith was my e n t h u s i a s t i c a s s i s t a n t throughout the f i e l d season. Ms. Peggy C a r t e r , Ms. S h e i l a Humphrey, Mr. B l a i r Smith, Dr. Benito Tan, Mr. Joop van Velzen and Mr. Alan Waters a l s o helped with v a r i o u s aspects of the f i e l d work. Mr. Joop van Ve l z e n and Dr. W i l f S c h o f i e l d k i n d l y i d e n t i f i e d many of the bryophytes and Mr. Trevor Goward helped me g r e a t l y with the l i c h e n i d e n t i f i c a t i o n s . Ms. Susan Phelps provided v a l u a b l e a s s i s t a n c e with computer problems. I am e s p e c i a l l y g r a t e f u l to my s i s t e r Ms. Marnie Andersen f o r her s k i l l e d p r e p a r a t i o n of the manuscript and to Mr. J . D. Dunlop, P r e s i d e n t of CANA I n d u s t r i a l C o n t r a c t o r s L t d . , x v i i i E dmonton, A l b e r t a f o r g e n e r o u s l y m a k i n g a v a i l a b l e w o r d p r o -c e s s i n g f a c i l i t i e s . I a l s o t h a n k Ms. L i n d a W i l s o n and Ms. P a t s y Quay f o r t h e i r h e l p i n p r e p a r i n g t h e m a n u s c r i p t a nd Ms. Eve C o u l s o n a n d Mr. T i m Mock f o r d r a f t i n g a n d r e p r o d u c t i o n o f some o f t h e f i g u r e s . F i n a n c i a l s u p p o r t f o r t h e f i e l d work was k i n d l y p r o v i d e d b y t h e R e s e a r c h B r a n c h , B r i t i s h C o l u m b i a M i n i s t r y o f F o r e s t s ; I t h a n k D r . Ted B a k e r , D r . D i c k A n n a s a n d Mr. Dave W i l f o r d f o r a r r a n g i n g t h i s f i n a n c i a l s u p p o r t . The M i n i s t r y o f F o r e s t s N o r t h C o a s t D i s t r i c t o f f i c e a n d P r i n c e R u p e r t / S m i t h e r s R e g i o n a l O f f i c e s p r o v i d e d l o g i s t i c a l s u p p o r t . F i n a l l y , I t h a n k t h e N a t u r a l S c i e n c e s a nd E n g i n e e r i n g R e s e a r c h C o u n c i l o f Canada who p r o v i d e d me w i t h a p o s t g r a d u a t e s c h o l a r s h i p d u r i n g my two y e a r s on campus. 1 CHAPTER I INTRODUCTION The development and application of ecological land c l a s s i f i c a t i o n in B r i t i s h Columbia over the past 30 years has contributed s i g n i f i c a n t l y to our knowledge of vegetation-environment relationships, p a r t i c u l a r l y within forest ecosystems. Since i t s introduction in B r i t i s h Columbia in the 1950's by V.J. Krajina (Botany Department, University of B.C.), the Biogeoclimatic c l a s s i f i c a t i o n system has been applied in many studies throughout B.C. (e.g., Krajina 1959, 1965, 1969, 1972; Brooke et a l . 1970; Kojima 1971; Wali and Krajina 1973; Klinka 1976; Annas 1977). An ecosystematic approach to land c l a s s i f i c a t i o n has been well recognized in B.C. as a means of organizing and communicating knowledge of our natural environment; as a predictive tool in land management and as a framework for research on the structure and functioning of ecosystems (Kimmins 1977). The Biogeoclimatic system has now been adopted and applied with some modifications by the Research Branch of the B r i t i s h Columbia Ministry of Forests in i t s c l a s s i f i c a t i o n of forest and range ecosystems, and management interpretations are being developed using this c l a s s i f i c a t i o n as a framework (Klinka 1977; Klinka et a l . 1979; Annas et a l . 1979; Mitchell and Green 1981; Pojar et a l . 1982). Several other land c l a s s i f i c a t i o n systems have also been applied in B r i t i s h Columbia. The Habitat-type c l a s s i f i c a t i o n 2 (Daubenmire 1952, 1968; P f i s t e r and Arno 1980) has been a p p l i e d by MacMillan B l o e d e l L t d . on Vancouver I s l a n d and the Queen C h a r l o t t e I s l a n d s (Packee 1976, 1979; Beese 1981) and the B i o p h y s i c a l c l a s s i f i c a t i o n system i s being a p p l i e d i n s e v e r a l areas throughout B.C. by the T e r r e s t r i a l S t u d i e s Branch of the B r i t i s h Columbia M i n i s t r y of Environment (Walmsley and van Barneveld 1977). Although these c l a s s i f i c a t i o n systems d i f f e r somewhat i n t h e i r approach to landscape s t r a t i f i c a t i o n and i n terminology, a l l of them attempt to d e f i n e and d e s c r i b e c l a s s i f i c a t i o n u n i t s (community types, h a b i t a t types, ecosystem a s s o c i a t i o n s , etc.) based on s t r u c t u r a l f e a t u r e s of the h a b i t a t ( v e g e t a t i o n , s o i l s , landforms, e t c . ) . Such a c l a s s i f i c a t i o n based on ecosystem s t r u c t u r e p r o v i d e s the framework i n which to study the f u n c t i o n a l aspects of ecosystems, such as the r a t e s and pathways of biogeochemical c y c l i n g and energy exchange, as w e l l as s u c c e s s i o n a l r e l a t i o n s h i p s . The c l a s s i f i c a t i o n i s a necessary f i r s t step s i n c e i t r e c o g n i z e s d i s t i n c t u n i t s which r e c u r i n the landscape. S u c c e s s i o n a l r e l a t i o n s h i p s between or w i t h i n these n a t u r a l u n i t s can then be studied and r e p r e s e n t a t i v e examples can be chosen f o r q u a n t i t a t i v e s t u d i e s of f u n c t i o n a l r e l a t i o n s h i p s . , 1.1 E c o l o g i c a l I n v e s t i g a t i o n s A l o n g N o r t h C o a s t a l B r i t i s h C o l u m b i a P r i o r to the 1970's, the f o r e s t and muskeg ecosystems which dominate the landscape along the outer coast of B r i t i s h 3 Columbia from northern Vancouver Island to the Alaska panhandle had received very l i t t l e ecological investigation. Early studies by Rigg (1914, 1917, 1925, 1940), Rigg and Thompson (1922), and Osvald (1933) concentrated on the description of vegetation and peat stratigraphy and successional relationships within bogs in Washington, Oregon, southwestern B.C. and southeastern Alaska. These early studies dealt l i t t l e with the s o i l s and vegetation of forest ecosystems or with the ecosystems along middle and north coastal B.C. More recent ecological studies of coastal forest and bog ecosystems in B.C. (Wade 1965; Krajina 1965, 1969; Mueller-Dombois 1959, 1965; Lesko 1961; O r l o c i 1961, 1964; Cordes 1972; Kojima and Krajina 1975; Klinka 1976; Klinka et a l . 1979, 1980) have been r e s t r i c t e d to the south coastal mainland and Vancouver Island. 8 Several palynological studies of bogs have been carried out in B r i t i s h Columbia, but these also have mainly been in south coastal regions (Hansen 1940; Mathewes 1973; Mathewes and Rouse 1975; Hebda 1977, 1979; Hebda & Rouse 1979). A noteable exception is the work of Heusser (1960) who produced pollen p r o f i l e s from several bogs in the Prince Rupert area and on the Queen Charlotte Islands as well as in Oregon, Washington and southeastern Alaska. Patterns of vegetation and s o i l development in the forest-bog complexes of Southeast Alaska described by Zach (1950), Lawrence (1958), Stephens et a l . (1970), Neiland (1971), and Ugolini and Mann (1979) appear to be more 4 pertinent to the ecosystems along the outer middle and north coast of B.C. than do most of the studies from the south coast. In 1976, the B r i t i s h Columbia Ministry of Forests began ecological sampling (vegetation, s o i l s , and mensurational data) in the Prince Rupert Forest Region (essentially the northwest quarter of the province). Access by boat and helicopter was provided into many coastal areas previously unstudied by s c i e n t i s t s . Reconnaissance and limited releve sampling in these areas resulted in the proposal of a new biogeoclimatic zone, the Coastal Cedars Pine - Hemlock Zone (CCPH); a lowland to montane outer coastal zone stretching from northern Vancouver Island to Southeast Alaska (Pojar and Annas 1980) and including the west coast of the Queen Charlotte Islands. Krajina (1969) included these areas within the Coastal Western Hemlock Biogeoclimatic Zone (CWH). This thesis was undertaken as part of the early description and c l a s s i f i c a t i o n of the proposed new zone. However, evaluation of the CCPH proposal was not an objective of the thesis. The Prince Rupert area, which represents one of the few accessible (by road) examples of the CCPH zone in B.C., proved to be ideal for this study since a variety of bog, forest and intermediate vegetation could be reached from Highway 16 or by boat on the many lakes within the area. 1.2 T h e s i s O b j e c t i v e s The objectives of this thesis were: 1. To c l a s s i f y and describe the major plant community 5 t y p e s and i d e n t i f y m a j o r t r e n d s i n v e g e t a t i o n and s o i l d e v e l o p m e n t a l o n g t o p o g r a p h i c t r a n s e c t s i n a r e p r e s e n t a t i v e a r e a o f t h e p r o p o s e d CCPH b i o g e o c l i m a t i c z o n e l . 2. To r e c o n s t r u c t t h e h i s t o r i c a l d e v e l o p m e n t o f a bog w o o d l a n d c o m m u n i t y t y p e u s i n g p o l l e n a n a l y s i s and p e a t s t r a t i g r a p h y . 3. To d e s c r i b e s u c c e s s i o n a l r e l a t i o n s h i p s w i t h i n t h e s t u d y a r e a by c o m b i n i n g p a l y n o l o g i c a l and s t r a t i g r a p h i c e v i d e n c e w i t h e m p i r i c a l e v i d e n c e f r o m p r e s e n t - d a y e c o s y s t e m s . 1 . 3 T h e s i s O u t l i n e The t h e s i s i s o r g a n i z e d as f o l l o w s : C h a p t e r 2 p r e s e n t s a l i t e r a t u r e r e v i e w o f t h e t e r m i n o l o g y and c o n c e p t s d e s c r i b i n g bog g e n e s i s , m o r p h o l o g y and n u t r i e n t r e l a t i o n s h i p s . I t a l s o p r o v i d e s d e f i n i t i o n s f o r t h e t e c h n i c a l t e r m s u s e d i n t h e t h e s i s . I n C h a p t e r 3 t h e s t u d y a r e a i s d e s c r i b e d and t h e e n v i r o n m e n t a l and e c o l o g i c a l f e a t u r e s o f t h e CCPH b i o g e o c l i m a t i c zone a r e o u t l i n e d . C h a p t e r 4 d e a l s w i t h t h e c l a s s i f i c a t i o n and d e s c r i p t i o n o f t h e m a j o r bog and f o r e s t c o m m u n i t y t y p e s w i t h i n t h e s t u d y a r e a . I d e c i d e d t h a t v e g e t a t i o n - e n v i r o n m e n t r e l a t i o n s h i p s c o u l d be b e s t e x a m i n e d by r e l e v e * s a m p l i n g a l o n g f o u r t r a n s e c t s w h i c h e x h i b i t e d t o p o g r a p h i c d i v e r s i t y and t r a v e r s e d a v a r i e t y -'-This p r o v i d e d t h e f r a m e w o r k f o r two s t u d i e s c o m p a r i n g t h e n u t r i e n t r e g i m e s o f s e l e c t e d c o m m u n i t y t y p e s u s i n g s o i l w a t e r c h e m i s t r y and f o l i a r a n a l y s i s . T h e s e two s t u d i e s w i l l be r e p o r t e d e l s e w h e r e . 6 o f m a t u r e c l i m a x e c o s y s t e m s , r a n g i n g f r o m open muskeg t o c l o s e d u p l a n d f o r e s t . V e g e t a t i o n d a t a f r o m 40 r e l e v e s were a n a l y z e d u s i n g b o t h s u b j e c t i v e t a b u l a r c o m p a r i s o n t e c h n i q u e s as w e l l a s more o b j e c t i v e o r d i n a t i o n t e c h n i q u e s ( p r i n c i p l e c o m p o n e n t s a n a l y s i s , r e c i p r o c a l a v e r a g i n g and p o l a r o r d i -n a t i o n ) . D e t a i l e d d e s c r i p t i o n s o f t h e v e g e t a t i o n , s o i l s and l a n d f o r m s c h a r a c t e r i z i n g 6 c o m m u n i t y t y p e s a r e p r e s e n t e d and v e g e t a t i o n - e n v i r o n m e n t r e l a t i o n s h i p s w i t h i n t h e s t u d y a r e a a r e d i s c u s s e d and c o m p a r e d w i t h o t h e r s t u d i e s . I n C h a p t e r 5 t h e h i s t o r i c a l d e v e l o p m e n t o f a bog w o o d l a n d c o m m u n i t y i s r e c o n s t r u c t e d u s i n g p o l l e n a n a l y s i s and p e a t s t r a t i g r a p h y . A 2 m p e a t c o r e was c o l l e c t e d f r o m a P i n u s  c o n t o r t a ^ - C h a m a e c y p a r i s n o o t k a t e n s i s - T h u j a p l i c a t a bog w o o d l a n d a l o n g one o f t h e s t u d y t r a n s e c t s and p r o c e s s e d i n t h e l a b f o r p o l l e n a n a l y s i s . A p r o f i l e o f p o l l e n / s p o r e p e r c e n t a g e s and a b s o l u t e p o l l e n c o n c e n t r a t i o n s was p r o d u c e d f o r t h e p e a t d e p o s i t and f r o m t h i s t h e v e g e t a t i o n h i s t o r y o f t h e bog w o o d l a n d was r e c o n s t r u c t e d . L o c a l e d a p h i c f a c t o r s as w e l l as l a r g e r s c a l e p a l e o c l i m a t i c f a c t o r s l i k e l y a s s o c i a t e d w i t h t h e d e v e l o p m e n t o f t h e e x i s t i n g bog w o o d l a n d a r e d i s c u s s e d . I n C h a p t e r 6, p r e s e n t - d a y f l o r i s t i c and e d a p h i c e v i d e n c e f r o m t h i s and o t h e r s t u d i e s a l o n g t h e n o r t h P a c i f i c c o a s t i s c o m b i n e d w i t h t h e p a l y n o l o g i c a l i n t e r p r e t a t i o n s i n a f i n a l d i s c u s s i o n o f e c o s y s t e m d y n a m i c s . S o u r c e s o f s p e c i e s n o m e n c l a t u r e a r e g i v e n on page 16. 7 CHAPTER 2 BOG TERMINOLOGY: A LITERATURE REVIEW 2.1 I n t r o d u c t i o n Much o f t h i s t h e s i s d e a l s w i t h d e s c r i p t i o n s o f b o g e c o s y s t e m s , b o g - f o r m i n g p r o c e s s e s and t h e d y n a m i c s o f b o g -f o r e s t s u c c e s s i o n . D i s c u s s i o n s o f b o g e c o s y s t e m s h a v e b e e n p l a g u e d b y t h e v a r i e t y o f t e r m i n o l o g y t h a t h a s a c c u m u l a t e d s i n c e t h e 1 6 t h c e n t u r y a n d p r o b a b l y e a r l i e r when p e a t l a n d s f i r s t f a s c i n a t e d n a t u r a l i s t s a n d s c i e n t i s t s (Gorham 1 9 5 3 ) . A r e v i e w o f t h e l i t e r a t u r e d e a l i n g w i t h p e a t l a n d s and t h e i r d e v e l o p m e n t p r o v i d e s s e v e r a l a l t e r n a t i v e t e r m i n o l o g i e s t o d e s c r i b e b o t h p e a t l a n d d e v e l o p m e n t a n d n u t r i e n t r e l a t i o n s h i p s ( T a n s l e y 1939; R i g g 1 940; S j o r s 1950; Gorham 1957; N e i l a n d 1 9 7 1 ; T a y l o r a n d S m i t h 1972; Moore and B e l l a m y 1 9 7 4 ) . B e c a u s e o f t h e l a c k o f s t a n d a r d i z a t i o n , e s p e c i a l l y b e t w e e n E u r o p e a n a n d N o r t h A m e r i c a n w o r k e r s , t h e p e a t l a n d e c o l o g i s t must c h o o s e t h e t e r m s h e f e e l s m o s t c o m f o r t a b l e w i t h and d e f i n e them a t t h e o u t s e t . I n t h i s t h e s i s , s e p a r a t e t e r m i n o l o g y w i l l be u s e d t o d e s c r i b e b o g g e n e s i s a n d m o r p h o l o g y ( s u r f a c e c o n f i g u r a t i o n , h y d r o t o p o g r a p h i c c i r c u m s t a n c e s f o r p e a t f o r m a t i o n ) and n u t r i e n t r e l a t i o n s h i p s , b e c a u s e i t i s s o m e t i m e s d i f f i c u l t t o r e l a t e t h e p h y s i c a l f e a t u r e s o f t h e b o g t o t h e a p p a r e n t n u t r i e n t c o n d i t i o n s , a s i n d i c a t e d e i t h e r b y t h e f l o r i s t i c c o m p o s i t i o n o r b y d i r e c t n u t r i e n t a n a l y s i s . 8 2 . 2 B o g G e n e s i s a n d M o r p h o l o g y I have a d o p t e d t h e scheme p r e s e n t e d by Moore and B e l l a m y (1974) b e c a u s e i t : 1) r e l a t e s p e a t l a n d ( m i r e ) d e v e l o p m e n t and p e a t t y p e s t o a h y d r o l o g i c a l b a l a n c e ; 2) r e c o g n i z e s t h e d i s t i n c t i o n b e t w e e n c l i m a t i c and e d a p h i c ( t o p o g r a p h y , m i n e r a l n u t r i t i o n ) f a c t o r s i n c o n t r o l l i n g p e a t a c c u m u l a t i o n ; and 3) u s e s s e p a r a t e t e r m i n o l o g y t o d e s c r i b e t h e m i n e r a l n u t r i t i o n and g e n e s i s o f p e a t l a n d s . . A b r i e f o u t l i n e o f t h e scheme f o l l o w s , t o g e t h e r w i t h d e f i n i t i o n s o f o t h e r t e r m s u s e d i n my d i s c u s s i o n o f p e a t l a n d e c o s y s t e m s . Moore and B e l l a m y (1974) d i s t i n g u i s h t h r e e c a t e g o r i e s o f p e a t t y p e s o r ' m i r e s y s t e m s ' ; p r i m a r y , s e c o n d a r y and t e r t i a r y . P r i m a r y p e a t s a c c u m u l a t e i n b a s i n s o r d e p r e s s i o n s . They d e v e l o p m a i n l y b e c a u s e t h e r e i s a b a s i n o r d e p r e s s i o n t h a t r e t a i n s w a t e r and c r e a t e s a ' t e m p l a t e ' f o r p r i m a r y p e a t a c c u m u l a t i o n . T h i s i n t u r n r e d u c e s t h e w a t e r r e t e n t i o n c a p a c i t y o f t h e r e s e r v o i r . P r i m a r y p e a t d e p o s i t s a r e f o u n d i n a l l e x c e p t t h e h o t t e s t and most a r i d r e g i o n s o f t h e e a r t h . S e c o n d a r y p e a t s a c c u m u l a t e beyond t h e p h y s i c a l c o n f i n e s o f t h e b a s i n . A t t h i s s t a g e t h e p e a t b e g i n s t o a c t as a r e s e r v o i r and i n c r e a s e s t h e w a t e r r e t e n t i o n c a p a c i t y o f t h e l a n d f o r m . T e r t i a r y p e a t s d e v e l o p a b o v e t h e p h y s i c a l l i m i t s o f t h e g r o u n d w a t e r . They a c t as a s p o n g e , h o l d i n g t h e i r w a t e r by c a p i l l a r i t y . T e r t i a r y p e a t s u s u a l l y h ave p e r c h e d w a t e r t a b l e s and r e c e i v e t h e i r m o i s t u r e p r i m a r i l y t h r o u g h p r e c i p i t a t i o n . The a c c u m u l a t i o n o f s e c o n d a r y and t e r t i a r y p e a t s i s l a r g e l y u n d e r c l i m a t i c c o n t r o l and t h e y a r e t h e r e f o r e r e s t r i c t e d i n d i s t r i b u t i o n t o r e g i o n s o f more humid a n d / o r c o o l e r c l i m a t e s 9 ( a r e a s o f l o w e r p o t e n t i a l e v a p o t r a n s p i r a t i o n ) . Moore and B e l l a m y (1974) d e s c r i b e t h e t e m p l a t e f o r p e a t l a n d f o r m a t i o n i n t e r m s o f t h e h y d r o l o g i c a l e q u a t i o n : i n f l o w + p r e c i p i t a t i o n = o u t f l o w + e v a p o r a t i o n + r e t e n t i o n U n d e r h u m i d c l i m a t e s ( > p p t . , <evap.) w a t e r w i l l be r e t a i n e d on a g r e a t e r v a r i e t y o f s i t e s , n o t o n l y i n b a s i n s and d e p r e s -s i o n s , b u t a l s o on f l a t o r g e n t l y s l o p i n g l o w l a n d s and e v e n on m o u n t a i n s l o p e s . C l i m a t i c f a c t o r s t h e m s e l v e s become t h e t e m p l a t e f o r p e a t a c c u m u l a t i o n ( a n d p e a t f o r m a t i o n becomes a z o n a l r a t h e r t h a n a t o p o e d a p h i c p r o c e s s ) . U n d e r s u c h c o n d i -t i o n s p e a t d e p o s i t s may c o n s i s t e n t i r e l y o f s e c o n d a r y and t e r t i a r y p e a t s . The d i v i s i o n o f p e a t t y p e s i n t o p r i m a r y , s e c o n d a r y and t e r t i a r y i s t h u s u s e f u l i n d i s t i n g u i s h i n g b e t w e e n c l i m a t i c and t o p o e d a p h i c c o n t r o l o f b o g f o r m a t i o n . Moore and B e l l a m y ( 1 9 7 4 ) h a v e u s e d t h i s c l a s s i f i c a t i o n o f p e a t t y p e s a s t h e b a s i s f o r a z o n a t i o n o f E u r o p e a n m i r e c o m p l e x e s . E l e v e n m i r e c o m p l e x z o n e s a r e d e l i n e a t e d i n a s o u t h e a s t - n o r t h w e s t d i r e c t i o n s t a r t i n g w i t h p r i m a r y and s e c o n d a r y m i r e s y s t e m s ( v a l l e y b o g s and f e n s ) i n t h e s o u t h e a s t ( B l a c k and C a s p i a n S e a s ) a n d p r o g r e s s i n g t h r o u g h a s e r i e s o f z o n e s r e p r e s e n t i n g a g e n e r a l i n c r e a s e i n d o m i n a n c e o f t h e t e r t i a r y p e a t component. M i r e s o f t h e w e s t c o a s t o f Norway, t h e m o u n t a i n s o f S c o t l a n d , n o r t h e r n E n g l a n d , W a l e s and I r e l a n d a r e i n c l u d e d w i t h i n t h e B l a n k e t M i r e Complex where " p e a t l a n d s c a n d e v e l o p d i r e c t l y o n m i n e r a l g r o u n d up t o a c o n s i d e r a b l e a n g l e o f s l o p e " . T h e s e a u t h o r s s t a t e t h a t i n t h e m o u n t a i n s o f 10 w e s t e r n I r e l a n d p e a t l a y e r s more t h a n two m e t e r s deep a r e p r e s e n t on s l o p e s o f 25 p e r c e n t . T h e s e b l a n k e t m i r e s c a n c o v e r e x t e n s i v e a r e a s . Bog f o r m a t i o n a p p e a r s t h e n t o be a z o n a l p r o c e s s ( c l i m a t i c a l l y c o n t r o l l e d ) i n t h e s e A t l a n t i c c o a s t a l a r e a s o f E u r o p e . The B l a n k e t M i r e Complex ( F i g . 1) p r o v i d e s a u s e f u l f r a m e w o r k i n w h i c h t o e x p l a i n t h e r e l a t i o n s h i p s among t h e v a r i o u s t y p e s o f bog e c o s y s t e m s o c c u r r i n g w i t h i n t h e s t u d y a r e a . T h i s c o n c e p t i s e s p e c i a l l y u s e f u l b e c a u s e i t d e s c r i b e s a c o m p l e x o f bog t y p e s d o m i n a t e d by b l a n k e t bog ( t e r t i a r y p e a t l a n d s l a r g e l y i n f l u e n c e d by c l i m a t e ) b u t a l s o i n c l u d i n g more l o c a l i z e d a r e a s o f p r i m a r y and s e c o n d a r y p e a t l a n d s ( b a s i n b o g s , f l a t b o g s , v a l l e y b o g s , f e n s ) as w e l l as a r e a s where t e r t i a r y p e a t s h a v e a c c u m u l a t e d o v e r 1° and 2° p e a t t y p e s ( t h e s e a r e t h e domed o r r a i s e d b o g s . ) E x a m p l e s o f a l l t h e s e p e a t l a n d t y p e s c a n be f o u n d w i t h i n t h e s t u d y a r e a b u t t e n d t o g r a d e i n t o one a n o t h e r . The many bog e c o s y s t e m s ( i n c l u d i n g b o t h open bogs and bog w o o d l a n d s ) a r e t h u s b e s t d e s c r i b e d as a c o m p l e x o f bog t y p e s d i f f e r i n g i n g e n e t i c d e v e l o p m e n t , s u r f a c e c o n f i g u r a t i o n and s p e c i e s c o m p o s i t i o n i n r e s p o n s e t o h y d r o l o g i c , g e o l o g i c and t o p o g r a p h i c f a c t o r s . The c o n c e p t o f t h e b l a n k e t bog c o m p l e x m u s t , h o w e v e r , be e x p a n d e d i n s c o p e t o p r o p e r l y c h a r a c t e r i z e e c o l o g i c a l c o n d i t i o n s w i t h i n t h e s t u d y a r e a . F o r e s t e d e c o s y s t e m s (bog w o o d l a n d s and u p l a n d f o r e s t s ) c o v e r much o f t h e l a n d s c a p e and t h e same p r o c e s s e s w h i c h c o n t r o l bog d e v e l o p m e n t a p p e a r t o i n f l u e n c e u p l a n d f o r e s t d e v e l o p m e n t u n d e r c e r t a i n c i r c u m -11 F i g . 1. P r o f i l e o f b l a n k e t m i r e complex show ing a l l t h e component p e a t t y p e s . A f t e r Moore and B e l l a m y ( 1 9 7 4 ) . 12 s t a n c e s . A r e c o g n i t i o n o f t h e r e l a t i o n s h i p s b e t w e e n t h e bog and u p l a n d f o r e s t e c o s y s t e m s i s c r u c i a l t o an u n d e r s t a n d i n g o f t h e z o n a l p r o c e s s e s o f s o i l and v e g e t a t i o n d e v e l o p m e n t . The m o s a i c o f e c o s y s t e m s w i t h i n t h e s t u d y a r e a i s t h e r e f o r e b e s t d e s c r i b e d as a " B l a n k e t B o g - U p l a n d F o r e s t C o m p l e x ' . 2 . 3 N u t r i e n t R e l a t i o n s h i p s So f a r I have d e a l t o n l y w i t h t h e g e n e t i c d e v e l o p m e n t o f p e a t l a n d s . R e g a r d i n g n u t r i e n t r e l a t i o n s h i p s w i t h i n t h e s e e c o s y s t e m s , I a g a i n p r e f e r t h e t e r m i n o l o g y o f Moore and B e l l a m y (1974) who hav e a d o p t e d a s y s t e m i n t r o d u c e d by t h e P o l i s h e c o l o g i s t , K u l c z y n s k i , i n 1949. K u l c z y n s k i ' s scheme e m p h a s i z e s t h e i m p o r t a n c e o f g r o u n d w a t e r m o b i l i t y i n c o n t r o l l i n g e d a p h i c c o n d i t i o n s w i t h i n p e a t l a n d s . He u s e s t h e t e r m s ' r h e o p h i l o u s ' , ' t r a n s i t i o n ' and ' o m b r o p h i l o u s ' t o d e s c r i b e t h e w a t e r r e g i m e o f p e a t l a n d s . R h e o p h i l o u s means ' l o v i n g t h e f l o w ' and d e s c r i b e s p e a t l a n d s t h a t have m o b i l e g r o u n d w a t e r s . O m b r o p h i l o u s means ' l o v i n g t h e r a i n ' and r e f e r s t o p e a t l a n d s t h a t have i m m o b i l e g r o u n d w a t e r s and t h u s r e l y on r a i n f a l l f o r t h e i r w a t e r and n u t r i e n t s u p p l y . T r a n s i t i o n p e a t l a n d s a r e i n t e r m e d i a t e b e t w e e n t h e two, p e r h a p s b e c a u s e o f i n s u f f i c i e n t g r o u n d w a t e r s u p p l y o r b e c a u s e t h e y a r e c h a n g i n g f r o m a r h e o p h i l o u s t o an o m b r o p h i l o u s w a t e r r e g i m e as p e a t a c c u m u l a t e s . I n t h e c l a s s i c s e q u e n c e o f w e t l a n d s u c c e s s i o n i n a b a s i n , t h e p r o g r e s s i o n f r o m p r i m a r y p e a t t h r o u g h t o t h e s e c o n d a r y and t e r t i a r y p e a t o f t h e r a i s e d bog s t a g e i s p a r a l l e l e d by t h e 13 s e q u e n c e f r o m a r h e o p h i l o u s t h r o u g h t r a n s i t i o n t o o m b r o p h i l o u s w a t e r r e g i m e , as t h e i n f l u e n c e o f f l o w i n g m i n e r a l g r o u n d w a t e r s l o w l y becomes o u t w e i g h e d by t h e i n f l u e n c e o f p r e c i p i t a t i o n f a l l i n g on t h e bog s u r f a c e . W i t h i n t h e b l a n k e t bog c o m p l e x , h o w e v e r , t h i s r e l a t i o n s h i p i s n o t a l w a y s as c l e a r , and p o c k e t s o f t r a n s i t i o n o r p e r h a p s r h e o p h i l o u s e c o s y s t e m s may o c c u r t h r o u g h o u t t h e d o m i n a n t l y o m b r o p h i l o u s b l a n k e t bog l a n d f o r m , w h e r e v e r l o c a l e d a p h i c f a c t o r s p e r m i t . F o r e x a m p l e , t h e most s l o p i n g p o r t i o n s o f b l a n k e t bogs may have w a t e r s p e r c o l a t i n g t h r o u g h them. E s p e c i a l l y where t h e s e bogs a r e s h a l l o w o v e r m i n e r a l - r i c h b e d r o c k , a t r a n s i t i o n o r w e a k l y r h e o p h i l o u s w a t e r r e g i m e may p r e v a i l and be m a n i f e s t e d i n t h e f l o r a . P o r t i o n s o f b l a n k e t bogs t h a t b o r d e r s t r e a m l e t s may a l s o have i m p r o v e d w a t e r r e g i m e s . The t e r m s 'ombrogenous* and ' s o l i g e n o u s ' ( H e i n s e l m a n 1 9 6 3 ; N e i l a n d 1971) o c c u r i n t h e l i t e r a t u r e and a r e a n a l o g o u s i n a g e n e r a l s e n s e t o o m b r o p h i l o u s and r h e o p h i l o u s r e s p e c t i v e l y . I p r e f e r t h e l a t t e r u s a g e , h o w e v e r , b e c a u s e t h e s u f f i x 'genous' r e l a t e s t o p e a t l a n d g e n e s i s and, as p r e v i o u s l y o u t l i n e d , t h e p a r a l l e l b e t w e e n g e n e s i s and n u t r i e n t c o n d i t i o n s c a n n o t a l w a y s be e a s i l y drawn. The t e r m ' t o p o g e n o u s ' w i l l be u s e d i n t h e g e n e t i c s e n s e o n l y , t o d e s c r i b e p e a t l a n d s w h i c h h a v e a p p a r e n t l y f o r m e d i n l o c a l d e p r e s s i o n s o r b a s i n s . T h i s t e r m d oes n o t r e l a t e t o s o u r c e s o f n u t r i e n t s o r w a t e r r e g i m e s d i r e c t l y . H e i n s e l m a n (1970) u s e s t h e t e r m s ' m i n e r o t r o p h i c ' and 1 o m b r o t r o p h i c 1 t h a t w e r e o r i g i n a l l y i n t r o d u c e d by t h e S w e d i s h 14 e c o l o g i s t Du R i e t z . T h e s e t e r m s a r e c o n s i d e r e d a n a l o g o u s t o r h e o p h i l o u s and o m b r o p h i l o u s r e s p e c t i v e l y , a l t h o u g h t h e y e m p h a s i z e t h e s o u r c e o f s o i l w a t e r r a t h e r t h a n i t s m o b i l i t y . B o t h s e t s o f t e r m s , h o w e v e r , a r e a n a l o g o u s i n t h e s e n s e t h a t t h e y make t h e d i s t i n c t i o n b e t w e e n l a t e r a l g o u n d w a t e r f l o w , and t h u s n u t r i e n t t u r n o v e r w i t h i n t h e e c o s y s t e m , as o p p o s e d t o i n p u t s f r o m a b o v e v i a p r e c i p i t a t i o n and a s u b s e q u e n t s t a g n a t i o n o f t h a t w a t e r once i t r e a c h e s t h e p e a t mass. A f u r t h e r a d v a n t a g e t o u s i n g t e r m s w h i c h make no r e f e r e n c e t o g e n e s i s i s t h a t t h e y c a n be more w i d e l y a p p l i e d t o e c o s y s t e m s o t h e r t h a n t r u e p e a t l a n d s . T h u s , I a l s o u s e t h e s e t e r m s t o d e s c r i b e w a t e r r e g i m e s w i t h i n t h e u p l a n d f o r e s t e c o s y s t e m s i n t h i s s t u d y b e c a u s e f o r e s t e s t a b l i s h m e n t a p p e a r s t o be i n f l u e n c e d by a s i m i l a r s e t o f p r o c e s s e s . A d i s t i n c t i o n m ust be made b e t w e e n t h e s e w a t e r r e g i m e t e r m s and t h e m o i s t u r e and n u t r i e n t r e g i m e s a p p l i e d by K r a j i n a (1969) a nd a d o p t e d by t h e B r i t i s h C o l u m b i a M i n i s t r y o f F o r e s t s i n t h e i r c l a s s i f i c a t i o n o f f o r e s t e c o s y s t e m s ( A p p e n d i x I ) . T h e s e t e r m s w i l l a l s o be u s e d i n t h i s s t u d y b u t o n l y t o r a t e e c o s y s t e m s on a r e l a t i v e s c a l e o f s o i l m o i s t u r e ( w i t h no r e f e r e n c e t o s o u r c e s o r r a t e s o f f l o w ) and n u t r i e n t s t a t u s w i t h i n t h e s t u d y a r e a . T h i s e d a t o p i c c l a s s i f i c a t i o n i s b a s e d on a b r o a d e r a s s e s s m e n t o f e d a p h i c and f l o r i s t i c c h a r a c t e r i s t i c s t h a n i s t h e w a t e r r e g i m e a s s e s s m e n t d e s c r i b e d a b o v e . The two a r e i n t i m a t e l y r e l a t e d h o w e v e r . F o r e x a m p l e , a h y g r i c / p e r m e s o t r o p h i c m o i s t u r e / n u t r i e n t r e g i m e w o u l d l i k e l y be a s s o c i a t e d w i t h a r h e o p h i l o u s ( o r m i n e r o t r o p h i c ) w a t e r 15 r e g i m e . A K r a j i n a - t y p e a s s e s s m e n t p l a c e s g r e a t e r e m p h a s i s on s t a t i c amounts o f s o i l w a t e r and n u t r i e n t s w h i l e K u l c z y n s k i ' s e m p h a s i z e s t h e d y n a m i c s o f t h e s y s t e m . 16 CHAPTER 3 LOCATION AND DESCRIPTION OF THE STUDY AREA 3.1 General D e s c r i p t i o n o f the C o a s t a l Cedars-Pine-Hemlock B i o g e o c l i m a t i c Zone The C o a s t a l C e d a r s - P i n e - H emlock B i o g e o c l i m a t i c Zone (CCPH) o c c u r s a s a n a r r o w b e l t , 30 t o 70 km w i d e , a l o n g t h e o u t e r c o a s t a l m a i n l a n d and o f f s h o r e i s l a n d s o f B r i t i s h C o l u m b i a f r o m n o r t h e r n V a n c o u v e r I s l a n d t o S o u t h e a s t A l a s k a , i n c l u d i n g t h e w e s t c o a s t o f t h e Queen C h a r l o t t e I s l a n d s . I t i s a l o w l a n d t o montane z o n e o c c u r r i n g f r o m s e a l e v e l t o a t l e a s t 600m e l e v a t i o n ( P o j a r a n d A n n a s 1 9 8 0 ) . K r a j i n a ( 1 9 6 9) i n c l u d e d t h e s e a r e a s w i t h i n t h e C o a s t a l W e s t e r n Hemlock Zone (CWH). K l i n k a e t a l . ( 1 9 7 9 , 1980) d e f i n e a w e t t e r M a r i t i m e N o r t h e r n s u b - z o n e o f t h e CWH z o n e o c c u r r i n g o n n o r t h e r n V a n c o u v e r I s l a n d and t h e a d j a c e n t m a i n l a n d t h a t a p p e a r s t o r e p r e s e n t t h e s o u t h e r n t r a n s i t i o n f r o m CCPH t o CWH. E c o s y s t e m s o c c u r r i n g i n S o u t h e a s t A l a s k a w h i c h a r e a n a l a g o u s t o many CCPH e c o s y s t e m s i n B.C. h a v e b e e n d e s c r i b e d b y A l a b a c k ( 1 9 8 0 ) , S t e p h e n s e t a l . ( 1 9 6 9 ) ( b o t h i n V i e r e c k and D y r n e s s 1980) and N e i l a n d ( 1 9 7 1 ) . T h i s z o n e may e x t e n d a s f a r n o r t h a s 56° N. l a t . ( P o j a r and A n n a s 1 9 8 0 ) . The c l i m a t e o f t h e CCPH i s c l a s s i f i e d a s C f b ( a f t e r K o p p e n ) . I t i s c o o l , m e s o t h e r m a l , e q u a b l e , p e r h u m i d , h y p e r o c e a n i c and r a i n y w i t h r e l a t i v e l y l i t t l e snow and f r e q u e n t p e r i o d s o f f o g ( P o j a r and Annas 1 9 8 0 ) . The g r a d u a l and c o n t i n u o u s a c c u m u l a t i o n o f o r g a n i c m a t t e r a t t h e g r o u n d s u r f a c e i s an i m p o r t a n t s o i l f o r m i n g p r o -17 cess within the CCPH zone. Wet s o i l conditions are maintained throughout the year even on freely drained upland s i t e s , as a res u l t of the cool, humid, rainy climate in combination with the thick surface organic accumulations common to the s o i l s of the zone. Scrub forests of Thuja p l i c a t a ^ (red cedar), Chamae- cyparis nootkatensis (yellow cedar), Tsuga heterophylla (western hemlock) and lesser amounts of Tsuga mertensiana (mountain hemlock) and Pinus contorta (shore pine) predominate on moderate slopes throughout the zone on wet F o l i s o l s or mineral s o i l s (Podzols, Gleysols, Brunisols) having thick (20-70cm) surface organic accumulations. Picea sitchensis (Sitka spruce), Abies amabilis (amabilis f i r ) and Alnus rubra (red alder) also occur, but are r e s t r i c t e d to better-drained, steep, c o l l u v i a l slopes and f l u v i a l landforms. Productive forest stands are mostly r e s t r i c t e d to these landforms. Bog and bog woodland ecosystems are ch a r a c t e r i s t i c on the le v e l and gently sloping t e r r a i n in the CCPH, and in some portions of the zone they also occur on considerable slopes (15 to 60%). 3.2 Ge o g r a p h i c L o c a t i o n o f th e Study T r a n s e c t s The study area is located between Prince Rupert and Rainbow Lake, 25 km inland along Highway 16 (Fig. 2). Fig. 3 ^•Species nomenclature follows Taylor and MacBryde (1977) for the vascular plants, Ireland et a l . (1980) for the mosses, Worley (1970) for the liverworts and Otto and Ahti (1967) for the 1ichens,except for the genus Cladonia which follows Thomson (1967). A species l i s t for the study area is presented in Appendix IV. 18 F i g . 2. L o c a t i o n s o f t h e f o u r s t u d y t r a n s e c t s n e a r P r i n c e R u p e r t , B r i t i s h C o l u m b i a a r e shown b y ) ^ The dashed l i n e r e p r e s e n t s t h e t r a n s i t i o n f r o m m e t a m o r p h i c (M) t o i g n e o u s (I) b e d r o c k . 19 F i g . 3. O b l i q u e a i r pho to o f the s t u d y a r e a ; l o o k i n g n o r t h w e s t o v e r Ra inbow L a k e . The a p p r o x i m a t e l o c a t i o n s o f the f o u r s t u d y t r a n s e c t s a r e i n d i c a t e d by a r r o w s , The haze between T3 and T4 i s no t f o g b u t r a t h e r e m i s s i o n s f r om t h e p u l p m i l l s o u t h o f P r i n c e R u p e r t . 20 i s an oblique a i r photo of the Rainbow Lake portion of the study area. F i e l d work was carried out over the summer (May -Sept.) of 1979. After an i n i t i a l ground and a i r recon-naissance of the area in March and May of 1979, together with examination of topographic maps, geologic maps, forest cover maps and a i r photographs, four transect locations were chosen for intensive study (Figs. 2 and 3). Transect #1 (Rainbow East or RE) begins at the northeast shore of Rainbow Lake and runs approximately 1.5 km in a southeast d i r e c t i o n . Transect #2 (Rainbow West or RW) begins on the northwest side of Rainbow Lake opposite transect #1 and extends 1.5 km in a northwest d i r e c t i o n p a r a l l e l to Highway 16. Transect #3 (Diana Creek or DC) extends 2.5 km in a northwest d i r e c t i o n from the Diana Creek road approximately 1 km south of Highway 16. Transect #4 (Hayes Mountain or HM) begins 0.5 km south of Oliver Lake Park along Highway 16 on Kaien Island and extends 2 km westward up Mt. Hayes to an elevation of 300 m. Sampling was mostly r e s t r i c t e d to the transects but in a few cases i t was necessary to locate plots at some distance from the transects to find examples of ecosystems large enough and homogeneous enough to sample. A l l releve sampling was done below 290 m elevation, although reconnaissance was done at higher elevations (up to 700 m) on Mt. Hayes to examine the montane and subalpine t r a n s i t i o n s . 21 3.3 P h y s i o g r a p h y and Geology M o s t o f t h e s t u d y a r e a i s i n c l u d e d w i t h i n t h e K i t i m a t R a n g e s o f t h e C o a s t M o u n t a i n s , a l t h o u g h much o f K a i e n I s l a n d w e s t o f Mt. Hayes ( i n c l u d i n g P r i n c e R u p e r t ) i s i n c l u d e d i n t h e H e c a t e L o w l a n d o f t h e C o a s t a l T r o u g h ( H o l l a n d 1 9 7 6 ) . The K i t i m a t R a n g e s , one o f t h r e e m a j o r d i v i s i o n s o f t h e C o a s t M o u n t a i n s , e x t e n d f r o m t h e N a s s R i v e r t o B u r k e C h a n n e l and B e l l a C o o l a R i v e r and a r e b o r d e r e d t o t h e n o r t h by t h e B o u n d a r y Ranges and t o t h e s o u t h by t h e P a c i f i c R a n g e s . They i n c l u d e t h e h e a v i l y g l a c i a t e d g r a n i t i c m o u n t a i n s o f t h e n e a r - s h o r e a r c h i p e l a g o and m a i n l a n d e x t e n d i n g f r o m s e a l e v e l t o 1875-2500 m. ( H o l l a n d 1 9 7 6 ) . The C o a s t a l T r o u g h f l a n k s t h e C o a s t M o u n t a i n s and e x t e n d s f r o m P u g e t Sound t o D i x o n E n t r a n c e . The H e c a t e D e p r e s s i o n c o m p r i s e s t h a t p o r t i o n o f t h e C o a s t a l T r o u g h b e t w e e n J o h n s t o n e S t r a i t and D i x o n E n t r a n c e , much o f w h i c h i s b e l o w s e a l e v e l . K a i e n I s l a n d i s i n c l u d e d i n t h e H e c a t e L o w l a n d , a b e l t o f l o w - l y i n g t o p o g r a p h y i n c l u d i n g t h e o u t e r m a i n l a n d c o a s t and a d j a c e n t i s l a n d s e x t e n d i n g f r o m P r i n c e R u p e r t t o n o r t h e r n V a n c o u v e r I s l a n d ( H o l l a n d 1 9 7 6 ) . The b e d r o c k g e o l o g y o f t h e s t u d y a r e a i s d e s c r i b e d by H u t c h i s o n (1966) and H u t c h i s o n e t a l . ( 1 9 7 9 ) . The a r e a b e t w e e n P r i n c e R u p e r t and D i a n a C r e e k , 18 km i n l a n d , i s d o m i n a t e d by t h i n l y b e dded m e t a s e d i m e n t a r y r o c k s o f P a l e o z o i c age t h a t r e p r e s e n t t h e s o u t h e r l y e x t e n s i o n o f t h e W r a n g e l l - R e v i l l a g i g e d o b e l t o f m e t a m o r p h i c r o c k s i n A l a s k a . T h i s map u n i t i s d o m i n a t e d by t h e h i g h e r g r a d e ( a l m a n d i n e a m p h i b o l i t e f a c i e s ) r o c k s c o n t a i n i n g h o r n b l e n d e b i o t i t e , 22 b i o t i t e garnet, amphibolite schists and gniesses, layered quartz feldspar schists and impure quartzites. Just east of Diana Creek there i s an abrupt t r a n s i t i o n to the igneous intrusive bedrock that characterizes the Coast Mountains. Poorly f o l i a t e d or massive quartz d i o r i t e dominate thi s portion of the study area. The t r a n s i t i o n from meta-morphic to igneous bedrock i s indicated in Fig. 2. The existence of these two contrasting bedrock types within the study area afforded the opportunity to examine the influence of bedrock on vegetation and s o i l development. Transects #1 and #2 are located over quartz d i o r i t e bedrock while Transects #3 and #4 occur over the metamorphic bedrock. Dominant landforms within the study area include organic blankets (>1 m) and veneers (<1 m) over bedrock or uncon-solidated mineral materials (bedrock-derived or f l u v i a l ) , c o l l u v i a l veneers and blankets, and f l u v i a l veneers and blankets. Extensive areas of g l a c i a l t i l l are uncommon in the study area 1 although small localized pockets were encountered. 3.4 C l i m a t e Monthly climatic s t a t i s t i c s from the Atmospheric Environ-ment station at the Prince Rupert airport (Atmospheric Environment Services, Canada Department of the Environment 1970) are summarized in Fig. 4. The climate of the Prince Rupert area is most simply described as cool and wet throughout the year. It i s influenced primarily by Maritime Polar and Maritime A r c t i c a i r masses and very occasionally during the winter by Continental A r c t i c a i r moving down the 23 F i g . 4. C l i m a t i c d i a g r a m f o r t h e P r i n c e R u p e r t a i r p o r t A t m o s p h e r i c E n v i r o n m e n t s t a t i o n . MONTH a S t a t i o n . b Height above sea l e ve l (metres) . -c Durat ion (years) o f obse rva t ions , ( [ad j j - Adjusted to the nearest long pe r i od s t a t i o n . ) d Mean annual temperature, e Mean annual p r e c i p i t a t i o n . f Mean d a i l y minimum temperature o f the c o l d e s t month, g Lowest temperature recorded . h Mean d a i l y maximum temperature o f the warmest month. i H ighest temperature recorded . k Curve (dotted) o f mean monthly temperature. 1 Curve o f mean monthly p r e c i p i t a t i o n . m Mean monthly r a i n > 100 mm. (black s ca l e reduced by 1/10). n Months with mean d a i l y minimum temperature < 0 ° C (b lack ) , o Months with abso lute temperature < 0 ° C ( f r o s t s occur ) , p Mean durat ion (days) o f f r o s t f ree pe r i od . 24 Skeena Valley from the i n t e r i o r (Wilson and Marsh 1975). The mean annual temperature at Prince Rupert is 7.7°C. August is the warmest month with a mean of 13.9°C. while January is the coldest month with a mean of 1.7°C. There are, on average, 199 f r o s t free days during the year at Prince Rupert. The average annual accumulated growing degree days (above 5°C) at Prince Rupert are 1359 (Aston 1978). However, the number of e f f e c t i v e growing degree days, a value that assumes an optimum temperature for growth ( i . e . , 25°C) and a decrease in growth as temperatures vary from th i s optimum (Coligada 1981) i s t y p i c a l l y lower than growing degree days for coastal l o c a l i t i e s . Mean annual p r e c i p i t a t i o n is high, ranging from 2414 mm to 2907 mm at the three Prince Rupert stations. Summer is the d r i e s t season (15% of the t o t a l precipitation) while autumn is the wettest (34% of the t o t a l ) . Only 4.5% of the t o t a l p r e c i p i t a t i o n f a l l s as snow. Mean r e l a t i v e humidity values (at 1630 hours) remain above 75% for every month except March, A p r i l and May (Kendrew and Kerr 1955). Annual potential evapotranspiration (PET) at Prince Rupert is low (156 mm). Maximum values occur in July (37.1 mm) while October through February have zero PET values ( B r i t i s h Columbia Ministry of Environment 1980). 25 CHAPTER 4 DESCRIPTION AND CLASSIFICATION OF COMMUNITY TYPES 4.1 I n t r o d u c t i o n I n t h i s c h a p t e r , t h e v e g e t a t i o n , s o i l and l a n d f o r m c h a r a c t e r i s t i c s o f t h e m a j o r bog and f o r e s t e c o s y s t e m u n i t s o c c u r r i n g a l o n g t h e f o u r s t u d y t r a n s e c t s a r e d e s c r i b e d . T h e s e e c o s y s t e m u n i t s w i l l be c a l l e d 'community t y p e s ' . I n t h e B r a u n - B l a n q u e t a p p r o a c h t o v e g e t a t i o n c l a s s i f i c a t i o n ( M u e l l e r - D o m b o i s and E l l e n b e r g 1974; W e s t h o f f and v a n d e r M a a r e l 1978) a v e g e t a t i o n u n i t i s r e f e r r e d t o as a com m u n i t y t y p e i f i t h a s n o t y e t been g i v e n s y s t e m a t i c r a n k o r h i e r a r c h i c a l s t a t u s w i t h i n t h e c l a s s i f i c a t i o n s y s t e m ( i . e . O r d e r , A l l i a n c e , A s s o c i a t i o n , S u b a s s o c i a t i o n e t c . ) . T h i s was c o n s i d e r e d t h e b e s t a p p r o a c h i n t h i s s t u d y b e c a u s e t h e v e g e t a t i o n was n o t s a m p l e d e x t e n s i v e l y enough t o d e f i n e i n d i v i d u a l a s s o c i a t i o n s i n many c a s e s . Some community t y p e s c o u l d be c o n s i d e r e d a s s o c i a t i o n s , b u t o t h e r s ( e s p e c i a l l y t h e o p e n bog c o m m u n t i t e s ) a r e s u f f i c i e n t l y v a r i a b l e t o be d e s c r i b e d as a l l i a n c e s . U n t i l more s a m p l i n g i s done, t h i s v a r i a b i l i t y i s b e t t e r d i s c u s s e d i n t e r m s o f t r e n d s w i t h i n e a c h o f t h e community t y p e s r a t h e r t h a n a t t e m p t i n g t o d e s c r i b e a l a r g e number o f a s s o c i a t i o n s b a s e d on one o r a few r e l e v e s . The o r d i n a t i o n o f r e l e v e s l e n d s i t s e l f t o t h i s a p p r o a c h b e c a u s e t h e r e l a t i o n s h i p s among s a m p l e p l o t s w i t h i n a community t y p e c a n be d i s c u s s e d i n t e r m s o f t h e i r r e l a t i v e p o s i t i o n on t h e s c a t t e r d i a g r a m s . 26 This approach follows the philosophy of Mueller-Dombois and Ellenberg (1974) "....we consider i t useful to maintain an unsystematic status for abstract vegetation communities in a l l cases where the emphasis is on intensive l o c a l vegetation studies. However, an h i e r a r c h i c a l scheme becomes very desirable where the emphasis l i e s on developing a vegetation synopsis at a more extensive geographical scale". The B.C. Ministry of Forests i s currently combining the releve' data from this study with data from other areas along the mainland coast and Queen Charlotte Islands to produce a more detailed ecosystem c l a s s i f i c a t i o n for the CCPH zone. Using the c l a s s i f i c a t i o n and ordination of the 40 sample plots and the description of community types as a framework, vegetation-environment relationships within the study area are discussed and compared with other studies of bog and forest ecosystems. 4 .2 F i e l d Methods Following the Braun-Blanquet approach (Westhoff and van der Maarel 1978), homogeneous examples of the major climax vegetation units were subjectively chosen for releve sampling along each of the study transects and a t o t a l of 40 releves were established. Plot size varied from 180 m2 (12 m x 15 m) in the open bog communities to 400 m2 (20 m x 20 m) in the closed forests. Smaller plot sizes were used in the open bog habitats because within-plot v a r i a b i l i t y ( i . e . , micro-community and tree species d i s t r i b u t i o n patterns) tended to occur over smaller areas in the bogs than in the forests. 27 I n o r d e r t o e n s u r e d a t a c o m p a t i b i l i t y w i t h t h e e c o l o g i c a l c l a s s i f i c a t i o n w ork c u r r e n t l y b e i n g done b y t h e B r i t i s h C o l u m b i a M i n i s t r y o f F o r e s t s , e c o s y s t e m d e s c r i p t i o n f o r m s s i m i l a r t o t h o s e b e i n g u s e d b y g o v e r n m e n t a g e n c i e s i n B.C. ( W a l m s l e y e t a l . 1980) w e r e u s e d f o r r e c o r d i n g d a t a w i t h i n e a c h o f t h e r e l e v e s . A c o m p l e t e d s e t o f f i e l d f o r m s f o r one r e l e v e i s p r e s e n t e d i n A p p e n d i x I I . G e o g r a p h i c and e n v i r o n m e n t a l i n f o r m a t i o n was r e c o r d e d on t h e s i t e d e s c r i p t i o n f o r m s a nd n o t e s w e r e made o n any o t h e r i m p o r t a n t o r u n u s u a l f e a t u r e s w i t h i n t h e s a m p l e p l o t . A c o m p l e t e s p e c i e s l i s t was made and e a c h s p e c i e s e v a l u a t e d u s i n g t h e 10 p o i n t D o m i n - K r a j i n a c o v e r - a b u n d a n c e s c a l e a n d a 4 - p o i n t v i g o u r s c a l e ( A p p e n d i x I I I ) . The v e g e t a t i o n was d i v i d e d i n t o 11 s t r a t a ( A p p e n d i x I I I ) and e a c h s t r a t u m e v a l u a t e d s e p a r a t e l y . T o t a l p e r c e n t c o v e r was a l s o e s t i m a t e d f o r e a c h s t r a t u m . S p e c i e s w h i c h w e re n o t i d e n t i f i e d i n t h e f i e l d w e r e g i v e n a number a n d c o l l e c t e d f o r l a t e r i d e n t i f i c a t i o n . V o u c h e r s p e c i m e n s o f most o f t h e s p e c i e s a r e l o c a t e d i n t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a H e r b a r i u m . Two s p e c i m e n s o f e a c h t r e e s p e c i e s p r e s e n t i n a p l o t w e r e c h o s e n f r o m t h e m a i n c a n o p y , a nd t h e i r a ge, h e i g h t and d i a m e t e r a t b r e a s t h e i g h t (dbh) d e t e r m i n e d . Two p r i s m c r u i s e p l o t s w e r e a l s o done w i t h i n t h e m a i n s a m p l e p l o t f o r b a s a l a r e a d e t e r m i n a t i o n s . W i t h i n e a c h p l o t one s o i l p i t was dug ( i n t h e c a s e o f m i n e r a l s o i l s and some o f t h e o r g a n i c s o i l s ) o r a p e a t c o r e 28 was t a k e n u s i n g a H i l l e r p e a t s a m p l e r . The s o i l h o r i z o n s were d e s c r i b e d a c c o r d i n g t o s t a n d a r d p r a c t i c e ( W a l m s l e y e t a l . 1980) a n d t h e humus f o r m s w ere d e s c r i b e d a n d c l a s s i f i e d t o s u b g r o u p u s i n g K l i n k a e t a l . ( 1 9 8 1 ) . The s o i l s w e r e c l a s s i -f i e d t o s u b g r o u p 1 a c c o r d i n g t o t h e Ca n a d a S o i l S u r v e y C o m m i t t e e (1978) a s w e l l a s t h e r e v i s i o n s r e c e n t l y p r o p o s e d f o r F o l i s o l s a n d f o l i s o l i c p h a s e s ( T r o w b r i d g e 1 9 8 1 ) . I t was a p p a r e n t d u r i n g t h e s a m p l i n g t h a t t h e d e s c r i p t i o n o f one "m o d a l " s o i l p i t d i d n o t p o r t r a y t h e o f t e n l a r g e e d a p h i c v a r i a b i l i t y w i t h i n e a c h p l o t . N o t e s w ere t h e r e f o r e made on t h e d e g r e e a n d n a t u r e o f t h i s v a r i a b i l i t y . I n r e l e v e s on d e e p e r o r g a n i c v e n e e r s a n d b l a n k e t s , t h r e e o r f o u r p e n e t r a -t i o n s w e r e made t h r o u g h o u t t h e p l o t ( u s i n g t h e H i l l e r s a m p l e r ) t o d e t e r m i n e t h e r a n g e i n d e p t h o f t h e d e p o s i t . Any u n c o n -s o l i d a t e d m i n e r a l s u b s t r a t e u n d e r t h e p e a t m a t e r i a l was d e s c r i b e d a n d s a m p l e d i n most c a s e s . S a m p l e s f r o m s o i l a n d humus h o r i z o n s w e re c o l l e c t e d a n d s t o r e d f o r f u t u r e a n a l y s i s . O n l y pH d a t a a r e i n c l u d e d i n t h i s t h e s i s . PH was d e t e r m i n e d a f t e r d i l u t i o n o f a i r - d r i e d s a m p l e s w i t h H2O ( J a c k s o n 1 9 5 8 ) . O r g a n i c h o r i z o n s r e q u i r e d g r e a t e r d i l u t i o n ( 1 : 2 0 f o r L and Of h o r i z o n s ; 1:10 f o r H and Oh h o r i z o n s ) t h a n d i d m i n e r a l h o r i z o n s ( 1 : 5 ) . 4 . 3 M e t h o d s o f D a t a A n a l y s i s The r e l e v e d a t a w e re a n a l y z e d u s i n g s e v e r a l t e c h n i q u e s . A t a b u l a r c o m p a r i s o n t e c h n i q u e f o l l o w i n g t h e b a s i c p r i n c i p l e s S o i l c l a s s i f i c a t i o n i s b a s e d on m o r p h o l o g i c a l f e a t u r e s o n l y . The v o n P o s t s c a l e o f d e c o m p o s i t i o n was u s e d t o c l a s s i f y o r g a n i c s o i l s . o f t h e B r a u n - B l a n q u e t a p p r o a c h ( M u e l l e r - D o m b o i s a nd E l l e n b e r g 1974) was u s e d t o g e t h e r w i t h t h r e e m a t h e m a t i c a l o r d i n a t i o n t e c h n i q u e s ; p o l a r o r d i n a t i o n ( C o t t a m e t a l . 1 9 7 8 ) , p r i n c i p a l c o m p o n e n t s a n a l y s i s ( O r l o c i 1966) and r e c i p r o c a l a v e r a g i n g ( H i l l 1 9 7 3 ) . 4.3.1 T a b u l a r Comparison A c o m p u t e r p r o g r a m d e v e l o p e d f o r t h e B.C. M i n i s t r y o f F o r e s t s ( K l i n k a and P h e l p s 1979) was u s e d t o p r o d u c e v e g e t a t i o n t a b l e s f r o m t h e r e l e v e d a t a ( A p p e n d i x V I ) . The 40 r e l e v e s w e r e t h e n s u b j e c t i v e l y c ompared w i t h t h e a i d o f t h e s e t a b l e s . By e x a m i n i n g s p e c i e s c o n s t a n c y (% o c c u r r e n c e i n a l l t h e r e l e v e s ) , f i d e l i t y ( d i s t r i b u t i o n among t h e r e l e v e s ) , c o v e r - a b u n d a n c e and v i g o u r , and by e x a m i n i n g t h e e n v i r o n m e n t a l c h a r a c t e r i s t i c s o f e a c h o f t h e r e l e v e s ( A p p e n d i x V I I I ) , t h e i n i t i a l f i e l d s t r a t i f i c a t i o n was t e s t e d and r e f i n e d . S i x c o m m u n i t y t y p e s w e r e t h e n d e f i n e d u s i n g a c h a r a c t e r i s t i c c o m b i n a t i o n o f s p e c i e s t o g e t h e r w i t h h a b i t a t f e a t u r e s . The i d e n t i f i c a t i o n o f c h a r a c t e r s p e c i e s f o l l o w s t h e p r o c e d u r e s o f B r o o k e e t a l . ( 1 9 7 0 ) and W e s t h o f f and v a n d e r M a a r e l ( 1 9 7 8 ) . C o n s t a n t s p e c i e s (C) a r e t h o s e w h i c h a r e p r e s e n t i n more t h a n 8 0 % o f t h e r e l e v e s o f a community t y p e . A c o n s t a n t s p e c i e s i s c o n s i d e r e d a c o n s t a n t d o m i n a n t (CD) i f i t s a v e r a g e c o v e r - a b u n d a n c e i n t h a t c o m m u n i t y t y p e i s 3 o r g r e a t e r . C h a r a c t e r s p e c i e s a r e f u r t h e r d e f i n e d a c c o r d i n g t o t h e i r f i d e l i t y . B r a u n - B l a n q u e t ( W e s t h o f f and v a n d e r M a a r e l 1978) d i s t i n g u i s h e s f i v e d e g r e e s o f f i d e l i t y : A. C h a r a c t e r T a x a E x c l u s i v e T a x a ( f i d e l i t y d e g r e e 5 ) : t a x a c o m p l e t e l y o r 30 almost completely c o n f i n e d to one v e g e t a t i o n u n i t . S e l e c t i v e Taxa ( f i d e l i t y degree 4): taxa with c l e a r p r e f e r e n c e f o r one v e g e t a t i o n u n i t but a l s o o c c u r r i n g i n other v e g e t a t i o n u n i t s . P r e f e r e n t i a l Taxa ( f i d e l i t y degree 3): taxa present i n s e v e r a l v e g e t a t i o n u n i t s , perhaps with about equal presence but with a higher combined e s t i m a t i o n value and/or with a higher v i t a l i t y (vigour) i n one p a r t i c u l a r v e g e t a t i o n u n i t . B. Companions: I n d i f f e r e n t Taxa ( f i d e l i t y degree 2) : taxa without pronounced pr e f e r e n c e f o r any v e g e t a t i o n u n i t . C. A c c i d e n t a l s Strange Taxa ( f i d e l i t y degree 1 ) : taxa having a d e f i n i t e presence optimum and mostly a l s o a cover-abundance optimum o u t s i d e the considered v e g e t a t i o n u n i t . These are o f t e n a c c i d e n t a l i n t r u d e r s from neigh-b o r i n g communities or r e l i c t s from a community that preceded i n s u c c e s s i o n . In a d d i t i o n to producing v e g e t a t i o n releve* t a b l e s , the M i n i s t r y of F o r e s t s computer program w i l l a l s o produce a v e g e t a t i o n summary or s y n t h e s i s t a b l e that combines the r e l e v e s of each v e g e t a t i o n u n i t and l i s t s a constancy value as w e l l as a mean cover-abundance value f o r each s p e c i e s w i t h i n t h a t u n i t (Appendix V I ) . Constancy i s d i v i d e d i n t o f i v e c l a s s e s : 1-20% ( I ) , 21-40% ( I I ) , 41-60% ( I I I ) , 61-80% (IV), and 81-100% (V) . T h i s t a b l e i s u s e f u l f o r i d e n t i f y i n g the 31 character species for each community type. D i f f e r e n t i a l species may also be i d e n t i f i e d and used to define further d i v i s i o n s within a vegetation unit (e.g., sub-associations within associations). As outlined in Section 4.1, community types are i d e n t i f i e d in this study without reference to systematic rank. In this case, certain species may be i d e n t i f i e d as d i f f e r e n t i a l species i f they appear to characterize a d i s t i n c t association within a broadly defined community type ( i . e . , an a l l i a n c e ) . 4 . 3 . 2 O r d i n a t i o n T e c h n i q u e s While the c l a s s i f i c a t i o n approach to vegetation analysis ( i . e . , the Braun - Blanquet approach) emphasizes the grouping of releves and the synthesis of discrete, abstract community types, ordination techniques emphasize the arrangement of samples (or species) along environmental gradients (either measured or inferred) (Whittaker 1978a and b). The purpose of ordination is to reduce the multi-dimensional variation e x i s t i n g among a group of sample plots and display i t in terms of the two or three most important dimensions. Most ordination techniques u t i l i z e c o e f f i c i e n t s of mathematical s i m i l a r i t y as well as matrix algebra, and the results are displayed using a scatter diagram usually of two dimensions (axes). Samples of r e l a t i v e s i m i l a r i t y in species composition, or species of r e l a t i v e s i m i l a r i t y in d i s t r i b u t i o n among the sample p l o t s , occur close to one another on the ordination. This enables the investigator to better v i s u a l i z e the data set and f a c i l i t a t e s communication of re s u l t s . 32 Although community types may be derived from an ordination of samples, the emphasis i s on the analysis of gradients rather than on the c l a s s i f i c a t i o n and character-i z a t i o n of communities. Used together, c l a s s i f i c a t i o n and ordination should r e s u l t in a better understanding of the var i a t i o n existing in the releve data and the relationships among community types. In a separate report (Banner 1980), three ordination techniques, polar ordination, p r i n c i p a l components analysis and r e c i p r o c a l averaging, were compared as to their usefulness in c l a r i f y i n g relationships among the 40 releves. This com-parison w i l l not be dealt with here and only those ordinations considered to be most useful w i l l be presented. The following outline of the concepts behind the three ordination techniques i s taken from Banner (1980). There is a multitude of references that give detailed conceptual and com-putational descriptions of these methods (Bray and Curtis 1957; Austin and O r l o c i 1966; Orloci 1966, 1975; Gi t t i n s 1969; H i l l 1973; Noy-Meir 1973; Mueller-Dombois and Ellenberg 1974; Noy-Meir et a l . 1975; Gauch et a l . 1977; Whittaker 1978a; Kenkell 1980). The emphasis w i l l be on the usefulness of each method in c l a r i f y i n g relationships rather than on the mathe-matics involved. The computer package ORDIFLEX (Gauch 1977), available from the Cornell Ecology Program Series, was used to apply the ordination techiques to the data. This program contains a l l three ordinations as well as a multitude of options for data 33 t r a n s f o r m a t i o n s , s t a n d a r d i z a t i o n s , and s a m p l e o r s p e c i e s d e l e t i o n s f r o m t h e a n a l y s i s . 4 . 3 . 2 . 1 P o l a r O r d i n a t i o n (PO) B r a y and C u r t i s (1957) d e v i s e d t h i s c o m p u t a t i o n a l l y s i m p l e and s t r a i g h t f o r w a r d method o f o r d i n a t i o n . S a m p l e s t h a t o c c u r a t o p p o s i t e e n d s o f e n v i r o n m e n t a l o r f l o r i s t i c g r a d i e n t s a r e c h o s e n as e n d p o i n t s , and d i s t a n c e m e a s u r e s a r e c a l c u l a t e d f o r e a c h e n d p o i n t - s a m p l e p a i r . The t h r e e i n t e r p o i n t d i s t a n c e s t h u s c a l c u l a t e d d e f i n e a t r i a n g l e . The p o s i t i o n o f t h e s a m p l e c a n t h e n be p e r p e n d i c u l a r l y p r o j e c t e d o n t o t h e o r d i n a t i o n a x i s b e t w e e n t h e two e n d p o i n t s . O t h e r e n d p o i n t s c a n be c h o s e n f o r a s e c o n d a x i s ( f r o m t h e s a m p l e s i n t h e c e n t r a l r e g i o n o f t h e f i r s t a x i s ) and a two d i m e n s i o n a l o r d i n a t i o n c a n be c o n -s t r u c t e d . S e v e r a l d i s t a n c e m e a s u r e s may be u s e d , and t h e c h o i c e o f e n d p o i n t s may be done s u b j e c t i v e l y o r be b a s e d on d i s t a n c e c a l c u l a t i o n s b e t w e e n e a c h s a m p l e p a i r i n t h e d a t a s e t . I n t h i s s t u d y , p e r c e n t a g e d i s t a n c e ( t h e most commonly u s e d m e a s u r e ) was u s e d and t h e c h o i c e o f e n d p o i n t s was b a s e d on t h e s e c o n d a r y m a t r i x o f d i s t a n c e c a l c u l a t i o n s ( a u t o m a t i c B r a y - C u r t i s m e t h o d ) . 4 . 3 . 2 . 2 P r i n c i p a l C o m p o n e n t s A n a l y s i s (PCA) P r i n c i p a l C omponents A n a l y s i s i s somewhat more a b s t r a c t and t h u s more d i f f i c u l t t o co m p r e h e n d . The e a s i e s t way t o u n d e r s t a n d i t i s i n g e o m e t r i c t e r m s . A p r i m a r y d a t a m a t r i x r e p r e s e n t s a two d i m e n s i o n a l a r r a y o f s a m p l e s and s p e c i e s . T h e s e same d a t a c a n be p o r t r a y e d as p o i n t s i n a m u l t i d i m e n s i o n a l s p a c e , e a c h p o i n t r e p r e s e n t i n g a 34 s a m p l e ( o r s p e c i e s ) . V e c t o r s g o i n g f r o m t h e o r i g i n t o t h e p o i n t s may a l s o be u s e d t o r e p r e s e n t t h e p o s i t i o n o f s a m p l e s i n t h e s p e c i e s s p a c e ( o r t h e s p e c i e s i n t h e s a m p l e s p a c e ) . The c o r r e l a t i o n b e t w e e n any two p o i n t s c a n be r e p r e s e n t e d by t h e c o s i n e o f t h e a n g l e b e t w e e n t h e two v e c t o r s . A s e c o n d a r y d a t a m a t r i x o f c o r r e l a t i o n c o e f f i c i e n t s c a n t h e n be c o n -s t r u c t e d . S u c h a r e p r e s e n t a t i o n w o u l d h a v e f a r t o o many d i m e n s i o n s t o v i s u a l i z e , and t h i s i s where PCA p l a y s a r o l e . The p u r p o s e o f PCA i s t o i l l u s t r a t e t h e d o m i n a n t and i m p o r t a n t d i m e n s i o n s o f v a r i a t i o n i n t h e a r r a y o f p o i n t s . I f most o f t h e v a r i a t i o n i n t h e d a t a s e t i s c o n c e n t r a t e d i n two o r t h r e e d i m e n s i o n s , PCA w i l l d i s p l a y t h i s u s i n g new a x e s w h i c h go t h r o u g h t h e l o n g e s t p o i n t c l u s t e r s . The d i r e c t i o n o f t h e f i r s t a x i s t h r o u g h t h e p o i n t s i s s u c h t h a t t h e v a r i a n c e o f t h e d i s t a n c e s o f t h e p o i n t s o f f t h e a x i s i s m i n i m i z e d . T h i s i s e q u i v a l e n t t o t h e l e a s t s q u a r e s l i n e t h r o u g h t h e p o i n t s . S u b s e q u e n t a x e s c a n be c o n s t r u c t e d t o a c c o u n t f o r r e m a i n i n g d i r e c t i o n s o f v a r i a t i o n i n t h e p o i n t c l u s t e r s . The amount o f v a r i a n c e r e p r e s e n t e d by e a c h o f t h e PCA a x e s i s t e r m e d t h e ' e i g e n v a l u e ' , w h i l e t h e ' e i g e n v e c t o r ' r e f e r s t o t h e d i r e c t i o n c o s i n e t h a t s p e c i f i e s t h e PCA a x i s . E i g e n v e c t o r s and e i g e n v a l u e s a r e c a l c u l a t e d ( u s i n g m a t r i x a l g e b r a ) f r o m t h e s e c o n d a r y c r o s s - p r o d u c t s m a t r i x d e r i v e d f r o m t h e o r i g i n a l d a t a m a t r i x . The n a t u r e o f t h e c r o s s - p r o d u c t s m a t r i x d e p e n d s on what s o r t o f a d j u s t m e n t s o r t r a n s f o r m a t i o n s a r e a p p l i e d t o t h e d a t a m a t r i x , and t h i s i n t u r n w i l l a f f e c t 35 the f i n a l d i s p l a y of p o i n t s about the PCA axes. If the o r i g i n a l data values are u n a l t e r e d ( i . e . , cover-abundance values from 1-9), then the cr o s s - p r o d u c t s matrix i s a d i s p e r s i o n matrix (PCA non-centered). S u b t r a c t i o n of the s p e c i e s means from each species value centres the data and a v a r i a n c e - covariance matrix i s c a l c u l a t e d (PCA c e n t e r e d ) . T h i s i m p l i e s that a s p e c i e s i s important only so f a r as i t s value i n a sample d i f f e r s from the average of that s p e c i e s i n a l l the samples. Cen t e r i n g by samples i s a l s o p o s s i b l e . If the s p e c i e s values are centered and subsequently d i v i d e d by the standard d e v i a t i o n f o r that s p e c i e s , the cros s - p r o d u c t s matrix i s a c o r r e l a t i o n matrix (PCA s t a n d a r d i z e d ) . T h i s has the e f f e c t of i n c r e a s i n g the importance of rare s p e c i e s or s p e c i e s t h a t have c h a r a c t e r i s t i c a l l y low cover v a l u e s . Noy-Meir (1973) and Noy-Meir et a l . (1975) examine the e f f e c t s of these and other data t r a n s f o r m a t i o n s on PCA o r d i n a t i o n s of s e v e r a l data s e t s . 4.3.2.3 R e c i p r o c a l Averaging (RA) Although M.O. H i l l i s u s u a l l y c r e d i t e d with i n t r o d u c i n g the technique of r e c i p r o c a l averaging, v i r t u a l l y the same technique under the name "Analyse F a c t o r i e l l e des Correspondances" was e a r l i e r d e s c r i b e d by J.P. B e n z e c r i and othe r French workers ( H i l l 1973). C o n c e p t u a l l y , RA i s r e l a t e d to the g r a d i e n t a n a l y s i s technique of weighted averages (Whittaker 1967), although computationally i t a l s o has s i m i l a r i t i e s to PCA. In the technique of weighted averages, s p e c i e s are given scores 36 according to their 'preferred', position along a gradient ( i . e . , 1 - 4 along a moisture gradient) and samples are scored according to the r e l a t i v e abundance and scores of the species that they contain. The r e s u l t i s a simple ordination of samples based on the known ecological preferences of the species which they contain. A similar technique is applied in RA except that the i n i t i a l species scores are ar b i t r a r y . The sample scores thus obtained are subsequently used to assign new species scores, which in turn are used to reordinate the samples. Each time thi s is done the scores are rescaled from 0-100. Eventually, and regardless of the i n i t i a l species scores, the scores w i l l s t a b i l i z e to a unique solution. A one dimensional ordination of both species and samples r e s u l t s . Second and higher axes are solved by choosing i n i t i a l scores near to the f i n a l ones for the f i r s t axis, as well as making other adjustments for the f i r s t axis solution ( H i l l 1973). The term 'reciprocal averaging' i s derived from the fact that the species scores are averages of the releve scores and the releve scores are averages of the species scores. It is possible to arrive at the same solution by using an eigenvector technique s i m i l a r to PCA. Hill(1973) and Gauch et a l . (1977) i l l u s t r a t e the RA procedure together with the conceptual and computational d i s t i n c t i o n s between RA and PCA. 4.3.2.4 Data A d j u s t m e n t s t o Produce t h e O r d i n a t i o n s . Before the species cover-abundance values were used to produce the ordinations, the following adjustments had to be 37 made to the raw f i e l d data. The veteran and Al tree layers were combined and only the bryophytes and lichens which were growing on humus were included in the analysis. Species recorded in the f i e l d as ' + ' were rescored as '1' and species occurring in only one releve were excluded from the analysis. These adjustments to the f i e l d data reduced the o r i g i n a l 286 species-layer combinations (191 species in 12 layers) to 178 (143 species in 7 layers) i n the 40 releves. 4.4 R e s u l t s 4.4.1 T a b u l a r Comparison and O r d i n a t i o n R e s u l t s Tabular comparison plus examination of the ordinations resulted in the d e f i n i t i o n and characterization of six community types from the forty r e l e v l s . Vegetation and envir-onmental tables for each of the community types are presented in Appendix VI and VIII respectively. Scatter diagrams of axes 1 and 2 for the PO, PCA (standardized) and RA ordinations of the 40 releves are presented in Figs. 5 - 7 . The PCA (standardized) and the RA releve' ordinations of the entire data set were found to be the easiest to interpret f l o r i s t i c a l l y and e c o l o g i c a l l y . The PO ordination (Fig. 5) i s somewhat bipolar with the open bogs at one extreme and the forest and woodlands at the other. The d i s t i n c t i o n s between the forest and woodland communities, however, are not as clear in this ordination as in the RA and PCA. The PCA (Fig. 6) and RA (Fig. 7) ordinations d i f f e r 38 F i g . 5. P o l a r o r d i n a t i o n o f r e l e v e s . The community t y p e s on t h i s and s u b s e q u e n t o r d i n a t i o n s a r e d e s i g n a t e d as f o l l o w s : E r i c a c e o u s s h r u b - T r i c h o p h o r u m - Sphagnum open bogs ( Q ) ^ Empetrum - V a c c i n i u m u l i g i n o s u m - R h a c o m i t r i u m d r y o u t c r o p s ( ( ^ ) ; P i n e - c y p r e s s - c e d a r - G a u l t h e r i a -F a u r i a bog wood lands ( Q ) ; Ceda r - c y p r e s s - hemlock -V a c c i n i u m - Cornus z o n a l u p l a n d f o r e s t s Ceda r -s p r u c e - hemlock - L y s i c h i t o n a l l u v i a l f o r e s t s (\^))> A m a b i l i s f i r - hemlock - c e d a r - Gymnocarp ium u p l a n d f o r e s t s / 3?A 1 0 0 . 0 -9 0 . 0 8 0 . 0 7 0 . 0 2 8 ) ( 3 8 6 0 . 0 5 0 . 0 y A X I S 4 0 . 0 3 0 . 0 1 0 . 0 0 . 0 | . . 0 . 0 0 © . 1 . . . I. 1 0 . 0 2 0 . 0 2 9 3 5 A ..1 1.(6) I I . . . 3 0 . 0 4 0 . . 5 0 . 0 6 0 . 0 X A X I S 1 5 A A A A .i i i i 7 0 . 0 8 0 . 0 9 0 . 0 1 0 0 . 0 39 F i g . 6 . PCA ( s t a n d a r d i z e d ) o r d i n a t i o n o f r e l e v e s , axes 1 (x ) and . 2 ( y ) . Community t y p e s a re d e s i g n a t e d as i n F i g . 5. 1 0 0 . 0 9 0 . 0 r 0 . 3 1 t I I ...I .W| I I I I 0 . 0 1 0 . 0 2 C . 0 3 0 . 0 4 0 . 0 50h—1 6 0 . 0 7 0 . 0 8 0 . 0 9 0 . 0 1 0 X A X I S F i g . 7. RA o r d i n a t i o n o f r e l e v e s , axes 1 (x ) and 2 ( y ) . Community t y p e s a r e d e s i g n a t e d as i n F i g . 5. 41 somewhat in that groups of releves are better defined as e n t i t i e s (community types) in the PCA ordination, whereas in the RA ordination, the releves are displayed in a more continuous c u r v i l i n e a r fashion. This i l l u s t r a t e s that RA is oriented more towards gradient analysis while PCA i s better suited to taxometric (data summarization) ordinations (Gauch 1977). The ordering of releves and species along the f i r s t RA axis was useful in producing a rearranged data matrix (Appendix VII). The species and releve's are ordered simul-taneously in thi s ordination and the resultant species-by-sample matrix, with the largest values concentrated along the diagonal, proved useful in examining species d i s t r i b u t i o n among the releves and aided in the i d e n t i f i c a t i o n of releve groups and character species. The order of releves in the vegetation tables (Appendix VI) i s that produced by the f i r s t axis RA ordination. The species, however, are ordered by decreasing order of abundance within each layer. The PCA (standardized) species ordinations is presented in Fig. 8. This ordination is useful in comparing the the ecological relationships of the species within the study area. In the scatter diagram (Fig. 8) d i r e c t i o n from the centroid r e f l e c t s the d i s t r i b u t i o n of each species among the sample plots while distance from the centroid r e f l e c t s the r e l a t i v e constancy and/or dominance of each species. Species that occur together on the ordination thus have similar ecological preferences and are l i k e l y to co-occur in the same 42 F i g . 8. PCA ( s t a n d a r d i z e d ) s p e c i e s o r d i n a t i o n , axes 1 and 2. Names a r e a b b r e v i a t e d u s i n g t h e f i r s t two l e t t e r s o f genus and s p e c i e s . Gene ra u n i d e n t i f i e d t o s p e c i e s a r e a b b r e v i a t e d u s i n g f i r s t f o u r l e t t e r s f o l l o w e d by ' s p . 1 . L a y e r d e s i g -, - n a t i o n s a r e i n d i c a t e d be low t h o s e s p e c i e s o c c u r r i n g i n more t han one l a y e r . S p e c i e s o f h i g h e s t c o n s t a n c y and dominance have been d i v i d e d i n t o e i g h t g roups a c c o r d i n g t o t h e i r d i s t -r i b u t i o n among t h e community t y p e s . Comple te s p e c i e s l i s t s f o r each g roup a r e p r e s e n t e d i n A p p e n d i x V. F u r t h e r e x p l a n -a t i o n i n t e x t (page 4 1 ) . 4-2. A 100-r 43 c o m m u n i t y t y p e ( s ) . S p e c i e s w h i c h show e i t h e r h i g h c o n s t a n c y o r h i g h d o m i n a n c e v a l u e s ( i . e . , t h o s e o c c u r r i n g f u r t h e s t f r o m t h e c e n t r o i d ) h a v e b e e n g r o u p e d a c c o r d i n g t o t h e i r o c c u r r e n c e i n s p e c i f i c c o m m u n i t y t y p e s . Some s p e c i e s g r o u p s a r e r e s t r i c t e d m a i n l y t o j u s t one c o m m u n i t y t y p e ( e x c l u s i v e o r s e l e c t i v e t a x a ) w h i l e o t h e r s may o c c u r i n two o r t h r e e b u t be p r e f e r e n t i a l f o r one. S p e c i e s l y i n g b e t w e e n an o u t l i n e d g r o u p and t h e c e n t r o i d o f t h e o r d i n a t i o n u s u a l l y o c c u r i n t h e same co m m u n i t y t y p e ( s ) as t h o s e w i t h i n t h e g r o u p b u t hav e l o w e r c o n s t a n c y o r d o m i n a n c e . They a r e t h e r e f o r e l e s s u s e f u l as c h a r a c t e r s p e c i e s f o r community t y p e s , a l t h o u g h some may p r o v e u s e f u l as d i f f e r e n t i a l s p e c i e s f o r more n a r r o w l y d e f i n e d a s s o c i a t i o n s w i t h i n c o m m u n i t y t y p e s . E i g h t s p e c i e s g r o u p s (A-H) have b e e n o u t l i n e d on t h e PCA o r d i n a t i o n ( F i g . 8 ) . S p e c i e s a r e a b b r e v i a t e d u s i n g t h e f i r s t two l e t t e r s o f t h e ge n u s and s p e c i e s . A c o m p l e t e l i s t o f s p e c i e s f o r e a c h g r o u p i s g i v e n i n A p p e n d i x V. The s i x community t y p e s w i l l be d e s c r i b e d i n t h e o r d e r i n w h i c h t h e y o c c u r on t h e f i r s t a x i s RA o r d i n a t i o n . 4.4.2 D e s c r i p t i o n o f C o m m u n i t y T y p e s 4.4.2.1 E r i c a c e o u s S h r u b - T r i c h o p h o r u m c e s p i t o s u m - Sphagnum s p p . - C l a d o n i a s p p . - S i p h u l a c e r a t i t e s o p e n b o g s . Open bog c o m m u n i t i e s b l a n k e t much o f t h e l e v e l t o g e n t l y s l o p i n g l o w l a n d s w i t h i n t h e s t u d y a r e a ( F i g s . 9 and 10) and a l s o o c c u r , t h o u g h l e s s commonly, on c o n s i d e r a b l e s l o p e s (up 44 F i g . 9. B l a n k e t bog - u p l a n d f o r e s t complex n e a r Rainbow L a k e . M o s i a c o f open b l a n k e t bog and bog w o o d l a n d ^ i n f o r e g r o u n d ; u p l a n d f o r e s t on h i 11 s i opes i n t h e d i s t a n c e . F i g . 10. P i n u s c o n t o r t a - e r i c a c e o u s s h r u b - T r i chopho rum• et spHosum Sphagnum s p p . - C l a d o n i a s p p . - S i p h u l a cexatrtes open b l a n k e t bog commun i t y ; r e l e v e 7. T r a n s i t i o n t o bog wood land i n b a c k g r o u n d . 4+f\ 45 to 60%). Islands of bog woodland often form a mosaic with the open bogs and the two community types intergrade as i l l u s t r a t e d in Fig. 9. The releve's of this community type form a d i s t i n c t group on the PCA and RA ordinations (Figs. 6 and 7). This is a r e f l e c t i o n of the d i s t i n c t i v e f l o r a of these habitats. Of the si x community types described, the open bogs have the largest number of exclusive and selective taxa. Although considerable v a r i a b i l i t y in f l o r i s t i c s exists among the open bog communities occurring along the transects, as a group they share the c h a r a c t e r i s t i c combination of species l i s t e d in Table 1. Most of the constant or dominant species are included in group A and B on the PCA species ordination (Fig.8). The species of group A are exclusive or selective for the open bogs while group B includes species occurring in both the open bogs and bog woodlands. Many of those species in group B (those closest to group A) are p r e f e r e n t i a l for the open bogs. Major d i s t i n c t i o n s in bog vegetation and surface features The most readily observable trend within the open bog community type is that from the r e l a t i v e l y flat-surfaced, homogeneous and f l o r i s t i c a l l y 'simple' herb-dominated bogs to the more heterogeneous and f l o r i s t i c a l l y complex, shrub-dominated bogs with mounded to hummocky surface topography. The 'simple' f l a t bogs (Figs. 11., 12) are most often r e s t r i c t e d to small confined areas surrounded by upland Table 1. Vegetative features and character species for Ericaceous shrub -Trichophorum cespitosum - Sphagnum spp. - Cladonia spp. - Siphula  ceratites open bog community type. General vegetative features - tree layers lacking - moderately wel developed shrub layer ( 7 - 81% cover; 5c = 42%) - wel developed herb layer (40 - 95% cover; • x = 80%) - wel  developed bryophyte and lichen layer (65 - 95% cover; x = 86%) Shrub layers: character species* Kalmia microphyla  Vaccinium uliginosum  Juniperus communis  Ledum groenlandicum  Pinus contorta Herb layer: character species CD2; preferential3 CD; + preferential CD; + selective CD (75% constancy) Trichophorum cespitosum CD; preferential Trientalis europaea CD; preferential Andromeda polifolia CD; preferential Coptis trifolia CD; preferential Vaccinium oxycoccos CD; selective Drosera rotundifolia CD; selective Gentiana douglasiana CD; exclusive Sanguisorba officinalis CD; preferential Eriophorum angustifolium CD; selective Empetrum nigrum CD; preferential Carex livida CD; exclusive Comus unalaschkensis CD Drosera anqlica exclusive Agrostis aequivalvis exclusive hytes and lichen layer: character species Sphagnum papilosum CD; preferential Sphagnum tenelum CD; exclusive Sphagnum rubelum CD; preferential Cladonia impexa CD; selective Cladonia gracilis CD; + preferential Rhacomitrium lanuginosum CD; + selective Siphula ceratites CD; exclusive Cladonia uncialis CD; exclusive Sphagnum fuscum D Cladonia ranqiferina + preferential •^Species are listed in order of decreasing constancy. The lists include only species of highest constancy and dominance as wel as exclusive, selective or preferential taxa of lower constancy and/or dominance. Refer to Appendix VI for complete vegetation tables. 2c = constant species (constancy of >^  80%) D = dominant species (average cover abundance value of >3) 3Refer to definitions of character taxa in text (page 29). 47 f o r e s t or bog woodland and are probably topogenous i n o r i g i n ( s m a l l b a s i n or d e p r e s s i o n bogs). The more complex bogs have an i r r e g u l a r s u r f a c e topography, occur on slopes as w e l l as f l a t s , and thus r e p r e s e n t the true blanket bog ( F i g s . 9, 10, 13, 14). Intermediates between these two extremes are common, however, and i t appears that true blanket bog and topogenous bog o f t e n c o a l e s c e so that the d i s t i n c t i o n between the two types of genesis i s masked. T e r t i a r y peat may blanket an area which i n some cases i s u n d e r l a i n by primary and secondary peat and i n others by min e r a l t e r r a i n (bedrock, g l a c i a l t i l l , e t c . ) . The 'blanket bog complex' concept o u t l i n e d i n Chapter 2 i s u s e f u l i n d e s c r i b i n g t h i s phenomenon. Ve g e t a t i o n and s u r f a c e f e a t u r e s of the f l a t bogs Releves 6, 14 and 24 r e p r e s e n t examples of f l a t or simple bogs ( F i g s . 11, 12). A l l have a p o o r l y developed shrub l a y e r . Juniperus communis, Ledum groenlandicum and Pinus c o n t o r t a , which are present i n most b l a n k e t bogs, are l a c k i n g or un-common i n the f l a t bogs. Two constant shrub s p e c i e s are Kalmia m i c r o p h y l l a and Vaccinium uliginosum. The herbaceous l a y e r i s u s u a l l y best developed i n these bogs; most of the c h a r a c t e r s p e c i e s f o r the open bog community type as a whole are abundant (Table 1). Constant species are Trichophorum cespitosum, T r i e n t a l i s europaea, Andromeda p o l i - f o l i a , C o p t i s t r i f o l i a , Vaccinium oxycoccos, Drosera r o t u n d i - f o l i a , Gentiana d o u g l a s i a n a , Empetrum nigrum, Cornus unalasch- k e n s i s , and Rubus chamaemorus. The bryophyte and l i c h e n l a y e r i s a l s o w e l l developed with 48 F i g . 1 1 . E r i c a c e o u s s h r u b - T r i c h o p h o r u m c e s p i t o s u m - Sphagum s p p . open f l a t bog commun i t y ; r e l e v e 6 . A b r u p t t r a n s i t i o n t o c e d a r - c y p r e s s - hemlock u p l a n d f o r e s t community i n b a c k g r o u n d . F i g . 1 2 . E r i c a c e o u s s h r u b - T r i c h o p h o r u m c e s p i t o s u m - Sphagnum s p p . open f l a t bog commun i t y ; r e l e v e * 14 . 42A 49 an o b v i o u s d o m i n a n c e o f Sphagnum s p e c i e s , i n c o n t r a s t t o t h e b l a n k e t bogs where Sphagnum i s a b u n d a n t b u t o f t e n n o t as c o n s i s t e n t l y d o m i n a n t as i n t h e f l a t b o g s . Sphagnum  p a p i l l o s u m i s t h e d o m i n a n t s p e c i e s b u t S_. t e n e l l u m , S_. r u b e l l u m , S_. l i n d b e r g i i and S_. f a l l a x a r e o f t e n a b u n d a n t as w e l l . R h a c o m i t r i u m l a n u g i n o s u m , a c o n s t a n t l y a b u n d a n t moss i n b l a n k e t b o g s , i s a b s e n t o r v e r y s p o r a d i c i n t h e f l a t b o g s . C l a d o n i a i m p e x a i s - t h e o n l y a b u n d a n t l i c h e n i n t h e f l a t b o g s . L i c h e n s s u c h as S i p h u l a c e r a t i t e s , C l a d o n i a g r a c i l i s v a r . e l o n g a t a , C. r a n g i f e r i n a and C. u n c i a l i s , c h a r a c t e r i s t i c a l l y a b u n d a n t i n b l a n k e t b o g s , a r e r e l a t i v e l y s c a r c e i n t h e f l a t b o g s . The f l a t bogs show c o n s i d e r a b l e v a r i a t i o n i n t h e r e l a t i v e w e t n e s s o f t h e i r s u r f a c e s . Some a r e s a t u r a t e d t o t h e s u r f a c e ( r e l e v e s 14 and 24) and show a r e l a t i v e a b u n d a n c e o f s p e c i e s s u c h as T r i c h o p h o r u m c e s p i t o s u m , S a n g u i s o r b a o f f i c i n a l i s , E r i o p h o r u m a n g u s t i f o l i u m , C a r e x 1 i v i d a , a n d Sphagnum t e n e l l u m w h i c h t e n d t o t o l e r a t e o r f a v o r t h i s w e t n e s s . O t h e r s ( r e l e v e 6) a p p e a r somewhat d r i e r a t t h e s u r f a c e and s p e c i e s s u c h as K a l m i a m i c r o p h y l l a , V a c c i n i u m u l i g i n o s u m , T r i e n t a l i s e u r o p a e a , V a c c i n i u m o x y c o c c o s , F a u r i a c r i s t a - g a l l i , C a r e x p l u r i f l o r a , R ubus chamaemorus, Sphagnum r u b e l l u m and Sphagnum f a l l a x become more d o m i n a n t . The g r o u p i n g o f c e r t a i n bog c o m m u n i t i e s i n t o ' s i m p l e ' f l a t bogs i s t h u s b a s e d more on s u r f a c e c o n f i g -u r a t i o n and o v e r a l l s t r u c t u r e o f t h e v e g e t a t i o n t h a n on s t r i c -t l y f l o r i s t i c s i m i l a r i t y , b e c a u s e c o n s i d e r a b l e v a r i a b i l i t y i n s p e c i e s c o m p o s i t i o n o c c u r s among t h e s a m p l e s o f t h i s bog t y p e . 50 The s o i l s of the f l a t bogs are d e s c r i b e d and co n t r a s t e d with those of the blanke t bogs on page 59. Ve g e t a t i o n and s u r f a c e f e a t u r e s of the blanket bogs The remainder of the open bog communities sampled are cons i d e r e d p o r t i o n s o f blanket bogs ( F i g s . 9, 10, 13, 14), although some may be r a i s e d bogs which have developed through the accumulation of t e r t i a r y peat over topogenous bogs. U n l i k e the f l a t bogs, blanket bogs are not confined to s m a l l , w e l l d e f i n e d areas but cover e x t e n s i v e areas of low r e l i e f and g e n t l e to moderate slo p e s (0 - 36% i n the bogs sampled). In comparison to the f l a t bogs, the blanket bogs have an i r r e g u l a r s u r f a c e due i n most cases to und u l a t i n g mounds and depressions d i f f e r i n g i n height by 10 - 30 cm and d i s t r i b u t e d i r r e g u l a r l y over the bog s u r f a c e . These su r f a c e i r r e g u -l a r i t i e s commonly r e s u l t from the hummock - forming growth c h a r a c t e r i s t i c s of a few Sphagnum s p e c i e s such as S. rubellum, S. fuscum and IS. imbricatum, as w e l l as s p e c i e s of C l a d o n i a l i c h e n s and the moss Rhacomitrium lanuginosum. In the case of the shallow bogs (<50 cm deep), the u n d e r l y i n g mineral sub-s t r a t e , which i s u s u a l l y bedrock, may s t i l l be i n f l u e n c i n g the s u r f a c e topography of the bog. Large Sphagnum mounds a l s o occur around the bases of t r e e s ( u s u a l l y Pinus c o n t o r t a ) s c a t t e r e d throughout the bogs ( F i g . 15). Bogs o c c u r r i n g on slo p e s o f t e n have elongated mounds o r i e n t e d at r i g h t angles to the s l o p e . Small, shallow pools 1 - 4 m2 i n s i z e and 10 - 50 cm deep are common f e a t u r e s of many of the bogs. If the pools 51 F i g . 1 3 . P i n u s c o n t o r t a - C h a m a e c y p a r i s n o o k a t e n s i s - e r i c a c e o u s s h r u b -T r i c h o p h o r u m c e s p i t o s u m - Sphagnum s p p . - C l a d o n i a s p p . -S i p h u l a c e r a t i t e s open b l a n k e t bog community nea r Ra inbow L a k e . F i g . 14 . P i n u s c o n t o r t a - e r i c a c e o u s s h r u b - T r i c h o p h o r u m c e s p i t o s u m -Sphagnum s p p . - C l a d o n i a s p p . - S i p h u l a c e r a t i t e s open b l a n k e t bog commun i t y ; r e l e v e 1 1 . 52 d e v e l o p i n s l o p i n g p o r t i o n s o f t h e b o g s , as t h e y o f t e n do, t h e y t a k e on t h e a p p e a r a n c e o f a s e r i e s o f t e r r a c e s , c i r c u l a r t o t e a r - d r o p - s h a p e d , w i t h 'dams' o f Sphagnum ( o f t e n c o v e r e d w i t h t u s s o c k s o f T r i c h o p h o r u m c e s p i t o s u m ) b u i l t up a c r o s s t h e d o w n s l o p e s i d e o f t h e p o o l s ( F i g . 1 6 ) . The ponds a p p e a r t o be l i n k e d i n s e r i e s t h r o u g h b o t h s u r f a c e and s u b - s u r f a c e s e e p a g e p a t h w a y s ; and t h e i r s u b s t r a t e , w h i c h c o n s i s t s o f an o o z y m i x t u r e o f o r g a n i c s e d i m e n t s and a l g a e , i s o f t e n more s u s p e n d e d and l o o s e on t h e u p s l o p e s i d e o f t h e pond where w a t e r and s e d i m e n t s e n t e r . R o o t e d a q u a t i c o r s e m i - a q u a t i c v e g e t a t i o n d o m i n a t e d by E r i o p h o r u m a n g u s t i f o l i u m i s u s u a l l y b e s t d e v e l o p e d a t t h e d o w n s l o p e end o f t h e pond a d j a c e n t t o t h e p e a t dam, where t h e s e d i m e n t a p p e a r s t o be more s t a b i l i z e d . The t e r r a c e d a s p e c t o f t h e s e s l o p i n g bog - p o o l c o m p l e x e s i s b e s t s e e n f r o m t h e a i r ( F i g . 1 7 ) . The bog a p p e a r s t o be s l o w l y o o z i n g d o w n s l o p e . I n e x t r e m e c a s e s w i t h i n t h e s t u d y a r e a , t h e bog s u r f a c e may c o n s i s t p r e d o m i n a n t l y o f w a t e r w i t h i s l a n d s o f ' d r y ' Sphagnum hummocks c o l o n i z e d by P i n u s c o n t o r t a , as i n t h e bog i l l u s t r a t e d i n F i g . 18 n e a r O l i v e r L a k e a l o n g H i g h w a y 16. I n t h i s c a s e , t h e r o a d b e d may h a v e i n t e r r u p t e d d r a i n a g e and r a i s e d t h e w a t e r t a b l e o f t h e bog. The v e g e t a t i o n o f t h e b l a n k e t bogs i s r e a l l y a m o s a i c o f m i c r o c o m m u n i t i e s w h i c h d e v e l o p i n r e s p o n s e t o , o r i n f a c t b r i n g a b o u t , t h e h e t e r o g e n e o u s s u r f a c e t o p o g r a p h y and m o i s t u r e c o n d i t i o n s t h r o u g h o u t t h e bog s u r f a c e . A l t h o u g h a d e t a i l e d c l a s s i f i c a t i o n o f t h e s e m i c r o c o m m u n i t i e s i s b e y o n d t h e s c o p e 53 F i g . 15 . L a r g e , r e d d i s h Sphagnum r u b e l l u m mound (no t snow-cove red ) a round t h e base o f P i n u s c o n t o r t a i n a b l a n k e t bog n e a r Rainbow L a k e . Pho to shows snow c o n d i t i o n s i n l a t e M a r c h , 1979 . F i g . 16 . S e r i e s o f t e r r a c e d ponds w i t h T r i c h o p h o r u m c e s p i t o s u m "dams" i n a s l o p i n g b l a n k e t bog wes t o f D i a n a C r e e k , t r a n s e c t 3 . Note l a r g e , w h i t e c o l o n i e s o f t h e l i c h e n S i p h u l a c e r a t i t e s . a A 54 F i g . 17. Sloping bog with ser ies of terraced ponds\ near Work Channel. Photo shows snow condi t ions in la te March, 1979. F i g . 18. Bog - pool complex near O l i ve r Lake. The const ruct ion of Highway 16 probably increased the water table in th i s bog. 55 o f t h i s s t u d y , a d e s c r i p t i o n o f t h e c h a r a c t e r i s t i c s o f bog v e g e t a t i o n w o u l d be i n c o m p l e t e w i t h o u t d i s c u s s i o n o f how s p e c i e s d i s t r i b u t i o n r e f l e c t s s u r f a c e t o p o g r a p h y and m o i s t u r e r e g i m e t h r o u g h o u t t h e bog c o m m u n i t i e s . The f o l l o w i n g d i s c u s s i o n i s b a s e d on f i e l d n o t e s made d u r i n g t h e r e l e v e ' s a m p l i n g and r e c o n n a i s s a n c e t h r o u g h o u t t h e s t u d y a r e a . A s i d e f r o m t h e r o c k o u t c r o p s , w h i c h a r e c o n s i d e r e d a d i s t i n c t c ommunity t y p e f r o m t h e b o g s , t h e mounds o r hummocks r e p r e s e n t t h e d r i e s t h a b i t a t s w i t h i n t h e b o g s . F o r t h e most p a r t t h e y a r e d o m i n a t e d by s p e c i e s o f Sphagnum. The d r i e s t hummocks u s u a l l y c o n s i s t o f S_. r u b e l l u m ( F i g . 15) and S_. f u s c u m . T h e s e two s p e c i e s c a n be d i s t i n g u i s h e d a t a d i s t a n c e by t h e i r c o l o r , t h e f o r m e r b e i n g q u i t e r e d d i s h o r r e d d i s h - g r e e n and t h e l a t t e r b r o w n i s h . S_. i m b r i c a t u m , an o r a n g e c o l o r e d s p e c i e s w i t h c o a r s e , r o b u s t l e a v e s , f o r m s somewhat w e t t e r hummocks as does S_. p a p i l l o s u m , a n o t h e r r o b u s t s p e c i e s o f f a i r l y w i d e d i s t r i b u t i o n w i t h i n t h e bog c o m m u n i t i e s . The o v e r a l l d o m i n a n c e o f Sphagnum i n t h e b l a n k e t bogs i s c o n s i d e r a b l y l e s s t h a n i n t h e f l a t b o g s . T h i s i s l a r g e l y due t o t h e i n c r e a s e d i m p o r t a n c e o f many o t h e r m o s s e s , l i v e r w o r t s a nd l i c h e n s t h a t a r e l a c k i n g i n t h e f l a t b o g s . R h a c o m i t r i u m  l a n u g i n o s u m o f t e n f o r m s e x t e n s i v e c o l o n i e s i n t h e d r y h a b i t a t s a s do many l i c h e n s ( F i g . 10, 13, 1 4 ) , e s p e c i a l l y C l a d o n i a  i m p e x a and C. r a n g i f e r i n a . C l a d o n i a u n c i a l i s , C. f u r c a t a , C. s y l v a t i c a , C. p s e u d o e v a n s i i and C. l e u c o p h a e a a r e o f t e n 56 p r e s e n t as w e l l . S h r u b d e v e l o p m e n t i s b e s t i n t h e d r i e s t p o r t i o n s o f b l a n k e t b o g . B o n s a i 1 P i n u s c o n t o r t a 0.5 t o 2 m i n h e i g h t , i s t h e m a j o r t r e e s p e c i e s . C h a m a e c y p a r i s n o o t k a t e n s i s a n d T h u j a p l i c a t a a r e l e s s f r e q u e n t . T s u g a h e t e r o p h y l l a a n d T. m e r t e n s i a n a o c c u r v e r y s p o r a d i c a l l y a n d show v e r y p o o r v i g o u r . O t h e r d o m i n a n t s h r u b s i n d r y h a b i t a t s i n c l u d e J u n i p e r u s communis, Ledum g r o e n l a n d i c u m and V a c c i n i u m  u l i g i n o s u m . G a u l t h e r i a s h a l l o n o c c u r s s p o r a d i c a l l y i n t h e s e h a b i t a t s . K a l m i a m i c r o p h y l l a , a l t h o u g h n o t r e s t r i c t e d t o t h e d r i e s t h a b i t a t s , o f t e n shows b e s t v i g o u r h e r e . H e r b l a y e r s i n t h e d r y m i c r o h a b i t a t s a r e o f t e n p o o r l y d e v e l o p e d a n d d o m i n a t e d b y l e a t h e r y l e a v e d and woody s p e c i e s ( a c t u a l l y d w a r f s h r u b s ) . Empetrum n i g r u m i s t h e most common s p e c i e s . L o i s e l e u r i a p r o c u m b e n s a n d V a c c i n i u m v i t i s - i d a e a a r e a p p a r e n t l y r e s t r i c t e d t o t h e s e h a b i t a t s a s w e l l ; b o t h s p e c i e s a r e o f t e n o f l o w a b u n d a n c e , t h e l a t t e r b e i n g more common i n t h e s h a d e d b o g w o o d l a n d s . L y c o p o d i u m a n n o t i n u m and L. c l a v a t u m o c c u r s p o r a d i c a l l y . S e v e r a l b r y o p h y t e s a n d l i c h e n s i n c l u d i n g P l e u r o z i u m  s c h r e b e r i , P o l y t r i c h u m j u n i p e r i n u m , B a z z a n i a s p p . , L o p h o z i a s p p . , D i c r a n u m h o w e l l i , D. m a j u s a nd C o r n i c u l a r i a a c u l e a t a , a l t h o u g h n e v e r a b u n d a n t i n t h e b o g s , o c c u r p r e d o m i n a n t l y o n t h e hummocks, o f t e n u n d e r t h e shade o f s h r u b s . 1 , 1 B o n s a i " d e s c r i b e s a d w a r f e d , o f t e n g n a r l e d and t w i s t e d g r o w t h f o r m . 57 S e v e r a l s p e c i e s show l i t t l e a p p a r e n t h a b i t a t p r e f e r e n c e t h r o u g h o u t t h e bogs and f o r m a component o f a l l e x c e p t t h e w e t t e s t m i c r o c o m m u n i t i e s . I m p o r t a n t among t h e s e a r e T r i c h o - phorum c e s p i t o s u m , Andromeda p o l i f o l i a , S a n g u i s o r b a o f f i c i - n a l i s , T r i e n t a l i s e u r o p e a , G e n t i a n a d o u g l a s i a n a , C o r n u s  u n a l a s c h k e n s i s , A g r o s t i s a e q u i v a l v i s , Rubus chamaemorus, T o f i e l d i a g l u t i n o s a , F a u r i a c r i s t a - g a l l i , V a c c i n i u m o x y c o c c o s and C a r e x p l u r i f l o r a . Many o f t h e s e s p e c i e s a r e more v i g o r o u s when g r o w i n g u n d e r t h e s h a d e o f s h r u b s i n t h e somewhat d r i e r h a b i t a t s ( i n t e r m e d i a t e b e t w e e n t h e wet d e p r e s s i o n s and d r i e r m o u n d s ) . M y r i c a g a l e becomes a d o m i n a n t s h r u b s p e c i e s i n some b l a n k e t bog c o m m u n i t i e s ( F i g . 1 9 ) . T h i s s p e c i e s i s u s u a l l y a s s o c i a t e d w i t h t r a n s i t i o n a l b o g - f e n c o m m u n i t i e s w i t h a m i n e r o -t r o p h i c r a t h e r t h a n o m b r o t r o p h i c w a t e r r e g i m e (Gorham 1957; R a t c l i f f e 1964; Wade 1965; W e l l s 1 9 8 1 ) . I n t h e s t u d y a r e a i t m o s t o f t e n c o l o n i z e s t h e e d ges o f s t r e a m l e t s m e a n d e r i n g t h r o u g h t h e b o g s . R e l e v e s 28, 38, 39 and 40 r e p r e s e n t bog c o m m u n i t i e s d o m i n a t e d by M y r i c a g a l e . A t t h e wet end o f t h e s c a l e t h e r e i s a s e r i e s o f h a b i t a t s a n d m i c r o c o m m u n i t i e s r a n g i n g f r o m open p o n d s t o wet d e p r e s s i o n s , and i n many c a s e s t h e s e a p p e a r t o r e p r e s e n t v a r i o u s s t a g e s i n pond s u c c e s s i o n ( F i g s . 20 - 2 3 ) . M e n y a n t h e s  t r i f o l i a t a i s common i n t h e d e e p e s t p o r t i o n s o f t h e p o n d s , s o m e t i m e s i n a s s o c i a t i o n w i t h N u p h a r l u t e a s u b s p . p o l y s e p a l u m , a l t h o u g h t h e l a t t e r i s more c h a r a c t e r i s t i c o f t h e l a r g e r p o n d s (>4 m 2) and s h a l l o w l a k e s . S c h e u c h z e r i a p a l u s t r i s i s o f 58 F ig . 19. A Myrica gale - dominated blanket bog community east of Rainbow Lake; releve' 28. F i g . 20. Sphagnum l i 'ndbergi i encroaching on a pond in a s lop ing blanket bog; releve' 30. The orange - colored Spagnum away from the water 's edge i s probably S_. imbricatum. Eriophorum angust -i f o l i um is growing in the pond. £74 59 F ig . 2 1 . A Sphagnum papi l losum hummock developing in a pond. S_. l i n d b e r g i i occurs at the water 's edge and Eriophorum  angust i fo l ium is growing p a r t i a l l y submerged in the pond. F i g . 2 2 . A l a t e r stage in pond succession in a 'b lanket bog." Several species inc lud ing Eriophorum augus t i fo l ium. Trichophorum  cespitosum, Drosera a n g l i c a , Sphagnum papi l losum and S iphula ce ra t i t e s have co lonized the o r i g i na l pond. 60 F i g . 23. Eriophorum angust i fo l ium (mainly) in a l a t e r stage of pond succession in blanket bog. F i g . 24. The l i chen S iphula ce ra t i t es (white) and the l i verwort Gymnocolea i n f l a t a (b lack ) ; common assoc iates in depressions subject to in te rmi t ten t submergence in open blanket bog/ The tussocks are Trichophorum cespitosum. 61 s p o r a d i c o c c u r r e n c e i n t h e p o n d s . E r i o p h o r u m a n g u s t i f o l i u m i s o f t e n t h e f i r s t ' s e m i - a q u a t i c ' p l a n t t o c o l o n i z e t h e s h a l l o w bog p o o l s and f o r m s e x t e n s i v e c o l o n i e s w i t h i n them ( F i g s . 20 - 2 3 ) . Dark b r o n z e p a t c h e s o f Sphagnum l i n b e r g i i a r e commonly s e e n e n c r o a c h i n g on po n d s ( F i g . 2 0 ) . T h i s s p e c i e s i s a b l e t o grow i n s t a n d i n g w a t e r , w h i l e Sphagnum t e n e l l u m and S_. compactum a r e s l i g h t l y l e s s t o l e r a n t o f s a t u r a t i o n and r e p l a c e S3, l i n b e r g i i i n l a t e r s t a g e s o f pond s u c c e s s i o n . S_. p a p i l l o s u m , a s p e c i e s o f f a i r l y w i d e d i s t r i b u t i o n i n t h e b o g s , may o c c u r as w e l l . I n h a b i t a t s s u b j e c t t o o n l y i n t e r m i t t e n t i n u n d a t i o n , t h e l i c h e n S i p h u l a c e r a t i t e s , a s p e c i e s v e r y c h a r a c t e r i s t i c o f t h e s e o c e a n i c b o g s , o f t e n f o r m s e x t e n s i v e mats ( F i g . 24) commonly i n a s s o c i a t i o n w i t h t h e l i v e r w o r t G y m n o c o l e a i n f l a t a a n d t h e sundews D r o s e r a a n g l i c a and D. r o t u n d i f o l i a ( F i g . 2 2 ) . D. r o t u n d i f o l i a a p p e a r s a b l e t o t o l e r a t e d r i e r c o n d i t i o n s t h a n D. a n g l i c a and t h u s h a s w i d e r d i s t r i b u t i o n w i t h i n t h e b o g s . D. r o t u n d i f o l i a a l s o o c c u r s i n bog w o o d l a n d s . The c l u b m o s s L y c o p o d i e l l a i n u n d a t a may a l s o o c c u r i n wet d e p r e s s i o n s and a l o n g t h e edge o f pon d s w i t h t h e ab o v e s p e c i e s . The s e d g e s R h y n c h o s p o r a a l b a and C a r e x 1 i v i d a c o l o n i z e wet d e p r e s s i o n a l a r e a s , t h a t i n many c a s e s a p p e a r t o h a v e d e v e l o p e d t h r o u g h s u c c e s s i o n f r o m open p o n d s . T r i c h o p h o r u m - c e s p i t o s u m b e g i n s t o o c c u r i n t h e s e h a b i t a t s as w e l l and d o m i n a t e s i n many o f t h e d r i e r m i c r o c o m m u n i t i e s w i t h i n t h e b l a n k e t b o g s . A few s p e c i e s s h o u l d be m e n t i o n e d t h a t a r e o f 62 l i m i t e d o c c u r r e n c e w i t h i n t h e s t u d y a r e a and a r e more t y p i c a l o f t r a n s i t i o n a l t o r h e o p h i l o u s p o r t i o n s o f b o g s . They a r e u s u a l l y f o u n d i n s h a l l o w (< 0.5 m deep) s l o p i n g b o g s a n d f e n s o r l o c a l i z e d i n m i n e r a l f l u s h e s ( F i g . 25) o r a l o n g c r e e k e d g e s i n t h e d e e p e r b l a n k e t b o g s . They i n c l u d e D o d e c a t h e o n  j e f f r e y i , Geum c a l t h i f o l i u m , E r i g e r o n p e r e g r i n u s and P e d i -c u l a r i s s p . The t a l l s e d g e s C a r e x s i t c h e n s i s and C. r o s t r a t a o c c u r v e r y s p o r a d i c a l l y i n t h e o m b r o p h i l o u s b o g s . They a r e much more t y p i c a l o f t h e m i n e r o t r o p h i c w e t l a n d s ( f e n s and m a r s h e s ) b o r d e r i n g l a k e s a n d m o v i n g w a t e r c o u r s e s . S o i l s a n d l a n d f o r m s o f t h e o p e n b o g c o m m u n i t i e s Open b o g c o m m u n i t i e s o c c u r most commonly on o r g a n i c b l a n k e t s (> 1 m i n d e p t h ) and ( l e s s commonly) on v e n e e r s (< 1 m i n d e p t h ) . S o i l s o f t h e f l a t b o g s s a m p l e d a r e F i b r i s o l s ( T y p i c and M e s i c ) w h i l e M e s i s o l s ( T y p i c , Humic and T e r r i c ) o c c u r i n a l l e x c e p t one o f t h e b l a n k e t b o g s . The e x c e p t i o n i s a n O r t h i c Humic R e g o s o l i n t h e s h a l l o w s l o p i n g b o g r e p r e s e n t e d b y r e l e v e 30. O r t h i h i s t o m o d e r s c o n s i s t i n g o f Om h o r i z o n s a t t h e s u r f a c e ( F i g . 26) a n d O r t h i h i s t o m o r s w i t h s p h a g n i c Of h o r i z o n s d o m i n a t i n g ( F i g . 2 7 ) , a r e t h e c h a r a c t e r i s t i c humus f o r m s i n t h e o p e n b o g s . Among t h e 12 o p e n b o g s s a m p l e d , p e a t d e p t h v a r i e d f r o m 40 t o 380 cm b u t was u s u a l l y g r e a t e r t h a n 100 cm. The s h a l l o w e s t b o g s o c c u r on s l o p e s ( r e l e v e 30) a n d o n l e v e l t o p o g r a p h y o v e r m e t a m o r p h i c b e d r o c k ( r e l e v e 2 2 ) . T e r r i c M e s i s o l s ( l i t h i c p h a s e ) o c c u r i n t h e l a t t e r . I n most c a s e s t h e o r g a n i c s o i l s o f t h e o p e n b o g c o m m u n i t i e s a r e u n d e r l a i n b y r o c k , o f t e n w i t h 63 F i g . 25. A mineral f l ush in a shal low, s lop ing blanket bog; r e l eve '30 . Dodecatheon j e f f r e y i (wide elongate leaves) character izes these minerotrophic hab i ta t s . F i g . 26. 0m horizon of an Orthihistomoder from an open blanket bog community: re leve 20. The sedges are Carex 1 i vi da and Rhynchospora a lba . 634 64 F i g . 27. Sphagnic Of horizon of an Orthihistomor from am open blanket bog; releve 14. The sedge i s Trichophorum cespitosum. F i g . 28. Shr inking sedimentary peat in a dr ied - up bog pond. 65 a 1 0 - 2 0 cm t h i c k sandy organo-mineral h o r i z o n immediately above the l i t h i c c o n t a c t . The v a r i a t i o n i n depth to bedrock i s s i g n i f i c a n t over s h o r t d i s t a n c e s . Probes taken throughout the 180 m2 p l o t s r e v e a l e d ranges i n depth of as much as 280 cm. Most p l o t s have a range of a t l e a s t 100 cm. T h i s v a r i a b i l i t y i n depth suggests t h a t wet depressions i n the u n d e r l y i n g hummocky bedrock landform could have acted as templates or n u c l e i f o r peat accumulation and bog formation. The hummocky topography would subsequently have been masked as these bogs expanded and co a l e s c e d over the high e r and b e t t e r d r a i n e d areas through the accumulation of secondary and t e r t i a r y peat. One of the bog communities sampled ( r e l e v e 12) i s under-l a i n by a s i l t y f l u v i a l d e p o s i t probably o r i g i n a t i n g from a nearby meandering stream that i s bordered by a sedge-dominated fen. The s o i l of t h i s bog i s a Humic M e s i s o l c o n s i s t i n g of Of and Om s u r f a c e h o r i z o n s with a dark humified sedge peat (Oh horizon) below 125 cm. This now ombrophilous bog i s probably topogenous i n o r i g i n , d e v eloping through s u c c e s s i o n from an o r i g i n a l l y r h e o p h i l o u s fen community. T h i s could c o n c e i v a b l y have r e s u l t e d from a r e d i r e c t i o n of the nearby stream and a subsequent removal of the n u t r i e n t - e n r i c h i n g moving water source. The t r u l y r h e o p h i l o u s fens and marshes were not sampled or d e s c r i b e d i n t h i s study. They are l o c a l i z e d w i t h i n the study area, r e s t r i c t e d to stream edges and lake margins. S o i l s of the open bog communities are f r e q u e n t l y s a t u r a t e d to w i t h i n 0 - 1 0 cm of the s u r f a c e although f r e e w a t e r i s o f t e n n o t e v i d e n t u n t i l some p r e s s u r e i s e x e r t e d . F r e e w a t e r may n o t a p p e a r f o r 5 - 1 0 m i n u t e s i n s h a l l o w e x c a v a t i o n s , i l l u s t r a t i n g t h e h i g h w a t e r r e t e n t i o n i n t h e s e o r g a n i c s o i l s . D u r i n g t h e month o f A u g u s t , 1979, t h e w e a t h e r was u n u s u a l l y d r y f o r t h e r e g i o n and many o f t h e bog s u r f a c e s t o o k on an u n n a t u r a l l y d u l l a p p e a r a n c e . Sphagnum hummocks w h i c h l o o k e d v e r y d r y r e m a i n e d m o i s t t o t h e t o u c h , h o w e v e r , and w a t e r c o u l d s t i l l be r e a d i l y s q u e e z e d f r o m them. The b o t t o m s o f p o n d s , c o n s i s t i n g o f a f i n e , o o z y , o r g a n i c s e d i m e n t and an a l g a l scum, s h r a n k and c r a c k e d i n a p o l y g o n a l p a t t e r n ( F i g . 28) d u r i n g t h i s d r y p e r i o d . N e i l a n d (1971) r e p o r t s s i m i l a r o b s e r v a t i o n s i n t h e bogs o f S o u t h e a s t A l a s k a . S h r i n k a g e v a l u e s o f 64 and 8 3 % have b e e n measured' on s a m p l e s o f s e d i m e n t a r y p e a t c o l l e c t e d f r o m t h e P a c i f i c c o a s t ( T a r n o c a i u n p u b l i s h e d d a t a ) and t h i s m a t e r i a l i s v e r y d i f f i c u l t t o r e w e t a f t e r i t has been d r i e d . A v e r a g e pH w i t h i n t h e r o o t i n g z o n e o f t h e open bog c o m m u n i t i e s i s v e r y l o w , r a n g i n g f r o m 2.5 t o 3.3 ( x = 2.8) among t h e 8 p r o f i l e s a n a l y z e d . The s h a l l o w s l o p i n g bog r e p r e s e n t e d by r e l e v e 30 had t h e h i g h e s t pH v a l u e . R e l a t i o n s h i p s among t h e r e l e v e s The r e l a t i o n s h i p s among t h e o p e n bog r e l e v e s i s b e s t i l l u s t r a t e d by t h e RA o r d i n a t i o n o f t h e bog and bog w o o d l a n d r e l e v e s i n F i g . 29. The f l a t bog r e l e v d s ( 2 4 , 14, 6) a r e s e p a r a t e d somewhat f r o m t h e b l a n k e t bog r e l e v e s , r e f l e c t i n g t h e f l o r i s t i c d i s t i n c t i o n s , o u t l i n e d a b o v e , b e t w e e n t h e s e two bog t y p e s . R e l e v e 6 s t a n d s o u t b e c a u s e i t i s s p e c i e s p o o r b u t 67 F ig . 29. RA ord inat ion of Er icaceous shrub - Trichophorum cespitosum -Sphagnum open bog communities ( Q ) a n c ' P l n e " cypress -cedar - Gaul ther ia - Fauria bog woodland communities ([ [ ) , axes 1 (x) and 2 (y). 100.0-128 9 0 . 0 8 0 . 0 - . - . (22) (u) 70 .0-60 .0 50.0-Y axis 40 .0 30.0 20 .0 10.0 o.ol.. 0.0 .1, © © © © .1 .1. 10.0 2 3 . j 30 .0 4 50.0 X AXIS 37 29 .1 . 35 3||rrj' 15 • I .1 60 .0 70.0 80.0 90 .0 1C0.C 68 a l s o p o s s e s s e s some h e r b s p e c i e s , s u c h as L y s i c h i t o n  a m e r i c a n u m , Maianthemum d i l a t a t u m and C o p t i s a s p l e n i i f o l i a t h a t a r e more c h a r a c t e r i s t i c o f t h e f o r e s t s . R e l e v e s 2 8 , 3 8 , 39 and 4 0 , c l u s t e r e d t o g e t h e r on t h e o r d i n a t i o n , r e p r e s e n t b l a n k e t bog c o m m u n i t i e s d o m i n a t e d by M y r i c a g a l e . The p o s i t i o n o f r e l e v e - 12 s u g g e s t s t e n d e n c i e s t o w a r d bog w o o d l a n d . P i n u s c o n t o r t a and P l e u r o z ium s c h r e b e r i , b o t h c o n s t a n t d o m i n a n t s i n t h e bog w o o d l a n d s , a r e e x c e p t i o n a l l y a b u n d a n t i n t h e o p e n bog r e p r e s e n t e d by r e l e v e 1 2 . R e l e v e 3 0 , f r o m t h e s h a l l o w s l o p i n g bog, had two u n i q u e s p e c i e s , D o d e c a t h e o n  j e f f r e y i and C a l t h a l e p t o s e p a l a . I t d o e s n ' t s t a n d o u t on t h e o r d i n a t i o n , h o w e v e r , s i n c e t h e s e two s p e c i e s o c c u r r e d i n o n l y one r e l e v e * and were t h u s e x c l u d e d f r o m t h e a n a l y s i s . 4.4.2.2 Empetrum nigrum - V a c c i n i u m u l i g i n o s u m - R h a c o m i t r i u m  lanuginosum - C l a d o n i a spp. o u t c r o p communities T h i s c o m m u n i t y t y p e ( F i g . 30) r e p r e s e n t s t h e d r i e s t e x t r e m e w i t h i n t h e s t u d y a r e a . I t i s o f l i m i t e d o c c u r r e n c e a l o n g t h e t r a n s e c t s and i s r e p r e s e n t e d by o n l y one r e l e v e . B e c a u s e o f l i m i t e d d a t a o n l y a b r i e f d e s c r i p t i o n w i l l be g i v e n h e r e . V e g e t a t i o n Many o f t h e s p e c i e s t h a t o c c u r i n t h e d r i e r m i c r o c o m m u n i t i e s o f t h e bogs d o m i n a t e on t h e s e o u t c r o p s ; t h i s e x p l a i n s t h e o r d i n a t i o n o f t h i s r e l e v e ( r e l e v e 1) b e t w e e n t h e bogs and bog w o o d l a n d s ( F i g s . 5 - 7 ) . M o i s t u r e s t r e s s ( p h y s i o l o g i c a l i n t h e bogs and p h y s i c a l on t h e o u t c r o p s ) i s common t o b o t h h a b i t a t s . 69 F i g . 30. Empetrum nigrum - Vaccinium uliginosum - Rhacomitrium lanuginosum - Cladonia spp. outcrop community; releve 1. F i g . 31. An Orth ic Regosol supporting the above outcrop community ( F ig . 30) ; re leve 1. (flu 70 D o m i n a n t s h r u b s a r e C h a m a e c y p a r i s n o o t k a t e n s i s , V a c c i n i u m  u l i g i n o s u m , G a u l t h e r i a s h a l l o n , J u n i p e r u s communis and P i n u s  c o n t o r t a . The h e r b l a y e r i s d o m i n a t e d by l e a t h e r y l e a v e d s p e c i e s s u c h as Empetrum n i g r u m , L o i s e l e u r i a p r o c u m b e n s , Andromeda  p o l i f o l i a , and V a c c i n i u m v i t i s - i d a e a . T r i c h o p h o r u m c e s p i - t o s u m , C o p t i s t r i f o l i a , C o r n u s u n a l a s c h k e n s i s , F a u r i a  c r i s t a - g a l l i and S a n g u i s o r b a o f f i c i n a l i s may o c c u r a s w e l l , a l t h o u g h t h e y u s u a l l y show p o o r v i g o u r i n s u c h h a b i t a t s . R h a c o m i t r i u m l a n u g i n o s u m i s t h e d o m i n a n t moss and numerous l i c h e n s s u c h as C l a d o n i a s y l v a t i c a , C. a m a u r o c r a e a , C. r a n g i f e r i n a and C_. g r a c i l i s v a r . e l o n g a t a a l s o d o m i n a t e t h e c r y p t o g r a m l a y e r . S o i l s and l a n d f o r m s O u t c r o p c o m m u n i t i e s o c c u r as i n c l u s i o n s w i t h i n t h e bogs on b e d r o c k e x p o s u r e s o r on l a r g e b o u l d e r s . S o i l s a r e s h a l l o w , w e l l d r a i n e d R e g o s o l s ( F i g . 31) o r F o l i s o l s , o f t e n r e d u c e d t o a few cm o f humus (Hemimors) o v e r r o c k ( n o n - s o i l ) . 4.4.2.3 Pinus c o n t o r t a - Chamaecyparis n o o t k a t e n s i s - (Thuja  p l i c a t a ) - G a u l t h e r i a s h a l l o n - F a u r i a c r i s t a - g a l l i -Pleurozium s c h r e b e r i - Sphagnum spp. bog woodlands. P i n e - c y p r e s s - c e d a r - G a u l t h e r i a - F a u r i a - P l e u r o z ium - Sphagnum bog w o o d l a n d c o m m u n i t i e s c h a r a c t e r i s t i c a l l y f o r m a m o s a i c w i t h t h e open bog c o m m u n i t i e s w i t h i n t h e s t u d y a r e a ( F i g s . 9, 32, 33 & 5 3 ) . The two community t y p e s s h a r e many o f t h e same s p e c i e s b u t a r e q u i t e d i s t i n c t i n o v e r a l l v e g e t a t i v e 71 F i g . 32. Complex of open blanket bog and bog woodland on the level to gentle slopes and upland fo res t on the steeper s lopes ; Tsimpsean Peninsula near Port Simpson, F i g . 33. The pine - cypress - cedar - Gaul ther ia - Faur ia -Pleurozium - Sphagnum bog woodland community type: t ransect 4. 7/A 72 s t r u c t u r e and v i g o u r . The bog w o o d l a n d s h a v e a t r e e c a n o p y a n d a much b e t t e r d e v e l o p e d t a l l s h r u b l a y e r t h a n t h e o p e n b o g s a l t h o u g h , a t b e s t t h e y w o u l d be c o n s i d e r e d v e r y s c r u b b y f o r e s t s . I n c o n t r a s t t o t h e o p e n b o g s , t h e b o g w o o d l a n d s p o s s e s s many s p e c i e s more t y p i c a l o f u p l a n d f o r e s t c o m m u n i t i e s . As a r e s u l t o f t h e c o m b i n a t i o n o f f l o r i s t i c e l e m e n t s f r o m b o t h t h e o p e n b o g s a n d u p l a n d f o r e s t c o m m u n i t i e s , b o g w o o d l a n d s h a v e t h e g r e a t e s t s p e c i e s d i v e r s i t y ( g r e a t e s t number o f s p e c i e s p e r u n i t a r e a ) o f a l l t h e c o m m u n i t y t y p e s d e s c r i b e d w i t h i n t h e s t u d y a r e a . On t h e PO, PCA a n d RA r e l e v e o r d i n a t i o n s ( F i g s . 5 - 7 ) t h e p i n e - c y p r e s s - c e d a r b o g w o o d l a n d s o c c u r b e t w e e n t h e o p e n b o g s and c e d a r - c y p r e s s - h e m l o c k - V a c c i n i u m - C o r n u s -R h y t i d i a d e l p h u s u p l a n d f o r e s t c o m m u n i t i e s . T h i s i l l u s t r a t e s t h e t r a n s i t i o n a l n a t u r e o f t h e bog w o o d l a n d s . V e g e t a t i o n The c h a r a c t e r i s t i c c o m b i n a t i o n o f s p e c i e s f o r t h e p i n e -c y p r e s s - c e d a r - b o g w o o d l a n d s i s l i s t e d i n T a b l e 2. The c o n s t a n t o r d o m i n a n t s p e c i e s a r e i n c l u d e d i n g r o u p s B, C, D, E and F o n t h e PCA s p e c i e s o r d i n a t i o n ( F i g . 8 ) . G r o u p B c o n t a i n s t h e s p e c i e s s h a r e d w i t h t h e o p e n b o g s . A few o f t h e s e ( t h o s e c l o s e s t t o g r o u p C) a r e somewhat p r e f e r e n t i a l f o r t h e b o g w o o d l a n d s . Group C c o n t a i n s t h e p r e f e r e n t i a l s p e c i e s f o r t h i s c o m m u n i t y t y p e . The s p e c i e s o f g r o u p D o c c u r w i t h a p p a r e n t l y e q u a l p r e f e r e n c e i n b o t h t h e b o g w o o d l a n d s and t h e c e d a r - c y p r e s s - h e m l o c k - V a c c i n i u m - C o r n u s - R h y t i d i a -73 Table 2. Vegetative features and character species for the Pinus contorta -Chamaecyparis nootkatensis - (Thuja plicata) - Gaultheria shalon -Fauria crista-galli - Pleurozium schreberi - Sphagnum spp. bog wood-land community type.  General vegetative features - very open, scrubby tree canopy (0-40% cover; x = 17%) - maximum height of tree canopy: +13 m - very wel developed shrub layer (85-95% cover; x = 94%) - diverse and wel developed herb layer (30-95% cover; x = 72%) - very wel developed bryophyte and lichen layer (50-99% cover; x = 90%) Tree layer: character species-*-Pinus contorta CD*; preferential^  Shrub layers: character species    Chamaecyparis nootkatensis  Thuja plicata  Tsuga heterophyla  Tsuga mertensiana  Gaultheria shalon  Ledum groenlandicum  Kalmia microphyla  Menziesia ferruginea  Vaccinium parvifolium Herb layer: character species Fauria crista-galli  Cornus unalaschkensis  Carex pluriflora  Linnaea borealis  Vaccinium vitis-idaea  Coptis trifolia  Coptis aspleniifolia  Sanguisorba officinalis  Empetrum nigrum  Lycopodium clavatum  Lycopodium annotinum  Trientalis europaea  Listera cordata  Vaccinium caespitosum  Toefieldia glutinosa Bryophyte and lichen layer: Pleurozium schreberi  Hylocomium splendens  Rhytidiadelphus loreus  Sphagnum rubelum  Sphagnum papilosum  Sphagnum girgensohni  Cladonia rangiferina  Cladonia gracilis  Sphagnum fuscum Ptilium crista-castrensis character species CD; prefrntial CD; preferential CD; preferential CD C CD CD; preferential CD CD C CD; selective CD CD; preferential CD; (preferential) CD; preferential C CD; (preferential) CD CD C C C C (preferential) (preferential) preferential preferential CD; selective CD CD CD CD CD; preferential C C preferential preferential •••Species are listed in order of decreasing constancy. The lists include only species of highest constancy and dominance as wel as exclusive selective or preferential taxa of lower constancy and/or dominance. Refer to Appendix VI for complete vegetation tables. 2c = constant species (constancy of >^  80%) D = dominant species (average cover abundance value of >3) 3Refer to definitions of character taxa in text (page 29). 74 d e l p h u s u p l a n d f o r e s t s w h i l e t h e s p e c i e s o f g r o u p E a r e p r e f e r e n t i a l f o r t h e l a t t e r . G r o u p F c o n t a i n s s p e c i e s u s u a l l y p r e s e n t i n a l l t h e f o r e s t and w o o d l a n d c o m m u n i t i e s . D w a r f e d , o f t e n f o r k e d and b o n s a i - l i k e i n d i v i d u a l s o f P i n u s c o n t o r t a a v e r a g i n g 10 m i n h e i g h t and 21 cm dbh make up t h e o p e n , s c r u b b y m a i n t r e e c a n o p y i n t h e bog w o o d l a n d s ( F i g . 3 3 ) . Maximum t r e e h e i g h t s e l d o m e x c e e d s 13 m. G r o w t h r a t e s a r e s l o w and m a i n c a n o p y t r e e s a r e u s u a l l y o v e r 200 y e a r s o l d . O t h e r t r e e s p e c i e s a r e m o s t l y r e s t r i c t e d t o s h r u b l a y e r s . T a l l s h r u b s (2 - 10 m) a r e d o m i n a t e d by t h e c o n s t a n t d o m i n a n t s P. c o n t o r t a , C. n o o t k a t e n s i s and T. p l i c a t a . T. h e t e r o p h y l l a and T. m e r t e n s i a n a a r e l e s s a b u n d a n t . A t h i c k , k n e e - h i g h and p r o f u s e l y b r a n c h e d l a y e r o f C. n o o t k a t e n s i s u s u a l l y d o m i n a t e s t h e l o w s h r u b s . E x c e p t f o r T. m e r t e n s i a n a , a l l o f t h e ab o v e s p e c i e s a r e c o n s t a n t d o m i n a n t s i n t h e low s h r u b l a y e r s as w e l l . O t h e r c o n s t a n t d o m i n a n t s i n t h e v e r y w e l l d e v e l o p e d s h r u b l a y e r s a r e G a u l t h e r i a s h a l l o n , Ledum g r o e n l a n d i c u m , K a l m i a  m i c r o p h y l l a s s p . o c c i d e n t a l i s and M e n z i e s i a f e r r u g i n e a . V a c c i n i u m p a r v i f o l i u m i s a c o n s t a n t n o n - d o m i n a n t s p e c i e s . V. o v a l i f o l ium and V. a l a s k a e n s e a r e l e s s a b u n d a n t and a p p a r e n t l y a r e r e s t r i c t e d t o t h e ' d r i e s t ' h a b i t a t s ( e l e v a t e d m i c r o s i t e s ) w i t h i n t h e bog w o o d l a n d s . T h e s e t h r e e V a c c i n i u m s p e c i e s a r e p r e d o m i n a n t l y f o r e s t s p e c i e s , w h e r e a s V. u l i g i n o s u m i s p r i m a r i l y an open bog s p e c i e s t h a t o c c u r s i n t h e s c r u b b i e s t e x a m p l e s o f bog w o o d l a n d . M y r i c a g a l e i s 75 a b u n d a n t i n some bog w o o d l a n d c o m m u n i t i e s and a b s e n t f r o m o t h e r s . As i n t h e open b o g s , i t s p r e s e n c e may r e f l e c t an i m p r o v e d n u t r i e n t s t a t u s . J u n i p e r u s communis i s r a r e i n t h e bog w o o d l a n d s . A d i v e r s e and w e l l d e v e l o p e d h e r b l a y e r t y p i f i e s t h e bog w o o d l a n d s . The mos t c h a r a c t e r i s t i c h e r b i s F a u r i a c r i s t a - g a l l i , a c o n s t a n t d o m i n a n t and s e l e c t i v e s p e c i e s . A l t h o u g h F a u r i a i s o f t e n a b u n d a n t i n t h e open bogs as w e l l , i t shows much b e t t e r v e g e t a t i v e v i g o u r i n t h e bog w o o d l a n d s . I t i s s e l d o m f o u n d f l o w e r i n g , h o w e v e r , e x c e p t i n t h e open b o g s . O t h e r c o n s t a n t d o m i n a n t s p e c i e s a r e C o r n u s u n a l a s c h k e n s i s , C a r e x p l u r i f l o r a ( p r e f e r e n t i a l ) , L i n n a e a b o r e a l i s (+ p r e f e r -e n t i a l ) , V a c c i n i u m v i t i s - i d a e a ( p r e f e r e n t i a l ) C o p t i s  a s p l e n i i f o l i a , ( p r e f e r e n t i a l ) , S a n g u i s o r b a o f f i c i n a l i s s s p . m i c r o c e p h a l a and Empetrum n i g r u m . C o n s t a n t s p e c i e s o f l o w e r d o m i n a n c e a r e C o p t i s t r i f o l i a , L y c o p o d i u m c l a v a t u m (+ p r e f e r e n t i a l ) , L. a n n o t i n u m (+ p r e f e r e n t i a l ) , T r i e n t a l i s  e u r o p a e a s s p . a r c t i c u s and L i s t e r a c o r d a t a . T o f i e l d i a g l u t i n o s a and V a c c i n i u m c a e s p i t o s u m a r e p r e f e r e n t i a l s p e c i e s o f l o w e r c o n s t a n c y ( 7 5 % ) and d o m i n a n c e . L e s s f r e q u e n t s p e c i e s w h i c h a r e , h o w e v e r , a b u n d a n t i n some bog w o o d l a n d c o m m u n i t i e s a r e L y s i c h i t o n a m e r i c a n u m , T r i c h o p h o r u m c e s p i t o s u m , B l e c h n u m s p i c a n t , E r i o p h o r u m a n g u s t - i f o l i u m , C a r e x a n t h o x a n t h e a , Rubus chamaemorus and M a i a n - themum d i l a t a t u m . S p e c i e s o f low ab u n d a n c e o r s p o r a d i c o c c u r r e n c e a r e D r o s e r a r o t u n d i f o l i a , V a c c i n i u m o x y c o c c o s , V e r a t r u m v i r i d e , A g r o s t i s a e q u i v a l v i s , C a r e x p a u c i f l o r a , C a r e x s i t c h e n s i s , P i n g u i c u l a v u l g a r i s , S t r e p t o p u s r o s e u s , G e n t i a n a d o u g l a s i a n a , P l a t a n t h e r a s t r i c t a , C a l a m a g r o s t i s - : n u t k a e n s i s , E r i g e r o n p e r e g r i n u s , Andromeda p o l i f o l i a ' and C a r e x l i v i d a . C l i n t o n i a u n i f l o r a , P l a t a n t h e r a d i l a t a t a v a r . a l b i f l o r a , Rubus p e d a t u s and S t r e p t o p u s a m p l e x i f o l i u s a r e c o n s i d e r e d a c c i d e n t a l i n t r u d e r s i n t h e bog w o o d l a n d s . P l e u r o z i u m s c h r e b e r i i s t h e c h a r a c t e r i s t i c s p e c i e s o f t h e v e r y w e l l d e v e l o p e d b r y o p h y t e and l i c h e n c o m m u n i t y o f t h e f o r e s t f l o o r . I t i s a c o n s t a n t d o m i n a n t and s e l e c t i v e s p e c i e s i n t h e bog w o o d l a n d s . The b r y o p h y t e f l o r a i s c o - d o m i n a t e d by t h e u p l a n d f o r e s t s p e c i e s H y l o c o m i u m s p l e n d e n s , R h y t i d i a - d e l p h u s l o r e u s and Sphagnum g i r g e n s o h n i i , a s w e l l as t h e bog s p e c i e s S_j_ r u b e l l u m and S_. p a p i l l o s u m . A l l a r e c o n s t a n t d o m i n a n t s . S_. f u s c u m and P t i l i u m c r i s t a - c a s t r e n s i s a r e p r e f e r e n t i a l a nd o f t e n a b u n d a n t s p e c i e s o f l o w e r c o n s t a n c y ( 7 5 % ) . The l i c h e n s C l a d o n i a r a n g i f e r i n a and C. g r a c i l i s v a r . e l o n g a t a a r e c o n s t a n t s p e c i e s ( a l t h o u g h s e l d o m d o m i n a n t ) i n t h e bog w o o d l a n d s . L e s s c h a r a c t e r i s t i c c r y p t o g a m s g r o w i n g on humus a r e D i c r a n u m h o w e l l i i , D. f u s c e s c e n s , P l a g i o t h e c i u m u n d u l a t u m , R h i z o m n i u m g l a b r e s c e n s , B a z z a n i a t r i l o b a t a , Hypnum c i r c i n a l e , Rhy t i d i a d e l p h u s , . t r i g u e t r u s , Sphagnum r e c u r v u m , C l a d o n i a - i m p e x a , C. l e u c o p h a e a and C. p s e u d o e v a n s i i . S p o r a d i c s i n c l u d e P l a g i o c h i l a a s p l e n i o i d e s , P e l l i a n e e s i a n a , P l e u r o z i a p u r p u r e a , B a z z a n i a t r i c r e n a t a , H e r b e r t a a d u n c a , L e p i d o z i a r e p t a n s , C l a d o n i a f u r c a t a , C . m u l t i f o r m i s , C . squamosa and C_. u n c i a l i s . 77 D e c a y i n g wood on t h e g r o u n d a s w e l l as s t a n d i n g d e a d s n a g s and l i v i n g t r e e s s u p p o r t s e v e r a l e p i x y l i c and e p i p h y t i c c r y p t o g a m s . A l e c t o r i a s a r m e n t o s a , A. f r e m o n t i i , H y p o g y m n i a  e n t e r o m o r p h a , F r u l l a n i a n i s q u a l l e n s i s (common on C. n o o t -k a t e n s i s ) and H e r b e r t a a d u n c a a r e u s u a l l y a b u n d a n t . O t h e r e p i x y l e s and e p i p h y t e s i n c l u d e A n t i t r i c h i a c u r t i p e n d u l a r  S c a p a n i a b o l a n d e r i , M y l i a a n o m a l a , M. t a y l o r i , I s o t h e c i u m  s t o l o n i f e r u m , P t i l i d i u m c a l i f o r n i c u m , M e t a n e c k e r a menz i e s i i , C e p h a l o z i e l l a d i v a r i c a t a , D o u i n i a o v a t a , J a m e s o n i e l l a a u t u m n a l i s , D i c r a n o w e i s i a c i r r a t a a n d I c m a d o p h i l a e r i c e t o r u m . T h i s l i s t i s n o t c o m p l e t e and many o f t h e s p e c i e s g r o w i n g on humus a l s o o c c u r as e p i p h y t e s o r e p i x y l e s . S o i l s and l a n d f o r m s S o i l s and l a n d f o r m s o f t h e bog w o o d l a n d s a r e s i m i l a r t o t h o s e o f t h e open b o g s : T y p i c and Humic M e s i s o l s and T y p i c F i b r i s o l s d e v e l o p e d p r e d o m i n a n t l y i n o r g a n i c b l a n k e t s . O r g a n i c v e n e e r s a r e l e s s common. A l l e x c e p t one o f t h e bog w o o d l a n d s s a m p l e d a r e u n d e r l a i n by b e d r o c k . The e x c e p t i o n ( r e l e v e 21) i s u n d e r l a i n by f l u v i a l m a t e r i a l s b e n e a t h 2 m o f p e a t . T h e r e i s as much v a r i a b i l i t y i n d e p t h t o b e d r o c k i n t h e bog w o o d l a n d p l o t s as i n t h e open b o g s . A r a n g e o f 100 cm i s common. O r t h i h i s t o m o r s , F e r m i h i s t o m o r s and O r t h i h i s t o m o d e r s a r e t h e most common humus f o r m s . The s o i l s u r f a c e i n t h e bog w o o d l a n d s i s d r i e r t h a n i n t h e o p e n bogs a l t h o u g h s o i l s a r e u s u a l l y s a t u r a t e d 10 t o 15 cm b e l o w t h e s u r f a c e . The ponds and wet d e p r e s s i o n s s o c h a r a c -78 t e r i s t i c o f t h e o pen bogs a r e much l e s s common i n t h e bog w o o d l a n d s . R o o t i n g z o n e pH v a r i e s f r o m 2.7 t o 2.9 i n t h e 5 p r o f i l e s a n a l y z e d . T h i s i s s i m i l a r t o most o f t h e open bog s o i l s w i t h t h e e x c e p t i o n o f t h e s h a l l o w s l o p i n g bogs t h a t have s l i g h t l y h i g h e r pH v a l u e s . The bog w o o d l a n d s l a c k t h e h i g h d i v e r s i t y o f m i c r o -c o m m u n i t i e s c h a r a c t e r i s t i c o f t h e o p e n b o g s . M i c r o t o p o g r a p h y i s more homogeneous and as a c o n s e q u e n c e t h e s o i l s u r f a c e i s r e l a t i v e l y d r y t h r o u g h o u t and p l a n t d i s t r i b u t i o n more u n i f o r m . The bog w o o d l a n d s a r e c l a s s i f i e d as s u b h y g r i c t o h y g r i c and o l i g o t r o p h i c t o s u b m e s o t r o p h i c i n t e r m s o f m o i s t u r e and n u t r i e n t r e g i m e s , r e s p e c t i v e l y ( A p p e n d i x I ) . They a r e c o n s i d e r e d o m b r o p h i l o u s w i t h r e g a r d t o w a t e r r e g i m e , p r o b a b l y r e l y i n g on p r e c i p i t a t i o n f o r most o f t h e i r m i n e r a l n u t r i t i o n . R e l a t i o n s h i p s among t h e r e l e v e s R e l e v e s 37 and 15 s t a n d o u t a t o p p o s i t e ends o f t h e bog w o o d l a n d o r d i n a t i o n s ( F i g s . 5, 6, 7 and 2 9 ) . R e l e v e 37 i s t r a n s i t i o n a l t o t h e o pen b o g s ; t r e e s p e c i e s a r e p r e s e n t as s h r u b s o n l y and a few u n d e r s t o r y s p e c i e s o c c u r ( s u c h as A ndromeda p o l i f o l i a , J u n i p e r u s communis and C l a d o n i a i m p e x a ) t h a t a r e more t y p i c a l o f o pen b o g s . The s o i l h e r e i s a Humic M e s i s o l , 50 t o 160 cm deep o v e r q u a r t z d i o r i t e b e d r o c k on an u p p e r s l o p e o f 12 t o 1 5 % . R e l e v e 15, on t h e o t h e r hand, t e n d s t o w a r d u p l a n d f o r e s t c o n d i t i o n s w i t h t a l l e r , more v i g o r o u s t r e e s and s c a t t e r e d s p e c i m e n s o f S t r e p t o p u s r o s e u s , R u b u s - p e d a t u s and P e l l i a n e e s i a n a as w e l l as d e p a u p e r a t e i n d i v i d u a l s 79 o f P i c e a s i t c h e n s i s a n d A b i e s a m a b i l i s a s l o w s h r u b s . S o i l s i n t h i s c o m m u n i t y a r e a m o s a i c o f s h a l l o w F o l i s o l s and H u m i s o l s 22 t o 70 cm d e e p , o v e r m e t a m o r p h i c b e d r o c k o n a m i d s l o p e o f 2 5%. T h i s s u g g e s t s a more m i n e r o t r o p h i c w a t e r r e g i m e . I t i s p o s s i b l e t h a t t h i s s a m p l e r e p r e s e n t s a d e g r a d e d u p l a n d f o r e s t c o m m u n i t y b e c a u s e a few l a r g e d e a d s n a g s a r e p r e s e n t . S u c c e s s i o n a l r e l a t i o n s h i p s among t h e co m m u n i t y t y p e s w i t h i n t h e s t u d y a r e a w i l l be d e a l t w i t h i n C h a p t e r 6. R e l e v e s 15, 2 1 , 3 1 , and 3, a l l o f w h i c h o r d i n a t e t o w a r d t h e u p l a n d f o r e s t r e l e v e s a r e l o c a t e d on m e t a m o r p h i c b e d r o c k o r f l u v i a l m a t e r i a l s d e r i v e d f r o m t h i s b e d r o c k . A l l e x c e p t r e l e v e 15 o c c u r o n o r g a n i c b l a n k e t s , h o w e v e r , a nd t h u s t h e i n f l u e n c e o f t h e u n d e r l y i n g b e d r o c k o n m i n e r a l n u t r i t i o n i s q u e s t i o n a b l e . U n d e r t h e s e c o n d i t i o n s , one w o u l d e x p e c t t h e i n f l u e n c e o f p r e c i p i t a t i o n f a l l i n g on t h e b o g s u r f a c e t o o u t w e i g h t h e i n f l u e n c e o f m i n e r a l s e e p a g e , e s p e c i a l l y o n s l o p e s o f l e s s t h a n 1 5%. 4.4.2.4 Th u j a p l i c a t a - Chamaecyparis n o o t k a t e n s i s - Tsuga h e t e r o p h y l l a - V a c c i n i u m ( p a r v i f o l i u m - o v a l i f o l i u m - a l a s k a e n s e ) - G a u l t h e r i a s h a l l o n - Cornus u n a l a s c h - k e n s i s - R h y t i d i a d e l p h u s l o r e u s - Hylocomium  sp l e n d e n s - Spaghnum g i r g e n s o h n i i c l i m a t i c c l i m a x u p l a n d f o r e s t s . The c e d a r - c y p r e s s - h e m l o c k - V a c c i n i u m - C o r n u s -R h y t i d i a d e l p h u s c o m m u n i t y t y p e ( F i g s . 34 - 3 7 ) , i s t h e most common u p l a n d f o r e s t t y p e w i t h i n t h e s t u d y a r e a and a p p e a r s t o b e s t r e p r e s e n t c l i m a t i c c l i m a x ( z o n a l ) v e g e t a t i o n i n t h i s F i g . 34. Looking west along transect 2, east of Rainbow Lake. Open blanket bog in foreground grad-ing into a pine - cypress - cedar -bog woodland and then into a cedar -cypress - hemlock upland fores t on far lower s lope. This grades into an amabil is f i r - hemlock -cedar fo res t on the upper s lope. 00 o F i g . 35. The cedar - cypress - hemlock -zonal fo res t on the midslope contrasts with the S i tka spruce a l l u v i a l f o res t at the toe of the s lope . F i g . 36. The cedar - cypress - hemlock -Vaccinium - Cornus - Rhyt id ia - delphus zonal fo res t community type; releve 33. F i g . 37. The cedar - cypress - hemlock -Vaccinium - Cornus - Rhy t id i a - delphus zonal f o r e s t community type; releve" 2. tf / A 82 p o r t i o n o f t h e CCPH z o n e . I t o c c u r s p r e d o m i n a n t l y on i m p e r f e c t l y d r a i n e d m o d e r a t e s l o p e s (15 - 30%)- b u t may a l s o d e v e l o p o n l e v e l g r o u n d o r s t e e p e r s l o p e s w here e d a p h i c f a c t o r s c o m p e n s a t e f o r t o p o g r a p h y a n d e n h a n c e o r i n h i b i t s o i l d r a i n a g e . V e g e t a t i o n The c h a r a c t e r i s t i c c o m b i n a t i o n o f s p e c i e s t y p i f y i n g t h e c e d a r - c y p r e s s - h e m l o c k - V a c c i n i u m - C o r n u s - R h y t i d i a -d e l p h u s c o m m u n i t y t y p e i s l i s t e d i n T a b l e 3. M o s t o f t h e s p e c i e s o f g r o u p s D, E, F a n d G a r e c o n s t a n t s o r d o m i n a n t s i n t h i s c o m m u n i t y t y p e ( F i g . 8 ) . Those o f g r o u p D o c c u r i n b o t h t h e b o g w o o d l a n d s a n d t h e z o n a l u p l a n d f o r e s t s w i t h e q u a l p r e f e r e n c e . The p r e f e r e n t i a l s p e c i e s f o r t h i s z o n a l f o r e s t c o m m u n i t y t y p e a r e c o n t a i n e d i n g r o u p E. The s p e c i e s o f g r o u p F a r e u b i q u i t o u s i n a l l f o r e s t a n d w o o d l a n d c o m m u n i t i e s a l t h o u g h t h o s e c l o s e s t t o g r o u p E may be s l i g h t l y p r e f e r e n t i a l f o r t h e c e d a r - c y p r e s s - h e m l o c k u p l a n d f o r e s t s ( i . e . g r e a t e r v i g o u r a n d / o r a b u n d a n c e ) . G r o u p G c o n t a i n s s p e c i e s s h o w i n g g r e a t e r p r e f e r e n c e f o r b o t h t h e c e d a r - s p r u c e - h e m l o c k -L y s i c h i t o n - R h i z o m n i u m a l l u v i a l f o r e s t s a s w e l l a s f o r t h e a m a b i l i s f i r - h e m l o c k - c e d a r - G y m n o c a r p i u m - R h i z o m n i u m u p l a n d f o r e s t s . T h e s e s p e c i e s a r e u s u a l l y c o n s t a n t d o m i n a n t s i n t h e z o n a l f o r e s t s as w e l l . T r e e c a n o p i e s o f t h i s c o m m u n i t y t y p e a r e g e n e r a l l y s c r u b b y a n d u n e v e n - a g e d , a v e r a g i n g 23 m i n maximum h e i g h t . M a i n c a n o p i e s a v e r a g e 15 m i n h e i g h t . H i g h s t e m d e n s i t y and s n a i l a v e r a g e dbh a r e t y p i c a l ( T a b l e 3 ) . S c a t t e r e d l a r g e d e a d Table 3. Vegetative features, stand characteristics and character species for the Thuja plicata - Chamaecyparis nootkatensis - Tsuga heterophyla -Vaccinium spp. - Gaultheria shalon - Comus unalaschkensis -Rhytidiadelphus loreus - Hylocomium splendens - Sphagnum girgensohni upland forest community type.  General vegetative features and stand characteristics - moderately closed, all-aged, somewhat scrubby tree canopy (40 - 90% cover; x = 64%) - maximum height of tree canopy: +22 m - average dbh: 25 - 60 cm (x = 34 cm) - number of stems per hectare: 215 - 1585 (X = 1013) - basal area per hectare: 52 - 104 m2 (x = 79) - very wel developed shrub layers (80 - 95% cover; x = 91%) - moderately wel developed herb layer (15 - 95% cover; X - 56%) - very wel developed bryophyte layer Tree layers: character species* Thuja plicata  Chamaecyparis nootkatensis  Tsuga heterophyla CD Tsuga mertensiana (C); Shrub layers: character species Tsuga heterophyla  Chamaecyparis nootkatensis  Thuja plicata  Gaultheria shalon  Menziesia ferruginea  Vaccinium parvifolium  Vaccinium ovalifolium  Vaccinium alaskaense Herb layer: character species Cornus unalaschkensis CD; Streptopus roseus CD Coptis aspleniifolia C(D) Listera cordata C Lvcopodium clavatum C Blechnum spicant CD Rubus pedatus CD Linnaea borealis C Lysichiton americanum (C)D Vaccinium vitis-idaea (C) Veratrum viride (C); Clintonia uniflora D; Listera caurina  Orthilia secunda  Calamagrostis nutkaensis Bryophyte and lichen layer: character species Rhytidiadelphus loreus CD; Hylocomium splendens CD; Sphagnum girgensohni  CD Rhizomnium glabrescens C(D) (90 - 98% cover; x = 95%) CD2; preferential3 CD; preferential preferential CD CD CD CD; preferential CD CD CD; preferential (C)D preferential preferential preferential (preferential) exclusive preferential preferential preferential -^Species are listed in order of decreasing constancy. The lists include only species of highest constancy and dominance as wel as exclusive selective or preferential taxa of lower constancy and/or dominance. Refer to Appendix VI for complete vegetation tables. 2c = constant species (constancy of >^  80%) i D = dominant species (average cover abundance value of >3) 3Refer to definitions of character taxa in text (page 29). 84 s n a g s a r e o f t e n p r e s e n t t h r o u g h o u t t h i s community t y p e . M a i n c a n o p y t r e e s a v e r a g e 250 y e a r s i n age. S p i k e - t o p p e d T. p l i c a t a and C_. n o o t k a t e n s i s u s u a l l y d o m i n a t e t h e u p p e r t r e e c a n o p y w i t h T. h e t e r o p h y l l a i n c r e a s i n g i n a b u n d a n c e i n t h e m a i n c a n o p y and s u p p r e s s e d t r e e l a y e r s . T. m e r t e n s i a n a i s o f t e n p r e s e n t i n t h e s u p p r e s s e d t r e e l a y e r s a s w e l l . P. c o n t o r t a , when p r e s e n t , i s u s u a l l y r e s t r i c t e d t o t h e m a i n c a n o p y o r d o m i n a n t t r e e l a y e r s . I n t e r e s t i n g l y , t h i s s p e c i e s o f t e n shows t h e b e s t g r o w t h o f a l l t h e t r e e s p e c i e s i n t h i s c o m m u n i t y t y p e ( s e e s e c t i o n 4 . 4 . 4 ) . B a s a l a r e a v a l u e s r a n g e f r o m 52 t o 104 m^/ha (x = 79) among t h e 9 r e l e v e " s ( T a b l e 3 ) . B a s a l a r e a v a l u e s a r e u s u a l l y h i g h e s t f o r T. p i i c a t a , f o l l o w e d by C. n o o t k a t e n s i s , T. h e t e r - o p h y l l a , P. c o n t o r t a and T. m e r t e n s i a n a . W i t h t h e e x c e p t i o n o f t h e s h a d e i n t o l e r a n t P. c o n t o r t a , w h i c h o n l y r e g e n e r a t e s i n v e r y s c r u b b y , open f o r e s t s , bog w o o d l a n d s and o p e n b o g s , a l l o f t h e ab o v e t r e e s p e c i e s s u c c e s s f u l l y r e g e n e r a t e w i t h i n t h i s c o m munity t y p e . T. h e t e r - ; o p h y l l a r e g e n e r a t i o n i s most a b u n d a n t , f o l l o w e d by C. n o o t - k a t e n s i s , T. p i i c a t a , and T. m e r t e n s i a n a . P i c e a s i t c h e n s i s and A b i e s a m a b i l i s a r e uncommon and when p r e s e n t a r e u s u a l l y o f e x t r e m e l y p o o r f o r m and v i g o u r . I n a d d i t i o n t o t r e e r e g e n e r a t i o n , t h e v i g o r o u s s h r u b l a y e r s a r e d o m i n a t e d by G a u l t h e r i a s h a l l o n , M e n z i e s i a  f e r r u g i n e a , V a c c i n i u m p a r v i f o l i u m , V. o v a l i f o l i u m and V. a l a s k a e n s e . G. s h a l l o n and V. o v a l i f o l i u m a r e c o n s i d e r e d p r e f e r e n t i a l s p e c i e s f o r t h i s c o m m u n i t y t y p e . Ledum g r o e n -85 l a n d i c u m a n d , t o a much l e s s e r e x t e n t , K a l m i a m i c r o p h y l l a s s p . o c c i d e n t a l i s s o m e t i m e s o c c u r i n t h e w e t t e s t and s c r u b b i e s t e x a m p l e s o f t h i s e c o s y s t e m . C l a d o t h a m n u s p y r o l i f l o r u s , Rubus  s p e c t a b i l i s a n d O p l o p a n a x h o r r i d u s a r e v e r y s p o r a d i c a n d o f p o o r v i g o u r . They a r e c o n s i d e r e d a c c i d e n t a l i n t r u d e r s . H e r b l a y e r s a r e u s u a l l y w e l l d e v e l o p e d and d o m i n a t e d by C o r n u s u n a l a s c h k e n s i s ( a w i d e s p r e a d s p e c i e s t h a t n e v e r t h e l e s s i s p r e f e r e n t i a l f o r t h i s c o m m u n i t y t y p e ) , S t r e p t o p u s r o s e u s , C o p t i s a s p l e n i i f o l i a , L i s t e r a c o r d a t a , L y c o p o d i u m c l a v a t u m , B l e c h n u m s p i c a n t , Rubus p e d a t u s and L i n n a e a b o r e a l i s . L y s i c h i t o n a m e r i c a n u m and V e r a t r u m v i r i d e a r e n o r m a l l y r e s t r i c t e d t o w e t s e e p a g e a r e a s . The l a t t e r i s c o n s i d e r e d p r e f e r e n t i a l f o r t h i s c o m m u n i t y t y p e . O t h e r common h e r b s p e c i e s o f l o w e r c o n s t a n c y (55 - 78%) i n c l u d e V a c c i n i u m v i t i s - i d a e a , C l i n t o n i a u n i f l o r a ( p r e f e r e n -t i a l ) , F a u r i a c r i s t a - g a l l i , L y c o p o d i u m a n n o t i n u m and L i s t e r a  c a u r i n a (+ p r e f e r e n t i a l ) . S p o r a d i c s p e c i e s (22 - 3 3 % c o n s t a n c y ) i n c l u d e C a l a m a g r o s t i s n u t k a e n s i s ( p r e f e r e n t i a l ) , Maianthemum d i l a t a t u m , O r t h i l i a s e c u n d a ( e x c l u s i v e ) and C a r e x  a n t h o x a n t h e a . R h y t i d i a d e l p h u s l o r e u s a n d H y l o c o m i u m s p l e n d e n s d o m i n a t e t h e v e r y w e l l d e v e l o p e d b r y o p h y t e l a y e r on t h e f o r e s t f l o o r . B o t h a r e c o n s i d e r e d p r e f e r e n t i a l s p e c i e s f o r t h i s c o m m u n i t y t y p e . Sphagnum g i r g e n s o h n i i i s a l s o a c o n s t a n t d o m i n a n t . T h i s s p e c i e s c a n h a v e a l a r g e i m p a c t o n s o i l m o i s t u r e r e g i m e b y f o r m i n g e x t e n s i v e s p o n g y mats w h i c h impede d r a i n a g e o v e r l a r g e a r e a s . I t i s u s u a l l y p r e s e n t t o some d e g r e e i n most o f 86 t h e f o r e s t c o m m u n i t i e s w i t h i n t h e s t u d y a r e a . O t h e r Sphagnum s p e c i e s a r e r e s t r i c t e d m a i n l y t o t h e open bogs and bog wood-l a n d s , a l t h o u g h s e v e r a l o c c u r s p o r a d i c a l l y i n t h e s c r u b b i e s t u p l a n d f o r e s t s . R h i z o m n i u m g l a b r e s c e n s i s a l s o a c o n s t a n t and s o m e t i m e s d o m i n a n t s p e c i e s i n t h i s c ommunity t y p e . M i n o r n o n - c o n s t a n t c o m p o n e n t s o f t h e f o r e s t f l o o r b r y o p h y t e f l o r a i n c l u d e P l a g i o - t h e c i u m u n d u l a t u m , D i c r a n u m h o w e l l i i , P l a g i o c h i l a a s p l e n - i o i d e s , P l e u r o z ium s c h r e b e r i , R h y t i d i a d e l p h u s t r i q u e t r u s and L e p i d o z i a r e p t a n s . S p o r a d i c s p e c i e s (< 33% c o n s t a n c y ) a r e P e l l i a n e e s i a n a , B a z z a n i a t r i c r e n a t a , B. t r i l o b a t a , H o o k e r i a  l u c e n s , L o p hoz i a s p p . , D i c r a n u m m a j u s , P t i l i u m c r i s t a - c a s t r e n s i s , M y l i a a n o m a l a and S c a p a n i a b o l a n d e r i . L i c h e n s o c c u r o n l y s p o r a d i c a l l y on t h e f o r e s t f l o o r and r a r e l y w i t h c o v e r - a b u n d a n c e v a l u e s o f g r e a t e r t h a n '+'. C l a d o n i a f u r c a t a , C. g r a c i l i s v a r . e l o n g a t a , C. l e u c o p h a e a , C. r a n g i f e r i n a and P e l t i g e r a a p h t h o s a a r e e x a m p l e s . H e r b e r t a a d u n c a i s a common e p i p h y t e . E x t e n s i v e g o l d e n b rown c l u m p s o f t h i s c h a r a c t e r i s t i c l i v e r w o r t f r e q u e n t l y d r a p e t r e e b r a n c h e s , s o m e t i m e s i n a s s o c i a t i o n w i t h A n t i t r i c h i a  c u r t i p e n d u l a and M e t a n e c k e r a m e n z i e s i i . O t h e r f r e q u e n t l y o c c u r r i n g s p e c i e s o f t h e u s u a l l y d i v e r s e and w e l l d e v e l o p e d e p i p h y t i c and e p i x y l i c b r y o p h y t e c o m m u n i t i e s i n c l u d e , i n a d d i t i o n t o many o f t h o s e c o l o n i z i n g t h e f o r e s t f l o o r , F r u l l a n i a n i s q u a l l e n s i s (common on C. n o o t k a t e n s i s ) , I s o t h e c i u m s t o l o n i f e r u m , C l a o p o d i u m b o l a n d e r i , D i c r a n e l l a  h e t e r o m a l l a , M y l i a t a y l o r i , R a d u l a b o l a n d e r i , B l e p h a r o s t o m a 87 t r i c h o p h y l l u m , C e p h a l o z i e l l a d i v a r i c a t a , P t i l i d i u m c a l i -f o r n i c u m , C a l y p o g e i a m u e l l e r i a n a , D i c r a n o w e i s i a c i r r a t a a n d Hypnum s u b i m p o n e n s . The b e a r d l i c h e n s A l e c t o r i a s a r m e n t o s a a n d A. f r e m o n t i i a r e a l s o common e p i p h y t e s . T h i s l i s t i s n o t c o m p l e t e s i n c e t h e s a m p l i n g e m p h a s i s was o n s p e c i e s o f t h e f o r e s t f l o o r . S o i l s a n d l a n d f o r m s S o i l s o f t h e c e d a r - c y p r e s s - h e m l o c k - V a c c i n i u m  C o r n u s - R h y t i d i a d e l p h u s c o m m u n i t y t y p e c h a r a c t e r i s t i c a l l y h a v e p e r m a n e n t l y w e t s u r f a c e o r g a n i c a c c u m u l a t i o n s ( L F H h o r i z o n s ) 20 t o 3 5 cm t h i c k ( r a n g e o f 18 t o 58 cm). T h e s e m o s t o f t e n o c c u r o v e r Humic a n d F e r r o - H u m i c P o d z o l s 2 t h a t h a v e d e v e l o p e d i n v e n e e r s o f c o a r s e l o a m y m i n e r a l m a t e r i a l ( w e a t h e r e d b e d r o c k ) ( F i g s . 38 a n d 3 9 ) . LFH h o r i z o n s a l s o o c c u r d i r e c t l y o v e r b e d r o c k . A c c o r d i n g t o t h e r e c e n t l y p r o p o s e d r e v i s i o n s t o t h e C a n a d i a n S y s t e m o f S o i l C l a s s i -f i c a t i o n ( T r o w b r i d g e 1 9 8 1 ) , most s o i l s c a n be c l a s s i f i e d a s f o l i s o l i c p h a s e s (15 t o 40 cm LFH h o r i z o n s ) o f Humic and F e r r o - H u m i c P o d z o l s , o r L i t h i c F o l i s o l s (>10 cm LFH h o r i z o n s d i r e c t l y o v e r r o c k ) . T e r r i c F o l i s o l s ( c o n s i s t i n g o f g r e a t e r t h a n 40 cm LFH h o r i z o n s o v e r u n c o n s o l i d a t e d m i n e r a l m a t e r i a l ) w e r e a l s o o b s e r v e d w i t h i n t h e s t u d y a r e a a l t h o u g h none were s a m p l e d . E x a m p l e s o f t h i s c o m m u n i t y t y p e w e r e a l s o d e s c r i b e d on " ^ C l a s s i f i c a t i o n o f Humic and F e r r o - H u m i c P o d z o l s i s b a s e d o n f i e l d i d e n t i f i c a t i o n o f Bh and B h f h o r i z o n s w h i c h h a v e n o t been c o n f i r m e d b y c h e m i c a l a n a l y s i s . CO oo F i g . 3 8 . An O r t h i c F e r r o - Humic Podzo l ( f o l i s o l i c phase ) s u p p o r t i n g a c e d a r - c y p r e s s - hemlock z o n a l f o r e s t commun i t y ; r e l e v e 4 . F i g . 3 9 . An O r t h i c Humic P o d z o l s u p p o r t i n g t h e c e d a r - c y p r e s s - hemlock z o n a l f o r e s t community o f F i g . 3 7 . ; r e l e v e 2. 89 o r g a n i c d e p o s i t s ( H u m i s o l s ) o v e r w h i c h t h i c k l a y e r s (35 t o 60 cm) o f f o r e s t humus have a c c u m u l a t e d ( F i g . 4 0 ) . L a y e r s o f s e d i m e n t a r y p e a t ( s i m i l a r t o what o c c u r s i n bog p o n d s ) and s e d g e r e m a i n s w i t h i n t h e s e H u m i s o l p r o f i l e s s u g g e s t t h a t t h e y a r e r e l i c s o f w e t l a n d e c o s y s t e m s t h a t were i n v a d e d by f o r e s t . The r e s t r i c t i o n o f mo s t o f t h e r o o t s t o t h e s u r f a c e f o r e s t humus l a y e r s o f t h e s e s o i l s i n d i c a t e s t h a t o v e r t i m e n u t r i e n t c y c l i n g has become r e s t r i c t e d t o t h e ' F o l i s o l ' t h a t h a s d e v e l o p e d o v e r w e t l a n d p e a t ( i . e . O r g a n o F o l i s o l , T r o w b r i d g e 1 9 8 1 ) . T h i s i s a n a l o g o u s t o t h e d e v e l o p m e n t o f F o l i s o l s o v e r u n c o n s o l i d a t e d m i n e r a l m a t e r i a l ( T e r r i c F o l i s o l s ) o r b e d r o c k ( L i t h i c F o l i s o l s ) . The a p p a r e n t t r a n s i t i o n f r o m f o r e s t humus (LFH h o r i z o n s ) t o w e t l a n d p e a t (Oh h o r i z o n s ) a t 80 cm d e p t h i n one o f t h e s e f o r e s t e d O r g a n o F o l i s o l s ( r e l e v e * 1 6 ; F i g . 40) was C-14 d a t e d a t 2310 +60 y e a r s (WAT 6 6 6 ) . T h i s s u g g e s t s t h a t t h e e x i s t i n g f o r e s t e c o s y s t e m h a s d e v e l o p e d w i t h i n t h e l a s t 2400 y e a r s . R e s i d u o h u m i m o r s , w i t h r e s i d u a l H h o r i z o n s d o m i n a t i n g , a r e t h e m o s t common humus f o r m o f t h i s c o m m u n i t y t y p e . O r t h i -h i s t o m o r s , R e s i d u o h e m i h u m i m o r s and ( r a r e l y ) A m p h i h e m i m o r s have b e e n d e s c r i b e d as w e l l . S o i l s o f t h i s c o m munity t y p e a r e o n l y s l i g h t l y l e s s a c i d i c t h a n t h o s e o f t h e p i n e - c y p r e s s - G a u l t h e r i a - F a u r i a - P l e u r o z ium bog w o o d l a n d s . PH v a l u e s w i t h i n t h e r o o t i n g zone r a n g e f r o m 2.9 t o 3.2 ( x = 3 . 0 ) . S o i l s a r e g e n e r a l l y i m p e r f e c t l y d r a i n e d w i t h s e e p a g e commonly o c c u r r i n g w i t h i n 40 cm o f t h e s u r f a c e . A m e s i c t o 90 F i g . 40. An Organo F o l i s o l cons i s t ing of 60 cm of fo res t humus (mostly H) over a 75 cm th ick Oh hor izon ; releve' 16. The whit ish horizon at 70-80 cm is sedimentary peat. This s o i l present ly supports a cedar - cypress - hemlock zonal upland fo res t community. Further explanat ion in text ( page 89, ). F i g . 41. A cedar - spruce - hemlock - Lys i ch i ton - Rhizomnium a l l u v i a l f o res t community; t ransect 4. °)0A 91 subhygric moisture regime and submesotrophic to mesotrophic nutrient regime (Appendix I) characterize t h i s community type. Relationships among the releves Samples of the cedar - cypress - hemlock - Vaccinium -Cornus - Rhytidiadelphus community type consistently ordinate between the more productive upland forest communities and the bog woodlands (Figs. 5 - 7 ) . The position of releve 20 suggests relationships with the more productive forests, Which i s understandable since t h i s community has developed on a steeper than average slope (45%) over metamorphic bedrock. Traces of T i a r e l l a t r i f o l i a t a and Oplopanax horridus, both in d i c a t i n g higher nutrient status, are present and the bryophyte f l o r a i s dominated by Rhizomnium glabrescens and P e l l i a neesiana rather than the usual dominants Rhytidiadelphus loreus and Hylocomium splendens. It i s interesting that the two communities (releves 16 and 33) developing on Humisols do not stand out on the o r d i -nations as s i g n i f i c a n t l y d i f f e r e n t f l o r i s t i c a l l y from the other samples of t h i s community type that have developed on Podzols. This emphasizes the major unifying role of humus i n vegetation development. It i s i n the humus that most of the very fine to medium roots occur and t h i s suggests that nutrient exchange takes place primarily i n the surface organic layers. 92 4.4 . 2 . 5 T h u j a p l i c a t a - P i c e a s i t c h e n s i s - T s u g a h e t e r o - p h y l l a - A l n u s r u b r a - ( A b i e s a m a b i l i s ) - L y s i c h i t o n  a m e r i c a n u m - G y m n o c a r p i u m d r y o p t e r i s - R h i z o m n i u m  g l a b r e s c e n s - P e l l i a n e e s i a n a a l l u v i a l f o r e s t s . The c e d a r - s p r u c e - h e m l o c k - L y s i c h i t o n - R h i z o m n i u m c o m m u n i t y t y p e o c c u r s on a l l u v i a l l a n d f o r m s a d j a c e n t t o c r e e k s a nd r i v e r s ( F i g s . 41 and 4 2 ) . I t i s uncommon a l o n g t h e s t u d y t r a n s e c t s and i s r e p r e s e n t e d by o n l y 3 r e l e v e s . V e g e t a t i o n The c h a r a c t e r i s t i c c o m b i n a t i o n o f s p e c i e s and s t a n d c h a r a c t e r i s t i c s f o r t h i s c o mmunity t y p e a r e l i s t e d i n T a b l e 4. C o n s t a n t a n d / o r d o m i n a n t s p e c i e s a r e i n c l u d e d i n g r o u p s F, G and H on t h e PCA s p e c i e s o r d i n a t i o n ( F i g . 8 ) . G r o u p F c o n t a i n s s p e c i e s u b i q u i t o u s i n a l l t h e f o r e s t and w o o d l a n d c o m m u n i t i e s . A few o f t h e s e s p e c i e s a p p e a r t o be p r e f e r e n t i a l f o r t h e a l l u v i a l f o r e s t s . T h i s c o m m u n i t y t y p e s h a r e s many o f t h e p r e f e r e n t i a l s p e c i e s o f t h e a m a b i l i s f i r - h e m l o c k - c e d a r - Gym n o c a r p i u m - R h i z o m n i u m f o r e s t s c o n t a i n e d i n g r o u p s G and H. T h u j a p i i c a t a and T s u g a h e t e r o p h y l l a d o m i n a t e t h e m a i n t r e e c a n o p y w i t h l e s s e r amounts o f A l n u s r u b r a and A b i e s  a m a b i l i s . The l a r g e s t and d o m i n a n t t r e e s a r e u s u a l l y P i c e a  s i t c h e n s i s a l t h o u g h t h i s s p e c i e s i s i n f r e q u e n t o r a b s e n t i n t h e m a i n c a n o p y and s u p p r e s s e d l a y e r s e x c e p t i n r e l a t i v e l y y o u n g s t a n d s . I n c o m p a r i s o n t o t h e c e d a r - c y p r e s s - h e m l o c k z o n a l f o r e s t s , t h e t r e e c a n o p y o f t h e a l l u v i a l f o r e s t community t y p e CO F i g . 4 2 . A c e d a r - s p r u c e - hemlock - F i g . 4 3 . A Rego Humic G l e y s o l s u p p o r t -: l y s i c h i t o n -Rhizomnium'comm- i n g the a l l u v i a l f o r e s t o f F i g , u n i t y ; r e l e v e 10 . 4 2 . ; r e l e v e 10 . 94 Table 4. Vegetative features, stand characteristics and character species for the Thuja plicata - Picea sitchensis - Tsuga heterophyla - Alnus rubra - (Abies amabilis) - Lysichiton americanum - Gymnocarpium dryopteris - Rhizomnium glabrescens  community type.  Pellia neesiana alluvial forest General vegetative features and stand characteristics (limited data; 3 plots) - closed tree canopy (65 - 80% cover; x = 77%) - maximum height of tree canopy: 30 - 32 m (x = 31) - average height of main canopy trees: 17 - 21 m (x = 20) - average stand dbh: 63 - 74 cm (x = 68) - number of stems per hectare: 252 - 301 (x = 281) - basal area per hectare: 88 - 125 m2 (x = 102) - moderate to very wel developed shrub layer (50 - 90% cover; x = 70%) - moderately wel developed herb layer (40 - 70% cover; x = 53%) - wel developed bryophyte layer (70 - 95% cover; x = 87%) - average height of main canopy trees: 17 - 21 m (x = 20) Tree layers: character species* Tsuga heterophyla  Thuja plicata  Alnus rubra  Picea sitchensis Shrub layers: character species Tsuga heterophyla  Menziesia ferruginea  Vaccinium parvifolium  Gaultheria shalon  Vaccinium alaskaense  Vaccinium ovalifolium  Rubus spectabilis Herb Layer: character species Cornus unalaschkensis  Rubus pedatus  Lysichiton americanum  Gymnocarpium dryopteris  Streptopus roseus  Blechnum spicant  Listera cordata  Coptis aspleniifolia  Mainthemum dilatatum  Clintonia uniflora Bryophyte and lichen layer: character species Sphagnum girgensohni  Hylocomium splendens  Rhizomnium glabrescens  Pellea neesiana  Plagiothecium undulatum  Plagiothecium laetum  Rhytiadiadelphus loreus Hookeria lucens CD2 CD; preferential3 CD; exclusive D; preferential CD CD; preferential CD; preferential D D preferential CD CD; + preferential CD; preferential CD; + preferential CD; + preferential CD C D D preferential D CD CD CD CD; preferential C C + preferential CD + preferential *Species are listed in order of decreasing constancy. The lists include only species of highest constancy and dominance as wel as exclusive selective or preferential taxa of lower constancy and/or dominance. Refer to Appendix VI for complete vegetation tables. 2c = constant species (constancy of ^> 80%) D = dominant species (average cover abundance value of >3) 3Refer to definitions of character taxa in text (page 29). i s g e n e r a l l y t a l l e r and much more v i g o r o u s w i t h l a r g e r a v e r a g e dbh, f e w e r stems p e r h e c t a r e and g r e a t e r b a s a l a r e a p e r h e c t a r e ( T a b l e 4 ) . M a i n c a n o p y t r e e s a v e r a g e 20 m (17 t o 21 m) i n h e i g h t and 60 cm dbh. D o m i n a n t t r e e s r e a c h a maximum h e i g h t o f 30 t o 32 m. S a m p l e t r e e s f r o m t h e m a i n c a n o p y a v e r a g e d 210 y e a r s o l d . S h r u b l a y e r s a r e m o d e r a t e l y t o v e r y w e l l d e v e l o p e d w i t h a b u n d a n t r e g e n e r a t i o n o f T s u g a h e t e r o p h y l l a . A b i e s a m a b i l i s a n d P i c e a s i t c h e n s i s r e g e n e r a t e o n l y s p o r a d i c a l l y and T h u j a  p i i c a t a i s u s u a l l y a b s e n t f r o m t h e s h r u b l a y e r s . C o n s t a n t d o m i n a n t s h r u b s s p e c i e s a r e M e n z i e s i a f e r r u g i n e a ( p r e f e r e n t i a l ) and V a c c i n i u m p a r v i f o l i u m ( p r e f e r e n t i a l ) . V a c c i n i u m a l a s k a e n s e and V. o v a l i f o l i u m a r e o f t e n d o m i n a n t as w e l l . The m o d e r a t e l y w e l l d e v e l o p e d h e r b l a y e r i s c o n s t a n t l y d o m i n a t e d by t h e p r e f e r e n t i a l s p e c i e s L y s i c h i t o n a m e r i c a n u m , G y m n o c a r p i u m d r y o p t e r i s , Rubus p e d a t u s and S t r e p t o p u s r o s e u s a s w e l l as C o r n u s u n a l a s c h k e n s i s and B l e c h n u m s p i c a n t . L i s t e r a c o r d a t a i s c o n s t a n t b u t n o t d o m i n a n t and C o p t i s  a s p l e n i i f o l i a , C l i n t o n i a u n i f l o r a and Maianthemum d i l a t a t u m a r e s o m e t i m e s d o m i n a n t ; M. d i l a t a t u m i s c o n s i d e r e d p r e f e r e n t i a l f o r t h e a l l u v i a l f o r e s t s . S p o r a d i c h e r b s i n c l u d e T i a r e l l a t r i f o l i a t a , S t r e p t o p u s a m p l e x i f o l i u s , L i s t e r a c o r d a t a , A t h y r i u m f i l i x - f e m i n a , H u p e r z i a s e l a g o , T i a r e l l a  t r i f o l i a t a , V e r a t r u m v i r i d e and V i o l a g l a b e l l a . The b r y o p h y t e l a y e r o f t h e f o r e s t f l o o r i s w e l l d e v e l o p e d and t y p i f i e d by t h e c o n s t a n t d o m i n a n t s Sphagnum g i r g e n s o h n i i , 96 H y l o c o m i u m s p l e n d e n s , R h i z o m n i u m g l a b r e s c e n s , P e l l i a n e e s i a n a ( p r e f e r e n t i a l ) and R h y t i d i a d e l p h u s l o r e u s . P l a g i o t h e c i u m  u n d u l a t u m , P. l a e t u m (+ p r e f e r e n t i a l ) , P ogonatum a l p i n u m , H o o k e r i a l u c e n s (+ p r e f e r e n t i a l ) , P l a g i o c h i l a a s p l e n i o i d e s , P t i l i u m c r i s t a - c a s t r e n s i s and D i c r a n u m h o w e l l i i a r e s o m e t i m e s a b u n d a n t as w e l l . S p o r a d i c b r y o p h y t e s o f t h e f o r e s t f l o o r i n c l u d e Sphagnum s q u a r r o s u m , C a l y p o g e i a i n t e g r i s t i p u l a , D i p l o p h y l l u m a l b i c a n s , L e p i d o z i a r e p t a n s , L o p h o z i a s p p . , P o g o n a t u m c o n t o r t u m and B a z z a n i a t r i c r e n a t a . M o s t o f t h e s p e c i e s o c c u r r i n g on d e c a y i n g wood and as e p i p h y t e s i n t h e c e d a r - c y p r e s s - h e m l o c k z o n a l f o r e s t c o m m u n i t y t y p e ( s e c t i o n 4.4.2.4) a r e a l s o f o u n d i n t h e f l u v i a l f o r e s t s , a l t h o u g h e p i p h y t i c and e p i x y l i c c o m m u n i t i e s a r e g e n e r a l l y n o t as d i v e r s e o r w e l l d e v e l o p e d . T h i s i s l i k e l y t h e r e s u l t o f f e w e r t r e e s p e c i e s , f e w e r d e a d s n a g s and d e a d -t o p p e d t r e e s and a c l o s e r c a n o p y i n t h e a l l u v i a l f o r e s t s . S o i l s and l a n d f o r m s Wet, g l e y e d , f i n e loamy t o s a n d y m i n e r a l s o i l s (Rego G l e y s o l s , G l e y e d F e r r o - H u m i c P o d z o l s ) w i t h Hydromor and Humimor humus d e v e l o p m e n t c h a r a c t e r i z e t h i s c ommunity t y p e ( F i g . 43). The m i n e r a l s o i l s a r e deep (> 100 cm) b u t i m p e r -f e c t l y t o p o o r l y d r a i n e d due t o t h e f l u c t u a t i n g w a t e r t a b l e o f t h e s e r i p a r i a n l a n d f o r m s . B u r i e d s u r f a c e o r g a n i c and m i n e r a l h o r i z o n s may o c c u r . T h e s e r e s u l t f r o m p a s t e p i s o d e s o f f l u v i a l d e p o s i t i o n . The f a i r l y t h i c k Mor humus f o r m s (26 t o 35 cm) i n t h e t h r e e r e l e v e s r e p r e s e n t i n g t h i s c o m munity t y p e s u g g e s t , h o w e v e r , t h a t t h e s e a l l u v i a l l a n d f o r m s a r e 97 p r e s e n t l y i n a c t i v e . A c t i v e a l l u v i a l l a n d f o r m s c h a r a c t e r -i s t i c a l l y show more e v i d e n c e o f b u r i e d h o r i z o n s c l o s e t o t h e s u r f a c e ( C u m u l i c R e g o s o l s ) , h a v e t h i n n e r Mormoder o r Moder humus f o r m s and g e n e r a l l y show g r e a t e r i n t e r m i x i n g o f m i n e r a l and o r g a n i c m a t e r i a l s a t t h e s o i l s u r f a c e t h a n was o b s e r v e d i n t h e t h r e e s a m p l e s . Humus l a y e r pH o f t h e two p r o f i l e s r a n g e s f r o m 2.1 t o 2.5 ( s i m i l a r v a l u e s were o b t a i n e d f o r t h e r o o t i n g z o n e ) . S u r p r i s i n g l y , t h e s e s o i l s a p p e a r t o be s l i g h t l y more a c i d i c t h a n t h o s e o f t h e much s c r u b b i e r z o n a l f o r e s t c o m m u n i t i e s . More s a m p l e s a r e r e q u i r e d , h o w e v e r , t o s u b s t a n t i a t e t h i s d i f f e r e n c e . T h i s c o m m u n i t y t y p e i s , c o n s i d e r e d m e s i c t o s u b h y g r i c and p e r m e s o t r o p h i c ( A p p e n d i x I ) . R e l a t i o n s h i p s among t h e r e l e v e s On t h e PCA and RA r e l e v e o r d i n a t i o n s ( F i g s . 6 and 7 ) , t h e t h r e e s a m p l e s r e p r e s e n t i n g t h e c e d a r - s p r u c e - h e m l o c k -L y s i c h i t o n - R h i z o m n i u m f l u v i a l f o r e s t c ommunity t y p e a r e w e l l d i s t i n g u i s h e d f r o m t h e c e d a r - c y p r e s s - h e m l o c k - V a c c i n i u m z o n a l f o r e s t c o m m u n i t y t y p e . The d i s t i n c t i o n on t h e o r d i -n a t i o n s b e t w e e n t h e f l u v i a l f o r e s t s and t h e a m a b i l i s f i r -h e m l o c k - c e d a r - G y m n o c a r p i u m u p l a n d f o r e s t s d e s c r i b e d b e l o w i s l e s s o b v i o u s , h o w e v e r . T h e s e two more p r o d u c t i v e f o r e s t c o m m u n i t y t y p e s s h a r e many o f t h e same c o n s t a n t d o m i n a n t s p e c i e s . 98 4.4.2.6 Abies amabilis - Tsuga heterophylla - Thuja p l i c a t a -Vaccinium (alaskaense-parvifolium) - Oplopanax  horridus - Gymnocarpium dryopteris - Rhytidiadelphus  loreus - Rhizomnium glabrescens - Plagiothecium  undulatum upland forests. The amabilis f i r - hemlock - cedar - Gymnocarpiurn -Rhizomnium upland forest community type i s r e s t r i c t e d to steep c o l l u v i a l slopes (45 - 80%) or more gradual slopes i n receiving positions at the toe of long steeper slopes (Figs. 44 - 47). The c h a r a c t e r i s t i c combination of species and stand c h a r a c t e r i s t i c s for t h i s community type are presented i n Table 5. Vegetation Species groups F, G and H (Fig. 8) contain the constant or dominant species of t h i s community type. Most of the pr e f e r e n t i a l and selective species are contained i n groups G and H. This forest type has the highest tree productivity of a l l the community types within the study area, with the possible exception of the a l l u v i a l forests (for Which there i s limited sampling). The closed tree canopy i s dominated by Abies  amabilis, Tsuga heterophylla and Thuja p l i c a t a . Mature i n d i -viduals of Picea sitchensis and Alnus rubra also occur but these are successional species seldom found regenerating. Chamaecyparis nootkatensis i s uncommon i n t h i s forest type and Pinus contorta i s absent. Average height of the main canopy ranges from 21 to 35 m (x = 24 m) and maximum canopy height to lO F i g . 4 4 . The a m a b i l i s f i r - hemlock - c e d a r - F i g . 4 5 . The a m a b i l i s f i r - hemlock - c e d a r -Gymnocarpium - Rhizomnium community Gymnocarp ium - Rh izomnium community t y p e . T h i s s t a n d i s r e l a t i v e l y young t y p e . T h i s s t a n d i s dom ina t ed by w e s t e r n and s p r u c e - d o m i n a t e d ; t r a n s e c t 3 . . hemlock and a m a b i l i s f i r ; t r a n s e c t 3 . o o F i g . 46. The Polystichum munitum associat-ion o f the' amabi 1 is f i r - hemlock -cedar - Gymnocarpium - Rhizomnium community type; releve 18. F i g . 47. The Polystichum munitum associ'at-ion of the amabil is f i r - hemlock -cedar - Gymnocarpium - Rhizomnium community type; t ransect 3. 101 Table 5. Vegetative features, stand characteristics and character species for the Abies amabilis - Tsuga heterophyla - Thuja plicata - Vaccinium spp. - Oplopanax horridus - Gymnocarpium dryopteris - Rhytidiadelphus  loreus - Rhizomnium glabrescens - Plagiothecium undulatum upland forest community type.  General vegetative features and stand characteristics - closed tree canopy (75 - 95% cover; x = 84%) - maximum height of tree canopy: 31 - 43 m (S! = 36) - average height of main canopy: 21 - 35 m (X = 24) - average stand dbh: 46 - 78 cm (x = 65) - number of stems per hectare: 124 - 569 (5J = 285) - basal area per hectare: 60 - 105 m2 (x = 82) - sparse to (usually) very wel developed shrub layer (20 - 95%; x = 56%) - sparse to wel developed herb layer (5 - 90% cover; X = 4%) - moderately to wel developed bryophyte layer (40 - 95% cover; X = 73%) Tree layers: character species* Abies amabilis Tsuga heterophyla  Thuja plicata  Picea sitchensis Shrub layers: character species Tsuga heterophyla  Abies amabilis  Vaccinium alaskaense  Menziesia ferruginea  Vaccinium parvifolium  Oplopanax horridus Herb Layer: character species Blechnum spicant  Cornus unalaschkensis  Rubus pedatus  Streptopus amplexifolius  Athyrium filix-femina  Gymnocarpium dryopteris  Streptopus roseus  Tiarella trifoliata  Coptis aspleniifolia  Thelypteris phegopteris  Viola glabella  Polystichum munitum  Huperzia selago Bryophyte and lichen layer: character species Rhytidiadelphus loreus  Rhizomnium glabrescens  Hylocomium splendens  Sphagnum girgensohni  Plagiothecium undulatum  Plagiochila asplenioides  Pellia neesiana  Poqonatum contortum  Conocephalum conicum • Stokesiella oregana CD2; preferential3 CD D (D) CD CD; preferential CD; preferential CD C CD; selective CD; preferential CD CD C ; preferential CD; selective CD; +_ preferential C ; preferential D; preferential D exclusive preferential D exclusive preferential CD CD CD CD CD; +_ preferential C ; preferential D; (^preferential) preferential exclusive exclusive *Species are listed in order of decreasing constancy. The lists include only species of highest constancy and dominance as wel as exclusive selective or preferential taxa of lower constancy and/or dominance. Refer to Appendix VI for complete vegetation tables. = constant species (constancy of >_ 80%) D = dominant species (average cover abundance value of >3) 3Refer to definitions of character taxa in text (page 29). 102 r a n g e s f r o m 31 t o 43 m (x = 36m). A l t h o u g h t r e e s a r e g e n e r a l l y t a l l e s t i n t h i s f o r e s t t y p e , t h e s t a n d s a r e r e l a -t i v e l y y o u n g , e s p e c i a l l y i n c o m p a r i s o n t o t h e z o n a l f o r e s t s and t h e bog w o o d l a n d s . M a i n c a n o p y s a m p l e t r e e s a v e r a g e 186 y e a r s i n age. A v e r a g e s t a n d d bh, number o f stems p e r h e c t a r e , and b a s a l a r e a p e r h e c t a r e d i f f e r l i t t l e f r o m t h e a l l u v i a l f o r e s t s ( T a b l e 5 ) . I n c o m p a r i s o n t o t h e c e d a r - c y p r e s s -h e m l o c k - V a c c i n i u m - C o r n u s - R h y t i d i a d e l p h u s z o n a l f o r e s t s , t h i s f o r e s t t y p e h a s c o n s i d e r a b l y l o w e r s t e m d e n s i t y , w i t h much l a r g e r a v e r a g e s t a n d d b h. B a s a l a r e a p e r h e c t a r e i s , h o w e v e r , o n l y s l i g h t l y g r e a t e r t h a n i n t h e z o n a l f o r e s t t y p e ( T a b l e s 3 and 5 ) . R e l a t i v e g r o w t h r a t e s o f e a c h o f t h e t r e e s p e c i e s w i t h i n t h e f o u r f o r e s t and w o o d l a n d c o m m u n i t i e s w i l l be c o m p a r e d i n s e c t i o n 4.4.4. S h r u b l a y e r s v a r y f r o m s p a r s e t o v e r y w e l l d e v e l o p e d , m a i n l y i n r e s p o n s e t o c a n o p y c l o s u r e . G e n e r a l l y t h e y a r e n o t a s w e l l d e v e l o p e d a s i n t h e z o n a l f o r e s t c o m m u n i t i e s . I n a d d i t i o n t o a b u n d a n t t r e e r e g e n e r a t i o n o f T s u g a h e t e r o p h y l l a and A b i e s a m a b i l i s ( p r e f e r e n t i a l ) , s h r u b l a y e r s c o n s i s t m a i n l y o f t h e c o n s t a n t d o m i n a n t s V a c c i n i u m a l a s k a e n s e ( p r e f e r e n t i a l ) , V. p a r v i f o l i u m , Menz i e s i a f e r r u g i n e a and O p l o p a n a x h o r r i d u s ( s e l e c t i v e ) . V a c c i n i u m o v a l i f o l i u m i s l e s s a b u n d a n t and T h u j a  p l i c a t a , P i c e a s i t c h e n s i s , Sambucus r a c e m o s a and G a u l t h e r i a  s h a l l o n a r e s p o r a d i c s h r u b s p e c i e s . H e r b l a y e r s a r e q u i t e v a r i a b l e b o t h i n d e g r e e o f d e v e l o p -ment and i n s p e c i e s c o m p o s i t i o n . U n d e r a c l o s e d c a n o p y and on s t e e p e r s l o p e s e x p e r i e n c i n g a c o n s t a n t downward movement o f 103 l i t t e r and d e b r i s , h e r b l a y e r s a r e s p a r s e w h e r e a s u n d e r a more o p e n c a n o p y and on more s t a b l e s l o p e s , h e r b l a y e r s c a n be w e l l d e v e l o p e d . C o n s t a n t d o m i n a n t s p e c i e s a r e B l e c h n u m s p i c a n t ( p r e f e r e n t i a l ) , C o r n u s u n a l a s c h k e n s i s , Rubus p e d a t u s , A t h y r i u m f i l i x - f e m i n a ( s e l e c t i v e ) , and G y m n o c a r p i u m d r y o p - t e r i s (+ p r e f e r e n t i a l ) . S t r e p t o p u s a m p l e x i f o l i u s ( p r e f e r e n t i a l ) and S_. r o s e u s a r e c o n s t a n t n o n - d o m i n a n t s p e c i e s . C h a r a c t e r s p e c i e s o f l o w e r c o n s t a n c y a r e C o p t i s  a s p l e n i i f o l i a ( o f t e n d o m i n a n t ) , T i a r e l l a t r i f o l i a t a ( p r e f e r e n t i a l ) , T h e l y p t e r i s p h e g o p t e r i s ( e x c l u s i v e ) , H u p e r z i a  s e l a g o ( p r e f e r e n t i a l ) , P o l y s t i c h u m m u n i t u m ( e x l u s i v e ) and V i o l a g l a b e l l a ( p r e f e r e n t i a l ) . The l a t t e r two s p e c i e s , e s p e c i a l l y P. m u n i t u m , a p p e a r t o c h a r a c t e r i z e a d i s t i n c t a s s o c i a t i o n ( F i g s . 46 and 47) on v e r y s t e e p c o l l u v i a l s l o p e s d e r i v e d f r o m m e t a m o r p h i c b e d r o c k ( r e l e v e s 18, 19, 3 2 ) . G. d r y o p t e r i s , T. t r i f o l i a t a and T. p h e g o p t e r i s , a l t h o u g h n o t r e s t r i c t e d t o t h e s e s i t e s , a l s o h e l p t o c h a r a c t e r i z e t h e P. muniturn a s s o c i a t i o n . O t h e r h e r b s p e c i e s o f l o w e r c o n s t a n c y and d o m i n a n c e i n t h i s c o m munity t y p e a r e L i s t e r a c o r d a t a , L. c a u r i n a and L y s i c h i t o n a m e r i c a n u m ( r e s t r i c t e d t o s e e p a g e c h a n n e l s ) . S p o r a d i c s p e c i e s i n c l u d e L y c o p o d i u m c l a v a t u m , Moneses  u n i f l o r a , G a l i u m b o r e a l e , A d i a n t u m p e d a t u m , C i r c a e a a l p i n a , C I i n t o n i a u n i f l o r a , C a r e x m e r t e n s i a n a , L u z u l a p a r v i f l o r a , Maianthemum d i l a t a t u m and T i a r e l l a , l a c i n i a t a . W i t h t h e e x c e p t i o n o f t h e f i r s t two s p e c i e s , t h e s e o c c u r o n l y i n t h e s a m p l e s o f t h e P. m u n i t u m a s s o c i a t i o n . 104 The b r y o p h y t e l a y e r o f t h e f o r e s t f l o o r v a r i e s f r o m m o d e r a t e l y t o v e r y w e l l d e v e l o p e d , a g a i n i n r e s p o n s e t o c r o w n c l o s u r e and d e g r e e o f s u r f a c e d i s t u r b a n c e . R h y t i d i a d e l p h u s  l o r e u s , R h i z o m n i u m g l a b r e s c e n s , H y l o c o m i u m s p l e n d e n s , Sphagnum  g i r g e n s o h n i i and P l a g i o t h e c i u m u n d u l a t u m (+ p r e f e r e n t i a l ) a r e a l l c o n s t a n t d o m i n a n t s p e c i e s . P l a g i o c h i l a a s p l e n i o i d e s ( p r e f e r e n t i a l ) , P e l l i a n e e s i a n a (+ p r e f e r e n t i a l ) , P o g o n a t u m  a l p i n u m (+ p r e f e r e n t i a l ) P. c o n t o r t u m ( p r e f e r e n t i a l ) , C o n o - c e p h a l u m . c o n i c u m ( e x c l u s i v e ) and S t o k e s i e l l a o r e g a n a ( e x c l u s i v e ) a r e c h a r a c t e r s p e c i e s o f l o w e r c o n s t a n c y a n d / o r d o m i n a n c e . S p o r a d i c b r y o p h y t e s on t h e f o r e s t f l o o r i n c l u d e D i c r a n u m  h o w e l l i i , H o o k e r i a l u c e n s , D i p l o p h y l l u m a l b i c a n s , P l a g i o - t h e c i u m l a e t u m , D i c r a n u m f u s c e s c e n s , A t r i c h u m u n d u l a t u m , S c a p a n i a b o l a n d e r i , B l e p h a r o s t o m a t r i c h o p h y H u m , B r a c h y t h e c i u m f r i g i d u m , D i c r a n e l l a h e t e r o m a l l a , D i c r a n u m  m a j u s , Hypnum c i r c i n a l e , L e p i d o z i a r e p t a n s , L e u c o l e p i s  menz i e s i i , L o p h o z i a s p p . , M i c r o l e p i d o z i a s e t a c e a and P o l y t r i c h u m formosum. The l i c h e n s , S t e r e o c a u l o n p a s c h a l e , P e l t i g e r a a p h t h o s a and C l a d o n i a b e l l i d i f l o r a , o c c u r r e d s p o r a d i c a l l y i n s a m p l e s o f t h i s c o m m u n i t y t y p e . Many o f t h e a b o v e c r y p t o g a m s a l s o o c c u r on r o t t e n wood and a s e p i p h y t e s , and most o f t h e e p i x y l e s and e p i p h y t e s l i s t e d i n t h e d e s c r i p t i o n o f t h e z o n a l f o r e s t c ommunity t y p e ( s e c t i o n 4.4.2.4) a l s o o c c u r i n t h e a m a b i l i s f i r - h e m l o c k -c e d a r - G y m n o c a r p i u m f o r e s t c o m m u n i t i e s . A l t h o u g h e p i p h y t i c c o m m u n i t i e s a r e much l e s s s t r i k i n g t h a n i n t h e z o n a l f o r e s t s , 105 numerous w i n d f a l l e n t r e e s on t h e s e s t e e p c o l l u v i a l s l o p e s p r o v i d e a b u n d a n t h a b i t a t f o r e p i x y l i c b r y o p h y t e s . S o i l s and l a n d f o r m s S o i l s show c o n s i d e r a b l e v a r i a b i l i t y ( F i g s . 48 and 4 9 ) . M o d e r a t e l y w e l l t o i m p e r f e c t l y d r a i n e d , f i n e loamy t o s a n d y , u s u a l l y s k e l e t a l m i n e r a l s o i l s (Humic and F e r r o - H u m i c P o d z o l s , R e g o s o l s ) o c c u r on c o l l u v i a l v e n e e r s and b l a n k e t s . L i t h i c F o l i s o l s (LFH d i r e c t l y o v e r r o c k ) , as w e l l as T e r r i c F o l i s o l s (>40 cm LFH o v e r m i n e r a l s o i l ) , o c c u r l e s s commonly. E x c e p t f o r two p l o t s l o c a t e d i n r e c e i v i n g p o s i t i o n s on t o e s l o p e s , a w a t e r t a b l e was a b s e n t f r o m t h e s a m p l e p i t s . S o i l s a r e u s u a l l y m o i s t t o wet and, c o n s i d e r i n g t h e h i g h r e g i o n a l r a i n -f a l l , a b u n d a n t l a t e r a l m i n e r a l s e e p a g e i s t o be e x p e c t e d on t h e s t e e p c o l l u v i a l o r b e d r o c k s l o p e s s u p p o r t i n g t h i s f o r e s t t y p e . Humus f o r m s a r e Humimors, Hemimors, Hemihumimors and H y d r o m o r s r a n g i n g i n d e p t h f r o m 10 t o 43 cm. M o s t o f t h e c o l l u v i a l s l o p e s s a m p l e d a p p e a r e d r e l a t i v e l y s t a b l e w i t h c o n s i d e r a b l e humus a c c u m u l a t i o n s . T h i n n e r humus l a y e r s w i t h e v i d e n c e o f m i n e r a l - o r g a n i c i n c o r p o r a t i o n ( A m phihumimors) w e r e d e s c r i b e d on more a c t i v e s l o p e s . Among t h e f i v e p r o f i l e s a n a l y z e d , pH v a r i e d f r o m 2.4 t o 3.0 (x = 2.7) f o r humus l a y e r s and 2.6 t o 3.1 f o r t h e r o o t i n g z o n e . T h e s e v a l u e s d i f f e r l i t t l e f r o m t h o s e o f t h e z o n a l f o r e s t s o i l s e x c e p t t h a t humus l a y e r s ( s u r p r i s i n g l y ) a p p e a r s l i g h t l y more a c i d i c i n t h e a m a b i l i s f i r - h e m l o c k - c e d a r -G y m n o c a r p i u m c o m m u n i t i e s . T h e s e s o i l s a p p e a r t o be s l i g h t l y 106 F i g . 48. A Typic F o l i s o l supporting an amabil is f i r - hemlock - cedar -Gymnocarpium - Rhizomnium upland fo res t community; releve" 9. F i g . 49. An Orth ic Ferro - Humic Podzol ( f o l i s o l i c phase) supporting an amabi l is f i r - hemlock - cedar - Gymnocarpium - Rhizomn- ium upland fo res t community; re leve 27. lobf) 10 l e s s a c i d i c t h a n t h o s e o f t h e a l l u v i a l f o r e s t s . More s a m p l e s a r e r e q u i r e d , h o w e v e r , t o p r o p e r l y c h a r a c t e r i z e t h e v a r i a t i o n i n pH v a l u e s . T h i s c o mmunity t y p e i s c o n s i d e r e d s u b m e s i c t o m e s i c and p e r m e s o t r o p h i c ( A p p e n d i x 1 ) . R e l a t i o n s h i p s among t h e r e l e v e s On t h e PCA and RA o r d i n a t i o n s , t h e a m a b i l i s f i r -h e m l o c k - c e d a r - G y m n o c a r p i u m f o r e s t c o m m u n i t i e s a r e w e l l d i s t i n g u i s h e d f r o m t h e z o n a l f o r e s t c o m m u n i t i e s ( F i g s . 6 and 7 ) . The a l l u v i a l f o r e s t s , h o w e v e r , a r e v e r y s i m i l a r f l o r i s -t i c a l l y t o t h e a m a b i l i s f i r - h e m l o c k - c e d a r - G y m n o c a r p i u m f o r e s t s and t h i s e x p l a i n s t h e i r p r o x i m i t y on t h e o r d i n a t i o n s . The t h r e e r e l e v e s r e p r e s e n t i n g t h e P o l y s t i c h u m muniturn a s s o c i a t i o n d e s c r i b e d a b o v e s t a n d o u t on t h e o r d i n a t i o n s . T h e s e r e l e v e s r e p r e s e n t t h e most v i g o r o u s s t a n d s s a m p l e d . A l l w e r e on s t e e p c o l l u v i a l s l o p e s (55 - 65%) o v e r m e t a m o r p h i c b e d r o c k . P_. m u n i turn, a b u n d a n t i n a l l t h r e e r e l e v e s , i s r e s t r i c t e d t o s u c h f r e e l y d r a i n e d b u t m o i s t and ( a p p a r e n t l y ) n u t r i e n t - r i c h h a b i t a t s i n t h e s t u d y a r e a . 4.4.3 Separate A n a l y s e s of Understory and O v e r s t o r y To e x a m i n e how w e l l s p e c i e s o f t h e u n d e r s t o r y v e r s u s t h o s e o f t h e o v e r s t o r y c h a r a c t e r i z e e a c h o f t h e c o m m u n i t y t y p e s , s e p a r a t e r e l e v e o r d i n a t i o n s w e re p e r f o r m e d u s i n g o n l y t h e s p e c i e s o f s p e c i f i c s t r a t a . The r e s u l t s o f t h e s e a n a l y s e s a r e p r e s e n t e d i n t h e f o r m o f r e l e v e s c o r e p r o f i l e s o f t h e f i r s t a x i s RA o r d i n a t i o n ( F i g s . 50 and 5 1 ) . F o u r s e p a r a t e a n a l y s e s o f t h e c o m p l e t e d a t a s e t (40 r e l e v e s ) w e re p e r f o r m e d 108 F i g . 5 0 . P r o f i l e s o f r e l e v e s c o r e s on t h e RA f i r s t a x i s o r d i n a t i o n by s t r a t u m o f a l l r e l e v e s . The community t y p e s a r e : 1) E r i c a c e o u s sh rub - T r i c h o p h o r u m - Sphagnum open b o g s ; 2) Empetrum - V a c c i n i u m u l i g i n o s u m - R h a c o m i t r i u m d r y o u t -c r o p s ; 3 ) ' P i n e - c y p r e s s - c e d a r - G a u l t h e r i a - F a u r i a bog w o o d l a n d s ; 4) C e d a r - c y p r e s s - hemlock - V a c c i n i u m -Cornus z o n a l u p l a n d f o r e s t s ; and 5) A m a b i l i s f i r - hemlock -c e d a r - Gymnocarp ium u p l a n d f o r e s t s t o g e t h e r w i t h t h e Ceda r - s p r u c e - hemlock - L y s i c h i t o n a l l u v i a l f o r e s t s ( 5 , 10 and 25) IOOI 50 o o CO c o < a)All Species b)Low Shrubs N c) Herbs 5 0 d)Bryophytes, Lichens and Seedlings |Y43 2 30I2 I3063200323 102 231 0 I I 210032 23 I I 3 I 2 4323012 I 3003200323I 0223 I 0II2 I0O3223 I I 3 I 8 0928744206179 83SI I 5 23 633 46 7005 967 54892 I 90928744206 I 79835 I I 5 2 3 63 3 4670059 67 54 89 2 Releve No. 109 F i g . 5 1 . P r o f i l e s o f r e l e v e s c o r e s on RA f i r s t a x i s o r d i n a t i o n by s t r a t u m o f f o r e s t and wood l and r e l e v e s . The community t y p e s a r e : 1) A m a b i l i s f i r - hemlock - c e d a r - Gymnocarp ium u p -l a n d f o r e s t s t o g e t h e r w i t h the Ceda r - s p r u c e - hemlock -L y s i c h i t o n a l l u v i a l f o r e s t s ( 5 , 1 0 and 2 5 ) ; 2) Ceda r -c y p r e s s - hemlock - V a c c i n i u m - Cornus z o n a l f o r e s t s and 3) P i n e - c y p r e s s - c e d a r - G a u l t h e r i a - F a u r i a bog w o o d -l a n d s 110 u s i n g a l l s p e c i e s ( F i g . 5 0 a ) , l o w s h r u b s ( F i g . 5 0 b ) , h e r b s ( F i g . 5 0 c ) , and b r y o p h y t e s a n d l i c h e n s g r o w i n g o n humus ( F i g . 50d) . S i n c e t h e o p e n b o g s c o n t a i n e d no t a l l s h r u b s ( e x c e p t s c a t t e r e d P. c o n t o r t a ) o r t r e e s , s e p a r a t e o r d i n a t i o n s u s i n g t h e s e l a y e r s w e r e n o t p e r f o r m e d on t h e e n t i r e d a t a s e t . F o u r a n a l y s e s o f t h e f o r e s t a n d w o o d l a n d r e l e v e s o n l y (27 r e l e v e s ) w e r e a l s o p e r f o r m e d ( F i g . 51) u s i n g a l l s p e c i e s ( F i g . 5 1 a ) , t r e e l a y e r s ( F i g . 51b) t a l l and l o w s h r u b s ( F i g . 51c) a n d h e r b s , b r y o p h y t e s and l i c h e n s ( F i g . 5 1 d ) . I n most o f t h e r e l e v e p r o f i l e s o f i n d i v i d u a l s t r a t a , t h e c o m m u n i t y t y p e s a r e a s r e a d i l y d i s t i n g u i s h a b l e (some o f them more s o ) a s i n t h e a n a l y s e s u s i n g a l l s p e c i e s . The t h r e e a l l u v i a l f o r e s t r e l e v e s a r e n o t c l e a r l y d i s t i n g u i s h e d f r o m t h e a m a b i l i s f i r - h e m l o c k - c e d a r - G y m n o c a r p i u m f o r e s t r e l e v e s o n any o f t h e p r o f i l e s . T h i s r e f l e c t s t h e f l o r i s t i c s i m i -l a r i t i e s b e t w e e n t h e s e two c o m m u n i t y t y p e s . A l t h o u g h a l l o f t h e c o m m u n i t y t y p e s a r e r e a d i l y r e c o g -n i z e d i n t h e f i e l d on t h e b a s i s o f t r e e s p e c i e s c o m p o s i t i o n a n d v i g o u r as w e l l a s o v e r a l l s t r u c t u r e o f t h e v e g e t a t i o n , t h e s e o r d i n a t i o n s o f s e p a r a t e s t r a t a i l l u s t r a t e t h e i m p o r t a n c e o f u n d e r s t o r y s p e c i e s c o m p o s i t i o n i n h e l p i n g t o c h a r a c t e r i z e e a c h o f t h e t y p e s . U n d e r s t o r y s p e c i e s ( i n c l u d i n g b r y o p h y t e s a n d l i c h e n s ) a r e t h u s u s e f u l i n d i c a t o r s o f t h e s o i l m o i s t u r e a n d n u t r i e n t r e g i m e s i n f l u e n c i n g t r e e s p e c i e s c o m p o s i t i o n and p r o d u c t i v i t y . T r e e s p e c i e s c o m p o s i t i o n o f t e n r e f l e c t s r e l a t i v e l y b r o a d e n v i r o n m e n t a l f a c t o r s a s w e l l a s c o n d i t i o n s a t t h e t i m e o f I l l e s t a b l i s h m e n t ( h i s t o r i c a l f a c t o r s ) . The u n d e r s t o r y , on t h e o t h e r h a n d , o f t e n r e f l e c t s t h e more s u b t l e , l o c a l i z e d and c u r r e n t e d a p h i c c o n d i t i o n s . T h u s , w i t h i n e a c h o f t h e b r o a d l y d e f i n e d c o m m u n i t y t y p e s , a s s o c i a t i o n s c a n o f t e n be r e c o g n i z e d t h a t r e f l e c t f i n e r d i s t i n c t i o n s i n e n v i r o n m e n t a l c h a r a c t e r -i s t i c s ( e . g . t h e P o l y s t i c h u m m u n i t u m a s s o c i a t i o n o f t h e a m a b i l i s f i r - h e m l o c k - c e d a r - G y m n o c a r p i u m community t y p e . We must a l s o a p p r e c i a t e t h e i n f l u e n c e t h a t t r e e c a n o p i e s h a v e on u n d e r s t o r y v e g e t a t i o n , h o w e v e r . Most u n d e r s t o r y s p e c i e s a r e i n f l u e n c e d t o some e x t e n t by n o n - e d a p h i c f a c t o r s s u c h as l i g h t , m i c r o c l i m a t e and n u t r i e n t c o n t e n t o f l i t t e r -f a l l , s t e m f l o w , and t h r o u g h f a l l , a l l o f w h i c h a r e i n t u r n i n f l u e n c e d by t h e s p e c i e s c o m p o s i t i o n and s t r u c t u r e o f t h e t r e e c a n o p y . 4.4.4 S i t e I n d i c e s and Growth C l a s s e s T a b l e 6 s u m m a r i z e s by community t y p e t h e m e n s u r a t i o n d a t a f r o m 149 m a i n c a n o p y s a m p l e t r e e s . S i t e i n d i c e s (100 y r s . ) w e r e d e r i v e d u s i n g h e i g h t / a g e c u r v e s o f t h e B.C. M i n i s t r y o f F o r e s t s ( H e g y i e t a l . 1 9 7 9 ) . E a c h s p e c i e s was t h e n c l a s s i f i e d a c c o r d i n g t o K r a j i n a ' s (1969) g r o w t h c l a s s e s . B e c a u s e t h e s a m p l e t r e e s a v e r a g e d 210 y e a r s i n age and s i t e i n d e x v a l u e s w e r e s o m e t i m e s b e l o w t h e l o w e r l i m i t o f t h e c u r v e s , t h e s e v a l u e s s h o u l d be c o n s i d e r e d as r o u g h c o m p a r i s o n s o n l y . As K l i n k a e t a l . (1979) p o i n t o u t , s i t e i n d e x v a l u e s d e r i v e d f r o m o l d g r o w t h f o r e s t s do n o t p o r t r a y t h e t r u e g r o w t h p o t e n t i a l on t h e s e s i t e s . T h e s e d a t a do i l l u s t r a t e , h o w e v e r , t h e r e l a t i v e l y h i g h J/ABLE 6. M e n s u r a t i o n d a t a f rom sample t r e e s o f the f o u r f o r e s t and w o o d l a n d community t y p e s . H e i g h t i s i n m e t e r s and dbh i n c e n t i m e t e r s . SI = s i t e i n d e x (100 y e a r s ) e s t i m a t e d f r om B .C. F o r e s t S e r v i c e * h e i g h t / a g e c u r v e s (Hegy i e t a l . 1 9 7 9 ) . GC = growth c l a s s a f t e r K r a j i n a ( 1 9 6 9 ) . P i n e - c y p r e s s - c e d a r - G a u l t h e n ' a Bog Woodlands T r ee S p e c i e s # o f A v e . Ave . A v e . S . I . T r e e s Age H t . dbh T. h e t e r o p h y l l a 4 149 6 .5 12 .7 3 T . p l i c a t a 5 282 11 .2 2 7 . 8 3 C. n o o t k a t e n s i s "6 236 1 1 . 0 2 2 . 7 3 P. c o n t o r t a 17 216 1Q.4 2 1 . 3 10" A . a m a b i l i s -F\ s i t c h e n s i s -T. m e r t e n s i a n a C e d a r - s p r u c e - h e m l o c k - a l d e r -L y s i c h i t o n A l l u v i a l F o r e s t s T r ee S p e c i e s # o f Ave . A v e . A v e . S.I T r e e s Age H t . dbh T. h e t e r o p h y l l a 6 226 2 6 . 4 4 3 . 2 18 T . p i i c a t a 6 172 2 2 . 9 4 6 . 2 17 C. n o o t k a t e n s i s •• - - - -P. c o n t o r t a - - - - -A. a m a b i l i s 2 264 2 5 . 3 3 7 . 5 17 P_. s i t c h e n s i s 4 213 2 5 . 3 5 2 . 6 17 T . m e r t e n s i a n a C e d a r - c y p r e s s - h e m l o c k - V a c c i n i urn -Cornus U p l a n d F o r e s t s G .C . # o f T r e e s A v e . Age A v e . H t . A v e . dbh S . I . G.C Vc 14 204 1 4 . 9 2 7 . 7 6 Vc Vc 18 230 1 6 . 9 3 4 . 5 6 Vc Vb 14 275 1 6 . 5 36 .1 6 Va IVb 8 248 16 .1 3 0 . 4 " 14 IVa -4 202 1 4 . 8 2 6 . 8 6 Vb A m a b i l i s f i r - h e m l o c k - c e d a r -Gymnocarpiurn U p l a n d F o r e s t s G .C . #of T r e e s A v e . Age A v e . H t . A v e . dbh S . I . G.C Vb 17 196 3 0 . 0 3 5 . 6 21 Va Vb 7" 220 2 6 . 6 5 5 . 8 17 Vb - 1 329 2 3 . 6 4 1 . 6 12? IVa' Vb 14 155 2 9 . 6 4 5 . 6 23 IVa Vb 2 125 3 4 . 2 4 4 . 2 30" IVa 113 g r o w t h r a t e s i n t h e a m a b i l i s f i r - h e m l o c k - c e d a r - Gymno- c a r p i u m f o r e s t s and a l l u v i a l f o r e s t s i n c o m p a r i s o n t o t h e z o n a l f o r e s t s and bog w o o d l a n d s . S i t e i n d i c e s , e v e n on t h e m o s t p r o d u c t i v e s i t e s a r e r e l a t i v e l y l o w , h o w e v e r , compared t o t h o s e r e p o r t e d by K r a j i n a (1969) f o r t h e most p r o d u c t i v e s i t e s i n t h e w e t t e r s u b z o n e o f t h e C o a s t a l W e s t e r n Hemlock z o n e . S i n c e y o u n g s e c o n d g r o w t h e x a m p l e s o f t h e community t y p e s a r e v e r y r a r e o r n o n - e x i s t e n t i n t h e s t u d y a r e a , t h e d e g r e e t o w h i c h t h e s e d a t a r e f l e c t t h e t r u e g r o w t h p o t e n t i a l o f t h e s e s i t e s i s n o t known. I n b o t h t h e p i n e - c y p r e s s - c e d a r bog w o o d l a n d s and t h e z o n a l c e d a r - c y p r e s s - h e m l o c k f o r e s t s , P i n u s c o n t o r t a h a s t h e h i g h e s t s i t e i n d e x v a l u e . T h e s e d a t a s u g g e s t t h a t P. c o n t o r t a i s t h e b e s t a d a p t e d t r e e s p e c i e s t o t h e wet s o i l s and s l o w r a t e s o f n u t r i e n t t u r n o v e r w h i c h c h a r a c t e r i z e t h e s e two c o m m u n i t y t y p e s . A l t h o u g h P. c o n t o r t a i s o f t e n d o m i n a n t i n bog w o o d l a n d s , b e c a u s e o f i t s s h a d e i n t o l e r a n c e i t i s u s a l l y p r e s e n t o n l y as s c a t t e r e d m a t u r e i n d i v i d u a l s i n t h e more c l o s e d z o n a l f o r e s t s . 4.5 D i s c u s s i o n 4.5.1 O r d i n a t i o n and C l a s s i f i c a t i o n A l t h o u g h o r d i n a t i o n and c l a s s i f i c a t i o n r e p r e s e n t two a l t e r n a t i v e a p p r o a c h e s t o t h e a n a l y s i s o f v e g e t a t i o n ( W h i t t a k e r 1978a and b) , t h e y a r e i n f a c t c o m p l e m e n t a r y . W h i l e a c l a s s i f i c a t i o n o f v e g e t a t i o n d a t a e m p h a z i s e s t h e q u a l i t a t i v e d i f f e r e n c e s b e t w e e n g e n e r a l i z e d u n i t s a b s t r a c t e d f r o m s e v e r a l s a m p l e p l o t s , o r d i n a t i o n e m p h a s i z e s t h e c o n t i n u a l 114 v a r i a t i o n among t h e s a m p l e p l o t s . M u e l l e r - D o m b o i s and E l l e n b e r g (1974) s u g g e s t t h a t c l a s s i f i c a t i o n p r o v i d e s t h e f i r s t o r d e r o f i n f o r m a t i o n w h i l e o r d i n a t i o n p r o v i d e s t h e f i n e s t r u c t u r e w i t h i n and among t h e c l a s s e s . I n t h i s s t u d y , t h e o r d i n a t i o n s w e r e u s e f u l i n o b j e c t i v e l y e v a l u a t i n g and r e f i n i n g t h e i n i t i a l f i e l d s t r a t i f i c a t i o n , and i n v i s u a l l y d i s p l a y i n g t h e r e l a t i o n s h i p s w i t h i n and among t h e c o m m u n i t y t y p e s . As t h e o r d i n a t i o n d i a g r a m s i l l u s t r a t e , t h e r e i s c o n s i d e r a b l e v a r i a b i l i t y among t h e s a m p l e s o f e a c h c o m m u n i t y t y p e , w i t h some s a m p l e s b e i n g t r a n s i t i o n a l b e t w e e n d i s t i n c t t y p e s . Had t h e s a m p l i n g method been s y s t e m a t i c ( s a m p l i n g a t r e g u l a r i n t e r v a l s a l o n g t h e t r a n s e c t s ) , r a t h e r t h a n s u b j e c t i v e , t h e o r d i n a t i o n s p r o b a b l y w o u l d have d i s p l a y e d e v e n more c o n t i n u o u s v a r i a t i o n b e t w e e n community t y p e s . I d e c i d e d , h o w e v e r , t h a t s u b j e c t i v e l y p l a c i n g s a m p l e s i n s e v e r a l e x a m p l e s o f e a c h e c o s y s t e m u n i t i d e n t i f i e d d u r i n g t h e i n i t i a l r e c o n n a i s s a n c e ( t h e B r a u n - B l a n q u e t method) w o u l d p r o v i d e a b e t t e r c h a r a c t e r i z a t i o n o f e a c h u n i t o v e r a l a r g e r g e o g r a p h i c a r e a . S y s t e m a t i c s a m p l i n g o f t h e same g e o g r a p h i c a r e a , a l t h o u g h more o b j e c t i v e , i s much more c o s t l y and t i m e c o n s u m i n g , and t h u s i m p r a c t i c a l when d e t a i l e d v e g e t a t i o n , s o i l , s i t e and m e n s u r a t i o n d a t a a r e c o l l e c t e d f o r e a c h r e l e v e . A p p l y i n g o r d i n a t i o n s t o t h e s e d a t a i s s t i l l a v a l u a b l e a i d t o t h e i n t e r p r e t a t i o n and c o m m u n i c a t i o n o f r e s u l t s h o w e v e r , p r o v i d i n g t h e l i m i t a t i o n s t h i s p l a c e s on t h e i n t e r p r e t a t i o n s a r e u n d e r s t o o d . 115 4.5.2 V e g e t a t i o n - Environment R e l a t i o n s h i p s I t i s o f t e n d i f f i c u l t t o c o r r e l a t e one o r two e n v i r o n -m e n t a l f a c t o r s w i t h s p e c i e s c o m p o s i t i o n and v i g o u r , b e c a u s e v e g e t a t i o n g e n e r a l l y r e f l e c t s s e v e r a l h a b i t a t f a c t o r s a c t i n g i n c o m b i n a t i o n , o f t e n w i t h one c o m p e n s a t i n g f o r a n o t h e r . The p o s i t i o n s o f r e l e v e s on t h e o r d i n a t i o n d i a g r a m s a r e t h u s b e s t e x p l a i n e d i n t e r m s o f a c o m p l e x g r a d i e n t i n v o l v i n g , a t t h e v e r y l e a s t , w a t e r and n u t r i e n t r e g i m e s and m i c r o c l i m a t e . A t one end o f t h e o r d i n a t i o n s l i e t h e open b o g s . T h e i r s o i l s a r e s a t u r a t e d t o t h e s u r f a c e and t h e y r e l y , t o a l a r g e e x t e n t , on p r e c i p i t a t i o n , r a t h e r t h a n l a t e r a l m i n e r a l s e e p a g e , f o r t h e i r n u t r i e n t s u p p l y ( i . e . , an o m b r o p h i l o u s r a t h e r t h a n r h e o p h i l o u s w a t e r r e g i m e ) . Due t o t h e e x t r e m e l y h i g h w a t e r a b s o r b i n g c a p a c i t y o f Sphagnum, i n c o m b i n a t i o n w i t h t h e g e n t l e s l o p e s o f many o f t h e b o g s , w a t e r movement t h r o u g h t h e o r g a n i c s o i l s i s v e r y s l o w ( R y c r o f t e t a l . 1975 a & b ) . The s l o w w a t e r movement c o n t r i b u t e s t o a l o w n u t r i e n t f l u x and o x y g e n d e p l e t i o n (Gorham 1 9 5 7 ; S p a r l i n g 1966; A r m s t r o n g and Boatman 196 7 ; C o l l i n s e t a l . 1978; M a r t i n and H o l d i n g 1 9 7 8 ) . The t e n d e n c y f o r Sphagnum t o a b s o r b c a t i o n s f r o m t h e s o i l s o l u t i o n and e x c h a n g e them f o r h y d r o g e n i o n s d e c r e a s e s pH and c o n t r i b u t e s t o low n u t r i e n t a v a i l a b i l i t y ( L u c u s and D a v i s 1 9 6 1 ; Moore and B e l l a m y 1 9 7 4 ) . U n d e r t h e s e c o n d i t i o n s m i c r o b i a l a c t i v i t y i s l i m i t e d ( M a r t i n and H o l d i n g 1 9 7 8 ) , s l o w i n g d e c o m p o s i t i o n r a t e s and n u t r i e n t t u r n o v e r . The wet o r g a n i c s o i l s become n e t a c c u m u l a t o r s o f o r g a n i c a l l y bound, and t h u s l a r g e l y u n a v a i l a b l e , n u t r i e n t s . 116 A l t h o u g h low s o i l t e m p e r a t u r e s a r e o f t e n a s s o c i a t e d w i t h bog s o i l s ( M a r t i n and H o l d i n g 1 9 7 8 ) , w i t h i n t h e s t u d y a r e a s o i l t e m p e r a t u r e i n t h e u p p e r 50 cm was u s u a l l y h i g h e r i n t h e o p e n bogs t h a n i n t h e a d j a c e n t s h a d e d f o r e s t s ( B a n n e r , u n p u b l i s h e d d a t a ) . The i n f l u e n c e o f b o t h b e d r o c k g e o l o g y and d e g r e e o f s l o p e on bog d e v e l o p m e n t i s s u g g e s t e d by t h e o c c u r r e n c e o f t h e s h a l l o w e s t bogs on s t e e p e r s l o p e s and o v e r m e t a m o r p h i c b e d r o c k . T h i s l i k e l y r e l a t e s t o h i g h e r c a t i o n l e v e l s a s s o c i a t e d w i t h t h e m e t a m o r p h i c b e d r o c k and a g r e a t e r i n f l u e n c e o f l a t e r a l m i n e r a l s e e p a g e i n t h e more s l o p i n g p o r t i o n s o f b l a n k e t b o g s . The a m a b i l i s f i r - h e m l o c k - c e d a r - G y m n o c a r p i u m -R h i z o m n i u m u p l a n d . f o r e s t s o c c u r a t t h e o p p o s i t e end o f t h e o r d i n a t i o n s . T h e s e h a b i t a t s , o f t e n on s t e e p s l o p e s , a r e m o d e r a t e l y w e l l t o i m p e r f e c t l y d r a i n e d w i t h a f r e q u e n t s u p p l y o f m i n e r a l s e e p a g e t h r o u g h t h e s o i l p r o f i l e c o n t r i b u t i n g t o h i g h e r n u t r i e n t i n p u t s and o x i d i z i n g c o n d i t i o n s . As a r e s u l t , d e c o m p o s i t i o n r a t e s and n u t r i e n t t u r n o v e r a r e e x p e c t e d t o be h i g h e r i n t h e s e h a b i t a t s . PH v a l u e s i n t h e s e f o r e s t s a r e s u r p r i s i n g l y l o w , and a r e c o m p a r a b l e t o t h o s e i n t h e open b o g s . O n l y a s m a l l number o f s a m p l e s were a n a l y z e d , h o w e v e r , and pH was d e t e r m i n e d on d r i e d s a m p l e s , w h i c h o f t e n y i e l d l o w e r v a l u e s t h a n f r e s h s a m p l e s ( F a r n h a m and F i n n e y 1 9 6 5 ; B a r t l e t t and James 1 9 8 0 ) . A l s o , t h e l a r g e r d i l u t i o n r a t i o s u s e d f o r o r g a n i c h o r i z o n s may have p r o d u c e d s l i g h t l y h i g h e r v a l u e s , t h u s d e c r e a s i n g t h e 117 d i f f e r e n c e s b e t w e e n t h e o r g a n i c and m i n e r a l s o i l s ( J a c k s o n 1 9 5 8 ) . The h i g h d e g r e e o f w e a t h e r i n g and l e a c h i n g and t h e h i g h o r g a n i c m a t t e r c o n t e n t o f t h e s e s o i l s may a l s o c o n t r i b u t e t o t h e low pH v a l u e s . PH d i d n o t d i f f e r s i g n i f i c a n t l y b e t w e e n t h e s o i l s o v e r i g n e o u s v s . m e t a m o r p h i c b e d r o c k . I n c l i m a t i c a l l y c o m p a r a b l e a r e a s o f S o u t h e a s t A l a s k a , s u r f a c e h o r i z o n s o f b o t h o r g a n i c and m i n e r a l s o i l s a r e g e n e r a l l y a c i d i c (Gas e t a l . 1966; D a c h n o w s k i - S t o k e s 1941) a l t h o u g h pH v a l u e s a r e g e n e r a l l y h i g h e r (3.5 - 5) t h a n t h o s e f o u n d i n t h i s s t u d y ( 2 . 5 - 3 . 5 ) . M i c r o c l i m a t i c d i f f e r e n c e s t h a t may i n f l u e n c e s p e c i e s c o m p o s i t i o n a l s o o c c u r b e t w e e n t h e o p e n bogs and u p l a n d f o r e s t s . The c l o s e d f o r e s t s a r e g e n e r a l l y c o o l e r and more h u m i d a t g r o u n d l e v e l w i t h s m a l l e r d a i l y t e m p e r a t u r e and h u m i d i t y f l u c t u a t i o n s t h a n t h e open bogs ( B a n n e r , u n p u b l i s h e d d a t a ) . A t t h e e x t r e m e end o f t h e o r d i n a t i o n a x e s l i e t h e most v i g o r o u s ( f o r t h e a r e a ) , s t e e p c o l l u v i a l s l o p e f o r e s t c o m m u n i t i e s on m e t a m o r p h i c b e d r o c k . T h e s e a r e c h a r a c t e r i z e d by a b u n d a n t P o l y s t i c h u m muniturn as w e l l as s e v e r a l o t h e r s p e c i e s s u g g e s t i n g h i g h n u t r i e n t s t a t u s ( s e e s e c t i o n 4 . 4 . 2 . 6 ) . T h i s i l l u s t r a t e s t h e c o n t r o l l i n g i n f l u e n c e o f b e d r o c k , l a n d -f o r m and t o p o g r a p h y on v e g e t a t i o n d e v e l o p m e n t . The i n f l u e n c e o f b e d r o c k c a n a l s o be s e e n i n t h e o r d i -n a t i o n s o f t h e bog w o o d l a n d and c e d a r - c y p r e s s - h e m l o c k u p l a n d f o r e s t r e l e v e s . R e l e v e s o c c u r r i n g c l o s e s t t o t h e most p r o d u c t i v e f o r e s t c o m m u n i t i e s a r e o f t e n t h o s e on m e t a m o r p h i c 118 b e d r o c k ( i . e . , r e l e v e s 15, 20, 2 1 , 3 1 ) . The p o s i t i o n o f t h e c e d a r - c y p r e s s - h e m l o c k -V a c c i n i u m - C o r n u s - R h y t i d i a d e l p h u s u p l a n d f o r e s t r e l e v e s on t h e o r d i n a t i o n s r e f l e c t s m o i s t u r e and n u t r i e n t c o n d i t i o n s i n t e r m e d i a t e b e t w e e n t h e bog w o o d l a n d s and t h e more p r o d u c t i v e u p l a n d f o r e s t c o m m u n i t i e s . T h i s c o m m u n i t y t y p e c h a r a c t e r i z e s m ost o f t h e f o r e s t e d m o u n t a i n s l o p e s w i t h i n t h e s t u d y a r e a . A l t h o u g h i t i s c o n s i d e r e d t h e c l i m a t i c c l i m a x o r z o n a l e c o -s y s t e m i n t h i s p o r t i o n o f t h e CCPH, t h e r e a r e p r o b l e m s i n a p p l y i n g t r a d i t i o n a l c o n c e p t s o f t h e c l i m a t i c c l i m a x i n t h i s z o n e . T h i s p o i n t w i l l be d i s c u s s e d i n C h a p t e r 6. U n d e r t h e i n f l u e n c e o f a p e r h u m i d , r a i n y c l i m a t e , f r e e d r a i n a g e c a n n o t be m a i n t a i n e d on t h e g e n t l e t o m o d e r a t e s l o p e s on w h i c h t h e z o n a l f o r e s t c o m m u n i t i e s most commonly o c c u r . The deep s u r f a c e o r g a n i c a c c u m u l a t i o n s and h i g h o r g a n i c m a t t e r c o n t e n t o f t h e m i n e r a l s o i l s c h a r a c t e r i z i n g t h e s e f o r e s t s m a i n t a i n wet s o i l c o n d i t i o n s t h r o u g h o u t t h e y e a r . N u t r i e n t a v a i l a b i l i t y i s p r o b a b l y low as a r e s u l t o f s l o w d e c o m p o s i t i o n r a t e s and s l o w n u t r i e n t r e l e a s e f r o m t h e s e s t a b l e , a c i d i c Mor humus f o r m s ( K l i n k a e t a l . 1 9 8 1 ) . A l s o , as o r g a n i c m a t t e r b u i l d s up a t t h e s o i l s u r f a c e , n u t r i e n t c a t i o n i n p u t s t h r o u g h m i n e r a l s e e p a g e may d e c l i n e and t h e u n d e r l y i n g m i n e r a l s u b s t r a t e becomes l e s s i m p o r t a n t as a s o u r c e o f p l a n t n u t r i e n t s . The r e s u l t i s a s c r u b b y f o r e s t w i t h f l o r i s t i c e l e m e n t s l i n k i n g i t w i t h b o t h t h e bog w o o d l a n d s and t h e more p r o d u c t i v e u p l a n d f o r e s t s . One o f t h e most i m p o r t a n t o v e r r i d i n g f a c t o r s i n f l u e n c i n g 119 v e g e t a t i o n development i n the blanket bog - upland f o r e s t complex w i t h i n the study area i s topography, p r i m a r i l y the degree of slope, through i t s c o n t r o l of drainage and water regimes. However, as i l l u s t r a t e d on the o r d i n a t i o n s , compen-s a t i n g f a c t o r s o f t e n outweigh the importance of s l o p e . The a l l u v i a l f o r e s t communities, f o r example, are maintained by a constant or i n t e r m i t t e n t flow of m i n e r a l seepage through f i n e t e x t u r e d s o i l s . Except where f l u v i a l landforms occur, f l a t or g e n t l y s l o p i n g t e r r a i n i s u s u a l l y covered i n open bog, bog woodland or scrub f o r e s t . However, open bogs and bog wood-lands are sometimes able to develop on c o n s i d e r a b l e slopes (up to + 40%), perhaps through expansion from l o c a l i z e d d e p r e s s i o n s or from adjacent f l a t t e r t e r r a i n . T h i s p o r t i o n of the study concentrated on the d e s c r i p t i o n of ecosystem u n i t s w i t h i n the study area and on t h e i r r e l a t i o n s h i p to one another i n the landscape. My d i s c u s s i o n of moisture and n u t r i e n t r e l a t i o n s h i p s i s thus based on s u b j e c t i v e e v a l u a t i o n of i n d i c a t o r s p e c i e s , s o i l morphology and landform c h a r a c t e r i s t i c s . U l t i m a t e l y , however, we need to q u a n t i f y these r e l a t i o n s h i p s with some f u n c t i o n a l s t u d i e s . Two such i n v e s t i g a t i o n s d e a l i n g with f o l i a r n u t r i e n t chemistry and seepage water chemistry i n s e l e c t e d community types were c a r r i e d out i n a s s o c i a t i o n with t h i s study (Banner, unpublished d a t a ) . The data are c u r r e n t l y being analyzed and the r e s u l t s w i l l be r e p o r t e d elsewhere. 120 4.5.3 Comparison W i t h O t h e r S t u d i e s i n B r i t i s h Columbia and S o u t h e a s t A l a s k a The b l a n k e t b o g - u p l a n d f o r e s t c o m p l e x d e s c r i b e d h e r e shows c l o s e r e l a t i o n s h i p s t o e c o s y s t e m s d e s c r i b e d i n S o u t h e a s t A l a s k a ( R i g g 1 9 2 5 ; D a c h n o w s k i - S t o k e s 1 9 4 1 ; Z a c h 1950; H e u s s e r 196 0 ; S t e p h e n s e t a l . 1970; N e i l a n d 1 9 7 1 ; V i e r e c k and D y r n e s s 1 9 8 0 ) . The most c o m p l e t e r e c e n t d e s c r i p t i o n o f bog and f o r e s t e c o s y s t e m s i n S o u t h e a s t A l a s k a i s t h a t o f N e i l a n d ( 1 9 7 1 ) . She d e a l s m a i n l y w i t h v e g e t a t i o n , a l t h o u g h g e n e r a l c h a r a c t e r i s t i c s o f p e a t p r o f i l e s and m i n e r a l s o i l s a r e a l s o p r e s e n t e d . H e r d e s c r i p t i o n s o f v e g e t a t i o n - e n v i r o n m e n t r e l a t i o n s h i p s a r e s i m i l a r i n most r e s p e c t s t o t h o s e d e s c r i b e d i n t h i s s t u d y . S e v e r a l common t a x a o f t h e P r i n c e R u p e r t f l o r a do n o t e x t e n d f a r i n t o S o u t h e a s t A l a s k a , h o w e v e r . T h u j a p l i c a t a , A b i e s  a m a b i l i s , G a u l t h e r i a s h a l l o n , C I i n t o n i a u n i f l o r a , D r o s e r a  a n g l i c a , L i n n a e a b o r e a l i s , R h y n c h o s p o r a a l b a and S i p h u l a  c e r a t i t e s , a l l r e l a t i v e l y a b u n d a n t i n t h e P r i n c e R u p e r t a r e a , d e c l i n e i n a b u n d a n c e n o r t h w a r d a l o n g t h e A l a s k a p a n h a n d l e . C o n v e r s e l y , P i c e a s i t c h e n s i s becomes a much more d o m i n a n t component o f t h e c l i m a x f o r e s t s i n S o u t h e a s t A l a s k a t h a n on t h e n o r t h c o a s t a l m a i n l a n d o f B r i t i s h C o l u m b i a . N e i l a n d (1971) r e l a t e s f o r e s t p r o d u c t i v i t y and s p e c i e s c o m p o s i t i o n t o a c o m b i n a t i o n o f f a c t o r s i n c l u d i n g s l o p e a n g l e and p o s i t i o n , t h e i n f l u e n c e o f bog d r a i n a g e w a t e r and p r o x i m i t y t o t h e o c e a n and l a r g e s t r e a m s . H i g h e s t f o r e s t p r o d u c t i v i t y i s a s s o c i a t e d w i t h m o i s t b u t n o t s t a g n a n t s o i l 121 c o n d i t i o n s , and t h e o c c u r r e n c e o f u n d e r s t o r y s p e c i e s c o n -s i d e r e d i n d i c a t i v e o f h i g h e r pH, h i g h e r n u t r i e n t a v a i l a b i l i t y , g r e a t e r n i t r i f i c a t i o n and h i g h e r o x y g e n l e v e l s . No c h e m i c a l d a t a t o s u b s t a n t i a t e t h e s e r e l a t i o n s h i p s a r e p r o v i d e d , h o w e v e r . N e i l a n d (1971) d e s c r i b e d s i m i l a r t r e n d s i n bog d e v e l o p -ment t o t h o s e o c c u r r i n g n e a r P r i n c e R u p e r t ; i . e . , f r o m t h e s m o o t h s u r f a c e d and f l o r i s t i c a l l y s i m p l e , h e r b - d o m i n a t e d , f l a t b o g s t o t h e c o m p l e x o f wet and d r y s u b c o m m u n i t i e s o f t h e b l a n k e t b o g s . She a l s o d e s c r i b e s t h e g r a d a t i o n f r o m r h e o -p h i l o u s t o o m b r o p h i l o u s w a t e r r e g i m e s a s s o c i a t e d w i t h s p e c i f i c bog e c o s y s t e m s ( N e i l a n d , h o w e v e r , u s e s t h e g e n e t i c t e r m s s o l i -g e n o u s and o m b r o g e n o u s ) . T h e r e a r e v e r y few p u b l i s h e d a c c o u n t s o f v e g e t a t i o n a n d s o i l s a l o n g t h e n o r t h c o a s t o f B r i t i s h C o l u m b i a . H e u s s e r (1960) p r o v i d e s g e n e r a l d e s c r i p t i o n s o f f o r e s t and bog v e g e -t a t i o n on t h e m a i n l a n d c o a s t as w e l l as on t h e Queen C h a r l o t t e I s l a n d s and C a l d e r a n d T a y l o r ( 1 9 6 8) p r o v i d e q u i t e d e t a i l e d f l o r i s t i c d e s c r i p t i o n s o f t h e m a j o r w e t l a n d and u p l a n d v a s c u l a r p l a n t c o m m u n i t i e s on t h e Queen C h a r l o t t e I s l a n d s . R e c e n t l y , L e w i s (1982) has p r o d u c e d an e c o l o g i c a l c l a s s i f i c a t i o n and map o f e c o s y s t e m / t e r r a i n u n i t s f o r T r e e F arm L i c e n c e #24 on t h e Queen C h a r l o t t e s . T h i s i n c l u d e s much o f M o r e s b y I s l a n d and a d j a c e n t s m a l l e r i s l a n d s . G e o g r a p h i c a l l y i s o l a t e d as t h e Queen C h a r l o t t e s a r e , w i t h u n i q u e g e o l o g i c and f l o r i s t i c c h a r a c t e r i s t i c s ( S u t h e r l a n d -B r own 1968; C a l d e r and T a y l o r 1 9 6 8 ) , t h e r e a r e n e v e r t h e l e s s many s i m i l a r i t i e s i n e c o s y s t e m c h a r a c t e r i s t i c s b e t w e e n t h e w e s t e r n and s o u t h e r n Queen C h a r l o t t e s and t h e n o r t h c o a s t a l m a i n l a n d . P o j a r and A n n a s (1980) i n c l u d e t h e w e s t c o a s t o f Graham I s l a n d , much o f M o r e s b y I s l a n d and K u n g h i t I s l a n d i n t h e C o a s t a l C e d a r s - P i n e - H emlock z o n e . Some o f t h e f o r e s t and s c r u b e c o s y s t e m s d e s c r i b e d by L e w i s (1982) on M o r e s b y I s l a n d a r e v e r y s i m i l a r i n s p e c i e s c o m p o s i t i o n and s o i l c h a r a c t e r i s t i c s t o t h o s e d e s c r i b e d i n t h i s s t u d y . M a j o r d i s t i n c t i o n s a r e t h e a b s e n c e o f A b i e s  a m a b i 1 i s and t h e more w i d e s p r e a d o c c u r r e n c e o f P i c e a s i t - c h e n s i s i n t h e p r o d u c t i v e f o r e s t e c o s y s t e m s on t h e C h a r l o t t e s . The c e d a r - c y p r e s s - h e m l o c k - V a c c i n i u m C o r n u s - R h y t i d i a - d e l p h u s u p l a n d f o r e s t and t h e p i n e - c y p r e s s - c e d a r bog w o o d l a n d community t y p e s have v e r y s i m i l a r c o u n t e r p a r t s on t h e Queen C h a r l o t t e s . However, s h r u b s a u c h as G a u l t h e r i a s h a l l o n and V a c c i n i u m s p p . and h e r b s s u c h as B l e c h n u m s p i c a n t have b e e n s e v e r e l y r e d u c e d t h r o u g h o v e r b r o w s i n g by i n t r o d u c e d d e e r ( P o j a r e t a l . 1980; L e w i s 1 9 8 2 ; P o j a r and B a n n e r 1 9 8 2 ) . T h e r e a r e some d i f f e r e n c e s i n b l a n k e t bog c h a r a c t e r i s t i c s b e t w e e n t h e Queen C h a r l o t t e s and t h e P r i n c e R u p e r t s t u d y a r e a . B l a n k e t bog d e v e l o p m e n t on much o f t h e w e s t c o a s t o f t h e C h a r l o t t e s i s s i m i l a r t o what I have o b s e r v e d on v e r y e x p o s e d o u t e r c o a s t a l l o c a t i o n s on t h e m a i n l a n d and o f f s h o r e i s l a n d s ( e . g . , C a l v e r t I s l a n d , M c C a u l e y I s l a n d ) a s w e l l as on t h e B r o o k s P e n i n s u l a on V a n c o u v e r I s l a n d . I n c o n t r a s t t o t h e r e l a t i v e l y deep b l a n k e t bogs t h a t p r e d o m i n a t e i n t h e s t u d y a r e a , t h e s e e x p o s e d bogs a r e v e r y s h a l l o w , o f t e n l e s s t h a n 50 123 cm o v e r b e d r o c k . I n s t e a d o f o r g a n i c s o i l s , g l e y e d m i n e r a l s o i l s p r e d o m i n a t e and Sphagnum i s r e l a t i v e l y uncommon. O t h e r m o s s e s s u c h as C a m p y l o p u s a t r o v i r e n s and R h a c o m i t r i u m l a n u - g i n o s u m and t h e l i c h e n S i p h u l a c e r a t i t e s p r e d o m i n a t e . S e v e r a l v a s c u l a r p l a n t s p e c i e s i n c l u d i n g D o d e c a t h e o n j e f f r e y i , E r i g e r o n p e r i g r i n u s , M i c r o s e r i s b o r e a l i s , P l a n t a g o m a c r o c a r p a and Geum c a l t h i f o l i u m a r e c h a r a c t e r i s t i c i n t h e s e s h a l l o w b l a n k e t b o g s . T h e s e s p e c i e s a r e uncommon i n t h e d e e p e r b l a n k e t bogs t h a t p r e d o m i n a t e i n t h e P r i n c e R u p e r t s t u d y a r e a . T h e y a r e r e s t r i c t e d m a i n l y t o t h e m o s t s l o p i n g and s h a l l o w p o r t i o n s o f b l a n k e t bogs ( e . g . , r e l e v e 3 0 ) . A r e l a t i v e l y m i n e r o t r o p h i c w a t e r r e g i m e l i k e l y p r e v a i l s i n t h e s h a l l o w b l a n k e t bogs and t h i s i s r e f l e c t e d i n t h e f l o r i s t i c d i f f e r e n c e s b e t w e e n them and t h e d e e p e r o m b r o t r o p h i c b l a n k e t b o g s . The d e e p e r bogs a p p e a r t o d e v e l o p on g e n t l e r t e r r a i n a n d u s u a l l y i n more p r o t e c t e d l o c a t i o n s a l o n g t h e c o a s t , and a r e t h u s p r o b a b l y l e s s i n f l u e n c e d by s e v e r e e x p o s u r e t o w i n d and w a t e r e r o s i o n t h a n a r e t h e s h a l l o w b l a n k e t b o g s . F r o m my own r e c e n t e x p e r i e n c e a l o n g t h e B r i t i s h C o l u m b i a c o a s t , e c o s y s t e m s s i m i l a r t o t h o s e d e s c r i b e d i n t h i s s t u d y a r e w i d e s p r e a d t h r o u g h o u t t h e CCPH z o n e . T h i s zone s p a n s more t h a n 5 d e g r e e s o f l a t i t u d e , h o w e v e r , and much v a r i a t i o n i n s p e c i e s c o m p o s i t i o n and e d a p h i c c h a r a c t e r i s t i c s c a n be e x p e c t e d . More n a r r o w l y d e f i n e d b i o g e o c l i m a t i c as w e l l as e c o s y s t e m a t i c u n i t s t h a t d e s c r i b e t h i s v a r i a b i l i t y w i l l h a ve t o be r e c o g n i z e d as a c l a s s i f i c a t i o n f o r t h e e n t i r e z one i s d e v e l o p e d . On n o r t h e r n V a n c o u v e r I s l a n d ( Q u a t s i n o Sound a r e a ) and 124 t h e a d j a c e n t m a i n l a n d , a W e t t e r M a r i t i m e N o r t h e r n s u b z o n e o f t h e C o a s t a l W e s t e r n Hemlock zone h a s been t e n t a t i v e l y d e f i n e d by K l i n k a e t a l . 1980. E d a p h i c and f l o r i s t i c c h a r a c t e r i s t i c s o f t h i s CWH s u b z o n e s u g g e s t t h a t i t r e p r e s e n t s t h e s o u t h e r n t r a n s i t i o n f r o m CCPH t o CWH. I n t h i s s u b z o n e , t h i c k o r g a n i c a c c u m u l a t i o n s (Humimors) o v e r m i n e r a l m a t e r i a l s a r e common on a l l s l o p e p o s i t i o n s and c o n t r i b u t e t o " m o i s t " c o n d i t i o n s t h r o u g h o u t t h e y e a r : " t h e s e m u c k - l i k e h o r i z o n s i n h i b i t down-w a r d p e r c o l a t i o n o f p r e c i p i t a t i o n a n d p r o v i d e many o f t h e s o i l s w i t h t e m p o r a r y o r p e r m a n e n t s e e p a g e c l o s e t o t h e g r o u n d s u r f a c e " ( K l i n k a e t a l . 1980, page 9 ) . N o r t h e r n m o s t V a n c o u v e r I s l a n d ( i . e . , n o r t h o f P o r t H a r d y - San J o s e f Bay) a p p e a r s t o b e l o n g t o t h e CCPH. T s u g a h e t e r o p h y l l a i s t h e p r e d o m i n a n t t r e e s p e c i e s and A b i e s a m a b i l i s and T h u j a p l i c a t a a r e common a s s o c i a t e s i n c l i m a t i c c l i m a x s t a n d s i n t h e p r o p o s e d W e t t e r M a r i t i m e N o r t h e r n CWH s u b z o n e . P i c e a s i t c h e n s i s and P i n u s c o n t o r t a a r e m i n o r s p e c i e s a t l o w e r e l e v a t i o n s (< 600 m) and C h a m a e c y p a r i s  n o o t k a t e n s i s and T s u g a m e r t e n s i a n a a r e more a b u n d a n t a t h i g h e r e l e v a t i o n s (>600m). C h a r a c t e r i s t i c u n d e r s t o r y s p e c i e s i n c l u d e G a u l t h e r i a s h a l l o n , V a c c i n i u m a l a s k a e n s e , V a c c i n i u m p a r v i f o - 1 i u m , V. o v a l i f o l i u m , Menz i e s i a f e r r u g i n e a , B l e c h n u m s p i c a n t , P o l y s t i c h u m m u n i t u m , C o p t i s a s p l e n i i f o l i a , L i s t e r a c a u r i n a , C a l a m a g r o s t i s n u t k a e n s i s , C a l t h a b i f l o r a (on p o o r l y d r a i n e d s i t e s ) , Sphagnum g i r g e n s o h n i i , M y l i a t a y l o r i and P l a g i o c h i l a  a s p l e n i o i d e s ( K l i n k a e t a l . 1 9 8 0 ) . A l l o f t h e s e s p e c i e s ( e x c e p t P. munitum) a r e common i n t h e CCPH zone f u r t h e r n o r t h 125 a l o n g t h e o u t e r c o a s t . I n t h e CCPH z o n e , h o w e v e r , t h e r e i s n o t t h e c l e a r d i s t i n c t i o n b e t w e e n submontane and montane v a r i a n t s d e s c r i b e d by K l i n k a e t a l . (1980) f o r t h e W e t t e r M a r i t i m e N o r t h e r n CWH s u b z o n e . I n s t e a d , montane and s u b a l p i n e e l e m e n t s commonly e x t e n d down t o s e a l e v e l i n t h e CCPH. A l s o , t h e r o l e o f T. p i i c a t a and C. n o o t k a t e n s i s i n c l i m a t i c c l i m a x s t a n d s e q u a l s o r s u r p a s s e s t h a t o f T. h e t e r o p h y l l a a t a l l e l e v a t i o n s , and f o r e s t p r o d u c t i v i t y d e c l i n e s c o n s i d e r a b l y i n t h e CCPH z o n e . 4.5.4 R e l a t i o n s h i p s W i t h O t h e r O c e a n i c Regions O c e a n i c b l a n k e t bog c o m p l e x e s have been d e s c r i b e d i n N e w f o u n d l a n d ( H e i k u r a i n e n 1968; Page 1 9 7 1 ; P o l l e t 1 9 7 2 ; P o l l e t and B r i d g e w a t e r 1 973; Rowe 1972; W e l l s 1 9 8 1 ; Z o l t a i 1 9 8 0 ) , S c a n d i n a v i a (Moore and B e l l a m y 1974) and t h e B r i t i s h I s l e s ( T a n s l e y 1 949; P e a r s a l l 1 9 5 6; R a t c l i f f e 1964; T a y l o r and S m i t h 19 7 2 ; Boatman and T o m l i n s o n 1 9 7 3 , 1977; Moore and B e l l a m y 1974; Boatman e t a l . 1 9 8 1 ) . Rowe (1972) s t a t e s t h a t "....many u p l a n d a r e a s and s l o p e s on t h e i s l a n d o f N e w f o u n d l a n d a r e b l a n k e t e d w i t h moss b o g s , a p o s s i b l e r e f l e c t i o n o f t h e h i g h m o i s t u r e s u r p l u s " . A c c o r d i n g t o W e l l s ( 1 9 8 1 ) , t h e c o o l summers, h i g h r a i n f a l l , h u m i d i t y and f r e q u e n c y o f f o g , i n c o m b i n a t i o n w i t h a c i d i c b e d r o c k and n u t r i e n t - p o o r s e e p a g e w a t e r s b r i n g a b o u t t h e e x t e n s i v e d e v e l o p m e n t o f b l a n k e t and s l o p e bogs i n e a s t e r n N e w f o u n d l a n d . B l a n k e t bogs c o v e r e n t i r e h i l l s and v a l l e y s o f t h e s o u t h e r n B u r i n ' and A v a l o n P e n i n s u l a s , w h i l e t h e s l o p e bogs a r e g e n e r a l l y l e s s e x t e n s i v e , u s u a l l y c o n f i n e d t o p o o r l y d r a i n e d 126 s l o p e s (5 - 15%) i n f o r e s t e d r e g i o n s . Where s l o p e bogs c o v e r l a r g e a r e a s t h e y a r e o f t e n d i f f i c u l t t o d i s t i n g u i s h f r o m t h e b l a n k e t b o g s . B o t h o f t h e s e bog t y p e s a r e u s u a l l y l e s s t h a n 2 m deep i n e a s t e r n N e w f o u n d l a n d ( W e l l s 1 9 8 1 ) . D e e p e r b l a n k e t bogs (+ 6 m) o c c u r a l o n g t h e n o r t h e a s t c o a s t ( P o l l e t and B r i d g e w a t e r 1 9 7 3 ) . I n c o n t r a s t t o t h e c o m p l e x o f o p e n b l a n k e t bogs and bog w o o d l a n d s c h a r a c t e r i z i n g t h e l e v e l t o g e n t l y s l o p i n g t e r r a i n a l o n g n o r t h c o a s t a l B r i t i s h C o l u m b i a , t h e s e l a r g e e x p a n s e s o f b l a n k e t / s l o p e bogs i n e a s t e r n N e w f o u n d l a n d a r e e s s e n t i a l l y t r e e l e s s . W e l l s (1981) d e s c r i b e s no f o r e s t e d bog o r f e n e c o s y s t e m s i n e a s t e r n N e w f o u n d l a n d and P o l l e t and B r i d g e w a t e r ( 1 9 7 3 ) s t a t e t h a t wooded p e a t l a n d s a r e s c a r c e i n c e n t r a l N e w f o u n d l a n d . However, b o t h Damman (1964) and H e i k u r a i n e n ( 1 9 6 8 ) d e s c r i b e wooded p e a t l a n d s t h a t a r e q u i t e w i d e s p r e a d i n some f o r e s t e d r e g i o n s o f N e w f o u n d l a n d , where t h e y o c c u r on l e v e l g r o u n d as w e l l as on s l o p e s . P i c e a m a r i a n a , L a r i x  l a r i c i n a and A b i e s b a l s a m e a a r e t h e m a j o r t r e e s p e c i e s i n t h e s e e c o s y s t e m s . Rowe (1972) i n c l u d e s a l l o f N e w f o u n d l a n d i n t h e B o r e a l F o r e s t R e g i o n o f C a n a d a . A l t h o u g h t h e r e a r e s e v e r a l s p e c i e s common t o t h e bog and f o r e s t e c o s y s t e m s o f b o t h t h e P a c i f i c and A t l a n t i c c o a s t s ( e . g . , Ledum g r o e n l a n d i c u m , T r i c h o p h o r u m  c e s p i t o s u m , many s p e c i e s o f Sphagnum, P l e u r o z ium s c h r e b e r i , R h a c o m i t r i u m l a n u g i n o s u m , M y l i a a n o m a l a ) , t h e r e a r e many b o r e a l f l o r i s t i c e l e m e n t s i n N e w f o u n d l a n d w h i c h a r e a b s e n t f r o m t h e t e m p e r a t e o c e a n i c P a c i f i c c o a s t ( e . g . , P i c e a m a r i a n a , 127 A b i e s b a l s a m e a , L a r i x l a r i c i n a , Chamaedaphne c a l y c u l a t a , K a l m i a a n g u s t i f o l i a and S a r r a c e n i a p u r p u r e a ) . Z o l t a i (1980) p r e s e n t s c l i m a t i c d a t a f o r e a c h o f t h e W e t l a n d R e g i o n s o f C a n a d a r e c e n t l y d e f i n e d by t h e N a t i o n a l W e t l a n d s W o r k i n g G r o u p . Mean d a i l y J a n u a r y and J u l y t e m p e r a t u r e s a r e +11°C and +3°C c o l d e r , r e s p e c t i v e l y , i n t h e A t l a n t i c O c e a n i c W e t l a n d R e g i o n ( e s s e n t i a l l y t h e s o u t h e r n A v a l o n and B u r i n P e n i n s u l a s i n N e w f o u n d l a n d ) t h a n i n t h e P a c i f i c O c e a n i c W e t l a n d R e g i o n ( t h e o u t e r n o r t h c o a s t o f B r i t i s h C o l u m b i a ) . I n a d d i t i o n , t h e A t l a n t i c O c e a n i c R e g i o n r e c e i v e s o n l y h a l f t h e a n n u a l p r e c i p i t a t i o n o f t h e P a c i f i c O c e a n i c R e g i o n and much more o f i t i s i n t h e f o r m o f snow. Page (1971) d e s c r i b e s s o i l f o r m i n g p r o c e s s e s and s o i l p r o p e r t i e s i n r e l a t i o n t o f o r e s t g r o w t h i n w e s t e r n and e a s t e r n N e w f o u n d l a n d t h a t a r e s i m i l a r i n many r e s p e c t s t o n o r t h c o a s t a l B r i t i s h C o l u m b i a . The w e t , c o o l c l i m a t e h a s r e s u l t e d i n t h e d e v e l o p m e n t o f h e a v i l y l e a c h e d P o d z o l s and B r u n i s o l s w i t h t h i c k o r g a n i c ( p e a t y ) a c c u m u l a t i o n s , and h i g h o r g a n i c m a t t e r i n c o r p o r a t i o n i n s u b s u r f a c e h o r i z o n s . G l e y s o l s and a c i d p e a t s a r e common as w e l l . Page (1971) s t a t e s t h a t s o i l d r a i n a g e and a e r a t i o n , o r g a n i c m a t t e r a c c u m u l a t i o n and m o i s t u r e r e t e n t i o n c h a r a c t e r i s t i c s a r e o f m a j o r s i g n i f i c a n c e t o f o r e s t g r o w t h o v e r a w i d e r a n g e o f s i t e s . He e m p h a s i z e s t h a t p h y s i c a l s o i l c o n d i t i o n s a r e l i m i t i n g g r o w t h and s u g g e s t s t h a t s i t e a m e l i o r a t i o n t e c h n i q u e s , s u c h as m i x i n g o f o r g a n i c and m i n e r a l m a t e r i a l s , y s h o u l d i m p r o v e m i c r o b i o l o g i c a l and c h e m i c a l a c t i v i t y t h e r e b y s p e e d i n g up t h e n u t r i e n t c y c l e and 128 i m p r o v i n g f o r e s t g r o w t h . As o u t l i n e d i n C h a p t e r 2, t h e B l a n k e t M i r e Complex z o n e o f Moore and B e l l a m y ( 1 9 7 4 ) i n c l u d e s t h e w e s t c o a s t o f Norway, t h e m o u n t a i n s o f S c o t l a n d , n o r t h e r n E n g l a n d , W a l e s and I r e l a n d a n d i s s i m i l a r i n many r e s p e c t s t o t h e b l a n k e t b o g - u p l a n d f o r e s t c o m p l e x i n t h e s t u d y a r e a . A c c o r d i n g t o a r e p r e s e n t a t i v e c l i m a t i c d i a g r a m f r o m Moore a n d B e l l a m y ( 1 9 7 4 ) , t h e a v e r a g e a n n u a l t e m p e r a t u r e and a v e r a g e a n n u a l p r e c i p i t a t i o n c h a r a c t e r i z i n g t h e E u r o p e a n B l a n k e t M i r e C o m p l e x a r e 10.2°C a n d 1439 mm r e s p e c t i v e l y . T h e r e a r e l e s s t h a n 10°C d i f f e r e n c e b e t w e e n t h e a v e r a g e t e m p e r a t u r e o f t h e w a r m e s t a nd c o l d e s t month, a n d e v e r y month e x c e p t M a r c h , A p r i l a n d May shows mean p r e c i p i t a t i o n g r e a t e r t h a n 100 mm. A l t h o u g h t h i s i s b o t h warmer and d r i e r t h a n P r i n c e R u p e r t ( F i g . 4 ) , i t s u g g e s t s a t e m p e r a t e o c e a n i c c l i m a t e c o m p a r a b l e t o t h a t o f n o r t h c o a s t a l B r i t i s h C o l u m b i a . I n a n e x c e l l e n t d e s c r i p t i o n o f t h e b o g e c o s y s t e m s o f S c o t l a n d , R a t c l i f f e (1964) s t r e s s e s t h e i m p o r t a n c e o f a n n u a l d i s t r i b u t i o n r a t h e r t h a n t o t a l a n n u a l p r e c i p i t a t i o n i n c o n -t r o l l i n g t h e d e v e l o p m e n t o f b l a n k e t b o g s . He s u g g e s t s t h a t t h e number o f "wet d a y s " ( a d a y w i t h >^  1 mm p p t . ) t o g e t h e r w i t h p o t e n t i a l e v a p o t r a n s p i r a t i o n ( e f f e c t i v e h u m i d i t y ) a r e t h e most u s e f u l c l i m a t i c p a r a m e t e r s f o r r e l a t i n g v e g e t a t i o n t o w e t n e s s o f c l i m a t e . The h u m i d c l i m a t e o f w e s t e r n I r e l a n d and S c o t l a n d r e s u l t s i n p e a t a c c u m u l a t i o n on s l o p e s o f l e s s t h a n 2 5 - 30% ( T a n s l e y 1949; R a t c l i f f e 1 9 6 4 ) . A l t h o u g h R a t c l i f f e d i s t i n g u i s h e s 129 b e t w e e n t h e r a i s e d and b l a n k e t bogs o f S c o t l a n d , he c o n s i d e r s t h e two t y p e s as end p o i n t s a l o n g a l i n e o f c o n t i n u a l v a r i a t i o n w i t h s e v e r a l i n t e r m e d i a t e s t h a t a r e d i f f i c u l t t o c l a s s i f y . He h y p o t h e s i z e s t h a t some e x t e n s i v e a r e a s o f b l a n k e t bog p r o b a b l y r e p r e s e n t t h e c o a l e s c e n c e o f many t o p o -g e n o u s bogs t h a t o r i g i n a l l y f o r m e d i n h o l l o w s i n t h e u n d e r -l y i n g g l a c i a l t i l l . A s i m i l a r o r i g i n was s u g g e s t e d f o r t h e P r i n c e R u p e r t b l a n k e t bogs ( s e e page 6 5 ) , a l t h o u g h b e d r o c k , r a t h e r t h a n g l a c i a l d e p o s i t s , u n d e r l i e most o f t h e s e b o g s . The g r a d u a l t r a n s i t i o n f r o m t h e deep o r g a n i c s o i l s o f b l a n k e t b ogs t o p e a t y , u p l a n d m i n e r a l s o i l s w h i c h o c c u r s i n t h e P r i n c e R u p e r t s t u d y a r e a i s a l s o d e s c r i b e d by R a t c l i f f e ( 1964) i n S c o t l a n d . The S c o t t i s h p e a t l a n d s have been d r a s t i c a l l y a f f e c t e d by human d i s t u r b a n c e s u c h as b u r n i n g , c l e a r i n g , p e a t c u t t i n g and d r a i n a g e . T h i s h i s t o r y o f d i s t u r b a n c e hampers any c o m p a r i s o n s o f v e g e t a t i o n and s o i l c h a r a c t e r i s t i c s w i t h t h e v i r g i n p e a t -l a n d s and f o r e s t s o f c o a s t a l B r i t i s h C o l u m b i a . F o r e x a m p l e , a l t h o u g h most o f t h e S c o t t i s h b l a n k e t bogs a r e t r e e l e s s , t h i s h a s p r o b a b l y r e s u l t e d f r o m r e l a t i v e l y r e c e n t f o r e s t c l e a r i n g ; " I t i s r e a s o n a b l e t o s u p p o s e t h a t , b e f o r e human i n f l u e n c e became s o w i d e s p r e a d , t h e e d g e s o f most b o g s , and e v e n t h e a c t u a l s u r f a c e s o f t h e l e s s wet b o g s , were a t l e a s t p a r t l y wooded up t o t h e a l t i t u d i n a l t r e e l i m i t " ( R a t c l i f f e 1 9 6 4 ) . I n c e r t a i n p a r t s o f S c o t l a n d some f o r e s t e d bogs o f P i n u s s y l v e s - t r i s do r e m a i n and R a t c l i f f e (1964) l i k e n s them t o t h e f o r e s t b ogs o f S c a n d i n a v i a . 130 The c o - d o m i n a n c e o f h e a t h e r s ( C a l l u n a and E r i c a ) t h r o u g h -o u t many o f t h e b l a n k e t b o g s i n S c o t l a n d and I r e l a n d h a s r e s u l t e d f r o m a d r y i n g - o u t o f t h e bog s u r f a c e t h r o u g h w i d e -s p r e a d b u r n i n g a n d d r a i n a g e ( R a t c i f f e 1964; T a n s l e y 1 9 4 9 ) . However, t h e l o w l a n d T r i c h o p h o r u m - E r i o p h o r u m b o g c o m m u n i t y , d e s c r i b e d b y R a t c l i f f e ( 1 9 6 4 ) a s one o f t h e most d i s t i n c t i v e o c e a n i c v e g e t a t i o n t y p e s c o n f i n e d t o w e s t e r n a r e a s o f h i g h r a i n f a l l and h u m i d i t y i n S c o t l a n d , shows c l o s e r e l a t i o n s h i p s t o t h e b l a n k e t b o g v e g e t a t i o n d e s c r i b e d i n t h i s s t u d y . The S c o t t i s h T r i c h o p h o r u m - E r i o p h o r u m b o g s a r e d o m i n a t e d b y Sphagnum s p p . , i n c l u d i n g most o f t h e s p e c i e s o c c u r r i n g i n t h e P r i n c e R u p e r t b o g s . S e v e r a l o t h e r f a m i l i a r c r y p t o g a m s o c c u r s u c h a s R h a c o m i t r i u m l a n u g i n o s u m , M y l i a a n o m a l a , M. t a y l o r i , P l e u r o z i u m s c h r e b e r i , C l a d o n i a i m p e x a and C. u n c i a l i s . The moss C a m p y l o p u s a t r o v i r e n s , t y p i c a l o f t h e s h a l l o w o u t e r c o a s t a l b l a n k e t b o g s i n B r i t i s h C o l u m b i a b u t r e l a t i v e l y uncommon i n t h e P r i n c e R u p e r t s t u d y a r e a , o c c u r s i n t h e w e s t e r n S c o t l a n d b l a n k e t b o g s . V a s c u l a r p l a n t s common t o b o g s o f b o t h S c o t l a n d a n d c o a s t a l B r i t i s h C o l u m b i a i n c l u d e T r i c h o p h o r u m c e s p i t o s u m , E r i o p h o r u m a n g u s t i f o l i u m , D r o s e r a  r o t u n d i f o l i a , D. a n g l i c a , R h y n c h o s p o r a a l b a , M e n y a n t h e s  t r i f o l i a t a , S c h e u c h z e r i a p a l u s t r i s , Empetrum n i g r u m and M y r i c a  g a l e . As i n t h e P r i n c e R u p e r t b o g s , M. g a l e i s l o c a l i n d i s t r i b u t i o n i n t h e S c o t t i s h b o g s . The s u r f a c e p a t t e r n o f d r y mounds a n d wet d e p r e s s i o n s a n d t h e a s s o c i a t e d d e v e l o p m e n t o f s e v e r a l d i s t i n c t m i c r o c o m m u n i t i e s t h r o u g h o u t t h e b o g s u r f a c e t h a t c h a r a c t e r i z e s t h e P r i n c e R u p e r t b l a n k e t b o g s i s a l s o 131 t y p i c a l o f b l a n k e t bogs i n t h e B r i t i s h I s l e s (Gorham 1957, R a t c l i f f e 1 9 6 4 ) . A l t h o u g h i t a p p e a r s t h a t h o l l o w s c a n e v e n t u a l l y g i v e r i s e t o hummocks a s was o b s e r v e d i n some P r i n c e R u p e r t bogs ( F i g . 20 - 2 3 ) , R a t c l i f f e (1964) s t a t e s t h a t u p w a r d g r o w t h i n hummock/hollow b l a n k e t bogs i s an i r r e g u l a r p r o c e s s w i t h some h o l l o w s p e r s i s t i n g i n t h e same p l a c e f o r v e r y l o n g p e r i o d s o f t i m e . He q u e s t i o n s t h e e v i d e n c e f o r a r e g e n e r a t i o n c o m p l e x o r c y c l i c a l pond/hummock s u c c e s s i o n ( W a t t 1947) i n S c o t t i s h b o g s . S e v e r a l a u t h o r s have d e s c r i b e d t h e p a t t e r n o f p o n d s and r i d g e s o r i e n t e d a c r o s s t h e s l o p e i n bogs o f t h e B r i t i s h I s l e s ( P e a r s a l l 1956; Boatman and A r m s t r o n g 1 9 6 8 ; Moore and B e l l a m y 1 974; Boatman e t a l . 1981; S m a r t 1982) as w e l l as i n b o r e a l p e a t l a n d s ( G l a s e r e t a l . 1 9 8 1 ) . The r e p o r t e d s u r f a c e p a t t e r n s a r e s i m i l a r i n some r e s p e c t s t o t h e p a t t e r n o f t e r r a c e d p o n d s and "dams" i n t h e P r i n c e R u p e r t b l a n k e t bogs ( F i g . 16 and 1 7 ) . P e a r s a l l (1956) s u g g e s t s t h a t s u c h s u r f a c e f e a t u r e s i n t h e b l a n k e t bogs o f S u t h e r l a n d , B r i t a i n c o u l d be r e l a t e d t o t h e f l u i d i t y o f s a t u r a t e d p e a t and a t e n d e n c y f o r i t t o f l o w down-s l o p e and s e t up p r e s s u r e r i d g e s o r i e n t e d a c r o s s t h e s l o p e . P o o l s may s u b s e q u e n t l y be f o r m e d i n " r i p s " i n t h e u p s l o p e t e n s i o n zone b e h i n d t h e p r e s s u r e r i d g e . P e a t d e s i c c a t i o n and s h r i n k i n g i n t h e t e n s i o n zone i s t h o u g h t t o p l a y a r o l e a s w e l l . P e a r s a l l ( 1956) c o m p a r e s a p e a t bog on a s l o p e t o "a f l u i d d r o p bounded by a s u r f a c e s k i n o f v e g e t a t i o n " . Moore and B e l l a m y (1974) d e s c r i b e a s i t u a t i o n i n M u c k l e M o s s , n o r t h e r n E n g l a n d where f o r e s t r y f e n c e s b u i l t a c r o s s t h e s l o p e 132 h a v e bowed and f r a c t u r e d as a r e s u l t o f d o w n s l o p e p e a t movement. They e s t i m a t e t h e r a t e o f movement a t 5 cm p e r y e a r . S p l i t p o o l s and w a t e r t r a c k s a r e t h o u g h t t o r e s u l t f r o m t h i s movement. As R a t c l i f f e (1964) p o i n t s o u t , t h e p o o l / r i d g e s y s t e m s i n many S c o t t i s h bogs a r e s u p e r f i c i a l l y s i m i l a r i n a e r i a l v i e w t o s o l i f l u c t i o n t e r r a c i n g and f u r r o w i n g i n l a v a f l o w s and g l a c i e r s . The o r i g i n o f t h e l o c a l t e r m " f l o w e " f o r many bogs i n t h e B r i t i s h I s l e s i s a p p a r e n t l y r e l a t e d t o t h e h i s t o r i c a l o c c u r r e n c e o f l a r g e p e a t s l i d e s (Moore and B e l l a m y 1974) . Boatman e t a l . (1981) r e l a t e s u r f a c e f e a t u r e s o f t h e S i l v e r F l o w e i n S c o t l a n d t o t h e t o p o g r a p h y o f t h e u n d e r l y i n g m i n e r a l m a t e r i a l r a t h e r t h a n t o f l u i d d y n a m i c s . G r o u p s o f p o o l s a p p e a r t o o v e r l i e b a s i n s w h i l e hummock/hollow p a t t e r n s o v e r l i e s l o p e s and t e r r a c e s . Boatman and A r m s t r o n g ( 1 968) s u g g e s t t h a t e l o n g a t e d p o o l s f o r m f r o m s m a l l e r p o o l s c o a l e s c i n g a l o n g t h e same c o n t o u r . G l a s e r e t a l . ( 1 9 8 1) p u t f o r t h a s i m i l a r h y p o t h e s i s f o r t h e f o r m a t i o n o f s t r i n g ( r i d g e ) / f l a r k ( p o o l ) p a t t e r n s i n t h e Red L a k e p e a t l a n d o f G l a c i a l L a k e A g a s s i z , M i n n e s o t a . The random f o r m a t i o n o f hummocks and h o l l o w s b r i n g s a b o u t t h e f o r m a t i o n o f p o o l s t h a t , t h r o u g h l a t e r a l f l o o d i n g , g i v e r i s e t o t h e e l o n g a t e p a t t e r n o f o f s t r i n g s and f l a r k s p a r a l l e l t o t h e c o n t o u r . As i n t h e P r i n c e R u p e r t b o g s , t h e c l o n a l g r o w t h o f T r i c h o p h o r u m c e s p i t o s u m i s i m p o r t a n t i n t h e f o r m a t i o n o f s t r i n g s o r "dams". I n t h e P r i n c e R u p e r t b o g s , e r o s i o n a l and s e d i m e n t a r y 133 p r o c e s s e s a p p e a r t o be a s s o c i a t e d w i t h t e r r a c e d p o n d s . M o s t p o n d s h a v e a s u b s t r a t e o f v e r y f i n e s e d i m e n t a r y p e a t w h i c h a p p e a r s t o be t r a n s p o r t e d by w a t e r f l o w b e t w e e n p o n d s . V e g e t a t i o n becomes e s t a b l i s h e d a t t h e d o w n s l o p e end o f t h e p o n d w h e r e s e d i m e n t s a r e more s t a b i l i z e d and t h i s a p p e a r s t o c o n t r i b u t e t o t h e f o r m a t i o n o f dams. A l t h o u g h t h e dams a r e somewhat e l o n g a t e d a n d o r i e n t e d a c r o s s t h e s l o p e , t h e p o o l s a r e c i r c u l a r o r t e a r - d r o p - s h a p e d r a t h e r t h a n e l o n g a t e a c r o s s t h e s l o p e . The t e r r a c e d s u r f a c e p a t t e r n s o f many o f t h e P r i n c e R u p e r t bogs s u g g e s t t h a t s l o w d o w n s l o p e movement may be i m p o r t a n t ( F i g . 1 7 ) , b u t t h e r e i s no d i r e c t e v i d e n c e f o r t h i s . 134 CHAPTER 5 HISTORICAL DEVELOPMENT OF A BOG WOODLAND: A PALYNOLOGICAL STUDY 5.1 O b j e c t i v e s o f the Study P a l y n o l o g i c a l s t u d i e s i n B r i t i s h C o l u m b i a , W a s h i n g t o n and A l a s k a h a v e d e a l t m o s t l y w i t h l a r g e s c a l e r e g i o n a l v e g e t a t i o n a n d c l i m a t i c c h a n g e s ( H a n s e n 1947a, 1955; H e u s s e r 1960; Mathewes a nd H e u s s e r 1 9 8 1 ) . However, s e v e r a l w o r k e r s h a v e d e m o n s t r a t e d t h a t p o l l e n p r o f i l e s r e f l e c t l o c a l s i t e f a c t o r s s u c h a s s o i l d e v e l o p m e n t , d r a i n a g e p a t t e r n s , s u c c e s s i o n , f i r e a n d s e a l e v e l c h a n g e s ( H a n s e n 1947b, Mathewes 1973; Mathewes an d Rouse 1975; Hebda a nd Rouse 1 9 7 9 ) , and p o l l e n a n a l y s i s h a s p r o v e n u s e f u l i n r e c o n s t r u c t i n g b o g d e v e l o p m e n t a n d v e g e t a t i o n h i s t o r y i m m e d i a t e l y a d j a c e n t t o a s a m p l i n g s i t e (Wade 1965; Hebda 1977, 1 9 7 9 ) . The o b j e c t i v e o f t h i s s t u d y was t o r e c o n s t r u c t t h e h i s t o r i c a l d e v e l o p m e n t o f a b o g w o o d l a n d c o m m u n i t y u s i n g p o l l e n a n a l y s i s a n d . p e a t s t r a t i g r a p h y . The p a t t e r n o f s u c c e s s i o n s u g g e s t e d b y t h i s p a l y n o l o g i c a l s t u d y i s t h e n r e l a t e d t o o t h e r s t u d i e s o f b o g s u c c e s s i o n and c l i m a t i c h i s t o r y i n c o a s t a l B r i t i s h C o l u m b i a and S o u t h e a s t A l a s k a . 5.2 L o c a t i o n and D e s c r i p t i o n o f the Study S i t e The b o g w o o d l a n d c h o s e n f o r t h e s t u d y i s l o c a t e d a t t h e e a s t end o f t h e Hayes M t n. s t u d y t r a n s e c t ( t r a n s e c t HM-4) a p p r o x i m a t e l y n i n e km s o u t h e a s t o f P r i n c e R u p e r t a l o n g H i g h w a y 16 ( F i g . 5 2 ) . The e l e v a t i o n o f t h e s i t e i s 30 m a . s . l . 135 F i g . 52. Locat ion of po l len p r o f i l e sampling s i t e near Mt. Hayes. The v e g e t a t i o n i m m e d i a t e l y s u r r o u n d i n g t h e s a m p l i n g s i t e ( r e p r e s e n t e d b y r e l e v e 21) c o n s i s t s o f s c a t t e r e d s c r u b b y P_. c o n t o r t a up t o 13 m t a l l d o m i n a t i n g t h e m a i n t r e e c a n o p y a n d s h r u b l a y e r s , t o g e t h e r w i t h C. n o o t k a t e n s i s and l e s s e r a m ounts o f T. p l i c a t a a n d T. h e t e r o p h y l l a ( F i g . 5 3 ) . D o m i n a n t u n d e r s t o r y s h r u b s a r e G a u l t h e r i a s h a l l o n , M e n z i e s i a f e r r u g i n e a , V a c c i n i u m p a r v i f o l i u m , a n d Ledum g r o e n -l a n d i c u m . K a l m i a m i c r o p h y l l a s s p . o c c i d e n t a l i s a n d M y r i c a  g a l e o c c u r s p o r a d i c a l l y . The d i v e r s e a n d w e l l d e v e l o p e d h e r b l a y e r i s d o m i n a t e d b y F a u r i a c r i s t a - g a l l i , V a c c i n i u m v i t i s - i d a e a , C o r n u s u n a l a s c h -k e n s i s , L i n n a e a b o r e a l i s , Maianthemum d i l a t a t u m , C a r e x p l u r i -f l o r a a n d s c a t t e r e d L y s i c h i t o n a m e r i c a n u m . P l e u r o z i u m s c h r e b e r i , H y l o c o m i u m s p l e n d e n s a nd Sphagnum  g i r g e n s o h n i i d o m i n a t e t h e w e l l d e v e l o p e d b r y o p h y t e l a y e r w h i c h i n c l u d e s r e l a t i v e l y m i n o r amounts o f P t i l i u m c r i s t a - c a s t r e n s i s , Sphagnum p a p i l l o s u m , S. r u b e l l u m , S_. f u s c u m , R h y t i d i a d e l p h u s l o r e u s and D i c r a n u m h o w e l l i i . S c a t t e r e d l i c h e n s , i n c l u d i n g C l a d o n i a g r a c i l i s a n d C l a d o n i a r a n g i f e r i n a , a l s o o c c u r w i t h t h e b r y o p h y t e s . S o i l s a t t h e s i t e a r e T y p i c and Humic M e s i s o l s ( C a n a d i a n S o i l S u r v e y C o m m i t t e e 1 9 7 8 ) , c o n s i s t i n g o f f i b r i c Sphagnum p e a t a t t h e s u r f a c e o v e r l y i n g more h u m i f i e d m e s i c and h u m i c h e r b a c e o u s a nd woody p e a t t o a d e p t h o f 200 cm ( F i g . 5 4 ) . F i n e s a n d g r a d i n g i n t o s i l t y l o a m u n d e r l i e s t h e p e a t d e p o s i t . T h i s m a t e r i a l p r o b a b l y o r i g i n a t e d f r o m a n e a r b y s t r e a m now l o c a t e d a p p r o x i m a t e l y 150 m t o t h e w e s t . A f o r e s t o f 137 F i g . 5 3 . The p i n e - c y p r e s s - c e d a r - G a u l t h e r i a - F a u r i a - P l e u r o z i u m -Sphagnum bog wood land community s u r r o u n d i n g the p o l l e n c o r e s a m p l i n g s i t e . J17S 138 F i g . 54 Mt. Hayes p o l l e n / s p o r e p r o f i l e MT. HAYES, PRINCE RUPERT, B C - A R B O R E A L P O L L E N (AP) NON ARBOREAL P O L L E N AND SPORES (NAP) • ABSOLUTE COUNT (AP • NAP) i i t i i t i t 10 S 10 20 SO IO 20 10 20 10 20 10 9 9 9 5 9 9 9 9 IO 10 20 30 40 SO 10 20 9 10 20 f ibr ic ( Sphagnum) pear herbaceous remains g^^ll mesic pear 1^ 1/"* | woody fragments 10 x Exaggeration of % mineral mater ia l T. p l i c a t a , P. sitchensis, T. heterophylla and A. rubra occurs on the a l l u v i a l materials bordering the stream and t h i s forest grades into the scrubby bog woodland surrounding the sampling s i t e . It i s also possible that an u p l i f t e d marine bench underlies the sampling s i t e . 5.3 Methods 5.3.1 F i e l d Methods A peat core 2 m in length was taken at the s i t e i n September, 1979 using a H i l l e r peat sampler. The core was separated into 10 cm samples that were sealed in p l a s t i c bags with a few drops of phenol (to prevent fungal contamination) and stored for three months before processing. A sample for carbon-14 dating was taken at the t r a n s i t i o n from organic to mineral sediment at the base of the core, using a piston sampler. This was frozen and l a t e r sent to the University of Waterloo, Ontario for C-14 determination. 5.3.2 Lab Methods Before being processed, each of the p r o f i l e samples was examined and the peat type described together with any recognizable s u b f o s s i l s . Ten cm3 subsamples were mixed with 6 - 8 exotic Lycopodium spore tablets, containing a known number of spores (Stockmarr 1971) and d i l u t e HCL to dissolve the tablet s . The solution was centrifuged for three minutes and the volume of the sample measured so that absolute pollen concentrations could be determined. The subsamples were then treated with 10% KOH followed by acetolysis (Faegri and Iverson 197 5). Samples containing mineral sediments were 140 t r e a t e d w i t h HF and h e a v y l i q u i d s e p a r a t i o n Z n B ^ was a l s o u s e d f o r some o f t h e l o w e r , m i n e r a l - r i c h s a m p l e s . The r e m a i n i n g p o l l e n s u s p e n s i o n s were b l e a c h e d , s t a i n e d w i t h s a f r a n i n and m o u n t e d o n s i l i c o n o i l s l i d e s . B e t w e e n 300 and 600 p o l l e n g r a i n s a n d s p o r e s w e re c o u n t e d f o r e a c h s a m p l e and p e r c e n t a g e s f o r e a c h t a x o n w e re c a l c u l a t e d and p l o t t e d ( F i g . 5 4 ) . A r b o r e a l p o l l e n (AP) p e r c e n t a g e s a r e p l o t t e d a s p a r t o f a s e p a r a t e sum w h i l e n o n - a r b o r e a l (NAP) p e r c e n t a g e s a r e p l o t t e d a s p a r t o f t h e t o t a l sum (AP + N A P ) . T h i s was done t o p r e v e n t p o l l e n - p r o l i f i c NAP t a x a f r o m p r o d u c i n g a b e r a t i o n s i n t h e t r e e s p e c i e s p e r c e n t a g e s . E x o t i c L y c o p o d i u m s p o r e s w e re c o u n t e d a l o n g w i t h t h e n a t i v e g r a i n s a n d s p o r e s f o r t h e c a l c u l a t i o n o f a b s o l u t e p o l l e n c o n c e n t r a t i o n s u s i n g t h e f o l l o w i n g f o r m u l a ( D a v i s e t a l . 1 9 7 7 ) : number o f _ n a t i v e g r a i n s c o u n t e d e x o t i c s a d d e d 3 — X 3 g r a i n s / c m e x o t i c s p o r e s c o u n t e d cm o f s e d i m e n t P o l l e n and s p o r e s w e re i d e n t i f i e d u s i n g a modern p o l l e n r e f e r e n c e c o l l e c t i o n t o g e t h e r w i t h a k e y t o t h e common p o l l e n a n d s p o r e t y p e s o f t h e P a c i f i c N o r t h w e s t p r e p a r e d b y R . J . Hebda o f t h e B r i t i s h C o l u m b i a P r o v i n c i a l Museum and G.E. Rouse a n d M. N o r t h , b o t h o f t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a ( u n p u b l i s h e d ) . 5 . 3 . 3 L i m i t a t i o n s o f the Methodology P o l l e n p r o f i l e a n a l y s i s T h e r e a r e l i m i t a t i o n s t o u s i n g p o l l e n d i a g r a m s f o r r e c o n -s t r u c t i n g v e g e t a t i o n h i s t o r i e s . T h e s e a r e d i s c u s s e d i n d e t a i l b y a n d D a v i s ( 1 9 6 3 ) a nd F a e g r i a n d I v e r s e n ( 1 9 75) a nd w i l l be 140A b r i e f l y s u m m a r i z e d h e r e . An a w a r e n e s s o f t h e s e l i m i t a t i o n s i s c r i t i c a l t o t h e r e a l i s t i c i n t e r p r e t a t i o n o f p o l l e n p r o f i l e s . B e c a u s e p l a n t s d i f f e r c o n s i d e r a b l y i n t h e i r p o l l e n p r o d u c t i o n and d i s p e r s a l m e c h a n i s m s , t h e r e may n o t a l w a y s be a d i r e c t r e l a t i o n s h i p b e t w e e n r e l a t i v e c o n c e n t r a t i o n s o f p o l l e n a n d r e l a t i v e a b u n d a n c e s o f t h e s e t a x a i n t h e s u r r o u n d i n g v e g e t a t i o n a t t h e t i m e o f d e p o s i t i o n . F o r e x a m p l e p i n e i s commonly o v e r - r e p r e s e n t e d i n p o l l e n p r o f i l e s w h e r e a s s p r u c e i s g e n e r a l l y u n d e r - r e p r e s e n t e d . T h i s r e f l e c t s d i f f e r e n c e s i n t h e r e l a t i v e amounts o f p o l l e n p r o d u c e d b y t h e s e t a x a . Some p l a n t s may n o t f l o w e r f o r p r o l o n g e d p e r i o d s and t h u s w i l l n o t b e r e p r e s e n t e d a t a l l i n t h e p o l l e n p r o f i l e . A r b o r e a l t a x a ( t r e e s ) p r o d u c e v e r y l a r g e amounts o f p o l l e n t h a t i s w i n d d i s p e r s e d o v e r l o n g d i s t a n c e s . P r o f i l e s o f t h e s e t a x a must be i n t e r p r e t e d d i f f e r e n t l y t h a n t h o s e o f n o n - a r b o r e a l , o f t e n i n s e c t - p o l l i n a t e d t a x a t h a t p r o d u c e s m a l l amounts o f l o c a l l y d i s p e r s e d p o l l e n . An u n d e r s t a n d i n g o f t h e s e f a c t o r s h e l p s t o s e p a r a t e t h e i n f l u e n c e o f l o c a l v e r s u s r e g i o n a l v e g e t a t i o n on t h e p o l l e n p r o f i l e . B a c k g r o u n d l e v e l s o f a r b o r e a l p o l l e n w i l l p e r s i s t i n t h e p r o f i l e e v e n i f t h e r e a r e no i n d i v i d u a l s c l o s e t o t h e s a m p l i n g s i t e . Low c o n c e n t r a -t i o n s o f p o l l e n f r o m n o n - a r b o r e a l t a x a , h o w e v e r , s u g g e s t t h a t t h e y w e r e p r e s e n t v e r y n e a r t h e s a m p l i n g s i t e . P o l l e n p r o f i l e s o f i n d i v i d u a l t a x a must n o t be i n t e r -p r e t e d i n d e p e n d e n t l y b u t r a t h e r i n r e l a t i o n t o t h e e n t i r e p r o f i l e . D e c l i n i n g p o l l e n c o n c e n t r a t i o n s o f s p e c i e s A may be a n a r t i f a c t o f a n i n c r e a s e i n s p e c i e s B r a t h e r t h a n a 140B r e f l e c t i o n o f r e a l d e c r e a s e s i n s p e c i e s A. N a r r o w z o n e s o f e x t r e m e l y h i g h c o n c e n t r a t i o n s o f p o l l e n may r e s u l t f r o m l o c a l i z e d d e p o s i t i o n e v e n t s , s u c h a s e n t i r e c a t k i n s f a l l i n g o n t o t h e p e a t o r s e d i m e n t s u r f a c e . D i f f e r e n t i a l p r e s e r v a t i o n o f p o l l e n a n d s p o r e s i s y e t a n o t h e r c o m p l i c a t i n g f a c t o r . Some t a x a w i l l be u n d e r - r e p r e -s e n t e d s i m p l y b e c a u s e t h e i r p o l l e n d o e s n o t p r e s e r v e w e l l . C h a n g e s i n r a t e s o f p e a t o r s e d i m e n t a c c u m u l a t i o n w i l l c l o u d t h e i n t e r p r e t a t i o n o f a b s o l u t e p o l l e n c o n c e n t r a t i o n s . P o l l e n d i a g r a m s a r e g e n e r a l l y i n t e r p r e t e d u s i n g a k n o w l -edge o f p r e s e n t - d a y e c o l o g i c a l r e l a t i o n s h i p s . The r e c o r d o f v e g e t a t i o n i s one a s p e c t o f t h e i n t e r p r e t a t i o n . A s e c o n d , p e r h a p s more i m p o r t a n t a s p e c t , i n v o l v e s r e l a t i n g t h e c h a n g e s i n v e g e t a t i o n t o c h a n g i n g e n v i r o n m e n t s . We assume ( p e r h a p s i n c o r r e c t l y ) t h a t s i m i l a r a u t e c o l o g i c a l and s y n e c o l o g i c a l r e l a t i o n s h i p s i n o p e r a t i o n t o d a y a p p l i e d t h o u s a n d s o f y e a r s a g o . B e c a u s e o f t h e s e many c o m p l i c a t i n g f a c t o r s , p a l y n o l o g y d o e s n o t y i e l d s t r a i g h t f o r w a r d , e a s i l y i n t e r p r e t a b l e r e s u l t s . I t n e v e r t h e l e s s i s a u s e f u l t e c h n i q u e t h a t , p r o v i d i n g c a r e f u l c o n s i d e r a t i o n i s g i v e n t o s u c h f a c t o r s a s d i f f e r e n t i a l p o l l e n p r o d u c t i o n , d i s p e r s a l and p r e s e r v a t i o n , c a n p r o v i d e a v a l u a b l e i n s i g h t i n t o t h e p a s t . P r o b l e m s a s s o c i a t e d w i t h t h e i n t e r p r e t a t i o n o f s p e c i f i c p o l l e n z o n e s o r t a x a w i l l be d i s c u s s e d i n s e c t i o n 5 .4 a n d 5 . 5 . R a d i o c a r b o n d a t i n g t e c h n i q u e s I n t h e t e c h n i q u e o f r a d i o c a r b o n d a t i n g , t h e r a d i o a c t i v i t y 140C o f c a r b o n - 1 4 i n t h e o r g a n i c s a m p l e i s m e a s u r e d and c o m p a r e d a g a i n s t a modern s t a n d a r d . The age o f t h e s a m p l e i s r e l a t e d t o t h e d i f f e r e n c e b e t w e e n t h e r a t i o s o f c a r b o n - 1 2 t o c a r b o n - 1 4 i n t h e s a m p l e and i n t h e a t m o s p h e r e . T h i s t e c h n i q u e i s b a s e d on t h e a s s u m p t i o n t h a t t h e r a t i o o f t h e s e two c a r b o n i s o t o p e s i n t h e a t m o s p h e r e h a s r e m a i n e d t h e same s i n c e t h e c a r b o n - 1 4 was a s s i m i l a t e d i n t o t h e l i v i n g t i s s u e o f t h e s a m p l e . How-e v e r , t h i s r a t i o h a s a p p a r e n t l y f l u c t u a t e d i n t h e p a s t and t h i s h a s r e s u l t e d i n d e v i a t i o n s o f r a d i o c a r b o n d a t e s f r o m t h e a c t u a l age o f s a m p l e s ( M i c h e l s 1 9 7 3 ) . A c o r r e c t i o n f o r t h i s d e v i a t i o n c a n be c a l c u l a t e d f r o m a c o r r e l a t i o n c u r v e o f C-14 age a g a i n s t known age o f b r i s t l e c o n e p i n e and S e q u o i a t r e e r i n g s a m p l e s ( M i c h e l s 1 9 7 3 ) . D e v i a t i o n s o f C-14 age e s t i m a t e s r a n g e f r o m -7 50 y e a r s (- i n d i c a t e s an u n d e r e s t i m a t i o n o f age) f o r s a m p l e s a p p r o x i m a t e l y 6000 y e a r s o l d t o +50 y e a r s ( o v e r -e s t i m a t i o n o f age) f o r v e r y r e c e n t s a m p l e s ( 2 0 0 - 4 0 0 y e a r s ) . A c c o r d i n g t o t h i s c a l i b r a t i o n c u r v e , t h e d a t e o f 2310 + 60 y e a r s f o r t h e O r g ano F o l i s o l on page 89 s h o u l d be f a i r l y a c c u r a t e . The d a t e o f 8710 + 210 y e a r s f r o m t h e b o t t o m o f t h e p o l l e n p r o f i l e , h o w e v e r , may u n d e r e s t i m a t e t h e a c t u a l age b y l e s s t h a n 700 y e a r s ( t h e d e v i a t i o n a p p a r e n t l y d e c r e a s e s a g a i n p a s t + 7000 y e a r s ; G.E. R o use, p e r s o n a l c o m m u n i c a t i o n ) . 5.4 R e s u l t s 5.4.1 P o l l e n and Peat S t r a t i g r a p h y The p e a t s t r a t i g r a p h y and p o l l e n f r e q u e n c i e s a r e p r e s e n t e d i n F i g . 54. F o u r m a j o r z o n e s h a v e b e e n i d e n t i f i e d i n t h e p r o f i l e . 141 Zone HM-I (200 - 190 cm; s a m p l e 20) The s e d i m e n t o f t h i s z one c o n s i s t s o f a g r i t t y ( f i n e s a n d y ) , b l a c k , h u m i c o r g a n o - m i n e r a l m a t e r i a l . B e l o w 200 cm t h i s g r a d e s i n t o g r a y i s h b r o w n , s i l t y l o a m . P o l l e n and s p o r e s w e re n o t c o u n t e d i n t h i s b a s a l m i n e r a l s e d i m e n t b e l o w 200 cm. A c a r b o n - 1 4 s a m p l e f r o m t h i s z o n e was d a t e d a t 8710 + 210 B.P. (WAT BAN2). P i n u s c o n t o r t a , A l n u s , and m o n o l e t e P o l y p o d i a c e a e ( f e r n s ) d o m i n a t e Zone I . T h e r e a r e m i n o r amounts o f C u p r e s s a c e a e ( T h u j a / C h a m a e c y p a r i s t y p e ) , T s u g a h e t e r o p h y l l a , P i c e a , and t r a c e s o f O p l o p a n a x h o r r i d u s , R o s a c e a e , and L y s i c h i t o n  a m e r i c a n u m . Zone HM-II (190 - 150 cm; s a m p l e s 19 - 16) T h i s i s t h e b e s t d e f i n e d zone i n t h e p r o f i l e , and i s c h a r a c t e r i z e d by d a r k , amorphous, h u m i c p e a t w i t h t r a c e s o f f i n e s a n d i n t h e l o w e r 20 cm. Woody and h e r b a c e o u s r e m a i n s a r e common t h r o u g h o u t . Zone I shows peak p o l l e n f r e q u e n c i e s o f P i c e a , A l n u s and L. a m e r i c a n u m t o g e t h e r w i t h h i g h b u t d e c l i n i n g s p o r e f r e q u e n c i e s o f m o n o l e t e f e r n s . The p e r c e n t a g e s o f T. h e t e r o p h y l l a and E r i c a c e a e p o l l e n b e g i n t o i n c r e a s e i n t h i s z o n e . B o t h P_. c o n t o r t a and Thu j a / C h a m a e c y p a r i s t y p e r e a c h t h e i r l o w e s t f r e q u e n c i e s i n Zone I I . The low f r e q u e n c i e s o f C u p r e s s a c e a e i n t h e l o w e r z o n e s may p r e s e n t p r o b l e m s i n i n t e r p r e t a t i o n s b e c a u s e b o t h r e d c e d a r and c y p r e s s p o l l e n i s b e l i e v e d t o p r e s e r v e p o o r l y , and f r e q u e n c i e s a r e c h a r a c t e r i s t i c a l l y l o w i n b o t t o m s e d i m e n t s ( H e u s s e r 1960, Mathewes 1 9 7 3 ) . However, C u p r e s s a c e a e g r a i n s 142 f o u n d i n d e e p e r s a m p l e s i n t h i s s t u d y a p p e a r e d t o be as w e l l - p r e s e r v e d as t h o s e f o u n d n e a r t h e s u r f a c e . I n c r e a s i n g f r e q u e n c i e s t o w a r d s t h e s u r f a c e p r o b a b l y r e p r e s e n t r e a l h i s t o r i c a l i n c r e a s e s r a t h e r t h a n b e t t e r p r e s e r v a t i o n . Mathewes (1973) came t o t h e same c o n c l u s i o n i n h i s i n t e r p r e t a t i o n s o f p o l l e n p r o f i l e s f r o m t h e Haney R e s e a r c h F o r e s t i n s o u t h c o a s t a l B r i t i s h C o l u m b i a . The C u p r e s s a c e a e p o l l e n f r e q u e n c i e s p r o b a b l y i n c l u d e m i n o r amounts o f J u n i p e r u s communis i n a d d i t i o n t o T h u j a and C h a m a e c y p a r i s b e c a u s e J u n i p e r u s a l s o o c c u r s w i t h i n t h e r e g i o n , i n o p e n muskeg and bog w o o d l a n d . O t h e r t a x a s u c h as A b i e s , B e t u l a , T s u g a m e r t e n s i a n a , M y r i c a , A c e r , R o s a c e a e , O p l o p a n a x , C y p e r a c e a e , A s t e r a c e a e , L i n n a e a b o r e a l i s , A p i a c e a e , L y c o p o d i u m , P t e r i d i u m a q u i l i n u m a nd Sphagnum o c c u r b u t v e r y s p o r a d i c a l l y o r i n v e r y l o w f r e q u e n c i e s i n t h i s z o n e . Zone H M - I I I (150-70 cm; s a m p l e s 15-8) The l o w e r 60 cm o f t h i s z one c o n s i s t o f b r o w n i s h - b l a c k m e s i c p e a t w h i c h g r a d e s upward i n t o b r o w n , f i b r i c Sphagnum p e a t i n t h e u p p e r 20 cm. H e r b a c e o u s r e m a i n s a r e a b u n d a n t t h r o u g h o u t t h i s z o n e and woody f r a g m e n t s o c c u r i n t h e l o w e r 20 cm. Zone I I I shows s i g n i f i c a n t i n c r e a s e s i n T h u j a/Chamae- c y p a r i s t y p e , T s u g a h e t e r o p h y l l a , C y p e r a c e a e , Sphagnum and P i n u s c o n t o r t a . P. c o n t o r t a r e a c h e s i t s peak i n t h e l o w e r p o r t i o n o f t h e z o n e , and d e c r e a s e s u p w a r d s . L y c o p o d i u m a l s o r e a c h e s i t s peak f r e q u e n c i e s ( 3 % ) and G e n t i a n a d o u g l a s i a n a , 143 C a r y o p h y l l a c e a e , M e n y a n t h e s t r i f o l i a t a , A b i e s , T s u g a m e r t e n - s i a n a and O p l o p a n a x a l l o c c u r s p o r a d i c a l l y o r i n low f r e q u e n c i e s t h r o u g h o u t t h e z o n e . E r i c a c e o u s p o l l e n i s p r e s e n t i n l ow f r e q u e n c i e s (< 5%) t h r o u g h o u t and t r a c e s ( a few g r a i n s ) o f A p i a c e a e , Rubus chamaemorus, C h e n o p o d i a c e a e and P o a c e a e a l s o o c c u r . The f r e q u e n c i e s o f A l n u s , L y s i c h i t o n , m o n o l e t e f e r n s and P i c e a a l l d e c l i n e f r o m l o w e r t o t h e u p p e r s e c t i o n s o f Zone I I I . Zone HM-IV ( 7 0 - 0 cm; s a m p l e s 7-1) The s u b s t r a t e c o n s i s t s o f brown, f i b r i c Sphagnum p e a t t h r o u g h o u t t h i s z o n e . F r a g m e n t s o f o t h e r moss s p e c i e s s u c h as P l e u r o z ium s c h r e b e r i , H y l o c o m i u m s p l e n d e n s , and P t i l i u m  c r i s t a c a s t r e n s i s o c c u r , e s p e c i a l l y i n t h e u p p e r p o r t i o n s o f t h e z o n e . T h e s e s p e c i e s a r e f a i r l y a b u n d a n t t o g e t h e r w i t h Sphagnum i n t h e s u r f a c e v e g e t a t i o n . T h u j a / C h a m a e c y p a r i s p o l l e n d o m i n a t e s Zone I V ; P. c o n t o r t a i n c r e a s e s t o w a r d s t h e s u r f a c e . M y r i c a , E r i c a c e a e , and G e n t i a n a a l l show p e a k s i n t h e u p p e r s e c t i o n o f t h i s z o n e ( 3 % , 6% and 3% r e s p e c t i v e l y ) , and C y p e r a c e a e c o n t i n u e t o o c c u r t h r o u g h o u t . T. h e t e r o p h y l l a , P i c e a , A l n u s , L y s i c h i t o n and m o n o l e t e f e r n s a l l c o n t i n u e t o d e c l i n e i n Zone I V . L y c o p o d i u m d r o p s o f f t o v e r y low f r e q u e n c i e s and Sphagnum s p o r e s d i s a p p e a r c o m p l e t e l y i n t h e u p p e r h a l f . 5.4 . 2 A b s o l u t e P o l l e n C o n c e n t r a t i o n s A b s o l u t e p o l l e n and s p o r e c o n c e n t r a t i o n s p e r cm^ o f s e d i m e n t a r e p l o t t e d on t h e r i g h t i n F i g . 54. The a b s o l u t e p o l l e n p r o f i l e shows e x t r e m e l y h i g h c o n c e n t r a t i o n s i n Zone I 144 (1600 x 1 0 3 m i c r o f o s s i l s / c m 3 ) f o l l o w e d by an i n i t i a l d e c l i n e (290 x 1 0 3 - 680 x 1 0 3 ) i n Zone I I . The l o w e r s e c t i o n s o f Zone I I I show a d r a s t i c d e c l i n e t o 40 - 50 x 1 0 3 f o l l o w e d by an i n c r e a s e i n u p p e r Zone I I I t o 230 x 1 0 3 and a r a t h e r i r r e g u l a r d e c l i n e t h r o u g h Zone I t o 80 x 1 0 3 a t t h e s u r f a c e . 5.5 D i s c u s s i o n 5.5.1 P o l l e n and Peat S t r a t i g r a p h y The p o l l e n and s p o r e a s s e m b l a g e o f t h e b a s a l zone s u g g e s t s t h a t , f o l l o w i n g t h e d e p o s i t i o n o f m i n e r a l s e d i m e n t s more t h a n 8500 y e a r s ago ( p r o b a b l y by c r e e k s o r r i v e r s d r a i n i n g o f f H ayes M t n . t o t h e w e s t ) , P. c o n t o r t a and A l n u s w e r e t h e m a i n c o l o n i z i n g t r e e s . T. p i i c a t a , C. n o o t k a t e n s i s , T. h e t e r o p h y l l a and P i c e a s i t c h e n s i s were a p p a r e n t l y l e s s i m p o r t a n t i n t h i s i n i t i a l p h a s e . The low p o l l e n p e r c e n t a g e s o f t h e s e t r e e s a r e p r o b a b l y due t o i n p u t f r o m t h e s u r r o u n d i n g r e g i o n a l f o r e s t s and t h u s r e p r e s e n t b a c k g r o u n d l e v e l s . A l s o , t h e i n t e r p r e t a t i o n o f t h i s m i n e r a l - r i c h b a s a l zone i s c l o u d e d by t h e p o s s i b i l i t y t h a t s t r e a m s and r i v e r s d r a i n i n g t h e s u r r o u n d i n g f o r e s t e d s l o p e s d e p o s i t e d p o l l e n and s p o r e s c a r r i e d i n t h e s e d i m e n t l o a d . The e x t r e m e l y h i g h c o n c e n t r a t i o n s o f m o n o l e t e P o l y -p o d i a c e a e s p o r e s s u g g e s t t h a t f e r n s were a s i g n i f i c a n t c omponent o f t h e u n d e r s t o r y v e g e t a t i o n o f t h i s e a r l y f o r e s t , and t h a t m o i s t and n u t r i e n t - r i c h c o n d i t i o n s p r e v a i l e d . The m i n e r a l s e d i m e n t s w o u l d have o r i g i n a t e d f r o m t h e b a s e - r i c h m e t a m o r p h i c b e d r o c k w h i c h d o m i n a t e s t h i s p o r t i o n o f t h e s t u d y 145 a r e a . I n t h e p r e s e n t day r e g i o n a l v e g e t a t i o n , f e r n s p e c i e s s u c h as P o l y s t i c h u m muniturn, A t h y r i u m f i l i x - f e m i n a , D r y o p t e r i s a s s i m i l i s and G y m n o c a r p i u m d r y o p t e r i s ( a l l i n s p e c i e s g r o u p H on t h e s p e c i e s o r d i n a t i o n ; F i g . 8) a r e most a b u n d a n t i n t h e more p r o d u c t i v e f o r e s t c o m m u n i t i e s o f a l l u v i a l l a n d f o r m s and s e e p a g e s l o p e s ( s e e s e c t i o n s 4.4.2.5 and 4 . 4 . 2 . 6 ) . B l e c h n u m s p i c a n t i s t h e o n l y f e r n s p e c i e s w h i c h i s a b u n d a n t i n t h e p o o r e r z o n a l f o r e s t s and bog w o o d l a n d s as w e l l as i n t h e more p r o d u c t i v e f o r e s t c o m m u n i t i e s . A l t h o u g h t h e h i g h e s t c o n c e n t r a t i o n s o f f e r n s p o r e s a r e a s s o c i a t e d w i t h t h e m i n e r a l - r i c h b a s a l s e d i m e n t s o f Zone I ( a g a i n s u g g e s t i n g e x t e r n a l i n p u t s v i a s t r e a m d e p o s i t i o n ) s i g n i f i c a n t c o n c e n t r a t i o n s (10 - 20%) p e r s i s t i n t h e p u r e l y o r g a n i c s a m p l e s i n t h e u p p e r p o r t i o n s o f Zone I I i n d i c a t i n g t h a t l o c a l p o p u l a t i o n s o f f e r n s w e r e p r e s e n t i n t h e d e p o -s i t i o n a l a r e a . I t i s p o s s i b l e t h a t t h e s t u d y s i t e i s l o c a t e d on an o l d m a r i n e t e r r a c e o v e r w h i c h f l u v i a l m a t e r i a l s were d e p o s i t e d f o l l o w i n g e mergence more t h a n 8500 y e a r s ago. The p r e s e n t e l e v a t i o n o f t h e s a m p l i n g s i t e i s 30 m above s e a l e v e l and f r o m h e r e t h e l a n d g r a d u a l l y s l o p e s i n t o M o r s e B a s i n a p p r o x i -m a t e l y one km t o t h e e a s t . D r a m a t i c s e a l e v e l c h a n g e s have b e e n r e p o r t e d by U g o l i n i and Mann (1979) a t L i t u y a B a y , S o u t h -e a s t A l a s k a . T h e s e w o r k e r s g i v e age e s t i m a t e s o f 2-3000 y e a r s f o r t e r r a c e s o f 26 m e l e v a t i o n and 6-8000 y e a r s f o r t e r r a c e s 80 m a b o v e and 5-600 m d i s t a n t f r o m t h e p r e s e n t s h o r e l i n e . H e u s s e r (1960) e s t i m a t e s , on t h e b a s i s o f a p o l l e n p r o f i l e , i 146 t h a t sea l e v e l s may have decreased by a t l e a s t 40 m s i n c e 7 500 B.P. a t P r i n c e Rupert. A r c h e a o l o g i c a l evidence from the P r i n c e Rupert Harbour i n d i c a t e s t h a t the present-day sea l e v e l has e x i s t e d f o r the l a s t 5000 years (Fladmark 1975). Although no m i c r o f o s s i l s suggestive of a marine h a b i t a t were observed i n the b a s a l sediments, more ext e n s i v e and deeper probing of the study s i t e would be r e q u i r e d t o d e t e r -mine the depth of the f l u v i a l m a t e r i a l s and the e x i s t e n c e of marine sediments beneath. The p o l l e n and spore f r e q u e n c i e s of Zone HM-II i n d i c a t e t h a t a f a i r l y wet and n u t r i e n t - r i c h spruce - western hemlock -a l d e r - L y s i c h i t o n - f e r n f o r e s t community became e s t a b l i s h e d f o l l o w i n g the i n i t i a l c o l o n i z a t i o n by shore pine, a l d e r and f e r n s . E r i c a c e o u s shrubs such as Vaccinium spp. and M e n z i e s i a c o u l d have been components of t h i s f o r e s t as w e l l . The decreased importance of P. c o n t o r t a i n Zone II i s p r o b a b l y due t o i t s shade i n t o l e r a n c e i n combination with edaphic f a c t o r s . Based on present day s p e c i e s autecology, the wet, humus-enriched, r e g o s o l i c s o i l s d eveloping from b a s e - r i c h a l l u v i u m would have favoured the growth of S i t k a spruce, red a l d e r , western red-cedar and western hemlock over pine (Fowells 1965; K r a j i n a 1969). As m i n e r a l s o i l s matured and humus l a y e r s b u i l t up, the P i c e a and Alnus component of t h i s a l l u v i a l f o r e s t a p p a r e n t l y decreased i n favour of T. h e t e r o p h y l l a and Thuja/Chamae-c y p a r i s . The s p e c i e s composition suggested by the p o l l e n / spore f r e q u e n c i e s i n the upper p o r t i o n s of Zone II i s s i m i l a r 147 t o t h a t of the present-day cedar - spruce - hemlock -L y s i c h i t o n - Rhizominum community type of i n a c t i v e f l u v i a l landforms ( s e c t i o n 4.4.2.5). /An example of t h i s community type (represented by r e l e v e 10) occurs adjacent t o a creek approximately 150 m west of the p o l l e n p r o f i l e sampling s i t e . I t i s p o s s i b l e t h a t a s i m i l a r f o r e s t community once surrounded the sampling s i t e or t h a t t h i s a l l u v i a l f o r e s t p r e s e n t l y l o c a l i z e d along the creek was once more widespread over the area. The organo-mineral and org a n i c m a t e r i a l s t h a t occur i n Zones I, I I , and the lower s e c t i o n s o f Zone I I I show s i m i l a r i t i e s t o the amorphous, w e l l h u m i f i e d 'Hi' and 'H' h o r i z o n s which c h a r a c t e r i z e the Humimor and Hydromor humus forms of many of the f o r e s t s o i l s w i t h i n the study area. High c o n c e n t r a t i o n s of woody remains i n the peat of Zones I and I I a l s o supports the hy p o t h e s i s t h a t a p r o d u c t i v e f o r e s t ecosystem once occupied the s i t e . That much of the amorphous and w e l l h u m i f i e d matrix of the lower four samples i s lig n e o u s (woody) i n o r i g i n i s suggested by obvious woody fragments and by the l a r g e decreases i n volume t h a t were noted f o l l o w i n g treatment with potassium hydroxide (KOH), used p r i m a r i l y f o r the removal of lig n e o u s m a t e r i a l s p r i o r t o p o l l e n p r e p a r a t i o n . The samples between 160 cm and 200 cm showed much l a r g e r volume decreases than d i d the upper samples. Marked changes i n e c o l o g i c a l c o n d i t i o n s are suggested by the c o n t r a s t i n g p o l l e n and spore assemblages of Zones II and I I I . A d e c l i n e i n n u t r i e n t s t a t u s i s i n d i c a t e d by the 148 d e c r e a s i n g f r e q u e n c i e s o f m o n o l e t e f e r n s , L y s i c h i t o n , P i c e a and A l n u s . T h e r e a r e c o r r e s p o n d i n g i n c r e a s e s i n t a x a w h i c h a r e more c h a r a c t e r i s t i c o f t h e s c r u b b y c e d a r - c y p r e s s -h e m l o c k - V a c c i n i u m - C o r n u s - R h y t i d i a d e l p h u s f o r e s t c o m m u n i t y t y p e ( S e c t i o n 4 . 4 . 2 . 4 ) . R e c a l l t h a t T h u j a , C h a m e a c y p a r i s and T. h e t e r o p h y l l a a r e t h e most a b u n d a n t t r e e s p e c i e s i n t h e s e s c r u b b y f o r e s t s , w i t h s h o r e p i n e o f t e n p r e s e n t i n t h e d o m i n a n t t r e e l a y e r . P i c e a s i t c h e n s i s and A b i e s a m a b i l i s a r e uncommon, u s u a l l y o n l y p r e s e n t a s s c a t t e r e d r e g e n e r a t i o n o f v e r y low v i g o u r . A l n u s r u b r a i s a l s o v e r y r a r e l y p r e s e n t b u t t h i s s p e c i e s p r o d u c e s c o p i o u s amounts o f p o l l e n and i s a l w a y s w e l l - r e p r e s e n t e d t h r o u g h o u t p o l l e n p r o f i l e s f r o m c o a s t a l B.C. ( H e u s s e r 1960; Mathewes 197 3 ; Mathewes and R ouse 1975; Hebda and Rouse 1 9 7 9 ) . T. m e r t e n s i a n a d o e s o c c u r i n t h e z o n a l f o r e s t s b u t i n r e l a t i v e l y low a m o u n t s . I n t h e p o l l e n p r o f i l e t h i s s p e c i e s r e a c h e s i t s peak i n Zone I I I b u t i s n e v e r p r e s e n t i n v e r y h i g h f r e q u e n c i e s . I n t e r m s o f t h e t r e e l a y e r s , t h e p o l l e n s p e c t r a o f Zone I I I t h u s c o r r e s p o n d s c l o s e l y t o t h e s p e c i e s c o m p o s i t i o n o f t h e p r e s e n t - d a y c l i m a t i c c l i m a x f o r e s t s . The p o l l e n s p e c t r a o f u n d e r s t o r y s p e c i e s i n Zone I I I a l s o show r e l a t i o n s h i p s w i t h t h e c e d a r - c y p r e s s - h e m l o c k c l i m a t i c c l i m a x f o r e s t s . The d e c l i n i n g f r e q u e n c i e s o f m o n o l e t e f e r n s and L y s i c h i t o n a r e p r o b a b l y t h e most s t r i k i n g f e a t u r e s o f t h i s z o n e . The f e r n f l o r a i s much l e s s d i v e r s e i n c l i m a t i c c l i m a x f o r e s t s t h a n i n t h e r i c h e r s e e p a g e and a l l u v i a l f o r e s t c o m m u n i t i e s . B l e c h n u m s p i c a n t i s t h e o n l y a b u n d a n t f e r n 149 s p e c i e s i n c l i m a t i c c l i m a x f o r e s t s . V a r i o u s s p e c i e s o f L y c o p o d i u m s u c h a s L. a n n o t i n u m and L. c l a v a t u m a r e common as w e l l . L y c o p o d i u m i n c r e a s e s i n f r e q u e n c y i n zone I I I . A l t h o u g h L y s i c h i t o n i s w i d e s p r e a d t h r o u g h o u t t h e s t u d y a r e a a n d i s o f t e n p r e s e n t i n c l i m a t i c c l i m a x f o r e s t s , i t i s commonly r e s t r i c t e d t o d r a i n a g e c h a n n e l s o r d e p r e s s i o n s w h e r e t h e r e i s m i n e r a l s o i l e x p o s u r e o r s e e p a g e o r i g i n a t i n g f r o m a m i n e r a l s o u r c e . I t i s much more a b u n d a n t , o f more u n i f o r m o c c u r r e n c e , and much more v i g o r o u s i n t h e mucky G l e y s o l s o r R e g o s o l s a s s o c i a t e d w i t h t h e r i c h e r , more p r o d u c t i v e s p r u c e -c e d a r - a l d e r a l l u v i a l f o r e s t s ( F i g . 5 5 ) . L y s i c h i t o n a p p e a r s t o n e e d a b o v e - a v e r a g e n u t r i e n t c o n d i t i o n s a s w e l l a s s a t u r a t e d s o i l c o n d i t i o n s i n o r d e r t o t h r i v e . L a w r e n c e (1958) d e s c r i b e s t h e r o l e o f L y s i c h i t o n i n p i t p o nd f o r m a t i o n d u r i n g s u c c e s s i o n f r o m f o r e s t t o muskeg. T h i s s p e c i e s a p p e a r s t o m a i n t a i n a f o o t h o l d d u r i n g f o r e s t d e t e r i o r a t i o n a nd muskeg d e v e l o p m e n t b y s h a d i n g o u t t h e Sphagnum g r o w i n g a r o u n d i t , t h u s c r e a t i n g deep p i t s W h e r e v e r i t o c c u r s i n t h e s c r u b f o r e s t o r muskeg. I n t h e s t u d y a r e a L y s i c h i t o n o c c u r s s p o r a d i c a l l y i n t h e o p e n muskeg a n d u s u a l l y o n l y i n l o c a l i z e d p a t c h e s i n t h e b o g w o o d l a n d and s c r u b f o r e s t s . B a s e d on i t s p r e s e n t day h a b i t a t p r e f e r e n c e s i n c o a s t a l f o r e s t a nd b o g e c o s y s t e m s , i t s d r a s t i c a l l y d e c l i n i n g f r e q u e n c i e s f r o m Zone I I t o Zone I I I h e l p t o c o n f i r m t h e d e t e r i o r a t i o n o f a p r o d u c t i v e f o r e s t t o s c r u b . The E r i c a c e a e p o l l e n s p e c t r u m i s o f l i m i t e d v a l u e i n t h e r e c o n s t r u c t i o n o f v e g e t a t i o n h i s t o r y s i n c e e r i c a d t e t r a d s a r e d i f f i c u l t t o i d e n t i f y t o s p e c i e s o r g e nus (Hebda 1977, 1979) 150 F i g . 5 5 . L y s i c h i t o n a m e r i c a n u m , an abundant s p e c i e s i n a l l u v i a l f o r e s t c o m m u n i t i e s i n the s t u d y a r e a . ISO A 151 and d i f f e r e n t s p e c i e s w i t h i n t h i s f a m i l y a r e p r e s e n t i n a l l c o m m u n i t y t y p e s i n t h e s t u d y a r e a . V a c c i n i u m a l a s k a e n s e , V. o v a l i f o l i u m , V. p a r v i f o l i u m and Menz i e s i a f e r r u g i n e a o c c u r i n a l l t h e f o r e s t and w o o d l a n d c o m m u n i t i e s w h i l e G a u l t h e r i a  s h a l l o n i s r e s t r i c t e d m o s t l y t o z o n a l f o r e s t s and bog w o o d l a n d . O t h e r s p e c i e s s u c h as Ledum g r o e n l a n d i c u m and K a l m i a m i c r o p h y l l a a r e r e s t r i c t e d t o v e r y s c r u b b y f o r e s t s , bog w o o d l a n d and o pen bog. I t i s t h e r e f o r e n o t s u r p r i s i n g t h a t t h e r e i s l i t t l e d i f f e r e n c e i n t h e f r e q u e n c y o f E r i c a c e a e p o l l e n b e t w e e n z o n e s I I , I I I , and I V . Hebda ( 1 9 7 7 , 1979) d i s t i n g u i s h e d t h r e e m a j o r s i z e r a n g e s o f E r i c a d t e t r a d s and was a b l e t o a s s o c i a t e t h e s e t e t r a d s i z e s w i t h d r y , medium and wet h a b i t a t s i n B u r n s Bog, S o u t h w e s t e r n B r i t i s h C o l u m b i a . The o n l y f o r e s t s p e c i e s he e x a m i n e d , how-e v e r , was G a u l t h e r i a s h a l l o n and i t was t h e r e f o r e n o t p o s s i b l e t o u s e t h e s e r e s u l t s t o d i s t i n g u i s h bog f r o m f o r e s t h a b i t a t s i n t h i s s t u d y . A l s o , Hebda (1977) c a u t i o n s a g a i n s t a p p l y i n g h i s m o d e l t o o t h e r bogs w i t h d i f f e r e n t s p e c i e s c o m p o s i t i o n and s e d i m e n t a r y r e g i m e s . The i n c r e a s e i n C y p e r a c e a e p o l l e n i n Zone I I I s u g g e s t s t h e d e v e l o p m e n t o f v e r y open f o r e s t c o n d i t i o n s . C a r e x a n t h o x - a n t h e a , C. l a e v i c u l m i s and C. p h y l l o m a n i c a o c c u r i n t h e s c r u b b i e s t z o n a l f o r e s t s , w h i l e numerous s e d g e s , i n c l u d i n g C. p l u r i f l o r a , C. p a u c i f l o r a , C. I i v i d a and T r i c h o p h o r u m  c e s p i t o s u m , i n a d d i t i o n t o t h e a b o v e t h r e e s p e c i e s o c c u r f r e q u e n t l y i n bog w o o d l a n d . The Sphagnum s p o r e p r o f i l e must be i n t e r p r e t e d w i t h 152 c a u t i o n s i n c e many s p e c i e s o f Sphagnum a r e r a r e l y f o u n d w i t h s p o r o p h y t e s a n d p r e s u m a b l y r e l y t o a l a r g e e x t e n t on a s e x u a l r e p r o d u c t i o n ( h o w e v e r , a few s p e c i e s s u c h as S_. g i r g e n s o h n i i a n d S. r u b e l l u m h a v e f r e q u e n t l y b e e n o b s e r v e d w i t h s p o r o p h y t e s i n t h e P r i n c e R u p e r t a r e a ) . The h i g h f r e q u e n c y o f Sphagnum s p o r e s i n Zone I I I i s u n d e r s t a n d a b l e b e c a u s e t h i s g e n us a p p e a r s t o be i n s t r u m e n t a l i n t h e p a l u d i f i c a t i o n o r 1 b o g i f i -c a t i o n ' o f f o r e s t s ( L a w r e n c e 1958, N e i l a n d 1 9 7 1 ) . S. g i r g e n -s o h n i i i s t h e common s p e c i e s i n c l o s e d u p l a n d f o r e s t s , w h e r e a s more l i g h t - r e q u i r i n g Sphagnum s p e c i e s s u c h a s S_. p a p i l l o s u m , S. r u b e l l u m , a n d S_. m a g e l l a n i c u m o n l y c o l o n i z e t h e most o p e n a n d s c r u b b y f o r e s t s . O p l o p a n a x i s r a r e l y p r e s e n t i n t h e s c r u b f o r e s t s e x c e p t i n t h e more n u t r i e n t r i c h m i c r o h a b i t a t s . I t s s p o r a d i c o c c u r r e n c e a n d l o w f r e q u e n c i e s t h r o u g h o u t t h e p r o f i l e s u g g e s t t h a t i t n e v e r was an i m p o r t a n t component o f t h e u n d e r s t o r y v e g e t a t i o n . The s p o r a d i c o c c u r r e n c e o f G e n t i a n a d o u g l a s i a n a p o l l e n t h r o u g h o u t Zones I I I and I V i s s i g n i f i c a n t s i n c e i n t h e p r e s e n t d a y v e g e t a t i o n t h i s s p e c i e s i s r e s t r i c t e d t o b o g w o o d l a n d o r o p e n b o g h a b i t a t s . I t s p r e s e n c e s u g g e s t s t h e d e v e l o p m e n t o f b o g g y m i c r o h a b i t a t s n o t f a r f r o m t h e s a m p l i n g s i t e . T h i s s p e c i e s i s i n s e c t - p o l l i n a t e d a n d p o l l e n t r a n s p o r t f r o m n e a r b y o p e n b o g s i s a p o s s i b i l i t y . T r a n s p o r t b y i n s e c t s may a l s o e x p l a i n t h e s p o r a d i c o c c u r r e n c e o f M e n y a n t h e s a n d N u p h a r i n t h e p r o f i l e . B o t h M. t r i f o l i a t a a n d N. l u t e a s u b s p . p o l y s e p a l a a r e i n s e c t - p o l l i n a t e d a n d a r e common i n t h e l a r g e r 153 p o n d s i n t h e muskeg. A few s u c h p o n d s o c c u r w i t h i n 100 m o f t h e s a m p l i n g s i t e . B a s e d on p r e s e n t - d a y s o i l - v e g e t a t i o n r e l a t i o n s h i p s w i t h i n t h e s t u d y a r e a ( C h a p t e r 4 ) , t h e c h a n g e s i n s p e c i e s c o m p o s i t i o n f r o m Zone I I t o Zone I I I p r o b a b l y p a r a l l e l t h e d e v e l o p m e n t o f p o d z o l i c and g l e y s o l i c s o i l s h a v i n g t h i c k , w e t , a c i d i c , Mor humus f o r m s s i m i l a r t o t h o s e f r e q u e n t l y a s s o c i a t e d w i t h t h e s c r u b b y c l i m a t i c c l i m a x f o r e s t c ommunity t y p e . The p e r i o d i c f l o o d i n g and d e p o s i t i o n o f f r e s h a l l u v i a l s e d i m e n t s t h a t was p r o b a b l y once i m p o r t a n t i n m a i n t a i n i n g s i t e f e r t i l i t y , w o u l d by t h i s t i m e have c e a s e d . M i n e r a l s e d i m e n t s and l a t e r a l m i n e r a l s e e p a g e w o u l d have become l e s s i m p o r t a n t i n s u p p l y i n g n u t r i e n t s t o t h e v e g e t a t i o n as o r g a n i c m a t t e r c o n t i n u e d t o b u i l d up a t t h e s o i l s u r f a c e . Low d e c o m p o s i t i o n r a t e s and s l o w r a t e s o f n u t r i e n t r e l e a s e f r o m t h e s e a c i d i c Mor humus f o r m s ( K l i n k a e t a l . 1981) w o u l d a l s o have c o n t r i -b u t e d t o d e c r e a s e d n u t r i e n t a v a i l a b i l i t y i n t h e s e s o i l s . The p o l l e n a s s e m b l a g e o f zone I V , p a r t i c u l a r l y f o r t r e e s and s h r u b s , shows c l e a r r e l a t i o n s h i p s w i t h t h e e x i s t i n g bog w o o d l a n d v e g e t a t i o n a t t h e s a m p l i n g s i t e . T h i s v e g e t a t i o n i s c h a r a c t e r i z e d by a m a i n c a n o p y o f s t u n t e d P. c o n t o r t a w i t h a b u n d a n t C. n o o t k a t e n s i s and s c a t t e r e d T. p l i c a t a and T. h e t e r o p h y l l a i n t h e s h r u b l a y e r s . The t r e e c o m p o s i t i o n c o r r e s p o n d s c l o s e l y t o t h e p o l l e n s p e c t r a o f c o n i f e r o u s t r e e s p e c i e s i n Zone I V . The low f r e q u e n c i e s o f P i c e a , T. h e t e r o - p h y l l a and A b i e s u n d o u b t e d l y r e f l e c t b a c k g r o u n d i n p u t s f r o m t h e s u r r o u n d i n g m o u n t a i n s l o p e s and n e a r b y a l l u v i a l f o r e s t . 154 M o u n t a i n h e m l o c k i s a b u n d a n t on n e a r b y Hayes Mtn. above 500 m and i s a l s o p r e s e n t as s c a t t e r e d i n d i v i d u a l s i n t h e s c r u b b y f o r e s t s , bog w o o d l a n d and muskeg a l l t h e way t o s e a l e v e l . E r i c a c e o u s s h r u b s s u c h as G a u l t h e r i a s h a l l o n , Ledum  g r o e n l a n d i c u m , V a c c i n i u m p a r v i f o l i u m and M e n z i e s i a f e r r u g i n e a a r e d o m i n a n t i n t h e u n d e r s t o r y v e g e t a t i o n s u r r o u n d i n g t h e s a m p l i n g s i t e . M i n o r amounts o f K a l m i a m i c r o p h y l l a and M y r i c a g a l e a l s o o c c u r . B o t h E r i c a c o u s and M y r i c a p o l l e n i n c r e a s e i n a b u n d a n c e i n t h e u p p e r s e c t i o n s o f Zone I V . The s l i g h t i n c r e a s e s i n A l n u s t o w a r d s t h e s u r f a c e must r e f l e c t i n p u t s f r o m t h e n e a r b y a l l u v i a l f o r e s t o r a l d e r t h i c k e t s a l o n g t h e h i g h w a y , b e c a u s e t h e r e i s no A l n u s i n t h e bog w o o d l a n d s u r r o u n d i n g t h e s i t e . D o m i n a n t h e r b s w i t h u n i f o r m d i s t r i b u t i o n t h r o u g h o u t t h e bog w o o d l a n d a r e F a u r i a c r i s t a - g a l l i , L i n n a e a b o r e a l i s , V a c c i n i u m v i t i s - i d a e a , C o r n u s u n a l a s c h k e n s i s , Maianthemum  d i l a t a t u m and C a r e x p l u r i f l o r a . H e r b s r e s t r i c t e d t o l o c a l i z e d p a t c h e s o r c l u m p s i n c l u d e L y s i c h i t o n a m e r i c a n u m , C o p t i s t r i - f o l i a , C. a s p l e n i i f o l i a , L y c o p o d i u m a n n o t i n u m , C a r e x s i t - c h e n s i s , C. l i v i d a , S a n g u i s o r b a o f f i c i n a l i s , T r i e n t a l i s  e u r o p e a , L i s t e r a c o r d a t a and D r o s e r a r o t u n d i f o l i a . E x c e p t f o r t h e d e c l i n i n g f r e q u e n c i e s o f L y s i c h i t o n , m o n o l e t e f e r n s and s l i g h t i n c r e a s e s i n C y p e r a c e a e , t h e p o l l e n and s p o r e s p e c t r a f o r h e r b s and v a s c u l a r c r y p t o g r a m s do n o t a d e q u a t e l y r e f l e c t t h e w e l l d e v e l o p e d h e r b a c e o u s v e g e t a t i o n a t t h e s a m p l i n g s i t e . T h i s i s p r o b a b l y b e c a u s e most o f t h e s e h e r b s p r o d u c e r e l a t i v e l y s m a l l amounts o f p o l l e n , and i n s e c t s , r a t h e r t h a n 155 w i n d , a r e t h e i r p r i n c i p a l p o l l e n v e c t o r . F u r t h e r m o r e , most o f t h e s e h e r b s have p a t c h y d i s t r i b u t i o n s and some s p e c i e s ( s u c h a s F a u r i a c r i s t a - g a l l i ) w h i c h a r e n o r m a l l y a b u n d a n t i n bog w o o d l a n d s e l d o m f l o w e r e x c e p t i n open b o g s . The d o m i n a n t b r y o p h y t e s o f t h e bog w o o d l a n d a r e Sphagnum  g i r g e n s o h n i i , P l e u r o z ium s c h r e b e r i , H y l o c o m i u m s p l e n d e n s , Sphagnum p a p i l l o s u m , R h y t i d i a d e l p h u s l o r e u s and P t i l i u m  c r i s t a - c a s t r e n s i s . O t h e r t h a n Sphagnum ( w h i c h has a l r e a d y b e e n d i s c u s s e d ) , b r y o p h y t e s a r e s e l d o m r e p r e s e n t e d i n p e a t p r o f i l e s e x c e p t as s u b f o s s i l r e m a i n s (Mathewes 1 9 7 3 ) , a few o f IS w h i c h w ere o b s e r v e d o n l y i n t h e s a m p l e s n e a r t h e s u r f a c e . T h a t t h e p o l l e n a s s e m b l a g e o f t h e s u r f a c e zone has c l o s e a f f i n i t i e s t o t h e bog w o o d l a n d v e g e t a t i o n i m m e d i a t e l y s u r r o u n d i n g t h e s a m p l i n g s i t e s u b s t a n t i a t e s t h e i n t e r p r e t a t i o n t h a t t h e p o l l e n p r o f i l e r e f l e c t s l o c a l v e g e t a t i o n h i s t o r y and s u c c e s s i o n . The r e l a t i v e l y m i n o r i n f l u e n c e o f t h e r e g i o n a l v e g e t a t i o n i s i n d i c a t e d by t h e low f r e q u e n c i e s o f t a x a s u c h as P i c e a , A l n u s , A b i e s , T s u g a h e t e r o p h y l l a , and T. m e r t e n s i a n a i n u p p e r Zone I V , i n s p i t e o f t h e i r a b u n d a n c e i n t h e n e a r b y a l l u v i a l f o r e s t and on t h e a d j a c e n t f o r e s t e d m o u n t a i n s l o p e s . 5.5.2 A b s o l u te P o l l e n C o n c e n t r a t i o n s I n t h e i n t e r p r e t a t i o n o f a b s o l u t e p o l l e n c o n c e n t r a t i o n s , f o u r m a i n f a c t o r s must be c o n s i d e r e d : r a t e s o f s u b s t r a t e a c c u m u l a t i o n , r a t e s o f p o l l e n d e p o s i t i o n , n a t u r e o f t h e s u b s t r a t e ( m i n e r a l o r o r g a n i c ) , and t h e s i g n i f i c a n t d e g r e e o f e r r o r i n h e r e n t t o t h e m e t h o d o l o g y ( S t o c k m a r r 1 9 7 1 ; Mathewes 1973 ; D a v i s e t a l . 1 9 7 7 ) . B e c a u s e o f t h e i n t e r a c t i o n o f t h e s e 156 f o u r f a c t o r s , t h e i n t e r p r e t a t i o n o f a b s o l u t e p o l l e n p r o f i l e s l e a v e s much room f o r s p e c u l a t i o n . A l t h o u g h m a r k e d f l u c t u a t i o n s i n a b s o l u t e m i c r o f o s s i l c o n c e n t r a t i o n s o c c u r t h r o u g h o u t t h e p r o f i l e , a n d some o f t h i s f l u c t u a t i o n may be a t t r i b u t e d t o e r r o r , t h e m a g n i t u d e o f t h e d e c l i n e f r o m t h e b o t t o m o f t h e p r o f i l e t o t h e s u r f a c e s u g g e s t s t h a t e c o l o g i c a l c o n d i t i o n s h a v e a l t e r e d d r a s t i c a l l y o v e r a p e r i o d o f 8700 y e a r s . H i g h e r p o l l e n a n d s p o r e c o n c e n t r a t i o n s i n Zones I and I I c o u l d h a v e r e s u l t e d i n p a r t f r o m t h e p r e v a l e n c e o f p o l l e n -p r o d u c t i v e t a x a i n t h e s e z o n e s , t o g e t h e r w i t h t h e r e l a t i v e l y l o w r a t e s o f o r g a n i c m a t t e r a c c u m u l a t i o n t h a t w o u l d p r o b a b l y h a v e b e e n a s s o c i a t e d w i t h t h e e a r l y a l l u v i a l f o r e s t . C o m p a c t i o n a n d g r e a t e r h u m i f i c a t i o n o f t h e l o w e r s e d i m e n t s w o u l d a l s o t e n d t o i n c r e a s e p o l l e n a n d s p o r e c o n c e n t r a t i o n s . I f e x t e r n a l p o l l e n a n d s p o r e i n p u t s f r o m t h e a l l u v i a l s e d i m e n t s p l a y e d a r o l e , t h e n t h e c o n c e n t r a t i o n s w o u l d a g a i n t e n d t o be h i g h e r i n t h e s e o r g a n o - m i n e r a l b a s a l s e d i m e n t s . The i n c r e a s e d r a t e s o f o r g a n i c m a t t e r a c c u m u l a t i o n , i n c o m b i n a t i o n w i t h d e c r e a s e d p o l l e n a n d s p o r e p r o d u c t i o n a s s o c i a t e d w i t h f o r e s t d e g r a d a t i o n a n d t h e g r a d u a l d e v e l o p m e n t o f muskeg, c o u l d p o s s i b l y e x p l a i n t h e l o w e r p o l l e n a n d s p o r e c o n c e n t r a t i o n s i n t h e u p p e r z o n e s o f t h e p r o f i l e . 5.5.3 Summary o f t h e Bog Woodland S u c c e s s i o n a l Development By c o m p a r i n g t h e p o l l e n a n d s p o r e a s s e m b l a g e s o f e a c h o f t h e f o u r z o n e s i n t h e p r o f i l e w i t h t h e s p e c i e s c o m p o s i t i o n o f e x i s t i n g c o m m u n i t i e s w i t h i n t h e s t u d y a r e a , a n d b y u t i l i z i n g 157 k n o w l e d g e o f p r e s e n t - d a y s o i l - v e g e t a t i o n r e l a t i o n s h i p s , a p o s s i b l e s e q u e n c e o f c h a n g i n g e c o l o g i c a l c o n d i t i o n s l e a d i n g t o t h e d e v e l o p m e n t o f t h e p r e s e n t bog w o o d l a n d e c o s y s t e m has been o u t l i n e d . T h i s s e q u e n c e b e g i n s w i t h P i n u s c o n t o r t a , A l n u s ( p r o b a b l y A. R u b r a ) and f e r n s w i t h p e r h a p s m i n o r amounts o f P i c e a ( p r o b a b l y P. s i t c h e n s i s ) , T s u g a h e t e r o p h y l l a and T h u j a  p l i c a t a / ( C h a m a e c y p a r i s n o o t k a t e n s i s ) c o l o n i z i n g t h e a l l u v i a l s e d i m e n t s more t h a n 8,500 y e a r s ago. T h i s c o l o n i z i n g p h a s e was s o o n r e p l a c e d by a p r o d u c t i v e f o r e s t o f m a i n l y P i c e a , A l n u s , T. h e t e r o p h y l l a , L y s i c h i t o n a m e r i c a n u m and f e r n s ( p r o b a b l y m a i n l y D r y o p t e r i s a s s i m i l i s , A t h y r i u m f i l i x - f e m i n a , G y m n o c a r p i u m d r y o p t e r i s , B l e c h n u m s p i c a n t and p o s s i b l y P o l y - s t i c h u m m u n i t u m ) . As a l l u v i a l d e p o s i t i o n c e a s e d , p e r h a p s as a r e s u l t o f c h a n g i n g s t r e a m c o u r s e s , r e g o s o l i c s o i l s d e v e l o p e d i n t o p o d z o l i c o r g l e y s o l i c s o i l s w i t h i n c r e a s i n g l y t h i c k humus a c c u m u l a t i o n s . As s o i l s m a t u r e d , a T h u j a / ( C h a m a e c y p a r i s ) -d o m i n a t e d f o r e s t s i m i l a r t o t h e p r e s e n t - d a y c e d a r - s p r u c e -h e m l o c k - L y s i c h i t o n - R h i z o m n i u m a l l u v i a l f o r e s t c o m m u n i t y t y p e g r a d u a l l y d e v e l o p e d . D e c l i n i n g n u t r i e n t a v a i l a b i l i t y and p o o r d r a i n a g e r e s u l t i n g f r o m t h e d e v e l o p m e n t o f t h i c k Mor humus f o r m s g a v e r i s e t o a f o r e s t s i m i l a r t o t h e c e d a r -c y p r e s s - h e m l o c k - V a c c i n i u m - C o r n u s - R h y t i d i a d e l p h u s c o m m u n i t y t y p e c o n s i d e r e d t h e p r e s e n t - d a y c l i m a t i c c l i m a x w i t h i n t h e s t u d y a r e a . C o n t i n u e d p a l u d i f i c a t i o n o f t h i s s c r u b b y f o r e s t r e s u l t e d i n s u c c e s s i o n t o t h e p i n e - c y p r e s s -c e d a r - G a u l t h e r i a - F a u r i a - P l e u r o z ium - Sphagnum bog wood-15 l a n d c o m m u n i t y c u r r e n t l y s u r r o u n d i n g t h e s a m p l i n g s i t e . T h i s s u c c e s s i o n a l s e q u e n c e s h o u l d n o t be c o n s i d e r e d as t h e g e n e r a l m o d e l f o r bog d e v e l o p m e n t w i t h i n t h e s t u d y a r e a . F i r s t o f a l l , i t i s b a s e d on o n l y one p r o f i l e and s e c o n d l y i f t h e many bogs and bog w o o d l a n d s u n d e r l a i n by b e d r o c k r a t h e r t h a n m i n e r a l s e d i m e n t s were p r e c e d e d by f o r e s t s , t h e s e f o r e s t s w o u l d p r o b a b l y n o t h a v e b e e n as v i g o r o u s o r o f t h e same c o m p o s i t i o n as t h o s e t h a t o c c u r r e d on a l l u v i a l l a n d f o r m s . 5.5.4 Comparison W i t h O t h e r S t u d i e s o f P a c i f i c C o a s t Bog Development S i m i l a r s u c c e s s i o n a l s e q u e n c e s t o t h a t j u s t o u t l i n e d have b e e n d e s c r i b e d e l s e w h e r e a l o n g t h e P a c i f i c C o a s t . The b e s t d o c u m e n t e d e x a m p l e i s a r e c e n t s t u d y by U g o l i n i and Mann ( 1 9 7 9 ) , w h i c h d e s c r i b e s an 8000 y e a r c h r o n o s e q u e n c e f r o m S i t k a s p r u c e f o r e s t t o muskeg on m a r i n e t e r r a c e s i n L i t u y a B a y , S o u t h e a s t A l a s k a . T h e s e a u t h o r s c o n c l u d e t h a t t h e accumu-l a t i o n o f s e s q u i o x i d e s i n t h e B h o r i z o n s o f t h e d e v e l o p i n g p o d z o l s c a u s e s t h e f o r m a t i o n o f p l a c i c h o r i z o n s w h i c h r e s t r i c t i n t e r n a l s o i l d r a i n a g e and i n i t i a t e a n a e r o b i c c o n d i t i o n s . T h e s e p r o c e s s e s c o m b i n e w i t h t h e h i g h p r e c i p i t a t i o n , c o o l t e m p e r a t u r e s and l o w e v a p o t r a n s p i r a t i o n r a t e s o f t h e r e g i o n t o l o w e r d e c o m p o s i t i o n r a t e s . Hence, o r g a n i c m a t t e r a c c u m u l a t e s more r a p i d l y a t t h e s o i l s u r f a c e . D r a i n a g e i s f u r t h e r impeded by t h e t h i c k s u r f a c e o r g a n i c h o r i z o n s . The P i c e a s i t c h e n s i s f o r e s t s w h i c h p r e d o m i n a t e on r e c e n t l y emerged t e r r a c e s a r e g r a d u a l l y r e p l a c e d by s c r u b b i e r f o r e s t s o f T. h e t e r o p h y l l a , C. n o o t k e n s i s , and T. m e r t e n s i a n a . P. c o n t o r t a - muskeg i s t h e 159 d o m i n a n t v e g e t a t i o n on t h e o l d e s t m a r i n e t e r r a c e s . S o i l d e v e l o p m e n t f o l l o w s a s e q u e n c e f r o m R e g o s o l s t h r o u g h P o d z o l s w i t h t h i c k s u r f a c e o r g a n i c h o r i z o n s (22 - 35 cm) and c e m e n t e d s u b s u r f a c e h o r i z o n s , t o p e a t s o i l s g r e a t e r t h a n 1 m d e e p . E v i d e n c e o f t h e p l a c i c h o r i z o n w h i c h p r e s u m a b l y i n i t i a t e d t h e p a l u d i f i c a t i o n p r o c e s s c a n n o t be f o u n d b e n e a t h t h e p e a t s o i l s b e c a u s e r e d u c i n g c o n d i t i o n s w i t h i n t h e m i n e r a l h o r i z o n s h a v e b r o u g h t a b o u t t h e s o l u t i o n o f t h e i r o n - c e m e n t i n g compounds. The s u c c e s s i o n o f f o r e s t and bog c o m m u n i t i e s on p r o g r e s s i v e l y o l d e r m a r i n e t e r r a c e s d e s c r i b e d by U g o l i n i and Mann (1979) c l o s e l y p a r a l l e l s t h e ^ s e q u e n c e p o r t r a y e d i n t h i s p a l y n o l o g i c a l s t u d y , a l t h o u g h t h e r o l e t h a t c e m e n t e d h o r i z o n s p l a y e d i n t h e H a y e s M t n . s e q u e n c e c a n n o t be d e t e r m i n e d . F o r e s t - bog s u c c e s s i o n a l r e l a t i o n s h i p s i n S o u t h e a s t A l a s k a were o r i g i n a l l y d i s c u s s e d by much e a r l i e r w o r k e r s . D a s c h n o w s k i - S t o k e s (1941) d e s c r i b e s bog s t r a t i g r a p h y p r o f i l e s n e a r K e t c h i c a n , P e t e r s b u r g and J u n e a u c o n t a i n i n g l a y e r s o f woody p e a t and b u r i e d stumps and r o o t s w h i c h he i n t e r p r e t e d as r e f l e c t i n g d r i e r , warmer c l i m a t i c i n t e r v a l s . B o t h Z a c h (1950) and L a w r e n c e (1958) d i s c u s s how muskeg d e v e l o p m e n t f o l l o w s f o r e s t e s t a b l i s h m e n t and e v e n t u a l l y f o r m s t h e c l i m a x e c o s y s t e m i n many p a r t s o f S o u t h e a s t A l a s k a . N e i l a n d (1971) made s i m i l a r o b s e r v a t i o n s , b u t , l i k e S t e p h e n s e t a l . ( 1 9 7 0 ) , s h e c o n c l u d e d t h a t i n some a r e a s i n S o u t h e a s t A l a s k a f o r e s t s a p p e a r e d t o be e x p a n d i n g a t t h e e x p e n s e o f b o g s . . T h i s t o p i c w i l l be f u r t h e r d i s c u s s e d i n C h a p t e r 6. On t h e b a s i s o f p o l l e n a n a l y s i s and s y n e c o l o g i c a l 16 s t u d i e s , Wade (1965) c o n c l u d e d t h a t Sphagnum bogs n e a r T o f i n o on s o u t h w e s t e r n V a n c o u v e r I s l a n d d e v e l o p e d t h r o u g h s u c c e s s i o n f r o m s p r u c e f o r e s t s f o l l o w i n g p o d z o l i z a t i o n and ' h a r d p a n 1 f o r m a t i o n i n t h e o u t w a s h o r a l l u v i a l d e p o s i t s b e n e a t h t h e s e b o g s . Sphagnum bogs a r e much l e s s w i d e s p r e a d i n t h e C o a s t a l W e s t e r n H e m l o c k and C o a s t a l D o u g l a s - F i r z o n e s ( K r a j i n a 1969) i n s o u t h c o a s t a l B r i t i s h C o l u m b i a t h a n i n t h e C o a s t a l C e d a r s - P i n e - Hemlock z o n e , and a r e r e s t r i c t e d m a i n l y t o b a s i n s and a r e a s o f low r e l i e f . The s l o p i n g b l a n k e t bogs common a l o n g t h e n o r t h c o a s t do n o t e x t e n d much f u r t h e r s o u t h t h a n n o r t h e r n V a n c o u v e r I s l a n d . 1 The most e x t e n s i v e bogs i n s o u t h c o a s t a l B r i t i s h C o l u m b i a a r e t h e r a i s e d ( o m b r o p h i l o u s ) bogs on t h e F r a s e r R i v e r Low-l a n d s ( R i g g 1925, O s v a l d 1933) i n t h e C o a s t a l D o u g l a s - F i r z o n e . Hebda (1977) s t u d i e d t h e h i s t o r i c a l d e v e l o p m e n t o f one o f t h e s e , B u r n s Bog, and d e v e l o p e d a m o d e l o f r a i s e d bog d e v e l o p m e n t f o r t h e F r a s e r R i v e r D e l t a . F o l l o w i n g e m ergence a p p r o x i m a t e l y 5,000 y e a r s B.P., t h e p o o r l y d r a i n e d d e l t a i c s e d i m e n t s were c o l o n i z e d by t h e a q u a t i c s S c i r p u s and C a r e x . T h r e e m a j o r s t a g e s f o l l o w e d t h i s i n i t i a l c o l o n i z a t i o n ; a s e d g e - g r a s s p h a s e , a M y r i c a - S p i r e a ( a n d l a t e r Ledum g r o e n - l a n d i c u m ) s h r u b p h a s e and f i n a l l y t h e p r e s e n t - d a y Sphagnum p h a s e . R a i s e d bog d e v e l o p m e n t on t h e F r a s e r D e l t a c o n t r a s t s s h a r p l y w i t h t h e bog w o o d l a n d d e v e l o p m e n t o u t l i n e d i n t h i s s t u d y . 5.5.5 S t u d i e s o f F o r e s t - Bog S u c c e s s i o n i n O t h e r R e g i o n s Much o f t h e a r e a now m a n t l e d i n b l a n k e t bog i n w e s t e r n 161 S c o t l a n d p r o b a b l y o n c e s u p p o r t e d f o r e s t on m i n e r a l s o i l s . R a t c l i f f e (1964) s t a t e s t h a t a b a s a l l a y e r w i t h t r e e r e m a i n s i s a c h a r a c t e r i s t i c f e a t u r e o f some b l a n k e t p e a t s i n w e s t e r n S c o t l a n d . M uskeg d e v e l o p m e n t r e s u l t i n g f r o m f o r e s t p a l u d i f i c a t i o n i s a p p a r e n t l y n o t r e s t r i c t e d t o o c e a n i c r e g i o n s . R o b i n s o n (1976) d e s c r i b e s s i m i l a r s u c c e s s i o n a l s e q u e n c e s i n b l a c k s p r u c e f o r e s t s on t h e c l a y b a n k s o f t h e O p a s a t i k a R i v e r i n t h e O n t a r i o c l a y b e l t , a p p a r e n t l y r e s u l t i n g f r o m Sphagnum e n c r o a c h m e n t f o l l o w i n g t h e f o r m a t i o n o f c l a y p a n s and t h e s u b s e q u e n t i m p e d e n c e o f d r a i n a g e . A c c o r d i n g t o S j o r s ( 1 9 7 6 ) , p a l u d i f i c a t i o n o f f o r e s t s , w o o d l a n d s and p r a i r i e s w h i c h began d u r i n g t h e m i d d l e p a r t o f t h e p o s t g l a c i a l p e r i o d (7 - 4,000 y e a r s B.P.) i s r e s p o n s i b l e f o r much o f t h e w o r l d ' s p r e s e n t - d a y p e a t l a n d a r e a s . I n h i s s t u d y o f G l a c i a l L a k e A g a s s i z p e a t -l a n d s i n M i n n e s o t a , H e i n s e l m a n (1963) p r e s e n t s e v i d e n c e f o r bog e x p a n s i o n f o l l o w i n g f o r e s t d e t e r i o r a t i o n w h i c h began a p p r o x i m a t e l y 4,300 B.P. I n much o f t h e b o r e a l f o r e s t r e g i o n , p e r m a f r o s t p l a y s a s i m i l a r r o l e t o c l a y p a n s and c e m e n t e d ( o r t s t e i n s , p l a c i c ) h o r i z o n s by c r e a t i n g an i m p e r m e a b l e l a y e r w h i c h impedes d r a i n a g e ( R o b i n s o n 1 9 7 6 ) . The i n t e r a c t i o n o f p e r m a f r o s t , p a l u d i f i c a t i o n and f o r e s t s i t e d e g r a d a t i o n i s i l l u s t r a t e d i n a s t u d y o f b l a c k s p r u c e f o r e s t s u c c e s s i o n on n o r t h s l o p e s i n t h e i n t e r i o r o f A l a s k a by H e i l m a n ( 1 9 6 6 , 1 9 6 8 ) . B i r c h - a l d e r f o r e s t s w h i c h d e v e l o p f o l l o w i n g f i r e on t h e s e n o r t h s l o p e s s o o n g i v e r i s e t o b l a c k s p r u c e f o r e s t s . The a c c u m u l a t i o n o f t h i c k Sphagnum c a r p e t s 162 u n d e r t h e s e c l o s e d s p r u c e c a n o p i e s i n s u l a t e t h e m i n e r a l s o i l , a l l o w i n g p e r m a f r o s t t o d e v e l o p c l o s e r t o t h e s u r f a c e . T h e s e Sphagnum l a y e r s impede d r a i n a g e and e f f e c t i v e l y b u r y s o i l n u t r i e n t s t o d e p t h s w here t e m p e r a t u r e s a r e t o o c o l d f o r r o o t g r o w t h and t h e m i n e r a l i z a t i o n o f o r g a n i c n i t r o g e n . The f i n a l s t a g e i n t h i s s u c c e s s i o n a l s e q u e n c e i s a p o o r l y g r o w i n g , c h l o r o t i c b l a c k s p r u c e - Sphagnum ( w a r n s t o r f i a n u m - m a g e l -l a n i c u m ) f o r e s t w i t h 37 - 80 cm o f p e a t a c c u m u l a t i o n . S t e a d i l y d e c l i n i n g n u t r i e n t a v a i l a b i l i t y t h r o u g h t h e s e s u c c e s s i o n a l s t a g e s was d e m o n s t r a t e d u s i n g f o l i a r a n a l y s i s . C h a n g e s i n m i c r o b i a l p o p u l a t i o n s and m e t a b o l i c a c t i v i t y i n t h e f o r e s t f l o o r s o f s i t e s u n d e r g o i n g p a l u d i f i c a t i o n h a v e b e e n d e m o n s t r a t e d by M a n n e r k o s k i ( 1 9 7 2 ) . H i s r e s u l t s showed d e c l i n i n g b a c t e r i a l and f u n g a l p o p u l a t i o n s t o g e t h e r w i t h l o w e r c a t a l a s e a c t i v i t y , CO2 r e l e a s e , O2 c o n s u m p t i o n and r e s p i r -a t o r y q u o t i e n t s i n t h e humus h o r i z o n s o f t h e s i t e s w h i c h were i n more a d v a n c e d s t a g e s o f p a l u d i f i c a t i o n . 5.5.6 P a l e o c l i m a t i c I n t e r p r e t a t i o n s T h u s f a r , I have i n t e r p r e t e d t h e s u c c e s s i o n a l s e q u e n c e p o r t r a y e d i n t h e p o l l e n p r o f i l e m a i n l y i n t e r m s o f l o c a l s i t e f a c t o r s and s o i l - f o r m i n g p r o c e s s e s . I w i l l now e x a m i n e t h e i n f l u e n c e t h a t l a r g e - s c a l e p a s t c l i m a t i c c h a n g e s may have had o n t h e s e p a t t e r n s o f v e g e t a t i o n and s o i l d e v e l o p m e n t . H e u s s e r (1960) d a t e s t h e h y p s i t h e r m a l i n t e r v a l , when c l i m a t i c c o n d i t i o n s w e r e warmer and d r i e r t h a n a t p r e s e n t , b e t w e e n 8,200 and 3,600 y e a r s B.P. a l o n g n o r t h c o a s t a l B r i t i s h C o l u m b i a . More r e c e n t l y , Mathewes and H e u s s e r (1981) u s e d 163 t r a n s f e r f u n c t i o n s , d e r i v e d from r e g r e s s i o n s of modern p o l l e n s u r f a c e samples and c l i m a t i c data, to r e c o n s t r u c t a 12,000 year r e c o r d of temperature and p r e c i p i t a t i o n f o r southwestern B r i t i s h Columbia. They d e f i n e an " E a r l y Holocene Xerothermal I n t e r v a l " as a p e r i o d of maximum temperature and minimum p r e c i p i t a t i o n o c c u r r i n g between 10,000 and 7,500 years B.P. T h i s was followed by a marked c o o l i n g from 7,000 to 6,000 years B.P. as w e l l as an i n c r e a s e i n p r e c i p i t a t i o n from 7,500 to 6,000 years B.P. to the present. Zone I (dated at 8710 years B.P.) of the Hayes Mtn. core corresponds with t h i s " E a r l y Holocene Xerothermal I n t e r v a l " and the t r a n s i t i o n from the P i c e a - Alnus - L y s i c h i t o n - P o l y -podiaceae assemblage of Zone II to the Pinus - Cupressaceae -Tsuga - Cyperaceae - Sphagnum assemblage of Zone I I I may correspond (although there i s no C-14 date) with the c o o l e r and wetter trend between 7,000 and 6,000 years B.P. The high a b s o l u t e p o l l e n / s p o r e c o n c e n t r a t i o n s of Zones I and II could be a t t r i b u t e d to h i g h e r p o l l e n p r o d u c t i v i t y and higher r a t e s of o r g a n i c matter decomposition a s s o c i a t e d with t h i s warmer, d r i e r c l i m a t i c i n t e r v a l . Durno (1961) s t r e s s e s the importance of c l i m a t e i n c o n t r o l l i n g the r a t e of peat accumulation and, by combining C-14 d a t i n g with p o l l e n s p e c t r a , has estimated peat accumulation r a t e s f o r the B r i t i s h I s l e s which correspond to major c l i m a t i c p e r i o d s . The e f f e c t of c l i m a t i c f l u c t u a t i o n s on v e g e t a t i o n and s o i l development i s expected to be more pronounced wherever c l i m a t i c c o n d i t i o n s are more l i m i t i n g , such as along north 164 c o a s t a l B r i t i s h C o l u m b i a and S o u t h e a s t A l a s k a . The c o o l i n g and w e t t i n g t r e n d b e t w e e n 7,000 and 6,000 y e a r s B.P., r e p o r t e d by Mathewes and H e u s s e r (1981) f o r s o u t h w e s t e r n B r i t i s h C o l u m b i a , may have b e e n s u f f i c i e n t t o t r i g g e r s u c c e s s i o n f r o m p r o d u c t i v e f o r e s t t o s c r u b a l o n g t h e n o r t h c o a s t , i n t u r n s e t t i n g t h e s t a g e f o r t h e w i d e s p r e a d d e v e l o p m e n t o f b l a n k e t bog c o m m u n i t i e s . A p p l i c a t i o n o f t r a n s f e r f u n c t i o n s , s i m i l a r t o t h o s e u s e d by Mathewes' and H e u s s e r ( 1 9 8 1 ) , t o g e t h e r w i t h more i n t e n s i v e C-14 d a t i n g o f n o r t h c o a s t a l p o l l e n p r o f i l e s w o u l d be w o r t h w h i l e , g i v e n t h e s e g e n e r a l c o r r e l a t i o n s b e t w e e n t h e Hayes M t n . p r o f i l e and t h e p a l e o c l i m a t i c r e c o n s t r u c t i o n f r o m s o u t h c o a s t a l B r i t i s h C o l u m b i a . 165 CHAPTER 6 ECOSYSTEM DYNAMICS T h i s chapter summarizes the s u c c e s s i o n a l r e l a t i o n s h i p s between bog and f o r e s t ecosystems w i t h i n the study area i n a co n c l u d i n g d i s c u s s i o n of ecosystem dynamics. The problem of d e f i n i n g the c l i m a t i c climax ecosystem i n the blanket bog -upland f o r e s t complex of the north P a c i f i c coast i s a l s o addressed. 6.1 C o n t r a s t i n g Evidence For Bog/Forest S u c c e s s i o n P a l y n o l o g i c a l evidence from t h i s study, as w e l l as e m p i r i c a l evidence from present-day ecosystems i n Southeast A l a s k a (Daschnowski-Stokes 1941; Zach 1950; Neiland 1971; U g o l i n i and Mann 1979) f o r p a l u d i f i c a t i o n or ' b o g i f i c a t i o n ' of p r o d u c t i v e f o r e s t , has already been summarized i n Chapter 5 (pages 156 - 158). Some f l o r i s t i c and edaphic c h a r a c t e r i s t i c s of community types d e s c r i b e d i n Chapter 4 may be i n t e r p r e t e d as evidence f o r a s i m i l a r s u c c e s s i o n a l sequence. The s c a t t e r e d occurrence of l a r g e , dead snags, as w e l l as s e v e r a l upland f o r e s t s p e c i e s such as P i c e a s i t c h e n s i s , Abies  a m a b i l i s , Streptopus roseus, Rubus pedatus and P e l l i a  neesiana, i n bog woodlands developed on F o l i s o l s or shallow o r g a n i c s o i l s (e.g., r e l e v e 15; page 78) suggest that the bog woodlands may r e p r e s e n t degraded upland f o r e s t communities. Where the cedar - cypress - hemlock zonal upland f o r e s t community type occurs on F o l i s o l s or peaty and f o l i s o l i c phases of m i n e r a l s o i l s , and s c a t t e r e d bog or bog woodland 166 e l e m e n t s s u c h as Ledum g r o e n l a n d i c u m , K a l m i a m i c r o p h y 1 1 a , F a u r i a c r i s t a - g a l l i , C a r e x s p p . , Sphagnum s p p . and P l e u r o z ium  s c h r e b e r i o c c u r , i n c i p i e n t bog d e v e l o p m e n t i s s u g g e s t e d . A l t h o u g h n a t u r a l f i r e s p l a y an i m p o r t a n t r o l e i n b o r e a l f o r e s t s by c h e c k i n g t h e p r o c e s s o f p a l u d i f i c a t i o n and s u c c e s s i o n f r o m f o r e s t t o muskeg ( H e i l m a n 1966, 1 9 6 8 ) , f i r e i s r a r e a l o n g t h e n o r t h c o a s t . S o i l d i s t u r b a n c e c a u s e d by n a t u r a l e v e n t s s u c h as mass w a s t i n g , blowdown, o r f l o o d i n g a p p e a r t o be i m p o r t a n t i n m a i n t a i n i n g p r o d u c t i v e f o r e s t s on t h e s t e e p e r m o u n t a i n s l o p e s and a c t i v e f l o o d p l a i n s . R e c a l l t h a t t h e c e d a r - c y p r e s s - h e m l o c k z o n a l f o r e s t c o m m u n i t y t y p e may a l s o d e v e l o p on deep o r g a n i c d e p o s i t s ( H u m i s o l s ) w i t h deep s u r f a c e a c c u m u l a t i o n s o f f o r e s t humus ( p a g e s 87 - 89; F i g . 4 0 ) . S u c h an o r g a n i c s e q u e n c e i n d i c a t e s t h a t t h e u n d e r l y i n g p e a t a c c u m u l a t e d i n a bog o r f e n e n v i r o n -ment b u t , t h r o u g h c h a n g e s i n d r a i n a g e , o r a g r a d u a l d r y i n g o u t o f t h e s u r f a c e l a y e r s ( p o s s i b l y r e s u l t i n g f r o m c l i m a t i c f l u c t u a t i o n s ) , c o n d i t i o n s s u b s e q u e n t l y became s u i t a b l e f o r i n v a s i o n by f o r e s t . C a r b o n - 1 4 d a t i n g o f one s u c h p r o f i l e i n d i c a t e d t h a t t h i s s u c c e s s i o n o c c u r r e d w i t h i n t h e p a s t 2,400 y e a r s ( s e e page 8 9 ) . I n t h i s as w e l l as o t h e r c a s e s , t h e u n d e r l y i n g o r g a n i c d e p o s i t had a c c u m u l a t e d i n a s h e d d i n g p o s i t i o n n e a r a r i d g e t o p , r a t h e r t h a n i n a d e p r e s s i o n o r b a s i n a s i n more c l a s s i c a l h y d r a r c h s u c c e s s i o n f r o m b a s i n bog t o f o r e s t . S u c h " u n o r t h o d o x " s u c c e s s i o n a l t r e n d s e m p h a s i z e t h e i m p o r t a n c e o f c l i m a t i c f a c t o r s as w e l l as t o p o g r a p h y i n s o i l d e v e l o p m e n t . 167 S t e p h e n s e t a l . ( 1 9 70) d e s c r i b e f o r e s t e d o r g a n i c s o i l s i n S o u t h e a s t A l a s k a t h a t have d e v e l o p e d t h r o u g h s i m i l a r s u c c e s s i o n a l s e q u e n c e s f r o m w e t l a n d t o f o r e s t . A c c o r d i n g t o t h e s e a u t h o r s t h i s s o i l t y p e c o v e r s 1,600,000 a c r e s ( A p p r o x i -m a t e l y 647,770 h e c t a r e s ) o r 10% o f t h e T o n g a s s N a t i o n a l F o r e s t , and p r o v i d e s e v i d e n c e f o r t h e d i m i n i s h e d e x t e n t o f p r e s e n t day muskeg i n S o u t h e a s t A l a s k a . 6.2 The S u c c e s s i o n Complex and t h e C l i m a t i c C l i m a x Ecosystem The f l o r i s t i c , e d a p h i c and p a l y n o l o g i c a l e v i d e n c e w h i c h h a s b e e n p r e s e n t e d h e r e and i n t h e s t u d i e s c i t e d f r o m S o u t h -e a s t A l a s k a c l e a r l y i l l u s t r a t e s t h a t t h e r e g i o n a l v e g e t a t i o n r e p r e s e n t s a s u c c e s s i o n a l c o m p l e x o r a d y n a m i c c o m p l e x o f e c o -s y s t e m s t h a t a r e i n c o n t i n u a l f l u x . G e n e r a l i z a t i o n s o f u n i -d i r e c t i o n a l s u c c e s s i o n a l t r e n d s c a n n o t accommodate t h e c o n t r a s t i n g e v i d e n c e f o r bog d e v e l o p m e n t f r o m f o r e s t as w e l l a s f o r e s t d e v e l o p m e n t f r o m bog. I t i s p r o b a b l e t h a t a l t h o u g h c l i m a t i c f l u c t u a t i o n i s a p r i m a r y c o n t r o l l i n g f a c t o r i n t h e d i r e c t i o n i n w h i c h s u c c e s s i o n w i l l p r o c e e d , o t h e r f a c t o r s s u c h a s t o p o g r a p h y , l a n d f o r m , n u t r i e n t r e g i m e and l o c a l d r a i n a g e p a t t e r n s a r e i m p o r t a n t s e c o n d a r y f a c t o r s . The t r a d i t i o n a l c o n c e p t o f t h e c l i m a t i c c l i m a x as a s t a b l e and s e l f - m a i n t a i n i n g e c o s y s t e m w h i c h e x p r e s s e s t h e m a c r o c l i m a t e o f t h e r e g i o n i s d i f f i c u l t t o a p p l y a l o n g t h e n o r t h c o a s t , w here t h e c o n t r o l l i n g e f f e c t s o f c l i m a t i c and e d a p h i c f a c t o r s a r e s o c l o s e l y i n t e r - r e l a t e d . M o s t e x i s t i n g c l i m a x e c o s y s t e m s a r e a p r o d u c t o f a s u c c e s s i o n w h i c h has been 168 allowed to proceed i n the absence of f i r e or other major d i s t u r b a n c e s f o r perhaps thousands of years, p o s s i b l y s i n c e e a r l y post g l a c i a l time. T a n s l e y (1920) and Whittaker (1974) s t r e s s the importance of r e g i o n a l prevalence i n d e f i n i n g the c l i m a t i c climax eco-system. I f we c o n s i d e r the ecosystem most common on the mid-slo p e p o s i t i o n s of moderate s l o p e s as best e x p r e s s i n g the macroclimate, then the cedar - cypress - hemlock - Vaccinium -Cornus - R h y t i d i a d e l p h u s upland f o r e s t community type which dominates on such s i t e s i n the study area should be considered the c l i m a t i c climax ecosystem. On the extreme outer coast of the mainland, on many of the o f f - s h o r e i s l a n d s and on p a r t s of the west coast of the Queen C h a r l o t t e I s l a n d s , however, s l o p i n g bog or bog woodland- predominates on such s i t e s and should probably be considered the c l i m a t i c climax ecosystem i n these p o r t i o n s of the CCPH zone. The r e c o g n i t i o n of the dynamics of these c o a s t a l eco-systems has important p r a c t i c a l i m p l i c a t i o n s . An under-s t a n d i n g of the s u c c e s s i o n a l processes c o n t r o l l i n g f o r e s t development or degradation, and the f a c t o r s i n f l u e n c i n g these p r o c e s s e s , becomes e s s e n t i a l i f we are i n t e r e s t e d i n manipu-l a t i n g the ecosystem to improve t r e e p r o d u c t i v i t y . 169 CONCLUSIONS Mathematical o r d i n a t i o n techniques such as p o l a r o r d i -n a t i o n , p r i n c i p a l components a n a l y s i s and r e c i p r o c a l averaging are u s e f u l i n o b j e c t i v e l y e v a l u a t i n g a s u b j e c t i v e c l a s s i f i -c a t i o n d e r i v e d by t a b u l a r comparison, and i n v i s u a l l y d i s p l a y i n g r e l a t i o n s h i p s w i t h i n and among community types. However, the l i m i t a t i o n s that s u b j e c t i v e sampling p l a c e s on the i n t e r p r e t a t i o n s must be recogn i z e d . The s i x community types d e s c r i b e d along the fo u r study t r a n s e c t s r e f l e c t the combined i n f l u e n c e of s e v e r a l e n v i r o n -mental f a c t o r s , i n c l u d i n g the o v e r r i d i n g e f f e c t of a c o o l , perhumid, r a i n y , hyperoceanic c l i m a t e ; s o i l drainage, through i t s i n f l u e n c e on s o i l water regimes and n u t r i e n t turnover; the predominant s o i l - f o r m i n g processes of org a n i c matter accumulation (both as f o r e s t humus and wetland peat) and p o d z o l i z a t i o n ; landform and topography through t h e i r e f f e c t on s o i l drainage; and bedrock geology through i t s e f f e c t on s o i l n u t r i e n t regime and drainage. S i m i l a r ecosystems to those d e s c r i b e d i n the P r i n c e Rupert study area occur i n Southeast A l a s k a , on the Queen C h a r l o t t e I s l a n d s and elsewhere on the outer coast of B r i t i s h Columbia north of Vancouver I s l a n d . T h i s blanket bog-upland f o r e s t complex of the north P a c i f i c coast shows many r e l a t i o n -s h i p s with the v e g e t a t i o n and s o i l s of the western B r i t i s h I s l e s and, to a l e s s e r extent, with e a s t e r n Newfoundland. P a l y n o l o g i c a l , edaphic and f l o r i s t i c evidence f o r the 170 development of muskeg from p r o d u c t i v e f o r e s t c o n t r a s t s with edaphic evidence f o r s u c c e s s i o n from wetland to f o r e s t i n the study area. I i n t e r p r e t the v e g e t a t i o n and s o i l s as a dynamic complex of s u c c e s s i o n a l l y r e l a t e d ecosystems. C l i m a t i c f l u c t u a t i o n s as w e l l as l o c a l edaphic f a c t o r s are l i k e l y important i n c o n t r o l l i n g the d i r e c t i o n of s u c c e s s i o n . Some ecosystems such as p r o d u c t i v e f o r e s t s on steep c o l l u v i a l s l o p e s are maintained by f r e e drainage and p e r i o d i c d i s t u r -bance caused by w i n d f a l l and c o l l u v i a l a c t i o n . R e g i o n a l p r e v a l e n c e appears t o be the best c r i t e r i o n t o use i n d e f i n i n g the c l i m a t i c climax ecosystem along the north P a c i f i c coast, where s u c c e s s i o n has been allowed t o proceed without d i s t u r b a n c e f o r thousands of years. 171 LITERATURE CITED Annas, A.M. 1977. Boreal ecosystems of the F o r t Nelson area of n o r t h e a s t e r n B r i t i s h Columbia. Ph.D. t h e s i s . U n i v e r s i t y of B r i t i s h Columbia, Vancouver, B.C., Canada. Annas, R.M., and R. Coupe, e d i t o r s . 1979. 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E d a t o p i c G r i d o f K r a j i n a (1969) A B C D E 0 1 2 3 4 5 6 7 8 M o i s t u r e Regime ( H y g r o t o p e s ) 0 v e r y x e r i c 1 x e r i c 2 s u b x e r i c 3 sub i f i es i c 4 m e s i c 5 s u b h y g r i c 6 h y g i " i c 7 s u b h y d r i c 8 h y d r i c N u t r i e n t Regime ( T r o p h o t o p e s ) A o l i g o t r o p h i c B s u b m e s o t r o p h i c C m e s o t r o p h i c D p e r m e s o t r o p h i c E s u b e u t r o p h i c t o e u t r o p h i c 184A APPENDIX II F i e l d Forms Used i n the Releve Sampling \ Appendix II. F i e l d forms used in the releve sampling. SITE DESCRIPTION FORM " NS J0Q9 -79 Tentative Veg. Name ^ . / • c ' - t f w ^ u i - /l&Kiim* - Ccx-n*. ^ - RU^tiiArlfib* - _ C , ^ j / > ^ r . ^ f>tf . ' ^ This fonn not to be copied; refer to definitions manual for al l terms used PAGE ^ OF ^> parameters which must be f i l l e d in are c i rc led ; an attempt should be made to f i l l in all other parameters. (TT) Project Identiftcaton fJCCf-j - A 1 S < © Date (Y/H/D) 0 NTS Sheet \OZ^~l<Z\J © Longitude £H ° \k ' 36 " \^~* 7. Location I4tt»j<: MT . • OlUtf Ik lAtan P/U^J. Qujfi+rf' 8. Photo number and Co-ord. f l ight l ine phot6"*no. (97) Aspect l^ff* * (lo) Slope lH~-XO percent (fT) Elevat ion. Plot number 0 Latitude 13,0' vg' 'MS"" ) W 12. Terrain c lass i f icat ion Texture Genet i c Material Super. Hod. Surface Exp. Hod. Proc. s 1 T E © V nap Unit 13. Physiographic Region /I Zone/Subzone 15. Landscape Type c lass i f i ca t ion system c lass i f i cat ion system 16. Vegetation Type 17. Soil Class i f icat ion c lass i f i cat ion system phase year " "T^f-iK filial family tex. Plot representing Ci\j~C' U n v " r 4 u Site position macro (2th) Site position meso (2K) Site surface shape (22^ Microtopography a. apex b. face c. upper slope (3t) middle slope e. lower slope f. valley f loor g. plain 23. Slope length 'aO crest Yjy upper slope c. middle slope d. lower slope e. toe f. depression g. level MlQ. /VYV (25^ Site position diagram {refer to data form no. concave, convex I^ . £.6 COI-AH-straight m 24 p-Slope length smooth s i ight ly mounded moderately mounded strongly mounded severly mounded extremely mounded ultra ' mounded /k«4. / i ' . j o j i U ( W ' ^ photo rol l no.7}'/1" J£K,lu Photo no. ^ W scale \Q0, Exposure type a. b. d. w1 nd Insolation frost cold air drainage saltspray atmospheric toxicity not applicable other 0/9 < 27. Ecological Moisture Regime 28. Nutrient Regime . .^9. Soil Moisture Subclass 30. a. very xeric a. b. xeric c. subxeric © d. submes1c cr: meslc e. 3 subhygrlc f. hygric h. subhydric i . hydrlc oligotrophic a. xeric submesotrophlc b. arid a mesotrophic c. subarid b permesotrophic d. semiarld c eutrophic e. subhumid d hypereutrophic f. humid e perhumld © subaquic i. aquic j . peraqulc Soil Temperature Class extremely cold very cold cold cool mild (3y Pervlousness a. rapidly b. moderately fc$ slowly Soil drainage a. very rapidly - b. rapidly c. well d. mod. well (e7) Imperfectly T. poorly g. very poorly ) Oepth to (cm) (c i rc le - 1f absent a, water table lcA«.fc-l /L-y\ £> b. rooting (effective) IO e Water 34. Flood hazard present a. frequent and Irregular absent b. frequent c. may be expected j k rare (eJ no hazard-root restr ict ing layer frozen layer _£) 37. 38. e. bedrock JO - D C O w . f. carbonate g. sa l in i ty Bedrock type {listed In order of dominance) _££ 1 ^ L _ Bedrock structure Coarse fragment Hthology a. type (1n order of dominance) Successional Status 40. Present Stage: PS, Y5, MS, OS, YC, YEC YCC, MC.(MCC) MEC, DC, NV Expected climax -Rate of succession ® slow b. moderate c. fast Veg. Plot Dim. ^ Q V Q O (metres) @ Surface Substrate _>? mixed c. none Cause of Stand Establishment Humus Form Class. WUXVYNA/\v*-«-\. SUBSTRATE % COVER • Organic Matter Decaying Wood /o-nT Bedrock' Cobbles and Stones Mi neral Soil Water Total 1001 44. Prof i le Status 45. Prof i le Deviation 46. Soil Mapping Unit a. modal a. solum thickness a. series b. variant b. colour b. family c. taxadjunct c. texture c. associate d. undecided d. drainage d. association e. other e. catena f. none f. 9-h. 1. complex land system land type other 47. 49. Soil name _ Prof i le No. ~Tvii)h foil it I 48. Associated soil Notes on Site Description (so) Project Coordinator (si) Agenc y (5y Type of Soil Sample (A ) Sampled ^ 1. Chemical a. fu l l b. partial B. No Sample Veg. Sampling Tech. . Physical a. fu l l b. partial 187 VEGETATION DESCRIPTION FORM PROJECT l.D. fl)CC H * M^C, PLOT NO. £7. /\Q\Q\°l\ PAGE T - OF V FLORISTIC LIST' STRATUM \ NOTES • c OJ c C i/> '_=' <U S t a. c o •— c ZZ O CM «* r— m 2 e r- 3 ca Si -J l/l CM CO r— __ -J _. O JZ t— m __ t-OJ <_> H *'c in a> O QJ _C to o V. +J 1/1 specify entries here (U-Height of top of stratafm) .2 / Number of dead snags Total percentage cover of each layer 20 77/ *f 7^ SPECIES LIST X I } -1./ fa J ' i I-1 7-1 U.- .v. •••:•/.•;. f-.3.2 •J, / • V ti y.a KM*— y.z. 1-7, a.5. 7,2 <...,-. 1.1 : ., . • w,. A'? # / • . .-, ......V liUr.:.. . -,-oW /. 2. r. ? A", y ? T . / -S3 W o IMI Wo I4s-Co lit (As IIS Comments on Vegetation Characteristics FLORISTIC LIST 51' 4 7? 11 3.2 Vi 1 7 -v U2 -r.-H 3 / 3 . • | » 7 * c 73 ^ •/»-, Afirust th* 121 ¥4 c : t ? " A HORIZON DESIG. aspect COLOUR 1 c X COLOUR 1 aspect COLOUR 2 COLOUR 2 3 o -n co j MOTTLES j MOTTLES O -o o "n o j MOTTLES aspect j MOTTLES COLOUR 1 j MOTTLES o co — o o o i/> § j MOTTLES 3> -O Tl -S > ROOTS I o 3 -n < ISt N n> ROOTS I 7 3 0 K o -i ROOTS I X o ROOTS I 3» -c -n «; > cr 30 O O o 3 "n < V) re » O I < O rs> 3 m — reaction rc method 1 I i !• c-\ m ° ° C ) 0-31 i } r HORIZON DESIG. i j >"„ K o OEPTH 1 t ^ \ o "\ \ o .-: -i "HICKNESS :M IE o" >s -'•-: • X "HICKNESS :M j ; X Dist . HORIZON BOUNDARY V _ t Form HORIZON BOUNDARY 0 COARSE FRAGMENT DESCRIPTION o Gravel <7.5cm COARSE FRAGMENT DESCRIPTION type Gravel <7.5cm COARSE FRAGMENT DESCRIPTION a* Cobbles 7.5-25cm COARSE FRAGMENT DESCRIPTION type Cobbles 7.5-25cm COARSE FRAGMENT DESCRIPTION « Stones COARSE FRAGMENT DESCRIPTION type Stones COARSE FRAGMENT DESCRIPTION r-FIELD TEXTURE GRADE PRIMARY STRUCTURE CLASS PRIMARY STRUCTURE KIND PRIMARY STRUCTURE KIND HOD. PRIMARY STRUCTURE 'GRADE SECONDARY STRUCTURE CLASS SECONDARY STRUCTURE KIND SECONDARY STRUCTURE KIND MOO. SECONDARY STRUCTURE J U N H H dry o o moist (/> wet Ln plast. —1 S S < ^ SOIL TEHP. •c TT •ON ib id ~>%W- H7~>(1 ioaroad wucy NOiidiaosaa nios 681 HORIZON DESIG. PORES HOR, POR. s H H CLAY FILMS EFFERVESC. SECONDARY CARBONATE DESCRIPTION SAL CONCf ETK NS CEt 1EN1 r. LAB SAMPLE NO. . Ab. Si ze Ori Dist Cont Mor Type Freq Thi Loc Deg. Reag. Cont Ab. Size Shape Consist. U M S Kind Ab. Size Loc Shape Cc lour c <a c; «t at V cr. <u o J Extent V F P A V F M C V H 0 R IN EX HTX CO DC S D C V I T F C M CS TN MTK TK VTK VH U M S F C M F M C Str. Spo o >. u o F C M F M . c-S 0 I p | Aspect J Extent V H R R 0 I | Aspect J Extent L / / j / )P-f S i / 1 7— 1 / 1 / 1 1 1 1 » V*' i i i i \A I I i i r I •?• \j 1 ! ! i i ! i i i i s \ < i i i I / I i I I I r 1 i ) , H A M / ' I | i \X\ 11 - — lit M s / i / i 1 jr I I i l t _ _ | 1 1 1—i / 1 1 r H — 1 — \ll n / i i / i A \ \\\\ y ' i > I I I..v1 1 I I _- | I 1 1 | | l?.H H — i — i — i — • • • i — i — i i i i i i i • • i i i > I I i I I I i i i i j [ 1 1 1 1 I I 1 1 1 1 1 1 i I I I i i [-. ; | | i j 1 1 I I I I 1 1 1 1 I I I 1 1 1 1 — 1 . — — i — i — i i — i — i — i I I i I , I i t i i i * i i i i t i I I i i i 1 1 I I ' I f f ' 1 1 1 1 1 1: . 1 | | 1 |—1| 1 T — H 1 1 1 1 •• I.S. I l 1 1 l 1 i 1 1 1 i I I i I I I 1 r l i i i I I I I 1 • t 1 1 1 1 l l t 1 HORIZON DESIG. ORGANIC MATERIAL DESCRIPTION Material Composition H 1 IK Hoody Material Hard S SH H VH JUL ita. VM Pyro. Index von Post Scale 1 General Comments on Soil Characteristics. • o 25 SCHEMATIC SOIL PROFILE 10 0 25 100 I HUMUS FORM PROFILE DESCRIPTION FORM '-°.*fttH-#Z&— p.ot N O _ I 2 I - . • • surveyor Hi lSQ«»\</~ ?oge f nf _ Hor. •Des. ! Depth Tnxrfness N o iBO'Jncory Min Max moisture Bulk Density Structure • Consistence Texture Boots Dist ronr Mineral Organic M ineral Organic Mineral Organic A D - Si Or 3s t L 0 / '1-s w u -. Y f f-< /.si i >-/'brie. A •A f 1 H-AM '7.2. IS- foj 3 M p>*H ' - f> M. 3 a /7.? o '4 ft F. F 1 I : - i 1 He /7< 3 i F rt ;*© 3 3 Hor. Des. Color Non-conforming Materials Soil Fauna Soil Microflora X-Sectkw. . ' O.-C'.-.'ir.gf -V' AS Mun. Not. L . j .<••; . £• 1 , 1 r u , 7 <3»r-<~, fteirU o-fpK"-7 • /• Hi, 7 • ^ ' // Iv ' • : v_ 1 Humus Fa'm; . |. Comments! 1 9 1 A MENSURATION/WILDLIFE DESCRIPTION PROJECT I.D. /JftU -/ASc. PLOT NO. SURVEYOR Q.^m.ftV>>^- J ft S/VYv^ fc-FORM T OF MtOJf CT *!0T Kb I |B| 1 TRU HO. IB|f« TOTAL •T|I«|I. V I c I • sojai O.t. B. «T«T"T» MTH. RCM-«*S 3 3 » o H S ST s 1 i 11 i! ii 8 . i i I 2 Oil (PP 'a a 0<l I6P I 5 1 l op /• <AH w* 1/1/72 /. Y /try. t °K J Vrto d /a \/\o /0\7 JW'C C\ \s\i\c i G\t 1 V>',c c\ 1 7i7T? ?> 0\t /ay> ,? \1\2V 1 i ; ! 1 I i --1 i * I 1 1 i - ----- -1 i t 1 I i t 1 1 7 1 1 i T 1 i r I l i t 1 i r 1 1 I t 1 i ; ! I i i 1 i 1 1 i i 1 1 T I i i 1 i ; I 1 1 ? 1 1 T | 1 I 7 3 TREE NO. TOTAL HEIGHT SPEC. D.B.H. c / c TOTAL AGE AGE CALCULATIONS HEIGHT CALCULATIONS COUNT. AGE B.H. COR. SUP. TOP BOT. TOTAL S.D. SLOPE H.D. HT. COR. <"i C/, 1 & 2 ,?,< ?77 • c s? ?l \9.l /-if r,a 2 1 ,s>3 3,2,2 VI °/ 7? O 17.1 IS* C ,r,, ^ l.f>T? • 7H-7T7 / L 4 if '5.0 - 1 h'c N.3 ft K7 ,3 1. 1 m IZ hh ff,1 IS'1 ft.l / c. , 7i6f/ 34 o . <? lo 23 '33 -'3 IS I I , T 1 , , f t 1 i f 1 , , ? • i I i , t 1 i . T 1 1 . t 1 I . t • • 1 • 1 , , T • i i t t , , T ,•••? 1 , , t • i t 1 , ? . , t • i , , t t , . f i i 1 9 1B UNGULATE SPECIES QUADRAT 1 2 3 4 S UNGULATE PELLET GROUP COUNTS Sightings Wildl i fe Rating . Season of use Domestic Range Condition: 1. Excellent 2. Good 3. Fair 4. Poor Miscellaneous Comments on Wildlife and Domestic Range Characteristics: General Miscellaneous Comments: APPENDIX III Cover-Abundance S c a l e , Vigour Scale and S t r a t a D e s i g n a t i o n s Used i n the Releve Sampling 192 Appendix I I I - Cover-abundance s c a l e , v i g o u r s c a l e and s t r a t a d e s i g n a t i o n s used i n the r e l e v e sampling. Domin - K r a j i n a Cover - Abundance S c a l e + s o l i t a r y , with ins i g n i f i c a n t cover 1 seldom, with i n s i g n i f i c a n t cover 2 very • s c a t t e r e d ' with cover • 1% 3 s c a t t e r e d , with cover of 1 - 4% 4 any number with cover of 5 - 9% 5 any number with cover of 10 - 24% 6 any number with cover of 25 - 32% 7 any number with cover of 33 - 49% 8 any number with cover of 50 - 75% 9 any number with cover of 76 - 99% 10 any number with complete cover (100 Vigour S c a l e 3 ' e x c e l l e n t 2 good 1 f a i r + poor 0 dead 193 Appendix I I I (cont'd) D e s c r i p t i o n of the twelve s t r a t a evaluated i n each r e l e v e . Stratum D e s c r i p t i o n V - Veteran t r e e s s i g n i f i c a n t l y o l d e r than the t r e e s o f the main canopy. A l - Dominant t r e e s extending above the main canopy (upper 5% o f h e i g h t d i s t r i b u t i o n p o p u l a t i o n ) A2 - Main canopy t r e e s A3 - Suppressed t r e e s not reaching the average l e v e l of the main canopy. B l - T a l l shrubs and t r e e r e g e n e r a t i o n between 2 and 10 m. h i g h . B2 - Low shrubs and t r e e r e g e n e r a t i o n l e s s than 2 m h i g h . C - Herbaceous and semi-woody s p e c i e s . DH - Bryophytes, l i c h e n s and s e e d l i n g s (<15 cm) growing on humus. DW - Bryophytes, l i c h e n s and s e e d l i n g s growing on r o t t i n g wood ( e p i x y l i c ) . DE - Epiphytes. DR - Species growing on rocks and boulders ( e p i l i t h i c ) DS - Species growing on bare s o i l ( e p i s o l i c ) 19 3A APPENDIX IV L i s t of Species Recorded i n the Study Area 194 Appendix IV - L i s t of s p e c i e s recorded i n the study a r e a 1 V a s c u l a r Plants Abies a m a b i l i s (Dougl. ex Loud.)Forbes ( a m a b i l i s f i r ) Adiantum pedatum L. subsp. a l e u t i c u m (Rupr.)Calder and T a y l o r (northern maidenhair fern) A g r o s t i s a e q u i v a l v i s ( T r i n . ) T r i n . (Alaska bent grass) Alnus rubra Bong, (red a l d e r ) Andromeda p o l i f o l i a L. subsp. p o l i f o l i a (bog rosemary) Athyrium f i l i x - f e m i n a (L.)Roth subsp. cycloglosum (Rupr.) C h r i s t e n s , i n H u l t . (common lady fern) Blechnum s p i c a n t (L.)Roth (deerfern) Calamagrostis canadensis (Michx.)Beauv. ( b l u e j o i n t small reedgrass) Calamagrostis nutkaensis ( P r e s l ) S t e u d . ( P a c i f i c reedgrass) C a l t h a l e p t o s e p a l a (DC.)var. b i f l o r a (DC.)G. Lawson (white marsh-marigold) Carex anthoxanthea P r e s l (sedge) Carex deweyana Schw. (Dewey's sedge) Carex l a e v i c u l m i s Meinsh. (smooth-stemmed sedge) Carex l i v i d a (Wahl.)Willd. (pale sedge) Carex m e r t e n s i i P r e s c o t t i n Bong, subsp. m e r t e n s i i (merten's sedge) Carex p a u c i f l o r a L i g h t f . (few-flowered sedge) Carex p l u r i f l o r a Hulten (many-flowered sedge) Carex r o s t r a t a Stokes i n Withering (beaked sedge) Carex s i t c h e n s i s P r e s c o t t i n Bong. ( S i t k a sedge) Carex s t y l o s a C.A. Mey. ( l o n g - s t y l e d sedge) Chamaecyparis n o o t k a t e n s i s (D. Don)Spach (yellow-cedar or cypress) C i r c a e a a l p i n a L. ( a l p i n e enchanters n i g h t shade) Cladathamnus p y r o l i f l o r u s Bong, (copperbush) C l i n t o n i a u n i f l o r a (Schult.)Kunth (blue-bead c l i n t o n i a ) C o p t i s a s p l e n i i f o l i a S a l i s b . (spleenwort-leaved goldthread) C o p t i s t r i f o l i a ( L . ) S a l i s b . ( t h r e e - l e a v e d goldthread) Cornus u n a l a s c h k e n s i s Ledeb. (bunchberry) Dodecatheon j e f f r e y i van Houtte subsp. j e f f r e y i ( J e f f r e y ' s s hooting s t a r ) Drosera a n g l i c a Huds. (g r e a t sundew) Drosera r o t u n d i f o l i a L. (round-leaved sundew) D r y o p t e r i s a s s i m i l i s S. Walker (spiny-wood fern) Empetrum nigrum L. (crowberry) E r i g e r o n p e r i g r i n u s (Pursh)Greene (subalpine fleabane) Eriophorum a n g u s t i f o l i u m Honck. (narrow-leaved c o t t o n grass) 195 Appendix IV - Cont'd F a u r i a c r i s t a - g a l l i (Menzies ex Hook.)Makino subsp. c r i s t a - g a l l i (deer cabbage) Galium boreale L. (northern bedstraw) G a u l t h e r i a s h a l l o n Pursh ( s a l a l ) Gentiana douglasiana Bong, (swamp gentian) Geum c a l t h i f o l i u m Menzies ex J.E. Smith i n Rees ( c a l t h a - l e a v e d avens) Gymnocarpium d r y o p t e r i s (L.)Newm. var. disjunctum (Rupr.)Ching (oak fern) Huperzia selago (L.)Bernh ( f i r clubmoss) Juniperus communis L. (common j u n i p e r ) Kalmia m i c r o p h y l l a (Hook.)Heller subsp. o c c i d e n t a l i s ( S m a l l ) T a y l o r and MacBryde (bog l a u r e l ) Ledum groenlandicum Oeder (Labrador tea) Linnaea b o r e a l i s L. (twinflower) L i s t e r a c a u r i n a P i p e r (northwestern twayblade) L i s t e r a c o r d a t a (L.)R. Br. i n W. A i t . ( h e a r t - l e a v e d twayblade) L o i s e l e u r i a procumbens (L.)Desv. ( a l p i n e azalea) Luzula p a r v i f l o r a (Ehrh.)Desv. ( s m a l l - f l o w e r e d wood rush) L y c o p o d i e l l a inundata (L.)Holub (bog clubmoss) Lycopodium annotinum L. ( s t i f f clubmoss) Lycopodium clavatum L. (running clubmoss) Lycopodium obscurum L. (ground pine) Lycopodium s i t c h e n s e Rupr. var. s i t c h e n s e (Alaska clubmoss) L y s i c h i t o n americanum Hulten and S t . John (skunk cabbage) Maianthemum d i l a t a t u m (Wood)Nels. and Macbr. (two-leaved f a l s e Solomon's s e a l ) Menyanthes t r i f o l i a t a L. (buckbean) Menziesia f e r r u g i n e a Smith ( f a l s e azalea) M i c r o s e r i s b o r e a l i s (Bong.)Schutlz-Bib. (apargidium) Monesis u n i f l o r a (L.)A. Gray (one-flowered wintergreen) Myrica gale L. (sweet gale) Nuphar l u t e a ( L . ) S i b t h . and Sm. subsp. p o l y s e p a l a (Engelm.) Beal (yellow pond l i l y ) Oplopanax h o r r i d u s ( J . E . Smith)Miq. ( d e v i l s club) O r t h i l i a secunda (L.)House (one-sided wintergreen) P a r n a s s i a f i m b r i a t a Konig var. f i m b r i a t a ( f r i n g e d grass of parnassus) P e d i c u l a r i s sp. L. (lousewort) P i c e a s i t c h e n s i s (Bong.)Carr. ( S i t k a spruce) 196 Appendix IV - Cont'd P i n g u i c u l a v u l g a r i s L. (common butterwort) Pinus c o n t o r t a Dougl. ex Loud. var. c o n t o r t a (shore pine) Plantago macrocarpa Cham, and S c h l e c h t . (Alaska p l a n t a i n ) P l a n t a n t h e r a d i l a t a t a ( P u r s h ) L i n d . ex L.C. Beck var. a l b i f l o r a (Cham.)Ledeb. (white r e i n o r c h i d ) P l a t a n t h e r a s t r i c t a L i n d l . ( s l e n d e r r e i n o r c h i d ) Polypodium g l y c o r r h i z a DC. Eaton ( l i c o r i c e f ern) Polystichum munitum ( K a u l f . ) P r e s l . (sword fern) P t e r i d i u m a q u i l i n u m (L.)Kuhn i n Decken subsp. aqui l i n u m (bracken fern) Rhynchospora a l b a (L.)Vahl (white-topped beak rush) Ribes bracteosum Dougl. ex Hook, ( s t i n k c u r r e n t ) Ribes l a x i f l o r u m Pursh ( t r a i l i n g b l a c k c u r r e n t ) Rubus chamaemoris L. (cloudberry) Rubus pedatus J.E. Smith ( f i v e - l e a v e d bramble) Rubus s p e c t a b i l i s Pursh (salmon b e r r y ) Sambucus racemosa L. subsp. pubens (A. Michaux)House (red e l d e r b e r r y ) Sanguisorba o f f i c i n a l i s L. subsp. microcephala (K.B. P r e s l ) C a l d e r and T a y l o r (great burnet) Scheuchzeria p a l u s t r i s L. subsp. americana ( F e r n a l d ) H u l t e n (scheuchzeria) S e l a g i n e l l a sp. Beauv. ( s e l a g i n e l l a ) Streptopus a m p l e x i f o l i u s (L.)DC. i n Lam. and DC. ( c l a s p i n g t w i s t e d s t a l k ) Streptopus roseus Mischx. (rosy t w i s t e d s t a l k ) Streptopus s t r e p t o p o i d e s (Ledeb.)Frye and Rigg var. b r e v i p e s ( B a k e r ) F a s s e t t (small t w i s t e d s t a l k ) Taxus b r e v i f o l i a Nutt. (western yew) T h e l y p t e r i s p h e g o p t e r i s ( L . ) S l o s s o n i n Rydb. (long beech fern) Thuja p l i c a t a Donn ex D. Don i n Lamb, (western red-cedar) T i a r e l l a l a c i n i a t a Hook, ( c u t - l e a v e d foamflower) T i a r e l l a t r i f o l i a t a L. ( t r i f o l i a t e - l e a v e d foamflower) T i a r e l l a u n i f o l i a t a Hook, ( u n i f o l i a t e - l e a v e d foamflower) T o f i e l d i a g l u t i n o s a (Michx.)Pers. ( s t i c k y f a l s e asphodel) Trichophorum cespitosum ( L . ) C . J . Hartm. subsp. cespitosum ( t u f t e d deer grass) T r i e n t a l i s europaea L. subsp. a r t i c a ( F i s c h . ex Hook.) Hulten (northern s t a r f l o w e r ) Tsuga h e t e r o p h y l l a (Raf.)Sarg. (western hemlock) 197 Appendix IV - Cont'd Tsuga mertensiana (Bong.)Carr. (mountain hemlock) Vaccinium alaskaense How. (Alaska b l u e b e r r y ) Vaccinium caespitosum Michx. (dwarf b l u e b e r r y ) Vaccinium o v a l i f o l i u m Sm. i n Rees ( o v a l - l e a v e d b l u e b e r r y ) Vaccinium oxycoccos L. (bog cranberry) Vaccinium p a r v i f o l i u m Sm. i n Rees (red huckleberry) Vaccinium uliginosum L. ( v e l v e t - l e a v e d blueberry) Vaccinium v i t i s - i d a e a L. subsp. minus (Lodd.)Hulten (mountain cranberry) Veratrum v i r i d e W. A i t . subsp. e s c h s c h o l t z i i (Gray)Love and Love ( f a l s e i n d i a n h e l l i b o r e ) V i o l a g l a b e l l a Nutt. i n T o r r . and Gray (yellow wood v i o l e t ) V i o l a p a l u s t r i s L. (purple v i o l e t ) Mosses and L i v e r w o r t s Anastrophyllum minutum (Schreb.)Schust. Andrea r u p e s t r i s Hedw. A n t h e l i a j uratzkana (Limpr.)Trev. A n t i t r i c h i a c u r t i p e n d u l a (Hedw.)Brid. Atrichum s e l w y n i i Aust. Atrichum undulatum (Hedw.)P. Beauv. Bazzania denudata ( T o r r e y ) T r e v . Bazzania t r i c r e n a t a (Wahlenb.)Trev. Bazzania t r i l o b a t a (L.)S. Gray Blepharostoma t r i c h o p h y l l u m (L.)Dum. Brachythecium f r i g i d u m (C. Muell.)Besch. C a l y p o g e i a i n t e g r i s t i p u l a Steph. Caly p o g e i a m u e l l e r i a n a ( S c h i f f n . ) K . M u l l C a l y p o g e i a t r i c h o m a n i s (L.)Corda Campylopus a t r o v i r e n s De Not. Cephaloz i a b i c u s p i d a t a (L.)Dum. Cephaloz i a media Lindb. C e p h a l o z i e l l a d i v a r i c a t a ( F r a n e ) S c h i f f n . Claopodium c r i s p i f o l i u m (Hook.)Ren. and Cord. Claopodium b o l a n d e r i Best Conocephalum conicum (L.)Dum. D i c r a n e l l a h eteromalla (Hedw.)Schimp. D i c r a n o w e i s i a c i r r a t a (Hedw.)Lindb. ex Milde Dicranum sp. Hedw. Dicranum fuscescens Turn. Appendix IV - Cont'd > Dicranum h o w e l l i i Ren. and Cord. Dicranum majus Sm. Dicranum undulatum B r i d . D i p l o p h y l l u m a l b i c a n s (L.)Dum. Dip l o p h y l l u m t a x i f o l i u m (Wahlenb.)Dum. Douinia ovata (Dicks.)Buch F r u l l a n i a n i s q u a l l e n s i s S u l l . Gymnocolea i n f l a t a (Huds.)Dum. Herberta adunca ( D i k s . ) S . Gray Hookeria lucens (Hedw.)Sm. Hylocomium splendens (Hedw.)B.S.G. Hypnum c i r c i n a l e Hook. Hypnum subimponens Lesq. Isothecium s t o l o n i f e r u m B r i d . J a m e s o n i e l l a autumnalis (DC.)Steph. (Persson, 1948) L e p i d o z i a reptans (L.)Dum. L e u c o l e p i s m e n z i e s i i (Hook.)Steere ex L. Koch Lophozia spp. Dum. Metaneckera m e n z i e s i i (Hook, ex Drumm.)Steere M i c r o l e p i d o z i a s e t a c e a . (Web.)Jorg. M y l i a anomala (Hook.)S. Gray M y l i a t a y l o r i (Hook.)S. Gray Odontoschisma denudatum (Nees)Dum. Paraleucobryum enerve (Thed. ex C.J. Hartm.)Loeske P e l l i a neesiana (Gott.)Limpr. P l a g i o c h i l a a s p l e n i o i d e s (L.)Dum. Pla g i o t h e c i u m laetum B.S.G. Pl a g i o t h e c i u m undulatum (Hedw.)B.S.G. P l e u r o z i a purpurea ( L i g h t f . ) L i n d b . Pleurozium s c h r e b e r i ( B r i d . ) M i t t . Pogonatum alpinum (Hedw.)Rohl. Pogonatum contortum ( B r i d . ) L e s q . P o l y t r i c h u m commune Hedw. Po l y t r i c h u m formosum Hedw. Po l y t r i c h u m juniperinum Hedw. P t i l i d i u m c a l i f o r n i c u m (Aust.)Und. and Cook P t i l i d i u m c i l i a r e (Web.)Hampe P t i l i u m c r i s t a - c a s t r e n s i s (Hedw.)De Not. Radula b o l a n d e r i G o t t . Rhacomitrium a c i c u l a r e (Hedw.)Brid. Rhacomitrium canescens (Hedw.)Brid. 199 Appendix IV - Cont'd Rhacomitrium heterostichum (Hedw.)Brid. Rhacomitrium lanuginosum (Hedw.)Brid. Rhizomnium glabrescens (Kindb.)Kop. R h y t i d i a d e l p h u s l o r e u s (Hedw.)Warnst. R h y t i d i a d e l p h u s squarrosus (Hedw.)Warnst. Rhy t i d i a d e l p h u s t r i q u e t r i s (Hedw.)Warnst. R i c c a r d i a l a t i f r o n s Lindb. Scapania b o l a n d e r i Aust. Sphagnum c e n t r a l e C. Jens, ex H. A r n e l l and C. Jens. Sphagnum compactum DC. ex fam. and DC. Sphagnum cuspidatum Ehrh. ex Hoffm. Sphagnum f a l l a x K l i n q g r . ( K l i n g g r . ) Sphagnum fuscum (Schimp.)Klinggr. Sphagnum g i r g e n s o h n i i Russ. Sphagnum imbricatum Hornsch. ex Russ. Sphagnum l i n d b e r g i i Schimp. ex Lindb. Sphagnum magellanicum B r i d . Sphagnum nemoreum Scop. Sphagnum p a p i l l o s u m Lindb. Sphagnum rubellum W i l s . Sphagnum squarrosum Crome Sphagnum ^ tenellum ( B r i d . ) P e r s . ex B r i d . S t o k e s i e l l a oregana ( S u l l . ) R o b i n s . Lichens A l e c t o r i a f r e m o n t i i Tuck A l e c t o r i a sarmentosa (Ach.)Ach. C e t r a r i a , .is land i c a (L.)Ach. C l a d o n i a sp. Wigg. C l a d o n i a a l p e s t r i s (L.)Rabh. C l a d o n i a amaurocraea ( F l o r k e ) S c h a e r . C l a d o n i a b e l l i d i f l o r a (Ach.)Schaer. C l a d o n i a f u r c a t a (Huds.)Schrad. C l a d o n i a g r a c i l i s ( L . ) W i l l d . v a r . elongata ( J a c q . ) F r . C l a d o n i a impexa Harm. C l a d o n i a leucophaea Abb. C l a d o n i a m u l t i f o r m i s Merr. C l a d o n i a pseudoevansii Asah. C l a d o n i a r a n g i f e r i n a (L.)Wigg. C l a d o n i a squamosa (Scop.)Hoffm. Appendix IV - Cont'd C l a d o n i a s y l v a t i c a (L.)Hoffra. C l a d o n i a t e n u i s (Florke)Harm. C l a d o n i a u n c i a l i s (L.)Wigg. C o r n i c u l a r i a a c u l e a t a (Schreb.)Ach. Hypnogymnia enteromorpha (Ach.)Nyl. Icmadophila e r i c e t o r u m ( L . ) Z a h l b r . L o b a r i a l i n i t a (Ach.)Rabenh. L o b a r i a oregana (Ruck.)Mull. Arg. P e l t i g e r a aphthosa ( L . ) W i l l d . S i h p u l a c e r a t i t e s (Wahlenb.)Fr. Sphaerophorus globosus (Huds.)Vain. Stereocaulon paschale (L.)Hoffm. ^-Nomenclature f o l l o w s T a y l o r and MacBryde (1977) f o r the v a s c u l a r p l a n t s , I r e l a n d et a l . (1980) f o r the mosses, Worley (1970) f o r the l i v e r w o r t s and Otto and A h t i (1967) f o r the 1ichens^except f o r the genus C l a d o n i a which f o l l o w s Thomson (1967) APPENDIX V Species Groups Outlined on the PCA Species Ordination (Fig. 8 . ) 201 Appendix V - L i s t of s p e c i e s f o r each s p e c i e s group o u t l i n e d on the PCA o r d i n a t i o n ( F i g . 8 ) . The s p e c i e s are l i s t e d a l p h a b e t i c a l l y by the four l e t t e r a b b r e v i a t i o n s used i n F i g . 8. Group A AGAE A g r o s t i s a e q u i v a l v i s ANPO Andromeda p o l i f o l i a CALI Carex l i v i d a CLIM C l a d o n i a impexa CLUN C l a d o n i a u n c i a l i s DRAN Drosera a n g l i c a DRRO Drosera r o t u n d i f o l i a ERAN Eriophorum a n g u s t i f o l i u m GEDO Gentiana d o u g l a s i a n a GYIN Gymnocolea i n f l a t a JUCO ' Juniperus communis PICO(B2) Pinus c o n t o r t a (low shrubs) RHAL Rhyncospora a l b a RHLA Rhacomitrium lanuginosum SICE S i p h u l a c e r a t i t e s SPLI Sphagnum l i n d b e r g i i SPTE Sphagnum tenellum TRCE Trichophorum cespitosum VAOX Vaccinium oxycoccus Group B CLGR C l a d o n i a g r a c i l i s CLRA C l a d o n i a r a n g i f e r i n a COTR C o p t i s t r i f o l i a EMNI Empetrum nigrum KAMI Kalmia m i c r o p h y l l a LEGR Ledum groenlandicum PICO(Bl) Pinus c o n t o r t a ( t a l l shrubs) SAOF Sanguisorba o f f i c i n a l i s SPFU Sphagnum fuscum SPPA Sphagnum p a p i l l o s u m SPRU Sphagnum rubellum TOGL T o f i e l d i a g l u t i n o s a TREU T r i e n t a l i s europaea VAUL Vaccinium u l i g i n o s u m 202 Appendix V - cont'd Group C CAPL Carex- p l u r i f l o r a CHNO(B2) Chamaecyparis n o o t k a t e n s i s (low shrubs) FACR F a u r i a c r i s t a - g a l l i LYAN Lycopodium annotinum PICO(A2, Bl) Pinus c o n t o r t a (main canopy, t a l l shrubs) PLSC Pleuroz ium s c h r e b e r i PTCR P t i l i u m c r i s t a - c a s t r e n s i s THPL(B2) Thuja p l i c a t a (low shrubs) VACA Vaccinium caespitosum VAVI Vaccinium v i t i s - i d a e a Group D CHN0(B1) GASH(B2) LIBO LYCL THPL(Bl) Chamaecyparis n o o t k a t e n s i s ( t a l l shrubs) G a u l t h e r i a s h a l l o n ( t a l l shrubs) Linnaea b o r e a l i s Lycopodium clavatum Thuja p l i c a t a ( t a l l shrubs) Group E CHNO(A2, A3) Chamaecyparis n o o t k a t e n s i s (main canopy and suppressed t r e e s ) TSME(A3, B l , B2) Tsuga mertensiana (suppressed t r e e s , t a l l and low shrubs) VEVI Veratrum v i r i d e Group F COAS C o p t i s a s p l e n i i f o l i a COUN Cornus u n a l a s c h k e n s i s HYSP Hylocomium splendens LICO L i s t e r a cordata LYAM L y s i c h i t o n americanum MEFE(B2) Mensi e z i a f e r r u q i n e a RHLO R h y t i d i a d e l p h u s l o r e u s SPGI Sphagnum q i r q e n s o h n i i THPL(A3) Thuja p l i c a t a (suppressed t r e e s ) TSHE(B2) Tsuga h e t e r o p h y l l a (low shrubs) VAOV(B2) Vaccinium o v a l i f o l i u m (low shrubs) VAPA(B2) Vaccinium p a r v i f o l i u m (low shrubs) 203 Appendix V - cont'd Group G BLSP Blechnum s p i c a n t LICA L i s t e r a c a u r i n a PLAS P l a g i o c h i l a a s p l e n i o i d e s PLUN Pla g i o t h e c i u m undulatum RHGL Rhizomnium g l a b r e s c e n s RUPE Rubus pedatus STRO Streptopus roseus THPL(A2) Thuja p l i c a t a (main canopy) TSHE(A2, A3, Bl) Tsuga h e t e r o p h y l l a (main canopy and suppressed t r e e s , t a l l shrubs) VAAL(B2) Vaccinium alaskaense (low shrubs) Group H ABAM(Al, A2, A3, B l , B2) Abies a m a b i l i s ( a l l t r e e and shrub l a y e r s ) ATFI Athyrium f i l i x - f e m i n a COCO Conocephalum conicum DRAS D r y o p t e r i s a s s i m i l i s GYDR Gymnocarpium d r y o p t e r i s OPHO(Bl, B2) Oplopanax h o r r i d u s ( t a l l and low shrubs) P e l l i a neesiana PENE PLLA POAL POCO POMU SARA ST AM STOR THPH TITR VIGL Plag i o t h e c i u m laetum  Pogonatum alpinum  Pogonatum contortum  Polystichum muniturn  Sambucus racemosa  Streptopus a m p l e x i f o l i u s  S t o k e s i e l l a oregana  T h e l y p t e r i s p h e g o p t e r i s  T i a r e l l a t r i f o l i a t a  V i o l a g l a b e l l a APPENDIX VI Complete V e g e t a t i o n Tables and V e g e t a t i o n Summary Tables f o r Each Community Type APPENDIX VI Comp le t e v e g e t a t i o n t a b l e s and v e g e t a t i o n summary t a b l e s f o r each community t y p e The v e g e t a t i o n t a b l e s f o r t h e s i x community t y p e s a r e p r e s e n t e d on pages 205-222 . S p e c i e s a r e l i s t e d i n o r d e r o f d e c r e a s i n g p r e s e n c e ( c o n s t a n c y ) v a l u e s w i t h i n each s t r a t u m . The s p e c i e s s i g n i f i c a n c e s c a l e s , v i g o u r s c a l e s and s t r a t a d e s i g n a t i o n s a r e e x p l a i n e d i n A p p e n d i x I I I . S p e c i e s p r e s e n c e o r c o n s t a n c y ( P ) , mean s p e c i e s s i g n i f i c a n c e (MS) and range o f s p e c i e s s i g n i f i c a n c e (RS) a r e summar ized f o r each o f the community t y p e s i n t h e t h r e e l e f t hand co lumms. The two l e t t e r s p r e c e e d i n g each r e l e v e number i n d i c a t e t h e s t u d y t r a n s e c t a l o n g w h i c h t h e r e l e v e o c c u r s ( see page 2 0 ) . The o r d e r o f r e l e v e s i s t h a t p r o d u c e d by t h e f i r s t a x i s RA r e l e v e o r d i n a t i o n ( A p p e n d i x V I I ) . The v e g e t a t i o n summary t a b l e i s p r e s e n t e d on pages 223-227 . In t h i s t a b l e , p r e s e n c e c l a s s and mean s p e c i e s s i g n i f i c a n c e v a l u e s f o r each s p e c i e s i s summar ized by communi ty t y p e . The p r e s e n c e ( o r c o n s t a n c y ) c l a s s e s a r e as f o l l o w s ; 1-20% ( I ) , 21-40% ( I I ) , 41-60% ( I I I ) , 61-80% ( I V ) , 81-100% ( V ) . S p e c i e s a r e l i s t e d a l p h a b e t i c a l l y . Appendix Vl-cont'd Vegetation Tables SYNSYSTEMATIC UNIT 001ErIcaceous shrub - Trichophorum - Sphagnum - Cladonia - Siphula Open Bogs PLOT NUMBER I SYNTHETIC | VALUES IRW IRE I 1 l | 28 R39 °22 RE IRW 381 7 RE IDC IRW IDC IHM I 14 I 24| 12| 301 G| ST.NO. S P E C I E S | P MS RS I S P E C I E S SIGNIFICANCE AND VIGOR c 1 PINUS CONTORTA 41 7 1 6 0-4 | * * * 1 1 ' I 2 KALMIA MICROPHYLLA 100 0 4 6 1 -6 1 1 3 2 4 2 4 2 2 1 4 2 5 7 3 1 3 t 6 2 2 2 5 2 3 VACCINIUM ULIGINOSUM 9 1 7 3 2 0-4 1 2 1 2 1 1 3 1 3 2 1 2 2 1 3 1 4 2 3 1 4 2 4 JUNIPERUS COMMUNIS 83 3 5 0 0-7 3 1 7 1 5 2 5 2 4 1 5 2 4 1 2 1 4 1 4 1 5 LEDUM GROENLANDICUM 83 3 3 5 0-5 1 1 2 1 4 2 4 2 + 1 4 2 2 + 1 1 1 1 5 2 PINUS CONTORTA - CONTORTA 75 0 3 8 0-6 4- + 4 + + 1 + + 3 + 4 + 3 + 6 4- 3 * 6 MYRICA GALE 33 3 5 1 0-8 8 2 5 2 6 2 7 2 7 CHAMAECYPARIS NOOTKATENSIS 33 3 2 5 0-5 + + 2 + + 1 5 + 8 THUJA PLICATA 25 0 1 9 0-4 4 + 3 + 4- + 9 TSUGA HETEROPHYLLA 25 0 + O 0-* + + + + + + 10 TRICHOPHORUM CESPITOSUM 100 0 7 1 2-8 5 2 5 2 6 2 6 2 4 2 6 2 8 2 8 2 8 2 8 2 7 2 2 2 1 1 TRIENTALIS EUROPAEA 100 0 4 5 1-8 3 1 2 1 1 2 2 2 2 1 2 2 4 2 3 1 2 1 4 2 2 1 8 2 12 ANDROMEDA POLYFOLI A 100 0 4 1 1-5 4 2 2 2 5 2 4 2 3 2 2 2 4 1 4 2 4 1 1 2 2 2 4 3 13 COPTIS TRIFOLIA 100 0 3 9 1-5 1 2 1 1 3 2 4 2 1 1 1 2 5 1 4 2 2 2 5 2 2 2 1 1 14 VACCINIUM OXYCOCCOS 100 0 3 9 + -7 1 2 + 1 2 2 3 2 1 2 1 2 4 1 2 2 1 2 1 2 1 2 7 3 15 DROSERA ROTUNDIFOLIA 100 0 3 6 1-S 5 3 1 2 2 2 2 3 2 1 2 4 1 3 2 3 2 1 1 1 2 4 1 16 GENT I ANA DOUGLAS I ANA 100 0 3 1 + -5 3 2 1 2 1 2 2 2 2 2 1 2 5 2 1 2 2 2 + 1 1 2 2 3 17 SANGUISORBA OFFICINALIS 9 1 7 5 6 0-8 7 2 4 1 7 2 6 2 4 1 4 1 6 2 5 1 4 1 4 1 4 1 18 ERIOPHORUM ANGUSTI FOLIUM 9 1 7 5 5 0-8 8 2 2 2 4 2 3 2 6 2 5 2 8 2 3 2 3 2 5 2 3 2 19 EMPETRUM NIGRUM 9 1 7 4 4 0-8 + 2 2 2 2 2 3 2 4 2 2 2 3 2 2 1 8 2 2 2 2 2 20 CAREX LIVIDA 91 7 3 3 0-4 1 2 3 2 3 2 4 2 2 2 3 2 4 2 3 1 3 2 2 2 2 2 2 1 CORNUS UNALASCHKENSIS 83 3 4 1 0-6 + 1 3 2 2 2 3 1 5 1 3 1 6 1 2 1 3 1 3 1 22 DROSERA ANGLICA 75 0 3 0 0-5 5 3 1 2 2 2 3 1 2 3 2 + 2 1 2 1 2 23 AGROSTIS AEOUIVALVIS 75 0 3 0 0-4 4 2 2 2 3 2 4 2 2 2 2 2 1 2 2 2 2 2 24 TOFI ELD IA GLUT INOSA 66 7 1 9 0-3 1 2 2 2 1 2 1 2 1 1 4- 1 3 2 3 2 25 RHYNCHOSPORA ALBA 58 3 5 1 0-7 5 2 7 2 7 2 6 2 4 2 1 4 2 26 FAURIA CRISTA-GALLI 58 3 3 3 0-5 • 3 + + 1 + 1 4 1 4 1 5 2 27 RUBUS CHAMAEMORUS 4 1 7 2 4 0-4 2 1 4 2 2 + 1 1 4 1 28 CAREX PLURIFLORA 33 3 4 5 0-9 1 2 3 2 2 2 9 2 29 PLATANTHERA DILI TATA 33 3 1 0 0-3 3 2 4 1 + 1 + 1 30 HUPERZIA SELAGO 33 3 + 3 0-2 + t 2 2 + 2 + 1 3 1 LYCOPODIUM ANNOTINUM 33 3 + O 0- + + 1 + 4 2 2 32 VACCINIUM CAESPITOSUM 33 3 + 0 0- + * + 4- 1 • + +• 1 33 LOISELEURIA PROCUMBENS 25 0 1 0 0-2 2 2 2 2 1 2 34 COPTIS ASPLENIFOLIA 25 0 + 4 0-2 1 2 + 1 2 1 35 LYCOPODIUM CLAVATUM 25 0 + 0 0- 1 + 1 1 + 1 36 CAREX SITCHENSIS 16 7 1 0 0-3 1 2 3 2 37 CAREX STYLOSA 16 7 1 0 0-3 1 2 3 2 38 DODECATHEON JEFFREY 1 8 3 + 9 0-3 3 2 39 GEUM CALTHIFOLIUM 8 3 + 9 0-3 3 1 40 LYSICHITON AMERICANUM 8 3 * 9 0-3 3 1 4 1 CAREX ROSTRATA 8 3 + 1 0-2 2 2 42 MA IANTHEMUM 01LATATUM 8 3 + 1 0-2 2 2 43 CAREX PAUCIFLORA 8 3 + 0 0- 1 1 2 44 ERIGERON PEREGRINUS 8 3 + 0 0- 1 1 2 45 LYCOPOOIELLA INUNDATA 8 3 +• 0 0- 1 1 2 SYNSY5TEMATIc UNIT 001ErIcaceous shrub - TrIchophorum - Sphagnum - Cladonia - Slphula Open Bogs PLOT NUMBER 1 SYNTHETIC 1 VALUES IRW IRE | II 1 28 ° 4 o "5. RE |RW 381 7 RE IDC IRW IDC IHM 1 141 241 12| 301 e| ST NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR DH 46 MENYANTHES TRIFOLIATA S 3 + 0 0- 1 1 2 47 PEOICULARIS 8 3 + 0 0- 1 1 2 48 PINGUICULA VULGARIS 8 3 • 0 0- 1 1 2 49 BLECHNUM SPICANT 8 3 + 0 0- + + * 50 LYCOPODIUM OBSCURUM 8 3 + 0 0- + * 2 51 LYCOPODIUM SITCHENSE 8 3 + 0 0-* + 52 PLATANTHERA STRICTA 8 3 + 0 0- + + . 2 53 SCHEUCHZERIA PALUSTRIS 8 3 + 0 0- + + 2 54 SELAGINELLA 8 3 + 0 0- + + 2 55 SPHAGNUM PAPILLOSUM 100 0 5 8 + -9 4 . 2 1 2 2 2 2 2 • . 2 5 2 7 2 9 3 7 2 2 2 2 2 8 . 56 SPHAGNUM TENELLUM 100 0 5 0 0-7 7 . 2 5 2 5 2 5 2 4 . 2 2 2 3 2 2 2 3 2 3 2 +• 57 SPHAGNUM RUBELLUM 100 0 4 5 + -7 2.2 1 2 1 2 1 2 1 .2 3 2 5 2 3 2 3 2 5 2 + 2 7 . 58 CLADONIA IMPEXA 91 7 5 0 0-8 4 . 3 5 5 4 . 4 3 2 2 3 2 8 3 59 CLADONIA GRACILIS 9 1 7 2 9 0-5 3 . 2 + 1 + . 1 5 2 + + 1 + 60 RHACOMITRIUM LANUGINOSUM 83 3 4 9 0-7 5.2 7 2 3 2 4 2 4.2 5 2 5 1 + 1 * 3 2 6 1 SIPHULA CERATITES 83 3 4 5 0-7 7 . 4 4 4 5.2 2 3 2 + 2 1 1 4 62 CLADONIA UNCIALIS 83 3 4 0 0-5 5. 2 3 3 4 . 2 5 2 + 2 4 1 63 SPHAGNUM FUSCUM 75 0 3 4 0-5 1 . 2 2 2 1 2 ».2 5 2 1 2 2 2 5 2 64 CLAOONIA RANG IFERINA 66 7 3 6 0-6 3. 2 3 3 3 . 2 6 2 1 65 SPHAGNUM LINDBERGH 58 3 4 1 0-7 4 2 3 2 7 . 2 2 2 4 2 2 2 3 . 66 PLEUROZIUM SCHREBERI 58 3 3 4 0-7 + + 2 4 + 2 1 1 7 + 2 67 GYMNOCOLEA INFLATA PINUS CONTORTA - CONTORTA 50 50 0 0 1 + 4 2 0-3 0-1 1 2 2 2 3 + . 1 + 2 1 + +• + 1 + + 2 + + . 2 68 SPHAGNUM COMPACTUM 4 1 7 3 6 0-6 4.2 4 2 + 2 3 2 6 2 69 CLADONIA FURCATA TSUGA HETEROPHYLLA 4 1 4 1 7 7 1 • 1 0 0-3 0- + 3. + . + + + + . 1 4- 2 + 70 CORNICULARIA ACULEATA 33 3 4 0- 1 1 1 . + 1 7 1 CLAOONIA LEUCOPHAEA 25 0 1 5 0-3 2 3 72 CLADONIA 25 0 + 0 0- 1 + 1 73 CLADONIA PSEUDOEVANSII 25 0 + 0 0-1 + + . 1 74 POLYTRICHUM JUNIPERINUM 25 0 + 0 0-1 + 2 1 + 75 MYLIA ANOMALA 25 0 + 0 0-* + + 76 SPHAGNUM IMBRICATUM 16 7 3 5 0-6 6 2 5 2 77 SPHAGNUM RECURVUM 16 7 3 4 0-7 1 2 7 . 78 CLADONIA SYLVATICA 16 7 1 8 0-4 3 4 . 79 LOPHOZIA 16 7 1 4 0-4 4.2 + 80 DICRANUM HOWELLI I 16 7 + 2 0-2 2 2 8 1 CETRARI A ISLANDICA 16 7 + 0 0-1 1 1 . 82 BAZZANIA TRICRENATA 16 7 + 0 0-* + + 83 CLADONIA BEL LID IF LORA 16 7 0 0-* *• f 84 DICRANUM UNDULATUM 8 3 + 9 0-3 3 85 CLADONIA TENUIS 8 3 • 1 0-2 2 . 86 DICRANUM SPECIES 8 3 + 0 0- 1 1 .2 87 HERBERTA ADUNCA 8 3 • 0 0- 1 1 88 PLEUROZIA PURPUREA 8 3 + 0 0- 1 1 2 89 PTILIUM CRISTA-CASTRENSIS 8 3 + 0 0- 1 1 90 SPHAGNUM MAGELLANICUM 8 3 0 0- 1 1 2 9 1 ANT ITRI CHI A CURT IPENDULA 8 3 + 0 0- + + . SYNSYSTEMATIC UNIT 001Er1caceous shrub - Tr1chophorum - Sphagnum - Cladonia - Slphula Open Bogs PLOT NUMBER 1 SYNTHETIC 1 VALUES IRW 1 11 R 28 *4o R 39 R 3 s | R W 7 RE IDC IRW IDC IHM I 14| 24| 12| 301 6| ST.NO. S P E C I E S | P MS RS I S P E C I E S SIGNIFICANCE AND VIGOR 92 BAZZANIA TRILOBATA 8 3 + 0 0- + 93 BLEPHAROSTOMA TRICHOPHYLLUM 8 3 + 0 0-CHAMAECYPARIS NOOTKATENSIS 8 3 + 0 0- + 94 CLADONIA SOUAMOSA 8 3 + 0 0- + 95 HYPNUM CIRCINALE 8 3 + 0 o- + 96 MYLIA TAYLORI 8 3 + 0 0- + 97 ODONTOSCHISMA DENUDATUM 8 3 0 0- + 98 P E L L IA NEESIANA 8 3 0 0- + 99 PLAGIOTHECIUM UNDULATUM 8 3 + 0 0- + 100 P T I L I D I U M C I L I A R E 8 3 + 0 0- + SYNSYSTEMATIC UNIT 002Empetrum - Vaccinium ullglnosum - Rhacomitrium - Cladonia Outcrops PLOT NUMBER 1 SYNTHETIC |RE 1 | VALUES 1 t| ST.NO. SPECIES | P MS RS | SPECIES SIGNIFICANCE AND VIGOR C OH 1 PINUS CONTORTA 100 0 2 4 2 -2 2 * 2 CHAMAECYPARIS NOOTKATENSIS 100 0 + 5 + - * 0 CHAMAECYPARIS NOOTKATENSIS 100 0 6 5 6 -6 6 + 3 VACCINIUM ULIGINOSUM 100 0 6 5 6 -6 6 3 4 GAULTHERIA SHALLON 100 0 5 5 5 -5 5 1 5 JUNIPERUS COMMUNIS 100 0 5 5 5 -5 5 1 PINUS CONTORTA too 0 2 4 2 -2 2 + 6 KALMIA MICROPHYLLA 100 0 1 5 1 - 1 1 2 7 LEDUM GROENLANOICUM 100 0 1 5 1 - 1 1 1 8 THUJA PLICATA 100 0 +• 5 + + + 9 TSUGA MERTENSIANA 100 0 + 5 + - 4 - + + 10 EMPETRUM NIGRUM too 0 7 5 7 -7 7 2 1 1 LOISELEURIA PROCUMBENS 100 0 4 5 4 -4 4 2 12 TRICHOPHORUM CESPITOSUM 100 0 4 5 4 -4 4 2 13 ANDROMEDA POLYFOLIA 100 0 2 4 2 -2 2 1 14 COPTIS TRIFOLIA too 0 2 4 2 -2 2 1 15 CORNUS UNALASCHKENSIS 100 0 2 4 2 -2 2 1 16 FAURIA CRISTA-GALLI 100 0 2 4 2 -2 2 1 17 SANGUISORBA OFFICINALIS 100 0 2 4 2 -2 2 2 18 VACCINIUM VITIS-IDAEA 100 0 2 4 2 -2 2 1 19 VACCINIUM CAESPITOSUM 100 0 1 5 1 - 1 1 1 20 BLECHNUM SPICANT 100 0 + 5 + - + + + 2 1 LINNAEA BOREAL IS too 0 5 + + 22 LYCOPODIUM CLAVATUM too 0 + 5 + - + + + 23 HERBERTA ADUNCA 100 0 4 5 - f - + + . 24 CLADONIA 100 0 1 8 1-1 1 . 25 CLADONIA SYLVATICA 100 0 7 5 7-7 7 . 26 RHACOMITRIUM LANUGINOSUM 100 0 7 5 7-7 7 . 3 27 CLADONIA AMOUROCRAEA 100 0 5 5 5-5 5 : 28 CLADONIA RANGIFERINA 100 0 3 4 3-3 3. 29 CLADONIA GRACILIS 100 0 2 4 2-2 2 . 30 HYLOCOMIUM SPLENDENS 100 0 2 4 2-2 2 . 31 RHYTIDIADELPHUS LOREUS 100 0 2 4 2-2 2 . 32 DICRANUM FUSCESCENS 100 0 1 5 1-1 1 . 33 STEREOCAULON PASCHALE 100 0 1 5 1-1 1 . 34 BAZZANIA TRICRENATA too 0 + 5 + -+ + . 35 DIPLOPHYLLUM ALBICANS too 0 5 + + . 36 HYPNUM CIRCINALE 100 0 5 + 37 ICMADOPHILA ERICETORUM 100 0 + 5 + -+ + . 38 LOPHOZIA 100 0 + 5 + -+ + . 39 MICROLEPIDOZIA SETACEA too 0 5 + -+ 40 MYLIA ANOMALA 100 0 + 5 + -+ + . 4 1 ODONTOSCHISMA DENUDATUM 100 0 + 5 + -+ 42 PARALEUCOBRYUM ENERVE 100 0 + 5 + + . 43 PLEUROZIUM SCHREBERI 100 0 + 5 + -+ + . 5YNSYSTEMATIC UNIT 002Empetrum Vaccinium ulIglnosum • Rhacomitrium - Cladonia Outcrops PLOT NUMBER 1 SYNTHETIC 1 RE 1 1 VALUES | 1| ST.NO. SPECIES | P MS RS | SPECIES SIGNIFICANCE AND VIGOR 44 SIPHULA CERATITES 45 SPHAGNUM RUBELLUM 46 ANDREA RUPESTRIS 47 ANTHELIA JURATZKANA DIPLOPHYLLUM ALBICANS LOPHOZIA MICROLEPIDOZIA SETACEA RHACOMITRIUM LANUGINOSUM SYNSYSTEMATIC UNIT OOSPInus - Chamaecyparis - Gualtherla - Fauria - Sphagnum Bog Woodlands PLOT NUMBER 1 SYNTHETIC | VALUES I"? RE IRW 291 8 HM IRW 3| 35 HM IDC 21| 31 ST.NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR A3 B1 B2 C 1 CHAMAECYPARIS NOOTKATENSIS 12 5 4 4 0-2 2 < 2 THUJA PLICATA 12 5 + 4 0-2 2 1 3 PINUS CONTORTA 87 5 5 2 0-7 5 1 7 1 5 1 5 1 4 1 4 1 2 2 THUJA PLICATA 37 5 1 5 0-3 1 4 2 1 3 1 CHAMAECYPARIS NOOTKATENSIS 12 5 1 1 0-3 3 1 THUJA PLICATA 37 5 4 6 0-2 4 4 4 1 2 1 CHAMAECYPARIS NOOTKATENSIS 25 0 4 8 0-2 1 4 2 1 4 TSUGA MERTENSIANA 12 5 4 4 0-2 2 1 PINUS CONTORTA 12 5 + 0 0- 1 1 1 5 TSUGA HETEROPHYLLA . 12 5 4 0 0-1 1 1 PINUS CONTORTA 100 0 5 4 2-7 5 4 3 4 4 4 5 1 7 1 6 1 5 1 2 1 CHAMAECYPARIS NOOTKATENSIS 100 0 5 0 2-5 2 4 4 1 5 4 4 1 5 1 4 1 4 1 5 1 THUJA PLICATA 100 0 4 2 2-5 3 1 4 4 5 4 2 4 3 1 2 1 4 1 4 1 TSUGA HETEROPHYLLA 87 5 2 2 0-3 + 1 1 1 3 1 1 1 3 1 1 1 2 1 TSUGA MERTENSIANA 62 5 1 8 0-3 4 1 2 1 2 2 4 1 3 1 6 MENZI ESI A FERRUGINEA 12 5 4 0 0- + 4 2 7 PICEA SITCHENSIS 12 5 4 0 0- + 4 1 CHAMAECYPARIS NOOTKATENSIS 100 0 8 6 6-9 8 4 8 1 9 4 9 4 8 4 9 4 7 1 6 1 8 GAULTHERIA SHALLON 100 0 7 0 2-8 2 1 5 1 6 1 8 1 7 2 6 2 7 2 7 2 THUJA PLICATA 100 0 4 2 2-5 3 4 4 4 4 4 2 4 5 1 3 4 3 1 3 1 9 LEDUM GROENLANDICUM 100 0 4 0 2-5 3 1 4 2 2 1 5 1 3 2 3 2 2 2 2 2 TSUGA HETEROPHYLLA 100 0 3 4 4-4 4 4 3 4 1 1 3 4 2 4 4 4 3 1 4 1 10 KALMIA MICROPHYLLA 100 0 3 3 4-4 3 2 3 2 4 2 3 1 1 2 4 2 3 3 3 3 PINUS CONTORTA 100 0 3 3 4-4 3 4 3 4 2 4 4 4 2 1 4 1 4 1 1 MENZIESIA FERRUGINEA 87 5 3 2 0-4 2 1 1 1 4 4 4 1 3 1 4 1 2 2 1 1 VACCINIUM PARVIFOLIUM 87 5 2 7 0-3 2 1 3 1 4 4 2 2 2 2 3 2 3 1 12 VACCINIUM OVALIFOLIUM 75 0 2 9 0-4 4 4 1 4 2 4 1 2 2 4 2 TSUGA MERTENSIANA 75 0 1 8 0-3 1 4 4 1 1 1 1 2 2 2 3 2 13 MYRICA GALE 50 0 3 6 0-5 5 2 4 2 4 1 4 2 14 VACCINIUM ALASKAENSE 50 0 3 6 0-5 3 1 3 2 5 2 4 2 15 VACCINIUM ULIGINOSUM 50 0 1 3 0-2 2 2 2 2 1 2 1 1 1G JUNIPERUS COMMUNIS 25 0 4 5 0-2 2 1 4 1 17 ABIES AMABILIS 25 0 4 0 0- + 4 4 4 1 PICEA SITCHENSIS 12 5 4 0 0- + 4 1 18 CAREX PLURIFLORA 100 0 4 5 1-7 3 2 1 3 7 2 2 2 1 2 2 2 4 2 3 3 19 FAURIA CRISTA-GALLI 100 0 6 0 *-8 4 3 7 3 a 3 4 1 4 2 3 2 5 3 8 3 20 CORNUS UNALASCHKENSIS 100 0 5 1 3-7 3 2 4 1 7 2 3 2 4 1 4 2 4 2 5 2 21 LINNAEA BOREALIS 100 0 3 6 1-4 1 2 2 2 4 2 4 2 2 2 3 2 2 2 4 2 22 VACCINIUM VITIS-IDAEA 100 0 3 3 1-4 3 2 2 2 4 2 3 2 2 2 3 2 1 2 3 2 23 COPTIS TRIFOLIA 100 0 2 2 + -3 2 2 1 3 3 2 4 2 1 2 2 2 1 2 2 2 24 COPTIS ASPLEN1F0LIA 87 5 3 4 0-5 4 2 3 2 2 2 2 2 2 2 2 2 5 2 25 SANGUISORBA OFFICINALIS 87 5 3 3 0-4 4 3 3 2 4 2 2 2 1 2 2 2 3 2 2G EMPETRUM NIGRUM 87 5 3 1 0-4 4 2 4 2 2 2 2 4 4 1 2 2 2 2 SYNSYSTEMATIC UNIT 003PInus - Chamaecyparis - Gualtherla - Fauria - Sphagnum Bog Woodlands PLOT NUMBER 1 SYNTHETIC | VALUES RE IRW IHM 29| 8| 3 HM |DC 2t| 31 ST.NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR DH 27 LYCOPODIUM CLAVATUM 87 5 2 7 0-4 1 2 2 2 4 2 1 .2 1 2 1 2 3 2 28 LYCOPODIUM ANNOTINUM 87 5 2 4 0-3 3 2 3 2 2.2 2 2 4 2 1 2 2 2 29 TRIENTALIS EUROPAEA 87 5 2 1 0-3 4 1 1 2 3 1 4 2 1 2 1 2 3 2 30 LISTERA CORDATA 87 5 1 0 0-1 4 1 1 4 . 1 1 1 4 2 4 1 1 1 31 LYSICHITON AMERICANUM 75 0 3 7 0-5 3 3 4 . 2 2 2 4 2 3 2 5 3 32 VACCINIUM CAESPITOSUM 75 0 3 0 0-3 3 2 3 1 3 3 2 2 3 2 3 2 33 TOFIELDIA GLUTINOSA 75 0 2 7 0-4 2 2 2 2 4 2 4 2 1 2 3 2 34 DROSERA ROTUNDIFOLIA 75 0 + 8 0-1 1 2 1 1 4 2 4 1 4 2 4 2 35 TRICHOPHORUM CESPITOSUM 62 5 5 1 0-8 8 2 5 2 4 2 2 2 3G BLECHNUM SPICANT 62 5 2 1 0-3 2 1 3 1 1 1 1 1 3 2 37 RUBUS CHAMAEMORUS 62 5 1 8 0-3 4 1 2 3 3.2 4 2 2 2 38 MA IANTHEMUM DILATATUM 50 0 1 7 0-3 3 . 2 2 2 2 2 4 2 39 VACCINIUM OXYCOCCOS 50 0 + 6 0-1 1 2 1 2 4 2 4 2 40 ERIOPHORUM ANGUSTIFOLIUM 37 5 2 6 0-4 4 2 3 2 3 2 41 VERATRUM VIRIDE 37 5 1 0 0-2 4 2 1 2 2 1 42 AGROSTIS AEOUIVALVIS 37 5 + 0 0- + 4 1 4 2 4 2 43 CAREX ANTHOXANTHEA 25 0 3 3 0-5 4 2 5 2 44 CAREX PAUCIFLORA 25 0 3 0 0-5 4 2 5 2 45 CAREX SITCHENSIS 25 0 2 0 0-4 4 2 1 3 46 PINGUICULA VULGARIS 25 0 4 5 0-2 4 2 2 3 47 STREPTOPUS ROSEUS 25 0 + 3 0-1 1 2 1 2 48 GENTIANA DOUGLASIANA 25 0 + 0 0-1 1 3 4 2 49 PLATANTHERA STRICTA 25 0 + 0 0-1 1 2 4 2 50 SELAGINELLA 25 0 + 0 0- + 4 2 4 2 51 CALAMAGROSTIS SPECIES CHAMAECYPARIS NOOTKATENSIS 12 12 5 5 1 + 9 4 0-4 0-2 2 2 4 3 52 ERIGERON PEREGRINUS 12 5 + 4 0-2 2 2 53 ANDROMEDA POLYFOLIA 12 5 4 0 0- 1 1 2 54 CALAMAGROSTIS NUTKAENSIS 12 5 4 0 0-1 1 2 55 CAREX LIVIDA 12 5 4 0 0- 1 1 2 56 CLINTONIA UNI FLORA 12 5 4 0 0-* 4 2 57 PLATANTHERA DILITATA 12 5 4 0 0- + 4 1 58 POACEAE SPECIES 12 5 4 0 0-* 4 2 59 RUBUS PEDATUS 12 5 4 0 0-* 4 2 60 STREPTOPUS AMPLEXIFOLIUS VACCINIUM ALASKAENSE 12 12 in m 4 4 0 0 0 - 4 0- + 4 4 1 6 1 PLEUROZIUM SCHREBERI 100 0 5 4 3-7 3 2 3 2 6 2 7 . 2 3 2 7 3 3 2 3 2 62 HYLOCOMIUM SPLENDENS 100 0 5 3 1-7 1 2 2 2 5 1 7 . 1 3 2 7 2 3 2 5 2 63 RHYTIDIADELPHUS LOREUS 100 0 4 6 1-5 1 2 5 2 4 2 5 . 1 3 2 3 2 4 2 4 2 64 SPHAGNUM RUBELLUM 100 0 4 0 + -S 5 2 3 2 5 2 2 . 1 2 1 2 4 2 1 2 65 SPHAGNUM PAPILLOSUM 100 0 3 8 1-4 4 2 4 2 4 2 2 . 4 2 3 2 1 2 2 2 66 SPHAGNUM GIRGENSOHNII 87 5 6 1 0-8 1 2 3 2 8 2 7 2 7 2 7 2 5 2 67 CLADONIA RANGIFERINA 87 5 2 0 0-3 2 3 2 1 4 1 4 68 CLADONIA GRACILIS 87 5 1 6 0-3 4 1 2 3 4 , 1 1 2 4 69 SPHAGNUM FUSCUM 75 0 3 7 0-5 3 2 5 2 1 . 4 2 1 2 3 2 70 PTILIUM CRISTA-CASTRENSIS 75 0 3 3 0-5 1 2 3 2 5 2 1 2 2 2 1 2 71 PLAGIOTHECIUM UNDULATUM 75 0 4 2 0 - 4 4 2 4 4 4 4 4 72 DICRANUM HOWELL I I 62 5 1 8 0-3 2 2 3 2 4 . 1 2 2 2 SYNSYSTEMATIC UNIT 003Pfnus - Chamaecyparis - Gualtherla - Fauria - Sphagnum Bog Woodlands PLOT NUMBER i SYNTHETIC VALUES 1^ R W s | H M 3 | ^ H ^ | D C 3 , ^1 ST.NO. S P E C I E S P MS RS 1 S P E C I E S SIGNIFICANCE"AND VIGOR • DW DR 73 RHIZOMNIUM GLABRESCENS 50 0 + 6 0-1 1 1 + 1 . 74 BAZZANIA TRILOBATA 50 0 4 3 0-1 1 .2 + + . 75 CLADONIA IMPEXA 37 5 2 1 0-4 4 . 2 . + 76 HYPNUM CIRCINALE 37 5 4 0 0-+ + , + + 77 RHYTIDIADELPHUS TRIOUETRUS 25 0 1 3 0-3 1 2 3 . 78 DICRANUM S P E C I E S 25 0 1 2 0-3 + , 3 2 79 CLADONIA LEUCOPHAEA 25 0 + 5 0-2 2 . + 80 CLADONIA PSEUDOEVANSII 25 0 4 5 0-2 + . 2. 81 SPHAGNUM RECURVUM 25 0 4 0 0- 1 + 82 CLADONIA FURCATA 25 0 4 0 0- + + . + 83 PLAGIOCHILA ASPLENIOIDES 25 0 4 0 0- + THUJA PLICATA 25 0 4 0 0 - 4 TSUGA HETEROPHYLLA 25 0 + 0 0 - 4 + . 84 SPHAGNUM NEMOREUM 12 5 4 5 0-8 8 3 VACCINIUM PARVIFOLIUM 12 5 1 1 0-3 3 . DROSERA ROTUNDIFOLIA 12 5 4 0 0-1 1.1 85 P E L L I A NEESIANA 12 5 + 0 0-1 1 . 86 PLEUROZIA PURPUREA 12 5 + 0 0-1 1. 87 BAZZANIA TRICRENATA 12 5 4- 0 0 - 4 88 CLADONIA 12 5 + 0 0- + 89 CLADONIA MULTIFORMIS 12 5 4 0 0 - 4 + , 90 CLADONIA SQUAMOSA 12 5 + 0 0-+ 91 CLADONIA U N C I A L I S 12 5 4 0 0-* 92 HERBERTA ADUNCA 12 5 4 0 0- + + . 93 L E P I D O Z I A REPTANS 12 5 4 0 0- + + MENZ I E S I A FERRUGINEA 12 5 4 0 0-* PINUS CONTORTA 12 5 4 0 0- + L E P I D O Z I A REPTANS 37 5 4 0 0 - 4 + 94 MYLIA ANOMALA 37 5 4 0 0-* + + . HYLOCOMIUM SPLENDENS 25 0 2 0 0-4 4 2 1 . 95 CLAOPODIUM C R I S P I F O L I U M 25 0 4 0 0- + + . + DICRANUM HOWELLII 25 0 4 0 0-* + + RHIZOMNIUM GLABRESCENS 25 0 4 0 0-+ + . 96 SCAPANIA BOLANDERI 25 0 4 0 0- + RUBUS CHAMAEMORUS 12 5 4 4 0-2 2 RHYTIDIADELPHUS LOREUS 12 5 4 0 0-1 FAURIA C R I S T A - G A L L I 12 5 4 0 0- + + HERBERTA ADUNCA 12 5 4 0 0- + + 97 ICMAOOPHILA ERICETORUM 12 5 4 0 0- + 98 ISOTHECIUM STOLONIFERUM 12 5 4 0 0- + 99 PLAGIOTHECIUM LAETUM 12 5 4 0 0- + + . PLAGIOTHECIUM UNDULATUM 12 5 4 0 0- + PLEUROZIUM SCHREBERI 12 5 4 0 0- + + . 100 P T I L I D I U M C I L I A R E 12 5 4 0 0- + + . P T I L I U M CRISTA-CASTRENSIS 12 5 4 0 0- + + RHYTIDIADELPHUS TRIOUETRUS 12 5 4 0 0- + 101 ANASTROPHYLLUM MINUTUM 12 5 4 0 0- + . 1 . I • SYNSYSTEMATIC UNIT OOSPInus - Chamaecyparis - Gualtherla - Fauria - Sphagnum Bog Woodlands PLOT NUMBER 1 SYNTHETIC 1 VALUES RE IRW 291 8 HM IRW s| 35 HM IDC 2 l | 31 ST.NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR 102 ANTHELIA JURATZKANA 12 5 + 0 o- 4- 4- . 103 CALYPOGEIA FI S S A 12 5 + 0 0- 4- 4- . 104 DIPLOPHYLLUM ALBICANS 12 5 + 0 0- 4- 4- . L E P I D O Z I A REPTANS 12 5 + 0 0- 4- + . 105 MICROLEPIDOZIA SETACEA 12 5 + 0 0-+ 106 RHACOMITRIUM LANUGINOSUM 12 5 + 0 0- + SCAPANIA BOLANDERI 12 5 + 0 0- •f + . SYNSYSTEMATIC UNIT 004Thuja - Chamaecyparis - Tsuga - Vaccinium - Cornus - Rhytidiadelphus Forests PLOT NUMBER 1 SYNTHETIC 1 VALUES IRE IDC | 2| 23 1. RW3 RE IRW IRW 13.1 "1 is HM IHM 1 17| 20| ST.NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR A1 A2 A3 B1 B2 1 CHAMAECYPARIS NOOTKATENSIS 55 6 4 1 0-5 4 1 2 1 5 1 4 1 5 1 2 THUJA PLICATA 33 3 2 g 0-5 1 1 1 1 5 2 3 TSUGA MERTENSIANA 1 1 1 2 7 0-5 5 2 4 TSUGA HETEROPHYLLA 1 1 1 1 7 0-4 4 2 5 PINUS CONTORTA 1 1 1 1 1 0-3 3 2 THUJA PLICATA 88 g 5 4 0-6 5 2 5 2 5 2 6 2 5 1 5 2 3 2 6 1 TSUGA HETEROPHYLLA 88 9 3 1 0-4 2 1 3 2 3 2 4 2 2 1 1 4- 2 1 3 2 CHAMAECYPARIS NOOTKATENSIS 77 8 4 g 0-6 + 1 6 2 6 1 1 1 4 + 3 1 5 1 PINUS CONTORTA 44 4 4 8 0-7 3 2 6 1 4 2 7 2 TSUGA MERTENSIANA 33 3 2 8 0-4 4 2 3 1 4 1 TSUGA HETEROPHYLLA 100 0 4 0 1-5 2 1 3 1 4 1 2 1 1 1 4 1 2 1 5 2 4 1 THUJA PLICATA 77 8 4 4 0-5 5 1 4 2 5 1 4 2 3 1 3 2 4 1 CHAMAECYPARIS NOOTKATENSIS 77 8 3 4 0-5 1 1 5 1 4 1 1 + 1 4- 1 1 3 4-TSUGA MERTENSIANA 77 8 2 2 0-3 + 1 3 2 3 1 2 1 + 1 2 1 2 1 PINUS CONTORTA 22 2 1 0 0-2 2 1 2 2 6 ABIES AMABILIS 1 1 1 1 1 0-3 3 2 7 PICEA SITCHENSIS 1 1 1 + 0 0- + 4- + TSUGA HETEROPHYLLA 100 0 5 6 4-6 5 1 4 1 5 1 5 1 4 1 6 2 6 2 6 2 5 1 CHAMAECYPARIS NOOTKATENSIS 88 9 3 7 0-5 2 1 4 1 4 1 2 + 5 1 + + 1 1 2 2 THUJA PLICATA 77 8 4 0 0-5 4 1 3 1 4 1 2 1 5 1 1 1 4 2 8 MENZIESIA FERRUGINEA 77 8 3 9 0-5 + 3 2 5 2 1 2 2 3 3 2 5 3 TSUGA MERTENSIANA 66 7 2 6 0-4 4 1 3 1 2 1 2 1 1 2 2 2 PICEA SITCHENSIS 33 3 + 3 0-1 + 1 1 1 1 g VACCINIUM PARVIFOLIUM ABIES AMABILIS 33 22 3 2 + + 3 0 0-1 0-1 1 3 + 1 1 2 + 1 2 1 10 VACCINIUM ALASKAENSE 22 2 + 0 0-1 + 2 1 2 11 TAXUS BREVIFOLIA 1 1 1 + 3 0-2 2 1 12 VACCINIUM OVALIFOLIUM PINUS CONTORTA 1 1 11 1 1 + + 0 0 0-1 0- + + 1 1 2 13 GAULTHERIA SHALLON 100 0 7 4 2-9 8 1 4 2 6 2 2 7 2 6 2 8 2 7 2 2 1 TSUGA HETEROPHYLLA 100 0 5 4 3-7 4 1 4 + 3 1 4 + 5 1 5 2 7 2 6 2 5 1 MENZIESIA FERRUGINEA 100 0 5 0 3-6 5 2 3 2 3 2 3 2 3 2 6 2 5 3 5 2 3 2 VACCINIUM PARVIFOLIUM 100 0 4 2 2-5 2 1 2 2 5 2 5 2 2 2 2 1 4 2 4 2 2 2 VACCINIUM OVALIFOLIUM 88 g 5 5 0-8 6 2 5 2 3 2 5 2 8 2 6 2 2 2 2 1 VACCINIUM ALASKAENSE 77 8 5 8 0-8 8 3 6 2 5 2 5 2 7 2 6 2 2 2 THUJA PLICATA 77 8 2 6 0-4 + 1 4 2 2 1 3 1 2 1 1 1 2 2 CHAMAECYPARIS NOOTKATENSIS 77 8 2 3 0-4 4 1 + 1 1 1 2 1 1 1 1 1 2 + TSUGA MERTENSIANA 77 8 2 0 0-4 + + 1 1 1 4- + + 1 4- 1 4 2 PICEA SITCHENSIS 55 6 + 3 0-1 1 1 + 1 + + + + + 1 14 LEDUM GROENLANDICUM ABIES AMABILIS 44 33 4 3 + + 9 0 0-2 0-* + + + + 1 1 1 + 1 2 2 3 1 15 CLADOTHAMNUS PYROLIFLORUS 22 2 + 0 0- + + 1 + 2 16 RUBUS SPECTABILIS 22 2 + 0 0- + + 1 4- 1 17 KALMIA MICROPHYLLA 1 1 1 + 0 0- + + 1 SYNSYSTEMATIC UNIT 004Thuja - Chamaecyparis - Tsuga - Vaccinium - Cornus - Rhytidiadelphus Forests PLOT NUMBER 1 SYNTHETIC 1 VALUES IRE IDC 1 21 23 °26 "Z3 RE IRW IRW 13l| 4| 16 HM IHM I 17| 20I ST.NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR C DH 18 OPLOPANAX HORRIDUS 11 1 + 0 0- + • I 19 CORNUS UNALASCHKENSIS i 100 0 6 7 3-8 8 . 2 4 2 3 2 6 2 6 2 8 3 8 2 5 2 4 2 0 STREPTOPUS ROSEUS j 100 0 3 1 + -4 3.2 1 2 + 2 4 1 1 2 3 2 1 2 4 2 4 21 C O P T I S A S P L E N I F O L I A ' 100 0 2 9 + -3 2.2 3 2 2 2 + 2 2 2 + 2 3 2 3 2 3 22 L I S T E R A CORDATA 100 0 2 4 1-3 3.3 2 2 1 2 1 1 1 2 2 2 3 2 2 2 1 23 LYCOPODIUM CLAVATUM too 0 2 2 + -3 1 . 2 2 3 2 2 1 2 2 2 3 2 2 2 1 2 4 24 BLECHNUM SPICANT 88 9 4 6 0-6 4 . 2 4 2 1 2 3 2 6 2 3 2 5 2 3 25 RUBUS PEDATUS 88 9 3 0 0-4 2 . 2 3 2 1 2 + 2 2 1 3 2 4 2 3 26 LINNAEA BOREALIS 88 9 2 8 0-4 2 . 2 2 2 1 2 3 2 2 2 4 2 2 2 + 27 L Y S I C H I T O N AMERICANUM 77 8 4 7 0-6 1 . 2 4 2 2 2 4 3 3 3 6 2 6 28 VERATRUM VIRIDE 77 8 2 6 0-4 + .2 3 2 2 2 2 2 1 2 4 2 2 29 VACCINIUM V I T I S - I D A E A 77 8 1 9 0-3 2 . 2 1 2 1 2 1 2 3 2 1 2 1 2 3 0 C L I N T O N I A UNI FLORA 66 7 4 3 0-7 + •1 5 2 + 2 1 2 7 2 4 31 FAURIA CRISTA-GALLI 66 7 2 1 0-3 3.3 3 2 2 2 1 2 2 3 + 1 32 LYCOPODIUM ANNOTINUM 55 6 1 7 0-3 1 2 2 2 + 2 2 2 3 2 33 L I S T E R A CAURINA 55 6 1 0 0-2 1 3 + 2 + 2 2 2 4 34 CALAMAGROSTIS S P E C I E S 33 3 2 3 0-4 2 2 3 2 4 35 MAIANTHEMUM DILATATUM 33 3 2 0 0-4 4 1 4 2 2 36 O R T H I L I A SECUNDA 22 2 1 3 0-3 2 2 3 2 37 CAREX ANTHOXANTHEA 22 2 + 7 0-2 1 .2 2 2 38 CALAMAGROSTIS NUTKAENSIS 1 1 1 2 7 0-5 5 2 39 AGROSTIS AEOUIVALVIS 1 1 1 + 3 0-2 2 2 40 PTERIDIUM AQUILINUM 1 1 1 4 3 0-2 2 2 4 1 CAREX LAEV1CULMIS 1 1 1 + 0 0-1 1 42 HUPERZ1A SELAGO 1 1 1 4 0 0- 1 1 43 RUBUS CHAMAEMORUS 1 1 1 + 0 0-1 1 1 44 T I A R E L L A TRIFOLIATA 1 1 1 + 0 0-1 1 45 COPTIS T R I F O L I A 1 1 1 4 0 0- + + 1 46 LOBARIA L I N I T A 1 1 1 4 0 0 - 4 + 2 47 PLATANTHERA D I L I TATA 1 1 1 4 0 0- + *.2 48 SANGUISORBA O F F I C I N A L I S 1 1 1 + 0 0- + + 1 49 STREPTOPUS AMPLEXIFOLIUS 1 1 1 + 0 0-* 2 5 0 T O F I E L D I A GLUTINOSA 11 1 4 0 0- + + 2 51 RHIZOMNIUM GLABRESCENS 100 0 4 1 • -6 + . 4 2 2 2 + 1 2 + 3 4 6 52 RHYTIDIADELPHUS LOREUS 100 0 8 2 + -9 8.2 8 3 8 3 9 2 8 2 6 2 9 2 8 2 4 53 HYLOCOMIUM SPLENDENS 100 0 7 2 2-9 8.2 4 2 7 3 9 2 4 2 6 2 8 2 7 2 2 54 SPHAGNUM GIRGENSOHNII 10O 0 5 4 2-7 4 . 4 2 2 2 3 2 5 2 7 2 2 2 7 3 6 55 PLAGIOTHECIUM UNDULATUM 77 8 1 2 0-2 4 . 4 1 + 2 1 1 2 1 2 2 56 DICRANUM HOWELLI I 66 7 1 3 0-2 1 . 4 2 + 1 2 2 57 PLAGIOCHILA ASPLENIOIDES 66 7 + 7 0- 1 4 2 4 2 + * 1 1 TSUGA HETEROPHYLLA 55 6 1 5 0-3 4 2 4 3 2 1 THUJA PLICATA 55 6 1 0 0-2 1 2 4 2 + 4 VACCINIUM PARVIFOLIUM • 44 4 3 4 0-6 4 2 2 6 2 2 2 58 PLEUROZIUM SCHREBERI 44 4 1 9 0-3 3. + 2 3 2 2 2 59 RHYTIDIADELPHUS TRIOUETRUS 44 4 1 7 0-3 3. 1 2 2 2 2 2 6 0 L E P I D O Z I A REPTANS 44 4 + 0 0-* + . 4 2 + 2 4 61 P E L L I A NEESIANA 33 3 2 2 0-4 + 2 3 4 SYNSYSTEMATIC UNIT 004Thuja - Chamaecyparis - Tsuga - Vaccinium - Cornus - Rhytidiadelphus Forests PLOT NUMBER 1 SYNTHETIC 1 VALUES IRE IDC | 21 23 R ^ RE IRW IRW 13 I 4| '6 HM IHM I 17| 20| ST.NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR DW OR 62 SPHAGNUM PAPILLOSUM 33 3 1 9 0-4 4 . 2 1 . + . 63 BAZZANIA TRICRENATA 22 2 + 0 0- + + . 64 BAZZANIA TRILOBATA 22 2 + 0 0- + + . + .2 65 CLADONIA FURCATA 22 2 + 0 0- + + . + . 66 CLADONIA GRACILIS 22 2 + 0 0- + + . + . 67 HOOKERIA LUCENS 22 2 + 0 0- + + , 68 LOPHOZIA 22 2 + 0 0- + + . + . 2 MENZIESIA FERRUGINEA 22 2 + 0 0- + + . 2 69 PELTIGERA APHTHOSA 22 2 + 0 0-* + . + 70 DICRANUM MAJUS 1 1 1 + 3 0-2 2 2 71 PTILIUM CRISTA-CASTRENSIS 1 1 1 3 0-2 2 . CHAMAECYPARIS NOOTKATENSIS 1 1 1 + 0 0- 1 1 72 HERBERTA ADUNCA 1 1 1 + 0 0-1 1 . TSUGA MERTENSIANA 1 1 1 + 0 0-1 1 73 CALYPOGEIA INTEGRISTIPULA 1 1 1 + 0 0-* + .' 74 CLADONIA LEUCOPHAEA 1 1 1 4- 0 0- + +•. 75 CLADONIA RANGIFERINA 1 1 1 + 0 0- + 4-76 MYLIA ANOMALA 1 1 1 + 0 0- + + . PICEA SITCHENSIS 1 1 1 + 0 o-+ + .2 77 SCAPANIA BOLANDERI 1 1 1 0 0-* 4- . 78 SPHAGNUM FUSCUM 1 1 1 4- 0 0- + + . 2 79 SPHAGNUM MAGELLANICUM 1 1 1 + 0 o-* TAXUS BREVIFOLIA 1 1 1 + 0 0- + 4- . HYLOCOMIUM SPLENDENS 77 8 2 0 0-3 3. + . + . + . 3 2 1 2 . SCAPANIA BOLANDERI 77 8 1 5 0-3 + . 2 2 + . 3 2 + . 4- , RHYTIDIADELPHUS LOREUS 66 7 3 2 0-4 4 . 4 . 4 2 3. BAZZANIA TRILOBATA 44 4 + 2 0-1 + . 1 2 4- . BAZZANIA TRICRENATA 44 .4 + 0 0-+ 4- , 4- . MYLIA ANOMALA 44 4 + 0 0- + + . + . + 2 4- , DICRANUM HOWELL I I 33 3 5 0-2 4- , 2 2 4- . 80 CLAOPODIUM CRISPIFOLIUM 33 3 0 0- + 4- . 4- . LEPIOOZIA REPTANS 33 3 0 0- + 4- , + 2 4- . PLAGIOTHECIUM UNDULATUM 33 3 + 0 0- + + . 4- , RHIZOMNIUM GLABRESCENS 33 3 O 0- + 4- . 4- 2 + . HERBERTA ADUNCA 22 2 + 0 0- + 4- . + 81 ISOTHECIUM STOLONIFERUM 22 2 + 0 0- + + . LOPHOZIA 22 2 + 0 o-* 4. 2 CLADONIA GRACILIS 1 1 1 0 0-1 1 . 82 ALECTORIA SARMENTOSA 1 1 1 + 0 0- + 83 DICRANUM SPECIES 1 1 1 + 0 0- + PLAGIOCHILA ASPLENIOIDES 1 1 1 + 0 0-* 4- . 84 PTILIDIUM CILIARE 1 1 1 + 0 0- + 4- 2 RHYTIDIADELPHUS TRIOUETRUS 1 1 1 + 0 0- + 85 SPHAEROPHORUS GLOBOSUS 1 1 1 + 0 0- + 4- . VACCINIUM VITIS-IDAEA 1 1 1 + 0 0- + 4-86 DIPLOPHYLLUM ALBICANS 11 1 + 0 0- + • • • I . | • • 87 DIPLOPHYLLUM TAXIFOLIUM '1 1 + 0 0- + • • • | l • • •SYNSYSTEMATIC UNIT 004ThuJa - Chamaecyparis - Tsuga - Vaccinium - Cornus - Rhytidiadelphus Fores PLOT NUMBER 1 SYNTHETIC 1 VALUES IRE IDC 1 2\ 3 3 W IRE IRW IRW 1 I 13| < H IG| IHM \i 1 '7I IM 1 301 ST NO. S P E C I E S | P MS RS 1 S P E C I E S SIGN! [FICANCE AND VIGOR HERBERTA ADUNCA 11.1 +.0 0-+ +. SCAPANIA BOLANDERI 11.1 *.0 0-+ +. SYNSYSTEMATIC UNIT 005ThuJa - Picea - Tsuga - Lysichiton - Rhizomnium - Pe l l la F l u v i a l Forests PLOT NUMBER 1 SYNTHETIC | VALUES 1HM IHM 1 <0| 5 V ST.NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR A3 B1 B2 C 1 PICEA SITCHENSIS 66 7 4 4 0-5 5 3 3 3 I 2 TSUGA HETEROPHYLLA 33 3 2 1 0-3 3 + TSUGA HETEROPHYLLA 100 0 5 8 4-7 7 2 5 1 4 2 3 THUJA PLICATA 100 0 5 7 3-6 6 2 3 2 6 2 4 ALNUS RUBRA 66 7 4 6 0-5 4 3 5 2 5 ABIES AMABILIS 33 3 4 9 0-6 6 2 TSUGA HETEROPHYLLA 100 0 5 5 4-6 5 2 6 1 4 1 ABIES AMABILIS 33 3 3 1 0-4 4 2 ALNUS RUBRA 33 3 2 1 0-3 3 2 THUJA PLICATA 33 3 1 2 0-2 2 2 PICEA SITCHENSIS 33 3 + 0 0- + + 1 TSUGA HETEROPHYLLA 100 0 5 5 4-6 6 2 4 1 5 1 6 MENZIESIA FERRUGINEA 100 0 5 1 2-6 6 3 2 2 4 2 7 VACCINIUM PARVIFOLIUM 66 7 3 1 0-4 1 3 4 2 PICEA SITCHENSIS 66 7 1 0 0- 1 1 3 + 1 ABIES AMABILIS 33 3 2 1 0-3 3 2 TSUGA HETEROPHYLLA 100 0 5 4 2-7 7 2 2 1 4 1 VACCINIUM PARVIFOLIUM 100 0 5 3 1-6 5 3 6 2 1 2 MENZIESIA FERRUGINEA 100 0 4 9 3-5 5 3 4 2 3 2 8 GAULTHERIA SHALLON 100 0 2 5 1-3 1 1 3 1 1 1 g VACCINIUM ALASKAENSE 66 7 4 6 0-5 4 2 5 2 10 VACCINIUM OVALIFOLIUM 66 7 4 6 0-5 4 2 5 2 11 RUBUS SPECTABILIS 66 7 + 1 0- + + 2 + 1 ABIES AMABILIS 33 3 2 1 0-3 3 2 12 OPLOPANAX HORRIDUS 33 3 + 0 0-* + 1 PICEA SITCHENSIS 33- 3 + 0 0- + + 1 13 RIBES BRACTEOSUM 33 3 + 0 0- + • 2 14 CORNUS UNALASCHKENSIS 100 0 5 8 4 7 7 2 5 2 A 2 15 RUBUS PEDATUS 100 0 5 4 3-6 6 2 3 2 5 3 16 LYSICHITON AMERICANUM 100 0 5 2 4-5 5 3 5 3 4 3 17 GYMNOCARPIUM DRYOPTERIS 100 0 4 3 2-5 2 2 2 3 5 2 18 STREPTOPUS ROSEUS 100 0 4 0 + -4 4 2 + 2 4 2 19 BLECHNUM SPICANT 100 0 3 7 2-4 2 2 3 2 4 2 20 LISTERA CORDATA | 100 0 1 7 + -2 1 2 + 2 2 2 2 1 COPTIS ASPLENIFOLIA | 66 7 4 2 0-5 1 2 5 3 22 MA IANTHEMUM DILATATUM 66 7 4 1 0-5 5 2 + 2 23 TIARELLA TRIFOLIATA 66 7 1 2 0-1 1 2 1 2 24 STREPTOPUS AMPLEXIFOLIUS 66 7 1 0 0-1 + 2 1 2 25 CLINTONIA UNI FLORA 33 3 3 1 0-4 4 2 VACCINIUM PARVIFOLIUM 33 3 2 1 0-3 3 2 26 LISTERA CAURINA 33 3 + 6 0-1 1 3 27 ATHYRIUM F'tLIX-FEMINA 33 3 + 0 0-* + 1 28 HUPERZIA SELAGO 33 3 + 0 0-+ + 2 •SYNSYSTEMATIC UNIT 005ThuJa - Picea - Tsuga - Lysichiton - Rhizomnium - Pe l l la F l u v i a l Forests PLOT NUMBER 1 SYNTHET | VALUES C [HM 1HM |DC 1 1 IO| 5| 25| ST.NO. SPECIES | P MS RS 1 SPECIES SIGNIFICANCE AND VIGOR 29 POACEAE SPECIES 30 TIARELLA UNIFOLI ATA 31 VERATRUM VIRIDE 32 VIOLA GLABELLA 33 SPHAGNUM GIRGENSOHNII 34 HYLOCOMIUM SPLENDENS 35 RHIZOMNIUM GLABRESCENS 36 PELL IA NEESIANA 37 PLAGIOTHECIUM UNDULATUM 38 PLAGIOTHECIUM LAETUM 39 RHYTIDIADELPHUS LOREUS 40 POGONATUM ALPINUM 4 1 HOOKERIA LUCENS 42 PLAGIOCHILA ASPLENIOIDES 43 BAZZANIA TRICRENATA 44 PTILIUM CRISTA-CASTRENSIS TSUGA HETEROPHYLLA 45 DICRANUM HOWELLII ABIES AMABILIS 46 ATRICHUM SELWYNII 47 DICRANUM SPECIES 48 SPHAGNUM SOUARROSUM VACCINIUM PARVIFOLIUM 49 CALYPOGEIA INTEGRISTIPULA 50 DIPLOPHYLLUM ALBICANS 51 LEPIDOZIA REPTANS 52 LOPHOZIA 53 POGONATUM CONTORTUM RHYTIDIADELPHUS LOREUS RHIZOMNIUM GLABRESCENS HYLOCOMIUM SPLENDENS 54 BAZZANIA TRILOBATA SPHAGNUM GIRGENSOHNII PLAGIOTHECIUM UNDULATUM HOOKER IA LUCENS LEPIDOZIA REPTANS THUJA PLICATA DICRANUM HOWELL 11 TSUGA HETEROPHYLLA BAZZANIA TRICRENATA 55 CLAOPODIUM CRISPIFOLIUM 56 DRYOPTERIS ASSIMILIS 57 HERBERTA ADUNCA PICEA SITCHENSIS PLAGIOCHILA ASPLENIOIDES 58 SCAPANIA BOLANDERI 33 3 + 0 o-+ 2 33 3 + 0 0- + 4- 2 33 3 + 0 0- + + 1 33 3 + 0 0-* + 2 100 0 5 2 3-6 6 3 3 3 4 2 100 0 4 7 2-5 4 3 2 3 5 2 100 0 4 5 4-4 4 3 4 3 4 2 100 0 3 9 3-4 3 3 3 4 2 100 0 2 1 1-2 2 2 2 3 1 2 100 0 1 0 + - 1 1 + 4-66 7 4 2 0-5 2 3 5 2 66 7 2 1 0-3 + 3 3 66 7 1 0 0-1 + 3 1 2 66 7 1 0 0-1 + 1 2 66 7 + 1 0- + + + 33 3 3 1 0-4 4 3 33 3 3 1 0-4 4 2 33 3 1 2 0-2 2 3 33 3 + 6 0-1 1 2 33 3 6 0-1 1 33 3 + 6 0-1 1 2 33 3 + 6 0-1 1 2 33 3 6 0-1 1 2 33 3 + 0 0- + 4-33 3 0 0- + + 33 3 + 0 0- + + 2 33 3 0 0-* + 33 3 + 0 0- + + 100 0 5 7 4-6 6 3 6 3 4 too 0 5 1 + -5 5 3 5 3 + 100 0 4 0 1-4 4 3 1 3 4 66 7 3 1 0-4 4 2 + 66 7 3 1 0-4 4 3 + 66 7 1 6 0-2 2 2 1 3 66 7 + 1 0-* + 2 + 3 66 7 + 1 0- + + 2 4-66 7 + 1 0- + + 2 4-33 3 3 1 0-4 4 2 33 3 6 0- 1 1 2 33 3 + 0 0- + + 33 3 + 0 0- + 4-33 3 + 0 0-* + 33 3 + 0 0- + + 33 3 0 0- + + 1 33 3 0 0-* + 33 3 + 0 0- + + SYNSYSTEMATIC UNIT OOGAbies - Tsuga - Thuja - Vaccinium - Gymnocarpium - Rhizomnium Forests PLOT NUMBER 1 SYNTHETIC | VALUES RW 34 HM IHM loc I 18 | 19 | 32 | ST.NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR A1 A2 A3 B1 B2 C 1 ABIES AMABILIS 42 9 4 5 0-6 5 3 4 2 6 3 2 THUJA PLICATA 42 9 4 2 0-5 5 3 4 + 5 3 3 PICEA SITCHENSIS 14 3 3 1 0-5 5 3 4 TSUGA HETEROPHYLLA 14 3 3 1 0-5 5 2 ABIES AMABILIS 85 7 5 5 0-7 4 3 4 3 5 2 7 3 6 2 6 3 TSUGA HETEROPHYLLA 85 7 5 1 0-6 5 2 4 1 4 2 6 3 5 2 4 2 THUJA PLICATA 42 9 4 8 0-6 5 3 6 1 5 3 PICEA SITCHENSIS 14 3 4 1 0-7 7 3 5 ALNUS RUBRA 14 3 2 0 0-4 4 1 TSUGA HETEROPHYLLA 100 0 5 6 4-6 4 2 6 1 5 1 5 1 6 1 5 1 5 1 ABIES AMABILIS 57 1 3 7 0-5 5 2 4 2 3 2 2 2 THUJA PLICATA 42 9 2 4 0-3 3 2 3 2 3 2 6 CHAMAECYPARIS NOOTKATENSIS 14 3 2 0 0-4 4 2 PICEA SITCHENSIS 14 3 2 0 0-4 4 2 TSUGA HETEROPHYLLA 100 0 5 8 4-8 8 2 5 1 4 1 7 2 5 2 4 1 4 1 ABIES AMABILIS 71 4 3 7 0-5 4 2 2 2 5 2 3 2 1 2 7 MENZIESIA FERRUGINEA 28 6 + 7 0-2 2 2 + 2 8 OPLOPANAX HORRIDUS 28 6 1 0-1 + 2 1 2 PICEA SITCHENSIS 14 3 + 0 0-1 1 4-THUJA PLICATA 14 3 + 0 0-1 1 1 9 VACCINIUM ALASKAENSE 14 3 + 0 0- + + 2 VACCINIUM ALASKAENSE too 0 5 1 2-6 6 3 6 2 2 '1 5 2 3 1 3 1 3 1 TSUGA HETEROPHYLLA 100 0 5 1 3-6 6 2 4 1 3 1 5 1 5 2 4 1 3 1 MENZIESIA FERRUGINEA 100 0 3 6 + -5 5 3 3 1 1 1 3 2 2 1 1 2 4- 1 10 VACCINIUM PARVIFOLIUM 100 0 2 5 1-3 3 2 2 1 2 1 2 2 2 1 2 2 1 1 OPLOPANAX HORRIDUS 85 7 4 1 0-6 2 3 1 2 2 2 1 2 4 2 6 2 ABIES AMABILIS 85 7 2 2 0-3 3 2 1 1 1 2 3 2 + 2 + 1 1 1 VACCINIUM OVAI.IFOLIUM 42 9 1 7 0-3 1 2 2 1 3 1 THUJA PLICATA 42 9 + 0 0- + + 1 + + 2 PICEA SITCHENSIS 28 6 0 0-+ + 1 4 1 12 SAMBUCUS RACEMOSA 28 6 0 0- + 4- 1 + 1 13 GAULTHERIA SHALLON 14 3 1 2 0-3 3 2 14 BLECHNUM SPICANT 100 0 5 5 1-8 8 3 4 2 1 2 6 2 3 2 1 1 6 3 15 CORNUS UNALASCHKENSIS 100 0 5 0 + -7 7 3 5 2 + 2 5 2 2 2 2 2 1 2 16 RUBUS PEDATUS 100 0 5 0 2-7 7 3 4 2 2 2 4 2 3 2 3 2 4 2 17 STREPTOPUS AMPLEXIFOLIUS 100 0 1 2 • - 1 1 3 1 2 + 2 1 2 * 2 4- 2 1 2 18 ATHYRIUM FILIX-FEMINA 85 7 3 2 0-5 + 2 + 1 1 1 2 2 4- 1 5 3 19 GYMNOCARPIUM DRYOPTERIS 85 7 3 1 0-4 4 3 + 1 2 2 2 2 3 2 3 2 20 STREPTOPUS ROSEUS 85 7 2 2 0-3 3 3 1 2 1 2 4- 2 3 2 1 2 21 TIARELLA TRIFOLIATA 7 1 4 4 5 0-6 6 3 3 2 2 2 2 2 5 2 22 COPTIS ASPLENIIFOLIA 71 4 3 0 0-4 3 2 3 2 . 2 4 2 1 2 23 DRYOPTERIS ASSIMILIS 7 1 4 2 3 0-4 1 3 + 1 1 . 2 1 2 4 2 24 THELYPTERIS PHEGOPTERIS 71 4 1 9 0-2 2 . 3 1 2 2 3 2 2 2 2 SYNSYSTEMATIC UNIT 006Ab(es - Tsuga - Thuja - Vaccinium - Gymnocarpium - Rhizomnium Forests PLOT SYNTHETIC RE RE 1 DC RW HM HM DC I NUMBER VALUES 9 36 27 34 18 19 32 | ST.NO. SPECIES. P MS RS SPECIES SIGNIFICANCE AND VIGOR 25 LISTERA CORDATA 71 4 1 8 0-3 3 . 3 2.2 1 . 2 4 . 2 4 . 2 26 LYSICHITON AMERICANUM 71 4 1 8 0-3 4 . 1 . 1 2. 1 3.3 4 . 1 • 27 LISTERA CAURINA 57 1 + 4 0-1 4.3 1 .3 + . 2 + .2 28 HUPERZIA SELAGO 57 1 4 0 0- + + .3 * .2 4 . 2 4.2 29 POLYSTICHUM MUNITUM 42 9 3 8 0-5 5 3 4 . 2 4 .2 30 VIOLA GLABELLA 42 9 1 4 0-3 1 2 *.2 3.3 31 LYCOPODIUM CLAVATUM 28 6 + 0 0- + + .3 * . 2 32 MONESES UNI FLORA 28 6 + 0 0- + 4.2 + .2 33 POACEAE SPECIES 28 6 + 0 0- + 4 2 4 . 2 34 GALIUM BOREALE 14 3 1 2 0-3 3 2 35 ADIANTUM PEDATUM 14 3 + 5 0-2 2 2 .36 CIRCAEA ALPINA 14 3 4 5 0-2 2 . 2 37 CLINTONIA UNI FLORA 14 3 4 5 0-2 2 . 2 38 CAREX MERTENSII 14 3 4 0 0-1 1 2 39 LUZULA PARVIFLORA 14 3 + 0 0-1 1 2 40 MAIANTHEMUM DILATATUM 14 3 + 0 0-* 4 2 41 TIARELLA LACINIATA 14 3 4 0 0-* 4 . 2 DH 42 RHYTIDIADELPHUS LOREUS 100 0 5 2 1-7 7 . 3 7.2 3 . 2 2.2 2 1 1 . 1 2 . 43 RHIZOMNIUM GLABRESCENS 100 0 5 1 3-5 4 . 3 5 . 2 3.2 4 . 2 5 2 5.3 5. 44 HYLOCOMIUM SPLENDENS 100 0 4 9 4-6 5 . 3 6 . 2 5.3 3 . 2 4 2 4 . 1 . 45 SPHAGNUM GIRGENSOHNII 100 0 3 3 1-4 4.3 3 . 2 1 . 4 . 2 1 2 1 .2 1 . 46 PLAGIOTHECIUM UNDULATUM 10O 0 3 1 4-4 4 . 2 4 . 4 . 3 1 . 2 2 2 1 . 1 4 . ABIES AMABILIS 100 0 + 8 4 - 1 + .2 4 . 4 . 4 . 1 2 + . 2 4 . 47 PLAGIOCHILA ASPLENIOIOES 100 0 + 8 4-1 1 . 2 4 . 4 . 4 . 4 4 . 2 4 . TSUGA HETEROPHYLLA 85 7 2 0 0-3 2 . 2 4 . 4 . 3 2 2 . 2 4 . 48 PELLIA NEESIANA 7 1 4 3 3 0-5 2.3 5.2 1 2 2.2 2 . 49 POGONATUM ALPINUM 57 1 1 2 0-2 2 . 3 1 2 4 . 2 1 . 50 POGONATUM CONTORTUM 42 9 4- 2 0- 1 4 . 4 2 1 . 51 DICRANUM H0WELLI1 42 9 4 0 0 - 4 4 . 4 . 4 . 52 HOOKERIA LUCENS 42 9 4 0 0- + 4 . 4 . 4 . 53 CONOCEPHALUM CONICUM 28 6 2 0 0-3 3 2 3 . 54 STOKES I ELLA OREGANA 28 6 1 0 0-2 2 2 1 . 2 55 DIPLOPHYLLUM ALBICANS 28 6 4 7 0-2 2 . 2 4 . VACCINIUM PARVIFOLIUM 28 6 4 1 0- 1 4 . 1 . 56 PLAGIOTHECIUM LAETUM 28 6 4. 0 0- + 4 . 4 . THUJA PLICATA 28 6 4 0 0- + + . 2 4 2 57 STEREOCAULON PASCHALE 14 3 2 0 0-4 4 . 2 58 DICRANUM FUSCESCENS 14 3 4 5 0-2 2 2 59 ATRICHUM UNDULATUM 14 3 4 0 0-1 1 2 60 PELTIGERA APHTHOSA 14 3 4 0 0- 1 1 . 2 61 SCAPANIA BOLANDERI 14 3 4 0 0- 1 1 . 2 62 BLEPHAROSTOMA TRICHOPHYLLUM 14 3 4 0 0- + 4 . 63 BRACHYTHECIUM ASPERRIMUM 14 3 4 0 0-* 4 . 64 BRACHYTHECIUM FRIGIDUM 14 3 4 0 0 - 4 4 2 65 CLADONIA BELLIDIFLORA 14 3 4- 0 0- + 4 . 66 DICRANELLA HETEROMALLA i 14 3 + 0 0- + 4 67 DICRANUM MAJUS i 14 3 4 0 0- + 4 . 68 HYPNUM CIRCINALE ' 14 3 + 0 0- + 4 . SYNSYSTEMATIC UNIT OOSAbles - Tsuga - Thuja - Vacclntum - Gymnocarpium - Rhizomnium Forests PLOT NUMBER 1 SYNTHETIC 1 VALUES IRE IRE 1 9\ 36 DC 27 RW 34 HM |DC I 19| 321 ST.NO. S P E C I E S | P MS RS 1 S P E C I E S SIGNIFICANCE AND VIGOR DW DR 69 LEPIDOZIA REPTANS 14 3 + 0 0- + + . 70 LEUCOLEPIS MENZIESII 14 3 + 0 0- + + . 71 LOPHOZIA 14 3 + 0 0- + MENZIESIA FERRUGINEA 14 3 0 + . 72 MICROLEPIDOZIA SETACEA 14 3 + 0 0- + + . OPLOPANAX HORRIDUS 14 3 + 0 0- + + . PICEA SITCHENSIS 14 3 + 0 0-+ + . 73 POLYTRICHUM FORMOSUM 14 3 + 0 0- + + . RHYTIDIADELPHUS LOREUS 100 0 4 7 +-5 5. 4 . 5. 5 . + . SCAPANIA BOLANDERI 100 0 1 7 +-3 3.2 4- . + . 2 . + . + . + . RHIZOMNIUM GLABRESCENS 85 7 4 1 0-5 2 . 5 . 4 . + . 4 . 4 . PLAGIOTHECIUM UNDULATUM 85 7 + 7 0-1 +. 1 . + . + . + . 1 + . DICRANUM HOWELLII 71 4 + 5 0-1 • . + . 1 . 4- . * . LEPIDOZIA REPTANS 57 1 1 4 0-3 +. + . 3. + . 74 BAZZANIA TRICRENATA 57 1 + 4 0- 1 1 . + . + . + . 75 MYLIA ANOMALA 57 1 + 0 0- + + . + . HYLOCOMIUM SPLENDENS 42 9 1 6 0-3 2 . 3. 76 CLAOPODIUM CRISPIFOLIUM 42 9 -*• 0 0- + + . + . + . PLAGIOCHILA ASPLENIOIDES 42 9 + 0 0- + + . 4- . + . THUJA PLICATA 42 9 + 0 0- + + . + . + . HOOKER I A LUCENS 28 6 + 1 0-1 1 . + . 77 BAZZANIA TRILOBATA 28 6 + 0 0- + + . PLAGIOTHECIUM LAETUM 28 6 0 0-+ + . + , SPHAGNUM GIRGENSOHNII 28 6 + 0 0- + + . + . 78 ATRICHUM SELWYNII 14 3 + 0 0- + + . DICRANUM FUSCESCENS 14 3 + 0 0- + + . 79 LOBARIA LINITA 14 3 + 0 0- + + . POGONATUM ALPINUM 14 3 + 0 0- + 80 POLYTRICHUM COMMUNE 14 3 + 0 0-* + . STOKESIELLA OREGANA 14 3 + 0 o-* TSUGA HETEROPHYLLA 14 3 + 0 0- + + . ATRICHUM SELWYNII 14 3 + .0 0- + + . BAZZANIA TRICRENATA 14 3 + .0 0- + + . DIPLOPHYLLUM ALBICANS 14 3 + .0 0- + + . MICROLEPIDOZIA SETACEA 14 3 + .0 0-* + , PLAGIOTHECIUM LAETUM 14 3 + .0 0- + + . POGONATUM ALPINUM 14 3 .0 0- + + . Appendix VI - cont'd Vegetation Summary Table SYNSYSTEMATIC UNITS 1 I 2 1 3 I * 5 1 G 1 SPECIES PRESENCE CLASS AND MEAN SPECIES SIGNIFICANCE ABIES AMABILIS II + 0 II +.0 II 4.9 V +.8 ADIANTUM PEDATUM I * . 5 AGROSTIS AEOUIVALVIS IV 3.0 I I + 0 I *.3 ALECTORIA SARMENTOSA I +.0 ALNUS RUBRA IV 4.6 I 2.0 ANASTROPHYLLUM MINUTUM I + 0 ANDREA RUPESTRIS V +.5 ANDROMEDA POLYFOLIA V 4 . 1 V 2 . 4 I + 0 ANTHELI A JURATZKANA V * 5 I + 0 ANTITRICHIA CURT IPENOULA I • .0 II • .0 ATHYRIUM FILIX-FEMINA V 3.2 ATRICHUM SELWYNII I I + .6 I + .0 ATRICHUM UNDULATUM 1 * .0 BAZZANIA TRICRENATA , * .0 V +.5 I + 0 III * .0 IV + . 1 III +.4 BAZZANIA TRILOBATA I • .0 III + 3 III +.2 IV 3 . 1 II +.0 BLECHNUM SPICANT I + .0 V + . 5 IV 2 1 V 4 . 6 V 3 . 7 V 5.5 BLEPHAROSTOMA TRICHOPHYLLUM I * .0 I +.0 BRACHYTHECIUM A5PERRI MUM I * .0 BRACHYTHECIUM FRIGIDUM I * 0 CALAMAGROSTIS NUTKAENSIS I + 0 I 2.7 CALAMAGROSTIS SPECIES I 1 9 II 2.3 CALYPOGEIA FISSA I + 0 CALYPOGEIA I NTEGRI ST IPULA I +.0 II * .0 CAREX ANTHOXANTHEA II 3 3 II +.7 CAREX LAEVICULMIS I * .0 CAREX LIVIDA V 3.3 I + 0 CAREX MERTENSII I * .0 CAREX PAUCIFLORA I • .0 I I 3 0 CAREX PLURIFLORA I I 4.5 V 4 5 CAREX ROSTRATA I + . 1 CAREX SITCHENSIS I t .0 11 2 0 CAREX STYLOSA I 1 .0 CETRARIA ISLANDICA I + .0 CHAMAECYPARIS NOOTKATENSIS I I 2.5 V 6 . 5 V 8 6 V 3 . 7 I 2.0 CIRCAEA ALPINA I * . 5 CLADONIA I I * .0 V 1.8 I + 0 CLAOONIA AMOUROCRAEA V 5 .5 CLADONIA BELLIDIFLORA I * .o I • .0 CLADONIA FURCATA I I I 1 . 1 II + 0 II +.0 CLADONIA GRACILIS V 2.9 V 2.4 V 1 6 II • .0 CLADONIA IMPEXA V 5.0 II 2 1 CLADONIA LEUCOPHAEA 11 I . 5 11 + 5 I + .0 CLADONIA MULTIFORMIS I + 0 CLADONIA PSEUDOEVANSII 11 * .0 11 + 5 CLAOONIA RANGIFERINA IV 3.6 V 3 . 4 V 2 .0 1 * .0 CLADONIA SQUAMOSA I • .0 I + .0 CLADONIA SYLVATICA I 1 .8 V 7 . 5 SYNSYSTEMATIC UNITS 1 | 2 | 3 | 4 | 5 | 6 | PRESENCE CLASS AND MEAN SPECIES SIGNIFICANCE CLADONIA TENUIS I + 1 CLADONIA UNCIALIS V 4 0 I + 0 CLADOTHAMNUS PYROLIFLORUS II 4 0 CLAOPODIUM CRISPIFOLIUM II 4 0 I I 4 0 I I 4 0 I I I 4 0 CLINTONIA UNIFLORA I 4 0 IV 4 3 II 3 1 I 4 5 CONOCEPHALUM CONICUM II 2 0 COPTIS ASPLENIFOLIA II + 4 V 3 4 V 2 9 IV 4 2 IV 3 0 COPTIS TRIFOLI A V 3 9 V 2 4 V 2 2 I 4 0 CORNICULARIA ACULEATA II 4 4 CORNUS UNALASCHKENSIS V 4 1 V 2 4 V 5 1 V 6 7 V 5 8 V 5 0 DICRANELLA HETEROMALLA I 4 0 DICRANUM FUSCESCENS V 1 5 I 4 5 DICRANUM HOWELLII I 4 2 IV 1 8 IV 1 3 II 3 1 IV 4 5 DICRANUM MAJUS I 4 3 I 4 0 DICRANUM SPECIES I 4 0 II 1 2 I 4 0 11 4 6 DICRANUM UNDULATUM I 4 9 DIPLOPHYLLUM ALBICANS V 4 5 I + 0 I 4 0 11 4 0 11 4 7 DIPLOPHYLLUM TAXIFOLIUM I 4 0 DODECATHEON JEFFREYI I 4 9 DROSERA ANGLICA IV 3 0 OROSERA ROTUNDIFOLIA V 3 6 IV + 8 DRYOPTERIS ASSIMILIS 11 4 0 IV 2 3 EMPETRUM NIGRUM V 4 4 V 7 5 V 3 1 ERIGERON PEREGRINUS I 4 0 I + 4 ERIOPHORUM ANGUSTIFOLIUM V 5 5 II 2 S FAURIA CRISTA-GALLI III 3 3 v 2 4 V s.o| IV 2 I I GALIUM BOREALE I 1 2 GAULTHERIA SHALLON V 5 5 V 7 0 V 7 4 V 2 5 I 1 2 GENT I ANA DOUGLASIANA V 3 1 II + 0 GEUM CALTHIFOLIUM I 4 9 GYMNOCARPIUM DRYOPTERIS V 4 3 V 3 1 GYMNOCOLEA INFLATA III 1 4 HERBERTA AOUNCA I 4 0 V 4 5 I 4 0 II 4 0 11 4 0 HOOKERIA LUCENS II 4 0 IV 1 0 111 4 0 HUPERZIA SELAGO II 4- 3 I 4 0 II 4 0 111 4 0 HYLOCOMIUM SPLENDENS V 2 4 V 5 3 V 7 2 V 4 7 V 4 9 HYPNUM CIRCINALE I 4 0 V + 5 II 4 0 I 4 0 ICMADOPHILA ERICETORUM v 4 5 I 4 0 1 ISOTHECIUM STOLONIFERUM I 4 0 II 4 0 I JUNIPERUS COMMUNIS v 5.0| V 5 5 II 4 5 1 KALMIA MICROPHYLLA I v 4 6| V t 5| V 3 3| I 4 o| 1 LEDUM GROENLANDICUM I v 3 5| V 1 5| V 4 01 111 4 9| 1 SYNSYSTEMATIC UNITS 4 | 5 | 6 | SPECIES PRESENCE CLASS AND MEAN SPECIES SIGNIFICANCE LEPIDOZIA REPTANS 11 4 0 111 4 0 IV 4 1 111 1 4 LEUCOLEPIS MENZIESII - I 4 0 LINNAEA BOREALIS V 4 5 V 3 6 V 2 8 LISTERA CAURINA III 1 0 11 4 6 III 4 4 LISTERA CORDATA V 1 0 V 2 4 V 1 7 IV 1 8 LOBARIA LINITA I 4 0 I 4 0 LOISELEURIA PROCUMBENS I I t 0 V 4 5 LOPHOZIA I 1 4 V 4 5 11 4 0 I I 4 0 I 4 0 LUZULA PARVIFLORA I 4 0 LYCOPODIELLA INUNDATA I 4 0 LYCOPODIUM ANNOTINUM II 4 0 V 2 4 III 1 7 LYCOPODIUM CLAVATUM II 4 0 V 4 5 V 2 7 V 2 2 11 4 0 LYCOPODIUM OBSCURUM I + 0 LYCOPODIUM SITCHENSE I 4 0 LYSICHITON AMERICANUM I 4 9 IV 3 7 IV 4 7 V 5 2 IV 1 8 MA IANTHEMUM DILATATUM I 4 1 111 1 7 11 2 0 IV 4 1 I 4 0 MENYANTHES TRIFOLI ATA I 4 0 MENZIESIA FERRUGINEA V 3 2 V 5 0 V 5 1 V 3 6 MICROLEPIDOZIA SETACEA V 4 5 I 4 0 I 4 0 MONESES UNI FLORA II 4 0 MYLIA ANOMALA II 4 0 V 4 5 II 4 0 111 4 0 III 4 0 MYLIA TAYLORI I 4 0 MYRICA GALE I I 5 t 111 3 6 ODONTOSCHISMA DENUDATUM I 4 0 • V 4 5 OPLOPANAX HORRIDUS I 4 0 II 4 0 V 4 1 ORTHILIA SECUNDA 11 1 3 PARALEUCOBRYUM ENERVE V 4 5 PEDICULARIS I 4 0 PELLIA NEESIANA I 4 0 I 4 0 11 2 2 V 3 9 IV 3 3 PELTIGERA APHTHOSA II 4 0 I 4 0 PICEA SITCHENSIS I 4 0 111 4 3 IV 4 4 II 4 0 PINGUICULA VULGARIS I 4 0 11 4 S PINUS CONTORTA - CONTORTA IV 3 8 V 2 4 V 5 4 111 4 8 IV PLAGIOCHILA ASPLENIOIDES 11 4 O IV 4 7 1 0 V 4 8 PLAGIOTHECIUM LAETUM I 4 0 V 1 0 II 4 0 PLAGIOTHECIUM UNDULATUM I 4 0 IV 4 2 IV t 2 V 2 1 V 3 1 PLATANTHERA DILI TATA II 1 0 I 4 0 I 4 0 PLATANTHERA STRICTA I 4 0 11 4 0 PLEUROZIA PURPUREA I 4 0 I 4 0 PLEUROZIUM SCHREBERI III 3 4 V 4 5 V 5 4 III 1 9 POACEAE SPECIES I 4 0 II 4 0 II 4 0 POGONATUM ALPINUM IV 2 t III 1 2 POGONATUM CONTORTUM II 4 0 III 4 2 POLYSTICHUM MUNITUM III 3 8 POLYTRICHUM COMMUNE I 4 0 POLYTRICHUM FORMOSUM I 4 0 POLYTRICHUM JUNIPERINUM II 4 0 SYNSYSTEMATIC UNITS I 3 | 4 | 5 | 6 | SPECIES PRESENCE CLASS AND MEAN SPECIES SIGNIFICANCE PTERIDIUM AOUILINUM I . 1 I I I + 3| I I PTILIDIUM CILIARE PTILIUM CRISTA-CASTRENSIS I I + + .0 .0 IV + 3 .0 .3 I + .0 .3 II 3 . 1 RHACOMITRIUM LANUGINOSUM V 4 .9 V 7 . 5 I + .0 RHIZOMNIUM GLABRESCENS III + S V 4 1 V 5 . 1 V 5 . t RHYNCHOSPORA ALBA III 5 1 RHYTIDIADELPHUS LOREUS V 2 4 V 4 .6 V 8 2 V 5 7 V 5 .2 RHYTIDIADELPHUS TRIOUETRUS 11 i . 3 III 1 7 RIBES BRACTEOSUM I I *- 0 RUBUS CHAMAEMORUS III 2 4 IV 1 .8 I + 0 RUBUS PEDATUS I + 0 V 3 0 V 5 4 V 5 0 RUBUS SPECTABILIS II + 0 IV + 1 SAMBUCUS RACEMOSA 11 4- 0 SANGUISORBA OFFICINALIS V 5 6 V 2 4 V 3 3 I + 0 SCAPANIA BOLANDERI II + 0 IV 1 5 11 + 0 V 1 7 SCHEUCHZERIA PALUSTRIS I + 0 SELAGINELLA I + 0 II + 0 SIPHULA CERATITES V 4 5 V + 5 SPHAEROPHORUS GLOBOSUS I + 0 SPHAGNUM COMPACTUM III 3 6 SPHAGNUM FUSCUM IV 3 4 IV 3 7 I + 0 SPHAGNUM GIRGENSOHNII V 6 1 V 5 4 V 5 2 V 3 3 SPHAGNUM IMBRICATUM I 3 5 SPHAGNUM LINDBERGH III 4 t SPHAGNUM MAGELLANICUM I 4 0 I + o SPHAGNUM NEMOREUM I 4 5 SPHAGNUM PAPILLOSUM V 5 8 V 3 8 II 1 9 SPHAGNUM RECURVUM I 3 4 11 + 0 SPHAGNUM RUBELLUM V 4 5 V + 5 V 4 0 SPHAGNUM SOUARROSUM II + 6 SPHAGNUM TENELLUM V 5 0 STEREOCAULON PASCHALE V 1 5 I 2 0 STOKES I ELLA OREGANA 11 1 0 STREPTOPUS AMPLEXIFOLIUS I + 0 I + 0 IV 1 0 V 1 2 STREPTOPUS ROSEUS 11 + 3 V 3 1 V 4 0 V 2 2 TAXUS BREVIFOLIA I + 3 THELYPTERIS PHEGOPTERIS IV 1 9 THUJA PLICATA II 1 9 V + 5 V 4 2 V 5 4 V 5 7 III 4 8 TIARELLA LACINIATA I + 0 TIARELLA TRIFOLIATA I + 0 IV 1 2 IV 4 5 TIARELLA UNIFOLIATA 11 + 0 TOFIELDIA GLUTINOSA IV t 9 IV 2 7 I + 0 TRICHOPHORUM CESPITOSUM V 7 t V 4 5 IV 5 t TRIENTALIS EUROPAEA V 4 5 V 2 1 TSUGA HETEROPHYLLA 111 + 0 V 3 4 V 5 6 V 5 8 V 5 8 TSUGA MERTENSIANA V + 5 IV 1 8 IV 2 2 VACCINIUM ALASKAENSE |III 3 IV 5 8 IV 4 G| V 5 'I SYNSYSTEMATIC UNITS 1 | 2 | 3 | 4 | 5 | 6 | SPECIES PRESENCE CLASS AND MEAN SPECIES SIGNIFICANCE VACCINIUM CAESPITOSUM II 4- 0 V 1 5 IV 3 0 VACCINIUM OVALIFOLIUM IV 2 9 V 5 5 IV 4 6 III 1 . 7 VACCINIUM OXYCOCCOS V 3 9 III + 6 VACCINIUM PARVIFOLIUM V 2 7 V 4 2 V 5 3 V 2.5 VACCINIUM ULIGINOSUM V 3 2 V 6 5 III 1 3 VACCINIUM VITIS-IDAEA V 2 4 V 3 3 IV 1 9 VERATRUM VIRIDE II I 0 IV 2 S II + 0 VIOLA GLABELLA II + 0 III 1 .4 227A APPENDIX VII Data Matrix f o r the F i r s t A x i s RA Releve O r d i n a t i o n s 228 A p p e n d i x V I I . Rea ranged d a t a m a t r i x based on t h e o r d e r i n g o f r e l e v e s and s p e c i e s by t h e f i r s t a x i s RA o r d i n a t i o n . The communi ty t y p e s a r e d e s i g n a t e d as i n A p p e n d i x V I I I . O © • , A ,0x0 124323012130(062003 23 lb223 10112100 3223113 180928 744206|l|79 33 5 115J2 363 34670(0596754892 1 4 5 CETR I SL 167 SPHA 1MB 58 DROS ANG 1 1 4 SPHA TEN 86 RH YN AL3 1 1 7 S I P r i CfcR 142 SPHA COM 1 4 6 CORN ACU 1 6 8 POLY JUN 143 GYMN INF 119 C L A D UNC 1 1 0 RH AC LAN 1 1 3 SPHA L IN 67 CARE L I V 6 9 AN OR POL 31 JUN I COM 59 GENT DOU 107 C A R E STY 1 1 6 C L A D IMP 63 ER IO ANG 81 L O I S PRO 133 CLAO SYL 57 DROS ROT 61 T R I C CES 52 V A C C OXY 55 AGRO AEQ 1 5 0 P INU CON 1 3 0 C L A D 1 3 5 0 3 ON OEN 3 4 V A C C ULI 54 SANG OFF 45 MYRI GAL 64 T R I E EUR 152 P L E U PUR 1 3 4 C L A O RAN 82 EMPE NIG 56 COPT TRI 1 5 0 C L A O PSE 112 SPHA RUB 33 KALM MIC 60 P L A T OIL 1 4 9 C L A O LEU 171 SPHA ReC 1 2 0 C L A O GRA 138 MYL1 AND 44 P INU CON 111 SPHA PAP 1 1 8 C L A O FUR 1 1 5 SPHA FUS 55 R JBU CHA 84 T J F I GLU 99 CARE S IT 32 L E O U GRO 144 C L A D BEL 1 6 9 C L A O SOU 89 P ING VUL 65 CARE PLU r l --1--1 -—65 51231-311-1 75554232331 -57764-1—4 744452311-4 44 1 3 6 -1--11 1 — 11 1— -123-1—1-2 52334251-41 573445511-3 —43 7--2 4-23| 13342343322 425432444124)2|1 375545 42-44-311221512112 1 3 43554432383 82436583353--22-1 -3--4 51223143311 55664688 8 872|4|854 112311421117 423422-12-2-1-1-1111 -111 1 1 1-1133123434 74764485444--856-7 321222432428(-jll3 323332 6 1-1223423-11341154 111-.-- 1 2--1 1 21131111-3232-1 111] 232274422122 2521221311212 12 211113533517153521111! 1344245336251(334311331 31-11 —23 1 32111151 111-1-1 1 1411343—63-412215797228H4442 3-1—1-11 1-21151125— --2--421 -12-111-133-1 13 1244142115— -1- - -1 — 1 3—29| 5|2-1 I 1 42—1 — 4—3-3-1 11—1111 - 2 - -11—11-11 1 1--1 62 21 1 |2|434-2123 5—4-1-4 -1113 |2|33242411| 4312 - 1 1 3 5-1 41-3] 12-31-2 [2 24-1-1 3| 41 1|3425 3 322|1112 1-2 1317212431 2--1 ' h l -1-1 •1 h-4— 1 h - 1 -— 1 h - 1 -11 •1-229 62 CARE PAU 101 ER IG PER 128 HERB AOU 100 PLAT su 140 LOPH 104 SEL A 83 VACC CAE 25 PI NU CON 136 HYPN CIR 68 FAUR CRI 131 PLEU SCH 172 CHAM NOD 170 SPHA MAG 151 01CR SPE 35 CHAM NOO 155 81EP T RI 87 LYCO ANN 137 BAZZ Trt I 36 THUJ PLI 85 HUPE SEL 70 VACC VIT 148 PTIL CRI 139 MICR SET 21 CHAM NOO 76 LfCO CL A 53 CORN UNA 6 PINU CO^ 71 LI NN BOR 154 DICR HOW 20 THUJ PLI 90 CARE ANT 38 TSUG MER 43 GAUL SHA 23 TSUG MER 103 MA I A OIL 72 COPT ASP 153 RH it TRI 147 TSUG HET 132 0 ICR rUS 121 SPHA Girt 74 VERA V H 123 HYLO SPL 73 LYSI A ME 122 RH rT LOR 175 MENZ F ER 105 CALA SPE 19 PI NU CON 105 CALA NUT 15 TSUG MER 141 DIPL AL3 126 LEPI RE? 37 TSUG HET 8 CHAM NOO 41 VACC OVA 42 VACC PAR 50 CLAD PYR 79 LIST COR 39 MENZ FER 16 CHAM NOO 88 POAC SPE 129 ST ER PAS 1243 2301 2 13010132003 23 1D22 3 I 0112100 3 223 I). 3 130928 74420617933 51152 363 3467 00596754892 -i H -4-u-- l - l — U 1 -1 41—11 1 3-114-- 1 U - U - - 7 1 1-1 1-21-5---1 1-1-1 —431 -1-2-1 1 Hi 1 1 II --1-1 13-3-3233| 253457652 11 — - l - l 45[24781 435813 32 1|33673733|3 6te8998976r4H-21l2 -1— 3 3-22112 111 1-1 -1 - -1344253331 213243231321113 135-121-1 11 1-32 3-1 12 11-11 1-— 1 —I 1-15-H 1-h i -l -4-121-23— I "1— 1 1 42 3212-1 -11 £ H 924< 12454544 112411 -53 623 3|2|3 47 3 44 45[8 4366885 4|757 51 5755442 H 1 2 4 4 2 3 2 4 2 2 1 3 2 231-12-1 1 138-77751 11- -2- -1-1-1412'. 1-1--12-1 -13 2-| 1 - . •425112 •13|122123211 1  -3647 '-421 -111 11222 |34523244U»342514-45|l 2 -14-'687 4322-12 1111-1-2311111-5|256876778 4697  7 2|13 3—1 12-2-13 H 3-221 13-1425 2 2 2 2 5 1 2 3 2 1 2 1 3 3 3 133122-h l l 321 •t ti 132-21-42 |2|12573735)8479468724256553111 31243 5142-43-665511243-1' 2|l 5453344|88898698l-4|-23 4f-2 2-5-2 1 1 3 3 - 2 1 2 2 -45543 4—125 — 3-11—11 131 3243414434 3|166-1435 65358622! 231 2233)225 52 2442|5632 21 2 221 11 I 3 2 l l l 2 3 2 l | l l 3 2 - 2 1 U •2114342 53333 6553|5453 13 3211 -21541-113-— h i 1-1 -1 - I 11 1-11— -11-1- -11-•4 • 1 1-45221 1-1-1 1 1-1— 133 154-1-4-2-1-132 1 » 2 3 5 7 2 7 6 | 6 3 4 3 1 4 4 l l l 1 •277352212 --2-11 1 1 55765172643' 230 124323012 130(0132003 23 l|022 3L 01121100 322 3 113 180928 744205 I 151 THUJ P L I 159 V A C C PAR 13 T H U J PLI 10 T S U G MER 40 V A C C ALA 2 CHAM N33 125 PL AG UNO 102 OR TH SEC 75 DL EC SP1 155 P E L T APH 7 T H U J P L I 22 T S U G H E T 77 C L IN UNI 4 7 P I C E S I T 156 PL AG ASP 27 P I C E S I T 29 V A C C ALA 24 MENZ F E R 1 7 4 C A L Y I N T 78 STRE ROS 1 T H U J P L I 1 6 3 SCAP BOL 127 OICR MAJ 46 A B I E A MA 91 L I S T CAU 1 6 4 P E L L NEE 3 0 V A C C PAR 124 R H I Z GLA 4 9 RUBU SPE 1 5 7 HDO< L U C 80 RUBU PEL) 14 T S U G HET 92 STRE AMP 9 T S U G HET 176 P I C E S I T 18 P I C E S I T 4 P I C E S I T 165 PL AG L AE 17 A B I E AM A 3 T S U G HET 95 MONE UNI 12 AL NU RUB 26 A B I E AMA 162 POGO A L P 96 G YMN DRY 158 A B I E AHA 98 T I AR TRI 11 Ad I E AM* 173 POGO CON 97 T H E L PHE 48 O P L O HOR 94 ORYO ASS 9 3 A T H Y r L 178 STCK ORE 5 A B I E AMA 28 O P L O HOR 109 V I C L GLA 108 P O L Y HUH 51 S AM8 RAC 177 CLINO CON 1-1-- l - h J-65 .485 457544 1}~-—4 2 I — 1 - 1 - - 1 -- l l l l l l l l l 1 1 -235162 421--1 .-41131 -1—5 4— 5 -1 L|—3 113311-L —11 — 11 — I .332—45122 114 h~ 1-1-1 ill 42 1113464445344 555 79 33 51132 3633467 00596 75 4892 3—3-5486557-— 2-• l i - i i i i i : • 1 - l h l l 2111—1 1-— 3-12 62 111 •1 - --12 545-43 34-2-3 3 3 -• 4-3—4--62-4-66255333 4-2545-11 11-12-12211411214 -23-23- 1-13|441-363532384146 316 1 1 1 • 12 31-555655366356-11313125455466656' 1 1-5117-1-—1-11 — 111-1 - I l l - I l l - -l | - l 1-11-— I 1-1 — | - 1 3 5 1-i 1 •1-131111314414131-21 1-5-1-1 1 If-11 ll--11—12 -11-l ( - l 3-1 1 11 — I 1 1|2 31-1234 31637 342123755444 1 123 1-1 I 1 1 111 11 4254334 1|2 342142 54|5546545 555 11111111 654 1 1 — 4 53—5 111—11 --5-4432-3 5 - - - 1 1 45 4— • 2-35-31 1132 111 1224- 152233 -11111111 16—13225 -4-465766 11 l - l -2 1222 -2-112145 -111- 1 — 4 -11-11215 + — 21-5 46 1 —I 1 113 544 1-1 -3-3 230A APPENDIX VIII Environment Summary Table f o r the F o r t y Releves 231 APPENDIX V I I I . E n v i r o n m e n t a l d a t a f o r the 40 r e l e v e s . E x p l a n a t i o n o f codes used i n the t a b l e o f e n v i r o n m e n t a l d a t a . Bed rock I Me i g n e o u s b e d r o c k ( m o s t l y " qua r t z d i o r i t e ) m e t a m o r p h i c . b ed rock ( m o s t l y s h i s t s and g n i e s s e s ) L and fo rm Ov o r g a n i c v e n e e r ( l e s s t han lm t h i c k ) ' Ob o r g a n i c b l a n k e t ( g r e a t e r t h a n lm t h i c k ) Sv s a p r o l i t e (decomposed r o c k ) v e n e e r Fv f l u v i a l v e n e e r Fb f l u v i a l b l a n k e t Cv c o l l u v i a l v e n e e r R bed rock i e : Ov Sv/R o r g a n i c v e n e e r o v e r s a p r o l i t e v e n e e r o v e r b e d r o c k S o i l Subgroup ( C a n a d i a n S o i l S u r v e y Commit tee 1978) T Y . F T y p i c F i b r i s o l ME. F M e s i c F i b r i s o l TY.M T y p i c M e s i s o l F I .M F i b r i c M e s i s o l HU.M Humic M e s i s o l T .M T e r r i c M e s i s o l TY.HU T y p i c Humiso l T.HU T e r r i c Humiso l TY.FO T y p i c F o l i s o l T .FO T e r r i c F o l i s o l ( T r o w b r i d g e 1981) 0 . F 0 Organo F o l i s o l ( T r o w b r i d g e 1981) O.R O r t h i c Regoso l CU.R C u m u l i c Regoso l 232 A p p e n d i x V I I I - c o n t ' d GL.R G l e y e d R e g o s o l ' O.HP O r t h i c Humic P o d z o l O.FHP O r t h i c F e r r o - H u m i c P o d z o l GL .FHP G l e y e d Fe r ro-Humic P o d z o l O.HFP O r t h i c H u m o - F e r r i c Pod. R.G Rego G l e y s o l R.HG Rego Humic G l e y s o l T e x t u r e o r d e c o m p o s i t i o n T e x t u r e ( m i n e r a l s o i l s o n l y ) S sand LS loamy sand SL sandy loam L loam SCL sandy c l a y loam FSCL f i n e sandy c l a y loam S i s i l t D e c o m p o s i t i o n ( o r g a n i c s o i l s o n l y ) Of f i b r i c p e a t Om m e s i c p e a t Oh humic p e a t Humus f o rm ( K l i n k a e t a l . 1981) 0 . HR O r t h i h e m i m o r M. HR Mycohemimor A . HR Amphihemimor 0 . HUR O r t h i h e m i h u m i m o r R. HUR Res i duohemihumi mor 0 . UR O r t h i h u m i m o r R. UR Res iduohumimor A . UR Amphihumimor H. YR Hemihydromor S. YR S a p r i h y d r o m o r 233 A p p e n d i x V I I I - c o n t ' d 0. IR O r t h i h i s t o m o r F. IR Fermi h i s t o m o r 0. ID O r t h i h i s t o m o d e r S. ID S a p r i h i s t o m o d e r 0. SL O r t h i s a p r i m u l1 Water T a b l e S A s a t u r a t e d t o s u r f a c e a b s e n t Symbols e n c l o s i n g each r e l e v e number deno te community t y p e s as f o l l o w s : O E r i c a c e o u s s h r u b - T r i c h o p h o r u m  Sphagnum open bogs Empretrum - V a c c i n i u m u l i g i n o s u m  R h a c o m i t r i u m d r y o u t c r o p s • A P i n e - c y p r e s s - c e d a r - G a u l t h e r i a -F a u r i a bog wood lands Ceda r - c y p r e s s - hemlock - V a c c i n i u m  Cornus z o n a l u p l a n d f o r e s t s O Ceda r - s p r u c e - hemlock - L y s i c h i t o n a l l u v i a l f o r e s t s A m a b i l i s f i r - hemlock - c e d a r - Gymnocarp ium u p l a n d f o r e s t s Append ix V I I I - c o n t ' d . T a b l e o f e n v i r o n m e n t a l d a t a f o r t h e f o r t y r e l e v e s ( p a r t 1) Releve # % (22) ® (24) 0 29 H m H 21 31 15 Slope {%) 5 8 2 0 2 5 8 0 10 0 36 0 100 13 15 8 5 18 0 15 25 Upslope dist.(m) 50 10 0 0 0 50 0 0 0 100 175 0 0 30 12 20 0 300 0 300 350 Bedrock I I I I He I I I Me I Me Me I I I I Me I Me Me Me Landform Ob R 0b R Ob Sv/R Ob Sv/R Ov R Ov-b R Ob R Ob R Ob Sv/R Ob F? Ov Sv/R Ob F? Sv R Ov-b R Ob R Ov-b Sv/R Ob ? Ov-b R Ob F Ov-b R Ov R Soil Subgroup HIJ H HU H FI M TY M T H TY M HU H TY F HE F HU H 0 HR TY F 0 R HU M TY M TY M TY F HU M HU M TY M TY/TY Hi/ FO Texture or decomp. 0m 0m Om Om Om Om Om Of Of Om Oh S Of SI Om Om Om Of Om Om Om Oh/SCL Coarse frag.(%) - - - - - - - - - - - - 50 - - - - - - - -Min s o i l depth (cm) - - -> ? - - - - - 14 -7-28 - - ? - - - - -0 horizon depth (cm) 100-270 100-250 180 100-250 40-90 50-240 100-375 100-380 150-200 200 30-40 300+ -50-160 170 25-200 110+ 50-220 195 50-100 0-70 Humus form 0 ID 0 ID 0 IR 0 IR F IR 0 IR 0 ID 0 IR 0 IR 0 IR 0 ID 0 IR 0 HR F IR F IR 0 ID 0 IR 0 ID 0 SL F IR 0 HUK I LFH horizons depth (cm) .2 .5 .5 .5 .5 .5 .2 .5 .5 .3 .5 .3 6-18 .5 .5 .3 .2 5 5 .5 .3 1 Water table (cm) S S S S S S 10 S S 5 S "5 A 10 10 10 10 10 15 10 0-10 Append ix V I I I - c o n t ' d . T a b l e o f e n v i r o n m e n t a l d a t a f o r t h e f o r t y r e l e v e s ( p a r t 2) Releve t A A A A A A A A A . ® ®<D <s> $ § y § § Slope(%) 60 50 32 0-30 22 65 15 22 45 0 5 4 80 4 0-50 50 55 50 65 Upslope d i s t . (ro) 45 35 0 0 100 100 10 20 50 0 0 0 100 250 500 700 120 140 700 Bedrock I Me Me I I I I Me Me Me Me Me I I. Me I Me Me Me Landform Sv R Sv R Sv R Ov R Sv R Sv R Ov Sv/R Ov R Sv R Fb Fb Fv Sv R Ov R Cv R Cv R Cv R Cv R Cv R So i l Subgroup 0 HP 0 HP 0 HP TY H TY FO 0 FHP 0 FO TY FO 0 HP R HG R G GL /C FHP / U TY R FO TY FO 0 HP GL R T FO 0 HP TY FO Texture or decomp. SL SL SL Oh SL SCL Oh SL SL Si SCL SCL LS S SL S SL FSCL L Coarse frag. (*) 40 50 70 - 50 55 - 40 15 -J 24 25 50 - 57 26 36 40 85 Min. s o i l depth(cm) 26 19 50 10 10 75 30 3 25 100 70 45+ 10 10 100 70 95 100 0-30 0 horizon depth(cm) - - - 73 - - 93 - - - - - - - - - - - -Humus form A HR R HUR R HUR 0 UR A HR R UR 0 UR R UR R UR s YR H YR 0 UR 0 HUR s YR R UR A UR 0 HR M HR 0 UR LFH horizons depth(cm) 34 27 18 35 20 26 58 23 18 33 26 35 20 50 20 10 43 20 14-33 Water table(cm) 20 35 A A 10 20 75 10 40 75 60 45 A 20 A 70 A A A 

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