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Plant communities and their standing crops on estuaries of the east coast of Vancouver Island Kennedy, Kathryn Ann 1982

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PLANT COMMUNITIES AND THEIR STANDING CROPS ON ESTUARIES OF THE EAST COAST OF VANCOUVER ISLAND by KATHRYN ANN KENNEDY B. Sc., U n i v e r s i t y of V i c t o r i a , 1974 The s i s Submitted i n P a r t i a l F u l f i l l m e n t the Requirements f o r the Degree of Master of Science i n THE FACULTY OF GRADUATE STUDIES (Department of P l a n t Science) We accept t h i s t h e s i s as conforming to the r e q u i r e d standard THE UNIVERSITY OF May, © Kathryn Ann BRITISH COLUMBIA 1982 Kennedy, 1982 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 a n 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 m a k e 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 m a y b e g r a n t e d b y t h e h e a d o f my d e p a r t m e n t o r b y 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 b e 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 Plant Science 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 2 0 7 5 W e s b r o o k P l a c e V a n c o u v e r , C a n a d a V 6 T 1W5 D a t e A p r i l 1, 1982 D E - 6 ( 2 / 7 9 ) ( i i ) A b s t r a c t The emergent p l a n t communities on eig h t e e n e s t u a r i e s on the east coast of Vancouver I s l a n d and one es t u a r y on the mainland coast of B r i t i s h Columbia are d e s c r i b e d and t h e i r d i s t r i b u t i o n mapped ( s c a l e 1 cm = 48 m, 1 cm = 158 m, 1 cm = 32 m). There are eleven types of e s t u a r i e s . Monthly measurements of n i t r o g e n (N) and s t a n d i n g crop by l i v i n g , dead, senescent and d u f f p o r t i o n s were made on e l e v e n communities i n f i v e e s t u a r i e s ; 1. Cowichan Carex l y n g b y e i and Juncus b a l t i c u s ; 2. Chemainus S a l i c o r n i a v i r g i n i c a and D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a ; 3. L i t t l e Qualicum Carex l y n g b y e i and P o t e n t i l l a p a c i f i c a - Carex  l y n g b y e i ; 4. Campbell Carex l y n g b y e i and P o t e n t i l l a p a c i f i c a - E l e o c h a r j s p a l u s t r i s ; and; 5. Salmon Carex l y n g b y e i , Deschampsia c e s p i t o s a - Carex l y n g b y e i and Poa p r a t e n s i s -A g r o s t i s a l b a v a r . s t o l o n i f e r a - P o t e n t i l l a p a c i f i c a . Root co r e s c o l l e c t e d monthly from f i v e p l o t s i n each community were grown i n the dark a t 20°C to measure root r e s e r v e s . K j e l d a h l n i t r o g e n (N) was determined f o r r e p r e s e n t a t i v e samples as a rough measure of the q u a n t i t y of p r o t o p l a s m i c c o n s t i t u e n t s as opposed to s t r u c t u r a l components. The h i g h e s t N v a l u e s occurred i n the l i v i n g p o r t i o n s with the h i g h e s t , 2.43%, o c c u r r i n g i n A p r i l i n the Cowichan Carex  l y n g b y e i community. The s t a n d i n g crops (gm m~2) are Cowichan Carex l y n g b y e i 588 Cowichan Juncus b a l t i c u s 754 Chemainus S a l i c o r n i a v i r g i n i c a 966 ( i i i ) C h e m a i n u s D i s t i c h l i s s p i c a t a - 1 4 3 7 G r i n d e l i a i n t e g r i f o l i a L i t t l e Q u a l i c u m C a r e x l y n g b y e i 1 5 0 4 L i t t l e Q u a l i c u m P o t e n t i l l a p a c i f i c a - C a r e x l y n g b y e i 7 7 0 C a m p b e l l C a r e x l y n g b y e i 4 8 7 C a m p b e l l P o t e n t i l l a p a c i f i c a -E l e o c h a r i s p a l u s t r i s 3 9 2 S a l m o n C a r e x l y n g b y e i 7 7 3 S a l m o n D e s c h a m p s i a c e s p i t o s a - C a r e x l y n g b y e i 1 0 9 2 S a l m o n P o a p r a t e n s i s - A g r o s t i s a l b a v a r . s t o l o n i f e r a - P o t e n t i l l a p a c i f i c a 7 7 2 T h e l a r g e s t s t a n d i n g c r o p s o c c u r i n t h e C a r e x l y n g b y e i , a n d C h e m a i n u s S a l i c o r n i a v i r g i n i c a a n d D i s t i c h l i s s p i c a t a -G r i n d e l i a i n t e g r i f o l i a c o m m u n i t i e s . W h i l e d r y m a t t e r i n t h e C a r e x l y n g b y e i c o m m u n i t i e s d i s a p p e a r e d b e t w e e n t h e g r o w i n g s e a s o n s i t a c c u m u l a t e d i n t h e S a l i c o r n i a v i r g i n i c a a n d D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a c o m m u n i t i e s . D r y m a t t e r a c c u m u l a t e d t o a l e s s e r e x t e n t i n t h e h i g h e r e l e v a t i o n c o m m u n i t i e s . E t i o l a t e d s h o o t s f r o m r o o t c o r e s g r e w i n t h e d a r k f r o m 0 t o a m a x i m u m o f 5 7 3 d a y s f o r t h e L i t t l e Q u a l i c u m P o t e n t i l l a p a c i f i c a - C a r e x l y n g b y e i c o m m u n i t y . R o o t r e s e r v e s a r e p e r i o d i c w i t h t h e d e c r e a s i n g t r e n d i n A p r i l a n d M a y c o i n c i d i n g w i t h t h e i n i t i a t i o n a n d r a p i d g r o w t h o f t h e c a n o p y . T h e p e a k i n s t a n d i n g c r o p i s f o l l o w e d b y a p e a k i n r o o t r e s e r v e s w i t h i n o n e m o n t h . I t i s p r o p o s e d t h a t s o m e c a r b o h y d r a t e s a n d m i n e r a l e l e m e n t s a r e t r a n s l o c a t e d , a n d r e a l l o c a t e d b e t w e e n t h e b e l o w a n d a b o v e g r o u n d s t r u c t u r e s . ( i v ) TABLE OF CONTEOTS Page Abstract ( i i ) Table of Contents (iv) List of Tables ( v i i ) List of Figures (xi) Acknowledgements ( x v i i i ) 1. Introduction 1 1.1 Background 5 1.2 Physiography of the east coast of Vancouver Island 7 1.3 Climate of the study areas on the east coast of 8 Vancouver Island 1.4 Hydrology of the river systems in the study areas 14 1.5 Fish in the river systems 17 Literature Review 19 3. The Plant communities of the estuarine marshes on the 22 east coast of Vancouver Island 3.1 I.fethods 22 3.2 Results and Discussion 24 3.2.1 Estuaries 4C 3.2.1.1 Coldstream 3.2.1.2 Cowichan 3.2.1.3 Chemainus 3.2.1.4 Nanaino 3.2.1.5 rTanoose-Bonell (v) Table of Contents (cont'd) Page 3.2.1.6 Englishmen 3.2.1.7 L i t t l e Qualicum 3.2.1.8 Big Qualicun 3.2.1.9 Courtenay 3.2.1.10 Oyster 3.2.1.11 Campbell 3.2.1.12 Salmon 3.2.1.13 Adam-Eve 3.2.1.14 Tsitika 3.2.1.15 KbfcLsh 3.2.1.16 Ninpkish 3.2.1.17 Cluxewe 3.2.1.18 Quatse 3.2.1.19 Kingcone 3.2.2 Dominant plant species 80 Estimating the production of dry natter of Vancouver 111 Island estuaries 4.1 Standing Crop 111 4.1.1 Methods 112 4.1.2 Results 114 4.1.3 Discussion 120 ( v i ) Table of Contents (cont'd) Page 4.2 Root Reserves 126 4.2.1 Methods 127 4.2.2 Results 128 4.2.3 Discussion 133 5. Conclusions 137 6. Literature cited. 141 Appendix 1. Location, in latitude and longtitude, 146 of the study areas. Appendix 2. Glossary of terns. 147 Appendix 3. Aerial photographs used in the 148 study. Appendix 4. List of plant species. 149 Appendix 5. Descriptionsnaps and area 160 determinations of the plant ccnmunities on the estuaries studied. Appendix 6. Graphs of the l i v i n g , dead, senescent 355 and d r i f f fractions in the plant communities studied. Appendix 7. Graphs of the number of days root - 394 cores from the plant conmunitieij studied grew in darkness. ( v i i ) L i s t of Tables Table Page 1. Population p r o j e c t i o n s f o r r e g i o n a l d i s t r i c t s 3 and t h e i r major urban cen t r e s , 1974-1996. 2. G e o l o g i c a l formations of the study areas. • 9 3. Mean temperature norms f o r s t a t i o n s nearest 11 the study areas. 4. Mean p r e c i p i t a t i o n norms f o r s t a t i o n s nearest 12 the study areas. " 5. H y d r o l o g i c a l i n f o r m a t i o n : watershed s i z e , 16 nainstem l e n g t h , number of t r i b u t a r i e s , gauging s t a t i o n , minimum monthly mean discharge (ens), maximum monthly mean discharge (ens), average A p r i l to September mean discharge (ens), mean annual discharge (ens), water d i v e r s i o n ( r e g u l a t e d or not) and suspended sediment c o n c e n t r a t i o n , p e r t a i n i n g to the study areas. 6. Economic f i s h species i n the r i v e r systems. 18 7. D e s c r i p t i o n of the plan t communities on the 162 Goldstream Estuary. 8. Area of each pla n t community on the Goldstream 169 Estuary. 9. D e s c r i p t i o n of the plant conmunities on the 172 Cowichan River Estuary. 10. Area of each p l a n t community on the Cowichan 131 River Estuary. ( v i i i ) Table 11. D e s c r i p t i o n of the p l a n t c o n n u n i t i e s on the Chemainus R i v e r E s t u a r y . 12. Area of each p l a n t community on the Chemanius R i v e r Estuary. 13. D e s c r i p t i o n of the p l a n t communities on the Nanaimo Ri v e r E s t u a r y . 14. Area of each p l a n t community on the Nanaimo Riv e r E s t u a r y . 15. D e s c r i p t i o n of the p l a n t communities on the Manoose- B o n e l l Creeks' Es t u a r y . 16. Area of each p l a n t community on the Nanoose-B o n e l l Creeks' Estuary. 17. D e s c r i p t i o n of the p l a n t communities on the Englishman R i v e r Estuary. 18. Area of each p l a n t community on the Englishman R i v e r Estuary. 19. D e s c r i p t i o n of the p l a n t communities on the L i t t l e Qualicum River Estuary. 2C. Area of each p l a n t community on the L i t t l e Qualicum R i v e r Estuary. 21. D e s c r i p t i o n of the p l a n t communities on the Big Qualicum River Estuary. 22. Area of each p l a n t community on the Big Qualicum River Estuary. 23. D e s c r i p t i o n of the p l a n t communities on the Courtenay River Estuary. Page 184 197 200 209 212 220 223 230 232 241 243 248 251 ( i x ) T a b l e ?A. A r e a o f e a c h p l a n t c o m m u n i t y on t h e C o u r t e n a y R i v e r E s t u a r y . 2 5 . D e s c r i p t i o n o f t h e p l a n t c o m m u n i t i e s on t h e O y s t e r R i v e r E s t u a r y . 2 6 . A r e a o f e a c h p l a n t c o m m u n i t y on t h e O y s t e r R i v e r E s t u a r y . 27 D e s c r i p t i o n o f t h e p l a n t c o m m u n i t i e s on t h e C a m p b e l l R i v e r E s t u a r y . 2 8. A r e a o f e a c h p l a n t c o m m u n i t y on t h e C a m p b e l l R i v e r E s t u a r y . 2 9 . D e s c r i p t i o n o f t h e p l a n t c o m m u n i t i e s on t h e Salmon R i v e r E s t u a r y . 3 0 . A r e a o f e a c h p l a n t c o m m u n i t y on t h e Salmon R i v e r E s t u a r y . 3 1 . D e s c r i p t i o n o f t h e p l a n t c o m m u n i t i e s on t h e Adam-Eve R i v e r s ' E s t u a r y . 3 2 . A r e a o f e a c h p l a n t c o m m u n i t y on t h e Adam-Eve R i v e r s ' E s t u a r y . 3 3 . D e s c r i p t i o n o f t h e p l a n t c o m m u n i t i e s on t h e T s i t i k a R i v e r E s t u a r y . 34. A r e a o f e a c h p l a n t c o m m u n i t y on t h e T s i t i k a R i v e r E s t u a r y . 3 5 . D e s c r i p t i o n o f t h e p l a n t c o m m u n i t i e s on t h e K o k i s h R i v e r E s t u a r y . (x) Table Page 36. Area of each plant community on the Kokish 314 River Estuary. 37. Description of the plant communities on the 316 Nimpkish River Estuary. 38. Area of each plant community on the Nimpkish 323 River Estuary. .39. Description of the plant communities on the 325 Cluxewe River Estuary. 40. Area of each plant community on the Cluxewe 332 River Estuary. 41. Description of the plant communities on the 334 Quatse River Estuary. 42. Area of each plant community on the Quatse 345 River Estuary. 43. Description of the plant communities on the 347 Kingcome River Estuary. 44. Area of each plant community on the Kingcone 353 River Estuary. 45. Simple l i n e a r c o r r e l a t i o n between species, and 354 flow, and p r e c i p i t a t i o n . 46. Standing crop by summing increments, peak value 115 and cleared plot for each of the 11 plant communities. ( x i ) L i s t of Figures Page 1. Diagram of the d e t r i t u s food web t y p i c a l of a 2 slough/channel t i d a l marsh h a b i t a t i n the Fraser River Estuary ( K i s t r i t z , 1978). 2. Population p r o j e c t i o n s f o r the Comox-Strathcona 4 ( + ) Manaino ( O ) i Cowichan V a l l e y ( x ) , and Mount Waddington (A) i'egional D i s t r i c t s , 1974-1990. 3. Map of Vancouver I s l a n d and adjacent mainland 6 coast showing the general l o c a t i o n s of the study areas. 4. D i s t r i b u t i o n of the pl a n t communities on the 1G1 Coldstream Estuary. 5. D i s t r i b u t i o n of the pl a n t communities on the 170 Cowichan River Estuary. 6. D i s t r i b u t i o n of the plant communities on the 182 Chemainus River Estuary. 7. D i s t r i b u t i o n of the plant communities on the 198 Nanaimo River Estuary. 8. D i s t r i b u t i o n of the plant communities on the 211 Nanoose- B o n e l l Creeks' Estuary. 9. D i s t r i b u t i o n of the plant communities on the 221 Englishman River Estuary. 10. D i s t r i b u t i o n of the plant communities on the 231 L i t t l e Oualicum River Estuary. 11. D i s t r i b u t i o n of the plant communities on the 242 Big Qualicum River Estuary. ( x i i ) L i s t of Figures (cont'd) IT. D i s t r i b u t i o n of the plant Courtenay River Estuary. 13. D i s t r i b u t i o n of the plant Oyster River Estuary. 14. D i s t r i b u t i o n of the plant Campbell River Estuary. 15. D i s t r i b u t i o n of the plant Salmon River Estuary. 16. D i s t r i b u t i o n of the plant Adam-Eve Rivers' Estuary. •17. D i s t r i b u t i o n of the plant T s i t i k a River Estuary. i n . D i s t r i b u t i o n of the plant Kokish River Estuary. 19. D i s t r i b u t i o n of the plant Nimpkish River Estuary. 20. D i s t r i b u t i o n of the plant Cluxewe River Estuary. 21. D i s t r i b u t i o n of the plant Quatse River Estuary. 22. D i s t r i b u t i o n of the plant Kingcome River Estuary. 23. Mean monthly dry weights ( dead, senescent and duff p the Cowichan Carex lyngbye Page communities on the 249 communities on the 262 communities on the 268 communities on the 278 communities on the 294 communities on the 303 communities on the 309 communities on the 315 communities on the 324 communities on the 333 communities on the 346 gm ,5m-2) of l i v i n g , 356 Iant fractions from i community. ( x i i i ) L i s t of F i g u r e s (cont'd) Page 24. Mean monthly dry weights (gm .5n~ 2) of l i v i n g , 358 dead, 358 senescent and duf f p l a n t f r a c t i o n s from the L i t t l e Q u a l i c u n Carex l y n g b y e i community. 25. Mean monthly dry weights (gm .5m~2) of l i v i n g , 360 dead, senescent and duf f p l a n t f r a c t i o n s from the Salmon Carex l y n g b y e i community. 26. Mean monthly dry weights (gm .5m~2) of l i v i n g , 362 dead, senescent and duff p l a n t f r a c t i o n s from the Cowichan Juncus b a l t i c u s community. 27. Mean monthly dry weights (gm .5m~2) of l i v i n g , 364 dead, senescent and duf f p l a n t f r a c t i o n s from the L i t t l e Qualicum P o t e n t i l l a p a c i f i c a - Carex  l y n g b y e i community. 28. Mean monthly dry weights (gm .5m - 2) of l i v i n g , 366 dead, senescent and duf f p l a n t f r a c t i o n s from the Campbell P o t e n t i l l a p a c i f i c a ~ S l e o c h a r i s  p a l u s t r i s community. 29. Mean monthly dry weights (gm .5m - 2) of l i v i n g , 368 dead, and duf f p l a n t f r a c t i o n s from the Chemainus S a l i c o r n i a v i r g i n i c a community. 30. Mean monthly dry v/eights (gm ,5m - 2) of l i v i n g , 370 dead and d u f f p l a n t f r a c t i o n s from the Chemanius D i s t i c h l i s s p i c a t a - G r i n d e l i a  i n t e g r i f o l i a community. 31. Mean monthly dry v/eights (gm .5n - 2) of l i v i n g , 372 dead, senescent and d u f f p l a n t f r a c t i o n s from the Campbell Carex l y n g b y e i community. ( x i v ) t of F i g u r e s (cont'd) Mean monthly dry v/eights (gn . 5 n - 2 ) of l i v i n g , dead, senescent and d u f f p l a n t f r a c t i o n s from the Salmon deschampsia c e s p i t o s a - Carex  l y n g b y e i community. Mean monthly dry v/eights (gm ,5m - 2) of l i v i n g , dead, senescent and d u f f p l a n t f r a c t i o n s from the Salmon Poa p r a t e n s i s - A g r o s t i s a l b a v a r . S t o l o n i f e r a - P o t e n t i l l a p a c i f i c a community. Mean monthly n i t r o g e n and ash f r e e dry weights (gm .5n - 2) of combined l i v i n g and senescent f r a c t i o n s i n the Cowichan Carex l y n g b y e i community. Mean monthly n i t r o g e n and ash f r e e dry weights (gn .5n~ 2) of the l i v i n g f r a c t i o n i n the Chemainus S a l i c o r n i a v i r g i n i c a community. Mean monthly n i t r o g e n and ash f r e e dry weights (gm .xm - 2) of the l i v i n g f r a c t i o n i n the Chemainus D i s t i c h l i s s p i c a t a - G r i n d e l i a  i n t e g r i f o l i a community. Mean monthly n i t r o g e n and ash f r e e dry weights (gm .5m - 2) of combined l i v i n g and senescent f r a c t i o n i n the L i t t l e Qualicum P o t e n t i l l a  p a c i f i c a - Carex l y n g b y e i community. Mean monthly n i t r o g e n and ash f r e e dry weights (gm .5m - 2) of combined dead and duf f f r a c t i o n s i n the Cowichan Carex l y n g b y e i community. ( X V ) L i s t of F i g u r e s (cont'd) Page 30. Mean monthly n i t r o g e n and ash f r e e dry v/eights 388 (gm .5m - 2) of combined dead and duf f f r a c t i o n s i n the Chemainus S a l i c o r n i a v i r g i n i c a community 40. Mean monthly n i t r o g e n and ash f r e e dry weights 390 (gm .5m - 2) of combined dead and d u f f f r a c t i o n s i n the Chemainus D i s t i c h l i s s p i c a t a - G r i n d e l i a  i n t e g r i f o l i a community. 41. Mean monthly n i t r o g e n and ash f r e e dry weights 392 (gm .5m - 2) of combined dead and duff p o r t i o n s i n the L i t t l e Qualicum P o t e n t i l l a p a c i f i c a -Carex l y n g b y e i community. 42. Exhaustion of root r e s e r v e s i n darkness: 129 maximum time (days) f o r (1) Carex l y n g b y e i , (2) Juncus b a l t i c u s , (3) Deschampsia c e s p i t o s a , and (4) S a l i c o r n i a v i r g i n i c a p l a n t c o n n u n i t i e s , and maximun time f o r (5) Agropyron spicatum, S t i p a  comata and Festuca s c a b r e l l a (Lobb, 1969), (6) Poa p r a t e n s i s and (7) Festuca r u b r a and A g r o s t i s t e n u i s (Marx, 1961). 43. The average number of days ro o t cores c o l l e c t e d 131 i n A p r i l (4), May (5), June (6), J u l y (7), August ( 8 ) , and October (10) grew i n the dark. 44. Exhaustion of root r e s e r v e s i n darkness f o r the 395 Cov/ichan Carex l y n g b y e i community: height (cn) ( x v i ) L i s t of F i g u r e s (cont'd) Page of regrown shoots and number of o v e r - w i n t e r i n g and regrowth shoots. 45. Exhaustion of ro o t r e s e r v e s i n darkness f o r the 397 Cowichan Juncus b a l t i c u s community: average time (days) and height (cm) of regrown shoots. 46. Exhaustion of root r e s e r v e s i n darkness f o r the 399 Chemainus S a l i c o r n i a v i r g i n i c a (S) and D i s t i c h l i s s p i c a t a - C r i n d e l i a i n t e g r i f o l i a (D) communities: average time ( d a y s ) . 47. Exhaustion of ro o t r e s e r v e s i n darkness f o r the 401 L i t l e Oualicum Carex l y n g b y e i community: average time (days), height (cm) of regrown shoots, and number of o v e r - w i n t e r i n g and regrowth shoots. 48. Exhaustion of root r e s e r v e s i n darkness f o r the 403 L i t t l e Qualicum P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i community: average time (days), average h e i g h t (cm) of regrov/n shoots of Lyngby's sedge and B a l t i c rush. 49. Exhaustion of ro o t r e s e r v e s i n darkness f o r the 405 Campbell Carex l y n g b y e i community: average time ( d a y s ) , h e i g h t (cm) of regrov/n shoots, and number of over - v/intering and regrowth shoots. 50. Exhaustion of root r e s e r v e s i n darkness f o r the 407 Campbell P o t e n t i l l a p a c i f i c a - E l e o c h a r i s ( x v i i ) L i s t of F i g u r e s (cont'd) Page p a l u s t r i s community: average t i n e (days), and height (cm) of regrown shoots. 5 1 . Exhaustion of r o o t r e s e r v e s i n darkness f o r the 409 Salmon Carex l y n g b y e i community: average time (da y s ) , height (cm) of regrown shoots, and number of o v e r - w i n t e r i n g and regrowth shoots. 52. Exhaustion of root r e s e r v e s i n darkness f o r the 411 Salmon Deschampsia c e s p i t o s a - Carex l y n g b y e i (D) and Poa p r a t e n s i s - A g r o s t i s a l b a var. s t o l o n i f e r a - P o t e n t i l l a p a c i f i c a (P) communities: average time ( d a y s ) . Ho-ps iVv Special Co I 'cecums , ( x v i i i ) Acknowledgements There are many people to whom I am indebted f o r t h e i r support and encouragement. In memory of N u r e t i n Keser, I would l i k e to express my a p p r e c i a t i o n f o r h i s t e a c h i n g me a i r photo i n t e r p r e t a t i o n , and f o r the time, thought and e x p e r t i s e he shared with me„ To my a d v i s o r , Dr. V.C. B r i n k , my most s i n c e r e thanks f o r h i s endless p a t i e n c e and f a i t h as w e l l as h i s e x p e r i e n c e , knowledge and time of which he gave f r e e l y . A s p e c i a l thanks to Dr. D a r y l Hebert who arranged f i n a n c i a l support through Dr. Don Eastman of the B r i t i s h Columbia F i s h and W i l d l i f e Branch and Jon S e c t o r of the B r i t i s h Columbia Lands Branch. D a r y l a l s o arranged f o r the use of the Nanaimo F i s h and W i l d l i f e Branch's a e r i a l photographs, f i e l d gear, and emergent v e g e t a t i o n data I c o l l e c t e d i n 1975. Besides l o g i s t i c a l a s s i s t a n c e D a r y l , George Reid and C. Lyons, of the Nanaimo F i s h and W i l d l i f e Branch, p r o v i d e d much a p p r e c i a t e d moral support. To my good f r i e n d Wayne Kale , who helped me i n the f i e l d , guided me through computer programs and was always w i l l i n g to d i s c u s s every aspect of t h i s work, a very s p e c i a l thanks. My s i n c e r e a p p r e c i a t i o n to everyone who helped i n the f i e l d and l a b ; Moira Lemon, Ron Fleming, Marika Townshend, Dr. L. L a v k u l i c h , Dr. A Bomke, Bev Herman, Mr. Reddy, Dr. B. F o s t e r , Dr. J . P o j a r , Dave Routledge, Kevan Wall, Lance Sundquist, L a u r e l Szasz, Dr. D. Shackleton, Stan Baker of the Cowichan V a l l e y N a t u r a l H i s t o r y S o c i e t y , the Nanaimo D i s t r i c t N a t u r a l i s t ' s Club, P h i l Capes, the M i t t l e n a t c h F i e l d N a t u r a l i s t S o c i e t y , and Judy Sachet. ( x i x ) The p r o d u c t i o n of t h i s t h e s i s would have been i m p o s s i b l e witho ut Jim Walker and Pat Young. Jim generously o f f e r e d the s e r v i c e s of the H a b i t a t P r o t e c t i o n D i v i s i o n of the V i c t o r i a F i s h and W i l d l i f e Branch and Pat typed the t h e s i s i n t o a word p r o c e s s o r . A very, very s p e c i a l thank you to both of you, and p a r t i c u l a r l y t o Pat, v/ho f i t t e d my t h e s i s i n t o her work l o a d . And to my husband Jim, f o r whom B a l t i c rush w i l l f o r e v e r be Juncus horrendus and to whom one of my hours means thr e e , and to our paren t s , thank you. - 1 -1. INTRODUCTION The impact of man's a c t i v i t i e s on e s t u a r i n e ecosystems has become a major concern (Sorenson, 1973) i n B r i t i s h Columbia. The e c o l o g i c a l values of the t i d a l marshes are widely r e c o g n i z e d (Odum, 1961) and have long been noted as v a l u a b l e h a b i t a t f o r s h o r e b i r d s , ducks and geese (Burgess, 1970, C h a t t i n , 1970, Burton, 1977). Recently i t has become e v i d e n t t h a t j u v e n i l e salmon, notab l y chum and chinook, are f e e d i n g i n sloughs and channels a s s o c i a t e d with t i d a l marshes (Dunford, 1975, Reimers 1973) ( F i g . 1.) F i s h e r i e s b i o l o g i s t s b e l i e v e the use of e s t u a r i n e marshes by j u v e n i l e salmon i s an important aspect of t h e i r l i f e h i s t o r y . The e s t u a r i e s on the east coast of Vancouver I s l a n d are p a r t i c u l a r l y noteworthy f o r they are p a r t of a l a r g e c o a s t a l s h e l f extending south from Campbell R i v e r t o V i c t o r i a . T h i s s h e l f i s r i c h i n b i o t a , eg. o y s t e r s , clams, crabs and algae as w e l l as f i s h and b i r d s . The abundance and d i v e r s i t y of f l o r a and fauna i s s t r i k i n g when compared to the lower d i v e r s i t y and p r o d u c t i v i t y of B r i t i s h Columbia's numerous f i o r d systems ( E l l i s , p e r s . comm.). The c o a s t a l s h e l f and a s s o c i a t e d lowlands p r o v i d e man with l e v e l b u i l d i n g s i t e s and i t i s here l o g storage and h a n d l i n g , c o a l mining, tourism, urban development, p o r t s and t r a n s p o r t a t i o n routes are co n c e n t r a t e d . The impact of human a c t i v i t y on the n a t u r a l systems i s d e s t r u c t i v e and e s c a l a t i n g . By the year 1996 the p o p u l a t i o n of major urban c e n t r e s , l o c a t e d on or adjacent to r i v e r d e l t a s , i s e x t r a p o l a t e d to be double that given i n the 1976 census (Table 1, F i g . 2). A s s o c i a t e d development p r e s s u r e s Detritus Particle igure 1. Diagram of the de t r i t u s food web t y p i c a l of a slough/channel t i d a l marsh habitat i n the Fraser River estuary ( K i s t r i t z , 1978). Table 1. P o p u l a t i o n s and po p u l a t i o n p r o j e c t i o n s f o r r e g i o n a l d i s t r i c t s and t h e i r major urban c e n t e r s , 1974 - 1996 Po p u l a t i o n c e n t r e s by Regional D i s t r i c t P o p u l a t i o n 1971 1976 Centre p o p u l a t i o n as a percent of r e g i o n a l d i s t r i c t 1971 1976 3 pop. P o p u l a t i o n 4 1996 I Comox-Strathcona R.D. 47345 55761 99646 1. Campbell R i v e r , DM2 10000 11787 21 21 20926 2. Comox, T-V 4055 5226 9 9 8968 3. Courtenay, C 7187 7566 15 14 13950 4. Sayward, VL 465 380 1 .7 698 II Cowichan R.D. 38988 45138 67277 1. Duncan, C 4388 3960 11 9 6055 II I Mount Waddington R.D. 10408 12306 18462 1. Port Hardy, DM 1777 3579 17 29 5354 IV Nanaimo R.D. 48006 60768 86426 1. Nanaimo, C 34029 39655 71 65 56177 2. P a r k s v i l l e , VL 2171 3158 5 5 4321 3. Qualicum Beach, VL 1245 1707 3 3 2593 1 B r i t i s h Columbia Research Management S e r v i c e s D i v i s i o n , 1974 and Census of Canada, 1976. 2 DM - d i s t r i c t m u n i c i p a l i t y T-V - town C - c i t y VL - v i l l a g e 3 Percentages c a l c u l a t e d to nearest u n i t from p o p u l a t i o n f i g u r e s . 4 P o p u l a t i o n c a l c u l a t e d u s i n g 1996 p o p u l a t i o n p r o j e c t i o n s and percentages. F i g u r e 2: Populations and population p r o j e c t i o n s f o r the Comox-Strathcona ( + ), Nanaimo (<» , Cowichan V a l l e y (X) and Mount Waddington (A) Regional D i s t r i c t s , 1974-1996 (from B.C. Research Management S e r v i c e s D i v i s i o n , 1974). - 5 -on the e s t u a r i n e systems w i l l i n c r e a s e d r a m a t i c a l l y . The need f o r i n f o r m a t i o n to enable those r e s p o n s i b l e to b e t t e r manage e s t u a r i n e ecosystems i s great now. The o b j e c t i v e s of t h i s study were set to p r o v i d e an i n f o r m a t i o n base and i d e n t i f y areas r e q u i r i n g f u r t h e r study. More s p e c i f i c a l l y they were to: 1. determine and d e s c r i b e the emergent p l a n t communities i n major e s t u a r i e s on the east coast of Vancouver I s l a n d as they p r e s e n t l y e x i s t . 2. attempt to i n d i c a t e the p r i n c i p a l p h y s i c a l f a c t o r s i n f l u e n c i n g p l a n t s p e c i e s d i s t r i b u t i o n . 3. estimate s t a n d i n g crop, determine i t s seasonal v a r i a t i o n s and q u a l i t y . 4. d e l i n e a t e root r e s e r v e p a t t e r n s i n the e s t u a r i n e communities. 1.1 Background Eighteen e s t u a r i e s i n t h i s study were s e l e c t e d from the east coast of Vancouver I s l a n d ; a n i n e t e e n t h estuary was s e l e c t e d from the mainland f i o r d system f o r comparative purposes. There were three c r i t e r i a g u i d i n g the choice of e s t u a r i e s . The f i r s t c r i t e r i o n was to o b t a i n a r e p r e s e n t a t i v e group from the many e s t u a r i e s , l a r g e and s m a l l , along the s h e l f ; the second, to o b t a i n a range of b i o p h y s i c a l f a c t o r s among the e s t u a r i n e ecosystems; and t h i r d , to i n c l u d e e s t u a r i e s under p r e s s u r e f o r development. Appendix 1 and F i g u r e 3 give the l o c a t i o n s of the study a r e a s . The nineteen e s t u a r i e s , by name are the Goldstream, Cowichan, Chemainus, Nanaimo, Nonoose-F i g u r e 3. Map of Vancouver I s l a n d and adjacent showing the general l o c a t i o n s of the mainland coast study areas. - 7 -B o n e l l , Englishman, L i t t l e Qualicum, Big Qualicum, Courtenay, Oyster, Campbell, Salmon, Adam-Eve, T s i t i k a , K okish, Nimpkish, Cluxewe, Quatse and Kingcome R i v e r ' s e s t u a r i e s . Each es t u a r y i s named a f t e r the p r i n c i p a l r i v e r ( s ) i n the d e l t a . For the sake of b r e v i t y , e s t u a r i e s are o f t e n r e f e r r e d to by the r i v e r ' s name onl y ( i . e . Cowichan, i n s t e a d of Cowichan R i v e r e s t u a r y ) , p a r t i c u l a r l y when d i s c u s s i n g p l a n t communities. They are d e a l t with i n the t e x t i n order from south to n o r t h . Terms are d e f i n e d i n the g l o s s a r y i n Appendix 2. 1.2 Physiography of the east coast of Vancouver I s l a n d E s t u a r i e s on the east coast of Vancouver Isla n d occur i n e i t h e r the Vancouver I s l a n d Ranges or the Nanaimo Lowlands. The Vancouver I s l a n d Ranges extend to the coast northwest of Kelsey Bay and there t i d a l marshes are l i m i t e d to the s h e l t e r e d mouths of g l a c i a t e d and g e n e r a l l y U-shaped v a l l e y s . The c o a s t l i n e r i s e s a b r u p t l y from the sea to the mountains except i n the Suquash Basin and N a h w i t t i Lowlands. These two p h y s i o g r a p h i c u n i t s have a g e n e r a l l y low r e l i e f with f l a t s and r o l l i n g h i l l s ; the e l e v a t i o n i n the Suquash Basin seldom exceeds 150 meters w h i l e the e l e v a t i o n i n the N a h w i t t i Lowlands i s g e n e r a l l y below 610 meters. The Kokish, Nimpkish and Cluxewe R i v e r s ' e s t u a r i e s occur i n the Suquash Basin and the Quatse occurs i n the N a h w i t t i Lowlands. E s t u a r i e s south of Kelsey Bay l i e w i t h i n the Nanaimo Lowland d e s c r i b e d by Holland (1964) as "a s t r i p of low l y i n g c ountry, below 2000 f e e t (610 meters) e l e v a t i o n , which extends southeastward f o r 175 m i l e s (282 k i l o m e t e r s ) along the east - 8 -coast of Vancouver I s l a n d from Sayward on Johnstone S t r a i t to Jordan R i v e r west o f V i c t o r i a . . . l a r g e l y u n d e r l a i n by sedimentary rocks of the Nanaimo Group of Upper Cretaceous age. It i s f l a n k e d on i t s western s i d e above the 2000 f o o t (610 meter) contour l i n e by the Vancouver I s l a n d Ranges... The lowland c o n s i s t s of many low, wooded c u e s t a - l i k e r i d g e s separated by narrow v a l l e y s . The n o r t h w e s t e r l y e l o n g a t i o n of .the r i d g e s of the G u l f I s l a n d s i s the r e s u l t of d i f f e r e n t i a l e r o s i o n of the Upper Cretaceous sedimentary r o c k s . The r i d g e s are u n d e r l a i n by hard sandstone and conglomerate beds, and the v a l l e y s are eroded i n s h a l e s and s o f t e r rocks or along f a u l t zones." A more recent treatment of the geology of the area i s that of M u l l e r (1971). Table 2 summarizes g e o l o g i c a l d e s c r i p t i o n s f o r the e s t u a r i e s s t u d i e d . Major f a u l t s occur i n the g e o l o g i c a l formations i n some of the study areas (e.g., K o k i s h and Nimpkish) and more than one bedrock u n i t i s p r e s e n t . The Kingcome i s at the mouth of a g l a c i a t e d r i v e r v a l l e y i n the g r a n i t i c P a c i f i c C o a s t a l Mountain Range. 1.3 Climate of the study areas on the east coast of Vancouver I s l a n d There are few c l i m a t o l o g i c a l s t a t i o n s l o c a t e d i n the e s t u a r i e s s t u d i e d and the weather r e c o r d s i n Tables 3 and 4 are from the nearest s t a t i o n s . In g e n e r a l , the c l i m a t e of the east coast of Vancouver I s l a n d becomes c o o l e r and wetter from south to n o r t h . South of Campbell R i v e r the e s t u a r i e s are i n the r a i n shadow of the mountains of the Vancouver I s l a n d Ranges. The Table 2. Ge o l o g i c formations i n the study a r e a s 1 Estuary Rock Unit Age D e s c r i p t i o n Goldstream Cowichan Nanoose-Bonell S i c k e r Group Late P a l e o z o i c V o l c a n i c t u f f and b r e c c i a of in t e r m e d i a t e composition with a r g i l l i t e and c h a r t o v e r l a i n by B u t t l e Lake limestone. The S i c k e r Group are metamorphosed and i n t e n s e l y deformed i n some areas Cowichan Chemainus Nanaimo Nanoose-Bonell Englishman L i t t l e Qualicum Bi g Qualicum Courtenay Oyster Cluxewe Late Mesozoic Sediments Late J u r a s s i c to Cretaceous Greywacke, sandstone, conglomerate and s h a l e , carbonaceous shale and c o a l Campbell Salmon Adam-Eve T s i t i k a Kokish Quatse Adam-Eve Karmutsen Formation Islan d I n t r u s i o n s T r i a s s i c and o l d e r (?) Middle to Late J u r a s s i c S l i g h t l y metamorphosed b a s a l t i c l avas c o n s i s t i n g of massive and amygdaloidal flows, p i l l o w l a v a s and a s s o c i a t e d b r e c c i a . Thin limestone members may occur near the top of the formation Large b a t h o l i t h s of g r a n o d i o r i t e to q uartz d i o r i t e composition with l o c a l stocks of quartz monzonite and g r a n i t e d i f f e r e n t i a t e s Table 2. (cont'd) Geologic formations i n the study areas Estuary Rock Unit Age D e s c r i p t i o n Kokish Bonanza Subgroup Late T r i a s s i c to Ear l y J u r a s s i c Upper d i v i s i o n s of commonly - red colored rhyodacite l a v a s , t u f f s , b r e c c i a s and i g n i m b r i t e s interbedded with b a s a l t i c andesite l a v a s , t u f f s and b r e c c i a s . Lower d i v i s i o n of b a s a l t i c to andesite t u f f , b r e c c i a and lava flows i n t e r -bedded with Lower J u r a s s i c greywacke and a r g i l l i t e Kokish Nimpkish Quatse Formation Late T r i a s s i c Massive gray to t h i n bedded black limestone, i n places with an upper part of calcareous greywacke and li m e -stone b r e c c i a Kokish Nimpkish Parson Bay Formation Late T r i a s s i c Thin bedded carbonaceous li m e -stone and interbedded t u f f s iFrom Muller (1971), and Keser and St. P i e r r e (1973) T a b l e 3 . Mean t e m p e r a t u r e n o r m s f o r s t a t i o n s n e a r e s t t h e s t u d y a r e a s ( A t m o s p h e r i c E n v i r o n m e n t , 1 9 7 3 ) T e m p e r a t u r e C O E s t u a r y S t a t i o n m a x . m i n . mean mean mean mean mean mean mean a n n u a l d a l l y a n n u a l d a i l y a n n u a l d a l l y d a i l y d a l l y m a x . d a l l y m i n . d a i l y temp temp temp m a x . m i n . G o l d s t r e a i n V i c t o r i a I n t . A i r p o r t J u l 1 6 . 4 J a n 2 . 9 9 . 6 J u l 2 1 . 9 1 3 . 9 J a n ~ 0 . 1 5 . 3 C o w i c h a n C o w i c h a n Bay J u l 1 7 . 3 Jan 2 . 2 9 . 6 J u l 2 2 . 2 1 3 . 3 J a n ~ 0 . 1 5 . 8 C h e m a i n u s C r o f t o n - - - -N a n a i m o D e p a r t u r e Bay J u l 1 8 . 3 Jan 3 . 5 1 0 . 6 J u l 2 3 . 2 1 4 . 5 J a n 0 . 8 6 . 6 E n g l i s h m a n P a r k s v i l l e C o u r t e n a y Comox A i r p o r t J u l 1 7 . 3 Jan 2 . 1 9 . 3 J u l 2 2 . 8 1 3 . 4 J a n ~ 0 . 8 5 . 2 C a m p b e l l C a m p b e l l J u l 1 7 . 4 Jan 1 . 3 8 . 9 J u l 2 3 . 5 1 3 . 2 J a n " 1 . 3 4 . 7 Q u a t s e P o r t H a r d y A i r p o r t A u g 1 3 . 8 Jan 2 . 4 7 . 9 A u g 1 7 . 3 1 1 . 2 J a n 0 . 0 4 . 7 Table 4 Mean p r e c i p i t a t i o n norms for s t a t i o n s nearest the study areas (Atmospheric Env i ronment , 1973) Go Idstream V i c t o r i a In t . A i r p o r t Cowichan Cowichan Bay Chemainus C ro f ton Nanaimo Departure Bay Englishman P a r k s v i I Ie Courtenay Courtenay Campbe 11 Beaver Harbour Campbe11 Quatse Ki ngcome P r e c i p i t a t i o n (mm) number of mean mean days with annual max mean m!n. mean A p r i l t o annual measurable t o t a l monthly monthly S e p t . mean r a i n f a l l r a i n p r e c i p i t a t i o n p r e c i p i t a t i o n p r e c i p i t a t i o n r a i n f a l l Roy - nearest coasta I main land s t a t i o n is southeast of KIngcome 811.0 905.5 963.4 886.5 875.0 1387.1 1435.9 1595.1 21 12.8 147 150 157 138 162 154 128 208 159 856.5 961.4 1029.0 929.6 952.2 1506.5 1538.5 1637.2 2205.2 Jan 146.3 Dec 163.6 Jan 175.8 Dec 157.0 Dec 159.8 Dec 261.4 Dec 270.3 Dec 230.4 Dec 329.2 Ju I 18.5 Ju I 2 1 . 6 Ju I 2 0 . 6 Ju I 26 .2 Ju I 28 .2 Ju I 3 3 . 8 Ju I 39.1 Ju I 51.6 Ju I 74 .4 183.9 2 0 9 . 3 221 .2 239 .3 2 4 8 . 9 3 2 0 . 0 3 3 4 . 8 481.8 635.3 - 13 -Goldstream, Cowichan and Chemainus, to some extent, are a l s o i n the r a i n shadow of the Olympic P e n i n s u l a r Mountains and are c o r r e s p o n d i n g l y d r i e r . Summers are long, c o o l and g e n e r a l l y dry, and w i n t e r s are s h o r t , m i l d and wet. It i s o f t e n r e f e r r e d to as "a cool-summer Mediterranean c l i m a t e " , based on the Koppen c l a s s i f i c a t i o n system (Ackerman, 1941). North of Courtenay the summers are long, c o o l and r e l a t i v e l y wet, and the winters are s h o r t , wet and m i l d . November, December and January are the wettest months. The Koppen c l a s s i f i c a t i o n system r e f e r s to i t as "a marine west coast c l i m a t e " . The Kingcome es t u a r y , where the o ceanic clouds s t r i k e the mountains and undergo a d i a b a t i c c o o l i n g i s n o t a b l y wetter and probably c o o l e r than Vancouver I s l a n d e s t u a r i e s . The mean annual d a i l y temperature ranges from a high of 10.6 °C a t Departure Bay t o 7.9°C a t Port Hardy a i r p o r t ; the maximum mean d a i l y temperatures range from 18.3°C a t Departure Bay (Nanaimo) t o 13.8°C a t Port Hardy a i r p o r t (Quatse). The mean annual r a i n f a l l i n c r e a s e s from 811.0 mm at V i c t o r i a I n t e r n a t i o n a l A i r p o r t to 1595.1 mm a t Port Hardy a i r p o r t . At Roy, the nearest s t a t i o n to the Kingcome, the mean annual r a i n f a l l i s 2112.8 mm, s u b s t a n t i a l l y g r e a t e r than that f o r the aforementioned s t a t i o n s on Vancouver I s l a n d . For a l l study areas, the month of maximum mean p r e c i p i t a t i o n i s e i t h e r December or January and the minimum i s J u l y . The A p r i l to September ( i . e . growing season) mean r a i n f a l l i n c r e a s e s from 183.9 mm at V i c t o r i a I n t e r n a t i o n a l A i r p o r t to 481.8 mm at P o r t Hardy a i r p o r t and 635.3 mm at Roy. On Vancouver I s l a n d , the number of f r o s t - f r e e growing days ranges from 322 at Cowichan Bay t o 280 at Campbell. - 14 -1.4 Hydrology of the r i v e r systems i n the study areas H y d r o l o g i c a l data (Table 5) f o r r i v e r s and t h e i r e s t u a r i e s are l i m i t e d and v a r i a b l e . Many r i v e r systems are not monitored and those t h a t are seldom have gauging s t a t i o n s i n t h e i r e s t u a r i e s to r e c o r d t i d a l i n f l u e n c e s . Most of the s t a t i o n s have not been r e c o r d i n g f o r the same l e n g t h of time and o f t e n the r e c o r d s are i n t e r m i t t e n t or d i s c o n t i n u e d . For example, only estimated J u l y v a l u e s were a v a i l a b l e f o r the Cluxewe and Quatse R i v e r s . Though records possess shortcomings they p r o v i d e t r e n d s . On the b a s i s of watershed s i z e (Table 5) the f i v e l a r g e s t r i v e r systems are the Nimpkish with a watershed of 1989 square k i l o m e t e r s f o l l o w d by the Kingcome a t 1456 sq. km, Campbell at 1432 sq. km, Salmon at 1380 s q . km and Cowichan at 834 sq. km. The f o u r s m a l l e s t are the Goldstream a t 70 sq. km, Quatse at 52 sq. km, B o n e l l at 47 sq. km, and Nanoose at 28 sq. km. The remainder range from 769 sq. km f o r the Puntledge and Tsolum, t r i b u t a r i e s of the Courtenay R i v e r , to 52 sq. km f o r the Quatse. The mainstem l e n g t h , number of t r i b u t a r i e s and mean annual d i s c h a r g e are g e n e r a l l y d i r e c t l y r e l a t e d to watershed s i z e . The l e n g t h of the Salmon R i v e r mainstem i s 82.1 k i l o m e t e r s f o l l o w e d by the Nanaimo at 80.5 km, Kigcome a t 74.4 km, Chemainus at 64.4 km, Nimpkish at 62.4 km, and Adam-Eve at 59.9 km. The s i x r i v e r s with s h o r t e s t mainstem le n g t h s are the Goldstream a t 11.1 km, Quatse a t 12.1 km, B o n e l l at 12.6 km, Nanoose at 14.3 km, Campbell at 9.7 km, and Courtenay at 2.4 km. The Chemainus has 62 t r i b u t a r i e s , the Adam-Eve and Salmon 55 each, the T s i t i k a 42, Cluxewe 36 and Englishman 27 down to 3 t r i b u t a r i e s f o r the Nanoose, and 5 f o r - 15 -the L i t t l e Qualicum. The s i x highest mean annual d i s c h a r g e s are 129.13 c u b i c meters per second (cms) f o r the Nimpkish, 98.83 cms f o r the Campbell, 66.54 cms f o r the Salmon, 54.35 cms f o r the Courtenay (Puntledge and Tsolum), 53.24 cms f o r the Cowichan and 41.91 cms f o r the Nanaimo. The s i x lowest mean annual d i s c h a r g e s are 7.39 cms f o r the Big Qualicum, 12.32 cms f o r the L i t t l e Qualicum, 13.14 cms f o r the Englishman, 14.39 cms f o r the Oyster, 17.98 cms f o r the Kokish and 18.75 cms f o r the Chemainus. The lowest flows occur i n August and September at a l l but the Coldstream s t a t i o n where low flows occur i n March. Maximum flows occur i n December and January at a l l but the Oyster, Coldstream and Nanoose s t a t i o n s . June i s the month of maximum d i s c h a r g e at the Oyster and Goldstream s t a t i o n s and A p r i l at Nanoose. During the growing season, A p r i l through September i n c l u s i v e , the mean d i s c h a r g e v a r i e s from 87.4 cms at the Campbell to .1 cms at Nanoose. The d i s c h a r g e r a t e s of the Goldstream, Cowichan, Nanaimo, Big Qualicum, Puntledge, Tsolum, Campbell and Salmon Ri v e r s are r e g u l a t e d i n many ways and by d i f f e r e n t agents; the r a t e of d i s c h a r g e v a r i e s depending on the agent. D e t a i l s may be obtained from the S p e c i a l Estuary S e r i e s r e p o r t s , C i t y of V i c t o r i a , F e d e r a l Department of F i s h e r i e s and Town of Sayward. The suspended sediment c o n c e n t r a t i o n of each r i v e r has been estimated though not measured. The r i v e r s south of Courtenay flow over the Late Mesozoic Sediments, S i c k e r Group and g l a c i a l t i l l , and have an estimated suspended sediment c o n c e n t r a t i o n of 0 - 50 m i l l i g r a m s per l i t e r (mg/1). The r i v e r s n orth of Courtenay mainly flow over b a s a l t i c l a v a s , g r a n i t e d i f f e r e n t i a t e s and g l a c i a l t i l l , and have an estimated Table 5. Hydrologies 1 informntion pertaining to the study areas Es tuary Watershed thins tern Tribu- Gauging 2 Min. monthly 2 Max. monthly 2 Avg. A p r i l Mean Annual size' length 1 taries' Station mean discharge mean discharge to Sept. Discharge sq. km. km. Nuntoer cms cms mean discharge cms oris Nirpkish 1989 62.4 24 081iF002 Aug 35.40 Jon 202.75 85.4 1926-1938 129.13 Ki ngecme 1456 74.4 24 - -• _ Car^'bel 1 1432 9.7 8 08HD003 Sept 66.83 Dec 132.24 87.4 1949-1970 98.83 Salnon 1330 82.1 55 08ID006 Aug 16.79 Deo 105.91 50.3 1956-1970 66.54 Covvi chan 834 45.5 23 08HA011 Aug 7.08 Deo 116.95 29.29 1960-1970 53.24 Courtenay (see 769 2.4 2 - - _ Puntledge and T s o l i n below) Nano ino 725 80.5 14 08HB034 Aug 5.13 Dec 90.05 22.8 1965-1976 41.91 .Ad mo-Eve 622 59.9 55 - - _ Pui;t 1 edge 518 18.1 1 0810)006 Aug 22.76 Dec 56.92 39.2 1914-1920. 1955-1957, 1964-1970.43.33 Ch era inus 378 64.4 62 08il'\001 Aug 1.16 Dec 38.51 9.6 1914-1970, 18.75 Ts i t i ka 376 38.6 42 08I1F004 18.6 Kok i sh 357 20.9 15 0811F001 Aug 3.68 • Dec 30.02 12.4 1927-1941 17.98 En>:l i shnan 287 35.4 27 08IB002 Sept 1.03 Dee 22.46 8.4 1913-1917, 1970 13.14 Tsolun ' 251 28.5 10 0810301 1 Aug .84 Dec 24.27 5.0 1914-1917. 1955-1957, 1964-1970 11.02 Li t t ie CXial i c i n 249 19.3 5 0RI1B0293 Aug 2.103 Jan 24.663 7.3 1960-1973 12.323 Oyster 181 43.5 22 081ID0023 Sopt 5.013 June 24.723 15.5 1914-1917 14.393 Big Ouil i c i n 145 25.1 23 08HR001 Aug 1.89 Jan 15.66 4.3 1913-1922, 1956-1970 7.39 Cluxewe 117 32.2 36 - - estimated 4.25 Ju l y 8, 1974' GoMst renin 70 11.1 6 08ND012 f.fer 6.23 June 140.45 - 1955-1970 39.07 CViatse 52 12.1 11 - — - estimated .42 Ju l y 5, 1974' Bonel1 47 12.6 - - - _ _ Nanoose 28 14.3 3 08W30393 Sept .023 Apr .603 .1 1970-1972 3 Water l ' 2 Diversion Suspended Sediment Concent rat ion m^ /1 No 0-50 No 201-400 Regulated since 1947 0-50 Regulated 0-50 Regulated since 1965 51-200 No C-50 Regulated since 1983 51-200 No 0-50 Regulated 0-50 No 51-200 No 0-50 No 0-r.o No 5I--JH0 Regulated since 1964 0-50 No 51-200 No 0-50 Regulated 51-200 No 0-50 Regulated 51-200 No 0-50 No 51-200 No 51-200 'Qr. iss . 1975 * Deportment of the Environment, 1971 3 Department of the Environment, 1974 •* S t i c h l ing, 1974 - 17 -c o n c e n t r a t i o n of 51 -200 mg/1. The Kingcome R i v e r , flows over g r a n o d i o r i t e and g l a c i a l t i l l and i s fed by l a r g e g l a c i e r s on the mainland c o a s t . It i s estimated to have a 201 - 400 mg/1 suspended sediment c o n c e n t r a t i o n . 1.5 F i s h L a t e l y , i t has become e v i d e n t t h a t t i d a l marshes are u t i l i z e d by j u v e n i l e salmon d u r i n g , what may be, a c r i t i c a l aspect of t h e i r l i f e h i s t o r y (Dorcey et a l . , 1973).- The f i s h s p e c i e s found i n the r i v e r systems f l o w i n g i n t o the e s t u a r i e s s t u d i e d i n c l u d e coho, chum, chinook, pink and sockeye salmon, s t e e l h e a d , and rainbow, c u t t h r o a t and D o l l y Varden t r o u t . H e r r i n g spawn i n s u b t i d a l waters of the Goldstream, Cowichan, Nanaimo, Nanoose-Bonell, Courtenay, Nimpkish and Cluxewe R i v e r s ' d e l t a s (Table 6 ) . D o l l y Varden, rainbow and c u t t h r o a t t r o u t are p r e s e n t i n a l l the r i v e r systems and D o l l y Varden and c u t t h r o a t may be found i n the e s t u a r i n e systems d u r i n g t h e i r l i f e h i s t o r i e s . Coho occur i n a l l e s t u a r i e s but the Kingcome; chum are p resent i n a l l but the Adam-Eve, Kokish, Nimpkish, Cluxewe and Kingcome; chinook are found i n a l l but the Nanoose, B o n e l l , Oyster, Adam-Eve, Cluxewe and Quatse; pink occur only i n the Courtenay, Salmon, Adam-Eve, T s i t i k a , Quatse and Kingcome; sockeye occur on l y i n the Englishman, L i t t l e Qualicum, Campbell, Cluxewe and Quatse R i v e r systems. Steelhead may be found i n a l l but the B o n e l l and Big Qualicum River systems. The s i z e , year and trends of the salmon runs i n each system may be obtained from Oguss e t . aJ.. (1975) and Environment Canada, F i s h e r i e s and Oceans salmon escapement r e c o r d s . Table 6. Economic Fish Species in the River Systems Estuary c Mainstem Length Ion Mains tern Length Accessible to Migrant r i s h km Coho Chum Chinook Pink Sockeye Steelhead Rainbow Cutthroat Holly Varden Herring Goldstream 11.1 - X X X X X X X X Cowichan River 45.5 45.5 x X X X X X X X Chemainus River 64.4 12.1 62.8 x X X X X X X Nanaimo River 80.5 64.4 X X X X X X X Nanoose Creek 14.3 4.8 X X X X X X X Bone 11 Creek 12.6 4.8 X X X X X X Englishman River 35.4 16.1 X X X X X X X X L i t t l e Qualicum River 19.3 11.6 X X X X X X X X Big Qualicum River 25.1 10.5 X X X X X X Courtenay River 2.4 2.4 X X X X X X X X X Puntledge River 18.1 - X X X X X X X Tsolum River 28.5 22.5 X X X X X X X Oyster River 43.5 35.4 X X X X X X X Campbell River 9.7 5.6 X X X X X X X X Salmon River 82.1 35.4 X X X X X X X X Adam-Eve Rivers 59.9 8.9 X X X X X X Tsi tika River 38.6 33.8 X X X X X X X X Kokish River 20.9 17.7 X X X X X X Nimpkish River 62.4 • - X X X X X X X Cluxeive River 32.2 20,9 X X X X X X X Quatse River 12.1 12.1 X X X X X X X X Ki ngcome River 74.4 - X X X X X X Oguss 1975 and G. Reid Pers. Comm. - 19 -2. LITERATURE REVIEW " S a l t marshes of the North American P a c i f i c Coast have been l i t t l e s t u d i e d and the r e l a t i o n s h i p s between s p e c i e s and t h e i r environment and among c o - e x i s t i n g s p e c i e s are poorly understood ( Z e d l e r , 1977)." MacDonald and Barbour (1974) found t h i s p a r t i c u l a r l y so f o r B r i t i s h Columbia; Calder and T a y l o r ' s (1968) d e s c r i p t i o n of s a l t marsh communities i n the Queen C h a r l o t t e I s l a n d s i s t h e i r only l i t e r a t u r e source. Most of the data p e r t a i n i n g to B r i t i s h Columbia's c o a s t a l marshes are found i n government r e p o r t s and u n i v e r s i t y theses; most focus on the Fraser and Squamish R i v e r s ' e s t u a r i e s . Burgess (1970) was the f i r s t to d e s c r i b e the composition of v e g e t a t i o n on Roberts and Sturgeon Banks of the F r a s e r R i v e r d e l t a . He found that s p e c i e s d i s t r i b u t i o n appeared to be determined by t i d a l f l o o d i n g , l o c a l drainage and p o s s i b l y by undemonstrated d i f f e r e n c e s i n s o i l and water s a l i n i t y . Seed p r o d u c t i o n was measured ; softstem bulrush and Lyngby's sedge produced the most seeds c l o s e l y f o l l o w e d by three square b u l r u s h . Forbes (1972) and McLaren (1972) mapped the d i s t r i b u t i o n of the f l o r a on the F r a s e r R i v e r d e l t a f o r e s h o r e , and i s l a n d s i n the south arm of the F r a s e r River, r e s p e c t i v e l y . The f o r e s h o r e v e g e t a t i o n of the F r a s e r R i v e r d e l t a was d e s c r i b e d again by Yamanaka (1975). He c o l l e c t e d p l a n t and s o i l samples along f o u r t e e n t r a n s e c t s running seaward from shore and determined an average dry matter y i e l d of 4.9 tons per h e c t a r e over an estimated 1901 h e c t a r e s . Ash, n i t r o g e n and l i g n i n were determined f o r p l a n t samples, and pH, o r g a n i c matter and e l e c t r i c c o n d u c t i v i t y were determined f o r s o i l samples. P l a n t s p e c i e s composition and average height of - 20 -spe c i e s were recorded. L a t e r s t u d i e s c o n f i n e d d e s c r i p t i o n s to s p e c i f i c areas on the F r a s e r R i v e r d e l t a and used a l a r g e r s c a l e f o r mapping. H i l l a b y and B a r r e t t (1976) used sys t e m a t i c p l o t s to d e s c r i b e the Roberts Bank s a l t marsh and i n d i c a t e the 4.6 meter t i d a l contour was the l i n e d e l i n e a t i n g the emergent (gumweed and s a l t b u s h ) and submergent ( s a l t g r a s s and salt w o r t ) communities. The Musqueam Marsh, below P o i n t Grey, was s t u d i e d by B e l l - I r v i n g (1977). P l a n t s p e c i e s d e n s i t y and hei g h t , and benthos were randomly sampled along four t r a n s e c t s . Heights of dominant s p e c i e s ranged f o r c a t - t a i l 128 - 220 cm, Lyngby's sedge 111 - 180 cm, and sof t s t e m b u l r u s h 102 - 234 cm. Moody (1978) r e l a t e d environmental f a c t o r s and s p e c i e s d i s t r i b u t i o n on the Brunswick P o i n t Marsh. She found 2.82 meters (above c h a r t datum) i s the lower l i m i t of emergent v e g e t a t i o n , s a l i n i t y and temperature i n f l u e n c e growth, and a peak aboveground phytomass of 909 gm m-2 f o r Lyngby's sedge, 397 gm m-2 f o r three square b u l r u s h , and 565 gm m~2 f o r maritime b u l r u s h . The Lyngby's sedge marsh on Woodward Is l a n d was s t u d i e d by K i s t r i t z and Yesaki (1979). They measured shoot growth, d e n s i t y , s t a n d i n g crop, root biomass, and carbon, n i t r o g e n and phosphorus content of t i s s u e over one year. Annual net primary p r o d u c t i o n was estimated to be 634 grams as h - f r e e dry weight per square meter (gm AFDW m~2). Net annual d e t r i t u s p r o d u c t i o n was estimated to be 435 gm AFDW m~2 and 62% of t h i s disappeared i n d i s s o l v e d and p a r t i c u l a t e o r g a n i c matter between September and June; the balance was probably b u r i e d by a l l u v i u m . Belowground biomass was approximately four times l a r g e r than aboveground standing crop. The emergent p l a n t communities of the Squamish Ri v e r e s t u a r y are d e s c r i b e d by Lim and Levings (1973). The - 21 -biomass of Lyngby's sedge communities ranged from 573-1657 gm dryweight (DW) m_2 (average 924 gm DW m -2). Levings and Moody (1976) measured net primary p r o d u c t i o n , over the p e r i o d l a t e March to mid-August, at 1322 gm DW m-2 on the c e n t r a l lobe of the Squamish R i v e r d e l t a . The emergent v e g e t a t i o n of other B r i t i s h Columbia e s t u a r i e s i s d e s c r i b e d i n the S p e c i a l Estuary S e r i e s , Task Force, and r e l a t e d environmental c o n s u l t a n t s r e p o r t s . The S p e c i a l Estuary S e r i e s l i s t s the f l o r a and maps the d i s t r i b u t i o n of major s p e c i e s or p l a n t communities f o r the Cowichan, Chemainus, Nanaimo, K i t i m a t , Campbell, Courtenay and Oyster R i v e r s ' e s t u a r i e s . Task f o r c e r e p o r t s provide much the same data f o r the Cowichan, Nanaimo, and Squamish R i v e r s ' e s t u a r i e s . Most of the i n f o r m a t i o n i n the S p e c i a l Estuary S e r i e s and Task Force r e p o r t s i s from government f i l e s and unpublished data. The d e t a i l of the maps v a r i e s from . d e l i n e a t i o n of vegetated areas through lumping v e g e t a t i o n types to p l a n t community o u t l i n e s . K i s t r i t z (1978) reviewed the l i t e r a t u r e on the r o l e of d e t r i t u s and the c y c l i n g of elements i n t i d a l marsh ecosystems. Much of the l i t e r a t u r e i s based on re s e a r c h c a r r i e d out i n s a l t marshes on the east coast of the United S t a t e s and i t s a p p l i c a b i l i t y to B r i t i s h Columbia i s l i m i t e d . There i s a l a r g e degree of v a r i a b i l i t y both w i t h i n and between marsh ecosystems as a r e s u l t of v a r i a t i o n i n g e o l o g i c a l , h y d r o l o g i c a l and c l i m a t o l o g i c a l f a c t o r s . He concluded there i s a need f o r data s p e c i f i c to west coast marshes. - 22 -3. THE PLANT COMMUNITIES OF THE ESTURARINE MARSHES ON THE EAST COAST OF VANCOUVER ISLAND The p l a n t communities i n t h i s study are composed of emergent p l a n t s p e c i e s that l i v e i n b r a c k i s h to s a l i n e water. The emergent p l a n t s , or v e g e t a t i o n , are v a s c u l a r p l a n t s that occur i n the i n t e r t i d a l zone, between the low and high high t i d e l i n e s , on r i v e r d e l t a s . There they are s u b j e c t to the f l o o d and ebb of the t i d e ; o f t e n submerged on the f l o o d t i d e , they "emerge" from the water on the ebb t i d e . Emergent v e g e t a t i o n i s a primary source of energy i n the e s t u a r i n e ecosystem and i s u t i l i z e d , e i t h e r d i r e c t l y or i n d i r e c t l y , by many organisms and most v i s i b l y by b i r d s , f i s h and w i l d l i f e . One of the i n i t i a l steps i n d e l i m i t i n g the energy pathways i s determining the p l a n t s p e c i e s and communities i n the e s t u a r y . T h i s s e c t i o n of the study d e s c r i b e s and r e p o r t s on the d i s t r i b u t i o n of p l a n t communities. It a l s o attempts to i d e n t i f y some of the major f a c t o r s i n f l u e n c i n g the p l a n t s p e c i e s d i s t r i b u t i o n . 3.1 Methods The emergent p l a n t communities are d e s c r i b e d a f t e r the a e r i a l photograph technique i n Mueller-Dombois (1974) and Daubenmire (1962). The technique was p a r t i c u l a r l y s u i t a b l e f o r t h i s study as i t i s comparatively f a s t e r to do than other procedures which made i t p o s s i b l e to d e s c r i b e the p l a n t communities on the nineteen e s t u a r i e s i n two f i e l d seasons (May to September, 1975 and 1976). T h i s would not have been - 23 -September, 1975 and 1976). T h i s would not have been p o s s i b l e using another technique. Though the a e r i a l photograph procedure does not produce q u a n t i t a t i v e data on p l a n t s p e c i e s , i t i s a s u i t a b l e technique f o r t h i s study because the work i s p r e l i m i n a r y and the technique i s r e l i a b l e , as shown by E i l e r s (1975). E i l e r s s t u d i e d Oregon's e s t u a r i n e marshes and found plant communities i d e n t i f i e d using t h i s technique were c l o s e l y r e l a t e d to those i d e n t i f i e d with SIMORD (an o r d i n a t i o n program based on the p h y t o s o c i o l o g i c a l r e f e r e n c e stand o r d i n a t i o n procedure of Whittaker (1967)). With s u b s t a n t i a t i o n of the technique f o r e s t u a r i n e marshes i t was p o s s i b l e to map the l o c a t i o n and extent of the p l a n t communities with confidence. The one disadvantage of the technique i s that the t r a n s i t i o n zone between communities may vary i n width; thus p l a n t communities presented should be c o n s i d e r e d as graduating from c e n t r a l c h a r a c t e r i s t i c s toward the margins. v a r i a t i o n s a s s o c i a t e d with p l a n t communities on a e r i a l photographs (Appendix 3) and f i e l d checking, i n the years 1975 and 1976. P l a n t s p e c i e s i n each community were catalogued, i n c l u d i n g more t e r r e s t r i a l s p e c i e s j u s t above the mean high water mark (Appendix 4 ) . Species were i d e n t i f i e d according to Hitchcock et a l . (1973). The dominant and subdominant p l a n t s p e c i e s , those with the h i g h e s t o c u l a r e s t i m a t i o n of percent coverage of a community, c h a r a c t e r i z e the community. F i e l d the S p e c i a l C o l l e c t i o n s D i v i s i o n at the U n i v e r s i t y of B r i t i s h The procedure i n v o l v e d determining t o n a l and t e x t u r a l notes c o n t a i n i n g the percent coverage estimates are on Columbia. - 24 -P l a n t communities i n each estuary were mapped from the a e r i a l photographs so that map and photos may be c o o r d i n a t e d . C o r r e c t i o n f o r a b e r r a t i o n was beyond the resources of t h i s study so, where p o s s i b l e , only the c e n t r e of each a e r i a l photograph was used to compose a map. The area of each community was c a l c u l a t e d and a simple l i n e a r c o r r e l a t i o n (Zar, 1974) was run between the t o t a l area of eleven dominant or subdominant s p e c i e s t y p i n g the communities and average A p r i l to September mean flows (Department of the Environment, 1974), and A p r i l to September mean r a i n f a l l (Atmospheric Environment, 1973). 3.2 R e s u l t s and D i s c u s s i o n The d e s c r i p t i o n s of the p l a n t communities i n nineteen e s t u a r i e s on the east coast of Vancouver I s l a n d r e q u i r e d the r e c o r d i n g of a l o t of d e t a i l and i t i s d i f f i c u l t to present the data here without a s u b s t a n t i a l l o s s of coherence i n the t e x t . T h e r e f o r e , the data i s presented i n t a b l e s 7 to 45 and f i g u r e s 4 to 22 i n Appendix 5. There i s a great deal of v a r i a t i o n i n the d e s c r i p t i o n and d i s t r i b u t i o n of the p l a n t communities i n the e s t u a r i e s s t u d i e d . The d e s c r i p t i v e v a r i a b i l i t y breaks the e s t u a u r i e s i n t o four main groups. In the f i r s t group of e s t u a r i e s v e g e t a t i o n i n d i c a t i v e of b r a c k i s h c o n d i t i o n s occurs around the main r i v e r channels. Lyngby's sedge i s found at the lowest e l e v a t i o n s and with i n c r e a s i n g e l e v a t i o n B a l t i c rush and c i n q u e f o i l . In areas d i s t a n t from the i n f l u e n c e of freshwater there i s an i n c r e a s e i n the occurence of p l a n t s p e c i e s i n d i c a t i v e of more s a l i n e - 25 -c o n d i t i o n s . S a l t w o r t i s found at the lowest e l e v a t i o n s and with i n c r e a s i n g e l e v a t i o n s a l t g r a s s , then s a l t b u s h , gumweed, meadow and mouse b a r l e y , and other g r a s s e s . In t h i s group maritime b u l r u s h r a t h e r than Lyngby's sedge i s present when the su b s t r a t e i s water s a t u r a t e d , and j o i n t e d rush, r a t h e r than B a l t i c rush, i s present when the s u b s t r a t e drainage i s poor. E s t u a r i e s i n t h i s group are the Cowichan, Chemainus, Nanaimo, Englishman, L i t t l e Qualicaum and Big Qualicum. In the second group of e s t u a r i e s , s p e c i e s i n d i c a t i v e of b r a c k i s h c o n d i t i o n s dominate the e s t u a r i e s . Lyngby's sedge i s dominant at the lowest e l e v a t i o n s of the vegetated i n t e r t i d a l zone. As the e l e v a t i o n i n c r e a s e s t u f t e d h a i r g r a s s then c i n q u e f o i l and a s s o r t e d s p e c i e s , mainly grasses, dominate. When the ecotone between marsh and f o r e s t i s p o o r l y drained bulrush and sweet g a l e are present, and when i t i s w e l l d r a i n e d , willow, twin-berry, Nootka rose and ninebark dominate. E s t u a r i e s i n t h i s group are the Courtenay, Oyster, Campbell, Salmon, Adam-Eve, T s i t i k a , Kokish, Nimpkish, Cluxewe, Quatse and Kingcome. In the t h i r d group of e s t u a r i e s , Lyngby's sedge occurs only i n channels where freshwater flows c o n s t a n t l y , and the marsh i s dominated by s a l i n e i n d i c t o r s p e c i e s . Saltwort i s dominant at the lowest e l e v a t i o n s i n the vegetated i n t e r t i d a l zone and s a l t g r a s s then s a l t b u s h , maritime p l a n t a i n and gumweed become dominant as the e l e v a t i o n i n c r e a s e s . S a l t marshes i n t h i s group are the Goldstream, Chemainus and Nanoose-Bonell. In the f o u r t h group of e s t u a r i e s Lyngby's sedge or sedge species are l i m i t e d to channels where freshwater flows c o n s t a n t l y and s a l i n e i n d i c a t o r s dominate. Saltwort dominates at the lowest e l e v a t i o n s and, with i n c r e a s i n g e l e v a t i o n , arrow-- 26 -grass becomes dominant and may be a s s o c i a t e d with t u f t e d h a i r g r a s s , Kentucky b l u e g r a s s , b a r l e y and other grasses at higher e l e v a t i o n s . S a l t marshes i n t h i s group are the Adam-Eve, Kokish and Cluxewe. The f o u r groups b r i n g out two p o i n t s . The f i r s t p o i n t i s that b r a c k i s h and s a l t marshes are d e s c r i b e d , the s a l t marshes being on the same d e l t a s as the b r a c k i s h marshes. The second p o i n t i s that the dominant p l a n t s p e c i e s at higher e l e v a t i o n s i n marshes north and south of Courtenay are d i f f e r e n t . In b r a c k i s h marshes south of Courtenay rushes, c i n q u e f o i l and some s a l t g r a s s dominate while marshes from Courtenay north are dominated by t u f t e d h a i r g r a s s and c i n q u e f o i l . And i n s a l t marshes south of Courtenay s a l t g r a s s , s a l t b u s h and gumweed dominate while north of Courtenay arrow-grass and grasses dominate. The two p o i n t s may be e x p l a i n e d by c o n s i d e r i n g the dominant p l a n t s p e c i e s . P l a n t s p e c i e s are known to i n d i c a t e t h e i r growing c o n d i t i o n s (Clements, 1939) and two i n d i c a t o r s p e c i e s are s a l t w o r t and Lyngby's sedge. The dominance of saltwo r t i n d i c a t e s s a l i n e c o n d i t i o n s and Lyngby's sedge i n d i c a t e s b r a c k i s h c o n d i t i o n s . Saltwort and Lyngby's sedge separate s a l t marshes from b r a c k i s h marshes and i d e n t i f y s a l i n i t y as a major f a c t o r i n f l u e n c i n g s p e c i e s d i s t r i b u t i o n . S a l i n i t y appears to be the f a c t o r s p l i t t i n g the i s l a n d i n h a l f as b r a c k i s h and s a l t marshes south of Courtenay have more s a l i n e i n d i c a t o r s p e c i e s dominant than do b r a c k i s h and s a l t marshes north of Courtenay. S a l i n i t y i s a w e l l known f a c t o r i n f l u e n c i n g species d i s t r i b u t i o n on b r a c k i s h and s a l t marshes. But s a l i n i t y i t s e l f i s a f u n c t i o n of many f a c t o r s and i t i s those f a c t o r s and t h e i r i n f l u e n c e on s a l i n i t y that are of i n t e r e s t . Consider the - 27 -f i r s t p o i n t again; that i s , there are b r a c k i s h and s a l t marshes because of a d i f f e r e n c e i n s a l i n i t y . A h y p othesis i s t h a t , on Vancouver I s l a n d , the d i f f e r e n c e i n s a l i n i t y i s due to stream flow, the s i z e of the d e l t a and freshwater c i r c u l a t i o n . When flows are low and/or i n t e r m i t t e n t i n the growing season, as they are on southeastern Vancouver I s l a n d , there i s l i t t l e freshwater to f l u s h the v e g e t a t i o n and s u b s t r a t e of s a l t s , and s a l i n i t i e s are h i g h e r . The l a r g e r the d e l t a the l e s s e f f e c t i v e the d i l u t i o n a f f e c t of freshwater. Freshwater c i r c u l a t i o n may a l s o be impeded by dykes, bermes or i s l a n d s . South of Courtenay the d i s t i n c t i o n between s a l t and b r a c k i s h marshes can be seen on the Chemainus R i v e r E s t u a r y . The Chemainus has a l a r g e d e l t a d i v i d e d i n t o s a l t and b r a c k i s h marshes by the i s l a n d s and dykes which impede freshwater c i r c u l a t i o n to the s a l t marsh on the southeastern lobe of the d e l t a . B o n s a l l Creek i s the only freshwater souce f o r the s a l t marsh. North of Courtenay the d i s t i n c t i o n i s seen on the Adam-Eve R i v e r s ' E s t u a r y . Here the s a l t and b r a c k i s h marshes are separated by a g r a v e l berme and i s l a n d s which r e s t r i c t freshwater c i r c u l a t i o n to the s a l t marsh on the e a s t e r n lobe. The freshwater source f o r the s a l t marsh i s an i n t e r m i t t e n t creek. ( S a l i n i t i e s i n s o i l cores c o l l e c t e d i n June 1976 from the marshes on the Chemainus, Cowichan, L i t t l e Qualicum and Campbell e s t u a r i e s ranged from 26.0 to 37.5 m i l l i micohms (MHO) f o r the s a l t marsh on the Chemainus and 1.79 to 15.4 m i l l i MHO f o r the b r a c k i s h marshes i n the other e s t u a r i e s . (A c o n d u c t i v i t y meter was used to measure the s a l i n i t i e s of f i l t r a t e s from s o i l p a s t e s ) . Reconsider the second p o i n t , that dominant p l a n t s p e c i e s at higher e l e v a t i o n s i n s a l t and b r a c k i s h marshes are i n d i c a t i v e - 28 -of more s a l i n e c o n d i t i o n s south of Courtenay than they are north of Courtenay. Here the hypothesis i s t h a t , on Vancouver Island, the d i f f e r e n c e i s due to p r e c i p i t a t i o n ranges from 857 mm at the V i c t o r i a I s l a n d A i r p o r t to 1507 mm at Courtenay. North of Courtenay i t i n c r e a s e s to 1637 mm at Port Hardy A i r p o r t and i t r a i n s as Por t Hardy 32% more of the year than i t does at V i c t o r i a . I f e e l r a i n f a l l moderates s a l i n i t y on b r a c k i s h and s a l t marshes and found there i s a s i g n i f i c a n t p o s i t i v e c o r r e l a t i o n between p r e c i p i t a t i o n and the area dominated by t u f t e d h a i r g r a s s and c i n q u e f o i l (Appendix 5 Table 45). Tufted h a i r g r a s s and c i n q u e f o i l being more abundant i n marshes north of Courtenay. South of Courtenay, where p r e c i p i t a t i o n i s lower, rushes, c i n q u e f o i l and s a l t g r a s s dominate at higher e l e v a t i o n s . The d e s c r i p t i v e v a r i a b i l i t y breaks the e s t u a r i e s i n t o the aforementioned four groups. However, when the p r i n c i p a l p h y s i c a l f a c t o r s i n f l u e n c i n g the d i s t r i b u t i o n , as w e l l as the d e s c r i p t i v e v a r i a b i l i t y , of the p l a n t communities are c o n s i d e r e d there are eleven groups of e s t u a r i e s . They, are: 1. Kingcome, 2. Goldstream, Chemainus, Nanoose-Bonell, 3. Adam-Eve, Kokish, Cluxewe, 4. Cowichan, Chemainus, Nanaimo, 5. Englishman, L i t t l e Oualicam, 6. Big Qualicum, 7. Courtenay, Oyster, Campbell, 8. Salmon, Quatse, 9. Kokish, Nimpkish, 10. Quatse, Adam-Eve and 11. T s i t k a . The eleven groups are based on the i n t e r a c t i o n of s i x p r i n c i p a l p h y s i c a l f a c t o r s . The f a c t o r s i n c l u d e : 1. time of maximum d i s c h a r g e , 2. the r e l a t i o n s h i p between a r i v e r ' s average A p r i l to September mean di s c h a r g e and the s i z e of the r i v e r ' s d e l t a , 3. mean annual t o t a l p r e c i p i t a t i o n , 4. the r e l a t i v e p r o t e c t i o n of a r i v e r d e l t a from wind and wave energy - 29 -of the sea, 5. the p a r t i c l e s i z e of the s u b s t r a t e , and 6. the d u r a t i o n and frequency of t i d a l i n u n d a t i o n . The f i r s t f a c t o r c o n s i d e r e d i s time of maximum d i s c h a r g e . The time of maximum di s c h a r g e f o r a r i v e r , s p r i n g or winter, i n f l u e n c e s the s a l i n i t y d u r i n g the growing season and the r a t e of a c c r e t i o n . During the s p r i n g f r e s h e t , the s a l i n i t y i s lowered by the l a r g e i n f l o w of freshwater, and the r a t e of a c c r e t i o n i s probably i n c r e a s e d when sediments are trapped i n the growing v e g e t a t i o n . R i v e r systems with a heavy s p r i n g f r e s h e t tend to o r i g i n a t e from g l a c i e r s and snow f i e l d s and carry l a r g e suspended sediment loads commonly r e f e r r e d to as g l a c i a l flow. G l a c i a l flow c o n t r i b u t e s to the r a t e of a c c r e t i o n on a d e l t a and g r e a t l y a l t e r s the l i g h t regime f o r inundated v e g e t a t i o n . R i v e r s with a heavy s p r i n g f r e s h e t a l s o have a l o t of water energy that can scour away p l a n t s thereby d e c r e a s i n g s p e c i e s d i v e r s i t y on the d e l t a s . In g e n e r a l , emergent veg e t a t i o n on d e l t a s s u b j e c t to s p r i n g f r e s h e t s i s not d i v e r s e and i s composed of b r a c k i s h i n d i c a t o r s p e c i e s d i s t r i b u t e d i n bands across the d e l t a . In r i v e r systems s u b j e c t to winter f r e s h e t s , the lower flows i n the growing season tend to r e s u l t i n higher s a l i n i t e s and lower r a t e s of a c c r e t i o n on the d e l t a s ; d u r i n g the winter, there i s o f t e n l i t t l e emergent v e g e t a t i o n to trap sediments and lower summer flows c a r r y s m a l l e r suspended sediment l o a d s . In g e n e r a l , emergent v e g e t a t i o n on d e l t a s s u b j e c t to winter f r e s h e t s i s composed of b r a c k i s h i n d i c a t o r s p e c i e s around the freshwater channels and, with i n c r e a s i n g d i s t a n c e from freshwater, an i n c r e a s i n g abundance of s a l i n e i n d i c a t o r s p e c i e s . - 30 -The second f a c t o r concerns the r e l a t i o n s h i p between a r i v e r ' s average A p r i l to September mean di s c h a r g e and the s i z e of the r i v e r ' s d e l t a . It appears that some r i v e r s c o n t i n u o u s l y inundate a l l or a l a r g e p o r t i o n of t h e i r d e l t a s during the spring f r e s h e t . T h i s long i n u n d a t i o n p e r i o d discourages the esta b l i s h m e n t of e x t e n s i v e stands of emergent v e g e t a t i o n which r e q u i r e d a y l i g h t exposure time each day of the growing season. It i s f e l t that where a d e l t a i s d i s p r o p o r t i o n a t e l y small r e l a t i v e to the r i v e r ' s average A p r i l to September mean di s c h a r g e the emergent v e g e t a t i o n w i l l not be d i v e r s e or extens i v e , and w i l l be r e s t r i c t e d to the highest e l e v a t i o n s i n the i n t e r t i d a l zone. The converse of t h i s f a c t o r i s a l s o important. When a d e l t a i s d i s p r o p o r t i o n a t e l y l a r g e r e l a t i v e to the r i v e r or creek's average A p r i l to September mean di s c h a r g e the d e l t a w i l l be c o l o n i z e d by p l a n t s p e c i e s considered i n d i c a t i v e of s a l i n e c o n d i t i o n s . B r a c k i s h i n d i c a t o r s p e c i e s w i l l be r e s t r i c t e d to freshwater channels. The t h i r d f a c t o r c o n s i d e r e d i s the mean annual p r e c i p i t a t i o n . The r o l e of p r e c i p i t a t i o n appears to l i e i n i t s i n d i r e c t e f f e c t s . In e s t u a r i n e marshes south of Courtenay, where the mean annual p r e c i p i t a t i o n i s low, l e s s than 1030mm, i t i s not enough to d i l u t e and wash away the s a l t s brought to the s o i l s u r f a c e by e v a p o t r a n s p i r a t i o n . Marshes i n t h i s p r e c i p i t a t i o n range w i l l u s u a l l y be dominated by p l a n t s p e c i e s i n d i c a t i v e of x e r i c and s a l i n e c o n d i t i o n s . In marshes north of Courtenay, p r e c i p i t a t i o n i s g r e a t e r than 1030mm, the r a i n f a l l appears to keep s a l i n i t e s r e l a t i v e l y low and the marshes are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h c o n d i t i o n s . - 31 -The f o u r t h f a c t o r c o n s i d e r e d i s p r o t e c t i o n of the r i v e r d e l t a from the wind and wave energy of the sea. The sub s t r a t e of r e l a t i v e l y p o o r l y p r o t e c t e d d e l t a s i s s u b j e c t to the e r o s i o n a l f o r c e s of the wind and waves. Hence, these d e l t a s tend to have l i m i t e d f i n e s u b s t r a t e and are mainly g r a v e l . The wind and waves may a l s o prevent s p e c i e s c o l o n i z a t i o n , and damage and uproot e s t a b l i s h e d p l a n t s . The dominant p l a n t s of many exposed d e l t a s are g r a s s e s . The deeper, f i n e s u b s t r a t e on r e l a t i v e l y w e l l p r o t e c t e d d e l t a s i s e s t a b l i s h e d to a g r e a t e r d i v e r s i t y of s p e c i e s and i n c l u d e s more f o r b s . The f i f t h f a c t o r i s the p a r t i c l e s i z e of the s u b s t r a t e . G e n e r a l l y , i t appears that the t h i c k e r the l a y e r of f i n e s the more d i v e r s e the p l a n t s p e c i e s , and, c o n v e r s e l y , the t h i n n e r the l a y e r of f i n e s the l e s s abundant and d i v e r s e , and more f i b r o u s r o o t i n g the p l a n t s p e c i e s . The s i x t h f a c t o r i s the d u r a t i o n and frequency of t i d a l i n u n d a t i o n . There are a few emergent p l a n t s that can s u r v i v e long p e r i o d s of in u n d a t i o n and a few t e r r e s t r i a l p l a n t s that can s u r v i v e i n f r e q u e n t i n u n d a t i o n . Lyngby's sedge and s a l t w o r t are the p r i n c i p a l emergent s p e c i e s that are known to t o l e r a t e i n u n d a t i o n from 38 to 62% of the d a y l i g h t hours during the growing season and S i t k a spruce i s known to t o l e r a t e f l o o d i n g f o r short p e r i o d s of time f o u r or f i v e times each year. Plants o c c u r r i n g at e l e v a t i o n s between the sedge and spruce are inundated f o r g e n e r a l l y l e s s than 30% of the d a y l i g h t hours during the growing season and i t i s i n t h i s range that the g r e a t e s t abundance of p l a n t s p e c i e s occur (Kennedy, 1978). The more ex t e n s i v e the range, the more abundant and d i v e r s e are the pl a n t s p e c i e s . The inundation to emergence r a t i o a l s o a f f e c t s the timing of the i n i t i a t i o n and c e s s a t i o n of growth, p a r t i c u l a r l y at the lov/er e l e v a t i o n s i n the i n t e r t i d a l zone. Lyngby's sedge w i l l i n i t i a t e growth as soon as i t s d a y l i g h t exposure t i n e i n c r e a s e s , i n A p r i l , and w i l l cease growth as d a y l i g h t exposure d e c l i n e s from August on. At higher e l e v a t i o n s , b r a c k i s h i n d i c a t o r species may complete t h e i r l i f e c y c l e very q u i c k l y , between A p r i l and June. Their i n i t i a t i o n of grov/th co-occurring w i t h f a v o r a b l e d a y l i g h t exposure and c e s s a t i o n of grov/th o c c u r r i n g with i n c r e a s i n g s a l i n i t y or xerophytic c o n d i t i o n s . I t appears that no one f a c t o r c o n t r o l s the composition or d i s t r i b u t i o n of a community. Rather i t i s a number of f a c t o r s and t h e i r i n t e r a c t i o n . These p h y s i c a l f a c t o r s i n t e r a c t to varying degrees and i n vari o u s ways to r e s u l t i n a m u l t i p l i c i t y of d i s t r i b u t i o n and compostion pa t t e r n s i n the plant communities on the e s t u a r i e s . However, there are general patterns among the e s t u a r i e s and these patterns lend themselves to a c l a s s i f i c a t i o n of e s t u a r i e s . A bi n u m e r i c a l key to the eleven groups of e s t u a r i e s i n t h i s study f o l l o w s . A Binumerical Key to the E s t u a r i e s l a . R iver's maximum discharge occurs i n the s p r i n g . 2 l b . River's maximum discharge occurs i n the wint e r . 7 2a. R i v e r ' s average A p r i l to September mean discharge 3 d i s p r o p o r t i o n a t e l y large r e l a t i v e to the s i z e of the d e l t a . 2b. River's average A p r i l to September mean discharge 4 p r o p o r t i o n a t e to the s i z e of the d e l t a . - 33 -3. Emergent v e g e t a t i o n i s not e x t e n s i v e . The marsh i s dominated by Carex l y n g b y e i communities which tend to occur as a f r i n g e where there i s s h e l t e r from f a s t moving water. On more e l e v a t e d areas of the d e l t a which are s u b j e c t to p e r i o d i c i n u n d a t i o n , the v e g e t a t i o n i s l i k e that found on the r i v e r ' s i s l a n d s . (eg. Skeena). 4a. Mean annual t o t a l p r e c i p i t a t i o n > 1540 mm. 5 4b. Mean annual t o t a l p r e c i p i t a t i o n < 1540 mm. 6 5. Emergent v e g e t a t i o n i s e x t e n s i v e on the d e l t a and i s dominated by Carex l y n g b y e i communities at the lowest e l e v a t i o n s i n the vegetated i n t e r t i d a l zone. E l e o c h a r i s  p a l u s t r i s i s dominant i n or adjacent to channels where freshwater flows and there i s some s u b s t r a t e drainage at low t i d e . With i n c r e a s i n g e l e v a t i o n , Deschampsia c e s p i t o s a dominates and may be found ^ n a. s o c i a t i o n with, at the lower e l e v a t i o n s of i t s range, Carex l y n g b y e i and then, with i n c r e a s i n g e l e v a t i o n , a s s o r t e d f o r b s and grasses (e.g. P o t e n t i l l a p a c i f i c a , T r i f o l i u m w o r m s k j o l d i i , Juncus  b a l t i c u s , A c h i l l e a m i l l e f o l i u m , Lathyrus p a l u s t r i s , Poa  p r a t e n s i s , Ranunculus orthorhynchus, A g r o s t i s t e n u i s , Elymus glaucus, Festuca s u b u l a t a , Bromus p a c i f i c u s ) . The number of a s s o c i a t e d s p e c i e s and p l a n t communities i n an estuary d e c l i n e s with i n c r e a s i n g l a t i t u d e . The extent of c h a n n e l i z a t i o n of the backshore i s dependent on the slope and s i z e of d e l t a ; the g e n t l e r the slope and the l a r g e r the d e l t a , the more channels. Dominant v e g e t a t i o n i n backshore areas appears to be r e l a t e d to s u b s t r a t e drainage which i s dependent on the extent of c h a n n e l i z a t i o n (see K r a j i n a - 34 -(1970) f o r a d e s c r i p t i o n o f p o s s i b l e c o m m u n i t i e s ) ( K i n g c o m e ) . 6. Emergent v e g e t a t i o n i s e x t e n s i v e on t h e d e l t a w i t h d o m i n a n t s on t h e f o r e s h o r e i n c l u d i n g S c i r p u s a m e r i c a n u s , S c i r p u s m a r i t i m u s , S c i r p u s a c u t u s , C a r e x l y n g b y e i and T y p h a  l a t i f o l i a . Towards t h e b a c k s h o r e , t h e v e g e t a t i o n i s d i v e r s e and i t s d i s t r i b u t i o n a p p e a r s t o be i n f l u e n c e d by s u b s t r a t e d r a i n a g e ; bog, f e n and swamp c o m m u n i t i e s a r e common away f r o m main c h a n n e l s , and b r a c k i s h marshes o f d i v e r s e s p e c i e s o c c u r a l o n g c h a n n e l s ( e . g . S c i r p u s a c u t u s , Typha l a t i f o l i a , C a r e x l y n g b y e i , C a r e x spp., E r i g e r o n s p p . , A s t e r s p . , S i d a l c e a h e n d e r s o n i i , J u n c u s s p p . , Sonchus  a r v e n s e , S a g i t t a r i s l a t i f o l i a , A l i s m a p l a n t a g o - a q u a t i c a , and A g r o s t i s s p p . ) . A d j a c e n t a r e a s o f ponded water a r e c o l o n i z e d by pondweeds ( e . g . Potamogeton s p p . Lemna m i n o r , Lemna t r i s u l c a , S p a r g a n i u m s p . , Nuphar sp.) ( e g . F r a s e r ) . 7a. R i v e r o r c r e e k ' s a v e r a g e A p r i l t o September mean 8 d i s c h a r g e i s d i s p r o p o r t i o n a t e l y s m a l l r e l a t i v e t o t h e s i z e o f d e l t a , a n d / o r the f r e s h w a t e r c i r c u l a t i o n may be impeded t o p a r t o f t h e d e l t a by p h y s i c a l s t r u c t u r e s . 7b. R i v e r ' s a v e r a g e A p r i l t o September mean d i s c h a r g e 11 p r o p o r t i o n a t e t o t h e s i z e o f d e l t a . 8a. Mean a n n u a l t o t a l p r e c i p i t a t i o n i n t h e r e g i o n o f t h e 9 e s t u a r y i s 660 -1540 mm. 8b. Mean a n n u a l t o t a l p r e c i p i t a t i o n i n t h e r e g i o n o f t h e 10 e s t u a r y i s > 154 mm. 9. C a r e x l y n g b y e i i s l i m i t e d t o c h a n n e l s where f r e s h w a t e r . f l o w s c o n s t a n t l y . S a l i c o r n i a v i r g i n i c a i s dominant a t t h e l o w e s t e l e v a t i o n s i n t h e v e g e t a t e d i n t e r t i d a l z o ne. W i t h - 35 -i n c r e a s i n g e l e v a t i o n D i s t i c h l i s s p i c a t a becomes dominant and s p e c i e s d i v e r s i t y i n c r e a s e s (eg. A t r i p l e x p a t u l a , Plantago maritima, Hordeum brachyantherum, Hordeum murinum, Plantago l a n c e o l a t a , A c h i l l e a m i l l e f o l i u m and G r i n d e l i a  i n t e g r i f o l i a ) . Grasses and shrubs become dominant towards the backshore and s p e c i e s d i v e r s i t y i s p r o p o r t i o n a t e to the s i z e of the d e l t a . (Goldstream, Chemainus, Nanoose-B o n e l l ) . 10. Stands of Carex l y n g b y e i or Carex spp. are l i m i t e d to channels where freshwater flows c o n s t a n t l y . S a l i c o r n i a  v i r g i n i c a i s dominant at the lowest e l e v a t i o n s i n the vegetated i n t e r t i d a l zone. With i n c r e a s i n g e l e v a t i o n , T r i g l o c h i n maritimum becomes dominant and may be a s s o c i a t e d with Carex l y n g b y e i and/or Deschampsia c e s p i t o s a at the higher e l e v a t i o n s w i t h i n i t s range. Grasses (Deschampsia c e s p i t o s a , Agropyron repens, Poa  p r a t e n s i s , Hordeum brachyantherum, A g r o s t i s t e n u i s ) tend to dominate at the hi g h e s t e l e v a t i o n s i n the marsh (Adam-Eve, Kokish, Cluxewe). 11a. Mean annual t o t a l p r e c i p i t a t i o n i n the re g i o n of the 12 es t u a r y i s 660 - 1540 mm. l i b . Mean annual t o t a l p r e c i p i t a t i o n i n the r e g i o n of the 19 estu a r y i s > 1550 mm. 12a. Mean annual t o t a l p r e c i p i t a t i o n i n the r e g i o n of the 13 es t u a r y i s 660 - 1030 mm. 12b. Mean annual t o t a l p r e c i p i t a t i o n i n the r e g i o n of the 18 estu a r y i s 1040 - 1540 mm. 13a. River d e l t a i s s h e l t e r e d . 14 13b. R i v e r d e l t a i s exposed. 17 - 36 -14a. R i v e r d e l t a i s embayed. 15 14b. R i v e r d e l t a i s mostly p r o t e c t e d by a s p i t . 16 15. At low t i d e , the r i v e r ' s flow i s mainly c o n f i n e d to one channel and v e g e t a t i o n i n d i c a t i v e of b r a c k i s h c o n d i t i o n s occurs around i t . In areas d i s t a n t from the i n f l u e n c e of freshwater d u r i n g low t i d e there i s an i n c r e a s e i n the occurrence of p l a n t s p e c i e s i n d i c a t i v e of more s a l i n e c o n d i t i o n s , p a r t i c u l a r l y where freshwater c i r c u l a t i o n i s impeded. In the vegetated i n t e r t i d a l zone, Carex l y n g b y e i i s dominant at the lowe""*" e l e v a t i o n s where i t i s f l u s h e d by freshwater on the ebbing t i d e . Juncus b a l t i c u s and P o t e n t i l l a p a c i f i c a become dominant as the e l e v a t i o n i n c r e a s e s . Juncus a r t i c u l a t u s may occur at t h i s e l e v a t i o n when the s u b s t r a t e i s not w e l l d r a i n e d . Should drainage of part or a l l of the d e l t a be impeded so that the s o i l i s always water s a t u r a t e d S c i r p u s maritimus w i l l be the dominant p l a n t s p e c i e s (Cowichan, Chemainus, Nanaimo). 16. The p l a n t communities are l o c a t e d landward of the s p i t which p r o t e c t s the v e g e t a t i o n from the worst wind and wave energy of the sea. Carex l y n g b y e i i s dominant at the lowest e l e v a t i o n . As the e l e v a t i o n i n c r e a s e s the sedge becomes codominant with g r a s s e s . Shrubs (e.g. Rosa nutkana, Rubus  s p e c t a b i l i s , Rubus l a c i n i a t u s , C y t i s u s s c o p a r i u s ) and g r a s s e s are dominant i n the backshore. V e g e t a t i o n landward of a s s o c i a t e d dunes or bermes i s l i k e that d e s c r i b e d under 9 (Englishman, L i t t l e Qualicum). 17. The emergent v e g e t a t i o n i s u s u a l l y c o n f i n e d to the streambed of the main channel and grows on sand and sandy g r a v e l s . Carex l y n g b y e i occurs at the lower e l e v a t i o n s and - 37 -does not grow v i g o r o u s l y . The channel s i d e s are s p a r s e l y covered by P o t e n t i l l a p a c i f i c a , Plantago l a n c e o l a t a , T r i f o l i u m pratense and Anthoxanthum odoratum. Alnus rubra occurs on the streambank. Emergent v e g e t a t i o n (e.g. Carex  lyngb y e i and Juncus b a l t i c u s ) w i l l occur i n d e l t a i c channels landward of beach bermes. Grasses occur between a berme and channels, and grasses and shrubs occur between the channels and f o r e s t margin (Big Qualicum). Carex l y n g b y e i i s dominant at the lowest e l e v a t i o n s of the vegetated i n t e r t i d a l zone. As the e l e v a t i o n i n c r e a s e s , the sedge's v i g o r and abundance decrease and Deschampsia  c e s p i t o s a becomes dominant. S c i r p u s americanus may occur i n t h i s e l e v a t i o n range when the s u b s t r a t e i s w e l l d r a i n e d , sandy and s a l i n e . At higher e l e v a t i o n s P o t e n t i l l a p a c i f i c a becomes dominant and a s s o c i a t e d s p e c i e s may i n c l u d e , T r i f o l i u m w o r m s k j o l d i i , E l e o c h a r i s p a l u s t r i s , E r i g e r o n  p h i l a d e l p h i c u s , Dodecatheon pulc h e l l u m , S i s y r i n c h i u m  a n g u s t i f o l i u m , C a s t i l l e j a l e v i s e c t a , S i d a l c e a h e n d e r s o n i i , Mimulus g u t t a t u s , V i c i a g i g a n t e a , Habenaria d i l a t a t a , D a c t y l i s glomerata, Phleum pratense, Poa p r a t e n s i s , Holcus  lana t u s , A g r o s t i s t e n u i s , Agropyron repens and Festuca p r a t e n s i s . When backshore areas adjacent to the t r e e l i n e are not w e l l d r a i n e d Myrica gale and S c i r p u s acutus or S c i r p u s microcarpus and Carex l y n g b y e i dominate. When backshore areas adjacent to the t r e e l i n e are w e l l d r a i n e d shrubs (e.g. Physocarpus c a p i t a t u s , S a l i x spp., Lon i c e r a  i n v o l u c r a t a and Rosa nutkana) dominate. P i c e a s i t c h e n s i s i s dominant i n the f o r e s t . The extent and s p e c i e s d i v e r s i t y of each p l a n t community i n t h i s type of estuary - 38 -i s p r o p o r t i o n a t e to the s i z e of the p r o t e c t e d area with good freshwater c i r c u l a t i o n , i . e . , the sma l l e r the pr o t e c t e d area with good freshwater c i r c u l a t i o n the l e s s e x t e n s i v e and d i v e r s e the p l a n t communities (Courtenay, Oyster, Campbell). 19a. River d e l t a i s r e l a t i v e l y w e l l p r o t e c t e d from wind and 20 wave energy of the sea. 19b. R i v e r d e l t a i s r e l a t i v e l y p o o r l y p r o t e c t e d from wind 27 and wave energy of the sea. 20a. River d e l t a i s deeply embayed. 21 20b. R i v e r d e l t a i s s h a l l o w l y embayed. 24 21a. A r e l a t i v e l y t h i c k l a y e r of f i n e 22 s u b s t r a t e o v e r l i e s g r a v e l . 21b. A r e l a t i v e l y t h i n l a y e r of f i n e 23 s u b s t r a t e o v e r l i e s g r a v e l . 22. Carex l y n g b y e i i s dominant at the lowest e l e v a t i o n s of the vegetated i n t e r t i d a l zone. As the e l e v a t i o n i n c r e a s e s the sedge's v i g o r and abundance decrease and Deschampsia  c e s p i t o s a becomes dominant. At higher e l e v a t i o n s P o t e n t i l l a p a c i f i c a i s a dominant s p e c i e s and may be a s s o c i a t e d with Poa p r a t e n s i s , A g r o s t i s a l b a var. s t o l o n i f e r a , Juncus b a l t i c u s , Agropyron repens, T r i f o l i u m  w o r m s k j o l d i i , A c h i l l e a m i l l e f o l i u m , S i d a l c e a h e n d e r s o n i i , P r u n e l l a v u l g a r i s and E r i g e r o n p h i l a d e l p h i c u s . In t h i s type of e s t u a r y , t" e extent and s p e c i e s d i v e r s i t y of each p l a n t community i s p r o p o r t i o n a t e to the area of d e l t a with good freshwater c i r c u l a t i o n , i . e . , the l a r g e r the area with good freshwater c i r c u l a t i o n the more e x t e n s i v e and d i v e r s e the p l a n t communities (Salmon, Quatse). - 3 9 -23. Carex l y n g b y e i i s dominant on sand and Deschampsia c e s p i t o s a i s dominant on g r a v e l at the lowest e l e v a t i o n s of the vegetated i n t e r t i d a l zone. With i n c r e a s i n g e l e v a t i o n , Deschampsia c e s p i t o s a dominates and may be found i n a s s o c i a t i o n with, at the lower e l e v a t i o n s of i t s range, Carex l y n g b y e i and then, with i n c r e a s i n g e l e v a t i o n , P o t e n t i l l a p a c i f i c a , Glaux maritima, T r i g l o c h i n maritimum, Agropyron repens, Hordeum brachyantherum, Poa p r a t e n s i s and A g r o s t i s t e n u i s (Kokish, Nimpkish). 24a. S p i t and/or berme occurs on the d e l t a . 25 24b. No s p i t and/or berme occurs on the d e l t a . 26 25. Deschampsia c e s p i t o s a i s dominant on g r a v e l and Carex  l y n g b y e i i s dominant on sand at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone. Deschampsia c e s p i t o s a becomes dominant as the e l e v a t i o n i n c r e a s e s . Near the t r e e l i n e , Deschampsia c e s p i t o s a may be codominant with P o t e n t i l l a p a c i f i c a , or Hordeum brachyantherum, T r i f o l i u m  w o r m s k j o l d i i , Plantago l a n c e o l a t a and Juncus e f f u s u s may be dominant. On some e s t u a r i e s , S a l i c o r n i a v i r g i n i c a , Plantago maritima and T r i g l o c h i n maritimum, species i n d i c a t i v e of r e l a t i v e l y s a l i n e c o n d i t i o n s , may be found at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone. Vegetation landward of a s s o c i a t e d bermes i s l i k e that d e s c r i b e d under 10 (Quatse, Adam-Eve). 26. Deschampsia c e s p i t o s a i s dominant on g r a v e l and Carex  l y n g b y e i i s dominant on sand at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone. With i n c r e a s i n g e l e v a t i o n , Deschampsia c e s p i t o s a dominates and may be found i n a s s o c i a t i o n with Carex l y n g b y e i at the lower e l e v a t i o n s of - 40 -i t s range and with T r i f o l i u m w o r m s k j o l d i i , A c h i l l e a  m i l l e f o l i u m , T r i g l o c h i n maritimum and Holcus lanatus adjacent to the f o r e s t . P i c e a s i t c h e n s i s is^ the dominant t r e e s p e c i e s on the d e l t a ( T s i t i k a ) . 27. Deschampsia c e s p i t o s a i s dominant on g r a v e l and Carex l y n g b y e i i s dominant on sand at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone. On r e l a t i v e l y l a r g e d e l t a s , S a l i c o r n i a v i r g i n i c a may be dominant at the lower e l e v a t i o n s . Deschampsia c e s p i t o s a , Agropyron repens, Poa  p r a t e n s i s , A g r o s t i s a l b a var. s t o l o n i f e r a , D a c t y l i s  glomerata and Bromus s i t c h e n s i s are dominant at higher e l e v a t i o n s . Where the d e l t a i s l a r g e and r e l a t i v e l y g e n tly s l o p i n g shrubs (e.g., Rubus s p e c t a b i l i s ) w i l l be dominant adjacent to the P i c e a s i t c h e n s i s f o r e s t (eg. west coast Vancouver I s l a n d ) . 3.2.1 E s t u a r i e s In doing t h i s study each estuary was surveyed in turn. During the course of the work i t became evident that the r o l e of p h y s i c a l f a c t o r s i n the composition and d i s t r i b u t i o n of the p l a n t communities i n the e s t u a r i e s was important. As a r e s u l t , the p h y s i c a l f a c t o r s have been d i s c u s s e d and the e s t u a r i e s have been d i v i d e d i n t o eleven groups on the b a s i s of the i n f l u e n c e of the p h y s i c a l f a c t o r s on the composition and d i s t r i b u t i o n of the p l a n t communities among e s t u a r i e s . It i s deemed u s e f u l to present a d i s c u s s i o n of each es t u a r y to give the nature of the p l a n t communities and the p h y s i c a l f a c t o r s i n f l u e n c i n g them on each e s t u a r y . 4/ 3.2.1.1 Coldstream Estuary The pl a n t communities of the Goldstream Estuary tend to be dominated by species i n d i c a t i v e of s a l i n e c o n d i t i o n s . These species i n c l u d e s a l t w o r t , s a l t g r a s s , B a l t i c rush and meadow b a r l e y . The abundance of the s a l i n e species increases with d i s t a n c e from the main streamflow. The b r a c k i s h water species (e.g., Lyngby's sedge) are most abundant i n the main channels of the stream. The v i g o r and abundance of Lyngby's sedge decreases with i n c r e a s i n g d i s t a n c e from the main streamflow. The Cytisus  scoparius - Rosa nutkana and Plantago l a n c e o l a t a - Rumex  s a l i c i f o l i u s -Hordeum brachyantherum - Chenopodium album communities occur on l a n d f i l l i n the backshore. These two communities are almost never flooded. The p l a n t species d i s t r i b u t i o n i s r e l a t e d to the three p r i n c i p a l p h y s i c a l f a c t o r s , d e l t a morphology, flow and p r e c i p i t a t i o n . The Coldstream d e l t a i s large and g e n e r a l l y elevated above the streamflow. Much of the d e l t a appears not to be s t r o n g l y i n f l u e n c e d by freshwater, probably due to the reduced streamflow as much of i t has been d i v e r t e d f o r urban use. The dominance of species i n d i c a t i n g p h y s i o l o g i c a l l y dry c o n d i t i o n s suggests that p r e c i p i t a t i o n during the growing season i s not great enough to reduce the predominantly s a l i n e c o n d i t i o n s . As the g l a c i e r s receded, streamflow was probably much greater which would account f o r the extensive and elevated d e l t a . Larger and more sediment laden flows formed the d e l t a and many of the channels i n i t today. It i s l i k e l y that the backshore f l o o d p l a i n , now i s o l a t e d from the estuary by the l a n d f i l l , was once a continuum of the wetland and was a - 42 -p o s s i b l e source of freshwater d i s t r i b u t i o n to the western h a l f of the e s t u a r y . 3.2.1.2 Cowichan River Estuary The emergent p l a n t communities of the Cowichan River Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h and b r a c k i s h s a l i n e c o n d i t i o n s . The b r a c k i s h water s p e c i e s (e.g., Lyngby's sedge, c i n q u e f o i l and B a l t i c rush) are most abundant north of the r a i l r o a d while the b r a c k i s h s a l i n e s p e c i e s (e.g., s a l t g r a s s , B a l t i c rush and s a l t w o r t ) tend to be most abundant south of the r a i l r o a d and w i t h i n some dyked areas. The p l a n t s p e c i e s d i s t r i b u t i o n i s r e l a t e d not only to the p r i n c i p a l p h y s i c a l f a c t o r streamflow but a l s o to man's i n f l u e n c e s on i t . The d i f f u s e d i s p e r s a l of an abundant streamflow i n an e s t u a r y "'"ends to maintain b r a c k i s h water c o n d i t i o n s which appear to be more p r o d u c t i v e than s a l i n e p r o d u c t i v e than s a l i n e water c o n d i t i o n s . However, human a c t i v i t y (eg. roads, r a i l w a y s , l a n d f i l l s and dykes) has a l t e r e d freshwater c i r c u l a t i o n i n the Cowichan R i v e r E s t u a r y . The road and r a i l r o a d to the lumber storage area appear to have a major i n f l u e n c e i n o b s t r u c t i n g freshwater c i r c u l a t i o n to the southern plant communities. As a r e s u l t , communities are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h s a l i n e c o n d i t i o n s . Dykes have also impeded freshwater c i r c u l a t i o n to emergent p l a n t communities, fragmenting and s a l i n i z i n g emergent v e g e t a t i o n stands. It i s probable that any f u r t h e r dredging, l a n d f i l l i n g or dyking on the Cowichan R i v e r Estuary could decrease the abundance of b r a c k i s h water s p e c i e s and the p r o d u c t i v i t y of the - 43 -estuary (Kennedy, 1978). As the g l a c i e r s receded, the streamflow of the Cowichan River was undoubtably gr e a t e r and more broadly d i s t r i b u t e d to form a continuous wetland from the present e s t u a r y , i n l a n d . T h i s wetland i s a l s o reduced, fragmented and a l i e n a t e d by roads, r a i l w a y , l a n d f i l l and dykes. 3.2.1.3 Chemainus R i v e r Estuary The emergent p l a n t communities on the Chemainus River d e l t a form a b r a c k i s h marsh to the northwest and a s a l t marsh to the southeast. The communities to the northwest, around the mouth of the Chemainus R i v e r , are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h (e.g., Lyngby's sedge) and p h y s i o l o g i c a l l y dry (e g., B a l t i c rush and v a r i o u s grasses) c o n d i t i o n s . Communities dominated by B a l t i c rush and v a r i o u s grasses tend to occur away from the main streamflow. The communities to the southeast, near the mouth of B o n s a l l Creek, are dominated by s p e c i e s i n d i c a t i v e of s a l i n e c o n d i t i o n s . These i n c l u d e s a l t w o r t , s a l t g r a s s and gumweed. Saltwort may be found at the lowest e l e v a t i o n s of the vegetated i n t e r t i d a l zone and, as the e l e v a t i o n i n c r e a s e s , s a l t g r a s s then gumweed may appear'. Maritime b u l r u s h , a s a l i n e water i n d i c a t o r , may be found in channels, w i t h i n the dyked areas, where the s u b s t r a t e i s s a l i n e and water s a t u r a t e d . The p l a n t s p e c i e s d i s t r i b u t i o n i s r e l a t e d to two p r i n c i p a l f a c t o r s , d e l t a morphology and streamflow. The d e l t a i s very l a r g e , extending south to the C r o f t o n pulp and paper m i l l , north to Chemainus Bay and east to the.Shoal I s l a n d s . The i s l a n d s have provided p r o t e c t i o n from the t i d a l c u r r e n t s of - 44 -S t u a r t Channel thus e n a b l i n g the d e l t a to b u i l d behind and around them. As the g l a c i e r s receded, the flow of the Chemainus River must have been g r e a t e r than present i n order to b u i l d the d e l t a . Indeed, o l d channels i n the a g r i c u l t u r a l areas, some supporting stunted Lyngby's sedge, suggest that the d e l t a was once more e x t e n s i v e l y f l o o d e d . The l e s s e r flow i s now d i r e c t e d , by the landform, mainly to the northwest and north, over the b r a c k i s h marsh and away from the saltmarsh to the southeast. Dykes and roads f u r t h e r i n h i b i t freshwater c i r c u l a t i o n to the southeast where the dominant s a l i n e i n f l u e n c e has r e s u l t e d in a s a l t marsh. Human a c t i v i t y has a l s o had an i n f l u e n c e on the d i s t r i b u t i o n of e s t u a r i n e f l o r a on the Chemainus River d e l t a . Two a c t i v i t i e s appear to have the most i n f l u e n c e . The f i r s t i s a g r i c u l t u r e which has been a c t i v e l y pursued f o r at l e a s t 100 years ( B o n s a l l , p e r s . comm.) on the d e l t a . The dyking and d r a i n i n g of land f o r farming has e l i m i n a t e d most of the backshore channels and, with t h e i r l o s s , the continuum between e s t u a r i n e wetland and f o r e s t has been broken. The second human a c t i v i t y i s the water f i l t r a t i o n s t a t i o n o u t f a l l at the southeastern end of the d e l t a . P e r i o d i c a l l y , the s t a t i o n cleans i t s f i l t e r s and f l u s h e s accumulated sediments through the o u t f a l l . It would appear that the sediments are a c c e l e r a t i n g a c c r e t i o n on the southeastern end of the d e l t a . A r e s i d e n t adjacent to the o u t f a l l r e p o r t s that he has had to d i g out h i s boat channel s i n c e the o u t f a l l was e s t a b l i s h e d . A l s o , he i n d i c a t e s beds of e e l - g r a s s and algae used to occur between the inner and outer Shoal Islands (McDonald, pers; comm.). T h i s area i s now an i n t e r t i d a l mudflat suggesting sediment - 45 -accumulation, probably from the f i l t r a t i o n s t a t i o n , the most immediate source. L a n d f i l l i n g between the C r o f t o n pulp and paper m i l l and outer Shoal I s l a n d s may f u r t h e r accentuate sedimentation and e l i m i n a t e s u b t i d a l f l o r a . 3.2.1.4 Nanaimo R i v e r Estuary The emergent p l a n t communities of the Nanaimo River Estuary tend to be dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h , p h y s i o l o g i c a l l y dry, and s a l i n e c o n d i t i o n s . I n i t i a l l y , there may not appear to be a p a t t e r n to the s p e c i e s d i s t r i b u t i o n though on c l o s e r examination of the data one i s suggested. The b r a c k i s h s p e c i e s (e.g. Lyngby's sedge and t u f t e d h a i r g r a s s ) appear to be more abundant and vigorous along main channels. The abundance and v i g o r of Lyngby's sedge i s an example. In the western channel, through which most of the Nanaimo River flows, Lyngby's sedge i s abundant and v i g o r o u s . In the main channels on the e a s t e r n h a l f of the estuary, Lyngby's sedge i s l e s s abundant, l i n i n g only the immediate edges of the channels, and i t i s l e s s vigorous being g e n e r a l l y h a l f as t a l l as the sedge i n the main, western channel. The s p e c i e s i n d i c a t i v e of p h y s i o l o g i c a l l y dry c o n d i t i o n s (e.g., B a l t i c rush, j o i n t e d rush and v a r i o u s grasses) most f r e q u e n t l y occur away from the main streamflow, toward the backshore and wit' i n dyked areas. J o i n t e d rush i s the most abundant dominant or subdominant on the e s t u a r y , c o v e r i n g 16.4 hect a r e s or 29 percent of the vegetated area s t u d i e d . It i s not known to occur as e x t e n s i v e l y or abundantly on any other estuary on Vancouver I s l a n d . It i s p o s s i b l e that j o i n t e d rush may have a p h y s i o l o g i c a l advantage - 46 -o v e r , f o r example, B a l t i c r u s h , when the g r o w i n g c o n d i t i o n s i n c l u d e a w a t e r s a t u r a t e d s u b s t r a t e and a p r o b a b l y i n c r e a s i n g s o i l s a l i n i t y d u r i n g t h e g r o w i n g s e a s o n . T h i s m i g h t e x p l a i n the u n i q u e d i s t r i b u t i o n p a t t e r n o f j o i n t e d r u s h on t h e Nanaimo R i v e r E s t u a r y . The s a l i n e i n d i c a t o r s ( e . g . , s a l t w o r t , s a l t g r a s s , s a l t b u s h and a r r o w - g r a s s ) t e n d t o become more a b u n d a n t away from the main s t r e a m f l o w and a t t h e l o w e r e l e v a t i o n s o f t h e v e g e t a t e d a r e a s t u d i e d . A r r o w - g r a s s o c c u r s most f r e q u e n t l y a l o n g t h e seaward edge o f t h e w e s t e r n i s l a n d where i t i s n o t s t r o n g l y i n f l u e n c e d by f r e s h w a t e r . S a l t w o r t becomes more abundant a t the l o w e r e l e v a t i o n s on t h e e a s t e r n h a l f o f t h e d e l t a where abundant s a l t w a t e r n e g a t e s f r e s h w a t e r i n f l u e n c e . The p l a n t s p e c i e s d i s t r i b u t i o n i s a g a i n r e l a t e d t o two p r i n c i p a l p h y s i c a l f a c t o r s , d e l t a m o r p h o l o g y and s t r e a m f l o w , and man's i n f l u e n c e on them. The e x t e n s i v e d e l t a i s bounded t o the s o u t h and e a s t by r o c k f o r m a t i o n s o f t h e Nanaimo Group ( M u l l e r , 1971) and t o t h e west and s o u t h w e s t by e l e v a t e d outwash d e p o s i t s o v e r b e d r o c k . The s i z e and e l e v a t i o n o f t h e d e l t a s u g g e s t t h a t i t was formed when t h e f l o w and s e d i m e n t l o a d o f t h e Nanaimo R i v e r were g r e a t e r , i . e . , i n p r o g l a c i a l t i m e s . Nowadays the f l o w i n t o t h e e s t u a r y i s r e g u l a t e d by u p s t r e a m dams and i t s d i s p e r s a l p a t t e r n a c r o s s t h e e s t u a r y i s i n f l u e n c e d by human a c t i v i t y ( i . e . , d y k i n g and d r e d g i n g ) . The dyke, a l s o s e r v i n g as a r o a d a l o n g t h e e a s t bank o f t h e Nanaimo R i v e r , i n h i b i t s f r e s h w a t e r f l o w , t h r o u g h c h a n n e l s v i s i b l e o v e r th e a g r i c u l t u r a l l a n d , t o t h e e a s t e r n h a l f o f t h e d e l t a . O t h e r d y k e s , g e n e r a l l y b r e a c h e d and o r i e n t e d i n v a r i o u s d i r e c t i o n s , would a p p e a r t o i m p o u n d w a t e r i n a r e a s . A number o f t h e s e a r e a s a r e d o m i n a t e d o r s u b d o m i n a t e d by j o i n t e d r u s h . P o s s i b l y t h e abundance o f - 47 -jo- in +ed rush i s r e l a t e d to dyking a c t i v i t y on the e s t u a r y . The d i s p e r s a l of freshwater across the e a s t e r n h a l f of the d e l t a i s f u r t h e r i n f l u e n c e d by dredging of the western channel of the r i v e r . The main streamflow i s drawn through the western channel where i t c r e a t e s b r a c k i s h c o n d i t i o n s s u i t a b l e f o r the growth of Lyngby's sedge on the western p o r t i o n of the d e l t a . At the b i f u r c a t i o n of the Nanaimo R i v e r , an a c c r e t i n g g r a v e l bar i n the e a s t e r n channel a s s i s t s , through d e f l e c t i o n , the d i r e c t i o n of flow to the western channel. T h i s f u r t h e r reduces freshwater d i s t r i b u t i o n to the e a s t e r n h a l f of the e s t u a r y . The g r a v e l bar has been known to r e p e a t e d l y i n c r e a s e i n s i z e and then wash out ( p e r s . comm. B. Smith). Three small channels occur on the i s l a n d between the east and west channels. At one time the channels on the i s l a n d may ' ave d i s t r i b u t e d freshwater to the i s l a n d ' s v e g e t a t i o n c r e a t i n g more b r a c k i s h , r a t h e r than the e x i s ^ ^ n g s a l i n e c o n d i t i o n s . P r e s e n t l y the channels are p a r t i a l l y blocked at the freshwater source and thus have a l i m i t e d i n f l u e n c e , however they do provide a p o t e n t i a l f o r s t u d i e s on the i n f l u e n c e of freshwater c i r c u l a t i o n on the d i s t r i b u t i o n of emergent p l a n t s p e c i e s . The continuum from s a l i n e to freshwater wetlands was probably very e v i d e n t before developments such as roads, dykes and l a n d f i l l a l t e r e d much of the backshore and f l o o d p l a i n . These developments appear to have broken the continuum and d i s r u p t e d the wetland food webs. In the past, the Nanaimo and Cowichan R i v e r V a l l e y s had the most e x t e n s i v e continuous wetland h a b i t a t s on the east coast of Vancouver I s l a n d . The remaining, " d i s j o i n t e d " e s t u a r i n e l a n d , back channels, meandering creeks and ponds are both remnants and r e l i c s of that wetland h a b i t a t . - 48 -The l o s s of the continum i s p a r t i a l l y r e f l e c t e d i n d e c l i n i n g numbers of i n d i v i d u a l s (e.g., s h e l l f i s h , salmon and waterfowl) at higher t r o p h i c l e v e l s of the food web. 3.2.1.5 Nanoose-Bonell Creeks' Estuary The emergent p l a n t communities of the Nanoose-Bonell Creeks' Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h s a l i n e and s a l i n e c o n d i t i o n s . The 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 of b r a c k i s h s a l i n e c o n d i t i o n s (e.g., Lyngby's sedge, s a l t w o r t , s a l t g r a s s and s a l t b u s h ) occur i n the main streamflow of Nanoose Creek. Lyngby's sedge, g e n e r a l l y considered i n d i c a t i v e of b r a c k i s h c o n d i t i o n s , has low v i g o r . T h i s low v i g o r may be a t t r i b u t e d to the higher s a l i n i t i e s suggested by i t s a s s o c i a t e d s p e c i e s . The s a l i n e s p e c i e s (e.g., s a l t w o r t , s a l t g r a s s , s a l t b u s h , s e a s i d e p l a n t a i n and s e a b l i t e ) occur throughout the lower e l e v a t i o n s of the vegetated study area where they are mainly i n f l u e n c e d by seawater. Toward the backshore, where in u n d a t i o n i s l e s s frequent and of short d u r a t i o n , the dominant species become more mesic. These s p e c i e s i n c l u d e Nootka rose, evergreen b l a c k b e r r y , thimbleberry, salmonberry, Canada t h i s t l e , v e l v e t - g r a s s , red a l d e r , ninebark and reed canarygrass. The p l a n t s p e c i e s d i s t r i b u t i o n i s r e l a t e d to the p r i n c i p a l p h y s i c a l f a c t o r , streamflow, and to human a c t i v i t y . The streamflows of Nanoose and B o n e l l Creeks are very small (Table 5) and B o n e l l Creek i s known to dry up i n the summer. Th i s small combined streamflow appears to provide l i t t l e freshwater i n f l u e n c e i n the e s t u a r y , r e s u l t i n g i n the predominance of s a l i n e h a b i t a t as i n d i c a t e d by the dominance of - 49 -s a l i n e i n d i c a t o r p l a n t s . Human a c t i v i t y (dyking and a g r i c u l t u r e ) appears to have a l t e r e d drainage p a t t e r n s and d i s p l a c e d n a t u r a l backshore p l a n t communities. P l a n t communities south of B o n e l l Creek d r a i n very slowly at low f d e . This drainage impediment appears to be a r e s u l t of dyking and probably r e s u l t s i n an i n c r e a s e i n the r e l a t i v e s a l i n i t y of the a f f e c t e d p l a n t communities. The backshore i s mainly used f o r hay crops and P h a l a r i s arundinacea dominates one backshore f i e l d that has not been c u l t i v a t e d r e c e n t l y . Forage crops d i s p l a c e the more d i v e r s e , n a t u r a l p l a n t communities prominent along the streambanks of Nanoose and B o n e l l Creeks. 3.3.1.6 Englishman R i v e r Estuary The emergent p l a n t communities of the Englishman River Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h , s a l i n e and p h y s i o l o g i c a l l y dry c o n d i t i o n s . The b r a c k i s h i n d i c a t o r species (e.g. Lyngby's sedge) occur at the lower e l e v a t i o n s of the vegetated study areas, adjacent to freshwater. The s a l i n e i n d i c a t o r s p e c i e s (e.g., s a l t w o r t and s a l t g r a s s ) a l s o tend to occur at the lower e l e v a t i o n s of the vegetated study area but away from the i n f l u e n c e of freshwater. The b r a c k i s h and s a l i n e growing c o n d i t i o n s o v e r l a p where the D i s t i c h l i s s p i c a t a - Carex  lyngbyei Community o c c u r s . In these areas the s a l i n i t y probably i n c r e a s e s through the growing season i n r e l a t i o n to d e c l i n i n g streamflow and p r e c i p i t a t i o n , and i n c r e a s i n g e v a p o t r a n s p i r a t i o n . In t h i s community, Lyngby's sedge i s able to i n i t i a t e growth and develop at the begining of the season when the s a l i n i t i e s are lowest. I t s reduced v i g o r suggests i t i s s t r e s s e d by the - 50 -i n c r e a s i n g s a l i n i t y . S a l t g r a s s i n i t i a t e s growth a f t e r Lyngby's sedge and appears able to develop to average v i g o r as the s a l i n i t i e s i n c r e a s e i n the community. Species i n d i c a t i v e of p h y s i o l o g i c a l l y dry c o n d i t i o n s (e.g., s a l t g r a s s , B a l t i c rush, gumweed and v a r i o u s grasses) occur toward the backshore where they are probably i n f r e q u e n t " " and s h a l l o w l y i n ndated by b r a c k i s h s a l i n e water. The more mesic p l a n t communitites i n the study area occur along the main channel of the r i v e r , i n the backshore and between the e a s t e r n dyke and r e s i d e n t i a l area. The main channel communities are dominated by red a l d e r b r o o m , evergreen b l a c k b e r r y and Nootka r o s e . The red a l d e r community i s only i n f l u e n c e d by freshwater i n the manner of most g r a v e l bar communities. The broom, evergreen b l a c k b e r r y and Nootka rose community i s growing on l o g d e b r i s which has probably elevated the s u b s t r a t e , r e s u l t i n g i n improved drainage and reduced i n u n d a t i o n . The Sonchus a r v e n s i s - C i r s i u m arvense and Holcus  lanatus - E p i l o b i u m a n g u s t i f o l i u m communities i n the backshore and w i t h i n t'"e dyked area are probably a r e s u l t of a dyke and l a n d f i l l . These appear to have e l i m i n a t e d t i d a l i n u n d a t i o n of the areas and have f a c i l i t a t e d t h e i r c o l o n i z a t i o n by mesic s p e c i e s . The adjacent land, o u t s i d e the dyke and u n a f f e c t e d by l a n d f i l l , i s c o l o n i z e d by emergent v e g e t a t i o n which suggests the dyked and l a n d f i l l e d areas might have supported emergent p l a n t s p e c i e s b e f o r e being a l t e r e d . The p l a n t s p e c i e s d i s t r i b u t i o n appears to be mainly i n f l u e n c e d by two f a c t o r s . The f i r s t i s streamflow and the second i s e x t e n s i v e human a c t i v i t y . The streamflow of the Englishman R i v e r today does not seem to be s u f f i c i e n t to - 51 -maintain b r a c k i s h c o n d i t i o n s throughout the e s t u a r y . T h i s i s i n d i c a t e d by the prominence of s a l i n e s p e c i e s with i n c r e a s i n g d i s t a n c e from the main flow. The d i s t r i b u t i o n of the flow i s i n f l u e n c e d by the second f a c t o r , human a c t i v i t y . Human a c t i v i t y i n the form of dykes, l a n d f i l l , dredge s p o i l d e p o s i t s , dredging, lo g h a n d l i n g and . r e s i d e n t i a l development have a l t e r e d freshwater c i r c u l a t i o n i n the e s t u a r y . Dykes, l a n d f i l l and dredge s p o i l d e p o s i t s tend to i n h a b i t freshwater d i s t r i b u t i o n and dredging c h a n n e l i z e s flow. The a c t i v i t i e s have stopped with the end of lo g h a n d l i n g i n the est u a r y and the p r o t e c t i o n of the r e s i d e n t i a l development; however, t h e i r e f f e c t s are s t i l l e v i d e n t . Evidence f o r r e s t r i c t e d freshwater c i r c u l a t i o n i s found i n the mesic p l a n t communities i n the dyked area and on the l a n d f i l l , and i n the i n c r e a s i n g frequency of s a l i n e p l a n t s as d i s t a n c e from the main streamflow i n c r e a s e s . The Englishman R i v e r d e l t a encompasses more area than was a c t u a l l y s t u d i e d and was probably formed as the g l a c i e r s receded when the r i v e r ' s streamflow and sediment load were much g r e a t e r . The study area i s only a small p o r t i o n of t h i s d e l t a ; however, i t appears that the marsh was l a r g e r when flows were p r o p o r t i o n a t e l y g r e a t e r . T h i s i s suggested by the presence of emergent p l a n t s p e c i e s i n some of the channels on the western d e l t a . These channels may have c o n s t i t u t e d the continuum between e s t u a r i n e and backshore wetlands before the western dyke was b u i l t . Now t h i s continuum i s d i s r u p t e d by the dyke and the v e g e t a t i o n on the western d e l t a appears to be e n t e r i n g mesic s e r a i stages. The p o s s i b l e r o l e of such backshore h a b i t a t w i t h i n a s p i t and dune d e l t a formation w511 be di s c u s s e d under the Cluxewe R i v e r E s t u r a r y . - 52 -3.2.1.7 L i t t l e Qualicum R i v e r Estuary The emergent p l a n t communities of the L i t t l e Qualicum River Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h and s a l i n e b r a c k i s h water c o n d i t i o n s . The s p e c i e s i n d i c a t i n g b r a c k i s h c o n d i t i o n s i n c l u d e Lyngby's sedge, c i n q u e f o i l and B a l t i c rush. Lyngby's sedge occurs at the lowest e l e v a t i o n s of the vegetated study area where i t i s inundated on each f l o o d t i d e . With i n c r e a s i n g e l e v a t i o n and b e t t e r s u b s t r a t e drainage the sedge i s a s s o c i a t e d with c i n q u e f o i l . In these areas i t i s l e s s v i g o r o u s than at lower e l e v a t i o n s . With i n c r e a s i n g e l e v a t i o n and r e l a t i v e l y b e t t e r s u b s t r a t e drainage, c i n q u e f o i l occurs with B a l t i c rush. At e l e v a t i o n s where f l o o d i n g i s probably i n f r e q u e n t and of s h o r t d u r a t i o n , t e r r e s t r i a l s p e c i e s occur with i n c r e a s i n g frequency. In these areas Nootka. rose i s the most abundant s p e c i e s . Nootka rose i s a l s o dominant on r e l a t i v e l y more e l e v a t e d land w i t h i n the dyked areas. At lower e l e v a t i o n s i n the dyked areas grasses are abundant and r e f l e c t e a r l i e r a g r i c u l t u r a l a c t i v i t i e s . S a l t g r a s s , the s p e c i e s i n d i c a t i n g s a l i n e b r a c k i s h c o n d i t i o n s , tends to be most abundant on the l e f t bank of the main channel. The p l a n t s p e c i e s d i s t r i b u t i o n i s r e l a t e d to the two p h y s i c a l f a c t o r s , e l e v a t i o n and s a l i n i t y . .The f i r s t , e l e v a t i o n , i n f l u e n c e s the i n u n d a t i o n : emergence r a t i o and s u b s t r a t e d r a i n a g e . The i n u n d a t i o n : emergence r a t i o i s g r e a t e r at lower e l e v a t i o n s and lowest at higher e l e v a t i o n s . Species v i a b l e a f t e r longer p e r i o d s of submergence (e.g., Lyngby's sedge) occur at the lower e l e v a t i o n s i n the vegetated study area while s p e c i e s i n c a p a b l e of s u r v i v i n g long p e r i o d s of i n u n d a t i o n (e.g., - 5 3 -Nootka rose) occur at the higher e l e v a t i o n s . It appears that there are more s p e c i e s able to withstand s h o r t e r p e r i o d s of submergence than there are s p e c i e s able to withstand longer-p e r i o d s of submergence. T h i s i s suggested by the i n c r e a s e i n number of s p e c i e s with i n c r e a s i n g e l e v a t i o n . The e l e v a t i o n a l s o i n f l u e n c e s s u b s t r a t e drainage i n t h a t , with i n c r e a s i n g e l e v a t i o n the s u b s t r a t e i s r e l a t i v e l y b e t t e r d r a i n e d at ebb t i d e . T h i s allows oxygenation of the p l a n t r o o t s i n the s u r f a c e s u b s t r a t e . Few p l a n t s p e c i e s are able to s u r v i v e when t h e i r r o o t s are i n the anaerobic environment of a water s a t u r a t e d s o i l . Lyngby's sedge i s one s p e c i e s that i s appa r e n t l y adapted and i t i s most abundant i n the water s a t u r a t e d s u b s t r a t e at the lowest e l e v a t i o n s i n the vegetated study area. The second p h y s i c a l f a c t o r a f f e c t i n g the p l a n t species d i s t r i b u t i o n , s a l i n i t y , appears to be most i n f l u e n t i a l on the l e f t bank of the main channel. There the dyke i n h i b i t s f l u s h i n g of the area by freshwater. Thus s a l t s brought to the s u b s t r a t e s u r f a c e by ev a p o r a t i o n are concentrated i n the s o i l ; s a l i n e growing c o n d i t i o n s r e s u l t , and s a l t g r a s s i s the dominant s p e c i e s . Through dyking, human a c t i v i t y has i n f l u e n c e d the d i s t r i b u t i o n of s a l t g r a s s . Other human a c t i v i t i e s have a f f e c t e d p l a n t s p e c i e s d i s t r i b u t i o n such as dredging, dredge s p o i l d e p o s i t i o n , l o g handl i n g and a g r i c u l t u r e . Dredging the main r i v e r channel has probable r e s u l t e d i n some c h a n n e l i z a t i o n and a l t e r a t i o n i n freshwater c i r c u l a t i o n . Dredge s p o i l placed on the dykes has accentuated t h i s c h a n n e l i z a t i o n . A l s o , dredge s p o i l d e p o s i t i o n behind the r i g h t bank dyke has a l t e r e d the in u n d a t i o n : emergence r a t i o of the p l a n t communities by i n c r e a s i n g the e l e v a t i o n . For example, the P o t e n t i l l a p a c i f i c a - 54 -- Carex l y n g b y e i Community, between two more e l e v a t e d areas c o l o n i z e d by the P o t e n t i l l a p a c i f i c a - Juncus b a l t i c u s Community and Nootka rose, dominates the southeast end of the dyke where r e l a t i v e l y more s p o i l has been put. I t i s p o s s i b l e that the dredging and dredge s p o i l d e p o s i t i o n a c t i v i t i e s may have been a s s o c i a t e d with main^a^ n1' ng " annel f o r log handling i n the estuary. Log h a n d l i n g no longer occurs i n the e s t u a r y . The e s t u a r y i s no longer "sed f o r a g r i c u l t u r e though the o l d farm f i e l d s , b u i l d i n g s and dykes are s t i l l e v i d e n t . It i s p o s s i b l e t h a t the f i e l d s may have been freshwater wetland at one time and thus formed a continuum between e s t u a r i n e wetland and upland v e g e t a t i o n . T h i s i s suggested by the channels i n the f i e l d s and p a r t i c u l a r l y the c o l o n i z a t i o n of a channel by b u l r u s h , a freshwater i n d i c a t o r s p e c i e s . T h i s i s a l s o i m p l i e d i n d e s c r i p t i o n s of d e l t a morphology by Coates (1972). The predominantly b r a c k i s h water growing c o n d i t i o n s on most of the e s t u a r y may seem incongrous with the preceding d i s c u s s i o n of a l t e r a t i o n s i n freshwater c i r c u l a t i o n due to dredging and dyking of the main channel. The maintenance of the b r a c k i s h water growing c o n d i t i o n s may be e x p l a i n e d by the occurrence of u p w e l l i n g ground water and l o c a l i z e d water c i r c u l a t i o n p a t t e r n s . The presence of water i n the channels i n the f i e l d s may support t h i s view. 3.2.1.8 Big Qualicum R i v e r Estuary The emergent p l a n t communities of the Big Qualicum R i v e r Estuary tend to be dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h water and - 55 -b r a c k i s h freshwater growing c o n d i t i o n s . The b r a c k i s h water i n d i c a t o r s p e c i e s , Lyngby's sedge, i s found at the lower e l e v a t i o n s , on f i n e r s u b s t r a t e s , where i t i s r e g u l a r l y inundated by •'"•'des. The two i n t r o d u c e d s p e c i e s i n d i c a t i v e of f a i r drainage and n o n - s a l i n e c o n d i t i o n s , red c l o v e r and r i b g r a s s , are found at r e l a t i v e l y higher e l e v a t i o n s on c o a r s e r s u b s t r a t e s where they are i n f r e q u e n t l y inundated. The p l a n t s p e c i e s d i s t r i b u t i o n i n t h i s e s t u a r y i s c l o s e l y r e l a t e d to two p r i n c i p a l p h y s i c a l f a c t o r s : e l e v a t i o n and s o i l p a r t i c l e s i z e . The f i r s t p h y s i c a l f a c t o r , e l e v a t i o n , i n f l u e n c e s the i n u n d a t i o n : emergence r a t i o of the s p e c i e s . The lower the e l e v a t i o n the h i g h e r the r a t i o and v i c e v e r s a . Lyngby's sedge appears to have the h i g h e s t i n u n d a t i o n : emergence r a t i o as i t occurs at the lowest e l e v a t i o n s of the vegetated study area, and red c l o v e r and r i b g r a s s appear to have the lowest i n u n d a t i o n : emergence r a t i o as they occur at a r e l a t i v e l y higher e l e v a t i o n i n the vegetated study area. The second p h y s i c a l f a c t o r , s o i l p a r t i c l e s i z e , may i n f l u e n c e s e v e r a l t h i n g s ; the s i z e of a species r o o t system, f e r t i l i t y and the degree of s u b s t r a t e d r a i n a g e . The root systems of the dominant p l a n t s p e c i e s on the Big Qualicum R i v e r Estuary seem to be e i t h e r r e l a t i v e l y l a r g e or s m a l l . The l a r g e r rhizomatous root systems seem to e x i s t i n f i n e r s u b s t r a t e s but seem to have d i f f i c u l t y i n g r a v e l s . T h i s seems to l i m i t d i s t r i b u t i o n of Lyngby's sedge, a s p e c i e s normally with w e l l developed rhizomes, on the widely d i s t r i b u t e d g r a v e l s u b s t r a t e s of the e s t u a r y . The dominant, s h o r t - l i v e d species with determinant semi-tap r o o t s (red c l o v e r (a n i t r o g e n - 56 -f i x e r ) and r i b g r a s s ) may c o l o n i z e g r a v e l with l i t t l e d i f f i c u l t y but are l i m i t e d to r e l a t i v e l y higher e l e v a t i o n s by t h e i r i n a b i l i t y to withstand i n u n d a t i o n . The r a p i d i t y of drainage i s a l s o r e l a t e d to the p a r t i c l e s i z e of the s u b s t r a t e . The l a r g e r the p a r t i c l e s i z e , and the higher the e l e v a t i o n , the b e t t e r the drainage and the d r i e r the grow'ng c o n d i t i o n s . Thus more mesic p l a n t s p e c i e s , such as sweet v e r n a l g r a s s , tend to c o l o n i z e the g r a v e l s at higher e l e v a t i o n s of the study area. Human a c t i v i t y has g r e a t l y i n f l u e n c e d the p l a n t s p e c i e s d i s t r i b u t i o n on the e s t u a r y . In 1976, one year a f t e r the survey, a c a t e r p i l l a r t r a c t o r was used to scour the r i v e r bed downstream of the b r i d g e and the v e g e t a t i o n on the r i v e r g r a v e l bars was removed i n the process (D. Morrison, p e r s . comm.). The es t a b l i s h m e n t of the campgrounds on both r i v e r banks, or u n s p e c i f i e d a c t i v i t y , r e q u i r e d some l a n d f u l l and/or s u b s t r a t e movement to f i l l o l d channels. No evidence of the o l d channels was observed. -However, that channels e x i s t e d i n the past i s i m p l i e d i n d e s c r i p t i o n s of d e l t a morphology by Coates (1972), and by comparing the Big Qualicum R i v e r d e l t a with s i m i l a r d e l t a s . These channels were vegetated with emergent plant species and represented the wetland h a b i t a t on the d e l t a . See the d i s c u s s i o n of the Oyster R i v e r Estuary f o r a comparable d e s c r i p t i o n of p o s s i b l e d e l t a s t r u c t u r e . 3.2.1.9 Courtenay R i v e r Estuary The emergent p l a n t communities of the Courtenay River Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h s a l i n e - 57 -and b r a c k i s h water c o n d i t i o n s . The b r a c k i s h s a l i n e i n d i c a t o r s p e c i e s (e.g., three-square b u l r u s h ) occurs on the southern side of Comox Harbour and the b r a c k i s h i n d i c a t o r s p e c i e s (e.g., Lyngby's sedge, c i n q u e f o i l and t u f t e d h a i r g r a s s ) occur along the main channel of the Courtenay R i v e r . T h i s d i s t r i b u t i o n of dominant p l a n t s p e c i e s appears to be mainly r e l a t e d to the d-5 s n e r s a l of the main streamflow i n the study area. Three-square b u l r u s h i s a b u n d a n t away from the main streamflow, and the i n f l u e n c e of freshwater, while the b r a c k i s h i n d i c a t o r s p e c i e s , p a r t i c u l a r l y Lyngby's sedge, are most abundant adjacent to the main streamflow, where the i n f l u e n c e of freshwater i s g r e a t e s t . The r e l a t i v e l y r e s t r i c t e d extent of three-square b u l r u s h w i t h i n a l a r g e r area of s u i t a b l e s a l i n i t y range i s r e l a t e d to the e l e v a t i o n of the d e l t a . The e l e v a t i o n i n f l u e n c e s s u b s t r a t e drainage which i s g e n e r a l l y i n s u f f i c i e n t f o r f ' r e e - s q u a r e b u l r u s h over a l l but the most e l e v a t e d areas w i t h i n i t s s a l i n i t y range. The d i s t r i b u t i o n of the dominant p l a n t s p e c i e s has a l s o been i n f l u e n c e d by human a c t i v i t y i n v o l v i n g dredging, dyking, marinas, dredge s p o i l d e p o s i t i o n , l a n d f i l l i n g , road c o n s t r u c t i o n , sewage treatment, an a i r s t r i p , a sawmill, a g r i c u l t u r e and a s s o r t e d urban developments. Dredging and dyking the main channel of the Courtenay R i v e r has accentuated c h a n n e l i z a t i o n and impeded freshwater d i s p e r s a l across the es t u a r y . T h i s i s suggested by the abundance of b r a c k i s h water i n d i c a t o r s p e c i e s around the main channel and t h e i r d e c r e a s i n g frequency of occurrence with d i s t a n c e from the channel. Dredging may a l s o i n f l u e n c e s p e c i e s d i s t r i b u t i o n by d e s t r o y i n g p l a n t s d u r i n g marina c o n s t r u c t i o n . Dredge s p o i l d e p o s i t i o n and - 5 8 -l a n d f i l l i n g may a l t e r s p e c i e s d i s t r i b u t i o n by i n c r e a s i n g the e l e v a t i o n to a height where the a f f e c t e d areas may be c o l o n i z e d by more t e r r e s t r i a l p l a n t s p e c i e s (e.g. m i l k - t h i s t l e , morning-g l o r y , willow dock and reed canarygrass) which d i s p l a c e emergent plant s p e c i e s . Emergent p l a n t s p e c i e s have a l s o been d i s p l a c e d on the d e l t a by the c o n s t r u c t i o n of roads, the sewage treatment p l a n t , the a i r s t r i p , a sawmill, and a s s o r t e d urban developments, and by a g r i c u l t u r a l c r o p s . The displacement of emergent v e g e t a t i o n i s suggested by the occurrence of emergent p l a n t s i n adjacent, u n a f f e c t e d areas. In the past, when the d e l t a was not h e a v i l y developed, the emergent p l a n t communities were probably very e x t e n s i v e . This i s suggested by two t h i n g s ; f i r s t , emergent v e g e t a t i o n i n the o l d channels i n what i s now a g r i c u l t u r a l land, and second, the l a r g e s i z e and e l e v a t i o n of the d e l t a . 3.2.1.10 Oyster R i v e r Estuary The emergent p l a n t communities of the Oyster River Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h water c o n d i t i o n s . These s p e c i e s i n c l u d e Lyngby's sedge, B a l t i c rush and c i n q u e f o i l . Lyngby's sedge i s g e n e r a l l y found at the lowest e l e v a t i o n s and B a l t i c rush at r e l a t i v e l y higher e l e v a t i o n s i n the vegetated study area. C i n q u e f o i l may be a s s o c i a t e d with e i t h e r . Lyngby's sedge may a l s o occur with w i l d r y e , near the f o r e s t and road, and B a l t i c rush may a l s o occur with s a l t g r a s s and bentgrass, near the road. - 59 -The d i s t r i b u t i o n of the emergent p l a n t s p e c i e s i s p r i n c i p a l l y r e l a t e d to two f a c t o r s , d e l t a morphology and human a c t i v i t y . The f i r s t , d e l t a morphology or development, i s r e l a t e d to the flow and sediment load of the Oyster R i v e r . The d e l t a i c d e p o s i t s extend from the U n i v e r s i t y of B r i t i s h Columbia Research Farm i n the northwest to the r e s i d e n t i a l developments i n the southwest which suggests the flow and ediment load of the Oyster R i v e r were g r e a t e r as the g l a c i e r s receded. The r i v e r ' s present flow and sediment load seem to have a r e l a t i v e l y l i m i t e d i n f l u e n c e on d e l t a development. In a d d i t i o n , i t appears that the vegetated study area i s a channel that was formed when the r i v e r flow was g r e a t e r . S i m i l a r channels occur i n the r e s e a r c h farm on the o p p o s i t e bank of the r i v e r . It i s proposed that these channels are remnants of a more e x t e n s i v e e s t u a r i n e wetland. T h i s e s t u a r i n e wetland probably s t a r r e d backshore of a beach berme or dune which p r o t e c t e d i t from the heavy wind- and wave a c t i o n i t would otherwise be exposed to along the open c o a s t l i n e . The wetland appears to have run p a r a l l e l with the s h o r e l i n e and had convoluted channels. The convoluted channels wove emergent and t e r r e s t r i a l v e g e t a t i o n and f o r e s t stands i n t o a network with two d i s t i n c t r e s u l t s . F i r s t , more s u r f a c e area was a v a i l a b l e f o r c o l o n i z a t i o n by emergent p l a n t s . And second, the weaving of emergent v e g e t a t i o n stands between more t e r r e s t r i a l v e g e t a t i o n and f o r e s t stands t e r r e s t r i a l v e g e t a t i o n and f o r e s t stands provided h a b i t a t f o r a l l t r o p h i c l e v e l s . The network may a l s o have f a c i l i t a t e d a l e s s p e r c e p t i b l e i n f l u e n c e on net primary p r o d u c t i v i t y . The convoluted channels enhanced net primary p r o d u c t i v i t y by making more energy sources and - 60 -energy r e c y c l i n g o p p o r t u n i t i e s a v a i l a b l e and by p r o v i d i n g a l a r g e s u r f a c e area f o r primary p r o d u c t i o n . The second p r i n c i p a l f a c t o r c o n t r o l l i n g the d i s t r i b u t i o n of emergent p l a n t s p e c i e s , human a c t i v i t y , a l s o appears to be r e l a t e d to d e l t a morphology i n so much as the wetland backshore of a beach berme or dune i s r e l a t i v e l y easy to develop. On the south bank of the Oyster River, human a c t i v i t y has e l i m i n a t e d channels by l a n d f i l l i n g and dredging to c r e a t e a r e c r e a t i o n a l r e s o r t (and p o s s i b l y other unknown a c t i v i t i e s ) . The p l a n t communities d e s c r i b e d occur i n the remaining channel which d r a i n s i n t o the marina through a c u l v e r t . The c u l v e r t and surrounding r e s o r t , r e s i d e n c e s and road r e s t r i c t wetland use by higher t r o p h i c l e v e l s . 3.2.1.11 Campbell R i v e r Estuary The emergent p l a n t communities of the Campbell River Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h water c o n d i t i o n s . These s p e c i e s i n c l u d e Lyngby's sedge, c i n q u e f o i l , common s p i k e - r u s h and t u f t e d h a i r g r a s s . The dominant s p e c i e s appear to be d i s t r i b u t e d r e l a t i v e to e l e v a t i o n ; Lyngby's sedge tends to be dominant at the lower e l e v a t i o n s , and with i n c r e a s i n g e l e v a t i o n s , c i n q u e f o i l and common sp i k e - r u s h , and t u f t e d h a i r g r a s s become dominant s p e c i e s . Sweet g a l e , hardstem b u l r u s h , red a l d e r , evergreen b l a c k b e r r y , broom, Nootka rose and salmonberry may be found adjacent to the f o r e s t . However, sweet gale and hardstem b u l r u s h occur i n freshwater areas where the s o i l i s p o o r l y d r a i n e d and water s a t u r a t e d , p o s s i b l y due to p e r i o d i c f l o o d i n g or the upwelling of ground water. The remaining s p e c i e s occur where the s o i l i s b e t t e r d r a i n e d . - 61 -The p l a n t s p e c i e s d i s t r i b u t i o n i s p r i m a r i l y r e l a t e d to two f a c t o r s , e l e v a t i o n and human a c t i v i t y . E l e v a t i o n i n f l u e n c e s the i n u n d a t i o n : emergence r a t i o and s u b s t r a t e drainage. The i n u n d a t i o n : emergence r a t i o i s g r e a t e s t at lower e l e v a t i o n s and lowest at hi g h e r e l e v a t i o n s . Species v i a b l e a f t e r long p e r i o d s of submergence (e.g., Lyngby's sedge) occur at the lower e l e v a t i o n s i n the vegetated study a r e a . Substrate drainage i s a l s o i n f l u e n c e d by e l e v a t i o n i n t h a t , with i n c r e a s i n g e l e v a t i o n the s u b s t r a t e i s r e l a t i v e l y b e t t e r d r a i n e d at ebb t i d e . T u f t e d h a i r g r a s s , c i n q u e f o i l and common s p i k e - r u s h are most abundant on the b e t t e r d r a i n e d s u b s t r a t e s at r e l a t i v e l y higher e l e v a t i o n s . The l i m i t e d d i s t r i b u t i o n of the emergent p l a n t s p e c i e s on the d e l t a of the Campbell R i v e r i s r e l a t e d to the second f a c t o r , human a c t i v i t y . Human a c t i v i t y tends to have d i s p l a c e d emergent p l a n t s p e c i e s i n almost a l l areas of the estu a r y , except on Indian Reserve No. 11. It appears that the emergent p l a n t stands are more abundant on the Indian Reserve because the a c t i v i t i e s , d redging, l a n d f i l l i n g , l o g ha n d l i n g , lumbering and roadways, are not as p r e v a l e n t t h e r e . These a c t i v i t i e s have tended to e l i m i n a t e or reduce the v i g o r of emergent p l a n t s i n a f f e c t e d a r e a s . In a d d i t i o n , S p i t Road may be i n c r e a s i n g the r a t e of s u c c e s s i o n i n the p l a n t stands south of the road as the road a c t s as a dyke, red u c i n g the i n u n d a t i o n : emergence r a t i o and improving the s u b s t r a t e drainage i n the a r e a . T 6 2 -3.2.1.12 Salmon R i v e r E s t u a r y The emergent p l a n t communities of the Salmon R i v e r E s t u a r y are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h water c o n d i t i o n s . These s p e c i e s i n c l u d e Lyngby's sedge, t u f t e d h a i r g r a s s , b e n t g r a s s , Kentucky b l u e g r a s s and c i n q u e f o i l . Lyngby's sedge occurs along the l e a d i n g edge of the marsh and i n channels . T u f t e d h a i r g r a s s i f more frequent adjacent and backshore of the sedge stands. It i s o f t e n a s s o c i a t e d with Lyngby's sedge and/or c i n q u e f o i l when i t occurs near channels, and with B a l t i c rush and/or Kentucky b l u e g r a s s when i t occurs away from channels. Bentgrass and Kentucky b l u e g r a s s are dominant towards the backshore and away from channels. The p l a n t s p e c i e s a s s o c i a t e d with bentgrass and Kentucky b l u e g r a s s are more mesic with i n c r e a s i n g d i s t a n c e from channels; adjacent to channels c i n q u e f o i l i s the most abundant a s s o c i a t e d s p e c i e s . C i n q u e f o i l g e n e r a l l y occurs a l l over the marsh from approximately the midpoint i n the d i s t r i b u t i o n of Lyngby's sedge to the t r e e l i n e . Hence, c i n q u e f o i l may be a s s o c i a t e d with many d i f f e r e n t p l a n t s p e c i e s . I t s v i g o r and abundance i s g r e a t e s t where the s u b s t r a t e i s moist and the s a l i n i t y i s freshwater b r a c k i s h . The p l a n t s p e c i e s d i s t r i b u t i o n i s r e l a t e d to two p h y s i c a l f a c t o r s , e l e v a t i o n and water c i r c u l a t i o n . E l e v a t i o n i n f l u e n c e s the i n u n d a t i o n : emergence r a t i o and s u b s t r a t e drainage. On the Salmon R i v e r Estuary, Lyngby's sedge i s dominant between .5 and 1.0 meter i n e l e v a t i o n . In t h i s e l e v a t i o n range i t i s exposed from 39 to 62 percent of the d a y l i g h t hours d u r i n g the growing - 63 -season, and grows i n water s a t u r a t e d s o i l . B a s i c a l l y , the sedge stands are inundated with b r a c k i s h water on every f l o o d t i d e . T ufted h a i r g r a s s i s dominant between 1.0 and 2.0 meters i n e l e v a t i o n . In t h i s e l e v a t i o n range i t i s e x p o s e d from 62 to 89 percent of the d a y l i g h t hours during the growing season, and the s o i l i s b e t t e r d r a i n e d than at lower e l e v a t i o n s . The growth form of t h i s grass f a c i l i t a t e s i t s e s t a b l i s h m e n t . F i r s t l y , the f i b r o u s root system i s able to anchor the t u f t s i n the f i r s t few centimeters of r e l a t i v e l y w e l l d r a i n e d s o i l . Secondly, the hummocks of grass surrounded by channels improve the ra t e and degree of drainage i n the stand. And t h i r d l y , the t u f t s d i r e c t r a i n f a l l to the r o o t s where i t decreases the s o i l s a l i n i t y . The d e c l i n e i n the frequency of occurrence of t u f t e d h a i r g r a s s with i n c r e a s i n g e l e v a t i o n i s p o s s i b l y r e l a t e d to a number of f a c t o r s i n c l u d i n g a s m a l l e r i n u n d a t i o n : emergence r a t i o , c o m p e t i t i o n from other p l a n t s p e c i e s , and d i s t a n c e from freshwater flow. Bentgrass and Kentucky bluegrass are dominant around 1.5 meters i n e l e v a t i o n where they are exposed f o r at l e a s t 85 percent of the d a y l i g h t hours during the growing season. The s u b s t r a t e i s r e l a t i v e l y very w e l l d r a i n e d . The grasses appear able to grow only where they are i n f r e q u e n t l y and inundated by b r a c k i s h freshwater and have a very w e l l d r a i n e d s o i l . The i n f r e q u e n t i n u n d a t i o n impedes competition from more t e r r e s t r i a l p l a n t s p e c i e s found at higher e l e v a t i o n s . C i n q u e f o i l occurs from around .8 to 2.0 meters i n e l e v a t i o n where i t i s exposed from around 50 to 89 percent of the d a y l i g h t hours during the growing season (Kennedy, 1978). The d i s t r i b u t i o n of c i n q u e f o i l does not appear to be l i m i t e d by the length of i n u n d a t i o n as i t i s subdominant over a wide range - 6 4 -of submergence times. However, i t does appear to be i n f l u e n c e d by s u b s t r a t e drainage and water c i r c u l a t i o n . At the lowest e l e v a t i o n s i n i t s d i s t r i b u t i o n i t s s t o l o n i f o r o u s root system a l l o w s i t to become e s t a b l i s h e d i n most s h a l l o w l y d r a i n e d s o i l s . As the drainage improves with e l e v a t i o n i t s abundance and v i g o r i n c r e a s e . Away from channels and at the h i g h e s t e l e v a t i o n s w i t h i n i t s d i s t r i b u t i o n , where i n u n d a t i o n becomes i n f r e q u e n t and s h o r t , and the s o i l i s w e l l d r a i n e d , the abundance and v i g o r of c i n q u e f o i l d e c l i n e s . C i n q u e f o i l , as e a r l i e r noted, i s observed to be a s p e c i e s adapted to a range of growing c o n d i t i o n s with i t s more vigorous growth o c c u r r i n g when the s u b s t r a t e i s moist and the s a l i n i t y i s freshwater b r a c k i s h . The f o u r dominant and/or subdominant p l a n t s p e c i e s on the Salmon R i v e r Estuary, Lyngby's sedge, t u f t e d h a i r g r a s s , Kentucky b l u e g r a s s and c i n q u e f o i l , a l s o tend to be the predominant emergent p l a n t s p e c i e s i n b r a c k i s h e s t u a r i n e marshes north of Kelsey Bay. While t u f t e d h a i r g r a s s , Kentucky bl u e g r a s s and c i n q u e f o i l can be found i n b r a c k i s h e s t u a r i n e marshes to the south, they are more abundant and dominant i n marshes north of Campbell R i v e r . T h i s apparent i n c r e a s e i n predominance i s a s s o c i a t e d with p h y s i c a l f a c t o r s i n c l u d i n g physiography and p r e c i p i t a t i o n . The e s t u a r i e s south of the Salmon River occur i n the p h y s i o g r a p h i c u n i t , the Nanaimo Lowlands, where d e l t a i c d e p o s i t s are more o f t e n sand than g r a v e l (the g r a v e l s are u s u a l l y s m a l l i n s i z e ) and are e x t e n s i v e along s h e l t e r e d c o a s t l i n e and i n coves and bays. E s t u a r i e s north of Say.ward to Kokish R i v e r occur i n the Vancouver I s l a n d Ranges. E s t u a r i e s i n t h i s u n i t occur at the s h e l t e r e d mouthes of v a l l e y s where - 65 -d e l t a i c d e p o s i t s are more o f t e n g r a v e l than sand, and the g r a v e l s are l a r g e i n s i z e . Thin l a y e r s of sand o v e r l a y the g r a v e l s toward the backshore. In g e n e r a l , the e s t u a r i e s north of Sayward have more g r a v e l i n the s u b s t r a t e and are not as l a r g e as e s t u a r i e s to the south. A l s o , i n g e n e r a l , p r e c i p i t a t i o n i n c r e a s e s towards the nor t h (see Table 4 ) . The physiography and p r e c i p i t a t i o n appear to i n f l u e n c e the abundance and dominance of p l a n t s p e c i e s i n four ways. F i r s t l y , the s m a l l e r the e s t u a r y the l e s s d i v e r s e the h a b i t a t and the fewer a s s o c i a t e d p l a n t s p e c i e s . Secondly, g r a v e l s may impede the growth of rhizomatous p l a n t s (e.g., Lyngby's sedge) while f i b r o u s and/or shallow r o o t i n g s p e c i e s (e.g., t u f t e d h a i r g r a s s , Kentucky b l u e g r a s s and c i n q u e f o i l ) grow more e a s i l y between and around r o c k s . T h i r d l y , g r a v e l s are b e t t e r d r a i n e d s o i l s r e s u l t i n g i n more a e r o b i c growing c o n d i t i o n s f o r r o o t s . F o u r t h l y , the high e r p r e c i p i t a t i o n tends to wash s a l t s out of the s u r f a c e s u b s t r a t e s and i s a s s i s t e d i n t h i s by the r e l a t i v e l y good drainage i n g r a v e l l y s o i l s . Thus, s o i l s a l i n i t i e s are g e n e r a l l y lower i n e s t u a r i e s north of Sayward and more glycophytes occur, and are more e x t e n s i v e , i n these northern e s t u a r i e s . 3.2.1.13 Adam-Eve R i v e r s ' Estuary The p l a n t communities of the Adam-Eve R i v e r s ' Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h and s a l i n e c o n d i t i o n s . The s p e c i e s i n d i c a t i n g b r a c k i s h c o n d i t i o n s , Lyngby's sedge and t u f t e d h a i r g r a s s , are dominant i n areas adjacent to the r i v e r . Saltwort, s e a s i d e p l a i n t a i n and arrow-- 66 -g r a s s , s p e c i e s i n d i c a t i v e of s a l i n e c o n d i t i o n s , are dominant at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone and on the e a s t e r n p o r t i o n of the d e l t a . The dominant s p e c i e s towards the backshore of the western p o r t i o n of the d e l t a are Kentucky bluegrass, c i n q u e f o i l , S i t k a spruce and western hemlock, while towards the backshore of the e a s t e r n p o r t i o n , the dominant sp e c i e s are t u f t e d h a i r g r a s s , s a l t w o r t , w i l d r y e and S i t k a spruce. The p l a n t s p e c i e s d i s t r i b u t i o n i s r e l a t e d to the two p r i n c i p a l p h y s i c a l f a c t o r s , the r i v e r s ' d i s c h a r g e during the growing season and d e l t a morphology. No d i s c h a r e f i g u r e s are a v a i l a b l e f o r the Adam or Eve R i v e r s , however, the dominance of s a l t w o r t along the l e a d i n g edge of the marsh i n d i c a t e s s a l i n i t i e s are r e l a t i v e l y high i n t h i s area during the growing season. High s a l i n i t i e s suggest the marine i n f l u e n c e i s g r e a t e r than the freshwater i n f l u e n c e . T h i s i s the small creek c r o s s i n g the e a s t e r n p o r t i o n of the d e l t a appears to have an i n t e r m i t t e n t flow. In areas of low flow or l i m i t e d freshwater i n f l u e n c e , p a r t i c u l a r l y at the lower e l e v a t i o n s of the vegetated study area, the marine i n f l u e n c e dominates, s a l i n i t e s are r e l a t i v e l y higher, and s p e c i e s t o l e r a n t of higher s a l i n i t i e s (e.g., s a l t w o r t , s e a s i d e p l a n t a i n ) occur. D e l t a morphology also appears to i n f l u e n c e the tendency towards higher s a l i n i t i e s . The r i g h t bank at the mouth of the r i v e r i n h i b i t s freshwater flow across the e a s t e r n p o r t i o n of the d e l t a . Reduced freshwater flow a c r o s s t h i s area heightens marine i n f l u e n c e , and i n c r e a s e s s a l i n i t i e s . The extent of the marsh a l s o appears to be a f u n c t i o n of d e l t a morphology. The marsh on the e a s t e r n p o r t i o n of the d e l t a probably e x i s t s due to p r o t e c t i o n a f f o r d e d - 67 -i t by the r i g h t bank of the r i v e r , i s l a n d and small s p i t of land along i t s f o r e s h o r e . The marsh i s e x t e n s i v e throughout the p r o t e c t e d p o r t i o n of the d e l t a . The western p o r t i o n may not be as e x t e n s i v e l y vegetated f o r two reasons. F i r s t l y , exposure of the western d e l t a to wind and water energy may i n h i b i t a c c r e t i o n at the lower e l e v a t i o n s which tends to r e s u l t i n low exposure: inundation r a t i o s u n s u i t a b l e f o r p l a n t s p e c i e s e s t a b l i s h m e n t . Secondly, emergent p l a n t s tend to c o l o n i z e g r a v e l s with d i f f i c u l t y and are probably d i s l o d g e d by wind and waves at the lower e l e v a t i o n s . Human a c t i v i t y , i n the form of log handling on the western p o r t i o n of the d e l t a , may emphasize some of the p h y s i c a l f a c t o r s i n f l u e n c i n g p l a n t s p e c i e s d i s t r i b u t i o n . It i s p o s s i b l e t h a t the l o g h a n d l i n g s i t e encourages the r i v e r to flow p r i n c i p a l l y i n the e x i s t i n g main channel i n s t e a d of d i f f u s i n g a c r o s s the marsh. The r i v e r i s s i t u a t e d so that i t would appear to have run through the l o g s o r t area at one time. Old channels at the south end of the l o g s o r t f u r t h e r t h i s s u g g e s t i o n . R e s t r i c t e d freshwater flow across the marsh i n c r e a s e s the tendency towards higher s a l i n i t i e s . Another r e s u l t of log handling i s the accumulation of beached logs on the backshore of the e a s t e r n p o r t i o n of the d e l t a . The logs have +ended to i n c r e a s e the e l e v a t i o n which improved the s u b s t r a t e drainage and c r e a t e d c o n d i t i o n s more s u i t a b l e f o r the establishment of w i l d r y e and S i t k a spruce. - 68 -3.2.1.14 T s i t i k a R i v e r E s t u a r y The emergent p l a n t c o m m u n i t i e s o f t h e T s i t i k a R i v e r E s t u a r y a r e d o m i n a t e d by s p e c i e s i n d i c a t i v e o f b r a c k i s h w ater c o n d i t i o n s . T h e s e s p e c i e s i n c l u d e L y n g b y ' s sedge and t u f t e d h a i r g r a s s . L y n g b y ' s sedge i s f o u n d on sandy s u b s t r a t e s a t t h e l o w e r e l e v a t i o n s o f t h e v e g e t a t e d s t u d y a r e a . T i i " + e d h a i r g r a s s i s f o u n d on g r a v e l a t l o w e r e l e v a t i o n s and on sandy s u b s t r a t e a t h i g h e r e l e v a t i o n s . At t h e t r e e l i n e , p l a n t c o m m u n i t i e s a r e d o m i n a t e d by w i l d r y e , c o w - p a r s n i p , g i a n t v e t c h and S i t k a s p r u c e s e e d l i n g s when l o g d e b r i s i s p r e s e n t . When i t i s a b s e n t t h e dominant p l a n t s a r e s p r i n g b a n k c l o v e r , y a r r o w , a r r o w - g r a s s and v e l v e t - g r a s s . The p l a n t s p e c i e s d i s t r i b u t i o n i s r e l a t e d t o two p r i n c i p a l p h y s i c a l f a c t o r s , e l e v a t i o n and s u b s t r a t e . At l o w e r e l e v a t i o n s L y n g b y ' s sedge and t u f t e d h a i r g r a s s d o m i n a t e ; L y n g b y ' s segdge i n sandy s u b s t r a t e s and t u f t e d h a i r g r a s s i n g r a v e l . T h e s e s p e c i e s a r e a d a p t e d t o the h i g h e r i n u n d a t i o n : emergence r a t i o a t t h i s e l e v a t i o n , however, t h e r h i z o m e o f Lyngby's sedge c o l o n i z e s sand more r e a d i l y t h a n g r a v e l s u b s t r a t e s , and t o l e r a t e s w ater s a t u r a t e d s o i l s b e t t e r t h an t u f t e d h a i r g r a s s , w h i c h c o l o n i z e s and grows f a i r l y v i g o r o u s l y on t h e b e t t e r d r a i n e d g r a v e l s ( d e t a i l s d i s c u s s e d p r e v i o u s l y ) . The w i l d r y e , g i a n t v e t c h , S i t k a s p r u c e and c o w - p a r s n i p i n l o g d e b r i s a l o n g t h e t r e e l i n e a r e a t an e l e v a t i o n where t h e s u b s t r a t e i s r e l a t i v e l y w e l l d r a i n e d and o t h e r w i s e s u i t a b l e f o r c o l o n i z a t i o n by t r e e s . However, i n f r e q u e n t f l o o d t i d e c o n d i t i o n s can c a u s e t h e l o g d e b r i s t o move and d i s l o d g e s e e d l i n g s . W i l d r y e i s a b l e t o s u r v i v e t h e i n f r e q u e n t f l o o d i n g and l o g movement, and becomes - 69 -e s t a b l i s h e d . Log d e b r i s has not accumulated on the d e l t a ' s c e n t r a l lobe and there the f o r e s t i s bordered by springbank c l o v e r , yarrow, arrow-grass and v e l v e t - g r a s s . Another p h y s i c a l f a c t o r t h a t appears to be a parameter i n the p l a n t s p e c i e s d i s t r i b u t i o n , p a r t i c u l a r l y on the e a s t e r n lobe of the d e l t a , i s streamflow. As present, the main flow of the T s i t i k a River i s i n the e a s t e r n channel and during high water the r i v e r erodes the e a s t e r n lobe ( p e r s . o b s e r v a t i o n based on s i t e i n s p e c t i o n s i n 1974 and 1976). The dominant s p e c i e s on the e a s t e r n lobe i s t u f t e d h a i r g r a s s which grows i n hummocks. The hummocky growth p a t t e r n i s c o n s i d e r e d to give l e s s r e s i s t a n c e to running water and may r e s u l t from i t . In areas of high water energy on other e s t u a r i e s t u f t e d h a i r g r a s s has a l s o been observed to grow i n hummocks. 3.2.1.15 Kokish R i v e r Estuary The emergent p l a n t communities of the Kokish River Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h and s a l i n e c o n d i t i o n s . The s p e c i e s i n d i c a t i v e of b r a c k i s h c o n d i t i o n s , Lyngby's sedge, t u f t e d h a i r g r a s s and c i n q u e f o i l , are l o c a t e d west of the r a i l w a y , at the r i v e r mouth. T h e i r d i s t r i b u t i o n and the p r i n c i p a l p h y s i c a l f a c t o r s i n f l u e n c i n g t h e i r d i s t r i b u t i o n are d i s c u s s e d ?n d e t a i l elsewhere (see d i s c u s s i o n s e c t i o n f o r each s p e c i e s and Salmon R i v e r E s t u a r y ) . The s p e c i e s i n d i c a t i v e of s a l i n e c o n d i t i o n s , s a l t w o r t , seaside p l a n t a i n , s a l t g r a s s and s a l t b u s h , are dominant east of the r a i l w a y . T h i s e a s t e r n p o r t i o n of the study area i s bounded by a r a i l w a y bed on the west and north, a l a n d f i l l on the east, - 70 -and the f o r e s t on the south. The area i s e n c i r c l e d except f o r a seaward opening through which sea water c i r c u l a t e s . The s a l i n e water c o n d i t i o n , the p r i n c i p a l p h y s i c a l f a c t o r governing t'e d i s t r i b u t i o n of the s p e c i e s i n the eastern p o r t i o n of the e s t u a r y , appears to be i n f l u e n c e d by human a c t i v i t y . T h i s human a c t i v i t y i s a s s o c i a t e d with the r a i l movement of logs to the log s o r t i n g area along the f o r e s e t beds of the d e l t a . The r a i l w a y bed i n h i b i t s freshwater flow to the east and t h e r e f o r e tends to promote the predominantly s a l i n e c o n d i t i o n s t h e r e . However, b r a c k i s h c o n d i t i o n s may be found around two channels i n t h i s area. The dominance of Lyngby's sedge, .a b r a c k i s h water i n d i c a t o r s p e c i e s , along the channels suggests that the channels c a r r y some freshwater. Since the e s t u a r y was surveyed i n 1975 a d r y l a n d s o r t has r e p l a c e d the water s o r t and outmoded the r a i l w a y e n c i r c l i n g the e a s t e r n h a l f of the e s t u a r y . Freshwater flow to t h i s h a l f of the estuary could be improved by breaching the r a i l w a y bed where the two channels used to flow from the r i v e r . I t i s expected that t h i s would reduce the s a l i n i t y and i n c r e a s e the v i g o r and abundance of the b r a c k i s h water p l a n t s p e c i e s . With m o d i f i c a t i o n s to the area's three openings, to c o n t r o l water q u a l i t y and c r e a t e an e n c l o s u r e , the e a s t e r n h a l f of the estuary might be used to rear salmonids ( p e r s . comm. Dr. M. Brownlee). 3.2.1.16 Nimpkish R i v e r Estuary The emergent p l a n t communities of the Nimpkish R i v e r Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h water - 71 -c o n d i t i o n s . These s p e c i e s i n c l u d e t u f t e d h a i r g r a s s , c i n q u e f o i l and Lyngby's sedge. T h e i r d i s t r i b u t i o n and the p r i n c i p a l p h y s i c a l f a c t o r s i n f l u e n c i n g t h e i r d i s t r i b u t i o n are d i s c u s s e d i n d e t a i l elsewhere (see d i s c u s s i o n s e c t i o n f o r each s p e c i e - and Salmon R i v e r E s t u a r y ) . At lower e l e v a t i o n s , s p i k e - r u s h i s a dominant s p e c i e s and, at higher e l e v a t i o n s on the two i s l a n d s , S i t k a spruce, Nootka rose, gooseberry and w i l d r y e are dominant. On the south bank a freshwater marsh i s dominated by sweet ga l e . The p r i n c i p a l p h y s i c a l f a c t o r s i n f l u e n c i n g the d i s t r i b u t i o n of these s p e c i e s are the i n u n d a t i o n : emergence r a t i o and s a l i n i t y . Favorable growing c o n d i t i o n s f o r s p i k e - r u s h are a r e l a t i v e l y small i n u n d a t i o n : emergence r a t i o and a b r a c k i s h freshwater s a l i n i t y range where the s u b s t r a t e i s sandy. S i t k a spruce, Nootka rose and gooseberry grow on w e l l d r a i n e d s u b s t r a t e s where they are i n f r e q u e n t l y and then only s h o r t l y inundated and the s a l i n i t y i s b r a c k i s h freshwater. On the downstream i s l a n d the s p e c i e s are e n c i r c l e d by w i l d r y e which probably p r o v i d e s some p r o t e c t i o n from s a l t spray. Wildrye i s a l s o dominant along the exposed south bank of the r i v e r . The grass seems to r e q u i r e growing c o n d i t i o n s o r i g i n a t i n g , on the Nimpkish R i v e r E s t u a r y , from a combination of three f a c t o r s , a very w e l l d r a i n e d g r a v e l s u b s t r a t e , i n f r e q u e n t , short i n u n d a t i o n , and s a l i n i t y from s a l t spray. Behind the w i l d r y e community, towards the backshore on the south bank, i s a marsh where sweet gale grows i n standing freshwater. Water c o l l e c t s i n a d e p r e s s i o n which may be an o l d r i v e r channel or may be the r e s u l t of e a r l i e r a c t i v i t y r e l a t i n g to l o g g i n g . Human a c t i v i t y , - 72 -p a r t i c u l a r l y l o g g i n g , has had other i n f l u e n c e s on the e s t u a r y . The abundance of a l i e n s p e c i e s such as v e l v e t - g r a s s , orchard grass, timothy and c r e s t e d dog's t a i l , suggest the area has been d i s t u r b e d . Indeed, the streambed g r a v e l d i s t r i b u t i o n , and shape and s i z e of the downstream i s l a n d i n d i c a t e past s u b s t r a t e a l t e r a t i o n , p o s s i b l y i n c o n j u n c t i o n with l o g movement. The presence of what appears to be an o l d log dump (see 0 on F i g u r e 19) on the south bank lends some credence to t h i s o b s e r v a t i o n . 3.2.1.17 Cluxewe R i v e r Estuary The emergent p l a n t communities of the Cluxewe R i v e r d e l t a are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h and s a l i n e c o n d i t i o n s . Species i n d i c a t i v e of b r a c k i s h water c o n d i t i o n s are most abundant on the e a s t e r n corner of the d e l t a , at the r i v e r mouth, and i n c l u d e Lyngby's sedge, t u f t e d h a i r g r a s s and c i n q u e f o i l . T h e i r d i s t r i b u t i o n and the p r i n c i p a l p h y s i c a l f a c t o r s i n f l u e n c i n g t h e i r d i s t r i b u t i o n are d i s c u s s e d i n d e t a i l elsewhere (see d i s c u s s i o n s e c t i o n f o r each s p e c i e s and Salmon R i v e r E s t u a r y ) . Species i n d i c a t i v e of s a l i n e water c o n d i t i o n s are most abundant west of the r i v e r mouth and i n c l u d e s a l t w o r t and arrow-grass. These s p e c i e s are mostly screened from the sea by a g r a v e l berme which i s c o l o n i z e d by S i t k a spruce and w i l d r y e . On a high t i d e , the s a l t w o r t and arrow-grass are covered with s a l t w a t e r that e n t e r s the low land through a breach i n the berme. The berme p r o t e c t s the western d e l t a from the d e s t r u c t i v e wave and wind energy of the sea thus promoting the e s t a b l i s h m e n t of p l a n t s . However, i t a l s o impedes freshwater flow from the Cluxewe R i v e r which tends to i n c r e a s e the s a l i n i t y - 73 -range. The higher s a l i n i t i e s are t o l e r a t e d best by s a l t w o r t and arrow-grass. The east corner of the western d e l t a i s dominated by g r a s s e s r a t h e r than s a l t w o r t and arrow-grass. T h i s i s probably a n a t u r a l s i t u a t i o n which has been enhanced by previous attempts at d r a i n i n g the area. Old dykes and drainage d i t c h e s appear to have improved drainage and reduced the frequency and extent of i n u n d a t i o n i n the east corner and c i n q u e f o i l , t u f t e d h a i r g r a s s , Kentucky b l u e g r a s s , meadow b a r l e y and wi l d r y e are dominant there probably i n response to the improved drainage and lower i n u n d a t i o n : emergence r a t i o . The dominance of meadow ba r l e y and w i l d r y e suggest the s u b s t r a t e s a l i n i t y may be high, p a r t i c u l a r l y i n the l a t t e r h a l f of the growing season, and may be c o n s i d e r e d i n d i c a t i v e of p h y s i o l o g i c a l l y dry growing c o n d i t i o n s . X e r o p h y t i c growing c o n d i t i o n s r e l a t i n g to good s u b s t r a t e drainage, s a l t s p r a y and i n f r e q u e n t f l o o d i n g i n f l u e n c e the dominance of S i t k a spruce and w i l d r y e on the g r a v e l berme. It i s proposed that the d e l t a of the Cluxewe R i v e r r e f l e c t s the kind of d e l t a morphology that e x i s t e d at the mouths of the Englishman, L i t t l e Qualicum, Oyster, and Campbell R i v e r s before human i n t e r v e n t i o n . A l l the r i v e r s appear, i n an e a r l i e r time, to have flowed p a r a l l e l to responses, both past and present, which c h a r a c t e r i z e d e p o s i t i o n a l areas of the east coast of Vancouver I s l a n d . The p h y s i c a l s i m i l a r i t i e s between the f i v e aforementioned d e l t a s may not be e v i d e n t . The only apparent c h a r a c t e r i s t i c common to the Englishman, L i t t l e Qualicum, Campbell and Cluxewe d e l t a s appears to be t h e i r s p i t s . The Englishman and Cluxewe d e l t a s have bermes as w e l l as s p i t s , and the L i t t l e Qualicum and Campbell d e l t a s do not. The Oyster d e l t a i s not s i m i l a r to the - 74 -others i n that i t does not have e i t h e r s p i t s or bermes. However, i t appears that the Oyster d e l t a has had bermes. Convoluted channels on the Oyster R i v e r d e l t a run p a r a l l e l and backshore of a t r e e d s t r i p of l a n d . T h i s t r e e d s t r i p i s considered to be an o l d berme. The b a s i s f o r t h i s c o n c l u s i o n i s the s i m i l a r i t i e s , i n c h a n n e l i z a t i o n p a t t e r n and l o c a t i o n of channels, between the Cluxewe and Oyster R i v e r s ' d e l t a s . It i s surmised that the morphology of the Big Qualicum R i v e r d e l t a was once l i k e that of the Oyster R i v e r d e l t a . The reasons f o r t h i s are i t s s i m i l a r l o c a t i o n on an open c o a s t l i n e and dark areas on a i r photographs. These dark areas are long and narrow and p a r a l l e l with the s h o r e l i n e which suggests o l d channels. While the convoluted c h a n n e l i z a t i o n p a t t e r n s , r i v e r flow p a r a l l e l to the s h o r e l i n e , s p i t s and bermes suggest low flows, the e x t e n s i v e d e l t a i c d e p o s i t s at each r i v e r mouth suggest l a r g e flows. T h i s i s not c o n s i d e r e d c o n t r a d i c t o r y . Instead, i t i s considered i n d i c a t i v e of a general trend toward sma l l e r flows. I t i s thought that as the g l a c i e r s the s h o r e l i n e before e n t e r i n g the sea, and the convoluted channels on the western d e l t a of the Cluxewe R i v e r are thought + o exemplify the c h a n n e l i z a t i o n p a t t e r n a s s o c i a t e d with t h i s p a r a l l e l flow. It appears that the r i v e r s flowed p a r a l l e l to the s h o r e l i n e because t h e i r energy was not great enough to overcome the opposing energy of the sea and flow i n t o i t . Instead, the sea f o r c e d the r i v e r s to flow along the s h o r e l i n e u n t i l they reached a breakout, p o i n t . T h i s might mean that a r i v e r flowed i n t o the waves from behind a s p i t . A s p i t and/or berme are a s s o c i a t e d with each of the above r i v e r s , and provide each r i v e r d e l t a with some p r o t e c t i o n from the wind and wave energy of the sea. A berme i s a n e a r l y h o r i z o n t a l - 75 -p o r t i o n of the beach or backshore formed by the d e p o s i t of m a t e r i a l by wave a c t i o n . Suspended m a t e r i a l i n the r i v e r s e t t l e s out upon c o n t a c t with s a l t w a t e r . Where the r i v e r ' s c u r r e n t i s s t r o n g i t c a r r i e s suspended m a t e r i a l i n t o the sea forming a fan shaped d e l t a , and where the c u r r e n t i s not strong the m a t e r i a l s e t t l e s out of suspension along the s h o r e l i n e at an e l e v a t i o n corresponding with the t i d e l i n e . When the r i v e r breaks through the berme a s p i t and berme or p a i r of bermes r e s u l t . The formation of s p i t s and/or bermes at the mouths of the Englishman, L i t t l e Qualicum, Oyster and Campbell R i v e r s probably o c c u r r e d at some time i n the past, when the amount of suspended m a t e r i a l i n the r i v e r s was higher, and t h e i r water energy was lower than the sea's. P r e s e n t l y , the s p i t s and bermes appear to be maintained by longshore d r i f t . Hale and McCann's ongoing work on the c o a s t a l geomorphology and dynamics of. the Georgia S t r a i t c o a s t l i n e i s expected to d e t a i l the p r o c e s s e s i n v o l v e d i n the e v o l u t i o n of c o a s t a l u n i t s such as s p i t s and bermes. I t i s a n t i c i p a t e d t h a t t h e i r work w i l l c l a r i f y concepts of c o a s t a l processes and that as the g l a c i e r s receded the r i v e r s ' flow and suspended m a t e r i a l were great enough to b u i l d the l a r g e d e l t a s and then d e c l i n e d u n t i l the r i v e r s ' d i d not have the energy to flow i n t o the sea. Hence, s p i t s and bermes were formed. Large flows are considered to have preceded s m a l l e r f l o w s . If i t was the other way around, l a r g e flows would have e r a d i c a t e d the s p i t s and bermes. Flov/s now are probably r e l a t i v e l y s l i g h t l y g r e a t e r than when the s p i t s and bermes v/ere formed. The morphology of the Englishman, L i t t l e Qualicum, Big Qualicum, Oyster, Campbell and Cluxewe R i v e r s ' d e l t a s encourages - 76 -development f o r two reasons. F i r s t l y the l a r g e , f l a t d e l t a s o f f e r ample space f o r human a c t i v i t y . The second i s that i t i s r e l a t i v e l y easy to dyke, d r a i n and l a n d f i l l channels surrounded by f o r e s t and p r o t e c t e d from the elements by s p i t s and bermes. The degree of development v a r i e s "from d e l t a to d e l t a but i n gen e r a l those d e l t a s with j u s t bermes, (e.g., the Big Qualicum and Oyster R i v e r s ' d e l t a s ) , and the southern e s t u a r i e s s u b j e c t to the g r e a t e s t p o p u l a t i o n p r e s s u r e s e x h i b i t the most ext e n s i v e a l t e r a t i o n s . The concept of "ext e n s i v e a l t e r a t i o n s " being r e l a t i v e to the t o t a l d e l t a i c land area. 3.2.1.18 Quatse R i v e r Estuary The emergent p l a n t communities of the Quatse River Estuary are dominated by s p e c i e s i n d i c a t i v e of b r a c k i s h and s a l i n e c o n d i t i o n s . These s p e c i e s i n c l u d e Lyngby's sedge, t u f t e d h a i r g r a s s , c o l o n i a l bentgrass, c i n q u e f o i l , s a l t w o r t and arrow-g r a s s . The d i s t r i b u t i o n of the s p e c i e -indicating b r a c k i s h c o n d i t i o n s , and the p r i n c i p a l p h y s i c a l f a c t o r s i n f l u e n c i n g t h e i r d i s t r i b u t i o n are d i s c u s s e d i n d e t a i l elsewhere (~ee d i s c u s s i o n s e c t i o n s f o r each s p e c i e s and Salmon R i v e r E s t u a r y ) . The s p e c i e s i n d i c a t i n g s a l i n e c o n d i t i o n s occur at the lowest e l e v a t i o n s i n the vegetated study area and away from the i n f l u e n c e of freshwater. Towards the backshore, grasses are dominant at lower e l e v a t i o n s , and t r e e s and shrubs are dominant at higher e l e v a t i o n s , p a r t i c u l a r l y on and adjacent to the roads and dykes. - 77 -E l e v a t i o n appears to be the p r i n c i p a l p h y s i c a l f a c t o r i n f l u e n c i n g the d i s t r i b u t i o n of the emergent p l a n t s p e c i e s on the Quatse R i v e r E s t u a r y . With i n c r e a s i n g e l e v a t i o n drainage improves and, i n a s s o c i a t i o n , the p l a n t s p e c i e s progress from s a l t w o r t to Lyngby's sedge; arrow-grass to Lyngby's sedge; t u f t e d h a i r g r a s s to t u f t e d h a i r g r a s s and c o l o n i a l bentgrass; c i n q u e f o i l to g r a s s e s ; to shrubs and f o r e s t . T h i s n a t u r a l p r o g r e s s i o n has been i n t e r r u p t e d by the road, dykes and i n d u s t r i a l park c o n s t r u c t i o n . The road and dykes have improved drainage to t h e i r south encouraging the e s t a b l i s h m e n t of more t e r r e s t r i a l p l a n t s p e c i e s (e.g., red a l d e r , salmonberry and v e l v e t - g r a s s ) which d i s p l a c e emergent s p e c i e s . The l a n d f i l l i n g and e a r t h moving f o r the I n d u s t r i a l Park have b u r i e d and removed other v e g e t a t i o n . The s o i l s a l i n i t y i s a secondary p h y s i c a l f a c t o r warranting comment. On the Quatse R i v e r Estuary, the s a l i n i t y appears to be r e l a t e d to e l e v a t i o n and p r e c i p i t a t i o n . During the growing season the l i m i t e d freshwater flow r e s u l t s i n a predominantly marine i n f l u e n c e , p a r t i c u l a r l y at the lowest e l e v a t i o n s where the s o i l i s probably always s a t u r a t e d . There i t i s s a l i n e and s a l t w o r t dominates. As the e l e v a t i o n i n c r e a s e s , the drainage improves and p r e c i p i t a t i o n d i l u t e s the s a l t s , r e d u c i n g the s o i l s a l i n i t y . Lyngby's sedge, t u f t e d h a i r g r a s s and other s p e c i e i n d i c a t i n g b r a c k i s h c o n d i t i o n s occur i n t h i s a r ea. - 78 -3.2.1.19 Kingcome Ri v e r Estuary The emergent plant communities of the Kingcome River Estuary are dominated by species i n d i c a t i v e of b r a c k i s h c o n d i t i o n s . These species i n c l u d e Lyngby's sedge and t u f t e d h a i r g r a s s . The d i s t r i b u t i o n of the species and the p r i n c i p a l p h y s i c a l f a c t o r s i n f l u e n c i n g t h e i r , d i s t r i b u t i o n are discussed i n d e t a i l elsewhere (see d i s c u s s i o n s e c t i o n s f o r each species and Salmon Ri v e r E s t u a r y ) . Unfortunately time and c o n d i t i o n s precluded an exhaustive survey of the d e l t a . Where p o s s i b l e , areas that were not examined i n the f i e l d were typed from a e r i a l photographs. These and areas that could not be typed are i n d i c a t e d by a question mark. A l l questionable areas are thought to be dominated by species from the Grass Family. Two observations r e l a t i v e to the Kingcome River Estuary are worthy of comment. The f i r s t , i s the presence of freshwater i n d i c a t o r s p e c i e s , spike-rush and w i l l o w . Spike-rush stands are found along the leading edge of the vegetation and w i l l o w stands on midstream i s l a n d s . The freshwater growing c o n d i t i o n s r e q u i r e d by spi'-e-rush are probably prevalent during the growing season, when the r i v e r i s i n f r e s h e t . During the remainder of the year, the r i v e r flow i s probably great enough to keep the s a l t wedge from pene t r a t i n g to the willow stands on the midstream i s l a n d s . The mean annual 2000 mm of p r e c i p i t a t i o n i n the region a l s o help keep s a l i n i t i e s low. The spring freshet and predominant freshwater c o n d i t i o n s year-round on the Kingcome River Estuary are almost opposite to c o n d i t i o n s i n e s t u a r i e s on - 79 -the east coast of Vancouver I s l a n d . There' the r i v e r s are i n f r e s h e t i n the winter, u s u a l l y December or January, and s a l i n i t i e s are highest i n the growing season when flows and p r e c i p i t a t i o n are low. The second observation i s the r e l a t i v e l y small number of species found on the estuary. On the Kingcome River Estuary, 42 plant species i n s i x communities cover 145 hectares. The plant communities on the Salmon River Estuary cover a comparable area but have twice the number of plant species as those on the Kingcome Ri v e r Estuary. On the Nanaimo River Estuary, the plant communities cover a l i t t l e more than o n e - t h i r d of the area covered by communities on the Kingcome River Estuary yet have twice the number of plant species. The d i f f e r e n c e i s considered to be r e l a t e d to two p h y s i c a l f a c t o r s , sedimentation and water energy. The Kingcome River d e l t a i s a c c r e t i n g and estimated suspended sediment concentrations f o r the r i v e r are two, four and, i n some ins t a n c e s , four hundred times greater than estimates f o r r i v e r s on the east coast of Vancouver Is l a n d . This sediment covers p l a n t s , i n t e r f e r i n g with l i g h t absorption f o r photosynthesis, and may bury p l a n t s . Only the h a r d i e s t plants (e.g., Lyngby's sedge and t u f t e d h a i r g r a s s ) have a chance of growing under these c o n d i t i o n s . Growing c o n d i t i o n s are made more adverse by the r i v e r ' s energy. It i s great enough at times to cut many, two meter deep channels through the southern, U-shaped i s l a n d . These channels are so numerous that they may be encountered with every step i n the t u f t e d h a i r g r a s s community. Such water energy i s capable of uprooting a l l but the most f i r m l y rooted p l a n t s (e.g., Lyngby's sedge and t u f t e d h a i r g r a s s ) - 80 -f u r t h e r l i m i t i n g the number of p l a n t s p e c i e s capable of becoming e s t a b l i s h e d on the d e l t a . 3.3.2 Dominant Pl a n t Species The p r e v i o u s d i s c u s s i o n i s based on the dominant plant s p e c i e s of communities as i n d i c a t o r s of t h e i r growing c o n d i t i o n s (Clements, 1939). For example, on the b a s i s of the dominance of s a l i n e i n d i c a t o r s p e c i e s on the Goldstream, the Goldstream i s 1. a s a l t marsh, 2. grou-ed with the Chemaimi<= and Nanoo e-Bonell s a l t marshes, and 3. has three p r i n c i p a l p h y s i c a l f a c t o r s ( l a r g e s i z e of d e l t a , low streamflow and low p r e c i p i t a t i o n ) i n f l u e n c i n g the d i s t r i b u t i o n of i t s p l a n t communities. As so much of the d i s c u s s i o n i s based on the dominant p l a n t s as i n d i c a t o r s of t h e i r growing c o n d i t i o n s an e x p l a n a t i o n of the p h y s i c a l f a c t o r s i n f l u e n c i n g the growth of the dominant plant s p e c i e s was u s e f u l . Another reason f o r the d i s c u s s i o n of the dominant s p e c i e s i n the communities i s man's l a c k of f a m i l i a r i t y with e s t u a r i n e p l a n t s . E s t u a r i n e v e g e t a t i o n i n i t s n a t u r a l form i s not much used by mankind p a r t l y because the v e g e t a t i o n i s in-ndated much of the time and human movements are very r e s t r i c t e d . It t h e r e f o r e seemed u s e f u l to d i s c u s s the dominant s p e c i e s which c h a r a c t e r i z e the e s t u a r i e s and t h i s d i s c u s s i o n f o l l o w s . Acer macrophyllum Pursh, F l . ( b r o a d l e a f maple) Native t r e e B r o a d l e a f maple i s a t r e e of lowlands, or bottomlands where permanent seepage has the s o i l wet f o r most of the v e g e t a t i v e season ( K r a j i n a , 1969/70). The presence of - 81 -subsurface water flow i n d i c a t e s medium to very r i c h n u t r i e n t regimes due to the i n f l u x of n u t r i e n t s i n the seepage water ( K l i n k a , .1977). Broadleaf maple i s a subdominant i n the Quercus  garryana - Acer macrophyllum community on the Nanaimo R i v e r E s t u a r y . Man's a c t i v i t i e s may have s u b s t a n t i a l l y reduced t h i s maple on other e s t u a r i e s f o r i t i s o f t e n a s s o c i a t e d with s o i l s used f o r a g r i c u l t u r a l purposes. A c h i l l e a m i l l e f o l i u m L. (yarrow) n a t i v e herb Yarrow, as a compositous herb, has some s a l t t o l e r a n c e . It u s u a l l y occurs on w e l l d r a i n e d s i t e s which are inundated f o r only sh o r t p e r i o d s of time and may be found i n a s s o c i a t i o n with grasses and c i n q u e f o i l . It i s present on almost a l l e s t u a r i e s and i s a dominant or subdominant on the Cowichan, Nanoose-B o n e l l , Englishman, Salmon, T s i t i k a and Cluxewe R i v e r s ' E s t u a r i e s . Yarrow might be dominant on more e s t u a r i e s i f more backshore areas remained. Agropyron repens (L.) Beauv. (couch g r a s s ) introduced from Europe Couch grass develops and reproduces i n the l a t t e r h a l f of J u l y through September, on w e l l - d r a i n e d s i t e s that tend to be inundated f o r r e l a t i v e l y s h o r t p e r i o d s of time. It seems to grow under p h y s i o l o g i c a l l y dry c o n d i t i o n s . The s p e c i e s l i f e h i s t o r y , tendency towards w e l l - d r a i n e d s i t e s , r e l a t i v e l y short i n u n d a t i o n p e r i o d , and a s s o c i a t e d s p e c i e s suggest t h i s . It i s a dominant and/or subdominant on the Cowichan, Nanaimo, Englishman, Salmon and Adam-Eve R i v e r s ' E s t u a r i e s . Some of the - 82 -s p e c i m e n s c o l l e c t e d a p p e a r t o be a h y b r i d o f A g r o p y r o n r e p e n s and E l y m u s g l a u c u s . A g r o s t i s a l b a L. ( b e n t g r a s s ) A g r o s t i s a l b a v a r . s t o l o n i f e r a L. ( b e n t g r a s s ) A g r o s t i s t e n u i s S i b t h . ( C o l o n i a l b e n t g r a s s ) n a t i v e and i n t r o d u c e d , g e n o t y p e s , t e m p e r a t e E u r a s i a a n d N o r t h A m e r i c a . B e n t g r a s s e s t e n d t o o c c u r i n t h e more e l e v a t e d o r b a c k s h o r e a r e a s where t h e s u b s t r a t e i s r e l a t i v e l y w e l l - d r a i n e d and i n u n d a t i o n , g e n e r a l l y , i s f o r o n l y s h o r t p e r i o d s o f t i m e . They a r e u s u a l l y f o u n d i n a s s o c i a t i o n w i t h o t h e r g r a s s e s and f o r b s . A g r o s t i s a l b a i s a d o m i n a n t a n d / o r s u b d o m i n a n t on t h e G o l d s t r e a m , N a n o o s e - B o n e l l , E n g l i s h m a n and O y s t e r R i v e r s ' E s t u a r i e s . A g r o s t i s a l b a v a r . s t o l o n i f e r a i s a d o m i n a n t and s u b d o m i n a n t on t h e S a l m o n R i v e r E s t u a r y and A g r o s t i s t e n u i s i s a d o m i n a n t a n d / o r s u b d o m i n a n t on t h e Nanaimo, K o k i s h , C l u x e w e and Q u a t s e R i v e r s ' E s t u a r i e s . A l n u s r u b r a Bong, ( r e d a l d e r ) n a t i v e t r e e Red a l d e r , a p i o n e e r s p e c i e s , i s g e n e r a l l y f o u n d i n t h e l o w l a n d s t o h i l l s a l o n g t h e e a s t c o a s t o f V a n c o u v e r I s l a n d . I n t h e e s t u a r i e s i t may o c c u r i n t h e b a c k s h o r e where t h e s o i l n u t r i e n t r e g i m e i s medium t o v e r y r i c h and t h e r e i s p e r m a n e n t s e e p a g e o r a v e r y h i g h w a t e r t a b l e . I t i s a d o m i n a n t a n d / o r - 83 -subdominant on the Englishman, Campbell and Quatse R i v e r s ' E s t u a r i e s . Anthoxanthum odoratum L. (sweet v e r n a l g r a s s ) introduced Sweet v e r n a l g r a s s may be found on very w e l l - d r a i n e d s i t e s such as shallow s o i l over g r a v e l or over bedrock. It i s almost never inundated but may be exposed to s a l t spray. It i s a subdominant on the L i t t l e Qualicum, Big Qualicum and and Cluxewe R i v e r s ' E s t u a r i e s . Arbutus m e n z i e s i i Pursh. (arbutus) n a t i v e tree Arbutus i s a subdominant on the Shoal I s l a n d s i n the Chemainus R i v e r E s t u a r y . Here the f o r e s t i s dry and the s o i l w e l l d r a i n e d , o f t e n shallow and n u t r i e n t poor with low s u p p l i e s of c a l c i u m and magnesium. Arbutus i s r e s t r i c t e d to very w e l l drained s i t e s ; never f a r from s a l t water, i t i s b e l i e v e d to be the most f r o s t s u s c e p t i b l e t r e e i n Canada ( K r a j i n a 1969/70). A s t e r h e s p e r i u s Gray (marsh a s t e r ) n a t i v e herb Marsh a s t e r occurs where c o n d i t i o n s are p h y s i o l o g i c a l l y dry. On the Goldstream Estu a r y , marsh a s t e r grows i n the r e l a t i v e l y more e l e v a t e d areas nearer the backshore. There the s u b s t r a t e i s r e l a t i v e l y w e l l - d r a i n e d , i n f r e q u e n t l y inundated, and p o s s i b l y s l i g h t l y s a l i n e . - 84 -A t r i p l e x p a t u l a L. ( s a l t b u s h ) n a t i v e herb Sa l t b u s h i s i n d i c a t i v e of s a l i n e or b r a c k i s h c o n d i t i o n s . It i s u s u a l l y most f r e q u e n t l y found i n the lower vegetated i n t e r t i d a l zone where i t i s r e g u l a r l y inundated. Saltbush may range over s i l t y sand to peaty s o i l s . It i s a subdominant on the Cowichan R i v e r Estuary, and a dominant on the Nanaimo and Kokish R i v e r s ' E s t u a r i e s . Barbarea o r t h o c e r a s Ledeb. ( w i n t e r c r e s s ) i n t r o d u c e d herb Winte r c r e s s i s a dominant on the Cowichan R i v e r E s t u a r y . I t s presence suggests a d i s t u r b e d s i t e such as the s u b s t r a t e sloughed from a dyke on the e s t u a r y . It occurs on w e l l - d r a i n e d s u b s t r a t e s which are not inundated. Other s p e c i e s , a l s o i n d i c a t i v e of dry, w e l l - d r a i n e d areas, which occur with w i n t e r c r e s s i n c l u d e ; Nootka rose, evergreen b l a c k b e r r y , white c l o v e r , Canada t h i s t l e and sour weed. Bromus c a r i n a t u s H. & A. (brome-grass) nat i v e Brome g e n e r a l l y occurs i n the backshore, away from major channels. Here i t i s inundated f o r r e l a t i v e l y short p e r i o d s of time and then probably by only r e l a t i v e l y s l i g h t l y b r a c k i s h freshwater. On the Salmon R i v e r Estuary, where i t i s a dominant, i t may be exposed from 85 to 90 percent of the d a y l i g h t hours dur i n g the growing season (Kennedy, 1978). It occurs on b e t t e r d r a i n e d s u b s t r a t e s with Kentucky b l u e g r a s s , white c l o v e r , r i b g r a s s and h a i r y c a t s - e a r . - 85 -Bromus m o l l i s L. ( s o f t brome-grass) i n t r o d u c e d S o f t brome i s a subdominant on the L i t t l e Qualicum River Estuary. It may be found on w e l l - d r a i n e d f i n e g r a v e l s u b s t r a t e behind dykes. I t i s not inundated but may be s u b j e c t to s a l t spray. It i s a s s o c i a t e d with other s p e c i e s commonly found on such dry s i t e s , i n c l u d i n g ; Nootka ro~e and sweet v e r n a l g r a s s . Carex l y n g b y e i Hornem. (Lyngby's sedge) nati v e Lyngby's sedge has a high frequency of occurrence on the nineteen e s t u a r i e s s t u d i e d being a dominant on a l l and a subdominant on eleven of the e s t u a r i e s . It may a l s o be present i n o ther e s t u a r i n e communities. It i s g e n e r a l l y found at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone where i t has a low exposure, high i n u n d a t i o n r a t i o . On the Salmon River Estuary, Lyngby's sedge occurs between .5 and 1.0 meter i n e l e v a t i o n where i t may be exposed from 39 to 62 percent of the d a y l i g h t hours during the growing season (Kennedy, 1978). At the lower e l e v a t i o n s of i t s d i s t r i b u t i o n Lyngby's sedge grows i n anaerobic, water s a t u r a t e d s o i l . T h i s i s p o s s i b l e due to the t r a n s p o r t of a i r from above to below ground organs where i t forms a l a y e r around the r o o t s and rhizomes reducing the anaerobic c o n d i t i o n s at the organ's s u r f a c e . As the r e l a t i v e e l e v a t i o n i n c r e a s e s , moving towards the backshore, the s u b s t r a t e becomes b e t t e r d r a i n e d , at l e a s t the f i r s t few c e n t i m e t e r s , and the number and frequency of p l a n t s p e c i e s o c c u r i n g with Lyngby's sedge i n c r e a s e . The s p e c i e s a s s o c i a t e d with i t tend to be s t o l o n i f e r o u s (eg. c i n q u e f o i l ) or shallow, f i b r o u s r o o t i n g s p e c i e s (eg. sea-milkwort) and use the upper few centimeters of r e l a t i v e l y b e t t e r d r a i n e d s u b s t r a t e . As s i t e c o n d i t i o n s vary - 86 -a s s o c i a t e d s p e c i e s vary. At s i t e s tending towards higher s a l i n i t e s Lyngby's sedge may occur with s a l t g r a s s or s a l t w o r t and on s i t e s with a r e l a t i v e l y b e t t e r d r a i n e d s u b s t r a t e i t may occur with t u f t e d h a i r g r a s s . In g e n e r a l , Lyngby's sedge i s i n d i c a t i v e of b r a c k i s h water c o n d i t i o n s and i t u s u a l l y occurs at the lowest vegetated e l e v a t i o n and along major channels where i t i s f l u s h e d by b r a c k i s h water. Fur t h e r study i s r e q u i r e d to determine i f Lyngby's sedge g e n e r a l l y l i n e s major channels because i t r e q u i r e s f l u s h i n g by b r a c k i s h freshwater or because channel l e v e e s are of an optimal e l e v a t i o n a l range and c l o s e to the modifying i n f l u e n c e of freshwater. Carex obnupta B a i l e y (slough sedge) n a t i v e Slough sedge i s a dominant on the Nimpkish R i v e r E s t u a r y . It occurs i n the more e l e v a t e d areas of the estuary,., u s u a l l y i n water s a t u r a t e d s o i l s , where i t i s inundated f o r short p e r i o d s of time by b r a c k i s h water. Carex r o s t r a t a Stokes (beaked sedge) n a t i v e Beaked sedge i s a dominant on the Cluxewe R i v e r E s t u a r y . It occurs with Lyngby's sedge i n a pool of running water adjacent to the t r e e l i n e . It i s i n d i c a t i v e of freshwater c o n d i t i o n s but i s t o l e r a n t of s l i g h t l y b r a c k i s h c o n d i t i o n s . Chenopodium album L. (pigweed) in t r o d u c e d herb from north temperate E u r a s i a Pigweed i s a subdominant on the g r a v e l f i l l i n f r o n t of the Nature House on the Goldstream Estuary. It i s o f t e n found on dry, w e l l - d r a i n e d , d i s t u r b e d s i t e s such as around the Nature - 87 -House. A s s o c i a t e d s p e c i e s are a l s o those of d i s t u r b e d s i t e s , i n c l u d i n g : willow dock, r i b g r a s s , broom, h a i r y c a t s - e a r , common t h i s t l e and Canada t h i s t l e . C i r s i u m arvense (L.) Scop. (Canada t h i s t l e ) i n t r o d u c e d herb from northern E u r a s i a Canada t h i s t l e , a s t r o n g l y rhizomatous s p e c i e s , i s a s p e c i e s of d i s t u r b e d s i t e s . On the Englishman R i v e r Estuary, where i t i s a subdominant, i t may be found i n the backshore and w i t h i n a dyked area. The backshore appears to have been a l t e r e d by human a c t i v i t y , p o s s i b l y dyking and/or log h a n d l i n g . It i s e x t e n s i v e l y d i s t r i b u t e d i n the dyked area because the area i s r e l a t i v e l y w e l l d r a i n e d and not t i d a l l y inundated. The presence of Nootka rose, e l d e r b e r r y , evergreen b l a c k b e r r y , broom, mil k -t h i s t l e , g r o u n d s e l , common t h i s t l e , v e l v e t - g r a s s and cheat with Canada t h i s t l e suggests the area i s tending towards " o l d f i e l d " communities. It i s l i k e l y that t h i s area would have been fl o o d e d b e f o r e the dyke was c o n s t r u c t e d , as i n d i c a t e d by the channels i n the dyked area and the f l o o d i n g of comparable, adjacent land o u t s i d e the dyke. Without the dyke Canada t h i s t l e would not be a subdominant i n t h i s a r ea. C y t i s u s s c o p a r i u s (L.) Lin k (broom) int r o d u c e d shrub Broom, an in t r o d u c e d s p e c i e s that d i s p l a c e s n a t i v e s p e c i e s , i s a subdominant and/or a dominant on the Goldstream, Chemainus, Nanaimo, Englishman and Campbell R i v e r s ' E s t u a r i e s . It i s o f t e n d i s t r i b u t e d through man's a c t i v i t i e s and may become e s t a b l i s h e d i n d i s t u r b e d areas, or along dykes, r a i l w a y s and roads, or around b u i l d i n g s on e s t u a r i e s . Broom occurs on - 88 -r e l a t i v e l y w e l l - d r a i n e d s u b s t r a t e that i s not inundated. It has some t o l e r a n c e of s a l t spray. D a c t y l i s glomerata L. ( o r c h a r d - g r a s s ) introduced from northern Europe c u l t i v a r Orchard-grass i s a dominant on the Salmon R i v e r Estuary and a subdominant on the Cluxewe R i v e r Estuary. It i s most often found toward the backshore where the s u b s t r a t e i s r e l a t i v e l y w e l l - d r a i n e d and i n u n d a t i o n i s probably shallow, f o r a short p e r i o d of time and of a more freshwater c o n d i t i o n . Deschampsia c e s p i t o s a (L.) Beauv. ( t u f t e d h a i r g r a s s ) native T u f t e d h a i r g r a s s , a circumboreal s p e c i e s , i s a dominant and/or a subdominant on twelve of the e s t u a r i e s s t u d i e d , i n c l u d i n g ; the Cowichan, Nanaimo, Courtenay, Campbell, Salmon, Adam-Eve, T s i t i k a , Kokish, Nimpkish, Cluxewe, Quatse and Kingcome R i v e r s ' E s t u a r i e s . The frequency of occurrence of t u f t e d h a i r g r a s s appears to be g r e a t e r i n e s t u a r i e s north of Courtenay and on the Kingcome R i v e r E s t u a r y . The f i b r o u s rooted grass grows i n dense, low t u f t s on r e l a t i v e l y w e l l - d r a i n e d s i l t e d sands and g r a y e l s , and may be inundated on each t i d e by b r a c k i s h water. The f i b r o u s root system i s b e t t e r s u i t e d than a rhizomatous ro o t system f o r growth i n c o a r s e r s u b s t r a t e s . On the T s i t i k a R i v e r Estuary, t u f t e d h a i r g r a s s i s the dominant s p e c i e s on two to four i n c h g r a v e l while the rhizomatous Lyngby's sedge i s the dominant s p e c i e s on an adjacent sand s u b s t r a t e . Another f a c t o r p o s s i b l y i n f l u e n c i n g the d i s t r i b u t i o n of t u f t e d h a i r g r a s s i s suggested by the simple l i n e a r c o r r e l a t i o n a n a l y s i s . The c o r r e l a t i o n c o e f f i c i e n t between the - 89 -t o t a l area on which tufted hairgrass i s either a dominant or a subdominant and the A p r i l to September mean r a i n f a l l for each estuary was s i g n i f i c a n t ; r=.7606 r 0.05(2),6 = ° « 7 0 7 (n=8) (Table 45). The interaction may be related to growth habit and p r e c i p i t a t i o n . The caespitose nature of the plant directs rain to the root system where i t may decrease the s a l i n i t y around the roots and hence increase vigor. The interaction of anatomical features, substra+e and p r e c i p i t a t i o n and their influence on the d i s t r i b u t i o n of tufted hairgrass requires further study. D i s t i c h l i s spicata (L.) Greene (saltgrass) native Saltgrass i s a subdominant and/or a dominant on the Goldstream, Cowichan, Chemainus, Nanaimo, Nanoose-Bonell, Englishman, L i t t l e Qualicum, Oyster, Salmon and Kokish Rivers' Estuaries. It has a higher frequency of occurrence on estuaries south of Courtenay where flows are low during the growing season (eg. Nanoose-Bonell) and .in estuarine areas where freshwater flow i s r e s t r i c t e d , for example, by dykes (eg. Cowichan). It is usually inundated on each tide and i s i n d i c a t i v e of higher s a l i n i t i e s though i t may occur in brackish conditions. Associated species may r e f l e c t variations in s i t e conditions. For example, on the Chemainus River Estuary, the D i s t i c h l i s  spicata - S a l i c o r n i a v i r g i n i c a Community i s found on a r e l a t i v e l y wet, poorly drained, peaty s o i l and the D i s t i c h l i s  spicata - G r i n d e l i a i n t e g r i f o l i a Community i s found on a better drained s o i l . - 90 -E l e o c h a r i s p a l u s t r i s (L.) R. & S. (Spike-rush) n a t i v e Spike-rush i s a subdominant and/or a dominant on the L i t t l e Qualicum, Courtenay, Campbell, Salmon and Nimpkish R i v e r s ' E s t u a r i e s . I t i s g e n e r a l l y found at r e l a t i v e l y high e l e v a t i o n s toward the backshore where i t i s s h a l l o w l y inundated on each t i d e . I t i s a br a c k i s h freshwater s p e c i e s . Elymus glaucus Buck, (blue w i l d r e) native Blue w i l d r y e i s a dominant on the Salmon Ri v e r Estuary. I t i s prominent i n the l a t t e r h a l f of the growing season r e l a t i v e l y w e l l - d r a i n e d s i t e s that are inundated f o r r e l a t i v e l y short periods of time by bra c k i s h freshwater. Elymus m o l l i s T r i n . ( w i l d r y e ) n a t i v e grass Wildrye i s a dominant on the Cowichan, Nanoose-Bonell, Englishman, L i t t l e Qualicum, Oyster, Salmon, Adam-Eve, T s i t i k a , Kokish, Nimpkish, Cluxewe and Kingcome R i v e r s ' E s t u a r i e s . I t i s a subdominant on the Courtenay, Cluxe~e and Kingcome Rivers' E s t u a r i e s . Wildrye i s g e n e r a l l y found toward the backshore, adjacent to t r e e l i n e s , on beach bermes'or elevated areas, such as wood debris p i l e s . The substrate i s u s u a l l y w e l l drained and very sandy. Wildrye i s i n u n d a t e d i n f r e q u e n t l y and t o l e r a n t of s a l t spray. Associated species i n d i c a t e a tendency toward more t e r r e s t r i a l h a b i t a t . These as s o c i a t e d species i n c l u d e S i t k a spruce, e l d e r b e r r y , salmonberry, w i l l o w dock and reed canarygrass. - 9 1 -Epilobium a n g u s t i f o l i u m L. (fireweed) n a t i v e herb Fireweed i s a dominant i n a dyked area of the Englishman Riv e r Estuary. The area i s not inundated and i s r e l a t i v e l y w e l l drained compared to adjacent, undyked land. The dominance of fireweed, a t e r r e s t r i a l , pioneer s p e c i e s , suggests the dyked area i s becoming c o l o n i z e d by t e r r e s t r i a l p l a n t s p e c i e s . F r i t i l l a r i a camschatcensis ( L . ) Ker.-Oawl. (chocolate l i l y ) n a t i v e Chocolate l i l y may be found on a number of the e s t u a r i e s s t u d i e d but i t i s a subdominant on the Salmon River Estuary. I t occurs on r e l a t i v e l y elevated, w e l l - d r a i n e d s u b s t r a t e s where i t i s emergent from 85 to 90 percent of the d a y l i g h t hours during the growing season (Kennedy, 1978). I t s higher frequency of occurrence on the Salmon River Estuary i s probably due to the abundance of r e l a t i v e l y elevated and w e l l - d r a i n e d areas. G a u l t h e r i a s h a l l o n Pursh ( s a l a l ) n a t i v e shrub S a l a l i s dominant i n the backshore area between a creek and s l a s h burn on the Quatse River Estuary. S a l a l i s an understory species of c o a s t a l f o r e s t s and f o r e s t margins. P o s s i b l y the G a u l t h e r i a s h a l l o n community c o l o n i s e s the area because i t i s r e l a t i v e l y w e l l drained and not inundated. Claux maritima L. (sea-nilkv/ort) n a t i v e herb Sea-milkwort i s a dominant on the Cowichan, Englishman and Salmon R i v e r s ' E s t u a r i e s and a subdominant on the Cowichan, Nanaimo and Adam-Eve R i v e r s ' E s t u a r i e s . I t i s g e n e r a l l y found - 92 -at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone where, on the Salmon R i v e r Estuary, i t may be exposed f o r 39 percent of the d a y l i g h t hours during the growing season (Kennedy, 1978). I t i s found i n b r a c k i s h to s a l i n e water c o n d i t i o n s as suggested by a s s o c i a t e d s p e c i e s such as Lyngby's sedge, an i n d i c a t o r of b r a c k i s h water c o n d i t i o n s , and s a l t w o r t , an i n d i c a t o r of s a l i n e water c o n d i t i o n s . It occurs on sandy and sandy g r a v e l s u b s t r a t e s . G l y c e r i a o c c i d e n t a l i s ( P i p e r ) N e l s . (western mannagrass) nati v e grass Western mannagrass i s a dominant on the Courtenay River E s t u a r y . I t occurs towards the backshore where i t i s i n f r e q u e n t l y inundated by freshwater. The a s s o c i a t e d s p e c i e s are t e r r e s t r i a l p l a n t s and i n c l u d e h a i r y c a t s - e a r , Canada t h i s t l e , miner's l e t t u c e , common h o r s e t a i l , f i e l d mint and meadow fesc u e . G r i n d e l i a i n t e g r i f o l i a DC. (gumweed) na t i v e herb Gumweed i s a dominant on the Chemainus, Nanoose-Bonell, Englishman and Salmon R i v e r s ' E s t u a r i e s and a subdominant on the Nanaimo and Englishman R i v e r s ' E s t u a r i e s . .It may be inundated r e g u l a r l y by s a l i n e to b r a c k i s h water. The s u b s t r a t e i s r e l a t i v e l y w e l l d r a i n e d and s a l i n e . Heracleum lanatum Michx. (cow-parsnip) n a t i v e herb Cow-parsnip i s a dominant on the L i t t l e Qualicum, Salmon, T s i t i k a and Kokish R i v e r s ' E s t u a r i e - and a subdominant on the Cluxewe and Kingcome R i v e r s ' E s t u a r i e s . It occurs i n the - 9 3 -ecotone between marsh and f o r e s t , or beach berme and f o r e s t , and i s a species of a damp to wet s u b s t r a t e . I t i s infrequently-inundated and then by freshwater. Holcus lanatus L. ( v e l v e t - g r a s s ) introduced from northern Europe Velvet-grass i s a dominant on the Englishman and Quatse Rivers' e s t u a r i e s and a subdominant on the Cowichan and T s i t i k a R i v e r s ' E s t u a r i e s . I t i s a freshwater species that i s i n f r e q u e n t l y inundated, and grows on r e l a t i v e l y w e l l drained s u b s t r a t e s . On the T s i t i k a River Estuary i t occurs i n the backshore, adjacent to the f o r e s t . On the Cowichan, Englishman and Quatse R i v e r s ' E s t u a r i e s i t occurs w i t h i n dyked areas and on l a n d f i l l . Associated species i n c l u d e c o l o n i z e r s of d i s t u r b e d s i t e s . Hordeum brachyantherum Nevski (meadow barl e y ) native grass Meadow barley i s a dominant and a subdominant on the Goldstream and Kokish R i v e r E s t u a r i e s , and a dominant on the Salmon, Adam-Eve and Cluxewe R i v e r s ' E s t u a r i e s . I t may be s h a l l o w l y inundated by b r a c k i s h to s a l i n e water as i n d i c a t e d by associated species such as t u f t e d h a i r g r a s s , an i n d i c a t o r of b r a c k i s h c o n d i t i o n s , and s a l t w o r t , an i n d i c a t o r of s a l i n e c o n d i t i o n s . Hordeum murinum L. (mouse barl e y ) native grass Mouse b a r l e y i s a dominant on the Goldstream Estuary and a subdominant on the Englishman River Estuary. I t i s a b r a c k i s h water species but appears to t o l e r a t e r e l a t i v e l y high s a l i n i t i e s - 94 -as i n d i c a t e d by the a s s o c i a t e d s p e c i e s s a l t w o r t , s a l t b u s h and s a l t g r a s s . Mouse b a r l e y i s u s u a l l y found on a wet s u b s t r a t e and i s inundated r e g u l a r l y by b r a c k i s h to s a l i n e water. Juncus a r t i c u l a t u s L. ( j o i n t e d rush) n a t i v e On the Nanaimo R i v e r E s t u a r y , j o i n t e d rush i s a dominant and a subdominant i n 50.7 percent of the emergent v e g e t a t i o n . It r a r e l y occurs on the other e s t u a r i e s s t u d i e d . B a l t i c rush, which may have a high frequency of occurrence on other e s t u a r i e s , o n l y covers 4.8 percent of the emergent v e g e t a t i o n on the Nanaimo R i v e r E s t u a r y . The reasons f o r the dominance of j o i n t e d rush on t h i s e s t u a r y are not c l e a r though previous a g r i c u l t u r a l p r a c t i s e s may have i n f l u e n c e d i t s d i s t r i b u t i o n . It occurs i n dense stands and i s e x t e n s i v e on the e s t u a r y . The s u b s t r a t e i s wet. P o s s i b l y o l d dykes and very shallow depressions i n the areas e s t a b l i s h e d to t h i s s p e c i e s impede drain a g e . The a s s o c i a t e d s p e c i e s i n d i c a t e j o i n t e d rush occurs i n b r a c k i s h to s a l i n e c o n d i t i o n s . A s s o c i a t e d s p e c i e s i n d i c a t i n g b r a c k i s h c o n d i t i o n s are Lyngby's sedge, t u f t e d hairgra<=~ and c i n q u e f o i l . A s s o c i a t e d s p e c i e s i n d i c a t i n g s a l i n e c o n d i t i o n s are s a l t w o r t , s a l t b u s h and s a l t g r a s s . Juncus b a l t i c u s W i l l d . ( B a l t i c rush) native B a l t i c rush i s a subdominant on the Kingcome R i v e r Estuary, a dominant on the Goldstream, Chemainus, Nanoose-B o n e l l , Courtenay, Salmon and Quatse R i v e r s ' E s t u a r i e s , and a subdominant and a dominant on the Cowichan, Nanaimo, Englishman, L i t t l e Qualicum, Oyster and Nimpkish R i v e r s ' E s t u a r i e s . It has a high frequency of occurrence on e s t u a r i e s south of Courtenay. - 9 5 -It i s , g e n e r a l l y , r e g u l a r l y and s h a l l o w l y inundated by b r a c k i s h water and the s u b s t r a t e i s r e l a t i v e l y w e l l d r a i n e d at low t i d e . The p i t h y shoots of B a l t i c rush suggest anatomical and p h y s i o l o g i c a l a d a p t a t i o n to a s p e c i a l environment. Juncus e f f u s u s L. (common rush) n a t i v e , circumboreal Common rush i s u s u a l l y c o n s i d e r e d to be a freshwater marsh s p e c i e s or a s p e c i e s common to the t r a n s i t i o n zone between marsh and c o a s t a l f o r e s t . The s u b s t r a t e may appear w e l l d r a i n e d , though there i s probably subsurface seepage, or the subs t r a t e may be wet to s a t u r a t e d . Common rush i s seldom inundated. It i s a dominant on the Cluxewe R i v e r Estuary but a l s o occurs on other e s t u a r i e s . There appear to be two p o s s i b l e reasons why common rush i s not abundant. F i r s t , the l i m i t e d area of the t r a n s i t i o n zone between a marsh and c o a s t a l f o r e s t on most of the e s t u a r i e s s t u d i e d , and second, the development of such zones. Juncus e n s i f o l i u s v a r . e n s i f o l i u s Wikst. (dagger-leaf rush) native Dagger-leaf rush i s a subdominant on the Quatse R i v e r E s t u a r y . Here i t occurs with c o l o n i a l bentgrass on a ' " e l l -drained s u b s t r a t e behind a dyke. It i s probably not inundated. Juncus g e r a r d i L o i s e l . (mud rush) n a t i v e Mud rush i s subdominant on the Adam-Eve R i v e r E s t u a r y . It occurs toward the backshore on a r e l a t i v e l y w e l l - d r a i n e d s u b s t r a t e . It i s s h a l l o w l y and s h o r t l y inundated by s a l i n e to - 96 -b r a c k i s h water. Two a s s o c i a t e d s p e c i e s i n d i c a t i v e of s a l i n e c o n d i t i o n s are s a l t w o r t and s a l t b u s h . J u n i p e r u s scopulorum Sarg. ( j u n i p e r ) n a t i v e tree J u n i p e r i s a dominant on the Chemainus R i v e r E s t u a r y . It i s not known to have a wide d i s t r i b u t i o n on the east coast of Vancouver I s l a n d and grows widely d i s p e r s e d . It p r e f e r s a d r i e r c l i m a t e , and w e l l d r a i n e d s u b s t r a t e s , c o n d i t i o n s which occur on the Shoal Islands and i n the backshore of the Chemainus R i v e r E s t u a r y . Myrica g a l e L. (sweet ga l e ) n a t i v e shrub Sweet g a l e , a s p e c i e s p o s s e s s i n g nodules f i x i n g atmospheric n i t r o g e n s y m b i o t i c a l l y , i s a dominant on the Campbell and Nimpkish R i v e r s ' E s t u a r i e s where i t occurs i n freshwater marshes adjacent to the backshore f o r e s t s . On the Campbell ftiver E stuary the marsh has much standing water; a s s o c i a t e d o l i g o t r o p h i c s p e c i e s i n c l u d e hardstem bulr u s h and h o r s e t a i l . On the Nimpkish R i v e r Estuary the marsh appears to be d r i e r with l e s s standing wa*er and a s s o c i a t e d s p e c i e s , i n d i c a t i v e of more mesic c o n d i t i o n s , i n c l u d e salmonberry, s e l f -h e a l , c r e e p i n g b u t t e r c u p , small bedstraw and water p a r s l e y . Oenanthe sarmentosa P r e s l . (water-parsley) herb Water-parsley i s a subdominant on the Courtenay and Quatse R i v e r s ' E s t u a r i e s . It occurs i n the backshore where i t may be i n f r e q u e n t l y inundated by b r a c k i s h freshwater. It may a l s o occur on other e s t u a r i e s on r e l a t i v e l y w e l l d r a i n e d s u b s t r a t e s where i t i s r e g u l a r l y , s h a l l o w l y and s h o r t l y - 97 -inundated. A s s o c i a t e d s p e c i e s h o r s e t a i l , c u r l y , d o c k , yarrow, sedge, b l a c k twin-berry and Pac i n c l u d e morning-glory, common bedstraw, meadow fescue, Lyngby's i f i c c r a b a p p l e . P a c h i s t i m a m y r s i n i t e s (Pursh) Raf. ( f a l s e box) n a t i v e shrub P a c h i s t i m a i s a dominant on the Shoal I s l a n d s i n the Chemainus R i v e r E s t u a r y . It occurs -in d r i e r c l i m a t e s and on well d r a i n e d s u b s t r a t e s . P h a l a r i s arundinaceae L. (reed canarygrass) n a t i v e and introduced Reed canarygrass i s a dominant on the Nanoose-Bonell Creeks' E s t u a r and a subdominant on the Courtenay River Estuary. It i s used in a g r i c u l t u r e as a hay crop and has probably escaped to the backshores of many of the e s t u a r i e s . It i s u s u a l l y inundated by freshwater i n winter but i s not very s a l t t o l e r a n t . The a s s o c i a t e d s p e c i e s tend to be mesic and i n c l u d e a l d e r , willow, Nootka rose, evergreen b l a c k b e r r y , Canada t h i s t l e , miner's l e t t u c e , rough bedstraw, white sweet c l o v e r , pearly e v e r l a s t i n g , water-hemlock, timothy and o r c h a r d - g r a s s . P i c e a s i t c h e n s i s (Bong.) C a r r . ( S i t k a spruce) n a t i v e t r e e S i t k a spruce i s a dominant on the Courtenay, Salmon, Adam-Eve, T s i t i k a , Nimpkish and Cluxewe R i v e r s ' E s t u a r i e s where i t occurs i n the backshore. It forms a s e r a i wet s u b s t r a t e stage i n f o r e s t s u c c e s s i o n to the climax Tsuga h e t e r o p h y l l a community f o r example, on the Salmon and Adam-Eve R i v e r s ' E s t u a r i e s . S i t k a spruce i s o f t e n found on d e l t a i c f l o o d p l a i n s i n the western hemlock b i o g e o c l i m a t i c zone where i t may grow on n u t r i e n t r i c h , - 98 -seepage s u b s t r a t e s . I t i s the s l i g h t l y s a l i n e c o n d i t i o n s and magnesium from the ocean spray ( K r a j i n a , 19G9/1970). only c o n i f e r that t o l e r a t e s i s thought to be able to use and b r a c k i s h t i d a l v/ater Plantago l a n c e o l a t a L. ( r i b g r a s s ) introduced herb from northwest Europe Plantago macrocarpa Cham. & Schlecht ( p l a n t a i n ) n a t i v e herb Ribgrass, an a l i e n s p e c i e s , has become a dominant and/or subdominant on the Goldstream, Big Qualicum, Salmon and Cluxewe R i v e r s ' E s t u a r i e s and p l a n t a i n , a n a t i v e , i s a dominant on the Eokish River Estuary. Ribgrass and p l a n t a i n tend to occur more f r e q u e n t l y toward the backshore, away from main channels, where they are s h o r t l y and s h a l l o w l y inundated by b r a c k i s h freshwater. The s u b s t r a t e i s r e l a t i v e l y porous and coarse. Plantago maritima L. (seaside p l a n t a i n ) n a t i v e herb Seaside p l a n t a i n i s a dominant on the Nanoose-Bonell, Englishman and Quatse R i v e r s ' E s t u a r i e s and a subdominant on the Adam-Eve and Kokish R i v e r s ' E s t u a r i e s . It i s found at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone where i t i s inundated during each f l o o d t i d e . It i s i n d i c a t i v e of s a l i n e to b r a c k i s h c o n d i t i o n s ; associated s a l t t o l e r a n t species are s a l t w o r t , s a l t b u s h , s a l t g r a s s , arrow-grass and gumweed. It may be found on sand to g r a v e l s u b s t r a t e s . Poa compressa L. (Canada bluegrass) introduced Canada bluegrass i s a subdominant species on the Cowichan - 99 -River E s t u a r y . It may be found where conditions, are b r a c k i s h and the s u b s t r a t e i s r e l a t i v e l y w e l l d r a i n e d . It i s probably inundated by the higher t i d e s . Poa p r a t e n s i s L. (Kentucky b l u e g r a s s ) introduced Kentucky b l u e g r a s s i s a dominant and/or a subdominant on the Englishman, Salmon, Adam-Eve and Cluxewe R i v e r s ' E s t u a r i e s . It tends to occur where the s u b s t r a t e i s w e l l d r a i n e d , c o n d i t i o n s are b r a c k i s h , and i n u n d a t i o n i s short and shallow, c o n d i t i o n s which appear to occur more f r e q u e n t l y i n northern e s t u a r i e s . Mesic s p e c i e s such as black twin-berry, P a c i f i c crabapple, salmonberry and Nootka rose are a s s o c i a t e d with Kentucky b l u e g r a s s towards the backshore, and f o r b s such as c i n q u e f o i l , springback c l o v e r , yarrow, t u f t e d h a i r g r a s s and c o l o n i a l bentgrass are a s s o c i a t e d with Kentucky bluegrass towards the f o r e s h o r e . P o t e n t i l l a p a c i f i c a Howell ( c i n q u e f o i l ) n a t i v e herb C i n q u e f o i l i s present on a l l the e s t u a r i e s s t u d i e d and i s a dominant and/or a subdominant on the Nanoo e - B o n e l l , Englishman, L i t t l e Qualicum, Big Qualicum, Courtenay, Oyster, Campbell, Salmon, Adam-Eve, Kokish, Nimpkish, Cluxewe, Quatse and Kingcome R i v e r s ' E s t u a r i e s . It i s a b r a c k i s h freshwater s p e c i e - and may be found on s i l t e d sand to g r a v e l from the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone to the backshore. T h i s broad d i s t r i b u t i o n exposes i t to a range of inundation, s a l i n i t y , s u b s t r a t e and drainage regimes. At lower e l e v a t i o n s i t may be inundated on each t i d e , and s u b j e c t to higher - 100 -s a l i n i t i e s and poor drainage. At higher e l e v a t i o n s i t may be inundated only s h o r t l y and s h a l l o w l y , and s u b j e c t to g e n e r a l l y lower s a l i n i t i e s and b e t t e r drainage. C i n q u e f o i l ' s broad d i s t r i b u t i o n over a wide range of h a b i t a t s may be r e l a t e d to i t s s t o l o n i f e r o u s h a b i t . The shallow r o o t i n g s t o l o n s root i n the f i r s t few centimeters of almost water s a t u r a t e d s o i l i n the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone. They are a l s o adapted to c o l o n i z i n g d r i e r g r a v e l s where the roots can penetrate between rocks and s t o l o n s b r i d g e r o c k s . A simple l i n e a r c o r r e l a t i o n between the t o t a l area on which c i n q u e f o i l i s e i t h e r a dominant or a subdominant and the A p r i l to September mean r a i n f a l l f o r each estuary suggests c i n q u e f o i l i s more abundant where p r e c i p i t a t i o n i s g r e a t e r ( r = .9217 r 0 . 0 5 ( 2 ) , 6 = 0»707 (n = 8 ) ) ( T a b l e 45). T h i s i s probably r e l a t e d to p r e c i p i t a t i o n reducing the s a l i n i t y a r ound the shallow r o o t i n g s t o l o n s . T h i s e f f e c t i s regarded as a f a c t o r i n the d i s t r i b u t i o n of c i n q u e f o i l , p a r t i c u l a r l y towards the backshore, i n e s t u a r i e s north of Courtenay. P t e r i d i u m aquilinum (L.) Kuhn (bracken) n a t i v e f e r n Bracken, a mesic s p e c i e s , i s a dominant i n the ecotone between-marsh and f o r e s t on the Oyster R i v e r Estuary. It occurs on r e l a t i v e l y w e l l d r a i n e d s u b s t r a t e and i s not inundated. A s s o c i a t e d s p e c i e s i n c l u d e f a l s e l i l y - o f - t h e - v a l l e y , marsh pea, sword f e r n , foamflower and w a l l l e t t u c e . Pseudotsuga m e n z i e s i i ( M i r b e l ) France (Douglas f i r ) n a t i v e t r e e Douglas f i r i s a dominant on the Chemainus, Nanaimo and - 101 -Oyster R i v e r s ' E s t u a r i e s . It i s found i n the backshore and on i s l a n d s i n the e s t u a r i e s where i t i s probably never f l o o d e d . The best growth of Douglas f i r occurs when the s u b s t r a t e i s moderately w e l l d r a i n e d , and the s o i l n u t r i e n t regime i s moderate but r i c h i n bases such as calcium and magnesium ( K r a j i n a , ' 1969/1970). A s s o c i a t e d s p e c i e s , p a r t i c u l a r l y Garry oak, on the Chemainus and Nanaimo R i v e r s ' E s t u a r i e s r e f l e c t d r i e r growing c o n d i t i o n s than on, f o r example, the Oyster River Estuary to the n o r t h . Pyrus f u s c a Raf. ( P a c i f i c crabapple) n a t i v e tree P a c i f i c crabapple, a small t r e e , i s a dominant on the L i t t l e Qualicum and Kokish R i v e r ' s E s t u a r i e - and a subdominant on the Chemainus and L i t t l e Qualicum R i v e r s ' E s t u a r i e s . It occurs on dykes and i n the backshore with more mesic species such as Garry oak, c a s c a r a , oregongrape, Douglas f i r , broom, s a l a l , t h i m b l e b e r r y and salmonberry. I n f r e q u e n t l y , i t may be s h o r t l y and s h a l l o w l y inundated with b r a c k i s h freshwater; i n t h i s case the s u b s t r a t e i s r e l a t i v e l y e l l d r a i n e d . Quercus garryana Dougl. (Garry oak) n a t i v e tree Garry oak i s a subdominant on the Chemainus R i v e r Estuary and a dominant on the Nanaimo R i v e r Estuary. I t grows at the foot of mountains i n areas of low annual t o t a l p r e c i p i t a t i o n (660 - 1020 mm). I t s f r o s t r e s i s t a n c e and shade t o l e r a n c e are low, and i t s n u t r i t i o n a l requirements are high, p a r t i c u l a r l y with regard to bases such as calcium and magnesium ( K r a j i n a , 1969/1970). Ribes sp. L. (gooseberry) n a t i v e shrub Gooseberry i s a dominant on the Nimpkish R i v e r Estuary where i t i s a s s o c i a t e d with S i t k a spruce, Nootka rose, red a l d e r , salmonberry, slough sedge and skunk cabbage. It i s i n f r e q u e n t l y f l o o d e d and then only s h o r t l y and s h a l l o w l y by b r a c k i s h freshwater, but s o i l moisture i s u s u a l l y maintained at high l e v e l s c o n t i n u o u s l y from groundwater sources or heavy frequent p r e c i p i t a t i o n . Rosa nutkana P r e s l . (Nootka rose) n a t i v e shrub Nootka rose i s a dominant and/or a subdominant on the Goldstream, Chemainus, Nanoose-Bonell, Englishman, L i t t l e Qualicum, Campbell, Salmon and Nimpkish R i v e r s ' E s t u a r i e s . It has a high frequency of occurrence i n e s t u a r i e s south of Courtenay. T h i s i s r e l a t e d to the higher a r i d i t y index south of Courtenay. Nootka rose occurs i n the backshore where the substrate i s d r i e r and may be s i l t e d sand to small g r a v e l . It i s i n f r e q u e n t l y or never inundated but appears to have some to l e r a n c e to s a l t spray. Rubus spp. L. ( b l a c k b e r r y , salmonberry) n a t i v e shrub Rubus spp. are dominants and/or subdominants on the Cowichan, Nanoose-Bonell, Englishman, L i t t l e Qualicum, Campbell, Nimpkish, Quatse and Kingcome R i v e r s ' E s t u a r i e s . B l a c k b e r r i e s tend to occur more f r e q u e n t l y on dykes and i n the backshore on the more southern e s t u a r i e s s t u d i e d and salmonberry tends to occur more f r e q u e n t l y i n the ecotone between marsh and f o r e s t on the more northern e s t u a r i e s . They may be found on moister or - 1 0 3 -seepage s u b s t r a t e s and are never, or very i n f r e q u e n t l y , s h o r t l y and.shallowly f l o o d e d . They appear to t o l e r a t e s a l t spray. Rumex s a l i c i f o l i u s Weinm. (willow dock) herb Willow dock i s a subdominant on the Goldstream and Courtenay R i v e r s ' E s t u a r i e s where i t i s found on d r i e r s u b s t r a t e s . It i s probably never or i n f r e q u e n t l y inundated by freshwater. It seems to t o l e r a t e s a l t spray. Ruppia maritima L. ( d i t c h - g r a s s ) n a t i v e herb D i t c h - g r a s s , a small herb, i s a dominant on the Campbell R i v e r Estuary and i s fo~nd on some of the other e s t u a r i e s s t u d i e d . It tends to occur i n shallow, standing or slow moving b r a c k i s h water. S u i t a b l e areas f o r the growth of d i t c h - g r a s s do not appear to be abundant or ex t e n s i v e on the e s t u a r i e s s t u d i e d . S a l i c o r n i a v i r g i n i c a L. ( s a l t w o r t ) n a t i v e herb Sa l t w o r t i s a dominant and/or a subdominant on the Goldstream, Chemainus, Nanaimo, Nanoose-Bonell, Englishman, Adam-Eve, Kokish, Cluxewe, and Quatse R i v e r s ' E s t u a r i e s . It may be found at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone, i s i n d i c a t i v e of s a l i n e c o n d i t i o n s , and i s probably inundated on each t i d e . Saltwort tends to occur on s i l t e d sand, sandy s i l t or, i n f r e q u e n t l y , peaty s o i l s that are always moist or wet. The a s s o c i a t e d s p e c i e s i n c l u d e s a l t g r a s s , seaside p l a n t a i n , s a l t b u s h , salt-marsh dodder, and gumweed which are a l s o c o n s i d e r e d i n d i c a t i v e of s a l i n e c o n d i t i o n s . - 104 -S a l i x spp. L. (willow) n a t i v e shrub Willow are present on many of the e s t u a r i e s studied and are dominant on the Kingcome R i v e r E s t u a r y . They may be found i n the backshore where i t i s i n f r e q u e n t l y flooded and then by freshwater. T h e i r presence i n d i c a t e s mesic to h y d r i c t r a n s i t i o n . Sambucus racemosa L. ( e l d e r b e r r y ) n a t i v e shrub E l d e r b e r r y i s a subdominant on the Kingcome R i v e r Estuary and occurs on many of the e s t u a r i e s s t u d i e d . It i s fo nd in the backshore, i n the ecotone between marsh and f o r e s t , where i t i s never or i n f r e q u e n t l y , s h o r t l y and s h a l l o w l y flooded by freshwater. The s u b s t r a t e i s probably moderately d r a i n e d . A s s o c i a t e d s p e c i e s tend to be mesic and i n c l u d e orange honeysuckle, b l a c k twin-berry, salmonberry, f i e l d mint, cow-pa r s n i p , w i l d r y e , blue w i l d r y e and or c h a r d - g r a s s . S c i r p u s acutus Muhl. (hardstem bulrush) native' Hardstem bulru s h i s a dominant on the Cowichan and L i t t l e Qualicum R i v e r s ' E s t u a r i e s and a subdominant on the Campbell River E s t u a r y . It i s a s p e c i e s of b r a c k i s h freshwater c o n d i t i o n - and occurs i n water s a t u r a t e d , s i l t e d sand from which the water d r a i n s very slowly on the ebb t i d e . It may be s h a l l o w l y inundated on each t i d e , though on the Campbell River Estuary, i t i s probably inundated by only the highest t i d e s . The l i m i t e d d i s t r i b u t i o n of hardstem b u l r u s h , on the east coast of Vancouver I s l a n d , i s r e l a t e d to the d e l t a s ' r e l a t i v e l y steeper slopes and b e t t e r drainage as compared to, f o r example, c e r t a i n areas of the F r a s e r R i v e r D e l t a . - 105 -S c i r p u s americanus Pers. (three-square bulrush) n a t i v e Three-square b u l r u s h i s a subdominant on the Cowichan River Estuary and a dominant on the Courtenay R i v e r E s t u a r y . It occurs at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone where i t i s inundated on each f l o o d t i d e . A s p e c i e s i n d i c a t i v e of s a l i n e to b r a c k i s h c o n d i t i o n s , i t occurs on r e l a t i v e l y w e l l -drained sandy s u b s t r a t e s , g e n e r a l l y out of the d i r e c t i n f l u e n c e of the r i v e r or major channels. The l i m i t e d d i s t r i b u t i o n of three-square b u l r u s h on the east coast of Vancouver I s l a n d i s r e l a t e d to the l i m i t e d area of r e l a t i v e l y w e l l - d r a i n e d sandy substrate i n the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone. By c o n t r a s t , i t occupies l a r g e areas of the F r a s e r River d e l t a f o r e s h o r e . S c i r p u s maritimus L. (seacoast bulrush) n a t i v e Seacoast b u l r u s h i s a dominant on the Chemainus, Nanaimo and Nanoose-Bonell R i v e r s ' E s t u a r i e s . It occurs i n standing water or water s a t u r a t e d s o i l that i s g e n e r a l l y very o r g a n i c and anaerobic, s m e l l i n g s t r o n g l y of hydrogen s u l p h i d e . It i s i n d i c a t i v e of s a l i n e b r a c k i s h c o n d i t i o n s as suggested by the copresence of s a l t w o r t and Lyngby's sedge, s p e c i e s i n d i c a t i v e of s a l i n e and b r a c k i s h c o n d i t i o n s . It may be inundated on each f l o o d t i d e . The l i m i t e d d i s t r i b u t i o n of seacoast b u l r u s h , on the east coast of Vancouver I s l a n d , i s r e l a t e d to the d e l t a s ' r e l a t i v e l y s t e e p e r s l o p e s and b e t t e r drainage as compared to, f o r example, areas of the F r a s e r R i v e r D e l t a where i t i s a common. - 106 -S c i r p u s microcarpus P r e s l . ( s m a l l - f r u i t bulrush) n a t i v e S m a l l - f r u i t b u l r u s h i s a dominant i n the ecotone between marsh and f o r e s t on the Courtenay River E s t u a r y . It i s i n d i c a t i v e of areas of freshwater seepage or shallow, standing water. A s s o c i a t e d s p e c i e s i n c l u d e c i n q u e f o i l , m i l k - t h i s t l e , morning g l o r y , common h o r s e t a i l , c u r l y dock, water-parsley, common forget-me-not and c o l o n i a l bentgrass. S i d a l c e a h e n d e r s o n i i Wats. (Henderson's checker-mallow) nat i v e herb Henderson's checker-mallow i s a subdominant on the Courtenay R i v e r Estuary and i t i s a l s o present on other e s t u a r i e s s t u d i e d . It occurs on the r e l a t i v e l y more e l e v a t e d areas toward the backshore where i t may be s h o r t l y and shallowly inundated by b r a c k i s h freshwater on higher t i d e s . The s u b s t r a t e i s u s u a l l y moist. The tendency to e s t a b l i s h i n more mesic h a b i t a t s and the b r i e f n e s s of any in u n d a t i o n are i n d i c a t e d by the d i v e r s i t y of a s s o c i a t e d s p e c i e s which i n c l u d e arrow-grass, c i n q u e f o i l , springbank c l o v e r , c u r l y dock, c a t - t a i l , willow dock, St. John's-wort, camas, shooting s t a r , bedstraw, S i b e r i a n miner's l e t t u c e , blue-eyed grass, and golden I n d i a n - p a i n t b r u s h . Sonphus a r v e n s i s L. ( m i l k - t h i s t l e ) i n t r o d u c e d herb M i l k - t h i s t l e i s a dominant and a subdominant on the Englishman R i v e r Estuary. The a s s o c i a t e d s p e c i e s i n d i c a t i v e of bra c k i s h c o n d i t i o n s are c i n q u e f o i l , r i b g r a s s and couch grass, and a s s o c i a t e d s p e c i e i n d i c a t i v e of s a l i n e c o n d i t i o n s are s a l t w o r t , seaside p l a n t a i n and s a l t g r a s s . It i s probably i n f r e q u e n t l y , and then only s h o r t l y and s h a l l o w l y , inundated by b r a c k i s h or s a l t water. The s u b s t r a t e i s r e l a t i v e l y w e l l -d rained but c o n t i n u o u s l y moist. S p e r g u l a r i a canadensis (Pers.) G. Don (saltmarsh sandspurry) herb Saltmarsh sandspurry i s a dominant and a subdominant on the Nanaimo and Kokish R i v e r s ' E s t u a r i e s . I t may be found at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l zone where i t i s i n u n d a t e d by each t i d e . It i s a s p e c i e s of b r a c k i s h s a l i n e c o n d i t i o n s and tends to occur on sandy, moist s u b s t r a t e s . Symphoricarpos albus (L.) Blake (snowberry) n a t i v e shrub Snowberry i s a subdominant on the Campbell River Estuary where i t i s found i n a logged c l e a r i n g i n the backshore. The area i s probably never flooded and i s c o l o n i z e d by mesic s p e c i e s , such as c a s c a r a , a l d e r , D o u g l a s - f i r , P a c i f i c crabapple, cedar, maple, Nootka rose, salmonberry, e l d e r b e r r y , gooseberry, hardhack, orange honeysuckle and oregongrape. Thuja p l i c a t a Donn. (western red cedar) nat i v e tree Western red cedar i s a dominant i n the backshore of the Quatse R i v e r Estuary. It i s probably never flooded but, i f so, then f o r only short p e r i o d s by freshwater. It grows in n u t r i e n t r i c h , seepage s o i l s . In the C o a s t a l Western Hemlock B i o g e o c l i m a t i c Zone the best growth of western red cedar occurs on s o i l s r i c h i n bases such as calcium and magnesium ( K r a j i n a 1969/1970). - 108 -T r i f o l i u m pratense L. (red c l o v e r ) i n t r o d u c e d herb Red c l o v e r , a legume, i s a subdominant on the Big Qualicum River E s t u a r y . It tends to occur along the strearnbank at the ecotone between mars' and f o r e s t where i t may be s h o r t l y and s h a l l o w l y inundated by freshwater on the f l o o d t i d e during the growing season. The s u b s t r a t e i s g r a v e l and w e l l drained. T r i f o l i u m w o r m s k j o l d i i Lehm. (springbank c l o v e r ) n a t i v e herb Springbank c l o v e r , a n a t i v e legume, i s a dominant and/or a subdominant on the Courtenay, T s i t i k a , Nimpkish, Cluxewe and Kingcome R i v e r s ' E s t u a r i e s . It i s i n d i c a t i v e of freshwater to b r a c k i s h freshwater c o n d i t i o n s and may be inundated by each f l o o d t i d e during the growing season. It i n v a r i a b l y occurs on r e l a t i v e l y w e l l drained s u b s t r a t e s . Being shallow r o o t i n g i t may be d e s s i c a t e d i n a w e l l drained s u b s t r a t e i f i t i s not r a i n e d on or inundated r e g u l a r l y . Springbank c l o v e r appears to have in c r e a s e d frequency of occurrence i n the e s t u a r i e s north of Courtenay where r a i n f a l l i s g r e a t e r during the growing season. T r i g l o c h i n maritimum L. (arrow-grass) n a t i v e herb Arrow-grass i s a dominant and/or a subdominant on the Cowichan, Nanaimo, Englishman, Adam-Eve, T s i t i k a , Cluxewe and Quatse R i v e r s ' E s t u a r i e s . It occurs on r e l a t i v e l y w e l l d r a i n e d , sandy s u b s t r a t e s where i t i s in-ndated on each f l o o d t i d e . It i s i n d i c a t i v e of b r a c k i s h to s a l i n e c o n d i t i o n s as suggested by the a s s o c i a t e d s p e c i e s t u f t e d h a i r g r a s s , Lyngby's sedge, and s a l t w o r t , s p e c i e s i n d i c a t i v e of b r a c k i s h and s a l i n e c o n d i t i o n s r e s p e c t i v e l y . - 109 -Tsuga h e t e r o p h y l l a (Raf.) Sarg. (western hemlock) n a t i v e tree Western hemlock i s a dominant on the Salmon, Adam-Eve and Quatse R i v e r s ' E s t u a r i e s . It grow best i n humid zones with long v e g e t a t i v e seasons and p o d z o l i z e d s o i l s . I t s n u t r i t i o n a l requirements i n c l u d e a w e l l balanced supply of n u t r i e n t s i n small q u a n t i t i e s and a continuous uptake of water provided by r a i n f a l l ( K r a j i n a 1969/1970). Typha l a t i f o l i a L. ( c a t - t a i l ) n a t i v e herb C a t - t a i l i s a dominant on the Cowichan, Nanaimo, L i t t l e Qualicum, Courtenay, Salmon and Nimpkish R i v e r s ' E s t u a r i e s . It i s i n d i c a t i v e of freshwater c o n d i t i o n s though i t i s known to t o l e r a t e very low s a l i n i t i e s . It tends to occur towards the backshore i n water s a t u r a t e d s o i l or standing water. There i t may be inundated on the higher f l o o d t i d e s during the growing season. I t s growth i s thought to be enhanced by changes i n the water l e v e l . V i c i a gigantea Hook, ( g i a n t vetch) n a t i v e herb Giant vetch i s a dominant on the T s i t i k a R i v e r Estuary and i s present on some of the other e s t u a r i e s s t u d i e d . It t r a i l s over other s p e c i e s such as w i l d r y e and young S i t k a spruce i n the ecotone between marsh and f o r e s t . It i s probably never inundated d u r i n g the growing season and t o l e r a t e s s a l t spray. It occurs on w e l l d r a i n e d , sandy s u b s t r a t e s . Z o s t e r a marina L. ( e e l - g r a s s ) n a t i v e herb E e l - g r a s s i s a dominant and a subdominant on the - 110 -Chemainus R i v e r E s t u a r y where i t may be found at the lower e l e v a t i o n s of the vegetated i n t e r t i d a l and s u b t i d a l zones. It i s a s u b t i d a l s p e c i e s as i t i s s u b j e c t to d e s i c c a t i o n when exposed. On the Chemainus R i v e r Estuary i t i s exposed f o r short periods of time i n t i d a l p o o l s . E e l - g r a s s occurs on sand or s i l t e d sand i n s a l i n e c o n d i t i o n s . It i s an important forage f o r waterfowl and probably f o r many other animals. - I l l -4. ESTIMATING THE PRODUCTION OF DRY MATTER OF VANCOUVER ISLAND ESTUARIES Annual p r o d u c t i o n of dry matter by communities i n e s t u a r i e s can be, i n terms of the world's v e g e t a t i o n systems, very high when elements of the e s t u a r i n e environment are s u i t a b l y combined. T h i s i s r e f l e c t e d i n the heavy occupancy of the major e s t u a r i n e systems of the world by man and other consumers i n the bio s p h e r e . It i s a l s o true that p r o d u c t i o n i n some communities or p a r t s of communities must be very low or n i l when elements such as l a c k of l i g h t or water movement prevent or s e v e r e l y l i m i t p l a n t e stablishment and development. However, i t i s only i n rec e n t decades that a t t e n t i o n has been d i r e c t e d by s c i e n t i s t s and t e c h n o l o g i s t s to pro d u c t i o n i n e s t u a r i e s and the b i o p h y s i c a l elements r e l a t i n g to i t . Methods of e s t i m a t i n g p r o d u c t i o n are not yet w e l l developed. In t h i s study, so l i t t l e was known about the North P a c i f i c e s t u a r i e s i t was b e l i e v e d that crude e s t i m a t e s of the annual p r o d u c t i o n of dry matter of major plant communities on e s t u a r i e s would be i n f o r m a t i v e and u s e f u l . 4.1 Standing Crop Three methods of determining standing crop were used i n t h i s study. The three methods are peak standing crop (Yamanaka, 1975), p e r i o d i c c o l l e c t i o n s (Levings and Moody, 1976) and cl e a r e d p l o t ( K i s t r i t z and Y e s a k i , 1979). Each method i n v o l v e s c o l l e c t i n g to ground l e v e l a l l the m a t e r i a l from a p l o t of known area (eg. one square meter) then d r y i n g and weighing i t to o b t a i n the dry weight (DW) of the matter produced i n the area. - 112 -In the peak standing crop method one c o l l e c t i o n i s made at what i s thought to be the time of maximum pr o d u c t i o n in the sampled stand. The time of sampling i s c r i t i c a l as e a r l y or l a t e c o l l e c t i o n s w i l l not repr e s e n t the peak. T h i s method does not take i n t o account the net changes i n standing crop or the presence of m a t e r i a l from previous growing seasons or other sources. In the p e r i o d i c c o l l e c t i o n s method, p l o t s are p e r i o d i c a l l y and s y s t e m a t i c a l l y sampled. T h i s method takes i n t o account the net changes i n the standing crop but does not e l i m i n a t e the weight c o n t r i b u t e d by the presence of m a t e r i a l from p r e v i o u s growing seasons or other sources. The c l e a r e d p l o t method removes the l a t t e r problem as the p l o t s are c l e a r e d before the sampling program begins. In the c l e a r e d p l o t method p e r i o d i c and sy s t e m a t i c sampling of the p l o t s allows the c a l c u l a t i o n of the net changes i n the standing crop. In t h i s study eleven p l a n t communities were sampled monthly from A p r i l to October i n c l u s i v e . The p l o t s c l i p p e d i n A p r i l were t r e a t e d as c l e a r e d p l o t s and r e c l i p p e d i n October to measure the c l e a r e d p l o t standing crop f o r the p e r i o d May to October. The l a r g e s t monthly weights were designated peak standing crops. The standing crop by p e r i o d i c c o l l e c t i o n s was determined by summing monthly increment and was designated standing crop by summing increments. 4.1.1 Methods Standing crop was obtained from eleven p l a n t communities i n f i v e e s t u a r i e s v i z . Cowichan, Chemainus, L i t t l e Qualicum, - 113 -Campbell and Salmon R i v e r s . The p l a n t communities are typed by dominants; Lyngby's sedge, B a l t i c rush, s a l t w o r t , s a l t g r a s s -gumweed, c i n q u e f o i l - L y n g b y 1 s sedge, c i n q u e f o i l - s p i k e rush, t u f t e d h airgrass-Lyngby' s^  -edge and Kentucky b l u e g r a s s -b e n t g r a s s - c i n q u e f o i l . Three t r a n s e c t s were placed across the e l e v a t i o n a l g r a d i e n t i n each community and f i v e , e q u i d i s t a n t l y spaced, .5 m2 p l o t s were placed one meter on each s i d e of each t r a n s e c t . F i v e p l o t s were sampled i n each of the months A p r i l (4), May ( 5 ) , June ( 6 ) , J u l y ( 7 ) , August (8) and October (10). In October the p l o t s c l i p p e d i n A p r i l were sampled again. The p l o t s were c l i p p e d to ground l e v e l with hand shears and the m a t e r i a l was separated i n t o l i v i n g , dead, senescent and duff f r a c t i o n s . A senescent f r a c t i o n was not c o l l e c t e d f o r the s a l t marsh communities f o r two reasons: a) the d i f f i c u l t y of sep a r a t i n g f r a c t i o n s of senescent m a t e r i a l from a p l a n t , and b) the l a t e August and October development of senescent m a t e r i a l . L i v i n g m a t e r i a l c o n s i s t e d of m a t e r i a l which grew i n 1976 and was s t i l l green; dead m a t e r i a l c o n s i s t e d of growth made before 1976 and was brown; senescent m a t e r i a l was t i s s u e which was produced i n 1976 and was l e s s than h a l f green; and duff was small p a r t i c l e s of amorphous m a t e r i a l on the s o i l s u r f a c e . Each p o r t i o n was bagged, l a b e l l e d and tr a n s p o r t e d to the U n i v e r s i t y of B r i t i s h Columbia where i t was d r i e d i n a Pot Hole D r i e r at 54°C. Depending on the p l a n t s p e c i e s t h i s took from 48 hours to 1.5 weeks. Dr i e d m a t e r i a l was weighed, a standing crop c a l c u l a t e d , and an a n a l y s i s of v a r i a n c e f o r weight by month sampled, and community done. - 114 -Subsamples were ground i n a Wiley M i l l using a 2 mm mesh. To reduce the volume, the f i v e r e p l i c a t e s were combined by equal amounts. Percent ash and n i t r o g e n were determined using methods i n Black et a l . (1965) and were used to c a l u c l a t e ash f r e e dry weight (AFDW) and weight of n i t r o g e n i n combined l i v i n g and senescent, and dead and duff AFDW f r a c t i o n s . 4.1.2 Results The three ways of c a l c u l a t i n g standing crop provide three sets of v a l u e s (Table 46). Standing crop by summing increments ranges from 392 gm m - 2 f o r the c i n q u e f o i l - s p i k e rush community on the Campbell R i v e r E s t u a r y to 1504 gm m~2 f o r the Lyngby's sedge community on the L i t t l e Qualicum R i v e r Estuary. Peak standing crop ranges from 613 gm m - 2 i n J u l y f o r the c i n q u e f o i l -s p i k e rush community on the Campbell R i v e r E s t u a r y " t o 2082 gm m - 2 i n August f o r the s a l t w o r t community on the Chemainus River E s t u a r y . Standing crop by c l e a r e d p l o t (May - October) ranges from 229 gm m - 2 f o r the c i n q u e f o i l - s p i k e rush community on the Campbell R i v e r Estuary to 726 gm m - 2 f o r the s a l t w o r t community on the Chemainus R i v e r E s t u a r y . The rank of each community v a r i e s by method. The a n a l y s i s f o r v a r i a n c e i s s i g n i f i c a n t (p<.05) f o r standing crop by summing increments and the l i v i n g and senescent f r a c t i o n s by community and sampling date. The dead and duff f r a c t i o n s are not s i g n i f i c a n t by sampling date. A m u l t i p l e range t e s t on standing crop y i e l d e d three subsets. The f i r s t subset c o n s i s t s of the communities not dominated by Lyngby's sedge; the second, the Lyngby's sedge communities on the Table 46. Standing Crop Estuary Corrmunity Cowichan Carex lyngbyei 588 July 1231 358 Cowichan Juncus balticus 754 August 1155 364 Chemainus Salicornia virginica 966 August 2082 726 Chemainus Distichlis spicata - Grindelia 1437 July 1420 519 integrifolia L i t t l e Qualicum Carex lyngbyei 1504 June 1746 443 L i t t l e Oualicum Potentilla pacifica - Carex 770 June 868 413 lyngbyei Campbell Carex lyngbyei 487 July 1223 527 Campbell Potentilla pacifica - Eleocharis 392 July 613 229 palustris Salmon Carex lyngbyei 773 July 1186 550 Salmon Deschampsia cespitosa - Carex 1092 July 1210 408 lyngbyei Salmon Poa pratensis - Agrostis alba var. 772 May 1002 342 stolonifera - Potentilla  pacifica Standing Crop Peak Standing Crop t (B^--B-(-_^)a Cleared Plot 1 Standing Crop May - Oct. gm m""^  month gm m - 2 gm m - 2 a. Bj- standing crop at time t ^t-1 standing crop at the previous sample time - 116 -Cowichan, Campbell and Salmon; and the t h i r d , the Lyngby's sedge community on the L i t t l e Qualicum. The l i v i n g f r a c t i o n of the standing crop i n each community (Appendix 6. F i g u r e s 23 to 33) +ends to r a p i d l y increase to a peak then r a p i d l y decrease, except f o r the Chemainus saltgrass-gumweed community which continued to slowly i n c r e a s e to October. Four communities reach t h e i r peak i n June, s i x i n J u l y , one i n J u l y and August, and one i n October. In general, b r a c k i s h marsh communities peak i n the f i r s t h a l f of the growing season and s a l t marsh communities i n the l a t t e r h a l f . B r a c k i s h marshes south of Campbell R i v e r peak i n June while those at Campbell R^ver and north peak i n J u l y . The dead f r a c t i o n of- the standing crop i n each community tends to decrease from A p r i l to October at d i f f e r e n t r a t e s f o r each community. The Lyngby's sedge communities appear to decrease q u i c k l y at the beginning of the growing season. The Cowichan and Campbell sedge communities d i d not have any dead to c o l l e c t from May on, and the L i t t l e Qualicum and Salmon d i d not have dead f r a c t i o n s a f t e r June. The dead f r a c t i o n f o r the Chemainus s a l t w o r t community i s i r r e g u l a r with s l i g h t i ncreases in June and August. A dead f r a c t i o n was c o l l e c t e d from the other communities each sampling time. The senescent f r a c t i o n of the standing crop i n each community i n c r e a s e s r a p i d l y from A p r i l on i n the four sedge communities and June on i n the remaining communities. The duff f r a c t i o n of the s + a n d i n g crop i n each community i s i r r e g u l a r . In g e n e r a l , as the l i v i n g f r a c t i o n of the standing crop i s i n c r e a s i n g to i t s peak, the dead f r a c t i o n i s d e c l i n i n g , the - 117 -senescent i n c r e a s i n g and the duff f l u c t u a t i n g . A f t e r reaching i t s peak t'-e l i v i n g f r a c t i o n d e c l i n e s and the senescent f r a c t i o n i n c r e a s e s r a p i d l y . In the Cowichan, L i t t l e Qualicum and Salmon Lyngby's sedge communities ( F i g u r e s 23, 24 and 25), the senescent f r a c t i o n exceeds the l i v i n g i n August. In the Cowichan B a l t i c rush, L i t t l e Qualicum c i n q u e f o i l - L y n g b y ' s "edge, and Campbell c i n q u e f o i l - s p i k e - r u s h communities ( F i g u r e s 26, 27 and 28), the l i v i n g f r a c t i o n peaks over three months (May, June and J u l y ) and the senescent f r a c t i o n appears to i n c r e a s e as r a p i d l y as the l i v i n g d e c l i n e s , exceeding the l i v i n g a f t e r August. The dead f r a c t i o n exceeds the l i v i n g i n A p r i l only on the Cowichan and Campbell but exceeds the l i v i n g f o r A p r i l and May on the L i t t l e Qualicum. In the Chemainus s a l t w o r t community ( F i g u r e 29) the dead f r a c t i o n i s g r e a t e r than the l i v i n g and both peak i n August. In the Chemainus saltgrass-gumweed community ( F i g u r e 30) the d e c r e a s i n g dead f r a c t i o n exceeds the i n c r e a s i n g l i v i n g u n t i l J u l y . In the Campbell Lyngby's sedge community ( F i g u r e 31) the l i v i n g peak spans May, June and J u l y and the senescent i n c r e a s e s though not as d r a m a t i c a l l y as i n pr e v i o u s e s t u a r i e s . In the Salmon t u f t e d h a i r g r a s s - L y n g b y 1 s sedge community ( F i g u r e 32) the dead f r a c t i o n exceeds the l i v i n g f r a c t i o n i n A p r i l , May and June and the dead peak i n A p r i l i s greater than the l i v i n g peak i n J u l y . In the Salmon Kentucky b l u e g r a s s - b e n t g r a s s - c i n q u e f o i l community ( F i g u r e 33) t'"e dead f r a c t i o n exceeds the l i v i n g i n A p r i l , May and June, and the dead peak i n A p r i l i s g r e a t e r than the l i v i n g peak i n June. In g e n e r a l , the dead f r a c t i o n exceeds the l i v i n g i n communities at higher e l e v a t i o n s and i n s a l t marshes. In communities at higher e l e v a t i o n s the dead f r a c t i o n decreases to - 118 -l e s s than the l i v i n g f r a c t i o n over the growing season, while i n s a l t marsh communities i t does not. The n i t r o g e n a n a l y s i s ranged from an A p r i l high of 2.43% i n the Cowichan Lyngby's sedge community to an October low of .54% i n the Campbell c i n q u e f o i l - spike rush community. Ranges f o r each f r a c t i o n are, f o r the l i v i n g f r a c t i o n , the aforementioned o v e r a l l high and October low of .64% i n the L i t t l e Qualicum c i n q u e f o i l - Lyngby's sedge community; for the dead f r a c t i o n , an October high of 1.61% i n the Salmon t u f t e d h a i r g r a s s - Lyngby's sedge community and an A p r i l low of .71% i n the L i t t l e Qualicum c i n q u e f o i l -Lyngby's sedge community; f o r the senescent f r a c t i o n , an A p r i l high of 1.26% i n the Salmon Kentucky b l u e g r a s s - bentgrass - c i n q u e f o i l community and an October low of .54% i n the Campbell c i n q u e f o i l - spike rush community; f o r the d u f f f r a c t i o n , an October high of 1.84% i n the Chemainus s a l t g r a s s - gumweed community and May low of .65% i n the Cowichan Lyngby's sedge community. In g e n e r a l , f o r a l l but the sedge communities, the nitrogen values of the l i v i n g f r a c t i o n s i n A p r i l are higher than the n i t r o g e n values f o r the senescent dead and duff f r a c t i o n s . However, the n i t r o g e n values f o r the l i v i n g f r a c t i o n s d e c l i n e to the lowest v a l u e s by October. The percent n i t r o g e n value of the senescent f r a c t i o n tends to decrease over time and, while the percent n i t r o g e n values of the dead and duff f r a c t i o n s a l s o decrease i n A p r i l and May, they i n c r e a s e from June ( J u l y ) on and have the h i g h e s t v a l u e s , p a r t i c u l a r l y the d u f f f r a c t i o n (around 1%). The range of values f o r the duff f r a c t i o n i s the s m a l l e s t of a l l the f r a c t i o n s . - 119 -The ash a n a l y s i s ranged from a May low of .74% f o r the l i v i n g f r a c t i o n of the L i t t l e Qualicum Lyngby's sedge community to an August high of 75.12% f o r the du f f f r a c t i o n of the Cowichan Lyngby's sedge community. Ranges f o r each f r a c t i o n are, f o r the l i v i n g f r a c i o n , the aforementioned o v e r a l l low and August high of 36.73% i n the Chemainus saltwort, community; f o r the dead f r a c t i o n , a J u l y low of 4.37% i n the c i n q u e f o i l -Lyngby's sedge, and May high of 50.97% i n the Lyngby's sedge communities on the L i t t l e Qualicum; f o r the senescent f r a c t i o n , an October low of 4.11% i n the Campbell c i n q u e f o i l - spike rush community and June high of 39.78% i n the L i t t l e Qualicum Lyngby's sedge community; f o r the duff f r a c t i o n , an August low of 14.55% and the aforementioned o v e r a l l h i gh. Though percent ash values f l u c t u a t e , i n g e n e r a l , they decrease from A p r i l to October. The lowest percent ash values occur i n the l i v i n g f r a c t i o n and i n c r e a s e through the senescent and dead f r a c t i o n s to the duff f r a c t i o n . The c a l c u l a t i o n of ash f r e e dry weight (AFDW) and n i t r o g e n values f o r the combination of l i v i n g and senescent f r a c t i o n s f o r each community r e s u l t s i n two types of graphs (Appendix 6, F i g u r e s 34 to 37). The f i r s t type, r a p i d r i s e to a peak, then r a p i d d e c l i n e of AFDW, occurs f o r the Lyngby's sedge communities on the Cowichan, L i t t l e Qualicum, Campbell and Salmon, the Chemainus s a l t w o r t community and the Campbell c i n q u e f o i l - spike rush community. The second type, continuous increase of AFDW, occurs f o r the remaining communities. There i s a decrease i n August before the i n c r e a s e c o n t i n u e s i n the three communities, Cowichan B a l t i c rush (where decrease occurs i n J u l y a l s o ) , L i t t l e Qualicum c i n q u e f o i l - Lyngby's sedge and - 120 -Salmon t u f t e d h a i r g r a s s - Lyngby's sedge. Nitrogen values i n c r e a s e and decrease with AFDW i n the Lyngby's sedge communities on the L i t t l e Qualicum, Campbell and Salmon, Chemainus s a l t w o r t communit and Campbell c i n q u e f o i l - spike rush community. In the Cowichan Lyngby's sedge community n i t r o g e n v a l u e s i n c r e a s e from August to October. In the remaining communities the n i t r o g e n values i n c r e a s e from A p r i l to October. The a p p l i c a t i o n of percent n i t r o g e n and AFDW v a l u e s to the combination of dead and duff weights f o r each community provides v a r i a b l e r e s u l t s though AFDW tends to decrease through the growing season (A^-endix 6, Fi g u r e s 38 to 41). Values f o r Lyngby's sedge communities d e c l i n e r a p i d l y , while values f o r the r e s t of the communities d e c l i n e comparatively s l o w l y . Weights are lowest, l e s s than 50 gms, f o r the sedge communities. The s a l t marsh communities have the hig h e s t weights ranging between 230 and 120 gms. AFDW i n the r e s t of the communities i s between 130 and 20 gms. Nitr o g e n values f l u c t u a t e with AFDW and are le s s i n October than i n A p r i l . 4.1.3. D i s c u s s i o n There i s c o n s i d e r a b l e v a r i a t i o n i n standing crop values by method and community. Of the three methods used, summing increments i s con s i d e r e d to be most r e p r e s e n t a t i v e . Summing increments between sampling p e r i o d s i s more e x a c t i n g than c o l l e c t i n g the peak standing crop which overestimates the value as i t does not account f o r the i n c l u s i o n of previous years' growth. The c l e a r e d p l o t method does not account f o r r e d u c t i o n s - 121 -i n standing crop at the end of the growing p e r i o d but i t e l i m i n a t e s the problem of previous years' growth and provides a d e f i n i t e time r e f e r e n c e over which to measure the standing crop. However, i n t h i s study, the c l e a r e d p l o t s were set up l a t e , i n May, and t h i s l i m i t e d the growth p e r i o d and r e s u l t e d i n the standing crop values being -nderestimated. A l l three methods have the same problems with c l i p p i n g to ground l e v e l , s o i l c o ntamination (as i n d i c a t e d by the percent ash v a l u e s ) , p o p u l a t i o n d e n s i t y v a r i a b i l i t y , sample s i z e , and l o s s due to l e a c h i n g , g r a z i n g and p h y s i c a l removal. None of the methods includes belowground o r g a n i c matter. The standing crop values obtained by summing increments (eg. Lyngby's sedge 487 to 1504 grams of dry weight per square meter (gm DW m - 2 ) ) compare to Levings and Moody's (1976) value of 1322 gm DW m - 2 f o r Lyngby's sedge in the Squamish R i v e r e s t u a r y and Moody's (1978) value of 909 gm DW m~2 f o r Lyngby's sedge i n the Brunswich P o i n t Marsh on the F r a s e r R i v e r d e l t a . D i r e c t comparisons between standing crop values f o r emergent plant communities along the P a c i f i c Northwest coast are d i f f i c u l t . D i f f e r e n t communities are sampled (eg. maritime bulrush) and d i f f e r e n t methods (eg. peak standing crop) are used. K i s t r i t z and Yesaki (1979) a l s o found " l a r g e temporal and s p a t i a l v a r i a t i o n s a s s o c i a t e d with measurements of marsh p l a n t s " and that there i s "a marked year to year v a r i a t i o n i n the timing and d u r a t i o n of growth..." In t h i s study t h i s i s seen i n the d i f f e r e n c e s i n the times of i n i t i a t i o n and c e s s a t i o n of growth among the p l a n t communites, f o r example, the b r a c k i s h marsh communities i n i t i a t e d growth i n l a t e M»r " and e a r l y A p r i l while - 122 -the s a l t marsh communities i n i t i a t e d growth i n l a t e A p r i l . K i s t r i t z (1978) a l s o found t h i s i n h i s l i t e r a t u r e review. The problems a s s o c i a t e d with the standing crop methodology, and r e p r o d u c a b i l i t y of the r e s u l t s lead to the c o n c l u s i o n that i t i s not a s a t i s f a c t o r y approach to measuring the dry matter c o n t r i b u t i o n s of emergent v e g e t a t i o n to the d e t r i t a l c y c l e i n the e s t u a r i n e food web. A l s o , the " q u a l i t y " of the dry matter c o n t r i b u t i o n s to the d e t r i t a l c y c l e , as i n d i c a t e d by the t o t a l n i t r o g e n content of each community, i s precluded by the problems with the standing crop technique and only one monthly measurement of n i t r o g e n f o r each community. While methodology and r e p r o d u c a b i l i t y problems with standing crop l i m i t the u s e f u l n e s s of the standing crop values., the f r a c t i o n and ash f r e e dry weight (AFDW) p o r t i o n of the study i n d i c a t e d i f f e r e n c e s between the p l a n t communities and t h e i r c o n t r i b u t i o n s to the d e t r i t a l c y c l e . By summing increments the four l a r g e s t standing crops oc c u r r e d on the L i t t l e Qualicum Lyngby's sedge community at 1504 grams per square meter (gm m - 2) followed by the Chemainus saltgrass-gumweed community at 1437 gm m~2 then the Salmon t u f t e d hairgrass-Lyngby's sedge at 1092 gm m - 2 and f i n a l l y the Chemainus s a l t w o r t community at 966 gm m - 2. The f r a c t i o n and ash f r e e dry weight data f o r each of these communities i n d i c a t e the r a t e s and timing of r e l e a s e of dry matter or p a r t i c u l a t e o r g a n i c matter to the e s t u a r i n e system. In the f r a c t i o n data the l i v i n g m a t e r i a l i n c r e a s e s from A p r i l to June or J u l y then decreases to October. The dead decreases from A p r i l to October. The senescent i n c r e a s e s from A p r i l to October and the duff i s i r r e g u l a r "nd f l u c t u a t e s aro"rtd the same g e n e r a l v a l u e s . In the sedge communities the dead p o r t i o n never exceded the l i v i n g and no dead was c o l l e c t e d a f t e r June. A l s o , the l i v i n g p o r t i o n increased and decreased very sharply i n both the portion and combined l i v i n g and senescent AFDW data. This does not occur i n the other communities and i n d i c a t e s that the sedge communities r e l e a s e matter to the d e t r i t a l c y c l e throughout the year, not j u s t i n the growing season, and the release appears to be a l l or nearly a l l the matter from the community. In the other communities, the dead p o r t i o n exceded the l i v i n g u n t i l June and was present u n t i l October, and the combined l i v i n g and senescent AFDWs increased to October. This i n d i c a t e s four things: 1. these communities release matter much more slowly than the sedge communities. 2. the matter released i s p r i n c i p a l l y from previous year's growth. 3. the release occurs mainly i n the growing season. 4. there i s a net accumulation of matter i n the communities. The accumulation of dead matter i n the s a l t marsh communities i s p a r t i c u l a r l y pronounced with the dead f r a c t i o n i n the saltwort community always exceeding the l i v i n g f r a c t i o n and the combined dead and duff AFDWs decreasing only s l i g h t l y . This i n d i c a t e s there i s l i t t l e matter released from the s a l t marsh communities to the estuarine system and matter accumulates i n the community. In the higher e l e v a t i o n communities the accumulated matter appears to be incorporated i n t o the substrate to such an extent that one may have to step up onto these communities from lower e l e v a t i o n communities. Alone, the standing crop values might suggest the sedge, higher e l e v a t i o n - 124 -t u f t e d h a i r g r a s s - s e d g e , and s a l t marsh s a l t w o r t and s a l t g r a s s -gumweed communities c o n t r i b u t e the g r e a t e s t amounts of matter to the d e t r i t a l c y c l e i n the e s t u a r i n e food web. However, the p o r t i o n and AFDW data p o i n t out that the r a t e and timing of r e l e a s e of matter to the e s t u a r i n e system v a r i e s between sedge, higher e l e v a t i o n b r a c k i s h marsh and s a l t marsh communities. The sedge communities have the highest r a t e of r e l e a s e and r e l e a s e a l l or almost a l l of the matter produced throughout the year. The higher e l e v a t i o n b r a c k i s h marsh communities slowly r e l e a s e some of the matter produced i n p r e v i o u s years dur i n g a growing season and accumulate matter through the remainder of the year. The s a l t marsh communities slowly r e l e a s e a l i t t l e matter i n the growing season and accumulate the r e s t . It appears that i n a b r a c k i s h marsh the emergent v e g e t a t i o n i s a source of p a r t i c u l a t e o r g a n i c matter f o r the d e t r i t a l c y c l e i n the food web with sedge r e l e a s i n g matter year ro~nd and higher e l e v a t i o n communities r e l e a s i n g matter only during the growing season when energy demands are probably at t h e i r h i ghest i n the e s t u a r y . However, i t appears that s a l t marshes c o n t r i b u t e l i t t l e p a r t i c u l a t e o r g a n i c matter to the d e t r i t a l c y c l e . Haines (1977) found t h i s i n cordgrass s a l t marshes at the U n i v e r s i t y of Georgia Marine I n s t i t u t e on Sapelo Isla n d and concluded that phytoplankton and t e r r e s t r i a l p l a n t m a t e r i a l were the probable major sources of p a r t i c u l a t e o r g a n i c matter i n s a l t marshes. Haines f e l t a major r e v i s i o n of ideas concerning the r o l e of s a l t marshes i n e s t u a r i n e p r o d u c t i o n might be r e q u i r e d . This study suggests any r e v i s i o n take i n t o account the d i f f e r e n c e s between b r a c k i s h and s a l t marshes. B r a c k i s h marshes r e l e a s e p a r t i c u l a t e o r g a n i c matter to the d e t r i t a l c y c l e i n the - 125 -e s t u a r i n e food web while s a l t marshes r e l e a s e l i t t l e . T e r r e s t r i a l p l a n t m a t e r i a l c o n t r i b u t e s to the c y c l e i n e s t u a r i e s i n e i t h e r marsh, but phytoplankton are probably more abundant in s a l t marshes due to the higher s a l i n i t i e s t h e r e . The side by side occurrence of many of Vancouver I s l a n d ' s east coast b r a c k i s h and s a l t marshes suggests t o t a l p r o d u c t i v i t y from a l l sources, emergent v e g e t a t i o n , t e r r e s t r i a l p l a n t m a t e r i a l and phytoplankton, w i t h i n the e s t u a r i n e ecosystem may be much higher than ever a p p r e c i a t e d . The d i f f e r e n c e s i n the r a t e s and timing of r e l e a s e of the p a r t i c u l a t e o r g a n i c matter to the d e t r i t a l c y c l e of the e s t u a r i n e food web separates sedge, s a l t and b r a c k i s h marsh communities and r a i s e s the q u e s t i o n , "Why are they d i f f e r e n t ? " There appear to be two p r i n c i p a l reasons. They are, one, mechanical damage and, two, p l a n t s p e c i e s anatomy. The f i r s t reason, mechanical damage, i s r e l a t e d to the d i s t r i b u t i o n of the p l a n t communities. P l a n t communities at lower e l e v a t i o n s i n the vegetated i n t e r t i d a l zone are s u b j e c t to the a c t i o n of waves twice d a i l y on t i d a l i n " n d a t i o n and t h i s a c t i o n breaks the plants down and c a r r i e s them i n t o the estuary over time. P l a n t communities at higher e l e v a t i o n s are s u b j e c t to only shallow inundation during much of the growing season. Wave energy d i m i n i s h e s with d e c r e a s i n g water depth so mechanical breakage from waves i s l e s s at higher e l e v a t i o n s , and the a b i l i t y of the waves to c a r r y m a t e r i a l i n t o the estuary i s reduced. The second reason, p l a n t s p e c i e s anatomy, i s r e l a t e d to the amount of s t r u c t u r a l m a t e r i a l i n the s p e c i e s . The amount of l i v i n g , as versus s t r u c t u r a l m a t e r i a l , i n the p l a n t communities s t u d i e d i s i n d i c a t e d by the n i t r o g e n v a l u e s . The highest amounts of l i v i n g - 126 -and lowest amounts of s t r u c t u r a l m a t e r i a l are i n the sedge communities, and the lowest l i v i n g and h i g h e s t s t r u c t u r a l amounts are i n the s a l t and higher e l e v a t i o n b r a c k i s h marsh communities. The higher s t r u c t u r a l component of the plant species of the s a l t and higher e l e v a t i o n b r a c k i s h marsh communities makes them more d i f f i c u l t to break down than the sedge communities . Sa l t w o r t and s a l t g r a s s , i n p a r t i c u l a r , have t h i c k c e l l w a l l s and the p l a n t s p e r s i s t from growing season to growing season. The high s a l i n i t i e s i n s a l t marshes may a l s o suppress m i c r o b i a l decomposers which were not i n h i b i t e d i n b r a c k i s h marshes as i n d i c a t e d by the duff n i t r o g e n values around one percent. In c o n c l u s i o n , the d i f f e r e n t r a t e s and timing of re l e a s e of the p a r t i c u l a t e o r g a n i c matter i n sedge, s a l t and higher e l e v a t i o n b r a c k i s h marsh communities appears to be r e l a t e d to the amount of mechanical a c t i o n the communities are s u b j e c t t o , and the p l a n t s p e c i e s decomposition r a t e s as i n f l u e n c e d by t h e i r anatomy and s a l i n i t y . 4.2 Root Reserves E s t i m a t i n g the belowground dry matter production i n e s t u a r i n e marshes pres e n t s as many problems as e s t i m a t i n g standing crop or aboveground dry matter p r o d u c t i o n . The t r a d i t i o n a l method of measuring belowground dry matter i n v o l v e s c o l l e c t i n g r o o t s and s e p a r a t i n g l i v e and dead m a t e r i a l . T h i s was i m p r a c t i c a l i n t h i s study as the communities i n v o l v e d formed root mats that made i t d i f f i c u l t to remove samples from the ground l e t alone p h y s i c a l l y separate l i v e and dead r o o t s . A l s o , i t i s d i f f i c u l t to d i s t i n g u i s h between dead and l i v i n g r o o t s , - 127 -and due to the d e n s i t y of the root mat, s e p a r a t i o n of the roo t s would r e s u l t i n a l o s s of m a t e r i a l , i n p a r t i c u l a r , the l i v i n g r o o t l e t s and root h a i r s which are considered to re p r e s e n t a major p o r t i o n of the l i v e weight. An a l t e r n a t i v e , p e r i o d i c c o l l e c t i o n s of known volumes of r o o t s was d i s c a r d e d due to the d i f f i c u l t y i n washing s o i l out of some of the root mats. One could not be sure how much was root and how much was s o i l and i f trends i n the values were dependent on changes i n the root reserves or s o i l content of the sample, p a r t i c u l a r l y when d i f f e r e n t communities were being examined. A novel, i n d i r e c t approach suggested by Burton and Jackson i n 1962 was used. The le n g t h of time i n t a c t cores of root and s o i l grew i n darkness was measured i n s t e a d . The time the root cores grew i n the dark and the a i r d r i e d weight of e t i o l a t e d c l i p p i n g s were used as a measure of the belowground energy r e s e r v e s f o r the p l a n t communities. The method allows f o r comparisons between e s t u a r i e s and g i v e s trends that are based on the p l a n t s and not on h a n d l i n g e r r o r s . 4.2.1 Methods Root cores were c o l l e c t e d from eleven p l a n t communities in f i v e e s t u a r i e s v i z . Cowichan, Chemainus, L i t t l e Qualicum, Campbell and Salmon R i v e r s . The p l a n t communities are typed by dominants; Lyngby's sedge, B a l t i c rush, s a l t w o r t , s a l t g r a s s -gumweed, c i n q u e f o i l - L y n g b y ' s sedge, c i n q u e f o i l - s p i k e rush, t u f t e d hairgrass-Lyngby's sedge, and Kentucky b l u e g r a s s -b e n t g r a s s - c i n q u e f o i l . One 9.9 cm i n diameter by 11.9 cm deep core was taken from each of f i v e .5 m2 c l i p p e d p l o t s i n each - 128 -community i n each of the months A p r i l ( 4 ) , May ( 5 ) , June ( 6 ) , J u l y (7), August (8) and October (10). The cores were tra n s p o r t e d to the U n i v e r s i t y of B r i t i s h Columbia where they were grown i n a dark room at ca. 20°C, given tap water every 15 days and c l i p p e d every 30 days u n t i l growth ceased. Dates of s e t t i n g out and t e r m i n a t i o n of growth, and a i r dry weight of e t i o l a t e d t i s s u e were recorded. Analyses f o r v a r i a n c e (ANOV) i n time and weight by communit and by month c o l l e c t e d , and between community and month c o l l e c t e d were done. Each month the h e i g h t s of two average shoots of Lyngby's sedge and/ or B a l t i c rush i n each p l o t c l i p p e d the previous month were recorded. In October, the number of shoots that regrew and the number of o v e r w i n t e r i n g shoots ( c a . 5 cm t a l l , blue-green, t a p e r i n g to a p o i n t , t h i c k c u t i c l e ) i n each p l o t c l i p p e d i n A p r i l through August i n c l u s i v e were counted. The number of shoots and o v e r w i n t e r i n g shoots i n the p l o t s c l i p p e d i n October were a l s o counted. A n a l y s i s f o r v a r i a n c e i n the number of regrown shoots and o v e r w i n t e r i n g shoots by time c l i p p e d was done. 4.2.2 Results The ANOV i s s i g n i f i c a n t (p<.05) i n a l l cases though b a r e l y - s o f o r weight. The same r e s u l t i s obtained when the sample p o p u l a t i o n c o n s i s t s only of communities dominated by Lyngby's sedge. A r o o t core from the c i n q u e f o i l and Lyngby's sedge community on the L i t t l e Qualicum R i v e r Estuary grew f o r the - 129 -300 200 CO >• < 100 1 2 3 4 5 6 7 PLANT COMMUNITY Figure 42. Exhaustion of root reserves i n darkness: maximum time the (1) Carex lyngbyei, (2) Juncus b a l t i c u s , (3) Deschampsia cespitosa and (4) S a l i c o r n i a v i r g i n i c a plant communities, and the maximum time (5) Agropyron  spicatum, Stipa comata and Festuca s c a b r e l l a (Lobb, 1969), (6) Poa prat e n s i s , and (7) Festuca rubra and Agrostis tenuis (Marx, 1961) grew i n the dark. - 130 -longest time i n the dark; 573 days. It was c o l l e c t e d on October 13, 1976, set out i n the dark room on October 21, 1976 and ceased growing on June 6, 1978. Fi g u r e 42 compares the maximum time root cores from four e s t u a r i n e p l a n t communities and three groups of grasses grew i n the dark. The e s t u a r i n e root cores grew 3.5 to 10 times longer than the g r a s s e s . The average time the root cores from the eleven communities grew i n the dark i s graphed i n Fi g u r e 43. The graph shows a drop i n the number of days the root cores grew i n the dark from A p r i l to May then an i n c r e a s e u n t i l August a f t e r which time i t d e c l i n e s . The Cowichan Lyngby's sedge community ( F i g u r e 44) reaches maximum time i n J u l y then stays r e l a t i v e l y s t a b l e between J u l y and October. The Cowichan B a l t i c rush community i s i n c r e a s e s from A p r i l to August then stays r e l a t i v e l y s t a b l e u n t i l October ( F i g u r e 45). The Chemainus s a l t w o r t community i s i r r e g u l a r though i t tends to d e c l i n e from A p r i l to October (Figure 46). The Chemainus s a l t g r a s s - gumweed community d e c l i n e s from A p r i l to June then i n c r e a s e s to October (Figure 46). The L i t t l e Qualicum Lyngby's sedge community i n c r e a s e s from A p r i l to October ( F i g u r e 47). The c i n q u e f o i l - L y n g b y ' s sedge community on the same estuary stays the same f o r A p r i l and May then i n c r e a s e s to October ( F i g u r e 48). The Campbell River Lyngby's sedge community i n c r e a s e s from A p r i l to August then d e c l i n e s s l i g h t l y to October ( F i g u r e 49). The c i n q u e f o i l - s p i k e rush community on the Campbell peaks i n J u l y and then stays r e l a t i v e l y s t a b l e u n t i l October ( F i g u r e 50). The Lyngby's sedge community on the Salmon i s l i k e that on the Campbell while the t u f t e d hairgrass-Lyngby's sedge and Kentucky b l u e g r a s s -b e n t g r a s s - c i n q u e f o i l communities on the Salmon i n c r e a s e (except - 131 -300 200 t o < 100 7 8 MONTH 10 Figure 43. The.average number, of days.root cores c o l l e c t e d i n A p r i l (4), May (5), June (6), July (7), August (8), and October (10) grew i n the dark. - 132 -f o r a decrease from A p r i l to.May f o r the t u f t e d h a i r g r a s s -Lyngby's sedge community) to August then d e c l i n e to October (Figures 51 and 52). The height of Lyngby's sedge one month a f t e r c l i p p i n g d e c l i n e s from A p r i l to August on the Cowichan; d e c l i n e s from June to August on, the L i t t l e Qualicum Lyngby's sedge community, d e c l i n e s from A p r i l to J u l y then i n c r e a s e s to August on the L i t t l e Qualicum c i n q u e f o i l - L y n g b y ' s sedge community d e c l i n e s A p r i l through J u l y then remains, r e l a t i v e l y s t a b l e i n August on the Campbell; and i n c r e a s e s from A p r i l to May then decreases to August on the Salmon. The height of B a l t i c rush one month a f t e r c l i p p i n g i s the same i n A p r i l and May, i n c r e a s e s to June, decreases to J u l y then remains r e l a t i v e l y s t a b l e to August on the Cowichan; decreases from A p r i l to May, i n c r e a s e s to June then decreases to August on the L i t t l e Qualicum; and decreases from A p r i l to May, remains r e l a t i v e l y s t a b l e to June then decreases u n t i l August on the Campbell. The number of Lyngby's sedge regrown shoots per month p l o t s c l i p p e d decreases from A p r i l to J u l y then i n c r e a s e s to August on the Cowichan; decreases from A p r i l to May, i n c r e a s e s to J u l y then decreases to August on the L i t t l e Qualicum; decreases from A p r i l to June then i n c r e a s e s to August on the Campbell; and decreases from A p r i l to June then remains r e l a t i v e l y s t a b l e to August on the Salmon. On a l l e s t u a r i e s , except f o r the L i t t l e Qualicum, the average number of shoots i n the p l o t s c l i p p e d i n October i s higher than the number of regrown shoots i n the p l o t s c l i p p e d p r e v i o u s l y . The number of Lyngby's sedge o v e r w i n t e r i n g shoots per month p l o t s c l i p p e d decreases from A p r i l to May, i n c r e a s e s to - 133 -J u l y then decreases to August on the Cowichan; decreases from A p r i l to May then i n c r e a s e s to August on the L i t t l e Qualicum; remains the same i n A p r i l and May, decreases to June then i n c r e a s e s to August on the Campbell; and decreases from A p r i l to June then i n c r e a s e s to August on the Salmon. The average number of o v e r w i n t e r i n g shoots i n the p l o t s c l i p p e d i n October i s higher or the same than f o r p l o t s c l i p p e d p r e v i o u s l y . 4.2.3 D i s c u s s i o n Burton and Jackson's method of e s t i m a t i n g the belowground dry matter p r o d u c t i o n does not o b t a i n the weight of belowground dry matter produced, r a t h e r , i t o b t a i n s two d i f f e r e n t measurements; 1. the number of days p l a n t s i n a root core w i l l grow i n the dark, that i s , u n t i l the energy r e s e r v e s of the p l a n t s ' r o o t s are exhausted, and 2. the a i r d r i e d weight of the e t i o l a t e d shoots produced. The measurements obtained using t h i s method are not comparable as the method has not been used on any other e s t u a r i n e p l a n t communities. However, the trends i n the time the root cores grew i n the dark are the same as K i s t r i t z and Y e s a k i ' s (1979) r e s u l t s by the t r a d i t i o n a l method. The r e s u l t s of t h i s experiment might be a comparable measure of the p r o d u c t i v i t y of the root system i f the gas exchange of the p l a n t s was monitored. Using gas exchange to measure r e s p i r a t i o n would allow the e x p r e s s i o n of the r e s u l t s i n the t r a d i t i o n a l dry weight of matter produced. A l s o , the problems of d i s t i n g u i s h i n g between l i v i n g and dead root m a t e r i a l , l o s s of m a t e r i a l and s o i l contamination would be avoided. - 134 -The r e s u l t s of the root r e s e r v e s p o r t i o n of t h i s study do not p o i n t out d i f f e r e n c e s i n belowground dry matter production between e s t u a r i n e marshes but do p o i n t out r e l a t i o n s h i p s between the below and above ground s t r u c t u r e s of the emergent p l a n t species when the l i v i n g f r a c t i o n of the standing crop s e c t i o n i s c o n s i d e r e d . The changes i n the root r e s e r v e s c o i n c i d e with canopy development. The decrease in the root r e s e r v e s from A p r i l to May c o i n c i d e s with the i n i t i a t i o n and r a p i d development of the canopy ( l i v i n g f r a c t i o n i n s e c t i o n 4.1 Standing c r o p ) , with the i n i t i a t i o n and r a p i d development of the canopy. The increase from May to August c o i n c i d e s with the i n c r e a s e i n s + a n d i n g crop, and the gradual decrease i n root r e s e r v e s a f t e r August occurs a f t e r the canopy d i e s back. What the root r e s e r v e s c o n s i s t of i s unknown but they are probably a combination of some min e r a l elements and carbohydrates. T h e i r storage l o c a t i o n ( s ) , and storage and t r a n s l o c a t i o n form(s) are unknown, as are the mechanism(s) and r a t e ( s ) of m o b i l i z a t i o n . However, i t appears that the root r e s e r v e s maintain the plant through unfavorable growing c o n d i t i o n s , such as winter, and provide energy f o r the r a p i d i n i t i a t i o n and development of the canopy as soon as the growing c o n d i t i o n s are f a v o r a b l e i n A p r i l . D i v e r s i o n of the root r e s e r v e s to the canopy r e s u l t s i n the May d e p l e t i o n but enables the p l a n t to make the most of the short d a y l i g h t low t i d e s i n A p r i l and May. Once the canopy i s e s t a b l i s h e d m a t e r i a l i s d i r e c t e d from the shoots to the r o o t s and the r e s e r v e s i n c r e a s e . The canopy d i e s back as the d a y l i g h t low t i d e s shorten and the p l a n t i s again maintained by the root r e s e r v e s u n t i l the next f a v o r a b l e growing season. The number and h e i g h t , one month a f t e r c l i p p i n g , of the Lyngby's sedge - 135 -stems that regrew on the p l o t s a f t e r c l i p p i n g help i l l u s t r a t e the root r e s e r v e s d i s c u s s i o n . The height decreases from A p r i l to August, however the p l o t s c l i p p e d i n A p r i l were no more than 10 cm s h o r t e r than the surrounding stands i n October. Regrowth on the p l o t s c l i p p e d i n A p r i l was p o s s i b l e due to the e x i s t i n g flow of r e s e r v e s from the r o o t s to the canopy which the stems were able to take advantage of f o r t h e i r regrowth. A f t e r May the flow was from the shoots to the r o o t s and i t would appear there was l i t t l e energy a v a i l a b l e to regrow stems. T h i s i s a l s o the case f o r the number of stems that regrew, however, i n s t e a d of decreasing to August they decrease to June or J u l y then i n c r e a s e to August. T h i s i s p a r a l l e l to the number of o v e r w i n t e r i n g shoots which decreased from A p r i l to June then i n c r e a s e d from June to August. The number of o v e r w i n t e r i n g stems produced on the c l i p p e d p l o t s appears to have been a f f e c t e d by the time the p l o t s were c l i p p e d . The time the p l o t s were c l i p p e d being synonymous with the s t a t e of the r o o t r e s e r v e s . Root r e s e r v e s would not have been r e p l e n i s h e d f o r p l o t s c l i p p e d i n May and June and the p l a n t s i n these p l o t s d i d not have the root r e s e r v e s to produce many o v e r w i n t e r i n g shoots Plants c l i p p e d i n J u l y and August had r e p l e n i s h e d t h e i r root r e s e r v e s and were able to produce o v e r w i n t e r i n g shoots a f t e r being c l i p p e d . The number of stems that regrew a f t e r being c l i p p e d i n c r e a s e d form June to August as a by-product of the formation of the o v e r w i n t e r i n g shoots. .Though t h i s i s not the t r a d i t i o n a l approach to q u a n t i f y i n g belowground matter i t obtained the same trends as K i s t r i t z and Yesaki (1979) d i d f o r Lyngby's sedge on the Woodward Island marsh on the F r a s e r River d e l t a , and i t i l l u s t r a t e s the i n t e r a c t i o n between the canopy and - 136 -r o o t s to q u i c k l y i n i t i a t e and maximize growth and s u r v i v e adverse c o n d i t i o n s i n the e s t u a r y . i - 137 -5. CONCLUSIONS The main c o n c l u s i o n s of t h i s study may be .summarized as fol l o w s : 1. There i s a great d e a l of v a r i a t i o n i n the species composition of p l a n t communities both w i t h i n and among e s t u a r i e s . 2. There are s a l t and b r a c k i s h marshes i n e s t u a r i e s on the east coast of Vancouver I s l a n d . 3. D i f f e r e n t p l a n t s p e c i e s dominate i n e s t u a r i n e marshes north and south of Courtenay. In b r a c k i s h marshes south of Courtenay Juncus spp., P o t e n t i l l a p a c i f i c a and D i s t i c h l i s  s p i c a t a dominate, while north of Courtenay Deschampsia  c e s p i t o s a and P o t e n t i l l a p a c i f i c a dominate. In s a l t marshes south of Courtenay D i s t i c h l i s s p i c a t a , A t r i p l e x p a t u l a and G r i n d e l i a i n t e g r i f o l i a dominate, while north of Courtenay T r i g l o c h i n maritimum and grasses dominate. 4. There are eleven types of e s t u a r i e s i n the study based on the i n t e r r e l a t i o n s h i p s of the s i x p h y s i c a l f a c t o r s ; 1. time of maximum d i s c h a r g e , 2. r e l a t i o n s h i p between a r i v e r ' s average A p r i l to September mean dis c h a r g e and the s i z e of the r i v e r ' s d e l t a , 3. mean annual t o t a l p r e c i p i t a t i o n , 4. the r e l a t i v e p r o t e c t i o n from wind and wave energy of the sea, 5. the p a r t i c l e s i z e of the s u b s t r a t e , and 6. the d u r a t i o n and frequency of t i d a l i n u n d a t i o n . - 138 -5. The standing crop f o r each community i s gm m-2 Cowichan Carex l y n g b y e i 5 8 8 Cowichan Juncus b a l t i c u s 754 Chemainus S a l i c o r n i a v i r g i n i c a 966 Chemainus D i s t i c h l i s s p i c a t a - G r i n d e l i a 1437 i n t e g r i f o l i a L i t t l e Qualicum Carex l y n g b y e i 1504 L i t t l e Qualicum P o t e n t i l l a p a c i f i c a - Carex 770 l y n g b y e i Campbell Carex l y n g b y e i 4 8 7 Campbell P o t e n t i l l a p a c i f i c a - E l e o c h a r i s 392 p a l u s t r i s Salmon Carex l y n g b y e i 773 Salmon Deschampsia c e s p i t o s a - Carex l y n g b y e i 1092 Salmon Poa p r a t e n s i s - A g r o s t i s a l b a v a r . 772 s t o l o n i f e r a - P o t e n t i l l a p a c i f i c a 6. The Carex l y n g b y e i communities have the highest standing crops. 7. The l i v i n g f r a c t i o n of the standing crop i s always l a r g e r than the senescent, dead or duff f r a c t i o n s i n the Carex  l y n g b y e i communities. 8. The l i v i n g f r a c t i o n i s exceeded by the dead f r a c t i o n f o r p a r t of the growing season i n the Cowichan Juncus b a l t i c u s , Chemainus D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a , - 139 -L i t t l e Qualicum P o t e n t i a l l a p a c i f i c a - Carex l y n g b y e i , Campbell P o t e n t i l l a p a c i f i c a - E l e o c h a r i s p a l u s t r i s , and Salmon Deschampsia c e s p i t o s a - Carex l y n g b y e i and Poa  p r a t e n s i s - A g r o s t i s a l b a var. s t o n i f e r a - P o t e n t i l l a  p a c i f i c a communities. 9. The l i v i n g f r a c t i o n i s exceeded by the dead f r a c t i o n f o r a l l of the growing season i n the Chemainus S a l i c o r n i a v e r g i n i c a community. 10. The ash f r e e dry weight of combined l i v i n g and senescent f r a c t i o n s i n the Carex l y n g b y e i , Chemainus S a l i c o r n i a  v i r g i n i c a and Campbell P o t e n t i l l a p a c i f i c a - E l e o c h a r i s  p a l u s t r i s communities r i s e s r a p i d l y to a peak then-d e c l i n e s . 11. The ash f r e e dry weights of combined l i v i n g and senescent f r a c t i o n s i n the Cowichan Juncus b a l t i c u s , Chemainus D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a , L i t t l e Qualicum P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i , Salmon Deschampsia  c e s p i t o s a - Carex l y n g b y e i , and Poa p r a t e n s i s - A g r o s t i s  alba var. s t o l o n i f e r a - P o t e n t i l l a p a c i f i c a communities r i s e r a p i d l y then l e v e l out. 12. The ash f r e e dry weights of the combined dead and duff f r a c t i o n s decrease through the growing season f o r a l l the communities. The Carex l y n g b y e i communities d e c l i n e most r a p i d l y as there i s no dead f r a c t i o n a f t e r May or June. - 140 -13. The percent n i t r o g e n content f o r a l l the communities was h i g h e s t i n A p r i l with the maximum being 2.43% f o r the l i v i n g f r a c t i o n of the Cowichan Carex l y n g b y e i community. The minimum was .54% f o r the senescent f r a c t i o n of the L i t t l e Qualicum P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i community. 14. The h i g h e s t n i t r o g e n values occur i n the l i v i n g f r a c t i o n s of each p l a n t community. 15. The r o o t r e s e r v e s vary through the growing season. They are low i n A p r i l and May, r a p i d l y i n c r e a s e to a peak i n August, then s l o w l y d e c l i n e to October and through the winter months. 16. The s p r i n g decrease i n root r e s e r v e s c o i n c i d e s with the i n i t i a t i o n of the aboveground growth. 17. The root r e s e r v e s are g r e a t e s t w i t h i n one month of the peak standing crop. 18. The extent of the root r e s e r v e s v a r i e s between plant communities. - 141 -6. L i t e r a t u r e C i t e d Ackerman, E.A. 1941. The Koppen c l a s s i f i c a t i o n of cli m a t e i n North America. In: Ceog. Rev., V o l . 31, 1941. p. 105-111. Atmospheric Environment. 1973. Canadian normals, V o l . 1, temperature 1941-1970. Atmospheric Environment, Environment Canada. Downsview, Ontario, Canada. 186 pp. Atmospheric Environment. 1973. . Canadian normals, V o l . 2, p r e c i p i t a t i o n 1941-1970. Atmospheric Environment, Environment Canada. Downsview, Ontario, Canada. 330 pp. B e l l - I r v i n g , R. 1977. The foreshore area of Musqueam Indian Reserve No. 2. Habitat P r o t e c t i o n D i r e c t o r a t e , F i s h e r i e s Management, P a c i f i c Region, Dept. of F i s h e r i e s and Environment, Vancouver. Black, C.A., ed. 1965. Methods of s o i l a n a l y s i s . Agronomy 9. Am. Soc. Agron. Madison, V,ris., U.S.A. B r i t i s h Columbia Research Management Services D i v i s i o n . 1974. B r i t i s h Columbia Population P r o j e c t i o n s 1974-1996. B r i t i s h Columbia Research C o u n c i l . Vancouver, Canada. Burgess, T.E. 1970. Foods and h a b i t a t of four a n a t i n i d s w i n t e r i n g on the Fraser d e l t a t i d a l marshes. M.Sc. Thesis, Dept. Zoology, U n i v e r s i t y of B r i t i s h Columbia. 124 pp. Burton, B.A. 1977. Ecology of l e s s e r snow geese w i n t e r i n g on the Fraser R i v e r D e l t a T i d a l Marshes. M.Sc. Thesis, Dept. of Animal Science, U n i v e r s i t y of B r i t i s h Columbia. Burton and Jackson, 1962. A method f o r measuring sod reserves. Agron. J . , 54: 53-55 Calder, J.A., and R.L. Taylor. 1968. Fl o r a of the Queen C h a r l o t t e I s l a n d s , part I. Systematics of the va s c u l a r p l a n t s . Canada Dept. of A g r i c u l t u r e Monogram 4(1). Ottawa. 659 pp. Census of Canada. 1976. Population p r e l i m i n a r y counts. S t a t i s t i c s Canada. Ottawa, Canada. Chapman, V.J. 1960. S a l t marshes and s a l t deserts of the world. Leonard H i l l . London. 392 pp. Chapman, V.J. 1964. Coastal v e g e t a t i o n . MacMillan Co., New York. C h a t t i n , J.E. 1970. Some uses of e s t u a r i e s by waterfowl and other migratory b i r d s . Northwest Estuary ;and Coastal Zone Symp., USDI Bureau of Sport F i s h and W i l d l i f e . P o r t l a n d , Oregan. pp. 108-118. - 142 -Clements, F.E. and V.E. S h e l f o r d . 1939. Bi o - e c o l o g y . John Wiley and Sons Inc. New York. Coates, D.R., ed. 1972. C o a s t a l Geomorphology. A proceedings volume of the T h i r d Annual Geomorphology Symposia S e r i e s , held at Binghamton, New York, September 28-30, 1972. S t a t e U n i v e r s i t y of New York. Binghamton, New York. Department of the Environment, 1971. To 1970: H i s t o r i c a l streamflow summary, B r i t i s h Columbia. Inland Waters D i r e c t o r a t e , Department of the Environment. Ottawa, Canada. 394 pp. Department of the Environment. 1974. H i s t o r i c a l streamflow survey, B r i t i s h Columbia, to 1973. Inland Waters D i r e c t o r a t e , Water Resources Branch, Department of the Environment. Ottawa, Canada. 694 pp. Dorcey, A.H.J., T.G. Northcote and D.V. Ward. 1978. Are the F r a s e r marshes e s s e n t i a l to salmon? Westwater Research Centre, L e c t u r e No. 1. U n i v e r s i t y of B r i t i s h Columbia. Dunford, W.E. 1975. Space and food u t i l i z a t i o n by salmonids i n marsh h a b i t a t s of the F r a s e r R i v e r E s t u a r y . M.Sc. T h e s i s , Department of Zoology, U n i v e r s i t y of B r i t i s h Columbia. E i l e r s , H. 1975. P l a n t s , p l a n t communities, net production and t i d e l e v e l s : the e c o l o g i c a l biogeography of the Nehalem s a l t marshes, Tillamook County, Oregon. Ph.D. T h e s i s , Oregon State U n i v e r s i t y . Forbes, R.D. 1972. A f l o r a l d e s c r i p t i o n of the F r a s e r R i v e r Estuary, and Boundary and Mud Bays. B.C. F i s h and W i l d l . Br., B.C. Dept. Recrea. and Conserv. Burnaby, B.C. 94 pp. Forbes, R.D. 1972. A d d i t i o n a l catalogue to "A f l o r a l d e s c r i p t i o n of the Fraser R i v e r estuary, and Boundary and Mud bays, B.C." B.C. F i s h and W i l d l . Br. Rept. Burnaby, B.C. 20 pp. Haines, E. 1977. The o r i g i n s of d e t r i t u s i n Georgia s a l t marsh e s t u a r i e s . Oikos 29:254-260. H i l l a b y , F.B. and D.T. B a r r e t t . 1976. Vegetation communities o f a F r a s e r R i v e r s a l t marsh. T e c h n i c a l Report S e r i e s Pac/T-76-14. Habi t a t P r o t e c t i o n D i r e c t o r a t e , F i s h e r i e s and Marine S e r v i c e Department of the Environment. Vancouver. Hitchcock, C.L. and A. C r o n q u i s t . 1973. F l o r a of the P a c i f i c Northwest. U n i v e r s i t y of Washington P r e s s . S e a t t l e , U.S.A. - 143 -H o l l a n d , S.S. 1964. Landforms of B r i t i s h Columbia, a p h y s i o g r a p h i c o u t l i n e . B.C. Dept. of Mines and Petroleum Resources B u l l e t i n 48. V i c t o r i a , B.C. 138 pp. I n g l i s , S i r C C . and T.J.F. Kestner. 1958. The long-term e f f e c t s of t r a i n i n g w a l l s , r e c l a m a t i o n , and dredging on e s t u a r i e s . Proc. I n s t . C i v i l Eng. 9, Pap. (6268). Kennedy, K.A. 1978. P l a n t communities of the Cowichan R i v e r E s t u a r y . _In The Cowichan R i v e r Estuary Task Force Report r e l e a s e d November 1980. Canada Department of Environment. Vancouver. Kennedy, K.A. 1978. P l a n t communities of the Salmon River E s t u a r y . C o n s u l t a n t s r e p o r t f o r LUPAC, MacMillan B l o e d e l . Nanaimo. Keser, N. and D. St. P i e r r e . 1973. S o i l s of Vancouver I s l a n d , a compendium. B r i t i s h Columbia Forest S e r v i c e Research Note No. 56. V i c t o r i a . K i s t r i t z , R. 1978. An e c o l o g i c a l e v a l u a t i o n of Fraser estuary t i d a l marshes: The r o l e of d e t r i t u s and the c y c l i n g of elements. T e c h n i c a l Report No. 15, Westwater Research Centre. U n i v e r s i t y of B r i t i s h Columbia. K i s t r i t z , R. and I. Y e s a k i . 1979. Primary p r o d u c t i o n , d e t r i t u s f l u x , and n u t r i e n t c y c l i n g i n a sedge marsh, Fraser R i v e r Estuary. T e c h n i c a l Report No. 17. Westwater Research Centre. U n i v e r s i t y of B r i t i s h Columbia. K l i n k a , K. 1977. Guide f o r t r e e s p e c i e s s e l e c t i o n and p r e s c r i b e d burning i n the Vancouver Forest D i s t r i c t . M i n i s t r y of F o r e s t s , F o r e s t S e r v i c e Research D i v i s i o n , Vancouver F o r e s t D i s t r i c t . Vancouver, B.C. K r a j i n a , V.J. and R.C. Brooke. 1969/70. Ecology of Western North America. Volume 2, Numbers 1 and 2. M i t c h e l l Press L i m i t e d . Vancouver, B.C., Canada. Lehmann, E.J. 1974. Sewage e f f e c t s i n marine and e s t u a r i n e environments. A b i b l i o g r a p h y with a b s t r a c t s . Gov. Repts. Announcements 74 (16): 40. Levings, C D . and A.I. Moody. 1976. Studies of i n t e r t i d a l v a s c u l a r p l a n t s , e s p e c i a l l y sedge (Carex l y n g b y e i ) , on the d i s r u p t e d Squamish R i v e r d e l t a , B r i t i s h Columbia. Environment Canada, F i s h e r i e s and Marine S e r v i c e T e c h n i c a l Report No. 606. Vancouver, B.C. 51 pp. Lim, P.G. and C. L e v i n g s . 1973. D i s t r i b u t i o n and biomass of i n t e r t i d a l v a s c u l a r p l a n t s on the Squamish D e l t a . F i s h e r i e s Research Board of Canada Manuscript Report S e r i e s 1219. P.E.I. Vancouver, B.C. - 144 -MacDonald, K.B. and M.G. Barbour. 1974. Beach and s a l t marsh v e g e t a t i o n of the North American P a c i f i c Coast. In: Reimold, R.J. and W.H. Queen, 1974. B i o l o g y of Halophytes. Academic Press. New York. 605 pp. McLaren, K.A. 1972. A v e g e t a t i o n study of the i s l a n d s and a s s o c i a t e d marshes i n the South Arm of the Fraser R i v e r , B.C., from the Dease I s l a n d Tunnel to Westham I s l a n d f o r e s h o r e . F i s h and W i l d l . Br., B.C. Dept. Recrea. and Conserv. Burnaby, B.C. 54 pp. and f i g u r e s . Moody, A.I. 1978. Growth and d i s t r i b u t i o n of the v e g e t a t i o n of a southern F r a s e r d e l t a marsh. M.Sc. T h e s i s , Dept. of P l a n t Science. U n i v e r s i t y of B r i t i s h Columbia. M u e l l e r - Dombois, D. and H. E l l e n b e r g . 1974. Aims and methods of v e g e t a t i o n ecology. John Wiley and Sons. New York. 547 pp. M u l l e r , J.E. 1971. G e o l o g i c a l Reconnaissance Map of Vancouver Is l a n d and G u l f I s l a n d s . G e o l o g i c a l Survey of Canada. Open F i l e Map 61. Odum, E.P. 1961. The r o l e of the t i d a l marshes i n e s t u a r i n e p r o d u c t i o n . U n i v e r s i t y of Georgia Mar. I n s t . C o n t r i b . (29). Oguss, E., A.F. Tautz, M.E. Anderson, S.M. S t e e l e . 1975. A p a r t i a l compendium of stream data f o r Vancouver I s l a n d . F i s h e r i e s T e c h n i c a l C i r c u l a r No. 16, B r i t i s h Columbia F i s h and W i l d l . Br. P r i t c h a r d , D.W. 1967. What i s an estuary: P h y s i c a l viewpoint, pp. 3-5. _In G.H. L a u f f , ed. E s t u a r i e s . American A s s o c i a t i o n of Advanced S c i e n c e . Reimers, P.E. 1973. The l e n g t h of r e s i d e n c e of j u v e n i l e f a l l chinook salmon i n Sixes R i v e r , Oregon. Research Report Oregon F i s h Commission. 4(2 ) : 1-43. Sorensen, J.C. 1971. A framework f o r i d e n t i f i c a t i o n and c o n t r o l of resource degradation and c o n f l i c t i n the m u l t i p l e use of the c o a s t a l zone. M. T h e s i s . U n i v e r s i t y of C a l i f o r n i a , B e r k l e y . S t i c h l i n g , W. 1974. Sediment loads i n Canadian R i v e r s . T e c h n i c a l B u l l e t i n No. 74. Inland Waters D i r e c t o r a t e , Water Resources Branch, Department of the Environment. Ottawa, Canada. 27 pp. Whittaker, R.H. 1967. Gradient a n a l y s i s of v e g e t a t i o n . B i o l . Rev. 49: 207-264. Yamanaka, K. 1975. P r o d u c t i v i t y of t i d a l marsh, Fr a s e r River f o r e s h o r e . M.Sc. 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. - 145 -Zar, J.H. 1974. B i o s t a t i s t i c a l A n a l y s i s . P r e n t i c e - H a l l , Inc. Englewood C l i f f s , New Jersey, U.S.A. 620 pp. Zedler, J.B. 1977. S a l t marsh community s t r u c t u r e i n the T i j u a n a Estuary, C a l i f o r n i a . E s t u a r i n e and C o a s t a l Marine Science 5: 39-53. - 146 -Appendix 1. Lo c a t i o n s , i n l a t i t u d e and l o n g i t u d e , of the study areas Estuary L a t i t u d e Longitude Goldstream 48° 29 * 10" 123° 32'50" Cowichan 48° 46 *10" 125° 38'15" Chenainus 48° 53 *40" 123° 41' 5" Nanaimo 49° 7 '50" 123° 53'30" Nanoose 49° 16 *15" 124° 11'45" B o n e l l 49° 15 ' 50" 124° 11'40" Englishman 49° 19 '4R" 124° 17'30" L i t t l e Qualicum 49° 21 1 ' 50" 124° 29'15" Bi g Qualicum 49° 23 1 '55" 124° 36'30" Courtenay 49° 36 ' i 125° Oyster 49° 45'12" 125° 7' 0" Campbell 49° 55 ' 35" 125° 15'35" Salmon 50° 15 1 '35" 125° 55 ' 30" Adam-Eve 50° 27 ' 45" 126° 16'50" T s i t i k a 50° 29'40" 126° 34'50" Kokish 50° 33 ' 42" 126° 52'15" Nimpkish 50° 35* 127° Cluxewe 50° 40'50" 127° 10'30" Ouatse 50° 41' 15" 127° 29'10" Ki ngcome 50° 55 ' 126° 20' - 147 -Appendix 2 Glossary of terms Emergent veg e t a t i o n - phanerogamic vegetation subject t o p e r i o d i c f l o o d i n g by the sea t i d e . In t h i s study the presence of e e l g r a s s , a s u b t i d a l p l a n t s p e c i e s , and rockweed, an algae, are noted when they occur with emergent ve g e t a t i o n . Estuary - semi-enclosed c o a s t a l bodies of water which have a f r e e connection with the open sea and v/ithin which sea water i s measurably d i l u t e d with freshwater from land drainage ( P r i t c h a r d , 1967). Estuarine marsh - lands covered by phanerogamic vegetation subject to p e r i o d i c f l o o d i n g by the sea t i d e . P h y s i o l o g i c a l drought - r e s u l t s from osmotic s t r e s s when the e x t e r n a l water p o t e n t i a l i s lowered below that of the pl a n t c e l l s . Standing crop - aboveground matter i n a s p e c i f i c area, u s u a l l y one square meter. S a l t marsh - lands covered by phanerogamic vegetation subject to p e r i o d i c f l o o d i n g by the sea t i d e (Chapman 1960, 1964). In t h i s study i t r e f e r s to marshes with high s o i l s a l i n i t i e s (> 20 p p t . ) . T i d a l marsh - land covered by p e r i o d i c f l o o d i n g by phanerogamic vegetation the sea t i d e . subject to Appendix 3. A e r i a l photographs used i n the study Estuary Goldstream B.C. 7401 No.166 1 Cowichan Ri v e r 67068-67105 .1 Chemainus River 67106-67139 1 Nanaimo Ri v e r 67159-67192 1 Nanoose-Bonell Creeks 67193-67202 1 Englishman Ri v e r 67203-67210 1 L i t t l e Qualicum River 67211-67219 1 Big Qualicum River 67220-G7227 1 Courtenay River 67263-67295 1 Oyster R i v e r G7296-67303 1 Campbell River 67304-67320 1 Salmon River 66881-66908 1 Adam-Eve River 66909-66923 1 T s i t i k a River 66924-66932 1 Kokish River 66933-66937 1 Nimpkish River 66938-66972 1 Cluxewe River . 66985-66903 1 Quatse River 66994-67007 1 Kingcome River B.C. 7270 No.076 1 Scale Date Source cm = 634m B.C. Government cm = 48 m March 22/73 P.S.C* cm = 48 m March 22/73 P.S.C. cm 48 m March 22/73 P.S.C. cm = 43 m March 22/73 P.S.C. cm 48 m March 22/73 P.S.C. cm = 48 m March 22/73 P.S.C. cm = 48 m March 22/73 P.S.C. cm = 48 m March 22/73 P.S.C. cm = 40 m March 22/73 P.S.C. cm = 48 m March 22/73 P.S.C. cm = 40 m March 22/73 P.S.C. cm = 48 ra March 22/73 P.S.C. cm = 48 m March 22/73 P.S.C. cm -- 48 m March 22/73 P.S.C. cm = 48 m March 22/73 P.S.C. cm = 48 m March 22/73 P.S.C. cm = 40 m March 22/73 P.S.C. cm = 158m B.C. Government • P a c i f i c Survey Corporation - 149 -Appendix 4 . L i s t of p l a n t s p e c i e s TREES Alnus rubra Bong. Pyrus fusca Raf. Pseudotsuga m e n z i e s i i ( M i r b e l ) Franco P i c e a s i t c h e n s i s (Bong.) C a r r . Tsuga h e t e r o p h y l l a (Raf.) Sarg. Populus t r i c h o c a r p a T. & G. Sorbus s c o p u l i n a Greene Cornus s t o l o n i f e r a Michx. Acer macrophyllum Pursh Thuja p l i c a t a Donn. Pinus c o n t o r t a v a r . c o n t o r t a Dougl. Quercus quarryana Dougl. Abies g r a n d i s (Dougl.) Forbes Pyrus malus L. Arbutus m e n z i e s i i Pursh. Rhamnus purshiana DC. Cornus n u t t a l l i i Aud. Juniperus scopulorum Sarg. Prunus sp. L. Prunus emarginata (Dougl.) Walp. SHRUBS  S a l i x sp. L. Crataegus d o u g l a s i i L i n d l . Rosa nutkana P r e s l . Rubus l a c i n i a t u s W i l l d . Red Alder P a c i f i c Crabapple Douglas F i r S i t k a Spruce Western Hemlock Cottonwood Mountain-ash Red-osier Dogwood Common Maple Western Red Cedar Lodgepole Pine Garry Oak Grand F i r Apple Arbutus ' Cascara Dogwood Junipe r C u l t i v a t e d Cherry Chokecherry Willow Black Hawthorn Nootka Rose Evergreen B l a c k b e r r y - 1 5 U -Symphoricarpos albus (L.) Blake C y t i s u s s c o p a r i u s (L.) Link Ribes sanguineum Pursh Rubus p a r v i f l o r u s Nutt. Rubus u r s i n u s Cham. & Schlecht L o n i c e r a i n v o l u c r a t a ( R i c . ) Banks Rubus s p e c t a b i l i s Pursh Qsmaronia c e r a s i f o r m i s (T. & G.) Greene G a u l t h e r i a s h a l l o n Pursh Vaccinium p a r v i f o l i u m Smith Vaccinium o v a l i f o l i u m Smith Ribes sp. L. A r c t o s t a p h y l o s nevadensis Gray S p i r a e a d o u g l a s i i Hook Rosa gymnocarpa Nutt. Physocarpus c a p i t a t u s (Pursh) Kuntze Myrica gale L. Sambucus racemosa L. H o l o d i s c u s d i s c o l o r (Pursh) Maxim. S a l i x hookeriana B a r r a t t M enziesia f e r r u g i n e a Smith Crataegus oxyacantha L. B e r b e r i s nervosa Pursh L o n i c e r a c i l i o s a (Pursh) DC. Amelanchier a l n i f o l i a Nutt. Rubus pedatus J.E. Smith S a l i x s c o u l e r i a n a B a r r a t t Snowberry Broom Red Currant Thimbleberry P a c i f i c B l a c k b e r r y Black Twin-berry Salmonberry Indian Plum S a l a l Huckleberry Blueberry Gooseberry K i n n i k i n n i c k Hardhack L i t t l e Wild Rose Ninebark Sweet Gale E l d e r b e r r y Ocean-spray Hooker Willow Mock Azalea Hawthorn Oregongrape Orange Honeysuckle S e r v i c e b e r r y F i v e l e a v e d Bramble Scouler Willow - 151 -B e r b e r i s a q u i f o l i u m Pursh Linnaea b o r e a l i s L. L o n i c e r a h i s p i d u l a ( L i n d l . ) Dougl. Pachistima m y r s i n i t e s (Pursh) Raf. Ribes sanguineum Pursh Ulex europaeus L. FORBS Myosotis l a x a Lehm. Ranunculus cymbalaria Pursh. Epilobium w a t s o n i i Barbey Lactuca b i e n n i s (Moench) Fern. Myosotis d i s c o l o r Pers. V i o l a sp. L. I r i s pseudacorus L. T r i e n t a l i s l a t i f o l i a Hook Cerastium arvense L. Medicago s a t i v a L. Delphinium m e n z i e s i i DC. Lomatium nu d i c a u l e (Pursh) C o u l t . & Rose S i l e n e n o c t i f l o r a L. Ranunculus a c r i s L. T e l l i m a g r a n d i f l o r u m (Pursh) Dougl. Montia p e r f o l i a t a (Donn) Howell Rumex maritimus L. Oregongrape Twinflower Hairy Honeysuckle Pachistima Red Currant Gorse Forget-me-not Seaside Buttercup Watson's Willow-herb T a l l Blue L e t t u c e Yellow and Blue Forget-me-not V i o l e t Yellow I r i s S t a r f l o w e r Mouse-ear Chickweed Medic Menzies 1 Delphinium D e s e r t - p a r s l e y C a t c h f l y Meadow Buttercup Fringecup Miner's L e t t u c e Seaside Dock - 1 5 2 -B e l l i s p e rennis L. Ranunculus orthorhynchus Hook T r i l l i u m ovatum Pursh T r i f o l i u m t ridentatum L i n d l . A chlys t r i p h y l l a (Smith) DC. Streptopus a m p l e x i f o l i u s (L.) DC. Habenaria d i l a t a t a (Pursh) Hook A t r i p l e x p a t u l a L. Suaeda maritima (L.) Dumort. S t e l l a r i a humifusa Rottb. A n g e l i c a sp. L. A r e n a r i a macrophylla Hook A r e n a r i a p a l u d i c o l a Robins A r e n a r i a s t r i c t a Michx. Barbarea v u l g a r i s R. Br. Cardamine i n t e g r i f o l i a (Nutt.) Greene Cerastium viscosum L. C l i n t o n i a u n i f l o r a ( S c h u l t . ) Kunth. C o l l i n s i a p a r v i f l o r a L i n d l . D i c e n t r a formosa (Andr.) Walp. Draba sp. L. Erythronium oregonum Applegate H e l i o t r o p i u m curassavicum L. L l o y d i a s e r o t i n a (L.) Sweet P l e c t r i s congesta ( L i n d l . ) DC. Polypodium s c o u l e r i Hook. & Grev. S a n i c u l a c r a s s i c a u l i s Poepp. Daisy Buttercup T r i l l i u m Sand C l o v e r V a n i l l a l e a f T w i s t e d - s t a l k Bog-candle Saltbush S e a b l i t e Spreading Starwort A n g e l i c a Sandwort Sandwort Slender Sandwort Wintercress B i t t e r c r e s s S t i c k y Chickweed C l i n t o n i a Blue-eyed Mary Bleeding Heart Draba D o g t o o t h - v i o l e t Seaside H e l i o t r o p e L l o y d i a Rosy Pink Polypody S a n i c l e - 153 -Sedum s p a t h u l i f o l i u m Hook. S t e l l a r i a c r i s p a Cham. & S c h l e c h t . Apocynum androsaemifolium L. Z o s t e r a marina L. Senecio s y l v a t i c u s L. C o c h l e a r i a o f f i c i n a l i s L. Montia s i b i r i c a (L.) Howell L i l a e o p s i s o c c i d e n t a l i s C o u l t . & Rose Polypodium g l y c y r r h i z a D.C. Eat. C o r a l l o r h i z a mertensiana Bong. T r i f o l i u m w o r m s k j o l k i i Lehm. F r a g a r i a c h i l o e n s i s (L.) Duchesne Convolvulus sepium L. Mimulus g u t t a t u s DC. Mentha s p i c a t a L. C i c u t a d o u g l a s i i (DC.) C o u l t . & Rose Lemna minor L. Typha l a t i f o l i a L. Montia p a r v i f o l i a (Moc.) Greene S i d a l c e a h e n d e r s o n i i Wats. M e l i l o t u s alba Desr. P r u n e l l a v u l g a r i s L. Ranunculus repens L. H e l i o p s i s h e l i a n t h o i d e s (L.) Sweet Sedum C r i s p e d Sandwort Dogbane E e l - g r a s s Groundsel Spoonwort S i b e r i a n Miner's L e t t u c e L i l a e o p s i s L i c o r i c e - f e r n Western C o r a l -r o o t Springbank C l o v e r C o a s t a l Strawberry Morning-glory Yellow Monkey-flower Spearmint Water-hemlock Duckweed C a t - t a i l Miner's Lettu c e Henderson's Checker-mallow White Sweet-c l o v e r S e l f - h e a l Creeping Buttercup Ox-eye Daisy - 1 5 4 -Camassia l e i c h t l i n i i (Baker) Wats. Dodecatheon pulchellum (Raf.) M e r r i l l L i l i u m columbianum Hanson Plantago major L. Plantago macrocarpa Cham. & Schl e c h t Lupinus sp. L. S i s y r i n c h i u m a n g u s t i f o l i u m M i l l . C a s t i l l e j a l e v i s e c t a Greenm. Mentha a r v e n s i s L. U r t i c a d i o i c a L. Galium t r i f i d u m L. Adenocaulon b i c o l o r Hook Oenanthe sarmentosa P r e s l . T i a r e l l a t r i f o l i a t a v a r . t r i f o l i a t a L. Taraxacum o f f i c i n a l e Weber Gymnocarpium d r y o p t e r i s (L.) Newm. Athyrium f i l i x - f e m i n a (L.) Roth. Blechnum s p i c a n t (L.) Roth Geum macrophyllum W i l l d . D i g i t a l i s purpurea L. Osmorhiza c h i l e n s i s H. & A. A l l i u m g e y e r i v a r . tenerum Jones C a p s e l l a b u r s a - p a s t o r i s (L.) Medic. Ranunculus uncinatus D. Don Galium sp. L. Galium bo r e a l e L. Cornus canadensis L. Camas Sh o o t i n g - s t a r Columbia L i l y Common P l a n t a i n P l a n t a i n Lupine Blue-eyed Grass Golden Indian-p a i n t b r u s h F i e l d Mint S t i n g i n g N e t t l e Small Bedstraw P a t h f i n d e r Water-parsley Foamflower Common Dandelion Oak-fern Lady-fern Deer-fern Avens Foxglove Sweet-root Geyer's Onion Shepherd's purse L i t t l e Buttercup Bedstraw Northern Bedstraw Bunchberry - 155 -L y s i c h i t u m americanum Hulten & St.John Skunk Cabbage S t e l l a r i a media (L.) C y r i l l . Chickweed A r a b i s g l a b r a (L.) Bernh. Towermustard Armeria maritima ( M i l l . ) W i l l d . Sea-pink Ruppia maritima L. D i t c h - g r a s s Asparagus o f f i c i n a l i s L. Asparagus P o t e n t i l l a p a c i f i c a Howell C i n q u e f o i l Plantago l a n c e o l a t a L. Ribgrass T r i g l o c h i n maritimum L. Arrow-grass Plantago maritima L. Seaside P l a n t a i n T r i f o l i u m repens L. White C l o v e r C i r s i u m arvense (L.) Scop. Canada T h i s t l e Hypochaeris r a d i c a t a L. Hairy Cats-ear A c h i l l e a m i l l e f o l i u m L. Yarrow C o t u l a c o r o n o p i f o l i a L. Brass Buttons Glaux maritima L. Sea-milkwort S a l i c o r n i a v i r g i n i c a L. Saltwort Sonchus a r v e n s i s L. M i l k - t h i s t l e G r i n d e l i a i n t e g r i f o l i a DC. Gumweed Ambrosia chamissonia (Less.) Greene Ragweed E r i g e r o n p h i l a d e l p h i c u s L. P h i l a d e l p h i a Fleabane Polygonum s p e r g u l a r i a e f o r m e Meisn. Spurry Knotweed Lactuca m u r a l i s (L.) Fresen. Wall L e t t u c e Rumex c r i s p u s L. Cur l y Dock Ranunculus repens L. Buttercup Creeping T r i f o l i u m pratense L. Red C l o v e r Equisetum arvense L. Common H o r s e t a i l - 156 -Pt e r i d i u m a q u i l i n u m (L.) Kuhn Lathyrus p a l u s t r i s L. Epilobium a n g u s t i f o l i u m L. Chenopodium album L. Anaphalis margaritacea (L.) B. & H. Ci r s i u m v u l g a r e ( S a v i ) Tenore Rumex a c e t o s e l l a L. Geranium molle L. Hypericum formosum H.B.K. Lotus c o r n i c u l a t u s L. Arctium minus ( H i l l ) Bernh. Barbarea o r t h o c e r a s Ledeb. Stachys cooleyae H e l l e r P e t a s i t e s f r i g i d u s (L.) F r i e s . T r i f o l i u m dubium S i b t h . A s t e r h e s p e r i u s Gray Rumex s a l i c i f o l i u s Weinm. Cuscuta s a l i n a Engelm. S p e r g u l a r i a canadensis (Pers.) G. Don Heracleum lanatum Michx. F r i t i l l a r i a camschatcensis' (L.) Ker.-Gawl, V i c i a gigantea Hook. Maianthemum d i l a t a t u m (Wood) Ne l s . & Macbr. Sium suave Walt. T r i e n t a l i s a r c t i c a F i s c h . Polystichum munitum ( K a u l f . ) P r e s l . Bracken Marsh Pea Fireweed Pigweed P e a r l y -e v e r l a s t i n g Common T h i s t l e Sour Weed Dovefoot Geranium St. John's-wort B i r d s f o o t - t r e f o i l Common Burdock Winte r c r e s s C o l l e y ' s Hedge-n e t t l e C o l t s f o o t Least Hop C l o v e r Marsh A s t e r Willow Dock Salt-marsh Dodder Canada Sandspurry Cow-parsnip Chocolate L i l y Giant Vetch F a l s e L i l y - o f - t h e -v a l l e y Water-parsnip S t a r f l o w e r Sword-fern - 157 -A q u i l e g i a formosa F i s c h . Galium asperrimum Gray Galium aparine L. Hieracium a l b i f l o r u m Hook. Ranunculus sp. L. Polygonum a v i c u l a r e L. B r a s s i c a campestris L. Symphytum asperum Lepech. Myosotis scorpioid.es L. GRASSES Holcus lanat u s L. D i s t i c h l i s s p i c a t a (L.) Greene Elymus m o l l i s T r i n . Phleum pratense L. A g r o s t i s alba L. P h a l a r i s aruridinacea L. Deschampsia c e s p i t o s a (L.) Beauv. D a c t y l i s glomerata L. Poa p r a t e n s i s L. Festuca o c c i d e n t a l i s Hook Bromus s e c a l i n u s L. Anthoxanthum odoratum L. Me l i c a subulata ( G r i s e b . ) S c r i b n . G l y c e r i a o c c i d e n t a l i s ( P i p e r ) Nels. A g r o s t i s alba v a r . s t o l o n i f e r a (L.) Red Columbine Rough Bedstraw Bedstraw White-flowered Hawkweed Buttercup Doorweed Common Mustard Rough Comfrey Common Forget-me-not V e l v e t - g r a s s S a l t g r a s s , Wildrye Timothy Bentgrass Reed Canarygrass Tuf t e d H a i r g r a s s Orchard-grass Kentucky Bluegrass Western Fescue Cheat Sweet V e r n a l g r a s s M e l i c Western Mannagrass Smith Bentgrass - 158 -Agropyron repens (L.) Beauv. Couch Grass Festuca rubra L. Red Fescue Elymus glaucus B u c k l . Blue Wildrye Hordeum murinum L. Mouse B a r l e y Hordeum brachyantherum Nevski. Meadow Ba r l e y Bromus s i t c h e n s i s T r i n . Brome-grass G l y c e r i a b o r e a l i s (Nash) Batch Northern Mannagrass A g r o s t i s t e n u i s S i b t h . C o l o n i a l Bentgrass Poa compressa L. Canada Bluegra Poa sp. L. Bluegrass Calamagrostis canadensis (Michx.) Beauv. B l u e j o i n t Reedgrass Bromus p a c i f i c u s Shear P a c i f i c Brome Bromus tectorum L. Brome-grass A i r a sp. L. Ha i r g r a s s Bromus inermis Leys. Smooth Brome Lolium perenne L. Ryegrass Festuca bromoides L. Fescue Festuca subulata T r i n . Nodding Fescue Festuca p r a t e n s i s Huds. Meadow Fescue H i e r o c h l o e odorata (L.) Beauv. Seneca Grass Cynosurus c r i s t a t u s L. Crested Dog's-t a i l F estuca sp. L. Fescue Bromus m o l l i s L. Soft Brome Bromus c a r i n a t u s H. &. A. Brome - 159 -SEDGES Carex l y n g b y e i Hornem. S c i r p u s microcarpus P r e s l . E l e o c h a r i s p a l u s t r i s (L.) R. & S. Carex pansa B a i l e y Carex p l u r i f l o r a Hulten S c i r p u s americanus Pers. Carex l e n t i c u l a r i s v a r . l i m n o p h i l a (Holm) Cronq. S c i r p u s acutus Muhl. S c i r p u s maritimus L. Carex o e d e r i Retz. Carex obnupta B a i l e y E l e o c h a r i s p a u c i f l o r a ( L i g h t f . ) Link Carex r o s t r a t a Stokes RUSHES Juncus b a l t i c u s W i l l d . Juncus e f f u s u s L. Juncus g e r a r d i i L o i s e l . Juncus acuminatus Michx. L u z u l a d i v a r i c a t a Wats. Juncus a r t i c u l a t u s L. Juncus e n s i f o l i u s v a r . e n s i f o l i u s Wikst, Juncus f a l c a t u s E. Meyer Lyngby's Sedge S m a l l - f r u i t B u l r u s h Spike-rush Sand-dune Sedge Sedge Three-square B u l r u s h Sedge Hardstem Bulrush Bulrush Green Sedge Slough Sedge Few-flowered Spike-rush Beaked Sedge B a l t i c Rush Common Rush Mud Rush Tapered Rush Woodrush J o i n t e d Rush Dagger-leaf Rush S i c k l e - l e a v e d Rush Luz u l a s p i c a t a (L.) DC. Spiked Woodrush - 160 -Appendix 5. D e s c r i p t i o n s , maps and area d e t e r m i n a t i o n s of the p l a n t communities on the e s t u a r i e s s t u d i e d . - 161 -Figure 4 . Map showing the d i s t r i b u t i o n of the plant communities on the Goldstream Estuary Legend Scale lcm=32m Plant Community 1 Plantago lanceolata-Rumex s a l i c i f o l i u s - H o r d e u m  brachyantherum- Chenopodiun album 2 C y t i s u s scoparius - Rosa nutkana 3 A g r o s t i s a l b a 4 Aster hesperius - Juncus b a l t i c u s 5 Hordeum brachyantherum 6 Carex lyngbyei 7 D i s t i c h l i s s p i c a t a - S a l i c o r n i a v i r g i n i c a S Hordeum murinum - Carex lyngbyei 9 S a l i c o r n i a v i r g i n i c a L t i d a l b u i l d i n g M N SL marina nature house s u b t i d a l road T a b l e 7. D e s c r i p t i o n o f t h e p l a n t c o m m u n i t i e s on t h e G o l d s t r e a m E s t u a r y , September, 1976. 1 P l a n t a g o l a n c e o l a t a - Rumex s a l i c i f o l i u s - Hordeum b r a c h y a n t h e r u m - Chenopodium album ?, C y t i s u s s c o p a r i u s - Rosa n u t k a n a 3 A g r o s t i s a l b a 4 A s t e r h e s p e r i u s - J u n c u s b a l t i c u s 5 Hordeum b r a c h y a n t h e r u m 6 C a r e x l y n g b y e i 7 D i s t i c h l i s s p i c a t a - S a l i c o r n i a v i r g i n i c a 8 Hordeum murinum - Carex l y n g b y e i 9 S a l i c o r n i a v i r g i n i c a F o l l o w i n g i s a l i s t o f t h e s p e c i e s i n each p l a n t community i n c l u d i n g d o m i n a n t s p e c i e s i n d i c a t e d by a D and subdominant s p e c i e s i n d i c a t e d by an S. TREES 1 Alnus rubra Bong. (Red A l d e r ) Pseudotsuga m e n z i e s i i ( M i r b e l ) Franco (Douglas F i r ) SHRUBS C y t i s u s s c o p a r i u s (L.) Link x (Broom) Rosa nutkana P r e s l . (Nootka Rose) Rubus u r s i n u s Cham. & Schlecht ( P a c i f i c B l a c k b e r r y ) S a l i x spp. L. (Willow) L o n i c e r a i n v o l u c r a t a (Rich.) Banks (Black Twin-berry) Rubus s p e c t a b i l i s Pursh (Salmonberry) Pyrus f u s c a Raf. ( P a c i f i c Crabapple) Crataegus d o u g l a s i i L i n d l . (Black Hawthorn) 2 Community 3 .4 5 x x D D x X TREES (cont.) Osmaronia c e r a s i f o r m i s (T.&G.) Greene (Indian Plum) FORBS Plantago l a n c e o l a t a L. (Ribgrass) Hypochaeris r a d i c a t a L. (Hairy Cats-ear) Chenopodium album L. (Pigweed) Anaphalis margaritacea (L.) B. & H. ( P e a r l y - e v e r l a s t i n g ) A c h i l l e a m i l l e f o l i u m L. (Yarrow) C i r s i u m vulgare (Savi) Tenore (Common T h i s t l e ) Rumex a c e t o s e l l a L. (Sour, weed) Geranium molle L. (Dovefoot Geranium) Rumex c r i s p u s L. (Cu r l y Dock) Community 1 2 3 4 5 6 7 8 9 S x x x S X X X X X X X X X X X X FORBS (cont.) P t e r i d i u m aquilinum (L.) Kuhn (Bracken) T r i f o l i u m pratense L. (Red Clover) Asparagus o f f i c i n a l i s L. (Asparagus) Hypericum formosum H.B.K. (S t . John's-wort) Arctium minus ( H i l l ) Bernh. (Burdock) C i r s i u m arvense (L.) Scop. (Canada T h i s t l e ) Barbarea orthoceras Ledeb. (Wintercress) Stachys cooleyae H e l l e r ( S t i n g i n g N e t t l e ) P e t a s i t e s f r i g i d u s (L.) F r i e s ( C o l t s f o o t ) P o t e n t i l l a p a c i f i c a Howell ( C i n q u e f o i l ) T r i f o l i u m repens L. (White Clover) Community 5 6 7 8 9 i i—1 FORBS (cont.) T r i f o l i u m dubium S i b t h . (Least Hop Clover) G r i n d e l i a i n t e g r i f o l i a DC. (Gumweed) E r i g e r o n p h i l a d e l p h i c u s L. ( P h i l a d e l p h i a Fleabane) A s t e r hesperius Grey (Marsh Aster) Plantago maritima L. (Seaside P l a n t a i n ) S a l i c o r n i a v i r g i n i c a L. (S a l t w o r t ) Rumex s a l i c i f o l i u s Weinm. (Willow Dock) T r i g l o c h i n maritimum L. (Arrow-grass) Glaux maritima L. (Sea-milkwort) Cuscuta s a l i n a Engelm. (Salt-marsh Dodder) S p e r g u l a r i a canadensis (Pers.) G. Don (Canada Sandspurry) Community 5 6 7 8 9 S D D X FORBS (cont.) Heracleum lanatum Michx. (Cow-parsnip) Epilobium w a t s o n i i Barbey (Watson's Willow-herb) A t r i p l e x p a t u l a L. (Saltbush) GRASSES D a c t y l i s glomerata L. (Orchard-grass) Poa p r a t e n s i s L. (Kentucky Bluegrass) Festuca o c c i d e n t a l i s Hook (Western Fescue) Bromus s e c a l i n u s L (Cheat) Anthoxanthum odoratum L. (Sweet v e r n a l g r a s s ) D i s t i c h l i s s p i c a t a (L.) Greene ( S a l t g r a s s ) Deschampsia c e s p i t o s a (L.) Beauv. (Tufted H a i r g r a s s ) Community 2 3 4 5 6 7 8 9 D GRASSES (cont.) M e l i c a subulata (Griseb.) S c r i b n . ( M e l i c ) A g r o s t i s alba L. (Bentgrass) Hordeum brachyantherum Nevski. (Meadow Bar l e y ) Hordeum murinum L. (Mouse Ba r l e y ) Agropyron repens (L.) Beauv. (Couch Grass) Community 1 2 3 4 5 x D x x S x D x X SEDGES Carex lyn g b y e i Hornem RUSHES Juncus b a l t i c u s W i l l d . ( B a l t i c Rush) - 169 -Table 8. Area of each p l a n t community on the Goldstream E s t u a r y Plant Community Area Percent ( h e c t a r e s ) (%) As t e r hesperius-Juncus b a l t i c u s 1.4 23.3 Hordeum murinum-Carex l y n g b y e i 1.1 18.3 C y t i s u s scoparius-Rosa nutkana .7 11.7 Hordeum brachyantherum .6 10.0 S a l i c o r n i a v i r i g i n i c a .6 10.0 A g r o s t i s a l b a .6 10.0 Carex l y n g b y e i .4 6.6 Su b t o t a l 5.4 89.9 Plantago lanceolata-Rumex s a l i c i f o l i u s -Hordeum braehyantherurn-Chenopodium album .3 5.0 D i s t i c h l i s s p i c a t a - S a l i c o r n i a v i r g i n i c a .3 5.0 TOTAL 6.0 99.9 - 1 7 0 -F i g u r e 5 . M a p s h o w i n g t h e d i s t r i b u t i o n o f t h e p l a n t c o m m u n i t i e s o n t h e C o w i c h a n R i v e r E s t u a r y -L e g e n d S c a l e l c m = 4 8 m P l a n t C o m m u n i t y 1 C a r e x l y n g b y e i 2 G l a u x m a r i t i m a 3 D y k e v e g e t a t i o n R u b u s l a c i n i a t u s - C y t i s u s s c o p a r i u s -R u b u s u r s i n u s 4 D i s t i c h l i s s p i c a t a - T r i g l o c h i n m a r i t i m u m - G l a u x m a r i t i m a -S c i r p u s a m e r i c a n u s 5 C a r e x l y n g b y e i - A g r o p y r o n r e p e n s 6 J u n c u s b a l t i c u s 7 T y p h a l a t i f o l i a 8 D i s t i c h l i s s p i c a t a 9 S c i r p u s a c u t u s 10 D i s t i c h l i s s p i c a t a - J u n c u s b a l t i c u s 11 P o a c o m p r e s s a - A c h i l l e a m i l l e f o l i u m - A t r i p l e x p a t u l a 12 C a r e x l y n g b y e i - D i s t i c h l i s s p i c a t a 13 B a r b a r e a o r t h o c e r a s - H o l c u s l a n a t u s 14 A g r o p y r o n r e p e n s - C i r s i u m a r v e n s e 15 E l y m u s m o l l i s 16 J u n c u s b a l t i c u s - C a r e x l y n g b y e i - D e s c h a m p s i a c e s p i t o s a A a g r i c u l t u r e d y k e HIM B l o g b o o m r a i l w a y - x X K x D d r e d g e d I i n d u s t r i a l r o a d -L t i d a l L B C l a w n b o w l i n g c l u b t r a n s e c t | _ 171 -F i g u r e 5 (cont.) LD log dump SL t i d a l b u i l d i n g s urban Table 9. D e s c r i p t i o n of the plant communities on the Cowichan River E s t u a r y , May 1976 1 Carex l y n g b y e i 2 Glaux maritima 3 Rubus l a c i n i a t u s - C y t i s u s s c o p a r i u s - Rubus u r s i n u s dyke v e g e t a t i o n 4 D i s t i c h l i s s p i c a t a - T r i g l o c h i n maritimum - Glaux maritima - S c i r p u s americanus 5 Carex l y n g b y e i - Agropyron repens 6 Juncus b a l t i c u s 7 Typha l a t i f o l i a 8 D i s t i c h l i s s p i c a t a 9 S c i r p u s acutus 10 D i s t i c h l i s s p i c a t a - Juncus b a l t i c u s 11 Poa compressa - A c h i l l e a m i l l e f o l i u m - A t r i p l e x p a t u l a 12 Carex lyng b y e i - D i s t i c h l i s s p i c a t a 13 Barbarea orthoceras - Holcus lanat u s 14 Agropyron repens - C i r s i u m arvense 15 Elymus moll i s 16 Juncus b a l t i c u s - Carex lyngbyei - Deschampsia c e s p i t o s a Following i s a l i s t of the species i n each plant community i n c l u d i n g dominant s p e c i e s i n d i c a t e d by a D and subdominant species i n d i c a t e d by an S. 1 TREES Pyrus fusca Raf. ( P a c i f i c Crabapple) Pseudotsuga m e n z i e s i i ( M i r b e l ) Franco (Douglas F i r ) Populus t r i c h o c a r p a T. & G. (Cottonwood) Alnus rubra Bong. (Red Alder) Acer macrophyllum Pursh. (Common Maple) Arbutus m e n z i e s i i Pursh (Arbutus) Community 9 10 11 1 SHRUBS C y t i s u s scoparius (L.) Link (Broom) Rubus urs i n u s Cham. & Schlecht ( P a c i f i c Blackberry) Rosa nutkana P r e s l . (Nootka Rose) Symphoricarpos albus (L.) Blake (Snowberry) Physocarpus c a p i t a t u s (Pursh) Kuntze (Ninebark) L o n i c e r a i n v o l u c r a t a (Rich.) Banks (Black Twin-berry) Sambucus racemosa L. ( E l d e r b e r r y ) Rubus l a c i n i a t u s W i l l d . (Evergreen Blackberry) Community 3 4 5 6 7 8 9 10 11 S S x S x X S FORBS P o t e n t i l l a p a c i f i c a Howell ( C i n q u e f o i l ) L i l a e o p s i s o c c i d e n t a l i s C o u l t . & Rose ( L i l a e o p s i s ) T r i g l o c h i n maritimum L. (Arrow-grass) Glaux maritima L. (Sea-milkwort) Plantago l a n c e o l a t a (Ribgrass) Epilobium a n g u s t i f o l i u m L. (Fireweed) T r i e n t a l i s l a t i f o l i a Hook ( S t a r f l o w e r ) P t e r i d i u m aquilinum (L.) Kuhn. (Bracken) Rumex c r i s p u s L. ( C u r l y Dock) A c h i l l e a m i l l e f o l i u m L. (Yarrow) T r i f o l i u m repens L. (White Clover) ommuni 9 10 FORBS (cont.) T r i f o l i u m pratense L. (Red C l o v e r ) S a l i c o r n i a v i r g i n i c a L. (Sal t w o r t ) E r i g e r o n p h i l a d e l p h i c u s L. ( P h i l a d e l p h i a Fleabane) Heracleum lanatum Michx. (Cow-parsnip) Typha l a t i f o l i a L. ( C a t - t a i l ) T r i f o l i u m w o r m s k j o l d i i Lehm. (Springbank C l o v e r ) Equisetum arvense L. (Common H o r s e t a i l ) C i r s i u m arvense (L.) Scop. (Canada T h i s t l e ) Plantago macrocarpa Cham. & S c h l e c h t ( P l a n t a i n ) Taraxacum o f f i c i n a l e Weber (Common Dandelion) F r i t i l l a r i a camschatcensis (L.) Kev.-Gawl. (Chocolate L i l y ) Community 4 5 6 7 8 9 10 11 12 13 14 15 16 x x x x x x x x x x x x x x D x X X X X x x x D x x x x X X X X FORBS (cont.) Rumex a c e t o s e l l a L. (Sour Weed) Plantago maritima L. (Seaside P l a n t a i n ) S t e l l a r i a media (L.) C y r i l l . (Chickweed) C o t u l a c o r o n o p i f o l i a L. (Brass Buttons) Barbarea orthoceras Ledeb. (Wintercress) Mentha a r v e n s i s L. ( F i e l d Mint) Lathyrus p a l u s t r i s L. (Marsh Pea) U r t i c a d i o i c a L. ( S t i n g i n g N e t t l e ) Myosotis d i s c o l o r Pers. (Yellow and Blue Foreget-me-not) V i o l a sp. L. ( V i o l e t ) Asparagus o f f i c i n a l i s L. (Asparagus) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 D x x x FORBS (cont.) Hypericum formosum H.B.K. ( S t . John's-wort) Cerastium arvense L. (Mouse-ear Chickweed) V i c i a gigantea Hook (Giant Vetch) Lemna minor L. (Duckweed) Ranunculus sp. L. (Buttercup) I r i s pseudacorus L. (Yellow I r i s ) Ranunculus repens L. (Creeping Buttercup) Camassia l e i c h t l i n i i (Baker) Wats. (Camas) Myosotis laxa Lehm. (Forget-me-not) B r a s s i c a campestris L. (Common Mustard) A t r i p l e x p a t u l a L (Saltbush) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 S FORBS (cont.) S t e l l a r i a humifusa Rottb. (Spreading Starwort) GRASSES D i s t i c h l i s s p i c a t a (L.) Greene ( S a l t g r a s s ) -Bromus tectorum L. (Brome-grass) A i r a sp. (H a i r g r a s s ) D a c t y l i s glomerata L. (Orchard-grass) Poa compressa L. (Canada Bluegrass) Agropyron repens (L.) Beauv. (Couch Grass) Deschampsia c e s p i t o s a (L.) Beauv. (Tu f t e d H a i rgrass) Holcus lanatus L. ( V e l v e t - g r a s s ) Elymus m o l l i s T r i n . (Wildrye) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 x x x S x D D x D x x x S x x D S x D GRASSES (cont.) Bromus inermis Leys. (Smooth Brome) SEDGES Carex l y n g b y e i Hornem. (Lyngby's Sedge) E l e o c h a r i s p a l u s t r i s (L.) R. & S. (Spike-rush) S c i r p u s acutus Muhl. (Bulrush) S c i r p u s americanus Pers. (Three-square Bulrush) S c i r p u s maritimus L. (Bulrush) Carex obnupta B a i l e y (Slough Sedge) RUSHES Juncus b a l t i c u s W i l l d . ( B a l t i c Rush) Juncus e f f u s u s L. (Common Rush) Community 1 2 3 4 5 6 7 8 9 10 11 12 . 13 14 15 16 x D x D x x x S x X x D x x S x x x x S D D 00 o X X Table 10. Area of each p l a n t community on the Cowichan R i v e r E s t u a r y P l a n t Community Area Percent ( h e c t a r e s ) (%) Carex l y n g b y e i 22.4 31.6 Juncus b a l t i c u s 19.5 27.5 D i s t i c h l i s s p i c a t a - Juncus b a l t i c u s 9.9 14.0 Agropyron repens - C i r s i u m arvense 8.4 11.8 Carex l y n g b y e i - D i s t i c h l i s s p i c a t a 4.2 5.9 S u b t o t a l 64.4 90.8 Juncus b a l t i c u s - Carex l y n g b y e i - 2.4 3.4 Deschampsia c e s p i t o s a D i s t i c h l i s s p i c a t a 1.3 1.8 Carex l y n g b y e i - Agropyron repens .9 1.3 Barbarea o r t h o c e r a s - Holcus l a n a t u s .7 1.0 Glaux maritima .5 .7 D i s t i c h l i s s p i c a t a - T r i g l o c h i n maritimum - .3 .4 Glaux maritima - S c i r p u s americanus Poa compressa - A c h i l l e a m i l l e f o l i u m - .2 .3 A t r i p l e x p a t u l a Typha l a t i f o l i a .1 .1 • Elymus m o l l i s • .1 .1 S c i r p u s acutus .01 .01 TOTAL 70.9 99.9 - 132 -F i g u r e 6. Map showing the d i s t r i b u t i o n of the p l a n t communities on the Chemainus River E s t u a r y Legend s c a l e lcm=48m Pl a n t Community 1 S c i r p u s maritimus 2 Pyrus f u s c a - Rosa nutkana 3 Carex l y n g b y e i - S c i r p u s maritimus 4 Pyrus f u s c a - Rosa nutkana - C y t i s u s s c o p a r i u s -Equisetum arvense 5 Arbutus m e n z i e s i i - Quercus garryana - Anthoxanthum odoratum 6 Juniperus scopulorum 7 Juncus b a l t i c u s 8 S a l i c o r n i a v i r g i n i c a l y n g b y e i 9 dyke v e g e t a t i o n C y t i s u s s c o p a r i u s - Rosa nutkana -GRAMINEAE 10 Pseudotsuga m e n z i e s i i - Juniperus scopulorum -Pachistima m y r s i n i t e s 11 S a l i c o r n i a v i r g i n i c a - D i s t i c h l i s s p i c a t a 12 S a l i c o r n i a v i r g i n i c a 13 D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a 14 Carex l y n g b y e i 15 S a l i c o r n i a v i r g i n i c a - Juncus b a l t i c u s 16 GRAMINEAE - Juncus b a l t i c u s 17 GRAMINEAE 18 Zostera marina 19 S a l i c o r n i a v i r g i n i c a - Z o s t e r a marina - GRAMINEAE - D i s t i c h l i s s p i c a t a - Carex - 183 A a g r i c u l t u r e r — | b u i l d i n g s , urban B log booms f | j dyke F f o r e s t K K tt r a i l w a y I i n d u s t r i a l • road I t r a n s e c t Table 11. D e s c r i p t i o n of the plant Communities on the Chemainus River Estuary, A p r i l 1975. 1 S c i r p u s maritimus 2 Pyrus fusca - Rosa nutkana 3 Carex l y n g b y e i - S c i r p u s maritimus 4 Pyrus f u s c a - Rosa nutkana - C y t i s u s s c o p a r i u s - Equisetum arvense 5 Arbutus m e n z i e s i i - Quercus garryana - Anthoxanthum odoratum 6 Juniperus scopulorum - GRAMINEAE 7 Juncus b a l t i c u s 8 S a l i c o r n i a v i r g i n i c a - D i s t i c h l i s s p i c a t a - Carex l y n g b y e i 9 C y t i s u s scoparius - Rosa nutkana - GRAMINEAE i 10 Pseudotsuga m e n z i e s i i - Juniperus scopulorum - Pachistima m y r s i n i t e s oo 11 S a l i c o r n i a v i r g i n i c a - D i s t i c h l i s s p i c a t a 1 12 S a l i c o r n i a v i r g i n i c a 13 D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a 14 Carex l y n g b y e i 15 S a l i c o r n i a v i r g i n i c a - Juncus b a l t i c u s 16 GRAMINEAE - Juncus b a l t i c u s 17 GRAMINEAE 18 Zos t e r a marina 19 S a l i c o r n i a v i r g i n i c a - Zostera marina Following i s a l i s t of the plant species in each community i n c l u d i n g dominant s p e c i e s i n d i c a t e d by a D and subdominant species i n d i c a t e d by an S. Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 TREES Abies grandis (Dougl.) Forbes x x (Grand F i r ) Acer macrophyllum Pursh x x (Common Maple) Alnus rubra Bong. x x (Red Alder) Arbutus m e n z i e s i i Pursh (Arbutus) Cornus n u t t a l l i i Aud. x (Dogwood) Juniperus scopulorum Sarg. x D D (Juniper) Prunus sp. L x x x ( C u l t i v a t e d Cherry) Prunus emarginata (Dougl.) Walp. x x x (Choke cherry) Pseudotsuga m e n z i e s i i ( M i r b e l ) Franco x x x D (Douglas F i r ) 00 TREES (cont.) Pyrus fusca Raf. ( P a c i f i c Crabapple) Quercus garryana Dougl. (Garry Oak) Rhamnus purshiana DC. (Cascara) Thuja p l i c a t a Donn. (Western Red Cedar) SHRUBS S a l i x hookeriana B a r r a t t (Hooker Willow) S a l i x s c o u l e r i a n a B a r r a t t (Scouler Willow) Amelanchier a l n i f o l i a Nutt. ( S e r v i c e b e r r y ) B e r b e r i s a q u i f o l i u m Pursh (Oregongrape) B e r b e r i s nervosa Pursh (Oregongrape) C y t i s u s s c o p a r i u s (L.) Link (Broom) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 S S x x S x x X X X X X I I- 1 00 x x a I X X X X X X X s S x D SHRUBS ( c o n t . ) G a u l t h e r i a s h a l l o n P u r s h ( S a l a l ) H o l o d i s c u s d i s c o l o r ( P u r s h ) Maxim. ( O c e a n - s p r a y ) L i n n a e a b o r e a l i s L. ( T w i n f l o w e r ) L o n i c e r a c i l i o s a ( P u r s h ) DC. (Orange H o n e y s u c k l e ) L o n i c e r a h i s p u d u l a ( L i n d l . ) D o u g l . ( H a i r y H o n e y s u c k l e ) L o n i c e r a i n v o l u c r a t a ( R i c h . ) Banks O s m a r o n i a c e r a s i f o r m i s ( T . & B.) G r e e ne P a c h i s t i m a m y r s i n i t e s ( P u r s h ) R a f . ( P a c h i s t i m a ) P h y s o c a r p u s c a p i t a t u s ( P u r s h ) K u n t z e ( N i n e b a r k ) R i b e s sanguineum P u r s h (Red C u r r a n t ) Rubus l a c i n i a t u s W i l l d . ( E v e r g r e e n B l a c k b e r r y ) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 x x X X X X x 1 CO •o X X I X X D X X SHRUBS (cont.) Rubus s p e c t a b i l i s Pursh (Salmonberry) Rubus urs i n u s Cham. & Schlecht ( P a c i f i c Blackberry) Rosa nutkana P r e s l . (Nootka Rose) Sambucus racemosa L. ( E l d e r b e r r y ) Symphoricarpos albus (L.) Blake (Snowberry) Ulex europaeus L. (Gorse) FORBS A c h i l l e a m i l l e f o l i u m L. (Yarrow) A n g e l i c a sp. L. (Angeli c a ) Apocynum androsaemifolium L. (Dogbane) Arctium minus ( H i l l ) Bernh. (Common Burdock) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 X X X S S x x S x X X X X X X X S X X X X X X FORBS (cont.) A r e n a r i a macrophylla Hook (Sandwort) A r e n a r i a p a l u d i c o l a Robins. (Sandwort) A r e n a r i a s t r i c t a Michx. (Slender Sandwort) Asparagus o f f i c i n a l i s L. ^  (Asparagus) A t r i p l e x patula L. (Saltbush) Barbarea orthoceras Ledeb. (Wintercress) Barbarea v u l g a r i s R. Br. (Wintercress) B e l l i s perennis L. (Daisy) Camassia l e i c h t l i n i i (Baker) Wats. (Camas) Cardamine i n t e g r i f o l i a (Nutt.) Green ( B i t t e r c r e s s ) C a s t i l l e j a l e v i s e c t a Greenm. (Golden Indian-paintbrush) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 x x X X X X X X X X X X I CO CD X I X X X FORBS (cont.) Cerastium arvense L. (Mouse-ear Chickweed) Cerastium viscosum L. (Sticky chickweed) Cirsium arvense (L.) Scop. (Canada Thistle) Cirsium vulgare (Savi.) Tenore (Common Thistle) C l i n t o n i a u n i f l o r a (Schult.) Kun (Clintonia) C o l l i n s i a p a r v i f l o r a L i n d l . (Blue-eyed Mary) Cuscuta salina Engelm. (Salt-marsh Dodder) Dicentra formosa (Andr.) Walp. (Bleeding Heart) Draba sp. L. (Draba) Epilobium angustifolium L. (Fireweed) Equisetum arvense L. (Common Horsetail) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 x x X X CD o X X X s X FORBS (cont.) E r i g e r o n p h i l a d e l p h i c u s L. ( P h i l a d e l p h i a Fleabane) Erythronium oregonum Applegate ( D o g t o o t h - v i o l e t ) F r a g a r i a c h i l o e n s i s (L.) Duchesne ( C o a s t a l Streawberry) Galium t r i f i d u m L. (Small Bedstraw) Geranium molle L. (Dovefoot Geranium) Glaux maritima L (Sea-milkwort) G r i n d e l i a i n t e g r i f o l i a DC. (Gumweed) Hel i o t r o p i u m curassavicum L. (Seaside H e l i o t r o p e ) Hypericum formosum H.B.K. (S t . John's-wort) Hypochaeris r a d i c a t a L. (Hairy Cats-ear) Lactuca m u r a l i s (L.) Fresen. (Wall Lettuce) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 x x X X X X X D X X X Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 FORBS (cont.) Lathyrus p a l u s t r i s L. x (Marsh Pea) L i l a e o p s i s o c c i d e n t a l i s C o u l t . & Rose x ( L i l a e o p s i s ) L l o y d i a s e r o t i n a (L.) Sweet x ( L l o y d i a ) Lomatium nudicaule (Pursh) C o u l t . & Rose x (De ser t-par s 1 ey) Maianthemum d i l a t a t u m (Wood) Nels. & Machr. x i ( F l a s e L i l y - o f - t h e - v a l l e y ) Oenanthe sarmentosa P r e s l . x 1 0 (Water-parsley) ' Montia p a r v i f o l i a (Moc.) Greene x (Miner's Lettuce) P e t a s i t e s f r i g i d u s (L.) F r i e s . x ( C o l t s f o o t ) Plantago l a n c e o l a t a L. x x x x x x (Ribgrass) Plantago macrocarpa Cham. & Schlecht x ( P l a n t a i n ) P l e c t r i s congesta ( L i n d l . ) DC. x (Rosy Pink) FORBS (cont.) Polypodium s c o u l e r i Hook & Grev. (Polypody) Polystichum munitum ( K a u l f . ) P r e s l (Sword-fern) P o t e n t i l l a p a c i f i c a Howell ( C i n q u e f o i l ) P t e r i d i u m aquilinum (L.) Kuhn (Bracken) Rumex a c e t o s e l l a L. (Sour Weed) Rumex c r i s p u s L. (Cu r l y Dock) S a l i c o r n i a v i r g i n i c a L. (Saltwort) Sedum s p a t h u l i f o l i u m Hook. (Sedum) S t e l l a r i a c r i s p a Cham. & Sch l e c h t . ( C r i s p e d Sandwort) Taraxacum o f f i c i n a l e Weber (Common Dandelion) T e l l i m a g r a n d i f l o r u m (Pursh) Dougl. (Fringecup) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 x I-X X X X X X X X X X X x x S D D x D x D x x x X X X X CD X FORBS (cont.) T r i e n t a l i s l a t i f o l i a Hook (S t a r f l o w e r ) T r i f o l i u m dubium S i b t h . (Least Hop Clover) T r i f o l i u m pratense L. (Red Clover) T r i f o L i u m repens L. (White Clover) T r i f o l i u m wormskjoldii Lehm. (Springbank Clover) T r i g l o c h i n maritimum L. (Arrow-grass) Typha l a t i f o l i a L. ( C a t - t a i l ) U r t i c a d i o i c a L. ( S t i n g i n g N e t t l e ) V i c i a gigantea Hook (Giant Vetch) Zostera marina L. ( E e l - g r a s s ) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 x x X X X X D S GRASSES Anthoxanthum odoratum L. (Sweet Ve r n a l g r a s s ) D a c t y l i s glomerata L. (Orchard-grass) D i s t i c h l i s s p i c a t a (L.) Greene ( S a l t g r a s s ) Elymus moll i s T r i n . (Wildrye) GRAMINEAE (Grasses) Hordeum brachyantherum Nevski. (Meadow Barley) SEDGES Carex l y n g b y e i Hornem (Lyngby 1s Sedge) S c i r p u s maritimus L. (Bulrush) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 S S D D x x x x S S x D D D D D x RUSHES Juncus b a l t i c u s W i l l d . ( B a l t i c Rush) Juncus e f f u s u s L. (Common Rush) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 D x D D x x x i }—*• co I - 197 -Table 12. Area of each p l a n t community on the Chemainus R i v e r Estuary Plant Community Area Percent ( h e c t a r e s ) (%) S a l i c o r n i a v i r g i n i c a - D i s t i c h l i s s p i c a t a 40.9 24.3 GRAMINEAE 27.7 16.4 D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a 25.0 14.8 S a l i c o r n i a v i r g i n i c a 19.0 11.3 Arbutus m e n z i e s i i - Quercus garryana - 16.5 9.8 Anthoxanthum odoratum Carex l y n g b y e i 14.7 8.7 GRAMINEAE - Juncus b a l t i c u s 7.8 4.6 S a l i c o r n i a v i r g i n i c a - D i s t i c h l i s s p i c a t a - 5.3 3.1  Carex l y n g b y e i S u b t o t a l 156.9 93.0 Juncus b a l t i c u s 3.5 2.1 S c i r p u s maritimus 3.3 2.0 Pseudotsuga m e n z i e s i i - Ju n i p e r u s 1.8 1.1 scopulorum - Pa c h i s t i m a m y r s i n i t e s Juniperus scopulorum - GRAMINEAE .9 .5 Carex l y n g b y e i - S c i r p u s maritimus .7 .4 Pyrus f u s c a - Rosa nutkana .5 .3 S a l i c o r n i a v i r g i n i c a - Zostera marina .4 .2 Pyrus f u s c a - Rosa nutkana - C y t i s u s .1 .1 sc o p a r i u s - Equisetum arvense S a l i c o r n i a v i r g i n i c a - Juncus b a l t i c u s .2 .1 Zoste r a marina . 1 .1 168.4 99.9 - 198 -Fi g u r e 7. Map showing the d i s t r i b u t i o n of the p l a n t communities on the Nanaimo R i v e r Estuary Legend Scale lcm=48m Plan t Community 1 Carex l y n g b y e i 2 S c i r p u s maritimus 3 T r i g l o c h i n maritimum 4 S p e r g u l a r i a canadensis 5 T r i g l o c h i n maritimum - S p e r g u l a r i a canadensis -S a l i c o r n i a v i r g i n i c a 6 Carex l y n g b y e i - Deschampsia c e s p i t o s a 7 Carex l y n g b y e i - Juncus b a l t i c u s 8 Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a - D i s t i c h l i s s p i c a t a 9 Agropyron repens - D i s t i c h l i s s p i c a t a - Juncus b a l t i c u s 10 Juncus a r t i c u l a t u s 11 A t r i p l e x p a t u l a - G r i n d e l i a i n t e g r i f o l i a 12 D i s t i c h l i s s p i c a t a 13 S a l i c o r n i a v i r g i n i c a - Glaux maritima 14 Carex l y n g b y e i - Deschampsia c e s p i t o s a - Juncus a r t i c u l a t u s 15 D i s t i c h l i s s p i c a t a - T r i g l o c h i n maritimum 16 D i s t i c h l i s s p i c a t a - Juncus a r t i c u l a t u s - A g r o s t i s t e n u i s 17 D i s t i c h l i s s p i c a t a - Juncus a r t i c u l a t u s 18 Quercus garryana - Acer macrophyllum 19 C y t i s u s s c o p a r i u s - 199 -20 P o t e n t i l l a p a c i f i c a 21 Pseudotsuga m e n z i e s i i 22 Typha l a t i f o l i a A a g r i c u l t u r e L t i d a l B l o g boom SL s u b t i d a l CD car dump j — j r£] u urban, b u i l d i n g s D dredged —14~ d y k e G g r a v e l GB g r a v e l bar I i n d u s t r i a l - 200 -Table 13. D e s c r i p t i o n of the p l a n t communities on the Nanaimo R i v e r E s t u a r y , June 1976 1 Carex l y n g b y e i 2 S c i r p u s maritimus 3 T r i g l o c h i n maritimum 4 S p e r g u l a r i a canadensis 5 T r i g l o c h i n maritimum - S p e r g u l a r i a canadensis -S a l i c o r n i a v i r g i n i c a Carex l y n g b y e i - Deschampsia c e s p i t o s a  Carex l y n g b y e i - Juncus b a l t i c u s Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a - D i s t i c h l i s  s p i c a t a Agropyron repens - D i s t i c h l i s s p i c a t a - Juncus b a l t i c u s  Juncus a r t i c u l a t u s A t r i p l e x p a t u l a - G r i n d e l i a i n t e g r i f o l i a  D i s t i c h l i s s p i c a t a S a l i c o r n i a v i r g i n i c a - Glaux maritima Carex l y n g b y e i - Deschampsia c e s p i t o s a - Juncus  a r t i c u l a t u s 15 D i s t i c h l i s s p i c a t a - T r i g l o c h i n maritimum 16 D i s t i c h l i s s p i c a t a - Juncus a r t i c u l a t u s - A g r o s t i s t e n u i s 17 D i s t i c h l i s s p i c a t a - Juncus a r t i c u l a t u s 18 Quercus garryana - Acer macrophyllum 19 C y t i s u s s c o p a r i u s 20 P o t e n t i l l a p a c i f i c a 21 Pseudotsuga m e n z i e s i i 22 Typha l a t i f o l i a F o l l o w i n g i s a l i s t of the s p e c i e s i n each p l a n t community i n c l u d i n g dominant s p e c i e s i n d i c a t e d by a D and subdominant s p e c i e s i n d i c a t e d by an S. 6 7 8 9 10 11 12 13 14 Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 TREES Quercus garryana Dougl. D x x (Garry Oak) Thuja plicata Donn. x (Western Red Cedar) Pyrus fusca Raf. x x x (Pacific Crabapple) Acer macrophyllum Pursh. S x (Common Maple) Alnus rubra Bong. x (Red Alder) Abies grandis (Dougl.) Forbes x S (Grand Fir) Pseudotsuga menziesii (Mirbel) Franco x D (Douglas Fir) Pyrus malus L. x (Apple) SHRUBS Cytisus scoparius (L.) Link x D (Broom) Symphoricarpos albus (L.) Blake (Snowberry) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 TREES (cont.) Rubus laciniatus Willd. (Evergreen Blackberry) Rubus spectabilis Pursh (Salmonberry) Crataegus oxyacantha L. (Hawthorn) Lonicera involucrata (Rich.) Banks (Black Twin-berry) Rosa nutkana Presl. (Nootka Rose) Vaccinium parvifolium Smith (Huckleberry) FORBS Potentilla pacifica Howell (Cinquefoil) Erigeron philadelphicus L. (Philadelphia Fleabane) Triglochin maritimum L. (Arrow-grass) Glaux maritima L. (Sea milkwort) x x x x x x x x x x x x S x x x x x x x D x x x x x x x x x x x D x D x x x x D x x x x x x x S x x x Corriminity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 FORBS (cont.) Achillea millefolium L. (Yarrow) x x x x x x Salicornia virginica L. x S x x x D x x x x x (Saltwort) F r i t i l l a r i a camschatcensis (L.) Ker-Gawl x (Chocolate L i l y ) x x Plantago lanceolata L. x x (Ribgrass) Plantago maritima L. x x x x x x x (Seaside Plantain) Plantago macrocarpa Cham. & Schlecht x x (Plantain) Camassia l e i c h t l i n i i (Baker) Wats. x (Camas) Spergularia canadensis (Pers.) G. Don x x x D S x S (Saltmarsh Sandspurry) Trifolium wormskjoldii Lehm. x x (Springbank Clover) Taraxacum officinale Weber x x (Common Dandelion) Grindelia i n t e g r i f o l i a DC. x x S x x x x x x (Gumweed) Cbmraunity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 FORBS (cont.) Pantago major L. x x (Common Plantain) Ruppia maritima L. (Ditch-grass) Trifolium repens L. (White Clover) Trifolium pratense L. (Red Clover) Trifolium dubium Sibth. (Least Hop Clover) Lotus corniculatus L. (Birdsf cot-trefoil) Rumex crispus L. (Curly Dock) Penanthe sarmentosa Presl. x (Water-parsley) Cotula coronopifolia L. x (Brass Buttons) Asparagus o f f i c i n a l i s L. x (Asparagus) Cirsium arvense (L.) Scop. (Canada Thistle) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 FORBS (cont.) Maianthemurn dilatatum (Wood) Nels. & Macbr. x (False Lily-of-the-valley) Ranunculus repens L. x (Creeping Buttercup) Lactuca muralis (L.) Fresen. x x (Wall Lettuce) Lathyrus palustris L. x (Marsh Pea) Prunella vulgaris L. x (Self-heal) x x x Hypochaeris radicata L x x x (Hairy Cats-ear) Vicia gigantea Hook x x (Giant Vetch) Ti a r e l l a t r i f o l i a t a var. t r i f o l i a t a L. x (Foamflower) Osmorhiza chilensis H. & A. . x (Sweet-root) Rumex acetosella L. x (Sour Weed) Typha l a t i f o l i a L. D (Cat-tail) FORBS (cont.) Atriplex patula L. (Saltbush) St e l l a r i a humifusa Rottb. (Spreading Starwort) Cirsium arvense (L.) Scop. (Canada Thistle) Epilobium watsonii Barbey (Watson's Willow-herb) Collinsia parviflora Lindl. (Blue-eyed Mary) GRASSES Poa pratensis L. (Kentucky Bluegrass) Dactylis glomerata L. (Orchard-grass) Di s t i c h l i s spicata (L.) Greene (Saltgrass) Deschampsia cespitosa (L.) Beauv. (Tufted Hairgrass) Agropyron repens (L.) Beauv. (Couch grass) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x x x x x D x x x x x x x x x x x x ' to o OS I X x X X X x D D D x x S x x x x x x x x x S D x x D x S x x x x D GRASSES (cont.) Holcus lanatus L. (Velvet-grass) Hordeum brachyantherum Nevski. (Meadow Barley) Hordeum murlnum L. (Mouse Barley) Phleum pratense L. (Timothy) Agrostis alba L. (Bentgrass) Festuca rubra L. (Red Fescue) Agrostis tenuis Sibth. (Colonial Bentgrass) Elymus glaucus Buckl. (Blue Wildrye) Poa compressa L. (Canada Bluegrass) Glyceria borealis (Nash) Batch (Northern Mannagrass) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 IS 19 20 21 22 X X X x x x S X X X to o x S X SEDGES Carex lyngbyei Hornem. (Lyngby's Sedge) Scirpus acutus Muhl. (Bulrush) Scirpus maritimus L. (Bulrush) Scirpus americanus Pers. (Three-square Bulrush) RUSHES Juncus balticus Willd. (Baltic Rush) Juncus articulatus L. (Jointed Rush) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 D x x x D D x x x x D S x x x x x D x x x O 00 D D S x x D x S x S D x x x - 209 -Table 14. Area of each p l a n t community on the Nanaimo R i v e r E s t u a r y Pl a n t Community Area Percent ( h e c t a r e s ) (%) Juncus a r t i c u l a t u s 16.4 29.1 Carex l y n g b y e i 9.2 16.3 D i s t i c h l i s s p i c a t a - Juncus a r t i c u l a t u s - 6.3 11.2 A g r o s t i s t e n u i s Carex l y n g b y e i - Deschampsia c e s p i t o s a - 4.2 7.4 Juncus a r t i c u l a t u s T r i g l o c h i n maritimum - S p e r g u l a r i a 4.1 7.3 canadensis S a l i c o r n i s v i r g i n i c a S a l i c o r n i a v i r g i n i c a - Glaux maritima 2.6 4.6 C y t i s u s s c o p a r i u s 2.2 3.9 Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a - 2.1 3.7 D i s t i c h l i s s p i c a t a D i s t i c h l i s s p i c a t a - Juncus a r t i c u l a t u s 1.7 3.0 D i s t i c h l i s s p i c a t a - T r i g l o c h i n maritimum 1.5 2.7 S p e r g u l a r i a canadensis 1.5 2.7 S u b t o t a l 51.8 91.9 Quercus garryana - Acer macrophyllum .9 1.6 Carex l y n g b y e i - Deschampsia c e s p i t o s a .9 1.6 Pseudotsuga m e n z i e s i i .8 1.4 Carex l y n g b y e i - Juncus b a l t i c u s .5 .9 A t r i p l e x p a t u l a - G r i n d e l i a i n t e g r i f o l i a .5 .9 S c i r p u s maritimus .4 .7 T r i g l o c h i n maritimum .3 .5 - 210 D i s t i c h l i s s p i c a t a Agropyron repens - D i s t i c h l i s s p i c a t a  Juncus b a l t i c u s P o t e n t i l l a p a c i f i c a Typha l a t i f o l i a TOTAL - 211 -F i g u r e 8. Map showing the d i s t r i b u t i o n of the p l a n t communities on the Nanoose-Bonell Creeks' Es t u a r y Legend Scale lcm=48 Pl a n t Community 1 Rosa nutkana - Rubus l a c i n i a t u s - Rubus p a r v i f l o r u s -Rubus s p e c t a b i l i s 2 Juncus b a l t i c u s 3 S c i r p u s maritimus 4 Carex l y n g b y e i 5 D i s t i c h l i s s p i c a t a - S a l i c o r n i a v i r g i n i c a 6 G r i n d e l i a i n t e g r i f o l i a - Elymus m o l l i s 7 S a l i c o r n i a v i r g i n i c a 8 Plantago maritima 9 A g r o s t i s a l b a 10 P o t e n t i l l a p a c i f i c a - A c h i l l e a m i l l e f o l i u m - GRAMINEAE 11 P h a l a r i s arundinacea A a g r i c u l t u r e *-*-*- r a i l w a y L t i d a l ... road S Sunday school camp I 1 b u i l d i n g s SL s u b t i d a l Table 15. D e s c r i p t i o n of the plant communities on the Nanoose-Bonell Creeks' September 16, 1976 1 Rosa nutkana - Rubus l a c i n i a t u s - Rubus p a r v i f l o r u s - Rubus s p e c t a b i l i s 2 Juncus b a l t i c u s 3 S c i r p u s maritimus 4 Carex ly n g b y e i 5 D i s t i c h l i s s p i c a t a - S a l i c o r n i a v i r g i n i c a 6 G r i n d e l i a i n t e g r i f o l i a - Elymus m o l l i s 7 S a l i c o r n i a v i r g i n i c a 8 Plantago maritima 9 A g r o s t i s a l b a 10 P o t e n t i l l a p a c i f i c a - A c h i l l e a m i l l e f o l i u m - GRAMINEAE 11 P h a l a r i s arundinacea Following i s a l i s t of the plant species i n each community i n c l u d i n g dominant by a D and subdominant species i n d i c a t e d by an S. TREES Alnus rubra Bong. (Red Alder) Pyrus fusca Raf. ( P a c i f i c Crabapple) SHRUBS S a l i x . sp. L. (Willow) Crataegus d o u g l a s i i L i n d l . (Black Hawthorn) Rosa nutkana P r e s l . (Nootka Rose) Rubus l a c i n i a t u s W i l l d . (Evergreen Blackberry) Symphoricarpos albus (L.) Blake (Snowberry) C y t i s u s scoparius (L.) Link (Broom) Ribes sanguineum Pursh. (Red Currant) Community 1 2 3 4 5 6 7 8 9 10 11 D x D S D x SHRUBS (cont.) Rubus p a r v i f l o r u s Nutt. (Thimbleberry) Rubus s p e c t a b i l i s Pursh. (Salmonberry) Physocarpus c a p i t a t u s (Pursh) Kuntze (Ninebark) FORBS P o t e n t i l l a p a c i f i c a Howell ( C i n q u e f o i l ) Plantago l a n c e o l a t a L. (Ribgrass) T r i g l o c h i n maritimum L. (Arrow-grass) Plantago maritima L. (Seaside p l a n t a i n ) T r i f o l i u m repens L. (White Clover) C i r s i u m arvense (L.) Scop. (Canada T h i s t l e ) Community 1 2 3 4 5 6 7 8 9 10 11 D D D x x x S D i to i X X D x X X X FORBS (cont.) Hypochaeris r a d i c a t a L. T r i f o l i u m pratense L. (Red C l o v e r ) Equisetum arvense L. (Common H o r s e t a i l ) P t e r i d i u m aquilinum (L.) Kuhn. (Bracken) Lathyrus p a l u s t r i s L. (Marsh Pea) Epilobium ang'ustifolium L. (Fireweed) A t r i p l e x p a t u l a L. (Saltbush) S p e r g u l a r i a canadensis (Pers.) G. Don (Sandspurry) Suaeda maritima (L.) Dumort. ( S e a b l i t e ) F r a g a r i a c h i l o e n s i s (L.) Duchesne (Wild Strawberry) Community 6 7 8 9 10 11 x x X X X X X S X X GRASSES Holcus lanatus L. ( V e l v e t - g r a s s ) D i s t i c h l i s s p i c a t a (L.) Greene ( S a l t g r a s s ) Elymus m o l l i s T r i n . (Wildrye) Phleum pratense L. (Timothy) A g r o s t i s alba L. (Bentgrass) P h a l a r i s arundinacea L. (Reed Canarygrass) Deschampsia c e s p i t o s a (L.) Beauv. (Tufted H a i r g r a s s ) D a c t y l i s glomerata L. (Orchard-grass) Hordeum brachyantherum Nevski (Meadow Barley) Hordeum murinum L. (Mouse Barley) Poa compressa L. (Canada Bluegrass) Community 1 2 3 4 5 6 7 8 9 10 11 x x x x S D S x S x D x x D x x i to l-» D °> i X X X X GRASSES Holcus lanatus L. ( V e l v e t - g r a s s ) D i s t i c h l i s s p i c a t a (L.) Greene ( S a l t g r a s s ) Elymus m o l l i s T r i n . (Wildrye) Phleum pratense L. (Timothy) A g r o s t i s alba L. (Bentgrass) P h a l a r i s arundinacea L. (Reed Canarygrass) Deschampsia c e s p i t o s a (L.) Beauv. (Tufted H a i r g r a s s ) D a c t y l i s glomerata L. (Orchard-grass) Hordeum'brachyantherum Nevski (Meadow Barley) Hordeum murinum L. (Mouse Barley) Poa compressa L. (Canada Bluegrass) Community 3 4 5 6 7 8 9 10 11 x S D S x S x D x x D S x x X X x D Community 1 2 3 4 5 6 7 8 GRASSES Bromus tectorum L. x (Brome-grass) Poa p r a t e n s i s L. (Kentucky Bluegrass) SEDGES Carex l y n g b y e i Hornem. t x D x (Lyngby's Sedge) Scirpus microcarpus P r e s l . ( S m a l l - f r u i t Bulrush) S c i r p u s maritimus L. (Bulrush) D x RUSHES Juncus b a l t i c u s W i l l d . ( B a l t i c Rush) Juncus e f f u s u s L. (Common Rush) Juncus a r t i c u l a t u s L. ( J o i n t e d Rush) Juncus e n s i f o l i u s v a r . e n s i f o l i u s Wikst. (Dagger-leaf Rush) Juncus f a l c a t u s E. Meyer ( S i c k l e - l e a v e d Rush) Community 6 7 8 9 10 11 S x x S x x x i x to co - 220 -Table 16. Area of each p l a n t community on the Nanoose-B o n e l l Creeks' Estuary P l a n t Community Area Percent ( h e c t a r e s ) (%) D i s t i c h l i s s p i c a t a - S a l i c o r n i a v i r g i n i c a 4.5 19.9 P o t e n t i l l a p a c i f i c a - A c h i l l e s m i l l e f o l i u m - 3.6 15.9 GRAMINEAE A g r o s t i s a l b a 3.1 13.7 P h a l a r i s arundinacea 3.0 13.3 Rosa nutkana - Rubus l a c i n i a t u s - Rubus 2.2 9.7 p a r v i f l o r u s - Rubus s p e c t a b i l i s Carex l y n g b y e i 2.1 9.3 S a l i c o r n i a v i r g i n i c a 2.0 8.8 S u b t o t a l 20.5 90.6 Juncus b a l t i c u s 1.5 6.6 G r i n d e l i a i n t e g r i f o l i a - Elymus m o l l i s .3 1.3 Plantago maritima .2 .9 S c i r p u s maritimus . 1 ._4 TOTAL 22.6 99.8 - 221 -F i g u r e 9. Map showing the d i s t r i b u t i o n of the p l a n t communities on the Englishman R i v e r E s t u a r y Legend Scale lcm=48m P l a n t Community 1 S a l i c o r n i a v i r g i n i c a - T r i g l o c h i n maritimum 2 D i s t i c h l i s s p i c a t a - S a l i c o r n i a v i r g i n i c a 3 D i s t i c h l i s s p i c a t a 4 D i s t i c h l i s s p i c a t a - Carex l y n g b y e i 5 G r i n d e l i a i n t e g r i f o l i a - A g r o s t i s a l b a 6 Sonchus a r v e n s i s - C i r s i u m arvense 7 Hoicus l a n a t u s - E p i l o b i u m a n g u s t i f o l i u m 8 Carex l y n g b y e i 9 dyke v e g e t a t i o n A g r o s t i s a l b a - Poa p r a t e n s i s -G r i n d e l i a i n t e g r i f o l i a 10 Carex l y n g b y e i - Juncus b a l t i c u s 11 Elymus m o l l i s 12 Glaux maritima - Plantago maritima - D i s t i c h l i s s p i c a t a 13 D i s t i c h l i s s p i c a t a - Hordeum murinum - P o t e n t i l l a p a c i f i c a 14 A g r o s t i s a l b a - P o t e n t i l l a p a c i f i c a 15 Sonchus a r v e n s i s - G r j n d e l i a i n t e g r i f o l i a - A g r o s t i s a l b a 16 Agropyron repens - A c h i l l e a m i l l e f o l i u m - G r i n d e l i a i n t e g r i f o l i a - C y t i s u s s c o p a r i u s 17 C y t i s u s s c o p a r i u s - Rubus l a c i n i a t u s - Rosa nutkana 18 Juncus b a l t i c u s - D i s t i c h l i s s p i c a t a 19 Alnus rubra - 222 -F i g u r e 9 (cont.) L t i d a l SL s u b t i d a l U I 1 | J urban dyke road Table 17. D e s c r i p t i o n of the plant communities on the Englishman R i v e r Estuary, August 1976. 1 S a l i c o r n i a v i r g i n i c a - T r i g l o c h i n maritimum 2 D i s t i c h l i s s p i c a t a - S a l i c o r n i a v i r g i n i c a 3 D i s t i c h l i s s p i c a t a 4 D i s t i c h l i s s p i c a t a - Carex l y n g b y e i 5 G r i n d e l i a i n t e g r i f o l i a - A g r o s t i s alba 6 Sonchus a r v e n s i s - C i r s i u m arvense 7 Holcus lanatus - Epilobium a n g u s t i f o l i u m 8 Carex l y n g b y e i 9 dyke v e g e t a t i o n A g r o s t i s a l b a - Poa p r a t e n s i s - G r i n d e l i a i n t e g r i f o l i a 10 Carex lyng b y e i - Juncus b a l t i c u s 11 Elymus moll i s 12 Glaux maritima - Plantago maritima - D i s t i c h l i s s p i c a t a 13 D i s t i c h l i s s p i c a t a - Hordeum murinum - P o t e n t i l l a p a c i f i c a 14 A g r o s t i s alba - P o t e n t i l l a p a c i f i c a 15 Sonchus a r v e n s i s - G r i n d e l i a i n t e g r i f o l i a - A g r o s t i s a l b a 16 Agropyron repens - A c h i l l e a m i l l e f o l i u m - G r i n d e l i a i n t e g r i f o l i a - C y t i s u s s c o p a r i u s 17 C y t i s u s s c o p a r i u s - Rubus l a c i n i a t u s - Rosa nutkana 18 Juncus b a l t i c u s - D i s t i c h l i s s p i c a t a 19 Alnus rubra F o l l o w i n g i s a l i s t of the species i n each plant community i n c l u d i n g dominant species i n d i c a t e d by a D and subdominant species i n d i c a t e d by an S. Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 TREES Pyrus f u s c a Raf. x x x x ( P a c i f i c Crabapple) Alnus rubra Bong. s (Red A l d e r ) Pseudotsuga m e n z i e s i i ( M i r b e l ) Franco x (Douglas F i r ) Thuja p l i c a t a Donn. (Red Cedar) Abies gran d i s (Dougl.) Forbes (Balsam F i r ) D SHRUBS Rosa nutkana P r e s l . (Nootka Rose) x D Sambucus racemosa ( E l d e r b e r r y ) Myrica g a l e L. (Sweet Gale) x SHRUBS (cont.) Rubus l a c i n i a t u s W i l l d . (Evergreen Blackberry) C y t i s u s s c o p a r i u s (L.) Link (Broom Ho l o d i s c u s d i s c o l o r (Pursh) Maxim. (Ocean-spray) Rubus u r s i n u s Cham. & Schlecht ( P a c i f i c Blackberry) Rubus s p e c t a b i l i s Pursh (Salmonberry) FORBS Rumex a c e t o s e l l a L. (Sour Dock) Ambrosia chamissonis (Less.) Greene (Ragweed) T r i f o l i u m w ormskjoldii Lehm. (Springbank Clover) Typha l a t i f o l i a L. (Common C a t - t a i l ) Ruppia maritima L (D i t c h - g r a s s ) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 D S D x FORBS (cont.) Asparagus o f f i c i n a l i s L. (Asparagus) Sonchus a r v e n s i s L. ( M i l k - t h i s t l e ) Lathyrus p a l u s t r i s L. (Marsh Pea) T r i f o l i u m pratense L. (Red C l o v e r ) E r i g e r o n p h i l a d e l p h i c u s L. ( P h i l a d e l p h i a Fleabane) F r a g a r i a c h i l o e n s i s (L.) Duchesne ( C o a s t a l Strawberry) S p e r g u l a r i a canadensis (Pers.) G. Don (Canada Sandspurry) A t r i p l e x p a t u l a L. (Saltbush) S t e l l a r i a humifusa Rottb. (Spreading Startwort) Oenanthe sarmentosa P r e s l . (Water-parsley) Senecio s y l v a t i c u s L. (Groundsel) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 x x x S x x x x x x x x x X X X X X X S x X X X X X X FORBS (cont.) Polygonum spergularlaeforme Meisn. (Spurry knotweed) Rumex s a l i c i f o l i u s Weinm. (Willow Dock) S a l i c o r n i a v i r g i n i c a L. (Pickleweed) T r i g l o c h i n maritimum L. (Arrow-grass) Glaux maritima L. (Sea-milkwort) G r i n d e l i a i n t e g r i f o l i a DC. (Gumweed) Epilobi u m a n g u s t i f o l i u m L. (Fireweed) Plantago maritima L. (Sea p l a n t a i n ) P o t e n t i l l a p a c i f i c a Howell ( C i n q u e f o i l ) Plantago l a n c e o l a t a L. (Ribgrass) C i r s i u m arvense (L.) Scop. (Canada T h i s t l e ) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 D D S x x x x x x x D x x x x x x x x x x x x x x x x x D x x x x x x S x S x • x D S x x x D x x x x x x x x x D x x x x x x x S D x x x x x x x x x x x S x S x x x S i to to -a i FORBS (cont.) C i r s i u m v u l g a r e (Savi) Tenore (Common T h i s t l e ) Rumex c r i s p u s L-(C u r l y Dock) A c h i l l e a m i l l e f o l i u m L. (Yarrow) Anaphalis margaritacea (L.) B. & ( P e a r l y - e v e r l a s t i n g ) GRASSES D i s t i c h l i s s p i c a t a (L.) Greene ( S a l t g r a s s ) Holcus l a n a t u s L. ( V e l v e t - g r a s s ) D a c t y l i s glomerata L. (Orchard-grass) Elymus moll i s T r i n . (Dune w i l d r y e ) Deschampsia c e s p i t o s a (L.) Beauv. (Tufted H a i r g r a s s ) Phalarus arundinacea L. (Reed Canarygrass) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 X X X X X X X X X X X S x X X X x D D D x x x x x x D D x x D x D x x x x D x x x X GRASSES (cont.) Hordeum murinum L. (Mouse B a r l e y ) A g r o s t i s alba L. (Bentgrass) Poa p r a t e n s i s L. (Kentucky Bluegrass) Poa sp. L. (Bluegrass) Agropyron repens (L.) Beauv (Couch Grass) Bromus s e c a l i n u s L. (Cheat) Horeurn brachyantherum Nevski (Meadow Barley) SEDGES Carex lyng b y e i Hornem. (Lyngby's Sedge) RUSHES Juncus b a l t i c u s W i l l d . (Rush) Community 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 x x x x x x S x x S x x x S x x D S x x x x S x x x x x x x S X X X X X X x x D D D x x x x x x x x x S x x x x x D - 230 -Table 18. Area of each p l a n t community on the Englishman R i v e r E s t u a r y . P l a n t Community Area Percent (Hectares) (%) Holcus l a n a t u s - Epilo b i u m a n g u s t i f o l i u m 3.7 14.9 Sonchus a r v e n s i s - C i r s i u m arvense 3.5 14.1 Carex l y n g b y e i 2.4 9.6 A g r o s t i s a l b a - P o t e n t i l l a p a c i f i c a 2.3 9.2 D i s t i c h l i s s p i c a t a - Carex l y n g b y e i 1.6 6.4 Carex l y n g b y e i - Juncus b a l t i c u s 1.5 6.0 Glaux maritima - Plantago maritima -D i s t i c h l i s s p i c a t a 1.5 6.0 Sonchus a r v e n s i s - G r i n d e l i a i n t e g r i f o l i a -A g r o s t i s a l b a 1.5 6.0 S a l i c o r n i a v i r g i n i c a - T r i g l o c h i n maritimum 1.3 5.2 Agropyron repens - A c h i l l e a m i l l e f o l i u m -G r i n d e l i a i n t e g r i f o l i a - C y t i s u s s c o p a r i u s 1.3 5.2 Juncus b a l t i c u s - D i s t i c h l i s s p i c a t a 1.1 4.4 D i s t i c h l i s s p i c a t a .8 3.2 Subtotal 22.5 90.2 Alnus rubra .7 2.8 Elymus m o l l i s .5 2.0 C y t i s u s s c o p a r i u s - Rubus l a c i n i a t u s - Rosa nutkana .5 2.0 G r i n d e l i a i n t e g r i f o l i a - A g r o s t i s a l b a .4 1.6 D i s t i c h l i s s p i c a t a - Hordeum murinum -P o t e n t i l l a p a c i f i c a .2 .8 D i s t i c h l i s s p i c a t a - S a l i c o r n i a v i r g i n i c a .1 .4 24.9 99.8 - 231 -F i g u r e 10. Map showing the d i s t r i b u t i o n of the p l a n t communities on the L i t t l e Qualicum R i v e r E s t u a r y . Legend Scale lcm=48m P l a n t Community 1 D i s t i c h l i s s p i c a t a 2 Pyrus f u s c a - Rosa nutkana 3 Elymus m o l l i s 4 P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i - E l e o c h a r i s p a l u s t r i s - Juncus b a l t i c u s 5 Typha l a t i f o l i a 6 Carex l y n g b y e i 7 P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i 8 P o t e n t i l l a p a c i f i c a - Juncus b a l t i c u s 9 Rosa nutkana 10 P o t e n t i l l a p a c i f i c a - Heracleum lanatum 11 GRAMINEAE 12 Rosa nutkana - Pyrus f u s c a - Rubus l a c i n i a t u s - Rubus s p e c t a b i l i s - Rubus p a r v i f l o r u s 13 S c i r p u s acutus 14 Rosa nutkana - Anthoxanthum odoratum - Bromus m o l l i s A a g r i c u l t u r e — — dyke F f o r e s t L t i d a l ,. - road R Canadian W i l d i f e S e r v i c e Reserve SL s u b t i d a l t r a n s e c t U urban b u i l d i n g s | j Table 19. D e s c r i p t i o n of the plant communities on the L i t t l e Qualicum R i v e r Estuary, June 1975. 1 D i s t i c h l i s s p i c a t a 2 Pyrus fusca - Rosa nutkana 3 Elymus m o l l i s 4 P o t e n t i l l a p a c i f i c a - Carex lyngbyei - E l e o c h a r i s p a l u s t r i s - Juncus b a l t i c u s 5 Typha l a t i f o l i a 6 Carex lyngbyei 7 P o t e n t i l l a p a c i f i c a - Carex lyngbyei 8 P o t e n t i l l a p a c i f i c a - Juncus b a l t i c u s , 9 Rosa nutkana co 10 P o t e n t i l l a p a c i f i c a - Heracleum lanatum i 11 GRAMINEAE 12 Rosa nutkana - Pyrus fusca - Rubus l a c i n i a t u s - Rubus s p e c t a b i l i s - Rubus p a r v i f l o r u s 13 Scirpus acutus 14 Rosa nutkana - Anthoxanthum odoratum - Bromus m o l l i s Following i s a l i s t of the plant species i n each community i n c l u d i n g dominant species i n d i c a t e d by a D and subdominant species i n d i c a t e d by an S. TREES Pyrus fusca Raf. ( P a c i f i c Crabapple) Acer macrophyllum Pursh (Common Maple) Thuja p l i c a t a Donn (Western Red Cedar) Pseudotsuga m e n z i e s i i ( M i r b e l ) Fran (Douglas F i r ) Pyrus malus L. (Apple) Picea s i t c h e n s i s (Borg.) Carr. ( S i t k a Spruce) SHRUBS Rosa nutkana P r e s l . (Nootka Rose) Symphoricarpos albus (L.) Blake (Snowberry) C y t i s u s scoparius (L.) Link (Broom) Lonicera i n v o l u c r a t a (Rich.) Banks (Black Twin-berry) SHRUBS (cont.) Sal ix sp. L. (Willow) Rubus s p e c t a b i l i s Pursh (Salmonberry) Rubus l a c i n i a t u s Willd. (Evergreen Blackberry) Rubus parv i f l o r u s Nutt. (Thimbleberry) FQRBS P o t e n t i l l a p a c i f i c a Howell ( C i n q u i f o i l ) Plantago lanceolata L. (Ribgrass) Trifolium wormskjoldii Lehm. (Springbank Clover) Hypochaeris radicata L. (Hairy Cats-ear) Triglochin maritimum L. (Arrow-grass) Trifolium dubium Sibth. (Least Hop Clover) Community 7 8 9 10 11 12 13 14 D D D x x x x x x x x x x FORBS (cont.) A c h i l l e a m i l l e f o l i u m L. (Yarrow) Rumex a c e t o s e l l a L. (Sour Weed) Ambrosia chamissonis (Less.) .Greene (Ragweed) Rumex c r i s p u s L. ( C u r l y Dock) Montia p e r f o j i a t a (Donn.) ftowell (Miner's L e t t u c e ) Barbarea orthoceras Ledeb. (W i n t e r c r e s s ) S i l e n e n o c t i f l o r a L. ( C a t c h f l y ) Ranunculus a c r i s L. (Meadow Buttercup) Oenanthe sarmentosa P r e s l . (Water-parsley) Glaux maritima L. (Sea-milkwort) Ranunculus c y n b a l a r i a Lursh. (Seaside Buttercup) Community 3 4 5 6 7 8 0 10 11 12 13 14 x x X X X X X 1 o FORBS (cont.) Geranium molle L. x (Dovefoot Geranium) Typha l a t i f o l i a L. ( C a t - t a i l ) T r i f o l i u m repens L. (White C l o v e r ) G r i n d e l i a i n t e g r i f o l i a DC. (Gumweed) T r i f o l i u m pratense L. x (Red C l o v e r ) T e l l i m a g r a n d i f l o r u m (Pursh) Dougl. x (Fringecup) Taraxacum o f f i c i n a l e Weber (Common Dandelion) F r i t i l l a r i a camschatcensis (L.) Ker.-Gawl (Chocolate L i l y ) E r i g e r o n p h i l a d e l p h i c u s L. ( P h i l a d e l p h i a Fleabane) Epilobium a n g u s t i f o l i u m L. (Fireweed) C i r s i u m arvense (L.) (Canada T h i s t l e ) Community 8 9 10 11 12 13 14 x x x x x x x x x x x x x x x x x x x x x x x 1 2 3 FORBS (cont.) V i c i a gigantea Hook. (Giant Vetch) S t e l l a r i a media (L.) C y r i l l . (Chickweed) Sisyrinchium a n g u s t i f o l i u m M i l l . (Blue-eyed Grass) Maianthemum dilatatum (Wood) Nels. St Macbr. (False L i l y - o f - t h e - v a l l e y ) Lathyrus p a l u s t r i s L. (Marsh Pea) L i l a e o p s i s o c c i d e n t a l i s Coult. & Rose ( L i l a e o p s i s ) Heracleum lanatum Michx. (Cow-parsnip) Sidal c e a hendersonii Wats. (Henderson's Checker-mallow) Medicago s a t i v a L. (Medic) P l e l i o p s i s h e l i a n t h o i d e s (L. ) Sweet (Ox-eye Daisy) B e l l i s perennis L. (Daisy) Community 4 5 6 7 8 9 x x x x x x 1 2 FORES (cont.) Lomatium nudicaule (Pursh) C o u l t . & Rose ( D e s e r t - p a r s l e y ) U r t i c a d i o i c a L. ( S t i n g i n g N e t t l e ) Delphinium m e n z i e s i i DC. (Menzies 1 Delphinium) A l l i u m g e y e r i v a r . tenerum Jones (Geyer's Onion) C o t u l a c o r o n o p i f o l i a L. x (Brass Buttons) C a p s e l l a b u r s a - p a s t o r i s (L.) Medic. x (Shepherd's-purse) GRASSES Elymus m o l l i s T r i n . (Wildrye) Bromus tectorum L. (Brome-grass) Holcus lanat u s L. ( V e l v e t - g r a s s ) D i s t i c h l i s s p i c a t a (L.) Greene D ( S a l t g r a s s ) Community 3 4 5 6 7 8 9 10 11 12 13 14 x X X D X 1 GRASSES (cont.) Poa p r a t e n s i s L. (Kentucky Bluegrass) Anthoxanthum odoratum L. (Sv/eet V e r n a l g r a s s ) P h a l a r i s arundinacea L. (Reed Canarygrass) Lolium perenne L. (Ryegrass) Festuca bromoides L. (Fescue) Bromus m o l l i s L. ( S o f t Brome) SEDGES Carex lyn g b y e i Hornem. x (Lyngby's Sedge) E l e o c h a r i s p a l u s t r i s (L.) R. & S. (Spike-rush) S c i r p u s microcarpus P r e s l . ( S m a l l - f r u i t Bulrush) S c i r p u s acutus Muhl. (Bulrush) Community 6 7 8 9 10 11 12 13 14 x x x S X X tc CO CD D D X X X D SEDGES (cont.) Carex p l u r i f l o r a Hulten (Sedge) Carex pansa B a i l e y (Sand-dune Sedge) Sc i r p u s maritimus L. (Bulrush) RUSHES Juncus b a l t i c u s W i l l d . ( B a l t i c Rush) Luzula s p i c a t a (L.) DC. (Spiked Woodrush) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 x x X D to o - 241 -Table 20. Area of each p l a n t community on the L i t t l e Qualicum R i v e r E s t u a r y P l a n t Community Area Percent ( h e c t a r e s ) (%) D i s t i c h l i s s p i c a t a 4.4 18.3 P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i 3.2 13.3 P o t e n t i l l a p a c i f i c a - Juncus b a l t i c u s 2.9 12.1 GRAMINEAE 2.9 12.1 Rosa nutkana - Anthoxanthum odoratum - 2.9 12.1 Bromus m o l l i s Carex l y n g b y e i 2.5 10.4 Rosa nutkana - Pyrus f u s c a - Rubus l a c i n i a t u s 2.1 8.8 - Rubus s p e c t a b i l i s - Rubus p a r v i f l o r u s Rosa nutkana 1.2 5.0 S u b t o t a l 22.1 92.1 Typha l a t i f o l i a .6 2.5 Pyrus f u s c a - Rosa nutkana .5 2.1 S c i r p u s acutus .5 2.1 Elymus m o l l i s .1 .4 P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i - .1 .4 E l e o c h a r i s p a l u s t r i s -Juncus b a l t i c u s P o t e n t i l l a p a c i f i c a - Heracleum lanatum .1. .4 TOTAL 24.0 100.0 - 242 -F i g u r e 11. Map showing the d i s t r i b u t i o n of the p l a n t communities on the Big Qualicum R i v e r Estuary Legend Scale lcm=48m P l a n t Community 1 Carex l y n g b y e i 2 Anthoxanthum odoratum - P o t e n t i l l a p a c i f i c a - Plantago  l a n c e o l a t a - T r i f o l i u m pratense boat launch camping ground dredged t i d a l s u b t i d a l b u i l d i n g road BL C D L SL Table 21. D e s c r i p t i o n of the plant communities on the Big Qualicum R i v e r Estuary. September, 1976. 1 Carex lyngbyei 2 Anthoxanthum odoratum - P o t e n t i l l a p a c i f i c a - Plantago l a n c e o l a t a - T r i f o l i u m pratense i CO I - 244 -Community 1 2 Trees Alnus rubra Bong. (Red A l d e r ) Acer macrophyllum Pursh (Common Maple) Populus t r i c h o c a r p a T. & G. (Cottonwood) Pyrus f u s c a Raf. ( P a c i f i c Crabapple) Shrubs Rubus s p e c t a b i l i s Pursh. x (Salmonberry) S a l i x sp. L. x (Willow) Rubus u r s i n u s Cham. & Schl e c h t x ( P a c i f i c B l a c k b e r r y ) Rosa nutkana P r e s l . (Nootka Rose) x - 245 -Forbs Lathyrus p a l u s t r i s L. (Marsh Pea) P o t e n t i l l a p a c i f i c a Howell (CinquefoTT) Plantago l a n c e o l a t a L. (Ribgra s s ) T r i f o l i u m pratense L. (Red C l o v e r ) V i c i a g igantea Hook (Giant Vetch) Rumex c r i s p u s L. (C u r l y Dock) Hypochaeris r a d i c a t a L. (Ha i r y Cats-ear) C i r s u i m arvense (L.) Scop. (Canada T h i s t l e ) Chenopodium album L. (Pigweed) E r i g e r o n p h i l a d e l p h i c u s L. ( P h i l a d e l p h i a Fleabane) Equisetum arvense L. (Common H o r s e t a i l ) G r i n d e l i a i n t e g r i f o l i a DC. (Gumweed) Anaphalis m a r g a r i t a c e a (L.) ( P e a r l y - e v e r l a s t i n g ) Plantago macrocarpa Chem. & ( P l a n t a i n ) Ranunculus repens L. (Creeping Buttercup) Glaux maritima L. (Sea-milkwort) Community 1 2 x x S S S x X X X X X X X B. & H. x Schlecht x x x - 246 -S e d g e C o m m u n i t y 1 2 G r a s s e s D e s c h a m p s i a c e s p i t o s a ( L . ) B e a u v , ( T u f t e d H a i r g r a s s ) H o r d e u m b r a c h y a n t h e r u m N e v s k i . ( M e a d o w B a r l e y ) " A n t h o x a n t h u m o d o r a t u m L . ( S w e e t V e r n a l g r a s s ) D i s t i c h l i s s p i c a t a ( L . ) G r e e n e ( S a l t g r a s s ) H i e r o c h l o e o d o r a t a ( L . ) B e a u v . ( S e n e c a G r a s s ) D a c t y l i s g l o m e r a t a L . ( O r c h a r d - g r a s s ) C y n o s u r u s c r i s t a t u s L . l ^ u r e s t e d D o g ' s - t a i 1) H o l c u s l a n a t u s L . ( V e l v e t - g r a s s ) F e s t u c a p r a t e n s i s H u d s . ( M e a d o w F e s c u e ) x x x x x C a r e x l y n g b y e i H o r n e n . ( L y n g b y ' s S e d g e ) S c i r p u s m i c r o c a r p u s P r e s l . ( S m a l l - f r u i t B u l r u s h ) E l e o c h a r i s p a l u s t r i s ( L . ) R . & S, ( S p i k e - r u s h ) D x - 247 -Community 1 2 Rush Juncus b a l t i c u s W i l l d . x x ( B a l t i c Rush) Juncus e f f u s u s L. x (Common Rush) - 248 -Table 22. Area of each p l a n t community on the Big Qualicum R i v e r Estuary P l a n t Community Area Percent (hectares) (%) Carex l y n g b y e i 1.3 50.0 Anthoxanthum odoratum - P o t e n t i l l a p a c i f i c a 1.3 50.0 Plantago l a n c e o l a t a - T r i f o l i u m  pratense TOTAL 2.6 100.0 - 249 -Fi g u r e 12. Map showing the d i s t r i b u t i o n of the p l a n t communities on the Courtenay R i v e r Estuary Legend Scale lcm=48m Pl a n t Community 1 S c i r p u s americanus 2 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i 3 S c i r p u s microcarpus - Carex l y n g b y e i 4 Carex l y n g b y e i 5 P o t e n t i l l a p a c i f i c a - T r i f o l i u m w o r m s k j o l d i i 6 P h a l a r i s arundinacea - Rumex s a l i c i f o l i u m - Elymus m o l l i s 7 Deschampsia c e s p i t o s a - E l e o c h a r i s p a l u s t r i s - P o t e n t i l l a p a c i f i c a 8 P i c e a s i t c h e n s i s 9 Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i 10 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i - S i d a l c e a h e n d e r s o n i i 11 Typha l a t i f o l i a 12 Oenanthe sarmentosa 13 G l y c e r i a o c c i d e n t a l i s A a g r i c u l t u r e B l o g boom D dredged I i n d u s t r i a l L t i d a l M marina SL s u b t i d a l U urban road i 1 b u i l d i n g F i g u r e 12 (cont.) P M u n i c i p a l i t y of Courtenay a i r s t r i p S M u n i c i p a l i t y of Courtenay sewage treatment p l a n t i to o I Table 23. D e s c r i p t i o n of the plant communities on the Courtenay River E s t u a r y , J u l y 1970. i 1 S c i r p u s americanus 2 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i 3 S c i r p u s microcarpus - Carex lyngbyei 4 Carex l y n g b y e i 5 P o t e n t i l l a p a c i f i c a - T r i f o l i u m wormskjoldii 6 P h a l a r i s arundinacea - Rumex s a l i c i f o l i u s - Elymus m o l l i s 7 Deschampsia c e s p i t o s a - E l e o c h a r i s p a l u s t r i s - P o t e n t i l l a p a c i f i c a 8 P i c e a s i t c h e n s i s 9 Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i 10 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i - S i d a l c e a h e n d e r s o n i i 11 Typha l a t i f o l i a 12 Oenanthe sarmentosa 13 G l y c e r i a o c c i d e n t a l i s Following i s a l i s t of the s p e c i e s i n each p l a n t community i n c l u d i n g dominant s p e c i e s i n d i c a t e d by a D and subdominant s p e c i e s i n d i c a t e d by an S. 1 2 TREES Alnus rubra Bong. (Red A l d e r ) Populus t r i c h o c a r p a T. & G. (Cottonwood) Pyrus fusca Raf. ( P a c i f i c Crabapple) P i c e a s i t c h e n s i s (Bong.) Carr. ( S i t k a Spruce) Sorbus s c o p u l i n a Greene (Mountain-ash) Cornus s t o l o n i f e r a Michx. (Red-osier Dogwood) SHRUBS Physocarpus c a p i t a t u s (Pursh) Kuntze (Ninebark) Sal ix sp. L. (Willow) Rubus u r s i n u s Chan. ? i Schlecht ( P a c i f i c B l a c k b e r r y ) Rosa gynnocarpa Nutt. ( L i t t l e Wild Rose) Community 3 4 5 6 7 8 9 10 11 12 13 D to O l to X SHRUBS (cont.) Spiraea d o u g l a s i i Hook (Hardhack) Rubus s p e c t a b i l i s Pursh (Salmonberry) C y t i s u s scoparius (L.) Link (Broom) Symphoricarpos albus (L.) Blake (Snowberry) Rosa nutkana P r e s l . (Nootka Rose) Lonicera i n v o l u c r a t a (Rich.) Banks (Black Twin-berry) Myrica gale L. (Sweet Gale) Community 8 9 10 11 12 13 x x X to CO X FORBS T r i g l o c h i n maritimum L. (Arrow-grass) S p e r g u l a r i a canadensis (Pers.) G. Don (Sandspurry) P o t e n t i l l a p a c i f i c a Howell ( C i n q u e f o i l ) Glaux maritima L. (Sea-milkwort) Lenna minor L. (Duckweed) Plantago maritima L. (Seaside P l a n t a i n ) T r i f o l i u m w ormskjoldii Lehm. (Springbank Clover) E r j g e r o n p h i l a d e l p h i c u s L. ( P h i l a d e l p h i a Fleabane) Sonchus a r v e n s i s L. ( M i l k - t h i s t l e ) Convolvulus sepium L. (Morning-glory) Equisetum arvense L. (Common H o r s e t a i l ) Community 1 2 3 4 .5 6 7 8 9 10 11 12 13 b X X X x D S x D x D S S x X X X D X X X X X X X X FORES (cont.) 1 Rumex c r i s p u s L. ( C u r l y Dock) Galium aparine L. (Bedstraw) Montia s i b i r i c a (L.) Hov/ell ( S i b e r i a n Miner's Lettuce) Epilobium a n g u s t i f o l i u m L. (Fireweed) Lupinus sp. L. -(Lupine) L i l a e o p s i s o c c i d e n t a l i s C o u l t . & Rose ( L i l a e o p s i s ) G r i n d e l i a i n t e g r i f o l i a DC. (Gumweed) Plantago macrocarpa Cham. & Schlecht ( P l a n t a i n ) S i s y r i n c h i u m a n g u s t i f o l i u m M i l l . (Blue-eyed Grass) C a s t i l l e j a l e v i s e c t a Greenm. (Golden I n d i a n - p a i n t brush) Community 2 3 4 5 G 7 8 9 10 11 12 13 x x x x x x x FORBS (cont.) Plantage l a n c e o l a t a L. (Ribgra s s ) V i c i a g i g a n t e a Hook. (Giant Vetch) C i r s i u m arvense (L.) Scop. (Canada T h i s t l e ) Typha l a t i f o l i a .L. ( C a t - t a i l ) Runex s a l i c i f o l i u s Wienm. (Willow Dock) Stachys cooleyae H e l l e r (Cooley's Hedge-nettle) T r i f o l i u n dubium S i b t h . (Least Hop Clover) Hypericum formosun H. R . K. ( S t . John's-wort) Hypochaeris r a d i c a t a L. (Hai r y Cats-ear) P t e r i d i u n aquilinum (L.) Kuhn (Bracken) A c h i l l e a n i l l e f o l i u m L. (Yarrow) Community 6 7 8 9 10 11 12 13 x x X x x x S S x x x x x X X to Oi X X 1 FORBS (cont.) Camassia quamash (Pursh) Greene (Camas) Dodecatheon pulchellum (Raf.) M e r r i l l ( S h o o t i n g - s t a r ) Ii i l i u m columbianum Hanson (Columbia L i l y ) Plantago major L. (Common P l a n t a i n ) P o t e n t i l l a p a c i f i c a Hov/ell ( C i n q u e f o i l ) Mentha s p i c a t a L. (Spearmint) As t e r hesperius Gray (Marsh Aster) Sium suave Walt. (Water-parsnip) Mentha a r v e n s i s L. ( F i e l d Mint) Heracleun 1anatum Michx. (Cow-parsnip) Oenanthe sarmentosa P r e s l . (Water-parsley) Community 8 0 10 11 12 13 x X X X X x x x x X X X X X X X X X x x x S x 1 FORBS (cont.) A t r i p l e x p a t u l a L. (Saltbush) Symphytum asperum Lepech. (Rough Comfrey) Myosotis s c o r p i o i d e s L. (Common forget-me-not) Epilobium w a t s o n i i Barbey (Watson's Willow-herb) C i c u t a d o u g l a s i i (DC.) C o u l t . & Rose (Water-hemlock) Habenaria d i l a t a t a (Pursh) Hook (Bog-candle) Lactuca b i e n n i s (Moench) Fern. ( T a l l Blue Lettuce) GRASSES Deschampsia c e s p i t o s a (L.) Beauv. ( T u f t e d H a i r g r a s s ) D i s t i c h l i s s p i c a t a (L.) Greene ( S a l t g r a s s ) D a c t y l i s glomerata L. (Orchard-grass) Community 2 3 4 5 6 7 8 9 10 11 12 13 x S x X X X X X X X X D x x x D x S 1 2 GRASSES (cont.) Elymus m o l l i s T r i n . (Wildrye) P h a l a r i s arundinacea L. (Reed Canarygrass) Phleum pratense L. (Timothy) G l y c e r i a o c c i d e n t a l i s ( P i p e r ) N e l s . (Western Mannagrass) Holcus l a n a t u s L. ( V e l v e t - g r a s s ) Festuca p r a t e n s i s Huds. (Meadow Fescue) A g r o s t i s t e n u i s S i b t h . ( C o l o n i a l Bentgrass) Agropyron repens (L.) Beauv. (Couch Grass) Hordeum brachyantherum Nevski (Meadow B a r l e y ) Community 3 4 5 6 7 8 9 10 11 12 13 S D x x x x x x SEDGES Carex lyn g b y e i Hornem. (Lyngby's Sedge) Sc i r p u s microcarpus P r e s l . ( S m a l l - f r u i t Bulrush) S c i r p u s acutus Muh1. (Bulrush) E l e o c h a r i s p a l u s t r i s (L.) R. ft S. (Spike-rush) S c i r p u s americanus Pers. (Three-square Bulrush) RUSHES Juncus b a l t i c u s W i l l d . ( B a l t i c Rush) Juncus e f f u s u s L. (Common Rush) Juncus acuninatus Michx. (Tapered Rush) Community 1 2 3 4 5 G 7 8 9 10 11 12 13 D S D x D x X X D X D x X x X X D x X - 261 -Table 24. Area of each p l a n t community on the Courtenay R i v e r E s t u a r y P l a n t Community Area Percent ( h e c t a r e s ) (%) Carex l y n g b y e i 10.7 48.6 S c i r p u s americanus 7.1 32.3 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a 1.1 5.0 Carex l y n g b y e i Oenanthe sarmentosa .7 3.2 Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a - .5 2.3 Carex l y n g b y e i  S u b t o t a l 20.1 91.4 p i c e a s i t c h e n s i s , .5 2.3 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a .5 2.3 Carex l y n g b y e i - S i d a l c e a h e n d e r s o n i i P h a l a r i s arundinacea - Rumex s a l i c i f o l i u s .3 1.4 Elymus m o l l i s P o t e n t i l l a p a c i f i c a - T r i f o l i u m w o r m s k j o l d i i .2 .9 Typha l a t i f o l i a .2 .9 S c i r p u s microcarpus - Carex l y n g b y e i .1 .5 G l y c e r i a o c c i d e n t a l i s .1 .5 Deschampsia c e s p i t o s a - E l e o c h a r i s .01 .04 p a l u s t r i s - P o t e n t i l l a p a c i f i c a  TOTAL 22.01 100.24 - 262 -F i g u r e 13. Map showing the d i s t r i b u t i o n of the p l a n t communities on the Oyster R i v e r E s t u a r y Legend Scale lcm=48m P l a n t Community 1 Carex l y n g b y e i 2 Elymus m o l l i s - Carex l y n g b y e i 3 Juncus b a l t i c u s - D i s t i c h l i s s p i c a t a - A g r o s t i s a l b a 4 Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a 5 Pseudotsuga m e n z i e s i i - P t e r i d i u m a q u i l i n u m 6 Elymus m o l l i s C campground CU c u l v e r t D dredged L t i d a l M marina SL s u b t i d a l b u i l d i n g road — • - 263 -Table 25. D e s c r i p t i o n of the p l a n t communities on the Oyster R i v e r Estuary, September 1976 1 Carex l y n g b y e i 2 Elymus m o l l i s - Carex l y n g b y e i 3 Juncus b a l t i c u s - D i s t i c h l i s s p i c a t a - A g r o s t i s a l b a 4 Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a 5 Pseudotsuga m e n z i e s i i - P t e r i d i u m aquilinum 6 Elymus m o l l i s F o l l o w i n g i s l i s t of the s p e c i e s i n each p l a n t community i n c l u d i n g dominant s p e c i e s i n d i c a t e d by a D and subdominant s p e c i e s i n d i c a t e d by an S. - 264 -Trees Pseudotsuga m e n z i e s i i ( M i r b e l ) Franco (Douglas F i r ) P i c e a s i t c h e n s i s (Bong.) C a r r . ( S i t k a Spruce) Pyrus f u s c a Raf. ( P a c i f i c Crabapple) Thuja p l i c a t a Donn. (Western Red Cedar) Tsuga h e t e r o p h y l l a (Raf.) Sarg. (Western Hemlock) Acer macrophyllum Pursh (Common Maple) Shrubs Rosa nutkana P r e s l . (Nootka Rose) Physocarpus c a p i t a t u s (Pursh) Kuntze (Ninebark) L o n i c e r a i n v o l u c r a t a (Rich.) Banks (Bla c k Twin-berry) Symphoricarpos a l b u s (L.) Blake (Snowberry) B e r b e r i s nervosa Pursh (Oregon grape) Community 1 2 3 4 5 D x x X X X X X X X X X X - 2 6 5 -Forbs Sonchus a r v e n s i s L. ( M i l k - t h i s t l e ) A c h i l l e a m i l l e f o l i u m L. (Yarrow) G r i n d e l i a i n t e g r i f o l i a DC. (Gumweed) Plantago l a n c e o l a t a L. ( Ribgrass) Maianthemurn d i l a t a t u m (Wood) N e l s . & Macbr. ( F a l s e L i l y - o f - t h e - v a l l e y ) Plantago m a r i t i m a L. (Seaside P l a n t a i n ) P o t e n t i l l a p a c i f i c a Howell ( C i n q u e f o i l ) L i l a e o p s i s o c c i d e n t a l i s C o u l t . & Rose ( L i l a e o p s i s ) P t e r i d i u m aquilinum (L.) Kuhn. (Bracken) Lathyrus p a l u s t r i s L. (Marsh Pea) Polystichum munitum ( K a u l f . ) P r e s l . (Sword-fern) T i a r e l l a t r i f o l i a t a v a r . t r i f o l i a t a L. (Foamflower) Lactuca m u r a l i s (L.) F r e s r . (Wall L e t t u c e ) Rumex c r i s p u s L. ( C u r l y Dock) V i c i a g i g a n t e a Hook (Giant Vetch) Chenopodium album L. (Pigweed) A t r i p l e x p a i u l a L. (Saltbush) - 266 -Grasses Elymus m o l l i s T r i n . (Wildrye) D i s t i c h l i s s p i c a t a (L.) Greene ( S a l t g r a s s ) Poa p r a t e n s i s L. (Kentucky Bluegrass) A g r o s t i s a l b a L. (Bentgrass) Hordeum brachyantherum N e v s k i . (Meadow Ba r l e y ) Deschampsia c e s p i t o s a (L.) Beauv, (T u f t e d H a i r g r a s s ) Community 1 2 3 4 5 6 x D x D Sedge Carex l y n g b y e i Hornem. (Lyngby's Sedge) D S Rush Juncus b a l t i c u s W i l l d . ( B a l t i c Rush) x S D - 267 -Table 26. Area of each, p l a n t community on the Oyster R i v e r Estuary P l a n t Community Area Percent ( h e c t a r e s ) (%) Carex l y n g b y e i .7 52.2 Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a .2 14.9 Pseudotsuga m e n z i e s i i - P t e r i d i u m  a q u i l i n u m .2 14.9 Elymus m o l l i s - Carex l y n g b y e i S u b t o t a l .1 7.5 1.2 89.5 Juncus b a l t i c u s - D i s t i c h l i s s p i c a t a -A g r o s t i s a l b a .1 7.5 Elymus m o l l i s TOTAL .04 3.0 1.34 100.0 - 268 -Fi g u r e 14. Map showing the d i s t r i b u t i o n of the p l a n t communities on the Campbell R i v e r E s t u a r y Legend Scale 1 cm = 48 m P l a n t Community 1 Carex l y n g b y e i 2 P o t e n t i l l a p a c i f i c a - E l e o c h a r i s p a l u s t r i s 3 Deschampsia c e s p i t o s a 4 Alnus rubra - Rubus l a c i n i a t u s - C y t i s u s s c o p a r i u s -Rosa nutkana - Rubus s p e c t a b i l i s 5 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a 6 Myrica g a l e - S c i r p u s acutus 7 Ruppia maritima 8 Alnus r u b r a - Rosa nutkana - Symphoricarpos albus B l o g boom b u i l d i n g | 1 D dredged I i n d u s t r i a l road L l a n d f i l l M marina P a i r s t r i p t r a n s e c t j U urban - 269 -Table 27. D e s c r i p t i o n o i the p l a n t communities on the Campbell R i v e r E s t u a r y , J u l y , 1976 1 Carex l y n g b y e i 2 P o t e n t i l l a p a c i f i c a - E l e o c h a r i s p a l u s t r i s 3 Deschampsia c e s p i t o s a 4 Alnus r u b r a - Rubus l a c i n i a t u s - C y t i s u s s c o p a r i u s -Rosa nutkana - Rubus s p e c t a b i l i s 5 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a 6 Myrica g a l e - S c i r p u s acutus 7 Ruppia maritima 8 Alnus r u b r a - Rosa nutkana - Symphoricarpos al b u s F o l l o w i n g i s a l i s t of the s p e c i e s i n each p l a n t community i n c l u d i n g dominant s p e c i e s i n d i c a t e d by a D and subdominant s p e c i e s i n d i c a t e d by an S. - 270 -1 Trees Rhamnus purs h i a n a DC. (Cascara) Alnus rubra Bong. (Red A l d e r ) Pseudotsuga m e n z i e s i i ( M i r b e l ) Franco (Douglas F i r ) Pyrus f u s c a Raf. ( P a c i f i c Crabapple) Thuja p l i c a t a Donn. (Western Red Cedar) Acer macrophyllum Pursh. (Common Maple) Shrubs Holodi s c u s d i s c o l o r (Pursh) Maxim. (Ocean-spray) Rubus l a c i n i a t u s W i l l d . (Evergreen B l a c k b e r r y ) C y t i s u s s c o p a r i u s (L.) L i n k (Broom) Myrica g a l e L. (Sweet gale) Rosa nutkana P r e s l . (Nootka Rose) L o n i c e r a i n v o l u c r a t a (Rich.) Banks (Bla c k Twin-berry) Rubus s p e c t a b i l i s Pursh. (Salmonberry) Physocarpus c a p i t a t u s (Pursh) Kunt: (Ninebark) Community 2 3 4 5 6 7 8 x S S s x S x X X S S x x D S S x s s e x - 271 -1 2 Shrubs (cont.) Sambucus racemosa L. ( E l d e r b e r r y ) Rubus p a r v i f l o r u s Nutt. (Thimbleberry) Rosa gymnocarpa Nutt. ( L i t t l e Wild Rose) Ribes sp. L. (Gooseberry) S p i r a e a d o u g l a s i i Hook (Hardhack) B e r b e r i s nervosa Pursh (Oregongrape) Vaccinium p a r v i f o l i u m Smith (Huckleberry) Symphoricarpos albus (L.) Blake (Snowberry) Amelanchier a l n i f o l i a Nutt. ( S e r v i c e b e r r y ) Rubus pedatus J.E. Smith ( F i v e l e a v e d Bramble) Rubus u r s i n u s Cham. & S c h l e c h t ( P a c i f i c B l a c k b e r r y ) L o n i c e r a c i l i o s a (Pursh) DC. (Orange Honeysuckle) S a l i x sp. L. (Willow) Community 3 4 5 6 7 8 x x S x X X X X S X X S S X Forbs P o t e n t i l l a p a c i f i c a Howell x ( P a c i f i c Silverweed) E r i g e r o n p h i l a d e l p h i c u s L. ( P h i l a d e l p h i a Fleabane) Glaux maritima L. (Sea-milkwort) S i s y r i n c h i u m a n g u s t i f o l i u m M i l l (Blue-eyed grass) Typha l a t i f o l i a L. (Common C a t - t a i l ) P r u n e l l a v u l g a r i s L. ( S e l f - h e a l ) C i r s i u m arvense (L.) Scop. (Creeping T h i s t l e ) A q u i l e g i a formosa F i s c h . (Red Columbine) V i c i a g i g a n t e a Hook. (Giant Vetch) Equisetum arvense L. (Common H o r s e t a i l ) Epilobium a n g u s t i f o l i u m L. (Fireweed) F r i t i l l a r i a camschatcensis (L.) Ker-Gawl (Chocolate L i l y ) Maianthemum d i l a t a t u m (Wood) N e l s . & Macbr. ( F a l s e L i l y - o f - t h e - v a l l e y ) T r i g l o c h i n maritimum L. x (Arrow-grass) L i l a e o p s i s o c c i d e n t a l i s C o u l t . x &. Rose ( L i l a e o p s i s ) S i d a i c e a h e n d e r s o n i i Wats. (Henderson's Checker-mallow) - 273 -Forbs (cont.) T r i f o l i u m repens L. (White Clover) T r i f o l i u m w o r m s k j o l d i i Lehm. (Springbank Clover) T r i f o l i u m pratense L. (Red C l o v e r ) C a s t i l l e j a l e v i s e c t a Greenm. (Golden Indian-paintbrush) A c h i l l e a m i l l e f o l i u m L. (Yarrow) Rumex c r i s p u s L. (C u r l y Dock) Plantago l a n c e o l a t a L. (Ribgrass) Hypochaeris r a d i c a t a L. (Hairy Cats-ear) Plantago maritima L. (Sea P l a n t a i n ) Plantago macrocarpa Cham. & Schlecht ( A l a s k a P l a n t a i n ) Montia p e r f o l i a t a (Donn) Howell (Miner's L e t t u c e ) Rumex a c e t o s e l l a L. (Sour Weed) Lupinus sp. L. (Lupine) T r i f o l i u m dubium S i b t h . (Least Hop Clover) Montia s i b i r i c a (L.) Howell ( S i b e r i a n Miner's L e t t u c e ) Galium sp. L. (Bedstraw) Community 2 3 4 5 6 7 8 x x x x x x x x x x X X x x x X X X X X X X X X X X X X - 274 -1 2 Forbs (cont.) Polystichum muni turn (Kaulf .) P r e s l . (Sword Fern) T r i l l i u m ovatum Pursh ( T r i l l i u m ) L i l i u m columbianum Hanson (Columbia L i l y ) T r i f o l i u m t r i d e n t a t u m L i n d l . (Sand C l o v e r ) Myosotis d i s c o l o r P ers. (Yellow and Blue Forget-me-not) Achlys t r i p h y l l a (Smith) DC. ( V a n i l l a l e a f ) P t e r i d i u m aquilinum (L.) DC. (Bracken) Streptopus a m p l e x i f o l i u s (L.) DC. ( T w i s t e d - s t a l k ) Convolvulus sepium L. (Morning-glory) Heracleum lanatum Michx. (Cow-parsnip) Habenaria d i l a t a t a (Pursh) Hook (Bog-candle) Oenanthe sarmentosa P r e s l . Hypericum formosum H.B.K. ( S t . John's-wort) Ranunculus orthorhynchus Hook (Buttercup) Lathyrus p a l u s t r i s L. (Marsh Pea) Dodecatheon p u l c h e l l u m (Raf.) M e r r i l l ( S h o o t i n g - s t a r ) Community 3 4 5 6 7 8 x x X X X X X X X X X X X X X X X X X X X X X - 275 -Community 1 2 3 4 5 Forbs (cont.) S p e r g u l a r i a canadensis x (Pers.) G. Don (Sandspurry) Ruppia maritima L. ( D i t c h - g r a s s ) Grasses Holcus l a n a t u s L. x x ( V e l v e t - g r a s s ) Deschampsia c e s p i t o s a (L.) x x D D Beauv ( T u f t e d H a i r g r a s s ) Hordeum brachyantherum Nevski x x (Meadow B a r l e y ) H i e r o c h l o e odorata (L.) x Beauv. ( V a n i l l a g r a s s ) Elymus m o l l i s T r i n . x (Dune Wildrye) A i r a sp. L. ( H a i r g r a s s ) Festuca sp. L. x x (Fescue) A g r o s t i s t e n u i s S i b t h . ( C o l o n i a l Bentgrass) x Sedges Carex l y n g b y e i Hornem. (Lyngby's Sedge) E l e o c h a r i s p a l u s t r i s (L.) R. & S. (Spike-rush) S c i r p u s microcarpus P r e s l . ( S m a l l - f r u i t Bulrush) S c i r p u s americanus P e r s . (Three-square Bulrush) S c i r p u s acutus Muhl. (Hardstem Bulrush) Carex l e n t i c u l a r i s v a r . l i m n o p h i l a (Holm) (Sedge) Cronq. Rushes Juncus a r t i c u l a t u s L. ( J o i n t e d Rush) Juncus b a l t i c u s W i l l d . ( B a l t i c Rush) Juncus e f f u s u s L. (Common Rush) - 277 -Table 28. Area of each p l a n t community on the Campbell R i v e r Estuary P l a n t Community Area Percent ( h e c t a r e s ) (%) Alnus r u b r a - Rubus l a c i n i a t u s - 10.3 31.9 C y t i s u s s c o p a r i u s - Rosa nutkana -Rubus s p e c t a b i l i s Deschampsia c e s p i t o s a 9.7 30.0 Carex l y n g b y e i 5.0 15.5 Alnus r u b r a - Rosa nutkana - 4.1 12.7  Symphoricarpos albus S u b t o t a l 29.1 90.1 P o t e n t i l l a p a c i f i c a - E l e o c h a r i s 2.2 6.8 p a l u s t r i s Deschampsia c e s p i t o s a - P o t e n t i l l a .9 2.8 p a c i f i c a M y rica g a l e - S c i r p u s acutus .1 .3 Ruppia maritima .04 .1 TOTAL 32.3 100.1 - -z/a -F i g u r e 15. Map showing the d i s t r i b u t i o n of the p l a n t communities on the Salmon R i v e r Estuary-Legend Scale 1 cm = 48 m P l a n t Community 1 Heracleum lanatum - Juncus b a l t i c u s 2 Carex l y n g b y e i 3 Juncus b a l t i c u s - Poa p r a t e n s i s - P o t e n t i l l a p a c i f i c a 4 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 5 E l e o c h a r i s p a l u s t r i s 6 Poa p r a t e n s i s - A g r o s t i s a l b a v a r . s t o l o n i f e r a -P o t e n t i l l a p a c i f i c a 7 Heracleum lanatum - P o t e n t i l l a p a c i f i c a 8 Elymus m o l l i s 9 Deschampsiua c e s p i t o s a - Carex l y n g b y e i 10 Deschampsia c e s p i t o s a - Poa p r a t e n s i s - Juncus b a l t i c u s 11 Rosa nutkana - P o t e n t i l l a p a c i f i c a - A g r o s t i s a l b a v a r . s t o l o n i f e r a - Poa p r a t e n s i s 12 Agropyron repens - Poa p r a t e n s i s - A c h i l l e a m i l l e f o l i u m 13 P o t e n t i l l a p a c i f i c a - A c h i l l e a m i l l e f o l i u m 14 D a c t y l i s glomerata 15 Deschampsia c e s p i t o s a - Juncus b a l t i c u s 16 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a 17 Elymus glaucus - Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a 18 D a c t y l i s glomerata - Deschampsia c e s p i t o s a 19 Plantago l a n c e o l a t a - F r i t i l l a r i a camschatcensis 20 Bromus c a r i n a t u s - Poa p r a t e n s i s - P o t e n t i l l a p a c i f i c a 21 Glaux maritima - D i s t i c h l i s s p i c a t a - 279 -22 A g r o s t i s a l b a v a r . s t o l o n i f e r a - Plantago l a n c e o l a t a 23 Hordeum brachyantherum 24 G r i n d e l i a i n t e g r i f o l i a 25 Typha l a t i f o l i a B l o g boom b u i l d i n g '— D dredged F f e r r y t e r m i n a l I i n d u s t r i a l dyke ||| L t i d a l LD l o g dump M marina road — S sewage treatment lagoon SL s u b t i d a l U urban t r a n s e c t | - 2 8 0 -Table 29. D e s c r i p t i o n of the p l a n t communities on the Salmon R i v e r E s t u a r y 1 Heracleum lanatum - Juncus b a i t i c u s 2 Ca:rex l y n g b y e i 3 Juncus b a l t i c u s - Poa p r a t e n s i s - P o t e n t i l l a p a c i f i c a 4 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 5 E l e o c h a r i s p a l u s t r i s 6 Poa p r a t e n s i s - A g r o s t i s a l b a v a r . s t o l o n i f e r a -P o t e n t i l l a p a c i f i c a 7 Heracleum lanatum - P o t e n t i l l a p a c i f i c a 8 Elymus m o l l i s 9 Deschampsia c e s p i t o s a - Carex l y n g b y e i 10 Deschampsia c e s p i t o s a - Poa p r a t e n s i s - Juncus b a l t i c u s 11 Rosa nutkana - P o t e n t i l l a p a c i f i c a - A g r o s t i s a l b a v a r . s t o l o n i f e r a - Poa p r a t e n s i s 12 Agropyron repens - Poa p r a t e n s i s - A c h i l l e a m i l l e f o l i u m 13 P o t e n t i l l a p a c i f i c a - A c h i l l e a m i l l e f o l i u m 14 D a c t y l i s glomerata 15 Deschampsia c e s p i t o s a - Juncus b a l t i c u s 16 Deschampsia c e s p i t o s a - P o t e n t i l l a p a c i f i c a 17 Elymus glaucus - Juncus b a l t i c u s - P o t e n t i l l a p a c i f i c a 18 D a c t y l i s glomerata - Deschampsia c e s p i t o s a 19 Plantago l a n c e o l a t a - F r i t i l l a r i a camschatensis 20 Bromus c a r i n a t u s - Poa p r a t e n s i s - P o t e n t i l l a p a c i f i c a 21 Glaux maritima - D i s t i c h l i s s p i c a t a 22 A g r o s t i s a l b a v a r . s t o l o n i f e r a - Plantago l a n c e o l a t a 23 Hordeum brachyantherum 24 G r i n d e i i a i n t e g r i f o l i a 25 Typha l a t i f o l i a F o l l o w i n g i s a l i s t of the s p e c i e s i n each p l a n t community i n c l u d i n g dominant s p e c i e s i n d i c a t e d by a D and subdominant s p e c i e s i n d i c a t e d by an S. TREES Alnus rubra Bong. (Red Alder) Pyrus fusca Raf. ( P a c i f i c Crabapple) Picea sitchensis (Bong.) Carr. (Sitka Spruce) Tsuga heterophylla (Raf.) Sarg. (Western Hemlock) Acer macrophyllum Pursh. (Cjornmon Maple) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 x D D to to I Cornnunity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 SHRUBS Lonicera involucrata (Rich.) Banks x x x x (Black Twin-berry) Rubus spectabilis Pursh. , S x x (Salmonberry) Rosa nutkana Presl. x D (Nootka Rose) Salix hookeriana Barratt x (Hooker Willow) Salix sp. (Willow) Cytisus scoparius (L.) Link (Broom) Sambucus racemosa L. S (Elderberry) Vaccinium parvifolium Smith x (Huckleberry) Holodiscus discolor (Pursh) Maxim. x (Ocean-spray) Rubus ursinus Cham. & Schlecht x (Pacific Blackberry) Osmaronia cerasiformis (T. & G.) Greene (Indian Plum) x SHRUBS (cont.) Rubus parviflorus Nutt. (Thimbleberry) FORBS F r i t i l l a r i a camschatcensis (L.) (Chocolate L i l y ) Ker.-Gawl. Trifolium pratense L. (Red Clover) Plantago lanceolata L. (Ribgrass) Heracleum lanatum Michx. (Cow-parsnip) Potentilla pacifica Howell (Cinquefoil) Triglochin rnaritimum L. (Arrow-grass) Sisyrinchium angustifolium M i l l . (Blue-eyed Grass) Trifolium wormskjoldii Lehm. (Springbank Clover) Plantago macrocarpa (Plantain) Cham. & Schlecht Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 X X X x x x x x x x x x x S S x S x D D x x x D x x x x x x S x S x D D x x x D x S x D S x S D x x S x x x x x S x X X X X X X X X X X X X X X X X Carinumty 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 FORBS (cont.) Achillea nallefolium L. x x x x x x x S S x x x x x x S (Yarrow) Lilaeopsis occidentalis x x (Lilaeopsis) Coult. & Rose Montia s i b i r i c a (L.) Howell x x x x (Siberian Miner's Lettuce) Vi c i a gigantea Hook. x x x x (Giant Vetch) Lathyrus palustris L. x x x (Marsh Pea) Ranunculus uncinatus D. Don x x x ( L i t t l e Buttercup) Epilobium angustifolium L. x x (Fireweed) Dodecatheon pulchellum S x x (Shooting-star) (Raf.) Merrill Sium suave Walt. x x x x x x x (Water-parsnip) C a s t i l l e j a levisecta Greenm. x x (Golden Indian-paintbrush) Pteridium aquilinum (L.) Kuhn x x (Bracken) FQRBS (cont.) Ranunculus repens L. (Creeping Buttercup) Geum macrophyllum Willd. (Avens) Plantago maritima L. (Seaside Plantain) Grindelia i n t e g r i f o l i a DC. (Gumweed) Trifolium repens L. (White Clover) D i g i t a l i s purpurea L. (Foxglove) Hypochaeris radicata L. (Hairy Cats-ear) Sidalcea hendersonii Wats. (Henderson's Checker-mallow) Prunella vulgaris L. (Self-heal) Petasites frigidus (L.) Fries (Coltsfoot) Arctium minus (Hill) Bernh. (Common Burdock) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 x x X X X D X X X X X X X X Cormiunity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 FORBS (cont.) Stachys cooleyae Heller (Cooley's Hedge-nettle) Qsmorhiza chllensis H. & A. (Sweet-root) Adenocaulon bicolor Hook (Pathfinder) Galium sp. L. (Bedstraw) Allium geyeri var. tenerum Jones (Geyer's Onion) Plantago major L. (Common Plantain) Barbarea orthoceras Ledeb. (Wintercress) Glaux maritima L. (Sea-mi lkwor t) Taraxacum officinale Weber (Common Dandelion) Rumex crispus L. (Curly Dock) Erigeron philadelphicus L. (Philadelphia Fleabane) x x x x x x x x x x x x x x x D x x x x x S x X X X X X x x x 1 2 FORBS (cont.) Maianthemum dilatatum (Wood ) Nels. & (False Lily-of-the-valley) Macbr. Cirsium arvense (L.) Scop. x (Canada Thistle) Cirsium vulgare (Savi) Tenore (Common Thistle) Galium trifidum L. (Small Bedstraw) Corallorhiza mertensiana Bong. (Western Coral-root) Polystichum munitun (Kaulf.) Presl. (Sword-fern) Gymnocarpium dryopteris (L.) Newm. (Oak-fern) Ti a r e l l a t r i f o l i a t a var. t r i f o l i a t a L. (Foamflower) Athyrium filix-femina (L.) Roth. (Lady-fern) Lactuca muralis (L.) Fresen. (Wall Lettuce) Rumex acetosella L. (Sour Weed) Community 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 x x x x X X X X to 00 00 X X X X X 1 2 3 4 5 6 7 FORBS (cont.) Blechnum spicant (L.) Roth x (Deer-fern) Spergularia canadensis (Pers.) G. Don x (Canada Sandspurry) Salicornia virginica L. (Saltwort) Equisetum arvense L. (Conmon Horsetail) Mentha arvensis L. (Field Mint) Capsella bursa-pastoris (L.) Medic. (Shepherd's-purse) Typha l a t i f o l i a L. (Cat-tail) Atriplex patula L. (Saltbush) Corrinunity 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 x x S to 00 CD X x S x GRASSES Phleum pratense L. (Timothy) Holcus lanatus L. (Velvet-grass) Deschampsia cespitosa (L.) Beauv. (Tufted Hairgrass) D i s t i c h l i s spicata (L.) Greene (Saltgrass) Elymus mollis Trin. (Wildrye) Hierochloe odorata (L.) Beauv. (Seneca Grass) Poa pratensis L. (Kentucky Bluegrass) Agrostis alba var. stolonifera L. (Bentgrass) Hordeum brachyantherum Nevski. (Meadow Barley) Br anus carinatus H. & A. (Brome-grass) Elymus glaucus Buckl. (Blue Wildrye) Conmunity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 x x x x x X X X X X x x x D D x D D x D x x S x D x x x x x S x x x x x x S D S x D S S x x x x x D x D S x x x D x x x x x x D x x x x D x x x x x D x x 1 2 3 4 5 6 7 8 GRASSES (cont.) Agropyron repens (L.) Beauv. x (Couch Grass) Agrostis tenuis Sibth. (Colonial Bentgrass) Festuca rubra L. x (Red Fescue) Dactylis glomerata L. x x (Orchard-grass) SEDGES Eleocharis palustris (L.) R. & S. x x x D (Spike-rush) Carex lyngbyei Hornem. x D x x (Lyngby's Sedge) Eleocharis pauciflora (Lightf.) Link x (Few-flowered Spike-rush) RUSHES Juncus balticus Willd. D D x x S x (Blatic Rush) Luzula divaricata Wats. x (Woodrush) Community 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 D x x x x D x D i to CD I x S x S x x x x D x x D D S x x - zwz -Table 30. Area of each plant corrinunity on the Salmon River Estuary Plant Cbmmunity Area Percent (hectares) {%) Picea sitchensis - Tsuga heterophylla 46.9 ' 29.5 Poa pratensis - Agrostis alba var. stolonifera -Potentilla pacifica 23.6 14.9 Carex lyngbyei 22.1 13.9 Deschampsia cespitosa - Carex lyngbyei 18.9 11.9 Deschampsia cespitosa - Potentilla pacifica 14.7 9.3 Elymus glaucus - Juncus balticus - Potentilla pacifica 14.4 9.1 Heracleum lanatum - Juncus balticus 6.1 3.8 Subtotal L46.7 92.4 Elymus mollis 2.5 1.6 Juncus balticus - Poa pratensis - Potentilla pacifica 2.2 1.4 Dactylis glomerata 1.5 .9 Rosa nutkana - Potentilla pacifica - Agrostis alba var. stolonifera - Poa pratensis 1.4 .9 Heracleum lanatum - Potentilla pacifica 1.2 .8 Agropyron repens - Poa pratensis - Achillea millefolium .8 .5 Plantago lanceolata - F r i t i l l a r i a camschatcensis .4 ' .3 Bromus carinatus - Poa pratensis - Potentilla pacifica .4 .3 Deschampsia cespitosa - Poa pratensis - Juncus balticus .3 .2 Glaux maritima - D i s t i c h l i s spicata .3 .2 Agrostis alba var. stolonifera - Plantago lanceolata .3 .2 Dactylis glomerata - Deschampsia cespitosa .2 .1 Hordeum brachyantherum .2 .1 Grindelia i n t e g r i f o l i a .2 .1 Eleocharis palustris .1 .06 - 293 -Plant Cbrrriunity Area Percent (hectares) (%) Deschampsia cespitosa - Juncus balticus .1 .06 Potentilla pacifica - Achillea millefolium .03 .02 Typha l a t i f o l i a .03 .02 TOTAL 158.9 100.2 - 294 -Figure 16. Map showing the distribution of the plant communities on the Adam-Eve Rivers' Estuary Legend Scale 1 cm = 48 m Plant Ccmmunity 1 Salicornia virginica - Triglochin maritimum - Plantago maritima -Glaux maritima 2 Carex lyngbyei 3 Deschampsia cespitosa - Carex lyngbyei - Glaux maritima -Potentilla pacifica 4 Poa pratensis - Potentilla pacifica 5 Picea sitchensis - Tsuga heterophylla 6 Picea sitchensis - Elymus mollis 7 Hordeum brachyantherum - Salicornia virginica 8 Deschampsia cespitosa 9 Deschampsia cespitosa - Juncus gerardii 10 Deschampsia cespitosa - Salicornia virginica 11 Deschampsia cespitosa - Agropyron repens ' 12 Picea sitchensis B log boom building D dredged I industrial ZZZZZ road L t i d a l zone SL subtidal Table 31. Description of the plant communities on the Adam-Eve Rivers' Estuary, June 1975. 1 Salicornia virginica - Triglochin maritimum - Plantago maritima -Glaux maritima 2 Carex lyngbyei 3 Deschampsia cespitosa - Carex lyngbyei - Glaux maritima -Potentilla pacifica 4 Poa pratensis - Potentilla pacifica 5 Picea sitchensis - Tsuga heterophylla 6 Picea sitchensis - Elymus mollis 7 Hordeum brachyantherum - Salicornia virginica 8 Deschampsia cespitosa 9 Deschampsia cespitosa - Juncus gerardii 10 Deschampsia cespitosa - Salicornia virginica 11 Deschampsia cespitosa - Agropyron repens 12 Picea sitchensis Following i s a l i s t of the species in each plant community including dominant species indicated by a D and subdominant species indicated by an S. - 296 -Commun i t y 1 2 3 4 5 6 7 Trees P i c e a s i t c h e n s i s (Bong.) C a m . D D ( S i t k a Spruce) A lnus rubra Bong. x (Red A Ider) Pyrus fusca R a f . x x ( P a c i f i c Crabapp le) Tsuga heterophy I la ( R a f . ) S a r g . D (Western Hemlock) Pseudotsuga m e n z i e s i i ( M i r b e l ) Franco (Doug las F i r ) Shrubs Rosa nutkana P r e s I. x x (Nootka Rose) Rubus s p e c t a b i l i s P u r s h . x x (Salmonberry) Gau I the r ia sha I Ion P u r s h . x (Sa la I) Vaccinium o v a l i f o l . i u m Smith x (B lueberry ) L o n i c e r a invo lucr .a ta ( R i c h . ) Banks x (Black T w i n - b e r r y ) Vaccin ium p a r v i f o l i u m Smith x (Huck leberry ) R ibes sp . L. x (Gooseberry) A rc tos taphy los nevadensis Gray (Ki nn ik i nn i ck ) Rubus urstnus Cham. & Sch lecht ( P a c i f i c B l a c k b e r r y ) x 1 Forbs S a l i c o r n i a v i r g i n i c a L. S (Sa Itwort) T r i g l o c h i n maritimum L. S (Ar row-grass ) P lantago m a r i t i m a L. S (Seas ide P l a n t a i n ) Glaux m a r i t i m a L. S (Sea-mi Ikwort) P o t e n t i I la pac i f i c a Howe I I (Ci nquefoi I) T r i f o L i u m wormskj o Idi i Lehm. (Springbank C l o v e r ) E p i l o b i u m a n g u s t i f o I imn L. (Fireweed) Ach i I lea m i l le fo I ium L. (Yarrow) F r i t i I l a r i a camschatcens is ( L . ) (Chocolate L i ly) G e r - G a w l . E r i g e r o n ph i lade Iph icus L. (Ph i lade Iph ia F leabane) V i c i a g igantea Hook (G iant Vetch) Heracleum lanatum Michx . (Cow-parsn ip) A r c t i u m minus (Hi I I) Be rnh . (Common Burdock) Maianthemum d,i latatum (Wood) Ne Is. & Macbr . ( F a l s e L i l y - o f - t h e - v a I ley) Ranuncu lus s p . L. x (But tercup) S i urn suave Wa It. (Water -parsn ip) - 297 -Commun i 2 3 4 5 6 7 x D x x x X X X S S x x S S D x x x S x x X X x x x x X X X X X X X X X X - 298 -Commun i t y 1 2 3 4 5 6 7 Forbs (cont inued) C i r s i u m arvense (Canada Th i s t le) Tr i e n t a I i s a r c t i c a F i sch (Sjtarf lower) P o l y s t ichum munltum ( K a u l f . ) P res I. (Sword - fern ) Aqu i legi a formosa F i s c h . (Red Co lumbi ne) Hypochaer is r a d i c a t a L. (Hai ry C a t s - e a r ) Anaphi l i s m.argar.itacea ( L . ) B. & H. (Pear l y - e v e r l a s t i n g ) V i c i a g igantea Hook (Giant Vetch) Galium asperrimum Gray (Rough Bedstraw) Gal ium apar ine L. (Bedstraw) La thy rus p a l u s t r i s L. (Marsh Pea) G r i n d e l i a i n t e g r i f o l i a DC. (Gumweed) S p e r g u l a r i a canadensis ( P e r s . ) (Sandspurry) G. Don Coch l e a r i a o f f i c i n a l i s L. (Spoonwort) Cuscuta s a l i n a Enge lm. ( S a l t - m a r s h Dodder) Mont ia s i b i r i c a ( L . ) Howell ( S i b e r i a n M i n e r ' s L e t t u c e ) L i l a e o p s i s o c c i d e n t a l i s Cou I t . & Rose (L i laeops is ) x - 299 -1 2 Forbs (cont inued) H ierac ium a l b i f l o r u m Hook (Wh i te - f l owered Hawkweed) Lactuca m u r a l i s ( L . ) F r e s e n . (Wa I I L e t t u c e ) Polypodium g l ycy r rh - i za D.C. E a t . ( L i c o r i c e - f e r n ) C o r a l l o r h i z a mertens iana Bong. (Western Cora l - r o o t ) F r a g a r i a c h i l o e n s i s ( L . ) Duchesne (Coas ta l St rawberry ) Rumex acetose I. la L. (Sour Weed) Commun i t y 3 4 5 6 7 8 9 10 11 12 x x X X X A t r i p lex patu la L. x x x x x x x (Sa l tbush ) Z o s t e r a mar ina L. x (Ee l - g r a s s ) S t e I l a r i a humfusa R o t t b . x x x x (Spreading Starwor t ) Ranuncu lus or thorhynchus Hook x (But tercup) - 300 -1 Grasses Deschampsia c e s p i t o s a (L . ) x (Tufted H a i r g r a s s ) Beauv. D i s t i c h l i s s p i c a t a ( L . ) Greene (Sa I tgrass) Poa p r a t e n s i s L. (Kentucky B l u e g r a s s ) E lymus mo I l i s Tr i n . (Wi Idrye) Ho leus lanatus L. (Ve I v e t - g r a s s ) Hordeum brachyantherum Nevsk i . (Meadow Bar ley) Fes tuca rubra L. (Red Fescue) Agropyron repens (L . ) Beauv. (Couch Grass) Fes tuca subu lata Tr i n. (Nodding Fescue) Commun i t y 2 3 4 5 6 7 8 9 10 1 1 1 2 D x x D D D D D x x x x D x x x x x x x D x S x x x x x D S Sedges Carex lyngbyei Hornem. (Lyngby's Sedge) Rushes Juncus g e r a r d i i L o i s e I. (Mud Rush) Table 32. Area of each plant cornmunity on the Adam-Eve Rivers' Estuary Plant Community Area Percent (hectares) (%) Salicornia virginica - Triglochin maritimum - 7.3 24.7 Plantago maritima - Glaux maritima Picea sitchensis - Elymus mollis 4.6 15.6 Deschampsia cespitosa - Salicornia virginica 4.4 14.9 Carex lyngbyei 3.1 10.5 Deschampsia cespitosa - Carex lyngbyei - Glaux 2.8 9.5 maritima - Potentilla pacifica Poa pratensis - Potentilla pacifica 1.8 6.1 Picea sitchensis - Tsuga heterophylla 1.3 4.4 Hordeum brachyantherum - Salicornia virginica 1.3 4.4 Subtotal 26.6 90.1 Deschampsia cespitosa 1.0 3.4 Deschampsia cespitosa - Juncus gerardii .8 2.7 Picea sitchensis .6 2.0 Deschampsia cespitosa - Agropyron repens .5 1.7 TOTAL 29.5 99.9 Figure 17. Map showing the distribution of the plant conrnunities the Tsitika River Estuary Legend Scale 1 cm = 48 m Plant Community 1 Carex lyngbyei 2 Carex lyngbyei - Deschampsia cespitosa 3 Elymus mollis - Vicia gigantea 4 Picea sitchensis - Heracleum lanatum - Vi c i a gigantea 5 Picea sitchensis 6 Deschampsia cespitosa 7 Trifolium wormskjoldii - Achillea millefolium - Triglochin maritimum - Holcus lanatus L t i d a l SL subtidal - 304 -Table 33. Description of the plant <xmiunities cn the Tsitika River Estuary, July, 1976. 1 Carex lyngbyei 2 Carex lyngbyei - Deschampsia cespitosa 3 Elymus mollis - Vicia gigantea 4 Picea sitchensis - Heracleum lanatum - Vi c i a gigantea 5 Picea sitchensis 6 Deschampsia cespitosa 7 Trifolium wormskjoldii - Achillea millefolium - Triglochin martimum - Holcus lanatus Following i s a l i s t of the species in each plant community including dominant species indicated by a D and subdominant species indicated by an S. - cSUO -Trees Picea sitchensis (Bong.) Carr. (Sitka Spruce) Shrubs Lonicera involucrata (Rich.) Banks (Black Twin-berry) Forbs Glaux maritima L. (Sea-milkwort) Potentilla pacifica Howell (Cinquefoil) Plantago maritima L. (Seaside Plantain) Spergularia canadensis (Pers.) G. Don (Sandspurry) Vi c i a gigantea Hook. (Giant Vetch) Galium asperrimum Gray (Rough Bedstraw) Heracleum lanatum Michx. (Cow-parsnip) Montia pa r v i f o l i a (Moc.) Greene (Miner's Lettuce) Montia s i b i r i c a (L.) Howell (Siberian Miner's Lettuce) Triglochin maritimum L. (Arrow-grass) Adenocaulon bicolor Hook. (Pathfinder) Community 1 2 3 4 5 D D x x x x x x X X X D D S' x x x x D S X X X Penanthe sarmentosa Presl. (Water-parsley) x x - 306 -Forbs (continued) T i a r e l l a t r i f o l i a t a var. unifoliata (Hook) (Foamflower) Trifolium wormskjoldii Lehm. (Springbank Clover) Achillea millefolium L. (Yarrow) Plantago lanceolata L. (Ribgrass) Rumex crispus L. (Curly Dock) Galium trifidum L. (Small Bedstraw) Atriplex patula L. (Saltbush) S t e l l a r i a humifusa Rottb. (Spreading Starwort) Corrinunity 1 2 3 4 5 x x X X Grasses Deschampsia cespitosa (L.) Beauv. (Tufted Hairgrass) Elymus mollis Trin. (Wildrye) Holcus lanatus L. (Velvet-grass) Hordeum brachyantherum Nevski. (Meadow Barley) x D x D x x x x X X X X X Grasses (continued) Festuca subulata Trin. (Nodding Fescue) Bromus pacificus Shear (Pacific Brome) Agrostis tenuis Sibth. (Colonial Bentgrass) Sedges Carex lyngbyei Hornem. . (Lyngby's Sedge) Rushes Juncus balticus Willd. (Baltic Rush) - 308 -Table 34. Area of each plant ccirrnTunity on the Tsitika River Estuary Plant Corrimunity Area Percent (hectares) (%) Carex lyngbyei - Deschampsia cespitosa 1.2 31.1 Deschampsia cespitosa 1.2 31.1 Carex lyngbyei .9 23.3 Elymus mollis - Vi c i a gigantea .4 10.4 Subtotal 3.7 95.9 Picea sitchensis .1 2.6 Trifolium wormskjoldii - Achillea millefolium - .04 1.0 Triglochin naritimum - Holcus lanatus Picea sitchensis - Heracleum lanatum - Vicia gigantea .02 .5 TOTAL 3.9 100.0 Figure 18. Map showing the distribution af the plant canrcunities on the Kokish River Estuary Legend Scale 1 cm = 48 m Plant Gcirimuriity 1 Deschampsia cespitosa - Potentilla pacifica 2 Carex lyngbyei 3 Deschampsia cespitosa - Potentilla pacifica - Plantago macrocarpa 4 Heracleum lanatum - Pyrus fusca 5 Elymus mollis 6 Atriplex patula - Spergularia canadensis 7 Salicornia virginica - Spergularia canadensis - D i s t i c h l i s spicata 8 Salicornia virginica - Potentilla pacifica - Plantago rnaritima 9 Hordeum brachyantherum 10 Deschampsia cespitosa - Potentilla pacifica - Hordeum brachyantherum - Agrostis tenuis 3 log boom x x x railway F ferry terminal ' I industrial Z Z road L t i d a l 0 o i l storage tank SL subtidal - 310 -Table 35. Description of the plant communities on the Kokish River Estuary, August 1975. 1 Deschampsia cespitosa - Potentilla pacifica 2 Carex lyngbyei 3 Deschampsia cespitosa - Potentilla pacifica - Plantago macrocarpa 4 Heracleum lanatum - Pyrus fusca 5 Elymus mollis 6 Atriplex patula - Spergularia canadensis 7 Salicornia virginica - Spergularia canadensis - D i s t i c h l i s spicata 8 Salicornia virginica - Potentilla pacifica - Plantago maritima 9 Hordeum brachyantherum 10 Deschampsia cespitosa - Potentilla pacifica - Hordeum  brachyantherum - Agrostis tenuis Following i s a l i s t of the species in each plant community including dominant species indicated by a D and subdominant species indicated by an S. - 311 -Trees A Inus rubra Bong. (Red A I der) P i c e a s i t c h e n s i s (Bong.) C a r r . ( S i t k a Spruce) Pyrus f u s c a R a f . ( P a c i f i c Crabapp Ie) Commun i t y 1 2 3 4 5 6 Shrubs Myr i ca g a l e L. (Sweet Ga Ie) Gau I t h e r i a s h a l l o n P u r s h . (Sa la I) Rubus spec tab i l i s P u r s h . (SaImonberry) L o n i c e r a i n v o l u c r a t a ( R i c h . ) Banks (B lack Twin -ber ry ) Forbs Potent i I la p a c i f i c a Howe II D x D x (Ci nquefoi I) G laux mar i t ima L. x x x (Sea-mi Ikwort) T r i f o l i u m w o r m s k j o l d i i Lehm. x (Springbank C l o v e r ) Ach i I lea m i l l e fo I ium L. x (Yarrow) Heracleum lanatum Michx . (Cow-parnsip) x D - 312 -Commun i ty 1 2 3 4 5 6 Forbs (cont inued) T r i g l o c h i n maritimum L. (Ar row-grass ) V i c i a g igantea Hook. (G iant Vetch) P lantago m a r i t i m a L. (Seas ide P l a n t a i n ) S i s y r i n c h i u m a n g u s t i f o l i u m M i l l . (B lue-eyed Grass) P lantago macrocarpa Cham. & Sch lecht ( P l a n t a i n ) Oenanthe sarmentosa Pres I. (Water -pars ley) F r i t i I l a r i a camschatcensis ( L . ) K e r . - G a w l (Chocolate L i ly) Maianthemum d i l a t a t u m (Wood) N e l s . & Macbr. (Fa Ise L i l y - o f - t h e - v a I ley) Hypochaeris rad i c a t a L. (Hairy C a t s - e a r ) Sa Hcorn ia v. irg i n i c a L. (Sa Itwort) C o c h l e a r i a o f f i c i n a l i s L. (Spoonwort) S p e r g u l a r i a canadensis ( P e r s . ) G. Don (Canada Sandspurry) S.tel l a r i a humifusa R o t t b . (Spreading Starwor t ) A t r ip lex patu la L. (Sa Itbush) x x D Grasses Deschampsia c e s p i t o s a ( L . ) Beauv. (Tufted H a i r g r a s s ) Hordeum brachyantherum Nevski (Meadow Bar ley) Poa pratens i s L. (Kentucky B l u e g r a s s ) Agropyron repens (L . ) Beauv. (Couch Grass) H i e r o c h l o e odorata ( L . ) Beauv. (Seneca Grass) Bromus s i t c h e n s i s T r i n . (Brome-grass) Ho Icus lanatus L. (Ve I v e t - g r a s s ) E lymus mo I l i s Tr i n . (Wl Idrye) D i s t i c h l i s s p i c a t a ( L . ) Greene (Sa I tgrass) A g r o s t i s t e n u i s S i b t h . ( C o l o n i a l Bentgrass ) ' Sedge Carex lyngbyei Hornem. (Lyngby 's ' Sedge) - 314 Table 36. Area of each plant ccranunity on the Kokish River Estuary Plant Community Area (hectares) Percent (%) Carex lyngbyei 2.0 26.2 Salicornia virginica - Potentilla pacifica -Plantago maritima 1.5 19.7 Hordeum brachyantherum 1.1 14.4 Deschampsia cespitosa - Potentilla pacifica 1.0 13.1 Deschampsia cespitosa - Potentilla pacifica -Plantago macrocarpa .7 9.2 Deschampsia cespitosa - Potentilla pacifica -Hordeum brachyantherum - Agrostis tenuis .7 9.2 Subtotal 7.0 91.8 Salicornia virginica - Spergularia canadensis -Dis t i c h l i s spicata .3 3.9 Elymus mollis .2 2.6 Heracleum lanatum - Pyrus fusca .1 1.3 Atriplex patula - Spergularia canadensis .03 .4 TOTAL 7.6 100.0 ^ 315 -Figure 19. Map showing the distribution of the plant communities on the Nimpkish River Estuary Legend Scale 1 cm = 48 m Plant Community 1 Deschampsia cespitosa - Potentilla pacifica 2 Juncus balticus - Trifolium wormskjoldii - Potentilla pacifica 3 GRAMINEAE - Juncus balticus - Rubus spectabilis 4 Eleocharis palustris 5 Deschampsia cespitosa - Trifolium wormskjoldii - Potentilla pacifica 6 Carex lyngbyei 7 Picea sitchensis - Rosa nutkana - Ribes sp. 8 Carex obnupta 9 Myrica gale 10 Deschampsia cespitosa - Potentilla pacifica - Juncus balticus 11 Typha l a t i f o l i a 12 Elymus mollis B barge L t i d a l 0 old log dump SL subtidal - 316 -Table 37. Description of the plant immunities on the Nimpkish River Estuary, August 1975. 1 Deschampsia cespitosa - Potentilla pacifica 2 Juncus balticus - Trifolium wormskjoldii - Potentilla pacifica 3 GRAMINEAE - Juncus balticus - Rubus spectabilis 4 Eleocharis palustris 5 Deschampsia cespitosa - Trifolium wormskjoldii - Potentilla pacifica 6 Carex lyngbyei 7 Picea sitchensis - Rosa nutkana - Ribes sp. 8 Carex obnupta 9 Myrica gale 10 Deschampsia cespitosa - Potentilla pacifica - Juncus balticus 11 Typha l a t i f o l i a 12 Elymus mollis Following i s a l i s t of the species in each plant community including dominant species indicated by a D and subdominant species indicated by an S. Trees Alnus rubra Bong. (Red Alder) Picea sitchensis (Bong.) Carr. (Sitka Spruce) Pyrus fusca Raf. (Pacific Crabapple) Shrubs Rubus spectabilis (Salmonberry) Myrica gale L. (Sweet Gale) Rosa nutkana Presl. (Nootka Rose) Ribes sp. L. (Gooseberry) Forbs Trifolium wormskjoldii Lehm. (Springbank Clover) Plantago maritima L. (Seaside Plantain) Community 1 2 3 4 5 6 7 8 9 10 11 12 D D D x D x x D i OJ I X X 1 Forbs (continued) Montia s i b i r i c a (L.) Howell x (Siberian Miner's Lettuce) Potentilla p a c i f i c a Howell S (Cinquefoil) Plantago macrocarpa Chem. & Schlecht (Plantain) Plantago lanceolata L. (Ribgrass) Triglochin maritimum L. (Arrow-grass) Spergularia canadensis (Pers.) G. Don. (Canada Sandspurry) Dodecatheon pulchellum (Raf.) Merrill (Shooting-star) Sisyrinchium angustifolium M i l l . (Blue-eyed Grass) Glaux maritima L. x (Sea-milkwort) Prunella vulgaris L. (Self-heal) Heliopsis helianthoides (L.) Sweet (Ox-eye Daisy) Community 2 3 4 5 6 7 8 9 10 11 12 S x x D x S x D x x x S X X X X X X X CO 1 Forbs (continued) Plantago lanceolata L. (Ribgrass) Hypochaeris radicata L. (Hairy Cats-ear) Ranunculus repens L. (Creeping Buttercup) Erigeron philadelphicus L. (Philadelphia Fleabane) Equisetum arvense L. (Common Horsetail) Achillea millefolium L. (Yarrow) Lilaeopsis occidentalis Coult. & Rose (Lilaeopsis) Galium t r i f idum L. (Small Bedstraw) Qenanthe sarmentosa Presl. (Water-parsley) Rumex crispus L. (Curly Dock) Athyrium filix-femina (L.) Roth. (Lady-fern) Community 6 7 8 9 10 11 12 x X X x x x x CO I—1 CD x x x Cornmunity 1 2 3 4 5 6 7 8 Forbs (continued) Lysichitum americanum Hulten & St. John x (Skunk Cabbage) Urtica dioica L. x (Stinging Nettle) Trifolium repens L. x (White Clover) Stachys cooleyae Heller x (Cooley's Hedge-nettle) Mentha arvensis L. x (Field Mint) Cochlearia o f f i c i n a l i s L. x (Spoonwort) Typha l a t i f o l i a L. (Cat-tail) Galium boreale L. (Northern Bedstraw) Atriplex patula L. (Saltbush) x 9 10 11 12 co CO o D x G r a s s e s D e s c h a m p s i a c e s p i t o s a ( L . ) Beauv . D ( T u f t e d H a i r g r a s s ) H o l c u s l a n a t u s L . ( V e l v e t - g r a s s ) P o a p r a t e n s i s L . ( K e n t u c k y B l u e g r a s s ) C a l a m a g r o s t i s c a n a d e n s i s ( M i c h x . ) Beauv . ( B l u e j o i n t R e e d g r a s s ) Bromus s i t c h e n s i s T r i n . ( B r o m e - g r a s s ) D a c t y l i s g l o m e r a t a L . ( O r c h a r d - g r a s s ) Ph leum p r a t e n s e L . ( T i m o t h y ) Hordeum b r a c h y a n t h e r u m N e v s k i (Meadow B a r l e y ) D i s t i c h l i s s p i c a t a ( L . ) Greene ( S a l t g r a s s ) E l y m u s m o l l i s T r i n . ( W i l d r y e ) Obmmunity 5 6 7 8 9 10 11 12 D x x D x S x x x x x x x x x x x x x x x x Cynosurus cristatus L. (Crested Dog's-tail) Agrostis tenuis Sibth. (Colonial bentgrass) Sedge Carex lyngbyei Hornem. (Lyngby's Sedge) Eleocharis palustris (L.) R. & S. (Spike-rush) Scirpus microcarpus Presl. (Small-fruit Bulrush) Carex oederi Retz. (Green Sedge) Carex obnupta Bailey (Slough Sedge) Rushes Juncus balticus Willd. (Baltic Rush) Juncus effusus L. (Common Rush) Juncus falcatus E. Meyer (Sickle-leaved Rush) Community 1 2 3 4 5 6 7 8 9 10 11 12 S S x x x x x x D x S x D x x x x x x x S x D S S x x D x x - 323 -Table 38. Area of each plant community on the Nimpkish River Estuary Plant Community Area Percent (hectares) (%) Juncus balticus - Trifolium wormskjoldii -Potentilla pacifica 2.3 28.6 Deschampsia cespitosa - Potentilla pacifica 1.2 14.9 Carex lyngbyei .9 11.2 Carex obnupta .8 10.0 Eleocharis palustris .6 7.5 Deschampsia cespitosa - Potentilla pacifica -Juncus balticus .6 7.5 Picea sitchensis - Rosa nutkana - Ribes sp. .6 7.5 Deschampsia cespitosa - Trifolium wormskjoldii -Potentilla pacifica .5 6.2 Subtotal 7.5 9.3.4 Myrica gale .3 3.7 GRAMINEAE - Juncus balticus - Rubus spectabilis .1 1.2 Elymus mollis .1 1.2 Typha l a t i f o l i a .03 .4 TOTAL 8.0 99.9 - 324 -Figure 20. Map showing the distribution of the plant cctrmunities on the Cluxewe River Estuary Legend Scale 1 cm = 48 m Plant Community 1 Carex lyngbyei 2 Carex lyngbyei - Deschampsia cespitosa 3 Potentilla pacifica - Trifolium wormskjoldii - Agrostis tenuis 4 Trifolium wormskjoldii - Deschampsia cespitosa - Poa pratensis -Potentilla pacifica 5 Deschampsia cespitosa - Potentilla pacifica 6 Hordeum brachyantherum - Trifolium wormskjoldii - Potentilla pacifica 7 Plantago lanceolata - Juncus effusus - Hordeum brachyantherum 8 Elymus mollis 9 Picea sitchensis 10 GRAMINEAE 11 Potentilla pacifica - Agrostis tenuis 12 Heracleum lanatum - Anthoxanthum odoratum - Dactylis glomerata 13 Poa pratensis - Salicornia virginica 14 Deschampsia cespitosa - Triglochin maritimum 15 Carex lyngbyei - Triglochin maritimum 16 Carex rostrata 17 Deschampsia cespitosa - Elymus mollis 18 Achillea millefolium - Poa pratensis - Elymus mollis C L SL campground t i d a l subtidal -H-+ dyke ZZZZ road - 3 2 0 -Table 39. Description of the plant cornrnunities on the Cluxewe River Estuary, July 1975. 1 Carex lyngbyei 2 Carex lyngbyei - Deschampsia cespitosa 3 Potentilla pacifica - Trifolium wormskjoldii - Agrostis tenuis 4 Trifolium wormskjoldii - Deschampsia cespitosa - Poa pratensis -Potentilla pacifica 5 Deschampsia cespitosa - Potentilla pacifica 6 Hordeum brachyantherum - Trifolium wormskjoldii - Potentilla pacifica 7 Plantago lanceolata - Juncus effusus - Hordeum brachyantherum 8 Elymus mollis 9 Picea sitchensis 10 GRAMINEAE 11 Potentilla pacifica - Agrostis tenuis 12 Heracleum lanatum - Anthoxanthum odoratum - Dactylis glomerata 13 Poa pratensis - Salicornia virginica 14 Deschampsia cespitosa - Triglochin maritimum 15 Carex lyngbyei - Triglochin maritimum 16 Carex rostrata 17 Deschampsia cespitosa - Elymus mollis 18 Achillea millefolium - Poa pratensis - Elymus mollis Following i s a l i s t of species in each plant community including dominant species indicated by a D and subdominant species indicated by an S. Trees Alnus rubra Bong. (Red Alder) Pyrus fusca Raf. (Pacific Crabapple) Picea sitchensis (Bong.) Carr. (Sitka Spruce) Tsuga heterophilla (Raf.) Sarg. (Western Hemlock) Shrubs Rubus spectabilis Pursh. (Salmonberry) Lonicera involucrata (Rich.) Banks (Black iVin-berry) Rosa nutkana Presl. (Nootka Rose) Ribes sanguineum Pursh. (Red Currant) Vaccinium parvif olium Smith (Huckleberry) Gaultheria shallon Pursh. (Salal) ~ Rubus parvif lorus Nutt. (Thimbleberry) Community 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 x x X X x D x x x x x x x x S x D X Forbs Potentilla p a c i f i c a Howell (Cinquefoil) Trifolium wormskjoldii Lehm. (Springbank Clover) Achillea millefolium L. (Yarrow) Plantago maritima L. (Seaside Plantain) Plantago lanceolata L. (Ribgrass) Salicornia v i r g i n i c a L. (Saltwort) Triglochin maritimum L. (Arrow-grass) Heracleum lanatum Michx. (Cow-parsnip) Equisetum arvense L. (Common Horesetail) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 x x D S D S x x D x S D x D x x x x x x x x x S S x S x x x x x P to o x D ' x x x x D S x x x x x x S S x Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Forbs (continued) Ranunculus sp. L. x (Buttercup) Stachys cooleyae Heller x (Cooley's Hedge-nettle) Oenanthe sarmentosa Presl. x (Water-parsley) Maianthemum dilatatum (Wood) Nels. & Macbr. x x (Fase Lily-of-the-valley) F r i t i l l a r i a camschatcensis (L.) Ker. - Gawl. x ' (Chocolate L i l y ) ^ Glaux maritima L. x x x (Sea-milkwort) Lathyrus palustris L. S x (Marsh Pea) Vic i a gigantea Hook x (Giant Vetch) Trifolium repens L. x x (White Clover) Community 1 2 , 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Forbs (continued) Plantago lanceolata L. x x x (Ribgrass) Fragaria chiloensis (L.) Duchesne x x (Coastal Strawberry) C a s t i l l e j a levisecta Greenra. x x (Golden Indian-paintbrush) Lilaeopsis occidentalis Coult. &. Rose x (Lilaeopsis) Trientalis arctica Fisch. x 1 (Starflower) w cc Galium aparine L. x , (Bedstraw) Atriplex patula L. . x x x x (Saltbrush) S t e l l a r i a humifusa Rottb. x x x (Spreading Starwort) Grasses Deschampsia cespitosa (L.) Beauv. (Tufted Hairgrass) Poa pratensis L. (Kentucky Bluegrass) Hordeum brachyantherum Nevski. (Meadow Barley) D i s t i c h l i s spicata (L.) Greene (Saltgrass) Holcus lanatus L. (Velvet-grass) Elymus mollis Trin. (Wildrye) Aira sp. (Hairgrass) Dactylis glomerata L. (Orhard-grass) Hierochloe odorata (L.) Beauv. (Seneca Grass) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 D x S D x x S x x D x D S x S x x x x S x x D x S x x x D D S S x x x D x x x x x D S x S Grasses (continued) Anthoxan odoratum L. (Sweet Vernalgrass) Agrostis tenuis Sibth. (Colonial Bentgrass) Sedges Carex lyngbyei Hornem. (Lyngby's Sedge) Scirpus microcarpus Presl. (Small-fruit Bulrush) Carex rostrata Stokes (Beaked Sedge) Rushes Juncus balticus Willd. (Baltic Rush) Juncus effusus L. (Common Rush) Community i 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 S S x x x x D x x x D D x x D D cc cc H D x X - 332 -Table 40. Area of each plant community on the Cluxewe River Estuary Plant Cbrnmunity Area (hectares) Percent (%) Poa pratensis - Salicornia virginica 16.2 31.3 Picea sitchensis 14.6 28.2 Elymus mollis 7.6 14.7 Carex lyngbyei 4.0 7.7 Carex lyngbyei - Deschampsia cespitosa 2.5 4.8 Deschampsia cespitosa - Triglochin iraxitimum 2.1 4.1 Subtotal 47.0 90.8 Potentilla pacifica - Agrostis tenuis 1.2 2.3 GRAMINEAE .9 1.7 Plantago lanceolata - Juncus effusus - Hordeum brachyantherum .5 1.0 Heracleum' lanatum - Anthoxanthum odoratum -Dactylis glomerata .5 1.0 Deschampsia cespitosa - Potentilla pacifica .4 .8 Hordeum brachyantherum - Trifolium wormskjoldii -Potentilla pacifica .3 .6 Deschampsia cespitosa - Elymus mollis .3 .6 Potentilla pacifica - Trifolium wormskjoldii -Agrostis tenuis .2 .4 Achillea millefolium - Poa pratensis - Elymus mollis .2 .4 Carex lyngbyei - Triglochin maritimum .1 .2 Carex rostrata .1 .2 Trifolium wormskjoldii - Deschampsia cespitosa — Poa pratensis - Potentilla pacifica .01 .02 Total 51.7 100.0 - 333 -Figure 21. Map showing the distribution of the plant communities on the Quatse River Estuary Legend Scale 1 cm = 48 m Plant Community 1 Carex lyngbyei - Potentilla pacifica 2 Carex lyngbyei 3 Deschampsia cespitosa - Carex lyngbyei - Potentilla pacifica 4 Carex lyngbyei - Agrostis tenuis - Oenanthe sarmentosa 5 Gaultheria shallon 6 Juncus balticus - Potentilla pacifica 7 Deschampsia cespitosa - Triglochin maritimum - Plantago maritima 8 Carex lyngbyei - Deschampsia cespitosa 9 Rubus spectabilis - Agrostis tenuis 10 Holcus lanatus - Agrostis tenuis 11 Agrostis tenuis - Juncus ensifolius var. ensifolius 12 Alnus rubra - Rubus spectabilis 13 Thuja plicata - Tsuga heterophylla 14 Deschampsia cespitosa - Agrostis tenuis - Potentilla pacifica 15 Carex lyngbyei - Triglochin maritimum 16 Salicornia virginica 17 Triglochin maritimum B log boom building H houseboat I industrial " | | | dyke L t i d a l ID log dump ZZZZ road SL subtidal - 334 -Table 41. Description of the plant comrminities on the Quatse River Estuary, July 1975. 1 Carex lyngbyei - Potentilla pacifica 2 Carex lyngbyei 3 Deschampsia cespitosa - Carex lyngbyei - Potentilla pacifica 4 Carex lyngbyei - Agrostis tenuis - Penanthe sarmentosa 5 Gaultheria shallon 6 Juncus balticus - Potentilla pacifica 7 Deschampsia cespitosa - Triglochin iraritimum - Plantago maritima 8 Carex lyngbyei - Deschampsia cespitosa 9 Rubus spectabilis - Agrostis tenuis 10 Holcus lanatus - Agrostis tenuis 11 Agrostis tenuis - Juncus ensifolius var. ensifolius 12 Alnus rubra - Rubus spectabilis 13 Thuja plicata - Tsuga heterophylla 14 Deschampsia cespitosa - Agrostis tenuis - Potentilla pacifica 15 Carex lyngbyei - Triglochin maritimum 16 Salicornia virginica 17 Triglochin maritimum Following i s a l i s t of species in each plant community including dominant species indicated by a D and subdominant species indicated by an S. Trees Thuja plicata Donn. (Western Red Cedar) Picea sitchensis (Bong.) Carr. (Sitka Spruce) Tsuga heterophylla (Raf.) Sarg. (Western Hemlock) Alnus rubra Bong. (Red Alder) Pinus contorta var. contorta Dougl. (Lodgepole Pine) Pyrus fusca Raf. (Pacific Crabapple) Shrubs Rubus spectabilis Pursh. (Salmonberry) Lonicera involucrata (Rich.) Banks (Black Twin-berry) Gcflimunity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 S x D S x x S S x D x x D x 1 CO to Ul X X X , x S S D S X X X X Community ^ 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Shrubs (continued) Gaultheria shallon Pursh. D x x (Salal) Menziesia ferruginea Smith S (Mock Azalea) Vaccinium parvifolium Smith S x x (Huckleberry) Rosa nutkana Presl. x x x (Nootka Rose) i Sambucus racemosa L. x x oo (Elderberry) g Salix sp. L. x x x 1 (Willow) Spiraea douglasii Hook x x (Hardhack) Physocarpus capitatus (Purch) Kuntze x (Ninebark) Rubus parviflorus Nutt. x (Thimbleberry) Forbs Hypochaeris radicata L . (Hairy Cats-ear) Glaux maritima L . (Sea-milkwort) Plantago maritima L . (Seaside Plantain) Sal icornia v i r g i n i c a L . (Saltwort) Rumex acetosel la L . (Sour Weed) Cirsium arvense (L . ) Scop. (Canada Thist le) Rumex crispus L . (Curly Dock) Lactuca muralis (L . ) Fresen. (Wall Lettuce) S t e l l a r i a media (L . ) C y r i l l . (Chickweed) Gc*imunity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 1 8 X X X X X S X x x x D S x x x x x x x D x x x Forbs (continued) Epilobium angustifolium L. (Fireweed) Arabis glabra (L.) Bernh. (Towermustard) Armeria maritima (Mill.) Willd. (Sea-pink) Cochlearia o f f i c i n a l i s L. (Spoonv/ort) Mentha arvensis L. (Field Mint) Blechnum spicant (L.) Roth (Deer-fern) Geum macrophyllum Willd. (Avens) Ranunculus repens L. (Creeping Buttercup) Ranunculus sp. L. (Buttercup) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 X X X X X X X X X X X X X Oornmunity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Forbs (continued) Potentilla pacifica Howell D S x D x x x x D (Cinquefoil) Plantago macrocarpa Cham. & Schlecht x x x x (Plantain) Trifolium wormskjoldii Lehm. x x x x S x x x (Springbank Clover) Triglochin rnaritimum L. x x x D x x x x S x D (Arrow-grass) , Achillea millefolium L. x x x x x x co (Yarrow) ^ i Heracleum lanatum Michx. x x x (Cow-parsnip) F r i t i l l a r i a camschatcensis (L.) Ker.-Gawl. x x x (Chocolate L i l y ) Oenanthe sarmentosa Presl. x S x x x x (Water-parsley) Maianthemum dilatatum (Wood) Nels. & Macbr. x x x x (False Lily-of-the-valley) Cbtnmunity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Forbs (continued) Erigeron philadelphicus L. x x (Philadelphia Fleabane) Montia s i b i r i c a (L.) Howell x x (Siberian Miner's Lettuce) Vicia gigantea Hook S x (Giant Vetch) Galium boreale L. x x x (Northern Bedstraw) Trientalis arctica Fisch. x x (Starflower) g Cornus canadensis L. x 1 (Bunchberry) Ti a r e l l a t r i f o l i a t a var. t r i f o l i a t a L. x (Foamflower) Lysichitum americanum Hulten & St. John x co (Skunk Cabbage) Pteridium aquilinum (L.) Kuhn (Bracken) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Forbs (continued) Trifolium pratense L. x x (Red Clover) Trifolium repens L. x x (White Clover) Spergularia canadensis (Pers.) G. Don x x x (Sandspurry) D i g i t a l i s purpurea L. x x (Foxglove) Prunella vulgaris L. x (Self-heal) Trifolium dubium Sibth. x (Least Hop Clover) Anaphalis margaritacea (L.) B. & H. x (Pearly-everlasting) Polystichum muniturn (Kaulf.) Presl. x (Sword-fern) Pteridium aquilinum (L.) Kuhn. x (Bracken) Forbs (continued) S t e l l a r i a humifusa Rottb. (Spreading Starwort) Atriplex patula L. (Saltbush) Grasses Deschampsia cespitosa (L.) Beauv. (Tufted Hairgrass) Holcus lanatus L. (Velvet-grass) D i s t i c h l i s spicata (L.) Greene (Saltgrass) Poa pratensis L. (Kentucky Bluegrass) Hordeum brachyantherum Nevski. (Meadow Barley) Agrostis tenuis Sibth. (Colonial Bentgrass) Community 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 x x S x x D D x x S D x x D x x CO to X X X X X X X X X x S x S D D x D Cbmmunity 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Grasses (continued) Hierochloe odorata (L.) Beauv. x (Seneca Grass) Agropyron repens (L.) Beauv. x x (Couch Grass) Sedges Carex lyngbyei Hornem. D D S D x D x x x D (Lyngby's Sedge) Scirpus microcarpus Presl. x x (Small-fruit Bulrush) Carex rostrata Stokes x (Beaked Sedge) Rushes Juncus balticus Willd. x x D (Baltic Rush) Juncus effusus L. x (Common Rush) OOTinumty 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Rushes (continued) Juncus ensifolius var. ensifolius Wikst. x x x S x (Dagger-leaf Rush) Juncus falcatus E. Meyer x (Sickle-leaved Rush) to - 345 -C Table 42. Area of each plant coniminity on the Quatse River Estuary Plant Corrmunity Area (hectares) Percent (%) Carex lyngbyei - Triglochin maritimum 6.6 28.7 Carex lyngbyei - Deschampsia cespitosa 2.8 12.2 Holcus lanatus - Agrostis tenuis 2.5 10.9 Alnus rubra - Rubus spectabilis 2.3 10.0 Deschampsia cespitosa - Agrostis tenuis -Potentilla pacifica 1.9 8.3 Salicornia virginica 1.9 8.3 Carex lyngbyei - Potentilla pacifica 1.1 4.8 Juncus balticus - Potentilla pacifica 1.0 4.3 Agrostis tenuis - Juncus ensifolius var. ensifolius .8 3.5 Subtotal 20.9 91.0 Thuja plicata - Tsuga heterophylla .6 2.6 Deschampsia cespitosa - Carex lyngbyei -Potentilla pacifica .5 2.2 Deschampsia cespitosa - Triglochin maritimum -Plantago maritima .4 1.7 Rubus spectabilis - Agrostis tenuis .3 1.3 Carex lyngbyei .1 .4 Gaultheria shallon .1 .4 Triglochin naritimum .1 .4 Carex lyngbyei - Agrostis tenuis -Oenanthe sarmentosa .01 .04 TOTAL 23.0 100.0 - 346 -Figure 22. Map showing the distribution of the plant communities on the Kingcome River Estuary Legend Scale 1 cm = 158.4 m Plant Community 1 Carex lyngbyei 2 Deschampsia cespitosa 3 Elymus mollis 4 Potentilla pacifica - Trifolium wormskjoldii - Juncus balticus 5 Salix sp. 6 Sambucus racemosa - Rubus spectabilis - Heracleum lanatum -Elymus mollis A agriculture I I I dyke B boat dock H Halliday's L logging company LD log dump Table 43. Description of the plant comiunities on the Kingcome River Estuary, July 1976. 1 Carex lyngbyei 2 Deschampsia cespitosa 3 Elymus mollis 4 Potentilla pacifica - Trifolium wormskjoldii - Juncus balticus 5 Salix sp. 6 Sambucus racemosa - Rubus spectabilis - Heracleum lanatum -Elymus mollis Following i s a l i s t of species in each plant community including dominant species indicated by a D and subdominant species indicated by an S. Trees Picea sitchensis (Bong.) Carr. (Sitka Spruce) Pyrus fusca Raf. (Pacific Crabapple) Shrubs Salix sp. L. (Willow) Lonicera c i l i o s a (Parsh) DC. (Orange Honeysuckle) Lonicera involucrata (Rich.) Banks (Black Twin-berry) Sambucus racemosa L. (Elderberry) Rubus spectabilis Pursh. (Salmonberry) Community 1 2 3 4 5 6 x x D x CO 00 S S Forbs (continued) Potentilla pacifica Howell (Cinquefoil) Sium suave Walt. (Water-parsnip) Triglochin maritimum L. (Arrow-grass) Plantago macrocarpa Cham. & Schlecht (Plantain) Glaux maritima L. (Sea-milkwort) Arctium minus (Hill) Reran. (Common Burdock) Trifolium wormskjoldii Lehm. (Springbank Clover) Erigeron philadelphicus L. (Philadelphia Fleabane) F r i t i l l a r i a camschatcensis (L.) Ker. - Gawl. (Chocolate L i l y ) CJommunity 2 3 4 5 6 x S x x x x x CO CD x x x Forbes (continued) Mentha arvensis L. (Field Mint) Galium sp. L. (Bedstraw) Maianthemum dilatatum (Wood) Nels. & Macbr. (False Lily-of-the-valley) Sonchus arvensis L. (Milk-thistle) Achillea millefolium L. (Yarrow) Heracleum lanatum Michx. (Cow-parsnip) Plantago major L. (Common Plantain) Penanthe sarmentosa Presl. (Water-parsley) Montia s i b i r i c a (L.) Howell (Siberian Miner's Lettuce) Community 1 2 3 4 5 6 x x x X X X X X X x x x x x x 1 CO Ol o X X X ^ X I X x x x X Forbes (continued) Lupinus sp. L. (Lupine) Ranunculus orthorhynchus Hook (Buttercup) lathyrus palustris L. (Marsh Pea) Grasses Festuca subulata Trin. (Nodding Fescue) Hordeum brachyantherum TTevski (Meadow Barley) Elymus mollis Trin. (Wildrye) Deschampsia cespitosa (L.) Beauv. (Tufted Hairgrass) Elymus glaucus Buckl. (Blue Wildrye) Hierochloe odorata (L.) Beauv. (Seneca Grass) Community 1 2 3 4 5 x x X x x x x x x X X D x x D x x x x x Ctomnunity 1 2 3 4 Grasses(continued) Festuca pratensis Huds. x x (Meadow Fescue) Agrostis tenuis Sibth. x x (Colonial Bentgrass) Dactylis glomerata L. (Orchard-grass) Bromus pacificus Shear x (Pacific Brome) Poa pratensis L. x (Kentucky Bluegrass) \ Sedge Carex lyngbyei Hornem. D D x (Lyngby's Sedge) Eleocharis palustris (L.) R. & R. x x (Spike-rush) Rush Juncus balticus Willd. (Baltic Rush) x S - Sob1 -Table 44. Area of each plant corrmunity on the Kingcome River Estuary Plant Community Area Percent (hectares) (%) Deschampsia cespitosa 74.2 51.2 Carex lyngbyei 41.6 28.7 P o t e n t i l l a p a c i f i c a - Trifolium wormskjoldii - 17.8 12.3 Juncus b a l t i c u s Subtotal 133.6 92.2 S a l i x sp. 8.5 5.9 Elymus m o l l i s 2.2 1.5 Sambucus racemosa - Rubus s p e c t a b i l i s - .5 .3 Heracleum lanatum - Elymus m o l l i s TOTAL 144.8 99.9 - 354 -Table 45. Simple linear correlation between species, and flow and precipitation Species r Flow r2-Precipitation r r 2 Agrostis spp. .2684 .0721 .4531 .2053 Carex lyngbyei .2026 .0411 .4528 .2050 Deschampsia cespitosa .4365 .1906 .7606* .5758 .05>P>.02 Di s t i c h l i s spicata .4944 .2444 .5624 .3163 Hordeum brachyantherum .1591 .0253 .4088 .1671 Hordeum murinum .2004 .0402 .4180 .1747 Juncus articulatus .0546 .00298 .1584 .0251 Juncus balticus .3682 .1356 .5192 .2696 Potentilla pacifica .1847 .0341 .9217** .8496 .001<P<.002 Salicornia'virginica .4311 .1858 .2670 .0713 Scirpus spp. .1366 .0187 .1377 .0190 n = 14 r0.05(2), 12 = 0- 532 n = 8 r0.05(2),6=0.707 - 355 -Appendix 6. L i v i n g , dead, senescent and duff f r a c t i o n s . F i g u r e s 23-41. The months i n the f i g u r e s are A p r i l ( 4 ) , May ( 5 ) , June (6), J u l y (7), August (8) and October (10). F i g u r e 23. Mean monthly dry weights (gm ,5m ) l i v i n g (.), dead (0) , senescent (A) and duff ( + ) pl a n t f r a c t i o n s from the Cowichan Carex lyngbyei community. S t a t i s t i c s f o r graph P o r t i o n Month Range Observations Minimum Maximum Std. Dev L i v i n g 4 5 6 7 8 10 13.1 140.7 45.5 168.5 78.5 8.5 5 5 5 5 5 5 34.9 80.3 213.5 116.4 0.0 4.4 48.0 221 .0 259.0 284.9 78.5 12.9 5.4 63.2 18.2 64.5 30.3 3.3 Dead 5 6 7 8 10 90.8 0.0 0.0 0.0 0.0 5 5 5 5 5 0.0 0.0 0.0 0.0 0.0 90.8 0.0 0.0 0.0 0.0 40.6 0.0 0.0 0.0 0.0 Senescent 4 5 6 7 8 10 0.0 20. 8 26.2 34.0 65.5 133.5 5 5 5 5 5 5 0.0 4.7 38.1 90.8 107.4 99.2 0.0 25. 5 64.3 124.8 172.9 232.7 0.0 8.7 11.0 12.7 26.4 53.0 Duff 5 6 7 8 10 105.5 28.2 17.4 21.1 0.0 5 5 5 5 5 0.0 0.0 0.0 0.0 0.0 105.5 28. 2 17.4 21.1 0.0 46.0 14.8 7.9 10.2 0.0 - 3 5 7 -F i g u r e 24. Mean monthly dry weights (gm .5m ) of l i v i n g (.), dead (0 ) , senescent (A) and duff (+) plant f r a c t i o n s from the L i t t l e Qualicum Carex lyngbyei community. S t a t i s t i c s f o r graph P o r t i o n Month Range Observations Minimum Maximum Std. De\ 4 213 .3 5 40. 6 253. 9 93. 8 5 19 .5 5 229. 7 249. 2 8. 3 L i v i n g 6 125 .4 5 188. 0 313. 4 50. 5 7 172 .8 5 83. 1 255. 9 70. 5 8 45 .8 5 101. 5 147. 3 22. 0 10 20 .6 5 6. 3 26. 9 8. 1 5 149 .2 5 0. 0 149. 2 69. 2 Dead 6 0 .0 5 0. 0 0. 0 0. 0 7 0 .0 5 0. 0 0. 0 0. 0 8 0 .0 5 0. 0 0. 0 0. 0 10 0 .0 5 0. 0 0. 0 0. 0 4 0 .0 5 0. 0 0. 0 0. 0 5 16 .7 5 22. 5 39. 2 7. 6 Senescent 6 106 .4 5 43. 7 150. 1 44. 1 7 160 .3 5 31. 8 192. 1 75. 6 8 102 .0 5 75. 2 177. 2 45. 6 10 151 .4 5 171. 7 323. 1 58. 5 5 110 .2 5 0. 0 110. 2 55. 1 6 83 .6 5 42. 4 126. 0 37. 6 Duff 7 248 .2 5 4. 3 252. 5 104. 5 8 105 .8 5 10. 9 116. 7 42. 6 10 57 .2 5 26. 1 83. 3 26. 0 GRRMS 0.0 34.875 69.75 104.63 139.5 174.38 209.25 244.13 279.0 F i g u r e 25. Mean monthly dry weights (gm .5m ) of l i v i n g (.), dead ( 0 ) , senescent (A) and duff (+) plant f r a c t i o n s from the Salmon Carex lyngbyei community. S t a t i s t i c s f o r graph P o r t i o n Month Range Observations Minimum Maximum Std. Dev, L i v i n g 4 5 6 7 8 10 57.2 42.9 77.4 36.5 46.8 52.6 5 5 5 5 5 5 46.1 111.4 169.1 187.8 127.4 2.4 103.3 154.3 246.5 224.3 174.2 55.0 21.5 17.6 31.5 16.2 18.4 22.9 Dead 5 6 7 8 10 36.4 0.0 0.0 0.0 0.0 5 5 5 5 5 0.0 0.0 0.0 0.0 0.0 36.4 0.0 0.0 0.0 0.0 16.6 0.0 0.0 0.0 0.0 co CTi O Senescent 4 5 6 7 8 10 0.0 14.3 13.6 38.6 56.9 63.1 5 5 5 5 5 5 0.0 2.7 15.7 41.4 53.8 124.9 0.0 17.0 29.3 80.0 110.7 188.0 0.0 5.9 5.6 16.9 22.8 23.0 Duff 5 6 7 8 10 61.2 27.6 27.4 33.7 35.0 5 5 5 5 5 20.4 15.3 19.6 19.2 2.2 81.6 42.9 47.0 52 .9 37.2 26.5 11.9 11.1 13.6 15.8 G R A M S D.O 25.315 5J.75 77.625 103.5 129.39 155.25 181.13 207.0 F i g u r e 26. Mean monthly dry weights (gm .5m ) l i v i n g (.), dead (0 ) , senescent ( A ) and d u f f (+) p l a n t f r a c t i o n s from the Cowichan Juncus b a l t i c u s community. S t a t i s t i c s f o r graph P o r t i o n Month Rang e Observations Minimum Maximum Std. De\ 4 16. 1 5 17 .0 33. 1 6 .6 5 51. 7 5 64 . 7 116. 4 20 .2 L i v i n g 6 46. 5 5 92 .9 139. 4 16 .8 7 37. 5 5 84 .6 122. 1 16 .6 8 45. 9 5 47 .7 93. 6 18 .8 10 29. 8 5 3 .7 33. 5 12 . 1 4 57. 4 5 88 .6 146. 0 21 .6 5 73. 8 5 28 .4 102. 2 30 .6 Dead 6 24. 2 5 37 .5 61. 7 9 .2 7 79. 0 5 43 .7 122. 7 33 .8 8 64. 3 5 26 .0 90. 3 27 .5 10 0. 0 5 0 .0 0. 0 0 .0 4 0. 0 5 0 .0 0. 0 0 .0 5 0. 0 5 0 .0 0. 0 0 .0 Senescent 6 0. 0 5 0 .0 0. 0 0 .0 7 1. 3 5 1 .5 2. 8 0 .5 8 46. 5 5 11 .3 57. 8 18 .9 10 90. 1 5 132 .0 222. 1 36 .8 5 73. 6 5 22 .3 95. 9 35 . 1 6 27. 2 5 , 26 .0 53. 2 12 .9 Duff 7 13. 9 5 9 .8 23. 7 5 .5 8 348. 1 5 9 .5 357. 6 151 .7 10 7. 8 5 14 .0 21. 8 2 .9 - 363 -F i g u r e 27. Mean monthly dry weights (gm . 5nf*) of l i v i n g (.), dead ( 0 ) , senescent (£>) and duff (+) plant f r a c t i o n s from the L i t t l e Qualicum P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i community. S t a t i s t i c s f o r graph P o r t i o n Month Range Observations Minimum Maximum S t d . Dev, L i v i n g 4 5 6 7 8 10 16.3 63.9 49.6 75.4 39.7 15.8 5 5 5 5 5 5 11.0 44.7 104.4 70.2 73.9 20.2 27.3 108.6 154.0 145.6 113.6 36.0 5.9 24.4 20.0 30.2 15.0 6.1 Dead 4 5 6 7 8 10 163.7 108.5 157.9 129.4 56.5 0.0 5 5 5 5 5 5 70.8 45.9 5.6 0.0 5.4 0.0 234.5 154.4 163.5 129.4 61.9 0.0 78.8 43.9 66.7 60.3 20.3 0.0 Senescent 4 5 6 7 8 10 0.0 0.0 0.0 27.4 42.9 125.7 5 5 5 5 5 5 0.0 0.0 0.0 15.5 20.1 98. 1 0.0 0.0 0.0 42.9 63.0 223. 8 0.0 0.0 0.0 10.8 15.6 49.9 Duff 5 6 7 8 10 24.4 37.6 59.2 114.7 18.1 5 5 5 5 5 16.3 5.2 0.0 14.2 9.9 40.7 42. 8 59.2 128.9 28.0 11.1 14.0 25.1 47.8 8.2 - 3 6 5 -F i g u r e 28. Mean monthly dry weights (gm .5m ) of l i v i n g (.), dead (0 ) , senescent and duff (+) plant f r a c t i o n s from the Campbell P o t e n t i l l a p a c i f i c a - E l e o c h a r i s p a l u s t r i s community. S t a t i s t i c s f o r graph P o r t i o n Month Range Observations Minimum Maximum S t d . Dev. L i v i n g 4 5 6 7 8 10 5, 46, 48, 77, 63, 28, 9 1 4 5 5 5 5 5 5 5 5 5 13.6 59.3 82.8 84.5 53.7 22.8 19.5 105.4 131.2 162.0 117.2 51.3 2.2 18.1 21 .2 29.8 24.6 10.9 Dead 4 5 6 7 8 10 19.6 9.9 38.7 22.8 0.0 0.0 5 5 5 5 5 5 29.6 11.0 3.6 3.1 0.0 0.0 49.2 20.9 42.3 25.9 0.0 0.0 8.8 3.8 16.3 9.7 0.0 0.0 Senescent 4 5 6 7 8 10 0.0 0.0 0.0 11.2 8.1 26.0 5 5 5 5 5 5 0.0 0.0 0.0 10.5 29.3 49.5 0.0 0.0 0.0 21.7 37.4 75.5 0.0 0.0 0.0 4.8 3.2 11.0 Duff 5 6 7 8 10 13.6 0.0 33.9 0.0 6.2 5 5 5 5 5 0.0 0.0 3.5 0.0 0.0 13.6 0.0 37.4 0.0 6.2 6.1 0.0 14.5 0.0 2.8 GRAMS F i g u r e 29. Mean monthly dry weights (gm ,5m-2) Q f l i v i n g (.), dead (0 ) , and d u f f (+) plant f r a c t i o n s from the Chemainus S a l i c o r n i a v i r g i n i c a community. P o r t i o n Month S t a t i s t i c s f o r graph Range Observations Minimum Maximum Std. Dev, L i v i n g 4 5 6 7 8 10 6.4 32.5 68.3 45.0 216.7 84. 1 5 5 5 5 5 5 3.5 54.4 49.6 136.3 72.1 39.8 9.9 86.9 117.9 181.3 288.8 123.9 2.6 12.8 25. 1 17.9 82.0 33.2 Dead 4 5 6 7 8 10 97.9 164.8 133.1 133.7 180.8 132.9 5 5 5 5 5 5 227.1 169. 1 223.0 221.3 284.4 106.3 325.0 333.9 356.1 355.0 465.2 239.2 41.2 64.3 53.0 49.9 86.1 56.9 Duff 5 6 7 8 10 63.1 39.9 0.0 0.0 0.0 5 5 5 5 5 3.4 17.3 0.0 0.0 0.0 66.5 57.2 0.0 0.0 0.0 24.8 15.3 0.0 0.0 0.0 F i g u r e 30. Mean monthly dry weights (gm .5m ) l i v i n g (.), dead (0 ) , and duff (+) plant f r a c t i o n s from the Chemainus D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a community. S t a t i s t i c s for graph P o r t i o n Month Range Observations Minimum Maximum Std. Dev, L i v i n g 4 5 6 7 8 10 35.8 56.8 72.1 39.3 138.8 47.7 5 5 5 5 5 5 1.0 22.1 59.0 135.6 93.3 151.4 36.8 78.9 131.1 174.9 232.1 199. 1 13.9 24.3 29.6 16.7 50.5 18.2 Dead 4 5 6 7 8 10 120. 1 89.4 94. 9 52.4 68.0 65.7 5 5 5 5 5 5 168.3 138.4 102.7 141.7 98.7 90.7 288.4 227.8 197.6 194. 1 166.7 156.4 52.7 39.1 38.6 23.1 26.4 28.0 Duff 4 5 6 7 8 10 0.0 31.8 32.4 18. 1 32.6 97.8 5 5 5 5 5 5 0.0 35.8 50.2 24.2 16.9 22. 1 0.0 67.6 82.6 42.3 49.5 119.9 0.0 13.1 12.2 6.6 12.3 37.4 - 371 -F i g u r e 31. Mean monthly dry weights (gm . 5m ) of l i v i n g (.), dead (0), senescent and d u f f ( + ) pl a n t f r a c t i o n s from the Campbell Carex l y n g b y e i community. P o r t i o n Month S t a t i s t i c s f o r graph Range Observations Minimum Maximum Std. Dev L i v i n g Dead Senescent Duff 4 5 6 7 8 10 5 6 7 8 10 4 5 6 7 8 10 5 6 7 8 10 115.7 122.2 131.5 61.8 79.2 13.5 0.0 0.0 0.0 0.0 0.0 0.0 14. 1 33.8 23.4 19.5 72. 1 20.8 5.0 42.7 37.0 11.9 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 18.8 123.3 115.2 184.3 125.0 2.1 0.0 0.0 0.0 0.0 0.0 0.0 8.6 10.0 40.3 47.5 64.5 4.9 12.2 10.0 13.0 7.8 134.5 245.5 246.7 246. 1 204.2 15.6 0.0 0.0 0.0 0.0 0.0 0.0 22.7 43.8 63. 7 67.0 136.6 25.7 17.2 52.7 50.0 19.7 43 50 64 25 ,3 2 ,5 .7 31.8 6.5 0.0 0.0 0.0 0.0 0.0 0.0 5.7 14.8 9.2 7.1 28.1 7.5 2.1 21.5 14.8 4.8 GRAMS F i g u r e 32. 2 Mean monthly dry weights (gm .5m ) of l i v i n g (.), dead (0 ) , senescent (£>) and duff (+) plant f r a c t i o n s from the Salmon Deschampsia c e s p i t o s a - Carex l y n g b y e i community. S t a t i s t i c s f o r graph P o r t i o n Month Range Observations Minimum Maximum Std. Dev, L i v i n g 4 5 6 7 8 10 6.9 36.1 24.9 131. 1 70.7 14.1 5 5 5 5 5 5 16.4 29.5 87.1 85.4 80.1 60.6 23.3 65.6 112.0 216.5 150.8 74.7 2.9 14.0 10.2 50.7 28. 8 5.5 Dead 4 5 6 7 8 10 148.8 70.6 49.6 99.8 88.5 45.5 5 5 5 5 5 5 115.9 86.0 86.8 63.3 36.8 18.3 264.7 156.6 136 163 125 63.8 59.2 28.9 20.9 43.8 37.3 16.8 Senescent 4 5 6 7 8 10 0.0 0.0 0.0 36.7 54.7 54.1 5 5 5 5 5 5 0.0 0.0 0.0 17.1 42.3 124.1 0.0 0.0 0.0 53.8 97.0 178.2 0.0 0.0 0.0 13.4 23.8 24.5 Duff 5 6 7 8 10 17.5 41.6 11.0 45.8 36.6 5 5 5 5 5 4.3 19.7 13.5 20. 1 "6.6 21.8 61.3 24.5 65.9 43.2 7.9 15.8 4.8 18.4 13.0 F i g u r e 33. Mean monthly dry weights (gm .5m ) of l i v i n g (.), dead (0 ) , senescent (^) and duff (+) plant f r a c t i o n s from the Salmon Poa p r a t e n s i s - A g r o s t i s a l b a var. s t o l o n i f e r a -P o t e n t i l l a p a c i f i c a community. S t a t i s t i c s f o r graph P o r t i o n Month Rang e Observations Minimum Maximum Std . Dev 4 10. 7 5 12. 9 23. 6 4 .5 5 45. 6 5 47. 9 93. 5 17 .6 L i v i n g 6 44. 1 5 63. 7 107. 8 17 .4 7 23. 6 5 63. 2 86. 8 9 .6 8 18. 1 5 51. 2 69. 3 7 .4 10 20. 0 5 11. 1 31. 1 7 .2 4 129. 9 5 66. 9 196. 8 49 .5 5 74. 5 5 104. 2 178. 7 29 .5 Dead 6 83. 9 5 62. 3 146. 2 33 .2 7 58. 2 5 41. 0 99. 2 22 . 1 8 23. 8 5 1. 0 24. 8 10 . 1 10 4. 3 5 0. 0 4. 3 1 .9 4 0. 0 5 0. 0 0. 0 0 .0 5 0. 0 5 0. 0 0. 0 0 .0 Senescent 6 0. 0 5 0. 0 0. 0 0 .0 7 16. 0 5 13. 2 29. 2 6 .3 8 20. 0 5 44. 1 64. 1 8 .8 10 44. 2 5 81. 3 125. 5 17 .7 5 30. 4 5 32. 2 62. 8 11 .9 6 35. 1 5 42. 7 77. 8 16 .8 Duff 7 18. 6 5 32. 4 51. 0 8 . 1 8 100. 2 5 26. 6 126. 8 37 .6 10 53. 5 5 42. 2 95. 7 22 .3 GRAMS 0.0 16.875 33.75 50.625 67.5 84.375 101.25 118.13 135.0 F i g u r e 34. Mean monthly ni t r o g e n (B) and ash free dry (8) weights (gm ,5m - 2) of combined l i v i n g and senescent f r a c t i o n s in the Cowichan Carex lyngbyei community. S t a t i s t i c s f o r the graph Month Range Observations Minimum Maximum Std. Dev, Nitr o g e n AFDW 4 0.6 5 1.7 2.3 0.3 5 4.3 5 2.4 6.7 1.9 6 1.2 5 5.0 6.2 0.5 7 2.7 5 3.7 6.4 1.1 8 0.6 5 2.2 2.8 0.2 10 3.1 5 2 .4 5.5 1.2 4 11.1 5 29.7 40.8 4.6 5 144.6 5 77.8 222.4 64.0 6 57.2 5 229.8 287.0 24.6 7 140.5 5 189.9 330.4 55.3 8 31.7 5 117.0 148.7 12.6 10 100.0 5 76.0 175.9 39.1 - 3?y -ID F i g u r e 35. Mean monthly nitrogen (B) and ash free dry (8) weights (gm .5m - 2) of the l i v i n g f r a c t i o n in the Chemainus S a l i c o r n i a v i r g i n i c a community. S t a t i s t i c s f o r the graph  Month Range Observations Minimum Maximum Std. Dev. 4 0.3 5 1.1 Ni t r o g e n 6 2.3 7 1.3 8 5.9 10 2.5 4 4.7 5 23.0 AFDW 6 47.2 7 32.5 8 137.1 10 61.8 5 0.2 0.5 0.1 5 1 .8 2 .8 0.4 5 1.7 4.0 0.9 5 3.8 5.1 0.5 5 2.0 7.9 2.2 5 1 .2 3.6 1.0 5 2.6 7.2 1.9 5 38.6 61.6 9.1 5 34.3 81.5 17.3 5 98.5 131.0 13.0 5 45.6 182.7 51.9 5 29.3 91. 1 24.4 F i g u r e 36. Mean monthly nitrogen (B) and ash free dry (8) weights (gm .5m - 2) of the l i v i n g f r a c t i o n in the Chemainus D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a community. S t a t i s t i c s f o r the graph Month Range Observations Minimum Maximum S t d . Dev. 4 1.6 5 1.8 Nitrogen 6 2.4 7 0.9 8 2.9 10 1.0 4 30.8 5 49.6 AFDW 6 60.8 7 30.9 8 119.6 10 42.4 5 0.0 1.6 0.6 5 0.7 2.5 0.8 5 1.9 4.3 1.0 5 3.0 3.9 0.4 5 1.9 4.8 1.0 5 3.2 4.2 0.4 5 0.9 31.7 11.9 5 19.3 68.9 21 .2 5 49.8 110.6 25.0 5 106.6 137.5 13.2 5 80.4 199.9 43.5 5 134.5 176.9 16.2 WEIGHT (GRAMSj 70.875 34.5 118.13 141.75 165.38 189.0 J L F i g u r e 37. Mean monthly n i t r o g e n (B) and ash free dry (8) weights (gm .5m~2) of combined l i v i n g and senescent f r a c t i o n s in the L i t t l e Qualicum P o t e n t i l l a p a c i f i c a - Carex  lyngb y e i community. S t a t i s t i c s f o r the graph Month Range Observations Minimum Maximum Std. Dev, Ni t r o g e n AFDW 4 0.5 5 0.4 0.9 0.2 5 1.7 5 1.2 2.9 0.7 6 1.1 5 2.2 3.3 0.4 7 1.5 5 1.5 2.9 0.6 8 1.2 5 1.4 2.7 0.5 10 2.6 5 2.3 4.9 1 .0 4 14.8 5 10.0 24.7 5.4 5 56.7 5 39.7 96.4 21.6 6 44.8 5 94.3 139.1 18.0 7 78.6 5 77.7 156.3 32.6 8 75.5 5 86.6 162.1 27.6 10 117.2 5 105.6 222.8 46.5 F i g u r e 38. Mean monthly ni t r o g e n (0) and ash f r e e dry (0) weights (gm .5m - 2) of combined dead and duff f r a c t i o n s i n the Cowichan Carex lyng b y e i community. S t a t i s t i c s f o r the graph Month Range Observations Minimum Maximum S t d . Dev 5 2.1 5 0.0 2.1 1.1 6 0.4 5 0.0 0.4 0.2 Nit r o g e n 7 0.2 5 0.0 0.2 0.1 8 0.3 5 0.0 0.3 0.1 5 66.5 5 0.0 66.5 30.9 AFDW 6 7.2 5 0.0 7.2 3.8 7 4.9 5 0.0 4.9 2.3 8 5.3 5 0.0 5.3 2.5 CO 00 WEIGHT (GRAMSJ 0.0 5.625 11.25 16.815 22.5 2A.125 33.15 39.315 45.0 *_| 1 1 1 I I L t 1 F i g u r e 39. Mean monthly nitrogen (0) and ash free dry (0 ) weights (gm .5m - 2) of combined dead and duff f r a c t i o n s i n the Chemainus S a l i c o r n i a v i r g i n i c a community. S t a t i s t i c s f o r the graph Month Range Observations Minimum Maximum Std. Dev. Nitrogen AFDW 4 2.7 5 6.2 8.8 1.1 5 4.1 5 5.4 9.5 1 .7 6 3.6 5 7.5 11.1 1.5 7 3.5 5 5.8 9.3 1.3 8 4.3 5 6.7 11.0 2.0 10 4.2 5 3.4 7.6 1.8 4 69.8 5 162.0 231.9 29.4 5 102.3 5 127.9 230.2 41.5 6 73.3 5 150.9 224.3 31.4 7 81.8 5 135.4 J 217.2 30.5 8 109.8 5 172.7 282.5 52.3 10 98.1 5 78.4 176.5 42.0 0.0 46.125 _J 32.25 WEIGHT (GRAMS) 138.38 184.5 230.63 ± 276.75 322.88 369.0 _J I 1 U 1 -2 co-co 00 CD F i g u r e 40. Mean monthly nitrogen (0) and ash f r e e dry (0) weights (gm .5m - 2) of combined dead and duff f r a c t i o n s i n the Chemainus D i s t i c h l i s s p i c a t a - G r i n d e l i a i n t e g r i f o l i a community. S t a t i s t i c s f o r the graph Month Range Observations Minimum Maximum Std. Dev. Nit r o g e n AFDW 4 3.5 5 4.9 8.4 1.5 5 3.3 5 5.6 9.0 1.2 6 1.7 5 4.8 6.4 0.7 7 1.5 5 4.3 5.8 0.7 8 2.7 5 3.8 6.5 1.0 10 3.3 5 4.5 7.7 1.3 4 107.2 5 150.2 257.4 47.0 5 93.4 5 154. 1 247.5 35.3 6 60.4 5 147.3 207.6 25.2 7 52.6 5 147.0 199.5 22.9 8 73.9 5 106.6 180.5 26.8 10 78.8 5 122.9 201.7 31.9 F i g u r e 41. Mean monthly n i t r o g e n (0) and ash free dry (0) weights (gm .5m - 2) of combined dead and duff f r a c t i o n s i n the L i t t l e Qualicum P o t e n t i l l a p a c i f i c a - Carex l y n g b y e i community. S t a t i s t i c s f o r the graph MOnth Range Observations Minimum Maximum Std. Dev. Nitrogen AFDW 4 2.3 5 1.0 3.3 1.1 5 2.8 5 1 .4 4.2 1.2 6 3.4 5 0.7 4.2 1.5 7 3.9 5 0.0 3.9 1.8 8 4.6 5 0.9 5.5 1.9 10 0.6 5 0.3 0.9 0.3 4 139.2 5 60.2 199.4 67.0 5 96.5 5 46.2 142.7 40.8 6 146.0 5 19.7 165.7 62.6 7 166.3 5 0.5 166.7 75.7 8 116.8 5 23.1 139.9 47.6 10 13.2 5 7.2 20.4 6.0 WEIGHT (GRAMSJ - 394 -Appendix 7. Graphs of the number of days root cores from the p l a n t communities s t u d i e d grew i n darkness. F i g u r e s 44-52, the months i n the f i g u r e s are A p r i l ( 4 ) , May ( 5 ) , June ( 6 ) , J u l y ( 7 ) , August (8) and October (10). - 395 -Figu r e 44. Exhaustion of root r e s e r v e s i n darkness f o r the Cowichan Carex l y n g b y e i community: average time (.), h e i g h t (x) of regrown shoots and number of o v e r w i n t e r i n g (•) and regrown (o) shoots. Note, f o r o v e r w i n t e r i n g and regrown shoots, p l o t s sampled i n October had not been c l i p p e d before. Regrown shoots f o r October are the number of shoots i n a p r e v i o u s l y u n d i p p e d .5m2 p l o t i n the sedge stand. - 397 -F i g u r e 4 5 . Exhaustion of root r e s e r v e s i n darkness f o r the Cowichan Juncus b a l t i c u s community: average time (.) and height (x) of regrown ' stems. - 398 -300 4 5 6 7 8 MONTH 10 - 399 -Fig u r e 46. Exhaustion of root r e s e r v e s i n darkness f o r the Chemainus S a l i c o r n i a v i r g i n i c a (S) and D i s t i c h l i s  s p i c a t a - G r i n d e l i a i n t e g r i f o l i a (D) communities: average time ( d a y s ) . - 401 -Fi g u r e 47. Exhaustion of root r e s e r v e s i n darkness f o r the L i t t l e Qualicum Carex l y n g b y e i community: average time (.), height of regrown shoots ( x ) , and number of o v e r w i n t e r i n g (O) and regrown (o) shoots. Note, f o r o v e r w i n t e r i n g and regrown shoots, p l o t s sampled in October had not been c l i p p e d b e f o r e . Regrown shoots f o r October are the number of shoots in a p r e v i o u s l y u n d i p p e d ,5m2 p l o t i n the sedge stand. - 403 F i g u r e 48. Exhaustion of root r e s e r v e s i n darkness f o r the L i t t l e Qualicum P o t e n t i l l a p a c i f i c a - Carex  l y n g b y e i community: average time (.), and average height (x) of regrown shoots of Lyngby's sedge (C) and B a l t i c rush ( J ) . - 404 -4 5 6 7 8 10 MONTH - 405 -Figure 49. Exhaustion of root reserves i n darkness f o r the Campbell Carex lyngbyei community: average time ( x ) , height of regrown shoots ( x ) , and number of overwintering (•) and regrown (o) shoots. Note, f o r overv/intering and regrown shoots, p l o t s sampled i n October had not been c l i p p e d before. Regrown shoots f o r October are the number of shoots i n a p r e v i o u s l y u n d i p p e d .5m2 p l o t i n the sedge stand. - 407 -F i g u r e 50. Exhaustion of root r e s e r v e s i n darkness f o r the Campbell P o t e n t i l l a p a c i f i c a - E l e o c h a r i s  p a l u s t r i s community: average time (.) and h e i g h t (x) of regrown stems. - 4 0 8 -300 4 5 6 7 8 MONTH I 10 - 4 0 9 -Figure 51. Exhaustion of root reserves i n darkness f o r the Salmon Carex lyngbyei community: average time (.), height of regrown shoots ( x ) , and number of ove r w i n t e r i n g (o) and regrown (o) shoots. Note, f o r o v e r w i n t e r i n g and regrown stems, p l o t s sampled i n October had not been c l i p p e d before. Regrown stems f o r October are the number of stems i n a p r e v i o u s l y u n d i p p e d ,5m2 p l o t i n the sedge stand. - 411 -F i g u r e 52. Exhaustion of root r e s e r v e s i n darkness f o r the Salmon Deschampsia c e s p i t o s a - Carex l y n g b y e i (D) and Poa p r a t e n s i s - A g r o s t i s alba v a r . s t o l o n i f e r a - P o t e n t i l l a p a c i f i c a (P) communities: average time (.). , - 412 -300 4 5 6 7 8 10 MONTH 

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