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Vegetation-environment relationships and plant community classification and ordination in British Columbia… Campbell, Alison 1986

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VEGETATION - ENVIRONMENT RELATIONSHIPS AND PLANT COMMUNITY CLASSIFICATION AND ORDINATION IN BRITISH COLUMBIA COASTAL SALT MARSHES by ALISON CAMPBELL B . S c . ( H o n s . ) , U n i v e r s i t y Of V i c t o r i a , 1 9 8 2 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES Botany Department We a c c e p t t h i s t h e s i s as c o n f o r m i n g t o the r e q u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH COLUMBIA October 1986 © A l i s o n C a m p b e l l , 1986 In 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 of the r e q u i r e m e n t s f o r an advanced degree a t the U n i v e r s i t y of B r i t i s h C olumbia, I agree t h a t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r agree 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 of t h i s t h e s i s f o r s c h o l a r l y purposes may be g r a n t e d by the Head of my Department or by h i s or her 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 or p u b l i c a t i o n of 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 not be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Department of Botany The U n i v e r s i t y of B r i t i s h Columbia 2075 Wesbrook P l a c e Vancouver, Canada V6T 1W5 Date: October 15, 1986 i i ^ A b s t r a c t The p l a n t communities of t h i r t e e n s a l t marshes a l o n g the c o a s t of B r i t i s h Columbia a r e d e s c r i b e d and mapped, and a l i t e r a t u r e r e v i e w on s a l t marsh e c o l o g i c a l s t u d i e s a l o n g the P a c i f i c c o a s t of N o r t h America i s p r e s e n t e d . C o a s t a l marshes i n B.C. were compared based on p l a n t s p e c i e s c o m p o s i t i o n . The p l a n t communities of the n o r t h e r n B.C. marshes i n t h i s study were s i m i l a r t o s o u t h e r n B.C. marshes ( e x c l u d i n g the F r a s e r R i v e r marshes) except t h a t they l a c k e d D i s t i c h l i s s p i c a t a and G r i n d e l i a i n t e g r i f o l i a as dominant s p e c i e s . The p l a n t communities on the Queen C h a r l o t t e I s l a n d s and n o r t h e r n m a i n l a n d d i f f e r e d s l i g h t l y from each o t h e r and from those on n o r t h e r n Vancouver I s l a n d . M u l t i v a r i a t e methods of a n a l y s i s ( p r i n c i p a l components a n a l y s i s , c a n o n i c a l c o r r e l a t i o n a n a l y s i s and m u l t i v a r i a t e a n a l y s i s of v a r i a n c e ) a p p l i e d t o the f i e l d d a t a i n d i c a t e d t h a t v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s i n the marshes ranged from s t r o n g t o weak. A l s o , a l t h o u g h e l e v a t i o n was i m p o r t a n t , f a c t o r s o t h e r than e l e v a t i o n were m a i n l y r e l a t e d t o p l a n t community p a t t e r n s i n the marshes. Most s p e c i e s were l o c a t e d t h r o u g h o u t the marshes (e. g . Carex l y n g b y e i , Deschampsia c e s p i t o s a ,  P o t e n t i l l a a n s e r i n a and T r i g l o c h i n maritimum ) w h i l e o t h e r s (e.g. Elymus m o l l i s ) showed r e s t r i c t e d d i s t r i b u t i o n s w i t h r e s p e c t t o e l e v a t i o n . U s i n g m u l t i v a r i a t e a n a l y s i s of v a r i a n c e , the p l a n t community c l a s s i f i c a t i o n s d e r i v e d s u b j e c t i v e l y i n the f i e l d f o r management purposes (based on a e r i a l photographs and p r i n c i p a l components a n a l y s i s ) were found t o e x p l a i n most ( > 84% ) of the v a r i a t i o n i n the v e g e t a t i o n d a t a . A comparison between the c l a s s i f i c a t i o n d e v e l o p e d i n t h i s s tudy and t h a t used by the B r i t i s h Columbia M i n i s t r y of Environment suggested t h a t the M i n i s t r y of Environment c l a s s i f i c a t i o n c o u l d be improved by f u r t h e r s u b d i v i d i n g on the b a s i s of s a l i n i t y . At the Yakoun and D a l a marshes, v a r i o u s o r d i n a t i o n methods ( p r i n c i p a l components a n a l y s i s , p r i n c i p a l c o o r d i n a t e s a n a l y s i s , r e c i p r o c a l a v e r a g i n g and d e t r e n d e d c o r r e s p o n d e n c e a n a l y s i s ) as w e l l as c l u s t e r a n a l y s i s were compared f o r t h e i r a b i l i t y t o i d e n t i f y the p l a n t communities and t o summarize v a r i a t i o n i n the d a t a . Detrended correspondence a n a l y s i s was the o n l y method t h a t d i s t i n g u i s h e d a l l the p l a n t communities r e c o g n i z e d i n the f i e l d . However, the f i r s t t h r e e axes o n l y e x p l a i n e d a s m a l l amount of the v a r i a t i o n i n the v e g e t a t i o n d a t a . P r i n c i p a l components a n a l y s i s u s i n g q u a n t i t a t i v e d a t a and a c o v a r i a n c e m a t r i x , and p r i n c i p a l c o o r d i n a t e s a n a l y s i s u s i n g q u a n t i t a t i v e d a t a a ccounted f o r t w i c e as much v a r i a t i o n i n the d a t a and i d e n t i f i e d t h r e e out of f o u r communities. A d e t a i l e d study of s o i l s - v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s a t the Yakoun and D a l a marshes showed t h a t weak r e l a t i o n s h i p s e x i s t e d between e l e v a t i o n and s o i l s v a r i a b l e s but a l l s o i l s v a r i a b l e s were i m p o r t a n t i n a c c o u n t i n g f o r the v a r i a t i o n i n the v e g e t a t i o n d a t a . i v T a b l e of C o n t e n t s A b s t r a c t i i L i s t of T a b l e s v i L i s t of F i g u r e s v i i i Acknowledgement i x Chapter I INTRODUCTION 1 Chapter I I LITERATURE REVIEW 7 2.1 STUDIES ON THE ECOLOGY OF PACIFIC COASTAL MARSHES ..7 2.1.1 BRITISH COLUMBIA 7 2.1.2 WASHINGTON 12 2.1.3 OREGON 13 2.1.4 CALIFORNIA 15 2.1.5 ALASKA 16 2.2 ZONATION 18 2.3 ENVIRONMENTAL FACTORS THAT INFLUENCE PLANT ZONATION 20 2.3.1 ELEVATION 20 2.3.2 TIDES 25 2.3.3 NUTRIENTS 26 2.3.4 SALINITY 31 2.3.5 ORGANIC MATTER 33 2.3.6 SOIL pH 34 2.3.7 SOIL TEXTURE 35 2.3.8 WATERLOGGING 36 2.3.9 COMPETITION 37 2.3.10 OTHER ENVIRONMENTAL FACTORS 38 Chapter I I I STUDY AREAS 40 3.1 DESCRIPTION OF STUDY AREAS 43 3.1.1 CLIMATE 43 3.1.2 SALINITY 45 3.1.3 OTHER ENVIRONMENTAL INFORMATION 46 3.1.4 DISTURBANCE 47 Chapter IV SAMPLING METHODS AND DATA COLLECTION 52 4.1 VEGETATION - ELEVATION SURVEYS 52 4.2 MAPPING OF PLANT COMMUNITIES 53 4.3 SOIL SURVEYS AND ANALYSIS 54 Chapter V DATA ANALYSIS 55 5.1 PLANT SPECIES COMPOSITION 55 5.2 VEGETATION - ELEVATION RELATIONSHIPS 55 5.3 COMPARISON OF PLANT COMMUNITY CLASSIFICATIONS 58 5.4 COMPARISON OF ORDINATION METHODS AND CLUSTER ANALYSIS 59 V 5.4.1 CLUSTER ANALYSIS 60 5.4.2 PRINCIPAL COMPONENTS ANALYSIS 61 5.4.3 PRINCIPAL COORDINATES ANALYSIS 62 5.4.4 RECIPROCAL AVERAGING 63 5.4.5 DETRENDED CORRESPONDENCE ANALYSIS 64 5.5 SOILS - VEGETATION - ELEVATION RELATIONSHIPS 64 Chapter VI RESULTS AND DISCUSSION 66 6.1 PLANT SPECIES COMPOSITION 66 6.2 VEGETATION - ELEVATION RELATIONSHIPS 68 6.3 COMPARISON OF PLANT COMMUNITY CLASSIFICATIONS 76 6.4 COMPARISON OF ORDINATION METHODS AND CLUSTER ANALYSIS 78 6.5 SOILS - VEGETATION - ELEVATION RELATIONSHIPS 99 Chapter V I I CONCLUSIONS ,109 LITERATURE CITED 112 APPENDIX A - RATING OF STUDY AREAS ACCORDING TO THE BRITISH COLUMBIA MINISTRY OF ENVIRONMENT 130 APPENDIX B - FREQUENCY OF PLANT SPECIES IN EACH MARSH, CALCULATED AS THE PERCENT OF QUADRATS OCCUPIED 133 APPENDIX C - CORRELATIONS OF PLANT SPECIES WITH PCA AXES I - I l l OF THE VEGETATION DATA 137 APPENDIX D - VEGETATION MAPS FOR STUDY AREAS 153 APPENDIX E - GRAPHS OF ELEVATION VERSUS PLANT SPECIES PERCENT COVER FOR EACH MARSH 179 v i L i s t of T a b l e s I . Review of l i t e r a t u r e on s a l t marsh v e g e t a t i o n -environment r e l a t i o n s h i p s 21 I I . Review of l i t e r a t u r e on c o r r e l a t i o n s between e l e v a t i o n and e n v i r o n m e n t a l f a c t o r s 23 I I I . C l i m a t e of study a r e a s 44 IV. E n v i r o n m e n t a l d a t a f o r study a r e a s 48 V. V e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s : R-square v a l u e s from l i n e a r r e g r e s s i o n s of PCA axes I - I I I on e l e v a t i o n . A l s o g i v e n a r e c a n o n i c a l c o r r e l a t i o n c o e f f i c i e n t s (Rc) r e l a t i n g t o t a l v e g e t a t i o n v a r i a t i o n (PCA I - I I I ) t o e l e v a t i o n 69 V I . V e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s : P r o p o r t i o n s of v e g e t a t i o n v a r i a t i o n e x p l a i n e d by the community c l a s s i f i c a t i o n s and the p e r c e n t v a r i a n c e e x p l a i n e d by PCA I - I I I f o r the t o t a l , p r e d i c t e d and r e s i d u a l components of the d a t a 74 V I I . Comparison of p l a n t community c l a s s i f i c a t i o n s : P r o p o r t i o n s of v e g e t a t i o n v a r i a t i o n e x p l a i n e d by the B r i t i s h Columbia M i n i s t r y of Environment c l a s s i f i c a t i o n and a more d e t a i l e d c l a s s i f i c a t i o n ..77 V I I I . Comparison of o r d i n a t i o n methods f o r the Yakoun R i v e r e s t u a r y : P e r c e n t v a r i a n c e e x p l a i n e d by the f i r s t t h r e e axes and p r o p o r t i o n s of v e g e t a t i o n v a r i a t i o n e x p l a i n e d by the community c l a s s i f i c a t i o n s 91 IX. Comparison of o r d i n a t i o n methods f o r the D a l a R i v e r e s t u a r y : P e r c e n t v a r i a n c e e x p l a i n e d by the f i r s t t h r e e axes and p r o p o r t i o n s of v e g e t a t i o n v a r i a t i o n e x p l a i n e d by the community c l a s s i f i c a t i o n s 92 X. C o r r e l a t i o n s of sample s c o r e s f o r axes I - I I I from o r d i n a t i o n s of the v e g e t a t i o n d a t a , f o r the Yakoun R i v e r e s t u a r y 94 X I . C o r r e l a t i o n s of sample s c o r e s f o r axes I - I I I from o r d i n a t i o n s of the v e g e t a t i o n d a t a , f o r the D a l a R i v e r e s t u a r y 95 X I I . C o r r e l a t i o n m a t r i x of s o i l s v a r i a b l e s , e l e v a t i o n and the f i r s t t h r e e PCA axes of a s o i l s and a v e g e t a t i o n o r d i n a t i o n , f o r the Yakoun R i v e r e s t u a r y 100 v i i X I I I . C o r r e l a t i o n m a t r i x of s o i l s v a r i a b l e s , e l e v a t i o n and the f i r s t t h r e e PCA axes of a s o i l s and a v e g e t a t i o n o r d i n a t i o n f o r the D a l a R i v e r e s t u a r y 102 XIV. Spearman rank c o r r e l a t i o n s between PCA axes I - I I I of the s o i l s and v e g e t a t i o n d a t a s e t s f o r the Yakoun R i v e r e s t u a r y 108 XV. Spearman rank c o r r e l a t i o n s between PCA axes I - I I I of the s o i l s and v e g e t a t i o n d a t a s e t s f o r the D a l a R i v e r e s t u a r y 108 v i i i L i s t of F i g u r e s 1. Map of study a r e a s 41 2. Graphs of c a n o n i c a l v a r i a t e a x i s I ( v e r t i c a l a x i s ) v e r s u s e l e v a t i o n i n metres above c h a r t datum ( h o r i z o n t a l a x i s ) 70 3. C l u s t e r a n a l y s i s of the Yakoun R i v e r e s t u a r y u s i n g the Minimum v a r i a n c e method and E u c l i d e a n d i s t a n c e 79 4. O r d i n a t i o n a n a l y s e s f o r the Yakoun R i v e r E s t u a r y 80 5. C l u s t e r a n a l y s i s of the D a l a R i v e r e s t u a r y u s i n g the Minimum v a r i a n c e method and E u c l i d e a n d i s t a n c e 85 6. O r d i n a t i o n a n a l y s e s f o r the D a l a R i v e r e s t u a r y 86 7. P r i n c i p a l components a n a l y s i s of s o i l s v a r i a b l e s f o r the Yakoun R i v e r e s t u a r y 106 8. P r i n c i p a l components a n a l y s i s of s o i l s v a r i a b l e s f o r the D a l a R i v e r e s t u a r y 107 i x Acknowledgement The f o l l o w i n g i n d i v i d u a l s and o r g a n i z a t i o n s have h e l p e d make t h i s t h e s i s p o s s i b l e , and I am ve r y g r a t e f u l t o a l l of them. Dr. Gary E. B r a d f i e l d , my t h e s i s s u p e r v i s o r , gave i n s t r u c t i o n i n da t a a n a l y s i s , a d v i c e and support t h r o u g h o u t . D r s . J.R. Maze and R. T u r k i n g t o n and N e i l Dawe of the Canadian W i l d l i f e S e r v i c e , P a c i f i c and Yukon R e g i o n , p r o v i d e d h e l p f u l c r i t i c i s m of the m a n u s c r i p t . The B r i t i s h Columbia M i n i s t r y of Environment and Parks ( e s p e c i a l l y Rodger Hunter) p r o v i d e d f i e l d equipment, a n a l y z e d s o i l s d a t a and f i r s t i n t r o d u c e d me t o the f a s c i n a t i n g w o r l d of s a l t marshes. The Canadian W i l d l i f e S e r v i c e , P a c i f i c and Yukon Re g i o n , p r o v i d e d v a l u a b l e f i e l d equipment as w e l l as f i n a n c i a l s u p p o r t . J i m Cryder was my h i g h l y competent f i e l d a s s i s t a n t . D r s . A. Ceska and R.T. O g i l v i e of t h e B r i t i s h Columbia P r o v i n c i a l Museum h e l p e d i n the i d e n t i f i c a t i o n of p l a n t s p e c i e s . Mr. F. Stephenson, F i s h e r i e s and Oceans, Canada, a s s i s t e d w i t h the c o n v e r s i o n of the l e v e l i n g survey t o e l e v a t i o n above c h a r t datum. Mrs. H e l e n O ' B r i e n , Mr. and Mrs. Cann, Mr. and Mrs. Bunn and Mr. B i l l Shephard were o u t s t a n d i n g h o s t s d u r i n g the work i n the f i e l d . Mr. J . M c M i l l a n p r o v i d e d t r a n s p o r t a t i o n t o the Ououkinsh e s t u a r y . My f a m i l y and Andrew p r o v i d e d s u p p o r t and encouragement t h r o u g h o u t . X T h i s study was s u p p o r t e d by g r a n t s from the N a t u r a l S c i e n c e s and E n g i n e e r i n g Research C o u n c i l of Canada. 1 I . INTRODUCTION S a l t marshes a r e d e f i n e d as a r e a s of l a n d b o r d e r i n g the se a , more or l e s s c o v e r e d w i t h v e g e t a t i o n and s u b j e c t t o p e r i o d i c i n u n d a t i o n by the t i d e (Chapman 1974). S a l t marshes commonly bor d e r on e s t u a r i e s d e f i n e d by P r i t c h a r d (1967) as s e m i - e n c l o s e d c o a s t a l b o d i e s of water which have a f r e e c o n n e c t i o n w i t h the open ocean and w i t h i n which sea water i s measurably d i l u t e d w i t h f r e s h water d e r i v e d from l a n d d r a i n a g e . S a l t marshes are found on most c o a s t s i n a r e a s s u p p l i e d w i t h r i v e r or marine sediments and s h e l t e r e d from d i r e c t a t t a c k by h i g h energy waves (Chapman 1974). They a r e worldwide i n d i s t r i b u t i o n a l t h o u g h a t t r o p i c a l l a t i t u d e s s a l t marshes t e n d t o be r e p l a c e d by mangrove swamps. There i s a s i m i l a r i t y i n appearance of s a l t marsh v e g e t a t i o n worldwide (most s p e c i e s b e l o n g t o a few c o s m o p o l i t a n genera) due t o s i m i l a r e n v i r o n m e n t a l f a c t o r s o p e r a t i n g i n t h e s e a r e a s ( R a n w e l l 1972; Long & Mason 1983). Chapman (1974) has d i v i d e d w o r l d s a l t marsh v e g e t a t i o n i n t o n i n e groups based on s u b s t r a t u m , t i d a l range and c l i m a t e . B r i t i s h Columbia f a l l s i n t o the West Coast of N o r t h America group which i s then d i v i d e d i n t o s u b a r c t i c , temperate and dry C a l i f o r n i a n subgroups. F r e y & Bason (1985) s e p a r a t e d the P a c i f i c c o a s t marshes i n t o a C e n t r a l P a c i f i c zone t h a t extends up t o Vancouver I s l a n d and a Western A l a s k a - S u b a r c t i c zone above Vancouver I s l a n d . Those zones a r e based on f l o r a , 2 s e d i m e n t s , o r g a n i s m s , topography, t i d a l range, wave and c u r r e n t energy and the t e c t o n i c and e u s t a t i c s t a b i l i t y of the c o a s t a l a r e a . A l o n g the P a c i f i c c o a s t , s a l t marshes a r e l i m i t e d t o p r o t e c t e d i n l e t s and the heads of f j o r d s . Only 10-20% of the west c o a s t of N o r t h America i s s u i t a b l e f o r s a l t marsh development compared t o 80-90% of the e a s t c o a s t . That i s due t o the a c t i v e t e c t o n i c n a t u r e (mountain b u i l d i n g and c o a s t a l u p l i f t ) of the west c o a s t which i s c h a r a c t e r i s e d by deep f j o r d s i n B r i t i s h Columbia and Southern A l a s k a and r e l a t i v e l y few bays and s m a l l l agoons i n C a l i f o r n i a , Oregon and Washington (Glooschenko 1980; F r e y & Bason 1985). In c o n t r a s t , the A t l a n t i c c o a s t a l p l a i n marshes a r e e s s e n t i a l l y c o n t i n u o u s on the low energy, submerging, c o a s t l i n e ( F r e y S. Bason 1985). S a l t marshes a r e e x t r e m e l y i m p o r t a n t a r e a s f o r f i s h and w a t e r f o w l and a r e a l s o u t i l i z e d h e a v i l y by man. Of the 10.8 m i l l i o n w a t e r f o w l of the P a c i f i c f l y w a y , 5.4 m i l l i o n use c o a s t a l r o u t e s . An e s t i m a t e d one m i l l i o n ducks and geese w i n t e r a l o n g B r i t i s h Columbia's c o a s t , and e s t u a r i n e w e t l a n d s a l o n g w i t h nearby f a r m l a n d a r e the p r i n c i p a l a r e a s u t i l i z e d by th e s e b i r d s f o r f e e d i n g and r e s t i n g (Burgess 1971; Hunter & Jones 1982). Ducks use t h e s e two a r e a s as a l t e r n a t e h a b i t a t s . Farmland i s p r e f e r r e d d u r i n g warm wet weather but e s t u a r i e s a r e i m p o r t a n t when f i e l d s a r e f r o z e n or d r y . A l l p a r t s of the marsh a r e i m p o r t a n t f o r f e e d i n g and l o a f i n g (Earner 1985). S a l t marshes and t h e i r a s s o c i a t e d e s t u a r i e s a r e a l s o i m p o r t a n t f o r salmon. A l l P a c i f i c salmon pass t h r o u g h an 3 e s t u a r y t w i c e i n t h e i r l i v e s ; f i r s t as j u v e n i l e s m i g r a t i n g from f r e s h water n u r s e r y a r e a s t o the ocean and s e c o n d l y as m a t u r i n g a d u l t s r e t u r n i n g from the ocean t o f r e s h water t o spawn. D i f f e r e n t s p e c i e s of salmon u t i l i z e d i f f e r e n t p a r t s of the s a l t marsh a t d i f f e r e n t t i m e s of the y e a r , f o r p h y s i o l o g i c a l t r a n s i t i o n from f r e s h t o s a l t w a t e r , as r e f u g i a from p r e d a t o r s , and as p r o d u c t i v e f o r a g i n g a r e a s (Dunford 1975; Dorcey e t a l . 1978; Levy & N o r t h c o t e 1982; Healey 1982; Simenstad e t a l . 1982). C o a s t a l w e tlands a l s o p r o v i d e f o r a g i n g a r e a s f o r mammals such as d e e r , e l k and bear (Fox & Nowlan 1978). S a l t marshes have one of the h i g h e s t a n n u a l net p r i m a r y p r o d u c t i v i t i e s (3000 g m~2) of a l l major ecosystems (Simmons 1981) . The s a l t marsh v e g e t a t i o n p r o v i d e s a c r u c i a l l i n k i n the e s t u a r i n e f o o d c h a i n by f o r m i n g d e t r i t u s which then s u p p o r t s i n v e r t e b r a t e s , f i s h and u l t i m a t e l y v e r t e b r a t e s . T i d a l marshes make up o n l y 2.3% of B r i t i s h Columbia's 27000 km c o a s t l i n e and many have been a f f e c t e d by d y k i n g , l o g s t o r a g e and h a n d l i n g , l a n d f i l l , d r e d g i n g and p o l l u t i o n (Hunter & Jones 1982) . E i g h t y p e r c e n t of B r i t i s h Columbia's p o p u l a t i o n l i v e s l e s s than 80 km from the c o a s t and e s t i m a t e s suggest t h a t two-t h i r d s of the e s t u a r i n e w e t l a n d s on the s o u t h e a s t c o a s t of Vancouver I s l a n d and the lower m a i n l a n d have been a f f e c t e d by man's a c t i v i t i e s . P a r t of the a t t r a c t i v e n e s s of e s t u a r i n e l o c a t i o n s f o r human s e t t l e m e n t l i e s i n t h e i r l e v e l b a c k s h o r e s , s h e l t e r e d h a r b o u r s , ample s u p p l i e s of f r e s h and t i d a l w ater, abundant f e r t i l e s o i l s and source of sand, g r a v e l and m i n e r a l s 4 (Fox & Nowlan 1978; Hunter & Jones 1 982) Dy k i n g has t r a d i t i o n a l l y been the major impact on s a l t marshes. On the F r a s e r R i v e r d e l t a , 70% (220ha) of former s a l t marsh and 30% (6260ha) of former t i d a l f r e s h water marsh and n e a r l y a l l (80ha) of the o t h e r f l o o d e d h a b i t a t has been l o s t by d y k i n g (Fox and Nowlan 1978). In C a l i f o r n i a , i t i s now d i f f i c u l t t o d i s t i n g u i s h n a t u r a l from u n n a t u r a l f e a t u r e s i n marshes because of t h e p e r v a s i v e e f f e c t s of d y k i n g ( Z e d l e r 1982). In Oregon, 80% of the s a l t marsh v e g e t a t i o n has been l o s t , m a i n l y by c o n v e r s i o n t o p a s t u r e ( F r e n k e l e t a l . 1981), w h i l e i n the U n i t e d S t a t e s as a whole, 35% of the f r e s h and s a l i n e w e t l a n d s have been l o s t . I n the N a t i o n a l E s t u a r y Study (U.S. F i s h and W i l d l i f e S e r v i c e 1970), the degree of m o d i f i c a t i o n of e s t u a r i e s i n the U n i t e d S t a t e s was c h a r a c t e r i z e d as f o l l o w s : s l i g h t m o d i f i c a t i o n - 27%, moderate m o d i f i c a t i o n 50%, s e v e r e m o d i f i c a t i o n - 23%. T h i s d e s t r u c t i v e t r e n d i s now b e i n g r e v e r s e d . I n some a r e a s such as the K o k i s h , Cowichan and Campbell R i v e r s on Vancouver I s l a n d and the F r a s e r R i v e r d e l t a , e f f o r t s a r e b e i n g made t o improve the h a b i t a t f o r f i s h and w a t e r f o w l and t o r e v e g e t a t e a r e a s a l t e r e d by man's a c t i v i t i e s (examples f o r B r i t i s h Columbia a r e documented by L e v i n g s 1980; Pomeroy e_t a l . 1981; Brownlee e t a l . 1984). Thus, because of t h e i r s c a r c i t y , importance t o f i s h and w a t e r f o w l , h i g h p r o d u c t i v i t y and the imminence of development, an i n c r e a s e d knowledge and u n d e r s t a n d i n g of s a l t marshes i s of g r e a t i m p o r t a n c e . That i s e s p e c i a l l y t r u e i n B r i t i s h C olumbia, 5 where l i t t l e p u b l i s h e d i n f o r m a t i o n i s a v a i l a b l e f o r s a l t m a r s h e s o t h e r t h a n t h o s e on t h e l o w e r m a i n l a n d a n d on t h e s o u t h 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 i s s t u d y , v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s , a n d p l a n t c o m m u n i t i e s i n s a l t m a r s h e s o f n o r t h e r n V a n c o u v e r I s l a n d , t h e n o r t h e r n m a i n l a n d o f B r i t i s h C o l u m b i a and t h e Queen C h a r l o t t e I s l a n d s were i n v e s t i g a t e d . 1. The p l a n t c o m m u n i t i e s o f t h i r t e e n a r e a s were d e s c r i b e d and mapped b e c a u s e most o f t h e m a r s h e s had n o t b e e n mapped b e f o r e , a n d maps p r o v i d e a v a l u a b l e t o o l f o r management o f t h e a r e a s . D i f f e r e n c e s b e t w e e n m a r s h e s b a s e d on p l a n t s p e c i e s c o m p o s i t i o n a n d p l a n t c o m m u n i t i e s were n o t e d . 2 . I n o r d e r t o i n v e s t i g a t e e n v i r o n m e n t a l f a c t o r s t h a t c o n t r o l p l a n t s p e c i e s d i s t r i b u t i o n s i n a m a r s h , t h e r e l a t i o n s h i p b e t w e e n v e g e t a t i o n a n d e l e v a t i o n was f o c u s s e d on b e c a u s e most p r e v i o u s s t u d i e s h a v e c o n s i d e r e d e l e v a t i o n t o be t h e most i m p o r t a n t f a c t o r c o n t r o l l 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 s o , e l e v a t i o n i s a g ood summary v a r i a b l e a s i t h a s been c o r r e l a t e d w i t h a number o f o t h e r e n v i r o n m e n t a l v a r i a b l e s ( e . g . s a l i n i t y , o r g a n i c m a t t e r , n u t r i e n t s a n d s o i l t e x t u r e ) a n d e l e v a t i o n i s one o f t h e c r i t e r i a u s e d f o r t h e B r i t i s h C o l u m b i a M i n i s t r y o f E n v i r o n m e n t c l a s s i f i c a t i o n o f e s t u a r i e s . V e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s were s t u d i e d u s i n g m u l t i v a r i a t e m e t h o d s o f a n a l y s i s ( p r i n c i p a l c o m p o n e n t s a n a l y s i s , c a n o n i c a l c o r r e l a t i o n a n a l y s i s a n d m u l t i v a r i a t e a n a l y s i s o f v a r i a n c e ) t o d e t e r m i n e t h e i m p o r t a n c e o f e l e v a t i o n i n a c c o u n t i n g f o r t h e v a r i a t i o n i n t h e v e g e t a t i o n d a t a and i n d i s t i n g u i s h i n g t h e p l a n t c o m m u n i t i e s . 6 3 . The s o i l s - v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s of two marshes were i n v e s t i g a t e d t o d e t e r m i n e which s o i l s v a r i a b l e s were c o r r e l a t e d t o e l e v a t i o n and which s o i l s f a c t o r s were i m p o r t a n t i n d e t e r m i n 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 s . 4 . A more d e t a i l e d p l a n t community c l a s s i f i c a t i o n , d e v e l o p e d i n t h i s s t u d y , was compared t o the B r i t i s h Columbia M i n i s t r y of Environment c l a s s i f i c a t i o n . T h i s was done because the M i n i s t r y of Environment c l a s s i f i c a t i o n o n l y r e c o g n i z e s t h r e e zones i n a marsh and t h i s may be o v e r s i m p l i f i e d . 5 . Most p r e v i o u s s t u d i e s have used u n i v a r i a t e methods of a n a l y s i s t o i n v e s t i g a t e p l a n t communities and v e g e t a t i o n environment r e l a t i o n s h i p s . That r e s t r i c t s the i n v e s t i g a t o r t o an a n a l y s i s of s i n g l e s p e c i e s or p l a n t c ommunities. In t h i s s t u d y , m u l t i v a r i a t e methods of a n a l y s i s were used: t h i s a l l o w s i n v e s t i g a t i o n of the v e g e t a t i o n as a whole ( a l l s p e c i e s • t o g e t h e r ) . D i f f e r e n t methods of o r d i n a t i o n ( p r i n c i p a l components a n a l y s i s , p r i n c i p a l c o o r d i n a t e s a n a l y s i s , r e c i p r o c a l a v e r a g i n g and d e t r e n d e d c o r r e s p o n d e n c e a n a l y s i s ) as w e l l as c l u s t e r a n a l y s i s were compared f o r t h e i r a b i l i t y t o d i s t i n g u i s h the p l a n t communities r e c o g n i z e d i n the f i e l d and t o e x p l a i n v a r i a t i o n i n the v e g e t a t i o n d a t a . 6. A l i t e r a t u r e r e v i e w on c o a s t a l marsh s t u d i e s a l o n g the P a c i f i c c o a s t of N o r t h America and e n v i r o n m e n t a l f a c t o r s t h a t 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 s i n these marshes i s p r e s e n t e d . The r e s u l t s of t h i s r e s e a r c h w i l l g i v e managers a more in f o r m e d b a s i s f o r d e c i s i o n making i n management of those v i t a l a r e a s . 7 I I . LITERATURE REVIEW An e x t e n s i v e l i t e r a t u r e e x i s t s f o r s a l t marsh v e g e t a t i o n on a w o r l d w i d e s c a l e . In t h i s s t u d y , o n l y l i t e r a t u r e on s a l t marshes of the P a c i f i c c o a s t of N o r t h America w i l l be r e v i e w e d . 2.1 STUDIES ON THE ECOLOGY OF PACIFIC COASTAL MARSHES 2.1.1 BRITISH COLUMBIA S t u d i e s of c o a s t a l marsh v e g e t a t i o n i n B r i t i s h Columbia a r e r e c e n t i n o r i g i n and most s t u d i e s have been done on the F r a s e r R i v e r e s t u a r y . Lack of i n f o r m a t i o n on e s t u a r i e s o t h e r than the F r a s e r i s c r i t i c a l as the F r a s e r R i v e r marshes have a somewhat d i f f e r e n t s p e c i e s c o m p o s i t i o n from o t h e r a r e a s i n B r i t i s h C o l umbia. The F r a s e r R i v e r i s not t y p i c a l of most B.C. r i v e r s d r a i n i n g t o the c o a s t . I t d r a i n s a v e r y l a r g e a r e a (228000 km 2), has 1500 ha of w e t l a n d s , 20000 ha of i n t e r t i d a l f l a t s and i s g r e a t l y m o d i f i e d by d y k i n g . In c o n t r a s t , the average c o a s t a l r i v e r i n B r i t i s h Columbia ( e . g . T a s h i s h R i v e r ) d r a i n s l e s s than 400 km 2, has 42 ha of i n t e r t i d a l f l a t s and 51 ha of marsh, and i s r e l a t i v e l y p r i s t i n e (Dawe & White 1986). C o a s t a l marshes i n 8 B r i t i s h Columbia range from r e l a t i v e l y f r e s h a r e a s such as the F r a s e r R i v e r , t o b r a c k i s h (<20 p a r t s per thousand s a l i n i t y ) and s a l t marshes (>20 p a r t s per thousand s a l i n i t y ) . In t h i s l i t e r a t u r e r e v i e w no d i s t i n c t i o n has been made among the t h r e e t y p e s . D e s c r i p t i v e s t u d i e s of p l a n t communities on the F r a s e r R i v e r have been r e p o r t e d by McLaren (1972), Forbes (1972) and H i l l a b y & B a r r e t (1976). P arsons (1975) s t u d i e d the v e g e t a t i o n of a F r a s e r R i v e r marsh and c o r r e l a t e d the v e g e t a t i o n p a t t e r n t o e l e v a t i o n , s o i l pH and s o i l c o n d u c t i v i t y . Burgess (1971) r e l a t e d p l a n t d i s t r i b u t i o n t o t i d a l f l o o d i n g , d r a i n a g e and s o i l and water s a l i n i t y . B r a d f i e l d & P o r t e r (1982) s t u d i e d the v e g e t a t i o n s t r u c t u r e and d i v e r s i t y of a F r a s e r R i v e r marsh and r e l a t e d v e g e t a t i o n zones t o the h y d r o l o g i c a l regime - i n u n d a t i o n g r a d i e n t and d r a i n a g e . H u t c h i n s o n (1982) c o n s i d e r e d e l e v a t i o n t o be the most i m p o r t a n t e n v i r o n m e n t a l 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 i n a F r a s e r R i v e r marsh, f o l l o w e d by water s a l i n i t y and s o i l t e x t u r e and m o i s t u r e . A number of M.Sc. t h e s e s have been w r i t t e n on F r a s e r R i v e r marshes. Yamanaka (1975) examined p r o d u c t i v i t y r e l a t i o n s and c o n s i d e r e d e l e v a t i o n and s o i l s a l i n i t y t o be i m p o r t a n t e n v i r o n m e n t a l f a c t o r s d e t e r m i n i n g z o n a t i o n . Moody (1978) a l s o examined p r o d u c t i v i t y as w e l l as the d e c o m p o s i t i o n of s e l e c t e d s a l t marsh p l a n t s . She found t h a t shoot d e n s i t y , r e p r o d u c t i v e shoot number, and the n i t r o g e n c o n t e n t of p l a n t s were r e l a t e d t o e l e v a t i o n , s a l i n i t y , and t e m p e r a t u r e . In Moody's t r a n s p l a n t e x p e r i m e n t s , a l l s p e c i e s grew be s t a t low s a l i n i t y . P o r t e r 9 (1982) s t u d i e d v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s i n t i d a l marshes of the F r a s e r R i v e r and d i v i d e d the marshes i n t o f r e s h and s a l i n e t y p e s . C o r r e l a t i o n s between e n v i r o n m e n t a l f a c t o r s were d i f f e r e n t i n the two t y p e s . Three r e p o r t s on the F r a s e r R i v e r E s t u a r y have been p u b l i s h e d by the Westwater Research C e n t r e a t the U n i v e r s i t y of B r i t i s h Columbia: N o r t h c o t e (1974) d e s c r i b e d the b i o l o g y of the F r a s e r r i v e r ; K i s t r i t z (1978) d e s c r i b e d the r o l e of d e t r i t u s and c y c l i n g of elements i n the e s t u a r y ; K i s t r i t z & Y e s a k i (1979) r e p o r t e d on p r i m a r y p r o d u c t i v i t y , 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 . A management p l a n f o r the F r a s e r R i v e r e s t u a r y was produced by the B r i t i s h Columbia and F e d e r a l Governments i n 1978. A r e p o r t on the h a b i t a t of the F r a s e r R i v e r was i n c l u d e d ( H a b i t a t Work Group 1978). On the lower m a i n l a n d , Lim & L e v i n g s (1973) s t u d i e d the d i s t r i b u t i o n and biomass of p l a n t s on the Squamish R i v e r d e l t a w h i l e L e v i n g s & Moody (1976) d e s c r i b e d v e g e t a t i o n communities, the p r o d u c t i v i t y of Carex l y n g b y e i and the e f f e c t s of d r e d g i n g and d y k i n g on the e s t u a r y . A management p l a n f o r the Squamish e s t u a r y was completed i n 1981 by the H a b i t a t Work Group. Kennedy (1982) s t u d i e d the p l a n t communities and s t a n d i n g c r o p s of e i g h t e e n e s t u a r i e s on Vancouver I s l a n d and one on the m a i n l a n d . She d i s t i n g u i s h e d e l e v e n t y p e s of e s t u a r i e s based m a i n l y on f a c t o r s t h a t i n f l u e n c e s a l i n i t y ( t i m e of maximum d i s c h a r g e , r e l a t i o n s h i p between d i 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 , p r e c i p i t a t i o n , p r o t e c t i o n from wind and wave 10 energy, s o i l t e x t u r e and t i d a l i n u n d a t i o n ) . Kennedy a l s o noted a g e n e r a l d i s t i n c t i o n i n the p l a n t communities o c c u r r i n g n o r t h of Courtenay from those o c c u r r i n g south of Courtenay on Vancouver I s l a n d . B r a c k i s h marshes s o u t h of Courtenay were dominated by Juncus spp., P o t e n t i l l a a n s e r i n a and D i s t i c h l i s  s p i c a t a ; s a l t marshes were c h a r a c t e r i z e d by 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 . Marshes n o r t h of Courtenay were dominated by Deschampsia c e s p i t o s a and P o t e n t i l l a  a n s e r i n a i n b r a c k i s h a r e a s ; marshes i n s a l i n e a r e a s c o n t a i n e d m a i n l y T r i q l o c h i n maritimum and g r a s s e s . Notes on the a u t e c o l o g y of each s p e c i e s were i n c l u d e d . Other s t u d i e s of Vancouver I s l a n d marshes i n c l u d e those of Ceska (1981) f o r the T s i t i k a R i v e r and van D i e r e n (1982) f o r the Somass R i v e r e s t u a r y . Dawe & White (1982, 1986) s t u d i e d p l a n t c ommunities, e n v i r o n m e n t a l f a c t o r s and p r o d u c t i v i t y a t the L i t t l e Q ualicum and N a n o o s e - B o n e l l e s t u a r i e s . They d e t e r m i n e d t h a t s a l i n i t y , e l e v a t i o n and s o i l t e x t u r e were the major f a c t o r s 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 c o n t r o l s the t y p e of marsh - f r e s h , b r a c k i s h or s a l t - w h i l e e l e v a t i o n and t e x t u r e d e termine th e s p e c i e s d i s t r i b u t i o n w i t h i n a marsh. The Cowichan e s t u a r y t a s k f o r c e r e p o r t (Environment and Land Use Committee S e c r e t a r i a t 1980) d e s c r i b e s the f l o r a and fauna of the e s t u a r y and g i v e s s u g g e s t i o n s f o r management. B r a d f i e l d & Campbell (1986) s t u d i e d v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s i n two dyked e s t u a r i e s (Cluxewe and K o k i s h ) of n o r t h e a s t e r n Vancouver I s l a n d . The r e l a t i o n s h i p between v e g e t a t i o n c o m p o s i t i o n and e l e v a t i o n tended t o be s t r o n g e r i n 11 the undyked a r e a s than i n the m o d i f i e d a r e a s i n s i d e the dykes. The a r e a s b e h i n d the dykes were a l s o l o c a t e d a t h i g h e r e l e v a t i o n s , p o s s i b l y due t o a b u i l d - u p of o r g a n i c matter because of a l a c k of f l u s h i n g from the r i v e r . A f i e l d - d e r i v e d community c l a s s i f i c a t i o n a c c o u n t e d f o r h i g h p r o p o r t i o n s of the v a r i a t i o n i n the v e g e t a t i o n d a t a , i n d i c a t i n g t h a t the communities were d i s t i n c t u n i t s . M u l t i v a r i a t e a n a l y s i s of v a r i a n c e was used t o show t h a t a l t h o u g h e l e v a t i o n was i m p o r t a n t , f a c t o r s o t h e r than e l e v a t i o n were m a i n l y r e s p o n s i b l e f o r p l a n t community f o r m a t i o n . C a l d e r & T a y l o r (1968) d e s c r i b e d the v e g e t a t i o n of s a l t marshes on the Queen C h a r l o t t e I s l a n d s . Two g e n e r a l t y p e s of marsh were r e c o g n i z e d , based on s a l i n i t y . Ten volumes p r e p a r e d by the E s t u a r y Working Group g i v e d e t a i l e d i n f o r m a t i o n on t e n B r i t i s h Columbia e s t u a r i e s . I n f o r m a t i o n i s p r e s e n t e d on g e o l o g y and s o i l s , c l i m a t e , h y d r o l o g y , oceanography, i n v e r t e b r a t e s , f i s h , f l o r a , w i l d l i f e , l a n d and water use, p o l l u t i o n and water q u a l i t y and e f f e c t s of development. E s t u a r i e s s t u d i e d were the F r a s e r (Hoos & Packman 1974), Squamish (Hoos & V o i d 1975), Skeena (Hoos 1975), Cowichan-Chemainus ( B e l l & K a l l m a n 1976a), Nanaimo ( B e l l & Kallman 1976b), K i t i m a t ( B e l l & K a l l m a n 1976c), Campbell ( B e l l & Thompson 1977), Courtenay ( M o r r i s e t a l . 1979), Somass ( M o r r i s & Leaney 1980), and B e l l a C o o l a (Leaney & M o r r i s 1981). The r e c e n t l y p u b l i s h e d B r i t i s h Columbia E s t u a r i n e I n f o r m a t i o n C a t a l o g u e (Hagen 1984a, 1984b) g i v e s r e s o u r c e i n f o r m a t i o n on a i r photos and maps a v a i l a b l e as w e l l as a b i b l i o g r a p h y of i n f o r m a t i o n f o r e s t u a r i e s on the lower m a i n l a n d -1 2 Sunshine Coast and e a s t c o a s t of V a n c o u v e r - I s l a n d . The B r i t i s h Columbia M i n i s t r y of Environment has been mapping the v e g e t a t i o n and s u b s t r a t u m - t y p e s of s e l e c t e d e s t u a r i e s s i n c e 1979, u s u a l l y a t a s c a l e of 1:5000 (Maps a v a i l a b l e from B.C. M i n i s t r y of Environment and P a r k s , V i c t o r i a , B.C.). A manual has been p u b l i s h e d on e s t u a r i n e h a b i t a t mapping, and a c l a s s i f i c a t i o n system f o r B r i t i s h Columbia s a l t marshes has been d e v e l o p e d (Hunter e t a l . 1983). Three c a t e g o r i e s of s a l t marshes a r e mapped - low, i n t e r m e d i a t e and h i g h marsh - based m a i n l y on e l e v a t i o n and s p e c i e s c o m p o s i t i o n . A l l major e s t u a r i e s i n B r i t i s h Columbia have been g i v e n a r a t i n g of importance w i t h r e s p e c t t o p r e s e r v a t i o n , p r o t e c t i o n and enhancement (see Appendix A ) . Wetlands were a s s e s s e d on the b a s i s of w a t e r f o w l , w i l d l i f e and f i s h usage, p r o d u c t i v i t y , s o c i a l f a c t o r s (how the p u b l i c p e r c e i v e s t h e area),--imminence of development and enhancement and r e h a b i l i t a t i o n p o t e n t i a l (Hunter et a l ^ 1 985) . 2.1.2 WASHINGTON R e l a t i v e l y few s t u d i e s on Washington S t a t e s a l t marshes were found i n the l i t e r a t u r e . D i s r a e l i & Fonda (1979) used a g r a d i e n t a n a l y s i s approach t o study a s a l t marsh i n B e l l i n g h a m Bay. They found t h a t marsh s p e c i e s were d i s t r i b u t e d m a i n l y 13 a c c o r d i n g t o e l e v a t i o n , l e n g t h of t i d a l submergence, s o i l t e x t u r e and s o i l m o i s t u r e . Burg e t a l . (1976, 1980) s t u d i e d the N i s q u a l l y s a l t marsh i n Puget Sound and d e t e r m i n e d e l e v a t i o n and water s a l i n i t y t o be the most i m p o r t a n t f a c t o r s 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 . The S k a g i t marsh was s t u d i e d i n g r e a t d e t a i l by Ewing (1982, 1983). He i n v e s t i g a t e d p l a n t communities and p l a n t response t o v a r i a t i o n i n s o i l t e x t u r e , o r g a n i c m a t t e r , s o i l t e m p e r a t u r e , s o i l water s a l i n i t y , s o i l redox p o t e n t i a l , e l e v a t i o n and time of i n u n d a t i o n . S a l i n i t y , e l e v a t i o n , t e x t u r e and s o i l redox p o t e n t i a l were ranked i n d e c r e a s i n g o r d e r of importance as the main f a c t o r s i n f l u e n c i n g s p e c i e s c o m p o s i t i o n . S e l i s k a r & G a l l a g h e r (1983) produced a g e n e r a l r e p o r t on the marshes of Washington and Oregon, d i s c u s s i n g the p h y s i c a l and c h e m i c a l environment, marsh d i s t r i b u t i o n , b i o t i c c o m munities, e c o l o g i c a l i n t e r a c t i o n s and management. 2.1.3 OREGON Many s t u d i e s have been un d e r t a k e n i n Oregon, b e g i n n i n g w i t h a d e s c r i p t i o n of the Coos Bay Region by House (1914). A k i n s & J e f f e r s o n (1973) s t u d i e d s e v e r a l marshes a l o n g the Oregon c o a s t and d i s t i n g u i s h e d e i g h t marsh-types r e l a t e d t o e l e v a t i o n , t e x t u r e , s a l i n i t y and d i s t u r b a n c e . J e f f e r s o n (1975) s t u d i e d p l a n t communities and s u c c e s s i o n i n n i n e t e e n s a l t marshes and d i s t i n g u i s h e d s i x v e g e t a t i o n t y p e s based on e l e v a t i o n , s u b s t r a t e 1 4 and s a l i n i t y . She a l s o worked out s u c c e s s i o n a l sequences based on z o n a t i o n and e x a m i n a t i o n of p a r t i a l l y decomposed p l a n t m a t e r i a l i n the s o i l . E i l e r s (1975, 1976) d e s c r i b e d p l a n t communities of the Nehalem marshes and r e l a t e d s p e c i e s and communities t o e n v i r o n m e n t a l f a c t o r s . He a l s o l o o k e d a t p r o d u c t i v i t y , the h i s t o r y of marsh development and the a b i l i t y t o d i s t i n g u i s h p l a n t communities on a i r p h o t o s . P l a n t s p e c i e s d i s t r i b u t i o n s were r e l a t e d m a i n l y t o e l e v a t i o n and t i d e l e v e l s . Liverman (1982) s t u d i 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 i n r e l a t i o n t o e n v i r o n m e n t a l f a c t o r s a t N e t a r t s S p i t . H i s purpose was t o examine methods of d e t e r m i n i n g the boundary between w e t l a n d and upl a n d v e g e t a t i o n . M i t c h e l l (1982) s t u d i e d s a l t marsh r e - e s t a b l i s h m e n t f o l l o w i n g dyke b r e a c h i n g a t the Salmon R i v e r . A d e c r e a s e i n abundance of u p l a n d s p e c i e s and an i n c r e a s e i n s a l t t o l e r a n t s p e c i e s were no t e d . B e f o r e b r e a c h i n g o c c u r r e d , the dyked marshes had lower s a l i n i t y , lower pH and lower e l e v a t i o n than undyked a r e a s . A f t e r b r e a c h i n g , s a l i n i t y and pH i n c r e a s e d . T a y l o r (1983) i n v e s t i g a t e d p l a n t communities i n a dyked and a d j a c e n t undyked marsh i n Coos Bay. The v e g e t a t i o n i n the two a r e a s was d i f f e r e n t and the a r e a b e h i n d the dyke had s u b s i d e d . McVay et a l . (1980) s t u d i e d t i d a l f r e s h water marsh e s t a b l i s h m e n t on dredge s p o i l s i n the Columbia R i v e r . 15 2.1.4 CALIFORNIA An e a r l y study of s a l t marshes i n C a l i f o r n i a i s P u r e r ' s (1942) i n v e s t i g a t i o n of s a l t marsh p l a n t s i n San Diego County. The v e g e t a t i o n was d i v i d e d i n t o t h r e e zones on the b a s i s of t i d a l submergence. S o i l and water s a l i n i t y and t i d e l e v e l s were r e l a t e d t o the d i s t r i b u t i o n of the p r i n c i p a l genera i n the s a l t marsh. Cooper (1926) and Hinde (1954) examined s a l t marsh v e g e t a t i o n i n San F r a n c i s c o Bay. Hinde i d e n t i f i e d t h r e e major v e g e t a t i o n a s s o c i a t i o n s which were based on t i d a l emergence and submergence. S t u d i e s on the v e g e t a t i o n of Newport Bay were r e p o r t e d by Stevenson (1954), Stevenson & Emery (1958) and V o g l (1966). Stevenson & Emery r e l a t e d z o n a t i o n t o t i d a l immersion and s o i l f a c t o r s ; V o g l i d e n t i f i e d t h r e e v e g e t a t i o n zones i n the marsh a l t h o u g h the zones d i d not have d i s c r e t e b o u n d a r i e s . Z e d l e r (1977) s t u d i e d the v e g e t a t i o n of the T i j u a n a r i v e r e s t u a r y and c o n c l u d e d t h a t the v e g e t a t i o n c o u l d not be d i v i d e d i n t o zones. Of s e v e r a l e n v i r o n m e n t a l f a c t o r s examined ( e l e v a t i o n , s a l i n i t y , o r g a n i c m a t t e r , b u l k d e n s i t y , t e x t u r e and pH) e l e v a t i o n was d e t e r m i n e d t o be the most i m p o r t a n t . Barbour (1970) d e s c r i b e d a s a l t marsh at Bodega Head, n o r t h of San F r a n c i s c o , w h i l e M a h a l l & Park (1976a, b & c) d e s c r i b e d the ecotone between S p a r t i n a a l t e r n i f l o r a and S a l i c o r n i a v i r g i n i c a i n San F r a n c i s c o Bay. Z o n a t i o n was r e l a t e d t o s a l i n i t y . Macdonald (1977) p u b l i s h e d a r e v i e w of s t u d i e s on C a l i f o r n i a n s a l t marshes which i n c l u d e d i n f o r m a t i o n on h i s t o r y , 16 e n v i r o n m e n t a l v a r i a b l e s , z o n a t i o n and the a u t e c o l o g y of seven s p e c i e s . He d i s t i n g u i s h e d t h r e e groups of C a l i f o r n i a n marshes -n o r t h e r n , c e n t r a l and s o u t h e r n . Two s t u d i e s on San F r a n c i s c a n s a l t marshes a r e those of F e l t o n (1978) on m i c r o c l i m a t i c f a c t o r s and W i n f i e l d (1980) on p r i m a r y p r o d u c t i v i t y and o r g a n i c carbon and i n o r g a n i c n i t r o g e n c y c l e s . Newby (1980) i n v e s t i g a t e d n u t r i e n t l e v e l s i n t i s s u e s of S p a r t i n a f o l i o s a and S a l i c o r n i a v i r g i n i c a w i t h r e s p e c t t o s p e c i e s d i s t r i b u t i o n . T i s s u e phosphorus had a s t r o n g c o r r e l a t i o n , w h i l e n i t r o g e n , sodium, c a l c i u m and i r o n had weak c o r r e l a t i o n s . In a study of a s a l t marsh t h a t has been dyked s i n c e 1899, E i l e r s (1980) found t h a t the a r e a b e h i n d the dyke had s u b s i d e d w i t h r e s p e c t t o e l e v a t i o n . Z e d l e r (1982) r e p o r t e d on s e v e r a l s o u t h e r n C a l i f o r n i a n s a l t marshes, i n c l u d i n g i n f o r m a t i o n on p h y s i o l o g y , g e o l o gy, h y d r o l o g y , c l i m a t e , v e g e t a t i o n , f i s h , w i l d l i f e and management. Z e d l e r a t t r i b u t e d the d i f f e r e n c e s between marshes m a i n l y t o d i f f e r e n c e s i n d i s t u r b a n c e and t i d a l f l u s h i n g . 2.1.5 ALASKA A number of s t u d i e s on A l a s k a n s a l t marshes have been r e p o r t e d . Cooper (1931) d e s c r i b e d the c o l o n i z a t i o n of m u d f l a t s f o l l o w i n g a r e c e n t g l a c i a l r e t r e a t a t G l a c i e r Bay. Hanson 17 (1951) d e s c r i b e d p l a n t communities and s o i l p r o f i l e s a t s i t e s i n s o u t h - c e n t r a l A l a s k a . Stephens & B i l l i n g s (1967) s t u d i e d p l a n t communities and e n v i r o n m e n t a l f a c t o r s (pH, c a t i o n exchange c a p a c i t y , exchangeable c a t i o n s and s o i l s p r o f i l e s ) i n C h i c a g o f I s l a n d , and r e l a t e d the d i s t r i b u t i o n of communities t o e l e v a t i o n and s o i l c h a r a c t e r i s t i c s . The Copper r i v e r d e l t a was s t u d i e d by Crow (1968, 1971). T h i s a r e a was u p l i f t e d a p p r o x i m a t e l y 1.9 m by an earthquake i n 1964. Decreases i n s a l i n i t y , pH and s e v e r a l c a t i o n s were n o t e d , a l o n g w i t h the i n v a s i o n of a number of n o n - s a l i n e s p e c i e s and sh r u b s . Z o n a t i o n was r e l a t e d t o e l e v a t i o n and s o i l t e x t u r e . Crow & Koppen (1977) d e s c r i b e d the p l a n t communities of a marsh i n Kachemak Bay. J e f f e r i e s (1977a) s t u d i e d the v e g e t a t i o n , water s a l i n i t y and p r o d u c t i v i t y of s a l t marshes a t seven s i t e s on the A r c t i c c o a s t of A l a s k a and Canada. A paper by Crow (1977) g i v e s g e n e r a l i n f o r m a t i o n about A l a s k a n s a l t marshes i n c l u d i n g d a t a on water s a l i n i t y , pH, t e x t u r e and e l e v a t i o n . S e r g i e f I s l a n d was s t u d i e d by Sparks e t a l . (1977), who d e s c r i b e d p l a n t community t y p e s , and d e l M o r a l & Watson (1978), who d e t e r m i n e d t h a t p l a n t communities were c o n t r o l l e d by t i d a l i n f l u e n c e and p h y s i o g r a p h y . V i n c e & Snow (1984) and Snow & V i n c e (1984) were c o n c e r n e d w i t h f a c t o r s c o n t r o l l i n g s p e c i e s d i s t r i b u t i o n and p l a n t z o n a t i o n on S u s i t n a F l a t s . In the f i r s t paper ( V i n c e & Snow 1984), e i g h t v e g e t a t i o n zones were r e c o g n i z e d t h a t d i f f e r e d w i t h r e s p e c t t o f l o o d i n g f r e q u e n c y , r a t e of s i l t a t i o n , s o i l o r g a n i c c o n t e n t , m o i s t u r e c o n t e n t , redox p o t e n t i a l and s a l i n i t y . S o i l s a l i n i t y 18 and w a t e r l o g g i n g were most i m p o r t a n t i n s e g r e g a t i n g the zones, but whether these f a c t o r s caused the z o n a l d i f f e r e n c e s was not d e t e r m i n e d . In the second paper (Snow & V i n c e 1984), e x p e r i m e n t s were und e r t a k e n t o study the e f f e c t s of s a l i n i t y , w a t e r l o g g i n g and s o i l t y pe on the f i v e dominant s p e c i e s i n the marsh. Pot e x p e r i m e n t s showed t h a t a l l f i v e s p e c i e s had o p t i m a l growth i n the w a t e r l o g g e d , low s a l i n i t y t r e a t m e n t . In t r a n s p l a n t e x p e r i m e n t s , the growth of the t h r e e h i g h marsh s p e c i e s was i n h i b i t e d i n s a l i n e s i t e s , w h i l e P u c c i n e l l i a  n u t k a e n s i s from the low marsh d i d b e t t e r i n the h i g h marsh; T r i g l o c h i n maritimum grew w e l l a t a l l s i t e s . That s u g g e s t s t h a t c o m p e t i t i o n p l a y s a r o l e i n s a l t marsh z o n a t i o n . Stone (1984) i n v e s t i g a t e d f o u r marshes near Juneau, c o n c l u d i n g t h a t s a l t marshes s h o u l d not be compared s o l e l y on the b a s i s of v e g e t a t i o n c o m p o s i t i o n owing t o c o n s i d e r a b l e d i f f e r e n c e s among the marshes. A c l a s s i f i c a t i o n of s a l t marshes based on p h y s i o g r a p h y was proposed t h a t took i n t o account the h i s t o r y and f u t u r e of the v e g e t a t i o n from a p h y s i o g r a p h i c v i e w p o i n t . 2.2 ZONATION Most a u t h o r s have found i t r e l a t i v e l y easy t o d i v i d e s a l t marsh v e g e t a t i o n i n t o zones, a l t h o u g h the zones o f t e n merge i n t o each o t h e r . In a stu d y of s a l t marshes i n Washington and 19 Oregon, S e l i s k a r & G a l l a g h e r (1983) found t h a t i n some marshes the zones were c l e a r w h i l e i n o t h e r s the zones were d i f f u s e . C l e a r z o n a t i o n a r i s e s when s p e c i e s a r e n a r r o w l y d i s t r i b u t e d and t h e i r b o u n d a r i e s do not o v e r l a p , w h i l e s p e c i e s t h a t can t o l e r a t e a wide range of e n v i r o n m e n t a l c o n d i t i o n s and oc c u r i n almost a l l p a r t s of the marsh tend t o cause b l u r r e d community b o u n d a r i e s (Ranwell 1972; Gray et a l ^ 1979; Z e d l e r 1982; V i n c e & Snow 1984). Z o n a t i o n i n s a l t marshes has o f t e n been c o n s i d e r e d t o i n d i c a t e d i f f e r e n t s t a g e s of s u c c e s s i o n (e.g. J e f f e r s o n 1975), but a r e c e n t study by Roozen & Westhoff (1985) d i s p u t e s t h i s . In a t h i r t y - y e a r study of a s a l t marsh i n the N e t h e r l a n d s , d i f f e r e n t a b i o t i c f a c t o r s were r e l a t e d t o the d i f f e r e n c e s between the zones; i n t h a t case z o n a t i o n was not e q u i v a l e n t t o s u c c e s s i o n . V e g e t a t i o n zones o f t e n d i f f e r from marsh t o marsh i n s p e c i e s - - c o m p o s i t i o n and r e l a t i o n s h i p s w i t h e n v i r o n m e n t a l f a c t o r s , but some common t r e n d s can be no t e d . The lower marsh tends t o be dominated by f a c t o r s a s s o c i a t e d w i t h submergence i n c o n t r a s t w i t h the upper marsh which i s l e s s f r e q u e n t l y submerged. The a b i l i t y t o t o l e r a t e t i d a l submergence and h i g h s a l i n i t y m a i n l y d e t e r m i n e s the lower l i m i t of a s p e c i e s on the marsh; c o m p e t i t i o n i s thought t o determine the upper l i m i t i n some c a s e s (Chapman 1974; R a n w e l l 1972; P i e l o u & Ro u t l e d g e 1976). 20 2.3 ENVIRONMENTAL FACTORS THAT INFLUENCE PLANT ZONATION Most a u t h o r s c o n s i d e r s u b s t r a t u m e l e v a t i o n , which d e t e r m i n e s t i d a l i n u n d a t i o n , t o be the most i m p o r t a n t 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 s a l t marsh v e g e t a t i o n (see Table I ) . S a l i n i t y , s o i l t e x t u r e , and s o i l m o i s t u r e a r e a l s o ranked h i g h l y . Other f a c t o r s of importance a r e n u t r i e n t s , o r g a n i c m a t t e r , pH, redox p o t e n t i a l , c o m p e t i t i o n , r a i n f a l l , t emperature of the a i r and w a t e r , e v a p o r a t i o n , l i g h t i n t e n s i t y , t u r b i d i t y , b i o t a , p h y s i o g r a p h y , chance and d i s t u r b a n c e . Some of those e n v i r o n m e n t a l f a c t o r s a r e c o n s i d e r e d below. I t s h o u l d be noted t h a t few g e n e r a l i z a t i o n s a r e apparent and the v e g e t a t i o n environment r e l a t i o n s h i p s a r e d i f f e r e n t i n each marsh s t u d i e d . 2.3.1 ELEVATION E l e v a t i o n per se does not c o n t r o l s p e c i e s d i s t r i b u t i o n s -i t i s a summary v a r i a b l e t h a t i s c o r r e l a t e d w i t h a number of e n v i r o n m e n t a l v a r i a b l e s which have a more d i r e c t p h y s i o l o g i c a l i n f l u e n c e (Table I I ) . E l e v a t i o n c o n t r o l s the amount and depth of submergence by t i d a l w ater. S a l i n i t y u s u a l l y d e c r e a s e s w i t h i n c r e a s i n g e l e v a t i o n , except i n a r e a s such as s o u t h e r n C a l i f o r n i a where r a i n f a l l i s low and e v a p o r a t i o n i s h i g h c r e a t i n g h y p e r s a l i n e c o n d i t i o n s i n the upper marsh. Table I: Review of literature oo salt marsh vegetation - environment relationships Author and Area Method of Analysis Distribution of vegetation mainly related to: Elevation Salinity Texture Moisture Other Chapman, 1938 England Purer, 1942 California Hinde, 1954 California Adams, 1963 eastern North America Stephens and Billings, 1967 Alaska Crow, 1968 Alaska Burgess, 1971 Fraser R., B.C. Obser. Obser. Obser. Obser. Obser. Obser. Obser. Gray and Bunce, 1972 P.C.A and Gray and Scott,' 1977 England Waisel, 1972 worldwide Ranwell, 1972 worldwide Akins and Jefferson, 1973 Oregon Jefferson, 1975 Oregon Parsons, 1975 Fraser R., B.C. Yamanaka, 1975 Fraser R., B.C. Eilers, 1975 Oregon Zedler, 1977 California Crow, 1977 Alaska del Moral and Watson, 1978 Alaska Moody, 1978 Fraser R., B.C. Obser. Obser. Obser. Obser. Obser. Obser. P.O. Obser. Obser. Obser. Correla-tions X X X X X X X X r a i n f a l l , tecp. ,evap., biota pH, cations nutrients, organic matter organic matter light, tenp. turbidity, disturbance disturbance pH organic matter sulphate, PH physiography tenperature "Cable I continued Author and Area Method Analysis Distribution of vegetation mainly related to? Elevation Salinity Texture Moisture Other Disraeli and Fonda, 1979 Washington Burg et a l . , 1980 Washington Porter, 1982 Fraser R., B.C. Hutchinson, 1982 Fraser R., B.C. AN0VA Obser. P.C.A. R.A. ANOVA Dawe and White, 1982 ANOVA L i t t l e Qualicum R., B.C. Bradfield and Porter, P.C.A 1982 Fraser R., B.C. van Dieren, 1982 Sanass R., B.C. Cooper, 1982 Zedler, 1982 California Liverman, 1982 Oregon R.A. Obser. Pot experiment Obser. Cluster, R.A. B.D.A. Obser. Ewing, 0982 and 1983 Washington Seliakar and Gallag-her, 1983 Wash, and Oregon Snow and Vince, 1984 ANOVA and Vince and Snow, 1984 Alaska Frey and Bason, 1985 Obser. worldwide Rozerna et a l . , 1985 Pot experiment X Dawe and White, 1986 ANOVA Nanoose-Bonell B. B.C. Bradfield and Campbell, 1986 Kokish and Cluxewe B.C. P.C.A. X X X nitrogen cations physiography disturbance nutrients PH redox potential nutrients chance redox potential organic matter carpetition Method of Analysis ANOVA - Analysis of Variance B.D.A. - Binary Discriminant Analysis C C A . - Canonical Correlation Analysis Cluster- Cluster Analysis Obser. - Observation P.C.A. - Principal Ocnponent Analysis P.O. - Polar Ordination R.A. - Reciprocal Averaging 2 X " Environmental factor Important Table II: He view of literature on correlations between elevation and environmental factors Author and Area As elevation increases, Salinity Moisture Ortranic matter Texture Nutrients Ranwell, 1964 England Stephens and Billings, 1967 Alaska Pigott, 1969 worldwide Gray and Bunce, 1972 England Ranwell, 1972 worldwide Parsons, 1975 leaser R., B.C. Eilers, 1975 Oregon Crow, 1977 Alaska Zedler, 1977 California Mac-Donald, 1977 California Disraeli and Fonda, 1979 Washington Seliskar, 1981 Oregon Liverman, 1982 Oregon Porter, 1982 Fraser R., B.C. Hutchinson, 1982 Fraser R., B.C. Ewing, 1982 Washington Zedler, 1982 California Dec. Dec. Dec. Dec. Dec. Dec. Dec. Dec. Dec. Dec. Inc. Long and Mason, 1983 1983 worldwide Vince and Snow, 1984 Alaska Ingold and Havill, 1984 England Dec. Dec. Inc. Dec. Inc. Inc. Inc. Inc Inc. Dec. Inc. Inc. Dec. Inc. Inc. Inc. Sand Dec, No No Gravel Dec. Dec. Sil t , Clay Inc. Clay Dec, Sil t Dec, Sand Inc. Sil t Inc. Sand Dec No Clay Dec Sil t Dec Si l t Inc. Sand Dec Sand Inc Dec. P,K,N Inc. Ca Dec. Cations Dec N,P Inc. Nutrients Inc. N,K Inc. Sulphide Dec. Dec. Decreases Inc. Increases No No Correlation 24 Lower e l e v a t i o n s n o r m a l l y have h i g h e r s o i l m o i s t u r e , lower o r g a n i c m atter c o n t e n t (except f o r Z e d l e r 1977, 1982 i n C a l i f o r n i a ) , and lower b u l k d e n s i t y . A e r a t i o n i s g r e a t e r a t h i g h e r e l e v a t i o n s owing t o h i g h e r l e v e l s of o r g a n i c matter and l e s s f r e q u e n t submergence. S o i l t e x t u r e i s o f t e n c o r r e l a t e d w i t h e l e v a t i o n : lower e l e v a t i o n s u s u a l l y have sandy s o i l s w h i l e h i g h e r e l e v a t i o n s have a l a r g e r p r o p o r t i o n of s i l t and c l a y . In C a l i f o r n i a , t h a t t r e n d i s r e v e r s e d ( Z e d l e r 1977, 1982; Macdonald 1977). The v e l o c i t y of the water i s reduced i n the upper marsh zones due t o i n c r e a s e d amounts of v e g e t a t i o n , f l a t n e s s of t e r r a i n and r e d u c t i o n of wave a c t i o n . That u s u a l l y causes f i n e sediment t o s e t t l e out i n the upper a r e a s (Crow 1977). There seems t o be no c o n s i s t e n t t r e n d between pH and e l e v a t i o n . Some s t u d i e s have found no c o r r e l a t i o n between the two, some have found h i g h e r pH a t low e l e v a t i o n s , w h i l e o t h e r s have found lower pH a t low e l e v a t i o n s (see T a b l e I I ) . S u l p h i d e c o n c e n t r a t i o n t e n d s t o be h i g h e s t a t low e l e v a t i o n s . N i t r o g e n , phosphorus and p o t a s s i u m i n c r e a s e w i t h i n c r e a s i n g e l e v a t i o n w h i l e o t h e r c a t i o n s d e c r e a s e . At low e l e v a t i o n s , d e n s i t y independent m o r t a l i t y ( m a i n l y from f l o o d i n g ) tends t o be dominant w h i l e a t h i g h e l e v a t i o n s d e n s i t y dependent m o r t a l i t y ( e.g. c o m p e t i t i o n f o r space and l i g h t ) i s more i m p o r t a n t . R e p r o d u c t i v e e f f o r t tends t o be h i g h e r a t low e l e v a t i o n s ( J e r l i n g 1985). S e l i s k a r (1985a, 1985b) found t h a t m o r p h o l o g i c a l c h a r a c t e r i s t i c s ( h e i g h t of p l a n t and f l o w e r i n g s h o o t s , l e a f number and w i d t h , i n t e r n o d e l e n g t h , 25 b r a n c h i n g , stem d i a m e t e r and a n a t o m i c a l stem structur-e-) of the same p l a n t s p e c i e s were d i f f e r e n t a t h i g h and low e l e v a t i o n s . That d i f f e r e n c e r e f l e c t e d a p l a s t i c response and was p r o b a b l y due t o s o i l m o i s t u r e which was h i g h e r a t low e l e v a t i o n s . T r a n s p l a n t e x p e r i m e n t s seem t o i n d i c a t e t h a t c o m p e t i t i o n i s more im p o r t a n t a t h i g h e l e v a t i o n ( S t a l t e r & Batson 1969; Snow & V i n c e 1984). 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 w i t h i n c r e a s i n g e l e v a t i o n (e.g. E i l e r s 1975; Dawe & White 1982; V i n c e & Snow 1984). 2.3.2 TIDES V a r i o u s t i d a l f a c t o r s i n f l u e n c e the d i s t r i b u t i o n of s a l t marsh v e g e t a t i o n . M e c h a n i c a l d i s t u r b a n c e due t o t i d a l a c t i o n , can d i s r u p t t h e s u b s t r a t e and uproot s e e d l i n g s ( R a n w e l l 1972). T i d a l submergence can change ambient t e m p e r a t u r e s , reduce oxygen and carbon d i o x i d e a v a i l a b i l i t y , and d e c r e a s e i n c i d e n t l i g h t , thus r e d u c i n g the p h o t o s y n t h e t i c and r e s p i r a t i o n r a t e s of the marsh p l a n t s . T i d a l f l u s h i n g i n f l u e n c e s s o i l and water s a l i n i t i e s and c o n t r o l s d i s s o l v e d n u t r i e n t and t r a c e element exchange between the marsh p l a n t s , s u b s t r a t e and water (Macdonald 1977). The v e r t i c a l range of the t i d e c o n t r o l s t i d a l f l o o d i n g depths and the v e r t i c a l e x t e n t of the s a l t marsh, w h i l e the form of the t i d a l c y c l e c o n t r o l s the fr e q u e n c y and d u r a t i o n of 26 submergence and emergence. The v e r t i c a l range of s a l t marsh f o r m a t i o n i s g r e a t e r i n l a r g e r t i d e range a r e a s ; i n a d d i t i o n , marshes i n a l a r g e t i d e range have more c l e a r l y zoned v e g e t a t i o n and s h a r p e r d r a i n a g e systems l o c a t e d a t r i g h t a n g l e s t o the s h o r e . Marshes w i t h a s m a l l t i d a l range have l e s s c l e a r l y zoned v e g e t a t i o n and s l u g g i s h d r a i n a g e which produces a complex network of w i n d i n g , many branched c r e e k s ( R a n w e l l 1972). R o u t l e d g e (1975) found t h a t the more n o r t h e r l y marshes of e a s t e r n Canada had wider zones p o s s i b l y due t o a g r e a t e r t i d a l range. 2.3.3 NUTRIENTS F o u r t e e n elements a r e u s u a l l y c o n s i d e r e d t o be e s s e n t i a l f o r p l a n t growth. S i x of those a r e used i n r e l a t i v e l y l a r g e q u a n t i t i e s and a r e thus r e f e r r e d t o as m a c r o n u t r i e n t s . They a r e n i t r o g e n ( N ) , phosphorus ( P ) , p o t a s s i u m ( K ) , c a l c i u m ( C a ) , magnesium (Mg), and s u l p h u r ( S ) . M i c r o n u t r i e n t s , or t r a c e e l e m e n t s , a r e u t i l i z e d i n s m a l l amounts and i n c l u d e i r o n ( F e ) , manganese (Mn), z i n c (Zn) and copper (Cu), boron ( B ) , molybdenum (Mo), c h l o r i n e ( C I ) , and c o b a l t ( C o ) . N u t r i e n t s can e n t e r a marsh through groundwater, r a i n , t i d a l f l o o d i n g and r i v e r s ; n u t r i e n t s l e a v e a marsh m a i n l y d u r i n g the ebb of t i d a l water ( V a l i e l a e t a l . 1978). S a l t marshes a r e u s u a l l y c o n s i d e r e d t o be n u t r i e n t s i n k s w i t h an abundance of 27 most e s s e n t i a l elements f o r p l a n t growth. Input from t i d a l water i s c o n s i d e r e d t o be the most i m p o r t a n t method of e n t r y (Pomeroy 1970; J e f f e r i e s e t a L 1979; Wolf & F u c i k 1981). Phosphorus i s n o r m a l l y i n abundance i n e s t u a r i n e waters owing t o r e l e a s e from c l a y m i n e r a l s i n the se d i m e n t s , i n p u t from r i v e r s , and d e c o m p o s i t i o n of o r g a n i c m a t t e r (Pomeroy e t a l . 1965; Whitney e_t a l . 1981). H i g h phosphate l e v e l s can a l s o o r i g i n a t e from sewage, f e r t i l i z e r s and i n d u s t r i a l e f f l u e n t s which e n t e r the e s t u a r y ( B e e f t i n k e t a l . 1977). Sources of p o t a s s i u m a r e c l a y p a r t i c l e s or sea water ( B e e f t i n k e_t a l . 1977); s o u r c e s of n i t r o g e n a r e sea water, r a i n , groundwater, o r g a n i c m a t t e r and n i t r o g e n f i x a t i o n by b a c t e r i a and b l u e - g r e e n a l g a e . Some p l a n t s , such as the legumes L a t h y r u s and T r i f o l i u m , can f i x t h e i r own n i t r o g e n . N i t r o g e n i s l o s t by d e n i t r i f i c a t i o n and t i d a l exchange ( B e e f t i n k 1977). The two p l a n t n u t r i e n t s t h a t a r e most l i k e l y t o be l i m i t i n g a r e n i t r o g e n and phosphorus ( P a t r i c k & De Laune 1977). In s a l t marshes, n i t r o g e n i s o f t e n l i m i t i n g i n h i g h marsh a r e a s owing t o the i n f r e q u e n c y of t i d a l submergence ( P i g o t t 1969; R a n w e l l 1972; Stewart e t a l . 1972, 1973; Macdonald 1977). N i t r o g e n o c c u r s i n many forms - e l e m e n t a l n i t r o g e n (N), n i t r i t e ( N 0 2 ) , n i t r a t e (N0 3) and ammonium (NH 4 ) . Ammonium i s the predominant form i n s a t u r a t e d sediment, whereas n i t r i t e and n i t r a t e a r e dominant i n a e r a t e d s o i l (De Laune e t a l . 1983). Ammonium i s the p r e f e r r e d form f o r p l a n t uptake (Pomeroy 1980). F e r t i l i z a t i o n e x p e r i m e n t s have been used t o det e r m i n e i f n i t r o g e n and phosphorus a re l i m i t i n g f a c t o r s i n s a l t marsh 28 p r o d u c t i o n . In one s t u d y , P i g o t t (1969) d e t e r m i n e d t h a t growth i n t he upper marsh was i n c r e a s e d by a d d i t i o n of n i t r o g e n and phosphorus, but the main response was t o n i t r o g e n . The p r o d u c t i o n of S p a r t i n a was i n c r e a s e d by n i t r o g e n f e r t i l i z a t i o n , but not by phosphorus, i n both the low and h i g h marsh ( V a l i e l a & T e a l 1974; V a l i e l a e t a l _ j _ 1976). V a l i e l a e t al^ (1975) f e r t i l i z e d an a r e a w i t h sewage sludge and found t h a t t h i s l e d t o i n c r e a s e d p r o d u c t i o n i n both low and h i g h marsh a r e a s ; f u r t h e r m o r e , changes i n s p e c i e s c o m p o s i t i o n a l s o o c c u r r e d ( e . g . S p a r t i n a a l t e r n i f l o r a e x c l u d e d S a l i c o r n i a s p p . ) . S t e w a r t e t a l . (1972, 1973) r e p o r t e d t h a t the growth of Suaeda m a r i t i m a i n the upper marsh appeared t o be n i t r o g e n l i m i t e d . Lower marsh s p e c i e s had h i g h e r n i t r o g e n c o n t e n t s and n i t r a t e r e d u c t a s e l e v e l s than t h o s e from the upper marsh. T h i s i n d i c a t e d t h a t more n i t r o g e n was a v a i l a b l e a t low e l e v a t i o n s . J e f f e r i e s (1977b) found t h a t s p e c i e s of the low marsh had f a s t e r growth r a t e s than t h o s e i n the upper marsh i n response t o n i t r o g e n a d d i t i o n . That was e x p l a i n e d as a r e s u l t of s e l e c t i o n f o r p l a n t s w i t h low growth r a t e s i n the upper marsh t h a t c o u l d s u r v i v e p e r i o d s of growth s t r e s s i n the summer when the s o i l i s h y p e r s a l i n e . C a r g i l l & J e f f e r i e s (1984) d e t e r m i n e d t h a t the a d d i t i o n of ammonium and n i t r a t e i n c r e a s e d p r o d u c t i v i t y e q u a l l y , whereas a d d i t i o n of phosphorus d i d not i n c r e a s e p r o d u c t i o n . A d d i t i o n of n i t r o g e n and phosphorus t o g e t h e r l e d t o h i g h e r p r o d u c t i v i t y than e i t h e r a l o n e , i n d i c a t i n g t h a t when the n i t r o g e n s u p p l y i s i n c r e a s e d , phosphorus becomes l i m i t i n g . T y l e r (1967) found t h a t 2 9 a d d i t i o n of n i t r o g e n as ammonium and phosphorus i n c r e a s e d p r o d u c t i v i t y w h i l e a d d i t i o n of n i t r o g e n as n i t r a t e had no e f f e c t . That may have been due t o the c l a y s o i l , which has a h i g h c a t i o n exchange c a p a c i t y , and can t h e r e f o r e r e t a i n ammonium (a c a t i o n ) b e t t e r than n i t r a t e (an a n i o n ) . Okusanya & Ungar (1984) found t h a t a d d i t i o n s of n u t r i e n t s under h i g h s a l i n i t y c o u l d overcome the s a l i n i t y problem. A d d i t i o n s of n u t r i e n t s i n c r e a s e d the c a t i o n c o n t e n t of the l e a v e s and enhanced growth. N u t r i e n t a v a i l a b i l i t y i s i n f l u e n c e d by s o i l t e x t u r e and o r g a n i c m a t t e r c o n t e n t . F i n e t e x t u r e d s o i l s w i t h o r g a n i c m atter have a h i g h c a t i o n exchange c a p a c i t y and a r e a b l e t o r e t a i n n u t r i e n t s . Broome e t a l . (1975) found t h a t on sandy s u b s t r a t e s , a d d i t i o n s of n i t r o g e n and phosphorus i n c r e a s e d y i e l d s , w h i l e on f i n e t e x t u r e d s e d i m e n t s , o n l y n i t r o g e n caused an i n c r e a s e i n p r o d u c t i o n . In c o n t r a s t t o t h o s e s t u d i e s , J e f f e r i e s & P e r k i n s (1977) found no change i n s t a n d i n g c r o p s and s p e c i e s c o m p o s i t i o n a f t e r p r o v i d i n g r e g u l a r a d d i t i o n s of n i t r o g e n , phosphorus and p o t a s s i u m over a f o u r t o f i v e y e a r p e r i o d . In i n t e r p r e t i n g the r e s u l t s of those e x p e r i m e n t s , c a u t i o n i s needed. Smart & Barko (1980) p o i n t e d out t h a t the r e s u l t s of n i t r o g e n l i m i t a t i o n s t u d i e s based on the response of v e g e t a t i o n t o n i t r o g e n f e r t i l i z a t i o n may be m i s l e a d i n g due t o secondary e f f e c t s of n i t r o g e n f e r t i l i z a t i o n on o t h e r a s p e c t s of the s a l t marsh ecosystem. A d d i t i o n s of n i t r o g e n can cause an i n c r e a s e i n d e c o m p o s i t i o n r a t e s and, t h e r e f o r e , m i n e r a l i z a t i o n of o t h e r 30 n u t r i e n t s from o r g a n i c m a t t e r . Adding n i t r o g e n may a l s o cause d e c r e a s e d r a t e s of n i t r o g e n f i x a t i o n , and a l t e r the i o n i c c o m p o s i t i o n and pH of the i n t e r s t i t i a l water. A l s o , changes i n v e g e t a t i o n c o m p o s i t i o n may not r e f l e c t the responses of d i f f e r e n t s p e c i e s t o n u t r i e n t s b u t , r a t h e r , the e f f e c t of i n t e r a c t i o n s between the component s p e c i e s when n u t r i e n t s a r e added ( J e f f e r i e s & P e r k i n s 1977). There i s a need t o e v a l u a t e the n i t r o g e n n u t r i t i o n of s a l t marsh p l a n t s w i t h o u t a l t e r i n g the sediment environment. Smart & Barko (1980) d i d t h i s by comparing the c o n c e n t r a t i o n of a n u t r i e n t i n p l a n t t i s s u e w i t h the minimum r e q u i r e d c o n c e n t r a t i o n of t h a t n u t r i e n t . U s i n g t h a t method, they d e t e r m i n e d t h a t the p r o d u c t i v i t y of D i s t i c h l i s and S p a r t i n a was l i m i t e d by n i t r o g e n but not by phosphorus. S u l p h u r may have an i m p o r t a n t i n f l u e n c e on p l a n t z o n a t i o n . I n g o l d & H a v i l l (1984), and H a v i l l e t a l ^ (1985) found t h a t s o i l s u l p h i d e was p r e s e n t o n l y i n low marsh a r e a s , and o n l y S a l i c o r n i a europaea was r o o t e d i n s u l p h i d e - c o n t a i n i n g s e d i m e n t s . In l i q u i d media, growth of a l l s p e c i e s except S a l i c o r n i a was i n h i b i t e d by s u l p h i d e . That s u g g e s t s t h a t S a l i c o r n i a i s p r e s e n t i n low a r e a s because i t i s the o n l y p l a n t t h a t can t o l e r a t e s u l p h i d e . S u l p h i d e i s produced i n an a n a e r o b i c environment where t h e r e i s a s u p p l y of s u l p h a t e and o r g a n i c m a t t e r adequate f o r s u l p h a t e - r e d u c i n g b a c t e r i a of the genus D e s u l p h i v i b r i o . S u l p h i d e i s r e l e a s e d i n t o the s o i l where i t may be taken up w i t h i r o n , c o p p e r , or manganese r e s u l t i n g i n the f o r m a t i o n of i n s o l u b l e i n o r g a n i c s u l p h i d e s . Some s u l p h i d e remains i n 31 s o l u t i o n as hydrogen s u l p h i d e . Cooper (1984) and S i n g e r & H a v i l l (1985) found no c o r r e l a t i o n between manganese and the p o s i t i o n of p l a n t s on the marsh, a l t h o u g h s p e c i e s responded d i f f e r e n t l y t o manganese a d d i t i o n . Rozema e_t a l . (1985b) d e t e r m i n e d t h a t s p e c i e s of the low marsh were more t o l e r a n t of h i g h l e v e l s of i r o n and manganese than s p e c i e s of the h i g h marsh. However, c a u t i o n i s needed i n i n t e r p r e t i n g t h o s e r e s u l t s , as i r o n and manganese t o x i c i t y depends on s o i l f a c t o r s , such as o v e r a l l n u t r i e n t s t a t u s and the p r e s e nce of s u l p h i d e . P r e c i p i t a t i o n of i r o n s u l p h i d e p r e v e n t s h i g h l e v e l s of s o l u b l e i r o n . 2.3.4 SALINITY S o i l s a l i n i t y a f f e c t s p l a n t s i n two main ways (Hayward & B e r n s t e i n 1958). F i r s t , the i n c r e a s e d osmotic c o n c e n t r a t i o n of s a l i n e s o i l s o l u t i o n s r e s t r i c t s the uptake of water by p l a n t r o o t s , and second, p l a n t s a b s o r b the c o n s t i t u e n t i o n s of the s a l i n e s o l u t i o n which may r e s u l t i n the t o x i c a c c u m u l a t i o n of a c e r t a i n i o n , or d e c r e a s e d a b s o r p t i o n of some e s s e n t i a l n u t r i e n t . The s a l i n i t y problem i s c o u n t e r a c t e d by the f o l l o w i n g s t r a t e g i e s (Chapman 1942; Stewart & Lee 1974; A l b e r t 1975): 1. S u c c u l e n c e - an i n c r e a s e i n sodium uptake i s compensated f o r by a c o r r e s p o n d i n g d i l u t i o n of the i n t e r n a l s o l u t i o n ( e . g . S a l i c o r n i a , P l a n t a g o m a r i t i m a , S p e r q u l a r i a ). 32 2. ^ — S a l t e x c r e t i o n mechanisms (e.g. D i s t i c h l i s s p i c a t a ,  Glaux m a r i t i m a ). 3. A c c u m u l a t i o n of p r o l i n e as a s o l u t e f o r i n t r a c e l l u l a r a djustment ( e . g . T r i g l o c h i n maritimum ). 4. Shedding of o l d s a l t - s a t u r a t e d l e a v e s (e.g. T r i g l o c h i n  maritimum, P l a n t a g o m a r i t i m a ). S o i l s a l i n i t y i s i n f l u e n c e d by a l a r g e number of f a c t o r s which i n c l u d e h e i g h t of t i d e , r a i n f a l l , e l e v a t i o n , p r o x i m i t y of c r e e k s and d r a i n a g e c o n d i t i o n s , s o i l t e x t u r e , p r e s ence or absence of v e g e t a t i o n , s l o p e of ground, d i s t a n c e from the s e a , r e l a t i o n of marsh t o major i n f l o w of f r e s h w a t e r , depth of s o i l water t a b l e , depth of s u b s u r f a c e s a l t d e p o s i t s , t e m p e r a t u r e , e v a p o r a t i o n and o r g a n i c matter (Chapman 1939, 1974; MacDonald 1977). There i s some debate as t o whether s o i l water s a l i n i t y i s more important---"•than t i d a l water s a l i n i t y i n d e t e r m i n i n g p l a n t s p e c i e s c o m p o s i t i o n and d i s t r i b u t i o n i n a marsh. J e f f e r s o n (1975) found t h a t s o i l s a l i n i t y was e q u a l t o t i d a l water s a l i n i t y but c o n s i d e r e d s o i l s a l i n i t y t o be more i m p o r t a n t because the r o o t s of p l a n t s a re c o n s t a n t l y immersed i n s o i l water w h i l e t i d a l submersion i s not c o n t i n u o u s . Two a r e a s c o u l d have d i f f e r e n t s o i l and water s a l i n i t i e s i f the s o i l t e x t u r e and/or e l e v a t i o n was d i f f e r e n t . Dawe & White (1982) c o n s i d e r e d t i d a l water s a l i n i t y t o be more i m p o r t a n t than s o i l s a l i n i t y i n d e t e r m i n i n g the s p e c i e s c o m p o s i t i o n of a marsh. S a l i n i t y reduces s o i l s t r u c t u r e (which d e c r e a s e s s o i l s t a b i l i t y ) and i n f l u e n c e s p l a n t n u t r i t i o n ( Z e d l e r 1977; 33 G a l l a g h e r 1980). A h i g h s a l i n i t y was found t o reduce the c a l c i u m and p o t a s s i u m c o n t e n t of l e a v e s of S p e r g u l a r i a (Okusanya & Ungar 1984). B e e f t i n k (1977) found t h a t p o t a s s i u m d e c r e a s e d w i t h d e c r e a s i n g s a l i n i t y because seawater has more p o t a s s i u m than f r e s h water. P l a n t s o f t e n have the h i g h e s t growth r a t e i n the l o w e s t s a l i n i t y (Barbour & D a v i s 1969; Smart & Barko 1980). C e r t a i n e c o t y p e s a r e more r e s i s t a n t than o t h e r s t o h i g h s a l i n i t i e s . For example, s a l t marsh p o p u l a t i o n s of A q r o s t i s s t o l o n i f e r a and F e s t u c a r u b r a grew b e t t e r i n s a l i n e c o n d i t i o n s ( i . e . e x h i b i t e d l e s s d e c r e a s e i n y i e l d ) than i n l a n d p o p u l a t i o n s (Hannon & Bradshaw 1968; T i k u & Snaydon.1971; Ahmad & Wainwright 1977; Ahmad e t a l . 1981). Most s p e c i e s r e q u i r e d i l u t i o n f o r g e r m i n a t i o n and p l a n t s u s u a l l y g erminate i n e a r l y s p r i n g when the s a l i n i t y i s low (Chapman 1939; P i g o t t 1969; Macke & Ungar 1971). S a l i n i t y i s e s p e c i a l l y i m p o r t a n t i n s o u t h e r n C a l i f o r n i a where t h e r e i s v e r y l i t t l e r a i n f a l l i n summer. E v a p o r a t i o n i s g r e a t e r than p r e c i p i t a t i o n , and h y p e r s a l i n e c o n d i t i o n s can o c c u r (Macdonald 1977). 2.3.5 ORGANIC MATTER Or g a n i c matter i s d e r i v e d from a number of s o u r c e s . P l a n t s on the marsh p r o v i d e shoot l i t t e r t o the s o i l s u r f a c e w h i l e 34 r o o t s and rhizomes add o r g a n i c m a t t e r from below the s u r f a c e . O r g a n i c m a t t e r i s a l s o t r a n s p o r t e d i n t o the e s t u a r i n e system from c o a s t a l waters and r i v e r s (Long & Mason 1983). Most carbon e n t e r s an e s t u a r y v i a the r i v e r ( H e i n l e & Flemer 1976; Naiman & S i b e r t 1978, 1979). Low r i v e r d i s c h a r g e r e l a t i v e t o b a s i n volume, low l e v e l s of t i d a l f l u s h i n g , and the p r e s e n c e of r e t e n t i o n s t r u c t u r e s such as woody d e b r i s , o y s t e r beds, and macroalgae w i l l t e n d t o i n c r e a s e the d e p o s i t i o n of o r g a n i c matter from r i v e r s (Naiman & S i b e r t 1978). O r g a n i c matter i s i m p o r t a n t f o r the r e t e n t i o n of water by the s o i l ; f o r p r o v i d i n g i o n exchange s i t e s which f u n c t i o n as r e s e r v e s of n i t r o g e n , phosphorus and o t h e r n u t r i e n t s ; and f o r s t a b i l i z i n g the s o i l s t r u c t u r e and f a c i l i t a t i n g s o i l a e r a t i o n (Broadbent 1965; A l l i s o n 1965). 2.3.6 SOIL pH The pH of the s o i l s o l u t i o n a f f e c t s p l a n t n u t r i t i o n by a l t e r i n g the a v a i l a b i l i t y of i o n s p e c i e s i n the s o i l medium. For example, the uptake of phosphorus i s h i g h e r i n a c i d s o l u t i o n because phosphorus i s a v a i l a b l e as H2.P0 4. W i t h low pH, fewer c a t i o n s but more a n i o n s a r e t r a n s p o r t e d i n t o p l a n t s . E f f e c t s of pH on the uptake of i o n s depend on t e m p e r a t u r e and s p e c i f i c i o n c o n c e n t r a t i o n ( W a i s e l 1972). D i s s o l v e d o r g a n i c carbon i n c r e a s e s w i t h i n c r e a s i n g pH and 35 d e c r e a s i n g redox p o t e n t i a l . S o l u b l e phosphorus, ammonium, i r o n and manganese i n c r e a s e w i t h d e c r e a s i n g redox p o t e n t i a l and d e c r e a s i n g pH (De Laune et a l . 1981). B e e f t i n k e t a l . (1977) found no c o r r e l a t i o n between pH and o t h e r s o i l c h e m i c a l p r o p e r t i e s ( c l a y , c a r b o n a t e , o r g a n i c m a t t e r , n i t r o g e n , p o t a s s i u m and p h o s p h a t e ) . Z e d l e r (1977) and Crow (1977) found no c o r r e l a t i o n between pH and s a l i n i t y . The pH of s a l t marsh s o i l i s u s u a l l y h i g h (6.0 t o 9.0) as a r e s u l t of a h i g h c a r b o n a t e c o n t e n t and f l o o d i n g w i t h i o n - r i c h marine or b r a c k i s h water ( B e e f t i n k e t a l . 1977; Rozema et a l . 1985a). S a l t marsh s o i l s w i t h no c a r b o n a t e s remain a l k a l i n e as l o n g as they a r e f l o o d e d w i t h sea water; however, t h e i r pH w i l l d r op t o v e r y low v a l u e s a f t e r the s o i l has been a e r a t e d . T h i s r e s u l t s from the o x i d a t i o n of s u l p h i d e s t o s u l p h u r i c a c i d w h i c h , i f c a r b o n a t e s a r e p r e s e n t , i s n e u t r a l i z e d t o gypsum (CaS0 4) or i r o n s u l p h a t e ( F e S 0 4 ) ( B e e f t i n k e t a l . 1977). The s o i l pH and s u l p h a t e a r e n e g a t i v e l y c o r r e l a t e d - the a c c u m u l a t i o n of s u l p h a t e i s s t i m u l a t e d by low pH ( W a i s e l 1972; Crow 1977). 2.3.7 SOIL TEXTURE F i n e - t e x t u r e d s o i l s p r o v i d e i o n exchange s i t e s which b u f f e r the s o i l a g a i n s t f l u c t u a t i o n s i n s a l i n i t y and cause g r e a t e r r e t e n t i o n of s a l t s (Long & Mason, 1983). F i n e t e x t u r e d s o i l s have a h i g h e r n u t r i e n t c o n t e n t because of a g r e a t e r c a t i o n 36 exchange c a p a c i t y ( B e e f t i n k et. a l . 1977; Smart & Barko 1978; G a l l a g h e r 1980; S e l i s k a r 1981; P o r t e r 1982; S e l i s k a r & G a l l a g h e r 1983) . Marshes on the open c o a s t l i n e may d e v e l o p on c o a r s e s u b s t r a t e s . In those a r e a s , f i n e p a r t i c l e s do not s e t t l e because of the s t r o n g water f l o w . Narrow-mouthed, deep embayments p r o v i d e more s h e l t e r a l l o w i n g d e p o s i t i o n of f i n e r p a r t i c l e s . Those c o n d i t i o n s r e s u l t i n s a l t marshes w i t h i n t r i c a t e d r a i n a g e systems on p o o r l y d r a i n e d c l a y and s i l t s o i l s ( R a n w e l l 1972). 2.3.8 WATERLOGGING Wate r l o g g e d s o i l s a r e c h a r a c t e r i z e d by low s t a b i l i t y and a n a e r o b i c c o n d i t i o n s ( R u s s e l l 1973; G a l l a g h e r 1980). W a t e r l o g g i n g a l s o causes c h e m i c a l changes as f o l l o w s : N0 3 i s reduced t o NH 4 and NH 3; C 0 2 , CO3 and HCO3 t o CH 4; SO^ t o HS" and 2.+ 2.+ UZS; Mn0 2 t o Mn and F e z 0 3 t o Fe (Ponnamperuma 1972; P a t r i c k & De Laune 1977) . Submergence of a s o i l causes a d e c r e a s e i n redox p o t e n t i a l , i n c r e a s e i n pH of a c i d i c s o i l s and d e c r e a s e i n pK of a l k a l i n e s o i l s (pH goes t o 7 ) , changes i n i o n i c s t r e n g t h and s h i f t s i n m i n e r a l e q u i l i b r i a (Ponnamperuma 1972). With w a t e r l o g g i n g , n i t r o g e n becomes l e s s a v a i l a b l e t o p l a n t s because of i n c r e a s e d d e n i t r i f i c a t i o n , whereas phosphorus 37 becomes more a v a i l a b l e (because of r e d u c t i o n of i r o n from F e P 0 4 t o Fe + P 0 4 ) ( P a t r i c k & De Laune 1977). Some s p e c i e s such as A g r o s t i s s t o l o n i f e r a a r e s t i m u l a t e d by w a t e r l o g g i n g w h i l e the y i e l d of o t h e r s p e c i e s (e.g. F e s t u c a r u b r a ) i s d e p r e s s e d ( D a v i s & S i n g h 1983). 2.3.9 COMPETITION Gray & S c o t t (1977) grew P u c e i n e l l i a , A g r o s t i s and F e s t u c a i n p a i r s i n a deWit replacement experiment w i t h v a r i o u s water l e v e l s , s o i l t y p e s and s a l i n i t i e s . P u c c i n e l l i a grew b e s t a t h i g h s a l i n i t i e s whereas A g r o s t i s grew be s t i n f r e s h water. The r e s u l t s s u g g e s t e d t h a t the d i s t r i b u t i o n p a t t e r n s of the t h r e e s p e c i e s were d e t e r m i n e d by t h e i r c o m p e t i t i v e a b i l i t i e s under v a r i o u s w a t e r l o g g i n g and s a l i n i t y c o n d i t i o n s . Barbour (1978) grew Jaumea carmosa and L o l i u m perenne s e p a r a t e l y and t o g e t h e r under d i f f e r e n t s a l i n i t y c o n d i t i o n s . L o l i u m was more a f f e c t e d by s a l i n i t y . At low s a l i n i t y the growth of Jaumea was d e p r e s s e d by L o l i u m , but a t h i g h s a l i n i t y t h e r e was no c o m p e t i t i o n e f f e c t . That s u p p o r t s the h y p o t h e s i s t h a t i n t o l e r a n t h a l o p h y t e s such as Jaumea a r e r e s t r i c t e d t o s a l t marshes because they a r e poor c o m p e t i t o r s w i t h g l y c o p h y t e s on non - s a l i n e s o i l s . S t a l t e r & Batson (1969) conducted t r a n s p l a n t e x p e r i m e n t s i n a South C a r o l i n a s a l t marsh w i t h i n f o u r v e g e t a t i o n zones. 38 S u r v i v a l and growth r a t e s s u g g ested t h a t s e v e r a l s p e c i e s c o u i d t o l e r a t e c o n d i t i o n s not found i n t h e i r u s u a l zones, e.g. S a l i c o r n i a v i r q i n i c a can grow i n h i g h e r a r e a s . They suggested t h a t s p e c i e s d i s t r i b u t i o n was l i m i t e d by c o m p e t i t i o n , s a l i n i t y and w a t e r l o g g i n g . Snow & V i n c e (1984) used t r a n s p l a n t s and pot e x p e r i m e n t s t o examine the e f f e c t s of w a t e r l o g g i n g and s o i l type on f i v e dominant marsh s p e c i e s . Pot e x p e r i m e n t s showed t h a t a l l f i v e s p e c i e s had o p t i m a l growth i n the w a t e r l o g g e d , low s a l i n i t y t r e a t m e n t . In t r a n s p l a n t e x p e r i m e n t s , growth of the t h r e e h i g h marsh s p e c i e s was i n h i b i t e d i n s a l i n e s i t e s , w h i l e P u c e i n e l l i a  n u t k a e n s i s from the low marsh d i d b e t t e r i n the h i g h marsh, and T r i g l o c h i n maritimum grew w e l l a t a l l s i t e s . That seemed t o i n d i c a t e t h a t c o m p e t i t i o n p l a y s a r o l e i n s a l t marsh z o n a t i o n . 2.3.10 OTHER ENVIRONMENTAL FACTORS J e r l i n g (1981) d e t e r m i n e d t h a t s u r v i v a l of P l a n t a g o  m a r i t i m a s e e d l i n g s was r e l a t e d t o m i c r o t o p o g r a p h y . Kemp & Cunningham (1981) found t h a t a t low l i g h t , the growth r a t e of D i s t i c h l i s s p i c a t a i n e n v i r o n m e n t a l chambers was reduced by h i g h s u b s t r a t e s a l i n i t i e s or by low t e m p e r a t u r e s , w h i l e a t h i g h l i g h t i n t e n s i t i e s , the growth r a t e s were not a f f e c t e d by temperature or s a l i n i t y . R a n w e l l (1972) reasoned t h a t wind may i n f l u e n c e p l a n t 39 h e i g h t and growth i n exposed s i t e s . Temperature f l u c t u a t i o n s i n s a l t marsh s o i l s can be extreme ( w i t h i n minutes s u b s t r a t e s u r f a c e t e m p e r a t u r e s can change from 30 t o 12 C) and t h i s may be a f a c t o r i n s p e c i e s d i s t r i b u t i o n s ( J e f f e r s o n 1975). 40 I I I . STUDY AREAS T h i r t e e n t i d a l marshes were s t u d i e d - n i n e on Vancouver I s l a n d , two on the n o r t h e r n B.C. m a i n l a n d , and two on the Queen C h a r l o t t e I s l a n d s (see F i g . 1). The s t u d y a r e a s were chosen a f t e r c o n s u l t a t i o n w i t h the B r i t i s h Columbia M i n i s t r y of Environment. A l l had h i g h v a l u e s i n the B.C. M i n i s t r y of Environment r a t i n g scheme (see Appendix A ) , r e p r e s e n t e d a s u b s t a n t i a l l a t i t u d i n a l range a l o n g the B.C. c o a s t and had not been p r e v i o u s l y s t u d i e d i n d e t a i l . P l a n t communities of the N i m p k i s h , Cluxewe and K o k i s h marshes had been d e s c r i b e d and mapped by Kennedy (1982) and the B.C. M i n i s t r y of Environment had mapped the v e g e t a t i o n and s u b s t r a t e of the K o k i s h R i v e r E s t u a r y . Most marshes were r e l a t i v e l y p r i s t i n e and a c c e s s i b l e . S i x of the a r e a s were sampled i n J u l y - August, 1985. The d a t a from seven a r e a s sampled i n J u l y - August, 1984 ( c o l l e c t e d w h i l e w o r k i n g as an a s s i s t a n t t o Dr. G.E. B r a d f i e l d ) were a l s o a v a i l a b l e f o r a n a l y s i s . The t h i r t e e n s t u d y a r e a s , l i s t e d by g e o g r a p h i c a r e a , a r e as f o l l o w s : Vancouver I s l a n d A) N o r t h e a s t c o a s t : 1. K o k i s h r i v e r e s t u a r y (1984) 2. Cluxewe r i v e r e s t u a r y (1984) 3 . N i m p k i s h r i v e r e s t u a r y (1984) 41 F i g . 1: Map of study areas B) Q u a t s i n o Sound - H o l b e r g I n l e t a r e a : 4. M a r b l e r i v e r e s t u a r y (1984) 5. Goodspeed r i v e r e s t u a r y (1984) 6. Waukwaas r i v e r e s t u a r y (1984) C) West c o a s t : 7. San J o s e f r i v e r e s t u a r y (1984) 8. Kaouk r i v e r e s t u a r y (1985) 9. Ououkinsh r i v e r e s t u a r y (1985) N o r t h e r n m a i n l a n d 10. D a l a r i v e r e s t u a r y (1985) 11. K i l d a l a r i v e r e s t u a r y (1985) Queen C h a r l o t t e I s l a n d s 12. Yakoun r i v e r e s t u a r y (1985) 13. Kumdis r i v e r e s t u a r y (1985) 43 3.1 DESCRIPTION OF STUDY AREAS 3.1.1 CLIMATE A l l the marshes f a l l i n t o the c o a s t a l Western Hemlock zone of B r i t i s h Columbia ( K r a j i n a 1965) where the c l i m a t e i s c l a s s i f i e d as equable mesothermal humid (Ackerman 1941). The c l i m a t e of c o a s t a l B r i t i s h Columbia i s s t r o n g l y d e t e r m i n e d by the e f f e c t s of the mountains and ocean. The mountains cause g r e a t e r p r e c i p i t a t i o n on t h e i r west s i d e compared t o the e a s t (Conner 1915; C h i l t o n 1980). That e f f e c t i s seen on Vancouver I s l a n d ( T able I I I ) , where the K o k i s h , N i m p k i s h and Cluxewe e s t u a r i e s , l o c a t e d on the e a s t c o a s t , have l e s s r a i n f a l l than the M a r b l e and Waukwaas e s t u a r i e s , l o c a t e d i n the c e n t e r of the i s l a n d . The Goodspeed, San J o s e f , Ououkinsh and Kaouk e s t u a r i e s , l o c a t e d on the west c o a s t , have t w i c e as much p r e c i p i t a t i o n as e s t u a r i e s on the e a s t c o a s t . The average t e m p e r a t u r e i s v e r y s i m i l a r i n a l l a r e a s on Vancouver I s l a n d . On the n o r t h e r n m a i n l a n d , the K i l d a l a and D a l a e s t u a r i e s have c o o l e r t e m p e r a t u r e s i n w i n t e r and h i g h e r t e m p e r a t u r e s i n summer than o t h e r a r e a s . Those two e s t u a r i e s are l o c a t e d a t the head of an i n l e t t h a t i s a c o n s i d e r a b l e d i s t a n c e i n l a n d ; hence, the c l i m a t e i s l e s s m a r i t i m e and more c o n t i n e n t a l (Conner 1915). 4 4 Table III: Climate of study areas (Environment Canada, 1981) Study Area (Reference station) Average temper-ature (C) Average rnaximum temper-ature (C Average rninimum temper-lature (C) Rainfall (cm) Snow (cm) Total precip-itation (cm) Kaouk, Ououkinsh (1) 8.8 12.5 5.1 327.7 4.0 331.9 Kokish, Nimpkish, Cluxewe (2) 8.5 11.6 5.4 147.4 7.85 155.5 Waukwaas, Marble (3) 8.6 12.2 4.9 184.8 7.4 192.1 Goodspeed (4) 5.4 8.2 2.7 282.4 27.2 315.7 San Josef (5) 8.7 10.9 6.3 256.6 6.39 262.9 Kildala, Dala (6) 6.3 9.7 2,8 179.4 33.1 218.0 Yakoun, Kumdis (7) 7.5 10.9 4.0 143.7 8.8 153.5 Reference Stations: 1. Quatsino and Kyuquot 5. Cape Scott 2. Alert Bay 6. Kildala 3. Coal Harbour 7. Port Clements 4. Holberg 45 Those a r e a s a l s o -have more s n o w f a l l than the o t h e r e s t u a r i e s and r a i n f a l l comparable t o mid Vancouver I s l a n d . The Yakoun and Kumdis e s t u a r i e s on the Queen C h a r l o t t e I s l a n d s have the same amount of p r e c i p i t a t i o n as e s t u a r i e s on the e a s t c o a s t of Vancouver I s l a n d but are s l i g h t l y c o o l e r s i n c e they are f a r t h e r n o r t h . 3.1.2 SALINITY No p u b l i s h e d i n f o r m a t i o n on s a l i n i t y was a v a i l a b l e f o r the s tudy a r e a s . No attempt was made t o c o l l e c t s a l i n i t y d a t a i n the f i e l d as s a l i n i t y i s so v a r i a b l e over the y e a r , and a r e a d i n g a t one p o i n t i n time i s not v e r y i n f o r m a t i v e . However, on the b a s i s of the s p e c i e s , c o m p o s i t i o n and o b s e r v a t i o n s i n the f i e l d , i t was noted t h a t the D a l a and K i l d a l a e s t u a r i e s a r e m a i n l y i n f l u e n c e d by f r e s h w a t e r . These two e s t u a r i e s a r e f e d by r i v e r s which c a r r y l a r g e volumes of water d i r e c t l y from g l a c i e r s . Both a r e l o c a t e d near the head of Douglas Chann e l , a p p r o x i m a t e l y 130 km from the o u t e r c o a s t . Because of the s t r o n g f r e s h w a t e r i n f l u e n c e , the head waters of the i n l e t f r e e z e over i n some w i n t e r s (Bunn, p e r s . comm. 1985). P l a n t s c h a r a c t e r i s t i c of low s a l i n i t y marshes ( Carex p l u r i f l o r a ,  Mentha a r v e n s i s , H i e r o c h l o e o d o r a t a , C a l l i t r i c h e s t a g n a l i s ,  L i m o s e l l a a q u a t i c a , L u p i n u s l i t t o r a l i s , L i l a e a s c i l l o i d e s , Poa  eminens and A n g e l i c a g e n u f l e x a ) a r e found t h e r e . 46 The N i m p k i s h e s t u a r y i s a l s o i n f l u e n c e d by a l a r g e volume of f r e s h water d i s c h a r g e d from the r i v e r . F r e s h water i n d i c a t o r p l a n t s p e c i e s found t h e r e a r e Carex p l u r i f l o r a , Carex obnupta,  Carex o e d e r i , Oenanthe sarmentosa, P l a n t a g o l a n c e o l a t a , Juncus  g e r a r d i , Juncus f a l c a t u s and M y r i c a g a l e . The K i l d a l a , D a l a and N i m p k i s h marshes had the l a r g e s t numbers of s p e c i e s , p r o b a b l y because of the f r e s h water e f f e c t s which a l l o w e d s p e c i e s o t h e r than h a l o p h y t e s t o e n t e r t h o s e a r e a s . The o t h e r e s t u a r i e s I examined c o n t a i n e d more e v i d e n c e of b r a c k i s h t o s a l i n e growing c o n d i t i o n s . At the Cluxewe and K o k i s h e s t u a r i e s , the dyked a r e a s were more s a l i n e than the undyked a r e a s , s i n c e d y k i n g impeded any f u r t h e r f l u s h i n g a c t i o n by the r i v e r s . I n those two marshes, s a l i n e i n d i c a t o r s p e c i e s such as S a l i c o r n i a v i r g i n i c a and T r i g l o c h i n maritimum were dominant i n s i d e the dyked a r e a s , whereas more f r e s h water i n d i c a t o r s p e c i e s such as Carex l y n g b y e i and Deschampsia  c e s p i t o s a were dominant o u t s i d e t h e dykes. 3.1.3 OTHER ENVIRONMENTAL INFORMATION The e l e v a t i o n ranges of marsh v e g e t a t i o n , numbers of p l a n t communities i d e n t i f i e d , a r e a s of marsh v e g e t a t i o n , l e n g t h s of r i v e r s , s u r f i c i a l g e o l o g y , bedrock m a t e r i a l s , l a r g e t i d a l ranges and numbers of p l a n t s p e c i e s f o r each a r e a examined are g i v e n i n T a b l e IV. 47 There a r e s i g n i f i c a n t c o r r e l a t i o n s between the e l e v a t i o n range of marsh v e g e t a t i o n and t i d a l range (r=0.52, p<0.05) and between the number of s p e c i e s and e l e v a t i o n range of marsh v e g e t a t i o n (r=0.85, p<0.05). The number of s p e c i e s was a l s o s i g n i f i c a n t l y c o r r e l a t e d t o the l e n g t h of the r i v e r (r=0.62, p<0.05). An e x p l a n a t i o n f o r t h i s i s t h a t a l o n g e r r i v e r has a l a r g e r volume of water, c r e a t i n g a more f r e s h - w a t e r i n f l u e n c e d marsh. T h i s c o u l d r e s u l t i n more s p e c i e s because the f r e s h water i n f l u e n c e would a l l o w s p e c i e s o t h e r than h a l o p h y t e s t o e n t e r the a r e a . 3.1.4 DISTURBANCE The s t u d y a r e a s were p u r p o s e l y chosen t o be as f r e e of d i s t u r b a n c e by man as p o s s i b l e . U n d i s t u r b e d a r e a s t h a t f i t the o t h e r c r i t e r i a f o r study s i t e s (no p r e v i o u s d e t a i l e d s t u d i e s , h i g h v a l u e s i n the B.C. M i n i s t r y of Environment r a t i n g scheme, r e p r e s e n t a t i v e of a l a t i t u d i n a l range a l o n g the B.C. c o a s t and a c c e s s i b l e ) were d i f f i c u l t t o f i n d and the a r e a s chosen had the l e a s t amount of d i s t u r b a n c e p o s s i b l e . The D a l a , K i l d a l a , O uoukinsh, N i m p k i s h , M a r b l e , and San J o s e f r i v e r e s t u a r i e s are r e l a t i v e l y u n d i s t u r b e d . The Kumdis e s t u a r y had h o r s e s and goats g r a z i n g on i t a t the time of s a m p l i n g . 4 8 Table IV: Environmental data for study areas Study Area Elevation range of marsh vege-tation (m) Number of plant communities Number of plant species Number of quadrats sampled Number of transects sampled Kaouk 1.8 6 26 168 11 Ououkinsh 2.1 3 28 191 12 dyked 2.2 5 24 136 10 Kokish undyked 2.1 5 20 100 8 Nimpkish 4.2 6 42 141 12 dyked 2.0 5 25 151 10 Cluxewe undyked 1.0 2 13 36 4 Waukwaas 3.7 5 32 277 17 Marble 2.6 3 34 143 9 Goodspeed .1.5 4 19 154 11 San Josef 1.8 4 18 37 3 Kildala 3.4 5 36 234 14 Dala 4.3 4 36 221 12 Yakoun 1.6 4 31 157 7 Kumdis 1.4 4 17 77 3 4 9 Table IV continued. Study Area Area of marsh vege-tation (ha) Length of river 1 (km) Surficial geology (Howes, 1981) Bedrock material (Holland, 1976 and Howes, 1981) Tidal range (m) (Fisheries and Oceans, 1985) Kaouk 43 17 fluvial sediments igneous, volcanic 5 Ououkinsh 18 13 fluvial sediments igneous, volcanic 5 dyked Kokish undyked 5 4 14 fluvial sediments igneous, volcanic 5 Nimpkish 13 97 fluvial sediments igneous, volcanic 5 dyked Cluxewe undyked Waukwaas 26 6 31 19 18 marine sediments t i l l and fluvial sediments sedimentary igneous, volcanic 5 4 Marble 20 27 t i l l sedimentary 4 Goodspeed 26 11 fluvial sediments igneous, volcanic 4 San Josef 4 25 fluvial sediments igneous, volcanic 5 Kildala 95 29 ? igneous 7 Dala 40 21 ? igneous 7 Yakoun 105 60 ? igneous, sedimentary 3 Kumdis 27 10 ? sedimentary 3 Marshall et al., 1977 and 1980, Brown et al., 1979, Brown and Musgrave, 1979 and Manzon and Marshall, 1981 50 The Goodspeed e s t u a r y , a d j a c e n t t o the town of H o l b e r g , was co v e r e d i n l o g d e b r i s . T h i s e s t u a r y has, no doubt, been a f f e c t e d i n s e v e r a l ways (e. g . l a n d f i l l , g r a v e l r e m o v a l ) . The Waukwass e s t u a r y i s l o c a t e d next t o a l a r g e copper/molybdenum mine which dumps l a r g e q u a n t i t i e s of s i l t i n t o the water. P a r t s of the e s t u a r y a r e g r a z e d by go a t s and c a t t l e . The Kaouk, Cluxewe and K o k i s h e s t u a r i e s have dykes a c r o s s them which were b u i l t f o r d i f f e r e n t purposes (see v e g e t a t i o n maps i n Appendix D). The Kaouk e s t u a r y i s c r o s s e d by a dyke c a r r y i n g a road t o the town of F a i r Harbour. Water e n t e r s and l e a v e s the dyked a r e a i n two p l a c e s - under a b r i d g e over the r i v e r and t h r o u g h a t i d a l c h a n n e l . The dyke b i s e c t i n g the K o k i s h marsh was c o n s t r u c t e d d u r i n g t h e e a r l y 1900's t o f a c i l i t a t e l o g h a n d l i n g i n the a r e a . I n t h e e a r l y 1940's, Canadian F o r e s t P r o d u c t s a c q u i r e d the l e a s e t o the s i t e , and i n 1954 the o r i g i n a l dyke was s t r e n g t h e n e d and a r a i l l i n e was i n s t a l l e d w i t h a t r e s t l e b r i d g e s p a n n i n g a s m a l l opening i n the seaward s e c t i o n of the dyke. The r a i l l i n e has now been removed and the a r e a i n s i d e the dyke i s no l o n g e r used f o r l o g s t o r a g e or h a n d l i n g . In the s p r i n g of 1985, p a r t of the dyke was removed i n hopes of i n c r e a s i n g the p r o d u c t i v i t y of t h e ar e a i n s i d e the dyke f o r f i s h and w a t e r f o w l . Three o l d dykes a r e p r e s e n t a t the Cluxewe e s t u a r y , p r o b a b l y b u i l t d u r i n g the e a r l y 1900's f o r a g r i c u l t u r a l purposes such as h a y i n g , g r a z i n g or c u l t i v a t i o n . Most of the dyked marsh i s l o c a t e d b e h i n d a beach berm which has a gap i n i t a l l o w i n g f o r t i d a l f l u s h i n g . The o n l y f r e s h water i n p u t i n t o the ar e a s 51 b e h i n d the dykes a t the Cluxewe and K o k i s h e s t u a r i e s i s from s m a l l c r e e k s and p r e c i p i t a t i o n . The Yakoun e s t u a r y had a s m a l l square (0.5 ha) dyke on i t , p r o b a b l y b u i l t f o r a g r i c u l t u r a l p u r p o s e s . 52 IV. SAMPLING METHODS AND DATA COLLECTION 4.1 VEGETATION - ELEVATION SURVEYS V e g e t a t i o n was sampled s y s t e m a t i c a l l y u s i n g 0.5 X 0.5 m q u a d r a t s p l a c e d every 5 m a l o n g t r a n s e c t s . A q u a d r a t s i z e of 0.25 m2 was chosen because i t e n a b l e d a l a r g e number of q u a d r a t s t o be sampled and p r o v i d e d d a t a s u i t a b l e f o r examining v a r i a t i o n among and w i t h i n communities. T r a n s e c t s were s u b j e c t i v e l y p l a c e d i n o r d e r t o sample the f u l l range of v e g e t a t i o n d i v e r s i t y ( b i s e c t v i s u a l l y d e t e r m i n e d v e g e t a t i o n b o u n d a r i e s ) and the e n t i r e e l e v a t i o n range. S y s t e m a t i c s a m p l i n g was chosen i n s t e a d of random samp l i n g because i t i s e a s i e r t o c a r r y out i n the f i e l d and p r o v i d e d s u i t a b l e d a t a f o r a n a l y s i n g v e g e t a t i o n environment r e l a t i o n s h i p s . (Moore et a l . 1970; G r e i g - S m i t h 1983). A l s o , because the methods of d a t a a n a l y s i s were used t o d e s c r i b e the v e g e t a t i o n , i . e . e r e c t r a t h e r than t e s t h y p o t h e s e s , random samp l i n g was not needed ( G r e i g - S m i t h 1983). At each q u a d r a t , a l l v a s c u l a r p l a n t s p e c i e s were i d e n t i f i e d and a s s i g n e d v a l u e s of 1, 2, 3, 4, or 5 d e s i g n a t i n g t h e i r o c c u r r e n c e i n one of f i v e a e r i a l coverage c l a s s e s (<5%, 6-25%, 26-50%, 51-75% and 76-100%). Nomenclature f o l l o w e d T a y l o r & MacBryde (1977). E l e v a t i o n was r e c o r d e d a t each q u a d r a t u s i n g a s u r v e y o r ' s l e v e l and r e l a t e d t o c h a r t datum based on a p r e d i c t e d 53 t i d a l h e i g h t model. The numbers of t r a n s e c t s and q u a d r a t s i n each e s t u a r y a r e g i v e n i n T a b l e IV. P l a n t specimens c o l l e c t e d f o r s p e c i e s i d e n t i f i c a t i o n were d e p o s i t e d i n the U n i v e r s i t y of B r i t i s h Columbia h e r b a r i u m ( e s t u a r i e s sampled i n 1984) and the B r i t i s h Columbia P r o v i n c i a l Museum h e r b a r i u m ( e s t u a r i e s sampled i n 1985). 4.2 MAPPING OF PLANT COMMUNITIES A f i e l d c l a s s i f i c a t i o n f o r the purpose of mapping v e g e t a t i o n f o r management was d e r i v e d i n the f i e l d by v i s u a l l y r e c o g n i z i n g zones t h a t c o u l d be d i s t i n g u i s h e d on a i r photos ( s c a l e 1:5000). The community-types were checked i n subsequent a n a l y s e s of the v e g e t a t i o n d a t a u s i n g p r i n c i p a l components a n a l y s i s , and minor m o d i f i c a t i o n s were made. The B r i t i s h Columbia M i n i s t r y of Environment c l a s s i f i c a t i o n system was a l s o used f o r comparison (Hunter e t a l . 1983). That r e p r e s e n t e d a g e n e r a l i z a t i o n of the f i e l d c l a s s i f i c a t i o n i n which some community-types (fo u n d a t the same e l e v a t i o n ) were combined t o g i v e a r e a s of low, i n t e r m e d i a t e and h i g h marsh. 54 4.3 SOIL SURVEYS AND ANALYSIS For the purpose of examining v e g e t a t i o n - s o i l s - e l e v a t i o n r e l a t i o n s h i p s , s o i l samples were c o l l e c t e d from marshes a t the D a l a and Yakoun R i v e r e s t u a r i e s . Those two marshes were chosen as they appeared t o have d i f f e r e n t v e g e t a t i o n - environment r e l a t i o n s h i p s i n the f i e l d and were l o c a t e d i n d i f f e r e n t g e o g r a p h i c a r e a s . The Yakoun R i v e r , on the Queen C h a r l o t t e I s l a n d s , has a s a l t - t o - b r a c k i s h marsh d r a i n i n g a l o w l a n d , boggy a r e a w i t h s t r o n g v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s and d i s t i n c t communities. The D a l a R i v e r marsh, on the n o r t h e r n m a i n l a n d , i s a f r e s h - t o - b r a c k i s h marsh w i t h a r i v e r d r a i n i n g a g l a c i e r . The v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s i n t h i s marsh ar e not as s t r o n g and the communities a r e not as d i s t i n c t . Three t o e i g h t s o i l samples each w e i g h i n g a p p r o x i m a t e l y 2 kg were randomly c o l l e c t e d w i t h i n each community-type. S o i l samples were a n a l y z e d by the B.C. M i n i s t r y of A g r i c u l t u r e S o i l s L a b o r a t o r y i n Kelowna, B.C. S o i l s were a n a l y z e d f o r t e x t u r e (hydrometer method), pH (CaCl;>), exchangeable Ca, Mg, K and Na, s o l u b l e Ca, Mg, K, and Na and c a t i o n exchange c a p a c i t y ( s p e c t r o p h o t o m e t r y ) , carbon (Leco or W a l k l e y - B l a c k method), t o t a l n i t r o g e n ( K j e l d a h l method), a v a i l a b l e phosphorus ( c o l o r i m e t r y ) and e l e c t r i c a l c o n d u c t i v i t y ( p o t e n t i o m e t r y and 1:2 s o i l : s o l u t i o n r a t i o ) . Some s o i l samples had a v e r y h i g h o r g a n i c m a t t e r c o n t e n t and s o i l t e x t u r e c o u l d not be a n a l y z e d i n those samples. 55 V. DATA ANALYSIS 5.1 PLANT SPECIES COMPOSITION Marshes were compared based on a p u r e l y s u b j e c t i v e e x a m i n a t i o n of s p e c i e s f r e q u e n c i e s ( c a l c u l a t e d as the p e r c e n t of q u a d r a t s o c c u p i e d ) , and on c o r r e l a t i o n s between s p e c i e s and PCA axes I - I I I from a n a l y s e s of the s p e c i e s coverage d a t a . The c o r r e l a t i o n s showed the importance of each s p e c i e s i n a c c o u n t i n g f o r o v e r a l l v a r i a t i o n i n the s e p a r a t e marshes. 5.2 VEGETATION - ELEVATION RELATIONSHIPS V e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s were examined i n t h r e e ways. F i r s t , p r i n c i p a l components a n a l y s i s and c a n o n i c a l c o r r e l a t i o n a n a l y s i s were used t o i n v e s t i g a t e the r e l a t i o n s h i p s between c o n t i n u o u s c o m p o s i t i o n a l changes i n the v e g e t a t i o n and e l e v a t i o n . Second, m u l t i v a r i a t e a n a l y s i s of v a r i a n c e was used t o examine how w e l l the communities r e c o g n i z e d were s e p a r a t e d e l e v a t i o n a l l y . T h i r d , graphs of the p e r c e n t c o v e r of i n d i v i d u a l s p e c i e s v e r s u s e l e v a t i o n were p l o t t e d . To i n v e s t i g a t e c o n t i n u o u s c o m p o s i t i o n a l changes i n the 56 v e g e t a t i o n , the raw v e g e t a t i o n d a t a from each marsh c o n s i s t i n g of s p e c i e s p e r c e n t c o v e r i n one of f i v e c o v e r c l a s s e s were s u b j e c t e d t o PCA. T h i s c r e a t e d a reduced number of u n c o r r e l a t e d v a r i a b l e s t h a t summarized the main t r e n d s of v e g e t a t i o n v a r i a t i o n w i t h i n each marsh. A t o t a l of f i f t e e n PCA's were performed. The dyked and undyked p o r t i o n s of the Cluxewe and K o k i s h marshes were a n a l y z e d s e p a r a t e l y because they had p r e v i o u s l y been found t o be d i f f e r e n t ( B r a d f i e l d and Campbell 1986). C a n o n i c a l c o r r e l a t i o n a n a l y s i s (CCA) i s a method of d e t e r m i n i n g the o v e r a l l l i n e a r c o r r e l a t i o n between two s e t s of v a r i a b l e s . Components i n each s e t of d a t a a r e e x t r a c t e d i n such a way t h a t the c o r r e l a t i o n between e q u i v a l e n t components of the two s e t s i s maximized. D a g n e l l e (1961) and Hughes (1961) d e s c r i b e two e a r l y a p p l i c a t i o n s of CCA t o v e g e t a t i o n d a t a . G i t t i n s (1985) p r o v i d e s a r e c e n t and comprehensive account of CCA. As a s t a t i s t i c a l method i n v o l v i n g h y p o t h e s i s t e s t i n g , CCA assumes t h a t the d a t a a r e n o r m a l l y d i s t r i b u t e d and random s a m p l i n g was used. For d e s c r i p t i v e , n o n - s t a t i s t i c a l usage, tho s e a s s u m p t i o n s may be waived ( G r e i g - S m i t h , 1983). CCA of the f i r s t t h r e e PCA axes vs e l e v a t i o n was used t o examine o v e r a l l v e g e t a t i o n - e l e v a t i o n c o r r e l a t i o n s f o r each marsh. C a n o n i c a l c o r r e l a t i o n c o e f f i c i e n t s (Rc) p r o v i d e d q u a n t i t a t i v e e x p r e s s i o n s of t h o s e r e l a t i o n s h i p s . A graph of the f i r s t c a n o n i c a l v a r i a t e of the v e g e t a t i o n vs e l e v a t i o n was a l s o produced f o r each a r e a . 57 To i n v e s t i g a t e community - e l e v a t i o n r e l a t i o n s h i p s w i t h i n each marsh, t h r e e s e p a r a t e r e g r e s s i o n a n a l y s e s of PCA I - I I I vs e l e v a t i o n were performed. That gave t h r e e v a r i a b l e s d e n o t i n g p r e d i c t e d changes i n the v e g e t a t i o n a c c o r d i n g t o e l e v a t i o n (PCA IP - H I P ) , and t h r e e r e s i d u a l v a r i a b l e s (PCA IR - I I I R ) which a c c o u n t e d f o r the v e g e t a t i o n v a r i a t i o n not p r e d i c t e d by e l e v a t i o n . The c o n c e p t u a l model used was V = E + R, where V = t o t a l v e g e t a t i o n v a r i a t i o n ( i . e . PCA I - I I I ) , E = v a r i a t i o n p r e d i c t e d by changes i n e l e v a t i o n , and R = v a r i a t i o n not acc o u n t e d f o r by e l e v a t i o n ( B r a d f i e l d & Campbell 1986). PCA I - I I I , PCA IP - H I P , and PCA IR - I I I R were then s u b j e c t e d t o t h r e e s e p a r a t e m u l t i v a r i a t e a n a l y s e s of v a r i a n c e (MANOVA's) t o examine the d i s t i n c t i v e n e s s of the communities. MANOVA i s a method of comparing groups based on more than one v a r i a b l e . The t o t a l v a r i a n c e i s p a r t i t i o n e d i n t o between-groups and w i t h i n - g r o u p s components. The W i l k s lambda t e s t compares the w i t h i n - g r o u p v a r i a n c e t o the t o t a l v a r i a n c e as f o l l o w s A=|W/T|, where the symbols denote the d e t e r m i n a n t of the r a t i o of w i t h i n - g r o u p s v a r i a n c e t o t o t a l groups v a r i a n c e ( P i m e n t a l 1979). E t a squared i s the one complement of W i l k s lambda ( °l = 1 - A ), and t h i s i s e q u i v a l e n t t o the p r o p o r t i o n of v a r i a n c e e x p l a i n e d by between-groups v a r i a n c e ( P i m e n t e l 1979). Assumptions of MANOVA f o r h y p o t h e s i s t e s t i n g a r e a normal d i s t r i b u t i o n , random s a m p l i n g , e q u a l v a r i a n c e of groups, and l i n e a r a d d i t i v e e f f e c t s of f a c t o r l e v e l s . In t h i s s t u d y , MANOVA was employed f o r d e s c r i p t i v e , n o n - s t a t i s t i c a l purposes o n l y and thes e assumptions were waived ( G r e i g - S m i t h , 1983). 58 MANOVA was u s e d t o - d e t e r m i n e t h e p r o p o r t i o n s o f v a r i a n c e a c c o u n t e d f o r by t h e p l a n t c o m m u n i t y c l a s s i f i c a t i o n s i n t h r e e s e p a r a t e c o m p o n e n t s o f t h e v e g e t a t i o n d a t a : t o t a l v e g e t a t i o n v a r i a t i o n ; v a r i a t i o n p r e d i c t e d by e l e v a t i o n ; a n d r e s i d u a l v a r i a t i o n u n r e l a t e d t o e l e v a t i o n . By c o m p a r i n g t h e r e l a t i v e a mounts o f v a r i a n c e e x p l a i n e d by t h e c l a s s i f i c a t i o n s i n t h e t h r e e s o u r c e s o f v e g e t a t i o n v a r i a t i o n , t h e s i g n i f i c a n c e o f e l e v a t i o n i n d e t e r m i n i n g t h e p l a n t c o m m u n i t i e s c o u l d be e x a m i n e d . R e l a t i o n s h i p s b e t w e e n f i v e s p e c i e s w i t h h i g h p e r c e n t c o v e r and f r e q u e n c y v a l u e s a n d e l e v a t i o n were v i s u a l l y a s s e s s e d by g r a p h i n g t h e mean s p e c i e s p e r c e n t c o v e r v a l u e v e r s u s e l e v a t i o n . The s p e c i e s s e l e c t e d were 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 ,  P o t e n t i l l a a n s e r i n a , T r i q l o c h i n m a r i t i m u m a n d E l y m u s m o l l i s . 5 . 3 COMPARISON OF PLANT COMMUNITY C L A S S I F I C A T I O N S The f i e l d c l a s s i f i c a t i o n o f p l a n t c o m m u n i t i e s d e r i v e d i n t h i s s t u d y was c o m p a r e d t o t h e B.C. M i n i s t r y o f E n v i r o n m e n t c l a s s i f i c a t i o n a p p l i e d t o t h e same d a t a . The f i r s t t h r e e PCA a x e s (a summary o f t h e v e g e t a t i o n d a t a ) were s u b j e c t e d t o MANOVA where t h e g r o u p s i n v e s t i g a t e d were t h e c o m m u n i t y - t y p e s . A c o m p a r i s o n was made b e t w e e n t h e W i l k s lambda v a l u e s f o r t h e two c l a s s i f i c a t i o n s . 59 5.4 COMPARISON OF ORDINATION METHODS AND CLUSTER ANALYSIS Two g e n e r a l s t r a t e g i e s are used t o d i s t i n g u i s h p l a n t communities and a n a l y z e v e g e t a t i o n - environment r e l a t i o n s h i p s : c l a s s i f i c a t i o n and o r d i n a t i o n (Gauch, 1982). C l a s s i f i c a t i o n i s used t o group s p e c i e s or p l o t s , whereas o r d i n a t i o n i s used t o examine r e l a t i o n s h i p s between v a r i a b l e s , such as p l o t s or s p e c i e s , t o reduce d i m e n s i o n a l i t y , and t o produce new, summary v a r i a b l e s f o r f u r t h e r a n a l y s i s . Some w i d e l y used o r d i n a t i o n methods ar e p r i n c i p a l components a n a l y s i s (PCA), p r i n c i p a l c o o r d i n a t e s a n a l y s i s (PCoA), r e c i p r o c a l a v e r a g i n g (RA) and d e t r e n d e d c o r r e s p o n d e n c e a n a l y s i s (DCA). There has been much debate i n the l i t e r a t u r e over whether t o c l a s s i f y or o r d i n a t e , and over which s p e c i f i c method of c l a s s i f i c a t i o n or o r d i n a t i o n t o use. R e c e n t l y , o r d i n a t i o n has been p r e f e r r e d s i n c e i t i s more s u i t a b l e f o r a n a l y s i n g the c o n t i n u o u s n a t u r e of v e g e t a t i o n - environment r e l a t i o n s h i p s ( A u s t i n 1985). N e v e r t h e l e s s , c l a s s i f i c a t i o n remains u s e f u l f o r management. C l a s s i f i c a t i o n and o r d i n a t i o n a r e complementary r a t h e r than a l t e r n a t i v e s t r a t e g i e s , and the d e c i s i o n of whether t o c l a s s i f y or o r d i n a t e depends on the o b j e c t i v e s of the s t u d y . In t h i s s t u d y , c l u s t e r a n a l y s i s and f o u r commonly used o r d i n a t i o n methods - PCA, PCoA, RA and DCA - were compared f o r the D a l a and Yakoun marshes u s i n g agreement w i t h the f i e l d c l a s s i f i c a t i o n as one c r i t e r i o n . A comparison between the v a r i o u s o r d i n a t i o n methods was a l s o done by i n v e s t i g a t i n g the 60 p e r c e n t v a r i a n c e e x p l a i n e d by the f i r s t t h r e e axes and the p r o p o r t i o n of v e g e t a t i o n v a r i a t i o n a c c o u n t e d f o r by the community c l a s s i f i c a t i o n . The sample s c o r e s from the d i f f e r e n t o r d i n a t i o n s were c o r r e l a t e d i n o r d e r t o determine the m a t h e m a t i c a l s i m i l a r i t i e s between the o r d i n a t i o n s . 5.4.1 CLUSTER ANALYSIS C l u s t e r a n a l y s i s i s a method of c l a s s i f i c a t i o n t h a t s t a r t s w i t h i n d i v i d u a l p l o t s and combines them t o form c l u s t e r s . The u s u a l method i s a g g l o m e r a t i v e and p o l y t h e t i c , where i n d i v i d u a l p l o t s a r e combined t o form c l u s t e r s based on a l l s p e c i e s p r e s e n t ( G r e i g - S m i t h 1983). In t h i s s t u d y , d i s t a n c e (or d i s s i m i l a r i t y ) between p l o t s was d e t e r m i n e d u s i n g E u c l i d e a n d i s t a n c e , and d i s t a n c e between c l u s t e r s was c a l c u l a t e d u s i n g the minimum v a r i a n c e method. In the minimum v a r i a n c e method, two p l o t s a r e u n i t e d whose f u s i o n y i e l d s the l e a s t i n c r e a s e i n w i t h i n - c l u s t e r d i s p e r s i o n . P r e v i o u s s t u d i e s had shown t h e s e two s t r a t e g i e s ( E u c l i d e a n d i s t a n c e and the minimum v a r i a n c e method) t o g i v e u s e f u l r e s u l t s ( P r i t c h a r d & Anderson 1971; P i m e n t e l 1979; P i e l o u 1984). 61 5.4.2 PRINCIPAL COMPONENTS ANALYSIS P r i n c i p a l components a n a l y s i s (PCA) i s an o b j e c t i v e method t h a t produces a reduced number of independent v a r i a b l e s or components, and a r r a n g e s p l o t s i n a graph. PCA was f i r s t i n t r o d u c e d f o r a n a l y s i s of v e g e t a t i o n d a t a by G o o d a l l (1954). The f i r s t component r e p r e s e n t s the g r e a t e s t amount of v a r i a t i o n e x p l a i n e d i n the v e g e t a t i o n d a t a . The second component i s o r t h o g o n a l t o the f i r s t • and a c c o u n t s f o r the second h i g h e s t amount of v a r i a t i o n i n the d a t a , and so on. In PCA, a secondary m a t r i x i s computed from the o r i g i n a l s p e c i e s by samples m a t r i x . T h i s secondary m a t r i x i s u s u a l l y e i t h e r a c o r r e l a t i o n or a c o v a r i a n c e m a t r i x . A c o r r e l a t i o n m a t r i x i s used when the v a r i a b l e s have been measured on d i f f e r e n t s c a l e s , t h u s r e q u i r i n g s t a n d a r d i z a t i o n of a l l the v a r i a b l e s t o the same s c a l e . A c o v a r i a n c e m a t r i x i s used when the v a r i a b l e s have the same s c a l e . Thus, PCA a n a l y z e s r e l a t i o n s h i p s between s p e c i e s t o d e r i v e a s p a t i a l c o n f i g u r a t i o n of the p l o t s . A ssumptions of PCA f o r h y p o t h e s i s t e s t i n g a r e t h a t the d a t a have a normal d i s t r i b u t i o n , s a m p l i n g i s random and t h e r e i s a l i n e a r r e l a t i o n s h i p between s p e c i e s abundances. However, Ga u s s i a n response c u r v e s of s p e c i e s t o e n v i r o n m e n t a l g r a d i e n t s a r e not l i n e a r and t h i s can l e a d t o a n o n l i n e a r r e l a t i o n s h i p between s p e c i e s . The end r e s u l t i s an a r c h or horseshoe e f f e c t i n the o r d i n a t i o n (Noy-Meir & A u s t i n 1970; A u s t i n & Noy-Meir 6 2 1971; B e a l s 1973; Gauch et a l . 1977; Peet 1980). In t h i s s t u d y , PCA was performed t h r e e t i m e s f o r the D a l a and Yakoun marshes, u s i n g b o t h a c o v a r i a n c e and c o r r e l a t i o n m a t r i x w i t h q u a n t i t a t i v e data ( s p e c i e s p e r c e n t c o v e r i n one of f i v e c o v e r c l a s s e s ) and presence - absence d a t a . 5.4.3 PRINCIPAL COORDINATES ANALYSIS P r i n c i p a l c o o r d i n a t e s a n a l y s i s (PCoA) i s v e r y s i m i l a r t o PCA except t h a t i t uses a d i s s i m i l a r i t y ( d i s t a n c e ) m a t r i x of samples by samples (Q a n a l y s i s ) , whereas PCA o p e r a t e s on a s p e c i e s by s p e c i e s m a t r i x (R a n a l y s i s ) . Thus, PCoA i s c o m p u t a t i o n a l l y more e f f i c i e n t than PCA when the number of v a r i a b l e s g r e a t l y exceeds the number of samples i n the d a t a . PCoA a l s o makes p o s s i b l e the a n a l y s i s of a wider range of d i s s i m i l a r i t y c o e f f i c i e n t s than i s a v a i l a b l e i n c o n v e n t i o n a l PCA. PCoA and PCA a r e prone t o the a r c h e f f e c t , but a d j u s t m e n t s can be a p p l i e d w i t h PCoA t h a t d i m i n i s h t h i s e f f e c t . For example, the " s t e p a c r o s s " method ( W i l l i a m s o n 1978) m i n i m i z e s the horseshoe e f f e c t by computing the d i s t a n c e between two samples t h a t have no s p e c i e s i n common by u s i n g o t h e r samples w i t h known d i s t a n c e s as s t e p p i n g s t o n e s . A r e l a t e d method termed f l e x i b l e s h o r t e s t p a t h adjustment was used h e r e . T h i s method r e c a l c u l a t e s the d i s t a n c e v a l u e s between p l o t s w i t h no o v e r l a p i n s p e c i e s c o m p o s i t i o n (SP-0 l e v e l ) , one or fewer 63 s p e c i e s i n common (SP-1) or two or fewer s p e c i e s i n common (SP-2) and so f o r t h b e f o r e the PCoA i s performed ( B r a d f i e l d & K e n k e l , i n p r e s s ) . In t h i s s t u d y , PCoA was done u s i n g q u a n t i t a t i v e and presence - absence d a t a and the e f f e c t of u s i n g the s h o r t e s t p a t h method a t the SP-1 l e v e l was i n v e s t i g a t e d . For the D a l a r i v e r e s t u a r y , PCoA was done on h a l f the number of p l o t s (odd numbered p l o t s ) because the computer program used c o u l d not tak e the f u l l number of p l o t s . A l s o , the s h o r t e s t p a t h method c o u l d not be performed as t h e r e were t o o many d i s s i m i l a r p l o t s i n the d a t a . Those problems d i d not a r i s e w i t h the Yakoun marsh. 5.4.4 RECIPROCAL AVERAGING The method was f i r s t i n t r o d u c e d t o o r d i n a t e v e g e t a t i o n d a t a by H i l l (1973). R e c i p r o c a l a v e r a g i n g i s v e r y s i m i l a r t o PCA except t h a t i t uses d o u b l y s t a n d a r d i z e d d a t a (by s p e c i e s and by sa m p l e s ) . L i k e PCA and PCoA, the method assumes a l i n e a r r e l a t i o n s h i p between s p e c i e s abundances and i s t h e r e f o r e prone t o the horseshoe or a r c h e f f e c t . As w e l l , the ends of the o r d i n a t i o n axes a re compressed i n r e l a t i o n t o the m i d d l e . 64 5.4.5 DETRENDED CORRESPONDENCE ANALYSIS T h i s i s a m o d i f i e d v e r s i o n of r e c i p r o c a l a v e r a g i n g proposed by H i l l (1979) and H i l l & Gauch (1980), t o m i n i m i z e the a r c h e f f e c t and the d i f f e r e n t i a l c o m p r e s s i o n of the o r d i n a t i o n axes. T h i s i s done by a d j u s t i n g the v a l u e s on the second o r d i n a t i o n a x i s . L i k e r e c i p r o c a l a v e r a g i n g , however, DCA remains s e n s i t i v e t o o u t l i e r s and d i s c o n t i n u i t i e s i n the d a t a ( H i l l & Gauch 1980). 5.5 SOILS - VEGETATION - ELEVATION RELATIONSHIPS The r e l a t i o n s h i p between the e l e v a t i o n g r a d i e n t and s o i l c h a r a c t e r i s t i c s a t the Yakoun and Dal a marshes was i n v e s t i g a t e d by c o r r e l a t i n g the s o i l s v a r i a b l e s w i t h e l e v a t i o n . P r i n c i p a l components a n a l y s i s was performed on a c o r r e l a t i o n m a t r i x of the s o i l s v a r i a b l e s from each marsh s e p a r a t e l y , t o reduce the number of v a r i a b l e s and g i v e an i n d i c a t i o n of which s o i l s v a r i a b l e s were i m p o r t a n t i n a c c o u n t i n g f o r the v a r i a t i o n i n the d a t a . The f i r s t t h r e e PCA axes were then c o r r e l a t e d t o e l e v a t i o n and the s o i l s v a r i a b l e s . In o r d e r t o i n v e s t i g a t e s o i l s - v e g e t a t i o n r e l a t i o n s h i p s i n the two s e p a r a t e marshes, a PCA of the v e g e t a t i o n data was performed. The f i r s t t h r e e PCA v e g e t a t i o n axes were then c o r r e l a t e d t o the s o i l s v a r i a b l e s and e l e v a t i o n t o g i v e an 65 i n d i c a t i o n of which e n v i r o n m e n t a l f a c t o r s were r e l a t e d t o the v e g e t a t i o n . Rank c o r r e l a t i o n s ( u s i n g Spearman's rho v a l u e ) were c a l c u l a t e d between the f i r s t t h r e e PCA axes of the v e g e t a t i o n and the s o i l s d a t a t o i n v e s t i g a t e the c o r r e l a t i o n between v e g e t a t i o n and s o i l s . A c a n o n i c a l c o r r e l a t i o n a n a l y s i s (CCA) p r o v i d e d a summary of the f i r s t t h r e e PCA v e g e t a t i o n axes based on maximum c o r r e l a t i o n w i t h e l e v a t i o n ( c a n o n i c a l v a r i a t e a x i s I ) and t h r e e s e p a r a t e rank c o r r e l a t i o n s were c a l c u l a t e d between the c a n o n i c a l v a r i a t e a x i s I and the f i r s t t h r e e PCA s o i l s a x e s . 66 V I . RESULTS AND DISCUSSION 6.1 PLANT SPECIES COMPOSITION A l l marshes examined c o n t a i n e d Deschampsia c e s p i t o s a , Carex  l y n q b y e i , P o t e n t i l i a a n s e r i n a , Glaux m a r i t i m a , Tr i g l o c h i n  maritimum and F e s t u c a r u b r a and most c o n t a i n e d A q r o s t i s  s t o l o n i f e r a , P l a n t a q o macrocarpa and Hordeum brachyantherum (See Appendices B & C ) . Other common s p e c i e s (found i n more than 8 out of 13 marshes) a r e Juncus a r c t i c u s , P l a n t a g o m a r i t i m a ,  A c h i l l e a m i l l e f o l i u m , S p e r q u l a r i a c a n a d e n s i s and T r i f o l i u m  w o r m s k j o l d i i . These s p e c i e s had h i g h f r e q u e n c y v a l u e s i n the marshes as w e l l as h i g h c o r r e l a t i o n s w i t h PCA axes I - I I I i n d i c a t i n g t h a t they a r e i m p o r t a n t i n e x p l a i n i n g v a r i a t i o n i n t h e v e g e t a t i o n d a t a . They can be c o n s i d e r e d t o be c h a r a c t e r i s t i c of n o r t h e r n B.C. c o a s t a l marshes. The marshes on the Queen C h a r l o t t e I s l a n d s and n o r t h e r n m a i n l a n d d i f f e r e d from those on Vancouver I s l a n d by the absence of C o c h l e a r i a o f f i c i n a l i s , Poa p r a t e n s i s , S a l i c o r n i a v i r g i n i c a and S i s y r i n c h i u m l i t t o r a l e . The Yakoun and Kumdis marshes on the Queen C h a r l o t t e I s l a n d s were m i s s i n g T r i f o l i u m w o r m s k j o l d i i as they a r e l o c a t e d j u s t n o r t h of i t s range ( C a l d e r & T a y l o r 1968). A p a r g i d i u m b o r e a l e was o n l y found on the Queen C h a r l o t t e I s l a n d s . The D a l a and K i l d a l a marshes on the n o r t h e r n m a i n l a n d 67 were i n f l u e n c e d by a l a r g e volume of f r e s h water from the g l a c i a l - f e d r i v e r s and had a l a r g e number of s p e c i e s not found i n the o t h e r marshes of t h i s s t u d y : C i c u t a d o u g l a s i i , Mentha  a r v e n s i s , Poa eminens, H i e r o c h l o e o d o r a t a , A n g e l i c a g e n u f l e x a ,  L u p i n u s l i t t o r a l i s , Ranunculus c y m b a l a r i a , Ga1iurn a p a r i n e , Siurn  suave, L i m o s e l l a a q u a t i c a , L i l a e a s c i l l o i d e s and C a l l i t r i c h e  s t a q n a l i s . They were m i s s i n g common s p e c i e s i n o t h e r marshes: P l a n t a g o m a r i t i m a , S p e r g u l a r i a c a n a d e n s i s and L i l a e o p s i s  o c c i d e n t a l i s . The N i m p k i s h marsh was a l s o i n f l u e n c e d by a l a r g e volume of f r e s h water and c o n t a i n e d a number of s p e c i e s not found i n the ot h e r study a r e a s : Ranunculus o c c i d e n t a l i s , Cynosurus c r i s t a t u s ,  Juncus q e r a r d i i , Juncus f a l c a t u s and C e r a s t i u m fontanum. S p e c i e s o n l y found on the Waukwaas marsh were 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 and Da n t h o n i a c a l i f o r n i c a .  G r i n d e l i a i n t e g r i f o l i a and D i s t i c h l i s s p i c a t a seem t o be c h a r a c t e r i s t i c of s o u t h e r n B.C. marshes o n l y . Kennedy (1982) noted t h a t marshes s o u t h of Courtenay on Vancouver I s l a n d were dominated by Juncus a r c t i c u s , P o t e n t i l l a a n s e r i n a , 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 q r i f o l i a whereas those n o r t h of Courtenay were dominated by Deschampsia  c e s p i t o s a , P o t e n t i l i a a n s e r i n a , T r i g l o c h i n maritimum and g r a s s e s . Other s t u d i e s of s o u t h e r n B.C. marshes a l s o show D i s t i c h l i s s p i c a t a and G r i n d e l i a i n t e g r i f o l i a t o be dominant s p e c i e s ( P a r s o n s 1975; P o r t e r 1982; H u t c h i n s o n 1982; Dawe & White 1982, 1986). Maps of p l a n t community-types (Appendix D) i n d i c a t e t h a t 68 t h e r e i s o f t e n a z o n a l sequence of p l a n t communities s p a t i a l l y s e g r e g a t e d a l o n g an e l e v a t i o n a l g r a d i e n t : a low marsh of Carex  l y n g b y e i i n f r e s h - t o - b r a c k i s h a r e a s , and S a l i c o r n i a v i r g i n i c a and T r i q l o c h i n maritimum i n more s a l i n e a r e a s ; an i n t e r m e d i a t e marsh c h a r a c t e r i z e d by Deschampsia c e s p i t o s a , Carex l y n g b y e i ,  P o t e n t i l l a a n s e r i n a and T r i g l o c h i n maritimum and an i n c r e a s e i n s p e c i e s d i v e r s i t y ; a h i g h marsh w i t h many s p e c i e s i n c l u d i n g A c h i l l i a m i l l e f o l i u m , P l a n t a g o macrocarpa, Juncus a r c t i c u s ,  Deschampsia c e s p i t o s a , Hordeurn brachyantherum and A g r o s t i s  s t o l o n i f e r a . These communities a r e s i m i l a r t o o t h e r s found a l o n g the c o a s t of B r i t i s h Columbia ( L e v i n g s & Moody 1976; Ceska 1981; van D i r e n 1982; Dawe & White 1982, 1986) but d i f f e r from communities a t the F r a s e r R i v e r (McLaren 1972; Forbes 1972; Parsons 1975; H a b i t a t Work Group 1978; P o r t e r 1982; B r a d f i e l d & P o r t e r 1982; H u t c h i n s o n 1982). At the F r a s e r R i v e r , S c i r p u s  pungens, S c i r p u s l a c u s t r i s and Typha l a t i f o l i a and i m p o r t a n t components, as w e l l as D i s t i c h l i s s p i c a t a and G r i n d e l i a  i n t e g r i f o l i a t h a t a r e found i n s o u t h e r n marshes o n l y . 6.2 VEGETATION - ELEVATION RELATIONSHIPS C a n o n i c a l c o r r e l a t i o n c o e f f i c i e n t s r e l a t i n g t o t a l v e g e t a t i o n v a r i a t i o n t o e l e v a t i o n ranged from 0.33 t o 0.94 (Ta b l e V ) . Those v a l u e s , a l o n g w i t h the graphs of c a n o n i c a l v a r i a t e a x i s I v e r s u s e l e v a t i o n ( F i g . 2 ) , i n d i c a t e t h a t i n some 69 Table V: Vegetation - elevation relationships: R-square values from l i n e a r regressions of PCA axes I - I I I on elevation. Also given are canonical correlation c o e f f i c i e n t s (Rc) r e l a t i n g t o t a l vegetation va r i a t i o n (PCA I - I I I ) to elevation. Study Area R-square values for PCA axes Rc I II I I I Kokish - undyked 0.68 0.09 0.03 0.89 Kokish - dyked 0.16 0.10 0.00 0.51 Cluxewe - undyked 0.38 0.00 0.02 0.63 Cluxewe - dyked 0.06 0.04 0.14 0.50 Nimpkish 0.21 0.02 0.02 0.50 Waukwaas 0.00 0.04 0.07 0.33 Marble 0.52 0.05 0.00 0.76 Goodspeed 0.00 0.14 0.21 0.59 San Josef 0.55 0.03 0.02 0.77 Kaouk 0.32 0.14 0.02 0.70 Ououkinsh 0.42 0.03 0.00 0.67 Yakoun 0.53 0.19 0.00 0.85 Kumdis 0.72 0.09 0.07 0.94 K i l d a l a 0.61 0.01 0.00 0.78 Dala 0.28 0.02 0.01 0.55 F i g . 2: Graphs of canonical variate axis 1 ( v e r t i c a l axis) versus elevation i n metres above chart datum (horizontal a x i s ) . Kokish - undyked i i i i i i j i i i *.t n *.« I ? i.o «.e s.t 1 4 i.i I I % » Kokish - dyked Cluxewe - undyked ~J~ I 1 1 1 1 1—I 1 I Cluxewe - dyked • . "71—7~—' 1 ' 1 ' 1 — - i ~ • • 1.1 It t.l 10 « • t t « 4 7.1 10 Nimpkish NOTE: Numbers plotted denote plant cxximunities referred to i n Appendix D. Least squares regression l i n e s through the points are also shown. 7 1 F i g . 2 continued. Waukwaas 5 I > Iff •: i ~T~ 1 1 1 1 1 1 1 1 I Marble . . ' I I 1 I | • •• I I 2.4 J 2 4 1 4 1 ) • 14 7.2 I t I I Goodspeed O i I I 24 3.2 4| 4 ( J | « 4 7.2 1.0 m San Josef ~->7 i i i 1 1 1 1 1 1 1 < • I t 2.4 J.2 40 4 | 5.1 I I 72 I I I I Ououkinsh i i i i i — i i i i i - i — i — i — i — i — i i — i i — i F i g . 2 continued. 7 2 Yakoun Kumdis "H 1 1 1 1 1 1 1—-' ' 1 ~T 1 1 1 1 1 1 1 1 1 1 08 I.S 24 3.2 4.0 48 5.6 6.4 T.2 8.0 «.8 01 1.6 M 3.2 4.0 I I 5.6 6.4 T.I 8.0 8.8 73 s a l t marshes ( e . g . K o k i s h - undyked, Yakoun and Kumdis r i v e r s ) , t h e r e i s a s t r o n g r e l a t i o n s h i p between v e g e t a t i o n and e l e v a t i o n w h i l e i n o t h e r marshes ( e . g . K o k i s h - dyked, Cluxewe - dyked, Waukwaas and Nimpkish) the r e l a t i o n s h i p i s weak. Those d i f f e r e n c e s a r e d i f f i c u l t t o e x p l a i n , a l t h o u g h i t i s i n t e r e s t i n g t o note t h a t the two a r e a s on the Queen C h a r l o t t e I s l a n d s had v e r y s t r o n g e l e v a t i o n - v e g e t a t i o n r e l a t i o n s h i p s . A l s o , a r e a s t h a t have been d i s t u r b e d ( e . g . Waukwaas, Cluxewe -dyked and K o k i s h - dyked) tended t o have poor e l e v a t i o n v e g e t a t i o n r e l a t i o n s h i p s . However, t h a t was not t r u e i n a l l c a s e s because the N i m p k i s h and D a l a e s t u a r i e s had weak v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s and tho s e a r e a s a r e r e l a t i v e l y p r i s t i n e . The R-square v a l u e s from l i n e a r r e g r e s s i o n s of PCA I - I I I on e l e v a t i o n ( T a b l e V) show t h a t i n most c a s e s PCA I has the h i g h e s t c o r r e l a t i o n w i t h e l e v a t i o n , w h i l e i n o t h e r marshes (Cluxewe - dyked, Waukwaas and Goodspeed) PCA I I I has the s t r o n g e s t c o r r e l a t i o n . Those t h r e e marshes a l l had some d i s t u r b a n c e . I t seems t h a t when u s i n g PCA, i t i s i m p o r t a n t t o i n v e s t i g a t e a t l e a s t the f i r s t t h r e e a x e s . The " t o t a l " v e g e t a t i o n v a r i a t i o n e x p l a i n e d by the community c l a s s i f i c a t i o n , was s e p a r a t e d i n t o v a r i a t i o n " p r e d i c t e d " by changes i n e l e v a t i o n , and " r e s i d u a l " v a r i a t i o n u n e x p l a i n e d by e l e v a t i o n (see T a b l e V I ) . The " t o t a l " v a r i a t i o n e x p l a i n e d by the p l a n t community c l a s s i f i c a t i o n was v e r y h i g h (0.84 - 0.99), i n d i c a t i n g t h a t the c l a s s i f i c a t i o n p r o v i d e d a good r e p r e s e n t a t i o n of the v a r i a t i o n i n the v e g e t a t i o n d a t a . 7 4 Table VX: Vegetation - elevation relationships: Proportions of vegetation v a r i a t i o n explained by the ccrrrnunity c l a s s i f i c a t i o n s and the percent variance explained by PCA I - I I I f o r the t o t a l , predicted and residual components of the data. Study Area Variation explained by the ccranunity classification Percent variation explained by PCA I - III Total 1-A Pred. 1-A Res. 1-A To t a l 1 2 Predicted Residual Kokish - undyked 0.99 0.89 0.83 72 29 43 Kokish - dyked 0.99 0.76 0.97 57 6 51 Cluxewe - undyked 0.87 0.64 0.67 79 19 60 Cluxewe - dyked 0.87 0.43 0.83 54 4 50 Nimpkish 0.84 0.50 0.72 47 5 42 Waukwaas 0.89 0.26 0.85 55 1 54 Marble 0.93 0.69 0.82 58 18 40 Goodspeed 0.90 0.32 0.82 66 7 59 San Josef 0.96 0.81 0.87 71 18 53 Kaouk 0.98 0.70 0.95 64 12 52 Ououkinsh 0.91 0.65 0.79 53 11 42 Yakoun 0.99 0.77 0.92 67 23 44 Kumdis 0.99 0.89 0.91 73 35 38 Kildala 0.97 0.77 0.88 58 19 39 Dala 0.96 0.43 0.89 57 9 48 Percent variance of vegetation data explained by PCA axes I - III. that proportion of (1) attributable to elevation, calculatecLas £r(product of the percent variance explained by the i PCA axis and the R value from linear regression of the i PCA axis on .elevation) vs. that proportion of (1) unrelated to elevation, calculated as „ £ r (product of the percent variance.explained by the i PCA axis and (1 - R )"frcm linear regression of the i PCA axis on elevation! NOTE: "Total" refers to the analysis using quadrat values on the f i r s t three axes from PCA as input variables in MANOVA; "Predicted" refers to the analysis using predicted values frcm regressions of PCA axes I - III on elevation; "Residual" refers to the analysis using residual scores from regressions of PCA axes I - III on elevation. 7 5 The v a r i a t i o n e x p l a i n e d by the community c l a s s i f i c a t i o n u s i n g " p r e d i c t e d " v a l u e s was s m a l l e r than t h a t e x p l a i n e d by the " r e s i d u a l " v a l u e s i n most c a s e s , which i n d i c a t e s t h a t f a c t o r s o t h e r than e l e v a t i o n a r e m a i n l y r e l a t e d t o p l a n t community p a t t e r n s . However, the p r e d i c t e d v a l u e s were s t i l l h i g h i n some c a s e s , which i n d i c a t e d t h a t e l e v a t i o n can be an i m p o r t a n t f a c t o r . The p r e d i c t e d v a l u e s show s i m i l a r t r e n d s t o the c a n o n i c a l c o r r e l a t i o n c o e f f i c e n t s . For example, K o k i s h undyked has a h i g h p r e d i c t e d v a l u e i n comparison t o the r e s i d u a l v a l u e and a h i g h Rc v a l u e i n d i c a t i n g t h a t t h e r e a r e s t r o n g v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s w h i l e Waukwaas has low p r e d i c t e d and Rc v a l u e s i n d i c a t i n g weak v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p s . I t i s i n t e r e s t i n g t o note t h a t the p r e d i c t e d v a l u e s were s i m i l a r t o the r e s i d u a l v a l u e s f o r the undyked a r e a s of the K o k i s h and Cluxewe marshes, whereas the r e s i d u a l v a l u e s were h i g h e r than p r e d i c t e d i n the dyked a r e a s . T h i s i n d i c a t e s t h a t the v e g e t a t i o n - e l e v a t i o n r e l a t i o n s h i p i s s t r o n g e r i n the undyked a r e a s than t h e dyked marshes (See B r a d f i e l d & Campbell 1986) . The p e r c e n t v a r i a t i o n e x p l a i n e d by the f i r s t t h r e e PCA axes u s i n g p r e d i c t e d v a l u e s was l e s s than the v a r i a t i o n e x p l a i n e d u s i n g r e s i d u a l v a l u e s ( T a b l e V I ) . That a l s o i n d i c a t e d t h a t a l t h o u g h e l e v a t i o n i s i m p o r t a n t , f a c t o r s o t h e r than e l e v a t i o n a r e m a i n l y r e l a t e d t o v e g e t a t i o n p a t t e r n s . Graphs of s p e c i e s p e r c e n t c o v e r v e r s u s e l e v a t i o n (Appendix E) i n d i c a t e d t h a t most s p e c i e s (e.g. Carex l y n q b y e i , T r i g l o c h i n  maritimum, Deschampsia c e s p i t o s a and P o t e n t i l l a a n s e r i n a ) were 76 l o c a t e d a t a l l e l e v a t i o n s t h r o u g h o u t the marsh whereas some s p e c i e s showed a ' r e s t r i c t e d d i s t r i b u t i o n w i t h r e s p e c t t o e l e v a t i o n ( e . g . Elymus m o l l i s ). Other s t u d i e s a l s o show P o t e n t i 1 1 a a n s e r i n a and T r i g l o c h i n  maritimum t o have a wide range w i t h r e s p e c t t o e l e v a t i o n and Elymus m o l l i s c o n f i n e d t o the h i g h marsh. Deschampsia c e s p i t o s a was found i n the h i g h marsh o n l y and Carex l y n g b y e i u s u a l l y i n the low marsh ( V o g l 1966; J e f f e r s o n 1975; J e f f e r i e s 1977a; D i s r a e l i & Fonda 1979; F r e n k e l e t a l . 1981; Liverman 1982; Kennedy 1982; S e l i s k a r & G a l l a g h e r 1983; V i n c e & Snow 1984). 6.3 COMPARISON OF PLANT COMMUNITY CLASSIFICATIONS The M i n i s t r y of Environment c l a s s i f i c a t i o n and the more d e t a i l e d f i e l d c l a s s i f i c a t i o n d e r i v e d i n t h i s t h e s i s a c c o u n t e d f o r s i m i l a r amounts of v a r i a t i o n i n the v e g e t a t i o n d a t a ( T a b l e V I I ) . In some c a s e s , however, the f i e l d c l a s s i f i c a t i o n e x p l a i n e d a l a r g e r amount of v a r i a t i o n . That seems t o i n d i c a t e t h a t the M i n i s t r y of Environment c l a s s i f i c a t i o n i s too g e n e r a l and c o u l d be r e v i s e d t o have more communities than low, i n t e r m e d i a t e and h i g h marsh. To improve the c l a s s i f i c a t i o n the low marsh c o u l d be d i v i d e d i n t o a low s a l i n e marsh dominated by S a l i c o r n i a  v i r g i n i c a and Tr i g l o c h i n mar itimum and a low f r e s h water -b r a c k i s h marsh dominated by Carex l y n g b y e i . 77 Table VTI: Comparison of plant comiiunity classifications: Proportions of vegetation variation explained by the British Columbia Ministry of Environment classification and a more detailed classification. Study Area B.C. Ministry of Envir-onment c l a s s i f i c a t i o n (1 -A) Detailed c l a s s i f i c a t i o n (1 -A) Kokish - undyked 0.93 0.99 Kokish - dyked 0.88 0.99 Cluxewe - undyked 0.78 0.87 Cluxewe - dyked 0.59 0.87 Nimpkish 0.73 0.84 Waukwaas 0.89 0.89 Marble 0.93 0.93 Goodspeed 0.76 0.90 San Josef 0.90 0.96 Kaouk 0.86 0.98 Ououkinsh 0.69 0.91 Yakoun 0.99 0.99 Kumdis 0.99 0.99 K i l d a l a 0.93 0.97 Dala 0.94 0.96 78 A l s o , the h i g h marsh c o u l d be d i v i d e d i n t o f r e s h and s a l i n e t y p e s . The p l a n t communities from the marshes i n t h i s s tudy (Appendix D) have a low e l e v a t i o n Carex l y n g b y e i community i n most c a s e s w h i l e a r e a s i n s i d e the dykes a t the Cluxewe and K o k i s h and a t the Waukwaas and Goodspeed marshes show the low s a l i n e T r i g l o c h i n maritimum and S a l i c o r n i a v i r q i n i c a community. Most marshes had s a l i n e - t o - b r a c k i s h h i g h marshes but the N i m p k i s h , D a l a and K i l d a l a had f r e s h - t o - b r a c k i s h h i g h marshes. Kennedy (1982) d i s t i n g u i s h e d b r a c k i s h from s a l t marshes on Vancouver I s l a n d and Dawe & White (1982, 1986), i n v e s t i g a t i n g two marshes on Vancouver I s l a n d , s t a t e d t h a t s a l i n i t y was i m p o r t a n t i n d e t e r m i n i n g the type of marsh - f r e s h , b r a c k i s h or s a l t . C a l d e r & T a y l o r (1968) r e c o g n i z e d two t y p e s on marshes on the Queen C h a r l o t t e I s l a n d s based on s a l i n i t y , and P o r t e r (1982) found f r e s h and b r a c k i s h marshes a t s i m i l a r e l e v a t i o n s a l o n g the F r a s e r R i v e r . 6.4 COMPARISON OF ORDINATION METHODS AND CLUSTER ANALYSIS The methods compared were ranked s i m i l a r l y based on r e s u l t s from a n a l y z i n g b oth the Yakoun and D a l a marshes. When the methods were compared f o r t h e i r a b i l i t i e s t o s e p a r a t e the p l a n t communities o b s e r v e d i n the f i e l d ( F i g s . 3 - 6 ) , d e t r e n d e d c o r r e s p o n d e n c e a n a l y s i s (DCA) was the o n l y method t h a t s e p a r a t e d a l l f o u r p l a n t communities f o r both marshes. . - 7 9 Fig. 3 : Cluster analysis of the Yakoun River estuary using the Minimum variance method and Euclidean distance. NOTE: Numbers on the graph, refer to plant ccmmunities given in Appendix D. 8 0 F i g . 4 : Ordination analyses f o r the Yakoun River estuary. a) P r i n c i p a l components analysis using q u a n t i t a t i v e d a t a and a covariance matrix * » * '» 4 «4 « " 4 4 4. 4 , 4 4 -4 4 4 4 , 4 » > -i 1 r-4 . 4 4 . 1 pctl 'i 10 (.1 ».j II • « II II b) P r i n c i p a l components analysis using q u a n t i t a t i v e d a t a and a correlation matrix » » » > > • > » i I 1 ' -I 1 1 1 1 •4 " I t I I I II l » NOTE: Numbers plotted denote plant communities referred to i n Appendix D. F i g . 4 continued. c) P r i n c i p a l components analysis using presence B-, absence data 4 4 4 , 4 1 I , I ' ** • » 1 1 1 1 r —i— 1 r-13 ».« ».I 4.0 F i g . 4 continued. 8 2 d) P r i n c i p a l coordinates analysis using q u a n t i t a t i v e d a t a e) P r i n c i p a l coordinates analysis using c u a n t i t a t i v e d a t a > and s h o r t e s t p a t h a d j u stment (SP=1) • > PI 4 « > F i g . 4 continued. 8 3 f ) P r i n c i p a l coordinates analysis using presence absence data . * > > 1 1 1 1 1 1 1 1 1 1 »C ••• -I J •* • -t 4 (0 I. ( I 1.2 II 11 g) P r i n c i p a l coordinates analysis using presence -absence data and s h o r t e s t p a t h a d j u stment CSP=1) > r 84 F i g . 4 continued. h) Reciprocal averaging using q u a n t i t a t i v e d a t a * • 4 i * • > » > i T 1 1 1 1 1 1 • , 1 . . -»•* pp-'j' ••« 10 • < | | ,.j , . , ^ i ) Detrended correspondence analysis using q u a n t i t a t i v e d a t a I i-l 6 ? > » I » • -i » .o ID o MI O C f i ) 85 F i g . 5: Cluster analysis of the Dala River estuary using the Mirdinum variance method and Euclidean distance. NOTE: Numbers on the graph ref e r to plant ccninunities given i n Appendix D, 8 6 F i g . 6; Ordination analyses f o r the Dala River estuary. a) P r i n c i p a l components analysis using q u a n t i t a t i v e d a t a and a covariance matrix i ' ' • » » 1 »* ' » » » » » » , ' . ' i * » r« i t « i i . a > I. -C.I? -1.1 I.It I.M I B • < b) P r i n c i p a l components analysis using q u a n t i t a t i v e d a t a and a correlation matrix 6* > » • i » * »} *• \ * • > i n i.o I It I H ..NOTE: Numbers plotted denote plant cotirajnities referred to i n Appendix D 6 continued. c) P r i n c i p a l components analysis using presence absence data 88 F i g . 6 continued d) P r i n c i p a l coordinates analysis using q u a n t i t a t i v e d a t a and half of the plots (odd numbered quadrats) r \ * * * • • i I I "I | i • . " M l " " " ^ IJ I « e) P r i n c i p a l coordinates analysis using presence -absence data and half of the plots (odd numbered quadrats) 89 F i g . 6 continued f ) Reciprocal averaging using q u a n t i t a t i v e d a t a CM"! ? 4 M * -1.0 •«.» IJ l.» 1.0 ».» »0 I I g) Detrended correspondence analysis using q u a n t i t a t i v e d a t a l-l o » » 4 4 « 7 J , ™ ? l>>*5 » * - ,» ' I , * , • »l » » • I > i i i — • — i — i i — •p.i in o J^IO o no.o fw.o 90 P r i n c i p a l components a n a l y s i s (PCA) u s i n g q u a n t i t a t i v e d a t a ( s p e c i e s p e r c e n t c o v e r i n one of f i v e c o v e r c l a s s e s ) and a c o v a r i a n c e m a t r i x and p r i n c i p a l c o o r d i n a t e s a n a l y s i s (PCoA) u s i n g q u a n t i t a t i v e d a t a s e p a r a t e d t h r e e out of f o u r communities f o r b oth marshes. For the Yakoun r i v e r , PCoA u s i n g q u a n t i t a t i v e d a t a and the s h o r t e s t p a t h method, r e c i p r o c a l a v e r a g i n g (RA) and c l u s t e r a n a l y s i s a l s o s e p a r a t e d t h r e e out of f o u r communities, w h i l e f o r the D a l a r i v e r , PCoA u s i n g p r e s ence - absence d a t a s e p a r a t e d a l l but two communities. The s e p a r a t i o n of the communities was a c h i e v e d i n 2 - d i m e n s i o n s w i t h DCA whereas the o t h e r methods r e q u i r e d t h r e e t o f o u r axes t o s e p a r a t e the communities. U s i n g the p e r c e n t v a r i a n c e e x p l a i n e d by the f i r s t t h r e e axes as a c r i t e r i o n f o r comparing o r d i n a t i o n methods, the r e s u l t s f o r the two marshes l e d t o i d e n t i c a l r a n k i n g s of the methods ( T a b l e s V I I I & I X ) . The o r d i n a t i o n methods, l i s t e d from h i g h e s t t o l o w e s t p e r c e n t v a r i a n c e e x p l a i n e d a r e : PCoA u s i n g q u a n t i t a t i v e d a t a ; PCA u s i n g q u a n t i t a t i v e d a t a and a c o v a r i a n c e m a t r i x ; PCoA w i t h q u a n t i t a t i v e d a t a and the s h o r t e s t p a t h method; PCoA w i t h p r e s ence - absence d a t a ; PCoA u s i n g presence -absence d a t a and the s h o r t e s t p a t h method; RA; PCA u s i n g q u a n t i t a t i v e d a t a and a c o r r e l a t i o n m a t r i x ; DCA; and PCA u s i n g p r e s e nce - absence d a t a . The v a r i a n c e e x p l a i n e d by the community c l a s s i f i c a t i o n s u s i n g the d i f f e r e n t o r d i n a t i o n methods ( T a b l e s V I I I & IX) was h i g h i n a l l c a s e s except f o r PCA u s i n g presence - absence d a t a . 91 Table VIII: Comparison of ordination methods for the Yakoun River estuary. Percent variance explained by the first three axes and proportions of vegetation variation explained by the community classifications. Ordination method Percent variance explained Variation explained by comvi>-unity classif-(1 -A) I II III Total I - III Principal components analysis Quantitative data Covariance matrix 35 24 8 67 0.99 Principal components analysis Quantitative data Correlation matrix 17 14 8 39 0.94 Principal components analysis Presence - absence data 11 7 6 24 0.73 Principal coordinates analysis. Quantitative data 38 21 10 69 0.99 Principal coordinates analysis. Quantitative dat Shortest path method a 43 18 4 65 0.99 Principal coordinates analysis. Presence -absence data 31 19 8 58 0.98 Principal coordinates analysis. Presence - absence data. Shortest path method 32 18 7 57 0.98 Reciprocal averaging Quantitative data 20 14 9 43 0.98 Detrended correspondence analysis Quantitative data 20 7 4 31 0.96 9 2 Table IX: Comparison of ordination methods f o r the Dala River estuary: Percent variance explained by the f i r s t three axes and proportions of vegetation va r i a t i o n explained by the community c l a s s i f i c a t i o n s . Percent variance explained Variation explained by comm-unity c l a s s i f -i c a t i o n (1 -A) Ordination method I II I I I Total I - I I I P r i n c i p a l components analysis Q u a n t i t a t i v e d a t a Covariance matrix 31 16 10 57 0.96 P r i n c i p a l components analysis Q u a n t i t a t i v e d a t a Correlation matrix 17 10 9 36 0.90 P r i n c i p a l components analysis Presence - absence data 9 6 5 20 0.87 P r i n c i p a l coordinates analysis. Q u a n t i t a t i v e d a t a 39 14 8 61 0.96 P r i n c i p a l coordinates analysis. Presence -absence data 28 13 10 51 0.95 Reciprocal averaging Q u a n t i t a t i v e d a t a 17 11 9 37 0.93 Detrended correspondence analysis Q u a n t i t a t i v e d a t a 17 10 2 29 0.92 93 I t i s i n t e r e s t i n g t o note t h a t a l t h o u g h DCA was the o n l y method t o d i s t i n g u i s h ( i n 2-dimensions) a l l the p l a n t communities r e c o g n i z e d i n the f i e l d , the f i r s t t h r e e axes e x p l a i n e d o n l y a s m a l l amount of the v a r i a t i o n i n the v e g e t a t i o n d a t a (29% f o r the D a l a marsh and 31% f o r the Yakoun marsh). PCA ( w i t h q u a n t i t a t i v e d a t a and a c o v a r i a n c e m a t r i x ) and PCoA ( q u a n t i t a t i v e data) s e p a r a t e d t h r e e out of f o u r p l a n t communities and a c c o u n t e d f o r t w i c e as much v a r i a t i o n i n the v e g e t a t i o n d a t a . The f i r s t t h r e e PCA axes e x p l a i n e d 57% of the v a r i a t i o n i n the d a t a f o r the D a l a marsh and 67% f o r the Yakoun; the t h r e e PCoA axes e x p l a i n e d 61% of t h e v a r i a n c e f o r the D a l a marsh and 69% f o r the Yakoun. C o r r e l a t i o n s between the f i r s t t h r e e axes of PCA, PCoA, RA and DCA u s i n g q u a n t i t a t i v e d a t a a r e g i v e n i n T a b l e s X & X I . The f i r s t axes a r e a l l h i g h l y c o r r e l a t e d . The second axes a r e c o r r e l a t e d f o r a l l methods except f o r DCA. T h i s i s because DCA a d j u s t s the v a l u e s of the second a x i s t o remove the a r c h e f f e c t ( H i l l & Gauch 1980). I t i s i n t e r e s t i n g t o note t h a t the second DCA a x i s was r e s p o n s i b l e f o r s e p a r a t i n g a l l the p l a n t communities. A l s o , a t the D a l a marsh, the second RA a x i s was not c o r r e l a t e d w i t h the axes of the o t h e r methods. In t h i s case the a n a l y s i s was s t r o n g l y i n f l u e n c e d by t h r e e p l o t s c o n t a i n i n g s p e c i e s not found i n any o t h e r p l o t s . The o n l y t h i r d axes t h a t were h i g h l y c o r r e l a t e d were PCA and PCoA. V a r i o u s s t u d i e s have compared the m e r i t s and f a u l t s of the c l a s s i f i c a t i o n and o r d i n a t i o n methods (See A u s t i n 1985). T a b l e X: C o r r e l a t i o n s of sample s c o r e s f o r axes I - I I I from o r d i n a t i o n s o f t h e v e g e t a t i o n d a t a , f o r the Yakoun R i v e r E s t u a r y . PCA ?CoA RA DCA I II I I I I I I I I I I II I I I I II I I I PCA I II I I I 1.0 .00 1.0 .00 .00 1.0 PCoA I II I I I -.99 -.02 .08 .00 -.99 -.06 .08 -.05 .94 1.0 .00 1.0 .00 .00 1.0 RA I II I I I -.94 -.13 .16 -.24 .90 .05 .00 -.14 -.34 .97 .12 .07 .24 -.92 .00 -.06 .15 -.30 1.0 .02 1.0 .00 .05 1.0 DCA I II I I I -.90 -.33 .13 .54 .14 -.66 .01 .06 -.07 .92 .31 .05 -.61 -.09 -.71 -.04 -.07 .00 .98 -.10 -.10 -.64 .00 .37 -.07 .12 .20 1.0 -.62 1.0 -.09 .04 1.0 NOTE: Quantitative data were used i n a l l cases. T a b l e X I : C o r r e l a t i o n s o f s a m p l e s c o r e s f o r a x e s I - I I I f r o m o r d i n a t i o n s o f t h e v e g e t a t i o n d a t a , f o r t h e D a l a R i v e r E s t u a r y . PCA PCoA RA DCA I II I I I I II I I I I II I I I I II I I I PCA I I I I I I 1.0 .00 1.0 .00 .00 1.0 PCoA I I I I I I -.95 -.27 -.01 -.24 .93 .03 .01 .04 -.98 1.0 .00 1.0 .00 .00 1.0 RA I I I I I I -.94 -.16 .00 .18 -.09 .00 -.50 .65 .41 .98 .10 .01 -.29 -.20 -.08 .35 .78 -.37 1.0 -.14 1.0 .38 -.06 1.0 DCA I II I I I -.90 -.31 -.03 .88 -.06 .11 -.49 -.06 -.80 .99 -.04 .03 -.87 -.28 -.12 .53 .11 -.78 .98 -.11 .24 -.86 .47 -.37 .52 -.10 .58 1.0 -.81 1.0 .48 -.37 1.0 NOTE: Q u a n t i t a t i v e data were used i n a l l cases. 96 C l u s t e r a n a l y s i s i s s u b j e c t i v e i n the sense t h a t c h o i c e s of parameters t o use must be made. Other problems w i t h c l u s t e r a n a l y s i s a r e t h a t the r e s u l t s may be d i f f i c u l t t o i n t e r p r e t w i t h l a r g e d a t a s e t s and many d o u b l e - z e r o matches, where s e v e r a l p l o t s have no s p e c i e s i n common, can produce groups w i t h low i n t e r n a l s i m i l a r i t y ( W i l l i a m s 1971). The problems of i n t e r p r e t a t i o n of the dendrogram and d i f f i c u l t y w i t h l a r g e d a t a s e t s were noted i n t h i s s t u d y . Problems w i t h PCA stem from the assumption of a l i n e a r r e l a t i o n s h i p between s p e c i e s abundances. That assumption i s u s u a l l y not c o r r e c t and can l e a d t o an a r c h e f f e c t i n the o r d i n a t i o n . A l t h o u g h the a r c h e f f e c t was seen i n t h i s s t u d y , PCA was s t i l l a b l e t o s e p a r a t e most of the p l a n t communities. O r l o c i ( 1966), W i e g l e b (1980) and P o r t e r (1982) a l s o found t h a t o r d i n a t i o n s u s i n g PCA gave i n t e r p r e t a b l e r e s u l t s . I n t h i s s t u d y , PCA u s i n g q u a n t i t a t i v e d a t a was more e f f e c t i v e i n s e p a r a t i n g the p l a n t communities and e x p l a i n i n g v a r i a t i o n i n the d a t a than p r esence - absence d a t a . That c o u l d be due t o the f a c t t h a t the communities o f t e n d i f f e r e d more i n the r e l a t i v e amounts of s p e c i e s p r e s e n t than i n the pr e s e n c e or absence of a p a r t i c u l a r s p e c i e s . O r l o c i (1966) found t h a t q u a n t i t a t i v e and presence - absence d a t a gave s i m i l a r r e s u l t s . B r a d f i e l d (1981) noted t h a t PCA u s i n g presence - absence, s t a n d a r d i z e d and u n s t a n d a r d i z e d d a t a gave v i s i b l y d i f f e r e n t o r d i n a t i o n s and emphasized d i f f e r e n t components of the v e g e t a t i o n . The presence - absence d a t a d e t e c t e d s p e c i e s - r i c h a r e a s , u n s t a n d a r d i z e d d a t a c o r r e s p o n d e d t o d i f f e r e n c e s i n v e g e t a t i o n age, and n o r m a l i z a t i o n 97 b y - — q u a d r a t s p l a c e d g r e a t e r emphasis on the d i s t r i b u t i o n of common s p e c i e s . The o r d i n a t i o n s were q u i t e s i m i l a r m a t h e m a t i c a l l y , as the f i r s t t h r e e axes were h i g h l y c o r r e l a t e d . PCoA i s a l s o prone t o the a r c h e f f e c t , a l t h o u g h the s h o r t e s t p a t h method has been found t o m i n i m i z e t h i s e f f e c t ( W i l l i a m s o n 1978; B r a d f i e l d & K e n k e l , i n p r e s s ) . However, i n t h i s s t u d y , PCoA w i t h and w i t h o u t the s h o r t e s t p a t h method showed the a r c h e f f e c t . T h i s r e s u l t may have been improved by u s i n g a h i g h e r S P - l e v e l . A l s o , the s h o r t e s t p a t h method s h o u l d p r o b a b l y be used w i t h a n o n - m e t r i c method such as non-metric m u l t i d i m e n s i o n a l s c a l i n g i n s t e a d of PCoA ( B r a d f i e l d & K e n k e l , i n p r e s s ) . PCoA u s i n g q u a n t i t a t i v e d a t a was s u p e r i o r t o PCoA u s i n g presence - absence d a t a . R e c i p r o c a l a v e r a g i n g i s prone t o both the a r c h e f f e c t and the c l u s t e r i n g of the p o i n t s a t the ends of the o r d i n a t i o n axes; d e t r e n d e d c o r r e s p o n d e n c e a n a l y s i s , a m o d i f i e d v e r s i o n of RA, was s p e c i f i c a l l y d e s i g n e d t o m i n i m i z e those e f f e c t s ( H i l l & Gauch 1980). For the Yakoun marsh, both RA and DCA had an a r c h e f f e c t a l t h o u g h the a r c h was l e s s pronounced f o r DCA. For the D a l a marsh, RA gave v e r y poor r e s u l t s , p o s s i b l y because of o u t l i e r s i n the d a t a ( t h r e e p l o t s had a v e r y d i f f e r e n t s p e c i e s c o m p o s i t i o n from the o t h e r s ) . Removing the o u t l i e r s from the a n a l y s i s may have improved the r e s u l t s . In t h e i r s t u d y of a F r a s e r R i v e r marsh, B r a d f i e l d & P o r t e r (1982) noted t h a t i n f r e q u e n t s p e c i e s had t o be e l i m i n a t e d from the d a t a b e f o r e a u s e f u l RA o r d i n a t i o n c o u l d be o b t a i n e d . DCA gave b e t t e r r e s u l t s than RA but was s t i l l s e n s i t i v e t o o u t l i e r s and had a s l i g h t 98 a r c h . D i f f e r e n t a u t h o r s v a r y as t o which o r d i n a t i o n method they c o n s i d e r t o be the most u s e f u l . The " b e s t " method has been r e c o g n i z e d by agreement w i t h a s i m u l a t e d d a t a s e t ; agreement w i t h the o r i g i n a l f i e l d o b s e r v a t i o n s ; or how i n t e r p r e t a b l e the r e s u l t s a r e ( e . g . how w e l l the communities a r e s e p a r a t e d ) . DCA i s u s u a l l y c o n s i d e r e d t o be the " b e s t " method f o l l o w e d by RA and then PCA. R e l a t i v e l y few s t u d i e s have compared PCoA: Clymo (1980) found t h a t PCoA gave " b e t t e r " r e s u l t s than PCA and RA, w h i l e K e n k e l & O r l o c i (1986) found t h a t RA and DCA were b e t t e r than PCoA. Most a u t h o r s c o n s i d e r RA t o be b e t t e r than PCA ( A u s t i n 1976a & b; Noy-Meir.& W h i t t a k e r 1977; Gauch e t a l . 1977; Fasham 1977; d e l M o r a l 1980; Clymo 1980; Gauch e t a l . 1981; Gauch & W h i t t a k e r 1982; P i e l o u 1984; A u s t i n 1985). P o r t e r (1982) found r e s u l t s from RA t o be e q u i v a l e n t t o PCA f o r v e g e t a t i o n d a t a and PCA t o be s u p e r i o r f o r e n v i r o n m e n t a l d a t a . Oksanen (1983) found t h a t RA was s i m i l a r t o PCA when the d a t a were s t a n d a r d i z e d . The r e s u l t s from t h i s s tudy d i f f e r . PCA s e p a r a t e d more communities than RA and e x p l a i n e d more v a r i a t i o n i n the v e g e t a t i o n d a t a . DCA i s c o n s i d e r e d t o be s u p e r i o r t o RA ( H i l l & Gauch 1980; Gauch e_t a l . 1981; Gauch 1 982; A u s t i n 1985) because i t removes the a r c h e f f e c t and compr e s s i o n of the a x i s ends. However, Oksanen (1983) found t h a t the r e s u l t s from RA were s l i g h t l y b e t t e r than DCA. K e n k e l & O r l o c i (1986) noted t h a t DCA was b e t t e r than RA i n some c a s e s but i n o t h e r s i t l e d t o a d i s t o r t i o n of the r e s u l t s . In t h i s s t u d y , DCA s e p a r a t e d the 99 p l a n t communities b e t t e r , but RA e x p l a i n e d more of the v a r i a t i o n i n the v e g e t a t i o n d a t a . K e n k e l & O r l o c i (1986) suggest t h a t none of the c u r r e n t l y a v a i l a b l e t e c h n i q u e s a r e a p p r o p r i a t e under a l l c i r c u m s t a n c e s . A number of o r d i n a t i o n methods s h o u l d be used t o a n a l y z e a g i v e n d a t a s e t ( O r l o c i 1978; Green 1979; G r e i g - S m i t h 1983) because d i f f e r e n t methods emphasize d i f f e r e n t a s p e c t s of the d a t a . In t h i s s t u d y , where the p l a n t communities are d i s t i n g u i s h e d more by the amount of a s p e c i e s than by presence - absence, PCoA and PCA u s i n g q u a n t i t a t i v e d a t a e x p l a i n e d more v a r i a t i o n i n the v e g e t a t i o n d a t a but DCA l e d t o a c l e a r e r s e p a r a t i o n of the c ommunities. 6.5 SOILS - VEGETATION - ELEVATION RELATIONSHIPS The r e s u l t s from c o r r e l a t i o n s between s o i l s v a r i a b l e s and e l e v a t i o n a t the Yakoun and D a l a marshes ( T a b l e s X I I and X I I I ) show t h a t a l a r g e number of s i g n i f i c a n t c o r r e l a t i o n s e x i s t and t h ey a r e somewhat d i f f e r e n t between the two marshes. The s o i l s - e l e v a t i o n c o r r e l a t i o n s common t o both marshes were a n e g a t i v e c o r r e l a t i o n between phosphorus and e l e v a t i o n and p o s i t i v e c o r r e l a t i o n s between e l e v a t i o n and exchangeable c a l c i u m and magnesium, c a t i o n exchange c a p a c i t y , carbon and n i t r o g e n . 100 Table XII: Correlation matrix of s o i l s variables, elevation and the f i r s t three PCA axes of a s o i l s and a vegetation ordination, for the Yakoun River estuary. E l e v - pH Ex. Ex. Ex. Ex. C.E.C. C N P E.C. S o l . a t i o n Ca Mg K Na Ca E l e v a t i o n 1.0 (171) pH (17) .26 1.0 Ex. Ca (17) .72 .08 1.0 Ex. Mg (17) .73 .04 .98 1.0 Ex. K (17) .64 .26 .86 .88 1.0 * •>(< * * Ex. Na (17) .65 -.02 .95 .98 .90 1.0 C.E.C. (17) .83 .22 .96 .96 .87 .92 1.0 * * 3k * * *, Carbon (17) .78 .14 .91 .93 .81 .88 .94 1.0 * * * * * * * Nitrogen .84 .18 .94 .94 .81 .89 .97 .94 1.0 ( 1 6 ) * * * * * * * Phosphorus -.58 .19 -.67 -.69 -.39 -.64 -.69 -.72 -.71 1.0 U 7 ) "* * * * * * E.C. (17) .32 -.37 .62 .65 .44 .70 .58 .47 .56 -.59 1.0 * ¥ S o l . Ca (17) .15 -.63 .52 .60 .36 .67 .41 .43 .42 -.48 .83 1.0 ^£ jff^ }j£ Q^E. ij^e ^ S o l . Mg (17) .33 -.48 .69 .76 .54 .81 .61 .61 .59 -.58 .86 .97 ^ ^ ^ ^ ^ ^ i*t^  ^ (^C 3ft: S o l . Na (17) .54 -.18 .84 .91 .77 .95 .81 .83 .79 -.63 .73 .80 S o l . K (17) .57 -.02 .88 .92 .84 .93 .83 .82 .81 -.51 .54 .62 Sand (6) .81 -.54 -.10 -.40 -.77 -.43 -.72 -.01 .22 .38 .03 .39 S i l t (6) -.85 .64 .06 .31 .76 .30 .60 -.15 -.39 -.21 -.13 -.44 Clay (6) -.08 -.26 .19 .52 .28 .69 .76 .73 .69 -.85 .43 .14 PCA I .78 .29 .84 .87 .83 .87 .90 .80 .92 -.62 .55 .40 Veg. (17) PCA II .31 .45 .19 .11 .17 -.04 .23 .28 .20 -.07 -.37 -.50 Veg. (17) PCA I I I .12 .14 -.11 -.11 -.06 -.12 -.07 -.12 -.04 .42 -.17 -.13 Veg. (17) * * * * * * * * PCA.I -.02 -.79 .83 .85 -.44 .84 .50 .92 .88 -.86 .97 .84 S o i l s (6) PCA II -.78 .55 -.00 .42 .86 .51 .84 .16 -.11 -.45 -.04 -.46 S o i l s (6) PCA I I I -.48 .06 .56 .25 -.21 .02 -.16 -.32 -.41 .14 .20 .26 S o i l s (6) 'sample s i z e * s i g n i f i c a n t at p±0.05 101 T a b l e X I I c o n t i n u e d : Vegetation S o i l s S o l . S o l . S o l . Sand S i l t Clay PCA PCA. PCA PCA PCA PCA Mg Na K 1 II I I I I i i i n S o l . Mg 1.0 S o l . Na .91 1.0 S o l . K .76* .92* 1.0 Sand .36 -.19 .54 1.0 S i l t -.43 .07 -.67 -.98 1.0 Clay .19 .56 .36 -.42 .23 1.0 * * * * * PCA I .57 .78 .7? -.93 .88 .55 1.0 Veg. PCA I I -.37 -.19 -.01 .58 -.44 -.80 .00 1.0 Veg. PCA I I I -.13 -.12 -.05 .47 -.33 -.79 .00 .00 1.0 Veg. * •* * PCA I .88 .95 .83 .04 -.17 .58 -.07 -.76 -.29 1.0 S o i l s *• * * PCA II -.41 .21 -.42 -.95 .90 .53 .95 -.61 -.66 .00 1.0 S o i l s PCA. III. .23 .16 -.33 -.24 .37 -.52 -.04 .00 .55 .00 .00 1.0 S o i l s T a b l e X I I I : 102 Correlation matrix of s o i l s variables, elevation and the f i r s t three PCA axes of a s o i l s and a vegetation ordination, for the Dala River estuary. E l e v - pH Ex. Ex. Ex. Ex. C.E.C. C N P E.C. S o l . a t i o n Ca Mg K Na Ca E l e v a t i o n 1.0 (201 pH (20) .15 1.0 Ex. Ca (20) .71 .14 1.0 Ex. Mg (20) .69* .18 .98* 1.0 Ex. K (20) .21 -.46* .52* .53* 1.0 * *. * Ex. Na (20) .40 .05 .59 .66 .57 1.0 * * * *- * C.E.C. (20) .69 .25 .88 .90 .52 .72 1.0 *fc # H V ^ Carbon (20) .71 .15 .84 .85 .56 .74 .93 1.0 ¥ y nf ^ y Nitrogen .66 .25 .87 .91 .56 .78 .96 .95 1.0 ( 2 0 ) * * Phosphorus -.48 .16 -.34 -.36 -.13 -.46 -.38 -.36 -.35 1.0 (20) E.C. (20) .08 -.90 .15 .14 .68 .31 .13 .20 .13 -.36 1.0 S o l . Ca (20) -.05 -.96 -.03 -.08 .53 .11 -.13 -.01 -.12 -.29 .92 1.0 * * * * S o l . Mg (20) -.04 -.96 -.01 -.05 .55 .11 -.10 .01 -.10 -.29 .94 .99 #. * * * *e S o l . Na (20) .22 -.62 .33 .32 .72 .56 .35 .42 .39 -.40 .78 .75 % -ft- ^ S o l . K (20) -.06 -.94 .00 -.04 .65 .12 -.07 .05 -.06 -.20 .93 .98 Sand C13) -.18 -.14 -.42 -.37 -.04 -.42 -.50 -.26 -.30 .51 -.24 -.02 S i l t (13) .14 .08 .37 .32 .10 .41 .45 .24 .26 -.53 .30 .08 Clay (13) .29 .31 .52 .45 -.16 .35 .57 .28 .38 -.34 .01 -.21 * * * * # • » • * * * PCA I .59 .61 .60 .60 -.12 .42 .62 .57 .61 -.51 -.36 -.45 Veg. (.20) PCA II .16 .02 .28 .29 .04 -.28 .15 .15 .15 .32 -.08 -.16 Veq. (20) PCA I I I -.07 .15 -.09 .04 '.19 .49 .22 .22 .27 -.03 .02 -.18 PCA I .70 .65 .82 .83 .11 .81 .93 .85 .91 -.38 -.20 -.49 S o i l s U 3 ) * * * PCA II -26 -.73 .26 .30 .56 .10 .25 .20 .13 -.46 .96 .85 S o i l s (13) PCA I I I -29 -.03 .22 .28 .58 .04 .15 .40 .33 .42 -.05 -.03 S o i l s (13) Sample s i z e ^ s i g n i f i c a n t at pi0.05 Table XIII continued: 1 0 3 Vegetation S o i l s S o l . S o l . S o l . Sand S i l t Clay PCA PCA. PCA PCA PCA PCA M 3 Na K i n m x Z I 1 X 1 S o l . Mg 1.0 S o l . Na .75 1.0 * * S o l . K .98 .77 1.0 Sand -.04 - .15 .06 1.0 * S i l t .10 .16 .03 -.99 1.0 Clay -.17 .09 -.34 -.85 .75* 1.0 * PCA I -.45 - .10 -.49 -.44 .40 .49 1.0 Veg. PCA I I -.13 - .26 -.10 .54 - .51 - .54 .00 1.0 Veg. PCA I I I -.18 .05 -.12 .26 - .20 - .44 .00 .00 1.0 Veg. * PCA I -.47 .23 -.52 -.56 .51 .64 .83 -.33 -.06 1.0 S o i l s * PCA I I .87 .49 .82 -.37 .42 .12 - .13 .12 .02 .00 S o i l s * * PCA I I I -.03 .17 .17 .68 - .66 - .60 - .14 .57 .23 .00 S o i l s 104 In a d d i t i o n , e l e v a t i o n at the Yakoun marsh was p o s i t i v e l y c o r r e l a t e d t o exchangeable p o t a s s i u m and sodium, s o l u b l e p o t a s s i u m and sodium and sand, and n e g a t i v e l y c o r r e l a t e d t o s i l t . The p o s i t i v e c o r r e l a t i o n between e l e v a t i o n and c a r b o n , n i t r o g e n and p o t a s s i u m agrees w i t h most s t u d i e s (see T a b l e I I ) . The n e g a t i v e c o r r e l a t i o n between e l e v a t i o n and phosphorus i s o p p o s i t e t o o t h e r s t u d i e s ( R a n w e l l 1964; P i g o t t 1969; Gray & Bunce 1972) as i s the c o r r e l a t i o n between e l e v a t i o n and c a l c i u m , magnesium and and sodium ( R a n w e l l 1964; Stephens & B i l l i n g s 1967). No p u b l i s h e d s t u d i e s were found on the c o r r e l a t i o n between c a t i o n exchange c a p a c i t y and e l e v a t i o n . The p o s i t i v e c o r r e l a t i o n t o sand and n e g a t i v e c o r r e l a t i o n t o s i l t a g r e e s w i t h some s t u d i e s (MacDonald 1977; Z e d l e r 1982; P o r t e r 1982) but d i f f e r s from o t h e r s (Gray & Bunce 1972; Crow 1977; D i s r a e l i & Fonda 1979; H u t c h i n s o n 1982). I t seems t h a t the o n l y g e n e r a l i z a t i o n s f o r s o i l s - e l e v a t i o n r e l a t i o n s h i p s based on t h i s and o t h e r s t u d i e s are t h a t the amounts of c a r b o n , n i t r o g e n and p o t a s s i u m i n c r e a s e w i t h i n c r e a s i n g e l e v a t i o n . The r e s u l t s of a p r i n c i p a l components a n a l y s i s of the s o i l s v a r i a b l e s f o r each marsh are g i v e n i n F i g u r e s 7 and 8. The f i r s t two PCA axes a c c o u n t e d f o r 88% of the v a r i a t i o n i n the d a t a f o r the Yakoun marsh and 70% f o r the D a l a . In both marshes, community 1 (Dala) and communities 1 and 2 (Yakoun) were s e p a r a t e d from the r e s t of the communities. Those communities a r e found a t the l o w e s t e l e v a t i o n s i n the marshes, so i t seems t h a t s o i l s of the lower marsh may d i f f e r from s o i l s 105 of the upper marsh. The f a c t t h a t a l l s o i l s v a r i a b l e s were s i g n i f i c a n t l y c o r r e l a t e d t o one or more of the f i r s t t h r e e PCA axes of the v e g e t a t i o n d a t a ( T a b l e s X I I and X I I I ) suggests t h a t a l l s o i l s v a r i a b l e s s t u d i e d a r e i m p o r t a n t i n d e t e r m i n i n g v a r i a t i o n i n the v e g e t a t i o n d a t a . The rank c o r r e l a t i o n s between s o i l s and v e g e t a t i o n ( T a b l e s XIV and XV) show t h a t f o r both marshes PCA I ( s o i l s ) was c o r r e l a t e d t o CV I ( v e g e t a t i o n w i t h maximum c o r r e l a t i o n t o e l e v a t i o n ) . For the Yakoun marsh, PCA I ( s o i l s ) was c o r r e l a t e d t o PCA I I ( v e g e t a t i o n ) and PCA I I ( s o i l s ) was c o r r e l a t e d t o PCA I ( v e g e t a t i o n ) . At the D a l a marsh, PCA I ( s o i l s ) was c o r r e l a t e d t o PCA I ( v e g e t a t i o n ) . That s u g g e s t s t h a t the s o i l s v a r i a b l e s a r e i m p o r t a n t i n d e t e r m i n i n g v a r i a t i o n i n the v e g e t a t i o n d a t a . F i g . 7 : P r i n c i p a l components analysis of s o i l s variables for the Yakoun River, estuary. 106 5= - 3 . 0 i - ? 0 - 1 . 0 PCA 1 - i — 0 . 0 — 1 — 1 . 0 - 1 — 2 . 0 I — 3 . 0 — I — 4 . 0 —I— S O 6 0 NOTE: Numbers plotted denote plant cramiunities referred to in Appendix D. Dense group of quadrats at origin includes cxxnmunities 3 and 4 . F i g . 8 : P r i n c i p a l components a n a l y s i s o f s o i l s v a r i a b l e s f o r t h e D a l a R i v e r e s t u a r y . 107 5~H a? 3 2 3 3 9 2 3 1 7— I 1 1 1 1 1 1 1 I 5 . 0 -4.0 - 3 . 0 - 2 . 0 - 1 . 0 0 . 0 1 . 0 2 . 0 3 . 0 4.0 5 0 PCR 1 NOTE: Numbers plotted denote plant ccmnunities referred to i n Appendix D. Dense group of quadrats at o r i g i n includes communities 3 and 4 . Table XIV: Spearman rank correlations between PCA axes I - III of the s o i l s and vegetation data sets for the Yakoun River estuary. CV I PCA I Veg. PCA Veg. II PCA Veg. I l l PCA I - S o i l s -0. 70 -0.06 -0. 81 -0. 17 PCA II • - S o i l s 0. 03 0.90 -0. 61 -0. 55 PCA III - S o i l s 0. 35 -0.29 0. 23 0. 52 NOTE: CV I is the fi r s t canonical correlation axis of the vegetation data. Table XV: Spearman rank correlations between PCA axes I - III of the s o i l s and vegetation data sets for the Dala River estuary. CV I PCA I Veg. PCA II Veg. PCA III Veg. PCA I - S o i l s 0. 75 0.67 -0.23 -0.04 PCA II - S o i l s 0. 16 0.06 0.00 -0.08 PCA III - S o i l s -0. 06 -0.18 0.48 0.36 NOTE: CV I is the fi r s t canonical correlation axis of the vegetation data. 109 V I I . CONCLUSIONS P l a n t s c h a r a c t e r i s t i c of n o r t h e r n B.C. c o a s t a l marshes are Deschampsia c e s p i t o s a , Carex l y n g b y e i , P o t e n t i l l a a n s e r i n a ,  Glaux m a r i t i m a , T r i g l o c h i n maritimum, F e s t u c a r u b r a , A g r o s t i s  s t o l o n i f e r a , P l a n t a q o macrocarpa, Hordeum brachyantherum, Juncus  a r c t i c u s , P l a n t a q o m a r i t i m a , A c h i l l e a m i l l e f o l i u m , S p e r g u l a r i a  c a n a d e n s i s and T r i f o l i u m w o r m s k j o l d i i . The marshes on the Queen C h a r l o t t e I s l a n d s and the n o r t h e r n m a i n l a n d d i f f e r e d s l i g h t l y from each o t h e r and from those on n o r t h e r n Vancouver I s l a n d . The K i l d a l a and D a l a marshes had 12 s p e c i e s not found i n o t h e r study a r e a s , and the Ni m p k i s h marsh had 5 s p e c i e s unique t o t h a t s i t e . The n o r t h e r n B.C. marshes d i f f e r from the s o u t h e r n ones by the absence of D i s t i c h l i s s p i c a t a and G r i n d e l i a i n t e g r i f o l i a as dominant s p e c i e s . Dominant s p e c i e s found o n l y on the F r a s e r R i v e r marshes a r e S c i r p u s pungens, S c i r p u s l a c u s t r i s and Typha  l a t i f o l i a . The p l a n t communities on the n o r t h e r n marshes were s i m i l a r t o o t h e r s a l o n g the c o a s t of B.C., but d i f f e r e d from the F r a s e r R i v e r marsh communities. The r e l a t i o n s h i p between v e g e t a t i o n and e l e v a t i o n i n the marshes ranged from s t r o n g t o weak, p o s s i b l y due t o f a c t o r s a s s o c i a t e d w i t h g e o g r a p h i c l o c a t i o n or d i s t u r b a n c e . The v a r i a t i o n " p r e d i c t e d " by e l e v a t i o n was l e s s than the v a r i a t i o n u n e x p l a i n e d by e l e v a t i o n i n most c a s e s . T h i s s u g g e s t s t h a t a l t h o u g h e l e v a t i o n i s i m p o r t a n t , f a c t o r s o t h e r than e l e v a t i o n a r e m a i n l y r e l a t e d t o p l a n t community p a t t e r n s . Some s p e c i e s 110 a r e l o c a t e d ! — a t a l l e l e v a t i o n s throughout the marsh (e.g. T r i g l o c h i n maritimum, Carex l y n g b y e i , Deschampsia c e s p i t o s a and P o t e n t i l l a a n s e r i n a ) w h i l e o t h e r s a re more r e s t r i c t e d ( e.g. Elymus m o l l i s ). The p l a n t community c l a s s i f i c a t i o n s a c c o u n t e d f o r l a r g e amounts of v a r i a t i o n i n the v e g e t a t i o n d a t a i n d i c a t i n g t h a t they p r o v i d e d good r e p r e s e n t a t i o n s of v a r i a t i o n of the v e g e t a t i o n d a t a . The f i e l d c l a s s i f i c a t i o n s d e s c r i b e d i n t h i s s t u d y a c c o u n t e d f o r the same or h i g h e r amounts of v a r i a t i o n i n the v e g e t a t i o n d a t a than the B r i t i s h Columbia M i n i s t r y of Environment c l a s s i f i c a t i o n . I t i s suggested t h a t the M i n i s t r y of Environment c l a s s i f i c a t i o n be s u b d i v i d e d i n t o low s a l i n e marshes, low b r a c k i s h - f r e s h marshes, an i n t e r m e d i a t e marsh ( t r a n s i t i o n z o n e ) , a h i g h s a l i n e marsh and a h i g h f r e s h b r a c k i s h marsh. DCA s e p a r a t e d ( i n 2 - d i m e n s i o n s ) a l l the p l a n t communities r e c o g n i z e d a t the Yakoun and D a l a marshes; the o t h e r o r d i n a t i o n methods r e q u i r e d more than 2 - d i m e n s i o n s t o s e p a r a t e communities. PCA ( q u a n t i t a t i v e d a t a and a c o v a r i a n c e m a t r i x ) and PCoA ( q u a n t i t a t i v e d ata) s e p a r a t e d t h r e e out of f o u r communities i n bot h marshes s t u d i e d . The methods w i t h the h i g h e s t p e r c e n t v a r i a n c e e x p l a i n e d by the f i r s t t h r e e axes were: PCoA ( q u a n t i t a t i v e d a t a ) , PCA ( q u a n t i t a t i v e d a t a and c o v a r i a n c e m a t r i x ) , PCoA ( q u a n t i t a t i v e data and s h o r t e s t p a t h method), PCoA (presence - absence d a t a ) and PCoA (presence - absence d a t a and s h o r t e s t p a t h method). Q u a n t i t a t i v e d a t a were more e f f e c t i v e than p r e s ence - absence d a t a i n s e p a r a t i n g the p l a n t communities 111 and e x p l a i n i n g v a r i a t i o n i n the v e g e t a t i o n d a t a . At the Yakoun and D a l a marshes, e l e v a t i o n was n e g a t i v e l y c o r r e l a t e d t o phosphorus and p o s i t i v e l y c o r r e l a t e d t o exchangeable c a l c i u m and magnesium, c a t i o n exchange c a p a c i t y , carbon and n i t r o g e n . E l e v a t i o n a t the Yakoun marsh was p o s i t i v e l y c o r r e l a t e d t o exchangeable p o t a s s i u m and sodium, s o l u b l e p o t a s s i u m and sodium, and sand and n e g a t i v e l y c o r r e l a t e d t o sand. The o n l y g e n e r a l i z a t i o n s f o r s o i l s - e l e v a t i o n r e l a t i o n s h i p s based on t h i s and o t h e r s t u d i e s a r e t h a t c a r b o n , n i t r o g e n and p o t a s s i u m i n c r e a s e w i t h i n c r e a s i n g e l e v a t i o n . R e s u l t s of a p r i n c i p a l components a n a l y s i s of the s o i l s d a t a showed t h a t the s o i l s of communities l o c a t e d a t low e l e v a t i o n s seem t o d i f f e r from t h o s e of communities l o c a t e d a t h i g h e l e v a t i o n s . A l l s o i l s v a r i a b l e s were s i g n i f i c a n t l y c o r r e l a t e d t o one or more of the f i r s t t h r e e PCA a x e s ; t h u s , a l l s o i l s v a r i a b l e s examined a r e p r o b a b l y r e l a t e d t o s p e c i e s d i s t r i b u t i o n s i n a marsh. Rank c o r r e l a t i o n s between PCA axes of s o i l s and v e g e t a t i o n showed t h a t a l l s o i l s v a r i a b l e s a r e i m p o r t a n t i n d e t e r m i n i n g v a r i a t i o n i n the v e g e t a t i o n d a t a . 1 12 LITERATURE CITED Ackerman, E.A. 1941. The Koppen c l a s s i f i c a t i o n of c l i m a t e s i n N o r t h A m e r i c a . Geogr. Rev. 31:105-111. Adams, D.A. 1963. 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E s t . C o a s t . Mar. S c i . 5:39-Z e d l e r , J.B. 1982. The e c o l o g y of s o u t h e r n C a l i f o r n i a s a l t marshes: a community p r o f i l e . FWS/OBS-81/54. D i v i s i o n of B i o l o g i c a l S e r v i c e s , U.S. F i s h and W i l d l i f e S e r v i c e , Washington, D.C. Appendix A: Rating of study areas according to the British Columbia Ministry of Environment (Hunter et a l . , 1985) Study Area Water-fowl Wild-l i f e Fish Produ-ctivity Social Faotoi Ininin-ence oi ievel-spment Enhanc. Rehab. Potent-i a l Total rating Kaouk 3 1 3 3 2 2 2 16 Ououkinsh 2 1 2 3 1 1 1 11 Kokish 2 0 2 1 3 2 3 13 Nimpkish 3 0 3 2 3 1 1 13 Cluxewe 3 1 3 3 3 2 2 17 Waukwaas 3 0 3 3 3 1 2 15 Marble 2 0 3 3 2 1 1 12 Goodspeed 2 0 2 3 3 1 2 13 San Josef 2 0 2 3 2 1 1 11 Dala, Kildala 3 1 3 3 2 2 1 15 Yakoun 3 1 3 3 3 1 2 16 Kumdis Bay 3 0 2 3 2 2 1 13 ^ DEFINITION OF RATING FACTORS RESOURCE VALUE RATINGS A. Waterfowl Waterfowl (ducks, geese and swans) numbers were used as an Ind icator of waterb l rd use o f an area. C r i t e r i a fo r ra t ing each area are l i s t e d below. Ratings C r i t e r i a 1 less than 200 waterfowl recorded on each survey of area 2 200 or more waterfowl recorded on at least one survey of area 3 200 or nore waterfowl recorded on three or «ore surveys of area B. Other W i l d l i f e This r a t i n g fac tor pertains to use o f the s i te by high p r o f i l e w i l d l i f e species. Species considered were: trumpeter swans (>70); peregrine fa lcons; sandh i l l cranes; . g r i z z l y bears; Roosevelt e lk ; harbour seals; and k i l l e r whales. 0 area does not receive s i g n i f i c a n t use by any of the species l i s t e d above 1 area receives s i g n i f i c a n t use by one or more of the species above C. Fish An est imation of the Importance of the area to f i s h was based pr imar i l y on salmon escapement records. The overr id ing assumption was that wetlands were Important to the wel l -being of f i s h . Although salmon were the major species considered, herr ing spawn s i tes and s h e l l f i s h areas were also considered Important and rated appropr ia te ly . Ratings C r i t e r i a 1 salmon escapement h i s t o r i c a l l y < 1500 2 salmon escapement h i s t o r i c a l l y 1500 - 7500 3 salmon escapement h i s t o r i c a l l y > 7500 Numbers Include the t o t a l of a l l species of salmon. D. Produc t iv i t y Produc t i v i t y general ly re fer red to the r e l a t i v e amount or proport ion of marsh and f i ne textured I n t e r t l d a l f l a t s present on a s i t e . General knowledge of the Ind iv idua ls Involved and the WB Coastal Photo Catalogue were used to derive these ra t i ngs . Ratings C r i t e r i a 1 s i tes having minimal areas of marsh and f ine textured I n t e r t l d a l f l a t s usual ly resu l t i ng from steep gradient streams and/or unprotected locat ions 2 s i tes Intermediate to 1 and 3 3 s i tes having propor t ionate ly high areas o f marsh and f l a t s usual ly associated wi th low gradient streams and protected locat ions £. Social Value Rating This r a t i n g r e f l e c t s the proximity of the wetland to human population centres and/or the publ ic appreciat ion o f the wetland. Rating C r i t e r i a 1 remote s i tes fo r which the general public Is perceived to have no special appreciat ion 2 intermediate to 1 and 3 3 s i tes located near a r e l a t i v e l y large centre of population and/or which the general publ ic perceive as having high natural resource values. F. Imminence o f Development This r a t i n g r e f l e c t s the current development pressures on the wetland. I t 1s an est imat ion of the p robab i l i t y o f development or fu ture development o f the area based on knowledge of BCHOE regional s t a f f . Rating C r i t e r i a 1 no e x i s t i n g or an t ic ipa ted development pressure 2 p r o b a b i l i t y o f development and/or s igni f icance of development considered moderate 3 high p robab i l i t y of development wi th p o t e n t i a l l y negative impacts on the f i s h and w i l d l i f e resource 6. Rehab i l i ta t ion and Enhancement Value This ra t ing indicates the current potent ia l of a s i t e fo r r e h a b i l i t a -tion/enhancement fo r f i s h and waterfowl . In keeping wi th BCMOE pol icy non-Impacted s i tes are not considered to have rehabil i tat ion/enhancement poten-t i a l . This r a t i n g Is based on the known state of the a r t o f management techniques and does not consider cos t /bene f i t . Rating C r i t e r i a 1 no r e h a b i l i t a t i o n or enhancement potent ia l ( p r i s t i n e , r e l a t i v e l y undisturbed or I r r e v e r s i b l y degraded) 2 moderate r e h a b i l i t a t i o n or enhancement potent ia l 3 product ive areas wi th development or disturbance amenable to current rehabil i tat ion/enhancement techniques 1 3 3 Appendix B: Frequency of plant species i n each marsh, calculated as the percent of quadrats occupied. KO NI CL MA WA GO SJ 0U KA KI DA YA KU Deschampsia c e s p i t o s a 39 50 28 59 32 39 51 71 53 73 70 52 44 Carex lyngbyei P o t e n t i l l a anserina Glaux maritima 41 43 22 55 39 69 59 50 66 66 49 4 Festuca rubra A g r o s t i s s t o l o n i f e r a 21 14 13 8 22 19 8 Plantago macrocarpa 20 Juncus a r c t i c u s Plantago maritima 52 58 17 24 36 29 3 14 2 32 8 26 27 24 19 24 A c h i l l e a m i l l e f o l i u m 10 19 9 22 8 Sper g u l a r i a canadensis 5 1 16 43 24 19 6 T r i f o l i u m wormskjoldii 11 50 7 26 11 5 L i l a e o p s i s o c c i d e n t a l i s - 28 - 31 Ast e r subspicatus - 3 3 - 13 C o c h l e a r i a o f f i c i n a l i s 2 1 2 29 16 13 16 5 4 47 34 52 20 43 16 16 24 37 54 59 86 29 19 19 34 18 19 33 42 41 39 44 3 5 33 40 T r i g l o c h i n maritimum 32 23 51 49 55 69 51 38 41 35 38 44 70 16 9 30 2 14 5 14 24 10 24 21 11 9 Hordeum brachyantherum 22 14 50 3 19 15 19 27 27 10 24 10 40 34 13 10 50 48 13 17 37 1 8 8 23 23 46 24 19 5 36 17 13 23 32 49 13 1 Legend: KO(Kokish), NI(Nimpkish, CL(Cluxewe), MA(Marble), WA(Waukwaas), GO(Goodspeed), SJ(San J o s e f ) , 0U(Ououkinsh), KA(Kaouk), K I ( K i l d a l a ) , DA(Dala), YA(Yakoun), KU(Kumdis) NOTE: Ncnenclature follows Taylor & MacBryde (1977). (+, species frequency <1%; -, species not found) 1 3 4 KO NI CL MA WA GO S3 OU KA KI DA YA KU P u c c i n e l l i a pumila 11 42 22 18 27 9 Elymus m o l l i s 11 15 11 8 14 Conioselinum pacificum 3 11 15 38 13 2 Scirpus cernuus 8 13 11 1 39 53 F r i t i l l a r i a camschatcensis l 12 8 S t e l l a r i a humifusa 17 - 17 13 1 10 16 C a s t i l l e j a sp. 11 5 Poa p r a t e n s i s 1 11 11 1 S a l i c o r n i a v i r g i n i c a 23 46 39 23 Sisyrinchium l i t t o r a l e 1 17 15 2 Galium t r i f i d u m 15 Maianthemum dilatatum 10 9 A t r i p l e x p a t u l a 44 Heracleum sphondylium 1 Plantago major E l e o c h a r i s p a l u s t r i s T r i e n t a l i s europaea Carex p l u r i f l o r a Dodecatheon pulchellum Holcus lanatus Calamagrostis nutkaensis Lathyrus p a l u s t r i s Elymus h i r s u t u s 34 1 1 17 10 2 18 23 24 14 10 6 12 11 1 36 31 17 135 KO NI. CL MA WA GO SJ OU KA KI DA YA KU C l a y t o n i a s i b i r i c a - 1 0 - 1 - - - - - - - 7 -Carex obnupta - 9 - 13 - - - - - - - 4 Hypochaeris r a d i c a t a - 5 - 1 1 _ _ _ _ _ _ _ _ P r u n e l l a v u l g a r i s j - 6 - 1 - - - - - - -Carex o e d e r i - 1 6 - 4 _ _ _ _ _ _ _ _ _ Myrica gale - 4 - 1 - - - - - _ _ _ _ C i c u t a d o u g l a s i i - - - - - _ _ _ _ 5 10 - -Mentha arvens i s - - - - - - _ _ _ 22 23 - -Poa eminens - _ - - - _ _ _ 18 20 - -Hierochloe odorata _ - - - _ _ _ _ _ g g _ _ Angelic a genuflexa - - - - - - - - - 41 1 9 - -Lupinus l i t t o r a l i s _ _ - - _ _ _ _ _ 7 4 _ _ Apargidium boreale _ _ _ _ _ _ _ _ _ _ _ q Oenanthe sarmentosa _ 4 _ _ - _ _ _ _ _ _ l _ Plantago l a n c e o l a t a - 1 8 - - + - - _ _ _ _ _ _ Ranunculus cymbalaria - - - - _ _ - - _ + 2 E l e o c h a r i s parvula _ _ _ _ - - _ _ 8 _ _ _ _ Galium aparine - - - - - - - _ _ _ . _ ! _ Li m o s e l l a aquatica _ - - - _ _ _ _ _ _ l _ _ L i l a e a s c i l l o i d e s _ _ _ _ _ _ _ _ _ _ _ _ _ S i urn suave - - - - - _ _ _ _ 2 - - -Ranunculus uncinatus - - _ _ _ _ _ _ _ _ _ 2 -V i c i a gigantea ! - - - _ _ _ _ _ _ _ _ _ KO NI Honkenya peploides ± Ranunculus o c c i d e n t a l i s - 3-Cynosurus c r i s t a t u s - 6 Juncus g e r a r d i i - 2 Juncus f a l c a t u s _ 4 Cerastium fontanum - 4 D a c t y l i s glomerata Scirpus v a l i d u s 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 Danthonia c a l i f o r n i c a Cirsium vulgare _ _ C a l l i t r i c h e s t a g n a l i s 1 3 7 Appendix C: Correlations of plant species with PCA axes I - H I of the vegetation data Kokish -Undyked Kokish -Dyked Cluxewe -Undyked Cluxewe Dyked -I PCA II I I I I PCA II I I I I PCA II I I I I PCA II I l l Deschampsia c e s p i t o s a .67 -.59 .41 .63 .39 .37 .91 .24 .08 .43 -.40 -.11 Carex lyngbyei -.81 -.31 .10 -.77 .57 .11 -.95 .22 .13 .25 .07 .07 P o t e n t i l l a anserina .57 -.45 -.50 .48 .27 .42 .07 -.71 .68 .41 .23 -.05 Glaux maritima -.08 .09 .17 .51 .59 .34 .17 -.20 .23 T r i g l o c h i n maritimum -.53 -.01 .28 .32 .70 .56 -.33 -.78 .47 Festuca rubra .75 .21 .08 .51 .18 .24 .75 .05 -.20 .15 .31 -.17 A g r o s t i s s t o l o n i f e r a .72 .41 .05 .46 .22 .28 .44 -.42 -.07 .39 .04 -.24 Plantago macrocarpa .45 .65 .28 .09 .02 -.02 Hordeum brachyantherum .29 -.16 -.18 .59 .30 .13 .63 -.15 -.22 -.10 -.56 -.75 Juncus a r c t i c u s .26 -.10 -.05 Plantago maritima -.01 -.18 .50 .10 -.21 .10 -.52 .15 -.03 A c h i l l e a m i l l e f o l i u m .80 .28 .14 .09 .02 -.02 .22 -.35 .02 .47 .30 .03 S p e r g u l a r i a canadensis -.14 -.09 -.06 -.32 -.05 .46 T r i f o l i u m wormskjoldii .69 .06 -.47 -.10 -.07 .16 .42 -.54 -.04 .29 .04 -.09 L i l a e o p s i s o c c i d e n t a l i s A s t e r subspicatus .21 .54 .05 C o c h l e a r i a o f f i c i n a l i s -.02 -.22 .12 .02 -.15 -.05 1 3 8 Nimpkish Marble Waukwaas Goodspeed I PCA II I I I I PCA II I I I I PCA II I I I I PCA II I I I Deschampsia c e s p i t o s a .69 .37 -.43 .25 .45 -.37 -.41 .73 -.31 .15 .73 -.25 Carex lyngbyei -.60 .05 -.59 -.65 -.20 -.68 -.68 .47 -.06 -.97 .02 -.01 P o t e n t i l l a anserina .67 -.36 -.38 .32 .43 -.26 .27 .60 .36 .04 .63 .06 Glaux maritima .31 -.25 .01 -.03 .47 .13 -.47 .52 -.14 .49 -.14 .32 T r i g l o c h i n maritimum -.16 -.38 .09 -.39 .83 .06 -.76 -.30 .54 .07 -.64 -.69 Festuca rubra .26 .29 .06 .16 .03 -.01 .07 .51 .05 .07 .33 .01 A g r o s t i s s t o l o n i f e r a .33 .11 -.06 .40 -.06 .02 .46 .48 .51 .24 .62 -.19 Plantago macrocarpa .38 .12 25 .33 -.17 -.17 .14 .11 .10 .23 .45 -.09 Hordeum brachyantherum .50 -.06 -.22 .13 .11 .06 .32 .48 .27 .20 .73 .03 Juncus a r c t i c u s .31 -.89 .07 .76 .43 -.07 .15 .61 .34 .28 .24 -.66 Plantago maritima .13 .05 -.02 -.15 .30 .13 -.58 .09 -.09 .38 -.33 .17 A c h i l l e a m i l l e f o l i u m .40 .29 .16 .70 -.27 -.13 .36 .20 .26 Spe r g u l a r i a canadensis -.08 .06 .07 .14 -.63 -.18 .29 -.46 .48 T r i f o l i u m wormskjoldii .69 -.10 .03 .56 .08 -.06 .41 .38 .43 .09 .36 -.02 L i l a e o p s i s o c c i d e n t a l i s -.41 -.34 .09 -.25 .59 .12 .19 -.11 .09 A s t e r subspicatus .07 .04 .04 .44 -.12 -.10 C o c h l e a r i a o f f i c i n a l i s -.08 .06 .07 -.01 .09 -.12 -.05 -.15 .00 San Josef Ououkinsh Kaouk I PCA II I I I I PCA II I I I I PCA II I I I Deschampsia c e s p i t o s a .93 .07 -.17 .11 -.43 -.84 .61 -.59 -.43 Carex lyngbyei -.57 .44 -.67 -.76 .20 -.18 -.88 .03 -.31 P o t e n t i l l a anserina .40 .56 .23 .51 .10 -.39 .55 -.08 -.57 Glaux maritima -.25 .05 .54 -.21 -.02 -.42 .24 .20 -.60 T r i g l o c h i n maritimum .15 .87 .40 -.41 .55 -.05 .06 .55 -.54 Festuca rubra .62 -.48 -.28 .63 .15 .03 .35 -.12 .41 A g r o s t i s s t o l o n i f e r a .46 -.36 -.22 -.01 -.01 -.02 .18 -.04 .03 Plantago macrocarpa .07 .31 -.09 .46 .41 -.01 .47 .70 -.04 Hordeum brachyantherum .55 .06 .02 .47 -.17 -.15 .47 -.34 -.24 Juncus a r c t i c u s .67 .60 -.10 .43 .77 .12 Plantago maritima -.08 .14 .48 -.26 -.04 -.17 -.04 .49 -.16 A c h i l l e a m i l l e f o l i u m .41 -.35 -.20 .18 -.21 .30 .19 .08 .34 Spe r g u l a r i a canadensis -.27 -.42 .51 -.15 -.02 .24 -.01 -.08 .24 T r i f o l i u m wormskjoldii .65 -.14 .00 .46 .08 .30 L i l a e o p s i s o c c i d e n t a l i s -.04 .38 .20 -.12 .04 -.01 .04 .33 -.12 Aster subspicatus .13 -.05 .14 .33 -.16 .61 Co c h l e a r i a o f f i c i n a l i s -.30 -.45 .49 -.07 -.09 .17 .05 -.12 .13 1 4 0 K i l d a l a Dala Yakoun Kumdis I PCA II I I I I PCA II I I I I PCA II I I I I PCA II I I I Deschampsia c e s p i t o s a .70 -.49 -.37 .69 -.63 .11 .76 -.52 .06 .91 -.03 -.27 Carex lyngbyei -.82 .13 -.35 -.93 -.13 .06 -.40. -.06 -.86 .05 -.28 .56 P o t e n t i l l a anserina .30 -.84 .16 .62 -.40 -.21 .61 -.32 .17 .56 .38 .36 Glaux maritima -.01 .06 -.15 .22 -.02 .31 .47 -.52 .15 .93 .15 -.09 T r i g l o c h i n maritimum .10 -.73 -.39 .22 -.61 .41 .39 -.78 -.18 .61 -.70 .15 Festuca rubra .33 .12 -.04 .34 .50 .50 .27 .38 .01 .32 .35 .41 A g r o s t i s s t o l o n i f e r a .06 .01 .04 .04 .05 -.14 .18 .31 .01 .48 .36 .42 Plantago macrocarpa .73 .38 -.44 .38 .19 .72 .45 .33 .10 .24 .28 .37 Hordeum brachyantherum .20 -.01 -.09 .35 -.04 -.12 .28 .15 .08 Juncus a r c t i c u s .19 .14 .01 .14 .01 .25 .61 .75 -.09 .48 .51 .60 Plantago maritima .31 -.50 .16 .67 -.17 -.58 A c h i l l e a m i l l e f o l i u m .70 .35 .14 .58 .51 .04 .36 .47 .12 S p e r g u l a r i a canadensis -.80 .05 .43 -.80 -.12 -.03 T r i f o l i u m wormskjoldii .30 .06 -.42 L i l a e o p s i s o c c i d e n t a l i s .06 -.35 -.15 .11 .16 .09 A s t e r subspicatus .64 .05 .20 .64 .45 -.23 .36 .44 .12 C o c h l e a r i a o f f i c i n a l i s 1 4 1 Kokish -Undyked Kokish -Dyked Cluxewe -Undyked Cluxewe Dyked I PCA II I I I I PCA II I I I PCA I I I I I I I PCA II I I I P u c c i n e l l i a purcila -.08 -.16 .01 -.70 .01 .11 Elymus m o l l i s .05 .11 -.74 .27 .04 -.65 .30 -.01 -.18 Conioselinum pa c i f i c u m .25 .60 .11 Scirpus cernuus F r i t i l l a r i a camschatcensis .18 .44 -.02 .20 .32 .03 S t e l l a r i a humifusa .37 -.16 -.01 .07 .28 .51 .57 .15 .01 .08 -.28 .31 i 1 C a s t i l l e j a sp. I 1 Poa p r a t e n s i s .09 .02 -.06 .50 .38 .10 S a l i c o r n i a v i r g i n i c a -.22 -.81 .37 -.91 .32 -.04 Sisyrinchium l i t t o r a l e .17 .43 .08 Galium t r i f i d u m Maianthemum dilatatum .28 .62 .13 A t r i p l e x patula .09 -.13 -.01 -.53 -.32 -.55 Heracleum sphondylium .23 -.03 -.05 .08 .01 -.04 .21 .17 .08 Plantago major E l e o c h a r i s p a l u s t r i s T r i e n t a l i s europaea Carex p l u r i f l o r a Dodecatheon pulchellum Holcus lanatus .08 .01 -.04 .14 .11 .05 Calamagrostis nutkaensis Lathyrus p a l u s t r i s .12 .11 .04 Elymus h i r s u t u s .24 .46 .10 i .27 .22 .06 1 4 2 Nimpkish Marble Waukwaas Goodspeed I PCA II I I I I PCA II I I I PCA I II I I I PCA I I I I I I P u c c i n e l l i a puniila -.23 -.48 .26 .28 -.37 .41 Elymus m o l l i s -.06 .17 .11 Conioselinum pac i f i c u m .32 .17 .27 .49 -.13 .08 Scirpus cernuus -.09 .22 .08 .10 -.27 -.08 F r i t i l l a r i a camschatcensis .14 -.06 .05 S t e l l a r i a humifusa -.50 .31 -.20 -.01 -.01 -.05 C a s t i l l e j a sp. .14 .10 .23 .28 -.10 .01 Poa p r a t e n s i s .27 .22 .21 .11 .01 .04 .27 .08 .14 S a l i c o r n i a v i r g i n i c a -.43 -.24 -.18 .47 -.43 .60 Sisyrinchium l i t t o r a l e .38 .25 .38 .50 -.12 -.07 .20 .08 .12 Galium t r i f i d u m .09 .11 .04 .06 .05 .14 Maianthemum dilatatum .18 -.08 -.05 .09 .01 .04 A t r i p l e x patula -.03 .33 -.15 Heracleum sphondylium Plantago major .25 .31 .19 -E l e o c h a r i s p a l u s t r i s -.40 -.30 .19 -.07 .26 -.18 T r i e n t a l i s europaea .11 .04 .11 .18 -.04 .05 Carex p l u r i f l o r a .04 .13 .17 Dodecatheon pulchellum .44 .19 .18 Holcus lanatus .14 .26 .39 •19 .09 .18 Calamagrostis nutkaensis Lathyrus p a l u s t r i s Elymus h i r s u t u s .07 -.02 .11 P u c c i n e l l i a pumila Elymus m o l l i s Conioselinum p a c i f i c u m Scirpus cernuus F r i t i l l a r i a camschatcensis S t e l l a r i a humifusa C a s t i l l e j a sp. Poa p r a t e n s i s S a l i c o r n i a v i r g i n i c a S isyrinchium l i t t o r a l e Galium t r i f i d u m Maianthemum dilatatum A t r i p l e x patula Heracleum sphondylium Plantago major E l e o c h a r i s p a l u s t r i s T r i e n t a l i s europaea Carex p l u r i f l o r a Dodecatheon pulchellum Holcus lanatus Calamagrostis nutkaensis Lathyrus p a l u s t r i s Elymus h i r s u t u s San Josef PCA I II I I I -.44 -.48 .47 Ououkinsh PCA I I I I I I -.23 .05 .30 .56 -.46 -.27 .33 -.59 .46 .26 .13 .08 -.12 .34 .18 .25 -.20 -.12 -.15 .04 -.05 .12 .06 .01 -.19 .05 .15 .20 .21 .10 .11 .11 .01 ! .08 .07 .10 143 Kaouk PCA I II I I I .02 -.08 .23 .32 -.20 .61 .28 .15 .24 .13 .12 -.05 .11 .03 .21 .04 -.10 .00 .06 -.06 .23 .01 -.05 .03 .16 .06 .08 .13 .01 .14 144 K i l d a l a Dala Yakoun Kumdis I PCA 11 I I I I PCA II I I I I PCA II I I I I PCA II I I I P u c c i n e l l i a pumila -.07 -.02 .12 i | Elymus m o l l i s .08 .01 .70 .19 .53 -.24 ! 1 Conioselinum pacificum .58 .20 .26 .24 .30 .28 .11 .19 .06 j i Scirpus cernuus -.93 -.04 .10 1 -.81 -.42 .17 F r i t i l l a r i a camschatcensis .26 .09 .19 .19 .41 .23 .07 .15 .05 S t e l l a r i a humifusa .31 -.39 .12 .56 -.12 -.42 C a s t i l l e j a sp. .22 -.08 .02 .14 .32 .10 .10 .07 .04 Poa p r a t e n s i s S a l i c o r n i a v i r g i n i c a S isyrinchium l i t t o r a l e Galium t r i f i d u m .25 -.05 .10 .17 .41 -.16 Maianthemum dilatatum .31 .20 .08 .22 .47 .21 A t r i p l e x patula • .05 -.04 -.07 Heracleum sphondylium .14 .02 .56 .11 .37 -.04 Plantago major -.09 .01 .01 .05 .15 -.04 E l e o c h a r i s p a l u s t r i s -.36 -.25 -.05 -.36 -.37 .10 T r i e n t a l i s europaea .47 .34 -.11 .30 .26 .63 Carex p l u r i f l o r a .39 .32 -.14 .14 .21 .49 .02 .06 -.17 Dodecatheon pulchellum .39 .26 -.20 .27 .18 .64 .10 .10 .10 Holcus lanatus ] Calamagrostis nutkaensis .31 .62 - .10 Lathyrus p a l u s t r i s .30 -.24 .00 .47 -.05 • -.46 Elymus h i r s u t u s .09 .03 • -.01 Kokish -Undyked Kokish -Dyked Cluxewe -Undyked Cluxewe -Dyked I PCA II I I I I PCA II I I I I PCA II I I I I PCA II I I I C l a y t o n i a s i b i r i c a Carex obnupta Hypochaeris r a d i c a t a P r u n e l l a v u l g a r i s Carex oederi Myrica gale C i c u t a d o u g l a s i i Mentha arvens i s c Poa eminens Hierochloe odorata A n g e l i c a genuflexa Lupinus l i t t o r a l i s Apargidium boreale Oenanthe sarmentosa Plantago l a n c e o l a t a Ranunculus cymbalaria E l e o c h a r i s parvula Galium aparine L i m o s e l l a aquatica L i l a e a s c i l l o i d e s Sium suave Ranunculus uncinatus V i c i a gigantea .07 -.02 -.22 146 Nimpkish PCA I II I I I C l a y t o n i a s i b i r i c a Carex obnupta Hypochaeris r a d i c a t a P r u n e l l a v u l g a r i s Carex oederi Myrica gale Cicuta d o u g l a s i i Mentha arvens i s Poa eminens Hierochloe odorata A n g e l i c a genuflexa Lupinus l i t t o r a l i s Apargidium boreale Oenanthe sarmentosa Plantago l a n c e o l a t a Ranunculus cymbalaria E l e o c h a r i s parvula Galium aparine L i m o s e l l a aquatica L i l a e a s c i l l o i d e s Sium suave Ranunculus uncinatus .21 .13 -.08 .13 .10 .17 .16 .13 .30 .18 .20 .43 .29 .06 .38 I-.08 .18 -.01 V i c i a gigantea .12 .04 -.18 .30 .38 .59 Marble PCA I I I I I I .06 .02 .09 .29 -.02 .29 .11 -.09 .04 .15 -.02 -.11 18 .04 .01 02 .10 .10 Waukwaas PCA I II I I I .18 .12 .23 Goodspeed PCA I II I I I .06 -.01 -.02 C l a y t o n i a s i b i r i c a Carex obnupta Hypochaeris r a d i c a t a P r u n e l l a v u l g a r i s Carex oederi Myrica gale C i c u t a d o u g l a s i i Mentha arvens i s Poa eminens Hierochloe odorata Ang e l i c a genuflexa Lupinus l i t t o r a l i s Apargidium boreale Oenanthe sarmentosa Plantago l a n c e o l a t a Ranunculus cymbalaria E l e o c h a r i s parvula Galium aparine L i m o s e l l a aquatica L i l a e a s c i l l o i d e s San Josef Sium suave Ranunculus uncinatus V i c i a gigantea PCA I I I I I I Ououkinsh PCA I II I I I Kaouk PCA I II I I I -.17 .26 -.04 C l a y t o n i a s i b i r i c a Carex obnupta Hypochaeris r a d i c a t a P r u n e l l a v u l g a r i s Carex oederi Myrica gale Cicuta d o u g l a s i i Mentha arvens i s Poa eminens Hierochloe odorata Ang e l i c a genuflexa Lupinus l i t t o r a l i s Apargidium boreale Oenanthe sarmentosa Plantago l a n c e o l a t a Ranunculus cymbalaria E l e o c h a r i s parvula Galium aparine L i m o s e l l a aquatica L i l a e a s c i l l o i d e s Sium suave Ranunculus uncinatus V i c i a gigantea K i l d a l a PCA I II I I I -.05 -.35 -.11 -.12 -.28 .04 Dala PCA I II I I I .33 -.03 .34 .33 .22 .03 .18 .16 .06 .70 .46 -.14 .21 .14 .06 .39 .25 -.14 .44 .19 -.08 .23 .23 .26 .34 .46 .61 .11 .35 .11 Yakoun PCA I II I I I .14 .39 -.24 09 .31 -.04 Kumdis PCA I I I I I I .09 .01 .01 .47 .26 .17 i .31 .43 .45 .06 .12 .03 .02 -.26 -.03 -.05 -.05 .01 .27 .17 -.20 -.09 -.01 .02 -.08 .02 .00 .15 .13 .06 149 Honkenya peploides Ranunculus o c c i d e n t a l i s Cynosurus c r i s t a t u s Juncus g e r a r d i i Juncus f a l c a t u s Cerastium fontanum D a c t y l i s glomerata Scirpus v a l i d u s 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 Danthonia c a l i f o r n i c a Cirsium vulgare C a l l i t r i c h e s t a g n a l i s Kokish -Undyked Kokish -Dyked Cluxewe -Undyked Cluxewe -Dyked PCA I II I I I PCA I II I I I PCA I II I I I PCA I II I I I .09 .09 -.07 .05 -.01 -.10 1 1 1 I I .09 .06 .04 j i 1 5 0 Nimpkish Marble Waukwaas Goodspeed I PCA II I I I PCA I II I I I PCA I II I I I PCA I II I I I Honkenya peploides Ranunculus o c c l d e n t a l i s .63 .18 .05 Cynosurus c r i s t a t u s .17 .20 .42 Juncus g e r a r d i i .04 .04 -.01 Juncus f a l c a t u s .10 .06 .17 Cerastium fontanum .14 .13 .30 D a c t y l i s glomerata Scirpus v a l i d u s .08 -.18 .02 G r i n d e l i a i n t e g r i f o l i a .11 .11 . 06 D i s t i c h l i s s p i c a t a -.04 -.10 -. 04 Danthonia c a l i f o r n i c a .06 -.01 -. 02 Cirsium vulgare .09 .03 . 05 C a l l i t r i c h e s t a g n a l i s Honkenya peploides Ranunculus o c c i d e n t a l i s Cynosurus c r i s t a t u s Juncus g e r a r d i i Juncus f a l c a t u s Cerastium fontanum D a c t y l i s glomerata Scirpus v a l i d u s 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 Danthonia c a l i f o r n i c a Cirsiuni vulgare C a l l i t r i c h e s t a g n a l i s San Josef PCA I II I I I Ououkinsh PCA I II I I I Kaouk PCA I II I I I 152 K i l d a l a Dala yakoun Kumdis PCA I II I I I PCA I I I I I I PCA I II I I I PCA I II I I I Honkenya peploides Ranunculus o c c i d e n t a l i s Cynosurus c r i s t a t u s Juncus g e r a r d i i Juncus f a l c a t u s ! | | Cerastium fontanum j i i D a c t y l i s glomerata : i i i ! | Scirpus v a l i d u s i i 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 Danthonia c a l i f o r n i c a i Cirsium vulgare C a l l i t r i c h e s t a g n a l i s -.09 .05 -.01 APPENDIX D: V e g e t a t i o n maps f o r study a r e a s . Legends f o r p l a n t communities g i v e dominant s p e c i e s i n each community (those s p e c i e s w i t h an average p e r c e n t cover > 10%). A v e g e t a t i o n map f o r the San J o s e f r i v e r e s t u a r y was not produced. Legend t o map symbols: B a ( b a r ) , C r ( c r e e k ) , D ( d y k e ) , F ( f o r e s t ) , F l ( m u d f l a t ) , R d ( r o a d ) , R e s ( r e s i d e n t i a l ) , S h ( s h r u b ) . KAOUK RIVER ESTUARY PLANT COMMUNITIES 1. Carex l y n g b y e i 2. C o c h l e a r i a o f f i c i n a l i s , Puce i n e l l i a p u m i l a , S p e r g u l a r i a  c a n a d e n s i s 3. Deschampsia c e s p i t o s a , Carex l y n g b y e i , P o t e n t i l l a a n s e r i n a ,  T r i g l o c h i n maritimum, Glaux m a r i t i m a , Hordeum brachyantherum 4. Carex l y n g b y e i , T r i g l o c h i n maritimum, P l a n t a q o m a r i t i m a ,  E l e o c h a r i s p a r v u l a , Juncus a r c t i c u s , Glaux m a r i t i m a , S c i r p u s  cernuus 5. P l a n t a g o macrocarpa, Juncus a r c t i c u s , P o t e n t i 1 1 a a n s e r i n a ,  T r i g l o c h i n maritimum, Glaux m a r i t i m a 6. Elymus m o l l i s , A s t e r s u b s p i c a t u s , Deschampsia c e s p i t o s a ,  P o t e n t i l i a a n s e r i n a , Juncus a r c t i c u s 155 OUOUKINSH RIVER ESTUARY PLANT COMMUNITIES 1. Deschampsia c e s p i t o s a , Carex l y n g b y e i , T r i g l o c h i n maritimum 2 . Juncus a r c t i c u s , P l a n t a g o macrocarpa, Deschampsia c e s p i t o s a ,  P o t e n t i 1 1 a a n s e r i n a , Tr i g l o c h i n maritimum, Tr i f o l i u m w o r m s k j o l d i 3. Elymus m o l l i s , Deschampsia c e s p i t o s a , T r i f o l i u m w o r m s k j o l d i i 1 5 7 OUOUKINSH RIVER ESTUARY VEGETATION MAP 1 5 3 KOKISH RIVER ESTUARY PLANT COMMUNITIES 1. Carex l y n g b y e i 2. 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 3. Carex l y n g b y e i , T r i g l o c h i n maritimum 4. Deschampsia c e s p i t o s a , Hordeum brachyantherum, A g r o s t i s  s t o l o n i f e r a , P o t e n t i l l a a n s e r i n a 5. I n s i d e dyke: Elymus m o l l i s , Deschampsia c e s p i t o s a , A g r o s t i s  s t o l o n i f e r a . O u t s i d e dyke: Elymus m o l l i s, T r i f o l i u m  w o r m s k j o l d i i , P o t e n t i l l a anser i n a 6. Carex l y n g b y e i , Deschampsia c e s p i t o s a , P o t e n t i l l a a n s e r i n a 7. Deschampsia c e s p i t o s a , P o t e n t i l l a a n s e r i n a , Hordeum  brachyantherum 8. Deschampsia c e s p i t o s a , A c h i I l e a m i l l e f o l i u m , P o t e n t i l l a  a n s e r i n a , P l a n t a g o macrocarpa, T r i f o l i u m w o r m s k j o l d i i 159 K O K I S H R I V E R E S T U A R Y V E G E T A T I O N M A P NIMPKISH RIVER ESTUARY PLANT COMMUNITIES Carex l y n g b y e i S p e r q u l a r i a c a n a d e n s i s , C o c h l e a r i a o f f i c i n a l i s Juncus a r c t i c u s Deschampsia c e s p i t o s a , P o t e n t i l l a a n ser i n a , Juncus a r c t i c u s M y r i c a g a l e , P l a n t a g o l a n c e o l a t a , Carex l y n g b y e i Elymus m o l l i s , P l a n t a g o l a n c e o l a t a , Deschampsia c e s p i t o s a N I M P K I S H R I V E R E S T U A R Y V E G E T A T I O N M A P 162 CLUXEWE RIVER ESTUARY PLANT COMMUNITIES 1. Carex l y n g b y e i , P o t e n t i l l a anser i n a 2. Deschampsia c e s p i t o s a , Tr i g l o c h i n maritimum, Carex l y n g b y e i ,  Hordeum brachyantherum 3. 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, Hordeum  brachyantherum, A t r i p l e x p a t u l a , P u c e i n e l l i a p u m i l a , P l a n t a g o  m a r i t i m a 4. Poa p r a t e n s i s , P o t e n t i l l a a n s e r i n a , A c h i l l e a m i l l e f o l i u m ,  Elymus h i r s u t u s 5. Elymus m o l l i s , P o t e n t i l l a a n s e r i n a , Deschampsia c e s p i t o s a ,  Hordeum brachyantherum 164 WAUKWAAS RIVER ESTUARY PLANT COMMUNITIES 1. Carex l y n g b y e i , Ruppia m a r i t i m a 2. Ruppia mar i t ima, S p e r q u l a r i a c a n a d e n s i s 3. S a l i c o r n i a v i r g i n i c a , S p e r q u l a r i a c a n a d e n s i s 4. T r i g l o c h i n maritimum, Carex l y n q b y e i , Deschampsia c e s p i t o s a 5. A g r o s t i s s t o l o n i f e r a , Juncus a r c t i c u s , Deschampsia c e s p i t o s a , P o t e n t i l l a a n s e r i n a W A U K W A A S R I V E R E S T U A R Y V E G E T A T I O N M A P 166 MARBLE RIVER ESTUARY PLANT COMMUNITIES 1 . Carex l y n g b y e i 2. T r i g l o c h i n maritimum, Deschampsia c e s p i t o s a , Carex l y n g b y e i ,  Juncus a r c t i c u s 3. P l a n t a g o macrocarpa, Juncus a r c t i c u s MARBLE RIVER ESTUARY VEGETATION MAP 168 GOODSPEED RIVER ESTUARY PLANT COMMUNITIES 1. Carex l y n g b y e i 2« T r i g l o c h i n maritimum, S a l i c o r n i a v i r q i n i c a 3. Tr i g l o c h i n maritimum, Carex l y n g b y e i 4. Deschampsia c e s p i t o s a , Carex l y n g b y e i , Juncus a r c t i c u s ,  T r i g l o c h i n maritimum G O O D S P E E D R I V E R E S T U A R Y V E G E T A T I O N M A P 1 6 9 SAN JOSEF RIVER ESTUARY PLANT COMMUNITIES P u c e i n e l l i a p u m i l a Carex l y n g b y e i , T r i q l o c h i n maritimum T r i g l o c h i n maritimum, Deschampsia c e s p i t o s a , Carex l y n g b y e i Deschampsia c e s p i t o s a , Elymus m o l l i s , A g r o s t i s s t o l o n i f e r a 171 KILDALA RIVER ESTUARY PLANT COMMUNITIES 1 . E l e o c h a r i s p a l u s t r i s 2. Carex l y n g b y e i 3. Deschampsia c e s p i t o s a , P o t e n t i 1 1 a a n s e r i n a , T r i g l o c h i n  maritimum, Carex l y n g b y e i , L a t h y r u s p a l u s t r i s , E l e o c h a r i s  p a l u s t r i s 4. Deschampsia c e s p i t o s a , P l a n t a g o macrocarpa, P o t e n t i l l a  a n s e r i n a , A s t e r s u b s p i c a t u s , A n g e l i c a g e n u f l e x a , A c h i l l e a  m i l l e f o l i u m , Carex l y n g b y e i , C o n i o s e l i n u m p a c i f i c u m , L a t h y r u s  p a l u s t r i s 5. Elymus m o l l i s , P o t e n t i l l a anser i n a , A s t e r s u b s p i c a t u s ,  A c h i I l e a m i l l e f o l i u m , Heracleum sphondylium, C o n i o s e l i n u m  p a c i f i c u m , Deschampsia c e s p i t o s a , A n g e l i c a g e n u f l e x a 172 KILDALA RIVER ESTUARY VEGETATION MAP 1 7 3 DALA RIVER ESTUARY PLANT COMMUNITIES 1. 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, P o t e n t i 1 1 a anser i n a 2. Deschampsia c e s p i t o s a , P o t e n t i l l a anser i n a , T r i g l o c h i n  maritimum, Carex l y n g b y e i 3. Deschampsia c e s p i t o s a , P o t e n t i l l a anser i n a , A s t e r  s u b s p i c a t u s , P l a n t a g o macrocarpa, A c h i I l e a mi l i e f o l i u m , F e s t u c a  r u b r a , L a t h y r u s p a l u s t r i s , T r i f o l i u m w o r m s k j o l d i i 4. P o t e n t i l i a a nser i n a , Elymus m o l l i s , A s t e r s u b s p i c a t u s ,  A c h i I l e a m i l l i f o l i u m , Deschampsia c e s p i t o s a , F e s t u c a r u b r a , Maianthemum d i l a t a t u m DALA RIVER ESTUARY VEGETATION MAP 175 YAKOUN RIVER ESTUARY PLANT COMMUNITIES 1. S c i r p u s c e r n u u s , S p e r q u l a r i a c a n a d e n s i s 2. Sc i rpus c e r n u u s , Carex l y n g b y e i , S p e r q u l a r i a c a n a d e n s i s ,  T r i g l o c h i n maritimum 3 . Deschampsia c e s p i t o s a , T r i g l o c h i n maritimum, P o t e n t i l i a  a n s e r i n a , Glaux m a r i t i m a 4. Juncus a r c t i c u s , Deschampsia c e s p i t o s a , C a l a m a g r o s t i s  n u t k a e n s i s, P o t e n t i 1 1 a a n s e r i n a , A p a r g i d i u m b o r e a l e , P l a n t a g o  macrocarpa 176 Y A K O U N RIVER ESTUARY V E G E T A T I O N M A P 177 KUMDIS RIVER ESTUARY PLANT COMMUNITIES 1. S p e r q u l a r i a c a n a d e n s i s , Ruppia m a r i t i m a , S c i r p u s cernuus 2. S c i r p u s c e r n u u s , T r i g l o c h i n maritimum, Carex l y n g b y e i ,  S p e r g u l a r i a c a n a d e n s i s , Ruppia m a r i t i m a 3. Deschampsia c e s p i t o s a , T r i g l o c h i n maritimum, Glaux m a r i t i m a ,  P l a n t a g o m a r i t i m a 4. Juncus a r c t i c u s , T r i g l o c h i n maritimum, Deschampsia c e s p i t o s a ,  P o t e n t i 1 1 a a n s e r i n a , A g r o s t i s s t o l o n i f e r a , Glaux m a r i t i m a ,  A p a r g i d i u m b o r e a l e , Carex l y n g b y e i 1 7 8 KUMDIS RIVER ESTUARY VEGETATION MAP Appendix E: Graphs o f e l e v a t i o n v e r s u s c o v e r f o r each marsh s p e c i e s p e r c e n t A. Carex l y n g b y e i 7 180 B. Deschampsia c e s p i t o s a 7 c . P o t e n t i l l a a n s e r i n a 1 8 1 7 1 8 2 7 -n-Tr ELEVATION (m) ! TTTJ • i ; l w M e NIMPKISH OUOUKlNSH ID 1 SAN JOSEF WAUKWAAS IT IP DALA KILDALA KUMDIS YAKOUN ELEVATION M fD H O fD 3 r t O O < fO s O o rtrrr ID" CLUXEWE UNDYKED CLUXEWE DYKED OOODSPEED KAOUK KOKISH UNDYKED w I—1 <^ 3 d 3 O in Tr KOKISH DYKED 

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