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Vegetation-environment relationships in the tidal marshes of the Fraser River Delta, British Columbia Porter, Glendon Leslie 1982

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VEGETATION-ENVIRONMENT RELATIONSHIPS IN THE TIDAL MARSHES THE FRASER RIVER D E L T A , BRITISH COLUMBIA by GLENDON L E S L I E PORTER B.Sc,  University  A THESIS SUBMITTED  Of B r i t i s h C o l u m b i a ,  IN PARTIAL FULFILMENT OF  THE REQUIREMENTS  FOR THE DEGREE OF  MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES D e p a r t m e n t Of  We a c c e p t to  this  the  as  conforming  standard  OF BRITISH COLUMBIA  October  ©  Botany  thesis  required  THE UNIVERSITY  1969  Glendon L e s l i e  1982  Porter,  1982  In  presenting  requirements  this for  British  Columbia,  freely  available  that  permission  scholarly or  by  copying not  an  for  for  or  Department  of  October  degree  that  extensive  the  her  without  Botany  7,  1982  by the  thesis  my w r i t t e n  Columbia  at  the  of  is  thesis  it  agree for  my D e p a r t m e n t  understood  financial  permission.  the  make  further  this  Head of It  for  I  of  U n i v e r s i t y of  shall  and s t u d y .  representatives. this  fulfilment  Library  copying  may be g r a n t e d  The U n i v e r s i t y of B r i t i s h 2075 Wesbrook P l a c e V a n c o u v e r , Canada V6T 1W5  Date:  partial  reference  o r p u b l i c a t i o n of  be a l l o w e d  in  advanced  I agree  purposes  his  thesis  gain  that shall  i i  ABSTRACT  The  literature  on  North  American  Pacific  Coast  tidal  marshes n o r t h of M e x i c o i s summarized. Vegetation marshes were  at  and  environmental  Ladner  analyzed  Marsh,  using  r e c i p r o c a l averaging between  species  environmental of  sand,  potassium,  and  and  clay);  local  and  tidal  found  to  correlation principal quality  be  environmental  the  sodium;  the  of  and  was  vegetation  elevation were  environmental  data  matrix)  data,  were  of the  ordination;  Reciprocal gave  root  e c o l o g i c a l gradients..  vegetation  not.  analysis  nitrogen,  Ordinations  correlation  i n exposing  in  selected  (percentage  square root t r a n s f o r m a t i o n  useful  components  a n a l y s i s was  The  and  matrix  with  Bay and  s t a n d a r d i z a t i o n (square  n o r m a l i z a t i o n , and  for their usefulness  texture  range).  b o t h t h e v e g e t a t i o n and  Normalization  analysis  s o i l concentration  D i f f e r e n t methods of d a t a  tested  Boundary  d i s t r i b u t i o n , and  These were: s o i l  magnesium,  by  transformation,  were  components  tidal  to i n v e s t i g a t e q u a n t i t a t i v e r e l a t i o n s h i p s  calcium,  p e r f o r m e d on sets.  principal  factors.  (standardized  from F r a s e r D e l t a  B r u n s w i c k P o i n t , and  performance  silt,  data  results  the  averaging of  but p r i n c i p a l  data  and  equivalent components  generally superior to r e c i p r o c a l averaging  in  the  ordinations.  m a r s h e s of t h e  study  area  separate  conspicuously  into  two  types  on b o t h  floristic  fresh-to-brackish western  Brunswick P o i n t ,  Brunswick main  Point  and a t  at  Agrostis Scirpus  alba;  (2.)  Agrostis patula;  Salicorn ia  Spergular ia  related  to  of  and  virginica  levels  of  alba  by:  the  in  were  (1.)  and  four  informally  Carex l y n g b y e i  and S c i r p u s m a r i t i m u s ; fluviat ile,  and  Distichlis  Triqlochin  Patterns  of  Sc i r p u s (5.)  spicata;  maritimum;  (8.)  performance  and  t i d a l m a r s h s p e c i e s were shown measured  and (3.)  Alisma plantago-aquatica; and  a  southeastern  W i t h i n each a r e a ,  groups  Carex l y n g b y e i  important  type  E q u i seturn'  alba,  canadensis.  distribution  sample  (4.)  (6.)  saline  respectively  criteria:  M a r s h and i n n o r t h e r n  Boundary Bay.  Agrostis  americanus;  validus, Atriplex  dominated  environmental  Ladner  and a  species-environment  recognized,  (7.)  type  and  environmental  factors.  to  be  TABLE OF CONTENTS  ABSTRACT  ii  L I S T OF TABLES  vii  L I S T OF FIGURES  ix  ACKNOWLEDGEMENT  xiii  1.  INTRODUCTION  1  2.  LITERATURE REVIEW: PACIFIC COAST TIDAL MARSHES  3  3.  2.1  General  Overview  3  2.2  California  5  2.3  Oregon  8  2. 4  Washington  11  2.5  Alaska  12  2.6  B r i t i s h Columbia  14  THE STUDY AREA  24  3. 1  Location  24  3.2  F o r m a t i o n And D e v e l o p m e n t Of The F r a s e r  3.3  Extent  3.4  Physical  Of T i d a l  Marshes  Environment  In The F r a s e r  Of The F r a s e r  Marshes  Delta  Delta  Delta  ....24 26  Tidal 29  3.4.1  Climate  29  3.4.2  R i v e r And M a r i n e I n f l u e n c e s  30  3.4.3  Tides  33  3.4.4  S e d i m e n t a t i o n And S u b s t r a t e  35  3.5  4.  5.  Transect  Locations  And S i t e  Descriptions  Of L o c a t i o n s  38  3.5.1  Selection  38  3.5.2  Ladner Marsh  38  3.5.3  Brunswick P o i n t  40  3.5.4  Boundary Bay  42  SOME FACTORS AFFECTING SPECIES DISTRIBUTIONS 4.1  Introduction  4.2  Tides  4.3  Salinity  4.4  Substrate  Nutrient  4.5  Substrate  Texture  52 52 ...52 56  Regime  61 65  SAMPLING METHODS AND DATA COLLECTION  68  5.1  Vegetation  68  5.2  Soil  69  5.3  L e v e l l i n g Survey  69  6.  SOIL ANALYSIS  71  7.  DATA ANALYSIS  73  8.  7.1  O r d i n a t i o n Methods  ..73  7.2  Treatment  Of The V e g e t a t i o n  7.3  Treatment  Of The E n v i r o n m e n t a l D a t a  Data  DISCUSSION Data S t a n d a r d i z a t i o n s  8.2  Ecological  Interpretation  8.2.1  Vegetation  8.2.2  E n v i r o n m e n t a l Data  8.2.3  Species-Environment  Plant  ...86 100  8.1  8.3  75  100 Of O r d i n a t i o n R e s u l t s  Data  Communities  .105 105 110  Diagrams  In The S t u d y A r e a  116 118  vi  9.  CONCLUSIONS  REFERENCES  123 126  APPENDIX  A.  NAMES OF SPECIES SAMPLED, WITH AUTHORITIES  APPENDIX  B.  ENVIRONMENTAL DATA  .140 142  vii  LIST OF TABLES  I.  Environmental  data summary f o r the study a r e a : mean  d a i l y temperature,  mean d a i l y maximum  mean d a i l y minimum temperature,  temperature,  mean t o t a l  precipitation II.  30  T y p i c a l c o n c e n t r a t i o n s (mg L " ) of macronutrient 1  c a t i o n s p l u s Na  +  and C l ~ i n s o i l  solution,  seawater,  and F r a s e r R i v e r water III.  Floristic  t a b l e : 64 s p e c i e s i n 103 sample p l o t s .  Group at l e f t , mainly at IV.  fresh-water i n f l u e n c e d ; group  r i g h t , mainly s a l t - w a t e r i n f l u e n c e d  76  Summary of r e s u l t s of PCA o r d i n a t i o n on square roots of  s p e c i e s cover data from a l l p l o t s . E i g e n v e c t o r  elements V.  62  i n the range  -0.1500 to +0.1500 not shown 106  Summary of r e s u l t s of PCA o r d i n a t i o n on s p e c i e s cover data from freshwater p l o t s . E i g e n v e c t o r elements  VI.  i n the range  -0.150 to +0.150 not shown ..107  Summary of r e s u l t s of PCA o r d i n a t i o n on normalized s p e c i e s cover data from freshwater p l o t s . E i g e n v e c t o r elements  i n the range -0.150 to +0.150  not shown VII.  Summary of r e s u l t s of PCA o r d i n a t i o n on s p e c i e s cover data from s a l t w a t e r p l o t s . E i g e n v e c t o r  1 08  vi i i  elements i n the range -0.150 to +0.150 not shown VIII.  Summary of r e s u l t s of PCA o r d i n a t i o n on environmental data from freshwater  and s a l t w a t e r  p l o t s together IX.  111  Summary of r e s u l t s of PCA o r d i n a t i o n on environmental data from freshwater  X.  plots  112  Summary of r e s u l t s of PCA o r d i n a t i o n on environmental data from s a l t w a t e r p l o t s  XI.  ..110  Product-moment c o r r e l a t i o n c o e f f i c i e n t s ronmental v a r i a b l e s  113 for e n v i -  (* = p < .05; ** = p < .01)  115  ix  LIST OF FIGURES  1. Map  of the study area  25  2. Ladner Marsh, v i c i n i t y of Transect J . Foreground: Lythrum s a l i c a r i a , Carex l y n g b y e i .  Background:  Populus t r i c h o c a r p a . Right: S c i r p u s v a l i d u s  39  3. Brunswick Point North, v i c i n i t y of Transect A. Carex lyngbyei  i s dominant. Conspicuous i n f l o r e s c e n c e s :  Lythrum s a l i c a r i a  (magenta); Sium suave (white)  4. Brunswick Point South, v i c i n i t y  of Transect  43  A.  T r i g l o c h i n mar i t imum forms i s o l a t e d clumps at marsh edge; clumps c o a l e s c e i n t o continuous stand  (right  background)  44  5. Brunswick Point South, Transect B. Scene i n l a t e winter  45  6. A e r i a l photograph of Brunswick P o i n t ,  showing  c i r c u l a r p a t t e r n i n g of marsh v e g e t a t i o n . Clumps are mainly Carex l y n g b y e i and S c i r p u s maritimus  46  7. Boundary Bay East, Transect E. Mixed grasses i n foreground; Carex lynqbyei  (yellow-green) i n mid-  p o r t i o n of t r a n s e c t ; S a l i c o r n i a  virginica,  T r i g l o c h i n mar itimum, Plantago maritima at f a r end of t r a n s e c t  (dark green)  8. Boundary Bay E a s t . T r i g l o c h i n mar itimum and  47  X  S a l i c o r n i a v i r q i n i c a , with patches of green and blue-green covered  a l g a e . Black  filamentous  reducing mud  is  by brownish diatom mat  49  9. Boundary Bay West, v i c i n i t y of Transect H. p a t u l a dominates the d r i f t w o o d zone i n F a r t h e r out, S a l i c o r n i a v i r q i n i c a y e l l o w i s h Cuscuta  Atriplex  foreground.  i s infested  by  salina  50  10. Boundary Bay West. G r i n d e l i a  integrifolia  flowers  amid the d r i f t w o o d 11. PCA  o r d i n a t i o n of samples using s p e c i e s cover  from freshwater 12. PCA  51 data  p l o t group  78  o r d i n a t i o n of samples using s p e c i e s cover  data  from s a l t w a t e r p l o t group 13. PCA  o r d i n a t i o n of samples using square r o o t s of  s p e c i e s cover data 14. PCA  79  from freshwater  p l o t group  80  o r d i n a t i o n of samples using square roots of  s p e c i e s cover data from s a l t w a t e r p l o t group 15. PCA  81  o r d i n a t i o n of samples using square r.oots of  s p e c i e s cover data from a l l p l o t s  (freshwater  group,  T r a n s e c t s A, D, J ; s a l t w a t e r group, T r a n s e c t s B,  C,  E, F, G, H) 16. PCA  o r d i n a t i o n of samples using normalized  cover data 17. PCA  from freshwater  84 species  from s a l t w a t e r p l o t group  o r d i n a t i o n of s p e c i e s and  cover data  species  p l o t group  o r d i n a t i o n of samples using normalized  cover data 18. RA  82  from freshwater  samples using  p l o t group  85 species 87  19. PCA o r d i n a t i o n of samples u s i n g environmental from  freshwater  data  ( T r a n s e c t s A, D, J) and s a l t w a t e r  (Transects C, G, H) p l o t groups  91  20. PCA o r d i n a t i o n of samples u s i n g environmental from  data  freshwater p l o t s  92  21. PCA o r d i n a t i o n of samples u s i n g environmental  data  from, s a l t w a t e r p l o t s  93  22. P l o t of cover c l a s s codes of A g r o s t i s a l b a on the o r d i n a t i o n of environmental data from plots.  freshwater  1 = <25% cover; 2 = 26-50%; 3 = 51-75%;  4 = 76-100%; • = absent  94  23. P l o t of cover c l a s s codes of S c i r p u s americanus the o r d i n a t i o n of environmental data from plots.  on  freshwater  1 = <25% cover; 2 = 26-50%; 3 = 51-75%;  4 = 76-100%; • = absent  95  24. P l o t of cover c l a s s codes of Carex o r d i n a t i o n of environmental plots.  l y n g b y e i on the  data from  freshwater  1 = <25% cover; 2 = 26-50%; 3 = 51-75%;  4 = 76-100%; • = absent  96  25. P l o t of cover c l a s s codes of S a l i c o r n i a v i r g i n i c a on the o r d i n a t i o n of environmental data from s a l t w a t e r plots.  1 = <25% cover; 2 = 26-50%; 3 = 51-75%;  4 = 76-100%; • = absent 26. P l o t of cover c l a s s codes of D i s t i c h l i s  97 s p i c a t a on  the o r d i n a t i o n of environmental data from s a l t w a t e r plots.  1 = <25% cover; 2 = 26-50%; 3 = 51-75%;  4 = 76-100%; • = absent  98  Plot  of cover  class  the o r d i n a t i o n plots.  of e n v i r o n m e n t a l  1 = <25%  4 = 76-100%;  c o d e s of T r i g l o c h i n  cover;  • = absent  data  m a r i t i m u m on  from  saltwater  2 = 26-50%; 3 = 51-75%;  ACKNOWLEDGEMENT  Many i n d i v i d u a l s have helped me with this p r o j e c t , and I thank them a l l . Some I want to s i n g l e out f o r s p e c i a l mention. Anna Scagel was my much-appreciated field a s s i s t a n t , and Andy Mackinnon a i d e d me w i t h the s o i l a n a l y s i s . M a r s h a l l Cant and Ted Wickman brought t h e i r t e c h n i c a l competence to the job of surveying the sample p l o t s . A l a r d Ages of the I n s t i t u t e of Ocean Sciences gave v a l u a b l e a s s i s t a n c e with the t i d a l data. Dr. L.M. L a v k u l i c h generously made a v a i l a b l e h i s l a b o r a t o r y f a c i l i t i e s i n the U.B.C. S o i l Science Department, where V a l M i l e s s u p p l i e d very h e l p f u l technical assistance. I am indebted to the members of my M.Sc. advisory committee, Drs. G.E. Bradfield, W.B. S c h o f i e l d , and J.R. Maze, f o r t h e i r helpfulness in many ways; and particularly to Dr. B r a d f i e l d , my thesis supervisor, for h i s assistance, advice, i n s t r u c t i o n , understanding, and p a t i e n c e . My parents have throughout, f o r which I am  been very supportive especially grateful.  T h i s study was supported by a grant from the N a t u r a l Sciences and E n g i n e e r i n g Research C o u n c i l of Canada.  1  1.  Tidal  marsh  INTRODUCTION  ecosystems,  belonging  as  they do n e i t h e r  e n t i r e l y t o the t e r r e s t r i a l environment nor to the marine, yet o v e r l a p p i n g the f r o n t i e r s of both, have long been a source interest  to  the d i k i n g ,  ecologists draining,  marshlands  ecosystems critical to  -- even as man's ambition has l e d to cultivating,  everywhere.  relationships  between  have  been  filling  Fortunately,  industrial the  or  the  activities  object  in  of  and  recent  marshlands  as  an  and p o l i t i c a l d e c i s i o n making.  natural  years of some  economic  e c o l o g i c a l c o n s i d e r a t i o n s are no longer  coastal  interactive  examination, with the r e s u l t that i t i s now  appreciate  of  possible  resource,  ignored  in  and  economic  There i s hope, t h e r e f o r e ,  that  remaining e s t u a r i n e marshes w i l l be v a l u e d as important l i v i n g systems, r a t h e r than being t r e a t e d as u s e l e s s wastelands. The  tidal  marshlands  of  the  F r a s e r e s t u a r y , as l o c a l  hunters and n a t u r a l i s t s have long known, are p r i n c i p a l and  feeding  areas  f o r migratory w a t e r f o w l ;  demonstrates  their  importance  well.  Not the l e a s t  1  recent  nesting research  to the f i s h e r i e s r e s o u r c e  importance of these marshlands,  2  as  however,  1  Burgess 1970, Burton 1977, Campbell et a l . 1972, H a b i t a t Work Group 1978, Hoos & Packman 1974, Leach 1972, Vermeer & Levings 1977.  2  Dorcey et a l . 1978, Dunford 1975, Levy & Northcote 1981, et a l . 1979, V a l i e l a & K i s t r i t z 1980.  Levy  2  is  their  the c i t y ,  value as w i l d e r n e s s . teeming  with  Here are vast areas very near  birdlife  and  vibrant  with  floral  splendor,  i n which the a n x i e t i e s of a c i v i l i z e d e x i s t e n c e seem  to  l i k e the f a l l i n g  recede The  present  study  vegetation-environment o b j e c t i v e s may (1.)  tide. i s concerned  with the e l u c i d a t i o n of  r e l a t i o n s h i p s i n the F r a s e r D e l t a .  The  be summarized as f o l l o w s :  To d e s c r i b e and compare  the  vegetation  of  three  F r a s e r D e l t a t i d a l marshes; (2.)  To  r e l a t e performance and d i s t r i b u t i o n of v a s c u l a r  p l a n t s p e c i e s to measured environmental (3.)  To assess the e f f e c t s of d i f f e r e n t  standardization  in  exposing  ecological  p r i n c i p a l components a n a l y s i s and It marsh to  variables;  i s hoped that any systems that may  researchers  establishment  and  and  result  managers  of  gradients,  reciprocal  increased  types  of  from t h i s study w i l l be  rehabilitation.  using  averaging.  understanding  concerned  data  with  tidal  tidal helpful marsh  3  2.  2.1  LITERATURE REVIEW: PACIFIC COAST TIDAL MARSHES  General  The  Overview  literature  on  tidal  marsh  ecosystems  and  their  vegetation  i s voluminous, and cannot be d e a l t with here in i t s  entirety.  T h i s review i s t h e r e f o r e r e s t r i c t e d  to  the  Pacific  Coast  of Canada and  geographically  the United S t a t e s .  comprehensive, g l o b a l treatments of s a l t marsh, and  other  coastal  vegetation,  Chapman (1960, 1974, The  area being  extent,  with  shoreline  is  headlands.  a  1976,  generally  From  Puget  Sound  Ranwell  and  north,  erosion  et a l . 1970).  and  to  Jefferson  be  quite  1974), and  bays, e s t u a r i e s and The  small,  beach and  (1972).  the  California  of  its  shelf;  the  by  rocky  coast  i s deeply  has been  subjected  subsequent  that  isostatic  our  favour marshes  (Macdonald & Barbour  commonly r e s t r i c t e d to  r i v e r mouths (Macdonald  1974, small  1977a).  s a l t marsh v e g e t a t i o n of the North American  P a c i f i c coast between Point Barrow, Alaska and Baja  much  Such c o n d i t i o n s do not  young  isolated —  marsh,  r e f e r r e d to  marked  c o a s t a l marsh development, with the r e s u l t tend  is  continental  a r c h i p e l a g o s , and  -glacial  (Mathews  narrow  steep-sloped  d i s s e c t e d i n t o f i o r d s and  uplift  and  reader  reviewed i s mountainous for  .usually  to rather recent  1977)  the  tidal  For  was  surveyed  Cabo San  by Macdonald & Barbour  Lucas, (1974).  4  For the same area, Macdonald (1977a) summarized the  knowledge  to that date of s a l t marsh and mangal v e g e t a t i o n , and a  t e n t a t i v e phytogeographical  s a l t marshes, groups:  based  arctic,  arid.  These  on  c l a s s i f i c a t i o n of P a c i f i c  regional  floras,  into  major  groups  correspond  Islands;  roughly  to:  northern  and and  and  southeastern Alaska and the Queen  British  Columbia, Washington, Oregon and  northern C a l i f o r n i a ; southern C a l i f o r n i a ; and Baja  California,  respectively.  theoretical  sample  A numerical a n a l y s i s of data  sites  (using  "expected"  supported  these groupings,  northern  and  latitude. southern now,  The  southern  but  species  indicated a  divisions  from  at  in  broadly  break  into  about 51 degrees north  scheme thus a s s i g n s F r a s e r D e l t a marshes to the  group.  ( I t would be i n t e r e s t i n g to redo the a n a l y s i s  using data from a c t u a l sample l o c a t i o n s . )  a  compilation  of a b s t r a c t s  Sixth Biennial International  1  In  general,  vegetation  the  i n the review  published area  are  indicated  from papers d e l i v e r e d at the  Estuarine  p u b l i s h e d i n E s t u a r i e s , v o l . 4,  of  lists)  major  Some d i r e c t i o n s i n c u r r e n t marsh r e s e a r c h  1  five  Coast  s u b a r c t i c , temperate, dry mediterranean,  western A l a s k a ; southern Charlotte  proposed  Research  Conference,  1981. literature  on  i s rather sparse.  C a l i f o r n i a and Alaska are the best-documented.  especially Eilers 1981, F r e n k e l 1981, 1981, O l i v e r & R e i l l y 1981, Onuf 1981, Zedler 1981.  t i d a l marsh The  marshes  Marshes of  Mahall 1981, M i t c h e l l Wolf & Fucik 1981,  5  the  Oregon,  described  Washington,  mainly  and  British  coasts  are  i n unpublished theses and government r e p o r t s  ( f r e q u e n t l y q u i t e obscure ones). review, the l i t e r a t u r e  In  the  remainder  of  this  i s surveyed g e o g r a p h i c a l l y : C a l i f o r n i a ,  Oregon, Washington, A l a s k a , and B r i t i s h  2.2  Columbia  Columbia.  California  The  first  major  published  ecological  t i d a l marsh v e g e t a t i o n i n C a l i f o r n i a (1942), who County,  i n v e s t i g a t i o n of  seems to be that of Purer  s t u d i e d twelve c o a s t a l s a l t marshes i n  in  the southern part of the s t a t e .  San  Diego  She d e s c r i b e d at  l e n g t h the ecology and anatomy of the nine p r i n c i p a l  vascular  s p e c i e s i n her study a r e a . The  marshes of Newport Bay,  were d e s c r i b e d by Stevenson and  Vogl  (1966).  a l s o i n southern  (1954), Stevenson  Stevenson  California,  & Emery  (1958),  & Emery (1958) recognized f i v e  p l a n t communities i n the marsh proper -- the Spartinetum,  the  Salicornietum,  the  the  Suaedetum,  D i s t i c h l i d e t u m -- and immersion  and  quantitatively marsh  could  dominated maritima  by and  the Monanthochloetum, and  r e l a t e d the community z o n a t i o n to  soil  factors.  Vogl  f o r frequency and cover, be  separated  Spart ina Salicornia  into  foliosa,  (1966)  and  found  tidal sampled  that  the  three zones: the lower zone the  virginica,  middle and  the  one  by  upper  S a l i c o r n i a v i r g i n i c a and Monanthochloe l i t t o r a l i s .  He  Bat i s one  by  found  6  that  the  zones  floristic  did  not  composition  gradients,  gradually  changed  along  sensu C u r t i s  (1977) i n v e s t i g a t e d the v e g e t a t i o n of the  salt  marsh,  at  the  southern  end  of the  c o a s t , in r e l a t i o n to measured environmental one-metre  elevation  gradient.  v a s c u l a r p l a n t s changed analysis  zonation.  did  She  Further  She  gradually  not  support  the  i n determining north,  Cooper  in  the  Palo  over  that dominance of  elevation; concept  of  rigorous intertidal the  (1926)  provided  a  role  Alto  San  in  brief early Francisco  A l t o area, Hinde (1954) i d e n t i f i e d  three major v e g e t a t i o n a s s o c i a t i o n s , d e s c r i b i n g them and  their  dominant s p e c i e s ( S a l i c o r n i a ambigua, D i s t i c h l i s s p i c a t a , Spartina  leiantha)  with respect to t i d e The was  in  terms  of t h e i r v e r t i c a l  and  distribution  levels.  s a l t marsh at Bodega Head, north  described  a  species d i s t r i b u t i o n s .  d e s c r i p t i o n of the marshes near Palo Also  factors  found  with  Tijuana  California  a l s o s t u d i e d s p e c i e s i n t e r a c t i o n s and  of competition  Bay.  that  (1951).  Zedler  data  but  environmental  c o n s i s t e n t with the continuum concept  & Mcintosh  Estuary  have d i s c r e t e boundaries,  of  San  Francisco,  by Barbour et a l . (1973), r e p o r t i n g on a  by Hamner (unpublished?),  who  sampled  species  cover  study along  elevational transects. In  a s e r i e s of three papers, Mahall  & Park (1976a, b,  d e s c r i b e d the ecotone between Spart ina f o l i o s a and  c)  Salicornia  7  virginica, two  corresponding  to the l e v e l of mean high water, i n  marshes with d i f f e r e n t  northern  San  Francisco  topographical Bay.  p r o d u c t i v i t y of both s p e c i e s frequent  for  T h e i r f i n d i n g s i n d i c a t e d : (1.)  is  open spaces, suggesting  from c o m p e t i t i o n , both  higher  the growing  season;  of higher  the  ecotone,  with  that the boundary r e s u l t s not  soil  salinity  there  a habitat  having  than that of S p a r t i n a  during  (3.) S p a r t i n a  is  less  tolerant  than  r a p i d s a l i n i t y changes, and much l e s s t o l e r a n t  root medium s a l i n i t i e s ;  important  at  (2.) S a l i c o r n i a occupies  considerably  of  low  but from poor environmental c o n d i t i o n s  species;  Salicornia  characteristics in  factor  affecting  the  (4.) s o i l a e r a t i o n i s not an distributions  of  the two  s p e c i e s about the ecotone; (5.)• i n h i b i t i o n of growth r e s u l t i n g from  tidal  immersion may be an important f a c t o r checking the  seaward advance of S a l i c o r n i a . Some recent with  tidal  Ph.D. d i s s e r t a t i o n s i n C a l i f o r n i a have  marsh  vegetation  community composition terms  of  Felton  (1978)  Francisco  energy  Bay  examined  chemistry,  microclimatic  marsh i n order  nature of the r a d i a t i o n a l and energy the  marsh, and the extent  affect  Tijuana vascular  aspects  Estuary:  rather  in  and n u t r i e n t c y c l i n g . conditions  in  a  San  to determine the d e t a i l e d exchanges  to which v e g e t a t i o n  the marsh m i c r o c l i m a t e .  functional  from the p o i n t of view of  or e c o l o g i c a l g r a d i e n t s , but  budgets,  salt  not  dealt  occurring  in  d i f f e r e n c e s can  W i n f i e l d (1980) s t u d i e d  three  of a s a l t marsh-estuarine ecosystem i n the primary  p l a n t s , organic  productivity  of  the  salt  carbon c y c l e , and i n o r g a n i c  marsh  nitrogen  8  cycle.  Newby  mineral  n u t r i e n t s and elements i n t i s s u e s of Spart ina  and  (1980)  Salicornia  investigated  virginica  distributions  in  a  marsh  phosphorus c o r r e l a t e d best species,  and  that  with  the  levels  respect  in  to  Humboldt Bay.  with  nitrogen,  of  fifteen foliosa  the  species  She found that  percent  cover  of  sodium,  calcium,  the  two  and  iron  c o r r e l a t e d only weakly. Macdonald  (1977b) compiled  a comprehensive  and  thorough  summary of what was known to that date of the p l a n t ecology of C a l i f o r n i a c o a s t a l s a l t marshes.  2.3  Oregon  the e a r l i e s t p u b l i s h e d Oregon  -- and probably  Coos  Oregon  Bay r e g i o n .  tidal  of t i d a l marsh v e g e t a t i o n i n  anywhere on the P a c i f i c Coast —  to be the paper by House the  study  (1914), who d e s c r i b e d the marshes The p r i n c i p a l  marsh  vegetation  are  tidal  marshes  describing  the  (1975) with  surveyed the  marsh  the  objectives  communities,  the  unpublished  tidal  inundation  She  determined  to marsh s p e c i e s previous  plant  Ph.D.  (1975).  v e g e t a t i o n of 19 Oregon of  determining  determining  r e l a t i o n s h i p s , and q u a n t i f y i n g the r e l a t i o n s h i p s and  of  r e f e r e n c e s , however, f o r  d i s s e r t a t i o n s of J e f f e r s o n (1975) and E i l e r s Jefferson  seems  and  successional of  salinity  distributions. communities at s e l e c t e d  9  s i t e s by examination the s o i l ,  e n a b l i n g the c o n s t r u c t i o n of  diagrams.  She  corresponded She  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  counting  a  and  approximately  time  frame  existing for  zonation  sequences.  marsh development by  dating  (yielding  dates  approach  to  the  she presented  characterization  communities  rather  of  than  e x i s t e n c e of environmental  association  vegetation  continua  was  of  (Nonetheless,  in  one another  along  Eilers  stands  plots  t r a n s e c t s u s u a l l y suggest  She  felt  terms  of  warranted  by  of  with  species  distinct  to  described  distributions  He  plotted for  species  as  identified  along  the marshes at Nehalem Bay,  environmental  e l e v a t i o n a l gradient dry  the  boundaries.  continuous  gradients  variation  and  weight  species May,  the  July,  distributions and  along the  September,  of  of  primary  using  index of p l a n t performance.  communities, p a r t l y with the a i d  in  by a s s e s s i n g  community s t r u c t u r e , together with an estimate of net productivity.  any  s p e c i e s d i s t r i b u t e d independently of  northern Oregon, by c o n s i d e r i n g the along  the  continua.)  (1975)  vegetation  for  d i s c o n t i n u i t i e s which precluded  homogeneous her  was  tables  m a n i f e s t a t i o n of continuous g r a d i e n t s , l e a d i n g r a t h e r formation  of  vegetation  marsh types comprising 29 d i s c r e t e communities. the  varve  410 and 770 years f o r mature h i g h marshes).  p h y t o s o c i o l o g i c a l , and  that  the  radiocarbon  Jefferson's  six  successional  found that the i n f e r r e d s u c c e s s i o n a l sequences  c l o s e l y with  obtained  detailed  a  He  reference-  10  stand  ordination  correspond  procedure;  these communities were found  c l o s e l y with major s i g n a t u r e  types  recognized  to on  c o l o u r and c o l o u r i n f r a - r e d a e r i a l photographs, p e r m i t t i n g the construction found  of a p h y t o s o c i o l o g i c a l map  that p l a n t  community  species  location,  of the marsh.  distributions,  seasonal  species  development,  Eilers  diversity, net  aerial  p r o d u c t i o n , and amount of net a e r i a l production exported a l l c l o s e l y r e l a t e d to e l e v a t i o n and a s s o c i a t e d t i d a l Vascular Bay,  on the  (1980), nr  2  year"  central  who  production  coast,  was  marshes).  He  v a l u e s on the order of 20 000 Two  Deschampsia  estuary, different  McVay  et  cespitosa  in an under  g  found A p r i l  nr  2  Hoffnagle from 400  year  and  (in  - 1  root standing  crop  2  seeded  Carex  in  on  the  fertilization  Mitchell  following  and  obnupta  location  different  g  g nr .  a l . (1980)  intertidal  t i d a l heights.  reestablishment estuary.  by  recent s t u d i e s have d e a l t with marsh establishment  restoration.  material  investigated  c a l c u l a t e d p r o d u c t i v i t y values ranging  validus  factors.  in s i x marshes in Coos  ( i n d i s t u r b e d marshes) to 1200  1  Sc i r p u s  p l a n t primary  were  dike breaching  transplanted sandy  Columbia regimes  (1982) s t u d i e d  or  salt  dredge River and  at  marsh  in the Salmon River  11  2.4  Washington  The marshes of Grays  Harbour,  on  Washington's  Pacific  Coast, were d e s c r i b e d by Messmer et a l . ( u n p u b l i s h e d ) , u s i n g a classification  system  adapted  from  i d e n t i f i e d t h i r t e e n communities  Jefferson  (1975).  They  w i t h i n seven marsh types,  and  presented graphs of s p e c i e s percent cover along an e l e v a t i o n a l g r a d i e n t , i n d i c a t i n g continuum-type the  distributions.  (Rarely i n  review area has a v e g e t a t i o n d e s c r i p t i o n system developed  by one author been a p p l i e d elsewhere by another, as here  --  with  the  result  d e s c r i b e d by d i f f e r e n t  that d i r e c t comparison  i n v e s t i g a t o r s i s often  The major p u b l i s h e d s t u d i e s of Washington are  those of Burg, Rosenberg  Bellingham  & Fonda (1979)  Bay,  north  a n a l y s i s approach. and  biomass  and  soil  submergence species  Puget  and  water.  a  brackish  Sound,  using  species  cover,  determined  measurements  and  periods.  Their  distributions  a  tidal  were  critical  tide  Below t h i s l e v e l ,  marshes & Fonda  marsh a  soil  gradient  salinity,  texture.  t i d a l data they c a l c u l a t e d results  indicated  in  frequency,  estimates; measured water t a b l e depth,  that  From tidal plant  r e l a t e d to a l l the environmental  f a c t o r s measured except s a l i n i t y noted  difficult.)  (1980).  examined  They recorded  moisture;  elevation  of  done  of marshes  & T r i p p (1976), D i s r a e l i  (1979), and Burg, T r i p p & Rosenberg Disraeli  was  (which was q u i t e low).  l e v e l at 274 cm above mean lower the marsh had  over  70  They low  submergence  12  days  per  year,  had  sandy,  poorly  drained  dominated by S c i r p u s americanus; above 274 fewer  cm,  than 50 submergence days per year, had  drained  soils,  and  (Notwithstanding  dominated  by  the  e l e v a t i o n seems to  siltier, Carex  type d i s t r i b u t i o n of  indicate  a  of  Rosenberg  Puget  analyzed  &  Tripp  (1976)  and  Sound.  They recorded  Tripp  &  southern  s p e c i e s cover values  and  the data by means of a computer-programmed v e r s i o n  field  a n a l y s i s , they  observation  and  ordination  twelve p l a n t a s s o c i a t i o n s , and  production  Curtis  (1957).  prepared a d e t a i l e d  of the d e l t a showing the extent  2.5  species  continuum-  the  &  obtained  better-  lyngbyei.  Burg,  the p o l a r o r d i n a t i o n method of Bray  map  had  species.)  Rosenberg (1980) s t u d i e d the N i s q u a l l y marsh at end  was  marsh  t h i s apparent ecotone, t h e i r p l o t of  importance against  Burg,  was  s o i l s , and  From  recognized vegetation  of each a s s o c i a t i o n .  estimates f o r e i g h t of the  of  They  associations.  Alaska  A Alaska  general was  description  provided  of  by Crow (1977),  hundreds of marshes i n the r e g i o n , The  earliest  following  who few  noted  that  have been  out  of  studied.  treatment of Alaskan t i d a l marshes seems to  be that of Cooper (1931), who mudflats  the P a c i f i c Coast marshes of  recent  described glacial  the  retreat  colonization at G l a c i e r  of Bay.  13  Puccinellia maritima came a  pumila  was  appearing community  the  primary  at s l i g h t l y higher dominated  brachyantherum,  by  colonist, elevations.  Plantaqo  and T r i g l o c h i n maritimum.  zone between the marsh proper  and  with  the  Glaux  Further  up  maritima,  Hordeum  There was  a meadow  forest,  dominated  by  Elymus a r e n a r i u s . Hanson  (1951) i d e n t i f i e d and d e s c r i b e d marsh communities  and  t h e i r zonation  sequences at s i t e s in Knik Arm  Bay  in south-central  Alaska.  Another Kachemak Bay (1977),  who  complexes".  marsh was  d e s c r i b e d by Crow & Koppen  identified  and  Net  production  aerial  described  community complex, the highest being mollis  -  Potentilla  in  vegetation  the  was 661  A list  report.  Crow  of  the  (1968,  the  "community  measured in  2  in  the  each Elymus  D e t r i t u s removal diatom 1971)  flora  described  was was the  of a l a r g e marsh complex i n the d e l t a of the Copper  R i v e r , in s o u t h - c e n t r a l A l a s k a , which was by  nine  g m~  a n s e r i n a complex.  found to be c l o s e to 100%. included  and Kachemak  earthquake  of  1964.  The  u p l i f t e d almost 2  m  r a i s e d marsh surface i s no  longer  f l o o d e d by even the h i g h e s t  being  d e s a l i n i z e d , with a consequent i n v a s i o n of what used to  be lower marsh higher marsh and  habitats  by  upland  zones.  Stephens & B i l l i n g s a  tidal  marsh  on  tides,  species  and  the  characteristic  (1967) r e p o r t e d a v e g e t a t i o n  Chichagof  I s l a n d in southeastern  Three major p l a n t communities, q u i t e d i s c r e t e  in  soil  is  of  the  study  of  Alaska.  appearance,  14  were s u b j e c t i v e l y to  highest)  i d e n t i f i e d , dominated (from lowest  by Carex  Elymus m o l l i s .  The  elevation  l y n g b y e i , Deschampsia atropurpurea, and  s o i l s of  each  community  were  described  p e d o l o g i c a l l y and c h e m i c a l l y . The  only  published  study  of  an Alaskan P a c i f i c  t i d a l marsh i n which data are analyzed o b j e c t i v e l y d e l Moral & Watson (1978), who the  delta  of  the s t a t e .  i n v e s t i g a t e d an i s l a n d marsh i n  by  clustering  method;  these  discriminant  analysis  using  discriminating  variables.  The  community  exception) were r e c o g n i z a b l e on a e r i a l mappable;  boundaries  into  one  another  discrete  species types  were  as  the  (with  one  photographs  types  were  imperceptibly.  data as i f they are d e r i v e d from v e g e t a t i o n of  groupings  an  and  thus  drawn between them, however, were  sometimes a r b i t r a r y because the  series  of  They i d e n t i f i e d e i g h t marsh community types by  supported  grade  i s that  the S t i k i n e R i v e r , i n the southeastern p a r t of  agglomerative  were  Coast  units,"  say  the  often  "While we  found  to  treat  the  consisting  of  authors, "such  a  i s not  always the case."  2.6  British  The Columbia Taylor  Columbia  earliest  published  literature  source  on  British  t i d a l marsh v e g e t a t i o n i s a p p a r e n t l y that of Calder & (1968),  who  p r o v i d e d a q u a l i t a t i v e d e s c r i p t i o n of the  s a l t marsh communities of the Queen  Charlotte  Islands.  (To  15  date  it  seems  that  no other work has been p u b l i s h e d  marshes of t h i s f l o r i s t i c a l l y Pojar coast a  interesting  (1974), working i n the  s a l t marsh community  function.  The p o p u l a t i o n  interspecific  association  constellation phenology, community biology  means  and  of  the  the r e p r o d u c t i v e  calculated  results  and  s t r u c t u r e and  determined,  was  was  L e v e l s of different  i n the form of a  levels,  dispersal  west  b i o l o g y of  for  presented  Polyploidy  parameters were  of each species  the  an aggregation index. were  diagram.  pollination  During  on  s t r u c t u r e of i n d i v i d u a l species  by  sizes  area  i n r e l a t i o n to community  characterized  quadrat  archipelago.)  Tofino  of Vancouver I s l a n d , s t u d i e d  on the  ecology, and  flowering and  the  various  reproductive  summarized.  the 1970's, information  was c o l l e c t e d i n many l o c a t i o n s by  on t i d a l marsh a  variety  of  vegetation government  agencies and p r i v a t e f i r m s ,  and by s e v e r a l graduate students.  Much  is  of  this  Environment  information Canada  summarized  publications  in  which  a  series  inventory  of the  environmental knowledge from a l l a v a i l a b l e sources f o r B r i t i s h Columbia species for  coastal lists,  tidal  estuaries  estuaries.  community  marshes have  (Bell  &  the  surveyed  volumes are planned): Fraser (Hoos & Void  publications  contain  d e s c r i p t i o n s , and maps, as a v a i l a b l e ,  within  been  These  estuaries. so  far  The  following  i n t h i s s e r i e s (more  (Hoos & Packman  1974),  Squamish  1975), Skeena (Hoos 1975), Cowichan and Chemainus  Kallman  1976a),  Nanaimo  (Bell  &  Kallman 1976b),  16  Kitimat  ( B e l l & Kallman  1977),  Courtenay  Leaney  (Morris  most.  (Bell  et a l . 1979), and  Squamish D e l t a marsh has Lim  &  Levings  vegetation  types,  important  species  Levings  different  &  Thompson  Somass (Morris &  in  with  1  been  recognized Carex  both  better  of  the  lynqbyei  geographical  growth  marsh,  includes  a  detailed  the  and  standing  2  day"  1  the  in  f o r net  g dry weight  g m"  ten most  measurements  A r e c e n t l y p u b l i s h e d management plan f o r  Estuary  as  f i n d i n g high values  up to 22.9  than  mapped  extent  of Carex: up to 1323 rates  studied  and  & Moody (1976) made production  areas  season" ,  (1973)  identifying  above-ground production  June.  Campbell  1980).  The  crop.  1976c),  m"  2  in late Squamish  environmental summary  (Habitat  Work Group 1981). The  p l a n t communities of  coast of Vancouver I s l a n d and described  and  photograph  interpretation  Kennedy  mapped  north  of  Courtenay  s p e c i e s , and species. north  of  those  one  by  marsh on the  Kennedy  and  found that at higher  marshes, a f l o r i s t i c  18 e s t u a r i n e marshes on the east  (1982),  associated  she  using  ground  were aerial  truthing.  l e v e l s in both b r a c k i s h and  salt  break occurred at Courtenay, with marshes dominated  more  by  brackish  indicator  south of Courtenay more by s a l i n e i n d i c a t o r  T h i s d i s c o n t i n u i t y she a t t r i b u t e d to higher Courtenay.  On  the b a s i s of community  Kennedy c l a s s i f i e d the e s t u a r i e s s t u d i e d i n t o which  mainland  believed  resulted  from  the  rainfall  composition  eleven  interaction  groups, of s i x  17  physical factors: between  time  of  maximum  discharge,  mean growing season d i s c h a r g e  and  relationship  s i z e of d e l t a , mean  annual t o t a l p r e c i p i t a t i o n ,  r e l a t i v e p r o t e c t i o n from wind  and  wave  frequency  and  energy, d i r e c t i o n and  substrate p a r t i c l e s i z e . and in  root  reserves  She  also  of t i d a l  inundation,  determined  standing  in monthly samples from eleven  crop  communities  five estuaries. The  delta  l a r g e s t t i d a l marsh syst-em in B.C.  of  the  Fraser R i v e r , and  the most abundantly d e s c r i b e d . are:  the  ecological  is  found  in  the  i t i s these marshes that  Some general  reference  overview of the d e l t a by Becker  the comprehensive summary of p u b l i s h e d and  unpublished  works (1971), sources  on the e s t u a r i n e environment by Hoos & Packman (1974), and review of e s t u a r i n e h a b i t a t i n the  context  of  a  are  the  management  plan by H a b i t a t Work Group (1978). Vegetation grouped  for  s t u d i e s done in the Fraser D e l t a area are here  convenience  i n t o two  c a t e g o r i e s : those d i r e c t e d  towards measurement of p r o d u c t i v i t y , biomass d i s t r i b u t i o n , nutrient  cycling;  compositional,  in  group, the  between Point Grey and dominance of the yields  of  490  2  g nr ,  up  those  concerned  with  floristic,  or s y n e c o l o g i c a l c h a r a c t e r i z a t i o n .  In the f i r s t productivity  and  or  major to 1819  or 4.9  Yamanaka  (1975)  measured  the F r a s e r D e l t a  primary  marshes  of  Crescent  Beach, and mapped the zones of  species. g nr  2  tonnes h a " , 1  He  year" , 1  estimated  foreshore  dry  matter  with an average y i e l d of  per year.  (This c o n t r a s t s  with  18  an average lower F r a s e r V a l l e y hay ha"  (Moody 1978).)  1  largest  S c i r p u s americanus not  snow  a  to  total  sampled  f a r behind  by  the  food producing  food  crop  He  He  with  wintering  at  various  ameri-  locations  then evaluated marsh areas for  attempted  rhizome  parameters,  (36%),  for  c a p a b i l i t y on the b a s i s of both  estimating  above-ground  tonnes  biomass of S c i r p u s  source,  geese.  and crude p r o t e i n l e v e l . of  3.8  (32%).  below-ground  preferred  frequented  method  standing  (1977), s t u d y i n g the food resource  geese,  canus ,  about  Yamanaka found Carex l y n g b y e i to make the  contribution  Burton  crop of  to  standing  but  could  rhizome d e n s i t y  find  a  reliable  crop from o b j e c t i v e  not  find  significant  correlations. The who  was  and  Brunswick  Point  concerned with  spatial  marsh was  primary  and  temporal  vegetation.  Moody  harvested  through  growing  the  productivity,  aerial  season,  and  shoot  numbers  contents  nitrogen  decomposition,  distributions  r a t e s , standing crops, and  s t u d i e d by Moody (1978),  of  biomass  was  densities,  the  marsh  periodically  able to r e l a t e growth reproductive  shoot  f o r the major s p e c i e s to such  v a r i a b l e s as s a l i n i t y ,  temperature,  year.  l a b o r a t o r y s t u d i e s i n d i c a t e d that  L i t t e r bag and  f l e s h y s p e c i e s such as fastest,  and  Transplantation observations  Carex  Triglochin lyngbyei  experiments  suggested  and  elevation,  maritimum  and  time  decomposed  of  soft, the  decomposed  the  slowest.  historical  and  present  a s u c c e s s i o n a l sequence commencing with  19  S c i r p u s americanus and S. maritimus. Ogwang  (1979)  disposition Island  i n two b r a c k i s h  and  Brunswick  marsh along harvested  investigated  aerial  Fraser  Point,  the P i t t R i v e r  phytomass Delta  related  s i t e s and years  to such v a r i a b l e s as  salinity,  substrate  nutrient  d i f f e r e n c e s between s p e c i e s which  a  Iona tidal  Fraser).  sequentially during  v a r i a t i o n s i n peak standing  and  the  and  at  and a l s o i n a freshwater  season, and extended the sampling over He  marshes,  (a t r i b u t a r y of  vegetation  production  He  the growing  three-year  period.  crop between s p e c i e s ,  climate, status.  water  regime,  He  found l a r g e  i n the timing of peak  production,  were r e l a t e d to such f a c t o r s as the presence or absence  of overwintered comprised  a  shoots.  high  He found that  proportion  routes  unimportant), export.  He  vegetation,  He i d e n t i f i e d  a  organic assayed  matter the  accumulation,  nutrient  and i n v e s t i g a t e d aspects  program  content  and  main  (fairly detritus  of the emergent  of n u t r i e n t  leaching. has  resulted  of s t u d i e s on the Fraser D e l t a t i d a l marshes  conducted by U.B.C.'s  Westwater  (1978)  literature  reviewed  the  f o r the shoot phytomass as g r a z i n g  A s e r i e s of r e p o r t s by K i s t r i t z and others from  phytomass  (up to 85%) of t o t a l phytomass;  the p r o p o r t i o n v a r i e d among s p e c i e s . disposition  belowground  the  Research on  Centre.  Kistritz  e c o l o g i c a l processes i n  t i d a l marshes, c o n c e n t r a t i n g  on primary p r o d u c t i o n ,  of  phytodetritus, estuarine  food webs, marsh biogeochemistry,  and  nitrogen c y c l i n g .  Kistritz  & Yesaki  (1979) and  the  role  Kistritz,  20  Hall  &  Yesaki  (1983)  reported  on  a  study  of  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 marsh shoot  dominated  by  Carex l y n g b y e i .  indicated  an  ash-free  exported  weight  was  (AFDW)  estimated at 435  i n t o the e s t u a r y , and  In the second category floristic  m;  net  2  sediment.  of s t u d i e s , the e a r l i e s t  detailed  studied  the  vegetation  a line-intercept  frequency  feeding ecology.  sampling  species  and  calculated seed  l y n g b y e i , S c i r p u s v a l i d u s and (1972)  conducted  seed  sources  production f o r ducks  a  vegetation  s u b j e c t i v e community d e s c r i p t i o n s , s p e c i e s and  McLaren (1972) p r o v i d e d  vegetation a  principal  indices. to  be  He  Carex  S. americanus.  foreshore marshes from Point Grey to Crescent  checklists,  Based  d i s t i n c t , divided  He mapped the d i s t r i b u t i o n s of the  the most important  Forbes  the  method, he d e s c r i b e d a  an upper marsh zone, f l o r i s t i c a l l y  by a low c l i f f .  of  marshes from Iona I s l a n d to Brunswick P o i n t ,  as part of a study of duck h a b i t a t and  and  was  d e s c r i p t i o n of the F r a s e r D e l t a marshes i s probably  delta front t i d a l  found  g  detritus  of which 62%  2  the balance  of 634  b u r i e d by  that of Burgess (1970), who  lower and  annual  g AFDW m~ ,  tissue Results  p r o d u c t i o n estimate  per  river  Monthly measurements of  n i t r o g e n and phosphorus were made.  annual net primary  dry  production  marsh  a  growth, d e n s i t y , standing crop, root biomass, and  l e v e l s of carbon,  on  in  primary  similar  maps  treatment  of  the  Beach, p r o v i d i n g  abundance  (not  marshes from Deas I s l a n d to Westham I s l a n d .  survey  very of  ratings  detailed). the  river  21  The by  s a l t marsh at Boundary Bay  Parsons  (1975),  communities, and  soil  who  relating  was  recognized  the  conductivity.  the o b j e c t of a six  plant  v e g e t a t i o n p a t t e r n to t i d e  levels  The  and  mapped  Tsawwassen  salt  d e s c r i b e d by H i l l a b y & B a r r e t t (1976), who of  The  assessed  in F r a s e r D e l t a marshes are those  by Hutchinson  (1982) and  Hutchinson delta  the degree  of  of  Island  with  the  to environmental  plots  was  substrate  were  mapped,  to  variation  texture.  The  and  The  separate  a  in  Species  types were  from  recognized.  distributions related  extent mean  salinity,  and  of marsh v e g e t a t i o n higher  low  water  l e v e l of mean lower high water (MLHW) appeared to lower  marsh, dominated by S c i r p u s americanus and  Potent i11a  Agrostis exarata. Triglochin  relating  procedure,  elevation,  lowest  S c i r p u s maritimus, from a higher latifolia,  of  parameters.  their  corresponded c l o s e l y to the l e v e l of (MHLW).  aim  the  determined by biomass measurement,  which seven main c l u s t e r s or v e g e t a t i o n  statistically  reported  (1982).  p l o t s were grouped by a c l u s t e r a n a l y s i s  types  environmental  recently  B r a d f i e l d & Porter  Lulu  v e g e t a t i o n composition  These  attempt  (1982) examined the b r a c k i s h marshes along  foreshore  composition  was  communities.  most d e t a i l e d s t u d i e s of v e g e t a t i o n and  relations  and  marsh  presence of each s p e c i e s in the marsh, but d i d not  t o d e f i n e or map  study  palustris,  Zones  mar itimum  marsh,  dominated  occurred  dominated  Di st i c h l i s by  Carex  at about MLHW.  by  Typha  spicata,  and  lyngbyei  and  The  low  cliff  22  d e s c r i b e d by Burgess (1970) was  found to occur at  about  this  level. Bradfield  &  s t r u c t u r e of a influence.  Porter  river  (1982)  marsh  Vegetation  i n v e s t i g a t e d the v e g e t a t i o n  under  data  predominantly  consisting  freshwater  of  cover  class  estimates were subjected to a c l u s t e r a n a l y s i s procedure p e r m i t t e d the r e c o g n i t i o n of seven community  types,  within  subgroups, r e f e r r e d  three main groups,  the  as  sedge  zone  in  f l o o d e d and d r a i n e d areas, a grass-willow zone along  crests  of l e v e e s , and a mixed f o r b zone i n areas of poor  drainage and high water t a b l e . species  to  corresponding to  s u b j e c t i v e l y r e c o g n i z a b l e v e g e t a t i o n zones: a regularly  which  mean  calculated  percent  and  cover  presented  For the seven community and  in  percent  tabular  types,  frequency  were  form, as were v a r i o u s  d i v e r s i t y components, i n c l u d i n g t o t a l number of s p e c i e s species  density  or  species  evenness.  alpha The  diversity,  beta  found,  diversity,  d i s t r i b u t i o n s of the community  were p l o t t e d along e l e v a t i o n a l . t r a n s e c t p r o f i l e s .  and types  In a d d i t i o n  to being c l u s t e r e d , the p l o t data were s u b j e c t e d to o r d i n a t i o n by means of p r i n c i p a l components a n a l y s i s averaging between  (RA). trends  environmental appeared  Ordination  to  in  compositional  gradients. be  results  related  The to  (PCA)  and  suggested variation  principal  reciprocal  relationships and  particular  ordination  s u b s t r a t e drainage and  axes  to t o t a l  p e r i o d of inundation, suggesting that these two components the  hydrologic  vegetation pattern.  regime  operate  independently  to  E l e v a t i o n above c h a r t datum was  of  control not  found  23  to be a r e l i a b l e p r e d i c t o r of  vegetation.  24  3.  3.1  STUDY AREA  Location  The the  THE  F r a s e r D e l t a i s l o c a t e d on the southeastern shore  Strait  of Georgia,  i n southwestern  B r i t i s h Columbia,  extends  from the c i t y of Vancouver southwards  States  border.  07' N by  I t s geographic  123° 05' W;  to  the  and  United  c e n t r e i s at approximately  the l o c a t i o n s sampled i n t h i s  of  study  49° are  a l l w i t h i n 12 km of that p o i n t ( F i g . 1).  3.2  Formation  and Development of the F r a s e r D e l t a  Southern  Georgia  Strait  and  the adjacent mainland  b u r i e d beneath P l e i s t o c e n e i c e u n t i l about (Mathews  et  a l . 1970).  Post-glacial  e s s e n t i a l l y complete by 8000 years shoreline  has  13 000  isostatic  before  years  ago  uplift  was  present,  and  (ibid.)  The d e l t a that e x i s t s today began to fan out from the  highlands  years ago, (980  at  what i s now  New  submerged  portions,  380  about 380  feet  (115 m)  square  with  ( i n areas where sediment accumulation  probably  a  gap  Westminster about 8000  covers an area of about  including  2  thickness of  and now  km ),  the  remained c l o s e to i t s present l e v e l during the  past 5500 years  in  lay  is  an  miles average  complete)  (Mathews & Shepard 1962).  25  Figure  1 - Map  of the study  area  26  The  a c t i v e f r o n t of the d e l t a extends some  western  perimeter,  between  i n a c t i v e f r o n t about Roberts  and Crescent  very g e n t l y i n c l i n e d the  dikes  slope.  The  The  Beach.  its  Point Roberts;  an  Point  Along the western d e l t a f r o n t , a  i n t e r t i d a l zone extends about 6  the southern  km  from  delta front  is  wide. F r a s e r River  delta  6  i s c u r r e n t l y d e p o s i t i n g about 700  m) 3  front,  of loose s i l t y representing  sand and  rapid  advance  occurs  mud  x  10  p.  s  annually  a load of about 21 x  tonnes ( d e r i v e d from Mathews & Shepard 1962 of  along  long faces southward between  e q u i v a l e n t zone along  cubic f e e t (20 x 10 the  Point Grey and  km  almost to the edge of the much steeper d e l t a f o r e -  about 4 km  at  13 km  23  1424).  A  10  6  zone  in the v i c i n i t y of the mouth of  the  main channel of the r i v e r ; there are areas, however, where the delta  f r o n t i s r e t r e a t i n g (Luternauer  measured  rate  c h a r t datum. foot  (91.4  &  of advance v a r i e s a c c o r d i n g A rate of 28 f e e t (8.5 m)  m)  Murray  contour  The  to the depth below  per year  over a 30-year p e r i o d was  Mathews & Shepard (1962); the r a t e was  1973).  at  the  300  determined  much lower at  by  shallower  depths.  3.3  Extent  The found  of T i d a l Marshes in the F r a s e r D e l t a  e x i s t i n g t i d a l marshlands of  mainly in three a r e a s :  1 km wide, along  the  western  the  Fraser  Delta  i n a broad b e l t , averaging delta  foreshore;  in  a  are about wide,  27  braided mouth;  portion  and  the  along  the  n o r t h shore  marsh v e g e t a t i o n its  are  mainly  generally  the  dikes;  base of  around  to  Boundary B a y ,  b r a c k i s h marshes marshes.  lies  p a r t of  From an Luternauer western  this  (1976)  preceding  25  years,  retreats.  Brunswick  One  Point,  90 ha f i r s t  km  to at  Narrow the  its its  fringes  main c h a n n e l  locations  to  regime,  1.5  that  been  area  form  marked by d r i f t w o o d ) in  the  are  marshland  limit  (1979)  seems  to  determined  from Westham I s l a n d the  however,  aerial  although  with  the  lands  of  lower  m below  limit  that  of  of  salt  may a c c o u n t  the  of  that  Scirpus  the  western  a  rapid  sand f l a t s in  americanus  Kellerhals of  stable  &  series marsh  p a r t of  over  were some l o c a l  apparently  an a r e a  Medley  marsh edge a l o n g  generally there  of  photographs,  a p p e a r e d on a e r i a l p h o t o g r a p h s  (in  flats  The l o w e r  Swinbanks  1.0  of  had  At Boundary B a y ,  evidence  limit  and c o n c l u d e d t h a t least  delta  for  difference.  where  vegetated  1978).  at  upper  dike.  concluded  front  the  several  examination  delta  densely  the  V a r i a t i o n s in t i d a l  least  tidal  close  1  Boundary Bay.  salinity:  lower marsh l i m i t a t  and  than  a g r i c u l t u r a l and r e s i d e n t i a l  v a r y a c c o r d i n g t o water  at  of  less  found elsewhere a l o n g  s u r r o u n d e d by p r o t e c t i v e  the  main c h a n n e l ,  distributaries.  The  is  river's  and i n a r a t h e r n a r r o w b a n d ,  widest, of  of  is about  and  was  1969  (Moody  (1962)  found  former beach  ridges  Murray of  advances  occupying  by  the the  growth  1948,  &  is  advancing  the  bay,  but  over receding  the in  28  t h e e a s t e r n p a r t , a s i n d i c a t e d by an about  0.7  estimated  m  high.  The  t o be a t l e a s t  active  erosional  r a t e of r e c e s s i o n i n t h i s  0.75 m i . ( 1 . 2 km) o v e r  cliff  a r e a was  the last  4350  years. The  total  estimated  area  of  the  delta  tidal  m a r s h e s h a s been  a t 2683 h a , o f w h i c h t h e w e s t e r n d e l t a  a c c o u n t f o r 1664 h a , t h e M a i n Arm r i v e r m a r s h e s Boundary  Bay  237  ha  (Yamanaka  mainly and  of  the delta  lands  782  marshes ha, and  1975, K i s t r i t z 1 9 7 8 ) .  m a r s h e s were more e x t e n s i v e b e f o r e diking  front  the a r r i v a l  of  The  Europeans;  f o r f l o o d c o n t r o l and r e c l a m a t i o n ,  d u r i n g t h e 1 8 9 0 ' s , e l i m i n a t e d a b o u t 221 ha o f s a l t m a r s h  629 ha o f f r e s h w a t e r m a r s h  activities  continue  to  construction,  since the  early  training river  walls  and  channels,  sedimentation,  alter  training described  species walls  and  in detail  by  several  jetties  Some  in  Levings  the  (1980)  of  erosion,  i n o t h e r s , and  the  Fraser and  of the  These a c t i v i t i e s  retreat of  The  jetties,  with dredging  distribution.  composition.  Other  marshlands.  patterns  h a v e l e d t o m a r s h g r o w t h i n some a r e a s , changes i n  a l . 1976).  of  coupled  modified  salinity  the  1900's,  causeways,  has  and  (Romaine e t  effects  Estuary Tamburi  of were  &  Hay  (1978). Much protected reserve.  of from Some  t h e t i d a l m a r s h a r e a o f t h e F r a s e r D e l t a i s now development areas,  under  some  form  though, . a r ed i r e c t l y  e x a m p l e by a i r p o r t d e v e l o p m e n t .  As i n d i c a t e d  of  government  threatened, f o r i n section  3.2,  29  however,  the  long-term  delta  outlook  itself for  i s growing r a p i d l y ,  the  marshes  is  one  so perhaps the of  continuing  expansion.  3.4  P h y s i c a l Environment of the F r a s e r D e l t a T i d a l Marshes  3.4.1  Climate  Summers in the study area are mostly sunny, d r y , and warm (seldom h o t ) ;  winters are d u l l ,  rainy,  and u s u a l l y q u i t e  Some major c l i m a t i c v a r i a b l e s are summarized i n Table The area l i e s w i t h i n  the  Csb  c l i m a t i c type in the c l a s s i f i c a t i o n The  mediterranean  designation  (mediterranean  is  a  than  in  the  1  subhumid)  system of Koeppen consequence  pronounced summer drought; mean temperatures are lower  I.  mild.  (1936). of  the  considerably  m e d i t e r r a n e a n - c l i m a t e s of Europe  (Hoos &  Packman 1974).  1  The data are from a s t a t i o n about 2 km east of the Brunswick Point marsh. Rose (1975) has pointed out, however, that there are m i c r o c l i m a t i c d i f f e r e n c e s between c o a s t a l wetlands and the adjacent d r y l a n d s , with the wetlands exhibiting a narrower range of temperatures both d i u r n a l l y and s e a s o n a l l y . Daytime maxima are lower as a r e s u l t of e v a p o r a t i o n and t r a n s p i r a t i o n , while n i g h t - t i m e minima are higher because of the greater heat storage c a p a c i t y of s a t u r a t e d s o i l .  30  Table I - Environmental data summary f o r the study a r e a : mean d a i l y temperature, mean d a i l y maximum temperature, mean d a i l y minimum temperature, mean t o t a l p r e c i p i t a t i o n  dan . Feb  Mar  Apr  8 . 6 11.8  May  <Jun  T  2. 3  4. 2  5. 7  Tmax  5. 2  7.6  9 . 5 12.6  0.7  1. 8  4 . 5  6 . 9  9.9  99  83  52  38  45  Tm i n  -0.7  P  1 15  Jul  Aug  Sep  Oct  Nov  Dec  5.9  4 . 3 9 . 4  9. 1  6 . 6 13.4  14.6  16.4  16 . 1 13.5  9.6  16 . 7 19.2  21.6  2 1.2 18.4  13.6  11.1  11.0  24  3 1  T = mean d a i l y t e m p e r a t u r e ( C ) Tmax = mean d a i l y maximum t e m p e r a t u r e T m i n = mean d a i l y minimum t e m p e r a t u r e P = mean t o t a l p r e c i p i t a t i o n (mm)  8.6  5. 7  2. 7  1 .9  5.3  50  99  125  142  903  (C) (C)  (Data source: Environment Canada, Atmospheric Environment V a n c o u v e r , B.C.; r e c o r d s f o r L a d n e r M o n i t o r S t a t i o n . )  3.4.2  The River  River and Marine  interacting and  elevational, different  Strait  The  influences  produce  and seasonal v a r i a t i o n  areas i n the d e l t a ;  the  .  Fraser  geographical, regime  of  i s the p r i n c i p a l  c h a r a c t e r of the marshes  in  localities. Fraser  River  drains  an area of about 233 000 km , 2  precipitation  form of snow (Ages & Woollard 1976). snow pack i n s p r i n g  rises  the  i n the s a l i n i t y  w i t h i n which about t w o - t h i r d s of the the  of  complex  this variation  c o n t r i b u t o r to the very d i f f e r e n t different  Service,  Influences  and opposing  Georgia  Year  rapidly  during  and  summer,  the  falls  in  With the m e l t i n g of river's  May with a pronounced  discharge  peak, c a l l e d the  31  freshet, July  i n l a t e May  or June; d i s c h a r g e remains  high  and August, d e c r e a s i n g to a low i n December.  discharges  (1912-1956 3  15 200 m s" 3  on May  1  out  in  a  600  m s" 3  i n summer; the peak flow recorded  1  31, 1948  highly  1  was  (ibid.).  Upon e n t e r i n g Georgia S t r a i t , fans  Mean d a i l y  average at Hope) vary between  in winter and 8800 m s~  through  the s i l t y brown r i v e r water  visible  plume over the c l e a r marine  water, spreading sometimes to the opposite s i d e of the S t r a i t , 30 km away (Tabata 1972); the Boundary Bay  (Swinbanks  plume  seldom  if  ever  enters  1979).  Since.the f r e s h r i v e r water i s l e s s dense than sea water, it  floats.  A  i n t e r f a c e between  certain  brackish;  of mixing takes place at the  the o u t f l o w i n g r i v e r water and the sea water  beneath, so that the f r e s h more  amount  in  surface  compensation  water for  gradually  the entrainment of sea  water in the outward-flowing water mass at the water  below  phenomenon  flows known  pronounced:  for  upstream. as  example,  Sand Heads i n August, measured  as  s a l i n i t y was upstream  as  0.0  the  surface,  salt  T h i s process g i v e s r i s e to the  salinity  wedge.  in a f a i r l y  salinity  becomes  to  a  The  effect  is  t y p i c a l o b s e r v a t i o n at depth  of  3.1  m  was  p a r t s per thousand ( p p t ) ; at 9 m depth, the  25 ppt. Annacis  The  wedge  Island,  has 10-15  Marsh, when r i v e r d i s c h a r g e i s low  observed  as  far  km upstream from Ladner  (Ages & Woollard 1976).  Detailed s a l i n i t y distributions were p l o t t e d by Ages  been  in  the  Fraser  Estuary  (1979) from o b s e r v a t i o n s made i n 1976 and  32  1977.  The  river  at  data  least  indicate  as  far  during  out  Sand Heads, s u r f a c e  during  this  brackish  estuary  at  discharge, mildly  the  waters  Surface  were  i n the  almost  substrate  Point  (1975) and  Swinbanks  range  August  of  Even as  and  0.0  far ppt  September,  penetrate  the  outer  t h e month of  lowest  0-4  recorded  of L a d n e r  and  area  in  lower  invariably  exceeded  December,  being  vicinity  Brunswick  by  i n the  were  seldom  occasionally  surface;  brackish) the  salinity  Beginning  salinities  u p s t r e a m as  waters  p e r i o d mid-May t o m i d - A u g u s t .  period.  slightly  surface  Steveston  fresh as  the  as  that  ppt  at  the  (still surface  only as  far  Marsh.  surface  water  were d e t e r m i n e d  salinities  by  Levings  ( 1 9 7 9 ) , whose r e s u l t s  in  the  & Coustalin  indicated a  mildly  t  brackish  environment  w e s t e r n p o r t i o n s of mildly  saline  southern  at  at  the  different  O'Connell  low  t i m e s of  (1975) and  from  during  freshet.  they  the  determined  ( w i t h one  In  mudflats, year  Swinbanks  the  salinity)  in northern  moderately  12-20  ppt that  brackish  salinity) surface  in  and to the  substrate  closely  to  surface  salinity  measurements  water  ebb.  the  Fraser  a  found  very  on  Boundary Bay  input  (ca.  Swinbanks  tide  ppt  marsh, and  corresponded  salinities On  the  environment  portion.  salinities  ( c a . 0-8  River area  salinity  anomalous r e a d i n g  in t i d a l  pools  low  (1979) i n d i c a t e t h a t i s minimal  or  of my  Boundary  values  ranging  of  at  39  ppt);  the  taken  tide  freshwater  nonexistent, Bay  West  from  23  highest  by  even  transect t o 33  ppt  readings  33  were  obtained  on  warm  days i n June.  S l i g h t l y lower values  (20-24 ppt) were recorded along a t r a n s e c t 5 km f u r t h e r p o s s i b l y r e f l e c t i n g the i n f l u e n c e of the minor r i v e r s  east,  emptying  i n t o Mud Bay.  3.4.3  The mainly  Tides  tides  in  the  semi-diurnal"  Hydrographic  study  (Ages  Service  area are d e s c r i b e d as "mixed, &  1981),  Woollard  which  complete  t i d a l o s c i l l a t i o n s d a i l y with  height  and  time  reaching  d e c l i n a t i o n of the moon  has  Hydrographic  S e r v i c e 1981).  In  when  summer,  f o r the  there  a r e "two  inequalities  greatest  passed  Canadian  values  i t s minimum"  both when  in the  (Canadian  the days are long and warm, the lowest  t i d e s occur d u r i n g the day circumstances  the  means  1976,  --  a  beneficial  marsh p l a n t s  combination  of  (and f o r b o t a n i s t s ) .  In  winter the lowest t i d e s occur at n i g h t . The  detailed  considerably study a r e a .  character  of  the  from one sampling l o c a l i t y Thus the t i d a l  tidal  curve  varies  to another w i t h i n the  range f o r l a r g e t i d e s i s 4.4 m a t  Crescent Beach i n Boundary Bay, with a mean water l e v e l of 2.3  34  m above c h a r t datum; delta  the  range  1  at Sand Heads on the a c t i v e  edge of the  i s 4.8 m, and the mean water l e v e l i s 2.9 m  above c h a r t datum (data source: Canadian Hydrographic 1981).  Moreover,  because of the strong summer peak i n F r a s e r  River d i s c h a r g e ( S e c t i o n 3.4.2), summer lower  river  Service  tide  levels  i n the  are higher than the l e v e l s elsewhere at the same  time, and a r e higher than winter t i d e l e v e l s  i n the same a r e a .  The e f f e c t  For example,  i s most pronounced on low t i d e s .  high water l e v e l s a t higher than  Deas  Island  (near  Ladner  June  Marsh)  are  than winter highs by about 0.4 m; June lows are higher winter  lows  Hydrographic  by  about  Service  1977).  h e i g h t s i s l e s s pronounced mouth;  at  Boundary  Bay  at  1.4  m  The  (data  from  Canadian  summer e l e v a t i o n of t i d a l  Steveston,  nearer  i t i s not s i g n i f i c a n t .  the  river  Winter low  water l e v e l s are at about the same l e v e l throughout the area, more or l e s s as a consequence  study  of the d e f i n i t i o n of c h a r t  datum. The sampling  considerable  variation  location  another  inconvenient,  since  to  1  within  tidal the  regime study  from one area  is  i t p r e c l u d e s any d i r e c t comparison with  respect to e l e v a t i o n of sample (My s o l u t i o n  in  plots  from  different  areas.  to t h i s problem i s e x p l a i n e d i n S e c t i o n 7.3.)  Chart datum i s d e f i n e d i n Canada as the plane of lowest normal tides (Canadian Hydrographic S e r v i c e 1981). T h i s i s not the same as c h a r t datum i n the U n i t e d S t a t e s , where the l e v e l of mean lower low water i s used (Tide T a b l e s 1976). Thus American c h a r t datum i s a l i t t l e higher than Canadian.  35  3.4.4  As very  Sedimentation and  indicated  Substrate  in Section  3.2, the F r a s e r River c a r r i e s a  l a r g e load of sediment i n t o the d e l t a each year; most  this  is  suspended load  (Ages & Woollard  1976).  Up t o 10% of  the suspended load may be c l a y , and the r e s t i s about sand and s i l t  (Mathews & Shepard 1962).  reaches  delta  the  during  Presumably i t i s a t sediment d e p o s i t i o n place,  this  the time  of  year  (Pretious, that  most  equally material 1972). of  the  i n the r i v e r and d e l t a - f r o n t marshes takes  the dense marsh v e g e t a t i o n  tidewaters  Most of t h i s  freshet  of  a c t i n g as a sediment t r a p as  r i s e over the marsh.  At Boundary Bay the sediment c o n t r i b u t i o n from the Fraser River  may  supplied  be by  depositing  slight the  silt  (Swinbanks  smaller and  Murray  of  differing  the  and  marsh  sediments are  Nicomekl  by e r o s i o n  sedimentological  give r i s e to d i f f e r i n g  Because of e r o s i o n , net or long-term Several  and  but  rivers,  of the c l i f f s at  itself  (Kellerhals  &  1969).  The area  Serpentine  clay,  Point Roberts and even  1979),  r a t e s of v e r t i c a l marsh a c c r e t i o n .  i t i s necessary to d i s t i n g u i s h between the  accretion  centimetres  regimes w i t h i n the study  of  mud  rate may  and be  the  short-term  deposited  s i n g l e season, only to be subsequently eroded  rate.  locally  away.  in a  Burgess  (1970) c i t e d the knowledge of l o c a l r e s i d e n t s that the surface of  the  Westham  Island  marsh had r i s e n by 4 f e e t (1.2 m) i n  36  under 35 years; i . e . about the same f i g u r e Brunswick  advance  Moody (1978) quoted  - 1  (perhaps d e r i v e d from the  Point.  accumulation.  3.5 cm y e a r .  same  sources)  Presumably these r a t e s represent short-term  Using the known  rate  of  and assuming a constant average  lateral  delta-front  s l o p e , Burgess  c a l c u l a t e d a long-term sedimentation r a t e of 0.004 mm)  per  year.  within  T h i s c a l c u l a t i o n would be an average  the  relatively  narrow  (1970)  feet  e n t i r e foreshore zone from the d i k e to the edge of front;  for  (1.2  f o r the  the  delta  s t r i p of t i d a l marsh,  however, the r a t e c o u l d w e l l be h i g h e r . At Boundary Bay, K e l l e r h a l s & Murray short-term  1  long-term  influences  Basan (1978). effect  on  by the  plants  sedimentation  and  Among the mechanisms they d i s c u s s e d of  vegetation  on  c u r r e n t competence and s e t t l i n g  the t r a p p i n g by m a c r o i n v e r t e b r a t e s matter, with subsequent The range  animals  there;  were reviewed by Frey & were:  wind-generated  impedance of c u r r e n t flow by v e g e t a t i o n , with of  over  1  accumulation w i t h i n t i d a l marshes i s thought to  be enhanced s i g n i f i c a n t l y  damping  r a t e of 0.42 mm y e a r "  4350 y e a r s .  Sediment  biotic  a  mean sedimentation r a t e of 5.0 mm y e a r " , but they  c a l c u l a t e d a much lower the l a s t  (1969) determined  the  waves; the  resulting  loss  out of suspended l o a d ; and of  suspended  particulate  d e p o s i t i o n i n the form of f e c e s .  a l l u v i a l sediments  of the F r a s e r D e l t a t i d a l marshes  i n t e x t u r e from sands t o  clayey  silts,  nonexistent development of m i n e r a l h o r i z o n s .  with  weak  to  The c o a r s e s t and  37  most  highly  sorted  sediments  are  found  at  Boundary Bay,  probably as a r e s u l t of both reworking by winter storm, waves, and  the absence of a major source of suspended mud  (Swinbanks  1979). At the highest  e l e v a t i o n s , s e v e r a l centimetres  m a t e r i a l may accumulate at the s u r f a c e . decomposing marsh v e g e t a t i o n accumulations (mainly  Zostera  Mineral of  of d r i f t w o o d  organic  The humus produced by  i s sometimes augmented  sediments may i n c o r p o r a t e matter,  consisting  considerable  of decaying r o o t s ,  In  prevail,  a  a  and  waterlogged this  a n a e r o b i c a l l y , g i v i n g r i s e to c o a l - b l a c k  In  from and  and  the  growth  storms  material.  sediments reeking  during  As d i s c u s s e d this  summer and f a l l  bury  of  of  by K e l l e r h a l s & results  of filamentous  surface,  green  followed  by  the a l g a l mats beneath a l a y e r of  Each varve thus represents  deposition.  river  which  layers  stratification  In s p r i n g and e a r l y summer the  surface.  decomposes  d i s p l a y s a pronounced varve-  blue-green a l g a l mats on the marsh  sand.  and  organic  (1962) f o r Boundary Bay,  winter  rhizomes,  environment,  material  l i k e s t r a t i f i c a t i o n : a l t e r n a t i n g narrow h o r i z o n t a l  Murray  amount  sulphide. some areas the s u b s t r a t e  mineral  local  marina) from o f f s h o r e .  conditions  hydrogen  by  and by washed-up r a f t s of e e l g r a s s  shoot bases, and a l g a l mats. reducing  of organic  algae  re-colonize  the  an annual c y c l e of growth  S t r a t i f i c a t i o n observed i n the marshes of the  mouth and d e l t a foreshore,  however, presumably  reflects  38  the  rather d i f f e r e n t annual c y c l e p r e v a i l i n g there,  sedimentation peaks with  3.5  Transect  3.5.1  which  the summer f r e s h e t .  -  Locations  in  and S i t e D e s c r i p t i o n s  S e l e c t i o n of L o c a t i o n s  Sampling  locations  were  selected  at  Ladner  Marsh,  Brunswick P o i n t , and Boundary Bay, a l l i n the M u n i c i p a l i t y  of  Delta,  of  British  representing variation  Columbia  ( F i g . 1), with  the wide range of v e g e t a t i o n a l  in  the  salt-water  objective  and  environmental  d e l t a marshes, from areas of mainly f r e s h -  water i n f l u e n c e i n and near mainly  the  influence  the  Fraser  on  River  Boundary  Bay  to  areas  and  of  Georgia  Strait.  3.5.2  Ladner Marsh  Ladner Marsh ( F i g . 2) was sampled near the north Ferry  Road.  Transect  A  single  transect perpendicular  J , extends 45 m from a swampy  thicket  end of  to the r i v e r , dominated  by  cottonwood, willows and red o s i e r dogwood to the bare mud of a slough.  The dominant s p e c i e s along  fluviatile,  the t r a n s e c t are Equisetum  A g r o s t i s a l b a , and Carex l y n g b y e i .  Somewhat l e s s  1  39  important are Scirpus palustris,  and O e n a n t h e  validus,  part  latifolia,  showy  inflorescences  of t h e m a r s h , and s e v e r a l m i n o r  present.  The  b a r e mud  colonized  by E q u i s e t u m  Eleocharis  sarmentosa. Lythrum s a l i c a r i a  s u a v e a r e c o n s p i c u o u s by t h e i r higher  Typha  species  a t t h e l o w e r edge o f t h e marsh  and  Sium  in  the  are  also  i s being  fluviatile.  Figure 2 - L a d n e r M a r s h , v i c i n i t y of T r a n s e c t J . Foreground: Lythrum s a l i c a r i a , Carex l y n g b y e i . Background: Populus t r i c h o c a r p a . Right: Scirpus validus  40  3.5.3  Brunswick Point  Brunswick Point  i s a p e n i n s u l a r marshland bounded on  north by Canoe Pass, a minor d i s t r i b u t a r y of the F r a s e r and  on the west and  gradient  south by Georgia  Strait.  from predominantly fresh-water  Transect  D samples the high marsh v e g e t a t i o n on the  lyngbyei,  Potentilla  pac i f i c a ,  E l e o c h a r i s p a l u s t r i s , A q r o s t i s a l b a , and Typha  lat i folia  present  include  and  Lythrum  Sium  suave,  Bidens  north  north  composed  Juncus  scattered  salicaria.  is a  south.  s i d e of the p o i n t , where the community i s mainly Carex  River,  Thus there  i n f l u e n c e on the  s i d e to b r a c k i s h or marine i n f l u e n c e on the  the  of  balticus, clumps  of  Other s p e c i e s a l s o  cernua,  and  Juncus  art iculatus. Transect  A,  the  from north to south.  longest, On  runs 1420  the r i v e r s i d e ,  m a c r o s s the p o i n t  bare  mud  is  being  c o l o n i z e d mainly by S c i r p u s americanus, and a l s o by E l e o c h a r i s palustris;  these  are  the  f r i n g e of t h i s marsh a r e a .  principal Higher up  and A q r o s t i s a l b a are the dominant  species  in the  ( F i g . 3), Carex  species  in  an  lowest  lyngbyei extensive  community which a l s o i n c l u d e s T r i g l o c h i n maritimum, S a g i t t a r i a latifolia,  Bidens cernua, L i l a e o p s i s o c c i d e n t a l i s , P o t e n t i l l a  p a c i f i c a , and common  in  several  channels.  other  species.  Ruppia  Towards the south end  ( F i g . 4), S c i r p u s maritimus becomes important. end,  the  mudflats  maritima  is  of t h i s t r a n s e c t At  the  south  are being c o l o n i z e d by Sc i rpus amer icanus  41  and  are  covered  Floodwaters from the  by  here  dense  mats  are f a i r l y  of  filamentous  algae.  f r e s h , sweeping around the p o i n t  river.  Transect B runs p e r p e n d i c u l a r to the dike i n an evidently  marine  influence  on  the  ( F i g . 5).  Nearest  the  dike  Distichlis  spicata  and  Atriplex  community composed mainly virginica,  mat  Tr i g l o c h i n and  dominant patula.  Distichlis  species  are  F u r t h e r out  spicata,  is a  Salicornia  At the lower  end of the  up i n t o clumps of S c i r p u s  maritimus  maritimum.  The mud  i s covered with an  i s being c o l o n i z e d by S p e r q u l a r i a canadensis.  out on the t i d a l  of  south s i d e of the p o i n t  and T r i g l o c h i n maritimum.  t r a n s e c t the marsh breaks and  of  the  area  f l a t , well  beyond  the  edge  of  algal  Further  the  marsh  proper, S c i r p u s americanus and S. maritimus are c o l o n i z i n g  the  mud. A  fourth  transect,  Transect  C,  i s l o c a t e d i n the same  general area as Transect B, but runs at an acute angle to dike,  from  the area of s a l t - w a t e r i n f l u e n c e towards the area  of fresh-water halophyte  influence.  The  f l o r a varies accordingly,  from  communities s i m i l a r to those d e s c r i b e d f o r Transect  B to a S c i r p u s maritimus maritimum transect  the  community  Sc i rpus  resembling  amer icanus  -  Triglochin  that i n the southern  p a r t of  A.  From the a i r , the Brunswick Point marsh ( F i g . 6) d i s p l a y s a prominent p a t t e r n of c i r c u l a r b l o t c h e s , which correspond dense  stands  of  mainly Carex l y n g b y e i or S c i r p u s  to  maritimus.  42  Presumably these stands are rhizomatously.  colonies  Johannessen  (1964)  that  have  considered  spread  such c i r c u l a r  c o l o n i e s at the edge of a marsh to be evidence of r a p i d expansion.  At  Brunswick  conspicuous in the higher, be  r e l i c t s of a previous  3.5.4  The  Boundary  of  P o i n t , where c i r c u l a r  Boundary Bay  112th  Street,  marsh was  East  sampled at two  Boundary Bay  study area in an area  marine i n f l u e n c e i s moderated by  i n t o the bay, the  part  of  large  species  high  the  the  l o c a t i o n s , here  West.  where the  marsh,  driftwood  to  next  G,  the to  deposits.  foot  predominantly  freshwater input  Three t r a n s e c t s , E, F, and  c h a r a c t e r i z e d by in  may  i s l o c a t e d near the  more or l e s s p e r p e n d i c u l a r  ' highest  are  episode of r a p i d expansion.  Delta,  ( F i g . 7).  features  o l d e r p a r t s of the marsh, they  East and  The  streams  marsh  Bay  Boundary Bay  c a l l e d Boundary Bay  out  from  nearby  extend south dike.  Here  the  dike,  The  principal  marsh i s u s u a l l y A t r i p l e x p a t u l a ;  important are Carex l y n g b y e i , S a l i c o r n i a v i r g i n i c a ,  is  also  Potentilla  p a c i f i c a , A g r o s t i s a l b a , and  Puceinellia  lyngbyei,  dense patches of Juncus q e r a r d i i ,  often  with  dominates  elevations, maritimum,  occasional the  middle  Distichlis Glaux  At  slightly  Carex  lower  s p i c a t a , Plantago maritima, T r i g l o c h i n  ma r i t i ma,  Salicornia virginica  levels.  nuttalliana.  Sperqularia  canadensis,  form a c h a r a c t e r i s t i c s p e c i e s  and  assemblage.  Figure 3 - Brunswick Point North, v i c i n i t y of Transect A. Carex lyngbyei i s dominant. Conspicuous i n f l o r e s c e n c e s : Lythrum s a l i c a r i a (magenta); Sium suave (white)  44  Figure 4 - Brunswick Point South, v i c i n i t y of Transect A. T r i g l o c h i n maritimum forms i s o l a t e d clumps at marsh edge; clumps coalesce i n t o continuous stand ( r i g h t background)  45  Figure  5 - Brunswick Point South, Transect l a t e winter  B.  Scene i n  Figure 6 - A e r i a l photograph of Brunswick P o i n t , showing c i r c u l a r p a t t e r n i n g of marsh v e g e t a t i o n . Clumps are mainly Carex lyngbyei and S c i r p u s maritimus  47  Figure 7 - Boundary Bay East, Transect E. Mixed grasses in foreground; Carex lyngbyei (yellow-green) i n m i d - p o r t i o n of t r a n s e c t ; 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, Plantago maritima at f a r end of t r a n s e c t Tdark green)  48  Several  other  in other  s p e c i e s a r e a l s o found.  saline-influenced  Spergularia  (Fig. 8). in  areas,  are  and  are  covered  colours  the  mud  canadensis  summer diatom bloom  The mudflats here, as  Zostera americana  being  colonized  by  by  a l g a l mats.  surface  A  pinkish-brown  and Ruppia maritima a r e p l e n t i f u l  channel bottoms and w a t e r - f i l l e d d e p r e s s i o n s .  As noted by  K e l l e r h a l s & Murray (1962), the marsh i n t h i s area i s eroding; I observed  several  centimetres  of  undercutting  during  one  season of f i e l d w o r k . The  Boundary  Bay  West  study area, at the foot of 72nd  S t r e e t , D e l t a , i s probably the most m a r i n e - i n f l u e n c e d tidal  marshes  thousand  in  this  were determined  study.  Salinities  of the  of 22 p a r t s per  f o r bay water sampled  here  at  high  t i d e i n summer. The higher marsh here, along T r a n s e c t H ( F i g s . 9, 10), characterized Salicornia  by  virginica,  integrifolia. Atriplex  o l d bleached  At  patula,  Salicornia lowest p a r t Plantago canadensi s,  Spergularia  middle and  i s densely of  the  mar i t ima, and  driftwood, marina,  elevations,  Distichlis  contiguous  with marsh  Salicorn ia  Tr i g l o c h i n  Atriplex patula, and  Grindelia  Salicornia  virqinica,  spicata  infested  is  predominate;  Cuscuta  salina.  i s characterized  virqinica,  maritimum.  The by  Spergular i a  Colonizing  s a n d f l a t s are clumps of S a l i c o r n i a and Plantago.  the  the  49  Figure 8 - Boundary Bay E a s t . 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 g i n i c a , with patches of f i l a m e n t o u s green and blue-green a l g a e . Black reducing mud i s covered by brownish diatom mat  Figure 9 - Boundary Bay West, v i c i n i t y of Transect H. A t r i p l e x p a t u l a dominates the d r i f t w o o d zone i n foreground. F a r t h e r out, S a l i c o r n i a v i r g i n i c a i s i n f e s t e d by y e l l o w i s h Cuscuta s a l i n a  51  Figure  10 - Boundary Bay West. G r i n d e l i a flowers amid the d r i f t w o o d  integrifoli  52  4.  4.1  SOME FACTORS AFFECTING SPECIES DISTRIBUTIONS  Introduction  Chapman (1938), environmental salinity,  factors  to  be  temperature,  rainfall,  soil,  influence.  categories In  tides,  Obviously these are a l l  in their  in general.  importance:  table,  and b i o t a .  Chapman's environmental marshes  major  water  some degree interdependent  tidal  of  drainage, a e r a t i o n ,  evaporation, to  i n a study of s a l t marshes, c o n s i d e r e d 10  seem  a p p l i c a b l e to  the present  study, I have  c o l l e c t e d data from four of them: b i o t a , t i d e s , s a l i n i t y , soil. be  Some of the other f a c t o r s  inferred  from  environmental  these.  The  and  ( i . e . drainage, a e r a t i o n ) can theoretical  relevance of my  data t o 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 s discussed  in the f o l l o w i n g four s e c t i o n s .  4.2  Tides  The course,  unique  phenomenon of  the t i d e s .  tidal  suspended  night  may  affecting  sediment,  exclude  habitats  D a y l i g h t f l o o d i n g reduces  by reducing the supply of carbon of  marsh  light.  oxygen  respiration.  from  is,  of  photosynthesis  d i o x i d e and, mainly  because  T i d a l f l o o d i n g during day or the  rooting  environment,  The f l o o d i n g t i d e may bathe the marsh  53  plants in saline substrate;  water  and  a f t e r the ebb  substrate s a l i n i t y .  increase  the  salinity  t i d e , r a i n f a l l or seepage may  Cool  floodwaters  may  then  expose  temperatures. receded pools  soil leaves  b r i e f and minutes and  light-blocking  the tide  sedimentation  elevations event,  and  mechanical s t r e s s .  patterns,  perhaps  and  d u r a t i o n of t i d a l  occurring  levels,  flooding  environmental  distributes  to a rather s l i g h t g r a d i e n t between  tidal  factors  of was  tidal that  for  a  few  flooding is daily  give  gradient  the e a r l y 1800's (Doty, 1957).  levels  on  i n the marsh, f l o o d i n g i s a  of i n t e r t i d a l organisms has  relation  to  mud  flooding  sometimes l a s t i n g most of the day.  relation  zonation  on  water  c a r r i e s away d e t r i t u s .  pronounced  A  least  shoots  once a month; at the lowest  corresponding  silt-laden  flow of water  plant  affects  infrequent  in frequency very  The  of  has  surface  coating  fine  highest  prolonged,  the s a l i n i t y of  a  subjects  the  tide  of  ebbing  propagules and  the  tide  increased  Evaporation  surfaces.  Tidal activity  increase  falling  sharply  after  sharply  water.  photosynthetic tides  to  on warm days  substantially  and  At  vegetation  Evaporation  may  commonly  the  the  reduce  suddenly and  reduce the temperature of the marsh p l a n t s ; the may  of  in  Differences rise the  to  a  marsh,  in elevation. and  the  vertical  been p o s t u l a t e d s i n c e at A  pioneering  study  of  marsh 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 to of  Johnson  &  York  (1915).  monographic treatment of a Long I s l a n d (N.Y.) s a l t marsh,  In  a  they  54  considered  the e f f e c t s of t i d a l o s c i l l a t i o n on  gas exchange,  soil  salinity,  In p a r t i c u l a r , they marsh  light  related  the  supply, and vertical  p l a n t s p e c i e s to both the frequency  tidal  f l o o d i n g , and  they  form  of  between  a  elevation  ratio and  competitive  total  total  interactions  factors.  time  of  the d u r a t i o n of  parameters exposed  submerged.  as  other  distributions and  summarized these  time  transpiration,  in  at  They  the  a given  considered  w e l l as a u t e c o l o g i c a l l i m i t s i n  explaining species d i s t r i b u t i o n s . On  the A t l a n t i c c o a s t , i n c l u d i n g the area  York's  study  described  above,  o s c i l l a t i o n s are n e a r l y equal coast tides  inundation  or  exposure,  in range,  Thus nor  Johnson  successive semidiurnal  there i s a marked i n e q u a l i t y (Frey & Basan 1978).  of  but  on  the  i n the range of neither  the  the  &  tidal  Pacific  successive  frequency  of  duration  of  cumulative  inundation or exposure, nor the d u r a t i o n of s i n g l e episodes inundation or exposure, changes smoothly  along  gradient.  c r i t i c a l elevations,  Rather,  there  are  certain  an  of  d e f i n e d by l e v e l s i n the t i d a l c y c l e , at which the these  parameters  jump  abruptly.  s e r i e s of s e v e r a l d i s t i n c t exposure predict)  with  intertidal  region.  first  its  own  The  zones,  characteristic  T h i s concept  result  biota,  p u b l i s h e d by Doty (1946) in connection  distributions the c r i t i c a l  of marine algae  in C a l i f o r n i a .  t i d e l e v e l s as: the lowest  tide  values  of  is a vertical  each  of c r i t i c a l  elevation  tide  (one within  the  levels  was  with the Doty  might  vertical  identified  level,  or  lower low water (LLLW); the mean l e v e l of the lower of the  low two  55  daily  low  tides,  or mean lower  l e v e l of the higher of the two low  water  (LHLW);  the  low water (MLLW); the  daily  highest  low t i d e s , or low  level  two d a i l y h i g h t i d e s , or low  lowest  higher  of the two d a i l y  t i d e s , or high higher low water (HHLW); the the  lowest  lowest  low  level  of  lower h i g h water (LLHW); the  l e v e l of the higher of the two  d a i l y high t i d e s , or  higher h i g h water (LHHW); and the h i g h e s t t i d e l e v e l , or  low high  higher h i g h water (HHHW). Swinbanks heights  varies  progressively lunar  day  years. tide  (1979)  pointed  according  to  out at  least  (24  h  50  min),  Even the  (hence the p e r i o d over recorded The  cycles  which  i s of c o n s i d e r a b l e critical  its  own  critical  m i n the F r a s e r D e l t a area  tidal  measurements  have  been  significance).  types of t i d a l  locations.  curves  there are so many of them that any b i o t i c  have d e s c r i b e d the c r i t i c a l  zone boundary  t i d e l e v e l concept  l e n g t h because of i t s obvious appeal and e x p l a i n i n g the observed  the i n t e r t i d a l marsh zone.  from  U n f o r t u n a t e l y , as Swinbanks  stands a good chance of c o i n c i d i n g rather c l o s e l y with  in  18.6  t i d e l e v e l s are u n i v e r s a l , and can be used  geographically distant  I  of  18.6-year t i d a l c y c l e i s s i g n i f i c a n t ,  c r o s s - c o r r e l a t e very d i f f e r e n t  noted,  four  tidal  1 lunar month, 1 year, and  Each of these c y c l e s g i v e s r i s e to levels.  the curve of  i n c r e a s i n g wavelength, having p e r i o d i c i t i e s of 1  p e r t u r b i n g t i d e l e v e l s by about 0.5  to  that  likely  one. at some  significance  z o n a t i o n of v a s c u l a r p l a n t s w i t h i n However, the concept  seems  to  me  56  to  be  of d o u b t f u l p r a c t i c a l u t i l i t y ,  of my own study, following:  for a  (1.) as  tide levels,  the  stated,  tide  over at l e a s t  18.6 years  what  such  as  for predicting critical  be  tide  observation,  correlations levels;  exists  with  field  (4.) i n t e r a c t i o n s  f a c t o r s and other environmental v a r i a b l e s (e.g.  distributional  s p e c i e s probably Largely  to?  may  information  s u b s t r a t e t e x t u r e , s a l i n i t y ) may be s i g n i f i c a n t species  which  responding  been i n f e r r e d e m p i r i c a l l y from f i e l d  between t i d a l  critical  s p e c i e s a u t e c o l o g i c a l l i m i t s and requirements  making i t u s e l e s s variables  the  l e v e l data base i s not everywhere adequate to  (3.)  mostly  including  there are a great many  are not w e l l enough understood, and has  reasons,  t i d e l e v e l s i s a species  the task, s i n c e o b s e r v a t i o n s required;  of  i n the context  some of which are bunched c l o s e l y together:  of s e v e r a l c r i t i c a l (2.)  variety  at l e a s t  limits;  (5.)  interactions  influence their d i s t r i b u t i o n a l  because  attempted to u t i l i z e  of  these  the c r i t i c a l  i n determining  reservations,  between  limits. I  have  not  in  this  t i d e l e v e l concept  study.  4.3  Salinity  In  the Fraser D e l t a , with  i t s strong g r a d i e n t  from f r e s h  water to s a l t water h a b i t a t s , the e f f e c t of s a l i n i t y on marsh p l a n t species d i s t r i b u t i o n s  i s r e a d i l y apparent.  tidal  57  Physiologically,  an  excess  of NaCl in the growth medium  can produce v a r i o u s t o x i c e f f e c t s i n higher p l a n t s , protoplasmic to  including  s w e l l i n g and changes i n enzyme a c t i v i t y ,  interference  in  respiration,  disturbance  of  leading nitrogen  a s s i m i l a t i o n , and a b n o r m a l i t i e s of p r o t e i n metabolism 1980).  Uptake  of  e s s e n t i a l n u t r i e n t ions may  the presence of NaCl ( i b i d . , Waisel  1972).  t o l e r a t e l e v e l s of NaCl that would k i l l The expressed subject  in the phenomenon of  an  of  extensive  (1972). respect  halophytes,  of  &  of  halophytism,  literature  species  halophyte  ecology  i s given  concise  specifically  metabolically,  to  salt  (1972) and symposium  in Reimold & Queen  of  halophytism  marshes  are found  out,  plant,  with  in Chapman  (1972).  must  pointed deal  with  three ways: " I t must s e l e c t i v e l y from  the  physiological  wider-ranging  treatments  is  i s d i s c u s s e d only  see the volumes by Waisel a  Useful,  which  1  of the  (1975);  As E p s t e i n (1969)  1  and  Gale  (1974) and Ranwell  elements  some  in  most p l a n t s .  For comprehensive reviews  Poljakoff-Mayber treatment  be reduced  a d a p t a t i o n of p l a n t s p e c i e s to s a l i n e environments i s  b r i e f l y here. ecology  Yet  (Larcher  [the  chemical  its  a  chemical  acquire  to  environment i n  essential  environment],  function  nutrient  i t must cope with  I am using the term "halophytism" to mean a d a p t a t i o n to a s a l i n e environment, without implying the degree of a d a p t a t i o n , or whether the a d a p t a t i o n J.s facultative or . o b l i g a t e . The opposite term i s "glycophytism", meaning a d a p t a t i o n to a freshwater environment.  58  elements present saline  i n excess, and  environment,  then,  problems (Queen 1975): nutrients NaCl),  from  (2.)  a  (1.)  within  acquiring  sufficient  its  internal  from  a  narrow  limits  essential  i o n i c mix  (high  balance  and  ionic  despite  c o n c e n t r a t i o n of NaCl i n the e x t e r n a l medium, water  In  p l a n t must overcome three main  medium with an unfavorable  maintaining  concentration  a  i t must a c q u i r e water."  the  (3.)  high  acquiring  an e x t e r n a l s o l u t i o n with a high osmotic  pressure  (low water p o t e n t i a l ) . With regard to the evidence  that  first  halophytes  discriminating  between  balance  were  Queen  better  essential  n o n e s s e n t i a l competitors The  problem,  than  nutrient  (mainly Na  (1975)  cited  glycophytes ions  and  their  and C I " ) . '  +  second problem, that of r e g u l a t i n g the i n t e r n a l and  d e a l t with  concentration  within  tolerable  i n three main ways: (a.) by  e x c e s s i v e or t o x i c  plant;  (c.)  increasing c e l l  by  succulence,  internal  by  the  for  method  s e g r e g a t i o n , to be c o n t r a d i c t o r y , but  Flowers  volume.  g e t t i n g r i d of excess 1972,  from  of s a l t s  endorsed by Dainty  (Waisel  (b.)  i.e. dilution  considered  et a l . (1977) reviewed the evidence view  be  i n t e r n a l s e g r e g a t i o n of  ions (mainly NaCl) w i t h i n the vacuole;  Queen (1974, 1975) (a.),  ionic  l i m i t s , may  by e x t e r n a l segregation of such i o n s , i . e . t h e i r removal the  at  Queen  (1979).  salt,  and  evidence  found  a  As for method ( b . ) , a c t u a l l y  s e v e r a l mechanisms  1975):  i t favorable,  (i.)  salt  may  glands  be —  cited highly  59  s e l e c t i v e e x c r e t o r s of Na leaching  from  leaves  and  +  (in  CI"  (in  e.g. Glaux); ( i i . )  e.g. A t r i p l e x ) ;  (iii.)  guttation;  ( i v . ) shedding of s a l t - c o n c e n t r a t e d l e a v e s or shoots Juncus g e r a r d i i ,  Atriplex,  Salicornia);  (v.)  ( i n e.g.  secretion  roots of s a l t s t r a n s l o c a t e d from shoots ( i n e.g.  by  Salicornia);  ( v i . ) accumulation of s a l t s i n s a l t h a i r s or bladders ( i n e.g. Atriplex).  Method ( c ) , the succulence s t r a t e g y , appears most  conspicuously a  specific  ( i n my  study area) i n S a l i c o r n i a ,  response  to  NaCl  i n which i t i s  (Poljakoff-Mayber 1975,  citing  Russian s o u r c e s ) . The led  t h i r d problem,  early  a c q u i r i n g water from a s a l i n e  medium,  r e s e a r c h e r s to s p e c u l a t e that halophytes endured a  water d e f i c i t ,  or " p h y s i o l o g i c a l drought".  T h i s idea  is  now  d i s c r e d i t e d ; halophytes are a b l e to maintain a f a v o r a b l e water potential  g r a d i e n t by osmoregulation, accomplished  ways (Gale 1975), Although potential  a  i n c l u d i n g NaCl .halophyte  uptake.  may  reduce  its  concentration halophyte  concentrations  below t o x i c l e v e l s . enzymes than  are  One  more  glycophyte  "there  citing is  different,  Flowers  no  evidence  in  their  et that  might  tolerant  enzymes.  c o n s i d e r e d the evidence c o n t r a d i c t o r y (1979),  internal  water  by t a k i n g i n s a l t , the need to maintain a f a v o r a b l e  g r a d i e n t must be balanced a g a i n s t the need to  that  in various  for  a l . (1977), enzymes  keep  the  s p e c u l a t e , then, of  high  Queen (1974, this,  but  was q u i t e of  NaCl  NaCl 1975) Dainty  definite:  halophytes  are  s e n s i t i v i t y to NaCl, from the enzymes of  60  other p l a n t s . "  Jefferies  (not v a c u o l a r ) Na about  100  mM,  (1973)  estimated  the  protoplasmic  c o n c e n t r a t i o n i n T r i g l o c h i n maritimum to be  +  which i s f a i r l y  low.  Various organic solutes,  however, are found i n halophyte protoplasm at about concentrations  for  osmoregulation organic  et a l . 1979). to  protect  of NaCl  (Dainty 1979,  i n v o l v e s the  (Stewart et a l . 1979). shows  that  very  few  i f . any  al.  Chapman 1977); most perform  1977,  salinity.  Barbour  (1970)  5  ppt.  Some  maximum growth has minimum  or  at  zero  reviewed the l i t e r a t u r e and  found  to be r e s t r i c t e d  to  salinities  common t i d a l marsh halophytes f o r which  been  maritima  (Pigott  (Flowers et  low  found  experimentally  zero s a l i n i t i e s are Juncus g e r a r d i i  Plantago  maritimum  best  (Chapman  1969).  1977),  to  and  patula,  Puce i n e l l i a  Ruppia maritima study  in  Triglochin  area,  conditions:  nuttalliana,  Distichlis  (Flowers et a l . 1977).  only  Salicornia  r e s t r i c t e d to s a l i n e h a b i t a t s  may  appear  Other  brackish  my to  halophytes  e.g.  Atriplex  spicata,  Of genera  at  1976,  (An a n e c d o t a l o b s e r v a t i o n from  much l a r g e r i n the l e s s s a l i n e areas.) growth  occur  (Rozema  study area i s that Tr i g l o c h i n maritimum p l a n t s  show optimal  salt-  or  that very few s p e c i e s appeared  grow  somehow  s a l t - s e n s i t i v e enzyme systems a g a i n s t high l e v e l s  s p e c i e s are a c t u a l l y o b l i g a t e halophytes  own  of  Jefferies  a l s o serve  tolerant  1979),  that  production  Stewart et a l . 1979,  Some of these compounds may  Much evidence now  above  right  osmoregulation; thus i t seems l i k e l y  i n halophyte c e l l s  osmotica  the  found i n  and my  be an o b l i g a t e halophyte,  (Ranwell  1972,  Chapman  1974);  61  even  for  contrary  this  genus,  though, there  (Flowers et a l . 1977).  Although most halophytes conditions  in  habitats  in  adaptations  the  laboratory,  nature. to  a  perform  Presumably  high-NaCI  the  In  Nutrient  tidal  determining carbon,  the  be  macronutrients,  as  to  expected  and  are  required  Seven  to of  in  elements,  i n t r a c e amounts.  the chemistry of the  play  a  major  plant species. six some  influenced  operating  i n t i d a l marsh h a b i t a t s : the the  Aside from  abundance  rooting  by  called by higher  calcium, and  chlorine,  are  environmental  factors  salt  of the  medium  s p e c i e s , e s p e c i a l l y t o x i c sodium; and the with  in  The a v a i l a b i l i t y of these n u t r i e n t  is  enriches  role  elements,  including  elements  floodwaters  competitive  environments.  oxygen,  required  other  strongly  halophytic  poor  p l a n t s : n i t r o g e n , phosphorus, potassium, sulphur, magnesium.  water  have a s i g n i f i c a n t  the  elsewhere,  distribution  hydrogen,  fresh  Regime  marshes,  r o o t i n g medium may  in  various  environment  a b i l i t y of halophytes i n g l y c o p h y t i c  Substrate  best  few are found i n f r e s h water  energy c o s t , no doubt c o n t r i b u t i n g  4.4  i s some evidence to the  content in  high  several water  ionic table,  p e r i o d i c f l o o d i n g , excludes oxygen from the i n t e r s t i t i a l  s o l u t i o n , g i v i n g r i s e to reducing there may be chemical g r a d i e n t s  conditions.  In  addition,  from the high marsh to the low  62  marsh  resulting  from the g r a d i e n t s  i n frequency  and d u r a t i o n  of f l o o d i n g . The  macronutrient  generally  made  cations  available  weathering of c l a y minerals the yet  nutrient  to  + +  ,  Mg ,  and  ++  terrestrial  (Etherington  K  plants  1975).  are  +  by the  Potassium  1969).  in soil  solution  i s often  very  +  saline  tidal  the F r a s e r River  low  In s a l t marshes, however, the macronutrient  c a t i o n s , i n c l u d i n g K , are s u p p l i e d i n high c o n c e n t r a t i o n s the  is  c a t i o n r e q u i r e d by p l a n t s i n g r e a t e s t q u a n t i t y ,  i t s concentration  (Epstein  Ca  floodwaters  (Table I I ) .  i s nutrient-poor  by  The f r e s h water of  by comparison.  Table II - T y p i c a l c o n c e n t r a t i o n s (mg L" ) of macronutrient c a t i o n s p l u s Na and C l " i n s o i l s o l u t i o n , seawater, and F r a s e r River water 1  +  I on spec i e s Ca Mg K Na Cl"  + + ++  +  +  Soil s o l u t ion  Seawater  2  Lower F r a s e r  River  3  1  1 36-560 46-168 27-39 23-667 39-710  400 1 300 400 1 1 000 20 000  Jan.  June  18 44 9 222 373  15 3.6 1 .4 2.0 3.9  F r i e d & Broeshart 1967. Ranges are from calcareous s o i l s . P i g o t t 1969 and Larcher 1980. Benedict, H a l l & Koch 1973. 1  acidic  soils  to  2 3  Much  the commonest  ions  in  seawater  a r e Na  +  and C l "  63  (Table most  I I ) , which are not only plants  essential 1980).  (Section  4.3), but  ions, p a r t i c u l a r l y K  Halophytes o b t a i n  spite  of  toxic in  +  also  and C a  what they  level  by  right  to  (Waisel  + +  from  1972, Larcher the  soil  in  The halophyte T r i g l o c h i n  maritimum has been shown to respond sodium  own  reduce the uptake of  need  high NaCl c o n c e n t r a t i o n s .  external  their  to  an  increase  a c t i v e l y increasing  i n the  i t s potassium  uptake (Parnham 1971), presumably f o r osmoregulation. Several nutrient  s t u d i e s have pointed  f o r t i d a l marsh halophytes  al.  1972, 1973; V a l i e l a & Teal  be  closely  related  containing  in  studies have no  to  compounds,  present at high grown  to nitrogen  Nitrogen metabolism may  tolerance;  certain  conditions  particularly  (Stewart et a l . 1979).  on n u t r i e n t l i m i t a t i o n i n t i d a l marshes  dealt  stress  to  requirement may t h e r e f o r e important  f i x a t i o n of  atmospheric  (Green & Edmisten  1978).  Nitrogen-fixing  mud s u r f a c e ;  be  Reported  seem  a l l to  dealt  with,  of  nitrogen  nitrogen  by  the  nitrogen  in  t i d a l marshes i s  blue-green  algae  and  1974, Jones 1974, P a t r i q u i n & Keddy blue-green algae are abundant on the  nitrogen-fixing bacteria  muds  and  be l e s s .  source  bacteria  anaerobic  when  with s a l t marshes; i n freshwater marshes there i s  salinity  An  nitrogen-  presumably osmoregulatory s o l u t e s , are  l e v e l s i n many halophytes,  saline  limiting  ( P i g o t t 1969; Stewart et  1974).  salt  as the  are  found  within  and a l s o i n a s s o c i a t i o n with the root  of many t i d a l marsh v a s c u l a r  plant  taxa  (Patriquin  the  systems &  Keddy  64  1978),  at l e a s t 22 of which are present  Fraser Delta. nitrogenase three  i n the marshes of the  To put matters i n p r o p o r t i o n , the l e v e l of root  activity  orders  of  found by P a t r i q u i n &  magnitude  greater  Keddy  one  to  the legume  Lathyrus  p a l u s t r i s than i n any of the other marsh s p e c i e s they  studied.  On the other hand, Green & Edmisten s u r f a c e mud, green p l a n t shoots, Mexico  in  was  (1974), who sampled water,  and dead shoots i n a Gulf  S p a r t i n a marsh, found t o t a l n i t r o g e n f i x a t i o n  up t o 1550 kg of n i t r o g e n per hectare an  order  of  magnitude  greater  commercial soybean or a l f a l f a The  nutrient status  influenced  by  the  This  permits  up,  leading  conditions. toxic  ferrous  bisulphide, is  found  tidal  effects  populations to  the  In  such  form,  than  which  soils  of waterlogging. the  of anaerobic  development  The high water .  marsh  sediments.  microorganisms to b u i l d  of  chemically  an environment, i r o n sulphur  i s greatly  reducing  i s present  i s present  as  or hydrogen s u l p h i d e , a l l h i g h l y t o x i c . as  ammonia  or  is  the f i x a t i o n r a t e f o r  marsh  oxygen from  and  r a t e s of  crops.  of  t a b l e e f f e c t i v e l y excludes  i n one month,  of  ammonium  i o n , forms  in i t s  sulphide, Nitrogen  which  are  u n a v a i l a b l e to many s p e c i e s . A  gradient  in  soil  oxygen s t a t u s from the high marsh,  where the r o o t i n g medium may be e s s e n t i a l l y lowest  marsh,  where  only  a  capable  of anaerobic  to the  t h i n surface l a y e r i s a e r o b i c ,  presumably a f f e c t s s p e c i e s d i s t r i b u t i o n s . are  aerobic,  Many marsh  species  metabolism when under oxygen s t r e s s  65  (Armstrong 1975); many can a s s i m i l a t e n i t r o g e n nitrate plants from  or  the  ammonium form.  i s the t r a n s p o r t the  shoots  rhizosphere the s o i l Some  to  the  v i a aerenchymatous  species, which  conservation  as  the  tissue  r o o t s , whence i t d i f f u s e s i n t o the ions  s o l u t i o n may be o x i d i z e d before  marsh  either  A common s t r a t e g y of wetland  ( i b i d . ) ; thus t o x i c reduced  adaptations,  flow  of oxygen  in  e.g. Juncus  presumably  approaching  in  they reach the p l a n t .  spp.,  function  show  xeromorphic  not  for  water  such, but rather to reduce the r a t e of water  towards the r o o t s , thus i n c r e a s i n g the time a v a i l a b l e f o r  o x i d a t i o n of p o t e n t i a l t o x i n s i n the rhizosphere The  a v a i l a b i l i t y of s o i l n u t r i e n t s to p l a n t s  considerably changing  (ibid.). is  by the s o i l pH, s i n c e i o n i c e q u i l i b r i a  pH.  Soil  d i f f e r e n c e s , with (Etherington  microflora  possible  1975).  also  are  consequences  Most v a s c u l a r  affected  shift  affected  with  by  f o r higher  pH  plants  p l a n t s have broad pH optima  in the range between weak a c i d i t y and weak a l k a l i n i t y  (Larcher  1980)., The pH of s a l i n e s o i l s i s u s u a l l y more or l e s s n e u t r a l but  may  be  affected  concentration  4.5  (Waisel  Substrate  Moody americanus species  in  by  leaching,  temperature,  and ion  1972).  Texture  (1978) and the  reported  Scirpus brackish  that  the  maritimus, Brunswick  habitats  of  Scirpus  the two major c o l o n i z i n g Point  marshes,  were  66  differentiated  by  substrate  on sandy s u b s t r a t e s , and established  on  a  t e x t u r e : S. amer icanus  S. maritimus in  sandy  substrate,  r e p l a c e d by S. maritimus i n apparent silt  silty  colonizes  areas.  S. americanus response  to  Once may  be  increasing  deposition. Substrate  texture probably  in s e v e r a l ways.  Coarse-grained  that  table w i l l  the  water  a f f e c t s species d i s t r i b u t i o n s s o i l s are more permeable,  f l u c t u a t e more, perhaps r i s i n g  the s u r f a c e on a f l o o d i n g t i d e ; t h i s does not l e s s permeable f i n e - g r a i n e d sediments Being  more  permeable,  b e t t e r d r a i n e d and  happen  to the  (Clarke & Hannon 1969).  however, the c o a r s e - g r a i n e d  aerated,  in  so  so that o x i d i z i n g  soils  conditions  are may  dominate. A  more  porous  soil  is  more s u b j e c t to the e f f e c t s of  l e a c h i n g by p r e c i p i t a t i o n or groundwater; fluctuate  more.  Presumably,  with  f l u c t u a t i o n between atmospheric and coarse-grained may  s o i l , other  the  aquatic  salinity  overall  may  greater  conditions  in  f a c t o r s such as temperature and  a pH  a l s o f l u c t u a t e more. C a t i o n exchange c a p a c i t y  particle  content;  thus  nutrient a v a i l a b i l i t y As pointed of  thus  i n c r e a s e s with  substrate  f o r marsh  out by J e f f e r i e s  fine-textured  water p o t e n t i a l .  s o i l s may  increasing  texture  fine-  may  influence  (1972), the matrix  potential  vegetation.  significantly  reduce the  Thus even though the osmotic  external  potentials  of  67  sandy may  and  be q u i t e  clayey  s o i l s may  different.  be s i m i l a r ,  the water p o t e n t i a l s  68  5.  5.1  SAMPLING METHODS AND DATA COLLECTION  Vegetat ion  Areas exhibit  of  marshland  were  selected  which  appeared  to  some v e g e t a t i o n a l or environmental d i s s i m i l a r i t i e s to  one another, and w i t h i n these areas t r a n s e c t s were l a i d out i n the d i r e c t i o n of evident or  assumed  (high e l e v a t i o n t o low e l e v a t i o n ; Metre-square  quadrats  were  environmental  f r e s h water t o s a l t water).  p l a c e d s u b j e c t i v e l y at i n t e r v a l s  along the t r a n s e c t s i n a manner intended to range of observed v a r i a t i o n .  gradients  Each quadrat  sample  the  full  l o c a t i o n was marked  with a wooden stake. Within  each  quadrat,  a l l vascular  s u b j e c t i v e l y assigned coverage code aerial  coverage  c l a s s e s of <1%,  76-95%, and 96-100%.  p l a n t s p e c i e s were  values  corresponding  to  1-5%, 6-25%, 26-50%, 51-75%,  Data were recorded f o r  a  total  of  64  s p e c i e s i n 103 quadrats. Plant specimens were c o l l e c t e d and  eventual deposition  Herbarium.  for species  identification  i n the U n i v e r s i t y of B r i t i s h  Columbia  69  5.2  Soil  At each p l o t one sediment for  (roughly  or  more  small  spadefuls  of  mineral  500 g) were c o l l e c t e d from the upper 15 cm  l a t e r p h y s i c a l and chemical  analysis.  Where  a  distinct  organic h o r i z o n had accumulated, a sample of t h i s m a t e r i a l was c o l l e c t e d as w e l l . Vegetation  and s o i l  sampling  were c a r r i e d out during the  summer of 1978.  5.3  L e v e l l i n g Survey  To  obtain  levelling sampling and  survey season.  visibility  DKM2  quadrat was  elevations performed  the  two  relative  quadrat and  f o l l o w i n g the  By means of a  Kern  e l e v a t i o n s were determined to an  p l u s or minus one c e n t i m e t r e .  elevations  thus  obtained  were  later  t o e l e v a t i o n s above l o c a l c h a r t datum, using one of  methods:  monuments  a  h o r i z o n t a l c o o r d i n a t e s of the quadrats  of approximately  converted  locations,  A e r i a l v e g e t a t i o n by t h i s time had d i e d down  were obtained and the quadrat  The  map  i n the winter  was thus g r e a t l y improved.  theodolite,  accuracy  and  (1.)  At  Boundary  Bay,  engineering  of known geodetic e l e v a t i o n were found  l e v e l s were t i e d to the l e v e l s  t h e i r geodetic e l e v a t i o n s converted  of  these  survey  nearby.  The  benchmarks,  to h e i g h t s above c h a r t  70  datum  at  White  Rock.  ( 2 . ) At  Ladner Marsh and Brunswick  P o i n t , o b s e r v a t i o n s were made of the times at which the level  rose above a r b i t r a r y benchmarks on s u c c e s s i v e days.  extrapolation the  water  from h o u r l y t i d e l e v e l r e c o r d s ,  benchmarks  1  above c h a r t datum at Steveston  the h e i g h t s  By of  ( f o r Brunswick  P o i n t ) and Deas I s l a n d ( f o r Ladner Marsh) were determined.  1  Supplied f o r Ladner Marsh by Environment Canada, Water Resources Branch, Inland Waters D i r e c t o r a t e , P a c i f i c and Yukon Region, Vancouver, B.C., and f o r Brunswick Point by I n s t i t u t e of Ocean S c i e n c e s , Sidney, B.C.  71  6.  Soil crumbled  SOIL ANALYSIS  samples were oven-dried at 100 C f o r 24 hours, with  a  rolling  pin.  Coarse  plant  separated from mineral sediments by  hand-picking  sieve.  little  Some  samples  had  very  remaining a f t e r r o o t s and rhizomes  were  then  m a t e r i a l was or  mineral  removed.  with  a  material For  this  reason and a l s o f o r reasons of economy, some samples were only partially The •  analyzed. f o l l o w i n g analyses were  Particle  performed:  size analysis,  i . e . d e t e r m i n a t i o n of percent  by weight of sand, s i l t ,  and c l a y , u s i n g the Bouyoucos  hydrometer, method (Day 1965). •  pH d e t e r m i n a t i o n on 1:1 s o i l : w a t e r Radiometer  pH  Meter  29  with  extract,  using  a  a Radiometer combined  glass/reference electrode. •  Electrical  conductivity  determination  saturation  paste,  using  a  with  a  conductivity  meter  conductivity •  Total  Radiometer Radiometer  on  soil  type  CDM2e  CDC  104  cell.  nitrogen  determination,  by a c i d d i g e s t i o n and  c o l o r i m e t r i c a n a l y s i s , using a Technicon  AutoAnalyzer  II . •  Determination  of  Ca  + +  ,  K, +  s o i l : w a t e r e x t r a c t , using a absorption  spectrophotometer.  Mg , ++  and  Perkin-Elmer  Na  +  306  i n 1:1 atomic  72  Except outlined  as  indicated,  in Lavkulich  (1978).  the  procedures  followed  were as  73  7.  7.1  Ordination  The  term  techniques  DATA ANALYSIS  Methods  "ordination" in  which  is  given  used  some  the  reveal  The  several  their relationships" (Orloci  o r d i n a t i o n methods used in t h i s study  category which Whittaker  analysis",  vegetation  in such a manner that t h e i r arrangenent w i l l about  or  as  denote  their  information  to one  such  to  of  useful  1978b).  ecology  entities,  r e l e v e s , "are ordered according properties  in  (1967) c a l l e d  fall  "indirect  i n which samples are arranged along  axes  into  gradient generated  by v e g e t a t i o n a l v a r i a t i o n . Principal  components  interpretation  of  introduced  ecology by Goodall  to  the redundancy summarizing  in  the  results  analysis, of  on  (1954).  In geometric terms, i t f i t s  of p o i n t s , r e p r e s e n t i n g coordinates the  n  number  space  of  by  axes,  ordination  l i n e s or planes to a c l o u d  i n d i v i d u a l s (e.g. sample p l o t s ) , whose  (e.g. s p e c i e s ) .  The  scores  first  f o r each  ordination  then corresponds to the d i r e c t i o n of maximum v a r i a n c e point  was  method reduces  vegetation  i n n-dimensional space are the  attributes  The  for  tests,  the o r i g i n a l v e g e t a t i o n  c o r r e s p o n d i n g to s p e c i e s , on a smaller axes.  developed  psychological  species-dimensional variation  first  in  of axis the  c l o u d ; each subsequent a x i s i s a l i g n e d i n the d i r e c t i o n  of g r e a t e s t  remaining v a r i a n c e .  Geometrically,  then, the axes  74  are o r t h o g o n a l .  However, s i n c e c e r t a i n s t a t i s t i c a l  especially  assumption of m u l t i v a r i a t e n o r m a l i t y , are not  met  by  the  ecological  biologically  of  (Beals  independent.  be represented deal  data  1973),  the  criteria,  axes  are  not  An e c o l o g i c a l g r a d i e n t may thus not  l i n e a r l y by PCA; t h i s has given r i s e t o a great  activity  by  mathematically-inclined  e c o l o g i s t s to  d i s c o v e r a b e t t e r method. A p o s s i b l e replacement f o r PCA i n is  reciprocal  earlier  averaging.  Like  ecological  PCA,  this  method was used  i n the s o c i a l s c i e n c e s ; the e a r l i e s t  applications  of t h i s method i n ecology  (1971) and H i l l  (1973).  ordination  English-language  were those by Hatheway  As d e s c r i b e d by H i l l ,  the  procedure  i s one of s u c c e s s i v e r e c i p r o c a l r e - c a l i b r a t i o n s of species and stand of  scores,  each  direct  i n which both are d e f i n e d and r e d e f i n e d i n terms  other. gradient  technique  of  that  i s thus r e l a t e d t o  and  to  Wisconsin  the  simultaneous  Whittaker's  weighted  school  averages  (Whittaker  RA i s s i m i l a r to PCA; i t s c h i e f  i t yields  ordinations,  method  analysis the  Computationally, is  The  paired  species  1978).  distinction and stand  " n e i t h e r of which has l o g i c a l pre-eminence"  (Hill  1973). The developed (1978a).  PCA and RA by  programs  employed  Dr. G.E. B r a d f i e l d  in  this  study  were  from d e s c r i p t i o n s i n O r l o c i  75  7.2  Treatment of the V e g e t a t i o n Data  R e c o g n i t i o n of p l o t groups: values  for  floristic by  the  64  species  table use  Bradfield  in  Initially,  the  cover  data  103 quadrats were organized i n a  (Table I I I ) .  T h i s procedure  was  facilitated  of a t a b u l a r s o r t i n g program w r i t t e n by Dr.  of  the  University  of  British  Columbia  G.E.  Botany  Department. It two  became evident that the p l o t s f e l l  floristically  comprises and  main  the p l o t s from western  from Ladner  designated all  distinct  the  Marsh  groups.  Point  The  (transects  A,  D,  and  "freshwater" group; the second  (transects  B,  first  group  and northern Brunswick Point  the Boundary Bay p l o t s p l u s the  Brunswick  rather neatly into  plots  C,  J ) , and  was  group i n c l u d e s  from  southeastern  E, F, G, and H), and  was  designated the " s a l t w a t e r " group. PCA  on unstandardized  complete  species  discontinuity level  of  in  cover species  between-habitat  data: value  As data  previously set  distributions, or  high  beta d i v e r s i t y  indicating  beta d i v e r s i t y .  1978), the data "freshwater" for  and  set  was  (Swan 1970; split  into  a  high  Since PCA  Whittaker the  the  d i s p l a y e d a marked  been shown to produce h i g h l y d i s t o r t e d o r d i n a t i o n s with  noted,  has  from  data  & Gauch  1973,  more  homogeneous  " s a l t w a t e r " subsets, i d e n t i f i e d p r e v i o u s l y ,  most o r d i n a t i o n procedures.  R e s u l t s of PCA  these freshwater and s a l t w a t e r cover data s e t s  o r d i n a t i o n s on are  shown  in  76  Table I I I - F l o r i s t i c t a b l e : 64 s p e c i e s i n 103 sample plots. Group at l e f t , mainly fresh-water i n f l u e n c e d ; group at r i g h t , mainly s a l t - w a t e r i n f l u e n c e d  A A J J J J J j j j J A A A A AA A D D D A D A D D D A A A A A A A A A A A A A A A A A A D A A B C C E E E E H H B E E EGGHHHHHF F B C C E G G G E G G F 3 H H C G S S 8 C C E F B E G B H H 8 B C G 5 9 6 9 7 1 5 : 3 - : 8 G 8 1 1 2 2 2 2 7 8 2 5 7 4 6 t .1 1 ) 1 1 2 2 2 1 1 2 13 1 1 2 2 0 3 3 2 1 7 8 8 1 3 6 7 8 5 2 7 8 2 3 6 9 1 1 5 3 4 3 3 4 5 6 5 4 9 8 9 5 7 4 1 1 1 1 2 J 5 P 4 1 9 8 7 6 2 3 1 1 2 1 08012 24673G7595 1348 0 9 2 O 1 0 01  EOUISETUM FLUVIATILE ALISMA PLANTAGO-AQUATICA SCIRPUS VALIOUS AGROSTIS ALBA CAREX LYNGBYEI ELEOCHARIS PALUSTRIS SAGITTARIA LATIFOLIA SCIRPUS AMERICANUS S C I R P U S MAR I T I M U S R U P P I A MAR I T I M A CHARA BRAUNII JUNCUS BALTICUS POTENTILLA PACIFICA TYPHA LATIFOLIA LYTHRUM SALICARIA SIUM SUAVE BIDENS CERNUA COTULA CORONOPIFOLIA TRIGLOCHIN MAR1TI1AUM SALICORNIA VIRGINICA DISTICHLIS 5PICATA SPERGULARIA CANADENSIS ATRIPLEX PATULA P L A N T A G O M A R I T I MA SPERGULARIA MARINA CU5CUTA SALINA PUCCINELLIA NUTTALLIANA GRINDELIA INTEGRIFOLI A G L A U X M A R I T I MA L1LA6A SCILLOIDES HORDEUM BRACHYANTHERUM JUNCUS GERARDII S P A 13 .  2223322;: 22221 1 4 4 * 2 1) 1 + 4 5 5 3 2 12 1 5 5 5 2 2 3 2 1 1 + 1 1 * * 3 3 5 5 5 4 5 1 - * +*1 2 3 35 4 4 6 5 * 4 6 2 6 6 2 3 1 5 6 2 * 13 1*1 * l 1** + 1 1 1 1 13 + 3 " 1 3 " 1+++ + 1 * 2 2 + 12+ 12 + 5 332 1 2 2 1 1522 164 14+* 2 11522+ 2 2 + 1+ B  2  22 2 3  2+1 + 4  5 ++5 1 2 2  + 3 5 + 1 2 2 2 2 1 2 2 14 1 1 1 + 2 2 + " 2 2 2 3 3 " + 2 1 5 +" 5 2 3 + 1 2 3 3 + 1 2 + 1 2 2 3 3 2 6 1 2 2 1 + 2 3 2 4 1 2 + - 4 2 1 " -+ + +3+2113+ 6342 626552 + * '25525526 1+  + + 122  +13 +3 33 +1+33422 422+ 21 4+ +22 232  2 12  1* 3 222  + +132  06NANTHE SARMENTOSA J U N C U S ART 1 C U L A T U S LILAEOPSIS OCCIDENTALIS AGROPVRON REPENS CALTHA ASARIFOLI A LIMOSELLA AQUATICA CALL ITRICHE SP . SPC2 S P G IO ACHILLEA MILLEFOLIUM HYGROHYPNUM LURIDUM LEPTOOICTYUM RIPARIUM ASTER EATONII SONCHUS ARVENSIS MYOSOTIS LAXA LATHYRUS S P . TRIFOLIUM OLIGANTHUM JUNCUS BUF0N1US MENTHA ARVENSIS DESCHAMPSIA CESPITOSA ELYMUS GLAUCUS ELYMUS MOLLIS FESTUCA ARUNOINACEA PUCCINELLIA NUTKAENSIS SPC3 SP01 SPE 1 POLYGONUM AVICULARE SP J 1 A MI M U l . U S G U T T A T U S SPJ4A  Legend: " = solitary individual, n e g l i g i b l e c o v e r ; + = s e v e r a l i n d i v i d u a l s , < 1% c o v e r ; 1 = 1-5% c o v e r ; 2 = 6-25% c o v e r ; 3 = 2 6 - 5 0 % c o v e r ; 4 = 5 1 - 7 5 % c o v e r ; 5 = 76-95% c o v e r ; 6 = 96-100% cover  Nomenc1 a t u r e i s a c c o r d i ng w i th author i t i es.  to  Hitchcock  et al  1969  See  Appendix  A for species  names  77  Figs.  11  and  ordinations resemblance A  are  (Unless  performed  otherwise using  PCA  a  ordination  on  the  PCA  cover data set  was  r e s u l t s proved u n s a t i s f a c t o r y , because many  of  standardizations:  method i n exposing affected  all  variance-covariance  complete  103 p l o t s were o v e r p r i n t e d on one Data  stated,  matrix.)  performed, but the  12.  by  The  another.  u s e f u l n e s s of an o r d i n a t i o n  ecological gradients  the  data  be  standardization  employed; hence d i f f e r e n t were t e s t e d on the data  may  considerably  or  transformation  s t a n d a r d i z a t i o n s and  transformations  i n order to  evaluate  their  effects.  These are d e s c r i b e d i n the remainder of t h i s s e c t i o n . Square weightings cover  I t was  noted  A c c o r d i n g l y , a means was  disparity  percent  between  high  cover v a l u e s , ranging  their  and  cover  square data  R e s u l t s of the PCA  root  the  high  information  being more or l e s s  ignored  sought to even out  somewhat  low cover v a l u e s .  Thus the  from 0 to 100,  square r o o t s , ranging  o r d i n a t i o n s on the saltwater  that the  on the p r i n c i p a l component axes were given to h i g h -  in lower cover values was  by PCA.  to  transformation:  s p e c i e s , l e a d i n g to the s u s p i c i o n that  supplied  the  root  were  from 0 to 10. transformed  matrices are given  o r d i n a t i o n on the square  transformed  R e s u l t s of freshwater  i n F i g s . 13 and root  PCA and 14.  transformed  complete cover data matrix are shown in F i g . 15. Normalization:  The  lowest  p o r t i o n s of the marsh, where  78  A20  J2 A17«* A 27  10 +  A16  J5  A1B  AXIS  1  J6 •  07  D3» . A 1 5 A11  A4, A28  A9  • A7  r  £ \ « A 2 9  J7  Aig'^o  •A  e  8  5 A3  A6  • 08  A10 •  D4»  •D6  Figure  11 - PCA o r d i n a t i o n of samples using s p e c i e s data from freshwater p l o t group  cover  79  G1 C2»  H4  6 f  G2 G9 • •H2 »H1  C7 R 1 l /  B  1  0  G10  ^W'E,  81 2  E6.*E8 E7»*G5» »H9 B5 H10 H 8 B7  H5  G6" B6  E-2  FA  E5  G8 G3 -3+  • B8 »B9  C.S F1 • B2 B3» C4  Figure  12 - PCA o r d i n a t i o n of samples using s p e c i e s cover data from s a l t w a t e r p l o t group  80  A,23 J1 . 4 2 0  1.0  »J2  A17  A 271  A26  A14  J5  J6  J3 J 9 A18  A13  D3  D2  -0.5 AO  0.5 J  • A 2 9 s.3,0 A 9 A*  8  A5  0 7 A7  • A 2 4  J4 -0.5 +  08 D5 A6  D4 A10 06  F i g u r e 13 - PCA o r d i n a t i o n of samples using square roots of s p e c i e s cover data from freshwater p l o t group  81  V m  C2  1.0  • H 3  C1 G9 H1  0.5  B10  %  C3  • B11 B12  G10  G7  F5 E9»  G2  H11  *C8 _ B 6  G5»  AXIS  0.5  1  E4  E6  G8  B8 B9 * .* * B 7 H10 • G6  G4  H9 E  1.0  H6  7  • E2  H8 G3  F2 B4  C4 B2  -0.5+ E5  F i g u r e 14 - PCA o r d i n a t i o n of samples using square r o o t s of s p e c i e s cover data from s a l t w a t e r p l o t group  C6  82  G10  G2 C3 i f * C6 •  0.4  F1  B3  F3  D8  J4 D7  D6 A22 0.4>  C5  A6 A18  H6  G3 D5  J 5  E5 J7  F4»  B4 H9  -t-  .  AXIS H 4 .  H8  H  Bp H5  E7  E6  G  B  ,  B*9  3  m  G 5  H11 %  H10.  0.4 "  1  • G9  G7  A29J8|  . • F 5 B12E9 E6 A30» * Bit* C8 B  C7»* ^ ^ A 4  A7 B  Al * ^ 1 3 AI9 . • ^••A13 »J6  5  A9 A28.| •  »A23  A14 A16 J1 A20. •A17  »A11  A24 A26  A 27 A23  F i g u r e 15 - PCA o r d i n a t i o n of samples using square roots of s p e c i e s cover data from a l l p l o t s (freshwater group, T r a n s e c t s A, D, J ; s a l t w a t e r group, T r a n s e c t s B, C, E, F, G, H)  83 flooding  effects  environment  are  for  most  plant  severe,  growth.  present  Releves  the PCA  diagrams,  rather  than  stands of s i m i l a r s p e c i e s composition A  means  was  therefore  stands with low t o t a l  sought  element  in  a  quadrat  value i n the quadrat) vector  in  the  standardization quadrat  was  ( i . e . the square  values  normalized  in  r a r e , low-cover generated  The  this  objective.  of the  squared  consequence in  a  quadrat cover  of  this  sparsely-vegetated  to cover a l l the bare areas,  the p r o p o r t i o n of t o t a l cover represented by  freshwater 16 and  the  R e s u l t s of and  PCA  ordinations  17. variance-covariance  preceding examples caused  species.  on  s a l t w a t e r cover data matrices  In c o n t r a s t ,  a  resemblance PCA  to ignore  resemblance  by the product-moment c o r r e l a t i o n c o e f f i c i e n t  stronger weighting to r a r e s p e c i e s . species  with  but higher cover v a l u e s .  root of the sum  C o r r e l a t i o n matrix: The used  on  v e c t o r ( i . e . each s p e c i e s cover  i s as i f the p l a n t s  are shown i n F i g s .  matrix  grouped  d i v i d e d by the l e n g t h of the  each s p e c i e s i n the quadrat. the  being  accomplished  quadrat).  changing  the o r i g i n  which would g i v e e q u a l i t y to  were to extend themselves  without  correspondingly  cover.  N o r m a l i z a t i o n of the data Each  are  from such areas were lumped near  scatter  hostile  P l a n t performance i s poor i n  such marginal h a b i t a t s , and cover values low.  a  It  was  felt  that  matrix gives rare  might p o t e n t i a l l y c o n t r i b u t e u s e f u l i n f o r m a t i o n to an  o r d i n a t i o n , so the data  were  analyzed  using  a  resemblance  84  A 20 A23  .08 +  J 5  .04+  ^ 5 , A 2 5 A13»«  Al 9  .08  -.12  J 9  D3  02 A18  J8  .04  A 5  A11 • A9  A29  A2.8*  VM A 30  D5  »A7  -.04+ 07  A22 .« D8 D4 • A6 • 06 A3  F i g u r e 16 - PCA o r d i n a t i o n of samples using normalized s p e c i e s cover data from freshwater p l o t group  A10  85  H3 H2»«G1  C2  G9  H4  .10  .05 + G7 E9« •F5 C1 B11,' E8  C  *B12 C3  G10 C6»  H10  G2  G5 G8  E4  H5  G4  G6 H9 C8 ,B6 B 8  VBS E6  C5  F3 B5 B4  Fl -.05  E2»« B3  G3  F i g u r e 17 - PCA o r d i n a t i o n of samples using normalized s p e c i e s cover data from s a l t w a t e r p l o t group  »B2  86  matrix of c o r r e l a t i o n c o e f f i c i e n t s . (not  shown)  featured  The  resulting ordinations  a very dense c l u s t e r of p l o t s near  the  o r i g i n , with a few o u t l i e r s around the edges. R e c i p r o c a l averaging; T h i s method may standardized  v e r s i o n of PCA  (Noy-Meir 1970,  i s c o n s i d e r e d here as a s t a n d a r d i z a t i o n . mainly  in  that  it  be  produces  regarded Hill  a  1973), so i t  It d i f f e r s  simultaneous  as  from  s p e c i e s and  PCA stand  ordinations. Initial  results  of  RA  ordinations  on  freshwater  and  s a l t w a t e r cover data matrices showed an extreme s e n s i t i v i t y to c e r t a i n data anomalies. species)  would  T y p i c a l l y , one or a few quadrats  appear as o u t l i e r s at one edge of the  diagrams, with the other quadrats  quadrats  from  7.3  By removing these  the data m a t r i c e s , g r e a t l y  r e s u l t s were obtained  scatter  (and s p e c i e s ) crowded  the o p p o s i t e edge of the diagrams.  (and  along  outlying  improved o r d i n a t i o n  ( F i g . 18).  Treatment of the Environmental  Data  S t a n d a r d i z a t i o n of e l e v a t i o n d a t a : E l e v a t i o n as such  has  nothing to do with the performance or d i s t r i b u t i o n of s p e c i e s ; rather  it  is  summarizes the  a  variable  levels  of  which  influence  a c t u a l l y act on the organism. is  c o n v e n i e n t l y estimates of  the  factors  and  which  In the. t i d a l marshes, e l e v a t i o n  mainly an index of the degree to which a p l a n t i s d i r e c t l y  87  J9 •+  Scival  J 6 • A14  Cotcor  > AO • A15  A13«  + Elepal  • J 8  A 2 5 Saglat  IChabra + • A 5  +  Alipla +  +  Equflu+«A27  Bidccr  A17 •  A23» A 2 0 * Scimar Rupmar  +  P u c n A12«  U  t 4  +  A  2  6  D3 T  y  p  l  +Lirnaqu +  ,  a  t  + +  Trimar  +  Agral b  J7  + A n g a r g  A11 • • A 2 9  +  +  J 5  +Calasa  Li locc +  • A 9  »A16  _»J2  +  Potpac  D2 • J 3  Si usua •fLytsal  A21  a  A18<  »A4 • A 3 0 > A 2 4 •A28  • J 4  K  D5  •  D7  A8» A 2 2 . D6 AS. Carlyn+. A 1 0 *  Legend:  + •  =  species sampl e p i o t s  s p e c i e s codes epithet;  = f i r s t 3 l e t t e r s o f g e n u s name + f i r s t 3 l e t t e r s o f s p e c i f i c s e e A p p e n d i x A f o r f u l l names ( P u c n u t = P u c c i n e l l i a nutkaensis)  F i g u r e 18 - RA o r d i n a t i o n of s p e c i e s and samples using s p e c i e s cover data from freshwater p l o t group  88  or i n d i r e c t l y body.  i n f l u e n c e d by the  Unfortunately,  water  level level  of  the  nearby  fluctuations  water  due to the  t i d a l c y c l e and to v a r i a t i o n s i n r i v e r flow do not f o l l o w  the  same  The  pattern  result good  throughout the study area ( S e c t i o n 3.4.3).  i s that a b s o l u t e e l e v a t i o n predictor  i s not l i k e l y to be  a  very  of p l a n t performance when comparing data from  d i f f e r e n t areas. To r e s o l v e t h i s problem, data  according  I  standardized  to l o c a l v a r i a t i o n s  in t i d a l  the range.  e l e v a t i o n s above c h a r t datum were d i v i d e d by the for  Thus the  tidal  range  l a r g e t i d e s at convenient t i d a l m o n i t o r i n g s t a t i o n s : Deas  Island  1  northern  for  the  and  Ladner  western  Marsh  plots,  the  Boundary  Bay p l o t s .  datum)  standardization reliability  to  100  for  2  and  Crescent  3  for  Beach"  range i n theory from  (the highest l o c a l water l e v e l ) .  procedure,  the  The s t a n d a r d i z e d e l e v a t i o n s are  expressed i n percentage u n i t s , and may (chart  Steveston  Brunswick Point p l o t s , Tsawwassen  the southeastern Brunswick Point p l o t s , for  elevation  however,  depends  for  0 The  its  on the j u d i c i o u s n e s s of c e r t a i n d e c i s i o n s -- such  1  Data o b t a i n e d from p r i n t o u t of 1978 hourly water l e v e l s at Deas I s l a n d Tunnel, s u p p l i e d by Environment Canada, Water Resources Branch, Inland Waters D i r e c t o r a t e , P a c i f i c and Yukon Region, Vancouver, B.C.  2  Data o b t a i n e d from p r i n t o u t of 1978 hourly water levels at . Steveston, supplied by I n s t i t u t e of Ocean S c i e n c e s , Sidney, B.C.  3  Data o b t a i n e d from Canadian Hydrographic S e r v i c e 1977, p. 18.  •  ibid.  89  as which t i d a l monitoring s t a t i o n to use as a over  what time p e r i o d to determine  reference,  the t i d a l  range.  and  Thus the  v a l u e s o b t a i n e d should be regarded as approximations. Ordination; Eight s o i l variables variable  were  selected  were: percentage and  variable  in  calcium, was  p a r t s per m i l l i o n  magnesium, • and  the  a  ordination.  by weight of sand, s i l t ,  concentration  potassium,  for  and  standardized  site  elevation  The s o i l  c l a y , and  variables nitrogen;  (ppm, or mg kg" ) of 1  sodium.  elevation  The  elevation  above c h a r t datum  d e s c r i b e d above. Initially, missing  were  a l l plots removed  from  data set thus d i f f e r e d set,  in  for  which  data  the data s e t . The  i n c h a r a c t e r from the  values  were  environmental  vegetation  data  which most of the values were zero ( i . e . s p e c i e s not  present).  With s p e c i e s data, t h i s  tends  cause  to  distorted  (Swan 1970; Whittaker data  any  ordinations,  i s almost  ordinations,  & Gauch 1973, 1978).  therefore,  unavoidable,  and  e s p e c i a l l y with PCA The  environmental  were free of at l e a s t t h i s one  source of c o n f u s i o n . The and  environmental  "saltwater"  data set was  subsets  on  the  split  into  same  basis  "freshwater" as  f o r the  v e g e t a t i o n data, and PCA and RA o r d i n a t i o n s were performed all  three  data  sets.  Since  the  variables  were  measured by the same u n i t s , a c o r r e l a t i o n matrix was the  PCA.  The  PCA r e s u l t s are shown i n F i g s .  The RA r e s u l t s were f a i r l y  on  not a l l used  in  19, 20, and 21.  s i m i l a r , but were not  as  easy  to  90  i n t e r p r e t and are not shown here. Display  of  ordinations:  spec i e s  cover  A major assumption  values  related  to  To demonstrate some  important  such a r e l a t i o n s h i p , species  were  Sc i rpus  and  amer icanus,  Distichlis ordination  Carex  cover  plotted  R e s u l t s are  plant  v a l u e s c o u l d be shown to  the l e v e l s of s e l e c t e d environmental  o r d i n a t i o n diagrams.  ordination  environmental  of t h i s study was that  s p e c i e s d i s t r i b u t i o n s and performance be  on  on  shown  values for  the environmental  for Agrostis  lyngbyei  ( F i g s . 22, 23, 24), and f o r  class  factors.  on  alba,  the freshwater  Salicornia  virginica,  s p i c a t a , and T r i g l o c h i n maritimum on the s a l t w a t e r ( F i g s . 25, 26, 27).  91  C 8  Q5 A17 A25»* • B8 A14  D6  J6 J4  A6  04  63  :  B12  A24  A19  A5  AXIS 1  A26» D2  B4 B2  D10  D8  AS  B.6  G2  A23  G5  A 28  Al  H*8 H10  A 21  A2  H1  D1  AO A3  G6 J2 J1  H11  F i g u r e 19 - PCA o r d i n a t i o n of s a m p l e s u s i n g e n v i r o n m e n t a l d a t a from f r e s h w a t e r ( T r a n s e c t s A , D , J ) and s a l t w a t e r ( T r a n s e c t s C , G , H) p l o t g r o u p s ,  92  ^25  A24  A5  A2B  A23 A1  A19  A2  A26 A14 J1  A17 •A10 AXIS 1  08  J2  A 21  A11  A8  A6 • J6  D2  DI  J4  D6 D4  Figure  20  -  PCA  o r d i n a t i o n of samples u s i n g data from f r e s h w a t e r plots  environmental  93  C8  G 3 B1  C5 B4 B2 B8  G2 AXIS  H8  G5  H1  H10  1  B6  B10  B12  G6  H11  Figure  21  - PCA  o r d i n a t i o n of samples u s i n g data from s a l t w a t e r p l o t s  environmental  94  4 1 AXIS 1  1  1  F i g u r e 22 - P l o t of cover c l a s s codes of A g r o s t i s a l b a on the o r d i n a t i o n of environmental data from freshwater p l o t s . 1 = <25% cover; 2 = 26-50%; 3 = 51-75%; 4 =76-100%; • = absent  AXIS  1  F i g u r e 23 - P l o t of cover c l a s s codes of S c i r p u s americanus on the o r d i n a t i o n of environmental data from freshwater p l o t s . 1 = <25% cover; 2 = 26-50%; 3 =51-75%; 4 = 76-100%; • = absent  96  AXIS  1  4  F i g u r e 24 - P l o t of cover c l a s s codes of Carex l y n g b y e i on the o r d i n a t i o n of environmental data from freshwater p l o t s , 1 = <25% cover; 2 = 26-50%; 3 = 51-75%; 4 =76-100%; • = absent  97  X  2  < AXIS 1  F i g u r e 25 - P l o t of cover c l a s s codes of S a l i c o r n i a v i r g i n i c a on the o r d i n a t i o n of environmental data from saltwater p l o t s . 1 = <25% cover; 2 = 26-50%; 3 = 51-75%; 4 = 76-100%; • = absent  2  98  AXIS 1  F i g u r e 26 - P l o t of cover c l a s s codes of D i s t i c h l i s s p i c a t a on the o r d i n a t i o n of environmental data from s a l t w a t e r plots. 1 = <25% cover; 2 = 26-50%; 3 = 51-75%; 4 = 76-100%; • = absent  4  AXIS  1  1 1  F i g u r e 27 - P l o t of cover c l a s s codes of T r i g l o c h i n maritimum on the o r d i n a t i o n of environmental data from saltwater p l o t s . 1 = <25% cover; 2 = 26-50%; 3 = 51-75%; 4 = 76-100%; • = absent  100  8.  8.1  DISCUSSION  Data S t a n d a r d i z a t i o n s  Square root t r a n s f o r m a t i o n ; As may be seen Fig.  11  with  F i g . 13  r e s u l t of using square points  on  the  and root  which  comparing  F i g . 12 with F i g . 14, the general transformed  data  was  that  the  s c a t t e r p l o t s were spread out more evenly.  p r a c t i c a l b e n e f i t of t h i s was a r e d u c t i o n plots  by  were  overprinted  in  the  by the computer.  number  A of  This effect  made l e g i b l e the o r d i n a t i o n of the complete cover data s e t , as shown i n F i g . 15. helpful  The use of t h i s t r a n s f o r m a t i o n may thus  f o r improving  the  readability  be  of h i g h l y congested  o r d i n a t i o n diagrams. The clumps  square of  between  spatial  clumps,  pattern  d i s t o r t i o n , at l e a s t percent  on  became  the  first  of t o t a l v a r i a n c e accounted  the  distinct  f i r s t . three  more  evident.  The  was preserved with r e l a t i v e l y minor  i s a l i t t l e lower with square for  previously  p l o t s i n d i s t i n c t , but trends i n the data, i . e . the  relationships overall  root t r a n s f o r m a t i o n made  and  second  The  f o r by the p r i n c i p a l axes  root transformed  axes  axes.  (transformed  data;  e.g. 61%  freshwater  data)  ordination  axes  w e l l with those obtained from untransformed  data,  compared with 70% (untransformed). The  variable  corresponded  weightings  on  the  101  at  l e a s t on the f i r s t  species  were  species  with  less  of  data  quite  dominate  different a tendency  the  the weightings  important  weightings, were somewhat  o r d i n a t i o n s , the  species  By a m p l i f y i n g  low-cover quadrats,  p l o t s being with  in  important  increased  resulted  in  easier  i n t e r p r e t a t i o n of the t h i r d a x i s .  Normalization: content  greatly  In the freshwater  of  ecological  to  transformed  more e q u i t a b l e . weighting  weighted  In the untransformed data, there was  a single  whereas  axes; on higher axes, the  sometimes  proportions. for  two  grouped together  respect  to  the  the  effective  normalization  according  proportionate  to  information  r e s u l t e d i n the  their  similarity  r e p r e s e n t a t i o n of  their  dominant s p e c i e s . As  with  the  square  root  t r a n s f o r m a t i o n , - the  broad  p a t t e r n s of the o r d i n a t i o n s were not changed ( c f . F i g s . 11  and  16,  The  12  percent  and  for the f i r s t  v a r i a n c e accounted for on the  slightly with  17), at l e a s t  reduced.  the  first  The and  and  second axes.  first  three  axes  was  s p e c i e s showing strongest c o r r e l a t i o n s second  axes  were  the  same,  though  d i f f e r e n c e s appeared in the t h i r d a x i s . In  F i g . 16,  apparent;  these  several groups  entities  characterized  important  species  americanus). ordination  distinct constitute  by  their  of  quadrats are  floristically  distinct  r e l a t i v e abundance of  ( A g r o s t i s a l b a , Carex l y n g b y e i , and  A similar pattern of  groups  non-normalized  of data  plots  is  found  ( F i g . 11), but  three  Scirpus in  there  the the  1 02  groups are  not  distinctly  separated.  The  species  govern  position  i s i n f l u e n c e d by r e l a t i v e s p e c i e s  ,case  of  same  principal  the a x i s o r i e n t a t i o n s i n e i t h e r case, but  normalized  normalized data.  data,  and  For example,  p l o t s j u s t below the l e f t proportionately  in  the  a b s o l u t e abundance with  non-  F i g . 16  abundance  plot  shows  a  cluster  of  end of the f i r s t a x i s : these are a l l  high i n S c i r p u s americanus.  In p l o t s A29 and  A30,  however, the a b s o l u t e cover of S c i r p u s americanus i s very  low;  they are l o c a t e d f a r out on a mudflat  amer icanus and Ruppia maritima 11,  however,  and  to the l e f t  which  they  the  of  similarity,  species  grounds  Typha  latifolia,  of  and  species  n o r m a l i z a t i o n procedure it  causes  between s i t e s  to  be  differences,  as  expressed  AO,  can  important  overlooked in  —  be  i.e. site plant be  to  among samples on the b a s i s of t h e i r  in  composition,  procedure floristic  on  in site productivity.  the  while  (Refer to  the  differences productivity performance. assess  the  resemblance  basis  of  their  N o r m a l i z a t i o n of data i s a  that enables the most e f f e c t i v e use to information,  floristic  c r i t i c i z e d ~ on  ecological  overall  not  only  of the p l o t s . )  relationships  resemblance  with  containing  performance.  However, the purpose of the o r d i n a t i o n may  floristic  plots  N o r m a l i z a t i o n thus emphasized  regardless  that  Fig.  at a l l i n common -- e.g. D3, a  Table III f o r the s p e c i e s composition The  In  p l o t s appear i n a t i g h t c l u s t e r , below  no  Eleocharis palustris•  Scirpus  occur, very s p a r s e l y .  of the o r i g i n , along with s e v e r a l  have  dense stand  two  where only  sacrificing  site  be  made  of  productivity  1 03  information. involve  An  the  optimal  analysis  ordination  of  strategy  both normalized and  might  thus  non-normalized  data. C o r r e l a t ion  matrix:  The  correlation  matrix  is  s t a n d a r d i z e d v a r i a n c e - c o v a r i a n c e matrix: each c o v a r i a n c e for  a  species  pair  the two s p e c i e s .  a  value  i s d i v i d e d by the standard d e v i a t i o n s of  In e f f e c t ,  the v a r i a b l e s are s t a n d a r d i z e d to  zero mean and u n i t v a r i a n c e . The  effect  correlation  of  values  then  plots  in  common.  highest  to the r a r e s t s p e c i e s , which The  principal  components  to the r a r e s t  were c l u s t e r e d t i g h t l y around the o r i g i n .  When the  from  obtained, but I found  the  i n only  data,  one  f o r on the major axes.  data,  three  axes accounted  two  quadrats)  ordinations  were  to i n t e r p r e t .  decreased  v a r i a n c e accounted first  or  improved  the r e s u l t s d i f f i c u l t  of the c o r r e l a t i o n matrix  the  the  majority  removed  Use  that  i n o r d i n a t i o n s i n which the great  r a r e s t s p e c i e s (those present were  was  gave the h i g h e s t a x i s weightings  species, resulting of  procedure  corresponded  had many zero values solution  this  the  amount  of  With the freshwater f o r only 34% of t o t a l  variance. The with  c o r r e l a t i o n matrix does not seem a p p r o p r i a t e f o r use  vegetation  data,  although  perhaps be obtained with data Under such  from  satisfactory very  r e s u l t s might  homogeneous  stands.  favourable c o n d i t i o n s , the c o r r e l a t i o n matrix might  1 04  be  recommended,  variance (Austin  i f axis  d i d not  g e n e r a t i o n on the b a s i s of maximum  produce  ecologically  meaningful  1969).  Rec i p r o c a l initial  averaging:  s p e c i e s and quadrat  disappointing, on the f i r s t  As  noted  in  Section  7.2,  o r d i n a t i o n s obtained with RA  second  were  a x i s , and a l l the other quadrats crowded together  axis).  Examination  species,  corresponding  high  which were themselves  species o r d i n a t i o n s .  unique  to  those  along  the  of the raw data r e v e a l e d that the  anomalous o u t l y i n g quadrats had very uncommon  the  with one or two quadrats appearing as o u t l i e r s  towards the other end (although with a good spread  species  results  cover  values  for  separated out i n the  When o u t l y i n g  plots  (and  p l o t s ) were removed from the data,  acceptable  ordinations  were  distortion  by  i s a c h a r a c t e r i s t i c problem with RA  outliers  (Gauch, Whittaker  obtained  ( F i g . 18).  & Wentworth 1977), but i t i s e a s i l y  This  dealt  wi t h . Reciprocal  averaging  some i n v e s t i g a t o r s 1977;  Robertson  e c o l o g i c a l data. ordination  has  been found s u p e r i o r to PCA by  (Austin 1976; Gauch, Whittaker 1978;  d e l Moral  Wentworth  1980) f o r the o r d i n a t i o n of  RA has been found t o  i n one dimension  &  (del Moral  produce  an  efficient  1980), and H i l l  (1973)  s t a t e d that when there i s a long f l o r i s t i c  g r a d i e n t , i t would  always  axis.  be presented l i n e a r l y on the f i r s t  c o n t r a s t , stands extreme on the f i r s t extreme on the f l o r i s t i c  With PCA, by  a x i s are not n e c e s s a r i l y  g r a d i e n t , and v i c e  versa.  Although  105  RA  may  yield  spurious r e s u l t s on the second  a x i s , with p l o t  p o s i t i o n r e f l e c t i n g only the degree of displacement first  axis  (Gauch,  Whittaker second  e c o l o g i c a l l y meaningful.  D e l Moral  usually  Whittaker high  RA  axis  can  sometimes  (1980) suggested  be the case when beta d i v e r s i t y  diversities,  be  that t h i s  i s low.  Gauch,  & Wentworth (1977) found RA much s u p e r i o r to PCA  beta  the  & Wentworth 1977), t h i s i s not  always the case, and the  will  from  at  and g e n e r a l l y p r e f e r a b l e at low beta  diversities.  8.2  E c o l o g i c a l I n t e r p r e t a t i o n of O r d i n a t i o n R e s u l t s  8.2.1  The from  V e g e t a t i o n Data  PCA o r d i n a t i o n of square  a l l 103  plots  Table IV) produced for  root transformed  a good  first-axis  separation,  23% of t o t a l v a r i a n c e , of the freshwater p l o t s  E,  F  ,G,  H)  on  the  left,  both  p l o t groups (Carex  freshwater p l o t s ; D i s t i c h l i s and  Salicornia virginica  accounting (Transects  (Transects  B,  on the b a s i s of a f l o r i s t i c  g r a d i e n t r e l a t e d to high cover v a l u e s of in  data  ( s c a t t e r diagram, F i g . 15; e i g e n v e c t o r s ,  A, D, J) on the r i g h t , and the s a l t w a t e r p l o t s C,  cover  community  dominants  l y n g b y e i and A g r o s t i s a l b a i n the spicata,  Tr i g l o c h i n  maritimum,  i n the s a l t w a t e r p l o t s ) .  The second  a x i s , however (14% of t o t a l v a r i a n c e ) , loaded p o s i t i v e l y  with  106  Carex  lyngbyei,  D i s t i c h l i s s p i c a t a , and A t r i p l e x p a t u l a , and  n e g a t i v e l y with S c i r p u s Agrostis  alba,  interpretation, resulting  americanus,  does and  not  may  from high beta  Scirpus  lend  largely  maritimus,  itself  to  reflect  the  such  and ready  distortion  diversity.  Table IV - Summary of r e s u l t s of PCA o r d i n a t i o n on square r o o t s of s p e c i e s cover data from a l l p l o t s . Eigenvector elements i n the range -0.1500 to +0.1500 not shown  Axis  % variance  Ranked e i g e n v e c t o r elements  Species  1  22.93  0.562 0.513 -0.179 -0.249 -0.283 -0.407  Agrostis alba Carex lyngbyei Atriplex patula T r i g l o c h i n maritimum Salicornia virginica Distichlis spicata  2  14.47  0.695 0.610 0.150 -0.116 -0.124 -0.238  Carex lyngbyei Distichlis spicata Atriplex patula Agrostis alba Scirpus maritimus Scirpus americanus  The  PCA  ordinations  on  the  freshwater  diagrams,, F i g s . 11, 13; e i g e n v e c t o r s , Table plots  on the f i r s t a x i s (accounting  in the o r d i n a t i o n of the high  V)  plots (scatter separate  f o r 32% of t o t a l  untransformed  data)  the  variance  between  those  i n Carex lyngbyei on the r i g h t and those high i n S c i r p u s  americanus on the l e f t . reveals  that  I n s p e c t i o n of the environmental  this f l o r i s t i c  g r a d i e n t corresponds  data  strongly to  107  elevation;  i t i s i n f a c t h i g h l y conspicuous  Brunswick  Point,  lyngbyei to  a  where  a  broad  belt  in  the  dominated  i n the upper and middle p a r t of the marsh  wide  field by  of  total  variance),  the  Carex  gives  f r i n g e of S c i r p u s americanus at the lower  On the second a x i s (28%  at  way  levels.  floristic  g r a d i e n t runs from high cover values of A g r o s t i s a l b a (top) to high  values  correspond  of  Carex  lyngbyei  with any of the  (bottom).  environmental  This  gradients  does  not  sampled,  though i t may be r e l a t e d to l o c a l v a r i a t i o n s i n topography and drainage,  with  the h i q h - A g r o s t i s group being  p o o r l y d r a i n e d areas richer,  and  where  where Scirpus  the  community  l o c a t e d i n more  is  floristically  maritimus tends to r e p l a c e Carex  lyngbyei.  Table V - Summary of r e s u l t s of PCA o r d i n a t i o n on s p e c i e s cover data from freshwater p l o t s . E i g e n v e c t o r elements i n the range -0.150 to +0.150 not shown  Axis  % variance  Ranked e i g e n v e c t o r elements  Species  1  31.59  0.912 0.232 -0.309  Carex l y n g b y e i Agrostis alba S c i r p u s americanus  2  27.50  0.893 0.229 -0.219 -0.293  Agrostis alba S c i r p u s maritimus S c i r p u s americanus Carex l y n g b y e i  In the (scatter  PCA  diagram,  ordination F i g . 16;  on  normalized  eigenvectors,  freshwater Table  data  V I ) , three  108  floristic the  nodal groups may be d i s t i n g u i s h e d .  second a x i s  usually,  At  the  i s a group of p l o t s r i c h i n A g r o s t i s  S c i r p u s maritimus; at the l e f t  i s a group of p l o t s r i c h  in Scirpus  other  plots  these s p e c i e s ,  alba and,  end of the f i r s t  americanus;  and  lower r i g h t corner i s a group of p l o t s high i n Carex The  top of  axis  i n the lyngbyei.  e i t h e r are dominated by some combination of  or are not r i c h i n any of them (the group  above and t o the l e f t  just  of the o r i g i n ) .  Table VI - Summary of r e s u l t s of PCA o r d i n a t i o n on normalized species cover data from freshwater p l o t s . E i g e n v e c t o r elements i n the range -0.150 to +0.150 not shown  Axis  % variance  1  27.89  2  22.04  The Figs.  Ranked e i g e n v e c t o r elements -  0.576 0.359 -0.719  Carex l y n g b y e i Agrostis alba S c i r p u s americanus  0.688 0.161 -0.184 -0.661  Agrostis alba Scirpus maritimus S c i r p u s americanus Carex l y n g b y e i  :  PCA o r d i n a t i o n s  on s a l t w a t e r data  12, 14; e i g e n v e c t o r s , Table  spicata-dominated  Species  plots  VII)  (right)  from  ( s c a t t e r diagrams,  separated plots  Distichlis  dominated  T r i g l o c h i n maritimum, S a l i c o r n i a  virginica,  canadensis  a x i s , and p l o t s dominated by  (left)  Atriplex patula  on  the f i r s t  (top) from  those  dominated  and  by  by  Spergularia  Salicornia  109  virginica,  Triglochin  Distichlis  maritimum,  Carex  lyngbyei,  s p i c a t a (bottom) on the second a x i s .  The  and/or  floristic  gradient  on the f i r s t a x i s does not seem to correspond with  gradient  i n any of the measured environmental f a c t o r s , whereas  the  second a x i s shows a strong  ordination  on  r e l a t i o n s h i p to e l e v a t i o n .  normalized s a l t w a t e r  in  the  of f i v e  plots  (including  the  Distichlis  lower r i g h t , while r e v e a l i n g greater  the group of p l o t s at l e f t , which  each  d e t a i l in  i n p a r t i c u l a r by i s o l a t i n g a group  were  dissimilar  other).  Taken  to  a l l other  together,  Atriplex  Distichlis species  patula;  a  group  dominated  by  --  Carex  The' RA  left alba  Atriplex  ordination  of  group  freshwater  plots a  dominated  ( F i g . 18) strong  The strong  separation  and  on both axes of  PCA o r d i n a t i o n , i n which these p l o t s f a l l swarm.  gradient  into  or  Scirpus from the  near  the  More i n t e r e s t i n g i s the arch-shaped sequence  extending from lower l e f t floristic  first-  Agrostis  validus-dominated p l o t s would not have been p r e d i c t e d  central  is  of the S c i r p u s americanus-dominated p l o t s at  from the p l o t s dominated by Carex lyngbyei at r i g h t .  other  Spergularia•  i n part to the PCA o r d i n a t i o n , with  separation  and  l y n g b y e i , Tr i g l o c h i n maritimum, S a l i c o r n i a  by S a l i c o r n i a , Tr i g l o c h i n , and  axis  dominated  s p i c a t a ; a group dominated by D i s t i c h l i s and  v i r g i n i c a , and S p e r g u l a r i a canadensis; and a  similar  plots  these o r d i n a t i o n  r e s u l t s suggest four nodal groups of p l o t s : a group by  The  data ( F i g . 17) added more  p l o t s to the A t r i p l e x group at the top and to group  a  to  lower  right,  which  follows  from high-Carex p l o t s at lower r i g h t  a  (high-  110  Table VII - Summary of r e s u l t s of PCA o r d i n a t i o n on s p e c i e s cover data from s a l t w a t e r p l o t s . E i g e n v e c t o r elements in the range -0.150 to +0.150 not shown  Axis  % variance  1  Ranked e i g e n v e c t o r elements  35.73  Species  0.941 -0.168 -0.176  Distichlis spicata Atriplex patula Salicornia virginica T r i g l o c h i n maritimum  0.869 -0.200 -0.264 -0.329  Atriplex patula Salicornia virginica Distichlis spicata T r i g l o c h i n maritimum  0.186  2  15.77  elevation)  through  a  floristically-rich  which A g r o s t i s i s prominent low-elevation  plots  (high  dominated  to by  s e r i e s of p l o t s i n  medium  elevations)  S c i r p u s americanus.  to  (The  dense c l u s t e r at r i g h t c e n t r e i s formed by uncommon, low-cover species.)  A linear  f l o r i s t i c g r a d i e n t i s curved  second o r d i n a t i o n dimension, but i s e f f e c t i v e l y than  8.2.2  The freshwater  somewhat  Data  ordination  on  environmental  and s a l t w a t e r p l o t s together  data  group from the s a l t w a t e r group on the f i r s t f o r 51% of t o t a l  variance,  from  the  ( s c a t t e r diagram, F i g .  19; e i g e n v e c t o r s , Table V I I I ) c l e a r l y separated  accounting  more  by PCA.  Environmental  PCA  displayed  here i n t o the  with  the freshwater  (horizontal) axis, the  two  groups  111  appearing on opposite s i d e s of the o r i g i n . weighted  negatively  were weighted The  second  on  Silt  and c l a y were  t h i s a x i s ; the other seven  positively,  especially  sodium  and  variables potassium.  ( v e r t i c a l ) a x i s , accounting f o r 23% of v a r i a n c e , i s  basically  textural  and  n u t r i t i o n a l , with sandy p l o t s toward  the bottom, and p l o t s h i g h i n c a t i o n s , s i l t ,  and  clay  toward  the t o p .  Table VIII - Summary of r e s u l t s of PCA o r d i n a t i o n on environmental data from freshwater and s a l t w a t e r p l o t s together  E i genvector Sand Silt C 1 ay N K Ca Mg Na Elev %  1  3  0 .3717 -0 .3813 -0 . 2781 0 .3101 0 . 4059 0 . 1967 0 . 35 13 0 . 4095 o . 2203  -o .3914 0 . 3462 0 . 4306 0 .0361 0 . 158 1 0 . 5224 0 . 4409 0 .2021 -0 . 0839  50.67  22.87  variance  The Fig.  2  -o . 1685 0 .0916 0 . 3282 0 . 5272 -0 .0439 -0 . 2366 -o . 1332 -0 .0470 o . 7069 13 . 24  4 -0. 1389 O. 1077 0. 1907 0. 1239 0.4342 -0. 5998 -0. 1276 0.4424 -0. 3964  6  -0. 0476 0. 1934 -0..3240 0.7287 -0.. 3708 0. 1060 -0..0109 0..0028 -0..4189  5 . 76  axis  (39%  of  total  separated  right,  low-sand,  the  environmental  weighted  data  0.95  0.13  ( s c a t t e r diagram, plots  along  the  v a r i a n c e ) mainly on the b a s i s of  high-cation  v a l u e s of the environmental  8  -0..0058 -0..0114 0 .0119 0..0123 -0 .0108 0 .0073 -0 . 2248 0..0314 -0 .6181 0 .2164 -0..0591 0 .5018 0 .0006 -0 .8044 0 . 7334 0 . 2299 O .0140 0 . 1605  1 .97  4 . 40  t e x t u r e and c a t i o n c o n c e n t r a t i o n : high-sand, at  7  0.. 2370 -0.. 5785 0..6578 0..1634 -0..2311 0..0787 -0..0332 -0..053 1 -0.. 2936  o r d i n a t i o n of the freshwater p l o t s  20/ e i g e n v e c t o r s , Table IX)  first  5  plots  v a r i a b l e s at  low-cation  at each  t a b l e i n Appendix B.)  left.  plots  (For the  plot,  consult  Only  sand was  p o s i t i v e l y ; the s t r o n g e s t negative weighting was  for  1 12  Table  IX - Summary of r e s u l t s of PCA o r d i n a t i o n on e n v i r o n m e n t a l d a t a from f r e s h w a t e r plots  E igenvector Sand Silt Clay N K Ca Mg Na Elev %  1  2  0. 3991 -0.3430 -0.3396 -0.2509 -0.4242 -O.3551 -0.3980 -0.2598 -0.1139  variance  0 . 3262 -0 . 2394 -0 . 3398 -0 . 3406 0 . 1729 0 . 3753 0 . 3706 0 . 3928 -0 . 3775  sodium,  On  generally matter  sandy,  at  -0 0 0 0 0 0 0  .0040 .4635 .0628 .0744 . 1222 . 1392 . 5903  13 . 42  ends of  first  the  21;  eigenvectors,  plots  separated  out  Brunswick  Point  axis  sand  contributed  at on  (30%  at  the  of  right,  on t h e total  plots  at  rich  in  cations  plots  first  the  Bay  (Transects  organic  (Transects  variance)  is  B  not  diagram, total  with  sandy,  plots  rich  are  thus  and H) from and  largely  w i t h c a l c i u m and magnesium c o n t r i b u t i n g the  is  (45% of  plots G  from  opposite  (scatter  axis  of  second.)  and l o w e r - e l e v a t i o n  left  were  top  at  and e l e v a t i o n ,  The B o u n d a r y right  clay  interpretation  saltwater  by t e x t u r e  strong  was a s e p a r a t i o n  same end of  the  variance),  and  plots  and  the  Table X ) ,  left.  plots  sand  and a t  total  nitrogen,  0.13  2 . 22  and  (Ecological  was d e t e r m i n e d  -0.0037 0.0032 0.0040 0.0848 -0.1989 -0.5951 0. 7626 -0.0815 -0.1040  -0. 0857 0. 1543 0. 5621 -0. 5461 0.. 3226 -0. 0166 0. 0467 -0..4980  Of  fine-textured  of  -o.0108  (28%  low-elevation  axis,  8  4 . 42  cation-rich,  l o a d i n g of  7  axis  elevation,  ordination  and c l a y  5.15  The r e s u l t  the  high-eleyation  -0. 1250 0. 2153 -0. 0579 -0. 1804 -0. 5161 -0. 0981 ' -0. 0719 0. 7216 0. 3167  negatively.  In  variance)  6  -0.. 4639 0..7581 -0,. 2788 0..0740 -0.. 1884 -0..0996 0..221 1 -0.. 1787  7.18  second  bottom. by the  second  -o..0243  high-elevation,  facilitated  silt  0. 0492 0. 0649 -0. 2048 0. 4157 0. 4202 -0..4735 -0. 3039 0.. 4248 -0..3241  magnesium,  weighted  generally  in  0 . 3623  weightings;  strongly  Fig.  5  -o .5115  the  calcium,  positive  4  28 . 14  39 . 34  potassium.  3  C).  the The  nutritional,  strongest  positive  1 13  l o a d i n g s and sand cation-poor  the  plots  only  H11  negative  loading.  The  sandy,  and G6 were thus separated out at the  bottom, and the s i l t y , c a t i o n - r i c h C8 was i s o l a t e d at the top. The  t h i r d o r d i n a t i o n a x i s (11% of t o t a l v a r i a n c e ;  strongly  separated  potassium  plots  potassium  high-nitrogen,  from  not  shown)  high-elevation,  low-nitrogen,  low-  low-elevation,  high-  p l o t s -- b a s i c a l l y an e l e v a t i o n g r a d i e n t .  Table X - Summary of r e s u l t s of PCA o r d i n a t i o n on environmental data from s a l t w a t e r p l o t s  E igenvector Sand Silt Clay N K Ca Mg Na Elev %  variance  1 0 . 4426 -0 . 4370 -O . 4275 0 . 2881 0 .3022 -0 .01 13 0 . 1406 0 .2817 O . 3960  -0 .2213 0 . 2216 0 . 2057 0 .0081 0 . 2344 0 . 5576 0 . 5742 0 .4010 0 .056 1  45 . 37  29 . 79  An examination be  3  2  helpful.  In  -0 . 1654 0 . 1803 0 . 1 152 0 . 7873 -0 . 4748 -0 .0103 -0 .0438 0 .0498 0 . 2777 10. 74  4  6  5  -0. 2426 0. 19 11 0..361 1 0..026 1 0.. 5839 -0..4274 -0.. 1288 -0 .0550 0.. 4800  0. 1037 -0. 2035 0. 1651 -0. 1397 -0. 1266' 0. 3720 0.. 1920 -0..6965 0. 4739  6 . 75  4 . 19  a l l the  datasets,  0.64  2 .48  magnesium  and  correlated,  potassium  The n u t r i e n t calcium  other  but  with  strongest  (positive),  reflecting  matter  peat  and  at  correlations the high  greater  with  E l e v a t i o n and n i t r o g e n  do not show such strong c o r r e l a t i o n s as the their  0.03  sand and s i l t / c l a y are  especially  with sodium.  -0. 0603 0..1415 -0. 1561 0..0299 0..2113 0.. 5929 -0 . 7346 0 , 0569 0 . 1093  (Table XI) may  s t r o n g l y or very s t r o n g l y n e g a t i v e l y c o r r e l a t e d . c a t i o n s are p o s i t i v e l y  0. 1939 -0. 5296 0. 6990 0..2117 0.. 1061 0. 1 133 -0..1712 0.. 1030 -0.. 2907  0. 0290 -0. 1 178 •o. 2056 -0. 4808 -0. 464 1 -0. 0586 -0. 1616 0..5053 0.. 4599  of the c o r r e l a t i o n matrices  8  7  are  accumulation  elevations.  In  the  variables, each  other  of  humic  freshwater  1 14  dataset,  sand i s n e g a t i v e l y c o r r e l a t e d with e v e r y t h i n g e l s e --  r e f l e c t i n g the  difference  nitrogen-poor,  lower-elevation  and  between  the  plots  the f i n e r - t e x t u r e d p l o t s , r i c h e r  matter,  that occur at higher  Point and Ladner Marsh and  sandy,  cation-poor,  along the Fraser  in  cations  and  e l e v a t i o n s i n northern  in  the  more  River  organic Brunswick  brackish  marsh  in  southern Brunswick P o i n t . In  the  saltwater  dataset,  however, sand i s p o s i t i v e l y  c o r r e l a t e d with e l e v a t i o n , potassium, suggesting plots  a  at  gradient  high  nitrogen-poor misleading, variation  lower-elevation saltier,  at  to  lower  r e f l e c t i n g as between  and  nitrogen,  from sandy, c a t i o n - r i c h , n i t r o g e n - r i c h  elevations plots  sodium,  finer-textured, elevations.  i t does  mainly  cation-poor,  T h i s i s somewhat the  geographical  the f i n e r - t e x t u r e d , perhaps more b r a c k i s h ,  Brunswick  higher-elevation  Point  marshes  marshes  at  and  the  sandier,  Boundary Bay.  In the  Brunswick Point marshes, t e x t u r e does become f i n e r towards the lower e l e v a t i o n s ; at Boundary Bay, however,true.  The  trends  the  opposite  i n n i t r o g e n and c a t i o n s with e l e v a t i o n are  not c l e a r , but g e n e r a l l y the h i g h e s t values do occur elevations. the  Apparently  saltwater  is  the p r i n c i p a l  "gradient"  PCA o r d i n a t i o n was g e o g r a p h i c a l .  at higher  exposed This  by  implies  that beta d i v e r s i t y was too high; the o r d i n a t i o n r e s u l t s would probably  be improved by o r d i n a t i n g  the  Brunswick  Point  and  Boundary Bay p l o t s s e p a r a t e l y . It  is  important  to  realize  that  the  results  of an  1 15  Table XI - Product-moment c o r r e l a t i o n c o e f f i c i e n t s f o r environmental v a r i a b l e s (* = p < .05; ** = p < .01)  1.  Combined Sand Silt Clay N K Ca Mg Na Elev  freshwater  1 .000 -O.979** -0.864** 0.374* 0.535** 0.004 0 . 275 0.506** 0.323*  1 .000 0.744** -0.410** -0.578** -0.029 -0.315* -0.542** -0.389**  Sand  Silt  Freshwater  3.  and  plots.  Sample  saltwater  1 .000 - 0 . 2 16 -0.327* 0.057 -0.12 1 -0.322* -0.097  size  1.000 0.441* 0.277 0.338 0.164 0.185 0.022  1.000 0.461* 0.326 0.112 0.165 -0.020  Elev  -0.237  0.062  0.416*  Sand  Silt  Clay  Sand Silt C1 ay N K Ca Mg Na Elev  1 .000 -0.992** -0.950** 0 . 380 0.390 -0.271 -0.055 0.248 O.585** Sand  Sample .  1.000 0.907** -0.352 -0.396 O. 269 0.062 -0.224 -0.608**' Silt  size  Sample s i z e  000 484** 163 433** 578** 59 1 * *  1 0 0 0 0  000 396** 772 * * 889** 3 19*  = 44.  1 0 0 0  K  N  000 860** 457** 008 Ca  1 000 0 815** 0 194 Mg  1 .000 O. 258 Na  = 27.  1.000 -0.909** -0.774** -0.408* -0.389* -0.167 -0.207 -0.006  plots.  1 0 0 0 0 O  C l ay  Sand Silt Clay N K Ca Mg Na  Saltwater  plots.  1 ooo 0 34 1 0 027 0 037 0 008 0 614** 0.614** N N  1 OOO 0 554** 0 697** 0 542** -0 053 -0.053 K K  1 000 0 961** 0 538** -0 091 -0.091 Ca Ca  1 000 0 640** -0 043 -0.043 Mg Mg  1.000 -0.192 -0.192 Na Na  1 000 0 7 11** 0 284  1 .000 0 . 438  = 17.  1 .000 -O.427 -0.348 0 . 257 0.036..-0.293 -0.485* C 1 ay  1 0 -0 0 0 0  OOO 066 027 147 360 608** N  1 0 0 0 0  000 178 1 514* 0 538* 0 494*. -0 K  000 912** 496* 003 Ca  Mg  Na  1.000 Elev  11 6  o r d i n a t i o n depend on the choice In  a  vegetation  selected;  in  of  variables  (Austin  1969).  o r d i n a t i o n , the v a r i a b l e s ( s p e c i e s ) are  an  environmental  ordination,  pre-  however,  the  i n v e s t i g a t o r makes a s u b j e c t i v e d e c i s i o n as to which v a r i a b l e s to  use.  Thus  a  different  choice  of v a r i a b l e s might have  produced very d i f f e r e n t o r d i n a t i o n r e s u l t s .  8.2.3  The  Species-Environment Diagrams  p r o j e c t i o n s of cover c l a s s codes of  species  on  ordinations  the  freshwater  and  important  saltwater  marsh  environmental  ( F i g s . 22-27) r e v e a l c l e a r p a t t e r n s  that  can  be  interpreted ecologically. Considering Agrostis  alba  (textural) along  first ( F i g . 22)  axis,  but  the second a x i s  pattern  for  the  freshwater s e r i e s , one  performs  seems  ( F i g . 24)  i t s performance p a t t e r n  low  sides  of  ( F i g . 23) The  i s more or l e s s the  d i s t r i b u t i o n pattern  reflect  patterns  of  of but'  Point,  species  S c i r p u s americanus i s found at  (top of diagrams) on both the north  Brunswick  The  is different.  i n the marsh i t s e l f .  elevations  first  i s s i m i l a r to that of A g r o s t i s ,  These o r d i n a t i o n p a t t e r n s zonation  the  (cationic, elevational, textural).  Sc i rpus amer icanus  lyngbyei  along  to form three performance zones  i n v e r s e of the A g r o s t i s p a t t e r n . Carex  evenly  sees that  extending  lower  than  and  south  Carex or  117  Agrostis.  At middle and upper  l e v e l s , Carex and A g r o s t i s  dominate,  but Carex outperforms A g r o s t i s at higher e l e v a t i o n s  ( f i n e r - t e x t u r e d , lower i n c a t i o n s , higher i n n i t r o g e n ; of  co-  bottom  diagrams). In  the  saltwater s e r i e s , S a l i c o r n i a v i r g i n i c a  shows i t s e l f plots  at  right.  to be mostly  left,  The  gradient  Examination  however,  the  Brunswick  Point  the  was  based  on  texture  and  of the data suggests that t e x t u r e i s  probably not the f a c t o r here;  from  but present i n a l l the Boundary Bay p l o t s at  first-axis  elevation.  absent  ( F i g . 25)  controlling two  S a l i c o r n i a does occur are  Salicornia  Brunswick the  two  Point  distribution  plots  highest  in  which  Brunswick  Point  . p l o t s i n the environmental d a t a s e t , suggesting a p o s s i b l e for  elevation  Brunswick  i n r e s t r i c t i n g the d i s t r i b u t i o n of S a l i c o r n i a at  Point.  Distichlis of  a x i s extremes,  marsh.  An  spicata  ( F i g . 26) seems to show an avoidance  which corresponds to i t s d i s t r i b u t i o n  important component of  middle l e v e l s , D i s t i c h l i s the  role  the  salt  i s outcompeted  marsh  i n the  flora  at  by A t r i p l e x p a t u l a at  h i g h e s t e l e v a t i o n s , and does not extend i t s e l f  l e v e l s where c o l o n i z a t i o n of the unvegetated f l a t s  to the low is  taking  place. Triglochin amplitude, gradient  maritimum  apparently extremes.  ( F i g . 27) shows a broad  tolerant  This  of  all  corresponds  observed i n the marsh: T r i g l o c h i n was  to  major the  found on  ecological ordination  distribution all  transects  118  except  Transect  r e v e a l s an the  J,  at  Ladner  Marsh.  interesting pattern: f a i r l y  Boundary  Bay  However, the f i g u r e  low,  uniform  cover  p l o t s ; more v a r i a b l e performance, with some  high cover values and  some absences, at Brunswick  Point.  may  be that the Brunswick Point marsh, c l o s e r to the  of  the  Fraser  River,  f l u c t u a t i o n s which may program.  This  in  might  is  It  influence  more s u s c e p t i b l e to environmental  not have been detected also  by  be an e x p l a n a t i o n  my  sampling  f o r the poorer  performance of S a l i c o r n i a at Brunswick P o i n t .  8.3  Plant Communities in the Study Area  In t h i s study, I have emphasized the continuous nature of v e g e t a t i o n a l v a r i a t i o n , rather than attempting  to d e f i n e p l a n t  a s s o c i a t i o n s , because I f e e l that the v e g e t a t i o n Delta t i d a l  Fraser  marshes i s best understood by using t h i s approach.  A number of other plant  of the  tidal  associations,  ecological  gradients,  supported  by  marsh s t u d i e s have rather with  available  than  individual  results  data.  attempted  that  Some  species,  tend  such  to  not  to  instances  fit to be were  mentioned in the L i t e r a t u r e Review. A  very  species  distinctive  forming successive  nonetheless a conspicuous  zonation  pattern,  with  e l e v a t i o n a l bands of dominance, i s and  fairly  universal  feature  tidal  marshes, i n c l u d i n g the F r a s e r D e l t a marshes.  there  is  a  very  clear  different  vegetational  and  of  Moreover,  environmental  1 19  disjunction  between  the  fresh  and  brackish  marshes on one hand and the s a l t marshes on the it  does  terms  not seem i n a p p r o p r i a t e  of  important  suggested field.  by  to d e s c r i b e  species-environment  ordination  results  marshes  Delta  other..  Thus  the v e g e t a t i o n i n nodes  that  are  and can be observed i n the  Such a d e s c r i p t i o n a l s o f a c i l i t a t e s  tidal  Fraser  comparisons  elsewhere that have been d e s c r i b e d  with  i n terms of  plant associations. Four of these nodal groups of samples may recognized lyngbyei  by  analysis  of the freshwater data:  - A g r o s t i s a l b a group on b e t t e r - d r a i n e d  to medium e l e v a t i o n s on s i l t y (2.)  an  drained  sites,  silty  tending  on s i l t y  low e l e v a t i o n s  nitrogen-rich;  plantago-aquatica  s i l t y c l a y loams. elevations,  Carex lyngbyei  nitrogen-  alba,  and  at medium to low e l e v a t i o n s on s i l t  f r e s h Ladner Marsh  nitrogen  medium  (4.) a group dominated  Scirpus v a l i d u s , Agrostis  data,  area.  four more groups emerge: (1.) an  A t r i p l e x p a t u l a group at the highest soil  to  f r e s h to b r a c k i s h areas on  c l a y loams t o sandy loams;  From the saltwater  lower  s i t e s at high  loams; (3.) a S c i r p u s americanus group at  in f a i r l y  loams i n the very  high  (1.) a Carex  to be more b r a c k i s h , at high  by Equisetum f l u v i a t i l e , Alisma  loams, sometimes  informally  A g r o s t i s a l b a - Scirpus maritimus group on l e s s w e l l  elevations  poor  be  elevations,  and a l s o o f t e n with driftwood, In a v a r i a n t Distichlis  of  this  spicata  - D i s t i c h l i s spicata  group  often  on sandy to at  co-dominates;  group  at  with  higher  somewhat (2.) a medium  120  elevations  on  loamy  concentrations; maritimum  sediments  (3.)  group  a  at  with high to medium n i t r o g e n  Salicornia  lower  e l e v a t i o n s , and  canadensis  variant  Salinity  Tr i g l o c h i n  Distichlis  variant  a Salicornia - Triglochin -  at  lower  canadensis group at low sands.  -  medium e l e v a t i o n s , d i s p l a y i n g  v a r i a n t s : a S a l i c o r n i a - Tr i g l o c h i n higher  virginica  elevations;  e l e v a t i o n s on  does  not  Spergularia  loams  seem to play an  at  Spergularia  (4.) a  sandy  two  to  loamy  important r o l e in  d i s t i n g u i s h i n g these groups. To p l a c e the  the Fraser D e l t a marshes in a r e g i o n a l  context,  above species-environment nodal groups were compared with  published  d e s c r i p t i o n s of  D e l t a and  neighbouring Washington  In  marshlands  in  the  Rosenberg  (1980)  d i s t r i b u t i o n s of drainage  but  can  Tripp  described which  features.  r e s u l t s suggest  &  that be  Their  (1976)  twelve  they  and  plant  related  to  floristic  these  Burg,  of Puget Tripp  associations, topographical  table  associations  interpreted  Fraser  State.  the N i s q u a l l y s a l t marsh at the southern end  Sound, Burg, Rosenberg  units,  nearby  and  are  & the and  ordination  not  discrete  as species dominance phases  along  an e l e v a t i o n a l g r a d i e n t .  T h e i r sequence i s  that  observed  p l o t s , with Spergular i a mar ina  in my  at  dominated  by  Salicornia  v i r g i n i c a , D i s t i c h l i s s p i c a t a , Carex l y n g b y e i ,  and  species  intermediate  (some  lowest  not  l e v e l s , and  elevations,  found  in  the  zones  to  pioneering  other  the  saltwater  similar  Fraser  a zone dominated by Festuca  Delta)  at  rubra  and  121  Carex  lyngbyei  important  at  highest  levels.  Atriplex  patula,  i n the F r a s e r D e l t a s a l t marshes, was not p r e s e n t .  Two important marsh  the  occur  in  species i d e n t i f i e d the  Fraser  in  Delta  the  only  in  Nisqually  the freshwater  marshes: Juncus b a l t i c u s and Deschampsia c e s p i t o s a . that  their  and  salt  Assuming  my s p e c i e s i d e n t i f i c a t i o n s are c o r r e c t , t h i s  i l l u s t r a t e s the f a c t that the e c o l o g i c a l r o l e s of s p e c i e s not  necessarily  the  same  at  different  community and h a b i t a t c o n d i t i o n s a l t e r F a r t h e r north, at Bellingham brackish  marsh  sites;  different  the r e a l i z e d  Bay, D i s r a e l i  similar  are  niche.  & Fonda (1979)  described  a  floristically  to the  Brunswick  Point marsh, with a lower zone dominated by S c i r p u s  americanus and an upper zone dominated by Carex l y n g b y e i • On the western s i d e of L u l u I s l a n d i n the Hutchinson the  Delta,  (1982) a l s o d e s c r i b e d a marsh of a s i m i l a r type t o  Brunswick  communities  Fraser  Point  by  marsh.  He  identified  seven  a h i e r a r c h i c a l c l u s t e r i n g procedure,  plant relating  t h e i r d i s t r i b u t i o n s t o e l e v a t i o n and s u b s t r a t e t e x t u r e . Brunswick P o i n t , the dominated higher,  by  Scirpus  Island  americanus  f i n e r - t e x t u r e d , lower-cation  lyngbyei and other Bradfield at  Lulu  &  marsh and  has  seven  lower  S. maritimus,  zone dominated  zone and  by  a  Carex  species. Porter  (1982) sampled much more e x t e n s i v e l y  Ladner Marsh than I d i d i n t h i s study,  recognize  a  As at  community  types  (about  and  were  equal  able  to  i n l e v e l of  122  discrimination analysis.  The  physiographic the  to  Carex  -  my  nodal  distributions  and drainage Agrostis  of these  factors.  from  presence  Only one  of  my  freshwater different  own marshes  study of  in character found  area.  of these  several  the  This  lower  cluster  types were r e l a t e d to  species  River  that are  the quite  from the marshes at the d e l t a f r o n t .  descriptions  in  the  marshes o u t s i d e of the lower F r a s e r  literature  River.  own  that were not  illustrates  Fraser  types,  in my  major  the area of the B r a d f i e l d & Porter study  found i n  have not  variants)  type, would be r e c o g n i z a b l e  data -- a r e s u l t of the within  group  of  I  similar  1 23  9.  (1.) square  Transforming  CONCLUSIONS  species  cover  r o o t s p r i o r to o r d i n a t i o n can  data  lead  v a l u e s to t h e i r  to  more  o r d i n a t i o n diagrams by spreading out the p o i n t s . the  bias  from  data trends  not  change i n p l o t axes may  overemphasis of high-cover otherwise  scores on the f i r s t  Normalization  o r d i n a t i o n exposes  PCA  expose  causing  axes,  of  major  though higher  than  absolute  information  of s p e c i e s cover value data p r i o r to  sample  thus made of f l o r i s t i c  for  s p e c i e s may  without  two  Reduction  be c o n s i d e r a b l y changed.  (2.)  rather  evident,  readable  relationships  species  abundances.  information,  is sacrificed.  based  while  on  relative  E f f e c t i v e use i s site  productivity  As an optimal o r d i n a t i o n str.ategy  v e g e t a t i o n data, the a n a l y s i s of both normalized and  non-  normalized data i s recommended. (3.)  Use  of  the  correlation  matrix  f o r o r d i n a t i o n of  v e g e t a t i o n data i s not recommended f o r general (4.)  Reciprocal  anomalous  samples,  averaging but  is  may  produce  o r d i n a t i o n s , at l e a s t along the f i r s t are  removed  from  the  data.  highly  data were  produced  to  vegetation  a x i s , when such  RA and PCA  inferior.  sensitive good  o r d i n a t i o n s of e q u i v a l e n t q u a l i t y , but the RA environmental  purposes.  samples  vegetation  ordinations  of  124  (6.)  Ordination  environmental worthwhile  of  gradients aid  to  directly; the  ordinations.  The  environmental  ordination  species  The in  the  of  can  suggest  the  On  the  Delta t i d a l  basis  Point  of  the  and  plots  the  Boundary Bay  nitrogen  and  cation  and  environmental  i n t o two  main groups:  salt  Boundary and  from  southeast  marshes Bay.  of  Each  environmental the north and Brunswick  and  western  marshes of Ladner Marsh; southeastern group  may  be  differences:  the  west Brunswick Point  Point  plots  from  the  plots.  Based  on f l o r i s t i c  field-recognizable  and  environmental  species-environment  with v a r i a n t s , are  i n f o r m a l l y recognized.  respectively  (i)  by:  A g r o s t i s a l b a and (iv)  plant  marshes were shown to be  floristic  the freshwater  by f l o r i s t i c  p l o t s , and  eight  and  representing  Marsh  (9.)  an  environmental f a c t o r s .  Fraser  Point  Brunswick  Ladner  on  site elevation.  other  subdivided  a  vegetation  data  r e p r e s e n t i n g the b r a c k i s h marshes of northern  Brunswick  be  r e l a t i o n s h i p s between  f a c t o r s , the sample p l o t s were separated one  can  of  species  r e l a t e d to s u b s t r a t e t e x t u r e , s u b s t r a t e  (8.)  results  exposes  performance and d i s t r i b u t i o n of dominant  the  l e v e l s , and  data  interpretation  projection  s p e c i e s performance and (7.)  environmental  distinctions,  sample  groups,  These are dominated  Carex lyngbyei and A g r o s t i s a l b a ; ( i i )  S c i r p u s maritimus;  Equisetum f l u v i a t i l e ,  ( i i i ) S c i r p u s americanus;  Scirpus validus, Agrostis alba,  and  1 25  A l i sma  plantaqo-aquat i c a ;  Atriplex  patula  Distichlis  spicata; (variants:  maritimum  -  patula  Salicornia  Distichlis  maritimum  Spergularia  canadensis.  ( v i ) Carex lyngbyei and  -  spicata,  virginica  -  Triglochin  and S a l i c o r n i a v i r g i n i c a -  Spergularia  canadensis);  (viii)  The marshes of the study area are s i m i l a r to other and  salt  marshes  that  have been d e s c r i b e d i n the  F r a s e r D e l t a and i n Washington, but are mostly the  (variant:  ( v i i ) 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  Tr i g l o c h i n  brackish  Atriplex  - D i s t i c h l i s spicata);  maritimum  (10.)  (v)  f r e s h F r a s e r River marshes.  dissimilar  to  1 26  REFERENCES  Ages, A. 1979. The s a l i n i t y i n t r u s i o n in the Fraser River: salinity, temperature and c u r r e n t o b s e r v a t i o n s , 1976, 1977. P a c i f i c Marine Science Report 79-14. Canada Dept. of F i s h e r i e s and Oceans, I n s t i t u t e of Ocean S c i e n c e s , Sidney, B.C. Ages, A. & A. Woollard. 1976. The t i d e s in the F r a s e r E s t u a r y . 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Northwest S c i . 41:178-183. Stevenson, R.E. 1954. The marshlands at Newport Bay (California). Ph.D. thesis, Oregon State University, C o r v a l l i s , Oregon ( c i t e d i n D i s r a e l i & Fonda 1979). Stevenson, R.E. & K.O. Emery. 1958. Marshlands at Newport Bay, California. A l l a n Hancock Foundation o c c a s i o n a l paper no. 20. Stewart, G.R., F. Larher, I . Ahmad & J.A. Lee. 1979. Nitrogen metabolism and salt-tolerance i n higher p l a n t halophytes. Pages 211-227 i n : J e f f e r i e s , R.L. & A.J. Davy, eds. E c o l o g i c a l processes in coastal environments. Blackwell S c i e n t i f i c P u b l i c a t i o n s , Oxford. Stewart, G.R., J.A. Lee & T.O. Orebamjo. 1972. Nitrogen metabolism of halophytes. I. N i t r a t e reductase a c t i v i t y i n Suaeda maritima. New P h y t o l . 71:263-267. Stewart, G.R., J.A. Lee & T.O. Orebamjo. 1973. Nitrogen metabolism of halophytes. I I . N i t r a t e availability and u t i l i z a t i o n . New P h y t o l . 72:539-546.  1 38  Swan, J.M.A. 1970. An examination of some o r d i n a t i o n problems by use of simulated v e g e t a t i o n data. Ecology 51:89-102. Swinbanks, D.D. 1979. Environmental f a c t o r s c o n t r o l l i n g f l o r a l zonation and the d i s t r i b u t i o n of burrowing and tube-dwelling organisms on F r a s e r D e l t a t i d a l f l a t s , B r i t i s h Columbia. Ph.D. t h e s i s , Dept. of G e o l o g i c a l Sciences, U n i v e r s i t y of British Columbia, Vancouver, B.C. Tabata, S. 1972. The movement of Fraser River-influenced s u r f a c e water in the S t r a i t of Georgia as deduced from a series of aerial photographs. P a c i f i c marine science report 72-6. Environment Canada, Water Management S e r v i c e , Marine Sciences Branch, P a c i f i c Region, V i c t o r i a , B.C. Tamburi, A. & D. Hay. 1978. An i n t r o d u c t i o n to r i v e r mechanics and the lower Fraser R i v e r . P u b l i c Works Canada. Tide Tables 1976. High and low water p r e d i c t o r s . West coast of North and South America i n c l u d i n g the Hawaiian I s l a n d s . Issued 1975. United States Dept. of Commerce, N a t i o n a l Oceanic and Atmospheric A d m i n i s t r a t i o n , N a t i o n a l Ocean Survey, R o c k v i l l e , Maryland. Valiela, D. & R.U. K i s t r i t z . 1980. Dependence of salmon on Fraser Estuarine marsh ecosystems: a simulation analysis. T e c h n i c a l report no. 18. Westwater Research Centre, U n i v e r s i t y of B r i t i s h Columbia, Vancouver, B.C. Valiela, I . & J.M.. T e a l . 1974. N u t r i e n t l i m i t a t i o n marsh v e g e t a t i o n . Pages 547-563 i n : Reimold, R.J. Queen, eds. Ecology of halophytes. Academic Press, New Vermeer, K. & CD. L e v i n g s . 1977. food h a b i t s of ducks on the Fraser B r i t i s h Columbia. Wildfowl 28:49-60.  in s a l t & W.H. York.  P o p u l a t i o n s , biomass and Delta intertidal area,  Vogl, R.J. 1966. 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Ph.D. t h e s i s , U n i v e r s i t y of C a l i f o r n i a , R i v e r s i d e , and San Diego State U n i v e r s i t y . Wolf, E.G. & K.W. Fucik. 1981. E c o l o g i c a l s i g n i f i c a n c e of primary production on two Alaskan marshes. E s t u a r i e s 4:262 (abstract). Yamanaka, K. 1975. Primary p r o d u c t i v i t y of the F r a s e r River d e l t a f o r e s h o r e : y i e l d estimates of emergent v e g e t a t i o n . M.Sc. thesis, Dept. of P l a n t Science, University of British Columbia, Vancouver, B.C. Zedler, J.B. 1977. S a l t marsh community s t r u c t u r e i n the T i j u a n a Estuary, C a l i f o r n i a . E s t u a r i n e and C o a s t a l Marine S c i . 5:39-53. Z e d l e r , J . 1981. A r i d region wetlands: d i s t u r b a n c e . E s t u a r i e s 4:262 ( a b s t r a c t ) .  susceptibility  to  1 40  APPENDIX A.  NAMES OF  SPECIES SAMPLED, WITH AUTHORITIES  A c h i I l e a m i l l e f o l i u m L. Aqropyron repens ( L T ) Beauv. A g r o s t i s a l b a L. var. a l b a A g r o s t i s a l b a L. var. p a l u s t r i s (Huds.) Pers. Alisma plantaqo-aquatica L. A s t e r eatoni i (Gray") Howell Atr i p l e x p a t u l a L. var. h a s t a t a (L.) Gray A t r i p l e x p a t u l a L. var. p a t u l a A t r i p l e x p a t u l a L. var. z o s t e r a e f o l i a (Hook.) C. L. H i t c h c . Bidens cernua L. C a l l i t r i c h e L. sp. C a l t h a a s a r i f o l i a DC. Carex lyngbyei Hornem. Chara brauni i Gm. C o t u l a coronopi f o l i a L. Cuscuta s a l i n a Engelm. Deschampsia c e s p i t o s a (L.) Beauv. D i s t i c h l i s s p i c a t a (L.) Greene E l e o c h a r i s p a l u s t r i s (L.) R. & S. Elymus glaucus Buckl. var. b r e v i a r i s t a t u s Davy Elymus m o l l i s T r i n . i n Spreng. Equisetum f l u v i a t i l e L. Festuca arundinacea Schreb. Glaux mar i t ima L. G r i n d e l i a i n t e g r i f o l i a DC. var. macrophylla (Greene) Cronq. Hordeum brachyantherum Nevski Hygrohypnum luridum CHedw.) Jenn. Juncus a r t i c u l a t u s L. Juncus b a l t i c u s W i l l d . Juncus bufonius L. Juncus g e r a r d i i L o i s e l . Lathyrus L. sp. Leptodictyum r i p a r i u m (Hedw.) Warnst. L i l a e a s c i l l o i d e s ( P o i r . ) Hauman L i l a e o p s i s o c c i d e n t a l i s C o u l t . & Rose .Limosella aquat i c a L. Lythrum s a l i c a r i a L. Mentha a r v e n s i s L. Mimulus g u t t a t u s DC. v a r . guttatus Myosotis laxa Lehm. Oenanthe sarmentosa P r e s l Plantago maritima L. Polygonum a v i c u l a r e L. Potent i 1 1 a pac i f i c a Howell P u c c i n e l l i a nutkaensis ( P r e s l ) Fern. P u c c i n e l l i a n u t t a l l i a n a (Schult.) A. S. H i t c h c . Ruppia maritima L. Sagittaria l a t i f o l i a Willd*  141  S a l i c o r n i a v i r g i n i c a L. S c i r p u s americanus Pers. S c i r p u s maritimus L. S c i r p u s v a l i d u s Vahl Sium suave Wa1t. Sonchus a r v e n s i s L. S p e r g u l a r i a canadensis (Pers.) G. Don S p e r g u l a r i a marina (L.) G r i s e b . T r i f o l i u m oliqanthum Steud. T r i g l o c h i n maritimum L. Typha l a t i f o l i a L. Zostera americana den Hartog  Nomenclature except: (Hedw.)  is  according  Chara braun i i Gm. Jenn.  and  to  Hitchcock  a l . 1969,  (a green a l g a ) , Hygrohypnum luridum  Leptodictyum  r i pa r i um  (mosses), and Zostera amer icana den Hartog nana Roth  et  i n Hitchcock et a l . ) .  (Hedw.)  (which keys  Warnst. to  Z.  1 42  APPENDIX B.  ENVIRONMENTAL DATA  Explanation of a b b r e v i a t i o n s : SAMPL = sample p l o t ( i n i t i a l l e t t e r d e s i g n a t e s t r a n s e c t ) ; SAND = percent by weight of sand in soil sample; SILT = percent by weight of s i l t i n s o i l sample; CLAY = percent by weight of c l a y in soil sample; TXTCLASS = t e x t u r a l c l a s s i f i c a t i o n of s o i l sample (L = loam, LS = loamy sand, S = sand, SCL = sandy c l a y loam, SiCL = s i l t y clay loam, SiL = silt loam, SL = sandy loam); COND = e l e c t r i c a l c o n d u c t i v i t y of s o i l sample i n mS cm" ; N = percent by weight of n i t r o g e n ( t o t a l ) i n s o i l sample; K = p a r t s per m i l l i o n of potassium i n s o i l sample; CA = p a r t s per m i l l i o n of c a l c i u m i n s o i l sample; MG = p a r t s per m i l l i o n of magnesium i n s o i l sample; NA = p a r t s per m i l l i o n of sodium i n s o i l sample; PH = pH of s o i l sample; ELEV = e l e v a t i o n of sample p l o t above l o c a l c h a r t datum; %ELEV = s t a n d a r d i z e d sample p l o t e l e v a t i o n as d e s c r i b e d i n s e c t i o n 7 . 3 . 1  SAMPL  SAND S I L T  CLAY  A 0 A 1 A 2 A 3 A 4 A 5 A 6 A 7 A 8 A 9 A10 A1 1 A12 A13 A 14 A15 A 16 A 17 A 18 A19 A20 A21 A22 A23 A24 A25 A26 A27 A28 A29 A30 B 1 B 2 B 3 B 4 B 5 B 6  27 ,. 5 60 25 . 60 27 .. 5 57 . 5 27 .. 5 57 . 5  12 15 15 15  25 . 15 .  TXTCLASS  . 5 SiL . SiL . SiL . SiL  60 15 . SiL 52 .. 5 32 .. 5 S i C L  COND  N  K  CA  MG  NA  5. 4 10 . 4 8. 4 5 .0  .04 .05 .06 . 10  31 .6 . 30. 45 . 50.  105 155 137 .5 82 .5  1 10 280 245 160  290 700 550 365  9 . 4 4. 5  .06 . 1 1  67 ,. 5 37 .. 5  280 40  580 120  625 425 ' 560  7 . 5 60  32 .. 5 S i C L  5 . 6-  . 14  72 ..5  140  275'  5. 65 . 7 .. 5 65 .  30. SiCL 27 .. 5 S i L - S i C L  8 .8 7. 8  . 14 . 12  77 .. 5 45 ,  272 . 5 285  710 525  855 325  . 13  57 .. 5 • 372.5  735  625  825 37 1 608 405 31 1 599 540 578 915 464 59 1 485 165 125 1115 867 840 905 896 670  415 703 608 7 14 877 1001 1380 1356 1335 864 837 706 1 130 5 10 8400 5272 7 100 6372 3240 5450  12 . 5 62 .. 5 25 .  SiL  9 .. 1  7 . 5 67 .. 5 2 5 .  SiL  8.1 1 1, 4 12 . 11 . 10. 14 . 2 12 . 21 . 18 . 14 . 15 . 4 16 . 23 . .25 . 46 . 39 . 39 42 40. 35  •i  30.  47 .. 5 22 . 5 L  22 . 5 57 ., 5 2 0 .  SiL  25 . 20. 7.5 22 .. 5  55 . 20. 57 . 5 22 .. 5 70. 22 . 5 55 . 22 ..5  SiL SiL SiL SiL  32 .. 57 .. 62 .. 27 .. 15 .  50. 17 . 5 S i L - L 32 . 5 10. SL 27 .. 5 10. SL 52 . 5 2 0 . SiL 65 . 20. SiL  5 5 5 5  20. 7 .5  55  25 .  SiL  65  27 .. 5 S i L - S i C L  . 14 . 14 . 12 . 12 . 17 . 13 . 12 .08 . 1 1 . 12 . 1 1 .06 .02 .02 . 24 . 24 . 19 .21 . 13 . 14  57 .. 5 39 . 9 39 . 91 .. 7 35. . 9 51 .. 3 67 . 5 61 .. 3 82 .. 5 40. 5 0 .. 2 42 ..6 57 ..5 70. 250. 180.. 6 217 ..5 405 .. 5 148 . 250.  460 209 335 231 . 6 135 . 2 295 . 1 237 . 5 254 4 15 240 301 . 3 246 .8 72 .5 162 . 5 315" 249 . 2 22.5 224 . 5 4 16 187 . 5  PH ELEV  5 .1 4. 4 4. 7 5.3 6.6 6.0  %ELEV  1 .82 2 . 22 2 . 53 2 .71 2 .73 2 . 59 2 .92 2 .80 2 .88 2 ,84 2 .88 2 .. 92 3 .00 . 3 .00 . 3 .. 1 1 3 .. 10 3 .. 20 3 .. 15 3 .. 20 3 .09 . 3 .08 3 .. 10 3 .03 . 2 .. 99 2 .68 . 2 .81 . 2 .91 . 2 .08 2 .. 37  44 .9 54 . 7 62 .6 67 . 0 67 . 5 63 .9 72 . 1 69 . 1 67 . 9 70 . 1 71 . 1 72 .0 74 . 2 74 . 2 76 ..7 76 .. 6 79 ..0 77 ..9 79 .. 1 76 . 4 76 ..0 76.. 4 74 .. 9 73 .. 9 66 .. 2 69 ..4 7 1 .9 . 51 . 3 58 ..4  3 .. 3 .. 3. 3. 3 .. 3.  8 0 .. 4 75 .. 1 72 ..8 69 .. 5 70..6 68 .. 4  75 50 39 24 29 19  1 43  SAMPL B 7 B 8 B 9 B 10 B1 1 B12 C 1 C 2 C 3 C 4 C 5 C 6 C 7 C 8 D 1 D 2 D 3 D 4 D 5 D 6 D 7 D 8 E 1 E 2 E 3 E 4 E 5 & 6 E 7 E 8 E 9 F 1 F 2 F 3 F 4 F 5 G 1 G 2 G 3 G 4 G 5 G 6 G 7 G 8 G 9 G10 H 1 H 2 H 3 H 4 H 5 H 6 H 7 H 8 H 9 H10 H1 1 J 1 J 2 d 3 J 4 J 5 0 6 u 7 . J 8 J 9  SAND S I L T  CLAY  TXTCLASS  COND  30.  S iCL  33 .  . 15  265 .  17 . 5 57 . 5 25 .  SiL  32 . 5  .08  27 . 5 1 25 . 27 . 5 17 . 5  L SiL SiL SIL  30.  .05  26 . 30. 41 . 36 . 38 . 46 . 40. 3 . 5 2 .3  . 13 . 28 . 19 .21 . 18 . 14 . 24 . 10  3 . 3 . 3 . 4. 5. 35 . 31 . 28 . 28 . 48 . 52 . 46 . 33 . 47 . 29 . 28 . 32 . 41 . 28 . 38 . 46 . 58 . 48 . 58 . 56 . 49. 40. 38 . 25 . 39 .  36 . 9 62 . 7 . 14 600. 56 . 3 . 26 76: 6 119. 5 . 26 83 . 3 154 . 2 . 23 92 . 5 235 . . 14 152 . 3 132 . 9 . 20 77 . 5 102 . 5 . 16 101 . 3 138 . 8 . 13 252 . 5 167 . 5 .21 320. . 32 6 2 5 . 370. 145. . 35 . 30 3 9 0 .. 4 233 . 220. 205 . .07 252 . 5 220. . 19 142 . 5 132 . 5 . 17 98 ..4 84 ., 8 .07 100. . 1 1 143 . 3 . 20 218 .. 7 • 109 . 3 84 .. 8 45 . 1 .09 590. 507 . 5 . 28 315 . . 23. 3 1 5 , 440. 332 .. 5 . 38 4 12.. 5 375 . . 26 415. 290. . 14 177 . 5 175 . 23 600. 350. 382 ,. 5 192 , 5 150. . 1 1 142 ,. 5 86 . 3 127 ,. 5 .09 282 ,. 5 292 . 5 .43  0.  70.  50. 22 . 5 70. 17 . 5 52 . 5 2 0 . 62 . 5 2 0 .  12 . 5 57 . 5 3 0 .  SiCL  5. 65 . 30. S iCL 17 . 5 65 . 17 . 5 S i L 2 . 5 72 . 5 25 . SiL 5.  67 . 5 27 . 5 S i L - S i C L  5.  65 .  30.  SiCL  10.  65 .  25 .  SiL  50.  37 . 5  12 . 5 L  30.  50.  20.  SiL-L  42 . 5 42 . 5  15 .  L  75 . 15 . 60. 27 . 5 57 . 5 3 0 .  10. SL 12 ..5 SL 15 . SL  75.  20.  5 ,  LS  80. 17 . 5 57 . 5 32 . 5 60. 30. 55 . 35 .  2 . 5 LS 10. SL 10. SL 10, SL  72 . 5 2 0 . 72 ..5 17 .5 . 60. 25 . 85 . 10.  7.5 10. 15 . 5  SL • SL SL LS  62 .. 5 22 . 5 15 SL 55 . 22 . 5 22 . 5 SCL 67 .. 5 10. 22 . 5 SCL 70.  15 ,  15  SL  60. 70.  22 . 5 17 .5 SL 17 .5 . 12 . 5 SL  82 . 5 7 . 5 10 LS 87 . 5 5 7.5 S 50 35 15 L 45 40 15 L 5  67 . 5 27 . 5 S i L - S i C L  2 . 5 72 . 5 25 10  SiL  72 . 5 17 . 5 S i L  1 3 8 7  5  N  K  CA  MG  NA  190.  690  5250  185 .  272 . 5  775  4650  157 . 5  227 . 5  660  4250  152 . 5  2 12. 5  700  2080  221 . 4 255 . 166 . 2 210. 8 285 . 31 . 38 . 7  278 . 8 322 . 5 253 . 3 450 . 4 577 . 5 25 . 54 . 1  951 965 823 1236 1625 58 129  . 72 83 . 62 . 57 . 58 . 52 . 1. 3 '1 .9 2 . 2 2 . 2 1. 8 3 . 2 2 . 4 2.8 1 .  .84 .61 . 18 . 26 . 13 .31 .06 • . 10 .01 .08 . 13 .08 .06 .03 .05  865  598 .  622 .5 799 490 182 . 5 6 8.5 6.2 6 .6 8. 1 •10 8.7 14 . 1 5.1  267 364 250 125 60 70 71 60 72 142 191 192 75  .5  .3 .5 .3 .5  PH ELEV %ELEV  6421 6000 6 . 3 5082 6143 7500 5 . 6 400 156  140 280 1 13 390 196 319 5. . 2 258 408 640 300 4 . 2 486 3403 395 3150 4 . 2 593 3900 645 4900 1285 9950 575 4350 5 . 5 974 8413 830 3890 3 .9 . 395 2600 495 3950 5 .•0 299 23 14 367 2620 820 496 1 4 14 2705 825 5500 3 . 9 1030 7500 4 . 2 1250 9650 4 ., 7 1523 10800 1035 8 5 0 0 4 .5 , 585 3550 4.,, 4 1350 10800 4 . 1 735 6995. 5 . 6 600 4200 405 2610 1060 8500 5 . 7 6 .0 5.6 2697  29906  5.3  5 1 140 5450 5 1 19310425 1040 9550 5 15 4600 5 60 17 70 65 5 86 20. 45 1 105 72 54 130 1 106 131 4 1 65 55 6 51 38  .3 .8 .O .5 . 4 .3 .8  3 . 20 68 . 7 3 . 1 1 66 . 8 3. 18 68 . 2 2 . 59 55 . 6 2 . 59 55 . 6 2 . 59 55 . 5 3 . 92 84 . 1 3 . 64 78 . 0 3 . 66 78 . 6 3 . 61 77 . 4 3 . 54 75 . 9 3 . 53 75 . 6 3 . 37 72 . 3 3 . 27 70. 2 3. 40 83 . 9 3 . 33 82 . 1 3 . 24 8 0 .. 1 3 , 48 85 ..9 3 . 19 78 . 8 3 .07 . 75 ,.9 3 . 06 75 ,. 5 2 .95 . 72 ,. 8  3 . 53 80 .0 3• , 22 . 72 .9 3 . 26 73 .6 3 . 13 70 . 9 3 . 20 72 . 5 3 . 13 70 . 8 2 .87 64 . 9 3 .05 69 . 1 3,. 54 80 . 1 3 .50 79 . 2 4 . 24 95 . 9 4 . 26 96 .4 4 . 28 96 . 8 4 . 22 95 .5 4 . 14 93 .7 4 . 15 93 .9 4 . 10 92 . 7 4 .07 92 . 2 4 .01 90 .7 3 . 93 88 . 9 3 . 76 84 . 8 3 . 45 77 . 2 3 .50 78 . 3 3 . 43 76 .6 3 . 42 76 .6 3 .04 68 . 0 2 .91 65 . 1 2 . 39 53 . 5 2 . 48  55 .5  

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