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Autecology of Blidingia minima var. Subsalsa (Chlorophyceae) in the Squamish River estuary, British Columbia Prange, Robert K. 1976

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AUTECOLOGY OF BLIDINGIA MINIMA VAR. SUBSALSA (CHLOROPHYCEAE) IN THE SQUAMISH RIVER ESTUARY, BRITISH COLUMBIA by ROBERT K. PRANGE B.Sc, Acadia University, Wolfville, Nova Scotia, 1973 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of BOTANY We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA MAY 1976 In p r e s e n t i n g t h i s t h e s i s in p a r t i a l f u l f i l m e n t o f the r e q u i r e m e n t s f o r an advanced degree at the U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r ee t h a t t he L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r ag ree t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y pu rpo se s may be g r a n t e d by the Head o f my Department o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Department o f Botany  The U n i v e r s i t y o f B r i t i s h Co l umb i a 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date A p r i l 29, 1976 / 1 i i ABSTRACT The autecology of the e s t u a r i n e a l g a , B l i d i n g i a minima v a r . subsalsa (Kjellman) Scagel (Chlorophyceae) was considered w i t h regard to i t s growth, re p r o d u c t i o n and d i s t r i b u t i o n . Studies were conducted on the Squamish R i v e r e s t u a r y , B r i t i s h Columbia, from May 1974 to J u l y 1975 and i n the l a b o r a t o r y from January to August 1975. The major environmental f a c t o r s considered were l i g h t , temperature, s a l i n i t y , n u t r i e n t s and d e s i c c a t i o n . The a l g a occurred on the Squamish R i v e r d e l t a i n the upper i n t e r t i d a l zone. Biomass, as estimated by percent cover, i n c r e a s e d i n the p e r i o d March to e a r l y May, then remained s t a b l e or decreased during s p r i n g runoff i n May and June, f i n a l l y i n c r e a s i n g to a maximum i n August. Increases i n percent cover were a s s o c i a t e d w i t h b r a c k i s h s a l i n i t y , h i g h l i g h t i n t e n s i t y , h i g h a i r temperature, c o n s i d e r a b l e 3 d e s i c c a t i o n , an absence of a l g a l competitors and p o s s i b l y favourable i o n r a t i o s . The three f a c t o r s i n v e s t i g a t e d i n the l a b o r a t o r y (temperature, s a l i n i t y and n u t r i e n t s ) i n t e r a c t e d i n t h e i r e f f e c t on net photosynthesis. Reproduction i n the l a b o r a t o r y occurred by r e l e a s e of q u a d r i -f l a g e l l a t e and isomorphic b i f l a g e l l a t e swarmers. Some b i f l a g e l l a t e swarmers fused and germinated, producing isomorphic p l a n t s . The p e r i o d of swarmer r e l e a s e was from January to e a r l y May. The a l g a was p e r e n n i a l but during the w i n t e r only i t s p r o s t r a t e b a s a l d i s c was present. Maximum v e r t i c a l d i s t r i b u t i o n was from 1.5 to 4.0 m above chart datum (lowest low wate r ) . Maximum percent cover occurred at aa. 3.25 m. The upper l i m i t appeared to be a s s o c i a t e d w i t h unfavourable osmotic c o n d i t i o n s , e.g. r a i n or d e s i c c a t i o n , and the lower l i m i t w i t h low l i g h t i n t e n s i t i e s . H o r i z o n t a l d i s t r i b u t i o n was l i m i t e d by absence of s a l t water on the freshwater s i d e and competition from Fucus distichus subsp. edentatus (De l a P y l a i e ) P o w e l l on the marine s i d e of the estuary. Blidingia minima v a r . subsalsa1s geographical and h a b i t a t d i s t r i b u t i o n was a l s o examined by reference to l i t e r a t u r e r e p o r t s and herbarium c o l l e c t i o n s . The species i s cosmopolitan, o c c u r r i n g i n every ocean except the Ind i a n and A n t a r c t i c w i t h most r e p o r t s from p o l a r and temperate r e g i o n s . The v a r i e t y occurs i n b r a c k i s h , marine and freshwater h a b i t a t s . TABLE OF CONTENTS ABSTRACT LIST OF TEXT TABLES LIST OF FIGURES LIST OF APPENDICES ACKNOWLEDGEMENTS INTRODUCTION FIELD METHODS Selection of Sampling Stations Physical and Chemical Sampling and Analyses Biological Methods LABORATORY MATERIALS AND METHODS Media General Culture Conditions Experimental Methods 1. Life History Study 2. Environment Factor Study FIELD RESULTS Description of Study Area 1. Introduction 2. Atmospheric Environment 3. Aquatic Environment Biology of Blidingia minima var. subsalsa and Associated Organisms 1. Annual Cycle of Percent Cover, Swarmer Release and Desiccation of B. minima var subsalsa 2. Production Measurements of B. minima var subsalsa 3. Seasonal Occurence of Other Dominant Organisms Benthic Algae Animals LABORATORY RESULTS Life History Study Environmental Factor Study V Page DISCUSSION 43 Environmental F a c t o r s , Growth, Reproduction and D i s t r i b u t i o n 43 1. L i g h t 45 2. Temperature 48 3. Osmotic E f f e c t s 52 S a l i n i t y 53 D e s i c c a t i o n 56 4. N u t r i e n t s 59 Systematic C o n s i d e r a t i o n of L i f e H i s t o r y Study 65 CONCLUSIONS 67 LITERATURE CITED 70 APPENDICES 79 LIST OF TEXT TABLES Table Page 1. Desi c c a t i o n at S t a t i o n I, II and I I I 3 2 v i i LIST OF FIGURES Fi g u r e Page 1. Sampling l o c a t i o n s i n Squamish R i v e r e s t u a r y , B r i t i s h Columbia. 8 2. Shore S t a t i o n s I on the west s i d e of the Squamish R i v e r t r a i n i n g dyke. 9 3. Shore S t a t i o n I I on the west shore of the C e n t r a l Channel mouth. 9 4. Shore S t a t i o n I I I at the mouth of Stoney Creek on the east shore of Howe Sound. 9 5. Frequency of t i d a l f l o o d i n g (%) above c h a r t datum (lowest low water) during the w i n t e r and summer. 21 6. A t t e n u a t i o n of l i g h t i n the top 3 m of water at S t a t i o n s I , I I and I I I f o r the p e r i o d J u l y 1974 to J u l y 1975. 23 7. Water temperature (°C) and s a l i n i t y (°/ ) at S t a t i o n s I , I I and I I I f o r the p e r i o d February 1974 to J u l y 1975. 24 8. N u t r i e n t concentrations at S t a t i o n s I , I I and I I I f o r the p e r i o d August 1974 to J u l y 1975. 26 9. Percent oxygen s a t u r a t i o n of s u r f a c e water at S t a t i o n s I , I I and I I I f o r the p e r i o d February 1974 to J u l y 1975. 28 10. Percent cover and v e r t i c a l d i s t r i b u t i o n of B. minima v a r . subsalsa at S t a t i o n s I , I I and I I I f o r the p e r i o d J u l y 1974 to J u l y 1975. 30 11. Representative net photosynthesis p r o f i l e s (mg C«gm ash-free dry wt ^ d a y 1 ) at B o t t l e Incubation S t a t i o n s I , I I and I I I f o r the p e r i o d May 1974 to J u l y 1975. 34 12. B i f l a g e l l a t e swarmer. 38 13. Q u a d r i f l a g e l l a t e swarmer. 38 14. The m u l t i c e l l u l a r stage f o l l o w i n g germination. 38 15. P r o s t r a t e d i s c stage. 38 16. P r o s t r a t e d i s c w i t h two u p r i g h t t h a l l i . 38 F i g u r e 17. I n i t i a l form of c e l l c e l l s of the p l a n t s . 18. F i n a l stage observed 19. Two way i n t e r a c t i o n s masses produced by the b a s a l i n c e l l masses. i n environment f a c t o r study. LIST OF APPENDICES Appendix I References used to determine occurences of B. minima and B. minima v a r . subsalsa I I Procedures used i n P h y s i c a l and Chemical Analyses I I I C u l t u r e Media IV Water Q u a l i t y R e s u l t s at S t a t i o n I , I I and I I I V Net Ph o t o s y n t h e t i c and R e s p i r a t i o n Rates (mg C* gm ash-free dry wt ^ d a y 1) at S t a t i o n I , I I and I I I VI A s s o c i a t e d a l g a l species at S t a t i o n s I , I I and I I I f o r the p e r i o d J u l y 1974 to J u l y 1975 VI I R e s u l t s of Environment Fact o r Study V I I I R e s u l t s of 3-Way A n a l y s i s of Variance (UBC ANOVAR) IX R e s u l t s of 2-Way A n a l y s i s of Variance (UBC MFAV) X ACKNOWLEDGEMENTS I wish to express my g r a t i t u d e to Dr. Janet R. S t e i n f o r her advice , guidance and f i n a n c i a l support, provided by funds from NRC grant A1035 and to Dr. C o l i n Levings who provided c o n s t r u c t i v e comments, unpublished data, f i e l d equipment and f a c i l i t i e s and s t a t i s t i c a l a dvice. I am a l s o g r a t e f u l to Dr. R.F. Scage l , Dr. P.G. H a r r i s o n and Dr. R. Foreman f o r t h e i r a s s i s t a n c e d u r i n g the study and t h e i r h e l p f u l suggestions concerning the p r e p a r a t i o n of t h i s t h e s i s . The a s s i s t a n c e given by Mr. M. Pomeroy i n the f i e l d work as w e l l as h i s h e l p f u l comments and suggestions i s g r e a t l y a p p r e c i a t e d . I a l s o thank Mr. A. Shearon f o r p r o v i d i n g a s s i s t a n c e i n the f i e l d and l a b o r a t o r y . 1 INTRODUCTION E s t u a r i e s and d e l t a s form an important t r a n s i t i o n zone between marine and freshwater h a b i t a t s (Kinne 1967, Odum 1971, Remane and S c h l i e p e r 1971). This t r a n s i t i o n zone provides a h a b i t a t necessary f o r the s u r v i v a l of many p l a n t and animal species such as salmon. Because of the mountainous nature of the B r i t i s h Columbia coast, the r i v e r d e l t a s along the coast are considered prime s i t e s f o r i n d u s t r i a l and port development. Such development u s u a l l y i n v o l v e s c o n s i d e r a b l e dyking and land f i l l i n g which a l t e r or destroy the e s t u a r i n e h a b i t a t to such an extent that the e s t u a r i n e species are destroyed or replaced. In t u r n t h i s reduces one of the renewable n a t u r a l resources and an important food i n d u s t r y , the salmon f i s h e r y . Squamish i s one of four d e l t a areas on the B.C. coast w i t h deep water, r a i l and f l a t l a n d to a l l o w b u i l d i n g of back-up f a c i l i t i e s f o r a deep sea p o r t (Waldichuk 1972). The other three are the F r a s e r d e l t a , K i t i m a t d e l t a and. P r i n c e Rupert. The Squamish R i v e r d e l t a and estuary i s being s t u d i e d i n t e n s i v e l y because i n 1972 t h i s area was d e s t i n e d to become a major i n d u s t r i a l p o r t , w i t h deep-sea s h i p p i n g f a c i l i t i e s , c o n t a i n e r and b u l k storage space and a major r a i l terminus f o r the B r i t i s h Columbia Railway. In 1972 government environmental impact s t u d i e s were i n i t i a t e d and that h a l t e d the c o n s t r u c -t i o n of a s h i p p i n g t e r m i n a l on the Squamish d e l t a . These environmental s t u d i e s have been continued i n c e r t a i n s e c t o r s to o b t a i n a sound s c i e n t i f i c background on e s t u a r i n e ecosystems (Hoos and Void 1975). P r e l i m i n a r y s t u d i e s show the most common and p r o d u c t i v e benthic a l g a on the d e l t a i s B l i d i n g i a minima v a r . subsalsa (Kjellman) Scagel 2 (Poraeroy and Stockner 1976, as Enteromorpha minima N S g e l i ex K U t z i n g ) . I t i s a l s o present i n other B r i t i s h Columbia e s t u a r i e s e.g. Toba (Toba I n l e t ) , Homathko (Bute I n l e t ) , Ocean F a l l s (Cousins I n l e t ) , K l i n a k l i n i (Knight I n l e t ) , K i t i m a t ( K i t i m a t Arm, Douglas Channel) (Levings, Pomeroy and Prange 1975; UBC Herbarium; personal c o l l e c t i o n ) and F r a s e r (Fraser R i v e r ) (Northcote, Ennis and Anderson 1975). B l i d i n g i a K y l i n (1947) i s based on Enteromorpha minima N S g e l i ex KUtzing (1849 p. 482). The holotype i s a specimen c o l l e c t e d by N a g e l i at Helgoland, Herb. Lugd. 938.69..168 ( B l i d i n g 1963). The c h a r a c t e r i s t i c s d i s t i n g u i s h i n g B l i d i n g i a from Enteromorpha a r e : BZidingia forms a hollow germination tube whereas Enteromorpha does not; B l i d i n g i a develops a p r o s t r a t e d i s c o i d h o l d f a s t l a c k i n g the non-septate r h i z o i d a l c e l l s that are t y p i c a l of Enteromorpha; and B l i d i n g i a has s m a l l e r v e g e t a t i v e c e l l s (3.5 - 8 ym diameter i n s u r f a c e view) compared to Enteromorpha (> 8 ym) ( B l i d i n g 1963). Sexual r e p r o d u c t i o n , commonly observed i n species of Enteromorpha, i s considered non-existent i n Blidingia^ s i n c e only q u a d r i f l a g e l l a t e zoospores without eyespots have been rep o r t e d . The s y s t e m a t i c p o s i t i o n of B l i d i n g i a i s not c l e a r l y d e f i n e d . I t i s a member of the Chlorophyceae (Chlorophyta) i n v a r i o u s a l g a l c l a s s i f i c a t i o n s (see reviews by Papenfuss 1955, Chapman 1964, Round 1963, 1971, Stewart and Mattox 1975). There i s disagreement con-cer n i n g the number of orders and f a m i l i e s , t h e r e f o r e a f f e c t i n g the p o s i t i o n of B l i d i n g i a . The genus i s u s u a l l y considered a member of the U l v a l e s ( B l i d i n g 1963, 1968, Gayral 1967, Stewart and Mattox 1975, Vinogradova 1969) even though the U l v a l e s can be considered w i t h the 3 Ulotrichales and has been reduced to a family in the Ulotrichales (Papenfuss 1960). When the family Monostromaceae is not recognized, Blidingia is lumped with Monostroma into the Ulvaceae (Scagel 1966). If the Ulvaceae and Monostromaceae are recognized, the genus may appear in the Ulvaceae (Chapman 1964, Chapman and Chapman 1973, as E. nana {Sommerfeldt} Sjostedt, Kylin 1947, Papenfuss 1960) or more commonly in the Monostromaceae (Scagel 1957, Bliding 1963, 1968, Gayral 1967, Vinogradova 1969). Bliding (1968) wisely cautions that any progress in the systematics of Blidingia will proceed only after more informa-tion about the anatomy and l i f e history of taxa belonging to the Ulvaceae and Monostromaceae are available. Preliminarily he places Blidingia as a "divergent" genus of Monostromaceae. Originally the genus contained one species and one variety (Kylin 1947). Scagel (1957) added a second variety B. minima var. subsalsa (Kjellman) Scagel. Prior to 1957 i t had been known as Enteromorpha miorocoooa f. subsalsa Kjellman (1883) and E. minima var. subsalsa (Kjellman) Doty (1947). Dangeard (1958) added a new species: B. marginata ( J . Agardh) P. Dangeard which had been originally named Enteromorpha marginata J. Agardh (1842) and also been called E. miorooooca3 E. canaliculata, and E. nana var. marginata (Bliding 1963). The most thorough examination of the genus is the critical survey by Bliding (1963, 1968). After reviewing structure in preserved and living specimens as well as reproduction and nomenclature, he arranged the genus into three species, two subspecies, and two varieties: 4 B. minima (N'ageli ex Kutzing) v a r . minima B. minima var. ramifera B l i d i n g B. marginata ( J . Agardh) P. Dangeard subsp. marginata B. marginata subsp. subsalsa (Kjellman) B l i d i n g B. chadefaudii ( J . Feldman) B l i d i n g I n the process, B l i d i n g added a t h i r d species B. chadefaudii which had p r e v i o u s l y been placed i n i t s own genus Feldmanodora Chadefaud (1957). Before 1957 i t was Enteromorpha micrococca and E. chadefaudii ( B l i d i n g 1963). He a l s o s p l i t B. minima v a r . subsalsa i n t o B. minima v a r . ramifera and B. marginata subsp. subsalsa. However, Scagel (1966) considers B. minima v a r . subsalsa synonymous w i t h B. minima var. ramifera. Vinogradova (1974) changes B. minima var. subsalsa to B. minima f. subsalsa and considers i t synonymous w i t h B. minima var. ramifera and B. marginata subsp. subsalsa. The drawing and d e s c r i p t i o n of E. nana v a r . minima ecad r i v u l a r i s (Chapman 1956) i n d i c a t e that i t may a l s o be a synonym f o r B. minima v a r . subsalsa. In a d d i t i o n to these taxa N o r r i s (1971) added a t h i r d v a r i e t y : B. minima v a r . vexata (S. & G.) J . N o r r i s that contains a f u n g a l symbiont. Although most p h y c o l o g i s t s may disagree on B l i d i n g 1 s arrangement of s u b s p e c i f i c taxa they have adopted the s p e c i f i c nomenclature. Only Chapman (1964) and Chapman and Chapman (1973) r a i s e s e r i o u s o b j e c t i o n s to B l i d i t i g ' s nomenclature. They contend the genus i s not s u f f i c i e n t l y d i s t i n c t from Enteromorpha and r e f e r the species B. minima to Enteromorpha nana. The combination i s E. nana because nana Sommerfeldt, i f considered as the same taxon, i s the o l d e r name ( B l i d i n g 1963). Since they do not r e f e r to the other two species I presume they consider B. marginata i s a v a r i e t y of E. nana (Chapman 1956) and B. chadefaudii is returned to the genus Feldmannodora (Chapman 1964). Using the taxonomy of Scagel (1957, 1966) and Norris (1971) which refer specifically to the local taxa, the alga at Squamish is B. minima var. subsalsa. The publications l i s t e d in Appendix I were used to determine the geographic distribution of B. minima (most workers do not consider varieties) as well as additional occurences and habitats of B. minima var. subsalsa including the synonyms discussed above. B. minima occurs in every ocean except the Indian and Antarctic and on every continent except the Antarctic. It occurs above the Tropic of Cancer and below the Tropic of Capricorn with the exception of the west coast of Africa where i t occurs as far south as Senegal, Lat. 15°N (Lawson and Price 1969). The most northern extreme is Spitzbergen, Lat. 79° 59'N (Kjellman 1883). The possible southern extreme is Tasmania (Womersley 1956) or South Island, New Zealand (Chapman 1956) both at ca. Lat. 47°S. In general, the habitat of B. minima var. subsalsa differs from the more saline habitats of the other subspecific taxa (see Appendix I ) . It i s found growing attached or free floating in some-what sheltered bays, lagoons, sloughs, on mudflats, and river mouths' in the high i n t e r t i d a l and occasionally in inland rivers and lakes (Norris 1971). B. minima var. subsalsa is reported in water temperatures from -22°C (Pt. Barrow, Biebl 1969) to at least +35°C (summer water temperature, Gulf of Mexico, Kapraun 1974) and s a l i n i t i e s ranging from freshwater (Yugoslavia, Bliding 1963; Faeroes Island, B<J>rgesen 1901; Japan, Hirose 1972; and California, Norris 1971) to at least normal sea water concentration (aa. 36°'/' Hope Island specimen, UBC No. 48842). 6 In order to i n v e s t i g a t e the ecology, i . e . seasonal p e r i o d i c i t y of growth, r e p r o d u c t i o n and d i s t r i b u t i o n , of a t y p i c a l e s t u a r i n e a l g a , f i e l d and l a b o r a t o r y s t u d i e s of B. minima var. subsalsa were undertaken on the Squamish R i v e r d e l t a from May 1974 to J u l y 1975. In the f i e l d study the environmental f a c t o r s measured or noted were: l i g h t , temperature, s a l i n i t y , d u r a t i o n of submergence and emergence ( d e s i c c a t i o n ) , n u t r i e n t s , -0 content of water, pH, s u b s t r a t e , turbulence, competition and p e r i o d i c i t y of these f a c t o r s . S tudies on the l i f e h i s t o r y and some environmental f a c t o r s (temperature, s a l i n i t y and n u t r i e n t s ) were conducted i n the l a b o r a t o r y from January to August 1975. 7 FIELD METHODS S e l e c t i o n of Sampling S t a t i o n s P r e l i m i n a r y i n v e s t i g a t i o n showed that B l i d i n g i a minima v a r . subsalsa was present i n the mouth of the Squamish R i v e r to the upper l i m i t of s a l t water i n t r u s i o n (ca. 49° 42'N, F i g . 1 ) , along the e n t i r e f r o n t of the r i v e r d e l t a , i n the C e n t r a l Channel, the Mamquam Channel and down the east shore of Howe Sound to Watts P o i n t ( F i g . 1 ) . B i o l o g i c a l sampling was done at three shore s t a t i o n s s e l e c t e d on the b a s i s of a c c e s s i b i l i t y and presence of the alg a ( F i g . 1 ) . Shore S t a t i o n I was on l a r g e rocks i n the mouth of the Squamish R i v e r on the west s i d e of the t r a i n i n g dyke t i p ( F i g . 2). Shore S t a t i o n I I was a group of p i l i n g s on the west shore of the C e n t r a l Channel mouth ( F i g . 3). Shore S t a t i o n I I I was a group of p i l i n g s l o c a t e d at the mouth of Stoney Creek on the east shore of Howe Sound ( F i g . 4 ) . The Mamquam Channel had been the intended l o c a t i o n but could not be used because of i n d u s t r i a l a c t i v i t y . B i o l o g i c a l f i e l d sampling i n c l u d e d measurement of p h o t o s y n t h e t i c and r e s p i r a t i o n r a t e s of B. minima v a r . subsalsa. The l o c a t i o n of the b o t t l e i n c u b a t i o n and shore s t a t i o n s d i f f e r ( F i g . 1 ) . The b o t t l e i n c u b a t i o n s t a t i o n s were s e l e c t e d on the b a s i s of p r o x i m i t y to shore s t a t i o n s , s u f f i c i e n t water depth (two or more metres), and a v a i l a b i l i t y of a permanent buoy or p i l i n g . B o t t l e s used i n p h o t o s y n t h e t i c and r e s p i r a t i o n r a t e experiments were attached to the p i l i n g ( S t a t i o n I and I I I ) or buoy ( S t a t i o n I I ) . P h y s i c a l and chemical water sampling s t a t i o n s v a r i e d , depending on the f a c t o r s measured. Water temperature, water s a l i n i t y and oxygen s a t u r a t i o n data were obtained from the sampling of Levings, 8 Figure 1. Sampling locations in Squamish River estuary, British Columbia. & indicates marshland To face page 8 F i g u r e 2. Shore S t a t i o n I on the west s i d e of the Squamish R i v e r t r a i n i n g dyke. F i g u r e 3. Shore S t a t i o n I I on the west shore of the C e n t r a l Channel mouth. Fi g u r e 4. Shore S t a t i o n I I I at the mouth of Stoney Creek on the east shore of Howe Sound. To face pace 9 McDaniel, C h r i s t i e , Pomeroy and Prange (1976). For each b o t t l e incubation s t a t i o n the data from the sample s t a t i o n c l o s e s t to i t was used. For St a t i o n I the nearest s t a t i o n was on the west bank of the Squamish River; f o r S t a t i o n II the nearest one was about 10 metres south of Shore S t a t i o n I I ; and f o r S t a t i o n I I I the nearest one was on the east shore of the Mamquam Channel, opposite F.M.C. Chemicals. In a d d i t i o n , one l i t r e water samples were obtained from the water surface at each b o t t l e incubation s t a t i o n . They were frozen the same day and l a t e r analyzed i n the laboratory f o r pH, nitrogen and phos-phorus concentration. Light attenuation i n the water was measured at the b o t t l e incubation s t a t i o n s . Only preliminary data on water movement were c o l l e c t e d from v i s u a l observations, p l a s t e r of p a r i s clod cards (Doty 1971) and current meter measurements. P h y s i c a l and Chemical Sampling and Analyses Measurements of selected p h y s i c a l and chemical f a c t o r s were made at monthly or bimonthly i n t e r v a l s . Water temperature, oxygen concentration and s a l i n i t y were measured monthly from February 1974 to J u l y 1975 at a l l three s t a t i o n s (Levings et al. 1976). One l i t r e surface water samples i n polyethylene b o t t l e s were c o l l e c t e d from August 1974 to Jul y 1975. Water l i g h t attenuation measurements were made on an i r r e g u l a r basis between June 1974 and July 1975. A table of methods used i n measuring the p h y s i c a l and chemical f a c t o r s i n t h i s study i s included i n Appendix I I . Water temperature and s a l i n i t y were measured in situ simultan-eously with a portable salinity-conductivity-temperature meter. Measurements at each s t a t i o n were taken at high t i d e at depths of 0, 1 and 2 metres. For c a l i b r a t i o n purposes the water s u r f a c e temperature was measured w i t h a mercury thermometer and s a l i n i t y samples, taken from a Van Dorn b o t t l e , were analyzed i n the l a b o r a t o r y on a c a l i b r a t e d salinometer. In a d d i t i o n , water from the Van Dorn b o t t l e was t r a n s f e r r e d to 300 ml B.O.D. b o t t l e s , f i x e d immediately and analyzed the same day f o r oxygen content. With a nomograph the percent oxygen s a t u r a t i o n of the water was determined. From the f r o z e n water samples the pH, N H 3 , NO3 , N 0 2 and P values were obtained i n the l a b o r a t o r y w i t h i n 72 hours a f t e r thawing. A d d i t i o n a l water q u a l i t y and h y d r o l o g i c a l data were obtained from Hoos and Void (1975). C l i m a t o l o g i c a l data f o r the Squamish estuary were obtained from the Atmospheric Environment S e r v i c e ' s c l i m a t o l o g y s e c t i o n i n Hoos and Void (1975). B i o l o g i c a l Methods I n i t i a l l y the e l e v a t i o n s above zero t i d e l e v e l (lowest low water) were marked w i t h f l u o r e s c e n t p a i n t i n 0.3 m i n t e r v a l s a t each shore s t a t i o n . The e l e v a t i o n s were determined u s i n g a carpenter's l e v e l w i t h a s t r i n g running h o r i z o n t a l l y through i t . With the s t r i n g h e l d h o r i z o n t a l l y over the water w i t h the l e v e l , the p o i n t at which the h o r i z o n t a l s t r i n g s t r u c k the shore was marked. At the same time the v e r t i c a l d i s t a n c e from the water s u r f a c e to the l e v e l was measured. Using a t i d e t a b l e or the t i d e gauge at the end of the Squamish R i v e r t r a i n i n g dyke the e l e v a t i o n of water s u r f a c e at the time of measurement and, t h e r e f o r e , the e l e v a t i o n of the marker on the shoreline could be determined. During each sampling of the shore stations collections were made of each algal species. Representative algal material was pressed on herbarium sheets for future reference. Some of the B. minima var. subsalsa sample was used for l i f e history study. A duplicate of each herbarium sheet was deposited i n the University of British Columbia Phycology Herbarium (UBC» Nos. 54330-54346). Algal nomenclature follows that of Scagel (1966) for Chlorophyceae, Widdowson (1972) for Phaeophyceae, Widdowson (1974) for Rhodophyceae and Prescott (1970) for Cyanophyceae. During low tide B.-minima var. subsalsa distribution and percent cover was determined at each elevation level using a 0.01 m2 quadrat (Station II and III) or a 0.06 m2 quadrat (Station I ) . The larger 0.06 m2 quadrat was used at Station I because the boulder substrate created patchy algal cover. Percent cover was determined by a subjective estimate with the quadrat in s i t u and from Kodachrome slides taken simultaneously. The vertical range of other benthic algae and any grazers at each station was recorded. At approximately bimonthly intervals, samples of the alga from one or more elevations at each station were placed i n air-tight plastic bags i n an ice chest and frozen the same day. They were later analyzed i n the laboratory for the degree of desiccation. Samples could not always be taken in the winter and spring at some elevations as not enough alga was available. The degree of desiccation was determined by weighing the collected sample ( f i e l d weight), soaking i t in d i s t i l l e d water for 24 hours and then reweighing after the excess water had been gently squeezed out (soaked weight). The f i e l d weight 13 as a percent of the soaked weight indi c a t e d the degree of des i c c a t i o n . Estimates of r e s p i r a t i o n and net photosynthetic rates of the alga were made by modifying the oxygen l i g h t - d a r k b o t t l e method of S t r i c k l a n d and Parsons (1972). T h a l l i r e l a t i v e l y f r ee of con-taminants were introduced i n t o each b o t t l e . The biomass was i n the 10-20 mg (ash-free dry weight) range. The incubation water was obtained from the surface at the b o t t l e incubation s t a t i o n s . Instead of an i n i t i a l b o t t l e , dark and l i g h t c o n t r o l b o t t l e s were used to account f o r the metabolic a c t i v i t y of the indigenous phyto- and zooplankton. Bottle p a i r s were suspended on a rope from a buoy at the b o t t l e incubation s t a t i o n s , at depths of 0.25 m, 0.5 m (c o n t r o l b o t t l e s ) , 1.0 m and 2.0 m. The incubation period was never more than four hours, usually between 1000 and 1400 hours. Biomass (ash-free dry weight) i n each b o t t l e was determined w i t h i n 24 hours. A f t e r the oxygen determination the t h a l l i were f i l t e r e d i n a M i l l i p o r e f i l t e r apparatus onto pre-ashed Whatman GF/C f i l t e r paper, oven-dried at 100°C f o r 24 hours and weighed. Then the sample was ashed at 500°C f o r four hours and weighed. The ash-free dry weight i s the d i f f e r e n c e between the oven-dried weight and the ashed weight. In order to compare incubations at d i f f e r e n t s t a t i o n s and on d i f f e r e n t days, the r e s p i r a t i o n rates and net photosynthetic rates were expressed as a d a i l y r a te. In order to do t h i s , i ncident s o l a r r a d i a t i o n was recorded with a B e l f o r t pyranometer. Photosynthetic and r e s p i r a t i o n rate values were expressed as mg Ogm ash-free dry wt *-day 1 using the equation (Pomeroy 1974): mg 02'1 ^O.3*0.278 mg C*gm ash-free dry wt **day 1 = ash-free dry wt*light where: mg 1 = oxygen concentration change (Strickland and Parsons 1972) 0.3 = correction factor to convert mg 02*£ 1 to mg 02*0.3 £ 1 (incubation volume) 0.278 = conversion factor for mg 0 2 to mg C assuming a PQ of 1.2 and RQ of 1.0 ash-free dry wt = weight of algal biomass used in incubation bottle ligh t incident solar radiation*incubation period 1 incident solar radiation*day 1 LABORATORY MATERIALS AND METHODS Media Two media were used in the laboratory: (1) filtered enriched Squamish seawater and (2) defined seawater. The method of prepar-ation for the two media is in Appendix III. In the l i f e history study both media were used. The salinity was maintained at 10-12°/ J J oo In the environmental factor study only defined seawater medium was used. General Culture Conditions Cultures in the l i f e history study and environmental factor study were maintained in two incubation chambers: (1) model T181, Controlled Environments Ltd., Winnipeg, Manitoba and (2) model RT-18B-5E Sherer Company Ltd., Marshall, Michigan. Light was supplied by General Electric 20 watt cool-white fluorescent lamps set in overhead pairs. The light intensity, as measured by a YSI model 65 radiometer, was 1.43x10 2 l y •min 520 ft-c). The light cycle was a 16:8 L:D photoperiod in both stud ies. The spectrum of light from 400—720 mm for the same light bulb type is given in Nordin (1974). Culture temperature in the l i f e history study was 10±1°C. In the environ-mental factor study the temperature was held at four levels: 5, 10, 15 and 20°C. Environmental Methods 1. Life History Study. Plants were collected at one or more of the three shore stations at Squamish every 2-3 wks. After being dried in the laboratory in a 20°C growth chamber overnight to simulate low tide water emersion, green portions of upright thalli in the size 16 range 0.3-1 mm x 10-60 mm were examined with a dissecting microscope. If there were no visible algal epiphytes, each portion was rinsed and placed in 5 cm x 1 cm covered pyrex petri plates with ca. 10 ml of culture medium. Within 24 h after immersion the culture was examined for swarmer numbers, size and flagella number. Morphology, size and colour of the thalli were also noted. The subsequent settlement, germination, and growth of the new generation were followed by examination of the settled swarmers semi-weekly for the first month and bi-weekly thereafter. The culture medium was changed once a week. Culture examination was terminated in August 1975 when the new thalli were ca. 1-2 mm high. 2. Environmental Factor Study. In order to determine the effect of several environmental factors on the net photosynthesis of the alga, a study was initiated in the summer of 1975. After examining the field results of the previous 12 months and considering the labor-atory equipment available, three factors were chosen for study: salinity, temperature, and phosphorus and nitrogen concentration. Salinity (°/ Q Q) a n c* temperature (°c) levels are shown in Fig. 19. Three nitrogen (KNO3) and phosphorus (KH2P0it) combinations (ug-at»£ 1) were used (Fig. 19): Nx 12.5 N and 1.25 P N2 25.0 N and 2.50 P N3 50.0 N and 5.00 P The plants used in this study were obtained from Shore Station II. The culture medium was defined seawater (Appendix III). Plants were grown in a l l possible combinations of the environmental factor levels listed above. Medium (100 m£) and ca. 5 gm (soaked weight) of alga 17 were added to a 250 ml erlenmeyer flask and incubated five days. Fresh medium was added on the second and fourth day. On the fifth day the algae in each treatment were placed in three clear 300 mil B.O.D. bottles and incubated for four hours in fresh culture medium of the same salinity and nutrient concentration. For each treatment there was a control bottle. The net photosynthetic rate of the alga in each of the three replicates was determined using equation 2. This is similar to equation 1 except there is a factor of 4 used to convert the four hour rate to an hourly rate. mg O2'£~1'0.3«0.278 ( 2 ) mg C*gm ash-free dry wt 1 ,h 1 = : ash-free dry wt*4 Using the results of this study, the effect of each environmental factor on the net photosynthetic rate was determined by using an Analysis of Variance Statistical Test. The statistics were performed on an IBM 370 (UBC Computing Centre) using two UBC computer programmes: ANOVAR for 3-way analysis of variance and MFAV for 2-way analysis of variance. 18 FIELD RESULTS Description of Study Area 1. Introduction. Most of the study area description is abstracted from Hoos and Void (1975) which should be referred to for a more detailed description. The Squamish River estuary (49°41'N, 123°10'W) is situated in the coastal western hemlock biogeoclimatic zone of British Columbia (Krajina 1970, 1973). It is ca. 48 km north of Vancouver at the head of Howe Sound, and may be classified a turbid outwash fjord using the criteria of Burell and Matthews (1974) (Fig. 1). In general, granite plutonic rock underlies the greater portion of the Squamish drainage system (Hoos and Void 1975). The inshore sediments of the Squamish delta are fine-to coarse-grained, containing large admixtures of s i l t and only small amounts of clay. The s i l t and clay content increases to the east as the effect of river current decreases. Silt and clay also accumulate in the sedge marshes of the mid and upper intertidal areas. The source of the sand is mainly volcanic or metamorphic. Sedimentation at the delta front is heavy with an average rate of delta front advancement approaching 6.0 m«yr - 1. 2. Atmospheric Environment. The Squamish estuary is in a moderate maritime environment. The precipitation regime is wet winter-dry summer with a monthly average over 25 cm between October and January inclusive and less than 8 cm between May and August inclusive (Hoos and Void 1975). Mean winter snowfall at Squamish is 145 cm and mean annual precipitation is 203 cm. 19 The maritime influence is reflected in the low range of temperature between cool summer and mild winter. Mean daily temper-ature is ca. 9°C with the mean monthly high in July (17°C) and mean monthly low in January (0°C) (Hoos and Void 1975). Winds are an important part of the atmospheric climate at Squamish. The north-south axis of the Squamish River Valley and Howe Sound restricts wind direction to a north-south axis. Between October and March the dominant wind direction is southerly due to Pacific storms approaching the mainland (Hoos and Void 1975). Frequently, cold arctic air builds up east of the Coast Mountains and suddenly surges down the Squamish Valley. These strong northerly winds are of short duration. During the summer, May to August, strong winds persist at night. The mean annual wind speed is 15 km per hour (Hoos and Void 1975). The pyranometer data indicated pronounced seasonal fluctuations in direct solar radiation at Squamish. The mean monthly radiation was ca. 231 lyday 1. The minimum was ca. 43 in January 1975 and the maximum was ca. 633 in June 1975. These data were based only on a short sampling period (August 1974 to July 1975). The radiation available for photosynthesis is approximately 47% of the direct solar radiation (Vollenweider 1974). Using sunrise and sunset times the mean daylength is 12.26 h with the longest daylength in June (16.27 h) and the shortest daylength in December (8.17 H) (Canadian Almanac and Directory 1974). 3. Aquatic Environment. Tides are of a mixed semi-diurnal type. Tide level varies from 0 to 4.7 m (Anonymous 1974). The mean tide 20 range is 3.2 ra. Strong winds or unusual river flow rates can change tide levels substantially. Diurnal inequality in time and height of succeeding tides results in low tides occurring during the day in the summer and during the night in the winter (Fig. 5). The Squamish River drains approximately 2500 km2 of the western slope of the Coast Mountains (Hoos and Void 1975). A 4.8 km long training dyke completed in 1972 now restricts the river's discharge to the west side of the dyke (Fig. 1). This dyke virtually removes any freshwater discharge through the Central Channel, thus allowing the transgression of seawater up the channel. Upon reaching the river mouth freshwater flows from west to east across the delta front. On reaching the eastern shore, the flow becomes southerly to Watts Point where i t reverses and flows from east to west. The long term (18 yrs) mean monthly flow of the Squamish River is 243 m^sec 1 with the maximum usually occurring in June (763 m3,sec *) during the spring runoff and the minimum usually in March (71 m3,sec 1) (Hoos and Void 1975). Flash floods can also occur in the autumn when heavy rains combine with sudden snow-thaw. During spring runoff and flash floods the flow rate is very rapid throughout the river's course and the water is heavily s i l t -laden (Hoos and Void 1975). The duration and intensity of the spring runoff in 1974 appeared to be greater than in 1975. This reduction was reflected in the salinity and light attenuation data. The turbidity of the estuary is directly related to the runoff of the Squamish River. The highly turbid condition is mostly confined to the upper brackish layer of 5 m or less. The rate of sedimentation 21 Figure 5. Frequency of tidal flooding (%) above chart datum (lowest low water) during the winter and summer. The vertical bar indicates the maximum vertical distribution of B. minima var. subsalsa during the period indicated. To face page 21 m Winter ( D e c 1 6 - 2 2 , I974) % Summer ( June 1 9 - 2 0 , I974) % 22 also Increases during high turbidity periods (spring runoff and flash floods) (Hoos and Void 1975). Light attenuation in the estuary water was affected by the turbidity. In general, light attenuation was greatest at the river mouth (Station I) and decreased towards the Mamquam Channel (Station III). During high runoff (May to August) almost a l l light was absorbed in the fir s t 2 m (Fig. 6). During autumn, winter and early spring, when runoff was lowest, light penetration was maximum (Fig. 6). The April 1975 light penetration was unusually low. This can be attributed to a spring phytoplankton bloom (M. Pomeroy pers. comm.). Because of the variation in runoff and winds there was marked seasonal variation in salinity distribution, horizontally and vertically (Fig. 7). The annual salinity cycle at a l l three stations was similar to the annual river flow cycle. Salinities were lowest during periods of high runoff (May to August) and highest during low runoff (December to February). Station I was influenced the most by river flow (Fig. 7). There was a surface layer of low salinity for most of the year. It was 3 m or more deep during the summer. The mean surface salinity was ca. 2-3°/ with a minimum of 0.0°/ in June and a maximum of 5.4°/ oo oo oo in January. Station II and III salinities were very similar (Fig. 7). Generally both had a low salinity layer during the summer months. During the winter months, when there was no stratification and runoff was low, the upper layer of water consisted of a mixed zone of uniform salinity. The mean surface salinity at Station II was 12.6°/ oo with a range of 1.2 to 27.9°/ . At Station III the mean surface oo salinity was 10.0°/ with a range of 1.4 to 26.7°/ J oo 6 oo 23 Figure 6. Attenuation of light in the top 3 m of water at Stations I, II and III for the period July 1974 to July 1975. Values are expressed as a percent of the radiation just above the surface of the water. To f ace page 23 Figure 7. Water temperature (°C) and salinity (°/ 0 0) at Stations I, II and III for the period February 1974 to July 1975. To face page 24 12-.8-4-o-\ F 1 M 1 A 1 M ' J 1 J ' A ' S ' O ' N ' D ' J ' F ' M ' A ' M ' J ' J Station U 3 0 -20-10 F ' M ' A ' M ' J ' J ' A ' S ' O ' N ' D ' J ' F ' M ' A 1 M~ r"J~ r"T 30H 20^ 10-F ' M ' A ' M ' J " J ' A ' S ' O ' N ' O ' J ' F ' M ' A ' M ' J ' J 25 Station I had the lowest mean temperature (6.2°C) with a range of 2.6°C-9.2°C (Fig. 7). The mean temperatures for Station II (7.6°C) and Station III (7.8°C) were similar; however, the range for Station II (5.0°C-9.9°C) was much smaller than for Station III (1.9°C-11.8°C) (Fig. 7). The water of the Squamish River is cold for most of the year because i t contains ice and snow melt from the Coast Mountains (Hoos and Void 1975). Station I temperature was depressed throughout the year. Station II and III temperatures were only affected by the river during high runoff. Generally the temperature variation at a l l three stations was very small and insignificant compared to changes in salinity, light and nutrient concentrations. The pH values were nearly neutral at a l l three stations through-out the year, ranging from a low of 6.65 to 8.13 (Appendix IV). Nitrite concentrations at a l l three stations were very low and varied l i t t l e throughout the year (Appendix IV). The nitrate values followed an annual cycle similar to salinity and river runoff (Fig. 8). At a l l stations values approached zero during summer and then reached a maximum in the winter. The winter maximum was greatest at Station II (29.6 ug-at«£ 1 ) . In April 1975 nitrate concentration suddenly dropped to zero during the spring phytoplankton bloom. Ammonia concentration (ug-at*£ 1) had two maxima and two minima at a l l three stations (Fig. 8). The concentration was high in the winter and late spring and dropped to 0 during the summer and early spring. At a l l three stations the ammonia concentration usually 26 Figure 8. Nutrient concentrations at Stations I, II and III for the period August 1974 to July 1975. To f ace page 26 10 S t a t i o n nr 27 increased and peaked before nitrate. At Station III ammonia concentration was always inversely related to the nitrate concentra-tion. This phenomenon might be related to decomposers associated with the abundant wood substrate in the Mamquam Channel. The concentration ranged from 0-5 pg-at*£ 1 at a l l three stations. Reactive phosphorus values varied l i t t l e among the three stations or throughout the year (Fig. 8). The concentration was never over 2 ug-at*£ dropping to 0 during the summer months. The Squamish River is not rich in nutrients. The low human population density, absence of agriculture, high flow rate and nutrient-poor parent material in the watershed keep nutrient concen-trations in the Squamish River low, except for high silica concentra-tions (Hoos and Void 1975). The Squamish River is relatively low in total carbonate alkalinity (<20 mg CaC03«£ 1 ) , especially when compared to sea water (100 mg CaC03'jf 2) (Cliff and Stockner 1973). Nutrient concentrations usually increased with salinity, e.g. below the halocline (Appendix IV). At a l l stations the water was either near saturation or super-saturated with oxygen throughout the year (Fig. 9). Supersaturation occurred most frequently during high runoff. Values ranged from a low of 77.5 in January to a high of 124 in April with percent satura-tion usually decreasing with depth. Such high oxygen saturation was due to well-mixed cold river water meeting warmer sea water. Preliminary observations of water motion suggested the following. Water motion was always high at Station I due to river flow. The annual regime followed the runoff regime. At Station II and III, water motion was not very great and highly irregular. Any water motion 28 Figure 9. Percent oxygen saturation of surface water at Stations I, II and III for the period February 1974 to July 1975. To face page 28 % F ' M ' A ' M ' J ' J ' A ' S ' O ' N ' D ' j ' F ' M ' A ' M ' j ' j Station H 80-F ' M ' A ' M ' J ' J ' A ' S ' O ' N ' D ' J ' F ' M ' A ' M ' J 1 J % 70-F ' M ' A ' M ' J ' J ' A ' S ' O ' N ' D ' J ' F ' M ' A ' M ' J ' j 29 was due to tidal ebb and flow or surface turbulence. There was no surf at any of the stations. Biology of Blidingia minima var. subsalsa and Associated Organisms 1. Annual Cycle of Percent Cover, Swarmer Release and Desiccation of B. minima var. subsalsa. In general, percent cover began to increase in March at Station I and in April to June at Station II and III (Fig. 10). The period of swarmer release was late February to early May. Percent cover and vertical distribution at a l l tidal elevations increased to a maximum in July-August at a l l stations. During this period of rapid development, very few swarmers were observed. Follow-ing August, abundance decreased at a l l elevations and vertical distribution narrowed as thalli turned yellow and deteriorated. Maximum percent cover at a l l three stations was at ca. 3.25 m above zero chart datum. The maximum range was 1.5-4.0 m above this datum. Maximum percent cover was always greatest on substrate facing south or east and smallest facing north or west. The south or east aspect had the highest light intensity and desiccation. On the east side of the dyke opposite Station I where there was no river flow, higher salinities, greater light penetration and less suspended sediment, the alga appeared only for a short period in late summer and early f a l l . The vertical distribution at Station I was not stable during the period April to July 1975 due to silting during spring runoff, which smothered portions of Station I (Fig. 10). The alga did not disappear completely during the study period at any station. During periods of low abundance in the winter the plants were reduced to basal discs and short encrusting thalli. As the percent cover increased, the upright 30 Figure 10. Percent cover and vertical distribution of B. minima var. subsalsa at Stations I, II and III for the period July 1974 to July 1975. At Station I, area indicated by dashed lines was under water at time of sampling. To face page page 30 31 portion of the thalli reappeared and grew up to lengths of 30 cm. Desiccation showed a similar pattern at a l l three stations (Table 1). High desiccation is represented in the table by low values because the field weight was low compared to the soaked weight when desiccation was high. During the period November to February no samples were taken because of insufficient algae. However, low solar radiation, low temperatures and high tides during daylight ensured minimum desiccation during this period. Desiccation was greater in the period April to September and generally increased with elevation. The most desiccated thalli weighed only 7% of the soaked weight. High percent cover of the alga from 3.0 to 3.3 m reduced desiccation during this period. 2. Production Measurements of B. minima var. subsalsa. The net photosynthetic and respiration rates varied considerably (Appendix V). Few obvious patterns in the rate measurement can be discerned. Profiles taken from bimonthly periods showing the annual cycle of net photosynthesis are illustrated in Fig. 11. At Station I net photosynthesis was usually maximum at the sur-face and decreased with depth. There was a subsurface maximum only in early autumn and late winter when light penetration was greatest. Highest total daily photosynthesis occurred in late summer, early autumn and early spring. During high runoff positive net photosynthesis was reduced to the first metre. Negative net photosynthesis at a l l depths was encountered in December and May. At Station II the trend was similar to Station I. However, net photosynthesis was lower in May to August 1974 and higher in May to July 1975. Again there were two minima but in December i t was not 32 Table 1. Desiccation at Station I, II and III. Values indicate field weight of thalli as a percent of soaked weight. Station I 1974 1975 Ht. July Sept Oct Nov to Feb Mar April May June July (m) 23 2 29 18 15 13 6 6 4.0 — 23(2)* — 3.7 — 21(1) — insuff. — — — 88(1) — 3.3 48(3) 30(2) -- alga 100(1) 37(2) 59(3) 92(1) 46(1) 3.0 — 34(1) 65(1) (oa. 100% 100(1) — — 85(1) 48(1) 2.7 64(3) 21(2) — at a l l — -- 58(1) 2.4 — 21(1) 100(1) heights) — — 97(2) 2.1 — 1.8 100(3) Station II 1974 1975 Ht. July Aug Oct Nov to Mar April May June July (m) 16 29 29 16 1 6 6 4.0 — 3.7 58(3) 30(1) 66(1) — 7(2) — 3.3 — — — Insufficient 27(1) 3.0 78(3) 53(1) 93(1) alga — — 48(1) 37(1) 2.7 — — — (oa. 100% at 2.4 — 53(1) 100(1) a l l heights) — -- — 55(1) 2.1 — 1.8 72(2) 82(1) — — 88(2) — * number of replicates in parentheses 33 Station III 1974 1975 Ht. July July Aug Oct Nov to Mar May May July (m) 5 30 29 29_ 1 .13 6 4.0 — 3.7 35(3) 14(2) 26(1) 47(1) — 21(1) — 3.3 — — — — Insufficient — — — 3.0 — 15(2) 20(1) 54(1) alga 2.7 59(3) — — - - (ca. 100% at 2.4 — 16(2) 30(1) 90(1) a l l heights) 2.1 — 1.8 -- — — — 60(2) 1.5 — — 20(1) — 34 Figure 11. Representative net photosynthesis profiles (mg Cgm ash-free dry wt-l.day-1) at Bottle Incubation Stations I, II and III for the period May 1974 to July 1975. The open circles indicate negative net photosynthesis. To face page 34 35 as low. Thus Station II net photosynthesis measurements reflected the strong influence of the river runoff cycle, i.e. the reduced runoff in the summer of 1975 allowed the net photosynthesis in water to increase. Station III net photosynthesis did not show the influence of river runoff as much. There was no definite minimum during high runoff. A subsurface maximum occurred in July 1974 as well as in the late winter-early spring period. Negative net photosynthesis was seen only during May to September 1974. Periods of high net photosynthesis occurred throughout the year, especially during late winter and spring. This coincided with high light penetration and high nutrient concentration. The December vertical profile was highly unusual and has not been considered in the results. 3. Seasonal Occurrence of Other Dominant Organisms. Benthic Algae: The number of benthic algal species that occurred with Blidingia minima var. subsalsa was never very high and at a l l stations no species was as abundant (Appendix VI). The highest number was six species at Station III in July 1975. Further from the delta B. minima var. subsalsa encountered an increased abundance of Fucus distichus subsp. edentatus (De La Pylaie) Powell and was gradually replaced. Fylaiella littoralis (Lyngbye) Kjellman occurred at a l l three stations. It was most abundant during late winter and early spring and seemed to prefer low light and high salinity. Since i t grew in Stoney Creek (Station III) into early summer, low light must be more limiting. Fucus distichus subsp. edentatus was found at a l l three stations. It definitely was most abundant at Station II and III when salinity was highest (December to April). When the study was started, very 36 l i t t l e Fucus was in the vicinity of Station II (June 1974). During 1975 i t was at Station II even during the summer, indicating the Central Channel was undergoing transition towards a more marine environment. In addition, Rivularia sp. and Rhodochorton purpureum (Lightfoot) Rosenvinge had also established themselves at Station II. Rhizoclonium implexum (Dillwyn) Kiitzing appeared at a l l three stations during high runoff and high solar radiation and temperature (April to August). Monostroma oxyspermum (Kiitzing) Doty also appeared most abundant at the same time of year but only at Station II and III, which indicated i t preferred a higher salinity than did Rhizoclonium implexum. In contrast to B. minima var. subsalsa, a l l of the algae mentioned above attained maximum coverage and highest vertical distribution on substrates facing north or west. This aspect decreased the light intensity, desiccation and temperature, especially during emersion. Animals: Only one benthic animal in its larval stage was con-sistently observed to be associated with Blidingia populations. It was a chironomid of the genus Saunderia (Camptocladius) as identified by Dr. C. Levings. The larvae were present in great abundance at a l l three stations from January to early May and occurred only when Blidingia was present, possibly using i t for protection from predators and a food source. Mytilus edulis (L.) and Balanus glandula Darwin were present at Station II and III but were not observed at Station II prior to autumn 1974. Both are firmly established now and are associated with F. distichus subsp. edentatus at both stations. 37 LABORATORY RESULTS Life History Study Greatest success in swarmer examination was experienced by removing the thallus from the culture and examining the edge of the thallus under a glass cover slip with a compound microscope at oa. 400x. The swarmer discharge season lasted from January to May. If swarmers were released they were most abundant in the first 24 h but active swarmers were s t i l l observed after 48 h. There were two tear-drop shaped swarmer types (Fig. 12, 13). One type was a small biflagellate swarmer oa. 3.75 ym long with two flagella of equal length (Fig. 12). The second was a quadriflagellate swarmer oa. 5.5-6.0 ym long with four equal flagella (Fig. 13). Fusion of two isomorphic biflagellate swarmers produced from the same thallus was observed frequently indicating the plants were monoecious. The quadriflagellate zygote produced through fusion could not be distinguished from other quadriflagellate swarmers. There was no morphological difference between thalli producing the biflagellate and quadriflagellate swarmers. Germination of zygotes, zoospores and possibly gametes could not be distinguished f rom one another because flagella were lost shortly after settlement. As reported by Bliding (1963) some germinating cells produced a hollow germination tube (Fig. 14). Cell division continued until a prostrate disc was formed (Fig. 15). The compact shape of the disc was not apparent in most plants. This may be attributed to culture conditions. The culture conditions might also have contributed to the granular appearance of some cells (Fig. 15). 38 Figure 12. Biflagellate swarmer. Figure 13. Quadriflagellate swarmer in end view. Figure 14. A multicellular stage following germination. Note hollow germination tube. Figure 15. Prostrate disc stage. The upright thallus is just beginning to form. Figure 16. Prostrate disc with two upright thalli. Figure 17. Initial form of cell masses produced by the basal cells of the plants. Figure 18. Final stage observed in cell masses. To face page 38 I 1 2 0 / im l 1 20 pm 39 Within two months an upright thallus was produced that was similar to thalli observed in the field (Fig. 16), indicating isomorphic generations. As the culture aged unusual groups of cells were produced by the basal cells of the plants (Fig. 17, 18). They were formed within the basal filaments and released. The size of the masses varied from 1-20 cells. The cells were spherical with a dense cytoplasm and thick outer walls (Fig. 18). Attempts to get these masses to grow were unsuccessful using the defined sea water medium, temperatures of 5 and 10°C, salinities of 10 and 30°/ and nutrient levels Ni oo A and N3. These cell masses were never observed in nature. Environmental Factor Study The results of the environmental factor study, 3-way ANOVA test and 2-way ANOVA test are in Appendix VII, VIII and IX respectively. The 3-way analysis of variance indicated significant interaction among a l l three factors. In order to determine interactions between any two factors, i.e. temperature vs salinity (3 nutrient levels), temperature vs nutrients (5 salinity levels) or salinity vs nutrients (4 temperature levels), refer to Appendix IX. The results indicate interactions are highly unpredictable and vary in degree of signif-icance. Salinity and temperature interact at a l l three nutrient levels. Temperature and nutrients interact at a l l five salinities. However, salinity and nutrients interact only at the lowest (5°C) and highest (20°C) temperatures. Because there is so much interaction i t is difficult to simplify and predict the alga's response to any combination of the 40 three factors. Figure 19 is a synthesis of the 2-way interaction data. There is some degree of error in each graph because the possible interaction with the third factor has been ignored in order to simplify the presentation. For example, on the temperature vs salinity graph each point is the mean of values at the three nutrient levels and the response surface of temperature vs salinity at each nutrient level would not be exactly the same as the one displayed. To determine the response surface at each nutrient level see the data in Appendix IX. Similarly, this also applies to the other two interactions in Fig. 19. Generally net photosynthesis increased with temperature and nutrient concentration and increased with salinity to a peak at 20°/ (Fig. 19A). The lowest salinity (0.25°/ ) had a drastic oo oo effect on net photosynthesis regardless of factor combinations. In the temperature vs salinity interaction net photosynthesis increased with salinity at a l l temperatures (Fig. 19A). However, net photo-synthesis did not always increase with temperature. At 5 and 12°/^ net photosynthesis increased at the two temperature extremes. In the temperature vs nutrient interaction maximum net photosynthesis shifted considerably with a change in either temperature or nutrient level (Fig. 19C). As temperature increased, optimum nutrient concen-tration also increased to a high at 20°C and nutrient level N3. The optimum was likely at a higher temperature and nutrient concentration because this was the highest level used for both factors. In the salinity vs nutrient interaction, depending on the nutrient concen-tration, net photosynthesis peaked at a different salinity (Fig. 19B). As nutrient concentration increased optimum salinity decreased. 41 Figure 19. Two way interactions in environmental factor study. A Temperature vs salinity interaction B Nutrient level vs salinity interaction C Nutrient level vs temperature interaction 42 If a l l three factors were varied, the net photosynthetic rate reflected interactions among a l l three factors (Appendix IX). As the nutrient concentration was increased, the optimum temperature increased and the optimum salinity decreased slightly. Photosynthetic rate at the optimum combination of temperature and salinity increased with nutrient concentration. At N^  the optimum combination was 10°C and 30°/ , at N2 20°C and 30°/ and at N3 20°C and 20°/ . Net oo ^ oo 3 oo photosynthesis was greatest at 20°C and 20°/ q q and N3. Minimum net photosynthesis was at 5°C and 0.25O/qo at a l l nutrient levels. The lowest photosynthetic rate measured was at 5°C, 0.25O/qo and N2 (Fig. 19). 43 DISCUSSION Environmental Factors, Growth, Reproduction and Distribution The annual cycle of standing biomass of Blidingia minima var. subsalsa as measured by cover estimation does not parallel net photosynthetic measurements. There are several possible deficiencies in the net photosynthetic measurements, which might explain this discrepancy. The water was saturated or supersaturated for a considerable part of the year. This can introduce errors in net photosynthetic measurements (Strickland and Parsons 1972, Pope 1975). The measurements were in an aquatic environment but Blidingia can be emersed for 75% or more of daylight hours in the summer. There is evidence that a considerable amount of the total primary produc-tivity in the marine intertidal occurs while the algae are exposed (Johnson, Gigon, Gulman and Mooney 1974). Therefore cover estimation will be considered the major growth parameter in the ensuing discussion. Since the alga occurs high in the intertidal zone, i t must integrate successfully the atmospheric and aquatic environment. Its high position in the intertidal suggests the factors determining its vertical distribution, growth and reproduction are purely physical and chemical (A.R.O. Chapman 1973). For benthic algae, Zaneveld (1969) considers "the principal factors affecting lit t o r a l zonation in a direct way are: exposure to waves, impact of waves, strength of tidal currents, turbulence, duration of emergence and submergence (desiccation), type of substrate, temperature, salinity, pH, 02-content, availability of nutrients, quantity and quality of light, moving ice, the periodicity of these factors and competition. 44 With increasing height in the eulittoral and supralittoral region there is a decrease in the time of submergence, an increase in the time of desiccation, an increase in intensity of light and a steeper gradient for salinity, temperature and pH." There are several reasons for excluding some bf these factors. The preliminary observations on wave action, tidal currents and turbulence suggest the effect is minimal except at Station I where strong river currents are important during submergence. During periods of high runoff, these currents are responsible for depositing s i l t and may be a major factor in determining the lower limit of the alga at Station I. The alga does not seem to have a substrate preference as i t grows on rocks, pilings, marsh plants, mud flats, driftwood or floating free in tide pools. The pH of the water is very stable except in tide pools (C. Levings pers. coram) and the oxygen content is always near saturation. Moving ice is never a problem because thick ice never occurs at Squamish. Competition is only important as the aquatic environment becomes more marine. Fucus distichus subsp. edentatus grows at the same elevation in marine water and as its biomass increases, B. minima var. subsalsa disappears. Of a l l the factors listed above, I consider the major factors in this study are: quantity and quality of light, temperature, salinity, availability of nutrients and desiccation. A l l of these factors are physical or chemical in nature. It cannot be assumed any of these factors operate independently on growth, reproduction and distribution. Interpretation of results is limited because of the many possible interactions. Where appropriate, the physiological and biochemical mechanisms that may be involved will be referred to 45 only as far as seems necessary for the understanding of the responses considered. 1. Light. Light serves as the energy source for photosynthesis and hence plays a dominant role in a l l photosynthetic plants. Their functional and structural responses are largely affected by light intensity, quality and exposure patterns (Hellebust 1970). During the study period maximum percent cover developed in July and August with a second maximum just prior to spring runoff. The minimum occurred in February, March and April. There was also a minor decrease in May and June during high runoff. Solar radiation and photoperiod maxima and minima were positively correlated with the extremes in maximum and minimum percent cover suggesting light plays a leading role in the alga's growth. The increased light attenuation associated with the lesser minimum in May and June supports this suggestion. Positive net photosynthesis was restricted to the surface during immersion in May and June. No laboratory experiment was done to determine the effect of light intensity and quality on increases in biomass but field results suggest a preference for or at least a tolerance to high solar radia-tion intensity of the normal solar spectrum. The vertical zonation and seasonality of the alga's biomass can be controlled in part by light. At its upper limit, the alga is exposed to high intensity, direct light. It is well known that high light intensity may be damaging to attached marine algae (Hellebust 1970). This is probably not the case with B. minima var. subsalsa because Biebl (1952a,b, 1956) found that algae normally exposed to high light intensity in the upper intertidal zone suffered no damage 46 after five hours of direct exposure to sunlight, 105 klux (oa. 0.63 lymin" 1, Strickland 1958). Low light intensities and changing light quality may be assoc-iated with the lower vertical limit of the alga because light intensity decreased and light quality changed with water depth. Since a photosynthetic alga relies on light as the energy source, i t will cease to grow and begin to degenerate beyond a certain lower tolerance limit or compensation level (Hellebust 1970). The light compensation level can be altered by environmental conditions. Temperature and heterotrophic utilization of dissolved organic substances (Hellebust 1970) as well as hydrostatic pressure (Zaneveld 1969) may be involved. For B. minima var. subsalsa these three factors may be ignored because the alga did not grow deep enough to be affected by temperature or hydrostatic pressure and there is no evidence that the alga is capable of heterotrophic utilization of dissolved organic substances. Green algae (Chlorophyta) due to their pigment content absorb effectively the blue and red wavelengths of visible radiation which are present in the upper layers of water (Zaneveld 1969, Levring 1968). In coastal waters, where transmission of red and blue light is reduced due to suspended particles and dissolved substances, the majority of Chlorophyta must be littoral forms (Levring 1968). At Squamish, where light transmission is heavily attenuated, visible radiation must be considered the major factor controlling the lower limit of B. minima var. subsalsa. Ion uptake is often light dependent and is closely interrelated with membrane potentials (Soeder and Stengel 1974). Brackish water species usually are quite hyperosmotic (Kinne 1967, Gessner and Schramm 1971) and an alga, especially an estuarine alga, relies on ion transport in osmoregulation and osmoadaptation (Gessner and Schramm 1971, Soeder and Stengel 1974) . In a l l algal cells studied, osmoregulation is controlled by mechanisms for active extrusion of sodium and active uptake of chloride and also active uptake of potassium in some cells (MacRobbie 1974) . There is also evidence for active uptake of phosphate, sulphate, bicarbonate and active efflux of hydrogen ions in some algae. Light is important because these active transport systems require energy, which ultimately must be produced by photosynthesis. Therefore i t is advantageous for B. minima var. subsalsa to maintain osmoregulation by maximizing photo-synthesis. The easiest way for the alga to accomplish this is to grow in the upper intertidal zone where the amount of light is greatest. The consequences of this are discussed in greater depth in the salinity section. Although i t was not examined in the field or tested in the environmental factor study, there is evidence that nutrient conditions as well as temperature may influence the relationship between light and growth in marine algae (Maddux and Jones 1964). In general, higher light intensities are required for maximal growth rates at optimal than at less favourable conditions of nutrition and temperature (Hellebust 1970). If this is true then i t would help to explain the increased percent cover of B. minima var. subsalsa during the spring when nutrients were abundant and periods of high light intensity were increasing and also the second increase in late summer when nutrients were in low abundance and periods of high light intensities were decreasing. 48 The alga's period of reproduction from January to early May occurred when visible radiation and photoperiod was increasing and light penetration in water was high. There was no attempt to study the relationship between light and photoperiod and reproduction. Hellebust (1970), in summarizing the information available, concludes that light intensity, light quality and photoperiod significantly affect the reproduction of unicellular as well as multicellular marine plants. Experiments on the effect of different light inten-sities on the growth of sporelings in Japan have shown that sporeling growth in green algae, except for Monostroma, is greatly retarded under low light (Zaneveld 1969). Arasaki (1953) demonstrated that Enteromorpha sporelings grow better under long day conditions, whereas sporelings of Monostroma grow best ini t i a l l y under medium day length, later under short day conditions. An analysis of its geographical distribution suggests that B. minima var. subsalsa prefers long day photoperiods because i t is usually found above or below the two tropics and rarely between. Therefore i t is reasonable to expect the period of reproduction and i n i t i a l increase in percent cover to occur from January to early May. 2. Temperature. Temperature should be considered as a possible factor controlling the growth, reproduction and distribution of B. minima var. subsalsa. In various algae temperature interacts with photosynthesis and respiration (Kanwisher 1966, Gessner 1970, Yokohama 1972) as well as ion transport and osmoregulation e.g. salinity interaction, (Soeder and Stengel 1974, this study) and nutrients (McCombie 1960, Maddux and Jones 1964, this study). 49 The temperature regime of the alga's environment is highly variable and unpredictable because i t resides in the upper intertidal zone. The daily and yearly tide cycle determines whether i t will be exposed to air or water temperature. At Squamish, water temper-ature had a pattern that followed solar radiation but with much less variation between maximum and minimum. Because of this coincident pattern, perhaps there is a similar relationship with growth, repro-duction and distribution. The environmental factor study indicated maximum net photosynthesis at water temperatures of 20°C or more. Only in tidal pools in mid-summer could temperatures of 20°C or more ever be reached at Squamish. Water temperatures in the range recorded at Squamish rarely increased net photosynthesis in the environmental factor study (Fig. 19). Pomeroy (1974) showed that there was very l i t t l e correlation between water temperature and growth of any alga in the Squamish estuary. Although water temperature exhibits very l i t t l e direct control over growth, the environmental factor study suggests i t may exert consider-able control through interaction with other factors, e.g. salinity and nutrients (Fig. 19). Air temperature had a wider range than water temperature. It followed a pattern similar to radiation and water temperature. Mean air temperature was lower than water temperature during periods of low percent cover and higher during periods of high percent cover. This was further accentuated by tidal cycles which exposed the alga during the night in winter and during the day in summer (Fig. 5). Air temperature probably exerted greater influence than water temperature. The optimum temperature in the environmental factor study was encountered 50 more often in air than in water. Temperatures of 20°C or more occurred during exposure in the summer when radiation was high and winds dried the thalli. It has long been observed that intertidal marine algae tolerate extremes of temperature variation during exposure although temper-ature is generally assumed to be one of the main factors determining the upper limit of algal growth (Zaneveld 1969). Indeed B. minima has been reported from water temperatures of -22°C to at least +25°C and I suspect the alga experiences even greater extremes while emersed. In intertidal algae resistance to frost and to high temperature has been correlated with resistance to desiccation (Dawson 1966, Kanwisher 1957) which in turn is physiologically equivalent to an ability to withstand changes in concentration of sea water (Dawson 1966). B. minima var. subsalsa is evidently eurythermal and i t is not surprising that the relatively moderate water temperature regime at Squamish has l i t t l e direct control of the alga's growth. The air temperature might have some control over the upper limit of the alga's vertical distribution when the temperature is too high for the alga to survive. The high degree of desiccation observed in B. minima var. subsalsa during the summer might help i t to withstand higher air temperature. This phenomenon has been observed in intertidal algae e.g. Bangia fusaopurpurea (Dillwyn) Lyngbye, Urospora penicilliformis (Roth) Areschoug (Biebl 1962) and Fucus vesiculosus L. (Schramm 1968). Although there are no general trends or rules (Gessner 1970) temperature can have quantative as well as qualitative effects on number and type of spores released in various algae (Dring 1974). Very l i t t l e can be said on control of reproduction in B. minima var. 51 subsalsa by temperature. . Together with temperature other important factors such as light intensities and photoperiod also vary in situ. The effect of temperature is not apparent in the field study or in the laboratory study. If temperature does influence reproduction increasing temperature, especially air temperature, would probably be correlated with increasing number of swarmers. As early as 1851, W.H. Harvey considered temperature to be the major factor affecting plant growth and geographical distribution. Since Setchell's studies (1915, 1917, 1920a, b) which demonstrated the relation of various algal species to the 0°, 10°, 15°, 20° and 25°C isotherms of the surface waters of the ocean, there is general agreement that temperature controls the geographical distribution of algae (Zaneveld 1969). Because the chemical milieu of an estuarine alga is not as uniform as that of marine algae, one should not expect the temperature effect to be as pronounced. Also, the equator-ward increase in temperature cannot be separated from a similar increase in intensity of light. By referring to water temperature and current maps in Kinne (1970) the southern extreme of the alga in the northern hemisphere is associated with southern extensions of cold water e.g. Louisiana (Kapraun 1974) and the west coast of Africa (Lawson and Price 1969). In the north Atlantic, Caribbean Sea and Mediterranean Sea, the southern limit is associated with the 20°C mean February isotherm and ca. 29°C mean August isotherm. In the remainder of its ocean di s t r i -bution i t has never been reported from water temperature higher than 20°C mean August isotherm. Toward the north pole i t extends as far as ca. 0°C mean February isotherm and ca. 5°C mean August isotherm. Toward the south pole the alga's limit occurs at ca. 12°C mean 52 February isotherm and ca. 9°C mean August isotherm. 3. Osmotic Effects. The salinity of the open ocean is fairly constant and so is the ionic composition. The salinity of estuaries often undergoes considerable fluctuation. The ionic composition and the relative proportions of other solutes may be subject to significant modifications that are not due solely to dilution (Khlebovich 1968, Kirch 1956). The effect of the Squamish River on ionic composition and ratios can be inferred from the study of two similar British Columbia fjords (Bute and Knight) by Kirsch (1956) and the review paper by Khlebovich (1968). In general, at salinities below 5°/QQ the Ca:Na, K:Na and Cl°/ :S°/ ratios increase. The cation ratio changes are oo oo not due to dilution, direct precipitation or experimental methods. The changes are due to cation exchange between the marine water and suspended clay particles introduced to the estuary by the river. Calcium is released by the clay and replaced by other cations, such as potassium, sodium or magnesium. The increase in the Cl°/ o o:S°/ o o ratio presumably occurs through a similar hydrochemical process not yet understood. The measurements of salinity in this study do not provide sufficient information about the true ionic environment of the alga. The salinity values can only give a general impression of the osmotic environment of the alga. B. minima var. subsalsa is subject to temporary emergence. During these periods, changes in osmotic conditions, as a result of rain or desiccation, are of considerable ecological importance (Gessner and Schramm 1971). The period of exposure and the atmospheric environmental conditions during exposure could be more important in 53 growth, reproduction and distribution than the period of immersion. Therefore desiccation (water loss) is included in this section as well as salinity. Salinity: In the Squamish estuary, while the alga was immersed i t was subjected to increasing salinity with depth of immersion. The annual salinity cycle in the top 2 m followed the river runoff cycle. Salinity was highest when algal percent cover was at a minimum and river runoff was lowest. When salinity was lowest, during high run-off, algal percent cover was not at a maximum. The reproductive period occurred when salinity had reached a maximum and was decreasing due to spring river runoff. There is an apparent correlation between salinity and growth, reproduction and distribution. Percent cover decreased at various low salinities and reproduction occurred at brackish salinities (5 to 30°/ ). This observation is misleading because the environmental oo factor study showed net photosynthesis was optimum between 20 and 30°/ (Fig. 19). The alga would experience such salinities during high river runoff only while emersed or at depths >2 m while immersed. Boyle and Doty (1949) demonstrated that certain algae could tolerate reduced salinities better i f the temperature was lowered. Results of the environmental factor study indicated this is not always true (Fig. 19). Net photosynthesis increased with low temperature only at 5 and 12°/ . At a l l other salinities i t stayed the same or decreased slightly. In order to understand the results discussed here, the effect of salinity on the alga's physiology must be considered. The main 54 regulatory device for counteracting detrimental effects of salinity is considered to be ion regulation. Most of the aquatic algae investigated tend to maintain an internal osmoconcentration somewhat higher than that of the surround-ing water. Most intertidal algae can tolerate concentration ranges of 0.1-3.0 times that of sea water (Biebl 1962). The internal osmo-concentration of marine algae and brackish water algae is usually hyperosmotic i.e. internal osmoconcentration (cell fluids) of marine algae generally ranges from 1.3-1.5 times seawater (Gessner and Schramm 1971). Most marine plants are "stenohyperosmotic" (Gessner and Schramm 1971). For example, in the euryhaline alga Chaetomorpha linwn (Muller) Kiitzing, turgor remained almost constant at 14.8-16.5 atmospheres over a salinity range (balanced a r t i f i c i a l sea water) from freshwater to 35°/ (Kessler 1959). oo Cellular osmoregulation in marine algae depends on the activity and specificity of inorganic ion pumps (Gessner and Schramm 1971, Soeder and Stengel 1974). These ion pumps involve active sodium efflux and chloride and potassium influx requiring energy in the form of ATP. Recent evidence shows that Na, Cl and K can act as activators of membrane-bound ATPases in marine phytoplankton and corn (Falkowski 1975, Maslowski and Komoszynski 1974). Membrane-bound ATPases have been demonstrated to support ion translocation although their mechanism remains obscure (Falkowski 1975). In a hyposmotic environment, such as the Squamish River mouth, the alga is subjected to the release of cell constituents into the ambient water. It has beem well known for a long time that calcium ions reduce the plasmotic permeability of other ions. This fact accounts for a 55 delay, and possibily a reduction, of damaging effects under hyposmotic conditions in the presence of calcium (Gessner and Schramm 1971). Calcium maintains membrane selectivity towards potassium and sodium (Kinne 1967). The ions that are important in osmoconcentration and osmoregula-tion are also the ions that undergo significant ratio changes at low salinity. B. minima var. subsalsa probably can maintain its pre-sence to the upper limit of salt water intrusion in the Squamish River mouth because the interaction of marine and freshwater would increase the Ca:Na and K:Na ratios. These modifications would permit the alga to endure frequent exposure to hyposmotic conditions. Further-more, its upper intertidal location permits i t to produce ATP through photosynthesis for ion transport. The absence of the alga on the more favourable east side of the dyke can be attributed to the absence of sufficient suspended clay particles to maintain favourable ion ratios. Unfavourable ion ratios might also explain the low percent cover of the alga in the winter when water salinity is high. During emersion the heavy winter rains wash away the seawater film normally present around the thalli. Rain is potentially a very serious threat to sur-vival, due in part to breakdown of ion transport (Kinne 1967, Gessner and Schramm 1971). In a study of Porphyra perforata J. Agardh, Eppley and Cyrus (1960) say heavy rain, because i t has low calcium content, would "almost certainly abolish membrane selectivity, decrease respir-ation, induce loss of cellular cations and result in high mortality i f the blades were exposed long enough". During the period of swarmer release immersion salinity was high-est but s t i l l quite variable (0-28°/^). Salinity around the thalli 56 during exposure would be increasing as radiation, photoperiod and exposure during daylight hours increased. The importance of salinity in reproduction is not very clear. In estuaries, decreasing salinity is usually related to reduced reproductive capacity of asexual reproduction (Nienhuis 1974, Russell 1971, Gessner and Schramm 1971). Russell (1971) states that only asexual reproduction occurs in several marine algae including B. minima (var. minima?). The occurrence of sexual reproduction in the laboratory in brackish salinities indicated this is not always true. According to Gessner and Schramm (1971) there is no doubt that salinity greatly affects sexual and asexual reproduction. However, there are few supportive facts in the literature and there is an urgent need for an analysis of the relation between salinity and algal reproduction (Gessner and Schramm 1971). Desiccation: As with many other intertidal algae (Gessner and Schramm 1971), one of the most important factors in the vertical distribution of B. minima var. subsalsa seems to be desiccation. Desiccation is controlled by the ebb and flow of the tide, by tempera-ture and by wind. Desiccation controls the environment of the alga while i t is emersed. Increase in percent cover commenced in the spring as desiccation increased. The period of reproduction occurred just prior to and during the increase in desiccation. A considerable portion of the increase in algal percent cover occurred in periods of high desiccation with maximum cover between 3.0-3.3 m. Measured desiccation at these levels during high percent cover were quite low (10% - 20%). The alga easily withstands high desiccation but requires frequent periods of 57 immersion. The morphology and physiology of the alga actually encouraged desiccation. Its hollow thallus was light, long and thin with many short branches. This increased surface area and thus desiccation. The thalli in mid-summer were loose and limp. The wind and sun quickly dried them. Evidently the plant did not exude mucus to retard desiccation as occurs in many members of the Fucaceae or Laminariaceae. The manner in which desiccation affects algal physiology is contentious. Part of the problem is due to inadequacies in method and materials (E.B. Tregunna pers. comm.). Johnson et at. (1974) point out that in various studies using different algae, some show significant increases in photosynthetic rate, others show rates equal to or below submerged rates and one showed a greatly reduced rate. Such variation in results could be due to more than one factor. Ogata and Matsui (1965) and Hammer (1968) believe that in studies of photosynthetic rate, changes in salinity, osmotic pressure, pH and also carbon dioxide supply, particularly in natural seawater, are inseparably associated. Since no study has considered a l l of these factors, i t is obvious why there are so many conflicting results. In the study by Johnson et al. (1974) the capacity to attain high photosynthetic rates in air is positively correlated with the vertical zonation of the algae studied (Prionitis lanoeolata Harvey, Ulva expansa (Setchell) S.&.G. Iridaea flaccida (S.&G.) Silva, Porphyra perforata, Fucus distichus L . , Endocladia muricata (Harvey) J. Agardh). Plants exposed 50-80% of the time, e.g. B. minima var. subsalsa, adapt to living under these conditions rather than relying on periods of submergence for their productivity (Johnson et al. 1974). Continuous desiccation 58 gradually depresses the photosynthetic rate. In Porphyra3 desiccation affects the photochemical reaction in photosynthesis (Fork and Hiyama 1972); drying stopped normal oxidation and reduction of cytochrome /. Respiration is depressed during desiccation in most algae but is enhanced by slight desiccation in some species (Ogata 1968, Chapman 1966). The recovery rate of respiration is positively correlated with vertical zonation of the algae studied. These studies suggest increase in percent cover of B. minima var. subsalsa under normal desiccating conditions is due to an increased photosynthetic rate and possibly a depressed respiration rate. The upper limit of algal distribution coincides with the approximate upper limit of tidal flooding (Fig. 5), beyond which desiccation is continuous. Desiccation should be considered the limiting factor at the upper level of the alga's vertical distribution. Osmotic effects, after temperature, seem to be the most important factor in determining geographical and habitat distribution. Although i t has been reported in marine and freshwater habitats, B. minima var. subsalsa is most common in brackish water. It survives in this habitat because i t can tolerate the changing osmotic environment. The fresh-water habitats in which i t occurs must possess a favourable ionic environment that allows i t to resist release of cellular constituents. The experiments by Baker (1909) show algae which can resist osmotic effects best, e.g. desiccation, grow more slowly while those which grow most quickly have the lowest tolerance to desiccation. B. minima var. subsalsa^ which fits into the f i r s t category, cannot outgrow other algae in the more favourable marine environment and eventually loses substrate to Fucus distichus subsp. edentatus. 4. N u t r i e n t s . There i s p r e s e n t l y good evidence that the f o l l o w i n g i n o r g a n i c elements ( i n a d d i t i o n to C, H and 0) are r e q u i r e d by one or more a l g a l s p e c i e s : N, P, K, Mg, Ca, S, Fe, Cu, Mn, Zn, Mo, S i , Na, Co, V, C l , B and I (O'Kelley 1974). Of these, B l i d i n g i a minima var. subsalsa r e q u i r e s at l e a s t N, P, Mg, Fe, Cu, Mn, Zn and Mo because these elements are considered to be r e q u i r e d by a l l algae (O'Kelley 1974). Each of these elements could be an important, f a c t o r i n determining the alga's growth, r e p r o d u c t i o n and d i s t r i b u -t i o n at any p a r t i c u l a r time or l o c a t i o n . Vitamins and growth r e g u l a t o r s may a l s o be important but cannot be discussed i n depth because of i n s u f f i c i e n t data. A l s o any e s s e n t i a l or n o n - e s s e n t i a l compound may be an important f a c t o r when i t i s present i n t o x i c c o n c e n t r a t i o n . Such p o l l u t a n t s may be placed i n one of the f o l l o w -i n g c a t e g o r i e s (FAO Dept. of F i s h e r i e s , F i s h e r y Research D i v i s i o n 1971): halogenated hydrocarbons, e.g. p e s t i c i d e s ; i n o r g a n i c chemicals e.g. heavy metals; organic chemicals, e.g. phenols; n u t r i e n t chemicals e.g. c o n t a i n i n g N or P; r a d i o a c t i v e chemicals; or petroleum. A f t e r carbon, n i t r o g e n and phosphorus are the most important elements i n an a l g a l c e l l . In the water, the C:N and C:P r a t i o i s u s u a l l y higher than i n l i v i n g organisms and n i t r o g e n or phosphorus g e n e r a l l y becomes the f i r s t l i m i t i n g n u t r i e n t . The two most common sources of n i t r o g e n used f o r growth of algae are n i t r a t e and ammonium ions (Morris 1974). N i t r i t e can a l s o be used but i t s t o x i c i t y at higher concentrations makes i t l e s s u s e f u l . For most algae, ammonium i s u t i l i z e d p r e f e r e n t i a l l y ( S y r e t t 1962) although there are exceptions (Morris 1974). The growth r a t e i s u s u a l l y the same on ammonium as i t i s on n i t r a t e ( S y r e t t 1962). 60 Nitrate, nitrite, ammonium (Morris 197A) and phosphate (Kuhl 1974) uptake is stimulated by light. In Skeletonema costatum3 a membrane-bound (nitrate, chloride)-activated ATPase apparently translocates nitrate across the plasmalemma (Falkowski 1975). In-side the cell 2 enzymes (nitrate reductase and nitrite reductase) reduce nitrate to ammonium, using energy (Morris 1974). Light is stimulatory because i t produces the necessary ATP and electron donors through photosynthesis. The implications of this interaction is discussed in the light section. The most common source of phosphorus is inorganic phosphate. The uptake of phosphate is an active light-dependent process result-ing from phosphate utilization in photophosphorylation as well as in other processes (Kuhl 1974). Active transport of ions, which require phosphorus in the form of ATP, is inhibited when phosphorus is not available (Kuhl 1974). Phosphate uptake may depend bn the simultaneous presence of any of Na, K or Mg (Healey 1973). This possible interdependence might restrict B. minima var. subsalsa's ability to maintain its internal osmoconcentration at the upper limit of its vertical distribution where i t is exposed to nutrient-poor river and rain water. The nutrient content of the freshwater input could be important in maintaining photosynthesis. Littoral algae show an increase in photosynthesis i f exposed to freshwater high in C, N and P, e.g. 6.7 meq*£ 1 alkalinity, 1.12 ug-afJl 1 P, 27 ug-at*£ 1 N, compared with their normal salinity (Zavodnik 1975). If they were exposed to distilled water photosynthesis was depressed. The Squamish River is low in carbonate alkalinity and nutrients, therefore its effect 61 would probably be to depress photosynthesis. This might also explain the depressed net photosynthesis at low salinity in the environmental factor study which used distilled water (Fig. 19B). Various algae are also capable of utilizing organic nitrogen, e.g. urea, amino acids or organic phosphorus, but such sources will not be considered in this study. In coastal waters, nitrogen is usually the first nutrient to limit phytoplankton growth (Ryther and Dunstan 1971), including local estuaries such as Puget Sound (Winter, Banse and Anderson 1975) and Fraser River (Takahashi, Fujii and Parsons 1973). The same nutrient should limit benthic algae as well as phytoplankton under the same conditions. Borowitzka (1972) found that "nitrates were one of the nutrients limiting growth of Ulva as well as unicellular algae and phosphates were not limiting". The limiting nutrient may change seasonally (Smayda 1974, Sullivan and Daiber 1975, Thayer 1974). There also may be seasonal variations in intensity of limitation by nutrients (Takahashi, Fujii and Parsons 1973, Smayda 1974), several nutrients limiting simultaneously and with a different intensity (Smayda 1974) and seasonal variation in limiting nutrient combinations (Smayda 1974). The availability of inorganic nutrients to an alga such as B. minima var. subsalsa is determined by water circulation and regen-eration which may be supplemented by sewage in some areas (Steemann Nielsen 1971). In the Squamish estuary, transport of relatively nutrient-rich subsurface water to the surface via the estuarine mechanism provides the major input of nutrients. The Squamish River provides very few nutrients. The phytoplankton, marsh plants, bacteria and other benthic algae compete with B. minima var. subsalsa for the upwelled nutrients. Thayer (1974) and Parker, Sibert and Brown (1975) give evidence that microbial immobilization of N and P during decomposition of organic matter may limit nutrient availability to algae. The annual cycle in nutrient concentration in an estuary in part may result from shifts in the equilibrium between microbial immobilization and remineralization (Thayer 1974). The immobilization and remineral-ization of nitrogen and phosphorus in plants, animals and microbes depends on the balance between the C:N:P ratio of the non-living matter in the estuary and the desired C:N:P ratio of the estuarine organisms (Parker et al. 1975, Ryther and Dunstan 1971, Steemann Nielsen 1971, Thayer 1974). Using an organism C:N:P ratio of 100:10:1.(Ryther and Dunstan 1971, Steemann Nielsen 1971, Thayer 1974), i f the C:N or C:P ratio in the non-living matter in the estuary is greater than 10:1 or 100:1, respectively, immobilization of inorganic nitrogen and phos-phorus will occur. Immobilization in this sense refers to uptake and preferential retention of nutrients by living cells. As heter-otrophic respiration releases the carbon as CO'2, remineralization will occur when the ratios are equal to or less than 10:1 or 100:1 respectively. Remineralization can be considered as the liberation of nutrients into the estuary from the estuarine organisms. During remineralization phosphorus is quickly liberated from dead material, whereas nitrogen is comparatively more refractory to decomposition (Menzel and Ryther 1964). At Squamish spring runoff and summer thermal stratification 63 combined with primary production by marsh plants, phytoplankton and benthic algae to reduce the concentration of nitrogen and some-times phosphorus to very low levels during the summer. As f a l l approached, salinity increased and the growing season ended for many plants, e.g. B. minima var. subsalsa, remineralization by microbes commenced. First ammonia and then nitrate increased to their maximum values during the winter. From August to November both nitrogen and phosphorus were low. The N:P ratio never exceeded 10:1 except in March (Station I) and January to March (Station II). The phosphorus concentration remained relatively stable at a l l stations. These data suggest nitrogen was a limiting factor throughout the year except perhaps during the period January to March. Possibly the more rapid remineralization of phosphorus helped to keep its con-centration stable. Nitrogen did not begin to remineralize until December because the abundant organic matter available in the estuary from dead marsh plants and detritus entering from the river encouraged immobilization by decomposing microbes. As nitrogen was remineralized i t appeared first as ammonia and then was oxidized to nitrate with the aid of bacteria. The nitrate maximum from January to March was much higher than the ammonium max-imum. At most times of the year ammonia concentration was usually lower than nitrate concentration possibly because of the availability of oxygen for oxidation and preferential uptake of ammonia by algae and other organisms. Blidingia minima var. subsalsa may have been limited in its growth from May to early f a l l by a low N:P ratio. Perhaps the desiccation during summer increased nutrient concentrations around 64 the thalli. Its high intertidal position would also increase its access to light and thereby increase its ability to absorb nitrogen and phosphorus. During the winter when remineralization was occurr-ing and the N:P ratio was greater than 10:1, nutrients should not have been an important factor. The results of the environmental factor study indicated maximum net photosynthesis at level N3 or higher. This level was never encountered at Squamish but i t may have occurred in marsh tide pools with high nutrient remineralization from the sediments. At Squamish there were several potential sources of pollutants that may have affected B. minima var. subsalsa: organic chemicals, . petroleum and inorganic chemicals. The organic chemicals were produced by wood debris from a lumber mill on the Mamquam Channel. The hydrogen sulphide and methane produced by decomposing wood debris (Hoos and Void 1975) may have discouraged growth of the alga in the Mamquam Channel. Petroleum losses from the numerous motorboats present in this channel also may have been involved. Losses of mercury from F.M.C. Chemicals (a chlor-alkali plant) at the head of the Mamquam Channel has produced mercury levels in benthic fauna as high as 13.4 ppm (27x the permissible level) (Hoos and Void 1975). However the growth of the alga was not adversely affected because i t was very abundant on the pilings around the plant. The reproductive period occurred when nitrate and phosphorus concentrations were maximum and N:P was greater than 10:1. This suggests the proper nutrient concentrations may be one of the factors initiating and maintaining the release of swarmers but further studies must be done to confirm this. 65 Although its vertical distribution was unaffected, the alga's horizontal distribution in the estuary was apparently restricted by the toxic conditions prevailing in the Mamquam Channel. Edwards (1972) in a study of three British estuaries, found B. minima equally distributed in a non-polluted and a nutrient-polluted estuary but reduced to a more saline distribution in an estuary polluted by industrial chemicals. A l l of the other algae associated with B. minima var. subsalsa at Squamish (except Fucus distichus subsp. edentatus and Rivularia ap.) were also mentioned in his study and were similarly affected. These results suggest the alga would maintain its growth, repro-duction and distribution i f enriched with nutrient chemicals but i f exposed to high concentrations of organic chemicals, inorganic chemicals or petroleum i t would not survive. These chemicals would act either through direct toxicity or immobilization or nutrient chemicals. Systematic Consideration of Life History Study Some phycologists believe Blidingia should remain within the genus Enteromorpha. Chapman and Chapman (1973) argue that the basal disc and small cells are also found in some species of Enteromorpha. Primary development somewhat similar to Blidingia is found in some species of Enteromorpha and Ulva (van den Hoek 1964). The primary development, considered a major taxonomic character in Blidingia as well as in Enteromorpha and Ulva, may depend on culture conditions and the nature of the substratum (van den Hoek 1964, C. Tanner pers. comm.). The significance of the cell masses observed in this study is 66 uncertain. Lokhorst and. Vroman (1974) encountered a similar phen-omenon in exhausted Ulothrix sonata (Weber and Mohr) Kiitzing cultures. They referred to them as true akinetes but were unsuccessful in attempts to make them germinate. Recently Tatewaki (1972, pers. comm. 1975) reported, in a Japanese population of B. minima (var. minima?), sexuality and an alternation of isomorphic generations as in most species of Entero-morpha and Ulva. Fusion of biflagellate swarmers and isomorphic gen-erations in this study indicates sexuality exists in B. minima var. subsalsa. Hori (1972), in a study of pyrenoids, demonstrated a pyrenoid structure for B. minima (?) that is similar to members of Ulvaceae (sensu Bliding 1968) and dissimilar to most of the Monostrom-aceae (sensu Bliding 1968). The evidence is accumulating in favour of placing Blidingia in the Ulvaceae and not in the Monostromaceae. Whether or not the genus is taxonomically distinct from Enteromorpha is a question that requires further investigation. CONCLUSIONS B l i d i n g i a minima var. subsalsa was present year-round in the Squamish River estuary and was associated with very few other benthic algae. Its maximum vertical distribution was from 1.5-4.0 m above chart datum (lowest low water), with maximum percent cover at ca. 3.25 m. Percent cover, reproduction and distribution varied throughout the study period. Percent cover was minimum in the winter and maximum in the summer, except for a slight decrease during spring runoff from the Squamish River. Annual light intensity was closely associated with the percent cover of the alga, which was highest under high light intensity and long photoperiods. Salinity affected the alga's percent cover when the alga was exposed frequently to freshwater (spring runoff) or heavy rain (winter). Increasing salinity in late autumn did not have any affect on its percent cover. As the alga's percent cover increased so did air temperature and desiccation A l l three factors were maximum during mid-summer. Apparently growth of the alga was not adversely affected by high air temperatures and desiccation as long as i t was frequently re-immersed in water. The absence of any competitive benthic algae or heavy grazers throughout the study period probably had a beneficial effect on the alga's abundance. There was also some suggestion that favourable ion ratios e.g. K:Na, Ca:Na, I Q O ' ' ^ ° IQO' w e r e important in increasing the alga's percent cover, especially in the mouth of the Squamish River. Nitrogen and phosphorus concentrations were low during periods of high percent cover and the N:P ratio was considerably less than 10:1. 68 Under these conditions nitrogen may have been limiting growth. In the laboratory, temperature, salinity and nutrients interacted in their effect on the net photosynthesis of the algae. Net photo-synthesis was greatest at 20°'/' , 20°C (highest temperature used in the study) and nutrient level N3 (highest nutrient level used in the study). This combination of factors would only occur at Squamish during low river runoff (brackish salinities), periods of emersion in the summer (high temperatures) and in areas of high nutrient input (marine water or sewage) or nutrient remineralization (tide pools). Minimum net photosynthesis was at 0.25O/qo and 5°C at a l l nutrient levels. This combination of salinity, temperature and nutrients would only occur at Squamish during high river runoff or exposure to rain while emersed (low salinity and nutrients) and in the winter or while immersed in water in summer (low temperatures). Algal reproduction occurred by swarmer release which began in January and ended in early May. Monoecious sexual reproduction by isomorphic biflagellate swarmers was observed in the laboratory. Quadriflagellate swarmers were also released and isomorphic generations were observed. Release of swarmers coincided with increasing photo-period, light intensity and light penetration into the water. During this period the influence of freshwater input from the Squamish River was at a minimum. Salinity values, although very variable, reached their maximum during swarmer release. Nutrient concentrations, which were associated with high salinities, were also maximum. The N:P ratio was maximum during the period of reproduction. The alga's vertical distribution was determined from the percent cover values. The upper limit was associated with the upper limit of tidal flooding. Beyond this point desiccation was too great. The lower limit was associated with low light intensities and perhaps low frequency of emersion. The horizontal distribution was limited on the freshwater side of the estuary by adverse osmotic effects due to low salinities and nutrients. 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Recent additions to the marine algal flora of Nahant, Massachusetts. Rhodora 77: 430-433. * Wilce, R.T. 1959. The Marine Algae of the Labrador Peninsula and Northwest Newfoundland (ecology and distribution). Nat. Mus. Canada. Bull. No. 158. 81 pp. * Womersley, H.B.S. 1956. A critical survey of the marine algae of southern. Australia. I. Chlorophyta. Aust. J. Mar. Freshwater Res. 7: 343-383. * Womersley, H.B.S. and A. Bailey. 1970. Marine algae of the Solomon Islands. Philos. Trans. R. Soc. London. Ser. B. 259: 257-352. Zaneveld, J.S. and W.M. Willis. 1974. The marine algae of the American coast between Cape May, New Jersey and Cape Hatteras, North Carolina. II. The Chlorophycophyta. Bot. Mar. 17: 65-81. * Zinova, A.D. 1967. Opredelitel' zelenykj, burykj i krasnykj vodoroslei yuzhnykh more! SSSR. Akademiya Nauk SSSR, Moskva. 396 pp. * Appendix II. Procedures used in Physical and Chemical Analyses 0 2 Salinity, in situ Salinity, in laboratory Water temperature, in situ . . . . Winkler titration (Strickland and Parsons, 1972) YSI Model 33 SCT meter Beckman RS5-3 SCT meter Hytech Inductive Salinometer Autosal Inductive Salinometer Mercury thermometer YSI Model 33 SCT meter Beckman RS5-3 SCT meter pH Orion pH meter N - NH phenol-hypochlorite method (Strickland and Parsons, 1972) N - reactive NO Cadmium reduction method 3 (Strickland and Parsons, 1972) N - reactive NO Sulphanilamide-azo dye method (Strickland and Parsons, 1972) P - reactive phosphorus Phospho-molybdate method (Strickland and Parsons, 1972) 86 Appendix III. Culture Media (a) Enriched Seawater 1. Seawater from Squamish (ca. 10-12°/ ) was filtered through 0.45 ym fi l t e r paper °° 2. To each litre was added 1 ml of 50 mg-at'l-1 N (KNOJ and 1 ml of 5 mg-at-1"1 P (KH^O^) (b) Defined Seawater Medium (National Eutrophication Research Program, 1974) 1. Basal Medium (35°/ ) oo Compound g.1-1 NaCI 23.48 Na2S0i+ 3.92 NaHC03 0.19 KC1 0.66 KBr 0.10 H3 B 03 0.03 MgCl2.6H20 10.61 SrCl2.6H20 0.04 CaCl2.2H20 1.47 ^0 to 1000 ml. 2. For Dilution to Various Salinities (4 1 batches) Salinity 0/ Basal medium Distilled Ho0 oo A2 il) (l) 30.00 3.430 0.570 20.00 2.290 1.710 12.00 1.370 2.630 5.00 0.570 3.430 0.25 0.028 3.972 To each 4 litres add: Na2 EDTA 1200 Vg *Trace metal mix 4 mis C^ompound mg» 500 ml*"1 H3BO3 92.8000 MnCl2«4 H20 208.0000 ZnCl2 16.0000 CoCl2.6 H20 0.7140 CuCl2.6 R^ O 0.0107 Na2Mo04.2 H20 3.6300 Filter through 0.45 urn membrane f i l t e r Add F e C l 3 (384 ug-4 l" 1) Add N (KNO3) and P (Kl^PO^) to make the nutrient levels Ni, N2 and N3: Nutrient level N(ug-af l" 1) P(yg-af l*"1) N x 12.5 1.25 N 2 25.0 2.50 No 50.0 5.00 88 Appendix IV. Water Quality Results at Station I, II and III (a) Station I Collection Date Analysis Date 1975 N(yg-N03 at-1 - 1 N02 ) NH3 P(yg-af 1 _ 1) Salinity (°/ ) v 1 oo' PH 11/Aug Ilk May 31 0.000 .073 0.000 0.277 1.1 7.45 6/Sept/74 May 29 0.346 .075 0.000 0.513 1.2 7.39 22/Dec /74 May 29 0.037 .068 1.643 1.202 3.4 8.13 22/Jan /75 May 29 5.614 .048 4.354 0.404 3.9 7.20 19/Mar /75 May 29 10.419 .045 0.449 0.555 2.1 6.81 *19/Mar /75 May 29 9.644 .023 0.000 1.396 31.6 7.85 14/May /75 May 31 8.751 .126 2.573 1.237 7.0 7.19 ll/June/75 July 31 4.215 .115 0.000 0.004 0.4 6.93 9/July/75 July 31 2.330 .191 4.416 0.131 1.1 7.11 (b) Station II 11/Aug /74 May 31 0.000 .126 0.602 1.855 0.8 7.22 6/Sept/74 May 29 0.000 .027 0.000 0.000 2.0 7.10 22/Dec /74 May 29 0.793 .068 1.304 0.303 3.6 6.65 22/Jan /75 May 29 29.715 .301 0.000 2.094 27.0 7.66 19/Mar /75 May 29 15.832 .023 0.000 1.102 7.4 7.96 *19/Mar /75 May 29 17.728 .041 0.000 1.312 22.6 7.74 31/Apr /75 May 31 0.000 .086 1.139 0.871 14.8 7.98 14/May /75 May 31 6.412 .147 2.901 0.000 9.4 7.68 ll/June/75 July 31 3.720 .095 0.000 0.015 2.4 7.35 20/June/75 July 31 2.188 .084 2.556 0.018 1.2 7.10 9/July/75 July 31 3.104 .143 0.359 0.023 1.6 6.92 89 (c) Station III Collection Date Analysis Date 1975 N(yg-N03 •at-1-1 N02 ) NH3 P(ug-at. I"1) Salinity pH (°/oo) 11/Aug /74 May 31 0.000 .065 1.040 1.725 0.7 6.98 6/Sept/74 May 29 0.894 .335 0.000 1.287 2.6 7.64 22/Dec /74 May 29 1.131 .080 4.057 0.521 4.3 7.88 22/Jan /75 May 29 6.194 .041 0.830 0.597 6.3 7.47 19/Mar /75 May 29 8.152 .018 0.034 0.471 6.0 7.69 *19/Mar /75 May 29 18.652 .050 0.152 1.623 30.4 7.87 14/May /75 May 31 5.596 .330 1.894 0.098 7.2 7.68 20/June/75 July 31 . 1.255 .098 2.514 0.009 1.1 7.11 9/July/75 July 31 1.848 .078 0.000 0.000 1.1 7.00 * Below the halocline Appendix V. Net Photosynthetic and Respiration Rates (mg Cgm ash-free dry wt - 1•day - 1) at Station I, II and III (a) Station I Date Depth (m) Net Photosynthesis Respiration Rate (1974) June 7 0 +00.63 -16.53 July 5 0 +10.50 - 4.75 1 + 9.45 - 3.50 2 + 3.70 - 5.30 Aug 11 0 +28.88 0.00 1 +26.70 - 4.04 2 + 9.52 - 2.88 Aug 27 0 +11.52 - 6.12 1 +11.95 -17.57 2 0.00 -27.04 Sept 2 0 +23.78 0.00 Oct 30 0 +16.87 - 4.06 1 +19.84 - 4.20 2 + 9.84 - 3.43 Dec 22 0 -35.18 -10.98 1 -30.33 -16.22 2 -31.51 -66.59 (1975) Jan 22 0 +17.58 -17.31 1 + 3.59 -17.04 2 - 8.15 -20.96 Feb 20 0 0.00 - 8.40 1 + 8.40 -10.28 2 0.00 - 6.49 Mar 19 0 +22.64 + 4.65 (?)* 1 + 7.71 -14.69 2 + 1.02 + 3.59 (?) May 14 0 - 1.85 -11.21 June 12 1 +15.06 -17.82 July 9 0 + 2.31 - 8.82 * (?) indicates value is questionable (b) Station II Date Depth (m) Net Photosynthesis Respiration Rate (1974) May 22 0 + 4.39 -16.92 June 3 0 -13.52 - 8.44 June 18 0 +3.00 - 6.78 July 19 0 + 4.42 -16.12 1 0.00 -16.42 2 5.27 -14.72 July 30 0 +3.44 - 9.20 Aug 28 0 +15.43 + 0.48 (?) 1 + 1.04 + 3.05 (?) Oct 30 0 +12.56 - 2.02 1 +13.12 - 2.78 2 +13.12 - 0.48 Dec 22 0 + 1.34 -14.66 1 -35.43 -21.73 2 -38.09 -22.14 (1975) Jan 22 0 +13.58 -14.48 1 + 8.63 -23.42 2 + 3.08 -13.35 Feb 20 0 +13.82 -14.48 1 +11.07 -12.14 2 - 4.48 - 8.69 Mar 19 0 +33.68 + 3.25 (?) 1 +44.61 - 2.06 2 +13.76 -25.74 Apr 16 0 +31.66 1 +23.40 -13.87 2 +18.26 May 14 0 +19.28 - 0.92 1 +11.48 - 7.20 2 +11.60 - 7.28 June 12 0 +23.61 -19.95 92 Date Depth (m) Net Photosynthesis Respiration Rate (1975) June 20 0 +20.38 -12.29 1 +15.04 -26.10 2 + 6.46 -15.40 Ju ly 9 0 +17.43 - 2.55 1 +12.18 - 3.21 2 + 6.42 - 3.75 (c) Station I I I (1974) May 27 0 -17.37 -20.13 June 19 0 +20.30 -10.70 1 + 1.00 -10.90 2 - 7.70 - 5.05 Ju ly 5 0 - 0.60 - 4.68 Ju ly 19 0 + 8.40 -23.28 1 +14.76 -14.08 2 + 1.30 -16.92 Aug 28 0 + 2.76 - 0.23 1 - 0.23 -12.42 2 - 1.80 - 6.78 Sept 13 0 0.00 - 6.39 1 0.00 - 0.72 2 - 3.06 0.00 Oct 30 0 +13.72 - 2.06 1 +11.48 - 0.98 2 + 5.32 - 5.51 Dec 22 0 + 3.41 - 8.94 1 +45.02 -14.16 2 +61.77 - 5.29 (1975) Jan 22 0 -16.29 1 + 2.43 -17.41 2 + 1.32 -15.04 Feb 20 0 + 7.58 - 7.38 1 +13.14 - 8.41 2 + 4.08 - 9.50 Date Depth (m) Net Photosynthesis Respiration Rate (1975) Mar 19 0 + 5.70 1 +32.35 - 3.35 2 - 5.52 -28.38 May 14 0 + 9.07 - 5.78 1 - 3.42 - 7.85 2 -14.56 - 5.19 June 20 0 +22.95 -24.59 1 + 7.84 -17.26 2 + 1.95 -14.52 July 9 0 + 9.70 -13.69 1 +10.41 -11.59 2 + 3.76 -14.71 Appendix VI. Associated algal species at Stations I, II and III for the period July 1974 to July 1975 1. Vertical bar indicates maximum vertical distribution 2. n.p. indicates not present 3. Dash (-) indicates station not sampled in that month 95 m 4.0-3.5-30-2.5-Station I n.p. _ _ n.p. _ n.p. Rhizoclonium implexum i n.p. n.p. J ' A ' S ' O ' N ' D ' J " F " M ' A " M ' J " J 4.0-3.5-Pyllaiella littoralis 3.0" 2.5-n.p. — n.p. _ n.p. T T T T T n.p. i 1 1 1 1 J A S O IM D J F M A M J J 4.0H 3.5H Fucus distichus subsp. edentatus 3.0 2.5-n.p. a p. n.p. n.p. n.p. n.p. n.p. n.p. T n.p. I I 1 1 1 1 1 1 1 1 r - r J A S O N D J F M A M J J Station JX 96 4.0-3.5-3.0-2.5-2.0-Monostroma oxyspermum 1 1 J A S N D J F M A M J 4.0-1 3.5-3.0-2.5-2.0-Rivularia sp. ap. ap. — n.p. — 1 1 1 1 r J A S 0 N D i — i ' — i — " 1—' r J F M A M J J 4.0^ Rhizoclonium implexum 35-3.0-2.5-2.0-a p. n.p. — n.p. i I n.p. n.p. n.p. n.p. - - — i 1 1 1 1 1 1 1 1 1 1 1 J A S O N D J F M A M J J 97 3.0-2.5^ 2.0-Petalonia fasc ia a p. n.p. _ 1 1 — n.p. n.p. n.p. n.p. n.p. n.p. n.p. n.p i 1 1 r i r 4.0- Rhodochorton purpureum 3.5-3.0-2.5-2.0-n.p. n.p. — n.p. — n.p. n.p. n.p. n.p. a p. n.p. n.p. 1 1 1 J A S 0 N D J 1 1 1 1 M A M J J Station III 4.0-3.5-3.0-2.5-2.0i Fucus distichus subsp. edentatus J A S O N ' D ' J ' F M A M J J 98 m 4.0-35-3.0 2.5 2.0 Monostroma oxyspermum I I _ n.p. _ n.p. n.p. n. p. n.p. _ J A S O N D J F M A M J J 4.0-1 3.5 3.0 2.5-2.0-Rhodochorton p u r p u r e u m — n.p. — a p . — J A ' S , O , N , D , J , F , M A ' M , , J , J 4.0- Rhizoclonium implexum 3.5-3.0 2.5-2.0-— n.p. — n.p. — a p. n.p. a p. n.p. a p . — .1 J ' A ' S ' O ' N ' D ' J ' F 1 M ' A ' M ' J J 35 Rivularia sp. 3.0- [I 2.5-2.0- — n.p. — a p . — ap. ap. a p. a p. n.p. — n.p. J ' A ' S ' O ' N ' D ' J ' F ' M ' A ' M ' J ' J 99 Appendix VII. Results of Environmental Factor Study. Net Photosyn-thesis (mg C'gm ash-free dry wt - 1 ,hr - 1) at 4 Temperatures (°C), 5 Salinities (°/ 0 0) and 3 Nutrient Concentrations (yg - a f l - 1 ) Temperature Salinity Nutrient Net Photosynthesis Level * Rate  5 0.25 Nx 0.2444 -0.0659 0.0000 N2 -0.1402 -0.4829 0.1819 N3 0.4128 0.0990 0.1203 5.00 Nx 1.4759 1.4915 1.2279 N2 1.5033 1.5870 1.7634 N3 1.0352 0.9798 1.4339 12.00 Nj 1.6461 1.6047 1.6351 N2 1.7473 1.0862 N3 1.9876 1.8796 2.1296 20.00 Nx 1.2501 1.1691 1.1559 N2 1.5280 1.6498 1.7343 * see Appendix III Temperature Salinity Nutrient Net Photosynthesis Level Rate 5 20.00 N 3 1.2431 1.0896 0.7048 30.00 Nj 1.0722 1.0455 1.1900 N 2 1.2669 1.3129 1.2536 N 3 1.1744 1.4741 1.0335 10 0.25 N x 0.5950 0.5331 1.0720 N 2 0.3880 -0.2055 0.3679 N 3 0.6022 0.2758 0.0837 5.00 Nx 0.7212 1.1379 0.8767 N2 0.8369 1.3937 1.2232 N 3 1.1863 0.8109 1.6018 12.00 Nx 1.8953 1.0510 1.2505 N2 0.9484 0.9267 0.7351 N3 1.7079 0.9855 1.4108 Temperature Salinity Nutrient Net Photosynthesis Level Rate 10 20.00 Nj 2.0920 1.1021 1.5649 N 2 1.6960 2.1396 1.6641 N 3 1.9624 1.8027 1.4566 30.00 Nj 1.9043 2.4111 1.8783 N2 1.8200 1.3053 1.5078 N 1.5976 1.6816 1.2841 15 0.25 Nx 0.6276 0.5402 0.6712 N2 0.8490 0.8353 0.9326 N 3 0.6530 0.8184 1.0017 5.00 1.3830 0.4324 1.2580 N2 0.6521 1.2959 1.1004 N3 0.6101 1.3216 1.5783 12.00 Nx 0.4318 0.8764 0.7018 Temperature Salinity Nutrient Net Photosynthesis Level Rate 15 12.00 N 2 1.0003 1.0454 1.6470 N 3 2.2326 1.1056 0.9442 20.00 Ni 1.5177 1.7238 1.9439 N 2 1.7886 2.2948 3.0770 N 3 1.4321 1.5544 1.6995 30.00 NX 1.7190 1.1053 N 2 2.1393 2.5242 1.7324 N 3 1.6418 1.7480 1.5611 20 0.25 Nx 0.3976 0.5864 0.6464 N 2 0.6870 0.9440 0.4717 N 3 0.5533 0.6303 5.00 Ni 0.6379 0.4141 0.9893 N 2 1.7822 1.8358 1.7422 Temperature Salinity Nutrient Net Photosynthesis Level Rate 20 5.00 N3 2.5199 2.3924 2.9568 12.00 Ni 0.9361 0.9087 1.1266 N2 1.7323 1.6439 N3 3.9539 3.1021 2..9981 20.00 Ni 1.0399 1.0442 1.3301 N2 1.7490 1.5455 2.0537 N3 3.0981 3.9778 3.4557 30.00 Ni 2.0241 1.5704 1.7754 N2 2.2375 2.4262 2.1052 N3 1.7230 1.4327 1.1788 104 Appendix VIII. Results of 3-Way Analysis of Variance (UBC ANOVAR) Analysis of Variance Table for Net Photosynthesis - overall mean is 1.330 Source No. Term Name D.F. Sum of Squares Mean Square Probabilityt 1 Temp 3 7.0962 2.3654 27.0028* 0.0000 2 Sal 4 36.8114 9.2029 105.0578* 0.0000 3 Temp x Sal 12 7.9082 0.6590 7.5231* 0.0000 4 Nut 2 3.8875 1.9437 22.1892* 0.0000 5 Temp x Nut 6 9.7271 1.6212 18.5070* 0.0000 6 Sal x Nut 8 5.7917 0.7240 8.2646* 0.0000 7 Temp x Sal x Nut 24 10.7522 0.4480 5.1143* 0.0000 8 Error Total 116 175 10.1614 92.1356 0.0876 * indicates value is significant with 95% confidence limits t probability value indicates probability that F value is solely due to random error 105 Appendix IX. Results of 2-Way Analysis of Variance (UBC MFAV) A. Temperature versus Salinity (at three levels of nutrients) i) Nutrient Level Ni Source D.F. Sum of Squares Mean Square F Probability T Temp 3 0.9096 0.3032 4.30* 0.0103 Sal 4 8.6375 2.1594 30.64* 0.0000 T x S 12 4.6505 0.3875 5.50* 0.0000 Error 39 2.7490 0.0705 Total 58 16.947 T \ S 0.25 5 12 20 30 Mean 5 0.059(3)t 1.398(3) 1.629(3) 1.192(3) 1.103(3) 1.076 10 0.733(3) 0.912(3) 1. 399(3) 1.586(3) 2.065(3) 1.339 15 0.613(3) 1.025(3) 0. 670(3) 1.779(3) 1.412(2) 1.067 20 0.543(3) 0.680(3) 0. 991(3) 1.138(3) 1.790(3) 1.029 Mean 0.487 1.004 1. 172 1.411 1.609 1.129 i i ) Nutrient Level N 2 Source Temp Sal T x S Error Total D.F. 3 4 12 38 57 Sum of Squares 3.0690 17.1150 3.9426 3.1350 27.2620 Mean Square 1.0230 4.2788 0.3285 0.0825 12.40* 51.87* 3.98* Probability 0.0000 0.0000 0.0005 * indicates value is significant with 95% confidence limits t number of replicates in parentheses T probability value indicates probability that F value is solely due to random error 106 T \ S 0.25 5 12 20 30 Mean 5 -0.147(3) 1.618(3) 1.417(2) 1.637(3) 1.278(3) 1.142 10 0.183(3) 1.151(3) 0.870(3) 1.833(3) 1.544(3) 1.116 15 0.872(3) 1.016(3) 1.231(3) 2.387(3) 2.132(3) 1.528 20 0.701(3) 1.787(3) 1.088(2) 1.783(3) 2.256(3) 1.640 Mean 0.402 1.393 1.251 1.910 1.803 1.355 i i i ) Nutrient Level N3 Source D.F. Sum of Squares Mean Square F Probability Temp 3 15.7080 5.2361 47.74* 0.0000 Sal 4 17.4850 4.3713 39.86* 0.0000 T x S 12 8.1507 0.6792 6.19* 0.0000 Error 39 4.2774 0.1097 Total 58 45.6220 T \ S 0.25 5 12 20 30 Mean 5 0.211(3) 1.150(3) 1.998(3) 1.012(3) 1.227(3) 1.120 10 0.321(3) 1.200(3) 1.368(3) 1.741(3) 1.521(3) 1.230 15 0.824(3) 1.170(3) 1.427(3) 1.562(3) 1.650(3) 1.327 20 0.592(2) 2.623(3) 3.351(3) 3.510(3) 1.442(3) 2.426 Mean 0.477 1.536 2.036 1.956 1.460 1.510 B. Salinity versus Nutrients (at four levels of temperature) i) Temperature Level 1 (5°C) Source D.F. Sum of Squares Mean Square F Probability Sal 4 14.2450 3.5613 95.26* 0.0000 Nut 2 0.0330 0.0165 0.44 0.6475 S x N 8 1.6030 0.2002 5.36 0.0000 Error 29 1.0842 0.0374 Total 43 16.9640 S \ N N l N2 N3 Mean 1 0.060(3) -0.147(3) 0.211(3) 0.041 2 1.398(3) 1.618(3) 1.150(3) 1.389 3 1.629(3) 1.417(2) 2.009(3) 1.714 4 1.192(3) 1.637(3) 1.012(3) 1.280 5 1.103(3) 1.278(3) 1.227(3) 1.203 Mean 1.076 1.142 1.120 1.112 ii ) Temperature Level 2 (10°C) Source D.F. Sum of Squares Mean Square F Probability Sal 4 10.4350 2.6088 26.51* 0.0000 Nut 2 0.0372 0.1857 1.89 0.1690 S x N 8 1.4510 0.1814 1.84 0.1077 Error 30 2.9520 0.0984 Total 44 15.2100 S \ N N l N 2 N3 Mean 1 0.733(3) 0.183(3) 0.321(3) 0.412 2 0.912(3) 1.151(3) 1.200(3) 1.088 3 1.399(3) 0.870(3) 1.368(3) 1.212 4 1.586(3) 1.833(3) 1.741(3) 1.720 5 2.065(3) 1.544(3) 1.521(3) 1.710 Mean 1.339 1.116 1.230 1.228 i i i ) Temperature Level 3 (15°C) Source D.F. Sum of Mean Squares Square F Probability Sal 4 8.2607 2.0652 14.38* 0.0000 Nut 2 1.5439 0.7720 5.37* 0.0103 S x N 8 1.3758 0.1720 1.20 0.3347 Error 29 4.1653 0.1436 Total 43 15.3460 S \ N N l N2 N3 Mean 1 0.613(3) 0.872(3) 0.824(3) 0.770 2 1.024(3) 1.016(3) 1.170(3) 1.070 3 0.670(3) 1.231(3) 1.427(3) 1.109 4 1.728(3) 2.387(3) 1.562(3) 1.892 5 1.412(2) 2.132(3) 1.650(3) 1.771 Mean 1.607 1.528 1.327 1.312 109 iv) Temperature Level 4 (20°C) Source D . F . Sum of Squares Mean Square F Probability Sal 4 12.3160 3.0789 43.99* 0.0000 Nut 2 14.1800 7.0899 101.29* 0.0000 S x N 8 10.2960 1.2870 18.39 0.0000 Error 28 1.9599 0.0700 Total 42 38.7510 S \ N Ni N2 N3 Mean 1 0.543(3) 0. 701(3) 0.592(2) 0.615 2 0.680(3) 1. 787(3) 2.623(3) 1.697 3 0.990(3) 1. 688(2) 3.351(3) 2.050 4 1.138(3) 1. 783(3) 3.510(3) 2.144 5 1.790(3) 2. 256(3) 1.442(3) 1.829 Mean 1.028 1. 640 2.426 1.682 C. Temperature versus Nutrient (at five levels of salinity) i) Salinity Level 1 (0.25°/ ) Source D . F . Sum of Squares Mean Square F Probability Temp 3 2.6547 0.8849 18.53* 0.0000 Nut 2 0.0512 0.0256 0.54 0.5920 T x N 6 0.7867 0.1311 2.75* 0.0367 Error 23 1.0983 0.0477 Total 34 4.5908 T\N Ni N2 N3 Mean 5 0.059(3) -0.147(3) 0.211(3) 0.041 10 0.733(3) 0.183(3) 0.321(3) 0.412 15 0.613(3) 0.872(3) 0.824(3) 0.770 20 0.543(3) 0.701(3) 0.592(2) 0.615 Mean 0.487 0.402 0.477 ii ) Salinity Level 2 <5°/oo> Source D.F. Sum of Mean Squares Square F Probability Temp 3 . 2.3643 0.7881 7.99* 0.0007 Nut 2 1.8183 0.9091 9.21* 0.0011 T x N 6 4.3954 0.7326 7.42* 0.0001 Error 24 2.3687 0.0987 Total 35 10.947 T\N Ni N 2 N3 Mean 5 1.398(3) 1.618(3) 1.150(3) 1.389 10 0.912(3) 1.151(3) 1.200(3) 1.088 15 1.024(3) 1.016(3) 1.170(3) 1.070 20 0.680(3) 1.787(3) 2.623(3) 1.697 Mean 1.004 1.393 1.536 1.311 I l l i i i ) Salinity Level 3 (12°/ ) Source D.F. Sum of Squares Mean Square F Probability Temp 3 4.9074 1.6358 12.58* 0.0001 Nut 2 5.3631 2.6816 20.63* 0.0000 T x N 6 5.2451 0.8742 6.72* 0.0004 Error 22 2.8601 0.1300 Toatal 33 18.3760 T \ N Nl N2 N 3 Mean 5 1.629(3) 1.417(2) 1.999(3) 1.714 10 1.399(3) 0.870(3) 1.368(3) 1.212 15 0.670(3) 1.231(3) 1.427(3) 1.109 20 0.990(3) 1.688(2) 3.351(3) 2.050 Mean 1.172 1.251 2.036 1.500 iv) Salinity Level 4 (20°/ ) oo Source D.F. Sum of Squares Mean Square F Probability Temp 3 3.5664 1.1888 .11.44* 0.0001 Nut 2 2.1932 1.0966 10.56* 0.0005 T x N 6 8.6906 1.4484 13.94* 0.0000 Error 24 2.4932 0.1039 Total 35 16.943 112 T \ N Ni N2 N3 Mean 5 1.192(3) 1.637(3) 1.012(3) 1.280 10 1.586(3) 1.833(3) 1.741(3) 1.720 15 1.728(3) 2.387(3) 1.562(3) 1.892 20 1.138(3) 1.783(3) 3.510(3) 2.144 Mean 1.411 1.910 1.956 1.759 v) Salinity Level 5 <30°oo> Source D.F. Sum of Squares Mean Square F Probability Temp 3 2.2184 0.7395 12.68* 0.0000 Nut 2 0.7076 0.3538 6.07* 0.0077 T x N 6 1.6019 0.2670 4.58* 0.0034 Error 23 1.3410 0.0583 Total 34 5.8689 T \ N Ni N2 N3 Mean 5 1.103(3) 1.278(3) 1.227(3) 1.203 10 2.065(3) 1.544(3) 1.521(3) 1.710 15 1.412(2) 2.132(3) 1.650(3) 1.771 20 1.790(3) 2.256(3) 1.442(3) 1.829 Mean 1.609 1.803 1.460 1.624 

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