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The distribution of diatoms in the surface sediments of British Columbia inlets Roelofs, Adrienne Kehde 1983

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THE DISTRIBUTION OF DIATOMS IN THE SURFACE SEDIMENTS OF BRITISH COLUMBIA INLETS by ADRIENNE KEHDE ROELOFS B.A., Michigan State University, 1968 M.Sc.r The University Of Michigan, 1979 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES Department Of Botany We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA A p r i l 1983 © Adrienne Kehde Roelofs, 1983 In presenting this thesis in p a r t i a l fulfilment of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t fr e e l y available for reference and study. I further agree that permission for extensive copying of t h i s thesis for scholarly purposes may be granted by the Head of my Department or by his or her representatives. It i s understood that copying or publication of this thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of Botany The University of B r i t i s h Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date: A p r i l 22, 1983 i i Abstract The purpose of the study was to examine the d i s t r i b u t i o n a l patterns of diatoms in the surface sediments of ten southern B r i t i s h Columbia i n l e t s with respect to oceanographic and hydrographic setting, and phytoplankton d i s t r i b u t i o n and productivity. The study area was divided on the basis of i n l e t type (high, medium, and low runoff), w i t h i n - i n l e t gradients, and zones (northern, c e n t r a l , and southern). A small group of species dominated the 95 sediment assemblages. There was a f a i r l y good co r r e l a t i o n between the biocoenoses and the thanatocoenoses in the sense that most of those species reported as dominants in the phytoplankton were also dominants in the sediment assemblages. However, there were discrepancies and these could not be explained on the basis of the r e l a t i v e s i l i c i f i c a t i o n of the diatom valves. Skeletonema costatum, usually considered a w e a k l y - s i l i c i f i e d , d i s s o l u t i o n - s e n s i t i v e species, was abundant in B r i t i s h Columbia sediments. Both the p a c i f i c a and the a e s t i v a l i s forms of Thalassiosira a e s t i v a l i s were abundant in the phytoplankton, but only the p a c i f i c a form was preserved well in the sediments. Thalassiosira  nordenskioeldii, which i s found in other sediment assemblages, was rare in most B r i t i s h Columbia sediments. The d i s t r i b u t i o n a l patterns of freshwater and marine l i t t o r a l species appeared to be ind i c a t i v e of river sources entering the estuarine system. The absolute abundance of diatoms in the sediment assemblages increased from the northern to the southern zone. Within the i n l e t s , both absolute abundance and primary p r o d u c t i v i t y i n c r e a s e d toward the mouth. E s t u a r i n e c i r c u l a t i o n d i d not appear to a l t e r s u b s t a n t i a l l y the s p a t i a l r e l a t i o n s h i p between the biocoenoses and the thanatocoenoses. In g e n e r a l , i n d i v i d u a l s p e c i e s and species-groups o f t e n e x h i b i t e d d i s t i n c t d i s t r i b u t i o n a l p a t t e r n s which c o u l d be r e l a t e d to i n l e t type, z o n a l , and w i t h i n - i n l e t p a t t e r n s . In p a r t i c u l a r , the p r i n c i p a l c o o r d i n a t e a n a l y s i s showed a zonal c o r r e l a t i o n between the dominant s p e c i e s i n the sediment assemblages, and primary p r o d u c t i v i t y , s a l i n i t y , and temperature in the s u r f a c e waters. Table of Contents Abstract i i L i s t of Tables v L i s t of Figures v i Acknowledgements v i i i I. INTRODUCTION 1 II. PHYSICAL AND BIOLOGICAL SETTING 7 PHYSICAL SETTING 7 BIOLOGICAL SETTING 14 II I . METHODS 18 SAMPLING PROCEDURES 18 CLEANING AND COUNTING 23 TAXONOMY 24 STATISTICAL ANALYSES 27 IV. RESULTS 30 I. LARGE-SCALE DIATOM DISTRIBUTIONAL PATTERNS 30 RECURRENT GROUP ANALYSIS 30 DIVERSITY 33 RELATIVE AND ABSOLUTE ABUNDANCES 36 1. Absolute Abundance Of Total Diatom Valves .36 2. Marine Planktonic Species 38 3. Marine L i t t o r a l Species 58 4. Freshwater Species 60 5. Summary Of Major D i s t r i b u t i o n a l Patterns ..62 PRINCIPAL COORDINATE ANALYSIS 63 II. DIATOM VALVE AND DINOFLAGELLATE CYST DISTRIBUTION 64 V. DISCUSSION 73 PART I 73 I. DIFFERENTIAL DISSOLUTION AND VERTICAL TRANSPORT 74 II. DISTRIBUTIONAL PATTERNS OF SELECTED SPECIES .104 SUMMARY OF PART I 134 PART II 1 37 ZONES 137 INLET TYPES 149 1. Sedimentation 151 2. Estuarine C i r c u l a t i o n 153 HOWE SOUND 161 SUMMARY OF PART II 165 VI. CONCLUSIONS 169 BIBLIOGRAPHY 173 APPENDIX A - SPECIES LIST 187 APPENDIX B - TAXONOMY 193 APPENDIX C - DISTRIBUTION OF SPECIES 197 APPENDIX D - RELATIVE ABUNDANCE OF SELECTED SPECIES 202 APPENDIX E - SITE DATA 207 APPENDIX F - PLATES 264 V L i s t of Tables 1. L i s t of s i t e s , s i t e numbers, and c o l l e c t i o n dates...19 2. D i s t r i b u t i o n of Chaetoceros spp. resting spores....26 3. Species l i s t for recurrent Groups A and B 32 4. The means and 95% confidence intervals of the t o t a l absolute abundance and d i v e r s i t y (H') and a comparison of the zones and i n l e t s using the Wilcoxon two-sample test 35 5. The means and 95% confidence intervals of the dominant species and a comparison of the r e l a t i v e and absolute abundances in the zones and i n l e t types using the Wilcoxon two-sample test 40 6. Relative abundance in Zones A, B, and C of Chaetoceros spp. resting spores 48 7. The means and 95% confidence i n t e r v a l s , and a comparison of r e l a t i v e and absolute abundances,and valve diameter using the Wilcoxon two-sample test...52 8. The means and 95% confidence i n t e r v a l s , and a comparison of r e l a t i v e and absolute abundances, and valve diameter using the Wilcoxon two-sample test...53 9. The means and 95% confidence intervals of the marine planktonic, marine l i t t o r a l and freshwater species, and a comparison of the r e l a t i v e and absolute abundances in the zones and i n l e t types 56 10. PCoA results 65 1 1. Comparison of the absolute abundance of diatom valves in oceanic and coastal sediments 139 12. Morphological data for Thalassiosira p a c i f i c a and T. a e s t i v a l i s 193 v i L i s t of Figures 1. B r i t i s h Columbia coast and Zones A, B, and C 9 2. Zones A and B 10 3 . Zone C 11 4. Generalized s p a t i a l pattern of annual phytoplankton production in the S t r a i t of Georgia 17 5. Zone A s i t e s 20 6. Zone B s i t e s 21 7. Zone C s i t e s 22 8. Recurrent groups and the intergroup relationships. ..31 9. Div e r s i t y 34 10. Total absolute abundance 37 1 1 . Skeletonema costatum. . . 39 12. Thalassiosira a e s t i v a l i s f. p a c i f i c a 45 13. Chaetoceros spp. resting spores 46 14. Thalassionema nitzschioides 49 15. Paralia sulcata 51 16. Marine planktonic species 55 17. Marine l i t t o r a l species 59 18. Freshwater species 61 19. Scatter diagram of quadrants on coordinate axes 1 and 2 66 20. Position of i n l e t s on coordinate axes 1 and 2 67 21. Position of i n l e t s and s i t e s outside i n l e t s on coordinate axes 1 and 2 68 22. D i s t r i b u t i o n of areolae in 10 ixm for Thalassiosira  a e s t i v a l i s f. pac i f ica from 30 s i t e s representing the entire study area 82 23. D i s t r i b u t i o n of areolae in 10 nm for Thalassiosi ra  a e s t i v a l i s f. pac i f ica from 11 s i t e s along the length of Howe Sound ( 1 974) . 83 24. Absolute abundance of Skeletonema costatum and depth of i n l e t s i t e s in Zone A 108 25. Absolute abundance of Skeletonema costatum and depth of i n l e t s i t e s in Zone B 109 26. Absolute abundance of Skeletonema costatum and depth of i n l e t s i t e s in Zone C 110 27. Total absolute abundance and depth of i n l e t s i t e s in Zone A 111 28. Total absolute abundance and depth of i n l e t s i t e s in Zone B 112 29. Total absolute abundance and depth of i n l e t s i t e s in Zone C 113 30. Relative and absolute abundances of Paralia sulcata in Zone A s i t e s 129 31. Relative and absolute abundances of Paralia sulcata in Zone B s i t e s 130 32. Relative and absolute abundances of Paralia sulcata in Zone C s i t e s 131 33. Gradients of selected factors from Zone A to C 138 34. Relative and absolute abundances (±95% confidence v i i i ntervals) of dominant species in Zones A, B, and C..141 35. Diversity and r e l a t i v e abundance of Skeletonema  costatum in Zone A i n l e t s 143 36. Dive r s i t y and r e l a t i v e abundance of Skeletonema  costatum in Zone B i n l e t s 144 37. Dive r s i t y and r e l a t i v e abundance of Skeletonema costatum in Zone C i n l e t s 145 38. Gradients of selected factors and PCoA diagram 146 39. Gradients of selected factors and PCoA diagram 147 40. Gradients of selected factors in a t y p i c a l i n l e t . ..150 41. Relative abundance of freshwater species along the length of Howe Sound (1974) 156 42. Relative abundance of marine l i t t o r a l species along the length of Howe Sound (1974) 158 43. Relative and absolute abundances of dominant species along the length of Howe Sound (1974) 160 44. Gradients of primary productivity and planktonic diatom abundance, and t o t a l absolute abundance along the length of Howe Sound (1974) 162 45. Mean monthly radiation values for the Howe Sound area. 163 46. Mean monthly discharge values for the Squamish River.164 v i i i Ac knowledgements I wish to thank G. Roelofs for moral support, editing, and wordprocessing, and Dylan and Tait Roelofs for the i r patience with and understanding of their mother through six years of graduate school. I am grateful to Drs. F.J.R. Taylor, G. Rouse, S. Calvert, and P.J. Harrison for editing and constructive c r i t i c i s m . This study was supported by National Science and Engineering Research Council of Canada Grant A-6137 to Dr. F.J.R. Taylor and by a Chevron Fellowship in Oceanography to the author. 1 I . INTRODUCTION The purpose of this study i s to examine the d i s t r i b u t i o n a l patterns of diatom valves in the surface sediments of southern B r i t i s h Columbia i n l e t s with respect to oceanographic and hydrographic setting, and phytoplankton d i s t r i b u t i o n and productivity. Diatoms are sensitive to changes in physical and chemical properties in their environment, many have d e f i n i t e geographical ranges, and their s i l i c e o u s frustules allow some to be preserved as f o s s i l s in sediments (Werner 1977). Extensive studies of the d i s t r i b u t i o n a l patterns of diatom assemblages in the sediments of the deep oceans and coastal zones has led to the conclusion that these patterns coincide with nutrient supply, primary productivity, phytoplankton d i s t r i b u t i o n , and oceanographic patterns (Kanaya and Koizumi 1966; L i s i t z i n 1971; Maynard 1976; Defelice and Wise 1981; DeVries and Schrader 1981; Sancetta 1981; Schuette and Schrader 1981; Tanimura 1981). The sediments of estuaries, however, have received l i t t l e attention. A relati o n s h i p has been shown between s a l i n i t y of the surface waters and diatom sediment assemblages in the Hudson Estuary, New York (Weiss et a l . 1978), and a similar relationship holds for Paralia (Melosira) sulcata in the Bay of Vigo, Spain (Margalef 1969). Neither of these estuaries i s s t r i c t l y comparable to the type found in B r i t i s h Columbia and the subject of t h i s study. The rel a t i o n s h i p between the diatom thanatocoenoses and some factors of the world's Ocean (e.g., primary productivity, 2 areas of upwelling, annual production of s i l i c a ) i s well established. In some cases, the results of diatom analyses are further correlated with the d i s t r i b u t i o n of other mi c r o f o s s i l s (e.g., foraminifera, Kanaya and Koizumi 1966; s i l i c o f l a g e l l a t e s , Kozlova and Mukhina 1967), and amorphous s i l i c a d i s t r i b u t i o n both in the surface waters and the sediments (Maynard 1976). Investigations of the d i s t r i b u t i o n of diatoms over a large area generally deal with a broad range of environmental paramaters. Although some diatoms are cosmopolitan, i t i s possible to establish diatom assemblages with limited geographical ranges which can be correlated with oceanographic and bio/geographic factors. These correlations are p a r t i c u l a r i l y useful in determining zonal s h i f t s of discrete water masses related to climatic change (Kanaya and Koizumi 1966).Although the thanatocoenosis i s not a mirror image of the biocoenosis (a major problem in some studies), the large scale and concordant results from numerous studies gives some confidence in the results of sediment assemblages. The d i f f e r e n t i a l d i s s o l u t i o n of diatom frustules has been studied in the laboratory (Lewin 1961; G r i l l and Richards 1964; Johnson 1974; Mikkelsen 1977) and in the f i e l d (Kozlova and Mukhina 1967; Schrader 1971; Parker et a l . 1977). The general conclusion of the f i e l d studies is that the thanatocoenoses are not a complete r e f l e c t i o n of the biocoenoses because of the select i v e d i s s o l u t ion of weakly s i l i c i f i e d f r ustules. Some studies (Kanaya and Koizumi 1966; Kozlova and Muchina 1967; Jouse et a l . 1971; Maynard 1976; Schuette and Schrader 1981; 3 Tanimura 1981), which compare the abundance of particular species, have shown, however, that the thanatocoenbses are a good representation of the biocoenoses. The concept of d i f f e r e n t i a l d i s s o l u t i o n can be misleading. A diatom frustule is made up of two valves and a g i r d l e region, which can be d i f f e r e n t i a l l y s i l i c i f i e d . In general, the g i r d l e region i s not as heavily s i l i c i f i e d as the valves. The taxonomy of diatoms i s usually based on valve morphology, so that i f a valve i s preserved the species can be i d e n t i f i e d . In some species (e.g., Rhizosolenia spp. or Ditylum b r i g h t w e l l i i ), although the valves and g i r d l e region are both weakly s i l i c i f i e d , a heavily s i l i c i f i e d , i d e n t i f i a b l e valvar spine or central process can be preserved. Other species (e.g., Chaetoceros spp. ) form heavily s i l i c i f i e d resting spores. Therefore, although i t is true that some weakly s i l i c i f i e d species i d e n t i f i e d in the phytoplankton are never found in the sediments, i t should be understood that a l l frustules are undergoing dissolution and that even weakly s i l i c i f i e d frustules may have a heavily s i l i c i f i e d , i d e n t i f i a b l e part or c e l l stage ( i . e . , resting spores). It has been estimated that, as a result of d i f f e r e n t i a l d i s s o l u t i o n , no more than 5% of the t o t a l diatom frustule content of the top 100 m of the open ocean reaches the sediments (Kozlova and Mukhina 1967) and that the s e t t l i n g period in a water column of 5000 m ranges from 30 days to decades, depending on the size of the frustule ( L i s i t z i n 1971). The bulk of the s i l i c a from the skeletons of microplankton appears to be 4 redissolved in the upper 1000 m of the water column (Calvert 1968; L i s i t z i n 1971). Dissolution of the frustule continues in the bottom waters and pore waters of the sediments, which may both be undersaturated with respect to amorphous s i l i c a (Lewin 1961; Fanning and Schink 1969; Calvert 1974). In order to explain the presence of some diatom valves which normally should have dissolved in the water column above deep ocean sediments, a number of accelerated forms of downward transport have been suggested, including fecal p e l l e t transport, aggregation of c e l l s , density inversion currents, and downwelling (Smayda 1971). Smayda considered f e c a l p e l l e t transport, which may also protect frustules from d i f f e r e n t i a l d i s s o l u t i o n , the most important. Depending on the feeding strategy of the zooplankton, diatom frustules can be fragmented or l e f t whole (Johannes and Satomi 1966; Wimpenny 1973; Urrere and Knauer 1981). Dissolution of frustules can also occur within the gut of some zooplankton (Schrader 1971; Smayda 1971), and reingestion of fecal p e l l e t s by other grazers can continue the process of fragmentation and d i s s o l u t i o n (Smayda 1971). The r e l a t i v e advantage ( i . e . , protection from dissolution of some frustules) and disadvantage ( i . e . , fragmentation and dissolution of some frustules) of zooplankton ingestion and f e c a l p e l l e t transport appear to have a similar result to that of the ultimate e f f e c t of d i f f e r e n t i a l d i s s o l u t i o n in the water column and sediments: weakly s i l i c i f i e d components of the frustule are destroyed. Another possible protective mechanism, seldom mentioned, is 5 the presence of a "casing" on the frustule. In the laboratory, two types of casings have been shown to retard d i s s o l u t i o n (Lewin 1961; Johnson 1974). Both l i v i n g and k i l l e d c e l l s (heat-treated, treated with protein-denaturing agents or organic solvents, or mechanically broken) dissolved more slowly than acid-cleaned valves, suggesting that some form of organic casing existed (Lewin 1961; Volcani 1981). Once organic material i s removed from the valves, i t appears that certain adsorbed inorganics (e.g., Fe and Al) can retard dissolution (Lewin 1961; Johnson 1974). This inorganic casing can be removed by chelators or acids, after which the rate of d i s s o l u t i o n increases u n t i l the inorganic casing reforms (Johnson 1974). Although the effectiveness of these coatings has been shown in the laboratory, their r e l a t i v e importance under natural conditions i s unknown. For example, Fe adsorbed on the valve could be removed under reducing conditions in anaerobic marine sediments, or the organic casing could be removed by bacteria (Lewin 1961). Studies of the diatoms in the sediments are usually concerned with the r e l a t i o n s h i p between the biocoenoses and thanatocoenoses, d i f f e r e n t i a l d i s s o l u t i o n , and the e f f e c t s of oceanographic factors on d i s t r i b u t i o n a l patterns. However, the investigation of B r i t i s h Columbia estuaries includes factors not commonly encountered in oceanic studies such as high sedimentation rates from g l a c i a l l y fed r i v e r s , and a r e l a t i v e l y larger proportion of freshwater and marine l i t t o r a l species. The geographical range of comparison i s more r e s t r i c t e d but i t 6 i s p o s s i b l e to examine a wide range of e s t u a r i n e types from high to low r u n o f f . In order to i n t e r p r e t the d i s t r i b u t i o n a l p a t t e r n s and r e l a t e them to b i o l o g i c a l and p h y s i c a l f a c t o r s of the o v e r l y i n g waters, a number of q u e s t i o n s must be c o n s i d e r e d . Is d i f f e r e n t i a l d i s s o l u t i o n as important i n e s t u a r i e s as i n the oceans? Can the gross d i s t r i b u t i o n a l p a t t e r n s of freshwater and marine l i t t o r a l s p e c i e s be c o n s i d e r e d i n d i c a t i v e of t h e i r source? Is i t v a l i d to compare the a b s o l u t e abundance of diatoms in the su r f a c e sediments with primary p r o d u c t i v i t y of s u r f a c e waters, given the e f f e c t of d i f f e r e n t i a l sedimentation r a t e s ? Does e s t u a r i n e c i r c u l a t i o n s e r i o u s l y change the s p a t i a l r e l a t i o n s h i p between the biocoenoses and the thanatocoenoses at the s c a l e of t h i s study? An i n v e s t i g a t i o n of the diatom d i s t r i b u t i o n i n the sediments and i t s r e l a t i o n s h i p to such f a c t o r s as n u t r i e n t s , l i g h t , primary p r o d u c t i v i t y , and s a l i n i t y should be v a l u a b l e i n a s s i s t i n g p a l e o e c o l o g i c a l i n t e r p r e t a t i o n s , p a r t i c u l a r l y i n the r e c o n s t r u c t i o n of such parameters as c u r r e n t and upwelling p a t t e r n s , zones of phytoplankton abundance, r i v e r and e s t u a r i n e flows, and paleotemperatures. 7 II. PHYSICAL AND BIOLOGICAL SETTING PHYSICAL SETTING Estuaries can be c l a s s i f i e d by various c h a r a c t e r i s t i c s (Lauff 1967; Dyer 1973; Olausson and Cato 1980); however, the most ci t e d d e f i n i t i o n i s : "An estuary is a semi-enclosed body of water which has a free connection with the open sea and within which sea water i s measurably d i l u t e d with freshwater derived from land drainage" (Cameron and Pritchard 1963). A l l B r i t i s h Columbia i n l e t s are generally p o s i t i v e , i . e . , freshwater inflow, derived from river discharge and p r e c i p i t a t i o n , exceeds the loss by evaporation (Pickard 1961). Most B r i t i s h Columbia i n l e t s are topographically similar to fjords, representing drowned arms of the sea, of g l a c i a l o r i g i n , overdeepened lo n g i t u d i n a l l y and have deep U-shaped cross-sections (Carter 1934; Peacock 1935; Fairbridge 1980). The depth may increase from the head (the inland end of the i n l e t ) to the outer s i l l at the mouth (the seaward end of the i n l e t ) , or the i n l e t may be a series of basins each separated by a s i l l (Carter 1934). There are a number of i n l e t s in the B r i t i s h Columbia mainland coast and in Vancouver Island. These i n l e t s are variously and unsystematically named "sound", " i n l e t " , and "arm". For t h i s paper " i n l e t " w i l l be used as a generic term. Pickard's (1961) review summarized the oceanographic features of i n l e t s in the B r i t i s h Columbia mainland coast. This study includes nine southern B r i t i s h Columbia i n l e t s 8 and one Vancouver I s l a n d i n l e t ( F i g s . 1-3). There are a l s o a few samples from the S t r a i t of Georgia, Queen C h a r l o t t e S t r a i t , and B u r r a r d I n l e t . P i c k a r d (1961) c l a s s i f i e d B r i t i s h Columbia i n l e t s on the b a s i s of the s u r f a c e s a l i n i t y d i s t r i b u t i o n . T h i s c l a s s i f i c a t i o n , with the i n l e t s s t u d i e d here i d e n t i f i e d , i s as f o l l o w s : Group A. I n l e t s with a low s a l i n i t y at the head 1. "high runoff i n l e t s " : s u r f a c e s a l i n i t y at head 0.1 -2% 0 s u r f a c e s a l i n i t y at mouth 5-20% o Secchi d i s k depth at head 0.3-1 m Knight I n l e t , Bute I n l e t , Howe Sound, and Toba I n l e t 2. "medium runoff i n l e t s " : head 1 - 1 1 %«, mouth 15-29%O Secchi d i s k 4-7 m J e r v i s I n l e t , Indian Arm, and S e c h e l t I n l e t Group B. I n l e t s with r e l a t i v e l y high s u r f a c e s a l i n i t y "low runoff i n l e t s " : head 18-29% D mouth 20-3 1 % e Secchi d i s k 6-10 m P e n d r e l l Sound, Hotham Sound, and Saanich I n l e t 9 130 125 Figure 1. B r i t i s h Columbia coast and Zones A, B, and C. F i g u r e 2. Zones A and B. BI = Bu r r a r d I n l e t , HS = Hotham Sound, NI = Narrows I n l e t , Sa = Salmon I n l e t , Se = Sechelt I n l e t , and TI = Texada I s l a n d . Figure 3. Island. Zone C. DS Desolation Sound and TI = Texada 1 2 Pickard (1961) does not include Hotham Sound or Saanich Inlet in his review. S a l i n i t y data from other studies (Gross 1967; Stockner and C l i f f 1975; Takahashi et a l . 1977) j u s t i f y placing both in Group B. For t h i s discussion, "Sechelt Inlet " w i l l be used as a generic term for the combined system of Sechelt Inlet, Salmon Inlet, and Narrows Inlet (Fig. 2). There is no s i l l between Sechelt Inlet and Salmon Inlet but there i s a shallow s i l l (10 m) between Sechelt Inlet and Narrows Inlet. Other c h a r a c t e r i s t i c s accompany the c l a s s i f i c a t i o n given above. High runoff i n l e t s usually have runoff from g l a c i e r s or snowfields. Medium runoff i n l e t s have l i t t l e or no contribution from g l a c i e r s . Low runoff i n l e t s have no g l a c i a l runoff. The high concentration of suspended sediment derived from g l a c i a l runoff in the high runoff i n l e t s reduces the Secchi disk depth (transparency) and results in suppression of phytoplankton production through l i g h t attenuation at the heads of the i n l e t s (Stockner et a l . 1977). River runoff in the i n l e t s is seasonal. River discharge in high runoff i n l e t s comes from summer melting of snow and g l a c i e r s . The minimum runoff i s during January-March and the maximum runoff is during May-July, with a secondary maximum in October due to direct p r e c i p i t a t i o n . The medium and low runoff i n l e t s derive their discharge from direct rather than stored runoff. The minimum runoff i s in July-September and the maximum is during the spring and winter months following the coastal p r e c i p i t a t i o n pattern. The water column can be divided into two main depth zones, 13 shallow and deep. The shallow zone i s f r e s h (from runoff) to b r a c k i s h water. The deep waters have a s a l i n i t y s i m i l a r to that of the deep water of the S t r a i t of Georgia. The shallow zone can f u r t h e r be subdivided i n t o s u r f a c e , and intermediate or mixing zones. In general, i n the surface water, temperature and s a l i n i t y increase toward the mouth. The c i r c u l a t i o n and water c h a r a c t e r s i t i c s i n the i n l e t s are mainly c o n t r o l l e d by freshwater run o f f , t i d e s , and winds. The c i r c u l a t i o n p a t t e r n i s often a h i g h l y s t r a t i f i e d , two-layer flow with entrainment (Bowden 1980). The surface seaward moving lay e r of r i v e r water e n t r a i n s s a l t water thereby i n c r e a s i n g the s a l i n i t y , the volume, and the v e l o c i t y of the upper l a y e r as i t moves toward the mouth. To maintain a s a l t balance, there i s a compensating current which moves toward the head below the r i v e r water l a y e r . This e s t u a r i n e c i r c u l a t i o n can a l s o be dr i v e n by winds. . Tides are semi-diurnal with a marked i n e q u a l i t y between the height and time of successive low waters. T i d a l range increases from south to north along the coast (P i c k a r d 1961). Winds c o n t r i b u t e to surface currents and mixing. Upwelling, which i s u s u a l l y a wind-induced phenomenon, i s a l s o caused by freshwater runoff (Hoos and Packman 1974). Upwelling zones occur mainly at the mouths of i n l e t s , although they can a l s o occur w i t h i n the i n l e t s themselves (Hutchinson and Lucas 1931; P i c k a r d 1961; Stockner et a l . 1977). The B r i t i s h Columbia i n l e t s are s t r i k i n g l y s i m i l a r to the Norwegian f j o r d s (Sakshaug and Myklestad 1973; Braarud 1976) 14 both in terms of oceanographic c h a r a c t e r i s t i c s and phytoplankton species composition. Unlike some Norwegian fjords, however, the s i l l s at the mouths of the studied i n l e t s are not shallow enough to r e s t r i c t deep water c i r c u l a t i o n , except in Saanich Inlet which has periodic anoxia in the bottom waters (Pickard 1961; Herlinveaux 1972). In Norway, another factor contributiong to anoxia i s the fact that the t i d a l range is so small that there is no ' t i d a l pumping' of new water over the s i l l s . T h e estuaries along the western A t l a n t i c coast (Pratt 1959; S i r o i s and Fredrick 1978; Marshall 1980) and Puget Sound (Winter et a l . 1975) also have similar species composition. Both the S t r a i t of Georgia and Johnstone S t r a i t , which connects the Queen Charlotte S t r a i t to the S t r a i t of Georgia (Figs. 1 and 3), are considered estuarine-type systems (Thomson 1976 and 1981; Stockner et a l . 1979). In general, s a l i n i t y of the surface water increases from the Fraser River area to the north, and from east to west in the S t r a i t of Georgia (Tully and Dodimead 1957; Waldichuck 1957). BIOLOGICAL SETTING A number of studies of the benthic and planktonic marine diatoms for the S t r a i t of Georgia, Puget Sound and contiguous waters have been published (Peck and Harrington 1897; Bailey and MacKay 1916; Hutchinson 1928; Gran and Angst 1931; Hutchinson and Lucas 1931; Phifer 1934 a and b; Legare 1957; Shim 1976). Many of the i n l e t s in t h i s study have also been investigated 15 ( B u r r a r d I n l e t : S t o c k n e r and C l i f f 1979; Hotham Sound: S t o c k n e r and C l i f f 1975; Howe Sound: S t o c k n e r e t a l . 1977; I n d i a n Arm: B u c h a n a n 1966; J e r v i s I n l e t : A. G. L e w i s , p e r s o n a l c o m m u n i c a t i o n , 1982; P e n d r e l l S o und: S t o c k n e r and C l i f f 1975; S a a n i c h I n l e t : T a k a h a s h i e t a l . 1 9 7 7 ) . The d o m i n a n t s p e c i e s o f t h e s t u d y a r e a ( S k e l e t o n e m a  c o s t a t u m , T h a l a s s i o s i r a s p p . , a n d C h a e t o c e r o s s p p . ) a r e s i m i l a r t o t h o s e f o u n d i n many t e m p e r a t e c o a s t a l w a t e r s a n d e s t u a r i e s ( s e e r e v i e w i n Smayda 1 9 8 0 ) . I n g e n e r a l , t h e t e m p o r a l s e q u e n c e o f e v e n t s f o r p h y t o p l a n k t o n , s u m m a r i z e d by H a r r i s o n e t a l . ( i n p r e s s ) , i s a s f o l l o w s : 1) The s p r i n g b l o o m b e g i n s b e t w e e n M a r c h and A p r i l . D o m i n a n t s p e c i e s a r e S k e l e t o n e m a c o s t a t u m , and T h a l a s s i o s i r a s p p . , f o l l o w e d by C h a e t o c e r o s s p p . The d e c l i n e o f t h e s p r i n g b l o o m c a n s o m e t i m e s be a s s o c i a t e d w i t h n u t r i e n t l i m i t a t i o n a n d s t r a t i f i c a t i o n o f t h e w a t e r c o l u m n . 2) I n t h e f a l l , a s m a l l e r b l o o m c a n o c c u r . I n h i g h r u n o f f i n l e t s , t h e d e c l i n e o f t h e s p r i n g b l o o m c a n a l s o be a s s o c i a t e d w i t h t h e f r e s h e t o f r i v e r s w h i c h commences b e t w e e n May a n d J u n e ( S t o c k n e r e t a l . 1 9 7 7 ) . P h y t o p l a n k t o n a t t h e h e a d o f t h e i n l e t a r e e x p o r t e d by t h e s e a w a r d - m o v i n g , f r e s h w a t e r s u r f a c e l a y e r a n d c a n be l i g h t - l i m i t e d by h e a v y s e d i m e n t l o a d s . The amount o f p h y t o p l a n k t o n t r a n s p o r t e d t o w a r d t h e h e a d by s u b s u r f a c e c o m p e n s a t i n g c u r r e n t s may be u n a b l e t o r e p l e n i s h t h e s u r f a c e l o s s e s . I n t h e S t r a i t o f G e o r g i a , a r e a s o f h i g h p r i m a r y p r o d u c t i v i t y a n d s t a n d i n g s t o c k h a v e been l o c a t e d i n t h e s o u t h e r n a n d n o r t h e r n e n d s , a t t h e mouth o f i n l e t s , a n d a d j a c e n t 1 6 to the F r a s e r R i v e r plume ( H a r r i s o n et a l . i n p r e s s ) . In g e n e r a l , primary p r o d u c t i v i t y i n c r e a s e s southward (Stockner et a l . 1979; F i g . 4). The v a r i a t i o n s i n t o t a l primary p r o d u c t i v i t y and standing stock can be a s s o c i a t e d with a number of f a c t o r s : l i g h t , n u t r i e n t s , g r a z i n g , s a l i n i t y , and s t a b i l i t y . 1 7 Figure 4 . Generalized production in the S t r a i t et a l . 1979) . . s p a t i a l pattern of annual phytoplankton of Georgia. (Reprinted from Stockner 18 I I I . METHODS SAMPLING PROCEDURES The term "surface sediments" i s somewhat ambiguous. In deep ocean studies (Maynard 1976; Sancetta 1981; Tanimura 1981), surface samples are usually those taken from the upper 1 to 2 cm of cores or surface grabs. These samples can represent a time span of 500 years or more (Maynard 1976). In the present study, an attempt was made to sample a much thinner surface layer. In the sediments of the study area, i t was often possible to v i s u a l l y descriminate the surface material, which was a glossy, dark brown layer, approximately 1 to 2 mm thick. In contrast, the underlying sediments were a d u l l , reddish-brown. In the c o l l e c t i o n of 49 of the 95 samples (collected by the author, and Drs. S. Calvert and J . Taylor), an e f f o r t was made to obtain only the glossy, dark upper layer. Since i t was so thin, a spatula was used to scrape numerous areas from the available sediment top in the box corer or Shipek grab, in order to obtain at least 1 g of sediment. These samples were then frozen. See Table 1 and Figs. 5-7 for dates, c o l l e c t o r s , and s i t e locations. The remaining 46 samples used in t h i s study were c o l l e c t e d previously by other workers. P. Dobell co l l e c t e d 21 of the samples using a Smith-Mclntyre grab, from which she took a subsample of the upper few centimeters to examine the di s t r i b u t i o n of di n o f l a g e l l a t e cysts. The sample from Hidden Basin was obtained with a suction sampler and consisted of 19 Table 1 . L i s t of s i t e s , s i t e numbers, and c o l l e c t i o n dates. s i t e s i t e number date co l l e c t e d by Agamemnon Channel 77-78 1981 A. Roelofs Burrard Inlet 44 1980 Dr. J. Thompson Burrard Inlet 92 1 981 Dr. J. Taylor Bute Inlet 7 1 977 P. Dobell Bute Inlet 89-91 1981 Dr. S. Calvert Cordero Channel 20 1 977 P. Dobell Desolation Sound 81 1 981 Dr. S. Calvert East Gorge Harbor 1 5 1976 P. Dobell Heriot Bay 19 1 977 P. Dobell Hidden Basin 95 1 977 P. Dobell Homfray Channel 83-84 1981 Dr. S. Calvert Hotham Sound 74-76 1981 A. Roelofs Howe Sound 27-43 1974 Dr. J. Thompson 45,53-58 Howe Sound 6 1976 P. Dobell Howe Sound 1-5 1 977 P. Dobell Indian Arm 11-12 1976 P. Dobell Indian Arm 93-94 1981 Dr. J. Taylor Je r v i s Inlet 21-26 1980 Dr. S. Calvert Je r v i s Inlet 64-73 1981 A. Roelofs Knight Inlet 8-10 1 977 P. Dobell Malaspina S t r a i t 79 1981 A. Roelofs Narrows Inlet 52 1981 Dr. S. Calvert Pendrell Sound 13-14 1 977 P. Dobell Pryce Channel 82 1 981 Dr. S. Calvert Queen Charlotte S t r a i t 18 1 977 P. Dobell Saanich Inlet 59-63 1 981 A. Roelofs Salmon Inlet 50-51 1981 Dr. S. Calvert Sechelt Inlet 46-49 1 981 Dr. S. Calvert S u t i l Channel 80 1981 Dr. S. Calvert Texada Island 1 6-1 7 1 976 P. Dobell Toba Inlet 85-88 1981 Dr. S. Calvert 20 Figure 5 . Zone A s i t e s . BI = Burrard Inlet (see Table 1 c o l l e c t i o n data). 21 Figure 6 . Zone B s i t e s . AC = Agamemnon Channel, HB = Hidden Basin, MA = Malaspina S t r a i t , and TI = Texada Island (see Table 1 for c o l l e c t i o n data). 22 Figure 7. Zone C s i t e s . CH = Cordero Channel, DS = Desolation Sound, EG = East Gorge Harbor, HC = Homfray Channel, HR = Heriot Bay, PC = Pryce Channel, PS = Pendrell Sound, and SC = S u t i l Channel (see Table 1 for c o l l e c t i o n data). 23 unconsolidated plankton d e t r i t u s . These samples were stored at 4°C. The remaining 25 samples were collected by Dr. J . Thompson (Ocean and Aquatic Sciences, Patrica Bay, B r i t i s h Columbia ) using a Shipek grab. The subsample was obtained by removing v e r t i c a l sections to the same depth with a p l a s t i c scoop. These samples were frozen and later freeze dried, with the exception of the sample from Burrard Inlet which was simply sealed in a j a r . There i s l i t t l e or no question that a l l these samples represent Recent sediments, regardless of sampling technique. They were coll e c t e d over a period of seven years (Table 1); however, with one exception (Howe Sound 1974), no attempt could be made to discriminate time intervals in any samples. In this study "surface sediments" is used as a generic term to represent Recent sediments c o l l e c t e d between 1974 and 1981. CLEANING AND COUNTING Before cleaning each sample, the dry weight was determined (approximately 1 gram). The sample was then treated with 30% H 20 2 and K 2 C r 2 0 7 to oxidize organic material, washed and centrifuged fiv e times to remove clay p a r t i c l e s , and d i l u t e d with d i s t i l l e d water to 50 ml. A 5 ml aliquot was d i l u t e d to 50 ml and a known volume (usually 0.1 ml) was placed on a cover s l i p with a micropipette. The cover s l i p was allowed to a i r dry and was then permanently attached to a s l i d e with Hyrax mounting medium. Duplicate s l i d e s were made for each sample. For large-scale d i s t r i b u t i o n a l patterns, approximately 200 24 valves were i d e n t i f i e d and counted (100 from each duplicate slide) from each sample. To determine variation in valve diameter of Paralia sulcata, approximately 20 valves were measured at each s i t e . Counting was done at 1000X power using a Zeiss phase contrast microscope. Broken valves consisting of more than one-half of one valve were counted as a whole valve. TAXONOMY The taxonomy of the majority of diatom species in the northwest P a c i f i c coastal region i s well established. Major references used to identify species were Cupp (1943), Gran and Angst (1931), Hasle (1978), Hendey (1964), Hustedt (1927-1966, 1930, 1955), Patrick and Reimer (1966, 1975), and Riznyk (1973). See Appendix A for a complete species l i s t and Appendix VI for photomicrographs of selected species. However, there were problems involving the d i s t i n c t i o n of Thalassiosira p a c i f i c a and T. a e s t i v a l i s , and Chaetoceros spp. resting spores. Both T_^  paci f ica and T^ a e s t i v a l i s have been included in phytoplankton studies in the S t r a i t of Georgia region and contigious waters (Stockner and C l i f f 1975 and 1976; Shim 1976: Taylor and Waters 1982; Harrison et a l . in press). This author does not consider these two separate species, but rather two forms of one polymorphic species. A detailed explanation of why T. p a c i f i c a and T_j_ a e s t i v a l i s are considered conspecific here i s given in Appendix B. The name T\_ a e s t i v a l i s has p r i o r i t y , and i f the species were combined and both forms considered v a l i d , their names would be T\_ a e s t i v a l i s f. a e s t i v a l i s and T. a e s t i v a l i s f. p a c i f i c a . 25 Many n e r i t i c diatoms form resting spores. Some resting spores are morphologically similar to their vegetative c e l l s (e.g., T. nordenskioeldi i , Syvertsen 1979), while others are di f f e r e n t (e.g., Leptocylindrus danicus , Hargraves 1976). In B r i t i s h Columbia sediments, Chaetoceros spp. are represented almost e n t i r e l y by resting spores, which are usually of the l a t t e r type. Not a l l Chaetoceros spp. are known to form resting spores, nor is the percentage of vegetative c e l l s which form resting spores known in a given population. Although Gran and Angst (1931) l i s t 32 Chaetoceros taxa and Shim (1976) 26, only 10 Chaetoceros spp. (Table 2) could be i d e n t i f i e d from resting spores in the sediments. One of the main problems in ide n t i f y i n g Chaetoceros spp. resting spores disassociated from their vegetative c e l l s i s that whereas one valve may be unique, the other valve may have no distinguishing c h a r a c t e r i s t i c s (e.g., C_^  lorenzianus and diadema ). When the two valves are separated, only one can subsequently be i d e n t i f i e d . The undifferentiated valve can then only be placed with the unknown Chaetoceros spp. resting spores. The group designated here as "Chaetoceros spp. resting spores" consists of the undifferentiated valves together with rounded, unpatterned valves similar to the resting spores of C. compressus and C_^  s o c i a l i s . To avoid confusion, the unidentified Chaetoceros spp. resting spores group w i l l be written "Chaetoceros u-spp. " Species with only one i d e n t i f i a b l e valve were probably underestimated in rel a t i o n to those species where both valves were i d e n t i f i a b l e (e.g., C. radicans). Given 26 Table 2 . D i s t r i b u t i o n of Chaetoceros spp. r e s t i n g spores. The number f o l l o w i n g the s p e c i e s code r e f e r s to the t o t a l number of v a l v e s and the number i n parentheses i s the number of s i t e s at which the sp e c i e s o c c u r r e d . C. a f f i n i s 399 (61) C. vanheurcki i 497 (88) C. d e b i l i s 440 (81) C^ diadema 51 (20) C. g r a c i l i s 1 (1 ) <LL l a u d e r i 37 (19) Chaetoceros u-spp. 804 (72) C. ra d i c a n s 2618 (93) C. s e i r a c a n t h u s 4 (2) C. vanheurcki i 34 (13) C. didymus 96 (30) C. lorenzJ 1 aniJS 6 (4) 27 these taxonomic problems, the Chaetoceros spp. were lumped together for most s t a t i s t i c a l analyses. STATISTICAL ANALYSES Relative and absolute abundances were calculated for each sample. Community d i v e r s i t y s t a t i s t i c s based on information theory were determined using the Information index (Shannon and Weaver 1949): where H' i s expressed as b i t s / i n d i v i d u a l , n i s the number of valves of the i ' t h taxon, N i s the t o t a l number of valves in the sample, and S i s the t o t a l number of taxa in the sample. A low number indicates that one taxon dominated the assemblage, whereas a high number indicates a r e l a t i v e l y even d i s t r i b u t i o n of taxa. P r i n c i p a l coordinate analysis (PCoA), an ordination technique, was also used to analyze the data (Bradfield 1981). PCoA, a type of multivariate analysis, has been used in other studies on diatom d i s t r i b u t i o n a l patterns (Mclntire and Moore 1977; Schuette and Schrader 1981). Because the o r i g i n a l matrix had fewer s i t e s (95) than species (216), a PCoA program was used. Had there been fewer species than s i t e s , a p r i n c i p a l component analysis program would have been used. Gower (1966) 28 has shown that these two analyses give the same r e s u l t s . The PCoA program was i n i t i a l l y run using a matrix consisting of 95 sites and 216 species. However, to minimize sampling error (Imbrie and Kipp 1971; Maynard 1976; Schuette and Schrader 1981) a l l those species which did not occur in quantities greater than 2% were excluded, and a second analysis was done using a matrix consisting of 95 s i t e s and 8 species. Standardization was accomplished using the following s i m i l a r i t y index: where P i j and Pin are the proportions of the i ' t h taxon in the j'th and n'th samples respectively, and S i s the t o t a l number of taxa. If the two samples being compared had no taxa in common, SI has a value of 0; i f the taxa present and their r e l a t i v e abundance were i d e n t i c a l , SI has a value of 1. Recurrent groups were determined using the index of a f f i n i t y (Fager 1957; Fager and McGowan 1963): I A = [ j / ( H A * N B ) / L J - 'kHV^ } where J i s the number of joint occurrences, NA i s the t o t a l 29 number of occurrences of species A, NB is the t o t a l number of occurrences of species B, and species are assigned to the l e t t e r s so that NA < NB. Species pairs for which this expression was equal to or greater than 0.50 were considered to show a f f i n i t y . The nonparametric, two-tailed Wilcoxon two-sample test (Sokal and Rohlf 1969) was used to compare t o t a l valves/gram dry weight (absolute abundance), d i v e r s i t y (H'), d i n o f l a g e l l a t e cysts to diatom valves, and r e l a t i v e and absolute abundance of individual species and species groups. The n u l l hypothesis was that the two samples had been drawn from populations having the same d i s t r i b u t i o n . The s t a t i s t i c i s given by: i P., where n, i s the sample size of the larger sample and n 2 that of the smaller sample, and R i s the sum of the ranks of n 2. The s t a t i s t i c C was compared with n, n 2 -C and the larger quantity was used for the test s t a t i s t i c Us. The significance test i s : _ _ |fTT_ where Tj i s a function of the number of variates t i e d in the j'th group of t i e s . 30 IV. RESULTS This chapter i s divided into two major sections: 1) large-scale diatom d i s t r i b u t i o n a l patterns, and 2) diatom valve and di n o f l a g e l l a t e cyst d i s t r i b u t i o n . I. LARGE-SCALE DIATOM DISTRIBUTIONAL PATTERNS Of the 213 diatom taxa i d e n t i f i e d from the 95 s i t e s , 39 were planktonic, 81 marine l i t t o r a l , 73 freshwater, and 20 could only be i d e n t i f i e d to genera. Three s i l i c o f l a g e l l a t e s were also i d e n t i f i e d . Seventy-five species occurred at only one s i t e and a t o t a l of 139 occurred at four or fewer s i t e s . Only seven species (Skeletonema costatum, T. a e s t i v a l i s f. p a c i f i c a , Thalassionema nitzschioides , Paralia sulcata, Chaetoceros  radicans, C. vanheurcki i , and C. d e b i l i s ) had a count of greater than two percent of the t o t a l count (20,955 t o t a l valves). S. costatum made up 25% and T. a e s t i v a l i s f. p a c i f i c a 19% of the t o t a l count. The eleven Chaetoceros spp. and one group of unidentified Chaetoceros spp. resting spores (= Chaetoceros u-spp. ) represented 24% of the t o t a l valve count. Chaetoceros  a f f i n i s , C. radicans, C. vanheurcki i , and C. d e b i l i s , accounted for 85% of the t o t a l Chaetoceros spp. resting spore count and 20% of the t o t a l valve count. RECURRENT GROUP ANALYSIS Recurrent group analysis of the 216 species and resulted in one group (A) of 15 diatoms and one s i l i c o f l a g e l l a t e , and one group (B) of three diatom species (Fig. 8; Table 3). Additionally, the largest group had two associated species 31 Group A 15 species 18/51 Group B 3 species 6/34 2 Assoc. Figure 8. Recurrent groups and the intergroup relationships. The fractions are the ratios of observed species-pair a f f i n i t i e s between groups to maximum number of possible a f f i n i t i e s . 32 Table 3 . Species l i s t for recurrent Groups A and B. P = planktonic species, ML = marine l i t t o r a l species, I-V = Shim's (1976) recurrent planktonic groups, 1 = North P a c i f i c n e r i t i c recurrent group (Venrick 1971), and 2 = seasonal phytoplankton d i s t r i b u t i o n in Puget Sound, Washington (Phifer 1934a). Group A Chaetoceros a f f i n i s P, I, 2 (May-Oct.) C. vanheurcki i P,2 (May-Sept.) C. d e b i l i s P, II, 1, 2 (May-Nov.) C. radicans P, I, 1, 2 (May-Oct.) Cocconeis costata ML C y c l o t e l l a s t r i a t a ML Distephanus speculum ( s i l i c o f l a g e l l a t e ) Ditylum b r i g h t w e l l i i P, V, 1, 2, ( a l l year) Paralia sulcata P, ML, 2 ( a l l year) Rhizosolenia spp. P, 1, 2 ( a l l year) Skeletonema costatum P, I, 1, 2 (May-Nov.) Thalassionema nitzschioides P, I, 1, 2 (May-Nov.) Thalassiosira angstii P, 1, 2, ( a l l year) T. eccentrica P, IV, 1, 2 ( a l l year) T. nordenskioeldi i P, I I , 2 (April-July) 1' a e s t i v a l i s f. p a c i f i c a P, III, 2 (April-June) Associated species Cocconeis disculoides ML Chaetoceros didymus P, I, 1, 2 (May-Oct.) Group B Leptocylindrus danicus P, 1, 2 (May-Nov.) Synedra tabulata ML Thalassiosira anguste-1ineata P, III 33 ( i . e . , species not included in the group but having a f f i n i t i e s only with members of the group.) In Group A, S. costatum, Thalassiosira a e s t i v a l i s f. p a c i f i c a , and T. nitzschioides occurred at a l l 95 s i t e s and Chaetoceros radicans occurred at 94 s i t e s . The mean occurrence of the group i s 77.5±SD13.5 s i t e s . C y c l o t e l l a s t r i a t a and Cocconeis disculoides , an associated freshwater species, did not occur at the s i t e s on Toba, Knight or Bute Inlets ( a l l high runoff i n l e t s ) . The other associated species, Chaetoceros  didymus, did not occur at the s i t e s in Knight Inlet. Group A contained the seven species (S. costatum, T. a e s t i v a l i s f. p a c i f i c a , T. nitzschioides, P. sulcata, C. radicans, C. d e b i l i s , C. vanheurcki i) which had individual counts of greater than two percent of the t o t a l count (20,955 valves). Either S. costatum, T. a e s t i v a l i s f. p a c i f i c a , or a Chaetoceros spp. was dominant at a l l 95 sites except at the head of Pendrell Sound where P. sulcata was dominant. Group B consisted of two planktonic and one marine l i t t o r a l species. A l l three species were absent in Knight Inlet and had a mean occurrence of 37.3±SD7.0 s i t e s . DIVERSITY (Fig. 9 and Table 4) In northern Zones B and C, d i v e r s i t y was s i g n i f i c a n t l y greater than in Zone A; there was no s i g n i f i c a n t difference between Zones B and C. Medium and low runoff i n l e t s had s i g n i f i c a n t l y greater d i v e r s i t y than high runoff i n l e t s but there was no s i g n i f i c a n t difference between medium and low 34 Figure 9. D i v e r s i t y . Shaded zones and i n l e t s represent s i g n i f i c a n t l y greater d i v e r s i t y (see Table 4). H = high runoff i n l e t s , M = medium runoff i n l e t s , L = low runoff i n l e t s , and A, B, and C are zones. 35 Table 4 . The means and 95% confidence intervals of the t o t a l absolute abundance and d i v e r s i t y ( H * ) and a comparison of the zones and i n l e t s using the Wilcoxon two-sample te s t . H R I = high runoff i n l e t s , M R I = medium runoff i n l e t s , L R I = low runoff i n l e t s , A = Zone A , B Zone B , C = Zone C , and v/gdw = valves/gram dry weight. t o t a l absolute abundance: zones A=7.7X10 7±2.0X10 7 v/gdw A > B P<0.10 B=5 . 6X10 7±1.3X10 7 A > C P<0.01 C=3.8X10 7±1.4X10 7 B > C p<0.05 t o t a l absolute abundance: i n l e t s HRI= 6 .0x10 7±1.6X10 7 v/gdw H R I - M R I p>0.80 MRI=6.3X10 7±2.2X10 7 H R I - L R I p>0.80 L R I = 6 . 3 X 1 0 7 ± 2 . 6 X 1 0 7 M R I - L R I p>0.80 d i v e r s i t y : zones A=2.9±0.1 b i t s / i n d i v i d u a l A < B P<0.001 B=3.7±0.1 A < C P<0.001 C=3.5±0.2 B - C p>0.60 d i v e r s i t y : i n l e t s H R I = 3.0±0.1 b i t s / i n d i v i d u a l H R K M R I p<0.00l M R I = 3.5±0.1 H R K L R I p<0.05 LRI=3.3±0.3 M R I - L R I p>0.60 number of s i t e s : zones and i n l e t s A n=41 H R I n=41 B n=31 M R I n=27 C n=23 L R I n=!0 36 runoff i n l e t s . Within a l l the i n l e t s , there was no consistent pattern that could be related to head-mouth orientation. The Information index (H') ranged from 4.37 b i t s / i n d i v i d u a l in Heriot Bay to 1.90 b i t s / i n d i v i d u a l in Howe Sound (1977). RELATIVE AND ABSOLUTE ABUNDANCES Within the i n l e t s and among the zones and i n l e t types, r e l a t i v e and absolute abundances (v/gdw = valves/gram dry weight) showed d i s t i n c t patterns of increase and decrease. The patterns of r e l a t i v e and absolute abundances of the dominant species (S. costatum, T. a e s t i v a l i s f. p a c i f i c a , and Chaetoceros spp. ) were often the same and may seem redundant. However, both s t a t i s t i c s were needed for interpretation because of processes (e.g., varying sedimentation rates) which can af f e c t the sediment assemblages. Relative abundance, a q u a l i t a t i v e s t a t i s t i c , was probably only s l i g h t l y affected. However, absolute abundance, a quantitative s t a t i s t i c , was most l i k e l y affected and should be interpreted with caution. The effects of these processes w i l l be discussed l a t e r . Appendix D l i s t s the abundance of the selected species and species-groups, and t o t a l absolute abundance at each s i t e . The s i t e s in each i n l e t are ranked in head to mouth order. 1. Absolute Abundance Of Total Diatom Valves (Fig. 10 and Table 4) The absolute abundance was greatest in Zone A and least in Zone C. Among the i n l e t types, there was no s i g n i f i c a n t difference in absolute abundance of diatom valves between high, medium, or low runoff i n l e t s . In a l l the i n l e t s , absolute 37 Figure 10. Total absolute abundance. The shaded zone represents s i g n i f i c a n t l y greater absolute abundance (see Table 4). H = high runoff i n l e t s , M = medium runoff i n l e t s , L = low runoff i n l e t s , V/GDW = valves/gram dry weight (absolute abundance), and A, B, and C are zones. 38 abundance increased toward the mouth or remained r e l a t i v e l y constant along the i n l e t . The only exception was Saanich Inlet, a low runoff i n l e t , where absolute abundance increased toward the head. The values of absolute abundance ranged from 3.4X10 8 v/gdw in Howe Sound (1974) to 7.2x10s v/gdw in Heriot Bay. 2. Marine Planktonic Species There were 39 marine planktonic species, plus three s i l i c o f l a g e l l a t e s , which represented 83% of the t o t a l number of valves. This group contained a l l the dominant species except Paralia sulcata, which i s tychopelagic (Cupp 1943). A. Skeletonema costatum (Fig. 11 and Table 5) The high r e l a t i v e and absolute abundance of Skeletonema  costatum in the 1974 Howe Sound s i t e s could, in some cases, a l t e r the results of the s t a t i s t i c a l analyses. The unusually high abundance of S. costatum in the 1974 Howe Sound samples w i l l be discussed l a t e r . Relative and absolute abundances of Zone A and high runoff i n l e t s were calculated with and without the 1974 Howe Sound s i t e s . Both r e l a t i v e and absolute abundances of S. costatum were greatest in Zone A and least in Zone C regardless of the presence or absence of the 1974 Howe Sound s i t e s . With the Howe Sound s i t e s included, the r e l a t i v e abundance of S. costatum was s i g n i f i c a n t l y greater in high runoff i n l e t s than in medium or low runoff i n l e t s , with no s i g n i f i c a n t 39 Figure 1 1 . Skeletonema costatum. Shaded zones and i n l e t s represent s i g n i f i c a n t l y greater r e l a t i v e and absolute abundances (see Table 5 ) . % = r e l a t i v e abundance, V/GDW = valves/gram dry weight (absolute abundance), H = high runoff i n l e t s , M = medium runoff i n l e t s , and L = low runoff i n l e t s , and A, B, and C are i n l e t s . 40 Table 5 . The means and 95% confidence inte r v a l s of the dominant species and a comparison of the r e l a t i v e and absolute abundances in the zones and i n l e t types using the Wilcoxon two-sample t e s t . HRI = high runoff i n l e t s , MRI = medium runoff i n l e t s , LRI = low runoff i n l e t s , A = Zone A, B = Zone B, C = Zone C, and v/gdw = valves/gram dry weight. Skeletonema costatum r e l a t i v e abundance: zones A=36.8±3.5% B=20.2±1 .5 C=11.2±2.5 A>B A>C B>C re l a t i v e abundance without Howe Sound 1974 A=32.8±6.3% A>B A>C re l a t i v e abundance: HRI=31.5±4.5% MRI=24.2±4.4 LRI=21.7±17.8 r e l a t i v e abundance HRI=19.9±6.4% i n l e t s without Howe absolute abundance: zones A=3.0x10 7±8.2x1 0 6 v/gdw B=1.2X10 7±3.3X10 6 C = 4.2x!06±1 .4x10s HRI>MRI HRI>LRI MRI-LRI Sound 1974 HRI-MRI HRI-LRI A>B A>C B>C absolute abundance without Howe Sound 1974 A=2.9x10 7±1.8x10 7 v/gdw A>B A>C absolute abundance: i n l e t s HRI=2.1X10 7±5.2X10 6 v/gdw MRI=1.9x10 7±1.2x10 7 LRI=1.8x10 7±1.0x10 7 absolute abundance without Howe HRI=7.6x10 6±4.1x10 6 v/gdw HRI-MRI HRI-LRI MRI-LRI Sound 1974 HRKMRI HRKLRI P<0.001 P<0.001 P<0.001 P<0.001 P<0.001 p<0.05 p<0.05 p>0.60 p>0.40 p>0.80 P<0.001 P<0.001 p<0.001 p<0.10 p<0.01 p>0.80 p>0.80 p>0.80 p<0. 10 p<0.10 41 Table 5 continued. Thalassiosira a e s t i v a l i s f. p a c i f i c a r e l a t i v e abundance: zones A=23.4±3.0% B=13.6±2.2 C=21.2±5.2 A>B A-C B<C P<0.001 p>0.60 P<0.01 re l a t i v e abundance: i n l e t s HRI=25.7±3.4% MRI=15.0±2.7 LRI=18.4±0.6 HRI>MRI HRI>LRI MRI-LRI P<0.001 p<0.05 p>0.40 absolute abundance: zones A=1.5x10 7±3.8X10 6 v/gdw B=7.6X10 6±2.1X10 6 C=6.3X10 6±2.1x10 s absolute abundance: i n l e t s HRI=1.3x10 7±3.3x10 6 v/gdw MRI=9.8X10 6±4.3X10 6 LRI=8.7X10 6±3.9X10 6 Chaetoceros spp. A>B A>C B-C H R I - M R I H R I > L R I M R I - L R I P<0.001 P<0.001 p>0.40 p>0.40 P<0.10 p>0.80 rela t ive A= 1 3 B=33 C=27 abundance: ,9±2.1% ,3±2.9 .614.4 zones A<B A<C B>C P<0.001 P<0.001 p<0.05 re l a t i v e abundance: i n l e t s HRI=15.1±2.4% MRI=28.8±4.8 LRI=27.0±4.5 HRI<MRI HRKLRI MRI-LRI P<0.001 P<0.001 p>0.60 absolute abundance: zones A=1 . 1X10 7±3.4X10 6 v/gdw B=1 .8x10 7±4.3X10 6 C=1.3x10 7±5.8x10 s absolute abundance: i n l e t s HRI=9.3X10 6±2.9X10 6 v/gdw MRI=1.6X10 7±3.1x10 s LRI=1.8x10 7±7.8x10 6 A<B A-C B>C H R K M R I H R I < L R I M R I - L R I P<0.01 p>0.40 p<0.02 P<0.01 p<0.05 p>0.60 42 Table 5 continued Thalassionema nitzschioides r e l a t i v e abundance: zones A=7.3±0.9% A - B p>0.80 B=7.3±1.5 A - C p>0.80 C=7.6±1.7 B - C p>0.80 r e l a t i v e abundance: i n l e t s HRI=7.6±1.1% H R I - M R I p>0.60 MRI=6.9±1.5 H R I - L R I p>0.40 LRI=6.4±2.0 M R I - L R I p>0.80 absolute abundance: zones A=5.6x10 6±1.8x10 5 A > B P<0.10 B=4.1x10 6±1.9X10 5 A > C p<0.0l C=2.9X10 6±2.4X10 5 B > C p<0.05 absolute abundance: i n l e t s HRI=4.6x10 6±1.8x10 5 H R I - M R I p>0.80 MRI=4.3X10 6±3.3X10 5 H R I - L R I p>0.80 LRI=4.0X10 6±5.2X10 5 M R I - L R I p>0.80 number of s i t e s : zones and i n l e t s A n=41 H R I n=41 B n=31 M R I n=27 C n=23 L R I n=lO number of s i t e s without Howe Sound 1974 A n=17 HRI n=17 43 d i f f e r e n c e between medium or low runof f i n l e t s . But when the 1974 Howe Sound s i t e s were excluded, there was no s i g n i f i c a n t d i f f e r e n c e i n r e l a t i v e abundance between any of the i n l e t types. There was a l s o no s i g n i f i c a n t d i f f e r e n c e i n abs o l u t e abundance of S. costatum between any of the i n l e t types when the 1974 Howe Sound s i t e s were i n c l u d e d . But without the Howe Sound s i t e s , a b s o l u t e abundance was s i g n i f i c a n t l y greater i n medium and low runoff i n l e t s than i n high runoff i n l e t s . In the comparisons between i n l e t types, the r e s u l t s which excluded the Howe Sound (1974) data were probably more r e p r e s e n t a t i v e of the d i s t r i b u t i o n a l p a t t e r n of S. costatum. The r e l a t i v e l y lower abundances of S. costatum i n the three Zone C high runoff i n l e t s were being s t a t i s t i c a l l y enhanced by the Howe Sound (1974) data. T h e r e f o r e , the i n l e t type d i s t r i b u t i o n a l p a t t e r n s f o r S. costatum i l l u s t r a t e d i n F i g . 11 were based on s t a t i s t i c a l comparisons which excluded the Howe Sound (1974) data. Within the i n l e t s , the p a t t e r n of S. costatum d i s t r i b u t i o n v a r i e d . In the low runoff i n l e t s , a b s o l u t e abundance i n c r e a s e d toward the mouth i n P e n d r e l l Sound and Hotham Sound, but toward the head i n Saanich I n l e t . In the medium runoff i n l e t s , a b s o l u t e abundance i n c r e a s e d toward the mouth i n Indian Arm and the S e c h e l t I n l e t system, although the hi g h e s t r e l a t i v e abundance was at the head.of Narrows I n l e t . In the 1980 J e r v i s I n l e t s i t e s , a b s o l u t e abundance i n c r e a s e d toward the mouth but remained r e l a t i v e l y constant i n the 1981 s i t e s . In the high runoff i n l e t s , the p a t t e r n v a r i e d f o r each i n l e t . The value s 44 fo r r e l a t i v e and absolute abundances f o r S. costatum ranged from 59.5% i n Howe Sound (1974) to 1.3% i n P e n d r e l l Sound, and 1.5x10 s v/gdw i n Indian Arm (1981) to 2.5x10" v/gdw i n P e n d r e l l Sound, r e s p e c t i v e l y . B. T h a l a s s i o s i ra a e s t i v a l i s f . pac i f i c a ( F i g . 12 and Table 5) The r e l a t i v e abundance of T. a e s t i v a l i s f . p a c i f i c a was s i g n i f i c a n t l y g r e a t e r i n Zones A and C than i n Zone B, while the ab s o l u t e abundance was s i g n i f i c a n t l y g r e a t e r i n Zone A than i n Zones B or C, with no s i g n i f i c a n t d i f f e r e n c e between Zones B and C. The r e l a t i v e abundance i n high runoff i n l e t s was s i g n i f i c a n t l y g r e a t e r than i n low or medium runoff i n l e t s . The a b s o l u t e abundance was gr e a t e r i n high than low runoff i n l e t s . W i t hin the i n l e t s , the abs o l u t e abundance of T. a e s t i v a l i s f . pac i f i c a i n c r e a s e d toward the mouth i n low runoff i n l e t s , except i n Saanich I n l e t . In the medium runo f f i n l e t s , a b s o l u t e abundance i n c r e a s e d toward the mouth in S e c h e l t I n l e t and Indian Arm (1976), remained r e l a t i v e l y constant i n J e r v i s I n l e t (1980 and 1981), but in c r e a s e d toward the head i n Indian Arm (1981). In the high runoff i n l e t s , r e l a t i v e abundance was hig h e s t at the head except i n Howe Sound (1974), but the p a t t e r n of the a b s o l u t e abundance v a r i e d . The values of r e l a t i v e and abs o l u t e abundance f o r T. a e s t i v a l i s f . pac i f i c a ranged from 59.1% i n Howe Sound (1977) and Knight I n l e t to 4.0% i n P e n d r e l l Sound, and 7.3x10 7 v/gdw i n Howe Sound (1974) to 7.6x10" v/gdw i n P e n d r e l l Sound, r e s p e c t i v e l y . 45 Figure 12. Thalassiosira a e s t i v a l i s f. p a c i f i c a . Shaded zones and i n l e t s represent s i g n i f i c a n t l y greater r e l a t i v e and absolute abundances (see Table 5). % = r e l a t i v e abundance, V/GDW = valves/gram dry weight (absolute abundance), H = high runoff i n l e t s , M = medium runoff i n l e t s , and L = low runoff i n l e t s , and A, B, and C are i n l e t s . 46 Figure 13. Chaetoceros spp. resting spores. Shaded zones and i n l e t s represent s i g n i f i c a n t l y greater r e l a t i v e and absolute abundances (see Table 5). % r e l a t i v e abundance, V/GDW = valves/gram dry weight (absolute abundance), H = high runoff i n l e t s , M = medium runoff i n l e t s , and L = low runoff i n l e t s , and A, B, and C are i n l e t s . 47 C. Chaetoceros spp. resting spores (Fig. 13 and Table 5) The Chaetoceros spp. were lumped together for some s t a t i s t i c a l analyses. As a group the re l a t i v e abundance of Chaetoceros spp. was greatest in Zone B and least in Zone A, while the absolute abundance was s i g n i f i c a n t l y greater in Zone B than in Zones C or A. Both the r e l a t i v e and absolute abundances were s i g n i f i c a n t l y greater in medium and low runoff i n l e t s than in high runoff i n l e t s . In low runoff i n l e t s , absolute abundance increased toward the mouth in Pendrell Sound and Hotham Sound but toward the head in Saanich Inlet. In medium and high runoff i n l e t s , absolute abundance usually increased toward the mouth. The values of r e l a t i v e and absolute abundances for Chaetoceros spp. ranged from 52.9% in Jervis Inlet (1980) to 1.9% in Howe Sound ( 1977), and 7.2x107 v/gdw in Hidden Basin to 1 . 1x10 5 v/gdw in Knight Inlet, respectively. The fiv e Chaetoceros spp. and the Chaetoceros u-spp. , a l l of which had a valve count of greater than 2% of the t o t a l Chaetoceros spp. count, had similar d i s t r i b u t i o n a l patterns in the three zones (Table 6). C. radicans, C. d e b i l i s and C. vanheurcki i tended to increase in r e l a t i v e abundance toward the mouth, whereas C. a f f i n i s tended to increase toward the head. C. didymus had a l o c a l i z e d d i s t r i b u t i o n a l pattern: i t was only found at the head in Pendrell Sound and Toba Inlet, and only at the middle and/or mouth in Hotham Sound, Howe Sound, and Jerv i s Inlet. D. Thalassionema nitzschioides (Fig. 14 and Table 5) 48 Table 6 . Relative abundance in Zones A, B, and C of Chaetoceros spp. resting spores with a valve count of >2% of the t o t a l Chaetoceros spp. resting spore count. A B C Chaetoceros a f f i n i s 11% 78% 1 1% Chaetoceros vanheurcki i 28 51 21 Chaetoceros d e b i l i s 17 54 29 Chaetoceros u-spp. 35 40 25 Chaetoceros radicans 26 44 30 Chaetoceros didymus 16 58 26 x±SD 22±8% 54±13% 24±7% 49 Figure 14. Thalassionema nitzschioides. Shaded zones and i n l e t s represent s i g n i f i c a n t l y greater r e l a t i v e and absolute abundances (see Table 5). % = r e l a t i v e abundance, V/GDW = valves/gram dry weight (absolute abundance), H = high runoff i n l e t s , M = medium runoff i n l e t s , and L = low runoff i n l e t s , and A, B, and C are i n l e t s . 50 The absolute abundance was greatest in Zone A and least in Zone C; however, there was no s i g n i f i c a n t d i s t r i b u t i o n a l pattern in the i n l e t types or within the i n l e t s for T. nitzschioides, one of seven species with a count of ^ 2% of the t o t a l count. The values of re l a t i v e and absolute abundances for t h i s species ranged from 17.8% in J e r v i s Inlet (1980) to 0.9% in Howe Sound (1974), and 4.7X107 valves/gram dry weight in Howe Sound (1974) to 4.7X10 3 valves/gram dry weight in Heriot Bay, respectively. E. Paralia sulcata (Fig. 15 and Tables 7-8) The greatest absolute abundance of P. sulcata was in high and low runoff i n l e t s . The greatest r e l a t i v e abundance was in Zone C and low runoff i n l e t s , the least in medium runoff i n l e t s . Within the i n l e t s , both r e l a t i v e and absolute abundances increased toward the mouth in high and medium runoff i n l e t s . In two of the low runoff i n l e t s (Pendrell Sound and Hotham Sound), r e l a t i v e abundance increased toward the head, while absolute abundance showed l i t t l e change from head to mouth. In the t h i r d low runoff i n l e t (Saanich I n l e t ) , r e l a t i v e abundance was highest at the head followed by an abrupt decrease after which i t increased toward the mouth; absolute abundance increased toward the head. In a l l medium and high runoff i n l e t s , P. sulcata constituted less than 1% of the t o t a l valves or was absent at the head. Excluding the 11 si t e s where P. sulcata was absent, the values of re l a t i v e and absolute abundances ranged from 0.4% in J e r v i s Inlet, Sechelt Inlet, and Indian Arm to 64.7% in Pendrell Sound, and 7.2x103 valves/gram dry weight in Heriot Bay 51 KNIGHT -H 7o V/GDW H 3.9 3.0-105 M 1.9 8.7x105 L 11.9 3.3*106 T O B A - H •PENDRELL- L J E R V I S - M CHELT-M HOWE H v.. V/GDW c 10.1 2.8*106 B 3.0 1.6 A 3.5 3.5 INDIAN-% = V/GDW SAANICH-L Figure 15. Paralia sulcata. Shaded zones and i n l e t s represent s i g n i f i c a n t l y greater r e l a t i v e and absolute abundances (see Table 5). % = re l a t i v e abundance, V/GDW = valves/gram dry weight (absolute abundance), H = high runoff i n l e t s , M = medium runoff i n l e t s , and L = low runoff i n l e t s , and A, B, and C are i n l e t s . 52 Table 7 . The means and 95% confidence i n t e r v a l s , and a comparison of r e l a t i v e and absolute abundances,and valve diameter using the Wilcoxon two-sample test. HRI = high runoff i n l e t s , MRI = medium runoff i n l e t s , LRI = low runoff i n l e t s , and LRI* = low runoff i n l e t s without the r e l a t i v e abundance (64.7%) of P. sulcata at the head of Pendrell Sound. Inlets Mean Significance Relative abundance HRI x=3.9±0.8% MRI x=1.9±1.3% LRI x=11.9±11.7% LRI* x=6.0±2.5% Comparison HRI>MRI p<0.0l HRI<LRI P<0.10 MRI<LRI P<0.10 HRKLRI* p<0.lO MRI<LRI* P<0.01 Absolute abundance HRI x=3.0x10 6±1.7x10 s v/gdw MRI X=8.7X10 5±4.0X10 5 v/gdw LRI x=3.3x10 6±1.3x10 s v/gdw Comparison HRI>MRI p<0.02 HRI-LRI p>0.80 MRI<LRI P<0.01 Valve diameters HRI x=12.5±0.3 Mm MRI x=12.7±0.6 Mm LRI x=13.7±0.9 Mm Comparison HRI-MRI p>0.60 HRI<LRI P<0.01 MRI<LRI p<0.05 Number of s i t e s : r e l a t i v e and absolute abundance (valve diameter) HRI n=37 (32) MRI n=l9 (8) LRI n=!0 (10) 53 Table 8 . The means and 95% confidence i n t e r v a l s , and a comparison of r e l a t i v e and absolute abundances, and valve diameter using the Wilcoxon two-sample test. A = Zone A, B = Zone B, C = Zone C, and C* = Zone C without the r e l a t i v e abundance (64.7%) of P. sulcata at the head of Pendrell Sound. Zones Mean Significance Relative abundance A x=3.5±0.7% B x=3.0±0.8% C x=10.1±6.2% C* x=7.3±2.8% Comparison A-B p>0.60 A<C p<0.05 B<C p<0.05 A<C* p<0.02 B<C* P<0.01 Absolute abundance A x=3.5x10 6±1.9x10 s v/gdw B x=1.6x10 6±7.4x10 5 v/gdw C x=2.8x10 6±1.3x10 6 v/gdw Comparison A>B p<0.02 A-C p>0.60 B-C p>0.20 Valve diameter A x= 1 2. 6±0 . 3 | im B x=12.9±0.7 fxm C x=12.9±0.6 /nm Comparison A-B p>0.60 A-C p>0.60 B-C p>0.80 Number of s i t e s : r e l a t i v e and absolute abundance (valve diameter) A n=39 (33) B n=24 (13) C n=20 (18) 54 to 3.3X10 7 valves/gram dry weight in Howe Sound (1974), respect i v e l y . A comparison of valve diameter showed that there was no si g n i f i c a n t difference between the zones, while valve diameter was s i g n i f i c a n t l y larger in low runoff i n l e t s . The largest mean diameters (>14 Mm) were found at the mouth of Je r v i s Inlet, the heads of Saanich Inlet and Pendrell Sound, in Desolation Sound, and in the outer basin of Howe Sound. The smallest mean diameters (^ 11 ixm) were at the heads of Howe Sound and Bute In l e t . F. Other planktonic species (Fig. 16 and Table 9) The remaining 28 marine planktonic species represented 9% of the t o t a l valve count and 11% of the planktonic species valve count. This group included three s i l i c o f l a g e l l a t e s (Dictyocha  f ibula, Dictyocha speculum and Ebria t r i p a r t i t a ) . Twelve of the species occurred at four or fewer s i t e s . Six species (Dictyocha  speculum, D i t y1um b r i g h t w e l l i i , Leptocylindrus danicus, Thalassiosira angsti i , T. eccentr ica, and T. nordenskioeldi i) had a t o t a l count of greater than 100 valves. These six species were di s t r i b u t e d throughout a l l three zones. See Appendix C for d i s t r i b u t i o n of individual species. The marine planktonic species, excluding the major dominants (S. costatum, T. a e s t i v a l i s f. p a c i f i c a , and Chaetoceros spp. ), had greatest r e l a t i v e and absolute abundances in Zone B. The r e l a t i v e and absolute abundances in medium runoff i n l e t s were s i g n i f i c a n t l y greater than in low 55 Figure 16. Marine planktonic species. Shaded zones and i n l e t s represent s i g n i f i c a n t l y greater r e l a t i v e and absolute abundances (see Table 9). % = re l a t i v e abundance, V/GDW = valves/gram dry weight (absolute abundance), H = high runoff i n l e t s , M = medium runoff i n l e t s , and L = low runoff i n l e t s , and A, B, and C are i n l e t s . 56 Table The means and 95% confidence intervals of the marine planktonic, marine l i t t o r a l and freshwater species, and a comparison of the r e l a t i v e and absolute abundances in the zones and i n l e t types using the Wilcoxon two-sample te s t . HRI = high runoff i n l e t s , MRI = medium runoff i n l e t s . LRI = low runoff i n l e t s , A = Zone A , B = Zone B , C = Zone C , and v/gdw = valves/gram dry weight. Marine Planktonic Species Excluding Dominant Species (Table 5) r e l a t i v e abundance: A=9.1±1.1% B=22.5±1.9 C=16.5±2.5 zones A < B A < C B > C P<0.001 p<0.001 P<0.001 re l a t i v e abundance: HRI=10.4±1.4% MRI=21.2±2.3 LRI=14.8±5.3 i n l e t s HRKMRI HRI-LRI MRI>LRI p<0.001 p>0.20 p<0.02 absolute abundance: zones A=6.9X10 6±2.1X10 6 v/gdw B=1.3X10 7±3.4X10 6 C=6.8x1u e±2.9xl0 6 A < B A - C B > C P<0.001 P<0.01 absolute abundance: i n l e t s H=5.9x10 6±1.7x10 6 v/gdw M=1 .2X10 7±2.8X10 6 L=8.6X10 6±4.1X10 6 HRI<MRI HRI-LRI MRI>LRI P<0.001 p<0.10 Mar ine L i t t o r a l Spec ies r e l a t i v e abundance: zones A=3.0±0.7% B=3.8±0.8 C=6.1±2.1 A - B A < C B < C p>0.20 P<0.01 p<0.05 r e l a t i v e abundance: HRI=3.2±1.0% MRI=2.9±0.6 LRI=4.7±1.8 i n l e t s HRI-MRI HRI-LRI MRKLRI p>0.80 p>0.20 p<0.10 absolute abundance: zones A=2.7X10 6±1.2x10 s v/gdw B=2.4X10 6±1.3X10 6 C=2.5X10 6±1.4x10 s A - B A - C B - C absolute abundance: i n l e t s H=2.0x10 6±8.7X10 5 v/gdw M=2.2X10 6±1.2x10 s L=2.1X10 6±6.8X10 5 HRI-MRI HRI-LRI MRI-LRI 57 Table 9 continued Freshwater Species r e l a t i v e abundance: zones A=3.1±1.0% B=1.6±0.4 C=2.5±1.1 r e l a t i v e abundance: i n l e t s HRI=3.1±0.8% MRI=1.5±0.4 LRI=1.4±0.5 A>B A-C B<C H R I > M R I H R I > L R I M R I - L R I P<0.01 p>0.60 p<0. 10 p<0.00.1 P<0.001 p>0.80 absolute abundance: zones A=2.1x10 6±5.7x10 5 v/gdw B=1.0x10 6±5.5x10 s C=7.2x10 5±2.7x10 5 A>B A>C B-C P<0.01 P<0.01 absolute abundance; i n l e t s H=1 .8x10 6±5.6x10 5 v/gdw M=9.5X10 5±4.0X10 5 L=9.1x10 5±4.3X10 5 H R I > M R I H R I > L R I M R I - L R I p<0.02 p<0.10 number of s i t e s : spec ies) A n=41 (35) B n = 31 (31 ) C n=23 (22) marine planktonic species (marine l i t t o r a l H R I M R I L R I n = 41 n = 27 n=1 0 (34) (27) (10) number of s i t e s : A n = 38 B n = 25 C n=l9 freshwater H R I n=37 M R I n=22 L R I n=8 58 runoff i n l e t s , with no difference between high and low runoff i n l e t s . Within the i n l e t s , there was no consistent head-mouth orientation. The values of r e l a t i v e and absolute abundances ranged from 32% in Sechelt Inlet to 3.7% in Howe Sound (1974), and 5.5X10 7 valves/gram dry weight in Hidden Basin to 1.9x1U5 valves/gram dry weight in Heriot Bay. 3. Marine L i t t o r a l Species (Fig. 17 and Table 9) There were 81 marine l i t t o r a l species i d e n t i f i e d which represented 10% of the t o t a l valve count. Paralia sulcata represented 44% of the t o t a l marine l i t t o r a l species count. P. sulcata was not included in the following s t a t i s t i c a l analyses. Thirty-one species (40% of the species) only occurred once, and a t o t a l of 53 species (65% of the species) occurred at four or fewer s i t e s . Seven of the 95 s i t e s had no marine l i t t o r a l species. Three species ( Cocconeis costata, Synedra  tabulata, and C y c l o t e l l a s t r i a t a ) had a t o t a l count of greater than 100 valves. These species were found in a l l three zones, but C_^  s t r i a t a was not found in Knight Inl e t , Bute Inlet, or Toba Inl e t . See Appendix C for d i s t r i b u t i o n of individual spec ies . The r e l a t i v e abundance was s i g n i f i c a n t l y greater in Zone C than in either Zones A or B, with no s i g n i f i c a n t difference between of Zones A and B. There was no s i g n i f i c a n t difference in absolute abundances between the zones or i n l e t types. The values for r e l a t i v e and absolute abundances ranged from 18.5% in East Gorge Harbor to 0.4% in Howe Sound (1977) and Indian Arm 59 7. V/GDW Figure 17. Marine l i t t o r a l species. Shaded zones and i n l e t s represent s i g n i f i c a n t l y greater r e l a t i v e and absolute abundances (see Table 9).% = r e l a t i v e abundance, V/GDW = valves/gram dry weight (absolute abundance), H = high runoff i n l e t s , M = medium runoff i n l e t s , and L = low runoff i n l e t s , and A, B, and C are i n l e t s . 60 (1976), and 2.2X10 7 valves/gram dry weight in Hidden Basin to 4.4x10" valves/gram dry weight in Knight Inlet. 4. Freshwater Species (Fig. 18 and Table 9) There were 73 freshwater species i d e n t i f i e d which represented 2% of the t o t a l valve count. Thirty-eight species (52% of the species) only occurred once, and a t o t a l of 55 species (75% of the species) occurred at four or fewer s i t e s . Thirteen of the 95 s i t e s had no freshwater species. Six species (C y c l o t e l l a o c e l l a t a , Diploneis s m i t h i i , Eunotia p e c t i n a l i s v. minor, F r a g i l a r i a pinnata, Gomphonema olivaceum, and T a b e l l a r i a  fenestrata) had a t o t a l count of greater than 20 valves. A l l six species were di s t r i b u t e d throughout the three zones except C. o c e l l a t a and G_^  olivaceum which were only found in Zones A and B. See Appendix C for d i s t r i b u t i o n of individual species. The r e l a t i v e abundance was s i g n i f i c a n t l y greater in both Zones A and C than in Zone B, with no s i g n i f i c a n t differences between of Zones A and C; absolute abundance was greatest in Zone A. As might be expected, the r e l a t i v e and absolute abundances in high runoff i n l e t s were s i g n i f i c a n t l y greater than in medium or low runoff i n l e t s , with no s i g n i f i c a n t difference between medium and low runoff i n l e t s . Within the i n l e t s , the r e l a t i v e abundance was greater near the freshwater source. The values of r e l a t i v e and absolute abundances ranged from 13.3% in Burrard Inlet to 0.4% in Sechelt Inlet, East Gorge Harbor, S u t i l Channel, and Howe Sound (1974), and 7.4x10s valves/gram dry 61 Figure 18. Freshwater species. Shaded zones and i n l e t s represent s i g n i f i c a n t l y greater r e l a t i v e and absolute abundances (see Table 9) % = r e l a t i v e abundance, V/GDW = valves/gram dry weight (absolute abundance), H = high runoff i n l e t s , M = medium runoff i n l e t s , and L = low runoff i n l e t s , and A, B, and C are i n l e t s . 62 weight in Hidden Basin to 1.7x10" valves/gram dry weight in Queen Charlotte S t r a i t . 5. Summary Of Major D i s t r i b u t i o n a l Patterns A. Dive r s i t y D i v e r s i t y in Zones B and C was greater than in Zone A. Medium and low runoff i n l e t s had greater d i v e r s i t y than high runoff i n l e t s . B. Absolute abundance of t o t a l diatom valves Zone A had the greatest absolute abundance of diatom valves and Zone C the le a s t . There was no difference between i n l e t types. C. Marine planktonic species 1) Skeletonema costatum Both r e l a t i v e and absolute abundances were greatest in Zone A and least in Zone C. Between i n l e t types, the results varied depending on whether or not the 1974 Howe Sound data were included. 2) Thalassiosira a e s t i v a l i s f. p a c i f i c a Both r e l a t i v e and absolute abundances were greatest in Zone A and r e l a t i v e abundance was greatest in high runoff i n l e t s . 3) Chaetoceros spp. Both r e l a t i v e and absolute abundances were greatest in Zone B and in medium and low runoff i n l e t s . 4) Thalassionema nitzschioides The absolute abundance was greatest in Zone A and least in Zone 63 C. 5) Paralia sulcata The greatest absolute abundance was in high and low runoff i n l e t s . The greatest r e l a t i v e abundance was in Zone C and low runoff i n l e t s , the least in medium runoff i n l e t s . Both r e l a t i v e and absolute abundances increased toward the mouth in high and medium runoff i n l e t s . The largest valve diameters were in low runoff i n l e t s . 6) Other planktonic species The r e l a t i v e and absolute abundances were greatest in Zone B and medium runoff i n l e t s . D. Marine l i t t o r a l species The r e l a t i v e abundance was greatest in Zone C and greater in low than medium runoff i n l e t s . E. Freshwater species The r e l a t i v e abundance was greater in Zones A and C than in Zone B, while absolute abundance was greatest in Zone A. Both r e l a t i v e and absolute abundances were greatest in high runoff i n l e t s . PRINCIPAL COORDINATE ANALYSIS The PCoA was run twice, once with a matrix of 216 species and 95 s i t e s , and once with seven species and the Chaetoceros u-spp. , a l l of which had counts of ^2% of the t o t a l count. The re s u l t i n g scatter diagrams from the two analyses were v i r t u a l l y i d e n t i c a l . 64 The ordination of the PCoA r e f l e c t s r e l a t i v e rather than absolute abundance as a result of standardization using the s i m i l a r i t y index. The f i r s t component explained 44% of the variance, and the second, 30%. Table 10 gives the ranking of the selected species and Chaetoceros u-spp. for the f i r s t two axes. Figure 19 shows the location of each s i t e on the graph (see Appendix E for species count at each s i t e ) . Figures 20 and 21 show the location of the i n l e t s and individual s i t e s outside the i n l e t s . Because a s i m i l a r i t y index was used to standardize the data, adjacent s i t e s are similar in terms of presence and r e l a t i v e abundance of species. Sites from Zone A tend to be on the l e f t half of the graph, while those from Zones B and C tend to be on the r i g h t . Factors (e.g., primary productivity and s a l i n i t y ) associated with the ordination pattern w i l l be discussed l a t e r . II . DIATOM VALVE AND DINOFLAGELLATE CYST DISTRIBUTION Twenty-one of the samples used in this study were co l l e c t e d by P. Dobell (1978) during a study on d i n o f l a g e l l a t e cyst d i s t r i b u t i o n from Recent marine sediments of B r i t i s h Columbia . She noted that the gradient of cysts/gram dry weight (absolute abundance) increased toward the mouth of the i n l e t s but did not s t a t i s t i c a l l y analyze the trends. She further noted that Operculodinium centrocarpum cysts (the dormant phase of Gonyaulax grindleyi ) were dominant at the majority of her 45 s i t e s and S p i n i f e r i t e s spp. cysts (the dormant phase of various Gonyaulax species) were dominant in the S t r a i t of Georgia s i t e s . 65 Table 10 . PCoA res u l t s . Ranking of product moment corr e l a t i o n c o e f f i c i e n t s between species and component axes. Component Ranking 0.675 0.540 0.379 0.369 0.307 0.225 0.084 0.036 -0. 120 -0.921 axis 1 Associated species Chaetoceros radicans  Chaetoceros vanheurcki i  Chaetoceros d e b i l i s  Paralia sulcata  Thalassionema nitzschioides  Chaetoceros a f f i n i s  Chaetoceros u-spp. Chaetoceros didymus  Thalassiosira p a c i f i c a Skeletonema costatum Component Ranking 0.901 0 0 0 0 -0 -0 -0 -0 -0 1 20 039 184 104 242 299 332 ,439 ,554 axis 2 Associated species Thalassiosi ra pac i f ica  Thalassionema nitzschioides  Paralia sulcata Chaetoceros d e b i l i s Chaetoceros  Skeletonema  Chaetoceros Chaetoceros  Chaetoceros a f f i n i s Chaetoceros radicans didymus  costatum u-spp. vanheurcki i 66 AXIS 2 '8 3« 32 . 2« 35- 46 33* 5K «12 -»n l22_ .39 AXIS 1 37* '57*44 •94 3 4 •4 «31 89 •47 88-49 * 77". 51 •62 •10 >91 •38 •54 .45 2L -30-t8 % •17 36 •61 *48 '85*9 •86' • 78. • 79 50 -90 -13 21-•2?8< -18*19 •63 • 93 . 52 56 92 58 95 75* 26 • 74 • 65 •59 •24 • «14 •71 •22 •66 , •67 ' 6 8 . «70»73 '25 .69 76 •15 '23 . '80 16 82 81-64 8 3' Figure 19. Scatter diagram of quadrants on coordinate axes 1 and 2. The numbers plotted refer to s i t e s (see Figs. 5-7). 67 F i g u r e 20. P o s i t i o n of i n l e t s on c o o r d i n a t e axes 1 and 2. (see F i g s . 5-7). 68 A X I S 2 Figure 21. Position of i n l e t s and s i t e s outside i n l e t s on coordinate axes 1 and 2. (see Figs. 5-7). AC = Agamemnon Channel, BI = Burrard Inlet, CC = Cordero Channel, DS = Desolation Sound, EG = East Gorge Harbor, HB = Hidden Basin, HC Homfray Channel, HR = Heriot Bay, MC = Malaspina S t r a i t , PC = Pryce Channel, QC = Queen Charlotte S t r a i t , SC = S u t i l Channel, TN = Texada Island north, and TS = Texada Island south. 69 A comparison of diatom valve and d i n o f l a g e l l a t e cyst d i s t r i b u t i o n and my analysis of Dobell's data established the following: 1) In four small, negligible runoff i n l e t s (Gorge Harbor, Pendrell Sound, Simon Sound, and Port Elizabeth) and one medium runoff i n l e t (Indian Arm), S p i n i f e r i t e s spp. cysts decreased in r e l a t i v e abundance from head to mouth, and 0. centrocarpum cysts increased. In two high runoff i n l e t s (Howe Sound and Knight I n l e t ) , S p i n i f e r i t e s spp. cysts increased in r e l a t i v e abundance from head to mouth, and 0. centrocarpum cysts decreased. This gradient pattern for the high runoff i n l e t s can be compared with phytoplankton and surface sediment data from high runoff Norwegian fjord s . In Hardangerfjord (Braarud 1976), Gonyaulax  gr i n d l e y i (the motile stage of the cyst 0. centrocarpum) decreased in abundance from head to mouth. In Trondheimsfjord, a sediment study (Dale 1976) showed that 0. centrocarpum decreased from head to mouth and the same pattern occurred in the phytoplankton (Sakshaug 1972). S p i n i f e r i t e s spp. cysts in Trondheimsfjiord increased from head to mouth in the surface sediments; however, i t was d i f f i c u l t to make a v a l i d comparison between the d i s t r i b u t i o n of the motile stage of the Gonyaulax  spinifera-group and the cysts known as Spini f e r i t e s sp.-group because of taxonomic problems (Dale 1976). This gradient s h i f t , where a species or species-group reverses i t s head to mouth r e l a t i v e abundance in r e l a t i o n to low or high runoff i n l e t s , was also seen in the r e l a t i v e abundance pattern of the tychopelagic diatom Paralia sulcata (discussed 70 above). In low runoff i n l e t s (Hotham Sound and Saanich I n l e t ) , P. sulcata decreased from head to mouth; while in medium and high runoff i n l e t s , the pattern was reversed. P. sulcata and S p i n i f e r i t e s spp. had their highest r e l a t i v e abundance at the heads of low runoff i n l e t s (65% in Pendrell Sound and 36% in Port Elizabeth). 0. centrocarpum had the highest r e l a t i v e abundance at the mouths of Indian Arm (85%)and Port Elizabeth (81%). P. sulcata was absent or in low abundance (<1%) and the d i n o f l a g e l l a t e cysts were always absent at the heads of high runoff i n l e t s . 2) There was a direct relationship between d i n o f l a g e l l a t e cysts/gram dry weight and depth, both of which increased from head to mouth in the low runoff type i n l e t s (Gorge Harbor, Pendrell Sound, Simon Sound, and Port Elizabeth) and one medium runoff i n l e t (Indian Arm). There was also a d i r e c t relationship between diatom valves/gram dry weight and depth, both of which increased from head to mouth, in two low runoff i n l e t s (Hotham Sound and Pendrell Sound), one medium runoff i n l e t (Indian Arm), and two high runoff i n l e t s (Bute Inlet and Toba I n l e t ) . 3) A comparison of zones using diatom valves and d i n o f l a g e l l a t e cysts was not completely possible because the majority of Dobell's 45 samples were from Zone C (29 s i t e s ) , six (four in Hidden Basin) were from Zone B, and nine (six in Howe Sound) were from Zone A. One sample was from Muchalat Inlet which was not in any of the zones. Zone B was excluded from th i s analysis but i t should be noted that in Hidden Basin the mean r e l a t i v e abundance of S p i n i f e r i t e s spp. and 0. centrocarpum was 71 x=40.3±SD11.5% and 11.8±SD2.6%, respectively. Using a l l (35) of Dobell's samples from Zones A and C, which included 19 of the samples used in the diatom analysis, Zone C was found to have a s i g n i f i c a n t l y greater absolute abundance of d i n o f l a g e l l a t e s (p<0.0l) than that of Zone A. Diatom absolute abundance was the reverse (p<0.0l). In contrast, the r e l a t i v e abundance of both S p i n i f e r i t e s spp. and 0. centrocarpum was s i g n i f i c a n t l y greater (p^O.01) in Zone A than in Zone C. Skeletonema costatum had the same pattern (p^O.001) and P. sulcata was the reverse (p<0.05). Dobell's- sample s i t e s included three high runoff i n l e t s (Howe Sound, Bute Inlet, and Toba I n l e t ) , two medium runoff i n l e t s (Indian Arm and Jervis I n l e t ) , and four low runoff type i n l e t s (Pendrell Sound, Gorge Harbor, Simon Sound, and Port Elizabeth). Medium run off i n l e t s were excluded from the analysis because there were only three samples. A l l the high runoff i n l e t s , Pendrell Sound and East Gorge Harbor, were used in the diatom analysis. The absolute abundance of d i n o f l a g e l l a t e cysts was s i g n i f i c a n t l y greater in low runoff i n l e t s than in high runoff i n l e t s (p^O.Ol). There was no s i g n i f i c a n t difference between i n l e t types for diatom absolute abundance. As Dobell noted, 0. centrocarpum was usually dominant within bays and i n l e t s . But i t should be further noted that in the low runoff type i n l e t s (Pendrell Sound, Gorge Harbor, Simon Sound, and Port Elizabeth) S p i n i f e r i t e s spp. , although not dominant except at the head of Port Elizabeth, always had a greater r e l a t i v e 72 abundance at the head, and 0. centrocarpum at the mouth. These analyses e s t a b l i s h e d the f o l l o w i n g c o n c l u s i o n s : 1) There was a d e f i n i t e head to mouth g r a d i e n t w i t h i n the i n l e t s f o r P. s u l c a t a , S p i n i f e r i t e s spp. , and 0. centrocarpum and these g r a d i e n t s change with respect to i n l e t type. 2) D i n o f l a g e l l a t e c y s t s had g r e a t e r a b s o l u t e abundance i n Zone C than i n Zone A, and diatom v a l v e s had a reverse p a t t e r n . 3) The a b s o l u t e abundance of d i n o f l a g e l l a t e c y s t s was g r e a t e r i n low runoff i n l e t s than i n high runoff i n l e t s but there was no d i f f e r e n c e f o r diatom a b s o l u t e abundance. 73 V. DISCUSSION The discussion section is divided into two parts: Part I deals with the relationship between the biocoenosis ( i . e . , species found in the water column) and the thanatocoenosis ( i . e . , species found in the sediments), both for B r i t i s h Columbia sediments and for areas which should t h e o r e t i c a l l y have similar sediment assemblages. The l o g i c a l expectation i s that dominant phytoplankton species w i l l become dominant species in the sediment assemblages; however, t h i s i s not always the case. As w i l l be seen, one of the major conclusions of t h i s study i s that the c a t c h - a l l explanation of d i f f e r e n t i a l d i s s o l u t i o n of weakly s i l i c i f i e d species i s simply not s u f f i c i e n t to explain the discrepancies between the biocoenoses and the thanatocoenoses. Part II deals with the diatom d i s t r i b u t i o n a l patterns in B r i t i s h Columbia sediments in r e l a t i o n to various factors (e.g., primary productivity, l i g h t , and upwelling). In general, the same correlations established in numerous other sediment studies are also exhibited in B r i t i s h Columbia . PART I Part I i s divided into two sections. The f i r s t section is a discussion of d i f f e r e n t i a l d i s s o l u t i o n and v e r t i c a l transport. Interesting cases of discrepancies between the biocoenoses and thanatocoenoses of Thalassiosira nordenskioeldii and the two forms of Thalassiosira a e s t i v a l i s are included in t h i s section. The second section i s a comparison of the biocoenoses and 74 the thanatocoenoses of recurrent groups and s e l e c t e d dominant and rare species i n the sediment assemblages. I. DIFFERENTIAL DISSOLUTION AND VERTICAL TRANSPORT A number of studies (Kanaya and Koizumi 1966; Kozlova and Mukhina 1967; Schrader 1971; Parker et a l . 1977; Schuette and Schrader 1981; Tanimura 1981) have noted that some diatom species i n the water column (the biocoenosis) are not found i n the sediments (the thanatocoenosis). The presence of diatoms i n the sediments has been as s o c i a t e d with some form of a c c e l e r a t e d v e r t i c a l t r a n s p o r t to the sediments, whereas the absence of diatoms i s u s u a l l y a s s o c i a t e d with d i f f e r e n t i a l d i s s o l u t i o n of weakly s i l i c i f i e d f r u s t u l e s . A c c e l e r a t e d v e r t i c a l t r a n s p o r t through the water column decreases the time i n which d i f f e r e n t i a l d i s s o l u t i o n can occur; however, d i s s o l u t i o n can continue i n the pore waters of the sediments (Lewin 1961; Fanning and Schink 1969; C a l v e r t 1974). The s i n k i n g r a t e s of diatoms can vary with l i f e stages and p h y s i o l o g i c a l s t a t e s . Resting spores of Chaetoceros spp. can sink approximately f i v e times f a s t e r than t h e i r v egetative stages (Bienfang 1981). Species under s i l i c a t e d e p l e t i o n sink f a s t e r than n u t r i e n t - r e p l e t e or N- or P-deplete c u l t u r e s (Bienfang et a l . 1982). The average s i n k i n g rate for whole-plankton assemblages i n the l a b o r a t o r y and i n the upper 12 m i n Saanich I n l e t was shown to be as l i t t l e as x=0.64±SD0.31 m/day and 0.72±SD0.08 m/day, r e s p e c t i v e l y (Bienfang 1981). At these r a t e s , i t would take nearly a year f o r diatoms to reach the deep basin (approximately 75 200 m) of Saanich Inlet. However, individual species may have faster sinking rates: for example, the mean sinking rate of Skeletonema costatum ranged from 0.30 to 1.35 m/day, with a maximum rate of 7.39 m/day (Smayda and Boleyn 1966). Sinking rates for fecal p e l l e t s ranged from 36-376 m/day (Smayda 1969; Bienfang 1981). At these rates, diatoms enclosed in fecal p e l l e t s could reach even the deepest studied s i t e (approximately 700 m in Homfray Channel and Bute Inlet) in a matter of days. Mixing, downwelling, and density inversion currents can also increase v e r t i c a l transport (Smayda 1970 and 1971; Walsby and Reynolds 1980). From a one year study in Saanich Inlet, Stephens et a l . (1967) noted that maximum deposition of phytoplankton material occurred two months after the spring bloom in which the most abundant organisms were diatoms. The quantity of thi s deposited material represented nearly half of the annual primary productivity. Although zooplankton grazing was noted, other factors were considered more important in rel a t i o n to increased phytoplankton sedimentation ( i . e . , aggregation, increased s t a b i l t i y of the water column, and increased sinking rates of diatoms related to senescence and nutrient l i m i t a t i o n ) . Another form of accelerated v e r t i c a l transport i s p a r t i c l e interaction, the products of which are variously c a l l e d agglomerates, floes, or marine snow. In a sediment trap study off the coast of C a l i f o r n i a , Urrere and Knauer (1981) noted that fecal p e l l e t s , diatoms, f l a g e l l a t e s and protozoans were associated with large amounts of flocculent material and that 76 t h e s e f l o e s c o u l d have been r a p i d l y s e d i m e n t e d . I n a n o t h e r s e d i m e n t t r a p s t u d y i n Howe So u n d , S y v i t s k i a n d M u r r a y (1981) i d e n t i f i e d ' a g g l o m e r a t e s ' ( l a r g e - g r a i n e d f l o e s c o n t a i n i n g d i a t o m f r u s t u l e s ) a s one o f f i v e t y p e s o f m a r i n e p a r t i c l e s . P a r t i c l e i n t e r a c t i o n was a s s o c i a t e d w i t h t h e h e a v y s e d i m e n t l o a d d u r i n g t h e f r e s h e t o f t h e S q u a m i s h R i v e r w h i c h e n t e r s a t t h e h e a d o f Howe So u n d . B i o l o g i c a g g l o m e r a t i o n a n d z o o p l a n k t o n p e l l e t i z a t i o n o f i n o r g a n i c f l o c c u l e s were c o n s i d e r e d t o be p r e d o m i n a n t i n t h e s e w a t e r s . A l t h o u g h many form's o f a c c e l e r a t e d v e r t i c a l t r a n s p o r t e x i s t , Smayda (1971) c o n s i d e r e d f e c a l p e l l e t t r a n s p o r t t h e most i m p o r t a n t . F e c a l p e l l e t f l u x e s h a v e been i m p l i c a t e d a s a p o s s i b l e m a j o r s o u r c e o f b o t h o r g a n i c a n d i n o r g a n i c m a t e r i a l t o d e e p - w a t e r s a n d s e d i m e n t s ( B o o t h a n d K n a u e r 1972; W i l s o n 1978; S u e s s 1980; U r r e r e a n d K n a u e r 1 9 8 1 ) . More i m p o r t a n t t o t h i s s t u d y i s t h e p r e s e n c e o f f r a g m e n t e d o r w h o l e empty f r u s t u l e s , o r i n t a c t c e l l s i n f e c a l p e l l e t s ( J o h a n n e s a n d S a t o m i 1966; Wimpenny 1973; U r r e r e a n d K n a u e r 1 9 8 1 ) . C o p r o p h a g y o f f e c a l p e l l e t s c a n p r o d u c e l a r g e r p a r t i c l e s w h i c h f u r t h e r i n c r e a s e s s e t t l i n g r a t e s ( H o n j o a n d Roman 1 9 7 8 ) . The p h y s i c a l s t a t e o f t h e d i a t o m ( i . e . , f r a g m e n t e d o r w h o l e empty f r u s t u l e s , o r i n t a c t c e l l s ) i s d i r e c t l y r e l a t e d t o d i s s o l u t i o n r a t e o f t h e f r u s t u l e . Numerous s t u d i e s h a v e e x a m i n e d d i s s o l u t i o n r a t e s o f a c i d - c l e a n e d f r u s t u l e s a n d e s t a b l i s h e d t h e f o l l o w i n g r e s u l t s : 1) f r a g m e n t a t i o n a n d i n c r e a s e d s u r f a c e a r e a i n c r e a s e d d i s s o l u t i o n r a t e s ( H u r d 1 9 7 2 ) ; 2) i n c r e a s e d pH and t e m p e r a t u r e a l s o i n c r e a s e d d i s s o l u t i o n r a t e s 77 (Lewin 1961; Lawson et a l . 1978); and 3) inorganic ions absorbed by the acid-cleaned frustules decreased d i s s o l u t i o n rates (Lewin 1961; Johnson 1974). The majority of the above authors recognized that the use of acid-cleaned frustules as opposed to l i v i n g c e l l s precluded d i r e c t comparison with the natural environment. By measuring the s i l i c i c acid concentration of the medium in which diatoms were allowed to dissolve, Lewin (1961) detected l i t t l e or no dissolution of s i l i c a from l i v i n g c e l l s , and showed that intact k i l l e d - c e l l s dissolved slower than acid-clea'ned valves. Lewin recognized and Nelson et a l . (1976) demonstrated, by using tracer techniques, that s i l i c a d i s s o l u t i o n of l i v i n g c e l l s is masked by s i l i c i c acid uptake. A l l diatoms probably have an organic "casing" (Stosch 1980; Volcani 1981), even though i t may not be apparent with electron microscopy. The chemical composition of the organic casing, silicalemma, and developing c e l l wall can be d i f f e r e n t among diatom species (Hecky et a l . 1973; Volcani 1981). Further, and most importantly, the r a t e - c o n t r o l l i n g step in the natural di s s o l u t i o n of the frustule in both the water column and the sediments may be the degradation rate of the organic casing (Hecky et a l . 1973). The persistence of the organic casing, given that i t may s i g n i f i c a n t l y decrease the dissolution rate of the frustule, is extremely important to sediment studies. Limiting the causal agent of d i f f e r e n t i a l preservation to the thickness of the s i l i c i f i e d f rustule is far too s i m p l i s t i c to explain the discrepancies between the biocoenoses and the 78 thanatocoenoses. The following observations from a number of studies demonstrate the complexity of this problem: 1) rapidly growing c e l l s deposit thinner frustules than slowly growing c e l l s (Lewin 1962) and the chemical composition of the organic casing appears to be related to the early development of the c e l l wall (Hecky et a l . 1973). C e l l s maintained in the dark at low temperatures have remained viable for as long as three years (Smayda and Mitchell-Innes 1974; Antia 1976). These species have slower growth rates, and given the above observations (Lewin 1962; Hecky et a l . 1973), may have thicker frustules and di f f e r e n t organic casings than when maintained under optimum growth conditions. Therefore, the same species under two separate conditions ( i . e . , rapid growth in the euphotic zone or slow growth in the aphotic zone or sediments) could conceivably have a d i f f e r e n t preservation p o t e n t i a l . 2) The v e r t i c a l d i s t r i b u t i o n of species in the water column may affe c t eventual preservation in the sediments. Venrick (1982) found two d i s t i n c t f l o r a l associations above and below a rapid t r a n s i t i o n region at approximately 100 m in the North P a c i f i c . Zooplankton grazing was predominant in the upper l e v e l . If zooplankton p e l l e t i z a t i o n i s a primary form of v e r t i c a l transport and preservation, species from the upper association w i l l be p r e f e r e n t i a l l y transported and possibly preserved in the sediments. 3) Some zooplankton have species- and s i z e - s p e c i f i c feeding strategies (Marshall and Orr 1955; Parsons et a l . 1977; Jewison et a l . 1981). Zooplankton fecal p e l l e t s may contain fragmented 79 frustules, empty but whole frustules, or intact c e l l s depending on feed strategy and passage time through the gut (Johannes and Satomi 1966; Smayda 1971; Wimpenny 1973; Urrere and Knauer 1981). As noted above, the state of the frustule i s d i r e c t l y related to the dissolution rate. The i n t e r i o r of fecal p e l l e t s i s probably anoxic (Wilson 1978) and axenic (Honjo and Roman 1978) and some c e l l s in fecal p e l l e t s can remain viable (Johannes and Satomi 1966). Therefore, even a weakly s i l i c i f i e d , intact c e l l transported to the sediments in a fecal p e l l e t may have a preservation advantage over a heavily s i l i c i f i e d empty or fragmented f r u s t u l e . 4) Stockner and Lund (1970) found viable c e l l s of Melosira  i t a l i c a in anoxic, freshwater sediments to a depth of 35 cm (195±12 years), and maintained c e l l s in the laboratory under anoxic conditions for three years. Intact c e l l s of Skeletonema  costatum and Paralia sulcata were found in the sediments of Saanich Inlet (personal observation), which i s p e r i o d i c a l l y anoxic. However, Hollibaugh et a l . (1981) examining survival and germination of Chaetoceros spp. resting spores from Saanich Inlet, suggest that spores lost v i a b i l i t y when exposed to anoxic conditions; they include no s p e c i f i c information. Preservation varies in the anoxic sediments of the Black Sea. Zgurovskaya (1979) examining shallow (30-50 m) surface sediments found Chaetoceros spp. resting spores and vegetative c e l l s of S. costatum, Thalassiosira eccentrica , and Thalassiosira spp. present and viable throughout the year. It was also noted that no empty frustules of S. costatum or 80 Chaetoceros spp. were found in the sediments. From deep-water cores, Maynard (1974) found very poor preservation of diatoms. She considered this surprising since diatom frustules were reported as being extremely abundant in bottom oozes from most parts of the sea (see Russian references in Maynard 1974), and suggested that there must be a high rate of s i l i c a d i s s o l u t i o n within the sediments. A similar lack of preservation with sediment depth does not appear to occur in Saanich Inlet (Gucluer and Gross 1964). 5) Different l i f e stages preserve d i f f e r e n t l y . Thickly s i l i c i f i e d resting spores (e.g., Chaetoceros spp. and Leptocylindrus danicus ) are often found in the sediments but the vegetative stages are very rare. Species which require some form of resting stage, whether i t i s a resting spore or a quiescent vegetative c e l l , may have a more dis s o l u t i o n resistant organic casing and/or a thicker frustule than species which constantly maintain a small population in the water column. If a species could maintain one c e l l per l i t e r , an undetectable amount at present, i t would be capable of reaching bloom proportions ( 1 0 6 c e l l s / l i t e r ) in seven days, given three d i v i s i o n s per day. Therefore, the presence or absence of a species in the sediments may be d i r e c t l y related to l i f e s trategies. A more h o l i s t i c view i s needed of the factors involved in the preservation of species in the sediments. Greater emphasis should be placed on understanding the role of the organic casing in preservation. Although weakly s i l i c i f i e d diatoms are more 81 l i a b l e to destruction by zooplankton and probably to dis s o l u t i o n , i f there i s a direct r e l a t i o n s h i p between the composition of the organic coating and c e l l wall thickness, a theory based on the thickness of the diatom frustule i s not s u f f i c i e n t to explain the disproportionate preservation of some medium to heavily s i l i c i f i e d diatoms in t h i s study. Two cases in point are the discrepancies between the phytoplankton records and the proportions in the sediments of the two forms of Thalassiosira a e s t i v a l i s and Thalassiosira nordenskioeldii. 1. Thalassiosira a e s t i v a l i s Appendix B is a detailed explanation of why Thalassiosira pac i f ica and T. aest i v a l i s are considered two forms of one polymorphic species. In thi s thesis, the two forms are referred to as T. a e s t i v a l i s f. a e s t i v a l i s and T. a e s t i v a l i s f. p a c i f i c a . The taxonomy of these forms i s important to thi s study because two separate but overlapping phytoplankton investigations (Shim 1976; Stockner and C l i f f 1976) state that both species were dominants for two years prior to the c o l l e c t i o n of 24 sediment samples in Howe Sound in 1974. However, only T. a e s t i v a l i s . f. pac i f ica was dominant in a l l the sediment samples including Howe Sound. Figure 22 shows the d i s t r i b u t i o n of areolae in 10 Mm from 300 T. a e s t i v a l i s valves counted from 30 sit e s representing every area in thi s study. The curve i s unimodal with a mean of 12 areolae in 10 nm. This f i t s into the T. a e s t i v a l i s f. p a c i f i c a range. Figure 23 i s the d i s t r i b u t i o n of areolae in 10 am in thi s species from 11 sediment samples (146 valves) along the length of Howe Sound. The mean i s 13 82 Figure 22. D i s t r i b u t i o n of areolae in 10 Mm for Thalassiosira  a e s t i v a l i s f. p a c i f i c a from 30 s i t e s representing the entire study area. 83 ^ 50 I LU i 1 " ' ' 1 9 10 11 12 13 U 15 16 17 18 19 20 AREOLAE IN 10um Figure 23. D i s t r i b u t i o n of areolae in 10 jum for Thalassiosira  a e s t i v a l i s f. p a c i f i c a from 11 s i t e s along the length of Howe Sound (1974).. 84 areolae in 10 jum which i s also within the T. a e s t i v a l i s f. p a c i f i c a range. Between May 1972 and July 1973, Shim (1976) sampled the southern S t r a i t of Georgia, which is contiguous with Howe Sound, in a study of the d i s t r i b u t i o n and taxonomy of marine planktonic diatoms. He states that T. a e s t i v a l i s was "very abundant," "occurred frequently," and had peak abundance in "late spring." T. p a c i f i c a had a "major population component in the spring, but was rarely observed from mid-summer to winter." This would appear to correspond with Phifer's (1934a) temporal d i s t r i b u t i o n (see Appendix B) except that Shim has some taxonomic problems with these forms. Shim's written descriptions and micrographs of T. pac i f ica and T. a e s t i v a l i s are both probably T. a e s t i v a l i s f. a e s t i v a l i s ; and Thalassiosira sp. B, which Shim separates out as a new species because the colonies are in a gelatinous mass, i s probably T. a e s t i v a l i s f. p a c i f i c a in fecal p e l l e t s . Between January 1973 and May 1975, Stockner and C l i f f (1976) examined phytoplankton succession and abundance in Howe Sound. The authors state that "dominant species in Howe Sound were a l l diatoms: Skeletonema costatum, T. a e s t i v a l i s , T. nordenskioeldi i , T. paci f ica and Chaetoceros spp. S. costatum and T. pac i f ica were dominants at a l l stations." A photograph of T. a e s t i v a l i s from a study of Pendrell Sound and Hotham Sound (Stockner and C l i f f 1975), where both T. p a c i f i c a and T. a e s t i v a l i s are l i s t e d as dominants, i s too poor to check the determination. The only real clue as to how Stockner and 85 C l i f f separated T. a e s t i v a l i s from T. p a c i f i c a i s in the Howe Sound study. The estimated c e l l volume of T. aest i v a l i s i s given as 7,000 ym3, and T. pac i f ica as 20,000 ym3. These r e l a t i v e volumes suggest that the species may have been separated by siz e , although the written descriptions (Gran and Angst 1931; Hasle 1978) show that both have the same size range. It could be argued that T. a e s t i v a l i s f. a e s t i v a l i s i s not a dominant in the sediments because 1) sediment samples are not representative of a period when thi s form was dominant, 2) i t was d i f f e r e n t i a l l y dissolved, and/or 3) i t was inaccurately i d e n t i f i e d . Sixty-one of the 95 sediment samples were coll e c t e d between August and November in five separate years; and 31 samples were taken in May of three years. Secondly, although components of the frustule of T. a e s t i v a l i s f. a e s t i v a l i s are considered to be more weakly s i l i c i f i e d than T. a e s t i v a l i s f. p a c i f i c a (Gran and Angst 1931; Hasle 1978), both have r e l a t i v e l y heavily s i l i c i f i e d valves. This author does not consider simple d i f f e r e n t i a l d i s s o l u t i o n , as a result of s e t t l i n g through a column of water, s u f f i c i e n t to explain the discrepancy between phytoplankton and sediment data. However, the propitious combination of d i f f e r e n t i a l d i s s o l u t i on and inaccurate taxonomy may explain the discrepancy. Shim (1976) found a pa r t i c u l a r Thalassiosira species, encased in what he described as a gelatinous mass, so abundant in the phytoplankton that he designated i t Thalassiosira sp. B. As has been noted, Thalassiosira sp. B i s most l i k e l y T. a e s t i v a l i s f. p a c i f i c a in fecal p e l l e t s . Shim further stated 86 that many "threads" ( f i b r i l s from marginal strutted processes) and numerous chloroplasts could be seen. Apparently T. a e s t i v a l i s f. p a c i f i c a can remain intact and possibly viable after passage through the zooplankter's gut. The presence of a "gelatinous mass" was not noted by Shim for either diatom which he described as T. p a c i f i c a or T. a e s t i v a l i s , both of which were probably T. aest i v a l i s f. a e s t i v a l i s . Therefore, i f T. a e s t i v a l i s f. p a c i f i c a were s e l e c t i v e l y grazed r e l a t i v e to T. aest i v a l i s f. a e s t i v a l i s , and T. a e s t i v a l i s f. pac i f ica maintained i t s organic casing within the fecal p e l l e t , i t would be doubly protected from di s s o l u t i o n and p r e f e r e n t i a l l y deposited in the sediments. The presence of f i b r i l s on the T. aest i v a l i s f. pac i f ica that Shim observed strongly suggests that i t may have been p r e f e r e n t i a l l y grazed. Jewison et a l . (1981) fed the copepod Calanus f inmarchicus "spined" and "unspined" Thalassiosira  weissflogi i (= T. f l u v i a t i l i s ) . The spined c e l l s were those which were allowed to maintain their chitan f i b r i l s , unspined c e l l s had their f i b r i l s removed by a g i t a t i o n . With f i b r i l s i ntact, the diameter of a spined c e l l appeared to be about 2.4 times greater than an unspined c e l l . The copepod preferred spined c e l l s presumably because of the i r apparently larger s i z e . There appears to be a seasonal separation of the two T. a e s t i v a l i s forms (see Appendix B). T. a e s t i v a l i s f. p a c i f i c a occurs during the spring which is the period of maximum zooplankton production in the S t r a i t of Georgia, and T. a e s t i v a l i s f. a e s t i v a l i s occurs during the late spring and 87 summer, when zooplankton production i s generally 10% or less of the peak spring production (Harrison et a l . in press). The temporal separation of the two forms and the r e l a t i v e zooplankton grazing pressure may also be important in rel a t i o n to seasonal temperature of the water column. Honjo and Roman (1978) demonstrated a di r e c t relationship between increasing degradation of copepod p e l l e t s and increasing temperature. Degradation was associated with increasing external colonization of bacteria; the inside of the p e l l e t appeared to be axenic. At 25°C, complete degradation of p e l l e t s in the dark, in seawater, occurred within 24 hours. At 0°C, p e l l e t s remained intact, with very l i t t l e b a c t e r i a l colonization, for 20 days, when the experiment was terminated. At 10°C and 15°C, degradation was reduced in r e l a t i o n to 25°C and ba c t e r i a l colonization was greater at 15°C than 10°C. These results suggest that fecal p e l l e t s produced in the spring when surface water temperature is approximately 8-10 oC (Pickard 1961) would degrade more slowly than p e l l e t s in the summer surface waters (approximately 10-15°C, Pickard 1961). The bottom waters remain at approximately 6-8°C throughout the year (Pickard 1961); however, the period a pe l l e t would remain in the r e l a t i v e l y warmer surface waters varies with a number of factors (e.g., s t a b i l i t y of the water column and size of the p e l l e t ) . Therefore, T. a e s t i v a l i s f. p a c i f i c a grazed in the spring may have a preservation advantage over T. a e s t i v a l i s f. a e s t i v a l i s in the summer. The dissolution of the s i l i c a frustules also varies with temperature. Assuming some c e l l s are stripped of their organic 88 casing by b a c t e r i a l action, Lewin's (1961) experiments using acid-cleaned frustules are relevant. She showed a di r e c t rel a t i o n s h i p between dissolution of s i l i c a from acid-cleaned frustules and increasing temperature. Heat-killed plankton assemblages dissolved more slowly at 6°C than at 19°C. Again, i t appears that a c e l l , empty or intact, could have a preservation advantage in cooler spring temperatures. That T. aest i v a l i s f. pac i f ica may have maintained i t s organic casing within fecal p e l l e t s , and occurred coincidentally with the peak abundance of possible predators and r e l a t i v e l y cooler temperatures, a l l suggest that T. a e s t i v a l i s f. p a c i f i c a may have been both p r e f e r e n t i a l l y grazed, preserved, and deposited in the sediments. This hypothesis appears to explain why T. a e s t i v a l i s f. p a c i f i c a i s both a dominant in the water column and the sediments, whereas T. aest i v a l i s f. a e s t i v a l i s is a dominant in the water column but extremely rare in the sediments. The r e l a t i v e thicknesses of the f r u s t u l e i s not relevant in t h i s case. Global d i s t r i b u t i o n patterns and sediment data for the two forms are limited. Hasle (1976 and 1978) described T. pac i f ica (= T. a e s t i v a l i s f. pacifica) as a cold to temperate water species r e s t r i c t e d to the northern hemisphere. In the Sea of Japan, Tanimura (1981) included T. p a c i f i c a in the l i s t of species dominant in the sediments and used in f l o r a l analysis, but made no further mention of i t . 89 Hasle (1978) stated that T. a e s t i v a l i s (= T. a e s t i v a l i s f . a e s t i v a l i s ) could be " i d e n t i f i e d with certainty only from northwest American coastal waters (Puget Sound, Washington and Vancouver Island)," and questioned reports of th i s species from numerous other areas ( i . e . , A t l a n t i c Ocean, Indian Ocean, P a c i f i c Ocean, Bay of Valparaiso and Bay of Conception in Chile, Gulf of Panama, Oslofjord in Norway, the North Sea, and the coastal waters of Peru, Chile, and South A f r i c a ) . Hasle (1978) further stated in the discussion of the d i s t r i b u t i o n of T. a e s t i v a l i s that "confusion with other species may ea s i l y take place i f too much emphasis i s l a i d on published drawings of colonies (Gran and Angst 1931; Cupp 1943)." The only mention of T. a e s t i v a l i s (= T. a e s t i v a l i s f. a e s t i v a l i s ) in sediments i s from the coastal upwelling area off South West A f r i c a (Schuette and Schrader 1981). The authors were aware that Hasle (1978) reserved judgement on a l l reported c i t i n g s of T. a e s t i v a l i s except those from the northwestern American coastal waters, but stated that their specimens f i t Hasle's amended description of T. a e s t i v a l i s . In the majority of their sediment samples, T. a e s t i v a l i s was most often represented by only a single specimen. At 12 s i t e s in or near Walvis Bay, i t comprised from 1 to 10% of the s i t e count. In the core study from Saanich Inlet (Gucluer and Gross 1964), neither form of T. a e s t i v a l i s was noted. However, in another core from Saanich Inlet examined by this, author, T. a e s t i v a l i s f. p a c i f i c a was found. The lack of either form of T. a e s t i v a l i s in the sediments 90 of other areas may be due to taxonomic confusion rather than d i f f e r e n t i a l d i s s o l u t i o n . 2. Thalassiosira nordenskioeldii T. nordenskioeldii has been reported as a dominant species in the water column during the spring and summer in the S t r a i t of Georgia and contiguous waters, and in lesser amounts throughout the rest of the year (Phifer 1934a; Buchanan 1966; Takahashi et a l . 1973; Shim 1976). It is a dominant of the spring bloom in many northern areas as a result of i t s short generation time under low l i g h t i n t e n s i t i e s and low temperatures (G u i l l a r d and Kilham 1977; Baars 1982). T. nordenskioeldi i i s a member of recurrent group A and i s found at 70 of the 95 s i t e s . However, there i s a discrepancy between the biocoenoses and the thanatocoenoses. Although a major dominant in the phytoplankton, T. nordenskioeldi i has a re l a t i v e abundance of ^ 3.5% in 77% of the sediment samples where i t occurred, and represents less than 2% of the t o t a l valve count. T. nordenskioeldi i can form resting spores. Syvertsen (1979) states that "the resting spore frustule of T. nordenskioeldii shows a close morphologic relationship to the vegetative fru s t u l e , " and "the most c h a r a c t e r i s t i c d i s t i n c t i o n i s the reduction of the foramina in the spore valve." However, Hasle (1978) notes that the foramina of vegetative c e l l s range from widely open to almost closed. Durbin (1978) states that the main difference i s that resting spore valves have "heavy sculpturing." A comparison of the various micrographs 91 demonstrates the confusion. In F i g . 11 (in Syvertsen 1979) and F i g . 18 (in Hasle 1978), the vegetative c e l l has open foramina, whereas the resting spore has almost closed foramina. Durbin (1978) i l l u s t r a t e s the vegetative c e l l (Fig. 3C) with almost closed foramina, and the resting spore (Fig. 3E) with open foramina. Figs. 17 and 20 (in Hasle 1978) show vegetative c e l l s with closed foramina. Whether or not the foramina are opened or closed i s obviously not a conservative c h a r a c t e r i s t i c . As Booth and Harrison (1979) have shown, Thalassiosira  eccentrica lacked a s i l i c a covering over the foramina under s i l i c a t e l i m i t a t i o n . The variation in the foramina covering of both vegetative c e l l s and resting spores of T. nordenskioeldi i may be a response to nutrient l i m i t a t i o n . The thickness of the resting spores i s also disputed. Hasle (1978) says the valves in resting spores are more coarsely s i l i c i f i e d ; Syvertsen (1978) says the spore g i r d l e i s s l i g h t l y more s i l i c i f i e d . These c h a r a c t e r i s t i c s may also be enviromentally induced. Regardless of these differences, the valves of both the vegetative c e l l s and resting spores can be considered moderately to heavily s i l i c i f i e d . The majority of T. nordenskioeldii resting spores are either endogenous spores ( i . e . , spore enclosed completely within parent c e l l ) or semi-endogenous spores ( i . e . , p a rtly enclosed within parent c e l l ) (Durbin 1978; Syvertsen 1979). The great advantage of endogenous or semi-endogenous spore formation i s that i t is easy to id e n t i f y the resting spores in culture. This advantage i s not available in sediment studies where natural 92 processes and/or cleaning processes separate the valves. Therefore, i t is impossible, at present, to determine whether the valves of T. nordenskioeldii in the B r i t i s h Columbia sediments are resting spores or vegetative c e l l s . The occurrence of resting spores or quiescent c e l l s in the sediments i s almost necessary to explain the repeated presence of T. nordenskioeldii in the annual spring bloom. In discussing the o r i g i n of the well-documented annual spring bloom of T. nordenskioeldi i in Narragansett Bay, Rhode Island (Pratt 1959; Smayda 1973), Durbin (1978) admitted the p o s s i b i l i t y of an allochthonous source but preferred the theory that the quiescent c e l l s survived in shallow areas where they would p e r i o d i c a l l y receive l i g h t . No allochthonous P a c i f i c Ocean source has been i d e n t i f i e d for the B r i t i s h Columbia area either (Venrick 1971; Taylor and Waters 1982). The discussion of the discrepancy between the proportion of T. nordenskioeldi i in the biocoenoses and the thanatocoenoses w i l l be limited to Saanich Inlet, where a large body of c o l l a t e r a l physical and b i o l o g i c a l information i s available. Throughout the study area, T. nordenskioeldi i i s reported as a dominant in the spring phytoplankton, and has a low abundance in the sediments; therefore, i t i s assumed that an explanation s u f f i c i e n t for Saanich Inlet may be considered indicative of the remainder of the study area. Saanich Inlet i s atypical in one respect: the bottom waters and sediments can be p e r i o d i c a l l y anoxic. However, the anoxia does not appear to aff e c t preservation since the abundance of T. nordenskioeldii in 93 Saanich Inlet sediments is similar to that in other i n l e t sediments. In order to explain the discrepancy between the proportion of T. nordenskioeldii in the biocoenoses and the thanatocoenoses in Saanich Inlet, six aspects need to be examined: 1) The autecology and preservation of vegetative c e l l s and resting spores of T. nordenskioeldii, 2) the presence of T. nordenskioeldii in the phytoplankton, 3) ambient nitrogen concentrations in the surface waters, 4) grazing pressure, 5) physical conditions and sedimentation, 6) the presence of T. nordenskioeldii in the sediments. 1) There are two excellent autecology studies of T. nordenskioeldii (Durbin 1978; Baars 1982). Baars (1982) found that the growth rate of T. nordenskioeldi i was related to temperature, l i g h t intensity, and day length. In general, he determined that T. nordenskioeldi i would be able to develop a bloom at low l i g h t i n t e n s i t i e s and temperatures between 0-6°C, although growth could s t i l l occur at 20-21°C. Baars includes a review of the d i s t r i b u t i o n of T. nordenskioeldii in the northern hemisphere which substantiates his findings that within a certain temperature range, growth i s inversely related to surface water temperatures. Durbin (1978) investigated the biology of resting spores of T. nordenskioeldii. Nitrogen l i m i t a t i o n was used to induce resting spore formation. Under low nitrogen concentrations, spore formation was inversely related to temperature: at 0°C and 5°C between 68 and 92% of the t o t a l c e l l s produced were resting 94 spores, at 10°C between 40 and 52%, and at 15°C no resting spores were produced. Once resting spores formed, the cultures were placed in the dark. Durbin noted that vegetative c e l l s soon died and disintegrated. V i a b i l i t y of resting spores was also inversely related to temperature: at 0°C resting spores remained viable for 576 days, at 5°C for 200 days, at 10°C for 95 days, and at 15°C for 60 days. 2) The spring bloom in Saanich Inlet begins in March or A p r i l (Stephens et a l . 1967; Huntley and Hobson 1978) and is usually i n i t i a l l y dominated by T. nordenskioeldii (Huntley and Hobson 1978). During A p r i l to May, 1975, Huntley and Hobson (1978) sampling at 0 and 5 m, found that T. nordenskioeldi i composed 76 and 65% of the t o t a l c e l l volume, respectively. In May 1981 and 1982 in the upper 10 m, T. nordenskioeldii represented 60-70% of the t o t a l c e l l count (personal observation). In A p r i l 1932, Phifer (1934c) sampling the upper 20 m in East Sound, Washington, found 70-80% of the T. nordenskioeldi i population in the upper 5 m. These data suggest that the majority of the T. nordenskioeldii population remains in the upper 10 m during the early spring bloom. 3) As noted above, low nitrogen concentrations, either n i t r a t e or ammonium, may induce resting spore formation in T. nordenskioeldi i (Durbin 1978; Syvertsen 1979). The combined data on nitrogen concentrations from a number of sources (Stephens 1967; Harrison et a l . in press) suggest that some form of nitrogen i s usually available in the study area. For example, as n i t r a t e levels decreased in the surface waters of 95 Saanich Inlet, ammonium level s increased (Huntley and Hobson 1978). The length of exposure to low nitrogen concentrations necessary to induce resting spore formation i s unknown. Large numbers of T. nordenskioeldi i resting spores have not been reported in the l i t e r a t u r e nor observed in B r i t i s h Columbia phytoplankton samples (F.J.R. Taylor, personal communication). These data suggest that resting spore formation in re l a t i o n to low nitrogen concentration may be very limited. 4) Herbivores are major members of the zooplankton in Saanich Inlet (Stephens et a l . 1967; Huntley and Hobson 1978). In phytoplankton samples taken in May 1981, the author saw zooplankton ingesting whole chains of T. nordenskioeldi i and observed fecal p e l l e t s containing only empty chains of T. nordenskioeldii. In t h i s case, the c e l l contents of T. nordenskioeldii were f u l l y digested although the frustule remained intact after passage through the zooplankter's gut. Based on four years of observations, Huntley and Hobson (1978) described the following successional pattern in r e l a t i o n to zooplankton grazing: 1) T. nordenskioeldi i dominates the early spring bloom beginning in A p r i l ; 2) the phytoplankton i s maintained at low levels toward the end of the early spring bloom (early June) by herbivore grazing; and 3) predation by medusae on herbivores allows a second spring bloom in late June, which i s dominated by Skeletonema costatum and Chaetoceros spp. 5) There are two periods of maximum deposition of sediment in Saanich Inlet: the f i r s t occurs between May to July and consists mainly of phytoplankton material and zooplankton fecal 96 p e l l e t s , and the second occurs between October to December and consists mainly of terrigenous material(Stephens et a l . 1967; Iseki et a l . 1980). Stephens et a l . (1967) noted that the f i r s t maximum of sedimented material occurred approximately two months after the spring phytoplankton bloom and the t o t a l quantity represented approximately half of the annual primary production. The lag between phytoplankton production and sedimentation of phytogenous material involved f i v e factors. 1) Although intensive zooplankton grazing was noted, i t was not thought to be d i r e c t l y related to increased sedimentation. 2) There was an increase in the size of suspended p a r t i c l e s between January and June, which was suggested to be related to aggregation of p a r t i c l e s or larger phytoplankton species. 3) Between May and July there was a decrease in s a l i n i t y of the surface waters associated with the freshet of the Fraser River; and as noted above, there i s a direc t relationship between decreased s a l i n i t y associated with the freshet in Howe Sound and increased aggregation of p a r t i c l e s (Syvitski and Murray 1981). 4) Increased sedimentation could be related to increased s t a b i l i t y of the water column, which would decrease v e r t i c a l mixing, and decreased density in the surface waters. The decreased density of the surface waters between May and July was established as an annual phenomenon by Herlinveaux (1962) based on 29 years of data. 5) Increased sedimentation may also be related to the increased sinking rate of senescent a l g a l c e l l s following the exhaustion of nitrat e in the surface layers. 97 In a study of biochemical changes of sedimented matter, Iseki et a l . (1980) noted that in the spring, decomposition was greatest in the upper 10 m. P a r t i c l e composition in the upper 10 m consisted of phytoplankton c e l l s , zooplankton carcasses and zooplankton fecal p e l l e t s , whereas in deeper waters the p a r t i c l e s consisted mainly of fecal p e l l e t s . Takahashi et a l . (1977) stated that frequent mixing or exchange of surface and deep waters occurred between spring and autumn and that these water movements introduced some phytoplankton seed populations and nutrients into the euphotic zone. 6) Despite the presence of large numbers of c e l l s in the phytoplankton, T. nordenskioeldi i i s rare in the sediments. From five Saanich Inlet surface sediment samples taken in May 1981, T. nordenskioeldii was absent in three samples and represented less than 2% of the t o t a l count in the other two samples. The lack of valves in three of the samples i s p a r t i a l l y a function of the counting technique. Counting approximately 200 valves from each sample only allows determination of dominant species. Counting larger number of valves per sample would probably increase the incidence of T. nordenskioeldi i . Nonetheless, i t would remain a rare spec ies . W.A. Dawson counted diatoms from 22 laminae in a 19m core representing 3100 (±200) years B.P. from Saanich Inlet (Gucluer and Gross 1964). Although T. nordenskioeldi i was l i s t e d as one of the i d e n t i f i e d species, i t was never referred to as one of 98 the dominant species. From another core taken in Saanich Inlet by Dr. S.E. Calvert, t h i s author counted approximately 200 valves from each of fiv e levels (approximately 20, 30, 50, 200, and 220 cm). At a l l l e v e l s , T. nordenskioeldi i represented <5% of the t o t a l valve count. The results from the two cores and the surface sediment samples suggest that T. nordenskioeldii i s probably always a rare species in the sediments. Given the above set of data, i t is possible to erect an hypothesis which appears to explain the discrepancy between the proportion of T. nordenskioeldii found in the water column and that found in the sediments. 1) -Vegetative c e l l s . The large population of T. nordenskioeldi i which exists during the spring bloom appears to be r e s t r i c t e d to the upper 10 m of the water column, which i s the layer where the highest l e v e l of decomposition occurs. C e l l s did not remain intact in fecal p e l l e t s or after being placed in the dark. This suggests that vegetative c e l l s are a short-term stage, readily susceptible to d i s s o l u t i o n once they leave the euphotic zone. In other words, vegetative c e l l s of T. nordenskioeldi i may be p r e f e r e n t i a l l y dissolved r e l a t i v e to other species which remain intact after passage through a zooplankter's gut or have a long-term quiescent vegetative stage. 2) Resting spores. In terms of l i f e strategies, T. nordenskioeldii may not require long-term survival of the 99 vegetative stage as a result of the a b i l i t y to produce resting spores, which are t h e o r e t i c a l l y the long-term survival stage. It appears that nitrogen concentrations may never be low enough to induce large-scale production of resting spores. When, toward the end of the spring bloom, some forms of nitrogen (e.g., nitrate) reach a low concentration, the temperature has increased to a l e v e l which may i n h i b i t the formation of resting spores. This suggests that production of resting spores may be very low. The temperature of the water column below 50 m remains at approximately 8°C throughout the year (Herlinveaux 1962). Assuming that a few of the valves found in the sediments were part of viable resting spores which could remain viable for approximately one year at 8°C (extrapolation from Durbin's 1978 data), then, given the upward v e r t i c a l mechanisms discussed above, T. nordenskioeldi i i s assured of su r v i v a l . In summary, the discrepancy between the biocoenoses and thanatocoenoses appears to have a two-part explanation: p r e f e r e n t i a l dissolution of the vegetative stage, and production and preservation of a low number of resting spores s u f f i c i e n t to ensure the continued existence of T. nordenskioeldi i . How can this discrepancy be interpreted in l i g h t of other sediment and phytoplankton d i s t r i b u t i o n studies of T. nordenskioeldii? In terms of general phytoplankton d i s t r i b u t i o n , T. nordenskioeldi i i s r e s t r i c t e d , with few exceptions, to the northern hemisphere above approximately 40°N (Hasle 1976; G u i l l a r d and Kilham 1977; Baars 1979). 100 More s p e c i f i c a l l y , the d i s t r i b u t i o n pattern follows the coastlines of the North At l a n t i c and P a c i f i c Oceans. Along the western A t l a n t i c coast, T. nordenskioeldii is a dominant in the phytoplankton from Long Island to the Gulf of Maine (Conover 1956; Pratt 1959; Riley and Conover 1967; Smayda 1973). In the eastern A t l a n t i c , T. nordenskioeldii is a dominant in the fjords, along the coast of Norway, and in the Norwegian Sea (Braarud et a l . 1974; Braarud 1976; Braarud and Nygaard 1980). In numerous other Central and North A t l a n t i c studies reviewed by G u i l l a r d and Kilham (1977), T. nordenskioeldi i i s not l i s t e d as a dominant in the phytoplankton. A similar phytoplankton d i s t r i b u t i o n pattern for T. nordenskioeldii occurs in the North P a c i f i c . T. nordenskioeldii is a dominant along the northwestern coast, through the Bering Sea, and along the northeastern coast (Kozlova and Mukhina 1967; Hasle 1976; Shim 1976; G u i l l a r d and Kilham 1977; Karentz and Mclntire 1977; Baars 1979). The zone in which T. nordenskioeldii is a dominant member of the phytoplankton in the North P a c i f i c has been designated the Subarctic by Kozlova and Mukhina (1967). The study area i s in t h i s zone. In a north-south transect through the northeast P a c i f i c , Venrick (1971) does not l i s t T. nordenskioeldii in any dominant oceanic or n e r i t i c phytoplankton assemblage. In transects across the Subarctic North P a c i f i c , Taylor and Waters (1982) showed that T. nordenskioeldi i was abundant in the western Gyre but was absent or had low abundance in the Alaskan Gyre, which 101 coincides with Venrick's e a r l i e r findings. In a study in the Bering Sea by Kozlova and Mukhina (1967), T. nordenskioeldi i was i d e n t i f i e d as a dominant n e r i t i c species in the surface phytoplankton, but between 500 m and 2000 m none of the dominant n e r i t i c species were found. This appears to be explained by the generalization made by Kozlova and Mukhina (1967), Jouse et a l . (1971), and L i s i t z i n (1971) that n e r i t i c species dissolve more readily in a long water column than oceanic species because they are more weakly s i l i c i f i e d , but i t does not explain the absence of T. nordenskioeldii from the thanatocoenoses in r e l a t i v e l y shallower areas. If depth of the water column were the only c r i t e r i o n , then T. nordenskioeldi i should be important in shallow-water assemblages but not in deep-water assemblages. Sediment studies from the North A t l a n t i c (Kolbe 1955; Maynard 1976) do not l i s t T. nordenskioeldii, probably because they are r e s t r i c t e d to deep-water areas. Resting spores of T. nordenskioeldii and other species were noted in the sediments of Narragansett Bay but neither viable spores nor empty frustules persisted in the sediments for more than a few weeks (Hargraves and French 1975). In the thanatocoenoses of the North P a c i f i c , T. nordenskioeldii is a member of the Arctoboreal diatom complex (Jouse et a l . 1971). The area of occurrence of the Arctoboreal diatom complex closely resembles the phytoplankton d i s t r i b u t i o n pattern of T. nordenskioeldii ( i . e . , along the ocean's northern margins including the study area). Kozlova and Mukhina (1967) c a l l a similar complex for the same area the Artie-Boreal 1 02 n e r i t i c species. The complex represents from 0.5 to 18.2% of the sediment assemblage. In a study of the d i s t r i b u t i o n of diatoms in the surface sediments of the Bering and Okhotsk Seas, Sancetta (1981) found that T. nordenskioeldii occurred throughout the area but was most important in Anadyr Bay and the Bering S t r a i t (both have a depth of approximately 50 m). The abundance of T. nordenskioeldii in the sediments of Anadyr Bay i s also noted by Kozlova and Mukhina (1967). Sancetta noted that the ins i g n i f i c a n c e of the Sea-Ice Assemblage ( i . e . , Nitzschia spp. and T. nordenskioeldi i) in the Sea of Okhotsk was probably due to the lack of shallow-water samples. Indeed, Jouse et a l (1971) state that the Arctoboreal diatom complex i s t y p i c a l of sediments in the insular shoals of the Sea of Okhotsk. The depth rela t i o n s h i p appears to be correct; T. nordenskioeldii i s important in the shallow-water assemblages of the Bering S t r a i t , Anadyr Bay, and the Sea of Okhotsk, but not in the deep-water assemblages of the Bering or Okhotsk Seas. A sediment study from the Sea of Japan (Tanimura 1981) adds another factor - temperature. The Sea of Japan has two d i s t i n c t temperature regions: a cold-current region along the northwest boundary, separated by a fron t a l zone from a warm-current region along the southeast boundary. Tanimura (1981) s t a t i s t i c a l l y demonstrated an inverse re l a t i o n s h i p between surface water temperature and the r e l a t i v e frequency of a three-species complex which included T. nordenskioeldii. The l a t t e r was found throughout the Sea of Japan sediments at a l l depths (maximum 103 depth approximatley 3000 m) but reached the highest abundance (5-15%) in the northern cold-current region (approximately 200-3000 m). In the southern warm-current (approximately 200-3000 m), where there were many more sediment samples taken than in the north, T. nordenskioeldii was present but not abundant. This shallow-water pattern i s similar to the B r i t i s h Columbia pattern where depths do not reach 1000 m. In another warm shallow-water area, the same pattern occurs. In the Yaquina Estuary, Oregon, an annual spring bloom of T. nordenskioeldii has been reported (Karentz and Mclntire 1977); yet an extensive investigation of t i d a l f l a t s sediments (Riznyk 1973) revealed no T. nordenskioeldi i , although other planktonic species were found. Temperature has several possible important e f f e c t s . F i r s t , temperature is d i r e c t l y related to the dissolution rate of s i l i c a (Lewin 1961). Second, b a c t e r i a l a c t i v i t y may be influenced by i t , thus causing a slower breakdown of an organic casing, i f present, at lower temperatures. Third, temperature i s inversely related to both production and v i a b i l i t y of resting spores (Durbin 1978). Although other factors (e.g., l i g h t and nutrients for growth of T. nordenskioeldi i and low nitrogen concentrations for resting spore production) are involved, the combination of depth and temperature may explain the varying preservation patterns of T. nordenskioeldi i in surface sediments. In summary, the following sediment patterns occur in the northern coastal zones where T. nordenskioeldi i i s a dominant in 104 the phytoplankton. In deep-water regions, few i f any vegetative c e l l s or resting spores of T. nordenskioeldi i reach the sediments; they are dissolved in the water column. In cold shallow-water regions (e.g., Anadyr Bay, the shoals of the Sea of Okhotsk, the northern Sea of Japan), T. nordenskioeldii has increased abundance in the sediments possibly as a result of the combination of increased resting spore production and decreased depth. In warmer shallow-water regions (e.g., the southern Sea of Japan and most of the B r i t i s h Columbia study area), T. nordenskioeldi i has a sediment pattern similar to deep-water regions possibly as a result of temperature i n h i b i t i o n of production of resting spores. II . DISTRIBUTIONAL PATTERNS OF SELECTED SPECIES Recurrent Groups A and B (Table 3) are good r e f l e c t i o n s of a common n e r i t i c and estuarine planktonic community. Skeletonema costatum, Thalassiosi ra spp. and/or Chaetoceros spp. are major components of the phytoplankton in many boreal and temperate coastal waters and estuaries (see review in Smayda 1980) and a major component of sediment assemblages in coastal waters and bays (DeVries and Schrader 1981; Garrison 1981; Schuette and Schrader 1981). Of the twelve planktonic species in Group A, eight are members of n e r i t i c planktonic recurrent groups established for the North P a c i f i c . In a study of phytoplankton d i s t r i b u t i o n in the Juan de Fuca S t r a i t and the southern S t r a i t of Georgia, Shim (1976) established f i v e recurrent groups. Ten of the species in Group A are in these groups. Shim's Group I constituted one of the 105 major components of the diatom community and usually occurred throughout the year at a l l stations and a l l depths. Group II were cold water species occurring from spring through late f a l l at a l l stations and a l l depths. Group III occurred only in the spring and summer, Group IV "frequently", and Group V in the spring through the f a l l , at a l l stations. The temporal d i s t r i b u t i o n of Group A species in Puget Sound, Washington (Phifer 1934a) which connects to the southern S t r a i t of Georgia, is l i s t e d in Table 3. In quantitative terms, there is good agreement between the thanatocoenoses and Shim's (1976) f i r s t four groups. Groups I-IV consist of 80 species and forms, 66% of which are also found in the sediment assemblages. Ten of the 80 species are Chaetoceros spp. , whose resting spores may be included in Chaetoceros u-spp. group. If these are removed, then there i s a 76% agreement between the thanatocoenoses and the biocoenoses. The dominant species (Skeletonema costatum, Thalassiosira  aest i v a l i s f. pac i f ica, Chaetoceros spp. as a group, Thalassionema nitzschioides, and Paralia sulcata) in the sediments are also the dominant species in the phytoplankton of the study area (Saanich In l e t : Takahaski et a l . 1977; Burrard Inl e t : Stockner and C l i f f 1979; Howe Sound: Stockner et a l . 1977; J e r v i s I n l e t : A.G. Lewis, personal communication, 1982; Pendrell Sound: Stockner and C l i f f 1975; Hotham Sound: Stockner and C l i f f 1975; and Indian Arm: Buchanan 1966; S t r a i t of Georgia: Legare 1957; Shim 1976; Stockner et a l . 1979). It i s not the presence or absence of these species in the surface 106 sediments, but rather their r e l a t i v e proportions, which determine their d i s t r i b u t i o n a l patterns. The d i s t r i b u t i o n of the dominant species in the sediments has unique patterns. To some extent these patterns can be associated with various factors (e.g., primary productivity or upwelling); but extensive analyses of these patterns are limited by i n s u f f i c i e n t phytoplankton data p a r t i c u l a r i l y for the northern waters. Since the patterns of absolute and r e l a t i v e abundance coincide to a large extent, only r e l a t i v e abundance w i l l be used for the discussion unless there i s a marked di fference. Skeletonema costatum S. costatum is a common, often dominant, n e r i t i c phytoplankton species found in both hemispheres (Hasle 1973; G u i l l a r d and Kilham 1977), yet i t i s almost never mentioned in sediment studies. S. costatum i s considered a d i s s o l u t i o n -sensitive species (Kozlova and Mukhina 1967; Schuette and Schrader 1979) and is not known to form resting spores (Smayda and Mitchel-Innes 1974). However, Hargraves and French (1975) note that S. costatum, found in the sediments of Narragansett Bay, are p h y s i o l o g i c a l l y similar to most diatom resting spores, although there i s no morphological difference. Dark survival appears to be inversely related to temperature; maximum survival of S. costatum was 4, 9, and 24 weeks at 20°C, 10°C, and 2°C, respectively (Antia 1976). There i s some taxonomic confusion between S. costatum and' S. tropicum . This author agrees with the conclusions of 1 07 Hasle 1s (1973) extensive study: the taxonomic status of S. tropicum i s unclear and i t cannot be distinguished from S. costatum at present. Kozlova and Mukhina (1967) state that S. costatum i s abundant in the North Tropical phytoplankton complex in the northern P a c i f i c in the 0-100 m layer, but most c e l l s dissolve between 300-500 m. I f , for the sake of argument, 500 m i s considered the c r i t i c a l d i s s o l u t i o n depth for S. costatum in the coastal zone of the North P a c i f i c , then S. costatum should only be found in the shallow sediments. Although S. costatum i s l i s t e d as abundant in the phytoplankton of the Bering and Okhotsk Seas (Guillard and Kilham 1977), i t i s not mentioned in shallow sediment studies for these areas (Jouse et a l . 1971; Sancetta 1981). In B r i t i s h Columbia sediment samples, which range between 15 and 706 m, S. costatum is a dominant, with r e l a t i v e abundance as great as 60%. If depth i s the c o n t r o l l i n g factor, there should be a negative relationship between abundance and depth. Instead, there i s a positive relationship within the i n l e t s , not only for S. costatum (Figs. 24-26; the gradients of r e l a t i v e abundance closely p a r a l l e l those of absolute abundance and are l i s t e d in Appendix D), but also for t o t a l diatoms (Figs. 27-29). This positive r e l a t i o n s h i p may be related to accumulation of fine p a r t i c l e s in deep basins, which w i l l be discussed in Part II, and primary productivity patterns. S. costatum increases in abundance from the northern Zone C to the southern Zone A (Fig. 11). Mean depth (± SD)in the 1 08 J I L M H M H ~ 1976 1981 Z INDIAN ARM SAANICH INLET _ l I 1 I I I I 1 L M H HOWE SOUND 1974 Figure 24. Absolute abundance of Skeletonema costatum and depth of i n l e t s i t e s in Zone A. A absolute abundance (v/gdw = valves/gram dry weight), D = depth, M = mouth of i n l e t , Md = middle, and H = head. 1 09 • » ' ' I I I 1 1 L . ^ JERVIS INLET .=1980 •=1981 HOTHAM SOUND f I • I I 1 1 1 L -H M H Md H NARROWS SECHELT SALMON INLET INLET INLET Figure 25. Absolute abundance of i n l e t s i t e s in Zone B. A valves/gram dry weight), D middle, and H = head. of Skeletonema costatum and depth = absolute abundance (v/gdw = depth, M = mouth of i n l e t , Md = 1 1 0 0 200 400 600 D M H KNIGHT INLET M Md BUTE INLET x LU Q 0 200 400 600 10? 10 6 105 f LU o < Q Z> m < 1 0 7 L U 3 10' o 6 OQ < 10= M TOBA INLET H M H PENDRELL SOUND Figure 26. Absolute abundance of Skeletonema costatum and depth of i n l e t s i t e s in Zone C. A absolute abundance (v/gdw valves/gram dry weight), D = depth, M = mouth of i n l e t , Md = middle, and H = head. 1 1 1 J 1 1 1 I L HOWE SOUND 1977 INDIAN ARM z> —I 1 1 I L M SAANICH INLET H J —I 1 1 1 I I I L M HOWE SOUND 1974 H Figure 27. Total absolute abundance and depth of i n l e t s i t e s in Zone A. A = absolute abundance (v/gdw = valves/gram dry weight), D = depth, M = mouth of i n l e t , Md = middle, and H = head. 1 1 2 Figure 28. Total absolute abundance and depth of i n l e t s i t e s in Zone B. A = absolute abundance (v/gdw = valves/gram dry weight), D = depth, M = mouth of i n l e t , Md = middle, and H = head. 1 1 3 E x »— Q_ UJ Q 100 300 500 700 h A 8 7 6 5 A 3 2 1»10l M H KNIGHT INLET M BUTE Md INLET 100 300 500 700 r A h D 8 7 6 5 3 H2 ",2x10 ,6 J TD cn LU O < < LU O LO CD < 1*10; M TOBA INLET H M H PENDRELL SOUND Figure 29. Total absolute abundance and depth of i n l e t s i t e s in Zone C. A = absolute abundance (v/gdw = valves/gram dry weight), D = depth, M = mouth of i n l e t , Md = middle, and H = head. 1 1 4 Zones A, B, and C i s 200±60 m, 400±170 m, and 390±220 m, respectively. Mean depth i s lowest and r e l a t i v e abundance of S. costatum is highest in Zone A; however, t o t a l diatom abundance and primary productivity (Stockner et a l . 1979) are also highest in Zone A. If depth were the only factor, Zones B and C should have a similar abundance since t h e i r mean depths are similar; yet both absolute and r e l a t i v e abundances of S. costatum are s i g n i f i c a n t l y greater in Zone B than in Zone C. The lack of a consistent inverse relationship between depth and abundance suggests that, in the B r i t i s h Columbia area, depth i s not the c o n t r o l l i n g factor in the preservation of S. costatum. The complexity of this preservation problem becomes evident with a consideration of other possible c o n t r o l l i n g factors: primary productivity, anoxic sediments,and population v a r i a t i o n . 1. In a study of surface sediments in the coastal area off Peru, Schuette and Schrader (1979) considered S. costatum as an indicator species of areas of high productivity induced by coastal upwelling. S. costatum had very low r e l a t i v e abundance in the Peruvian sediments (^1%) and was found in surface sediments as deep as 3000 m. The correlation between increased r e l a t i v e abundance and primary productivity, as noted above, i s also found in B r i t i s h Columbia sediments. 2. In a varved sediment core from Saanich Inlet, which i s p e r i o d i c a l l y anoxic, S. costatum has a r e l a t i v e abundance as high as 80% in the l i g h t laminae, and i s also found in the dark laminae (<35%) (Gucluer and Gross 1964). In comparison, in the Gulf of C a l i f o r n i a , laminated diatomaceous sediments are found 1 1 5 on the slopes of the basins where they intersect the oxygen minimum in the water column (Calvert 1964). The depth of the water column above the laminated sediments is approximately 400-1400 m. S. costatum i s not found in the sediments, but i t i s dominant in the phytoplankton (Calvert 1964 and 1966). Of course, temperature may be involved in the discrepancy noted above, as temperature i s d i r e c t l y related to dissolution (Lewin 1961). However, i f temperature were the only c o n t r o l l i n g factor, S. costatum should be abundant in the sediments of the cold Bering and Okhotsk Seas; and i t i s not. 3. The last factor, and most d i f f i c u l t to assess, i s population v a r i a t i o n . S. costatum i s one of the most studied diatom species. The a v a i l a b i l i t y of this extensive l i t e r a t u r e and the presence of the species in numerous culture c o l l e c t i o n s allowed Hasle (1973) to determine that there i s some variation in diameter and pervalvar axis of S. costatum populations. Diameter not only varies geographically but also seasonally (Hasle 1973) and during growth in culture (Smayda and Boleyn 1965; Harrison et a l . 1977). Genetic, physiological, and dark survival differences have also been shown between seasonal and geographical populations of S. costatum (Hargraves and French 1975; Gallagher 1980 and 1982). If these population variations are also related to preservation of S. costatum, and, by implication, other species, then comparison of sediment assemblages for d i f f e r e n t areas may be very d i f f i c u l t . There is no information on this problem. In summary, the d i s t r i b u t i o n a l pattern of S. costatum in 1 16 B r i t i s h Columbia sediments seems to be related to primary productivity patterns of the zones. There is no consistent pattern within i n l e t s or between i n l e t types. Dissolution with depth does not appear to be a c o n t r o l l i n g factor. It i s unknown why there is a good co r r e l a t i o n for S. costatum between the biocoenoses and the thanatocoenoses in B r i t i s h Columbia sediments when there i s a very poor correlation in other coastal areas. Thalassiosi ra a e s t i v a l i s f. pac i f ica The d i s t r i b u t i o n of T. aest i v a l i s f. pac i f ica appears to be related to both zones and i n l e t type (Fig. 12). The greatest absolute abundance of T. a e s t i v a l i s f. p a c i f i c a i s in Zone A, which i s probably related to the higher primary productivity in th i s area. The greatest r e l a t i v e abundance is in Zones A and C, and in high runoff i n l e t s . There are no high runoff i n l e t s in Zone B. This suggests that some factor common to both Zones A and C and high runoff i n l e t s may be a f f e c t i n g the d i s t r i b u t i o n of T. a e s t i v a l i s f. pac i f i c a . One p o s s i b i l i t y i s that fro n t a l boundaries associated with increased nutrients occur in both the southern and northern ends (Zones A and C) of the S t r a i t of Georgia, these having been shown to be areas of high chlorophyll concentrations (Harrison et a l . in press). Another factor is that high runoff i n l e t s usually have runoff from g l a c i e r s or snowfields, which can affect the temperature, s a l i n i t y , and transparency. In the coastal upwelling areas off South West A f r i c a , T. a e s t i v a l i s in the sediments was associated with cold, n u t r i e n t - r i c h upwelled waters (Schuette and Schrader 1981). 1 17 Ignoring the taxonomic problems, the d i s t r i b u t i o n a l patterns of T. a e s t i v a l i s f. p a c i f i c a in the B r i t i s h Columbia sediments and Schuette and Schrader's (1981) results suggest that temperature and nutrients may be the c o n t r o l l i n g factors. Although the r e l a t i v e thickness of the frustule i s not s u f f i c i e n t to explain the preservation of some species, there is a strong correlation between preservation and heavily s i l i c i f i e d species. Three of the dominants (Thalassionema nitzschioides, Chaetoceros spp. resting spores, and Paralia sulcata) of the B r i t i s h Columbia sediments are heavily s i l i c i f i e d . Thalassionema nitzschioides If one had to choose a species which was consistently abundant in both the biocoenoses and the thanatocoenoses on a global scale, T. nitzschioides would seem to be a good choice. It has a world-wide d i s t r i b u t i o n and i s often a dominant in both the phytoplankton and the sediments ( G u i l l a r d and Kilham 1977). In the North P a c i f i c , T. nitzschioides is considered a cosmopolitan phytoplankton species (Venrick 1971) and i s a member of the Subtropical thanatocoenosis in the P a c i f i c (Jouse et a l . 1971) and the A t l a n t i c (Maynard 1976). In the southern S t r a i t of Georgia, Shim (1976) recorded T. nitzschioides as one of the most common and frequent species in the phytoplankton. Three studies associate the sediment d i s t r i b u t i o n of T. nitzschioides with "warmer" and/or "more salin e " surface waters (Weiss et a l . 1978; Sancetta 1981; Tanimura 1981). 1 18 Temperature and s a l i n i t y are r e l a t i v e ; for example, "warmer" in the Sea of Okhotsk is 10-12 °C (Sancetta 1981), which in B r i t i s h Columbia would be a "cooler" temperature. In B r i t i s h Columbia sediments, T. nitzschioides i s also dominant in both the phytoplankton and the sediments, and absolute abundance is greatest in the southern Zone A, decreasing northward to Zone C (Fig. 14). In general, temperature of the surface waters decreases to the north, while s a l i n i t y increases in the same sense. This suggests a temperature c o r r e l a t i o n ; however, primary productivity also decreases northwards. In the sediments off South West A f r i c a , T. nitzschioides i s associated with high productivity upwelling areas. Therefore, the greatest abundance of T. nitzschioides in B r i t i s h Columbia sediments appears to be correlated with the higher primary productivity, higher temperatures, and lower s a l i n i t y of Zone A. Chaetoceros spp. resting spores There are numerous species in the genus Chaetoceros , many of which are present and dominant in the phytoplankton in both hemispheres (Guillard and Kilham 1977). Many Chaetoceros spp. form resting spores and i t i s these resting spores rather than the vegetative frustules which are usually found in the sediments. As discussed in the Methods section, there are d e f i n i t e taxonomic problems in the i d e n t i f i c a t i o n of Chaetoceros spp. resting spores. Possibly as a result of these problems, Chaetoceros spp. resting spores are often lumped together in sediment studies. As a group Chaetoceros spp. resting spores preserve well in the sediments and often dominate assemblages 1 19 (Kozlova and Mukhina 1967; G u i l l a r d and Kilham 1977; Schuette and Schrader 1979 and 1981; Sancetta 1981). It has been suggested that resting spores are either long-term or overwintering stages (Durbin 1978) or short-term survival stages (Garrison 1981), and formation i s either a response to l i g h t l i m i t a t i o n (Durbin 1978), nutrient l i m i t a t i o n (Garrison 1981; Hollibaugh et a l . 1981), or photoperiod (Hollibaugh et a l . 1981). Garrison (1981) found that Chaetoceros spp. resting spores, many of which are also found in B r i t i s h Columbia , were present both in the sediments and the water column of Monterey Bay, C a l i f o r n i a , throughout the year, and formation of resting spores usually occurred between spring and f a l l . As a group, Chaetoceros spp. resting spores in the sediments have been associated with high primary productivity (Schuette and Schrader 1979 and 1981; Sancetta 1981), upwelling systems (Schuette and Schrader 1979 and 1981), and large fluctuations in surface water temperatures from spring to f a l l (Sancetta 1981). In the B r i t i s h Columbia sediments, ten Chaetoceros spp. resting spores were i d e n t i f i e d to species. The relat i o n s h i p between the biocoenoses and the thanatocoenoses varies from f a i r to good. In the phytoplankton of the southern S t r a i t of Georgia and the contiguous waters of Puget Sound, Shim (1976) l i s t s 26 and Gran and Angst (1931) 32 Chaetoceros taxa. Not a l l the Chaetoceros spp. l i s t e d by Shim (1976) and Gran and Angst (1931) are known to form resting spores and 16% of the resting 1 20 spores in the sediments could not be i d e n t i f i e d to species. The five Chaetoceros spp. resting spores, each of which have a count of >2% of the t o t a l resting spores count (Table 6), are a l l l i s t e d as common vegetative c e l l s in the phytoplankton (Shim 1 976) . There i s a good co r r e l a t i o n for C. radicans between the biocoenoses and the thanatocoenoses. Out of the 26 species and forms that Shim (1976) i d e n t i f i e d , only C. radicans was l i s t e d as "very abundant" in the phytoplankton. In the sediments, C. radicans represents 52% of the t o t a l resting spores count. There i s also a good cor r e l a t i o n between the biocoenoses and the thanatocoenoses for resting spores as a group. Chaetoceros spp. as a group are dominant members of the B r i t i s h Columbia phytoplankton p a r t i c u l a r l y during summer months (Buchanan 1966; Shim 1976; Stockner et a l . 1977 and 1979; Takahashi et a l . 1977; Huntley and Hobson 1978), and resting spores as a group are a dominant member of the sediment assemblages. Although resting spores are dominants in both the dark and l i g h t laminae of a core from Saanich Inlet (Gucluer and Gross 1964), their greatest abundance i s in the l i g h t laminae which corresponds to their peak phytoplankton abundance in summer months. Since the d i s t r i b u t i o n a l pattern throughout the study area of individual Chaetoceros spp. resting spores i s very similar to that of resting spores as a group (see Table 2), the following discussion w i l l only refer to resting spores as a group. 121 Chaetoceros spp. resting spores are the only dominant in the sediment assemblages which have consistent d i s t r i b u t i o n a l patterns in a l l three categories: within the i n l e t s , i n l e t types, and zones. 1) Within the i n l e t s . As a group Chaetoceros spp. resting spores increase in absolute abundance toward the mouth in a l l the studied i n l e t s except Saanich Inlet (see Appendix D). Two exceptions are C. a f f i n i s , which tends to increase toward the head, and C. didymus , which is l o c a l i z e d ; these two represent 8% and 2% of the t o t a l resting spores count, respectively. The w i t h i n - i n l e t pattern, including that in Saanich Inlet, exactly corresponds to the generalized s p a t i a l pattern of annual phytoplankton production established from a number of sources by Stockner et a l . (1979)(Fig. 4). This pattern also corresponds with upwelling which usually occurs at the mouth of i n l e t s (Pickard 1961). This association between abundance of Chaetoceros spp. resting spores in the sediments and high primary productivity, sometimes associated with upwelling, has been established in other areas (Schuette and Schrader 1979 and 1981; Sancetta 1981). 2) Inlet types and zones. Unlike the d i s t r i b u t i o n within the i n l e t s , which is predictable given the patterns established in other areas, the variation between the i n l e t types and zones may be more t y p i c a l of fjord-type estuarine systems. Chaetoceros spp. resting spores have their highest r e l a t i v e and absolute abundance in Zone B and in medium and low runoff i n l e t s (Fig. 13). This zonal pattern d i f f e r s from that of t o t a l 1 22 absolute abundance and some of the other dominant species which have a higher abundance in Zone A, the area of higher primary productivity. There appears to be a simple explanation for the resting spores pattern which is related to the successional pattern and the basis of Pickard's (1961) separation of i n l e t types. Seasonal succession in B r i t i s h Columbia usually consists of a spring bloom of Skeletonema costatum, Thalassiosi ra spp. and Chaetoceros spp. , followed by a mixed Chaetoceros spp. assemblage (sometimes accompanied by S. costatum), during summer and f a l l (Buchanan 1966; Shim 1976; Stockner and C l i f f 1976; Takahaski et a l . 1977; Huntley and Hobson 1978; Stockner et a l . 1979) . This successional pattern i s t y p i c a l of a number of areas (Gu i l l a r d and Kilham 1977; Garrison 1979 and 1981; Smayda 1980) . Besides varying surface s a l i n i t i e s at the head, Pickard (1961) also separates the i n l e t s on the bases of runoff: stored runoff i n l e t s ( i . e . , the high runoff i n l e t s in this study) have peak runoff during late May through June, while dir e c t runoff i n l e t s ( i . e . , the medium and low runoff i n l e t s in th i s study) have peak runoff which follows the coastal r a i n f a l l patterns ( i . e . , in A p r i l and September through December). The r e l a t i v e volume of runoff in the direc t runoff i n l e t s i s considerably lower than in stored runoff i n l e t s , as evidenced by the r e l a t i v e l y higher s a l i n i t y at the heads of medium and low runoff i n l e t s . Estuarine c i r c u l a t i o n i s in part a result of th i s runoff and i s p a r t i c u l a r l y powerful during peak runoff periods 1 23 ( i . e . , the freshet). The result of the freshet in relation to the phytoplankton is documented in Howe Sound, a high runoff i n l e t (Stockner et a l . 1977). In general, a spring bloom occurs along the length of Howe Sound. The decline of the bloom corresponds with the peak runoff of the freshet. Phytoplankton are removed from the upper and middle reaches of the i n l e t by the seaward moving upper surface layer. The result i s that the summer Chaetoceros spp. period i s limited to the stations at the mouth of the i n l e t . The pattern i s di f f e r e n t in medium and low runoff i n l e t s ( i . e , the di r e c t runoff i n l e t s ) . In two medium runoff i n l e t s (Jervis I n l e t : A. G. Lewis, personal communication, 1982; Indian Arm: Buchannan 1966) and Saanich Inlet (a low runoff i n l e t : Takahashi et a l . 1977: Huntley and Hobson 1978; Hollibaugh et a l . 1981), the summer Chaetoceros spp. period can occur along the length of the i n l e t . The combination of successional and runoff patterns appears to explain the d i s t r i b u t i o n of Chaetoceros spp. resting spores in the sediments of the i n l e t types and zones. Medium and low runoff i n l e t s have the highest abundance of resting spores because Chaetoceros spp. can po t e n t i a l l y produce resting spores over a greater area. Zone B has the highest abundance of resting spores because i t contains two of the three medium runoff i n l e t s , one low runoff i n l e t , and no high runoff i n l e t s . 1 24 Paralia sulcata There have been two previous studies of the d i s t r i b u t i o n of P. sulcata in estuarine surface sediments. In the Bay of Vigo (Spain), Margalef (1969) showed a positive c o r r e l a t i o n between valve diameter and primary productivity, and a negative co r r e l a t i o n between valve diameter and s a l i n i t y of the surface waters. In the Hudson Estuary (U.S.A.), a positive relationship was shown between the dominance of P. sulcata in the sediments and s a l i n i t y of the surface waters (Weiss et a l . 1978). According to the MacDonald-Pfitzer rule (Drebes 1977), the size of diatoms should decrease with each successive vegetative d i v i s i o n . At a c r u c i a l minimum size and under special environmental conditions, auxosporulation occurs and results in restoration of maximum valve s i z e . Observations of auxosporulation suggest that i t is primarily a sexual process (see review on sexuality in Drebes 1977). Vegetative c e l l enlargement can also occur; however, some diatoms can undergo unlimited vegetative d i v i s i o n without s i g n i f i c i c a n t size v a r i a t i o n (Paasche 1973a; Drebes 1977). Based on the d i s t r i b u t i o n of P. sulcata in the sediments, Margalef (1969) suggested that discrete size morphs existed as a result of d i f f e r e n t i a l selection by several environmental factors. A number of studies have related the d i s t r i b u t i o n of, and changes in, valve size in the biocoenoses and thanatocoenoses to environmental factors and physiological properties of diatoms: nutrients (Taylor 1966; Paasche 1973a; Hecky and Kilham 1974; Harrison et a l . 1977; Malone 1980), grazing (Wimpenny 1973), 1 25 photosynthetic rate (Taguchi 1976), turbulence (Bellinger 1977), v e r t i c a l water movement (Semina 1972), equatorial upwelling (Burckle and McLaughlin 1977), respiration rate (Laws 1975), sinking rate (Smayda 1970), s a l i n i t y (Williams 1964), and temperature (Arrhenius 1952; Kolbe 1954). P. sulcata i s a cosmopolitan, marine species found predominantly in l i t t o r a l and n e r i t i c sediments, but i t also occurs in the phytoplankton ( i . e . , i t i s tychopelagic) (Cupp 1943; Mclntire and Moore 1977). The thick-walled, d i s s o l u t i o n -resistant, s i l i c e o u s valves of P. sulcata are usually well-preserved in the sediment and i t has been reported as a dominant in a number of studies (Maynard 1976; Sancetta 1981; Schuette and Schrader 1981). P. sulcata is found in the l i t t o r a l and s u b l i t t o r a l zones of the study area (personal observation) and there are many reports from other areas of the species being benthic (see review in Mclntire and Moore 1977; Sullivan 1978). P. sulcata is predominantly benthic, but i t can be a major constituent of the f a l l and winter phytoplankton (Conover 1956; Riley 1967; Shim 1976). Conover (1956) found that P. sulcata grew best in low l i g h t , with enrichment, and at temperatures "of at least 7°C". By maintaining a numerically constant population throughout the year in the phytoplankton, P. sulcata becomes dominant in the f a l l and winter by default as the spring bloom species are absent or in low numbers (Riley and Conover 1967; Shim 1976). In the study area and adjacent waters P. sulcata i s found throughout the year in the phytoplankton (Phifer 1934a; 126 Legare 1957;Buchanan 1966; Shim 1976) and at a l l depths between 0-175 m (Phifer 1934b; Shim 1976). The general d i s t r i b u t i o n of P. sulcata in the phytoplankton in the B r i t i s h Columbia i n l e t s appears to be strongly related to s a l i n i t y . In the southern section of the S t r a i t of Georgia, the d i s t r i b u t i o n of P. sulcata was negatively correlated with the low s a l i n i t y of stations in the Fraser River plume (Shim 1976). The surface s a l i n i t y in the Fraser River plume varies from 0-20%» and in the S t r a i t of Georgia from 24-29%*. P. sulcata i s not found in fresh water and was not reported in a study of the periphytic and planktonic algae of the lower Fraser River (Northcote et a l . 1975). In a three year study of the B r i t i s h Columbia coastal waters (Dilke et a l . 1979; Perry et a l . 1981), P. sulcata was only reported twice within the S t r a i t of Georgia (near Texada Island);however, i t was reported at 21 stations in contiguous waters north of the S t r a i t of Georgia (Discovery Passage, Johnstone S t r a i t , Queen Charlotte S t r a i t , and Hecate S t r a i t ) and once southwest of the S t r a i t of Georgia (Juan de Fuca S t r a i t ) . The s a l i n i t y of these s i t e s was very similar ( 30. 79±SD0 . 90%a) . There was no s i g n i f i c a n t difference (p>0.80) between the number of c e l l s / l i t e r from f a l l and winter samples (October-February, n=12) and spring and summer samples (April-September, n=12). As a predominantly benthic species (Mclntire and Moore 1977), the presence of P. sulcata in the phytoplankton i s dependent upon some form of v e r t i c a l transport. The areas north and south of the S t r a i t of Georgia have strong t i d a l mixing, 1 27 upwelling, and storm-induced mixing (Phifer 1934b; Parsons 1965; Thomson 1976) . P. sulcata was a major dominant in the dark laminae of a varved sediment core from Saanich Inlet (Gucluer and Gross 1964). The l i g h t laminae were considered to represent maximum diatom production in the spring and summer (Gucluer and Gross 1964); therefore, the dark laminae would represent the f a l l and winter season when Paralia sulcata i s known to be a dominant. In the Hudson Estuary, increasing dominance of P. sulcata in sediment assemblages was associated with increasing s a l i n i t y of the surface waters (Weiss et a l . 1978). Likewise, in the high and medium runoff i n l e t s in B r i t i s h Columbia , both s a l i n i t y and r e l a t i v e abundance of P. sulcata increase toward the mouth. In the low runoff i n l e t s , s a l i n i t y can be p e r i o d i c a l l y higher at the head and the r e l a t i v e abundance of P. sulcata i s highest at the head. As discussed above (see Results section), this gradient s h i f t , where a species or species-group reverses i t s head to mouth r e l a t i v e abundance in relation to i n l e t type, i s also seen in d i n o f l a g e l l a t e sediment d i s t r i b u t i o n a l patterns. In the S t r a i t of Georgia, r e l a t i v e abundance increases northward as does s a l i n i t y , from Burrard Inlet (x=0.9±SD0.1 % ) , which i s p e r i o d i c a l l y influenced by the Fraser River, to north of Texada Island (19.5%). The r e l a t i v e abundance of Zone C i s s i g n i f i c a n t l y higher than either Zones A or B. This may be the result of both the higher s a l i n i t y of Zone C and an influx of P. sulcata from contiguous coastal waters. 1 28 Absolute abundance of diatoms in surface sediments of oceanic and coastal zones has been p o s i t i v e l y correlated with primary productivity (Jouse et a l . 1971; Maynard 1976; Schuette and Schrader 1981). In the high and medium runoff i n l e t s , primary productivity i s highest at the mouth, where there may be upwelling. In the low runoff i n l e t s , primary productivity can be higher at the head. The d i s t r i b u t i o n of absolute abundance of P. sulcata follows t h i s pattern in the high and medium runoff i n l e t s and to some extent in the low runoff i n l e t s (Figs. 30-32) . Rivers from g l a c i a l runoff carry a large sediment load which results in the d i l u t i o n of sediment assemblages in the upper reaches of the high runoff i n l e t s and in the Fraser River area. The low absolute abundance of P. sulcata at the heads of high runoff i n l e t s may be in part a result of the sediment d i l u t i o n e f f e c t but low primary productivity and s a l i n i t y are also factors. Absolute abundance i s low at the heads of medium runoff i n l e t s which also have lower s a l i n i t y and primary productivity but lack a heavy sediment load. The high absolute abundance of P. sulcata in Zone A (Fig. 15) may be the result of the r e l a t i v e l y higher primary productivity in Zone A. In the sediments of the Bay of Vigo, Margalef (1958,1969) found that the valve diameter of P. sulcata increased with decreasing s a l i n i t y and increasing primary productivity. In the B r i t i s h Columbia i n l e t s , t h i s was not the case: valve diameter increased with increasing s a l i n i t y and increasing primary productivity. 1 29 Figure 30. Relative and absolute abundances of Paralia sulcata in Zone A s i t e s . R = r e l a t i v e abundance, A = absolute abundance (v/gdw = valves/gram dry weight), M = mouth of i n l e t , Md = middle, and H = head. 130 "D u 1 0 o 3 m < LU I — 3 _ l O LO 2io6 10 : 7 . J E R V I S INLET 1981 M H HOTHAM SOUND 1981 h_R_ A R H M NARROWS INLET i10 5 'o H Md H SECHELT SALMON INLET 1981 INLET 10 - 5 0 . o LU O < Q Z 3 CD < LU > LU or Figure 31. Relative and absolute abundances of Para l i a sulcata in Zone B s i t e s . R = r e l a t i v e abundance, A = absolute abundance (v/gdw = valves/gram dry weight), M = mouth of i n l e t , Md = middle, and H = head. 131 Figure 32. Relative and absolute abundances of Paralia sulcata in Zone C s i t e s . R = r e l a t i v e abundance, A = absolute abundance (v/gdw = valves/gram dry weight), M = mouth of i n l e t , Md = middle, and H = head. 1 32 Margalef (1969) notes other studies which show a relationship between increasing valve size of diatoms and decreasing temperature but could not find t h i s relationship for P. sulcata in Recent sediments in the Bay of Vigo. In the B r i t i s h Columbia i n l e t s , the smallest diameters are associated with the lowest temperatures (low nutrient g l a c i a l runoff at the heads of i n l e t s and the Fraser River). The range of P. sulcata valve diameter in the Bay of Vigo i s greater (10-64 nm) than in the B r i t i s h Columbia i n l e t s (8-29 Aim) . However, the mean temperatures of the B r i t i s h Columbia surface waters (Pickard 1961) are less than those of the Bay of Vigo (Margalef 1969), approximately 10°C and 17°C, respectively. In sediment core studies from the Bay of Vigo (Margalef 1956,1969), the mean diameter of P. sulcata increased with sediment depth. This increase was associated with cooler past climates. P. sulcata i s a dominant member of the dark laminae in the varved sediments of Saanich Inlet (Gucluer and Gross 1964). Thus, a core study in Saanich Inlet i s needed to investigate the relationship between valve diameter and paleotemperature in B r i t i s h Columbia . Rare Species There are a few rare species in B r i t i s h Columbia sediments which exhibit interesting d i s t r i b u t i o n a l patterns and have been l i s t e d as dominant or included in indicator assemblages for other areas. Nine Coseinodiscus spp. (see Appendix A) were i d e n t i f i e d from B r i t i s h Columbia sediments. Together these species 1 33 represent 0.5% of the t o t a l diatom count. Many of these species have been l i s t e d in sediment studies from both hemispheres (Guillard and Kilham 1977). A l l are l i s t e d as common but not abundant in the phytoplankton of the S t r a i t of Georgia, with the exception of C_^  perforatus which i s l i s t e d as rare (Shim 1976). In B r i t i s h Columbia sediments, these species had their highest abundance in Zones B and C, and in medium runoff i n l e t s . T halassiosira (Coscinodiscus) eccentrica i s a member of the Northboreal sediment diatom complex in the P a c i f i c (Jouse et a l . 1971), and i s considered a cosmopolitan plankton species in the northeast P a c i f i c (Venrick 1971). It i s an important member of North A t l a n t i c sediment assemblages (Gui l l a r d and Kilham 1977). Shim (1976) l i s t s T\_ eccentrica as very common but not abundant in the B r i t i s h Columbia phytoplankton. In B r i t i s h Columbia sediments, T\_ eccentr ica i s most abundant in Zone C and in high runoff i n l e t s . C y c l o t e l l a s t r i a t a i s common in the sediments of the Gulf of C a l i f o r n i a (Calvert 1966; Round 1967). In the B r i t i s h Columbia phytoplankton, Cj_ s t r i a t a i s common in f a l l and winter (Shim 1976). In B r i t i s h Columbia sediments, C_^  s t r i a t a i s almost e n t i r e l y r e s t r i c t e d to Zones A and B, and i s absent from sediments in Bute Inlet, Toba Inlet, and Knight Inlet, a l l of which are high runoff i n l e t s in Zone C. 1 34 SUMMARY OF PART I 1) The concept of d i f f e r e n t i a l d i s s o l u t i o n should be expanded to i n c l u d e not only the r e l a t i v e t h i c k n e s s o f . t h e f r u s t u l e but a l s o such f a c t o r s as the p h y s i c a l s t a t e of the diatom ( i . e . , fragmented, or whole empty f r u s t u l e s , or i n t a c t c e l l s ) , the orga n i c c a s i n g , l i f e stages, p h y s i o l o g i c a l and g e n e t i c d i f f e r e n c e s i n p o p u l a t i o n s of the same s p e c i e s , the r e l a t i o n s h i p between the v e r t i c a l d i s t r i b u t i o n of phytoplankton and zooplankton, and zooplankton feeding p r e f e r e n c e s . 2) T h a l a s s i o s i r a a e s t i v a l i s f. p a c i f i c a i s abundant i n the sediment assemblages of B r i t i s h Columbia and T. a e s t i v a l i s f. a e s t i v a l i s i s r a r e ; yet both are dominant i n the phytoplankton and appear e q u a l l y s i l i c i f i e d . The e x p l a n a t i o n fo r t h i s d i s c r e p a n c y appears to be r e l a t e d to the temporal d i s t r i b u t i o n of the two forms: T. a e s t i v a l i s f . p a c i f i c a c o i n c i d e s with peak abundance of p o s s i b l e p r e d a t o r s and r e l a t i v e l y c o o l e r temperatures. A l s o , there i s the p o s s i b i l i t y t h a t T. a e s t i v a l i s f . p a c i f i c a may remain i n t a c t w i t h i n f e c a l p e l l e t s . 3) T h a l a s s i o s i r a n o r d e n s k i o e l d i i i s a dominant i n the phytoplankton and u s u a l l y r a r e i n . t h e sediment assemblages of B r i t i s h Columbia . C e l l s and r e s t i n g spores are m o r p h o l o g i c a l l y very s i m i l a r . The e x p l a n a t i o n f o r the di s c r e p a n c y between the biocoenoses and the thanatocoenoses i s not c l e a r . R e l a t i v e l y warmer s u r f a c e temperatures may be i n h i b i t i n g p r o d u c t i o n of 1 35 resting spores in most of the study area. 4) The recurrent groups are t y p i c a l of n e r i t i c and estuarine communities seen in many areas. The dominant species in the sediment assemblages are also dominants in the phytoplankton. 5) Skeletonema costatum i s usually considered to be a diss o l u t i o n sensitive species and i s reported in very low abundance in other sediment studies. In B r i t i s h Columbia , S. costatum has high abundance in both the phytoplankton and the sediment assemblages. The reason for thi s good cor r e l a t i o n i s unknown; depth does not appear to be a c o n t r o l l i n g factor. The d i s t r i b u t i o n a l patterns of S. costatum p a r a l l e l those of the primary productivity patterns between the zones. 6) The greatest abundance of Thalassiosira a e s t i v a l i s f. p a c i f i c a i s in Zones A and C, and in high runoff i n l e t s . This pattern coincides with increased nutrient supply at front a l boundaries in the southern and northern ends (Zones A and C) of the S t r a i t of Georgia, and the r e l a t i v e l y lower temperatures of high runoff i n l e t s . This corresponds with a similar pattern described for coastal upwelling areas off South West A f r i c a . 7) Thalassionema nitzschioides is a dominant in both the phytoplankton and the sediment assemblages in B r i t i s h Columbia and many other areas. The greatest abundance i s in Zone A, the least in Zone C; thi s gradient p a r a l l e l s that for primary productivity, t o t a l absolute abundance in the sediments, and temperature. 8) The d i s t r i b u t i o n a l pattern of Chaetoceros spp. resting spores may be unique to fjord-type estuarine systems. The 1 36 highest abundance i s i n Zone B, i n medium and low runoff i n l e t s , and u s u a l l y at the mouth of i n l e t s . The i n c r e a s i n g abundance toward the mouth c o i n c i d e s with primary p r o d u c t i v i t y , s a l i n i t y , and temperature gradients w i t h i n the i n l e t s . The r e l a t i v e l y lower abundance i n high runoff i n l e t s , and therefore i n Zones A and C which contain a l l the high runoff i n l e t s , seems to be r e l a t e d to the removal of phytoplankton from w i t h i n the i n l e t by the freshet during the pe r i o d of peak Chaetoceros spp. r e s t i n g spore production. 9) The d i s t r i b u t i o n a l p a t t e r n of P a r a l i a s u l c a t a i s p o s i t i v e l y c o r r e l a t e d with primary p r o d u c t i v i t y and s a l i n i t y both i n B r i t i s h Columbia and other areas. In both B r i t i s h Columbia and the Bay of Vigo, valve diameter increased with i n c r e a s i n g primary p r o d u c t i v i t y ; however, the c o r r e l a t i o n with s a l i n i t y was negative i n the Bay of Vigo and p o s i t i v e i n B r i t i s h Columbia . 1 37 PART II The d i s t r i b u t i o n a l patterns of diatoms in the B r i t i s h Columbia sediments can be associated with a number of factors. The discussion of the relationships w i l l be divided into two sections: zones and i n l e t types. The Howe Sound (1974) set of samples have unique aspects which w i l l be discussed at the end of Part I I . ZONES Figure 33 shows the general variation of a number of factors between the zones (Hutchinson and Lucas 1931; Legare 1957; Pickard 1961; Stockner et a l . 1979). These trends, with the exception of s a l i n i t y , are reflected by the t o t a l absolute abundance of the sediment assemblages. Nutrients may also be associated with these patterns as the Fraser River (Zone A) contributes up to 30% of the t o t a l available nitrogen in the S t r a i t of Georgia (Harrison et a l . in press). However, nutrients are usually at a saturating l e v e l except during May to September when nitrogen may be l i m i t i n g (Shim 1976; Harrison et a l . in press). In the world's oceans, the pattern of quantitative d i s t r i b u t i o n of diatom valves in the surface sediments resembles the patterns of annual production of s i l i c a and primary productivity ( L i s i t z i n 1971). The highest abundances, both in the phytoplankton and in the surface sediments, are in the Antarctic zone. Table 11 i s a comparison of diatom absolute abundance in oceanic and coastal surface sediments. The mean 138 >-< LO LU cn 3 t— < cr LU CL LU o 3 Q o or o_ >-on < cr 0L LU Q LU LO Q t N B ZONE A Figure 33. Gradients of selected factors from Zone A to C. Direction of arrows indicates increasing values. 139 Table 11 . Comparison of the absolute abundance of diatom valves in oceanic and coastal sediments, v/gdw = valves/gram dry weight. P a c i f i c Ocean (Jouse et a l . 1971) B r i t i s h Columbia coast 25-50x106 v/gdw mid-ocean North P a c i f i c 5-25x106 v/gdw Atla n t i c Ocean (Maynard 1976) Subtropical zones 0-1X10 6 v/gdw Equatorial zone 0-10X106 v/gdw Coastal upwelling zone off South West Af r i c a (Schuette and Schrader 1981) 2-300X106 v/gdw Indian Ocean (Kozlova 1971) 2-10X10 6 v/gdw Southern Ocean (Jouse et a l . 1971) (Kozlova 1971) (Maynard 1976) >100X106 v/gdw 90->200X106 v/gdw 10-1,000x106 v/gdw 1 40 absolute abundance (±95% confidence intervals) of the three zones of the study area (A=77x10 6±20x10 6 v/gdw, B=56x10 6±13x10 6 v/gdw, C=38x10 6±14x10 6 v/gdw) correspond well with the range of values (Table 11) for the B r i t i s h Columbia coast established by Jouse et a l . (1971). The l a t i t u d i n a l d i s t r i b u t i o n of the dominant species in the B r i t i s h Columbia sediments i s similar to that found in the phytoplankton in Norway. In the near-coastal waters (62°N-69°N), Braarud and Nygaard (1980) i d e n t i f i e d two diatom s o c i e t i e s : the 'Skeletonema costatum society' and the 'mixed spring diatom society.' The S. costatum society i s t o t a l l y dominated by S. costatum. The mixed spring diatom society contains the same species but was dominated by Chaetoceros spp. and Thalassiosira spp. during fiv e of the eight sampling periods between 1968-1971. The dominant Thalassiosi ra spp. in the Norwegian waters is T. nordenskioeldi i . The S. costatum society was dominant in the southern coastal waters, but the mixed spring diatom society dominated in the north. In B r i t i s h Columbia sediments, S. costatum dominates in the southern Zone A, T. a e s t i v a l i s f. p a c i f i c a in Zone C, and Chaetoceros spp. dominate in the northern Zones B and C (Fig.34). The a b i l i t y of S. costatum to overwhelmingly dominate the annual phytoplankton cycle has been noted in Hardangerfjord, Norway (Braard 1976), and in the Long Island Sound, Narragansett Bay, and the Gulf of Maine (Gran and Braarud 1935; Pratt 1959; Riley and Conover 1967). No one factor has been established to explain the extraordinary importance of S. costatum in these 141 +s T J L +s s —i— S+ S * —i-To — + c c-+-j i i i 1 i — •4-+ To R „ C - c—+• s -*-s t - T ' T To + C L ^ — C j i 1 1 L 10 6 10 7 10 8 0 10 20 30 AO A B S O L U T E R E L A T I V E A B U N D A N C E (*/•) A B U N D A N C E (v/gdw) Figure 34. Relative and absolute abundances (±95% confidence intervals) of dominant species in Zones A, B, and C. S = Skeletonema costatum, S+ = S. costatum without Howe Sound (1974) data, T = Thalassiosira a e s t i v a l i s f. p a c i f i c a , C = Chaetoceros spp. resting spores, To = t o t a l absolute abundance" v/gdw = valves/gram dry weight, and A, B, and C are zones.. 1 42 areas (Smayda 1980). The inverse relationship between d i v e r s i t y and the r e l a t i v e abundance of S. costatum (Figs. 35-37) may be a r e f l e c t i o n of the a b i l i t y of S. costatum to dominate assemblages in B r i t i s h Columbia . Diversity i s greatest in Zones B and C (Fig. 9), while the abundance of S. costatum i s greatest in Zone A (Fig. 11). No other species has a consistent relationship with d i v e r s i t y . The f i r s t axis of the p r i n c i p a l coordinate analysis (PCoA) can be related to several of the l a t i t u d i n a l gradients (Figs. 38-39). S. costatum, which dominates in Zone A, i s in the l e f t half of axis 1; and Chaetoceros spp. , which dominates in Zones B and C, are in the right half (see Table 10 for c o r r e l a t i o n c o e f f i c i e n t s ) . The assemblages of both Saanich Inlet and Sechelt Inlet l i e between the two s o c i e t i e s . The d i s s i m i l a r i t y between the sediment assemblages in Sechelt Inlet and Jervis Inlet indicated by the PCoA may be related to the highly turbulent t i d a l jet which enters Sechelt Inlet across the shallow s i l l between the two i n l e t s (Lazier 1963). The s i t e s are c l e a r l y not d i s t r i b u t e d on the basis of i n l e t type. Pickard (1961) separated the i n l e t s on the basis of surface s a l i n i t y , and Legare (1957) stated that "the chief factor a f f e c t i n g the general d i s t r i b u t i o n of plankton (in the S t r a i t of Georgia) is the s a l i n i t y gradient." In the sense that the s a l i n i t y does increase from Zone A, where the Fraser River contributes 50-80% of the t o t a l freshwater runoff (Harrison et a l . in press), to Zone C, axis 1 can be associated with a 143 -J 1 1 l i i i • i M H HOWE SOUND 1974 Figure 35. Diversity and r e l a t i v e abundance of Skeletonema  costatum in Zone A i n l e t s . D = d i v e r s i t y , R = r e l a t i v e abundance, M = mouth of i n l e t , Md = middle, and H = head. 1 4 4 M H .E HOTHAM SOUND « I I I I 1 I 1 L. H M H Md H NARROWS SECHELT SALMON INLET INLET INLET Figure 36. Diversity and r e l a t i v e abundance of Skeletonema  costatum in Zone B i n l e t s . D = d i v e r s i t y , R = r e l a t i v e abundance, M = mouth of i n l e t , Md = middle, and H = head. 1 45 LU z < -z. 3 m < LU > LU cr M H KNIGHT INLET 2 0 r M Md BUTE INLET M H TOBA INLET M H PENDRELL SOUND to >-I— cr LU > Figure 37. Diversity and r e l a t i v e abundance of Skeletonema  costatum in Zone C i n l e t s . D = d i v e r s i t y , R = r e l a t i v e abundance, M = mouth of i n l e t , Md = middle, and H = head. 1 46 ^ SALINITY _ TEMPERATURE _ PRIMARY PRODUCTIVITY _ V/GDW IN SEDIMENTS ZONE A ZONES B AND C f l 8 3 Skeletonema costatum Chaetoceros spp. < ^ « Figure 38. Gradients of selected factors Direction of arrows indicates increasing valves/gram dry weight. and PCoA values. diagram. V/GDW = 147 _ ^ S A L I N I T Y _ T E M P E R A T U R E P R I M A R Y PRODUCTIV ITY _ V / G D W IN S E D I M E N T S Z O N E A Skeletonema costatum Z O N E S B A N D C Chaetoceros spp. <s / B E C H E L T V / X \ A C > * T N A X I S 1 SALMON—•) A I \ / B I / ' / * M C •cc ;QC HR / <^AANICH Hc\ S C • ' N A R R O W S \ 1 H B ^ \ / A X I S 2 P C * EG * *TS Q . C L 10 (/> O a> <_> o —^< O Figure 39. Gradients of selected factors and PCoA diagram. Direction of arrows indicates increasing values. AC = Agamemnon Channel, BI = Burrard Inlet, CC = Cordero Channel, DS = Desolation Sound, EG = East Gorge Harbor, HB = Hidden Basin, HC = Homfray Channel, HR = Heriot Bay, MC = Malaspina S t r a i t , PC = Pryce Channel, QC = Queen Charlotte S t r a i t , SC = S u t i l Channel, TN = Texada Island north, TS = Texada Island south, and V/GDW = valves/gram dry weight. 1 48 s a l i n i t y gradient. The higher s a l i n i t i e s are in the northern S t r a i t of Georgia and the s i t e s from th i s area are on the far right of axis 1. Axis 1 can also be related to the other factors which vary between the zones (Figs. 38-39). The l e f t side of axis 1 consists mainly of Zone A s i t e s which have r e l a t i v e l y higher primary productivity, phytoplankton abundance, surface water temperatures, and absolute abundance of diatoms in the sediments. The right side of axis 1 consists mainly of Zone B and C s i t e s which have, with the exception of s a l i n i t y , r e l a t i v e l y lower valves for these factors. It is d i f f i c u l t to relate axis 2 (Figs. 38-39) to anything other than the r e l a t i v e d i s t r i b u t i o n of T. a e s t i v a l i s f. p a c i f i c a and Chaetoceros spp. Zone B s i t e s and the S t r a i t of Georgia s i t e s , which are dominated by Chaetoceros spp. , are in the lower half of the diagram (Fig. 39). With the exception of Howe Sound (1974), the high runoff i n l e t s , which are for the most part dominated by T. a e s t i v a l i s f. p a c i f i c a , are in the upper ha l f . The Zone A si t e s overlap the two halves. There are also s i g n i f i c a n t d i s t r i b u t i o n a l patterns related to zones for marine l i t t o r a l and freshwater species but these are d i r e c t l y related to i n l e t type and w i l l be discussed below in the section on estuarine c i r c u l a t i o n . In summary, there are d e f i n i t e d i s t r i b u t i o n a l patterns related to the zones. The co r r e l a t i o n between primary productivity and absolute abundance of diatoms in the sediments i s almost predictable given the results of other sediment studies ( L i s i t z i n 1971; Maynard 1976; Sancetta 1981). A more 1 49 detailed understanding of phytoplankton d i s t r i b u t i o n in B r i t i s h Columbia sediment i s needed before the s i m i l a r i t y between Norwegian phytoplankton and B r i t i s h Columbia sediment assemblages can be explained. The r e l a t i v e influence of s a l i n i t y and temperature i s not well understood; but some species discussed in Part I appear to be influenced by these factors. INLET TYPES As noted above, PCoA does not c l e a r l y separate the assemblages on the basis of i n l e t type. Nor i s there any s i g n i f i c a n t difference between absolute abundance in the sediments between the i n l e t types. Diversity i s highest in medium and low runoff i n l e t s which may be related to the summer diatom populations which can occur along the length of these i n l e t s , while summer populations appear to be r e s t r i c t e d to the mouth of high runoff i n l e t s . There are some species associations with p a r t i c u l a r i n l e t types: 1) marine planktonic species have their highest abundance in medium runoff i n l e t s , 2) marine l i t t o r a l species in low and high runoff i n l e t s , 3) freshwater species in high runoff i n l e t s , 4) Chaetoceros spp. resting spores in medium and low runoff i n l e t s , and 5) Thalassiosira aest i v a l i s f. pac i f ica in high and medium runoff i n l e t s . There are gradients within the i n l e t s . Figure 40 i l l u s t r a t e s the d i s t r i b u t i o n of several parameters in a t y p i c a l B r i t i s h Columbia i n l e t . The greatest primary productivity, absolute abundance in the sediments, and often depth, a l l occur 150 < DEPTH <Z SALINITY < . TEMPERATURE <= PRIMARY PRODUCTIVITY <= V/GDW IN SEDIMENTS MOUTH HEAD Figure 40. Gradients of selected factors in a t y p i c a l i n l e t . Direction of arrows indicates increasing values. V/GDW = valves/gram dry weight. 151 at the mouths of the i n l e t s except in Saanich Inlet where a l l three coincide at the head (Lebrasseur 1954; Stockner and C l i f f 1975; Stockner et a l . 1979). High primary productivity and abundance of phytoplankton have been associated with nu t r i e n t - r i c h upwelling zones (Garrison 1979; Tont 1981). Upwelling in the study area usually occurs at the mouth of i n l e t s or near islands (Pickard 1961). Scale does not appear to affe c t the good cor r e l a t i o n between upwelling and the abundance of diatoms in the sediments. The r e l a t i v e l y smaller upwelling zones in B r i t i s h Columbia have a similar c o r r e l a t i o n to that in the larger upwelling zones along the south west coast of South America and A f r i c a (Schuette and Schrader 1979 and 1981). There are two factors which are probably a f f e c t i n g the sediment assemblages: varying sedimentation rates and estuarine c i r c u l a t i o n . 1. Sedimentation The sediment load in r i v e r s , p a r t i c u l a r l y g l a c i a l l y - f e d , high runoff r i v e r s , a f f e c t s the sediment assemblages by d i l u t i n g the rain of plankton. Two sediment samples from the head of Bute Inlet were e s s e n t i a l l y barren of diatoms, most probably because of sediment d i l u t i o n . Only a few of the i n l e t s have sedimentation accumulation data: Saanich Inlet 4-6 mm/year (Gucluer and Gross 1964) and 1.7 mm/year (Stephens et a l . 1967), and Bute Inlet and Jervis Inlet 0.35 mm/year (Pickard and Giovando 1960). Sediment d i s t r i b u t i o n studies (Lewis 1976, 1979) show that 1 52 a greater percentage of the s i l t - s i z e d p a r t i c l e s s e t t l e in the upper reaches of Jervis Inlet and Knight Inlet. Howe Sound, Knight Inlet, and Bute Inlet have the most comprehensive data. In the upper reaches of Howe Sound organic and inorganic sedimentation rates range from 0.3 to 28.2 g/m2/day and 6.8 to 1304.4 g/m2/day, respectively (Syvitski and Murray 1981). Along the length of Knight Inlet (head to mouth), summer sedimentation rates range from 3,700 g/m2/day (200 cm/yr) to 6 g/m2/day, and winter rates from 33 g/m2/day (26.8 cm/yr) to 0.1 g/m2/day (0.04 cm/yr) (Farrow et a l . in press). These rates were considered by the authors to be similar to those in Bute Inl e t . The sedimentation rates dropped markedly within 20 km of the head (Knight Inlet i s 80 km in length) both in the summer: 3,700 g/m2/day to 20 g/m2/day; and winter: 33 g/m2/day to 0.3 g/m2/day (Farrow et a l . in press). Increased sedimentation rates at the head of i n l e t s very l i k e l y decrease the absolute abundance of the sediment assemblages. This does not change the pattern of increasing absolute abundance toward the mouth, nor the co r r e l a t i o n with primary productivity which also increases toward the mouth. However, dire c t comparison of absolute abundance values at the head of i n l e t s should not be made because of the possible ef f e c t s of varying sedimentation rates. 1 53 2. Estuarine C i r c u l a t i o n The general temporal sequence of events for the phytoplankton in an i n l e t was discussed above in d e t a i l . B r i e f l y , the spring bloom begins between March and A p r i l and in high runoff i n l e t s continues within the whole i n l e t u n t i l the freshet commences between May and June. Phytoplankton at the head are removed by the seaward, freshwater surface layer. The effect of this process on the sediment assemblages i s that diatoms from the phytoplankton at the head may become part of the sediment assemblages at the mouth. Added to th i s possible mouthward horizontal transport i s the focusing of diatoms in deep basins. Most diatoms are in the s i l t size range (4-62 ym) and as fine p a r t i c l e s tend to be focused in the deep basins of the i n l e t s . This relationship between accumulation of diatoms and deeper waters has been shown in Monterey Bay, C a l i f o r n i a (Garrison 1981) and Saanich Inlet (Gucluer and Gross 1964). There i s a positive relationship between depth and t o t a l absolute abundance in a l l the studied i n l e t s (Figs. 27-29). Theoretically, the high absolute abundance at the mouth of i n l e t s could be the result of p a r t i c l e focusing in the deepest basins, which tend to be near the mouth, and estuarine c i r c u l a t i o n transporting the phytoplankton toward the mouth. However, Syvitski and MacDonald (in press) noted that sediment p a r t i c l e s have nearly v e r t i c a l descent paths in Howe Sound; although before deposition, the travel distances up and down i n l e t of a given p a r t i c l e may be large. Three sets of data suggest there i s a good s p a t i a l c orrelation between the 1 54 biocoenoses and the thanatocoenoses, and that physical processes are not seriously a l t e r i n g the sediment assemblages. A. Freshwater species There are two major sources of freshwater species to the sediments: the Fraser River, which contributes 50-80% of the t o t a l freshwater runoff to the S t r a i t of Georgia (Harrison et a l . in press); and rivers within the i n l e t s . Thirty of the 73 freshwater species found in the sediment assemblages were also found in a study of the periphytic and planktonic algae of the lower Fraser River (Northcote et a l . 1975)(see Appendix C for d i s t r i b u t i o n of freshwater species). Twelve of these species were only found in Howe Sound, Burrard Inlet, and Indian Arm (Zone A), and six other species were found in both Zones A and B. The remaining twelve species were found in a l l three zones and had 92 t o t a l occurrences, of which 71 were in Zones A and B. This suggests that the Fraser River may be a major source of freshwater species for Zones A and B. In Zones A and B, the large number of marine occurrences (114) of freshwater species also found in the Fraser River, compared to the low number of occurrences (39) of 22 species not found in the Fraser River, supports t h i s suggestion. In a l l the i n l e t s , the r e l a t i v e abundance of freshwater species tends to increase toward the freshwater source. (See Appendix D for head to mouth abundance of selected species and species-groups.) In Saanich Inlet, the highest r e l a t i v e abundance i s at the mouth, which receives sediment from the 155 Fraser and the Cowichan Rivers (Gross 1967). In Howe Sound (Fig. 41) the highest r e l a t i v e abundance i s at the head, the lowest i s in the middle section, and then the r e l a t i v e abundance increases again near the mouth. This pattern is also seen in Jervis Inlet, Knight Inlet, Toba Inl e t . The highest abundance can be associated with the rive r at the head of the i n l e t . The second highest abundance, which occurs at the mouth, i s either the result of sediment focusing, or, in the case of i n l e t s in Zones A and B, a combination of sediment focusing and allochthonous deposits from the Fraser River. The highest r e l a t i v e abundance of freshwater species in a l l the s i t e s i s in Burrard Inlet which receives a heavy sediment load from the Fraser River (Stockner and C l i f f 1979). The highest absolute abundance i s in Zone A which i s probably indicative of the influence of the Fraser River. Zones A and C, and high runoff i n l e t s have s i g n i f i c a n t l y greater r e l a t i v e abundances of freshwater species (Table 9 and F i g . 18). This is the result of the high runoff r i v e r s in these zones and i n l e t s . Likewise, the sediments of the Hudson Estuary, New York (Weiss et a l . 1978), a freshwater C y c l o t e l l a assemblage dominated at the head. The d i s t r i b u t i o n a l patterns of the freshwater species are indicative of their freshwater sources. Although there i s evidence that estuarine c i r c u l a t i o n , sediment focusing, and allochthonous species a l t e r the pattern somewhat, they do not t o t a l l y obscure the basic patterns. 1 56 L U O M H Figure 41. Relative abundance of freshwater species along the length of Howe Sound (1974). R = r e l a t i v e abundance, D = depth, M = mouth of i n l e t , and H = head. 157 B. Marine l i t t o r a l species The marine l i t t o r a l species appear to be indicative of the brackish water deltas at the mouth of r i v e r s . Zone C and low runoff i n l e t s have s i g n i f i c a n t l y greater r e l a t i v e abundance of marine l i t t o r a l species (Table 9 and F i g . 17). The s i t e s which have >10% r e l a t i v e abundance are a l l in Zone C, except Hidden Basin which i s in Zone B. Zone C may have a higher r e l a t i v e abundance because a greater number of sheltered, shallow s i t e s (e.g., East Gorge Harbor, Heriot Bay, and Pendrell Sound) were sampled in this zone. In the high runoff i n l e t s , the highest r e l a t i v e abundance i s at the head. The ri v e r s at the heads have large, shallow, p e r i o d i c a l l y brackish water deltas which are probably the source of the marine l i t t o r a l species. The mouth of Saanich Inlet also has a high r e l a t i v e abundance, presumably from the delta sources of the Fraser and Cowichan Rivers. Only six of the 81 marine l i t t o r a l species found in the sediments were also found in the Fraser River (Northcote et a l . 1975). The majority of their occurrences are in Zones A and B. C y c l o t e l l a s t r i a t a occurred at 66 sites,, only six of which were in Zone C. Again using Howe Sound (Fig. 42) as an example of a pattern also seen in Je r v i s Inlet and Toba Inlet, there i s a s i m i l a r i t y to freshwater species d i s t r i b u t i o n . Marine l i t t o r a l species tend to be deposited in areas influenced by delta sources. C. Dominant planktonic species 158 Figure 42. Relative abundance of marine l i t t o r a l species along the length of Howe Sound (1974). R = r e l a t i v e abundance, D = depth, M = mouth of i n l e t , and H = head 159 The large set of samples (24) from Howe Sound (1974) i s p a r t i c u l a r l y valuable because i t has concurrent phytoplankton data. The seasonal succession and species composition for a two year period (1973-1974) in Howe Sound showed that Thalassiosira spp., s p e c i f i c a l l y T. a e s t i v a l i s f. p a c i f i c a and T. a e s t i v a l i s f. a e s t i v a l i s , were the major dominants at most stations from March-July in 1973, and Skeletonema costatum from April-November in 1974 (Stockner et a l . 1977). During the 1974 spring bloom, primary productivity and diatom abundance were greatest at the mouth. At the two stations at the mouth of Howe Sound, S. costatum was a co-dominant with Chaetoceros spp. from August-November 1974. The Howe Sound sediment samples were taken in November 1974 and S. costatum was dominant at a l l 24 s i t e s . Figure 43 shows the absolute abundance of S. costatum, T. a e s t i v a l i s f. p a c i f i c a , and Chaetoceros spp. from the sediment assemblages along the length of Howe Sound. There i s a good c o r r e l a t i o n with Stockner et a l . ' s (1977) phytoplankton data. Skeletonema costatum i s always dominant, and T. a e s t i v a l i s f. p a c i f i c a and Chaetoceros spp. resting spores have the i r highest abundance at the mouth. These data demonstrate, l i k e that for freshwater and marine l i t t o r a l species, that there is a good s p a t i a l c o r r e l a t i o n between the biocoenoses and the thanatocoenoses. As discussed above, the actual values of absolute abundance may be misleading as a result of the possible d i l u t i o n e f f e c t . In the case of Howe Sound, both the mouth and the head receive increased 1 60 c n 5< _ mm < < 10c J: 5 0 UJ u jg ~ g 30 J D Lum C r < 101 M H Figure 43. Relative and absolute along the length of Howe Sound a e s t i v a l i s f. p a c i f i c a , S Chaetoceros spp. resting spores, v/gdw = valves/gram dry weight, D and H = head. abundances of dominant species (1974). T = Thalassiosira  Skeletonema costatum, C To = Total absolute abundance, = depth, M = mouth of i n l e t , 161 sediment loads which would decrease absolute abundance to varying degrees. Sediment focusing may also increase the absolute abundance in deep basins. Yet there remains a good corr e l a t i o n between the gradients of high absolute abundance in the sediments, primary productivity, and phytoplankton abundance (Fig. 44). Estuarine c i r c u l a t i o n and sediment focusing do not appear to be seriously changing the sediment assemblages. HOWE SOUND The existence of concurrent phytoplankton data for the Howe Sound (1974) set of sediment samples allows for an interesting c o r r e l a t i o n with varying yearly radiation l e v e l s and the determination of the temporal period these samples represent. Stockner et a l . (1977) noted that, largely as a result of lowered l i g h t l e v e l s in the spring of 1974, the spring bloom in Howe Sound was much later in 1974 than in 1973. A comparison of the radiation l e v e l s for the f i r s t f i v e months of 1973, 1974, and 1977 (Fig. 45) shows that radiation was markedly lower. S. costatum has a low l i g h t demand (Chan 1978). It was this c h a r a c t e r i s t i c of low l i g h t demand along with others (e.g., high maximum growth rate and low s a l i n i t y tolerance) which was used to explain the dominance of S. costatum in the winter-spring blooms of Hardangerfjord, Norway (Braarud 1974) and Narragansett Bay, Rhode Island (Smayda 1973). The low s a l i n i t y tolerance of S. costatum i s also applicable. Mean monthly discharge of the Squamish River at the head of Howe Sound was greater in 1974 (Fig. 46). The r e l a t i v e l y higher radiation and lower Squamish River 1 62 PRIMARY PRODUCTIVITY PLANKTONIC DIATOM ABUNDANCE < M H Figure 44. Gradients of primary productivity and planktonic diatom abundance, and t o t a l absolute abundance along the length of Howe Sound (1974). Direction of arrows indicates increasing values. A = absolute abundance, v/gdw = valves/gram dry weight, D = depth, M = mouth, and H = head. 1 63 Figure 45. Mean monthly radiation values for the Howe Sound area. Dots represent average normal values. (Data from Monthly Radiation Summary, Environment Canada.) 1 64 Li i L_ i i 1 1 1 ' M A M J J A S O N Figure 46. Mean monthly discharge values for the Squamish River. Dots represent average normal values. (Data from H i s t o r i c a l Streamflow Summary, Water Survey Canada.) 1 65 discharge in 1973 and 1977 (Figs. 45-46) correspond to the dominance of Thalassiosira spp. in Stockner et a l . ' s (1977) 1973 seasonal succession data and the Howe Sound (1977) sediment assemblages. Stockner et a l . ' s (1977) phytoplankton data show Thalassiosira spp. dominant in 1973 and Skeletonema costatum in 1974. The sediment samples were taken in November 1974 and S. costatum was dominant at a l l 24 s i t e s . This strongly suggests that the 1974 sediment samples represent only that year. Howe Sound does not have varved sediments so that consecutive years dominated by the same assemblages would be indistinguishable. Only an unusual year, such as 1974, could be used as a temporal marker. The cor r e l a t i o n between S. costatum and low radiation and/or s a l i n i t y levels in 1974 suggest that a core study in Howe Sound could be worthwhile. SUMMARY OF PART II 1) Zones. The gradient of primary productivity and temperature between the zones i s p o s i t i v e l y related to t o t a l absolute abundance of the sediment assemblages; the gradient of s a l i n i t y is negatively related. The range of absolute abundance in B r i t i s h Columbia sediments i s similar to that reported for the B r i t i s h Columbia coast in another study. The l a t i t u d i n a l gradients of the dominant species in B r i t i s h Columbia sediments are similar to that of the phytoplankton in Norwegian coastal waters. The l a t i t u d i n a l gradients are evident in the PCoA ordination: on axis 1, Zone A 1 66 s i t e s , which are dominated by Skeletonema costatum, are in the l e f t half of the graph; and s i t e s from Zones B and C, dominated by Thalassiosi ra a e s t i v a l i s f. pac i f ica and Chaetoceros spp. resting spores, are to the right of Zone A s i t e s . The d i s t r i b u t i o n of s i t e s on axis 1 can be related to the gradients of s a l i n i t y , temperature, and primary productivity. Axis 2 appears to be related to the r e l a t i v e d i s t r i b u t i o n of T. a e s t i v a l i s f. p a c i f i c a and Chaetoceros spp. resting spores. 2) Inlet types. Inlet types are not grouped together by the PCoA. There i s no s i g n i f i c a n t difference in absolute abundance between the i n l e t types. Div e r s i t y is highest in medium and low runoff i n l e t s , which i s probably related to runoff patterns and the a b i l i t y of the freshet in high runoff i n l e t s to remove phytoplankton from the surface waters. Some speices and species-groups have associations with p a r t i c u l a r i n l e t types. Within the i n l e t s , as with the zones, there are gradients of primary productivity, s a l i n i t y , and temperature, a l l of which increase toward the mouth with some exception in low runoff i n l e t s . The pattern of absolute abundance in the sediments usually coincides with these gradients. However, two physical processes may be a f f e c t i n g the sediment assemblage patterns. F i r s t , sedimentation rates vary throughout the study area, the highest rates being near the head of high runoff i n l e t s and in areas influenced by the Fraser River. The sediment load a f f e c t s the sediment assemblages to varying degrees by d i l u t i n g the rain of plankton with sediment p a r t i c l e s . This d i l u t i o n effect 167 probably results in decreased absolute abundance values p a r t i c u l a r l y in areas of heavy sediment loading. Between s i t e comparisons of absolute abundance values are not v a l i d as a result of the probable d i l u t i o n e f f e c t . However, i t appears v a l i d to compare the gradient of absolute abundance as i t coincides with known primary productivity gradients regardless of i n l e t type. Second, estuarine c i r c u l a t i o n may be aff e c t i n g the sediment assemblages by transporting species from the head mouthward along the i n l e t . This possible effect i s coupled with sediment focusing in deep basins, which tend to be near the mouth of i n l e t s . However, d i s t r i b u t i o n a l patterns in the sediments of the dominant phytoplankton species in the Howe Sound (1974) set of samples, and that of freshwater and marine l i t t o r a l species throughout the study area suggest that there is a good s p a t i a l r e l a t i o n s h i p between the biocoenoses and the thanatocoenoses. Physical processes do not seem to be seriously a l t e r i n g the sediment assemblages in B r i t i s h Columbia . 3) Howe Sound . The set of samples from Howe Sound (1974) is unique because i t has concurrent phytoplankton data. Thalassiosira spp. were dominant in the phytoplankton during 1973, Skeletonema costatum during 1974. The samples were taken in November 1974 and S. costatum was dominant at a l l 24 s i t e s , which suggests that the sediment assemblages represent only one year. There were lower radiation and higher river discharge levels in 1974, which may allow a future co r r e l a t i o n in core 1 68 studies in Howe Sound with abundance of S. costatum in sediment assemblages. 169 VI. CONCLUSIONS Four questions were asked in the Introduction. F i r s t , i s d i f f e r e n t i a l d i s s o l u t i o n as important in estuaries as i t i s in the oceans? In the sense that neither the ocean nor the B r i t i s h Columbia thanatocoenoses is a mirror image of the biocoenoses, the broad answer i s yes. Of greater interest is the relationship between possible c o n t r o l l i n g factors and d i f f e r e n t i a l d i s s o l u t i o n . The assertion of many studies, that species or species-groups ( i . e . , n e r i t i c species) do not reach deep-ocean sediments because they are weakly s i l i c i f i e d and p r e f e r e n t i a l l y dissolve in deep water, i s not s u f f i c i e n t to explain the discrepancies between the biocoenoses and the thanatocoenoses in B r i t i s h Columbia . Skeletonema costatum, usually considered to be a di s s o l u t i o n - s e n s i t i v e species, i s abundant in B r i t i s h Columbia sediments; Thalassiosira  nordenskioeldi i , which produces heavily s i l i c i f i e d resting spores, i s rare; and Thalassiosira a e s t i v a l i s f. p a c i f i c a i s abundant in the sediments while Thalassiosi ra a e s t i v a l i s f. a e s t i v a l i s i s rare, yet both are dominant in the phytoplankton and appear to be equally s i l i c i f i e d . Numerous factors (e.g., temperature, depth, grazing pressure, the organic casing, l i f e stages, and/or the physical condition of the c e l l ) appear to be involved in the pr e f e r e n t i a l deposition and preservation of some species over others. At present the o r i g i n a l question r e a l l y has no answer; our understanding of the processes involved in d i f f e r e n t i a l d issolution i s simply too limited, p a r t i c u l a r l y with regard to the presence of an organic 170 wall. Second, can the gross d i s t r i b u t i o n a l patterns of freshwater and marine l i t t o r a l species be considered indicative of their source? The answer here i s a q u a l i f i e d yes. Both freshwater and marine l i t t o r a l species appear to be indicative of r i v e r sources entering the estuarine system. Within the i n l e t s , these species have their highest abundance r e l a t i v e l y near their source. However, at the mouth of i n l e t s , p a r t i c u l a r l y those in Zones A and B, some freshwater species found in the sediments probably originate from the Fraser River. Third, i s i t v a l i d to compare absolute abundance of diatoms in the surface sediments with primary productivity of surface waters, given the effect of d i f f e r e n t i a l sedimentation rates? The d i l u t i o n effect related to sediment loading varies throughout the B r i t i s h Columbia study area so that exact values of absolute abundance should not be compared between s i t e s . However, a comparison of the gradients of these values i s v a l i d : in the zones, the absolute abundance in the sediment assemblages increases from the northern Zone C to the southern Zone A as does primary productivity; within the i n l e t s , both absolute abundance and primary productivity usually increase toward the mouth. Fourth, does estuarine c i r c u l a t i o n seriously change the relationship between the biocoenoses and the thanatocoenoses at the scale of t h i s study? The answer appears to be no. The d i s t r i b u t i o n of freshwater and marine l i t t o r a l species in the sediment assemblages and the dominant marine phytoplankton 171 species in the Howe Sound (1974) s i t e s , a l l indicate a good sp a t i a l relationship between the biocoenoses and the thanatocoenoses. Are the thanatocoenoses, therefore, a good r e f l e c t i o n of the biocoenoses in B r i t i s h Columbia ? In general, the answer i s yes, since the dominant members of the sediment assemblages are also dominants in the phytoplankton. Recurrent groups found throughout the study area are t y p i c a l of many estuarine systems; and the s p a t i a l relationship appears to be maintained. F i n a l l y , can these d i s t r i b u t i o n a l patterns be related to oceanographic factors? The sediment assemblages were grouped for analysis on the basis of i n l e t type, zonal, and within-inlet patterns. In general, the species and species-groups examined often exhibited d i s t i n c t d i s t r i b u t i o n a l patterns related to these categories. The most consistent pattern was the cor r e l a t i o n between abundance and primary productivity, usually associated with minor upwelling zones, which has also been demonstrated in other areas. On the other hand, the d i s t r i b u t i o n a l pattern of Chaetoceros spp. resting spores in the B r i t i s h Columbia sediments may be unique to estuarine systems. High runoff i n l e t s have a markedly lower abundance of Chaetoceros spp. resting spores probably as a result of the freshet which removes the phytoplankton from within the i n l e t during the period of peak Chaetoceros spp. resting spores formation. S a l i n i t y and temperature vary within the i n l e t s and between i n l e t types and zones, and some diatom d i s t r i b u t i o n a l patterns can be correlated with these gradients. In p a r t i c u l a r , 1 72 the PCoA analysis shows a zonal c o r r e l a t i o n between the dominant species in the sediment assemblages and three aspects ( i . e . , primary productivity, s a l i n i t y , and temperature). General d i s t r i b u t i o n studies, such as th i s one, can be used as a data base for companion studies investigating paleoecology, po l l u t i o n e f f e c t s , c i r c u l a t i o n patterns, and phytoplankton d i s t r i b u t i o n . In p a r t i c u l a r , the results of th i s study suggest six areas of further research: 1) an autecology study of Thalassiosira a e s t i v a l i s to determine why T. a e s t i v a l i s f. p a c i f i c a i s dominant in sediment assemblages while T. aest i v a l i s f. a e s t i v a l i s i s rare; 2) a study of Thalassiosira  nordenskioeldii to determine i f temperature i s c o n t r o l l i n g resting spores formation in B r i t i s h Columbia waters; 3) an investigation of the l a t i t u d i n a l planktonic gradients exhibited by the dominant members of the sediment assemblages; 4) a d i s t r i b u t i o n a l study of freshwater species in surface waters and sediments to determine the r e l a t i v e influence of the Fraser River on sediment assemblages in Zones A and B; and 5) a comparison of yearly radiation data and Squamish River discharge data with the abundance of Skeletonema costatum in sediment cores from Howe Sound to determine i f S. costatum can be used as a temporal marker in unvarved sediments; and 6) an investigation of the role of the organic wall component in preservation. 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Oceanology 19:720-724. 187 APPENDIX A - SPECIES LIST The species l i s t i s ordered alphabetically by the three l e t t e r code used for each species in the s i t e counts (see Appendix D). Habitats are designated as FW (freshwater), ML (marine l i t t o r a l ) , and MP (marine plankton). S designates s i l i c o f l a g e l l a t e s . Numbers refer to references found at the end of th i s appendix. acp acs ahb ahc ahd ahe ahf ahh ahk ahl aho ahr ahs ahw ahx ahy ame amo amp amr ams amt ast ats bat bda bdl cac cbm cbn cbs ceo eds cha chd che chg chl chm splendens (Shadbolt) Ralfs ML 11 senarius Ehrbg. ML 1,2 Ag. ML 1,5 conspicua A. Mayer FW 7,13 lanceolata v. dubia Grun. FW 7 . (Breb.) Grun. FW 4,7 f r a q i l a r i o i d e s Peterson FW 13 hauckiana Grun. ML 1,4 kryophila Peterson FW 13 lemmermanni Hust. FW 6,13 Act inoptychus  Actinoptychus  Achnanthes brevipes  Achnanthes Achnanthes  Achnanthes lanceolata  Achnanthes  Achnanthes  Achnanthes Achnanthes  Achnanthes longipes Ag. Achnanthes 1inearis f. Achnanthes spp. Achnanthes a f f i n i s Grun. FW 4 Achnanthes microcephala (Kuetz.) Grun, subsalsoides Hust. ML 13 eunotia CI. ML 5 ovali s (Kuetz.) Kuetz. FW 4 Greg. ML 1,5 Greg. ML 5 ML 1 ,8 curta H.L.Sm. FW 7 FW 4 Achnanthes  Amphora  Amphora  Amphora  Amphora  Amphora  Amphora Ast e r i o n e l l a Act inocyclus Bacteriasrtum proteus  granulata spp. truncata (Greg, spp. s u b t i l i s ) CI. ML 5 ) Ralfs MP 12,13 (= Odontella aur i t a (Greg, spp. MP 3 Biddulphia aur i t a (Lyngb.) Breb, (Lyngb.) C.Ag.) ML 1,3,15 Biddulphia l o n g i c r u r i s G r e v i l l e MP 2 5 15 .Campylosira cymbelliformis (Schm.) Grun. ex V.H. ML 1 5 Cymbella mmuta Hilse ex Rabh. FW 7 — ' bymbelll f p l T ~ V ' s i l e s i a c a < B l - i s c h ex Rabh.) Reim. FW 7 Cocconeis costata Greg. ML Cocconeis disculoides Hust, Chaetoceros a f f i n i s Lauder Chaetoceros d e b i l i s CI. MP (Ehrbg.) g r a c i l i s Schuett lauderi Ralfs MP Chaetoceros diadema Chaetoceros Chaetoceros 1 2 ML 1 MP 3, 2,11 CI. MP 13 2 1 1 1 2 MP 3 3,11,12 Chaetoceros spp. resting spores 188 chr Chaetoceros radicans Schuett MP 2,11,12 chs Chaetoceros seiracanthus Gran MP 8,11 chv and chc Chaetoceros vanheurckii Gran MP 11,12 chy Chaetoceros didymus Ehrbg. MP 2,11,12 chz Chaetoceros lorenzianus Grun. MP 3,11,12 c l a Caloneis a l p e s t r i s (Grun.) C l . FW 7 cmg Coscinodiscus marqinatus Ehrbg. MP 3,11,12 coa Cocconeis clandestina A.Sm. ML 5 coc Coscinodiscus curvatulus Grun. MP 3,5,11 cod Coscinodiscus o c u l u s - i r i d i s Ehrbg. MP 12 cof Cocconeis f l u v i a t i l i s Wallace FW 4,7 col Cocconeis placentula Ehrbg. FW 4 com Coscinodiscus curvatulus v. minor (Ehrbg.) Grun. MP 13 con Coscinodiscus nitidus Greg. ML 1,5,11 coo Coscinodiscus obscurus A.Sm. MP 3,12 cop Coscinodiscus perforatus Ehrbg. MP 11,12 cor Coscinodiscus radiatus Ehrbg. MP 8,12 cos Coscinodiscus s t e l l a r i s Roper MP 11,12 cot Coscinodiscus c e n t r a l i s v. p a c i f i c a Gran and Angst MP 11,12 cov Coscinodiscus perforatus v. p a v i l l a r d i (Forte) Hust. MP 11 cow Coscinodiscus w a i l e s i i Gran and Angst MP 2,11,12 cps Cocconeis pseudomarqinata Greg. ML 13 csb Cocconeis s u b l i t t o r a l i s Hendey ML 8 esc Cocconeis scutellum Ehrbg. ML 1,5 cse Cocconeis placentula v. euqlypta (Ehrbg.) Cl.FW 7 csm C y c l o t e l l a spp. esp Cocconeis pelluc ida Hantz. FW 4 ess Cocconeis spp. cya C y c l o t e l l a comta (Ehrbg.) Kuetz. FW 4 eye Cymbella microcephalia Grun. FW 4,7 cym Cymbella mexicana (Ehrbg.) C l . FW 4,5 cyo C y c l o t e l l a ocellata Pant. FW 4 cys C y c l o t e l l a s t r i a t a (Kuetz.) Grun. ML 3,4,12 c y t C y c l o t e l l a s t e l l i q e r a Cl.and Grun. FW 4 dia Diatoma anceps (Ehrbg.) Kirchn. FW 4 die Diatoma elonqatum (Lyngb.) C.Ag. ML 1,4,6 d i f Dictyocha f i b u l a Ehrbg. MP S 16 dih Diatoma hiemale (Lyngb.) Heib. FW 4,6 dim Diatoma hiemale v. mesodon (Ehrbg.) Grun. FW dis Dictyocha speculum (Ehrbg.) Haeckel MP S 16 di t Ditylum brightwelli i (West.) Grun. MP 2,11,12 dmm Dimereqramma minor (Greg.) Ralfs ML 5 dpb Diploneis crabro Ehrbg. ML 5,8 dpc Diploneis smithi i v. c o n s t r i c t a Heiden FW 13 dpp Diploneis spp. dps Diploneis smithi i Breb. FW 4,5 dpt Diploneis c f . peterseni Hust. FW 13 ebt Ebria t r i p a r t i t a (Schum.) Lemmermann MP S 16 ept Epithemia turqida (Ehrbg.) Kuetz. ML 5 eua Eunotia arcus Ehrbg. FW 4 eud Eunotia sudetica 0.Muell. FW 4 e u e Eunotia exigua fBreb.ex Kuetz.) Rabh. FW 4 eul Eunotia polydentula Brun. FW 13 eup Eunotia p e c t i n a l i s v. minor (Kuetz.) Rabh. FW 4,7 189 eus Eunotia spp. fra F r a g i l a r i a capucina Desm. FW 4 frb F r a g i l a r i a b r e v i s t r i a t a Grun. FW 4 frc F r a g i l a r i a construens (Ehrbg.) Grun. FW 4 f r i F r a g i l a r i a bicapitata A.Mayer FW 7 f r l F r a g i l a r i a leptostauron (Ehrbg.) Hust. FW 4 frn F r a g i l a r i a construens v. venter (Ehrbg.) Grun. FW 4 frp F r a g i l a r i a pinnata Ehrbg. FW 4,5 f r s F r a g i l a r i a construens v. subsalina Hust. FW 13 frv F r a g i l a r i a vaucheriae (Kuetz.) Peters (= Synedra vaucheriae Kuetz.) FW 4,7 fry F r a g i l a r i a c y l i n d r i c u s Grun. (= Nitzschia c y l i n d r i c u s (Grun.) Hasle) MP 3 gld Glyphodesmis distans (Greg.) Grun.ex V.H. ML 8 goa 'Gomphonema acuminatum Ehrbg. FW 4 goc Gomphonema constrictum Ehrbg. FW 4 goo Gomphonema olivaceum (Lyngb.) Kutz. FW 4 gos Gomphonema spp. gra Grammatophora c f . a r c t i c a CI. ML 13 grc Grammatophora oceanica v. macilenta (W.Sm.) Grun. ML 1 grg Grammatophora anqulosa Ehrbg. ML 1,5 grm Grammatophora marina TLynqb.) Kutz. ML 1,5 gro Grammatophora oceanica Ehrbg. ML 1 grx Grammatophora maxima Grun. ML 5 9Y f Gyrosiqma f a s c i o l a (Ehrbg.) CI. ML 1,4 9Y9 Gyrosiqma s t r i q i l i s (W.Sm.) CI. ML 12 gys Gyrosiqma spenceri i (W.Sm.) C l . ML 1,4 gyt Gyrosiqma t e n u i r o s t r i s (Grun.) C l . ML 12 hna Hannaea arcus (Ehrbg.) Patr. FW 7 hnx Hannaea arcus v. amphioxys (Rabh.) Patr. FW 7 hyr Hyalodiscus radiatus (O'Meara) Grun. ML 1 hys Hyalodiscus s u b t i l i s B a i l . ML 1,12,16 led Leptocylindrus danicus C l . MP 2,12 l i e Licmophora ehrenberqi i (Kutz.) Grun. ML 1,5 l i g Licmophora g r a c i l i s (Ehrbg.) Grun. ML 1 mec Meridion c i r c u l a r e (Grev.) C.Ag. FW 4 mla Melosira ambiqua~"Grun. ) 0.Muell. FW 4 mid Melosira distans (Ehrbg.) Kuetz. FW 4 mlg Melosira granulata (Ehrbg.) Ralfs FW 4 mli Melosira islandica 0.Muell. FW 4,12 mlm Melosira moniliformis (Mull.) Ag. ML 1 mis Melosira sol (Ehrbg.) Kuetz. MP 12 mlt Melosira i t a l i c a (Ehrbg.) Kutz. FW 4,12 mse Mastogloia exigua Lewis ML 1,5 neb Neidium bisulcatum v. subundulatum (Grun.) Reim. FW 7 nva Navicula/Achnanthes sp. nvd Navicula d i g i t o - r a d i a t a (Greg.) Ralfs ML 5 nvf Navicula forcipata Grev. ML 1,9 nvg Navicula palpebralis Breb. ex Wm.Sm. ML 8 nvj Navicula r o s t e l l a t a v. major A.C1. ML 5 nvk Navicula subtilissima C l . FW 4 nvl Navicula lanceolata(Ag.) Kuetz. FW 7 nvp Navicula pupula f. capitata Skv.and Meyer FW 4 nvq Navicula c f . annexa Hust. FW 13 190 nvr Navicula rhynchocephala Kuetz. FW 4 nvs Navicula spp. nvt Navicula directa v. oceanica Karsten ML 5 nvu Navicula pupula Kuetz. FW 4 nvv Navicula c f . d i v e r s i s t r i a t a Hust. ML 9 nvy Navicula pseudony Hust. ML 5,9 nvz Navicula c f . microcephala Grun. FW 13 nza Nitzschia acuminata (W.Sm.) C l . and Grun. ML 1,5 nzb Nitzschia bilobata W.Sm. ML 8 nzc Nitzschia microcephala Grun. FW 4,10 nzd Nitzschia delicatissima C l . ML 11 nzf Nitzschia p a c i f i c a Cupp ML 11 nzg Nitzschia anqularis W.Sm. ML 1,5' nzi Nitzschia sigma W.Sm. ML 5,8 nzl Nitzschia lonqissima (Breb.) Ralfs ex Pr i t c h . ML 1 nzm Nitzschia marqinulata Grun. ML 5 nzn Nitzschia punctata (W.Sm.) Grun. ML 1,5 nzo Nitzschia s o c i a l i s Ralfs ML 1,5 nzp Nitzschia paradoxa (Gmelin) Grun. ML 11 nzr Nitzschia granulata Grun. ML 5 nzs Ni tzschia spp. n z t Nitzschia t r y b l i o n e l l a Hantz. ML 4,9 nzu Nitzschia vermicularis (Kuetz.) Grun. FW 4,12 nzv Nitzschia b r e v i r o s t r i s Hust. ML 9 nzw Nitzschia pungens Grun. in C l . and Moller ML 12 nzx Ni tzschia c f . majuscula Grun. ML 14 opm Opephora martyi Herib. FW 4 opp Opephora p a c i f i c a (Grun.) Petit ML 1 opr Opephora marina TGreq.) Petit ML 1,5 ops Opephora schwartzii (Grun.) Petit in P e l l a t i n ML 5 par Paralia sulcata (Ehrbg.) Cl.(= Melosira sulcata (Ehrbg.) Kuetz.) ML 1,12 pib Pinnularia biceps Greg. FW 4 pic Pinnularia cruciformis (Donkin) C l . ML 5,8 pirn Pinnularia microstauron (Ehrbg.) C l . FW 4 p l l Pleurosiqma lonqum C l . ML 12 pis Plaqioqramma staurophorum (Greg.) Heib. ML 1,5 raa Rhaphoneis angustata Pant. ML 5 ras Rhaphoneis spp. rcc Rhoicosphenia curvata (Kuetz.) Grun. ML 1,5 res Rhoicosphenia spp. rhz Rhizosolenia spp. MP rom Rhopalodia musculus (Kuetz.) Muller sec Sceptroneis caducea Ehrbg. ML 13 skc Skeletonema costatum (Grev.) C l . MP 2,11 sta Stephanodiscus astraea (Ehrbg.) Grun. FW-4 stm Stephanodiscus astraea v. minutula (Kuetz.) Grun. FW 4 sts Stephanopyxis spp. MP 3,4,5,12 suf S u r i r e l l a fastuosa v. recedens (A.Sm.) C l . MP 11 sui S u r i r e l l a intermedia A.C1. ML 12 suo S u r i r e l l a ovata Kuetz. ML 5 sya Synedra acus Kuetz. FW 4 syf Synedra f a s c i c u l a t a v. truncata (Grev.) Patr. FW 4,5,7 syr Synedra rumpens Kuetz. FW 4 191 sys Synedra spp. syt Synedra tabulata (Ag.) Kuetz. ML 1,5 syu Synedra ulna (Nitz.) Ehrbg. FW 4 taf Thalassiothrix frauenfeldi i Grun. MP 3,11 tas Thalassionema spp. tbf T a b e l l a r i a flocculosa (Roth) Kuetz. FW 4 tbn T a b e l l a r i a fenestrata (Lyngb.) Kuetz. FW 4 the Thalassiosira eccentrica (Ehrbg.) C l . MP 2 thd Thalassiosi ra dec ipiens (Grun.) Joergensen MP 3,12 the Thalassiosira anguste-lineata (A.Sm.) G.Fryxell and Hasle MP 12 thg Thalassiosira angsti i (Gran) Makarova MP 2,12 thn Thalassiosira nordenskioeldii C l . MP 3,12 thp Thalassiosi ra a e s t i v a l i s f. pac i f ica MP tnz Thalassionema nitzschioides Hust. MP 1,2 tra Trachyneis aspera (Ehrbg.) C l . ML 1,8 tr s Tropidoneis spp. ML 11 t x l Thalassiothrix longissima C l . and Grun. MP 3,11 References 1. Mclntire, CD., and W.W. Moore. 1977. Marine l i t t o r a l diatoms: ecological considerations. In: The Biology of Diatoms (ed. D. Werner) pp.333-371. Blackwell S c i e n t i f i c Pubis., Oxford. 2. Venrick, E.L. 1971. Recurrent groups of diatom species in the North P a c i f i c . Ecology 52:614-625. 3. G u i l l a r d , R.R.L., and P. Kilham. 1977. The ecology of marine planktonic diatoms. In: The Biology of Diatoms (ed. D. Werner) pp. 372-469. Blackwell S c i e n t i f i c Pubis., Oxford. 4. Stein, J.R. and CA. Borden. 1978. Checklist of freshwater algae of B r i t i s h Columbia. Syesis 12:3-37. 5. Riznyk, R.Z,. 1973. I n t e r s t i t i a l diatoms from two t i d a l f l a t s in Yaquina Estuary, Oregon, U.S.A. Botanica mar. 16:113-138. 6. Patrick, R. 1977. Ecology of freshwater diatoms and diatom communities. In: The Biology of Diatoms (ed. D. Werner) pp. 284-332. Blackwell S c i e n t i f i c Pubis., Oxford 7. Patrick, R. and C.W. Reimer. 1966. The Diatoms of the United States. Vol.1, 688 pp. Acad. Natn. S c i . Philadelphia, Monograph 13. 1 92 8. 1975. Vol. 2, Part 1. 213 pp. Acad. Natn. S c i . Philadelphia, Monograph 13. 9. Hendey, N.I. 1964. An Introductory Account of the Smaller Algae of B r i t i s h Coastal Waters. Bacillariophyceae (Diatoms). 317 pp. HMSO, London. 10. Hustedt, F. 1955. Marine l i t t o r a l diatoms of Beaufort, North Carolina. B u l l . Duke Univ. mar. Stn. 6:1-67. 11. Hustedt, F. 1930. Ba c i l i a r i o p h y t a . In: Die Susswasser-Flora Mitteleuropas (ed. A. Pascher). 468 pp. Gustav Fisher, Jena. 12. Cupp, E.E. 1943. Marine plankton diatoms of the west coast of North America. B u l l . Scripps Instn. Oceanogr. tech. Ser. 5:1-238. 13. Shim, J.H. 1976. D i s t r i b u t i o n and taxonomy of planktonic marine diatoms in the S t r a i t of Georgia. Ph.D. thesis, Univ. B r i t i s h Columbia, Vancouver. 248 pp. 14. Hustedt, F. 1927-1966. Die Kieselalgen Deutschlands, Oesterreichs und der Schweiz. In: Kryptogamen-Flora 7, Part I, II, III (ed. L. Rabenhorst) 7(1): 920pp, 1927-1930; 7(2): 845 pp., 1931-1959; 7(3): 816 pp., 1961-1966. 15. Peragallo, H., and M. Peragallo. 1897-1908. Diatomees marines de France et des d i s t r i c t s maritimes voisines. 540 pp. M.J. Tempere, Grez-sur-Loing. 16. Gran, H.H., and E.C. Angst. 1931. Plankton diatoms of Puget Sound. Pubis. Puget Sound mar. b i o l . Stn. 7:417-519. 17. Gemeinhardt, K., and J . S c h i l l e r . 1930. S i l i c o f l a g e l l a t a e und Coccolithineae. In: Kryptogamen-Flora 10 (ed. L. Rabenhorst). 273 pp. 1 93 APPENDIX B - TAXONOMY Centric diatoms in the genera Thalassiosira , C y c l o t e l l a , Coscinodiscus , and Stephanodiscus have been shown to exhibit some p l a s t i c i t y of valve morphology in response to changing environmental conditions. Polymorphism has been attributed to changes in s i l i c o n concentrations (Becher et a l . 1966; Hasle et a l . 1971; F r y x e l l and Hasle 1972; Paasche 1973b; Booth and Harrison 1979), nitrogen and phosphorus concentrations and temperature (Syvertsen 1977), and s a l i n i t y (Schultz 1971). Valve morphology can also change as a result of the inverse relationship between the number of areolae in 10 Mm and valve diameter in some species (Hasle 1978; Theriot and Stoermer 1981). These polymorphic a b i l i t i e s have caused considerable confusion in taxonomy of these genera. During the summer of 1928, Dr. H. H. Gran taught a course on diatoms at the b i o l o g i c a l station at Friday Harbor, Washington. At that time, he found a new species, T. p a c i f i c a , which was abundant during March-April of 1927 and 1928. He returned to Friday Harbor in the summer of 1930 and discovered another new species in July, T. a e s t i v a l i s . Both new species were formally described by Gran and Angst (1931). Between May 1, 1931 and A p r i l 30, 1932, L. D. Phifer, a student from Gran's 1928 diatom course, made an intensive study (six or seven samples a month) of the seasonal d i s t r i b u t i o n of planktonic diatoms at Friday Harbor (Phifer 1934a). He noted that T. a e s t i v a l i s occurred "from late June continuously through August and then intermittently u n t i l late November," and that T. p a c i f i c a occurred "from May to mid-June, 1931, and throughout A p r i l , 1932," i . e . , there appeared to be a d i s t i n c t temporal separation, T. p a c i f i c a occurring in the spring, T. a e s t i v a l i s in the summer and f a l l . Thalassiosira nordenskioeldi i occurred in the interim between T. pac i f ica and T. a e s t i v a l i s ; Phifer stated that T. nordenskioeldii occurred "constantly from mid-May to August, reaching a maximum...in early July." Gran and Angst (1931) noted that T. aest i v a l i s "has some likenesses to T. nordenskioeldii," and both Shim (1976) and Cupp (1943) expressed d i f f i c u l t y in distinguishing between the two. The o r i g i n a l descriptions (Gran and Angst 1931) contain the seeds of the subsequent confusion (see Table 12 for a comparison Gran and Angst state that "T. pac i f ica i s distinguished from T. a e s t i v a l i s by the rounded corners of the c e l l s seen in g i r d l e view, by the coarse structure of the valve, and by equal thickening of a l l sutures in the g i r d l e zone." The f i r s t c h a r a c t e r i s t i c i s the shape of the mantle. T. a e s t i v a l i s (Gran and Angst 1931) i s described as having a "beveled" mantle, while in the f i r s t sentence of the description of T. p a c i f i c a the mantle i s described as "rounded." In the second sentence, the mantle of T. p a c i f i c a i s described as 1 94 Table 12 . Morphological data for Thalassiosira p a c i f i c a and T. a e s t i v a l i s . diameter valve areolae marginal (/um) i n 10 /xm i n 10 Mm face mantle Gran and Angst (1931) T. a e s t i v a l i s 26-55 20 3.5 T. p a c i f i c a 15-46 12-15 4 Cupp (1943) T. a e s t i v a l i s 20-45 20-22 Hasle (1978) T. a e s t i v a l i s 14-46 16 >20 4(3-5) !• p a c i f i c a 7-46 10-14 (18) >20 5(4-6) Shim (1976) T. a e s t i v a l i s 21 19 T. p a c i f i c a 22-46 15-18 4-6 195 "beveled." The i l l u s t r a t i o n s for both species show beveled mantles. Hasle (1978) states that "the angle between valve face and mantle of T. p a c i f i c a may be less sharp than in T. aestivalis"(emphasis mine). Shim (1976) describes the mantle of T. p a c i f i c a as rounded in one sentence, and beveled in another. Obviously, the shape of the mantle cannot be used as a distinguishing c h a r a c t e r i s t i c . The second c h a r a c t e r i s t i c i s the coarse structure of the valve in T. pac i f i c a . In the o r i g i n a l description (Table 12), the difference between the number of valve areolae in 10 /_m on the valve face of the two species i s discrete. However, Hasle (1978), using the holotype material, found that T. pac i f ica had "10-18 (mostly 10-14) areolae in 10 um on valve face" and T. a e s t i v a l i s had "about 18 areolae in 10 Mm." She further notes that T. p a c i f i c a "specimens larger than 25 Mm have 10-12 areolae in 10 Mm." This may be a function of the inverse relationship between areolae number and valve diameter ( i . e . , smaller valves w i l l have more areolae in 10 Mm). Shim (1976) states that T. p a c i f i c a had "15-18 areolae in 10 nm" and T. a e s t i v a l i s had "about 19 areolae in 10 Mm." In the present material T. p a c i f i c a had 8-18 (mostly 12-13) areolae in 10 Mm. The t h i r d d i s t i n c t i o n between the two species in the o r i g i n a l descriptions i s the equal thickening of a l l sutures in the g i r d l e zone. Hasle (1978) simply states that her investigation does not include information on these structures. However, she does note that the "valve wall" of T. p a c i f i c a can be "more strongly s i l i c i f i e d than in T. a e s t i v a l i s . " It appears, then, that none of the o r i g i n a l distinguishing c h a r a c t e r i s t i c s seems to be s u f f i c i e n t l y discrete to distinguish the species with confidence. Hasle (1978) states that "the main d i s t i n c t i o n between T. pac i f ica and T. a e s t i v a l i s i s , however, more manifest in the areolae array. The areolae of T. aest i v a l i s are in straight r a d i a l rows, mostly in a large number of f a i r l y narrow sectors.... In contrast, the areolae of T. p a c i f i c a are in straight or s l i g h t l y curved tangential rows or in r a d i a l rows p a r a l l e l to the median row of much wider sectors...." T. p a c i f i c a can also have a linear areolae pattern lacking sectors. Gran and Angst (1931) noted that both species have straight rows p a r a l l e l to the median row, and e a r l i e r in her paper Hasle states T. a e s t i v a l i s also has the same c h a r a c t e r i s t i c . Oddly enough, the c h a r a c t e r i s t i c that Hasle (1978) considers the "main d i s t i n c t i o n " between T. p a c i f i c a and T. a e s t i v a l i s ( i . e . , the areolae array) i s one of the main c h a r a c t e r i s t i c s that was used to determine that Coseinodiscus  i n f l a t u s and C_j_ tumidus are conspecific when they were combined into T_j_ tumida (Hasle et a l . 1971). There i s the same intergrading of morphological c h a r a c t e r i s t i c s in C_^  i n f l a t u s and C. tumidus as in T. p a c i f i c a and T. a e s t i v a l i s . The authors state that coarsely s i l i c i f i e d C_j_ tumidus has "valve areolae in 1 96 straight tangential or eccentric curved rows," while the weakly s i l i c i f i e d C_^  i n f l a t u s has "valve areolae arranged in sectors." Further, they note that the coarsely s i l i c i f i e d species can have a lineatus structure and can have fewer areolae in 10 nm. It i s curious that, on the basis of the same c h a r a c t e r i s t i c s , Hasle chooses to maintain T. pacif ica and T. a e s t i v a l i s as separate species while making C_^  inf latus and C_^  tumidus conspecific. There are further s i m i l a r i t i e s between T. p a c i f i c a and T. a e s t i v a l i s (Hasle 1978): there i s a similar range of marginal processes in 10 nm (Table 12), the location of the labiate process is the same, the height of the valve mantle i s 2-3 areolae, and the number of marginal ribs in 10 Mm i s half the number of valve areolae for both. T. p a c i f i c a has 5-6 processes per sector, and T. a e s t i v a l i s 2-3; however, th i s is a function of sector width, not process placement. There are 3-5 s t r i a e between processes regardless of sector width. The extreme intergrading of c h a r a c t e r i s t i c s suggests that these are not two d i s t i n c t species but rather two forms of one polymorphic species. Although both forms may coexist, probably in inverse proportions, there also appears to be a temporal separation which investigators should note. Unknowingly, Hasle (1978) demonstrated the temporal separation shown by Gran and Angst (1931) and Phifer (1934a). The 27 photo- and micrographs (Hasle 1978) used to i l l u s t r a t e T. p a c i f i c a represent samples taken in February-March in B r i t i s h Columbia , Washington, and Norway. The samples for the 17 photo- and micrographs for T. a e s t i v a l i s were taken in May-July in B r i t i s h Columbia and Washington. Two other temporal examples e x i s t : 1) T. a e s t i v a l i s was isolated from the phytoplankton in June in Saanich Inlet (Hollibaugh et a l . 1980), and 2) Thalassiosira p a c i f i c a was a member of the spring bloom for two consecutive years in the Yaquina Estuary, Oregon (Karentz and Mclntire 1977). If the two forms are to be maintained, there should be some discrete morphological c h a r a c t e r i s t i c . For h e u r i s t i c purposes, these forms w i l l be separated using areolae numbers. On the basis of p r i o r i t y , T. a e s t i v a l i s i s the species name ( i . e . , described e a r l i e r in the same publication). Valves with less than 16 areolae in 10 Mm w i l l be designated " f . p a c i f i c a " , while those with greater than 16 areolae in 10 Mm w i l l be " f . a e s t i v a l i s " . This i s an admittedly a r b i t r a r y separation; however, i t is similar to Gran and Angst's (1931) o r i g i n a l descriptions. 1 97 APPENDIX C - DISTRIBUTION OF SPECIES This appendix is a description of the d i s t r i b u t i o n of the marine planktonic, marine l i t t o r a l , and freshwater species. The marine planktonic species l i s t does not include the dominant species (Skeletonema costatum, Thalassiosira a e s t i v a l i s f. pac i f ica, Thalassionema nitzschioides, or Chaetoceros spp. 5" and the marine l i t t o r a l species l i s t does not include Paralia  sulcata. The key to the 3-letter species code is in Appendix A. The number following each species refers to the t o t a l number of si t e s at which the species occurs. Marine planktonic species Zones A, B, and C (numbers in parentheses refer to number of s i t e s in Zones A and B) cmg 11 (10) led 44 (31) coc 10 (3) mis 2 (1 ) coo 13 (8) taf 23 (12) cop 14 (12) the 65 (48) cos 11 (9) thd 18 (12) cot 10 (5) the 30 (22) di f 4 (3) thg 51 (37) dis 70 (50) thn 70 (52) d i t 89 (71 ) t x l 9 (3) fry 2 (1) Zones B and C only ats 3 bdl 4 ebt 4 J e r v i s Inlet, Malaspina S t r a i t , and Agamemnon Channel only cod 1 cor 3 cow 1 Howe Sound only cov 1 East Gorge Harbor only suf 1 Heriot Bay only com 1 Marine l i t t o r a l species Zones A, B, and C (numbers in parentheses refer to number of s i t e s in Zones A and B) acp 29 (24) nzb 4 (3) acs 26 (19) nzg 5 (3) ahb 4 (3) nzn 2 (1) 198 ahh 21 (14) ame 5 (2) amr 6 (4) bda 19 (12) ceo 73 (51) eds 38 (30) esc 28 (14) cys 66 (60) dmm 2 (1) dpb 2 (1) grc 4 (2) grm 8 (6) gro 4 (3) mlm 19 (16) nvd 5 (4) nvf 4 (3) nvt 4 (1) nza 10 (5) Zones B and C only cac 2 coa 3 con 3 gld 2 grg 2 hyr 7 hys 7 nvy 6 ops 2 Zones A and B only nzd 5 rcc 2 suo 3 Howe Sound only die 1 grx 1 gyf 1 gyg 2 gyt 1 l i e 3 l i g 1 nvv 1 Sechelt Inlet only aho 1 nvq 1 Toba Inlet only ahy 1 csb 1 nv j 1 pic 1 nzo 1-5 (11) nzp 9 (7) nzw 4 (2) opp 4 (2) opr 13 (11) pis 11 (8) rom 2 (1) syt 38 (26) tra 5 (2) nzf 1 nz i 1 nzl 3 nzm 1 nzv 1 raa 1 sui 2 199 Homfray Channel only amp 1 Hotham Sound only amt 1 Jer v i s Inlet only gra 1 mse 1 East Gorge Harbor only cps 1 gys 1 Malaspina S t r a i t only ept 1 Heriot Bay only t r s 1 Burrard Inlet only nva 1 nzt 1 S u t i l Channel only sec 1 Saanich Inlet only nzv 1 nzx 1 Hidden Basin only p l l 1 Freshwater species Zones A, B, and C (numbers in parentheses refer to number of s i t e s in Zones A and B) ahc 10 (7) mec 4 (3) ahk 2 (1) nvk 2 (1 ) col 14 (10) opm 8 (7) cse 2 (1) stm 12 (11) esp 5 (3) syf 3 (1 ) cyt 9 (8) syu 7 (6) dih 8 (7) tbf 10 (7) dps 17 (9) tbn 1 1 (10) eup 12 (5) frb 3 (2) frc 5 (4) frp 24 (22) f rv 8 (5) hna 6 (4) 200 Zones B and C only dim 3 dpc 1 Zones A and B only ahe 2 cyo 1 1 f r n 2 goo 1 7 mlg 3 mlt 4 Zone A only f r l 3 syr 2 Burrard I n l e t only ahw 1 mid 1 cya 1 nzc 1 eye 1 nzu 1 dpt 1 Howe Sound only ahd 1 ahf 1 ahr 1 ahx 1 cbm 2 cof 1 d i a 1 eua 1 eue 1 f r i 1 f r s 2 goa 1 hnx 1 mla 1 mli 4 neb 1 nvp 1 nvu 1 s t a 1 sya 1 J e r v i s I n l e t only a h l 1 amo 2 eud 1 f r a 1 pirn 1 P e n d r e l l Sound only c l a 1 Toba I n l e t only cbn 1 e u l 1 Saanich I n l e t only cym 1 Cordero Channel only n v l 1 201 Knight I n l e t only goc 1 nvr 1 nvz 1 p i b 1 202 APPENDIX D - RELATIVE ABUNDANCE OF SELECTED SPECIES This appendix contains the r e l a t i v e abundance for selected species and species-groups, and the t o t a l absolute abundance at each s i t e . Sites are ranked head to mouth. See Figs. 5-7 for s i t e locations. SK = Skeletonema costatum TH = Thalassiosira aest i v a l i s f. pac i f ica CH = Chaetoceros spp. resting spores PR = Paralia sulcata TZ = Thalassionema nitzschioides MP = marine planktonic species ML = marine l i t t o r a l species FW = freshwater species V/GDW = valves/gram dry weight 203 ZONE A SITE SK TH CH PR TZ MP ML FW V/GDW NO. Howe Sound 1974 53 40.6 1 5 15 0 4.3 3.7 8.2 9.4 2.7x107 27 59.5 15.5 4.6 1 .4 4.5 4.3 0 7 6.0x107 29 50.6 17.7 13 3.4 6.3 7.9 1 .2 0.8 7.0X10 7 28 52.8 16.4 11.6 1 .9 7.5 4.2 0.5 2.8 5.3x107 45 38.6 19.9 20.2 0.8 6.8 7.2 2 1 .6 6.6x107 31 45.9 22.3 13.0 1 .7 6.4 11.7 1 .7 0.8 5.7X10 7 32 37. 1 30.8 9.7 3 8.9 8.1 1 .6 0.4 5.3X10 7 33 37 26.9 1 1 3.7 9.6 9 1 .4 1 .4 6.1x107 56 47. 1 13.7 15.2 3.9 6.4 5.2 1 5.5 9.9x107 37 36.8 23.5 9.9 3.4 8.1 9.5 0.8 2 7.1X10 7 35 40.2 30. 1 9.1 2.9 3.3 8.5 4.5 1 .5 7.9X10 7 38 26.5 22.3 10.7 6 1 4 9.7 5.2 1 .9 6.2x107 57 32.4 20. 1 17.1 4.9 11.8 8.2 0 5.5 6.1X10 7 34 37.4 23.2 12.4 2 13.3 9.2 3 2 6.4x107 36 22.7 21.4 16.8 9.6 1 4 10.4 4.3 2.1 3.4x108 30 50.4 15.8 11.3 2.1 10 5.9 2.9 0.4 1.2x10s 39 39. 1 25.2 10.5 2.5 8.4 8.5 3.4 2.4 7.9x107 40 50.7 17.4 10.8 5.6 4.7 9.2 1 3.8 9.1x107 41 44. 1 15.9 12.7 4 7 5.3 2.2 5.7 8. 1x107 42 34.6 18.7 21.5 6.1 0.9 8.2 3.4 5.6 8.7X10 7 58 30.6 12.6 24.9 5 11.3 8.4 1 .9 4.4 6.1X10 7 43 38 19.7 18.8 5.2 7.5 10.3 1 .5 0.9 6.1x1O7 54 27.4 19 19.5 4.9 8.4 8.7 4.2 4.7 1.0x108 55 31.5 24.5 15.5 1 .5 6.5 10 7.5 2 4.8x107 Howe Sound 1977 1 15.3 59. 1 6.8 0.9 2.1 11.1 0.4 5.2 1.3x107 2 36.4 32.7 8.4 2.3 9.3 8.1 0 0 3.3X10 7 3 32.9 32.9 9.3 6. 1 7 13.2 0 1 .4 4.1x1O7 4 59 25 1 .9 0.9 3.8 6.5 0 0 1.9x107 5 17.7 48.8 2.4 7.2 6.2 15.7 1 0 2.3x107 6 23.6 40.4 11.1 7.2 5.3 11.4 0 0.5 3.8X10 7 Indian Arm 1981 94 40.2 22.3 10.4 0.4 5.7 9.2 6.8 1 .6 2.3X10 8 93 55.8 13.8 9.2 1 .3 2.9 11.7 3.7 1 .2 2.7X10 8 Indian Arm 1976 1 1 31.3 32.2 6.3 0 3.4 24.3 1 .5 0.5 2.1x107 1 2 41.3 27.2 8.5 0.9 5.8 14.9 0.4 0.8 4.3X10 7 Burrard Inlet 44 37.7 25.5 10.8 1 .0 6.4 5.1 2 10.5 2.3x107 92 37. 1 12.5 13.6 0.8 7.1 8.9 7.9 13.3 4.2x107 Saanich Inlet 59 37.7 6.1 30.8 6.9 8.2 6.9 2.1 0.9 1.2x10s 60 35.5 17.5 25.5 3.1 6.1 9.9 2.1 1 .2 1.2x1O8 204 SITE SK TH CH PR TZ MP ML FW V/GDW NO. 63 28 .1 19.3 31 . 2 4.8 7.5 4.9 3 0.8 8 . 0 X 1 0 7 61 2 0 . 6 23.4 24 .3 5 .1 1 0.7 8.3 5.7 1 .9 5 . 0 X 1 0 7 62 8.4 31.6 25 .3 6 . 2 1 0.7 13.1 3.4 2 . 2 2 . 2 X 1 0 7 ZONE B SITE SK TH CH PR TZ MP ML FW V/GDW NO. Jerv i s Inlet , 1980 26 36.4 1 0.5 38 .8 0 1 .4 13.7 1 .5 3 3.6 X 1 0 25 23.5 5 .1 52 .9 0 8 . 1 18 1 .7 0 4. 1 X 1 0 7 24 19.1 16.1 40 . 0 0.4 7.8 23.9 2 . 2 0 3 . 8 X 1 0 7 23 9 . 2 1 1 . 5 41 . 2 5.5 14 . 2 19.8 3.4 1 2 . 1x10 7 2 2 16.7 13.1 31 . 2 4.5 14.9 1 2 . 9 2.8 1 2.2x1 0 7 2 1 3.7 14.5 26 .3 1 0.7 17.8 27.4 2.4 0.8 2.3x1 0 7 J e r v i s Inlet 1981 66 25.2 11.9 42 0 5.8 21 . 5 0.8 1 . 6 4.8x107 67 23.6 9.6 34.6 0. 9 3.5 25 5. 1 2. 5 5.6X10 7 69 21.3 6.8 41 .7 0 2.6 23. 4 2.1 3. 8 2.4x107 64 22.5 8.7 47.5 0. 4 6.7 15. 2 2.4 0. 8 4.5X10 7 65 29.4 10.4 40.2 0 5.6 22. 8 0.8 1 . 6 6.5X10 7 70 21.4 7. 1 41.1 0. 8 4.6 25. 1 5.8 0. 8 3.4X10 7 71 18.5 13.9 36 0. 9 9.3 15. 6 6.6 1 4.7x107 73 23 7.1 40. 1 0. 8 6.7 23. 3 3.2 2. 5 7.5x107 72 16.6 15.6 36.4 1. 9 7.6 21 . 7 3.5 2. 3 6.9x107 68 22.2 9.6 35.5 2. 9 7.5 16. 3 4.5 2. 8 5.9X10 7 Hotham Sound 76 1 1 9.6 34.7 6. 4 5.9 31 . 8 3.8 0 6.0x107 75 34.3 9.8 29.5 2 2 20. 8 6 0. 5 6.7x107 74 25.5 11.5 29.4 4. 7 6.8 19 3 2. 2 1.0x108 Agamemnon ( Channel 78 10.5 18.2 31.9 0. 5 11.8 25. 6 5.1 0 1.7x107 77 17.1 27.5 24.2 0. 5 4.7 27. 8 7.7 0 5.9X10 7 Malaspina : Stra i t 79 16.9 17.4 25.9 7 12.7 17. 1 4.8 1 . 4 7.2x107 Sechelt Inlet 52 45.5 8 22.3 0 3.3 22. 6 2.4 0. 5 5.0x107 46 31.7 26.2 21.3 0 3 20. 3 2 0 1.0x108 47 12.3 21 .5 26.4 0. 9 12.3 24. 2 3.2 0 5.7X10 7 48 12.4 17.9 23.5 0. 9 10.7 29. 7 4.4 0. 9 9 . 0 X 1 0 7 49 17.1 27. 1 16.8 0. 4 3.8 31 . 6 3.2 0. 4 2.5x107 50 18.6 17.6 32.4 0. 5 4.3 27. 5 0.5 0 8 . 3 X 1 0 7 51 14.9 20.7 23.2 0. 5 6.3 32 2.8 1. 9 1.9x1O7 Hidden Basin 95 19.9 9.7 34.5 2.7 1.3 26.1 10.7 3.5 2.1x10 205 SITE SK TH CH PR NO. Texada Island South 16 7.2 8.2 31 .3 15.4 ZONE C SITE SK TH CH PR NO. Bute Inlet 89 22.4 31.5 15 . 1 0.4 90 15.5 16.7 27 . 1 2.1 91 7.9 26 29 .3 6 7 1 .4 37.3 28 .2 15.9 Cordero Channel 20 13.4 18.2 20 .3 19.9 Toba Inlet 88 16.6 27 1 3 .7 0.4 87 19.6 31 .8 20 . 1 0 86 17.1 22. 1 31 .2 2.3 85 12.9 22.6 27 .2 3.2 Pryce : Channel 82 10.6 1 1 42 .7 3.1 Homfray Channel 84 9.2 18 37 .9 1 .8 83 8.6 5.9 46 . 1 3.2 Desolat ion Sound 81 3.3 6.5 41 .5 3.7 Pendrell Sound 1 3 1 .3 4 8. 4 64.7 1 4 14.2 10.9 31 .2 15.2 East Gorge Harbor 15 11.6 4.5 39.7 15.6 Heriot Bay 19 6.3 15.9 31.3 5.3 S u t i l Channel 80 5.5 10.9 37.7 6.3 Texada Island North 17 11.9 22.9 22.4 19.5 TZ MP ML FW V/GDW 12.5 9.7 3.2 0.5 3.4X10 7 TZ MP ML FW V/GDW 12.5 12.1 7.4 5 15.1 24 21.9 11.2 4.8 2.9 3.3 0.5 1 .7 0.8 2.9 0 1.9x107 6.0X10 7 7.7X10 7 1.5X10 7 10 6.9 4.3 1 .8 4. 1X10 7 2.2 4.2 8.1 7.8 1 9 16.3 14.4 18.2 14.9 2.9 3.4 7.6 9.2 4.2 1 .9 2.4 1.3x107 2.8X10 7 3.6X10 7 5.0x107 12.8 15.1 4.3 1 .3 1 . 1 x 1 0 88.8 7.2 21.9 25.8 4.2 4.3 0 2 7 . 5 X 1 0 6 7.7x107 11.2 21.4 10 1 .9 5.8X10 7 1 .3 4.7 10.6 23.2 11.9 5.7 0 1 .5 1.9x106 1.3x107 2.7 16.3 18.5 0.4 6.6x107 1 27 13.7 3.9 7.2x10s 7.6 2.0. 1 9.8 0.4 1.2x10s 9.5 10.4 1 2.4 2.3x107 206 SITE SK TH CH PR TZ MP ML FW NO. Knight Inlet 8 3.4 59. . 1 2.5 0 17.2 8.8 1 8 9 18.9 31 . ,5 29.3 9 4. 1 5.7 1 0 10 17.7 32. , 1 20. 1 9.6 4.3 15.7 0 1 V/GDW 4.4x10 4.2x10 1.7x10 Queen Charlotte S t r a i t 18 9.2 20.3 31.3 11.5 12.9 9.7 3.2 0.5 3.4x10 207 APPENDIX E - SITE DATA This appendix contains the valve counts for species at each of the 95 s i t e s . The key below i s an alphabetic l i s t of the i n l e t s . The numbers refer to the s i t e numbers. Sites are ordered numerically Agamemnon Channel 77-78 Burrard Inlet 44,92 Bute Inlet 1977 7 Bute Inlet 1981 89-91 Cordero Channel 20 Desolation Sound 81 East Gorge Harbor 15 Heriot Bay 19 Hidden Basin 95 Homfray Channel 83-84 Hotham Sound 74-76 Howe Sound 1974 27-43, 45, 53-58 Howe Sound 1976 6 Howe Sound 1977 1-5 Indian Arm 1976 11-12 Indian Arm 1981 93-94 •> Jervis Inlet 1980 21-26 Jervis Inlet 1981 64-73 Knight Inlet 8-10 Malaspina S t r a i t 79 Narrows Inlet 52 Pendrell Sound 13-14 Pryce Channel 82 Queen Charlotte S t r a i t 18 Saanich Inlet 59-63 Salmon Inlet 50-51 Sechelt Inlet 46-49 S u t i l Channel 80 Texada Island 16-17 Toba Inlet 85-88 208 Site 1 Howe Sound head 1977 depth = 283m d i v e r s i t y = 2.42 valves/gram dry weight = 1.3x107 COP 1 DIT 6 THP 1 39 CHL 4 SKC 36 EUA 2 CHD 1 TNZ 5 DIH 2 DIS 1 CHR 6 CHC 5 GOA 4 PAR 2 CBS 1 THN 5 HNX 3 ACS 1 NVS 1 THC 3 RHZ 2 CYS 4 ML I 1 SITE 2 Howe Sound head 1977 depth = 200m d i v e r s i t y = 2.50 valves/gram dry weight = 3.3X10 7 THP 70 SKC 78 TNZ 20 CHR 15 DIS 1 PAR 5 GOS 4 CHD 3 CYS 4 THN 4 DIT 4 RHZ 4 CSM 2 Site 3 Howe Sound head 1977 depth = 274m d i v e r s i t y = 2.75 valves/gram dry weight = 4.1X10 7 THP 70 SKC 70 PAR 13 CHR 12 209 RHZ 5 DIT 4 TNZ 15 LED 2 THG 3 THC 1 CSM 2 CCO 4 ML I 2 CHE 5 COP 1 CHA 3 FRC 1 Site 4 Howe Sound middle 1977 depth = 164m d i v e r s i t y = 1.90 valves/gram dry weight = 1.9X10 7 THP 53 SKC 1 25 DIS 1 CSM 5 TNZ 8 THG 1 CHZ 1 THN 3 DIT 6 CHC 3 CBS 1 EUS 1 PAR 2 RHZ 2 Site 5 Howe Sound mouth 1977 depth = 65m d i v e r s i t y = 2.53 valves/gram dry weight = 2.3X10 7 THP 102 THC 9 DIT 10 PAR 15 TNZ 13 ACS 1 THN 3 CCO 6 CSM 5 SKC 37 GRM 1 CHE 3 RHZ 2 CHR 2 Site 6 Howe Sound mouth 1976 210 depth = 146m d i v e r s i t y = 2.66 valves/gram dry weight = 3.8X10 7 THP 84 SKC 49 PAR 15 CHR 23 RHZ 4 DIT 7 TNZ 1 1 LED 1 CYS 1 THG 1 THD 3 THC 2 THN 2 CSM 1 CCO 1 CMG 1 COC 1 DPS 1 Site 7 Bute Inlet mouth 1977 depth = 700m d i v e r s i t y = 2.56 valves/gram dry weight = 1.5X10 7 TNZ 1 1 BDA 1 DIT 3 CCO 5 THP 82 PAR 35 RHZ 4 CHC 2 CHR 60 THD 2 DIS 5 THG 2 THC 4 SKC 3 CBS 1 SITE 8 Knight Inlet head depth = 260m d i v e r s i t y = 2.21 valves/gram dry weight = 4.4x10s NVA 1 1 THP 120 EUP 6 THC 4 SKC 7 TNZ 35 NZA 2 DIH 2 CCO 1 21 1 PIB 1 CHD 2 NVR 1 CHR 3 NVZ 3 FRV 1 TXL 2 GOC 1 TBN 1 Site 9 Knight Inlet middle depth = 360m d i v e r s i t y = 2.63 valves/gram dry weight = 4.2X10 7 PAR 2 0 THP 7 0 SKC 42 TNZ 9 CHR 52 DIS 7 CHD 13 THC 5 AHS 1 CSC 1 COA 1 THG 1 Site 10 Knight Inlet mouth depth = 245m d i v e r s i t y = 3.02 valves/gram dry weight = 1.7X10 7 COO 4 THP 67 PAR 20 THN 1 3 SKC 37 CHD 1 7 CYT 2 TNZ 9 CHR 24 CCO 7 THC 2 DIT 1 DIS 3 RHZ 2 CHG 1 SITE 11 Indian Arm head 1976 depth = 106m d i v e r s i t y = 2.73 valves/gram dry weight = 2.1X10 7 DIS 29 THP 67 CHD 10 212 SKC 65 THE 5 TNZ 7 THC 1 CBS 2 THN 5 CHC 1 BDA 2 CHR 2 COS 2 CYS 5 LED 1 TBF 1 DIT 1 NZS 1 ACP 1 Site 12 Indian Arm mouth 1976 depth = 230m d i v e r s i t y valves/gram dry weight THP 61 CHR 10 DIS 21 TNZ 1 3 SKC 92 CHD 9 CYS 5 COO 2 CSS 1 PAR 2 THG 3 HYS 1 OPM 1 DIT 2 FRP 1 Site 13 Pendrell Sound head depth = 80m d i v e r s i t y = 2.44 valves/gram dry weight = 1.9x10s PAR 50 THP 9 CYS 3 CHA 5 CDS 2 GRM 5 SKC 3 RHZ 3 PLS 4 CHR 8 CCO 4 GLD 3 CHC 3 = 2.51 = 4.3X10 7 213 NVS 1 HYR 10 CMG 1 LED 1 RAS 2 NVY 1 CHY 3 TNZ 3 THN 1 COT 1 COO 1 DPB 1 CSC 1 Site 14 Pendrell Sound mouth depth = 410m d i v e r s i t y = 4.03 valves/gram dry weight = 1.3X10 7 SKC 30 DPS 1 THP 23 PAR 32 CHZ 2 CHM 14 FRB 1 DIS 4 CCO 4 CHA 1 1 HYR 2 CHC 8 CHD 2 CHR 28 CHE 1 CSC 1 GRG 1 CYS 1 PLS 1 LED 3 COC 1 COP 2 CON 1 RHZ 1 DIT 3 ACS 2 OPP 1 ACP 2 TNZ 10 THN 12 THC 4 CLA 1 GRM 1 Site 15 East Gorge Harbor 214 depth = 15m d i v e r s i t y = 4.17 valves/gram dry weight = 6.6X10 7 THP 10 SKC 26 CHR 39 TNZ 6 PAR 35 THC 4 THG 1 SYT 1 GYS 5 DIS 5 DIF 1 THN 4 MLM 1 DPS 1 CPS 1 CCO 2 CHA 18 RHZ 2 CHD 8 CHC 1 1 NZP 3 CHM 8 SUF 1 OPR 7 CYS 2 ACS 1 DIT 1 AHS 2 NZB 2 COC 1 DPP 1 CHL 2 LED 6 CHV 3 CSC 1 AME 2 Site 16 Texada Island south depth = 400m d i v e r s i t y = 3.85 valves/gram dry weight = 3.4x107 THP 17 SKC 15 CHR 42 TNZ 26 PAR 32 LED 3 CHC 9 CHD 10 CHS 3 CHM 2 THN. 3 215 THC 3 DIS 8 DIT 5 COC 1 FRC 1 CSC 2 CDS 6 CCO 3 THG 2 CYS 3 GRM 3 BDA 2 TAF 2 RHZ 4 ACS 1 Site 17 Texada Island north depth = 400m d i v e r s i t y = 3.40 valves/gram dry weight = 2.3X10 7 THP 48 SKC 25 CHR 25 TNZ 20 PAR 41 FRC 4 COO 2 CHD 5 THC 4 CHC 12 RHZ 1 ACS 2 THN 2 CHM 5 DIT 3 NVS 1 LED 3 DPS 1 COT 1 CCO 2 THG 1 DPP 1 TAS 1 Site 18 Queen Charlotte S t r a i t depth = 120m d i v e r s i t y = 3.31 valves/gram dry weight = 3.4x10s THP 44 SKC 20 CHR 53 TNZ 28 PAR 25 THC 4 216 COT 5 CHD 9 CHY 3 CHC 3 COC 3 NVS 1 FRP 1 DIT 1 CYS 1 AHS 1 HYS 2 CDS 4 CCO 2 DIS 1 CSM 5 CSC 1 Site 19 Heriot Bay depth = 22m d i v e r s i t y = 4.37 valves/gram dry weight = 7.2x10s THP 33 SKC 13 CHR 39 TNZ 2 PAR 1 1 THC 1 4 CHM 5 AMS 3 SYT 1 COT 1 CHV 1 1 DPS 6 DPC 2 MLM 8 RHZ 2 CHA 1 GLD 2 COP 1 DIS 6 CHD 4 COO 1 TRS 1 ACS 2 CYS 3 CSC 1 CCO 4 NVT 1 COC 4 LED 2 CON 1 BDA 2 TRA 1 MLS 2 217 COM 2 ROM 1 HYR 2 ACP 1 CHC 5 CDS 4 CHY 1 TXL 2 SITE 20 Cordero Channel depth = 120m d i v e r s i t y = 3.28 valves/gram dry weight = 4.1X10 7 THP 42 SKC 31 PAR 46 CHR 42 TNZ 23 DIS 2 CHD 8 CHC 1 DIT 2 CDS 10 CCO 4 CSS 1 CSM 6 THG 2 THN 1 FRP 2 RHZ 1 COO 1 GOS 1 TAF 1 FRY 1 NVL 2 CHA 1 SITE 21 Jervis Inlet mouth 1980 depth = 308m d i v e r s i t y = 3.70 valves/gram dry weight = 2.3X10 7 THP 35 SKC 9 CHR 2 9 TNZ 43 DIS 1 STS 3 PAR 26 CHC 27 CDS 2 CHD 1 1 CHA 10 CYS 1 1 RHZ 4 218 CHM 1 5 CSC 1 GRM 1 DIT 4 PLS 1 COW 1 TBF 1 FRC 1 CMG 1 ACS 1 DPP 2 AHS 1 CHE 1 SITE 22 Jervis Inlet mouth 1980 depth = 585m d i v e r s i t y = 3.50 valves/gram dry weight = 2.2X10 7 THP 2 9 SKC 37 CHR 49 TNZ 33 CHA 9 COP 1 CHC 12 CHD 7 CSM 3 ACP 2 CHM 1 DIT 1 0 CYS 3 PAR 1 0 OPM 1 CDS 3 HYR 1 AHS 1 DIS 1 CCO 3 CMG 1 AHL 1 NVS 1 GOS 1 RHZ 2 SITE 23 Jerv i s Inlet middle 1980 depth = 443m d i v e r s i t y = 3.79 valves/gram dry weight = 2.1X10 7 THP 25 SKC 20 CHR 46 TNZ 31 CYS 6 GRA 1 219 CHA 14 RCS 1 CHC 14 CSC 1 GRO 1 HYR 1 NZP 1 PAR 12 DPS 1 DIT 1 1 THC 3 CHD 1 0 RHZ 1 CHY 4 CSM 3 NVS 1 COC 1 CMG 1 TBF 1 CCO 1 CHM 2 GOS 1 DIS 1 CDS 2 SITE 24 J e r v i s Inlet middle 1980 depth = 547m d i v e r s i t y = 3.39 valves/gram dry weight = 3.8X10 7 THP 3 7 SKC 44 CHR 45 TNZ 1 8 DIT 4 HYR 2 CHA 20 CHD 22 CHE 4 CCO 5 CYS 1 2 CHC 1 THN 3 RHZ 1 THC 2 ACP 2 PAR 1 DPP 2 NZP 1 CMG 1 THG 2 COO 1 Site 25 Je r v i s Inlet middle 1980 220 depth = 532m d i v e r s i t y = 3.46 valves/gram dry weight = 4.1X10 7 THP 12 SKC 55 CHR 43 TNZ 19 CHD 21 CSM 3 CHA 22 CHC 12 DIS 4 THG 2 CYS 2 ACP 2 CHM 20 CHL 1 RHZ 2 STS 1 CHY 5 NZP 1 DIT 3 CBS 1 EUS 2 CSC 1 LED 5 Site 26 Jervis Inlet head 1980 depth = 314m d i v e r s i t y = 3.18 valves/gram dry weight = 3.6X10 7 THP 22 SKC 76 CHR 36 TNZ 3 CHA 19 CHC 6 TBF 1 CHM 10 CCO 1 CHD 9 CSM 4 GRO 1 EUS 1 FRV 2 FRA 2 DIT 3 THN 1 NZP 1 SYS 3 FRN 1 CYS 2 RHZ 1 ACP 1 CHL .1 221 AHS 1 CSS 1 Site 27 Howe Sound head 1974 depth = 233m d i v e r s i t y valves/gram dry weight THP 34 SKC 131 CHR 4 TNZ 1 0 THC 1 MEC 1 CYT 1 AST 2 PAR 3 TBN 2 CHD 1 NEB 1 FRN 2 GOO 2 CYS 1 CHM 2 AHC 3 DIT 2 CHC 3 STM 1 FRP 2 THE 1 THN 2 THG 2 DIS 1 CSM 5 Site 28 Howe Sound middle 1974 depth = 193m d i v e r s i t y valves/gram dry weight THP 3 5 SKC 113 CHR 15 ) TNZ 16 CHS 1 CHM 5 CYS 4 DIT 1 FRS 1 PAR 4 RHZ 1 CHD 2 NZP 1 DIH 2 CCO 2 THG 1 CHC 2 2.42 6.0X10 7 2.54 5.3x107 222 EUS 2 FRL 1 NVS 1 CSM 1 CYT 2 LED 1 SITE 29 Howe Sound middle 1974 depth = 164m d i v e r s i t y valves/gram dry weight THP 42 SKC 120 CHR 1 1 TNZ 1 5 DIS 1 CHY 4 EUS 1 CSM 2 CHM 1 0 CSP 1 CMG 2 TAF 2 CYS 5 AHK 1 CHD 2 DIT 3 CHC 3 CHA 1 CDS 1 PAR 8 LIE 2 SITE 30 Howe Sound middle 1974 depth = 243m d i v e r s i t y = 2.63 valves/gram dry weight = 1.2x10 THP 38 SKC 121 CHR 16 TNZ 24 THC 4 SYT 2 CHD 2 AHS 2 CHM 6 CDS 3 CYS 2 PAR 5 DIT 1 CHC 3 CCO 1 TAF 1 CBS 1 = 2.62 = 7 . 0 X 1 0 7 223 THE 1 NVA 2 STM 1 DIS 1 LIE 2 RHZ 1 SITE 31 Howe Sound middle 1974 depth = 229m d i v e r s i t y = 2.72 valves/gram dry weight = 5.7X10 7 THP 52 SKC 107 CHR 12 TNZ 15 THC 1 CCO 3 PLS 1 CHA 6 CHC 3 THN 3 CHM 7 PAR 4 DIS 1 CYS 4 OPM 1 DIT 2 RHZ 4 COO 1 GYG 1 AHB 2 STM 1 CHY 2 Site 32 Howe Sound middle 1974 depth = 243m d i v e r s i t y = 2.70 valves/gram dry weight = 5.3x!07 THP 73 SKC 88 CHR 1 2 TNZ 21 CHD 6 STM 1 RHZ 5 DIT 3 CHC 2 NZB 2 CSC 1 TXL 1 PAR 7 CYS 2 COO 1 CHA 3 224 NZP 1 CCO 3 LED 1 CSM 4 SITE 33 Howe Sound middle 1974 depth = 244m d i v e r s i t y valves/gram dry weight THP 59 SKC 81 CHR 12 TNZ 21 THC 1 NZI 2 COS 1 CCO 3 GRM 1 CHC 4 FRP 1 CYS 3 PAR 8 THN 2 RHZ 1 CHM 1 DIT 4 CHD 2 CSM 3 SYF 2 CHA 1 AST 2 CHY 2 CHL 2 SITE 34 Howe Sound middle 1974 depth = 251m d i v e r s i t y valves/gram dry weight THP 47 SKC 76 CHR 13 TNZ 27 CHA 4 RHZ 1 CHM 3 CHV 4 PAR 4 DIE 1 ML I 1 DIF 1 DIT 4 GOO 3 CCO 1 CDS 2 2.92 6.1X10 7 2.92 6.4X10 7 225 NZS 1 THN 2 CYS 2 CSC 1 GOS 2 LIG 2 CHC 1 Site 35 Howe Sound middle 1974 depth = 179m d i v e r s i t y = 2.83 valves/gram dry weight = 7.9X10 7 THP 63 SKC 84 CHR 7 TNZ 7 CHD 1 RHZ 1 CYS 6 AHH 1 FRL 1 THN 1 PAR 6 CHC 3 DIS 2 CHM 4 DIT 3 LIE 2 CSM 3 GYG 1 GOO 2 NZS 1 GYT 1 CHL 2 NZL 2 NZD 2 CCO 1 CHE 2 Site 36 Howe Sound middle 1974 depth = 243m d i v e r s i t y = 3.47 valves/gram dry weight = 3.4X10 8 THN 4 THC 2 THP 49 SKC 52 CHR 22 TNZ 32 CHD 1 NZA 2 SUI 1 DIT 4 PAR 22 226 ACS 1 CYS 1 NZG 2 CHC 4 OPR 1 RHZ 2 CHM 9 CHL 1 THE 5 HNA 1 CCO 1 MLT 1 CHY 1 CDS 2 SYA 2 THG 1 CHA 1 FRV 1 NZD 1 Site 37 Howe Sound middle 1974 depth = 250m d i v e r s i t y = 3.05 valves/gram dry weight = 7.1X10 7 THP 55 SKC 86 CHR 13 TNZ 1 9 STM 1 CHM 2 CHD 3 CYS 4 CSM 13 RHZ 3 CHC 4 PAR 8 THN 6 NVS 1 DIT 5 CBM 1 FRV 1 NZS 1 CYT 1 ACS 1 NZL 1 CCO 2 GOO 1 THC 1 CHV 1 Site 38 Howe Sound middle 1974 depth = 246m d i v e r s i t y = 3.39 valves/gram dry weight = 6.2X10 7 227 THP 48 SKC 57 CHR 18 TNZ 30 RHZ 5 CHD 3 PAR 13 DIT 4 NZD 1 NZB 1 NZP 1 AHS 1 CCO 1 STM 1 NVS 3 NZS 2 CDS 5 TBF 1 GOO 2 SUI 1 THN 5 DIS 3 CSM 2 CYS 1 NZL 2 LED 1 THE 1 CHM 2 SITE 39 Howe Sound middle 1974 depth = 124m d i v e r s i t y = 2.87 valves/gram dry weight = 7.9X10 7 THP 60 SKC 93 TNZ 20 CHR 1 3 RHZ 6 CCO 3 STA 1 PAR 6 THN 2 CHM 9 THC 3 CSM 2 CHE 2 ; CHC 1 FRB 2 SYT 4 DIT 3 GOO 2 THG 1 DIH 1 MLM 4 228 SITE 40 Howe Sound middle 1974 depth = 221m d i v e r s i t y = 2.67 valves/gram dry weight = 9.1X10 7 THG 39 SKC 108 CHR 9 TNZ 10 CHY 4 CHA 5 RHZ 3 MLM 1 FRB 1 CHC 1 NZM 1 PAR 1 2 DIT 3 CYT 2 CHM 4 CSM 1 DIS 2 HNA 1 THC 1 GOO 4 AST 1 Site 41 Howe Sound middle 1974 depth = 242m d i v e r s i t y = 3.03 valves/gram dry weight = 8.1X10 7 THP 36 SKC 100 CHR 18 TNZ 16 THC 1 CHA 1 PAR 9 CSM 7 DIS 1 CHC 2 GOO 4 RHZ 3 AST 1 ML I 3 ACS 2 DIT 3 CYT 2 CCO 1 CHM 6 NVF 2 HNA 2 CHD 2 CYS 2 AHH 1 229 DPS 1 SYU 1 Site 42 Howe Sound middle 1974 depth = 230m d i v e r s i t y = 3.27 valves/gram dry weight = 8.7X10 7 THP 40 SKC 74 CHR 32 THC 1 THN 1 TNZ 2 PAR 1 3 NVS 1 CYT 3 AST 1 CHE 1 CSC 2 DIT 7 RHZ 2 CHC 3 GRO 1 AHS 1 THG 2 CHM 8 AHH 1 CCO 2 GYF 1 CHD 2 MLM 1 BDA 1 MEC 4 CBM 1 CYS 1 OPM 2 AHF 2 THD 1 Site 43 Howe Sound mouth 1974 depth = 256m d i v e r s i t y = 2.87 valves/gram dry weight = 6.1X10 7 THP 42 SKC 81 CHR 30 TNZ 16 THN 3 DIS 1 PAR 1 1 CYS 3 NVF 1 230 CHM 2 CHA 4 DIT 4 CHD 3 CCO 4 CHC 1 CDS 1 RHZ 3 FRP 2 NZB 1 Site 44 Burrard Inlet east 1980 depth = 60m d i v e r s i t y = 3.08 valves/gram dry weight = 2.3X10 7 THP 52 SKC 7 7 CHR 17 TNZ 1 3 NZC 3 PAR 2 CYS 2 TBF 1 DIT 2 CHC 3 NVS 1 NZT 2 DIS 3 CYT 2 SYS 1 AHS 2 AHE 1 CYC 2 AHW 2 FRP 3 MLD 2 CCO 1 MLM 1 DPS 1 CSE 1 THG 1 CHA 2 TBN 1 CDS 1 CYA 2 Site 45 Howe Sound middle 1974 depth = 128m d i v e r s i t y = 2.93 valves/gram dry weight = 6.6X10 7 THP 47 SKC 91 231 CHR 23 TNZ 1 6 THC 1 THN 2 COO 1 NVS 2 CHM 23 DIT 3 RHZ 5 DIH 1 DIS 3 CHC 1 NVF 1 FRP 1 CDS 2 CHA 1 FRV 1 AHH 2 CSM 5 PAR 2 NZS 1 CYO 1 Site 46 Sechelt Inlet mouth depth = 279m d i v e r s i t y =3.11 valves/gram dry weight = 1.0x10 THP 53 SKC 64 CHR 13 TNZ 6 THN 9 CHL 5 RHZ 1 CHD 2 CHM 15 AHB 1 DIS 6 DIT 5 BDA 1 LED 5 ACP 1 CHC 2 AMS 2 CHA 6 CYS 4 MLM 1 Site 47 Sechelt Inlet middle depth = 258m d i v e r s i t y = 3.84 valves/gram dry weight = 5.7X10 7 232 THP 47 SKC 27 CHR 27 TNZ 27 THC 3 THN 16 NZN 2 CHD 2 CBS 1 CHA 7 GOS 1 DIS 5 LED 2 CYS 4 THG 2 DIT 6 CHM 1 3 MLM 2 CHC 6 RHZ 1 CHY 3 STS 1 ACP 1 THD 3 AMS 1 PAR 2 CDS 1 CSM 5 BDA 1 Site 48 Sechelt Inlet middle depth = 235m d i v e r s i t y = 3.98 valves/gram dry weight = 9.0x107 THP 42 SKC 29 CHR 27 TNZ 25 THC 2 THN 19 CYS 13 CHD 8 DIT 1 1 DIS 6 BDA 2 CSM 4 THD 1 PAR 2 CMG 2 RHZ 3 CHC 12 DMM 2 CBS 1 233 THG 3 NVS 2 CHY 4 CDS 1 CHA 4 LED 3 NVQ 2 FRP 2 GRG 2 ACP 1 Site 49 Sechelt Inlet head depth = 107m d i v e r s i t y = 3.70 valves/gram dry weight = 2.5X10 7 THP 65 SKC 41 CHR 18 TNZ 9 THC 5 THN 21 CYS 16 CHA 5 DIS 12 CHC 3 CHD 5 CSM 5 NZS 1 CHM 4 CMG 1 CCO 5 NVS 3 ATS 1 PAR 1 FRP 1 RHZ 2 THG 1 CHY 5 OPR 2 THD 1 CDS 2 ACS 1 DIT 1 MLM 1 HYS 2 Site 50 Salmon Inlet middle depth = 260m d i v e r s i t y = 3.54 valves/gram dry weight = 8.3X10 7 THP 37 SKC 39 234 CHR 27 TNZ 9 THC 2 THN 26 CHM 16 EBT 2 PAR 1 CHD 18 COO 3 CYS 5 ATS 2 DIT 4 LED 1 CHC 3 DIS 5 CCO 3 CHA 4 NZS 1 STS 1 MLM 1 Site 51 Salmon Inlet head depth = 220m d i v e r s i t y = 3.68 valves/gram dry weight = 1.9X10 7 THP 46 SKC 33 CHR 1 5 TNZ 1 4 THN 37 THC 1 DPB 1 CHM 19 DIT 6 CHD 7 CYS 1 1 EUS 2 DIS 3 CHA 3 ACS 2 AHC 2 COO 1 ACP 1 COP 1 AHE 1 CHC 3 THG 2 PAR 1 RHZ 1 CHY 4 CCO 2 TBN 1 MLM 2 235 Site 52 Narrows Inlet head depth = 84m d i v e r s i t y =3.11 valves/gram dry weight = 5.0x107 THP 17 SKC 97 CHR 5 TNZ 7 THN 8 THC 1 CHY 5 CHM 15 DIS 13 DIT 6 CHA 5 CYS 4 ACP 3 CHD 15 CHL 1 EBT 2 MLM 1 NVS 1 FRP 1 CCO 1 THG 1 RHZ 1 AHO 1 CHC 2 Site 53 Howe Sound head 1974 depth = 220m d i v e r s i t y = 3.22 valves/gram dry weight = 2.7X10 7 THP 31 SKC 84 CHR 2 TNZ 9 SYT 5 NVD 7 CHM 28 DIT 4 CHA 1 RHZ 1 CSS 1 NVP 1 AST 3 AHX 2 COF 2 GOO 2 TBN 3 AMR 1 NZV 3 EUE 1 236 FRP 1 RAA 1 EUP 2 NVS 1 MLT 2 CSM 3 SYS 1 DIH 2 THC 1 CCO 1 SYU 1 THN 2 Site 54 Howe Sound mouth 1974 depth = 235m d i v e r s i t y = 3.68 valves/gram dry weight = 1.0x10 THC 1 THP 4 3 SKC 62 CHR 25 TNZ 19 COP 1 CHD 1 PAR 1 1 NVD 2 DIT 5 SYT 1 NVS 2 CYO 4 FRP 1 CHM 1 1 CHC 6 CHA 1 CDS 1 THG 2 NZS 4 LED 2 GOO 2 DIS 1 TAF 1 NW 1 NVT 1 AHC 1 TBN 1 RHZ 2 NZG 2 BDA 1 NVU 1 CCO 1 CYT 1 NZF 1 AHS 2 237 Site 55 Howe Sound mouth 1974 depth = 99m d i v e r s i t y = 3.31 valves/gram dry weight = 4.8X10 7 THP 49 SRC 63 CHR 13 TNZ 13 DIT 9 RHZ 2 MLM 2 CHD 2 AHH 4 CHC 4 DIS 1 CYS 4 AMR 1 AME 2 GRX 1 THD 2 PAR 3 CYO 2 CHM 12 SYT 2 AHS 2 COL 1 NZO 2 COT 1 FRP 1 CCO 1 OPR 1 Site 56 Howe Sound middle 1974 depth = 245m d i v e r s i t y valves/gram dry weight THP 28 SKC 96 CHR 16 TNZ 13 CHC 2 RHZ 2 CSM 3 GOO 4 NZS 1 CHM 12 CCO 1 AHR 3 PAR 8 DIT 3 THD 2 CYO 2 DIS 1 = 2.87 = 9.9X10 7 238 CYS 1 CHD 1 THG 1 FRS 2 NVD 2 Site 57 Howe Sound middle 1974 depth = 245m d i v e r s i t y = 3.22 valves/gram dry weight = 6.1X10 7 THP 41 SKC 66 CHR 6 TNZ 24 CHM 17 CYS 4 PAR 10 DIT 7 CHC 7 COT 1 CHD 2 CHY 2 MLA 1 AHD 2 CCO 2 SYR 1 GOS 1 GOO 2 CHA 1 NZS 2 DIA 2 STM 1 COL 1 CYO 1 Site 58 Howe Sound mouth 1974 depth = 250m d i v e r s i t y = 3.35 valves/gram dry weight = 6.1X10 7 THP 28 SKC 68 CHR 27 TNZ 25 THN 1 PAR 1 1 CYO 1 NZO 3 CYS 5 GOO 3 CHD 1 CHM 21 CHC 6 239 COT 1 CSM 2 DIT 5 FRI 1 FRC 1 CCO 4 RHZ 3 FRP 1 STM 1 SYU 1 ACS 1 THE 1 Site 59 Saanich Inlet head depth = 212m d i v e r s i t y = 2.82 valves/gram dry weight = 1.2x10 THP 14 SKC 87 CHR 54 TNZ 1 9 CCO 1 CHM 14 PAR 16 CSM 1 RHZ 4 DIT 6 CHD 2 SYT 3 LED 1 DPT 2 CHC 1 AMR 1 DIS 4 BDA 1 Site 60 Saanich Inlet middle depth = 227m d i v e r s i t y = 2.99 valves/gram dry weight = 1.2X10 8 THN 3 THP 40 SKC 81 CHR 39 TNZ 14 THC 3 CCO 5 PAR 7 CHM 12 THE 1 PLS 1 RHZ 5 240 CHC 5 DIT 3 SYU 1 CHA 2 CYO 1 SYT 3 CSC 1 FRP 1 Site 61 Saanich Inlet middle depth = 187m d i v e r s i t y = 3.40 valves/gram dry weight = 5.0X10 7 THP 50 SKC 44 CHR 28 TNZ 23 THC 1 CHC 21 DIT 10 PAR 1 1 COL 1 CCO 3 NZR 1 NZA 1 FRP 2 SYT 5 RHZ 1 NZX 2 CHA 1 DIS 1 STM 1 CSM 1 CHD 2 BDA 1 CYS 1 CDS 1 OPR 1 Site 62 Saanich Inlet mouth depth = 62m d i v e r s i t y = 3.46 valves/gram dry weight = 2.2X10 7 THP 71 SKC 19 CHR 17 TNZ 24 THC 4 THN 4 CHM 32 PAR 14 AHH 1 241 SYT 2 CCO 7 CHD 3 FRP 2 LED 2 DIS 1 CHC 3 CYS 1 CHA 2 OPR 1 THE 3 DIT 3 NZA 1 RHZ 2 CDS 1 CYM 1 NZD 2 COL 2 Site 63 Saanich Inlet middle depth = 223m d i v e r s i t y = 3.07 valves/gram dry weight = 8.0X10 7 THP 44 SKC 64 CHR 41 TNZ 1 7 THC 1 CHC 28 PLS 1 DIT 4 PAR 1 1 NZS 1 RHZ 3 OPR 1 SYT 2 STM 1 CCO 3 CHD 2 COL 1 AHH 2 GRM 1 Site 64 Jer v i s Inlet middle 1981 depth = 540m d i v e r s i t y = 3.34 valves/gram dry weight = 4.5X10 7 THP 21 SKC 54 CHR 69 TNZ 16 CHC 26 242 THN 7 CHA 1 1 CYS 1 CHY 2 CHD 6 DPS 1 DIT 5 PAR 1 ACS 1 NVA 2 RHZ 2 THE 3 DIS 5 NZS 1 AHH 2 EUD 1 SYT 1 ACP 1 CDS 1 Site 65 Jervis Inlet middle 1981 depth = 550m d i v e r s i t y = 3.27 valves/gram dry weight = 6.5X10 7 THP 24 SKC 68 CHR 48 TNZ 1 3 THN 5 CHM 10 CHA 24 CHD 6 CHC 4 DIS 8 DIT 1 LED 2 COR 1 FRP 1 CYS 3 COL 1 ACP 1 TBF 1 CHL 1 THE 3 RHZ 3 ACS 1 THG 1 AMO 1 Site 66 Jervis Inlet head 1981 depth = 330m d i v e r s i t y = 3.32 243 valves/gram dry weight = 4.8x107 THP 27 SKC 57 CHR 56 TNZ 13 THN 8 CHM 9 CHD 5 CHA 17 CHV 3 RHZ 5 DIS 4 CHC 2 DIT 2 CYS 5 MLT 1 CYO 1 THG 1 ACS 1 CHL 3 COL 1 CSM 3 ACP 1 FRP 1 Site 67 Je r v i s Inlet head 1981 depth = 285m d i v e r s i t y = 3.88 valves/gram dry weight = 5.6X10 7 THP 22 SKC 54 CHR 48 TNZ 8 THC 2 THN 1 1 CHM 10 DIS 6 AMR 6 CHD 3 CDS 1 SYT 1 CHA 7 PAR 2 CHC 5 FRP 1 THE 2 CHL 1 COT 1 CYS 6 CCO 2 ACP 2 NVY 1 DIT 5 244 DIF 1 THG 2 MLG 1 GOO 4 CHE 5 LED 2 CSM 2 COS 3 COA 1 RHZ 1 Site 68 Jervis Inlet mouth 1981 depth = 677m d i v e r s i t y = 3.89 valves/gram dry weight = 5.9X10 7 THP 2 3 SKC 53 CHR 51 TNZ 18 THC 1 THN 1 CHM 13 DIS 4 COS 3 CHC 10 CCO 5 CHE 2 NVS 2 LED 3 CYS 3 PAR 7 THG 2 CHA 4 CDS 1 DIF 1 MLG 1 DIT 4 THE 4 CHD 4 ACP 1 AHH 2 GOO 2 CSM 2 SYT 3 COA 1 CHZ 1 FRV 1 TBN 1 OPR 2 ACS 1 AMO 1 PIM 1 245 Site 69 Jervis Inlet head 1981 depth = 366m d i v e r s i t y = 3.76 valves/gram dry weight = 2.4X10 7 THC 1 THP 16 SKC 50 CHR 59 TNZ 6 THN 4 CHM 1 1 CYO 4 CHD 7 CHA 8 CYS 6 MLM 1 CHL 1 DIT 3 THE 5 CSM 4 DPS 1 ACS 2 CCO 4 STS 1 DIS 6 TAF 2 RHZ 1 CHC 12 PLS 1 LED 13 ACP 1 TBN 2 DIH 1 HNA 1 NZS 1 Site 70 Jervis Inlet middle 1981 depth = 503m d i v e r s i t y = 3.99 valves/gram dry weight = 3.4X10 7 THP 17 SKC 51 CHR 49 TNZ 1 1 THN 2 CHM 17 PLS 1 DIS 4 THE 3 CHA 9 PAR 2 DIT 9 LED 4 246 TRA 1 CHC 6 SUO 1 NVY 2 ACP 3 CHL 3 RHZ 3 MSE 1 SYT 3 CHD 8 GOS 2 CHV 4 CCO 2 CSP 1 TAF 6 CDS 2 CHY 3 COR 1 FRP 1 CYS 5 THG 1 Site 71 Je r v i s Inlet middle 1981 depth = 567m d i v e r s i t y = 3.85 valves/gram dry weight = 4.7X10 7 THP 30 SKC 40 CHR 42 TNZ 20 THC 1 THN 2 CHM 1 1 DIH 1 DIT 5 LED 1 CCO 1 CHC 7 TAF 1 CHD 7 STS 1 NVA 1 RHZ 5 THE 2 SYT 4 CHY 1 PLS 1 CSM 4 CYS 2 CHA 10 NZO 3 AHH 1 GOS 2 247 DIS 1 PAR 2 CSP 1 HYS 3 THG 1 RCC 2 Site 72 Jer v i s Inlet middle 1981 depth = 677m d i v e r s i t y = 3.90 valves/gram dry weight = 6.9X10 7 THP 33 SKC 35 CHR 39 TNZ 1 6 THC 1 THN 5 CHM 1 4 PAR 4 SYT 1 DIT 9 CHC 5 RHZ 2 OPR 1 COS 3 CHY 3 CHA 6 CHD 7 CHV 1 EUP 1 TAF 5 COL 2 CHL 2 ACP 1 CDS 1 THG 2 LED 4 CYS 2 COP 1 SUO 1 ACS 1 NZA 1 AHC 2 Site 73 Jer v i s Inlet middle 1981 depth = 685m d i v e r s i t y = 3.70 valves/gram dry weight = 7.5X10 7 THP 17 SKC 55 CHR 50 TNZ 16 248 THC 2 THN 4 CHM 23 CHA 14 COL 2 HYS 1 RHZ 5 LED 8 PAR 2 DIT 10 TBN 1 THE 3 COS 2 CHC 4 CHV 1 NZO 2 SYT 2 CAC 1 CHD 2 THG 2 ACP 1 GOS 1 EUP 3 CHY 2 COP 1 CYS 1 NVY 1 Site 74 Hotham Sound mouth depth = 612m d i v e r s i t y = 3.74 valves/gram dry weight = 1.0x10s THP 27 SKC 60 CHR 36 TNZ 1 6 THN 3 CHM 13 ACP 2 PAR 1 1 THD 6 DIT 9 AHH 1 GOO 2 SYT 3 CHC 10 CYS 7 THG 3 LED 6 CHD 3 COL 1 CHY 3 CCO 1 249 CSM 2 CHA 4 CMG 1 DPS 2 RHZ 2 CDS 1 Site 75 Hotham Sound middle depth = 556m d i v e r s i t y = 3.48 valves/gram dry weight = 6.7X10 7 THP 20 SKC 70 CHR 33 TNZ 4 THN 2 CHM 8 THC 1 DIT 9 MLM 1 CYS 4 CHD 5 CHA 10 ACS 1 COS 1 LED 5 THD 2 PAR 4 THE 1 NZO 1 SYT 5 DIS 5 CHC 4 DPS 1 ACP 3 RHZ 1 CCO 1 OPR 1 THG 1 Site 76 Hotham Sound head depth = 373m d i v e r s i t y = 3.99 valves/gram dry weight = 6.0x107 THP 21 SKC 24 CHR 43 TNZ 13 THC 2 THN 10 CHM 1 1 DIT 10 250 RHZ 3 CHD 3 CHC 12 CHA 20 LED 4 TAF 5 HYS 5 PAR 14 CDS 1 AMT 1 THD 2 NZO 1 CCO 3 COS 3 CYS 3 DIS 1 THG 1 COP 1 CHL 1 CHV 1 Site 77 Agamemnon Channel south depth = 271m d i v e r s i t y = 3.81 valves/gram dry weight = 5.9X10 7 THP 58 SKC 36 CHR 1 7 TNZ 1 0 AHB 1 MLS 2 PAR 1 LED 5 DIT 2 DIS 10 GRM 1 GRC 1 THE 3 CHA 1 9 THD 3 CHC 5 SYT 3 AHH 1 CHD 6 TAF 1 CYS 3 CHE 1 DPS 2 CSC 1 CCO 2 CYO 1 RHZ 3 HYS 3 251 THG 1 CDS 1 NZO 1 ACP 2 ACS 1 COP 1 CHY 3 Site 78 Agamemnon Channel north depth = 194m d i v e r s i t y = 3.70 valves/gram dry weight = 1.7X10 7 THP 40 SKC 2 3 CHR 34 TNZ 26 THN 4 CHM 12 PAR 1 THC 1 COP 1 CHC 7 BDL 1 CHD 7 OPM 2 FRY 7 CYS 7 DIT 4 CHA 6 LED 1 1 COD 1 DIS 2 TXL 1 1 RHZ 1 THE 2 CHE 2 COT 1 ACP 2 SYT 2 CHY 2 CSC 3 CDS 2 BDA 1 TAF 3 NZA 1 Site 79 Malaspina S t r a i t depth = 271m d i v e r s i t y = 3.97 valves/gram dry weight = 7.2x017 THP 37 SKC 36 252 CHR 24 TNZ 27 THC 2 THN 2 CHM 1 1 DIM 1 PAR 15 THE 1 COR 2 CHD 3 NZO 1 BDA 1 CYS 3 TAF 4 NVS 1 DIT 4 CHC 8 CHA 3 COP 2 RHZ 2 OPM 1 DIS 2 SYT 1 CCO 2 CDS 2 CSC 3 ACP 1 CHE 2 CHY 4 EPT 1 GOS 2 THD 1 ACS 1 Site 80 S u t i l Channel depth = 256m d i v e r s i t y = 4.14 valves/gram dry weight = 1.2x10s THP 26 SKC 13 CHR 52 TNZ 18 THC 4 THN 7 CHM 12 CHC 22 CHD 4 SCC 1 NZA 1 NVS' 1 NZN 1 RHZ 10 PAR 15 253 GRC 3 SYT 2 TAF 4 LED 5 AHH 1 ACS 1 DIS 4 NZO 1 BDA 1 DIT 6 PLS 1 CSM 3 TRA 1 CDS 7 STM 1 CCO 3 AMR 1 THE 3 OPR 1 AHB 1 THG 1 Site 81 Desolation Sound depth = 512m d i v e r s i t y = 4.22 valves/gram dry weight = 5.8X10 7 THP 1 4 SKC 7 CHR 54 TNZ 24 THC 2 THN 6 CHM 14 AMR 2 GRC 1 DIT 6 CCO 4 CHC 1 1 TAF 4 RHZ 4 SYT 7 PAR 8 THG 4 THE 1 THD 4 CDS 3 CSM 4 NZA 1 BDA 1 CYS 2 DPS 1 COL 2 CHA 3 •254 BDL 1 TRA 1 CHD 7 OPM 1 CSC 3 LED 4 DIS 1 ACP 1 OPP 1 Site 82 Pryce Channel depth = 497m d i v e r s i t y = 3.74 valves/gram dry weight = 1.1xl0 8 THP 24 SKC 2 5 CHR 60 TNZ 29 THC 4 THN 4 CHM 19 CHD 9 THE 3 DIT 4 CSC 2 CHE 3 PAR 7 COT 2 LED 1 CHC 6 CDS 2 CCO 3 SYT 3 TAF 3 DIS 1 NZG 1 BDL 1 NZO 1 THG 2 RHZ 3 RAS 1 DMM 1 DPS 2 CSP 1 Site 83 Homfray Channel south depth = 706m d i v e r s i t y = 4.10 valves/gram dry weight = 7.7X10 7 THP 13 SKC 19 CHR 55 255 TNZ 1 6 THC 1 THN 1 1 CHM 18 CSP 2 CHA 6 LED 5 DIT 9 CHE 4 GOS 2 AMP 1 CSC 1 PAR 7 RHZ 6 CHD 13 CHC 6 DIS 3 BDA 1 THG 1 THD 2 COP 1 TAF 3 CCO 3 THE 3 CDS 2 DPS 1 SYT 3 EUP 1 NVS 1 NZA 1 Site 84 Homfray Channel north depth = 671m d i v e r s i t y = 3.90 valves/gram dry weight = 7.5X10 6 THP 39 SKC 20 CHR 39 TNZ 1 9 THC 2 CHM 21 THN 5 PAR 4 RHZ 7 CHC 1 1 DIT 6 THG 2 DIS 6 THD 1 NVS 1 CHD 6 CHZ 2 NVY 1 256 CHL 2 TAF 6 CCO 1 GRO 1 CHE 1 CSC 2 AME 2 LED 5 SYT 2 NVT 1 EBT 2 Site 85 Toba Inlet mouth depth = 501m d i v e r s i t y = 3.92 valves/gram dry weight = 5.0X10 7 THP 49 SKC 28 CHR 35 TNZ 17 THC 2 THN 4 CHM 15 DIT 2 HNA 1 THG 4 STS 1 LED 4 COL 1 CSC 1 AHH 2 CCO 4 SYT 6 TAF 4 EUP 1 THE 3 PAR 7 CHC 2 DIS 6 ACS 2 CHA 3 COS 1 DPS 2 CSB 1 OPS 1 CHD 3 OPR 1 NZA 1 CHV 1 AME 1 RHZ 1 257 Site 86 Toba Inlet middle depth = 475m d i v e r s i t y = 3.56 valves/gram dry weight = 3.6X10 7 THP 49 SKC 38 CHR 38 TNZ 18 THC 2 THN 9 CHM 19 DIS 4 CCO 3 RHZ 6 CHC 6 CHD 3 CSC 1 EUP 3 PAR 5 NVS 2 THG 2 TAF 3 COC 1 CHV 1 CHA 2 TBF 1 SYT 3 AHH 1 HYR 1 ACP 1 Site 87 Toba Inlet middle depth = 405m d i v e r s i t y = 3.43 valves/gram dry weight = 2.8X10 7 THP 68 SKC 42 CHR 22 TNZ 9 THC 3 THN 12 CHM 1 5 DIT 3 DIS 5 CHC 2 TAF 2 TXL 3 TBF 1 MEC 2 COC 2 NVS 2 CSE 2 THG 2 258 HNA 1 CAC 3 CHD 4 EBT 1 NVD 1 CSC 1 AHH 1 COS 1 EUP 1 FRV 2 CCO 1 Site 88 Toba Inlet head depth = 238m d i v e r s i t y = 4.01 valves/gram dry weight = 1.3X107 THP 61 SKC 38 CHR 14 TNZ 5 THC 2 THN 13 CHM 8 TBF 3 NVY 1 CCO 1 BDA 1 SYT 2 EUP 4 CBN 1 ATS 1 NZP 2 AHC 3 BAT 12 PAR 1 DIT 1 NVA 1 CHY 9 ACP 3 NZO 5 SYF 4 NVT 3 DIM 3 MLM 3 AHH 4 NVK 1 NVF 3 PIC 2 AHY 2 THD 3 NVJ 3 EUL 2 GOS 1 259 Site 89 Bute Inlet middle depth = 630m d i v e r s i t y = 3.26 valves/gram dry weight = 1.9X10 7 THP 7 3 SKC 52 CHR 16 TNZ 29 THC 1 THN 4 CHM 7 AMS 1 DIS 8 PAR 1 DIM 1 NZW 5 EUP 2 NZO 2 THE 6 CHY 6 AHH 3 CHA 3 TXL 1 AHC 1 SYT 1 TAF 1 CCO 1 CHD 3 THD 1 BDL 2 DIT 1 Site 90 Bute Inlet middle depth = 640m d i v e r s i t y = 3.73 valves/gram dry weight = 6.0X10 7 THP 40 SKC 37 CHR 38 TNZ 29 THC 2 THN 2 CHM 16 CHA 2 DIS 21 THE 4 OPS 1 LED 7 CHD 6 TXL 4 RHZ 3 PAR 5 CCO 5 260 BDA 3 CHE 1 THG 1 SYF 1 ACS 1 DIT 2 TAF 2 CHC 1 GOS 1 AHH 2 COL 1 CHY 1 Site 91 Bute Inlet mouth depth = 644m d i v e r s i t y = 3.79 valves/gram dry weight = 7.7X10 7 THP 56 SKC 17 CHR 34 TNZ 16 THC 3 THN 5 CHM 8 AHC 2 PAR 1 3 CHY 5 CON 1 DIS 13 TXL 4 CHE 4 THE 6 NZW 1 SYU 1 LED 3 CHC 3 SYT 2 NZG 2 FRV 1 CHD 8 DIT 1 CCO 1 COC 1 COL 1 CHA 1 CSC 1 AHK 1 Site 92 Burrard Inlet west 1981 depth'= 60m d i v e r s i t y = 3.71 valves/gram dry weight = 4.2x107 261 THP 30 SKC 89 CHR 21 TNZ 17 THC 1 THN 4 CHM 6 THE 2 RHZ 2 CYO 3 SUO 1 PAR 2 NZD 3 SYT 2 MEC 2 CYS 1 OPR 2 TBN 6 NVG 1 NVK 1 LED 2 AHH 1 CHD 1 CHA 2 CHL 2 MLT 1 THG 2 AHC 1 MLG 1 NZU 4 GOO 7 SYU 2 FRL 3 DIT 3 CDS 4 CSM 1 FRP 1 NZW 3 NZO 2 CHC 1 Site 93 Indian Arm mouth 1981 depth = 225m d i v e r s i t y = 2.58 valves/gram dry weight = 2.7xl0 8 THP 33 SKC 134 CHR 13 TNZ 7 THC 3 THN 5 CHM 6 DIS 11 262 PAR 3 CSM 3 CCO 4 SYT 1 CYS 1 RHZ 1 DIT 1 SYU 1 CHE 2 EUP 1 CHD 1 CSC 1 AHH 1 OPP 1 MLM 4 NVD 1 TBN 1 Site 94 Indian Arm head 1981 depth = 212m d i v e r s i t y =3.11 valves/gram dry weight = 2.3x10 THP 51 SKC 92 CHR 10 TNZ 1 3 THN 1 CHM 9 THE 1 DIS 8 CHC 1 CYS 5 THG 1 CSC 2 EUP 1 CSM 3 NZO 1 NZW 1 NZS 4 CHD 4 DIT 2 MLM 4 SYT 6 CCO 1 OPP 1 BDA 1 PAR 1 AHC 1 LED 1 STM 1 SYR 1 NVA 1 263 Site 95 Hidden Basin depth = 6m d i v e r s i t y = 4.01 valves/gram dry weight = 2.1x10 THP 22 SKC 45 CHR 25 TNZ 3 THC 1 THN 7 CHM 15 CHE 5 AHC 1 CDS 1 PLL 1 CHD 7 CCO 9 CHC 1 1 MLM 1 FRP 1 LED 10 THG 1 NZO 1 GRC 1 SYT 8 NZG 1 AME 3 OPM 6 COO 2 CHA 8 RHZ 1 ROM 1 COS 1 PAR 6 TXL 1 PLS 2 DIS 2 CHV 3 CHL 2 CMG 1 NVS 1 CSC 2 TRA 1 CHY 2 RCC 2 COP 2 264 APPENDIX F - PLATES Scale = 10 nm Plate J_ Figs. 1-2. Skeletonema costatum. F i g . 3. Thalassionema nitzschioides. Figs. 4-5. Paralia sulcata. F i g . 6. Ditylum brightwelli i . F i g . 7. Rhizosolenia sp. Plate2 Resting spores. Figs. 1-2. Chaetoceros vanheurcki i . F i g . 3. C. diadema. F i g . 4. C. lorenzianus. F i g . 5. Chaetoceros a f f i n i s . F i g . 6. C. lauderi. F i g . 7. C. radicans. F i g . 8. C. d e b i l i s . F i g . 9. C. didymus. Plate3 Figs. 1-2. Thalassiosira a e s t i v a l i s f. p a c i f i c a . Figs. 3-4. T. nordenskioeldi i . F i g . 5. Dictyocha speculum. F i g . 6. C y c l o t e l l a s t r i a t a . 266 P L A T E 2 2 6 7 

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