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Study of dinoflagellate cysts from recent marine sediments of British Columbia Dobell, Patricia Elda Rose 1978

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STUDY OF DINOFLAGELLATE CYSTS FROM RECENT MARINE SEDIMENTS OF BRITISH COLUMBIA by PATRICIA ELDA ROSE DOBELL B.Sc, University of British Columbia, 1976 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF . MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES Department of Biology We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA October, 1978 © P a t r i c i a E lda Rose D o b e l l , 1978 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of Brit ish Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of (5 I'OIOCJ <j The University of Brit ish Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date Qcrobe/v 10 ltln2 / i i ABSTRACT Viable cysts collected from natural sediments were induced to excyst. Ten cyst-theca relationships, f i r s t established elsewhere, were confirmed for British Columbia (B.C.). These were: Gonyaulax  tamarensis, Protoperidinium aspidotum, P. claudicans, P. conicoides, P. conicum, P_^_ cf. denticulatum, P. leonis, P. oblongum, and Pj^ punctu- latum. Five' cyst-theca relationships were established for the f i r s t time: Peridiniopsis cf. hainanensis, Protoperidinium sp. nov., P.  thorianum, and two apparently new species of Gonyaulax. P. pentagonum was found to have a cyst different from the cyst of this species in the Atlantic. Forty-five samples from Recent sediments were collected along the coast of B.C. Twenty-three of the samples had very few cysts. Hidden Basin was the chief source of viable cysts for the excystment experiments. Ten cyst-based taxa were described from the sediment samples. These were: Operculodinium centrocarpum, the cyst of Scrippsiella  faeroense (= Micrhystridium bifurcatum), Spiniferites belerius, S.  bentori, S. bulloideus, S. elongatus, S. membranaceus, S. nodosum, and S. ramosus. Tanyosphaeridium sp. has been recorded previously as the cyst of Polykrikos schwarzi. Two new cyst-based taxa are described for the f i r s t time. These are a cyst of Protoperidinium sp., and Spiniferites "sp. A". / i i i Cyst assemblages in the Recent sediments of B.C. were similar to many temperate estuarine and neri t i c areas. Some cysts which are characteristic of these areas in other regions, have not yet been found i n B.C. The relative importance of some cysts also varies from that found in similar sediments elsewhere. The dominance of Operculodinium centrocarpum in many of the cyst assemblages, including B.C., is a pattern typical of temperate estuarine conditions. Some cysts appear to be characteristically associated with fjord environments. • Scrippsiella faeroense, for example, has been found in Norwegian fjords and Scottish sea lochs as well as some B.C. fjords and inlets. / i v TABLE OF CONTENTS Page Tit l e Page i Abstract i i . Li s t of Figures & Plates v i Explanation of Figures v i Explanation of Plates v i i List of Tables xv Acknowledgements x v i i Part I-Introduction 1 A - Cysts and Thecae 6 1 B - Sediment Analysis 5 C - Cysts and Life Cycles 6 Part II - Materials and Methods 9 A - Field methods 9 B - Laboratory Methods 9 1 T Palynological methods 9 2 - Method for absolute counts using exotic pollen 12 3 - Procedure for hatching viable cysts 13 4 - Photography 14 5 - Sand and Carbon analysis 14 6 - Statistics 19 ./v /v Part III - Results 19 A - Sediment analysis 19 1 - Tables 20 2 - Correlations 32 B - Excystment Results 36 C - Cysts in Sediments 78 Part IV - Discussion 105 A - The role of cysts in l i f e cycles 105 B - The archeopyle 108 C - The cyst-theca relationship 113 D - The distribution of cysts in sediments 116 E - The Spiniferites/Operculodinium ratio 131 Part V - Summary and Conclusions 133 A - Excystment experiments 133 B - Cysts in sediments 134 Plates and Explanations 137 Bibliography 166 .. . . / v i / v i LIST OF FIGURES AND PLATES  EXPLANATION OF FIGURES Figure 1 - I l l u s t r a t i o n of suction sampler used to obtain samples from Hidden Basin (#1) and Theodosia I n l e t (#2). Figure 2 - Map of the P a c i f i c Northwest. Figure 3 - Enlarged from Figure 2; northernmost s t a t i o n s are i n d i -cated. Figure 4 - Enlarged from Figure 2; stati o n s from Bute I n l e t and the northern Island channels are ind i c a t e d . Figure 5 - Enlarged from Figure 2; southernmost sta t i o n s are i n d i -cated. Figure 6 - Scatter diagram of cysts/gm vs. % carbon. Figure 7 - Scatter diagram of the % carbon vs. the % sand. Figure 8 - Cluster diagram of body length vs. body width of S p i n i - f e r i t e s ramosus cysts from various sample s i t e s of t h i s coast. Figure 9 - Intercalary plates of various Protoperidinium spp. are shown (s t i p p l e d ) ; o u t l i n e of cyst archeopyle shape of each species i s superimposed. / v i i EXPLANATION OF PLATES Page Plate 1 -Figure 1-4 P e r i d i n i o p s i s c f . hainanensis; 1. v e n t r a l view; 2. dorsal view; 3. a p i c a l view; 4. a n t a p i c a l view. Figure 5 Protoperidinium claudicans, dorsal view. Figure 6-8 P. conicum; 6. aberrant cyst; 7. dorsal view of theca; 8. v e n t r a l view of theca. Figure 9-11 P. aspidotum; 9. ventral'view; 10. dorsal view; 11. a p i c a l view. Figure 12,14,15 P^ c f . denticulatum; 12. ve n t r a l view; 14. antapical view; 15. a p i c a l view. Figure 13 P_^_ l e o n i s , v e n t r a l view. Figure 16 P^ oblongum, v e n t r a l view. Figure 17-19 P_^  thorianum; 17. v e n t r a l view; 18. a p i c a l view; 19. the cyst. Plate II -Figure 1-5 Protoperidinium pentagonurn; 1. v e n t r a l view; ./ v i i i / v i i i 2. dorsal view; 3. the cyst; 4. apical view; 5. antapical view. Figure 6-13 P^.sp.; 6. ventral view; 7. dorsal view; 8. apical view; 9. right oblique view; 10. l e f t side view; 11. oblique dorsal view; 12. the s u l c a l plates; 13. the cyst. Figure 14-17 Zygabikodinium lenticulatum; 14. a p i c a l view; 15. antapical view; 16. ventral view; 17. dorsal view. Plate I I I -Figure 1,2 Protoperidinium thorianum;!. aberrant c e l l ; ventral view; 2. dorsal view. Figure 3-6 Gonyaulax sp.2; 3. ventral view; 4. dorsal view; 5. isolated plate with thecal patterning; 6. apical view. Figure 7-9 P_. punctulatum; 7. ventral view; 8. dorsal view; 9. apical view. Figure 10,11 G_. sp. 1. 10. apical view; 11. ventral view. Plate IV -Figure 1,2 S p i n i f e r i t e s "sp. A"; l i n e drawing; 1. right . . . / i x / i x l a t e r a l view; 2. l e f t l a t e r a l view. Figure 3 Unusual cyst of unknown a f f i n i t i e s . Figure 4,5 nodosum; 4. l e f t l a t e r a l view; 5. r i g h t l a t e r a l view. Plate V -Figure 1,2 Tanyosphaeridium sp. cyst of ?Polykrikos; 1. " r u f f l e " not well developed; 2. " r u f f l e " w e l l developed. Plate VI -Figure 1-3 Protoperidinium claudicans; 1. dorsal view of c e l l ; 2. the cyst; 3. a cyst from palynolo-g i c a l l y treated material. Figure 4,5 P_^_ thorianum; 4. v e n t r a l view; 5. do r s a l view. Figure 6-8 P_^  sp.; 6. v e n t r a l view; 7. r i g h t oblique view; 8. r i g h t l a t e r a l view. Figure 9,10 P. punctulatum; 9. ve n t r a l view; 10. v e n t r a l hypotheca view. Figure 11 P. conicoides;-ventral view. Figure 12 P. l e o n i s ; v e n t r a l view. • * • • / x /x Plate VII -Figure 1-3 Zygabikodinium lenticUlatum; 1. a p i c a l view; 2. v e n t r a l view; 3. oblique a n t a p i c a l view. Figure 4 Protoperidinium leonis c y s t . Figure 5 lenticulatum cyst. Figure 6,7,10 P^ conicum; 6. normal cyst; 7. aberrant cyst; 10. c e l l fror.! cyst i n F i g . 7. Figure 8,9 P. oblongum; 8. cyst; 9. cyst of d i f f e r e n t morphology. Figure 11,12 P^ _ c f . denticulatum; 11 cyst; 12. operculum from cyst i n fig."11. Plate VIII -Figure 1,2 Protoperidinium aspidotum; 1. cyst; 2. r i g h t oblique v e n t r a l view of c e l l . Figure 3,4,7 P_^_ c f . denticulatum; 3. epitheca; 4. hypotheca; 7. c e l l p a i r s . Figure 5,6 P. pentagonum; 5. v e n t r a l view; 6. cyst. Figure 8-10 P e r i d i n i o p s i s c f . hainanensis; 8. cyst; 9. / x i / XI a p i c a l view of c e l l ; 10. an t a p i c a l view. Figure 11,12 S p i n i f e r i t e s elongatus; 11. v i a b l e cyst; 12. cyst from p a l y n o l o g i c a l l y treated material. Plate IX -Figure 1 Protoperidinium thorianum; aberrant c e l l , oblique v e n t r a l view. , Figure 2,3 Gonyaulax tamarensis; 2. v i a b l e cyst; 3. empty cyst. Figure 4-8 G^ sp. 2; 4. v i a b l e c e l l ; 5. v e n t r a l view; 6. oblique v e n t r a l view; 7. dorsal view; 8. cyst, oblique v e n t r a l view. Figure 9-12 G^ sp. 1; 9. cyst, a p i c a l view; 10. v e n t r a l view of c e l l ; 11. oblique r i g h t l a t e r a l view; 12. v e n t r a l view. Plate X Figure 1,2 S p i n i f e r i t e s ramosus; 1. dorsal view; 2. v e n t r a l view. Figure 3,6 S c r i p p s i e l l a faeroense; 3. cyst, o p t i c a l section; 6. cyst, side view. x n / x i i Figure 4,5 S p i n i f e r i t e s bentori var. truncata; 4. dorsal view; 5. v e n t r a l view. \ Figure 7-9 S_;_ membranaceus; 7. dorsal view; 8. v e n t r a l view; 9. o p t i c a l section. Figure 10 S_^_ b e l e r i u s ; d i s t o r t e d c y s t . Figure 11,12 S. bentori; 11 dorsal view; 12. o p t i c a l section. Plate XI -Figure 1,2,4 S p i n i f e r i t e s sp. A (?Hystrichosphaera c f . dentata Gocht); 1. oblique dorsal view; 2. oblique v e n t r a l view; 4. antapical view. it » Figure 3,6 S^ sp. A; 3. dorsal view of cyst with long processes; 6. ve n t r a l view. Figure 5 Protoperidinium 1cyst of unknown a f f i n i t i e s . II " Figure 7,8 S. sp. A; 7. dorsal view; 8. operculum has f a l l e n i n s i d e the cyst. Figure 9. Tanyosphaeridium sp. (?), v i a b l e . Plate XII -Figure 1 Operculodinium centrocarpum. ./ x i i i / x i i i a II Figure 2,3 S p i r i i f e r i t e s sp. A; 2. cyst with narrow four-sided archeopyle; 3. o p t i c a l section. it " Figure 4,5 S. sp. A; 4. antapical view; 5. a p i c a l view. Figure 6 ii " S. sp. A; o p t i c a l section, processes are unusual. Figure 7 sp. A, operculum i n place. Figure 8 S_^_ sp. A; cyst s i m i l a r to F i g . 7. Plate XIII -Figure 1,2 S p i n i f e r i t e s ramosus; 1. dorsal view; 2. an t a p i c a l view. Figure 3,4 Operculodinium centrocarpum; 3. e n t i r e cyst with archeopyle; 4. enlarged view of processes. |l H Figure 5-7 sp. A; 5. fenestrate bases of process; 6. t r i f u r c a t e process with b i f i d t i p s and fenestrate base; 7. inset, close view of microgranular wall seen i n F i g . 5 and 6. Plate XIV Figure 1,2 Tanyosphaeridium sp.; 1. l a t e r a l view; 2. a p i c a l view. . / x i v / x i v Figure 3, 4 sp.; 3. processes are flattened out; 4. apical view. •/ xv /xv LIST OF TABLES Table 1 Summary of p h y s i c a l and b i o l o g i c a l parameters measured at each sample s i t e . l a Tide- and current-scoured channels and bays l b S t r a i t of Georgia l c Fjord environments Id I n l e t s without s i l l s l e Very sheltered i n l e t s Table II Summary of r e s u l t s of -product-moment c o r r e l a t i o n c o e f f i -c i e n t s . Table III A comparison of cyst morphology of aspidotum (Fukuyo et a l . , 1977) and 0. centrocarpum (Wall & Dale, 1968a). Table IV Cyst assemblages associated with B l environments — "temperate, low to medium s a l i n i t y , seasonally s t r a t i f i e d (Wall et a l . , 1977). Table V Cyst assemblages associated with A2 environments — "mild to cool temperate, low s a l i n i t y , moderately s t r a t i f i e d (Wall e t a l . , 1977). Table VI A comparison of cyst assemblages associated with A2 environ-ments & t h e - S t r a i t of Georgia s i t e s (Table l b ) . ...../ x v i / x v i Table VII Cyst assemblages associated with A l environments "cool-temperate, brackish, s t r a t i f i e d e s tuaries." (Wall et a l . , 1977) . / x v i i / x v i i ACKNOWLEDGEMENTS I am deeply indebted to Dr. F.J.R. Taylor for h i s support, advice, and kindness while t h i s thesis was being c a r r i e d out. Special thanks are due to Dr. Glen Rouse for h i s e f f o r t s and a s s i s -tance on my behalf. I would also l i k e to thank Dr. P.J. Harrison, Dr. J . Berger, and Dr. J . Murray who gave h e l p f u l advice while members of my advisory Committee. I wish to thank a l l my colleagues and friends who gave me invaluable support throughout the duration of t h i s study: Carol Pollock, Bev Hymes, Adrian Smith, J o l i e Mayer, and Rosemary Waters. Some of the samples were c o l l e c t e d by Dave Turpin, B i l l McKay, and P r . Taylor. Ms. Dru S u l l i v a n d i d the scanning electron micrographs. . The R.V. Vector and R.V. Pandora were used i n the c o l l e c t i o n of a l l the samples of t h i s study. This study was supported by a grant from the Imperial O i l Company of Canada. F i n a n c i a l assistance was also provided by National Research Council Grant A-6137 to Dr. F.J.R. Taylor. PART I INTRODUCTION This study consists of two par t s . The f i r s t i s the study of the r e l a t i o n s h i p between d i n o f l a g e l l a t e cysts and the thecate c e l l s which a r i s e from t h e i r germination; the second i s a p a l y n o l o g i c a l survey of sediments from approximately 35 s i t e s on the B r i t i s h Columbia coast between Vancouver Island and the mainland. A. CYSTS AND THECAE Many d i n o f l a g e l l a t e cysts along with p o l l e n , fungal spores, chitinous and . l i g n i f i e d remains survive the strong chemical treatments used i n the routine p a l y n o l o g i c a l preparation of sediments. The occurrence of cysts as f o s s i l s has been known for as long as p o l l e n . I n i t i a l l y i t was not known that many of these f o s s i l cysts had a f f i n i t i e s with d i n o f l a g e l l a t e s . However, some c l o s e l y resemble vegetative c e l l s as they occur today and thus t h e i r a f f i n i t i e s with d i n o f l a g e l l a t e s were never i n doubt. I t was believed for some time that these types of remains were f o s s i l i z e d thecae of vegetative c e l l s and not cysts. Other m i c r o f o s s i l s with uncertain or unknown a f f i n i t i e s were, and s t i l l are, assigned to the group known as " A c r i -tarchs" ( E v i t t , 1963). "Hystrichospheres" ( E v i t t , 1961) are another group of "enigmatic bodies" ( E v i t t & Davidson, 1964) whose a f f i n i t i e s remained uncertain f o r many years. F i n a l l y , i t was discovered that they too had dino-/2 /2 f l a g e l l a t e a f f i n i t i e s but as cysts and not as thecae. Hystricho-spheres and other d i n o f l a g e l l a t e cysts indicate t h e i r a f f i n i t i e s i n the following ways: 1. The excystment aperture, c a l l e d an archeopyle ( E v i t t , 1961), i s angular. I t s shape and p o s i t i o n can be r e l a t e d to a p a r t i c u l a r thecal plate (or serie s of plates) of the thecate c e l l . 2. Spines or other sculpture elements e x h i b i t a pattern which " r e f l e c t s " the tabulation of the vegetative c e l l ( E v i t t , 1961). Even though a c r i t a r c h s survive p a l y n o l o g i c a l treatment as we l l , they do not possess the c h a r a c t e r i s t i c s described above. Also, i f an opening i s present i t i s nondescript. Cysts were f i r s t seen i n natural populations i n the l a t e 1800's (F.J.R. Taylor, pers. comm.). Later, cysts i n laboratory cultures of d i n o f l a g e l l a t e s were f i r s t observed by Braarud (1945). He observed spiny spheres occurring i n a culture of Protoceratium reticulatum Claparede & Lachmann (= Gonyaulax g r i n d l e y i Reinecke). No r d l i (1951) found " r e s t i n g " cysts of Gonyaulax polyedra Stein i n a plankton sample from a Norwegian f j o r d . Some were enclosed i n t y p i c a l thecae making i d e n t i f i c a t i o n c e r t a i n . I d e n t i f i c a t i o n to species was not possible when the cyst alone was present. Later these cysts were found to belong to the f o s s i l species Lingulodinium  machairophorum (Deflandre & Cookson) Wall (Wall, 1967). At the time N o r d l i (1951) thought that these cysts resembled a "dubious genus" ..../3 /3 Trochischia Kutz.(most of which were l a t e r determined to be copepod eggs) . Neither Braarud nor N o r d l i made the connection that the spiny spheres they were seeing i n laboratory cultures and plankton samples also occurred as f o s s i l i z e d bodies i n aquatic sediments. I t remained for E v i t t & Davidson (1964) to do t h i s . The l a t t e r authors found r e s t i n g cysts of Gonyaulax polyedra, G. d i g i t a l i s (Pouchet) Kofoid, and Protoperidinium leonis (Pavillard) Balech i n net samples and were able to r e l a t e them to the thecate stage because many of the cysts were enclosed i n more or less i n t a c t thecae. Both G^ _ polyedra and G.  d i g i t a l i s had spiny cysts. By d i s s o c i a t i n g thecal plates from around these cysts they were able to confirm the theca-cyst r e l a t i o n s h i p previously hypothesized.by E v i t t (1961) ; i . e . , the spines or processes extending from the cyst correspond to p a r t i c u l a r plates of the thecate c e l l . They also found empty cysts with archeopyles i n a C a l i f o r n i a t i d e p o o l . P. leonis was found to have smooth cysts which c l o s e l y resemble thecate c e l l s i n shape. Wall (1965) i s o l a t e d v i a b l e cysts from d e t r i t a l sludge obtained from the bottom of Great Harbour near Woods Hole and from s e t t l i n g trays i n the laboratories where the seawater source was also the l o c a l harbour. He described a number of cysts i n p a l y n o l o g i c a l terms and was able to show that modern d i n o f l a g e l l a t e s form cysts with a f o s s i l h i s t o r y . Culture experiments on vi a b l e cysts showed that the hystrichosphere Hystrichosphaera bentori Rossignol gave r i s e to the /4 /4 thecate c e l l d i g i t a l i s . Thus d i g i t a l i s has been shown to be related to i t s cyst through encystment by Evitt & Davidson (1964) who found the cysts inside intact thecae, and through excystment by Wall (1965) who induced viable cysts to excyst. Wall (1965) hypothesized that the cysts he found had a f f i n i t i e s with two modern families. Hystrichosphaera (0. Wetzel) Deflandre & Cookson and Nematosphaeropsis Deflandre & Cookson were of the Gonyau-lacaceae; Deflandrea Eisenack resembled the Peridiniaceae. Deflandrea i s characterized by having an inner capsule which is absent from cysts of modern Protoperidinium so they are not truly synonymous. . Wall & Dale (1968a), continuing the i n i t i a l success of Wall (1965), were able through incubation experiments with viable cysts to relate over 30 cysts to their thecate stages. This provided defini-tive proof of cyst-theca relationships by a means other than finding cysts with intact thecae around them. Since then additional experi-ments with viable cysts have led to confirmation of other cyst-theca relationships (Wall & Dale, 1969; Wall & Dale, 1970; Wall, 1975; Dale, 1977; Anderson & Wall, 1978). In the f i r s t part of this study, dinoflagellate cysts isolated from unconsolidated plankton debris principally from Hidden Basin on Nelson Island were incubated and cyst-thecae relationships are examined. /5 /5 B. SEDIMENT ANALYSIS The main focus of the second part of t h i s study i s to record the types arid numbers of cysts present and to attempt to r e l a t e these to some ph y s i c a l parameters measured at each s i t e . The presence of modern d i n o f l a g e l l a t e cysts i n marine sediments i s b a s i c a l l y the r e s u l t of two f a c t o r s . The species must be present as part of the phytoplankton i n the water above the sediments for t h e i r cysts to be found i n the sediments. Cysts, once formed, behave as non-motile d i s c r e t e p a r t i c l e s which are subject to the same factors as currents and water mass movements which a f f e c t f i n e s i l t p a r t i c l e s (Dale, 1976). By necessity t h i s study does not deal with the planktonic influence on cyst assemblages i n sediments. Quantitative data on the phyto-plankton present i n most of the locations studied i s simply not a v a i l -able. The only published information a v a i l a b l e i s a record of presence or absence of species (Wailes, 1937). C a t t e l l (1969) provided records of some species although he does give the seasonal occurrence and abundance of many common ones. Moreover, studies which have de a l t with the r e l a t i o n s h i p of cysts to planktonic c e l l s i n the past (Reid, 1975; Wall et a l . (1977) have shown i t to be very complex. Wall et a l . (1977) have done an extensive review of most of the p a l y n o l o g i c a l work on recent sediments up to 1976. Using f a c t o r ana-l y s i s and other s t a t i s t i c a l measures they were able to make some sense /6 of a large mass of data which has been accumulating over the years. V i r t u a l l y a l l the information used i n their analyses was from the North and South A t l a n t i c Ocean and the Mediterranean Sea. The only P a c i f i c samples were from offshore Peru. The present study i s one of several done i n the North P a c i f i c . Sasada e_t a l . (1975) studied sediments from the Fukuyama area of Japan, an area with a history of red tides. They found a number of peridiniacean and gonyaulacacean cysts which could not be incu-bated successfully. Fukuyo ^ t al.(1977) studied the cyst-theca relationship of Pro toperidinium aspidotum (Balech) Balech(misidenti-f ied as _P. minutum (Kofoid) Loeblich) from Japanese waters. Heusser & Balsam (1976) included dinocyst counts i n a study of core tops taken from the continental margin from C a l i f o r n i a to Alaska, including the shelf zone west of Vancouver Island. Rather than try to c l a s s i f y environmental climates and "ecologic species groups" l i k e those of Wall et al.(1977) an attempt was made to apply their findings to the B r i t i s h Columbia coast. They were able to relate groups of dinocyst species to s p e c i f i c types of marine environments. Thus the focus of this part of the study w i l l be to relate environments and species groups found i n the North East P a c i f i c to those found i n the A t l a n t i c regions. C. CYSTS AND LIFE CYCLES Dinoflagellates have l i f e cycles which have features i n common .11 n with many other p r o t i s t groups. The vegetative c e l l i s usually motile, haploid and reproduces asexually by oblique f i s s i o n . Cyst formation i n p r o t i s t s , i n c l u d i n g d i n o f l a g e l l a t e s , has been known for many years. Various functions have been a t t r i b u t e d to cysts i n c l u d i n g resistance to adverse conditions, d i s p e r s a l of the species, reproduction, diges-t i o n , and dormancy ( C o r l i s s & Esser, 1974). C i l i a t e s such as Wood- r u f f i a metabolica form cysts when t h e i r food supply, Paramecium, gets low (Johnson & Evans, 1941). S o i l amoebae such as Acanthamoeba form cysts as a regular part of t h e i r l i f e cycle; optimum conditions for the motile stage can deteriorate quickly i n s o i l s (Sleigh, 1973). Other a l g a l groups such as the Chlorophyta, are known to form cysts some of which are apparently f o s s i l i z a b l e (Boalch & Parke, 1971). Di n o f l a g e l l a t e s form more than one kind of cyst. Occasionally vegetative c e l l s w i l l undergo a process c a l l e d ecdysis;. the theca i s cast o f f and the cytoplasm rounds up- to form a thin-walled "ecdysal" cyst (Dale, 1977; Turpin e t a l . , 1978) (= p e l l i c l e cysts of Anderson & Wall, 1978). These cysts form very quickly when adverse changes i n environment occur; they are temporary and not very sturdy (Anderson & Wall, 1978). In batch cultures these cysts are commonly seen when the cultures become senescent. Although these cysts have not been s i g n i -f i c a n t p a l y n o l o g i c a l l y , there i s some evidence that the material making up the wall can r e s i s t a c i d treatments (Loeblich, 1970). Ecdysal cysts are proximal, ( i . e . , they form close to the inner wall of the theca). I f they do have a f o s s i l record most would probably not be /8 identifiable as dinoflagellates. Hystrichospherid- and theca-like dinoflagellate cysts are different from ecdysal cysts. Their composition is such that they are preserved in palynologically treated samples. The purposes of such cysts in the l i f e cycle of dinoflagellates is not yet established. These cysts tend to form toward the end of some blooms in late summer (Eren, 1969). The stimulus for this type of cyst formation is presently unknown, but "crowding" has been advanced by Eren (1969) as one possibility. He saw cysts of Protoperidinium cineturn fa. westii in natural samples. The cysts, once formed, sink out of the surface waters and eventually accumulate in the sediments. One possibility is that hystrichospherid- and theca-like cysts are zygotic, (i.e., resulting from the fusion of two "haploid" vegetative c e l l s ) . Sexual fusion leading to cyst formation is known for freshwater species such as Peridinium cinctum f. ovoplanum, P.  w i l l e i , and P. gatunense (Pfiester, 1975, 1976, 1977), Gymnodinium  pseudopalustre and Wolozynskia apiculata (Von Stosch, 1973), and Cera- tium spp. (Von Stosch, 1972; Wall & Evitt, 1975). Marine ceratia undergo sexual fusion but do not form cysts (Von Stosch, 1965; Wall & Evitt, 1975). Some marine dinoflagellates which form zygotic cysts are known (Von Stosch, 1972).Helgolandinium subglobosum (Von Stosch, 1969) forms temporary cysts. Gonyaulax tamarensis Lebour also forms zygotic cysts (Turpin, e_t a l . , 1978) . None of these species in which /9 l i f e cycle studies have been c a r r i e d out, form cysts with a known f o s s i l record; most are c e l l u l o s i c i n nature and are not preserved i n sediments. Ceratium cysts have been found i n varved sediments up to seven years o l d (Wall & E v i t t , 1975). PART II MATERIALS AND METHODS A. FIELD METHODS Sediment samples were c o l l e c t e d from various locations i n B r i t i s h Columbia. Fjords, sheltered i n l e t s and bays, channels, and r e l a t i v e l y open waters of the S t r a i t of Georgia were a l l sampled using e i t h e r a Smith-Maclntyre grab sampler or a snapper grab sampler, both of which sample the upper few centimeters of surface sediments. Samples were stored i n p l a s t i c bags at 4°C u n t i l ready for use. In two sheltered bays, Hidden Basin (#1) and Theodosia I n l e t (#15) a s p e c i a l suction sampler was developed (Figure 1) i n order to c o l l e c t the unconsoli-dated plankton debris which could not be c o l l e c t e d by routine s e d i -ment samplers. B. LABORATORY METHODS 1. PALYNOLOGICAL METHODS A l l sediments were processed using a technique which allowed the c a l c u l a t i o n of absolute cyst (or pollen) d e n s i t i e s . Two subsamples were removed at random from a sediment sample. Both were placed i n / IP / i o FIGURE 1 The suction sampler used to obtain samples of d e t r i t a l material i n Hidden Bay (#1) and Theodosia I n l e t (#15) . A - The pi s t o n i s set as when a sample has been c o l l e c t e d . The t r i g g e r i s l i f t e d . B - The pi s t o n i s set primed to c o l l e c t a sample. The t r i g g e r i s shown i n place. / l l graduated t e s t tubes and centrifuged f o r three minutes at 3000 rpm; the volume of sediment i n each tube was then recorded. One t e s t tube sample was a i r dried to obtain the dry weight of the known volume of sediment. The other t e s t tube of sediment was treated using standard p a l y n o l o g i c a l techniques (Barss & Williams, 1973) which varied accord-ing to the type of sediment being processed but generally the proce-dure was as follows: mixed with 10% HCl for one hour mixed with 40% HF overnight screening through a 30 urn mesh screen (the f r a c t i o n passing through the sieve i s discarded) a c e t o l y s i s ( i . e . , a 9:1 s o l u t i o n of a c e t i c anhydride and H2SO^ i s added to the sediments and b o i l e d f o r 1 - 1 0 min.) 40% HCl' (heated) for sediments observed to have a large mineral content heavy l i q u i d separation for sediments with a large sand component ; ZnBr. s o l u t i o n at 3.26 s p e c i f i c gravity i s added to the sample; l i g h t p a r t i c l e s l i k e cysts and po l l e n f l o a t on the surface and sand p a r t i c l e s sink 10% KOH (heated) f o r sediments with l i g n i f i e d component as observed through a microscope. /12 /12 The residue remaining a f t e r the a c i d treatments was treated for two min i n a 50% s o l u t i o n of commercial bleach, stained with sa f r a n i n (or l e f t unstained), and then passed through alcohol washes to be stored i n t e r t i a r y b u t y l alcohol (TBA). 2. METHOD FOR ABSOLUTE COUNTS USING EXOTIC POLLEN (Adapted from R. Hebda' 1977) The exotic p o l l e n used i n a l l s l i d e s was pine p o l l e n (Pinus  contorta) stained with methyl green. Ten m i c r o l i t r e s of a known concentration of t h i s stained exotic p o l l e n i n TBA were added to a drop of s i l i c o n e o i l on a glass s l i d e . The stained residue from the p a l y n o l o g i c a l treatment was topped up to a measured volume of TBA, usually 1 or 2 ml. The residue was suspended and a subsample was qui c k l y taken, usually 25 or 50 u l , and added to the s i l i c o n e o i l s l i d e with the exotic p o l l e n already present. The alcohol was allowed to evaporate leaving the number of cysts and green p o l l e n countable per volume or weight of sediment. A cover s l i p was added and p a r a f f i n used to s e a l the edges and make a permanent s l i d e . The following formula was used for c a l c u l a t i o n of the number of cysts/gm or ml of sediment. A.X . M Y N D = G D = cysts/gm of dry sediment. /13 /13 A = cysts counted on one s l i d e y = e x o t i c p o l l e n counted on the same s l i d e as cysts X = exotic p o l l e n added to one s l i d e M = volume of TBA and residue i n t e s t tube (ul) N = volume of TBA and residue added to s l i d e (ul) G = dry weight of o r i g i n a l sediment With t h i s formula any part of the s l i d e can be scanned u n t i l the desired number of cysts has been counted. Where possible at l e a s t 200 cysts were counted for each sample s i t e . 3. PROCEDURE FOR HATCHING VIABLE CYSTS Unconsolidated sludge was c o l l e c t e d from Hidden Basin ( s i t e #1) using a s p e c i a l l y designed vacuum sampler or by S.C.U.B.A. d i v i n g (sample #1, #15). The sludge was passed through graded sieves (Buckbee-Meers) and the 90-20 um s i z e f r a c t i o n was saved. Any cysts or other paly-nomorphs smaller than 20 um were discarded. Cysts were picked up with a micro-pipette from t h i s material and washed i n f i l t e r e d seawater to clean o f f some of the debris which clung to the cysts. Four cysts were placed i n four separate drops of seawater (each about 0.5 ml) i n small disposable p e t r i dishes. They were incubated at 17°C and i n a 2 14:10 (light:dark) c y c l e . Light i n t e n s i t y was .35 me/m: /sec,. The drops with one cyst i n each were examined d a i l y by means of an inverted microscope. When excystment occurred, i n d i v i d u a l thecae and cysts could be c l e a r l y associated. /14 /14 4. PHOTOGRAPHY Photomicrographs were obtained using e i t h e r a Zeiss inverted microscope with camera attachment, or a Zeiss compound microscope. Film used was Kodak te c h n i c a l panatomic X. Pictures were p r i n t e d on I l f o r d photographic paper #3 or #4. 5. SAND AND CARBON ANALYSIS Carbon analysis was one of several p h y s i c a l parameters measured at each sample s i t e . The values obtained plus other measurements such as sand or clay content are useful to help explain differences seen i n cyst d i s t r i b u t i o n i n various areas. Sand content was measured by washing a weighed, sediment sample through a 62.5 jum mesh s i z e sieve and weighing the f r a c t i o n retained. The f r a c t i o n passing through mesh of 62.5 urn size i s the s i l t and clay p o r t i o n . The amount of carbon i n the sediment was analysed for each sample s i t e using a Leco Carbon Analyzer. E i t h e r 015 or 0.25 gm of dry sediment were measured in t o a small c r u c i b l e . The amount used depended on the amount of carbon the sample was expected to contain. The sample was mixed with approximately 0.5 ml of ir o n and t i n p e l l e t s . These metals provide enough mass f o r the ent i r e sample to be i g n i t e d , completely o x i d i z i n g a l l carbon present to CO^ . The gases given o f f are c o l l e c t e d and passed through concentrated KOH which removes the CQ2. The volume of gas i s then measured with /15 /15 FIGURE 2 Map of the P a c i f i c Northwest. Inset are locations of other maps with sample s i t e s . /17 FIGURE 4 Enlarged from Figure 2; stations from Bute I n l e t and the northern Island channels are in d i c a t e d . ..-./•18 FIGURE 5 /18 Enlarged from Figure 2; southernmost stations arc indicated. • /19 /19 the CO^  missing. This value is multiplied by a correction factor and the weight of the sample to obtain a value for the percentage of carbon originally present in the sample. 6. STATISTICS The product-moment.correlation coefficient, " r " , was calculated to determine i f there was a correlation between the change in the percen-tage of sand with the change in the percentage of carbon in each sample site, and for the change in the percentage of carbon with the absolute cyst numbers (cysts/gm dry weight of sediment). A high value of "r" (or a value of "r" above 0.372) indicates that the two factors (parameters) are highly correlated. PART III  RESULTS A. SEDIMENT ANALYSIS The results of sediment analysis and associated environmental chara-cters are presented in tabular form. The sample sites are divided into five categories, with one table for each category. The stations are numbered in the order in which they were collected over the two-year study period. Each station has a name locating i t s position on the Pacific coast rather than using latitude and longitude figures, and a brief description is given for each sample site. The description i s the basis for placing each site in i t s respective category. The environ-mental description as per Wall et a l . (1977) i s the one which f i t s each sample site as defined by those authors. The depth of each site i s .../20 /20 given i n meters. The percent of sand obtained by s i e v i n g i s given to the nearest percent. The percent of carbon as measured by a Leco Carbon Analyser i s given without any rounding o f f of values. Cysts/gm of dry weight i s given. Numbers over 1000 were rounded o f f to the nearest 100. The dominant cyst species i s given; i t i s the s i n g l e species which i s numerically most abundant. The percent S p i n i f e r i t e s i s the percentage of a l l S p i n i f e r i t e s cysts from each s i t e , whereas the percent Operculodin-ium i s the percentage of CK_ centrocarpum plus a l l i t s morphological v a r i a n t s . Davey & Rogers (1975) found that i n t h e i r study s i t e changes i n these values plus changes i n the Spiniferites/Operculodinium r a t i o could be r e l a t e d to c e r t a i n environmental f a c t o r s . The s i g n i f i c a n c e of the changes i n these values here i s analysed i n the discussion. The r a t i o of Foraminifera/cysts includes only those forams with chitinous inner t e s t s , because the CaCO of the outer s h e l l i s dissolved by the 3 HCl used i n t r e a t i n g the samples. Hence any forams without the inner l i n i n g are dissolved away. These values are not discussed i n the text but have been recorded here for future reference. 1. TABLE SUMMARY OF PHYSICAL AND BIOLOGICAL PARAMETERS MEASURED AT EACH  SAMPLE SITE (Table 1) Environmental Type 1 (Table la) - Tide- & Current-Scoured Channels  and Bays The f i r s t environmental type to be shown i s defined as channels or open bays scoured by currents and t i d e s . The sediments have high gravel /21 121 TABLE 1A TIDE- AND CURRENT-SCOURED CHANNELS AND BAYS Queen C h a r l o t t e S t r a i t Channel I s l a n d S t r a i t Sunderland Channel O k i s a l l a Channel Boundary Bay Owen Bay H e r i o t Bay Kanish Bay c u r r e n t - s c o u r e d channel c u r r e n t - s c o u r e d channel c u r r e n t - s c o u r e d channel c u r r e n t - s c o u r e d channel sandy beach open bay open bay open bay to n) S 5 Lght S iters 5 g c Depth (m % Sand % Carbon Cysts/gra of sediire 120 87 U .46 60 120 57 1.05 110 204 84 0.82 80 120 95 0.69 3 0 92 0.5 0 10 / 0.33 340 22 86 0.63 290 35 66 1.31 265 Spiniferites  ramosus Operculodinium  centrocarpum 0. centrocarpum  O. centrocarpum / Q. centracarpum Q. centracarpum / 15 24 15 / 58 24 30 / / 0.25:1 1:1 0.5:1 temperate estuarine ./22 /22 and sand content. The sand content ( i . e . , the amount of material retained by a 62.5 um mesh s i z e s i e v e ) , varied from 57 - 94% dry weight. Sand from s i t e #17 (Owen Bay) was not measured because the sample was too small. Sample depths varied from 120 - 204 meters for the channels, whereas depths of the open bays ranged from 10 - 35 meters. The dominant cyst species i s 0_^  centrocarpum. The absolute cyst numbers i n a l l these areas i s low being between 0 - 300 cysts/gm. The number of cysts that could not be i d e n t i f i e d i s high i n a l l these l o c a -tions . This appears mainly the r e s u l t of the action of coarse sand p a r t i c l e s on the palynomorphs, many of which are broken or t a t t e r e d beyond recognition. The percentage of carbon i s l e s s than 1% except f o r Channel Island S t r a i t (#16) and Kanish Bay (#42) . However these two areas also had les s sand than the others. Environment Type 2 (Table lb) - S t r a i t of Georgia Environmental type 2 consists of the open regions of the S t r a i t of Georgia. I t i s somewhat s i m i l a r to the environmental regime desig-nated B l by Wall e t a l . (1977) ( i . e . , coastal-temperate, low to medium s a l i n i t y , seasonally s t r a t i f i e d ) , e . g . the Middle A t l a n t i c Bight region. However the S t r a i t of Georgia d i f f e r s from t h i s region i n that i t i s not true continental s h e l f because of the presence of Vancouver Island to the west. I t i s permanently s t r a t i f i e d due to r i v e r outflow and thus also has some resemblance to A2 environments ( i . e . temperate-./23 /23 TABLE l b STRAIT OF GEORGIA Station number Station Name Location Description Environmental designa-tion of Kali et al.,1977 Depth (meters) % Sand | % Carbon Cysts/gram dry weight of sediment Dominant Species in 0) 4J u V c a to % Operculodinium +J (0 cr c o <u »l «* Ratio Spiniferites/ Operculodinium Ratio microforams/ total cysts 2 south o f Texada I s l a n d p a r t o f S t r a i t the A2«-B1 400 2 2.01 1000 S p i n i f e r i t e s 40 33 5 1.2: 1 .05:1 ramosus 3 Gambier Bay p a r t o f the A2-B1 146 1 2 .20 4500 S. ramosus 60 9 4 6.6: 1 .06: 1 S t r a i t 6 n o r t h o f Texada I s l a n d open t o S t r a i t the A2-B1 400 1 3.55 4200 S. ramosus 44 27 3 1.6: 1 .02:1 21 Howe S o u n d - C o l l -ingwood Channel open to S t r a i t the A2-B1 65 4 1.91 1500 S. ramosus 58 20 5 3:1 0.2:1 44 behind Hernando I s l a n d p a r t o f S t r a i t the A2-B1 260 8 3.59 10,400 S. ramosus 35 31 3 1.1: 1 0 . 3:1 * temperate e s t u a r i n e to temperate c o a s t a l ./24 / 2 4 estuarine regions) . The depth of these sample s i t e s ranges from 6 5 - 4 0 0 meters with the sand content being low ( 1 - 8 % ) . Carbon content i s 2 . 0 1 - 5 . 2 % . Absolute cyst numbers are a l l moderately high, ranging from 1000 -1 0 , 5 0 0 cysts/gm. S a l i n i t i e s are generally 2 6 % o a t the surface to 3 1 % o on the bottom. In these areas there i s not a marked seasonal f l u c t u a -t i o n i n these values. The sample s i t e north of Texada Island ( # 6 ) has a s l i g h t l y higher s a l i n i t y of 2 9 % « . Although Gambier Bay ( # 3 ) and Collingwood Channel ( # 2 1 ) are part of Howe Sound they appear to be more influenced by the S t r a i t water than the f j o r d conditions of the Sound. The dominant species i n a l l the sample s i t e s i s S p i n i f e r i t e s  ramosus (Ehrenberg) Loeblich S Loeblich; sometimes i t comprises over h a l f the cysts. This i s i n contrast to most other sample s i t e s where 0. centrocarpum i s dominant. Environment type 3 (Table Ic) - Fjords Fjords are c h a r a c t e r i s t i c of t h i s environmental type. The s i t e s summarised here are s i m i l a r to regime A l (Wall et a l . , 1 9 7 7 ) , i . e . cool-temperate, brackish water, s t r a t i f i e d , often anoxic on the bottom. Bute, Knight, Muchaletand J e r v i s Inlets (figure 2-5) are a l l c l a s s i f i e d as f j o r d s , being long, deepest at the entrance, and with a shallow s i l l at the entrance r e s t r i c t i n g water movement between the o / 2 5 /25 TABLE l c FJORD ENVIRONMENTS Station Number Station Name Location Description Environmental Designa-tion of Wall et al.,1977 Depth (meters) % Sand % Carbon ! Cysts/gram dry weight of sediment 1 Dominant Species : % Spiniferites % Operculodinium 3 V rr c o w Ratio Spiniferites/ \ Operculodinium ! Ratio microforams/ j total cysts 7 Bute Inlet - fjord Al* 700 2 2.15 600 0. centrocarpum 20 49 0 0 4 1 .16:1 entrance 8 Bute Inlet - - fjord Al 275 1.5 0.85 0 / / / / / / head 9 Bute Inlet - fjord Al 600 26 0.93 0 / / / / / / r.iddle 11 Knight Inlet - fjord Al 245 64 2.11 450 0^ centrocarpum 10 22 8 0 .5 :1 1.3:1 entrance 12 Knight Inlet - fjord Al 360 7 2.42 170 0. centrocarpum 12 42 0 0 .25:1 .23:1 middle 13 Knight Inlet - fjord Al 260 51 0.17 0 / / / / / / head 20 Howe Sound - in fjord Al 274 4 0.85 160 0. centrocarpum 15 59 4 0.25:1 0.3:1 fjord portion 22 Howe Sound fjord Al 200 10 2.85 330 centrocarpum 26 30 5 111 0.3il 23 Howe Sound - fjord Al 283 2 1.14 30 / / / / / / head 24 Muchalet Inlet fjord Al 375 11 / 400 0. centrocarpum 17 24 0 0.7:1 .07:1 19 Howe Sound . fjord Al 164 1 0.93 50 0_ centrocarpum 12 59 6 0.2:1 0.5:1 46 Jervis Inlet - fjord Al 325 32 1.59 4050 0 centrocarpum 35 37 2 1: 1 .02:1 entrance •temperate fjords ./26 /26 LIST OF TRENDS IN PHYSICAL PARAMETERS OF FJORDS Physical parameter Range of Values at the; Head Middle Entrance Sand content 2 - 51% 4 - 26% 1 - 64% Carbon Content 0.17 - 1.14% 0.85 - 2.42% 0.93 - 2.15% Cyst numbers 0 - 27/gm 0 - 170/gm 50 - 4050/gm ./27 /27 f j o r d and the outside waters. Howe Sound (Figure 5) i s a combined fjord-embayment (Stockner et a l . , 1977) . Samples #20, #22, #23 are i n the f j o r d portion; sample #19 i s i n the embayment portion but i s s t i l l influenced by the r i v e r runoff at the head. Sand content i s highly v a r i a b l e from the head to the entrance .of a l l these f j o r d s but with no d e f i n i t e pattern of increase or decrease. Q u a l i t a t i v e l y the samples at the heads of the fjords (#8, #13, #23) contained large amounts of c l a y . Carbon content showed a s l i g h t ten-dency to increase towards the entrance. The trend f o r increasing cyst numbers towards the entrance i s more obvious than the trends for carbon (Table I I ) . The dominant cyst species i n a l l cases i s O.  centrocarpum. The s a l i n i t y of the surface waters of these fjords i s always low e s p e c i a l l y i n spring. Below f i v e meters depth the s a l i n i t y varies from 25 - 31 %<>with no h a l o c l i n e . Environment type 4 (Table Id) - Inlets without S i l l s Table Id summarises the values f o r i n l e t s without s i l l s . The length of these i n l e t s varies from 58 km (Indian Arm) to 8 km (Forward Harbour). Environmental regime A2 (Wall et a l . , 1977) i s s i m i l a r , i . e . cool-temperate, low s a l i n i t y , moderately s t r a t i f i e d to v e r t i c a l l y homogeneous /28 /28 TABLE Id INLETS WITHOUT SILLS Station Number Station Name Location Description Environmental designation of Wall et al., 1977 Depth (meters) •0 s * % Carbon Cysts/gram dry weight of sediii.ent Dominant species j % Spiniferites a •H T> 0 *3 u u <y c-ol s« % S. elongatus Ratio Spiniferites/ Operculodinium \ Ratio microforams/total cysts 4 Indian Arm - head sheltered i n l e t A2 * 106 22 4.48 3700 0. centrocarpum 8 65 4 0.1:1 L 1 J : 1 5 Bur.tzen Bay in Indian Arm sheltered i n l e t A2 230 19 1.45 6500 0. centrocarpum 2 86 1 .04:1 01:1 31 Pendrell Sound -head in l e t A2 80 32 1.44 18,300 0. centrocarpum 29 30 18 1:1 .05:1 32 Pendrell Sound -entrance i n l e t A2 410 7 4.45 21,50'. 0. centrocarpum 36 27 12 1.3:1 .008:1 33 Simoon Sound — Inlet A2 13 22 8.12 830 0. centrocarpum 25 32 2 0.7sl .4:1 head 34 Simoon Sound -entrance in l e t A2 54 17 8.24 97,000 0. centrocarpum 6 81 0.5 0.07:1 J . l i l 36 Port Elizabeth -head in l e t A2 38 81 CO 150 Protoperidinium SPP- 36 4 0 9:1 ).2:1 37 Fort Elizabeth -entrance i n l e t A2 120 2 4.23 45,000 0. centrocarpum 7 61 0.3 0.13:1 .05:1 39 Topaze Harbour sheltered i n l e t A2 27 6 4.08 920 Protoperidinium SPP- 15 14 3 1.1:1 0.1:1 40 43 Forward Harbour Ramsay Arm sheltered i n l e t A2 26 13 2.71 530 0. centrocarpum 25 48 3 0.5:1 / Inlet A2 350 20 7.07 4300 0. centrocarpum 33 47 0.7 0.7:1 / *temperate estuarine /29 /29 Sand content ranges from 2 - 81%. A s l i g h t gradient i s evident with more sand being present at the head than near the entrance (e.g. Pendrell Sound). Carbon content tends to increase towards the entrance as w e l l . Values range from 1.44 - 8.24%. Absolute cyst numbers show a tendency to increase toward the entrance. Pendrell Sound shows t h i s tendency only s l i g h t l y , but i n Simoon Sound and Port E l i z a b e t h t h i s trend i s pronounced. In Indian Arm cyst numbers are lower near the entrance. Absolute cyst numbers vary widely from 145 - 97,000 cysts/gm. Sample #34 (Simoon Sound) has the highest value of a l l the sample s i t e s . This i s v i r t u a l l y a monospecific assemblage (81%) of 0^ centrocarpum. Sample #5 (Buntzen Bay) also contains a high percentage (85%) of t h i s species. 0. centrocarpum i s the dominant species i n a l l these s i t e s except f o r Port E l i z a b e t h (#36) and Topaze Harbour (39) which has Protoperidinium spp. dominant. Protoperidinium spp. includes cysts of a l l species of t h i s genus (e.g. P^ l e o n i s , P. conicum, P. claudicans). S p i n i f e r i t e s cysts comprise 2 - 36% of the assemblages. Sample #31 (Pendrell Sound-head) has the l a r g e s t percentage of S. elongatus cysts at 18%. /30 /30 Environment type 5 (Table le) - Very Sheltered Inlets Both. Hidden Basin and Theodosia I n l e t were sampled several times over the two year study i n t e r v a l . In Hidden Basin the material used f o r the various analyses was unconsolidated planktonic debris rather than sediment. The absolute cyst numbers numbers var i e d each time as d i d the dominant species. Hidden Basin (#1) i s an unusual study s i t e f o r several reasons. I t i s very sheltered. The entrance i s narrow, being about 50 meters wide and obstructed by an i s l a n d which narrows i t even more. I t i s also shallow; at low t i d e the entrance i s impassable by boat. The bay then widens into a t r i a n g u l a r shape and i s about 1/2 km long. Theodosia I n l e t (#15) i s very sheltered, being at one end of another long curving i n l e t ; thus i t i s far removed from the S t r a i t of Georgia. There i s a r i v e r at i t s head. This area has several logging camps along i t s shores and also i t i s used as a booming ground. The bottom of the i n l e t i s covered with sunken logs and large amounts of bark, needles,' leaves and other t e r r e s t r i a l plant debris due both to the logging a c t i v i t i e s and to the r i v e r runoff which c a r r i e s a l o t of t h i s material (and sand) into the i n l e t . This s i t e (as i n Hidden Basin) had a r e l a t i v e l y t h ick mat of accumulated planktonic debris on the bottom although not as great as at Hidden Basin. The highest values for carbon were recorded f o r the d e t r i t a l samples while the one sediment sample was much lower. /31 /31 TABLE l e VERY SHELTERED INLETS 0 Station Number Station Name Location Description Environmental designation Wall et al^, 1977 Depth (meters) % Sand % Carbon Cysts/gram dry weight of sediment Dominant Species % Spiniferites \ Operculodinium CO ±J O C 0 0) »l of Ratio. Spiniferites/ Operculodinium Ratio microforams/total cysts 1. Hidden Basin sheltered bay -entrance onto Strait of Georgia A2* 6 1 7.33 4200 Protoperidinium spp. 27 13 4 2a 03:1: 14 Hidden Basin s .a. s. a. s. a. s .a. 6.6 34,000 Spiniferites ramosus 55 8 2 6.0:1 .04:1 28 Hidden Basin* s. a. s .a. s.a. s .a. 6.39 9 BOO Scrippsiella faeroense 40 14 4 2 i l .02:1 45 Hidden Basin s .a. a .a. s .a. s .a. 7.52 6300 Protoperidinium spp. 39 12 6 3:1 .07:1 15 Theodosia Inlet sheltered i n l e t -at head of long in l e t A2 6 .5 17.05 60 0. e c u L r u c i rpuin 17 50 0 0.3ll ,03:1 29 Theodosia Inlet s .a. s. a. s .a. s .a. 16.5 490 0. centrocarpum 20 58 2 0.3:1 / 30 Theodosia Inlet s .a. s .a. s .a. s .a. 2.72 200 0. centrocarpum 5 76 0 0.07:1 1:5 26 West Gorge Harbour sheltered bay A2 10 86 0.37 70 0. centrocarpum 19 51 6 0.3:1 ).15:1 27 East Gorge Harbour s .a. s .a. 15 14 1.81 2100 0. centrocarpum 30 40 2 0.8:1 ).06:1 * temperate estuarine ./32 /32 In p a l y n o l o g i c a l l y treated material the dominant species was 0.  centrocarpum (see the discussion f o r further comment on t h i s i n l e t ) . Forward Harbour was not included i n t h i s category because i t i s not as sheltered. Strong t i d a l currents are present through the entrance which i s narrow, deep and long. Gorge Harbour (#26, #27) i s an i n l e t with a narrow entrance but i t lacks the shallow s i l l of Hidden Basin; thus planktonic debris does not accumulate here. This harbour i s an anchorage for many boats; there are homes a l l along the shore; at the time the samples were c o l l e c t e d there were innumerable j e l l y f i s h i n the water (none outside the harbour however). I t i s not known what e f f e c t , i f any, these factors would have on the cyst assemblages i n the sediments. That these disturbances would a f f e c t the phytoplankton and i n d i r e c t l y the cyst population i s a p o s s i b i l i t y but the extent or the manner of the change i s unknown. 2. CORRELATIONS Two c o r r e l a t i o n s were c a r r i e d out on the data from sample s i t e s here. These were a c o r r e l a t i o n of changes i n sand content with carbon content of sediments and a c o r r e l a t i o n of changes i n carbon content with absolute cyst numbers. Other co r r e l a t i o n s were not attempted because s t a t i s t i c a l analysis of environmental data was not the main focus of t h i s study and also because only a few environmental para-meters were measured at each sample s i t e . /33 /33 TABLE II SUMMARY OF RESULTS OF PRODUCT-MOMENT CORRELATION COEFFICIENTS 1. % sand vs. % carbon - r = 0.55 df = 40 p = 0.5; r = 0.305 p = 0.01; r = 0.393 2. % carbon vs. cysts/gm - r = 0.47 df = 38 p = 0.05; r = 0.325 p = 0.01; r = 0.418 /34 /34 2BD-T-FIGURE 6 24.0-+-1.0 Cysts per grom(log) 20 Scatter diagram of cysts/gm vs. the % carbon from each sample s i t e . /35 FIGURE 7 240-200-+-16J0 -H 4.0" 20 °/o land (arc ii n) Scatter diagram of the % carbon vs. the % sand from each sample s i t e . /36 In t h i s t e s t " r " can have a value of +1 for p e r f e c t p o s i t i v e c o r r e l a t i o n between the two variables to -1 f o r p e r f e c t negative corre-l a t i o n . (Table II ) Summary of Results of Product-Moment C o r r e l a t i o n C o e f f i c i e n t s The r e s u l t s show that the percentage of sand i s s i g n i f i c a n t l y , negatively correlated with the percentage of carbon. Such a r e s u l t i s not unexpected. Carbon f r a c t i o n s of sediments tend to be part of the f i n e material. When sand content i s high the amount of fine material i s i n e v i t a b l y low (figure 7). Sand i s not always a useful index of the organic ( i . e . , carbon), content of sediments. Some sediments can contain large amounts of clay which i s also inorganic and i n such sediments carbon i s low even though the percentage of sand may be low as w e l l . The " r " value f o r the percentage of carbon vs. cysts/gm was s i g n i f i c a n t (p = 0.01) and p o s i t i v e i n d i c a t i n g that when carbon tended to be high cyst numbers also tended to be high. Again t h i s i s not an unexpected r e s u l t . Cysts, being organic, form part of the carbon content of sediments, (figure 6) B. EXCYSTMENT RESULTS Viable cysts were i s o l a t e d from fresh d e t r i t a l sludge material from Hidden Basin. The cysts were i s o l a t e d and placed i n a drop of seawater as already described i n the methods. On excystment the cysts /37 /37 could then be correlated with p a r t i c u l a r thecate vegetative c e l l s . The r e s u l t s of excystment experiments are described below. Nine hundred and f i f t y v i a b l e cysts were i s o l a t e d over the two year study. period. Of these, 404 s u c c e s s f u l l y excysted which i s 42% of the t o t a l . I n i t i a l l y a n t i b i o t i c s were used to r i d the cysts of b a c t e r i a but t h i s treatment did not improve the rate of successful excystments and was discontinued. C h a r a c t e r i s t i c s important i n i d e n t i f y i n g the species are l i s t e d . Size ranges (where possible) are given for both c e l l s and cysts from t h i s region and, for comparison, ranges for the species from other regions are given as w e l l , most often from S c h i l l e r (1937). General comments are made about each species. Records of presence i n the phytoplankton of t h i s coast, any unusual feature of the c e l l , cyst, or excystment process, ecology and other notes of i n t e r e s t are men-tioned. GENUS GONYAULAX - TYPICAL TABULATION: 3-6', 0-4a, 6", 6c, 6'1 ' , 5-10s, 5-6"', l p , 1 " " GONYAULAX SP. 1 Thecate Stage Descriptive features: (Plate 111, f i g . 10,11; Plate IX, f i g . 10-12) The c e l l i s nearly as broad as i t i s long. O v e r a l l i t appears somewhat misshapen due to the extreme displacement of the g i r d l e m /38 /38 The epitheca has convex sides; the angle subtended by the two sides at the apex is 90° or slightly greater. There are four apical plates; plate 4' is crescent-shaped with straight ends. There i s an extra plate (la) at the l e f t margin of the sulcus (Pi. 3, f i g . 11) . Whether i t touches the apex or not is uncertain but i f i t does i t would be plate 5"'. Plate 6" is quadrangular; the upper margin is straight to accommodate Plate l a (or 5"). The girdle is descending (left-handed), displaced by three times i t s own -width, is deeply excavated, with l i s t s that appear to have serrated edges. There is no overlap of the girdle edges at the sulcus. The sulcus zigzags slightly; i t indents the hypotheca deeply. The hypotheca is roundly squared; the right antapical horn i s slightly lower than the l e f t . Plate 1'11 is linear and rectangular. The rest of the tabulation is typical for the genus. The theca is somewhat delicate; the plate pattern consists of rows of pores with uneven connecting lines. Size: Body Length 44-56 urn mean - 52 urn Body width 40-50 urn mean - 48 urn N = 4 /39 /39 Cyst Stage Cyst name; S p i n i f e r i t e s sp. Descriptive features; (Plate IX, f i g . 9) The body i s nearly s p h e r i c a l with a pronounced a p i c a l boss; the wall i s granular. T h e tabulation i s t y p i c a l for the genus. The processes have fenestrate bases (Plate XIII, f i g . 5). They are t r i f u r c a t e with b i f i d ends and are often bent over onto the cyst w a l l . The processes around the lower paragirdle margin are connected 'by a low membranous septum which looks l i k e a r u f f l e . Membranous septa occur between the processes around the operculum and the apex. Their development can be highly v a r i a b l e from being along the para-sutures to being between only a few selected processes. With respect to the processes these cysts resemble S^ bentori Rossignol as i l l u s t r a t e d i n Eisenack (page 525, band I I , 1971). The parasulcus i s r e l a t i v e l y s t r a i g h t and broad and broader at the antapex than at the apex. The pa r a s u l c a l p l a t e l e t s are not c l e a r l y defined! l 1 1 ' i s l i n e a r . The archeopyle i s large; i t i s a d i s t o r t e d quadrangle i n shape. The operculum consists of two pieces. Size: Transdiameter - 47-67 pm N = 4 ./40 Ao Due to typographical error, there is no page 40. / 4 1 General remarks: These cysts appear to be quite v a r i a b l e i n the following charac-t e r i s t i c s : - length of the processes - whether the processes stand out or are bent over onto the cyst wall - the degree of development of sutural septa Another cyst type very s i m i l a r to the above species has an archeopyle which i s even larger and i s f i v e - s i d e d (Plate XII, f i g . 7, 8). GONYAULAX SP 2 Thecate Stage Descriptive features: (Plate I I I , f i g . 3-6; Plate IX, f i g . 4-7) The c e l l i s roundly oval; the epitheca has convex sides; a small a p i c a l horn i s present. The e p i t h e c a l tabulation appears to be s i m i l a r to that for Gonyaulax sp- 1 including the presence of an extra small plate ( i a ) on the l e f t margin of the sulcus (PI. 3, f i g . 3). The g i r d l e i s descending (left-handed), displaced by one and a h a l f to two times i t s own width, moderately excavated, with small l i s t s . The hypotheca i s rounded i n ventral view; the tabulation i s t y p i c a l for the genus. The theca i s d e l i c a t e with small scattered pores which are occasionally joined by f i n e l i n e s (PI. I l l , f i g . 5). /42 /42 Size: Length - 49-57 urn . Width - 44-53 um N = 3 General remarks: This c e l l has v i r t u a l l y i d e n t i c a l tabulation to Gonyaulax sp. 1 as far as could be determined. I t d i f f e r s from i t i n the following ways: - theca i s more d e l i c a t e - hypothecal o u t l i n e i s more rounded - g i r d l e displacement i s not as great - o v e r a l l body d i s t o r t i o n i s absent Because of the r e l a t i v e l y few numbers of c e l l s which have been excysted so f a r i t i s not known whether the differences between Gonyaulax sp. 1 and 2 are to be expected i n a normally varying popu-l a t i o n of one species, or whether they are two d i s t i n c t species. Cyst Stage Size: Transdiameter -Processes -49-55 um 15-18 pm . .../43 /43 This cyst i s i d e n t i c a l to the cyst of Gonyaulax sp. 1. The only d i s c e r n i b l e difference i s i n the processes which stand erect rather than curving over towards t h e i r bases as they do i n G-^  sp. 1 (Plate IX, f i g . 8). GONYAULAX TAMARENSIS Lebour Thecate Stage References used: Lebour, 1925: 95, PI. 14, Figs. l a - d . S c h i l l e r , 1973: 299, F i g . 306a-d. Anderson & Wall, 1978: 224-234, Figs. Turpin et a l . , 1978: 235-238, Figs. 1-9. Descriptive features: The c e l l i s roundly oval, s l i g h t l y longer than wide. Plate 1' i s f a i r l y broad. The g i r d l e i s descending(left-handed), displaced one girdle-width, deeply excavated, with well-developed l i s t s . The sulcus broadens p o s t e r i o r l y and reaches the centre of the antapex. The theca i s smooth. There are numerous golden-brown chloroplasts present. Siz e : B.C. Anderson & Wall. S c h i l l e r Length - 23-30 um f 36 um Width - 23-26 pn 35 um N = 7 ./44 /44 General remarks: This species tends to occur i n brackish water. I t commonly occurs i n pa i r s i n catenate fashion, occasionally i n f o u r s , and r a r e l y i n a chain of eight c e l l s . I t i s commonly t o x i c and forms red t i d e s . Taxonomic d i f f i c u l t i e s have occurred with t h i s species and another much l i k e i t c a l l e d Gonyaulax excavata (Braarud) Balech (= G^ _ tamarensis var. excavata). G^ tamarensis has been distinguished by Anderson & Wall (1978) by the presence of an a p i c a l pore near the r i g h t margin of the f i r s t a p i c a l p l a t e although Lebour does not mention a pore i n the o r i g i n a l d e s c r i p t i o n . excavata does not have t h i s pore. The species found here (Turpin et a l . , 1978)lacks the pore as w e l l but i s s t i l l being c a l l e d G^ _ tamarensis. Cyst Stage Descriptive features: (Plate IX, f i g s . 2, 3) The cyst i s an " a c r i t a r c h - l i k e " nondescript oval body with an a p i c a l excystment aperture. The wall i s colourless and smooth with no d i s t i n g u i s h i n g features ( P i . IX, f i g . 3). Size: B.C. Anderson & Wall Length - 30-40 um 43-72 urn Width - 24-29 urn 26-39 um N=7 ./45 /45 General remarks: Pr i o r to excystment the cysts are t y p i c a l l y bilayered, oval, with an orange pigment spot and numerous starch granules (PI. IX, f i g . 2). The cysts are much larger than the vegetative c e l l s . Anderson & Wall (1978) found several excysted c e l l s with two posterior f l a g e l l a . This evidence plus the larger size of the cysts compared to the c e l l s supports the conclusion that these are zygotic cysts. Work done in B.C. (Turpin jet al.,1978) showing G. tamarensis undergoing sexual fusion to form large immobile zygotic cysts also supports this conclusion. GENUS PERIDINIOPSIS - TYPICAL TABULATION 3*, l-2a, 6", 3-6c, 6s, 5'" , 2"" PERIDINIOPSIS CF.HAINANENSIS (Nie) Taylor, 1976; 131 Thecate Stage Synonym:- Diplopsalis hainanensis Nie, 1943: 13, f i g s . 20-25. Descriptive features: (Plate I, f i g . l-4;Plate V I I I , f i g . 9,10). The body i s spherical; apical plates are three i n number and com-pletely surround the apex and the closing p l a t e l e t ; plate 1' i s dia-mond-shaped and nearly as wide as i t i s long; plate l a i s well back from the closing p l a t e l e t . The g i r d l e i s s l i g h t l y ascending (right-.../46 /46 handed), being displaced about k of i t s own width; the g i r d l e has l i s t s without "supporting r i b s . The body i s smooth with r e l a t i v e l y few i n d i s t i n c t pores. Size: B. C. Japan Transdiameter - 42 pm 39-78 pm (Nie) Apex to antapex - 38 pm N = 3 General remarks; The thecate stage of t h i s species i s not known to occur i n the phytoplankton here but has been found on excystment on three occa-sions so f a r . Nie (1948) found that i t was always i n as s o c i a t i o n with D i p l o p s a l i s l e n t i c u l a Bergh and P e r i d i n i o p s i s asymmetrica Mangin. These l a t t e r species are known to occur i n these waters (Wailes, 1937). These c e l l s are very s i m i l a r to P^ asymmetrica except f o r the lack of the small anterior i n t e r c a l a r y p l a t e , a c h a r a c t e r i s t i c of t h i s species. Plate 5" of these c e l l s has an angularity such that i t looks l i k e i t might give r i s e to an i n t e r c a l a r y p l a t e , but (at l e a s t i n the laboratory) f a i l e d to do so (PI. 1, f i g . 3). Af t e r excystment the c e l l ecdysed and t h i s process involved the attachment of the theca to the substrate by means of a strong tubular gelatinous structure which arose from a portion of the dorsal hypotheca (PI. I, f i g . 2). The union of t h i s structure with the substrate could • /47 /47 not be severed although the theca was separated from i t readily. The actual plate involved in this attachment was not apparent. Motile living cells have not as yet been observed, only thecae. Cyst Stage: (Plate VIII, figs. 8-10) Descriptive features: The cyst is small, about 43 um in diameter and covered with short (2 um) simple spines (PI. VIII, f i g . 8). The cysts are characteris-t i c a l l y dark gray before excystment occurs. The excystment aperture consists of a s p l i t nearly encircling the whole cyst (see arrow in PI. VIII, f i g . 8). GENUS PROTOPERIDINIUM - TYPICAL TABULATION 4', 2-4a, 7", 3-6c, 4-7c, 5'", 2 " " PROTOPERIDINIUM ASPIDOTUM (Balech) Balech Thecate Stage Basionym: Peridinium aspidotum Balech, 1964a:. 23, P i . 2, figs. 15 Synonym: Protoperidinium minutum sensu Fukuyo et a l . , 1977: 11, figs. 5-14 non P. minutum (Kofoid) Loeblich. Descriptive features: (Plate I, f i g . 9-11; Plate VIII, f i g . 2) The c e l l has a rotund shape with a small apical horn; the two anterior intercalary plates are asymmetrical being notably unequal in ./48 /48 s i z e . Plate 1" i s much smaller than 7" ( P i . I, f i g . 10). The r i g h t s u l c a l plate gives r i s e to a r e l a t i v e l y large f i n on i t s l e f t border. The g i r d l e i s not displaced, not excavated, with small l i s t s . S i z e : B.C. Japan Argentina Transdiameter - | 31-47 um (mean - 40 pm) 29-33 pm 34-35 pm N = 14 General remarks: This species i s very s i m i l a r i n shape to Pj_ minutum (Kofoid) Loeblich and can be confused e a s i l y with the l a t t e r unless one looks at i n d i v i d u a l plates c l o s e l y noting the asymmetry and s i z e differences of the key plates as noted above. The si z e of excysted c e l l s from B.C. i s s i g n i f i c a n t l y l a rger than that of c e l l s from other areas. Fukuyo et a l . (1977) has evidently m i s i d e n t i f i e d P^ aspidotum c a l l i n g i t P^ minutum. His drawings c l e a r l y show the unequal pre-cingular and i n t e r c a l a r y p l a t e s . Motile c e l l s of P^ _ aspidotum do occur i n the phytoplankton of t h i s coast and has been examined with the scanning electron microscope (F.J.R. Taylor, pers. comm.). Cyst Stage Cyst name: none •. ../49 /49 References used: Fukuyo et a l . , 1977: 11, f i g s . 2a, 3. Descriptive features: (Plate VIII, f i g . 1) The body i s s p h e r i c a l ; the processes are simple spines covering the t e s t wall i n no d i s c e r n i b l e pattern. The t e s t wall between the spines i s smooth; the archeopyle i s an elongate hexagon and presumed to be i n t e r c a l a r y . Size: B.C. Japan Transdiameter -Processes -34-52 um, 28-32 um long 2-7 um 4-5 um N = 14 General remarks: The cysts which produce P^ aspidotum i n t h i s region are s i m i l a r to those from Japan. Wall & Dale (1968a) have excysted I\_ minutum from cysts which are quite d i f f e r e n t . The d i f f e r i n g cyst character-i s t i c s further confirms that these two s i m i l a r c e l l s are d i s t i n c t species. The m i s i d e n t i f i c a t i o n by Fukuyo et a l . (1977) explains the difference i n cyst morphology. These differences were of concern to him because he thought that h i s r e s u l t s showed that two d i f f e r e n t cysts could a r i s e from the same thecal types. He was also concerned that the cyst might be confused with Operculodinium centrocarpum (Deflandre & Cookson)Wall, The differences between the two cysts seem to be s u f f i c i e n t to separate them r e l i a b l y (Table III). /50 /50 TABLE I I I A COMPARISON OF CYST MORPHOLOGY OF PERIDINIUM ASPIDOTUM (FUKUYO ET AL.,1977) AND OPERCULODINIUM CENTROCARPUM (WALL & DALE, 1968a) Feature P. aspidotum 0. centrocarpum Size - -Archeopyle intercalary, hexagonal precingular, subtrapezoidal Processes simple apices, 3-5-um f i b r i l l a r apices, 1/5- 1/4 long test diameter i n length Process base not stated s t r i a t e d Test wall smooth granular Composition unknown sporopollenin-type material Fossils Recent fresh material Mesozoic, Tertiary, Recent only treated sediments ./51 /51 PROTOPERIDINIUM CLAUDICANS (Paulsen) Balech Thecate Stage Basionym: Peridihium claUdicans Paulsen References used: Lebour, 1925: 123, P i . 25, f i g s . l a - d . Wailes, 1937: 36, f i g . 109. S c h i l l e r , 1937: 249, f i g s . 250 a-g. Descriptive features: (Plate I, f i g . 5; Plate VI, f i g . 1) This c e l l c l o s e l y resembles Protoperidinium oblongum ( A u r i v i l l i u s ) Cleve (see p. 51), but may be distinguished from i t by plate 2a often being i n contact with precingular plates 3" and 4" rather than j u s t 4" (penta), and by the c e l l being r e l a t i v e l y more rotund. In the l o c a l population the antapical horns are divergent. Plate 2a i s often displaced to the l e f t side of the c e l l . S i z e : B.C. S c h i l l e r Length - 70 - 125 um (mean - 79pm) 51 - 105 pm Width - 50 - 70 um (mean - 62 pm) N = 6 General comments: The s i z e range of t h i s species overlaps with that of Vj_ oblongum; thus s i z e cannot be used to help d i s t i n g u i s h them. /52 /52 These c e l l s are r e l a t i v e l y common to abundant i n the phytoplank-ton of t h i s coast (Wailes, 1937). C e l l s required one to s i x days of incubation at 17°c to excyst. They were small on f i r s t excysting but grew quickly. One c e l l grew from 84 to 125 um long i n f i v e days. I They contain o i l storage pro-ducts ranging i n colour from yellow to deep red. Presumably they s t a r t to use these up as soon as they emerge. The c e l l s normally l i v e only a few days, although one c e l l survived 12 days i n ES medium (Provasoli, 1957) i n which diatoms had begun to grow. Cyst Stage Cyst name: none References used: Wall, 1965: 307, f i g . 16. Wall & Dale, 1966: 351, P i . 1, f i g . R. Wall & Dale, 1968a: 273, PI. 1, f i g . 30, P i . 2, f i g s . 1, 2, PI. 3, f i g . 12. Davey & Rogers, 1975: 217, PI. 1, f i g . 10. Descriptive features: (Plate VI, f i g . 2, 3) The cysts from t h i s region are s i m i l a r to those found on the A t l a n t i c coast (Wall & Dale, 1968a). The shape i s cordate with rounded apex and antapical horns. The e n t i r e cyst i s covered with short (2 jam) simple spines i n no d i s c e r n i b l e pattern. /53 /53 The archeopyle has been presumed to be i n t e r c a l a r y p a r t l y because of i t s p e r i d i n i a n a f f i n i t i e s . In appearance however, i t i s more sub-a p i c a l or even a p i c a l e s p e c i a l l y i f the operculum i s l o s t (Plate VI, f i g . 3). General remarks: These cysts, as for a l l Recent Protoperidinium cysts, survive only poorly i n standard palynogical treatments. In fresh material they and other cysts of t h i s genus are abundant. They were found i n only a few treated samples (1, 6, 11, 24, 36). Wall et a l . (1977) included these cysts i n a general "Peridinium spp." category because t h e i r numbers alone were too low to have any s i g n i -f i c a n t i n d i c a t o r value. PROTOPERIDINIUM CONICOIDES (Paulsen) Balech Thecate Stage Basionym: Peridinium conicoides Paulsen References used: Lebour, 1925: 112, PI. 20, f i g s . 2a-d. Wailes, 1937: 33, f i g . 97. S c h i l l e r , 1937: 231, f i g . 228a-d. Wall & Dale, 1968a: 277, PI. 2, f i g . 28, PI. 3, f i g . 26. Descriptive features: (Plate VI, f i g . 11) The c e l l i s rhombic i n v e n t r a l view with s l i g h t l y convex sides; the ..../54 /54 g i r d l e i s c i r c u l a r and median, moderately excavated, with f i n e l i s t s ; the sulcus broadens p o s t e r i o r l y and twists l e f t j u s t below the g i r d l e ; the hypotheca has two small hollow spines on the antapical horns. Size: B.C. S c h i l l e r . Wailcs Length -Width -75 um 48-60 pn 44-55 pn . 64 um N = 1 General remarks: Only o n e - c e l l has been excysted from the B.C. region. Its s i z e was larger than the stated range ( S c h i l l e r , 1937) f o r t h i s species but other-wise appeared as described. Wailes (1937) has recorded i t s presence i n the phytoplankton here. Cyst Stage Cyst name: Chytroeisphaeridia simplica Wall References used: Wall & Dale, 1968a: 277, P i . 2, f i g . 29, 30, PI. 3, f i g . 27, 28. Descriptive features: The cyst i s the same as that described by Wall & Dale (1968a); i . e . , a brown sphere with a s i x - s i d e d elongate hexagonal archeopyle presumed to be i n t e r c a l a r y . /55 /55 General remarks: Wall & Dale (1968a) d i d not excyst c e l l s of t h i s species. Rather they found cysts with i n t a c t thecae around them and thus established the cyst-theca r e l a t i o n s h i p that way. An indentation i n i t h e cyst wall where i t adjoins the ve n t r a l sulcus of the c e l l was not noted i n the cyst found here. PROTOPERIDINIUM CONICUM (Gran) Balech ... Thecate State Basionym: Peridinium conicum (Gran) Ostenfeld & Schmidt ... References used: Lebour, 1925: 111, P i . 19, f i g s . l a - d . Wailes, 1937: 34, f i g . 100. S c h i l l e r , 1937: 233, f i g s . 229a-j. C a t t e l l , 1969: PI. 6, f i g . 43. Taylor, 1976: 139, PI. 33, f i g s . 361, 362. Descriptive features: (Plate I, f i g . 7, 8; Plate VII, f i g . 10) The c e l l i s nearly symmetrical around the l o n g i t u d i n a l axis and only s l i g h t l y f l a t t e n e d dorso-ventrally (P_;_ pentagonum i s more com-pressed) . The epitheca and hypotheca are nearly equal i n height; the epitheca i s t r i a n g u l a r i n ve n t r a l view with s t r a i g h t sides. The sulcus reaches beyond the centre of the hypotheca; t h i s distinguishes the c e l l from P^ pentagonum which has a shorter sulcus. The sutures /56 /56 between 1' and 2', and 1" and 2" plus t h e i r analogues on the r i g h t form a s t r a i g h t l i n e from apex to g i r d l e (a c h a r a c t e r i s t i c shared only with IV pentagonum). Size: . B.C. S c h i l l e r Length 66-76 um 70-80 um General remarks: One excysted c e l l was more elongated than normal (PI- VII, f i g . 10) but was s t i l l within the reported s i z e range. This species has been recorded i n the phytoplankton of B r i t i s h Columbia (Wailes, 1937; Buchanan 1966; C a t t e l l , 1969). Cyst Stage Cyst names: Mu l t i s p i n u l a quanta Bradford, 1974: 3067, f i g s . 5-7 Figured as P_^_ conicum cysts by: Wall, 1965: 308, f i g s . 17, 23. Wall & Dale, 1968a: 273, P i . 2, f i g s . 3-5, PI. 3, f i g . 13 Dale, 1976: 50, PI. 1, f i g . 17. General remarks: (Plate I, f i g . 6; Plate VII, f i g . 6, 7) This cyst was present i n small numbers i n only seven treated samples (11, 14, 17, 27, 36, 39, 44). A l l of these locations are s h e l -tered bays or i n l e t s . Others have found i t to be present i n many estuarine-coastal locations but i n low numbers. (Wall et a l . , 1977) ./57 /57 They have found i t to be a cosmopolitan " e s t u a r i n e ' n e r i t i c " species. Dale (1976) states that these and other Protoperidinium cysts are badly af f e c t e d by p a l y n o l o g i c a l treatments and are probably under-represented i n a l l treated Quaternary sediments. In comparing cyst numbers i n fresh and acid-treated material from Hidden Basin (station #14) i t can be confirmed that many of these cysts are destroyed by palynologic treatment. Thus i t i s questionable what value they might have i n treated sediments as in d i c a t o r s of environmental conditions. When acid-treated these cysts were i d e n t i c a l i n appearance to those p i c t u r e d by Bradford (1974) and quite d i f f e r e n t to those seen i n fresh material. Size: B.C. Wall & Dale L e f t to r i g h t - 50-65 um 46- 71 um Dorsal to ve n t r a l - 44-47 um 28- 52 um Apex to antapex - 35-51 um 28- 52 um N = 5 Cysts required two to f i v e days to excyst at 17 c. Only P^ _ conicum has been excysted i n t h i s region although others have found s i m i l a r cysts to give r i s e to both P^ conicum and P_^_ nudum (Wall & Dale, 1968a) . One cyst was aberrant i n having spines c l o s e l y adherent to the /58 /58 t e s t w a l l . The operculum was present although displaced and could be seen to consist of two pieces ( P i . 1, f i g . 6; PI. 7, f i g . 7). Dale (1976) shows a cyst with the operculum i n place but c o n s i s t i n g of only one piece. PROTOPERIDINIUM CF. D E N T I C U L A T U M (Gran & Braarud) Balech Thecate Stage Basionym: Peridinium denticUlatum Gran & Braarud References used: Jorgensen, 1900: 40-41. Gran & Braarud, 1935: 381, f i g . 58. Wailes, 1937: 31, f i g . 88. S c h i l l e r , 1937: 519, f i g . 606a-c. Braarud, 1940: 143-144, f i g . l a - c . Wall & Dale, 1968a: 277, PI. 4, f i g . 3. Descriptive features: (Plate I, f i g . 12, 14, 15; Plate VIII, f i g . 3, 4, 7) The epitheca i s c o n i c a l with s t r a i g h t sides; the apex has an unusually elongated pore p l a t e , a c h a r a c t e r i s t i c shared with P^ abei (Paulsen) Balech and Pj_ ventricum (Abe) Balech. There are two a n t e r i o r i n t e r c a l a r i e s . The g i r d l e i s descending (left-handed) about one girdle-width and i s deeply excavated. The hypotheca i s nearly f l a t p o s t e r i o r l y . This c h a r a c t e r i s t i c i s true for s i n g l e c e l l s or the /59 /59 upper c e l l of a p a i r . The theca i s d e l i c a t e . The c e l l s are nearly always joined i n p a i r s although they may be found s i n g l y . S i z e : B.C. S c h i l l e r Length - 32-36 um (mean - 33 um) / Width - 36-46 um (mean - 42 um) 56-76 um N = 8 General remarks: These c e l l s excyst as a s i n g l e rather elongate c e l l (51 um long). with a d e l i c a t e theca and a descending (left-handed) g i r d l e of about one g i r d l e width. Further tabulation was impossible to see. Within 24 hours t h i s c e l l divided to form a c e l l p a i r which was nearly the same s i z e as the o r i g i n a l s i n g l e c e l l . The c e l l p a i r divided once more within the next 24 hours to form two p a i r s s t i l l nearly the same s i z e as the o r i g i n a l s i n g l e c e l l over a l l . The tabulation on one theca showed the t h i r d a p i c a l p l a t e (3 1) to be s l i g h t l y more rounded and larger (PI. I, f i g . 15) than the one figured i n the o r i g i n a l drawings. Otherwise the epit h e c a l tabu-l a t i o n was consistent with that shown by Gran & Braarud (1935) . When f i r s t described only si n g l e c e l l s were mentioned and the species was characterized l a r g e l y on the basis of shape. Later an undesignated student of Braarud's found s i m i l a r c e l l s ..../60 /60 i n a Norwegian phytoplankton sample but they were attached i n p a i r s , hypotheca to hypotheca. Braarud (1940) stated that Wailes' (1937) i d e n t i f i c a t i o n of P.  decepiens Jorgensen from B r i t i s h Columbia was more l i k e l y P^ denticulatum. Wailes' drawings unequivocally show the c e l l p a i r s attached base to base. Braarud's drawings of P. denticulatum show the c e l l s apparently attached base to base. However he d i d not draw i n the s u l c i which, i f they had been seen, would be more d e f i n i t i v e proof of t h i s unusual, mode of attachment. Wall & Dale (1968a) i d e n t i f i e d s i m i l a r c e l l s as P^ denticulatum from excystment experiments using the shape of the theca, and the occurrence of the c e l l s i n p a i r s . They d i d not mention the mode of attachment of the c e l l s i n the p a i r s . However the photograph by Wall & Dale (1968a) (PI. 4, f i g . 3) c l e a r l y shows a p o s t e r i o r flagellum i n such a p o s i t i o n as to preclude the p o s s i b i l i t y of a base to base attachment for these c e l l s . They d i d not comment on t h i s anomaly. In excystment experiments done here the c e l l s were i d e n t i f i e d on the basis of epi t h e c a l tabulation, the shape of the c e l l s , and t h e i r p a i r i n g , although they were found to be attached epitheca to hypotheca. Thus the excysted c e l l s from here are very s i m i l a r to those excysted c e l l s from Woods Hole. Technically they d i f f e r from the c e l l s described by Gran & Braarud (1935) as denticulatum because of t h e i r mode of attachment being apex to base and not base to base /61 /61 and therefore they cannot be d e f i n i t e l y assigned to P. denticulatum. There are two possible explanations: e i t h e r the c e l l s found here and by Wall S Dale (1968a) are not P_^_ denticulatum but a new species with s i m i l a r morphology but a d i s s i m i l a r mode of attachment, or an error was made i n the e a r l i e r observations. C e l l s i n p a i r s have a shape which s u p e r f i c i a l l y makes them look l i k e they are attached base to base. However closer examination of the s u l c i and the arrangement of the f l a g e l l a precludes t h i s p o s s i b i l i t y , at l e a s t for the c e l l s found, here. I t i s suspected that an erro r was made i n s t a t i n g that these c e l l s attach i n a base to base fashion but t h i s cannot be demonstrated. Cyst Stage Cyst name: Chytroeisphaeridia cariacoensis Wall Descriptive features: (Plate VII, f i g . 11, 12) These cysts are small brown smooth-walled spheres with a hexa-gonal archeopyle presumed to be i n t e r c a l a r y . Size: B.C. Woods Hole Mean diameter 56 um 56 um N = 8 General remarks: The cysts from t h i s region are i d e n t i c a l to those described by . . ./62 /62 Wall & Dale (1968a) for P^ denticulatum from the east coast of North America. They are s i m i l a r to cysts produced by other Protoperidinium species such as P. avellana (and P. thorianum (Paulsen) Balech although i t s cyst wall i s s t r i a t e d ) . PROTOPERIDINIUM LEONIS (Pavillard) Balech Thecate Stage Basionym: Peridinium leonis P a v i l l a r d References used: Lebour, 1925: 112, PI. 21, f i g . l a - d . Wailes, 1937: 33, f i g . 98. S c h i l l e r , 1937: 236, f i g . 236a-m. Wall & Dale, 1968a: 276, PI. 3, f i g . 22. Taylor, 1976: 141, P l . 33, f i g . 369. Descriptive features: (Plate I, f i g . 13, Plate VI, f i g . 12) The f i r s t a p i c a l plate i s 'ortho'. Although s i m i l a r to P.  pentagonum and P^ conicum i t can be distinguished by the zigzag nature of the suture running from apex to g i r d l e between plates 1' and 2' and 1" and 2" and the analogues on the r i g h t ( Pl. VI, f i g . 12). The f i r s t and seventh precingular plates are quadrangular. The g i r d l e i s very s l i g h t l y descending (left-handed) being displaced by less than one girdle-width, moderately excavated and with l i s t s . The hypotheca has two small s o l i d spines on the antapical horns, the a n t a p i c a l furrow i s reached by the sulcus. The thecal plates are covered with /63 /63 short spiny processes which are l a r g e l y connected up in t o l i n e a r markings; some areas are f i n e l y r e t i c u l a t e . Size: B.C. S c h i l l e r Length - 61-65 pm (mean - 64 pm) . 65-95 pm Width - 61-46 pm (mean - 63 pm) 75-80 pm N = 10 General remarks: One c e l l which came from a t y p i c a l cyst had a small aberration i n the ve n t r a l g i r d l e region (Plate VI, f i g . 12). Cyst Stage Cyst name: none References used: Wall & Dale, 1968a: 276, Pi.' 2, f i g s . 18-21. Descriptive features: (Plate VII, f i g . 4). The cysts from t h i s region are i d e n t i c a l to those described by Wall & Dale (1968a). They are pentagonal i n ve n t r a l view, tapering to two antapical horns; a ve n t r a l s u l c a l region deeply indents between them. The g i r d l e region i s defined by a broad ridge j u s t above the antapical region. The cyst w a l l i s smooth, dark brown, with an i n t e r c a l a r y , subtriangular archeopyle. /64 /64 Size: B.C. Woods Hole Length - 62-74 um 52-72 um Width - 52-74 um 53-76 .pm General remarks: P. leonis cysts were present i n treated sediment samples i n low numbers. The thecate stage i s generally known here and i s very common (Wailes, 1937; C a t t e l l , 1969). In untreated Hidden Basin material P.  leonis cysts were abundant although i t was d i f f i c u l t to get many to excyst. PROTOPERIDINIUM OBLONGUM ( A u r i v i l l i u s ) Parke and Dodge, 1976 Thecate Stage Basionym: Peridinium divergens var. oblongum A u r i v i l l i u s Synonym(s): Peridinium oceanicum var. oblongum ( A u r i v i l l i u s ) Paulsen Peridinium oblongum ( A u r i v i l l i u s ) Cleve .... References used; Lebour, 1925: 121, PI. 24, f i g s . l a - c . Wailes, 1937: 36, f i g . 107. S c h i l l e r , 1937. 260, f i g s . 257c-e, g, g l , g2. Wall & Dale, 1968a: 272, PI. 1, f i g . 22, 24, 25, PI. 3, f i g . 9-11. /65 /65 Descriptive features: (Plate I, f i g . 16) It i s d i f f i c u l t to d i s t i n g u i s h t h i s species from the morpholo-g i c a l l y widely va r i a b l e group of c e l l s known as Protoperidinium oceani- cum . In P. oblongum most authors have used the following character-i s t i c s to i d e n t i f y i t as a species separate from the others of the complex: siz e - body length less than 170 um, lack of a d i s t i n c t a p i c a l horn, width approximately 2/3 of length. Size: B.C. Woods Hole Length - 93-121 pm (mean - 116 um) 77-132 pm Width - 65 - 98 um (mean N = 6 - 76 um) 52-80 pm General remarks: This species i s known to occur i n B.C. waters. P. oceanicum and P. depressum also occur (Wailes, 1937; C a t t e l l , 1969). Cyst Stage Cyst name: none References used: Wall & Dale, 1968a: 272, P l . 1, f i g . 23, 29. Davey & Rogers, 1975: P l . 1, f i g . 11. /66 /66 Descriptive features: (Plate VII, f i g . 8, 9) The cyst wall i s smooth, the cyst shape varies from pentagonal with a p i c a l and antapical horns to cordiform with rounded ends. The archeopyle i s i n t e r c a l a r y and sub-triangular i n shape. Size : B.C. Woods Hole Length -Width -74-84 pm (mean - 81 pm) 62-84 pm (mean - 77 pa) 70-N = 74 pm 10 (mean - 73 pm) 56-74 pm (rhombic (mean -cysts) 65 pm) General remarks: These cysts are seen only o c c a s i o n a l l y i n treated samples. Their appearance i s a l t e r e d somewhat from those i n fresh material; the wall i s very t h i n , i t has a crumpled look and often the shape i s d i s t o r t e d . They were abundant i n fresh Hidden Basin material; i n fac t they were the most common cysts found there. Cysts described by Wall & Dale (1968a) as being horned (form 1), have not been found here. The cysts are sa i d by Wall & Dale (1968a) to resemble the f o s s i l genus Deflandrea except a f o s s i l record f o r these p a r t i c u l a r Proto- peridinium cysts i s absent. ./67 /67 PROTOPERIDINIUM PENTAGONUM (Gran) Balech Thecate Stage Basionym: Peridinium pentagonum Gran References used: Lebour, 1925: 112, P l . 20, f i g s . l a - e . Abe, 1927: 409, f i g . 28. Wailes, 1937: 34, f i g . 101. . S c h i l l e r , 1937: 241, f i g . 242a-e. Wall & Dale, 1968a: 274, P l . 2, f i g s . 8-10, P l . 3, f i g . 15. C a t t e l l , 1969: P l . 7, f i g . 50. Descriptive features: (Plate I I , f i g . 1, 2, 4, 5; Plate VIII, f i g . The c e l l i s f i v e - s i d e d i n v e n t r a l view and i n a p i c a l view the l e f t side i s smaller than the r i g h t ; i t has weakly developed antapical horns tipped with small spines. I t i s moderately dorso-ventrally com-pressed. The g i r d l e i s descending (left-handed) about one g i r d l e -width, and moderately excavated with narrow f i n e l y - r i b b e d l i s t s . The sulcus i s shorter than i n P^ _ conicum and does riot reach to the post-e r i o r margin of the hypotheca. The sulcus i s rounded at the antapical end. The sutures between 1' and 2', and 1" and 2" plus t h e i r analogues on the r i g h t side form a r e l a t i v e l y s t r a i g h t l i n e from apex to g i r d l e (P l . I I / f i g . 1) and are ridged . This c h a r a c t e r i s t i c i s shared only with P_;_ conicum. /68 /68 Size: B.C. Schiller Length - 79--120 um (mean - 100 Um) .75-100 um Width - • 74--106 pm (mean - 89 pm) 75-100 pm N = = 9 General remarks: The presence of P^ pentagonum has been recorded from the phy-toplankton in B.C. waters and is f a i r l y common (Wailes, 1937; Cattell, 1969) . Cells excysted from Hidden Basin material have an antapical plate (homologous to the posterior sulcal plate of other Protoperidinium species) which is smaller than the same plate shown by Lebour (1925)-(PI. 2, f i g . 5). Excystment occurred within two days of viable cysts being kept o at 17 c. Excysted cells were characteristically deeply coloured with orange-brown o i l droplets. They did not contain chloroplasts. Schiller (1937) reported golden chloroplasts but they are not present . Cells frequently divided within one day of excystment. Further-more, they often ecdysed without dying so that in time as many as five thecae could result from a single c e l l . Maximum survival time for excysted cells was five days. ./69 /69 The l a r g e s t c e l l s which excysted were larger than the maximum length, as recorded by S c h i l l e r (1937). Cyst Stage Cyst name: none References used: Wall & Dale, 1968a: 274, P l . 2, f i g . 29, 30, P l . 3, f i g . 28. Descriptive features: (Plate I I , f i g . 3; Plate VIII, f i g . 6) Smooth brown spheres with a single-layered outer w a l l : cyst contents were dark brown and granular, enclosed within an inner hyaline envelope. The hyaline envelope was e a s i l y seen when p u l l e d away from the outer wall i n empty cysts. Occasionally cysts are noted which are more pyriform i n shape. The archeopyle i s an elongate hexagon and presumably i n t e r c a l a r y . The operculum often remains attached and i s i n one piece. Size: Transdiameter 65-93 pn (mean - 78 um) N = 11 General remarks: The cysts of P^ pentagonum from t h i s region are very d i f f e r e n t from those described from Woods Hole by Wall & Dale (1968a). They found /70 /70 that the cysts for t h i s species were pentagonal i n v e n t r a l view with rounded antapices and a blunt apex, colourless, and decorated ran-domly with spinules. The cysts were 72 um long and 76 pm wide. How-ever, the archeopyles of both cysts are s i m i l a r . PROTOPERIDINIUM PUNCTULATUM (Paulsen) Balech Thecate Stage Basionym: Peridinium punctulatum Paulsen Synonym(s): Peridinium subinerme var. punctulatum (Paulsen) S c h i l l e r References used: Lebour, 1925: 123, text f i g . 37. Wailes, 1937: 37, f i g . 110. S c h i l l e r , 1937: 245, f i g . 245a, b. Wall & Dale,'1968a: 276/ P i . 2, f i g . 25, PI. 3, f i g . 24. Descriptive features: (Plate I I I , f i g . 7-9; Plate VI, f i g . 9, 10) This c e l l i s s i m i l a r to P^ subinerme (Paulsen) Loeblich i n basic shape but d i f f e r s i n several other respects. I t i s broader than long. The sulcus i s narrow, not widening p o s t e r i o r l y as i n P^ subinerme.. The second a n t e r i o r i n t e r c a l a r y i s usually penta, sometimes hexa. Antapical horns are as i n P_^_ subinerme. The theca i s f i n e l y punctate and possesses small scattered spines. ..../71 /71 Size: B.C. S c h i l l e r Length -Width -66-70 um 58-107 um 60 um 50-103 pm N = 2 General remarks: Two c e l l s have been excysted from Hidden Basin m a t e r i a l . In both cases most of the dorsal epitheca was l o s t on ecdysis and thus i d e n t i -f i c a t i o n was based on features of the theca other than epi t h e c a l plates. Excysted c e l l s of punctulatum from B r i t i s h Columbia are much lar g e r than the range given by Wailes (1937) (34-38 um length and breadth). Such a large discrepancy casts doubt on Wailes' observa-tions and i t i s u n l i k e l y that he was measuring the same species. Cyst Stage Cyst name: none References used: Wall & Dale, 1968a: 276, P l . 2, f i g . 27, P l . 3, f i g . 25. Descriptive features: The cyst i s a brown sphere. B.C. specimens were 52 pm diameter. The archeopyle i s presumed to be i n t e r c a l a r y . I t i s an elongate hexagon, much l i k e that found i n P. pentagonum (Gran) Balech or P^ /72 /72 conicoides (Paulsen) Balech). General remarks: P r i o r to excystment these cysts are v i r t u a l l y i n d i s t i n g u i s h a b l e from the cysts of several other Protoperidinium species as was also noted by Wall & Dale (1968a). They state that these cysts are s i m i l a r to those of P_^_ avellana (Meunier) Balech and P^ denticulatum (Gran & Braarud) Balech, but see further comments i n the discussion. PROTOPERIDINIUM SP. NOV. Thecate Stage Diagnosis: This species i s very s i m i l a r to Protoperidinium nux (Abe) Balech but d i f f e r s from i t i n lacking an a p i c a l pore, having a g i r d l e d i s -placement greater than one girdle-width, being nearly twice as large, and having a f i n e l y rugose thecal surface. Size: Transdiameter - 48- 54 um (mean -- 51 um) Length - 53- 67 um (mean -- 59 um) N = = 8 Holotype - 51 x 54 um D i s t r i b u t i o n - thecae derived from excystment of cysts from fresh Hidden Basin material ...113 /73 Description: (Plate 11, f i g . 6-12; Plate VI, f i g . 6-8) The c e l l i s roundly rhombic i n v e n t r a l view; the a p i c a l end i s more pointed than the antapical end. There i s only s l i g h t dorso-ve n t r a l f l a t t e n i n g . The length i s s l i g h t l y greater than the width. The epitheca i s coni c a l with s l i g h t l y angular, convex sides; there i s no a p i c a l pore present; p l a t e 1' i s 'ortho' (four-sided). There are two anterior i n t e r c a l a r y p l a t e s , the l e f t smaller than the r i g h t . There are seven precingular p l a t e s . Plate 1" i s smaller than p l a t e 7" as a r e s u l t of the g i r d l e displacement. The g i r d l e i s descending (left-handed), displaced by s l i g h t l y more than one girdle-width; i t i s moderately excavated without conspicuous l i s t s . The sulcus i s narrow near the g i r d l e and widens towards the antapex; i t does not reach the antapical p r o f i l e when seen i n f u l l v e n t r a l view;;it i s moderately indented. The surface i s covered with small p a p i l l a e which may be arranged i n rows. No chloroplasts are present. Cyst Stage Cyst name: none ./74 /74 Description: (Plate I I , f i g . 13) The cyst i s a smooth dark brown sphere with a s i x - s i d e d arche-opyle presumed to be i n t e r c a l a r y . The operculum i s i n one piece. Size: Transdiameter 47-56 pm N = 8 General remarks: Cysts of t h i s new species are d i f f i c u l t to d i s t i n g u i s h from those of P^ punctulatum, P. thorianum and others p r i o r to excystment. Brown sp h e r i c a l cysts l i k e those of t h i s species have been r e f e r r e d to the genus Chytroeisphaeridia Sarjeant by Wall & Dale (1968a). These cysts are abundant i n fresh Hidden Basin material but rare i n treated material. PROTOPERIDINIUM THORIANUM (Paulsen) Balech Thecate Stage Basionym: Peridinium thorianum Paulsen References used: Lebour, 1925: 108, P i . 17, f i g s . 2a-f. Wailes, 1937: 31, f i g . 91. S c h i l l e r , 1937: 142, f i g s . 143a-e. Descriptive features: (Plate I, f i g . 17, 18; Plate I I I , f i g . 1, 2; Plate VI, f i g . 4, 5) ./75 /75 The c e l l has a rhombic shape i n v e n t r a l view. The g i r d l e has l i n e a r markings and i s moderately excavated, descending (left-handed) about one girdle-width. The sulcus i s s t r a i g h t and narrow and does not reach the centre of the hypotheca. The theca i s highly rugose, the plates being covered by conspicuous p a p i l l a e . Size: B.C. S c h i l l e r Transdiameter 62-70 um 56-70 pm N = 7 General remarks: This species has been previously recorded from B.C. waters by Wailes (1937). One excysted c e l l was smaller than usual, being only 55 um wide. Part of the theca had the d i s t i n c t i v e p a p i l l a e of a normal c e l l but seven of the e p i t h e c a l plates were smooth. The sulcus was wide and deeply indented the hypotheca. The side of the epitheca with the smooth plates was concave whereas the normal side was convex. (Plate I I I , f i g . 1 , 2 ) . Cyst Stage Cyst name: none Descriptive features: (Plate I, f i g . 19) These are s p h e r i c a l and dark brown. The outer w a l l i s s i n g l e /76 . - /76 layered and occasionally a second inner hyaline layer can be observed shrunken away from the outer cyst w a l l . The cyst diameter i s remark-ably constant, being 56-57 um i n a l l those measured (N = 11). The archeopyle i s somewhat unusual being nearly seven-sided, and presumed to be i n t e r c a l a r y . General remarks: These cysts are very s i m i l a r to the cysts of several other thecate c e l l s . P r i o r to excystment i t i s d i f f i c u l t to t e l l them apart. The f a c t that P^ _ thorianum has t h i s kind of cyst i s confirmation of a previously unknown cyst-theca r e l a t i o n s h i p . GENUS ZYGABIKODINIUM - TYPICAL TABULATION 3' , l-2a, 7", 3c, 4s, 5" ' , 1' ' ' ' ZYGABIKODINIUM LENTICULATUM Thecate Stage Synonyms: D i p l o p s a l i s l e n t i c u l a f. minor P a v i l l a r d D i p lopeltopsis minor (Paulsen) P a v i l l a r d ... References used: Lebour, 1922: 801, f i g s . 11-15. Lebour, 1925: 102, f i g . 2a-e. Wailes, 1937: 29, f i g . 84. S c h i l l e r , 1937: 105, f i g . 96a-e. Wall & Dale, 1968a: 280, f i g . 7 Taylor, 1976: 169, P i . 28, f i g s . 295, 297, 300. /77 /77 Descriptive features: Plate I I , f i g . 14-17; Plate VII, f i g . 1-3) The c e l l i s c i r c u l a r i n a p i c a l view, l e n t i c u l a r i n v e n t r a l view. There are seven precingular plates ( P l . 2, f i g . 14) rather than s i x as i n Diplopsalis;there are two anterior i n t e r c a l a r i e s , the f i r s t (2a, P l . 2, f i g s . 14, 16) i s small, diamond-shaped and on the l e f t side. The f i r s t a p i c a l plate ( l 1 ) i s c h a r a c t e r i s t i c a l l y narrow. The g i r d l e i s not displaced, not indented and has transverse l i s t s with very f i n e r i b s . There i s only one large antapical plate ( P l . 2, f i g . 15). The-theca i s covered with r e l a t i v e l y conspicuous pores ( P l . 2, f i g . 14). General remarks: Only one c e l l has been found i n excystment experiments. Its diameter was 47 pm which was within the stated range of 40-55 pm f o r the species ( S c h i l l e r , 1937). I t has been previously recorded from the phytoplankton of the B.C. coast (Wailes,1937). Cyst Stage Cyst name: none. References used: Wall & Dale, 1968a: 280, P l . 4, f i g s . 3, 21, 22. Descriptive features: (Plate VII, f i g . 5) The cyst for t h i s thecate species i s s i m i l a r to that described by Wall & Dale (1968a). I t i s a brown sphere; the archeopyle i s a /78 /78 s p l i t along the cyst margin i n the region of the upper g i r d l e . Reflected tabulation can be seen on the cyst wall to a greater or le s s e r extent once the cyst i s empty. C. CYSTS IN SEDIMENTS Many cysts described i n t h i s study were observed only i n paly-nologic samples. Each cyst species i s of necessity presented under the p a l e o n t o l o g i c a l name except where the thecate stage i s known and i t s name has p r i o r i t y . Diagnostic characters are l i s t e d . Size ranges are given for cyst species from B.C. and, for comparison, s i z e ranges are given for the cyst species from Eisenack (1964, 1971) and other sources where po s s i b l e . General remarks about cyst species include any d e t a i l s of i d e n t i f i c a t i o n problems, any c h a r a c t e r i s t i c s which make i d e n t i f i c a t i o n uncertain, d i s t r i b u t i o n and r e l a t i v e abundance, and other notes of i n t e r e s t . Throughout t h i s study the terminology for cyst features has been based on the recommendations of E v i t t et a l . (1976). GENUS OPERCULODINIUM - TYPICAL TABULATION " Not disti n g u i s h a b l e OPERCULODINIUM CENTROCARPUM (Deflandre & Cookson) Wall Cyst Stage Synonym(s): Baltisphaeridium centrocarpum (Deflandre & Cookson) Gerlach ... ./79 /79 References used: Wall, 1967: 111, P l . 16, figs. 1, 2, 5. Wall & Dale, 1967: 352, P l . 1, f i g . M. Wall & Dale, 1968a: 272, P l . 1, figs. 20, 21. Wall & Dale, 1968b: 326, P l . 1, f i g . 10-12. Eisenack, 1971: 867 (Bd. II) Reid, 1974: 594, P l . 2, figs. 10, 11. Dale, 1976: 53, P l . 1, f i g . 3. Descriptive features: (Plate XII, f i g . 1; Plate XIII, f i g . 3, 4) The cyst is globular, more or less e l l i p s o i d a l , covered with numerous straight slender processes. The surface i s granular between the spines using light microscopy. The processes are solid with radiat-ing f i b r i l s at their point of insertion (Pl- XIII, f i g . 4). Under SEM (scanning electron microscope) their apices can be seen to be fringed by the same sort of f i b r i l s which under light microscopy appear as a cup-like termination. The processes are typically 1/5 to 1/4 the cyst diameter in length. The archeopyle is roundly quad\-irangular and i s presumed to correspond to the third dorsal precingular plate (Wall, 1967). S i z e : B.C. Woods Hole Caribbean North Sea Transdiameter - 31-45 um (mean - 42) 33-37 pm (Wall & Dale, 40-50 pm 33-48 pm 1968a) (Wall, 1967) (Reid, 1974) Process length - 2-12 um N = 58 6-16 pm y. . 7-14 pm ./80 /80 General remarks: The general morphology of t h i s cyst i s highly v a r i a b l e . The wal l can be thick or t h i n , smooth or granular, the processes vary from being very long to absent (Harland, 1973 and present m a t e r i a l ) . 0. psilatum Wall appears to be the smooth species of the genus. 0. israelianum (Rossignol) Wall has shorter spines, equivalent to 1/10 or le s s of the diameter of the body. Reid (1974) d i d not f i n d psilatum i n samples around the B r i t i s h I s l e s although t h i s cyst species gives r i s e to the same thecate species as centrocarpum, i . e . Protoceratium reticulatum = Gonyaulax  g r i n d l e y i (Von Stosch, 1969). He included 0^ israelianum with h i s 0.  centrocarpum counts without giving a reason. 0. psilatum has not been found on the B.C. coast although cysts without spines and a granular rather than smooth w a l l do occur. 0.  israelianum has been noted as w e l l . A l l the types which occur here from the spineless granulated forms to the " t y p i c a l " forms have been com-bined with centrocarpum because the existence of gradations between them makes t h e i r assignment to species very d i f f i c u l t . These cysts are over-represented i n Recent sediments of th i s region, when compared with the numbers of the thecate stage i n the plankton (F.J.R. Taylor,pers. comm.) a s i t u a t i o n also found i n Norway(D al e> 1976). 0^ centrocarpum i s the dominant cyst species i n most of the ./81 /81 sample locations here. Two samples (#34, #37) had very high absolute cyst numbers (reaching 97,000 cysts/gram) i n which t h i s cyst was v i r t u a l l y monospecific. The geochromologic range f or t h i s species i s Eocene-Recent. Thecate Stage Name: Gonyaulax g r i n d l e y i Reinecke Synonym(s): Protoceratium reticulatum (Claparede & Lachmann) II B u t s c h l i References used; Wailes, 1937: 26, f i g . 73. Wall & Dale, 1968a: 272, P l . 1, f i g . 19, P l . 3, f i g s . 7, 8. General remarks: This species has not been excysted from any cysts i n t h i s region. I t i s present but uncommon i n the plankton (Wailes, 1937). Occa-s i o n a l l y G^ g r i n d l e y i forms red tides of b r i e f duration i n South A f r i c a (Reinecke, 1967) . I t i s also common i n the plankton of northern European waters, Greenland, and the Mediterranean ( S c h i l l e r , 1937). GENUS PROTOPERIDINIUM - TYPICAL TABULATION As f o r the thecate stage CYST OF PROTOPERIDINIUM SP. ./82 /82 Cyst Stage Cyst name: none Descriptive features: (Plate XI, f i g . 5) The body has a p e r i d i n o i d shape; there i s an a p i c a l horn and two antapical horns. The paragirdle i s not descending and i s defined by a f l a t b e l t - l i k e border around the midline of the cyst; The parasulcus i s indented deeply. The cyst wall i s two-layered. The inner layer i s smooth; the outer layer i s thrown i n t o a s e r i e s of folds and ridges to form a complex pattern of predominantly l o n g i t u d i n a l s t r i a t i o n s . This outer layer appears as though draped over the inner l a y e r . The archeopyle i s i n t e r c a l a r y , located on the dorsal side towards the apex. The shape i s very s i m i l a r to that of P_^  oblongum cysts ( P l . VII, f i g . 9). Size: Length - 50-61 um Width - 75-80 um N = 4 General remarks: This cyst has unknown p e r i d i n o i d a f f i n i t i e s . Although several v i a b l e cysts were i s o l a t e d from fresh Hidden Basin material, none su c c e s s f u l l y excysted. In treated samples they were seen r a r e l y i n /83 /83 Hidden Basin and one other northern i n l e t (#39, Topaze Harbour). GENUS SCRIPPSIELLA - TYPICAL TABULATION Indistinguishable i n the cyst CYST OF SCRIPPSIELLA FAEROENSE ( P a u l s e n ) B a l e c h & S o a r e s Cyst Stage Basionym: Peridinium faeroense Paulsen Synonym: Micrhystridium bifurcatum Williams References used: Dale, 1976: 59, PI. 1, f i g . 19. Dale, 1977: 241, f i g s . 30-35. Descriptive features: (Plate X, f i g s . 3, 6) The cysts range from s p h e r i c a l to oval. The processes have b i f i d t i p s ; there are occasional b i f u r c a t i o n s along the length of the pro-cesses while others are not b i f u r c a t e d . No excystment aperture was noted i n any of the cysts. S i z e : B.C. Dale (1976) Length - 24-33 pm (mean - 29 pm) diameter - 19-36 pm Width - 23-28 pm (mean - 25 pm) Processes - 2-6 pm N = 20 1-8 pm ./84 /84 General remarks: These cysts are d i s t i n c t i v e and were o r i g i n a l l y described as an a c r i t a r c h Micrhystridium bifurcatum (Dale, 1976). The cyst-theca r e l a t i o n s h i p has been documented through incubation experiments by Dale (1977). Although v i a b l e cysts of t h i s species from B.C. have excysted, a p o s i t i v e i d e n t i f i c a t i o n of the r e s u l t a n t theca i s not yet possible because the c e l l s are very small and they are very s i m i l a r to several other S c r i p p s i e l l a spp. These cysts survive palynological. treatment i n contrast to those of S c r i p p s i e l l a trochoidea (Stein) Loeblich I I I , which are s i m i l a r i n appearance but calcareous (Wall et a l . , 1970). GENUS SPIHIFERITES- TYPICAL TABULATION As f o r Gonyaulax SPINIFERITES BELERIUS Reid, 1974: 596, P i . 2, figs.' 12, 13 Cyst Stage Synonym(s): Pterosperma ovum Gaarder, 1954: 16, f i g . 14 References used: Reid,, 1974: l o c . c i t . Wall & Dale, 1968a: 270, P i . 1, f i g . 14. Descriptive features: (Plate X, f i g . 10) The body i s oval, sometimes nearly diamond-shaped, with an a p i c a l boss. The surface i s smooth and t h i n . Gonal processes are ./85 /85 t r i f u r c a t e with b i f i d t i p s , formed from parasutural septa. A high p o s t e r i o r "trumpet-like" process at the junction of 1" and 2' 1' i s c h a r a c t e r i s t i c . There may also be a high antapical flange. The para-cingulum i s displaced (descending, left-handed) one to three widths. Size: B.C. Reid Length - 35-45 pm (mean - 39 pm) 35-42 pm Width - 27-38 pm (mean - 33 pm) 28-37 pm Processes - 9-12 pm N = 11 7-10 pm N = 15 General remarks: These cysts are small, d e l i c a t e , and frequently misshapen i n samples from B.C. The presence of the t y p i c a l trumpet-shaped process a r i s i n g from the v e n t r a l paraplates 1" and 2 1'' i s considered to be diagnostic. Wall & Dale (1968a) i l l u s t r a t e a cyst they c a l l S^ bulloideus which gave r i s e to a c e l l resembling Gonyaulax scrippsae Kofoid. They state that S^ bulloideus varies i n i t s sculpturing from being "predominately spinous to e n t i r e l y membranous". Reid (1974) however, states that the Wall & Dale cyst i s S^ b e l e r i u s . Reid (1974) considers Pterosperma ovum Gaarder to be synonymous with S^ b e l e r i u s . However, Wall & Dale (1968a) believed P^ _ ovum to be a synonym of S^ elongatus Reid. ..../86 /86 I t seems that b e l e r i u s could be a membranous "end member" of a cyst complex a l l g i v i n g r i s e to the same thecate species, Gonyaulax  scrippsae Kofoid. S_^_ bulloideus (Deflandre and Cookson) Sarjeant and S. elongatus Reid are known to be part of t h i s complex. The dilemma of having several cyst taxa producing a sing l e thecate taxon i s causing confusion elsewhere ( i . e . with Gonyaulax s p i n i f e r a and G.  d i g i t a l i s ) . SPINIFERITES BENTORI Rossignol Cyst Stage References used: Wall, 1967: 298, f i g s . 1-5. Wall & Dale, 1967: 350, P l . 1, f i g . B. Wall, 1967: 101, P l . 14, f i g . 4, text f i g . 2. Harland, 1968: 542, f i g s . 3, 4, 6, 7, 14-18. Eisenack, 1971: 525. Wall & Dale, 1968a: 269, P l . 1, f i g . 2-5. Reid, 1974: 598, P l . 2, fig s . . 14-16, text f i g . 3. Morzadec-Kerfourn, 1977: 159, P l . 1, f i g s . 1, 2, P l . 3, f i g s . 1, 4. Descriptive features: (Plate X, f i g . 11, 12) The cyst body i s ovoid with a pronounced a p i c a l boss. Paraplate boundaries are marked by low parasutural l i s t s and gonal, and occa-s i o n a l l y parasutural processes. The processes have fenestrate bases ..../87 /87 and are t r i f u r c a t e with b i f i d t i p s . The parasulcus i s almost s t r a i g h t and widens s l i g h t l y p o s t e r i o r l y . Paraplate 6" i s tr i a n g u l a r ; l ' " ' i s long and t h i n , shaped as a poorly defined rectangle. The body wall i s smooth i n B.C. specimens, or i s l i g h t l y granular elsewhere. Size: B.C. Rossignol(1962) Reid(1974) Wall & Dale(1968a) Length - 50-64 p i 63- 73 p 58-69 p 48-55 p (dimension mean - 54 p not specified) N = 9 N = 15 Width - 40-53 p i 45-•60 p 52-55 p mean - 46 p Pro- 6-15 p i 15-•20 P 0-20 p 2-15 p cesses -St r a t i g r a p h i c range i s Pleistocene-Recent. General remarks: In the o r i g i n a l d e s c r i p t i o n 6" i s designated as being hexa- or pentagonal. Subsequently 6" was described as t r i a n g u l a r (Wall & Dale, 1968a; Harland, 1968) as i n the specimens found here. The thecate stage (Gonyaulax d i g i t a l i s ) has a t r i a n g u l a r 6" as w e l l . Paraplate 1' 1 1 i s small and l i n e a r (Wall & Dale, 1968a) which i s c h a r a c t e r i s t i c of the thecate stage as w e l l . Rossignol (1962) does not mention t h i s p l a t e i n the o r i g i n a l d e s c r i p t i o n . Harland (1968, f i g . 7, pg. 544) overlooked 1*'' and consequently, m i s i d e n t i f i e d the remaining paraplates i n the s e r i e s . Further, the area corresponding to the po s t e r i o r s u l c a l p late i s numbered as 6 , , , . Rossignol's •••./88 /88 tabulation appears to be correct. S. bentori from Brittany was shown to have processes with palmate ends by SEM rather than trifurcate ends with b i f i d tips as seen by the light microscope (Morzadec-Kerfourn, 1977, P l . 1, f i g . 1, 2). The fenestrate bases are used to distinguish this species from S^ ramosus but in the latter author's SEMs no well developed fenestrations are evident. These fenestrations are also not clearly seen in the local material. However, they are s t i l l recognizable by the "flanges" on the processes. Wall & Dale (1968a) have shown that these cysts give rise to motile cells which they called Gonyaulax d i g i t a l i s (Pouchet) Kofoid. T h i s cyst is relatively common only in Hidden Basin material, but occurs infrequently in a few other lo c a l i t i e s (27, 33, 37). Viable cysts were not found in Hidden Basin material although the species is known to occur in B.C. (Wailes, 1937). SPINIFERITES BULLOIDEUS (Deflandre & Cookson) Sarjeant Cyst Stage References used: Deflandre & Cookson, 1955: 264, p l . 5, figs. 3, Wall, 1965; 300, f i g . 6. Wall, 1967: 100, text f i g . 2. Wall & Dale, 1967: 352, P l . 1, f i g . K. Wall & Dale, 1968a: 271, P l . 1, f i g . 15. /89 /89 Harland, 1968: 544, f i g . 20 Eisenack, 1971: 533. Reid, 1974: 600, PI. 2, f i g s . 3, 17-19. Dale, 1976: 49, f i g . 4. Morzadec-Kerfourn, 1977: 159, PI. 1, f i g . 12; P i . 3, f i g . 6. General remarks: S. bulloideus was f i r s t described as being a small, d e l i c a t e rounded hystrichospherid cyst. Body diameter i s 30-37 pm with processes 10-15 pm long. Over time t h i s d e s c r i p t i o n has been modified so that now confusion has arisen between bulloideus and ramosus (Ehrenberg) Loeblich & Loeblich. The s i z e range for ramosus has been given at 30-50 um, 32-56 pm, and 34-41 pm depending on the l o c a t i o n and age of the sample (Davey et a l . , 1966). Harland (1968) distinguished S_^  bulloideus from ramosus on the basis of i t s smaller s i z e and s p h e r i c a l t e s t . bulloideus was c i t e d as approximately 35 pm i n diameter (some up to 40 pn) whereas S.  ramosus was 64 pm long, 65 pm wide with processes 16-20 pm long. Reid (1974) gives a very d e t a i l e d d e s c r i p t i o n of bulloideus. Unfortunately the diagnosis of ramosus i s much les s d e t a i l e d . The s i z e range f o r Reid's cysts i s 41-46 pn long and 19-27 um wide, with /90 /90 processes 8-16 um long. Wall (1967) provides a d e s c r i p t i o n of bulloideus and states that i t i s s i m i l a r to ramosus except that the t e s t i s l e s s than 40 um (dimension not specified) . Wall et_ a l . (1977) found bulloideus and Sj^ ramosus to be distinguishable by s i z e and also by other unspe-c i f i e d c h a r a c t e r i s t i c s . The d i s t r i b u t i o n o f these two cyst types i n various marine sediments also helped to separate them.. Davey & Rogers (1975) state that they could not d i s t i n g u i s h these two species other than by s i z e and thus i n t h e i r study c a l l e d a l l cysts with t h i s morphology, ramosus var. ramosus. The s i z e range was 30-40 pm iri diameter. In marine sediments from B.C., cysts of t h i s morphology have a s i z e range of 34-49 pm i n length and 28-43 pm i n width ( f i g . 8) which occurs within the stated range for both bulloideus and ramosus. On the basis of both d e s c r i p t i v e features and s i z e ranges, i t i s not possible to state unequivocably that the species occurring i n B.C. waters i s b u l l o i d e u s . To set a l i m i t of 40 pm as the maximum transdiameter of S. bulloideus and everything over that s i z e as S. ramosus i s a r b i t r a r y and a r t i f i c i a l l y cuts a population of cysts with a widely varying range of morphology including s i z e . Because of t h i s dilemma a l l species i n t h i s region are r e f e r r e d to as ramosus except for those few cysts which more nearly f i t the o r i g i n a l d e s c r i p t i o n . /91 /91 FIGURE 8 55-r E 3 c 25-25 Width in um 35 Cluster diagram showing length (in um) vs. width (in pm) of central body of S p i n i f e r i t e s ramosus from B.C. ./92 /92 SPINIFERITES ELONGATUS Reid, 1974: 602, P l . 3, f i g s . 23, 24 Cyst Stage References used: Wall & Dale, 1968a: 271, P l . 1, f i g . 16. Harland & Downie, 1969: 236, P l . 7, f i g . 4. Harland, 1973: 235, P l . 1, f i g s . 4-6. Dale, 19(76: 45, P l . 1, f i g . 8. Descriptive features: (Plate VIII, f i g s . 11, 12) An elongate oval cyst with wide f l a r i n g parasutural septa; the septa vary i n height from high f l a n g e - l i k e antapical processes to low gonal processes i n the paragirdle zone; the paragirdle zone i s des-cending (left-handed), displaced by one girdle-width; the parasulcus, p a r a l l e l to the l o n g i t u d i n a l axis, increases to three times i t s width p o s t e r i o r l y . Size: B.C. Reid (1974) Length - 50-59 um (mean - 54 pm) 40-59 pm Width - 29-38 um (mean - 33 um) 26-42 um Antapical pro- 6-15 um (mean - 12 um) 12-16 pm cess length -N = 25 N = 15 General remarks: Cysts of S^ _ elongatus occurred i n most of the treated samples although generally i n low numbers. In the Pendrell Channel sample (#31) /93 /93 these cysts form 18% of the cyst assemblage, the highest percentage of t h i s species attained i n any of the samples. Harland (1973) described t h i s species as a new species of S p i n i - f e r i t e s ( S p i n i f e r i t e s sp. nov.) although he d i d not name i t . His photographs c l e a r l y show the cysts to be S^ elongatus. Wall e_t a l . (1977) c l a s s i f i e d t h i s cyst species as temperate-estuarine as i t i s r e s t r i c t e d to those regimes. Dale (1976) stated that t h i s species appears to be r e s t r i c t e d to temperate (including c o l d water) regimes. Its d i s t r i b u t i o n i n the sediments of t h i s coast tends to confirm that i t can be used as an in d i c a t o r species for temperate-estuarine conditions. This cyst has been observed to give r i s e to a c e l l 'resembling' Gonyaulax scrippsae Kofoid (Wall & Dale, 1968a). However t h i s c y s t -theca r e l a t i o n s h i p has not been confirmed and remains ten t a t i v e . SPINIFERITES MEMBRANACEUS (Rossignol) Sarjeant Cyst Stage Basionym: Hystrichosphaera membranacea Rosignol References used: Wall, 1967: 102, P i . 14, f i g s . 14, 15. Eisenack, 1971: 559. Reid, 1974: 605, P i . 3, f i g s . 28-31. Morzadec-Kerfourn, 1977: 159, PI. 3, f i g . 5. /94 /94 Descriptive features: (Plate 10, f i g . 7-9) The cyst has an ovoid body; surface scabrate to microgranular with membranous su t u r a l septa. Gonal projections are t r i f u r c a t e while mar-g i n a l processes are b i f u r c a t e . This species i s characterized by a strong antapical flange at the juncture between paraplates 1"" and 4' 1 1. The g i r d l e i s displaced by two times i t s own width. The a p i c a l region has a crown of processes joined by high septa. The archeopyle i s dorsal 3", .quadrangular w i t h rounded corners. Size: B.C. Reid Rossignol (1962) Length - 35-44 um 34--44 p 57 p (mean - 41 p ) Width - 35-44 um 34--43 p 50 p (mean - 39 p ) Processes - 12-14 um 12--17 p 20--25 p (mean - 13 p ) N = 5 N = = 16 General remarks: This cyst was observed only o c c a s i o n a l l y i n Hidden Basin material and r a r e l y i n a few other locations,(6, 27, 31, 32). No archeopyle was seen i n any of the cysts. This species has a c h a r a c t e r i s t i c a ntapical flange and i n that respect i s s i m i l a r to S^ m i r a b i l i s (Rossignol) Sarjeant. The remainder of the processes are d i f f e r e n t i n the two species and supposedly makes /95 /95 them r e a d i l y d i s t i n g u i s h a b l e . In membranaceus the processes are weak and membranous and formed from the membranous septa d e l i m i t i n g paraplate margins. In S. m i r a b i l i s the processes are numerous, stout, and r i g i d . Nonetheless d i f f i c u l t y has been encountered i n d i s t i n g u i s h i n g these two species both i n material from t h i s region and apparently- i n English i n t e r g l a c i a l deposits examined by Harland & Downie (1969) (Reid, 1974). Most species found here with a strong antapical flange are c a l l e d S. membranaceus. I t i s possible that S^ m i r a b i l i s cysts were included with them. S. m i r a b i l i s cysts were not found i n B.C. sediments and since they are common i n Recent sediments (Wall, 1967) i t i s l o g i c a l to assume that t h e i r seeming absence may be due to m i s i d e n t i f i c a t i o n ( i . e . , they are counted i n with S^ membranaceus). membranaceus has not yet been li n k e d to any thecate species of d i n o f l a g e l l a t e • SPINIFERITES"SP. A " Cyst Stage Descriptive features: (Plate XI, f i g s . 1, 2, 4, Plate XII, f i g s . 2-8; Plate XIII, f i g s . 5-7) The body i s s p h e r i c a l to ovoidal with a pronounced a p i c a l boss; the wall i s microgranular ( P l . 12, f i g . 7). The processes vary from being long and s t r a i g h t to short and misshapen; they a l l have fene-s t r a t e bases ( P l . XIII, f i g . 5). Most appear to be t r i f u r c a t e with /96 /96 b i f i d t i p s . Some of these cysts have processes which remain s o l i d and slender along t h e i r length while others have hollow trumpet-like bases u n t i l branches occur, then the branches are slender and s o l i d with b i f i d t i p s (PI. XII, f i g . 6). Membranous septa u n i t i n g the processes are va r i o u s l y developed; others have membrane-like flanges j o i n i n g a l l the processes to give the appearance of a r u f f l e ( P i . XI, f i g . 1, 2, 4) along most of the parasutures; i n others they are low and more r i d g e - l i k e . The paragirdle i s narrow, edged by gonal t r i f u r c a t e processes with a few shorter b i f u r c a t e processes along the paraplate margins, descending (left-handed) and displaced two to three times i t s width. A l l the cysts of t h i s group are characterized by having an exceptionally large archeopyle. The operculum when present i s composed of two pieces. The archeopyle varies from being broad and f i v e - s i d e d to narrower and four-sided ( P i . XII, f i g . 2-8). E v i t t (1967) had a c l a s s i f i c a t i o n f o r such archeopyles, c a l l i n g them 2P, meaning they are composed of two precingular paraplates (see the d i s -cussion on archeopyles for further comments). Size: Diameter - 53-76 um (mean - 61 pm) Processes - 10-22 pm (mean - 15 pm) N = 12 General remarks: The cysts of t h i s general morphology are being dealt with as a ./97 /97 group for the following reasons: they are s i m i l a r morphologically; but the differences are great enough to d i s t i n g u i s h them i n t o several types. Whether they should be dealt with as a new genus having a number of species or whether they are merely a population of cysts showing a range of morphological v a r i a t i o n to be expected i n a varying environment i s unknown at t h i s time. Two cysts of t h i s morphology have been excysted.to give r i s e to Gonyaulax sp. 1 and sp. 2. One cyst of t h i s group resembles Hystrichosphaera dentata Gocht e s p e c i a l l y i n the development of the processes ( P l . XI, f i g . 1 , 2 ) . However, the archeopyle i s t y p i c a l f o r t h i s group. SPINIFERITES NODOSUM Wall, 1967: 101, P l . 14, f i g s . 7-9. Cyst Stage Descriptive features: (Plate IV, f i g s . 4, 5) These cysts are s i m i l a r to those described by Wall (1967). The body i s ovoid; paraplate areas are i d e n t i f i e d by low (1 um or less) parasutural septa and gonal processes. The processes are small, ei t h e r t r i f u r c a t e or b i f u r c a t e and curve strongly towards t h e i r own bases o r ij.e along the body surface to form a small pad or bump. ./98 /98 Size: B.C. Wall (1967) Length - 40-65 pm (mean - 49 pm) 31-62 pm Width - 33-56 pm (mean - 43 pm) 38-52 pm N = 5 General remarks: Rossignol (1964) described a taxon, Sj^ bentori var. truncata characterized by reduced processes which appear interrupted i n t h e i r development or as i f they had been severed. The only apparent d i f f e r -ence between S^ bentori var. truncata and S^ nodosum i s i n the s p e c i f i c appearance of the processes. In S. nodosum they do not appear severed but rather are misshapen and crushed. There are cysts i n fresh Hidden Basin material which f i t the des c r i p t i o n of both nodosum and S^ bentori var. truncata. There are also cysts resembling S_^_ elongatus Reid, but with reduced septa and spines. I t could be c a l l e d S^ elongatus var. nodosum. The cysts with reduced spines occur (in low numbers) with those with normal spines. S. nodosum was the only one of t h i s group found i n treated samples from other B.C. s i t e s (6, 17, 27, 31, 32, 37) although always i n very low numbers. Thecate Stage General remarks: Wall & Dale (1968a) found that cysts of S p i n f e r i t e s nodosum . . ./99 /99 gave r i s e to Gonyaulax d i g i t a l i s (Pouchet) Kofoid. This species i s known to a r i s e also from cysts of S^ b e n t o r i . (Wall & Dale, 1968a). Reid (1974) considered S_^_ nodosum a junior synonym of Sj_ bentori mostly because of the shared a f f i n i t y with Gj_ d i g i t a l i s but also because the only d i f f e r e n c e between them was i n the processes. Viable cysts of S^ nodosum have not been found i n B.C. coastal sediments. In f a c t , G\ d i g i t a l i s has not as yet been found to a r i s e from any cysts from t h i s region although the species has been reported (Wailes, 1937). SPINIFERITES RAMOSUS (Ehrenberg) Loeblich S Loeblich, 1966: 56-57. Cyst Stage Basionym: Hystrichosphaera ramosa Ehrenberg Synonym(s): Hystrichosphaera furcata (Ehrenberg) 0. Wetzel ... S p i n i f e r i t e s ramosus var. ramosus (Davey & Williams, 1966) References used: Davey et a l . , 1966: 33, P l . 1, f i g s . 1, 6; P l . 3, f i g . 1, text f i g . 8. Wall, 1967: 99, P l . 14, f i g . 1, 2. Harland, 1968: 540, f i g . 21. Harland & Downie, 1969: 232, P l . 7, f i g s . 3, 7. Wall & Dale, 1970: 49, P l . 1, f i g . 1-4, 6-15. /100 /100 Eisenack, 1971: 583. Harland, 1973: 231, P l . 1, f i g s . 9, 10. Davey & Rogers, 1975: 223, P l . 1, f i g . 5. Morzadec-Kerfourn, 1977: 159, P l . 1, f i g s . 3-7; P l . 3, f i g . 3. Descriptive features: (Plate X, f i g . 1, 2) Ovoidal c e n t r a l body,circular i n polar view; gonal processes are t r i f u r c a t e or b i f u r c a t e with b i f i d ends. Parasutural processes are b i f u r c a t e with b i f i d ends. Parasutural crests are o u t l i n e d by low ridges. 6' i s t r i a n g u l a r . The paragirdle i s descending (left-handed) approximately one width. The p a r a s u l c a l region i s weakly i n c l i n e d , narrow a n t e r i o r l y , broadening p o s t e r i o r l y , and does not indent the cyst w a l l . Size: B.C. Harland (1973) Davey et a l . (1966) Length - 34-49 pm (mean - 40 pm) 32-60 um (mean - 46.5 um) see below Width - 28-43 pm (mean - 34 pm) 30-50 um (mean - 39.8 um) Processes 11-22 um N = 20 11-30 pm N = 40 Barremain Cenomanian London Clay. Length - 34-41 pm 30-50 pm 32-56 pm Processes 5-13 pm 7-27 pm 11-20 um N = 2 N = 13 N = 9 ./101 /101 General remarks: This cyst has been a source of confusion because of i t s resem-blance to bulloideus (Deflandre & Cookson) Sarjeant. The o r i g i n a l drawing by Ehrenberg i s unfortunately i n l a t e r a l view. Also the o r i -g i n a l d e s c r i p t i o n , emended by Davey e_t a l . (1966) i s not as d e t a i l e d as the d e s c r i p t i o n given by Reid (1974) for b u l l o i d e u s . However a point by point comparison of these two cyst species leaves few chara-c t e r i s t i c s i n which they d i f f e r . Transdiameter i s the main c r i t e r i o n used by most authors to d i s t i n g u i s h the species (Wall, 1967; Harland, 1968; Wall et a l . , 1977) . In Reid's d e s c r i p t i o n the upper s i z e l i m i t f o r cysts of bulloideus i s over 40 pn, the c u t o f f point for most authors. Other authors (Davey & Rogers, 1975) have noted that the s i z e range of ramosus as o r i g i n a l l y described by Davey ejt a l . (1966) includes the s i z e range given f o r Sj_ bulloideus. Wall & Dale (1970) claim that a cyst s i m i l a r to ramosus gives r i s e to a c e l l r e f e r r a b l e to Gonyaulax sp i n f e r a , whereas bulloideus gives r i s e to G. scrippsae (Wall & Dale, 1968a). The S_^_ ramosus cyst giving r i s e to G^ s p i n i f e r a (Wall & Dale, 1970) i s s a i d to have a descending (left-handed) paragirdle of two widths. Wall (1967) however, describes S. ramosus as having a d i s -placement of s l i g h t l y more than one paragirdle width which i s s i m i l a r to the displacement seen i n Sj_ b u l l o i d e u s . The cysts from B.C. have /102 /102 one width of displacement and thus are not dis t i n g u i s h a b l e from S.  bulloideus on that b a s i s . However they are not i d e n t i c a l to the cysts of Wall & Dale (1970) e i t h e r . Size, paragirdle displacement, and other d e s c r i p t i v e features are inadequate to properly d i f f e r e n t i a t e these two species i n B.C. sediments. That they need to be distinguished i s obvious e s p e c i a l l y i n view of the f a c t that two d i s t i n c t thecate species emerge on excystment. As has been pointed out i n the S_^_ bulloideus general comments section, S^ ramosus i s the designation of a l l cysts of t h i s general morphology regardless of s i z e . GENUS TANYOSPHAERIDIUM - TYPICAL TABULATION Unknown i n the cyst TANYOSPHAERIDIUM SP. Cyst Stage References used: Davey et a l . , 1966: 98, PI. 6, f i g . 7; P i . 3, f i g . 3. Wall & Dale, 1968a: 281, PI. 4, f i g . 28 (as Po l y k r i k o s ) . Eisenack, 1971: 1021. Wall, Dale & Harada , 1973: 26, P i . 1, f i g . 23, 24. Dale, 1976: 51, P i . 1, f i g . 18 (as Polykrikos). ./103 /103 Reid & Harland, 1977: 169, P l . 2, f i g . 1-3 (as Polykrikos). Descriptive features: (Plate XI, f i g . 9; Plate XIV, f i g . 1-4) The cyst has an elongate c e n t r a l body, smooth endophragm, t h i n periphragm and forms the processes; i n the spaces between the pro-cesses the periphragm i s folded. The processes are c y l i n d r i c a l with connected i n a network by low ridges. In some cysts the process apices f l a t t e n out along the ridges to form a lacy f a n - l i k e network ( P l . XIV, f i g . 3, 4). The processes and ridges do not form an obvious pattern although they are regular. In the l i g h t microscope the c e n t r a l body appears to be s k i r t e d by a r u f f l e of processes which i s more or le s s developed (Pl. V, f i g . 1, 2). This e f f e c t i s not apparent, however, by SEM. The archeopyle i s a p i c a l . The opening i s somewhat ta t t e r e d as i f i t has been merely torn open ( P l . XIV, f i g . 4). An obvious oper-culum i s not v i s i b l e although one cyst has a f l a p present ( P l . XIV, blunt apices l i k e c a u l i f l o w e r heads ( P l . XIV, f i g . 1, 2) and are f i g . 2) . Size: B.C. Length - 50-88 um o v e r a l l Width - 44-56 pm o v e r a l l N = 6 ./104 /104 General remarks: Tanyosphaeridium has a f o s s i l record from the Hauterivian (Lr. Cretaceous) - Danian (Paleocene) and to date has not been reported from younger sediments. Specimens encountered here d i f f e r from the d e s c r i p t i o n of the genus i n the character and arrangement of the processes which are usually arranged i n a "more or les s c i r c u l a r manner around the c e n t r a l body" (Davey et a l . , 1966,.p. 98). Assignment of these modern cysts to t h i s genus i s therefore t e n t a t i v e . They may i n f a c t belong to a previously undescribed genus. However the cyst does bear close resemblance to Tanyosphaeri- dium tenerum Benedek (Eisenack, 1971: 1030a). The cyst i s common i n fresh Hidden Basin material but r a r e l y seen i n treated m a t e r i a l . I t occurs i n very low numbers i n those locations where d i v e r s i t y of cysts i s high. "Motile" Stage References used: Wailes, 1937: 13, f i g . 38. S c h i l l e r , 1937: 551 (vol. 1), f i g . 580a-c. Reid, 1975: 595, table 3. Dale, 1976: 59 PI. 1, f i g . 18. Descriptive features: The excysted c e l l i s d e l i c a t e , athecate, and colourless,from 50-102 pm long depending on the s i z e of the cyst; i n shape i t i s . . . / 1 0 5 /105 roundly oval, s l i g h t l y c o n i c a l at the apex; the g i r d l e i s c l o s e r to the apex than the base and divides the c e l l into approximately 1/3 -2/3 portions. The sulcus has not been c l e a r l y seen. General remarks: In the f a l l of 1976 these c e l l s excysted with ease and several p o l a r o i d photographs of the o f f s p r i n g were obtained. Since then excystment attempts have been unsuccessful. On the basis of these few photographs i t i s impossible to confirm that these cysts give r i s e to mature Polykrikos schwarzi B u t s c h l i as previously stated by Dale (1976) . The i n i t i a l excysted c e l l bears no resemblance to P.  schwarzi but whether i t matures into i t remains to be seen. PART IV DISCUSSION A. THE ROLE OF CYSTS IN LIFE CYCLES Many f o s s i l d i n o f l a g e l l a t e s were o r i g i n a l l y thought to be the remains of thecae. Many of the f o s s i l forms known today are remarkably theca-like e.g. P e r i d i n i t e s Lefevre. Later i t became c l e a r ( E v i t t , 1961; E v i t t & Davidson, 1964; Wall, 1967) that many of the bodies r e f e r r e d to as hystrichospheres were cyst remains of d i n o f l a g e l l a t e s . Even those resembling thecae were not thecae but cysts of vegetative c e l l s . ./106 /106 The r o l e of cysts i n the l i f e cycle of these organisms l e d to speculation. Were they for r e s t i n g or over-wintering? Were they a r e s i s t a n t stage? Or, were they a sexual stage? I t was and s t i l l i s important to discover t h e i r r o l e and the conditions under which they form. I f function and the f a c t o r s , environmental or otherwise, lead-ing to t h e i r formation are known, then understanding t h e i r d i s t r i b u -t i o n i n r e l i c t sediments becomes p o s s i b l e . Some of the f a c t s now known about l i f e cycles have been summarised i n the Introduction (Section C). Some freshwater d i n o f l a g e l l a t e s undergo sexual fusion to form zygotic cysts under conditions of nitrogen depletion ( P f i e s t e r , 1975, 1976, 1977). Marine Protoperidinium, however, are l a r g e l y non-photo-synthetic and thus are not amenable to such i n v i t r o manipulations. Cyst formation by many marine d i n o f l a g e l l a t e s i s e i t h e r unknown (Wall & E v i t t , 1975) or spontaneously occurs i n culture for reasons unknown (Braarud, 1945). Hystrichospherid-like cysts such as those produced by Gonyaulax s p i n i f e r a and others have never been found i n culture under any conditions. The excystment of c e l l s of G^ tamarensis from smooth, oval cysts from Hidden Basin sediments independently confirmed s i m i l a r observations of Dale (1977) and Anderson & Wall (1978). Sexual fusion i n t h i s species was found to be enhanced i n nitrogen-depleted cultures (Turpin et a l . , 1978). They form l a r g e , i n i t i a l l y somewhat lumpy zygotic cysts ./107 /107 which i n time are believed to come to resemble the cysts found i n natural environments. These v i a b l e cysts from sediments have been observed to produce a c e l l with two po s t e r i o r f l a g e l l a (Anderson & Wall, 1978) and are thus seen to be zygotic. The two independent studies on t h i s organism have thus c l a s s i f i e d the r o l e of p o t e n t i a l l y f o s s i l i z a b l e cysts i n i t s l i f e - c y c l e . In most marine d i n o f l a g e l l a t e s which are known to produce cysts, experimental proof of the ro l e of such cysts remains undetermined. Size of cysts and post-excystment c e l l s could i n f e r that they are zygotic. C e l l s which have ar i s e n from cysts from t h i s l o c a l i t y tend to be i n the upper siz e range f o r the species and sometimes even l a r g e r . tamarensis cysts and the post-excystment c e l l are much larger than the vegetative c e l l s (Turpin et a l . , 1978; Anderson & Wall, 1978). Meiosis i n d i n o f l a g e l l a t e s has been i n f e r r e d to occur e i t h e r within zygotic cysts ( P f i e s t e r , 1977) or i n the post-excystment c e l l (Anderson & Wall, 1978). In c e l l s which excysted here c e l l d i v i s i o n commonly occurred within one or two days of excystment, e s p e c i a l l y with P^ denticulatum and P^ _ pentagonum. These species are both non-photosynthetic and t h e i r e a r l y d i v i s i o n must be due to factors other than the surroundings i n t o which they excyst ( i . e . a drop of enriched seawater su i t a b l e f o r the growth of photosynthetic c e l l s o n l y ) . Most c e l l s did not appear to grow much. Thus the d i v i s i o n i s not l i k e l y to be the r e s u l t of the c e l l reaching the optimum s i z e for f i s s i o n to ./108 /108 occur. Because these post-excystment c e l l s tend to be somewhat large and also because of the tendency to divide under conditions which are les s than optimum for m i t o t i c d i v i s i o n , one might i n f e r that these c e l l s are zygotic. This i s an area for future research. B. THE ARCHEOPYLE A number of Protoperidinium species produce brown s p h e r i c a l cysts which are morphologically i d e n t i c a l to each other with l i t t l e s t r u c t u r a l d e t a i l . I d e n t i f i c a t i o n i s possible only when excystment occurs, since both the thecate c e l l and the cyst with an archeopyle are needed for i d e n t i f i c a t i o n . The species of Protoperidinium known to produce t h i s cyst are so f a r of two types: those with three a n t e r i o r i n t e r c a l a r y plates and those with only two anterior i n t e r -c a l a t e s . The f i r s t group contains P_^_ punctulatum (Wall & Dale, 1968a), P. conicoides (Wall & Dale, 1968a) and P. pentagonum. Their archeopyles are generally elongate hexagons corresponding i n shape to the second anterior i n t e r c a l a r y plate of the theca ( P i . I I , f i g . 2, 3). In P.  punctulatum plate 2a i s sometimes penta and the photograph by Wall & Dale (1968a, PI. 2, f i g . 27) shows the archeopyle resembling t h i s f i v e -sided shape. These are r e f e r r e d to as archeopyle type I ( E v i t t , 1967) . Often ./109 /109 the operculum remains attached at the cingular end. Type I archeo-pyles occur from the middle Jurassic-Recent. They occur i n Deflandrea, W e t z e l i e l l a and other le s s w ell known ones. A modern example i s P.  leonis ( E v i t t , 1967) . p^ _ leonis cysts are d i s t i n c t i v e , however, and are not part of the above group. P_^_ oblongum and P^ claudicans (Wall & Dale, 1968a) are believed to have type I archeopyles but i n r e a l i t y they are subapical and i n the case of V\_ claudicans i t can appear to by a p i c a l ( P l . VI, f i g . 3) although i t i s considered to be i n t e r -calary as are a l l the archeopyles of Protoperidinium known to date. Archeopyle shape can i d e n t i f y the thecal plate or plates to which i t corresponds because they both have s i m i l a r shape. The only thecal plate of these c e l l s which resembles a s l i g h t l y elongated hexagon i s p l a t e 2a; thus one can state that t h i s p l a t e corresponds to the archeo-pyle . The p e r i d i n o i d group with two anterior i n t e r c a l a r i e s i s larger than the f i r s t group. I t includes: P. aveliana (Wall & Dale, 1968a), P. denticulatum (Wall & Dale, 1968a), P_^  sp. nov., and P^ thorianum, The archeopyles produced by t h i s group are s i g n i f i c a n t l y d i f f e r -ent from those of the f i r s t group. They are roughly hexagonal and h o r i z o n t a l l y elongate. The upper edge i s more or le s s concave. This concavity varies between species of the group ( f i g . 9). These are type 21 archeopyles of E v i t t (1967) and correspond to the two anterior i n t e r c a l a r y plates of the theca. According to E v i t t (1967) the opera-./110 /no culum should be composed of two pieces but so f a r there has been no published evidence that these opercula are made up of more than one piece. The degree of f i t of the archeopyle to the two plates varies between species but i s constant within species. With P^ thorianum the shape corresponds c l o s e l y ; with P^ sp. nov. the f i t i s moderately good; with P^ denticulatum the f i t i s poor ( f i g . 9)•• The lower c i n -gular margin of these archeopyles appears as a s t r a i g h t l i n e whereas • the margin of the thecal plates i s angular. Thus the o u t l i n e of the two plates together has 10 sides and not seven as has the archeopyle. I t i s possible that the archeopyle of t h i s group of c e l l s corres-ponds to only one of the thecal p l a t e s , e i t h e r l a or 2a. However, the obvious concavity of the upper margin of the archeopyle precludes t h i s p o s s i b i l i t y more than the p o s s i b i l i t y of i t corresponding to two p l a t e s . E v i t t (1967) indicated (p. 29) that type 21 archeopyles are unknown i n the f o s s i l record a f t e r the Lower Cretaceous. However he admitted that h i s information may be incomplete. I t i s of i n t e r e s t to note that Protoperidinium aspidotum (Balech) Balech has an elongate hexagonal archeopyle (Fukuyo ejt a l . , 1977). The cyst, a round sphere, i s covered with short simple spines. The c e l l has two a n t e r i o r i n t e r c a l a r i e s and i t i s possible that the archeopyle of t h i s species i s a 21 even though the cyst i s d i f f e r e n t ./111 / I l l FIGURE 9 A Outlines of the i n t e r c a l a r y plates ( la & 2a) of several species are shown with the s t i p p l e d pattern. Superimposed i n heavy black i s the outli n e of the cyst archeopyle of each species. B Stippled outlines of i n t e r c a l a r y p l a t e (2a) of several species having three i n t e r c a l a r i e s are indicated. The cyst archeopyles of these species are superimposed. A B P. sp- nov. P. pentagonum P. thoria num P. c o n i c o i d e s P. den t i c u latum P. p u n c t u l a t u m P. asp idotum /112 /112 from the other 21 cysts already discussed. A group of Spiniferites cysts from this region appear to be unusual in the type of archeopyle they possess (Pl. XII, fig.2,5). 7, 8). The cysts have basically two types of archeopyles - one is rather broad and six-sided and the operculum is in two pieces (Pl. XII, f i g . 5, 7, 8; P l . XI, f i g . 1); the other is narrower, four-sided, with a two-piece operculum as well (Pl. XII, f i g . 2). According to Evitt (1967) this archeopyle type is a 2P and corresponds to the third and fourth precingular plates of the thecate species. The geo-chromologic range of 2P archeopyles is Upper Jurassic-Neogene (Evitt, . 1967) . Gonyaulacysta is one example. Bitectatodinium tepikiense Wilson, one of several Recent cysts giving rise to Gonyaulax spinifera has a large archeopyle which is probably 2P as well (Dale, 1976). The modern cysts from this region differ from each other in size and process development. The operculum often remains attached and is joined to the cyst wall along the apical edge. The two opercular pieces tend to stay together but can be dissociated quite readily. Two viable cysts of this type excysted and were designated Gonyaulax species 1 and species 2 (see results section). Evitt (1967) pp. 440, drew the possible thecal tabulation of a Gonyaulax c e l l with a 2P cyst archeopyle. Plate 6" is shown to be rather large and quadran-gular . The two species from cysts with 2P archeopyles here have a quandrangular 6" plate rather than the much reduced triangular 6" seen ./113 /113 i n Gonyaulax c e l l s with a P archeopyle. E v i t t (1967) was drawing a hypothetical case but i n t h i s instance i t f i t s the r e a l i t y very w e l l . The group of S p i n i f e r i t e s found here with t h i s unusual configura-t i o n needs c a r e f u l work to so r t out the number of new species or the morphological v a r i a t i o n of the group as a whole. Such an i n v e s t i g a -t i o n was beyond the scope of t h i s study. C. THE CYST - THECA RELATIONSHIP This i s not a complete treatment of t h i s subject as i t has been dealt with i n d e t a i l by other authors ( E v i t t , 1961; 1963/ 1976; Wall & Dale, 1968a). A p a r t i c u l a r d i n o f l a g e l l a t e species always produces the same cyst type(s). This p r i n c i p l e has not been v i o l a t e d i n any excystment experiments c a r r i e d out to date (Wall, 1965; Wall & Dale, 1968a; Wall & Dale, 1970) . However, d i f f e r e n t cysts can appear to produce the same thecate species. Gonyaulax s p i n i f e r a (Clap. & Lachm.) Dies, or c e l l s so s i m i l a r to i t that i d e n t i f i c a t i o n has been very d i f f i c u l t , has excysted from f i v e to s i x cyst types each of which has a separate pale o n t o l o g i c a l name. These are Bitectatodinium tepikiense Wilson, Nematosphaeropsis labyrinthus (Ostenfeld) Reid, Planinosphaeridium  membranaceus (Rossignol) Sarjeant, S p i n i f e r i t e s m i r a b i l i s (Rossignol) Sarjeant, and S_^  scabratus (Wall) Sarjeant (adapted from Dale, 1976). Cyst morphology has been presumed to be r e l a t i v e l y constant within /114 /114 species. Processes can grow a f t e r excystment has occurred; thus length of ornamental elements has not been a r e l i a b l e c r i t e r i o n for i d e n t i f i c a t i o n (Wall & Dale, 1968a). Whether process morphology i s subject to environmental e f f e c t s and thus i s highly v a r i a b l e i s unknown. Because spiny cysts of G. s p i n i f e r a , f o r example, have never been produced under laboratory conditions, the degree of v a r i a t i o n possible i n one cyst population r e s u l t i n g from one species i s unknown. Sometimes the c r i t e r i a used to e s t a b l i s h a new species of cyst are minor and p r o l i f e r a t i o n of new names needs to give way to a more sensible, more d i f f i c u l t c l a s s i f i c a t i o n of cysts where allowance i s made for population v a r i a t i o n . The p r i n c i p l e stated above, i . e . one species always produces the same type(s) of cyst, appeared to have been v i o l a t e d when Fukuyo et a l . (1977) described Protoperidinium minutum (Kofoid) Loeblich as having a cyst d i f f e r e n t from that described by Wall & Dale (1968a). Close examination of t h e i r accurate drawings showed that they had a c t u a l l y found P_^_ aspidotum (Balech) Balech. They are very s i m i l a r species but differences i n c e r t a i n key plates a i d i n i d e n t i f i c a t i o n (see the r e s u l t s s e c t i o n ) . P. pentagonum (Gran) Balech on t h i s coast has been found to produce a cyst d i f f e r e n t from that found by Wall & Dale (1968a). Careful examination of the thecal plate pattern of the B.C. species has confirmed that i t i s indeed P^ pentagonum judging by c e r t a i n key /115 /115 c h a r a c t e r i s t i c s . Balech (1949) describes a new species c a l l e d P. pentagonoides which i s very s i m i l a r to P^ pentagonum. P. pentagonoides i s generally a l i t t l e smaller," s u l c a l plates are d i s t i n c t i v e ; the g i r d l e i s without displacement; plate 4' i s shorter than 2'; plate 4" adjoins 1'; 2' adjoins both 1' and 2" making an unusual four-cornered suture. In p l a t e 2, f i g . 8 (Wall & Dale, 1968a). i t i s possible that the theca shown i s P^ pentagonoides. The g i r d l e does not show the d i s p l a c e -ment that i s t y p i c a l of P_^_ pentagonum and t h i s i s one of the h e l p f u l features to d i s t i n g u i s h between these two species. If the species on the east coast i s P_^_ pentagonoides then the basic p r i n c i p l e that one species produces the same cyst type(s) remains i n t a c t . However i f i t i s P^ pentagonum, then one species is_ producing d i f f e r e n t cyst types. One conclusion to be drawn from t h i s i s that there might be a P a c i f i c and A t l a n t i c population of c e l l s with s i m i l a r morphology but d i f f e r e n t evolutionary h i s t o r i e s . Another conclusion i s that cyst morphology i s not an immutable unchanging g e n e t i c a l l y determined character, but i s subject to environmental influences; thus one i s seeing P a c i f i c and A t l a n t i c populations of cysts where each type i s r e f l e c t i n g subtle environmental influences which might not have any e f f e c t on the morphology of the vegetative c e l l s . U n t i l i t i s known f o r c e r t a i n that the eastern population i s not P. pentagonoides, t h i s dilemma w i l l remain unresolved. ./116 /116 D. THE DISTRIBUTION OF CYSTS IN SEDIMENTS Wall e_t a l . (1977) discuss the factors c o n t r o l l i n g cyst d i s t r i -bution i n a l l the areas of t h e i r study. They note two trends; the f i r s t being an inshore-offshore trend, the second being a l a t i t u d i -n a l trend. Because a l l the s i t e s on t h i s coast are v i r t u a l l y i n the same l a t i t u d e the second trend w i l l be ignored. Most s i t e s f or t h i s study are i n estuarine environments so a s i g n i f i c a n t inshore-offshore trend i s not r e a l l y expected e i t h e r . They found some cyst species r e s t r i c t e d to offshore sediments, Leptodinium being an example. Leptodinium species have not been found i n any of the samples here. Other cyst species had propor-t i o n a l changes from nearshore to offshore, i . e . , species had peak abundances i n d i f f e r e n t areas. Factors which contribute to the trends i n cyst d i s t r i b u t i o n were divided i n t o b i o l o g i c a l and hydrodynamic s e t s . Hydrodynamic boundaries of water masses marked by temperature and s a l i n i t y changes can act as boundaries for cyst d i s t r i b u t i o n as w e l l . The hydrodynamic boundaries of the Middle A t l a n t i c Bight, f o r example, are useful i n explaining the d i s t r i b u t i o n of cysts i n that region. B i o l o g i c a l factors contributing to cyst assemblages are para-phrased below from Wall e_t a l . (1977) . They hypothesize that cyst /117 / l l 7 assemblages r e f l e c t the preference of c e r t a i n "species groups" f o r c e r t a i n types of water. Factors which cause species to p r e f e r cer-t a i n water types are a l l those b i o l o g i c a l parameters which define a niche for a species. F o s s i l i z a b l e d i n o f l a g e l l a t e s , i . e . , Gonyaulax, Protoperidinium, S c r i p p s i e l l a (some) are generally adapted to unstable estuarine-n e r i t i c environments while those that do not form cysts, e.g. Ceratium, are found i n more stable regions such as the open ocean. Thus cyst production may be seen as a l i f e - h i s t o r y strategy of adaptation to unstable, unpredictable regions. The cyst "end-types" species, i . e . , those d i s t i n c t cyst morpho-types with r e s t r i c t e d occurrence i n sediments, are seen as having become adapted to more l o c a l i s e d , more stable conditions. Therefore these cyst species p o s s i b l y o f f e r the most value i n paleo-ecological i n t e r p r e t a t i o n . Wall ejt a l . (1977) c l a s s i f i e d the marine environments of t h e i r study area into f i v e main d i v i s i o n s . Only a few of these d i v i s i o n s can be applied to the study s i t e s of t h i s coast. System A l i s defined as "cool-temperate, brackish, s t r a t i f i e d e stuaries" exemplified by fjords both here and i n Norway. System A2 i s "mild to cool temperate, low s a l i n i t y , moderately s t r a t i f i e d to v e r t i c a l l y homogeneous estuaries." An example i s Chesapeake Bay of the east coast of the United States of America. B l i s defined as "temperate, low to medium /118 /118 s a l i n i t y , seasonally s t r a t i f i e d subsystems", (more offshore) e.g. the Middle A t l a n t i c Bight. These three "environmental-climatic" d i v i s i o n s are the only ones po s s i b l y applicable to temperate l a t i -tudes and the conditions found i n most of the study s i t e s here. Some d i s t i n c t i v e cyst-based taxa were found to be r e s t r i c t e d i n t h e i r d i s t r i b u t i o n within temperate regimes. These were c l a s s i f i e d within the environmental-climatic d i v i s i o n s . d e f i n e d . e a r l i e r . There are only two cyst species having value i n paleo-ecological analysis which are found here, S p i n i f e r i t e s elongatus Reid and Sj_ bentori Rossignol (with short spines). Regions which were scoured by currents and tid e s (Table la) were very low i n cysts and unsuitable f o r the study of such micro-f o s s i l s . A l l these areas had fewer than 1 0 0 0 cysts/gm. The dominant cyst species was centrocarpum with fewer numbers of Protoperidinium spp., ramosus, and S c r i p p s i e l l a faeroense and more r a r e l y P T  conicum was found. These species were seen only i n those areas where there were more than 2 0 0 cysts/gm. Wall et a l . ( 1 9 7 7 ) do not have an environmental c l a s s i f i c a t i o n f o r such areas. I t i s obvious that fine-grained sediment p a r t i c l e s , i n c l u d i n g cysts and p o l l e n , are c a r r i e d away by such currents and tides to be deposited i n deeper, more t r a n q u i l locations (Dale, 1 9 7 6 ) . These regions of Table l a are b i o l o g i c a l l y s t r e s s f u l and are "high-energy" environments. 1 . BI ENVIRONMENTS Although the s i t e s i n the S t r a i t of Georgia (Table lb) bear . . . /119 /119 some resemblance to type A2 environments, they are much deeper. For that reason they w i l l f i r s t be considered as possible type Bl environments. The s i t e s from the S t r a i t of Georgia were character-i s t i c a l l y dominated by S p i n i f e r i t e s ramosus (Table IV), a species which was found by Wall et al.(1977) to be dominant ( or especially abundant) i n estuarine systems A3, and A4, and oceanic system D2 - a l l regions of warmer water than are found i n B.C. The S t r a i t sites also bore some resemblance to A2 environments. Table V summarises the cyst assemblages i n such s i t e s . The cyst assemblages found to be associated with A2 environments i n the A t l a n t i c regions are d i s s i m i l a r to the S t r a i t of Georgia samples much as the Bl assemblages are; i.e. the dominant species d i f f e r as do the r e l a t i v e values of the common and rare species. The common and rare species found i n the S t r a i t occurred i n B l and Al environments of the A t l a n t i c but their r e l a t i v e importance varied. Sc. faeroense i s not mentioned i n any of their samples as a cyst of importance. However i t was common i n many si t e s here and was a dominant species i n one Hidden Basin sample ( see Table l e ) . This .. ./120 /120 TABLE IV Wall et a l . (1977) B.C. (1978) Dominant: 0. centrocarpum *S. ramosus Common: *S. bulloideus 0. centrocarpum S. mirabilis Protoperidinium spp. P. conicum Sc. faeroense P. pentagonum S. elongatus Rare: Nematosphaeropsis balcombiana S. membranaceus Tectatodinium pellitum Wall * The S. ramosus found in B.C. is probably the same cyst species designated as S^ bulloideus on the east coast (see the results section for comments). S. ramosus thus has a comparable cyst distribution to S_^  bulloideus as i t is defined elsewhere. Cyst assemblages associated with Bl environments - "temperate, low to medium salinity, seasonally s t r a t i f i e d ..." (Wall et a l . , 1977) • /121 /121 TABLE V Dominant: Wall et a l . (1977) 0. centrocarpum or grouped P^ spp. B.C. (1978) S. ramosus Common: P. conicum 0. centrocarpum S. elongatus P. spp. S. mirabilis Sc. faeroense, S. elongatus Rare: S. bulloideus S. membranaceus • A comparison of cyst assemblages associated with A2 environments and the Strait of Georgia sites (Table lb). ./122 /122 TABLE VI Wall et a l . (1977) B.C. (1978) Dale (1976) Dominant 0. centrocarpum 0. centrocarpum Sc. faeroense Common S. bulloideus Protoperidinium sp. (incld. grouped P. spp.) P. conicum P. pentagonum Protoperidinium spp. 0. centrocarpum S. cf. ramosus (Harland, 1973) Rare: N. balcombiana T. pellitum S. ramosus P. conicum S. elongatus S. elongatus N. balcombiana Specimen density 3,416 - 16,106 0 - 4,050 1000 - 21,252 Cyst assemblages associated with Al environments - "cool-temperate, brackish, s t r a t i f i e d estuaries." Wall et a l . , 1977 /123 TABLE VII Wall et a l . (1977) B.C. (1978) Dominant: O. centrocarpum or 0. centrocarpum, a few with Proto-grouped J?. spp. peridinium spp. Common: P. conicum S. elongatus S. mirabilis S. ramosus Sc. faeroense Rare: S. bulloideus S. elongatus Cyst assemblages associated with BI environments - "temperate, low to medium salinity, seasonally s t r a t i f i e d ..." (Wall et a l . 1977). ./124 /124 cyst was important i n Trondheimsfjord of Norway (Dale, 1976). This cyst, o r i g i n a l l y thought to be of the genus Impletosphaeridium Morgenroth by Reid (1975) has been found to be dominant i n S c o t t i s h sea lochs as well (Reid & Harland, 1977) . N. balcombiana and T_^_ p e l l i t u m have not been found i n any of the study s i t e s of t h i s region. Hydrodynamic factors which influence cyst assemblages of the Middle A t l a n t i c Bight such as shoreward incursions of warm oceanic water and upwelling do not occur i n the S t r a i t of Georgia because of the presence of Vancouver Island to the west. The general c i r c u l a t i o n pattern of the S t r a i t i s counter-clockwise with seawater entering from the south through Juan de Fuca S t r a i t ( f i g . 2) and from the north around a l l the small islands located there. The Fraser River i s a major contributor of freshwater so that net flow i s outward to the P a c i f i c (Waldichuk, 1957). The S t r a i t i s enclosed by islands both to the south and the north and thus i s somewhat l i k e a b i g bowl without the large oceanic connection of the Middle A t l a n t i c Bight. The d i f f e r -ences i n the hydrodynamic patterns between these two s i t e s i s s u f f i -c i e n t to explain the differences i n cyst assemblages i n both areas and to preclude putting them i n the same category. The S t r a i t samples, e s p e c i a l l y when examined through t h e i r cyst assemblages, are not of A2 environmental type e i t h e r although hydro-dynamically they are s i m i l a r . I t would seem that these S t r a i t s i t e s are s p e c i a l and require an environmental-climatic c l a s s i f i c a t i o n not /125 /125 defined by Wall et_ a l . (1977) . 2. A l ENVIRONMENTS The f j o r d s of t h i s coast are environmentally s i m i l a r to the fjor d s of Norway. Fjords of B.C. have a c h a r a c t e r i s t i c configura-t i o n ; they are long and narrow and tend to zigzag. There i s a s i l l near the entrance and the water behind the s i l l i s deep, deeper than the regions outside the s i l l . Fjords and longer i n l e t s such as Indian Arm (#4, #5) have low s a l i n i t i e s at the surface. There i s a "lens" of freshwater present which i s most pronounced with the spring runoff. This layer of low s a l i n i t y water i s deeper and less s a l i n e at the head of the i n l e t but with entrainment becomes shallower and s a l t i e r towards the entrance. Below f i v e meters t h i s e f f e c t i s not apparent. Bute and Knight Inlets plus Howe Sound are a l l fed by g l a c i a l melt water and as a consequence the water at the head i s milky from g l a c i a l " f l o u r " . This opacity has a strong e f f e c t on phytoplankton i n that i t i s v i r t u a l l y o b l i t e r a t e d i n such areas (Stockner et a l . , 1977). The reduced phytoplankton plus a high sedimentation rate from the s i l t - l a d e n water may adequately explain the lack of cysts i n the sediments at the head. A gradient of cyst numbers increasing towards the entrance of a f j o r d has been noted elsewhere (Dale, 1976). The f a c t that the head i s shallower and that cysts and p o l l e n acting as fine-grained sediment p a r t i c l e s accumulate i n deeper waters has ./126 /126 also been noted before (Dale, 1976) . Fjords, as environmental-climatic d i v i s i o n type A l , are associated with a p a r t i c u l a r cyst assemblage (Table VI) . P. pentagonum cysts on t h i s coast are included with Protoperidinium spp. cysts. The cysts of t h i s species from the A t l a n t i c are d i s -t i n c t and also d i f f e r e n t from the cysts of t h i s species here. S.  elongatus i s considered to be rare here; Dale (1976) records i t s presence i n Trondheimsfjord but Wall ejt a l . (1977) apparently do not consider i t as part of t h e i r f j o r d assemblages. Protoperidinium spp. i s designated here as including a l l Proto- peridinium cysts except P^ conicum. This d e f i n i t i o n d i f f e r s from that of Wall e_t a l . (1977) . Their group of Protoperidinium spp. included only those cysts which were morphologically s i m i l a r spheres. The dominance of 0^ centrocarpum i s t y p i c a l for many temperate regions besides fjords (Harland, 1973; Reid, 1974). Dale (1976) found Sc. faeroense to be the dominant species i n Trondheimsfjord. The f j o r d s of t h i s region appear to have fewer species than one would expect but those that are present are s i m i l a r to those from Norwegian fj o r d s (Dale, 1976; Wall et a l . , 1977). Specimen d e n s i t i e s here were much lower than those seen elsewhere. Dale (1976) rejected any samples he found with le s s than 1000 cysts/gm as being unsuitable. In the fj o r d s here the s i t e s were analysed /127 /127 whether they had large numbers of cysts or not and also regardless of the percentage of sand i n them. If a l l samples with les s than 1000 cysts/gm had been rejected only one sample would have been acceptable (Je r v i s , #46). Future work i n t h i s region can at l e a s t begin by avoiding the upper portions of B.C. fjords as sample s i t e s of r i c h cyst assemblages. 3. A2 ENVIRONMENTS Indian Arm (#4, #5) has the c i t y of Vancouver near i t s entrance. The shores of t h i s i n l e t are more populated than many of the other i n l e t s and t h i s i s ind i c a t e d i n the p o l l e n assemblage found i n the sediments; there i s more angiosperm p o l l e n than i s seen i n more i s o -l a t e d areas. S a l i n i t i e s f o r Indian Arm are a l i t t l e lower than e l s e -where, being about 27%o from f i v e meters to the bottom. There i s a "lens" of freshwater on the surface i n the spring runoff much l i k e that found i n f j o r d s . Many of the A2 environments (Table Id) are r e l a t i v e l y uninhabited i n l e t s . The p o l l e n i n the sediments from these areas i s a character-i s t i c North P a c i f i c c o n i f e r f o r e s t assemblage (Dr. G. Rouse, pers. comm..). A few have logging companies operating along t h e i r shores. Topaze (#40) and Forward (#39) Harbours are both used as booming grounds. The amount of r i v e r runoff a f f e c t i n g these northern i n l e t s i s highly v a r i a b l e . Indian Arm i s moderately high whereas Pendrell Sound .../128 /128 (#31, #32) i s low. Forward Harbour has none. Hidden Basin represents an unusually stable marine environment; t h i s s t a b i l i t y i s r e f l e c t e d i n the great numbers and d i v e r s i t y of the cyst assemblage which i n turn must be a r e f l e c t i o n of the same trends i n the phytoplankton. The hypothesis advanced by Wall e_t a l . (1977) that the most s p e c i a l i z e d cyst morphotypes are l o c a l i z e d i n the most stable environments i s supported by the evidence i n Hidden Basin. This area, more than any other, had not only the most diverse group of cysts but also had more cyst species that d i d not occur anywhere e l s e . The unusual S p i n i f e r i t e s cysts with the two-part opercula, f o r example, were seen only r a r e l y i n other s i t e s , but i n Hidden Basin they achieved the greatest morphological v a r i a t i o n . Nearly a l l A2 s i t e s on t h i s coast had cyst assemblages dominated by centrocarpum and i n several areas such as Station #5, #34, t h i s cyst species was an overwhelming dominant. (Table VII) The study s i t e s here comparable to A2 environments elsewhere, tend to have s i m i l a r species but t h e i r r e l a t i v e importance d i f f e r s , a s i t u a t i o n found f o r A l and BI environments as w e l l . Also here more than i n other environmental zones there are notable absences of expected cyst-based taxa such as m i r a b i l i s and Bitectatodinium  tepikiense. The dominant cyst species are s i m i l a r f or A2 environments of both coasts; the common ones d i f f e r . S^ elongatus, although seen i n many /129 /129 samples here never became as abundant as i t d i d f o r some east coast samples. The maximum achieved here was 18% i n Pendrell Sound (#34) whereas i n Somes Sound, Maine i t reached 38% of the assemblage. This species i s considered to be rare here. P. conicum was seen very r a r e l y i n a l l the A2 samples from t h i s coast. Sc. faeroense cysts were commonly seen i n a l l s i t e s and i n one sample (Hidden Basin, #28) i t was the dominant species. This p a r t i c u l a r cyst species i s not discussed by Wall et a l . (1977). S^ m i r a b i l i s has not yet been seen i n samples from t h i s coast. Hidden Basin was sampled several times over the two-year study period. The absolute cyst numbers varied greatly between samples and the dominant species d i f f e r e d three out of four times. The v a r i a t i o n seen i n absolute cyst counts i s expected as a r e s u l t of the techniques used to prepare the samples. I t has also been shown (Dale, 1976) that p a l y n o l o g i c a l chemicals are p a r t i c u l a r l y destructive of Protoper- idinium cysts, one of the s i g n i f i c a n t cyst groups from t h i s s i t e . Because i t i s unknown as to which season or even which year the cysts formed i n each Hidden Basin sample, the s i g n i f i c a n c e of the change i n the dominant cyst species also remains unknown. The sample method made I t d i f f i c u l t to be c e r t a i n that the freshest phytoplankton d e t r i t u s was being sampled each time. The f l o c c u l e n t layer which accumulates on the bottom of the i n l e t was sampled at the bottom, a portion which was anoxic and which had po s s i b l y been subjected to greater disturbance by b i o l o g i c a l factors such as grazing by c i l i a t e s /130 /130 or r e c y c l i n g by b a c t e r i a than the top part of the layer. These factors may also be the reason the amount of carbon was much less than expected. The age of the sampled portion i s uncertain as well; whether the layer i s conserved year a f t e r year or renewed every year i s unknown. The carbon content of plankton debris from Theodosia I n l e t was the highest of a l l the sample s i t e s . However the carbon content i n the consolidated sediment (#30) was more l i k e that found i n other s i t e s . I t i s l i k e l y that the large amount of woody plant debris i n the plank-ton samples was a f a c t o r i n t h e i r high carbon content. A l l samples from t h i s i n l e t had very low cyst numbers, an unexpected r e s u l t i n view of the high carbon content found there. The dominant cyst species was centrocarpum. Examination of fresh material showed that there were large numbers of small spiny calcareous cysts much l i k e those produced by S c r i p p s i e l l a sweenyi Balech ex Loeblich (Wall et a l . , 1968b) and Sc. trochoidea (Stein) Loeblich I I I (Wall tat a l . , 1970). Being calcareous these cysts do not survive immersion i n HCl,one step i n the routine p a l y n o l o g i c a l treatment of these samples. Both samples from Gorge Harbour were dominated by CK_ centrocarpum, a r e s u l t consistent with the other A2 environments of t h i s coast. I t i s to be expected that the east sample within the basin (#27) has larger cyst numbers than the west (#26) because i t i s deeper. The values f o r sand content and carbon content support the p o s s i b i l i t y that the fin e sediment i s moving to the deeper portions of the i n l e t . /131 /131 In conclusion one can say that most of the s i t e s of t h i s coast f a l l i n t o the A2 category as defined by Wall et a l . (1977) . In comparison to the east coast the cyst assemblages associated with t h i s environmental type i n B.C. d i f f e r i n r e l a t i v e numbers and impor-tance of the various species, although the i d e n t i t y of the species i s s i m i l a r (only one species seems to be missing). E. THE SPINIFERITES/OPERCULODINIUM RATIO Davey & Rogers (1975) found that 0^ centrocarpum cysts appeared to be associated with the westward flowing warm-water Aghulus current whereas S_^_ ramosus appeared to be associated with the cold-water Benguela current o f f the west coast of southern A f r i c a . Wall et a l . (1977) disagree with the above authors. They believe that a high r a t i o of Operculodinium over S p i n i f e r i t e s cysts indicates a n e r i t i c rather than estuarine influence at the time that the cysts were formed. They further state that the r a t i o has no r e a l e c o l o g i c a l s i g n i f i c a n c e but may have more s i g n i f i c a n c e i n r e l a t i o n to the o r i g i n of sediments: "a high r a t i o r e f l e c t s d e r i v a t i o n of ... f i n e - g r a i n e d material from n e r i t i c d epositional or paleo-depositional s i t e s whereas a low r a t i o r e f l e c t s d e r i v a t i o n of cysts from paleoestuarine s i l t s and c l a y s " . The term n e r i t i c i s usually applied to regions of a c o a s t l i n e with l e s s than 200 meters depth. Estuarine environments are thus a sub-category of n e r i t i c environments. Wall et a l . (1977) use these /132 /132 two words, estuarine and n e r i t i c , as though they were separate and therefore equal. The use of "estuarine vs. n e r i t i c influences" i s d i f f i c u l t to understand i n view of the f a c t that the meaning of these two terms i n that context i s somewhat obscure. In t h i s study area Operculodinium cysts tended to be the domi-nant species i n most s i t e s although there were a few exceptions to t h i s , notably a l l the s i t e s located i n the S t r a i t of Georgia. Operculodinium here had a d i s t r i b u t i o n not unexpected i n view of what i s known of i t s d i s t r i b u t i o n on the east coast of North America. Centres of d i s t r i b u t i o n were found for i t i n A l , A2 and BI environ-ments i n the east coast samples. I t i s s u r p r i s i n g that i t i s not a dominant species i n the S t r a i t samples. S. bulloideus (= S^ ramosus here) had centres of d i s t r i b u t i o n mainly i n estuarine environments A2, A3, A4, A5, and A8, A2 being the only one of these found here. This species was not a dominant one i n A l environments (fjords) e i t h e r here or on the east coast, although the species i s present i n such areas, i n low r e l a t i v e numbers. I t i s usually not important i n coastal systems (such as BI environments). The dominance of S^ ramosus i n the S t r a i t of Georgia samples may be an i n d i c a t i o n that these areas are more estuarine than n e r i t i c , i . e . more A2 than BI. However the cyst assemblage of these s i t e s i s not much l i k e those of A2's e i t h e r . I t seems that on the basis of i t s cyst assemblage the S t r a i t of Georgia should have a separate environ-./ 133 /133 mental c l a s s i f i c a t i o n besides those defined by Wall ejt a l . (1977) i n t h e i r study. PART V SUMMARY AND CONCLUSIONS A. EXCYSTMENT EXPERIMENTS There have been ten cyst-theca r e l a t i o n s h i p s i d e n t i f i e d here confirming reports from elsewhere. Cysts of Gonyaulax tamarensis from B.C. waters were excysted. Protoperidinium aspidotum was excysted from cysts s i m i l a r to those found by Fukuyo e_t a l . (1977) , who mis-i d e n t i f i e d the species as P^ minutum. Most of the other excysted c e l l s were Protoperidinium spp. and the cyst-theca r e l a t i o n s h i p i s unchanged from that found by Wall & Dale (1968a). These are P^ _ c l a u - dicans , P. conicoides, P. conicum, P. denticulatum, P. l e o n i s , P. oblon- gum, and P^ punctulatum. Zygabikodinium lenticulatum was the only cyst-theca r e l a t i o n s h i p confirmed here for those cysts whose archeo-pyle i s a s p l i t around the paragirdle region. Five new cyst-theca r e l a t i o n s h i p s from B.C. were established. Two Gonyaulax species excysted from new cysts. Both c e l l s and cysts have not been previously described i n the phytoplankton or sediments of t h i s coast. P e r i d i n i o p s i s c f . hainanensis was excysted on several occasions. I t i s a species previously unrecorded from the phyto-plankton here. Protoperidinium sp. nov. i s a species not described ./134 /134 before. thorianum i s known to occur i n the phytoplankton but i t s cyst stage was previously not known. The cyst found here to give r i s e to pentagonum on t h i s coast i s not the same as the cyst found to give t h i s species from the east coast. B. CYSTS IN SEDIMENTS Fo r t y - f i v e sediment samples were c o l l e c t e d from various s i t e s along the coast of B.C. Of these 23 were very low i n cysts and of no p a l e o n t o l o g i c a l value. Seven of the 45 were repeated samples of two loca t i o n s , Hidden Basin (four samples) and Theodosia I n l e t (three samples). Hidden Basin was the s i t e of fresh d e t r i t a l material for the excystment experiments. There are ten cyst species described from the sediments of t h i s coast which have been found elsewhere i n Recent sediments. These are Operculodinium centrocarpum, Sc. faeroense c y s t , S p i n i f e r i t e s  b e l e r i u s S. b e n t o r i , S. bulloideus, S. elongatus, S. membranaceus, S. nodosum and S^ ramosus. Tanyosphaeridium sp. has been recorded i n sediments from elsewhere but as the cysts of Polykrikos. In t h i s study i t i s t e n t a t i v e l y assigned to a f o s s i l genus because i t f i t s the d e s c r i p t i o n c l o s e l y . There were two new cyst species described from t h i s region which have not been described elsewhere. In both cases a thecate stage /135 /135 i s unknown for them. These are Protoperidinium sp. nov. (cyst), and S p i n i f e r i t e s "sp. A". The cysts found i n sediments of t h i s region are commonly found i n many areas o f temperate estuarine and n e r i t i c conditions (Davey & Rogers, 1975; Dale, 1976; Harland, 1974; Wall et a l . , 1977). The cyst assemblages are s i m i l a r to those of Quaternary s e d i -ments of the A t l a n t i c . The number of species appears to be lower than expected. Some cysts which should be present i n these temperate assemblages have not yet been found. In many cases the thecate stage i s known to occur i n the phytoplankton. In some cases common cysts here are apparently rare species on the east coast. Differences i n r e l a t i v e numbers of cyst types i s less s i g n i f i c a n t than the presence or absence of some. The major feature of cyst assemblages here i s the dominance, of Operculodinium centrocarpum. This pattern i s t y p i c a l f o r temperate estuarine areas. The presence of such cyst assemblages from the N.E. P a c i f i c i s further confirmation of patterns already established f o r temperate estuarine conditions on the east coast of North America and Europe (Wall et. a l . , 1977) . Dale (1976) found s i m i l a r c y st assemblages i n Trondheimsfjord of Norway but also found an unknown cyst now known to be that of Scripp- s i e l l a faeroense. This cyst i s s i g n i f i c a n t not only i n fjords of /136 /136 Norway and sea lochs of Scotland, but also i n the fjords and some sheltered i n l e t s of t h i s coast. ./137 /137 PLATE I Figure 1-4 Peridiniopsis cf. hainanensis; 1. ventral view; 2. dorsal view, tubular connection between these and substrate i s illustrated; 3. apical view, plate 5" has an angularity (arrow) such that i t looks like i t might give rise to an intercalary plate but has failed to do so; 4. antapical view. (Bar = 50 um for a l l f i g s . ) . Figure 5 Protoperidinium claudicans. The divergence of the antapical horns i s exaggerated. Figure 6-8 P. conicum, 6. aberrant cyst. The operculum, in two pieces, i s shown slightly displaced within the cyst; 7. dorsal view of theca; 8. ventral view, c e l l i s elongated. Figure 9-11 P. aspidotum. 9. ventral view, unequal plates 1" and 6" are indicated; 10. dorsal view. 11. apical view, unequal intercalary plates are indicated. Figure 12, 14, 15 P_^_ cf. denticulatum. 12. ventral view, one c e l l pair. The sulci and cytoplasmic inclusions help to confirm the apex to base mode of attachment (shape somewhat inaccurate); 14. antapical ./138 /138 view; 15. apical view. Plate 3' is more rounded and larger than the same plate in the original drawings. Plate 4" plus one girdle plate is missing. Figure 13 Figure 16 Figure 17-19 P. leonis. ventral view. P. oblongum, ventral view. P. thorianum. 17. ventral view; papillate sculpturing is indicated on one plate; 18. apical view; 19. the cyst. The operculum i s attached, inner hyaline envelope is indicated (arrow). ./139 /140 PLATE II Figure 1-5 Protoperidinium pentagonum; 1. ventral view; 2. dorsal view; 3. cyst with elongated archeopyle and attached operculum; 4. apical view; S. antapical view. The posterior sulcal plate (sp) homologue is smaller than the same plate figured by Lebour (1925) . (Bar = 20 um) Figure 6-13 P^ sp. nov. 6. ventral view; 7. dorsal view showing the unequal intercalary plates; 8. apical view; 9. right oblique view; 10. l e f t side view; 11. oblique dorsal apical view; 12. the sulcal plates; sa - anterior sulcal, sd -right sulcal, sm - median sulcal; sp - posterior sulcal; as - l e f t sulcal; spa - posterior acce-ssory; t - transitional; 13. the cyst with the oper-culum attached. (fig. 6-11, Bar = 20 um: f i g . 12, Bar = 10 um) Figure 14-17 Zygabikodinium lenticalatum; 14. apical view. Note the small diamond-shaped 2a plate. The thecal pores are relatively conspicuous in this species; 15. antapical view. There is only one large anta-pical plate; 16. ventral view; 17. dorsal view. (Bar = 20 um) /141 141 /142 PLATE III Figure 1, 2 Protoperidinium thorianum. 1. aberrant c e l l , v e n t r a l view showing misshapen 1' and broad s u l -cus; 2. dorsal view (reversed) showing p l a i n and p a p i l l a t e p l a t e s . Figure 3-6 Gonyaulax species 2. 3. v e n t r a l view, extra p l a t e (la or 5') i s indicated (arrow); 4. dorsal view; 5. i s o l a t e d p l a t e with thecal patterning; 6. a p i c a l view. Figure 7-9 P. punctulatum. 7. v e n t r a l view; 8. dorsal view. Missing plates have been dotted i n ; 9. a p i c a l v i e w . Figure 10,11 G. species 1. 10. a p i c a l view with 4' indicated; 11. v e n t r a l view. The arrow indicates the extra p l a t e ( l a ) . Quadrangular 6" i s indicated; l i n e a r l ' 1 ' 1 i s indicated. (Bar = 20 pm for each species e x c l . f i g . 5). ./143 143 /144 PLATE IV Figure 1 Figure 2 Line drawing of S p i n i f e r i t e s "sp. A" cyst, r i g h t l a t e r a l view. The p a r a s u l c a l region can been seen as well as t r i a n g u l a r paraplate 6". Stippled areas are obscuring debris. (Bar=20um f o r a l l f i g s . ) Same cyst as i n figure 1, l e f t l a t e r a l view plates 2" and 3" are v i s i b l e . Figure 3 Figure 4, 5 Unusual cyst of unknown a f f i n i t i e s seen r a r e l y i n a few northern i n l e t samples. This possibly i s a cyst reworked from e a r l i e r sediments. I t stained very darkly with the safranin. S p i n i f e r i t e s nodosum; l e f t and r i g h t l a t e r a l views. The processes are much reduced gonal nubs of material. ./145 1 4 5 /146 PLATE V Figure 1, 2 Tanyosphaeridium sp. ; cyst of ?Polykrikos schwarzi. Two d i f f e r e n t examples showing varied development of the median r u f f l e - l i k e processes. The ridges on the cyst w a l l are not i n any apparent pattern. (Bar = 20 um) ./147 147 /148 PLATE VI Figure 1-3 Figure 4, 5 Protoperidinium claudicans; 1. the c e l l i n dorsal view; 2. the cyst recently excysted to give the c e l l i n f i g . 1. The subapical archeopyle with attached operculum i s v i s i b l e ; 3. cyst from palyno-l o g i c a l l y treated material. The archeopyle appears completely a p i c a l i n t h i s cyst. (Bar = 20 um i n a l l fi g s . ) P. thorianum; 4. v e n t r a l view. P a p i l l a e on the theca are prominent; 5. dorsal view with two i n t e r c a l a r y plates (one i s missing). Figure 6-8 P. sp. nov.; 6 ventral view; 7. oblique view of r i g h t hypotheca; l e f t and p o s t e r i o r s u l c a l plates (ss, ps) are e a s i l y seen; 8. r i g h t l a t e r a l view. Figure 9, 10 P. punctulatum; 9. ventral view. Most of the epitheca i s l o s t when the c e l l ecdyses; 10. v e n t r a l hypotheca view. Figure 11 Figure 12 P. conicoides, ventral view. P. leonis showing obvious aberration at the g i r d l e region of p l a t e 1' (arrow). This aberration was found i n only one specimen. Zigzag suture between 1* & 2" and 1" & 2" i s v i s i b l e ( a r r o w ) . /149 /150 PLATE VII Figure 1-3 Zygabikodiniuiri lenticulatum; 1. a p i c a l view. Narrow 1' i s v i s i b l e . Pores on the theca are conspicuous; 2. v e n t r a l view. The f i n i s v i s i b l e i n the s u l -c a l region; 3. oblique antapical view showing the large s i n g l e antapical p l a t e . (Bar = 20 um i n a l l f i g s . ) Figure 4 Protoperidinium leonis cyst with unknown i n c l u -sion . Figure 5. Figure 6,7,10 Cyst of Z^ _ lenticulatum. The displacement oh the r i g h t side represents the archeopyle s p l i t chara-c t e r i s t i c f o r t h i s species. P. conicum; 6. normal cyst; 7. aberrant cyst with the spines c l o s e l y adherent to the body w a l l . The operculum i s i n two pieces (arrows); 10. C e l l , elongated,from cyst i n f i g . 7. Figure 8,9 P. oblongum; 8. cyst with subapical archeopyle; 9, cyst of d i f f e r e n t morphology, operculum shows c h a r a c t e r i s t i c shape. Figure 11,12 P. c f . denticulatum; 11. cyst with archeopyle; cyst i s also known as Chytroeisphaeridia cariacoensis 12. operculum from cyst of f i g . 11. ../151 /152 PLATE VIII Figure 1,2 Figure 3,4,7 Protoperidinium aspidotum; 1. cyst, archeopyle not v i s i b l e ; 2. c e l l , oblique r i g h t v e n t r a l view. (Bar = 20 pm i n a l l f i g s . ) c f . denticulatum; 3. epitheca; 4. hypotheca; 7. c e l l p a i r s . A l l four c e l l s arose from one parent c e l l over a period of two days. Cyto-plasmic in c l u s i o n s (arrows) confirm an apex to base mode of attachment. Figure 5,6 Figure 8-10 Figure 11,12 P. pentagonum; 5. v e n t r a l view; 6. cyst with archeopyle. P e r i d i n i o p s i s c f . hainanensis; 8. cyst. Archeo-pyle i s a s p l i t around the paragirdle region (arrow); 9. a p i c a l view of c e l l ; 10. antapical view. The dent i n the c e l l (arrow) has d i s t o r t e d the suture between the two antapical p l a t e s . S p i n i f e r i t e s elongatus; 11. viable cyst showing diagnostic antapical process; 12. cyst i n r i g h t l a t e r a l view as i t appears i n treated m a t e r i a l . ./153 /154 PLATE IX-Figure 1 Figure 2,3 Figure 4-8 Figure 9-12 Protoperidinium thorianum ( o p t i c a l l y reversed); aberrant c e l l , oblique v e n t r a l view showing the wide sulcus. (Bar = 20 pm i n a l l f i g s . ) Gonyaulax tamarensis; 2. v i a b l e cyst with starch granules and pigment spot; 3. empty cyst with a p i c a l excystment aperture; inner hyaline envelope i s v i s i b l e (arrow). G. species 2; 4. viable c e l l , v e n t r a l view; 5. v e n t r a l view showing d e l i c a t e theca; 6. oblique v e n t r a l view showing quadrangular 6" (arrow); 7. dorsal view; 8. cyst, oblique v e n t r a l view. Paraplate 6" i s centered. The two-part operculum i s v i s i b l e with an unusual l a t e r a l attachment. The spines are short. G. species; 9. cyst, a p i c a l view, showing two-part operculum with an a p i c a l attachment. The spines are much reduced; 10. v e n t r a l view; 11. oblique r i g h t l a t e r a l view. Extra p l a t e ( l a or 5') i s indicated (arrow); 12. v e n t r a l view; d i s t o r t e d shape of the c e l l i s apparent. Figure 10-12 i n Normarsky i l l u m i n a t i o n . ./155 /156 P L A T E X Figure 1,2 Figure 3,6 Figure 4, 5 Figure 7-9 Figure 10 S p i n i f e r i t e s ramosus; 1. dorsal view; 2. v e n t r a l view. (Bar = 20 jam i n a l l f i g s . ) S c r i p p s i e l l a faeroense; 3. cyst i n o p t i c a l section showing short b i f i d spines; 6. cyst, side view. S. bentori c f . var. truncata; 4. dorsal view; 5. v e n t r a l view. Triangular 6" i s e a s i l y seen. S. membranaceus; 7. dorsal view; 8. v e n t r a l view; 9. o p t i c a l section showing c h a r a c t e r i s t i c anta-p i c a l flange and other membranous processes. S. b e l e r i u s , d i s t o r t e d . The c h a r a c t e r i s t i c p o s t e r i o r process i s indicated (arrow). Figure 11,12 S. bentori; 11. dorsal view; 12. o p t i c a l section showing f l a n g e - l i k e antapical process. Figure 11 and 12 i n Normarsky i l l u m i n a t i o n . ./157 /158 Figure 1,2,4 PLATE XI S p i n i f e r i t e s sp. A (?Hystrichosphaera c f . dentata Gocht), oblique dorsal view showing large two-part operculum i n place within a "reduced" archeo-pyle; 2. oblique v e n t r a l view. Numerous short processes are united along parasutural margins by membranous r u f f l e - l i k e flanges; 4. antapical view showing r u f f l e around i t s border. (Bar = 20 pm i n a l l f i g s . ) Figure 3,6 S. sp. A; 3. cyst has long t r i f u r c a t e processes with b i f i d t i p s . The two-part operculum i s v i s i b l e (arrow); 6. ve n t r a l view. Figure 5 Figure 7,8 Protoperidinium cyst of unknown a f f i n i t i e s . S. sp. A ; 7. dorsal view showing very round archeopyle. The c h a r a c t e r i s t i c processes are clearly, seen; 8. same cyst showing two-part operculum that has f a l l e n i n s i d e the cyst. Figure 9 Tanyosphaeridium sp. ; (?) v i a b l e . ./159 1 5 9 /160 PLATE XII Figure 1 Figure 2,3 Operculodinium centrocarpum with short peg-like processes. Archeopyle i s v i s i b l e . (Bar = 20 pm i n a l l fig s . ) S p i n i f e r i t e s sp. A; 2. cyst with narrow four-sided archeopyle and two-part operculum; 3. o p t i c a l section showing short processes with trumpet-like bases and short branches beyond. A p i c a l boss i s v i s i b l e . Figure 4,5 S."sp. A; 4. antapical view; 5. a p i c a l view. Two-part operculum can be seen. Figure 6. Figure 7 S. sp. A • o p t i c a l s ection showing another v a r i a -t i o n i n the processes. These have! trumpet-like bases with slender t r i f u r c a t e branches bent back over toward the cyst w a l l . S^ sp. A; with t y p i c a l operculum. Fenestrations of the process bases are c l e a r l y seen as i s the microgramular w a l l . Figure 8 S. sp. A; the processes along the lower paragirdle margin are united by low membranous septa. ./161 1 6 1 /162 PLATE XIII • Spiniferites ramosus; 1. dorsal view. "Reduced" archeopyle with narrow membranous border is obvious. Two layers of the cyst wall are dis-tinguishable. Bi- and trifurcate processes have clearly b i f i d tips; 2. antapical view. Low septa delimit parasutural boundaries. (Bar = 20 um) Operculodinium centrocarpum; 3. quadrangular rounded archeopyle is typical. The f i b r i l l a r nature of the cyst wall is seen. The processes have f i b r i l l a r bases, the tips are flattened out. The arrow indicates the region of enlargement of f i g . 4; 4. enlarged view of the processes. Hooklets on the spine tips as described by Wall (1967) are not readily apparent. Some processes appear to be fused. (Fig. 3, Bar = 20 um; Fig. 4, Bar = 2 um) Sj_ " s p . A"; 5. Fenestrate bases of the processes; 6. trifurcate process with b i f i d tips and fenestrate base; 7. Inset-close view of the microgranular wall seen in f i g . 5 and 7 (top). (Fig. 5»Bar = 1 um; Fig. 6, Bar = 2 pm; Fig. 7, Bar = 0.1 pm) /163 /164 PLATE XIV Tanyosohaeridium sp.; 1. l a t e r a l view of cyst. The processes are c y l i n d r i c a l with blunt t i p s l i k e cauliflower heads and connected by a network of low ridges; 2. view of the a p i c a l archeopyle. A f l a p - l i k e operculum i s present. (Bar = 20 pm i n a l l f i g s . ) T. sp.; 3. i n t h i s cyst the processes are f l a t t e n e d out at the apices to form a lacy f a n - l i k e network; 4. the archeopyle i s a t a t t e r e d opening as i f i t had been ripped. An operculum i s not r e a d i l y d i s t i n g u i s h a b l e . ./165 165 /166 BIBLIOGRAPHY Abe, T.H. 1927. Report of the B i o l o g i c a l Survey of Mutsu Bay. 3. 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