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Analipus japonicus (Harv.) Wynne (Phaeophyta): studies of its biology and taxonomy 1980

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ANALIPUS JAPONICUS (HARV.) 1YNNE (PHAEOPHYTA) z STUDIES OF ITS BIOLOGY AND TAXONOMY by •HEHDI ALISON NELSON B . S c , U n i v e r s i t y of A u c k l a n d , New Z e a l a n d , 1975 B.Sc. (Hons.), V i c t o r i a U n i v e r s i t y o f W e l l i n g t o n , Ne Z e a l a n d , 1976 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n THE FACULTY OF GRADUATE STUDIES (Department o f Eotany) Me a c c e p t the t h e s i s as conforming t o the r e q u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH COLUMBIA 27 November 1980 (c) Wendy A l i s o n N e l s o n , 1980 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f the r e q u i r e m e n t s f o r an advanced degree at the U n i v e r s i t y o f B r i t i s h C olumbia, I agree t h a t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and stud y . I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y purposes may be g r a n t e d by the Head o f my Department o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Depa rtment The U n i v e r s i t y o f B r i t i s h Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date -28 -XI- SO i i A b s t r a c t A c u l t u r e and l i g h t microscope study was made of the development, morphology and anatomy of Analipus j a p o n i c u s (Harv.) Wynne (Phaeophyta) . I t s l i f e h i s t o r y was observed i n c u l t u r e and f o l l o w e d i n the f i e l d over a p e r i o d of two years. C y t o l o g i c a l s t u d i e s employing microspectrophotometry to measure r e l a t i v e DNA content confirmed the l i f e h i s t o r y t o be an a l t e r n a t i o n of isomorphic g e n e r a t i o n s - v e r i f y i n g the work of Abe (1935a,b, 1936) which has been questioned i n the l i t e r a t u r e . The u n i l o c u l a r p l a n t s proved t o be d i p l o i d . M e i c s i s occurs i n the u n i l o c u l a r sporangia, r e s u l t i n g i n the f o r m a t i c n of h a p l o i d unispores. Haploid p l u r i l o c u l a r p l a n t s were the only p l a n t s of A.japonicus found i n B r i t i s h Columbia. A survey of herbarium specimens and c o l l e c t i o n s o u t s i d e B r i t i s h Columbia showed t h a t only p l u r i l o c u l a r p o p u l a t i o n s cccur i n most of the e a s t e r n p o r t i o n of A. j a p o n i c u s 1 geographic range {Alaska, B.C., Washington, Oregon, C a l i f o r n i a ) . U n i l o c u l a r p l a n t s occur i n Japan and i n one s i t e i n C a l i f o r n i a . P r i o r to t h i s study the r a r i t y of t h i s stage of A.japonicus' l i f e h i s t o r y had not been r e a l i s e d . The ecology of A.japonicus i n B r i t i s h Columbia was s t u d i e d at a s i t e i n the S t r a i t of Georgia and at two s i t e s i n Earkley Sound on the west co a s t of Vancouver I s l a n d . A comparison was made of A.japonicus and the i i i s u b a n t a r c t i c a l g a Ca e p i d i u m a n t a r c t i c u m J.&g. . ( t o which A . j a p o n i c u s has been l i k e n e d i n t h e l i t e r a t u r e ) . . A l t h o u g h many s i m i l a r i t i e s e x i s t between them, i t i s c o n c l u d e d t h a t t h e s e p l a n t s a r e g e n e r i c a l l y d i s t i n c t . The a f f i n i t i e s o f C a e p i d i u m w i l l r e m a i n u n c l e a r u n t i l l i f e h i s t o r y d a t a a r e a v a i l a b l e . I t i s c o n c l u d e d from an e v a l u a t i o n o f t h e o r d e r E a l f s i a l e s ( i n t o which A . j a p o n i c u s has been r e c e n t l y p l a c e d ) t h a t t h i s o r d e r s h o u l d not be r e c o g n i s e d . , The p l a c e m e n t o f A . j a p o n i c u s i n t o t h e B a l f s i a c e a e i s e v a l u a t e d and c o n s i d e r e d i n a p p r o p r i a t e . The use o f t h e f a m i l y H e t e r o c h o r d a r i a c e a e f o r A . j a p o n i c u s and t h e p l a c e m e n t o f t h i s f a m i l y i n t h e E c t o c a r p a l e s a r e a d v o c a t e d . . i v TABLE OF CONTENTS ABSTRACT i i LIST OF TABLES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v i i i LIST OF FIGURES i x ACKNOWLEDGEMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . x v i GENERAL INTRODUCTION . 2 Terminology ....................................... 5 CHAPTER 1 Development, morphology and anatomy of Analipus j a p o n i c u s (Harv.) Wynne . 7 I n t r o d u c t i o n , 7 M a t e r i a l s and Methods 9 Results and D i s c u s s i o n ............................10 Germination of spores and growth i n c u l t u r e .....10 Morphology and development c f the c r u s t 12 Development of e r e c t axes and l a t e r a l branches . 1 5 Reproductive Organs .19 Conclusions .............................28 CHAETER 2 The phenology and d i s t r i b u t i o n of Ana l i p u s -japonicus (Harv.) Wynne i n the eas t e r n P a c i f i c .....53 I n t r o d u c t i o n .53 M a t e r i a l s and Methods , 56 1.A. C o l l e c t i o n s and Observations i n B r i t i s h Columbia 56 1. B. C o l l e c t i o n s and Observations o u t s i d e B r i t i s h Columbia 57 2. Herbarium Survey 58 Results 59 V 1.A. C o l l e c t i o n s and O b s e r v a t i o n s i n B r i t i s h C o l u m b i a . . . . . . 5 9 1. B. C o l l e c t i o n s and O b s e r v a t i o n s o u t s i d e B r i t i s h C o l u m b i a 62 2. H e r b a r i u m S u r v e y 64 D i s c u s s i o n . . 6 5 CHAPTER 3 L i f e h i s t o r y and c y t o l o g i c a l s t u d i e s o f A n a l i p u s j a p o n i c u s (Harv.) Wynne 87 I n t r o d u c t i o n .87 M a t e r i a l s and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 A. . C u l t u r e Work . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 B. C y t o l o g i c a l Methods .94 R e s u l t s . 9 7 A. . C u l t u r e Work . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 B. C y t o l o g i c a l R e s u l t s . . . . . . . . . . . . . . . . . . . . . . . . 1 0 0 D i s c u s s i o n 102 CHAPTER 4 A u t e c o l o g y o f A n a l i p u s -japonicus a t B a t h I s l a n d , B r i t i s h C o l u m b i a 113 I n t r o d u c t i o n , 1 13 M a t e r i a l s and Methods . . . . . . . . . . . . . . . . . . . . 1 1 4 Q u a d r a t s and T r a n s e c t s 114 C l e a r i n g E x p e r i m e n t . . . . . . . . . . . . . 1 1 6 M o n t h l y C o l l e c t i o n s . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 6 P h o t o g r a p h i c Q u a d r a t s 117 S e t t l e m e n t P l a t e s . . . . 1 1 7 B e s u l t s 118 F i e l d S i t e . . . . . 1 1 8 Quadrats and Trans e c t s 119 C l e a r i n g Experiment ...........120 Monthly C o l l e c t i o n s ............................120 Photographic Quadrats ..121 Settlement P l a t e s .....122 General Observations ...........................122 D i s c u s s i o n ...123 CHAPTER 5 A comparison of Caepidium antarcticum j.Ag. and Analipus japonicus (Harv.) Wynne (Phaeophyta) .. 159 I n t r o d u c t i o n 159 M a t e r i a l s and Methods .............................162 Results and D i s c u s s i o n 163 Morphology . 163 Crustose System .......164 Erect Axis .............166 Bladders .......................................168 The s y s t e m a t i c a f f i n i t i e s of C.antarcticum and i t s r e l a t i o n s h i p t o A. japonicus ....... . ...... 169 CHAPTER 6 An e v a l u a t i o n of the R a l f s i a l e s , R a l f s i a c e a e , and the taxonomic p o s i t i o n of Ana l i p u s japonicus (Harv.) Wynne ...........189 The R a l f s i a l e s 189 The R a l f s i a c e a e ........195 The taxonomic p o s i t i o n of Analipus japonicus (Harv.) Wynne 20 2 GENERAL DISCUSSION 20 7 v i i REFERENCES ......215 APEENDICES ..........231 Appendix I ........................................231 Appendix I I ..235 Appendix I I I 237 Appendix IV ....................................... 239 Appendix V ........241 Appendix VI .............................. 242 Appendix VII .......243 LIST OF TABLES D e s c r i p t i v e measures o f A . j a p o n i c u s sampled from August 1978 t o J u l y 1979 a t t h r e e s i t e s i n B r i t i s h C o l u m b i a 86 Summary o f q u a d r a t c l e a r i n g e x p e r i m e n t ...............158 i x LIST OF FIGURES Fi g u r e 1a. S e t t l e d p l u r i s p o r e s of ft. japonicug ........ 30 Figure 1b. S e t t l e d unispores of fl.japonicus 30 F i g u r e 1c. Germinating p l u r i s p o r e s 30 F i g u r e 1d. Developing d i s c s (from p l u r i s p o r e s ) with evacuated (e) and non-evacuated (n) germination tubes 30 F i g u r e 1e. D i s c s developed from unispores showing empty spore wall (s - arrow) and h a i r (h) .30 F i g u r e 1f. Developing d i s c detached from base of d i s h with c l u s t e r of h a i r s (h) , f i l a m e n t (f) and r h i z o i d s (rh) 30 F i g u r e 2a. Primary d i s c (pd) with f i l a m e n t s (f) and secondary d i s c s (sd) 32 F i g u r e 2b. Developing c r u s t s on s u r f a c e of c o v e r s l i p . C r u s t s w i t h secondary d i s c s i n d i c a t e d by arrows . . . 3 2 F i g u r e 2c. D e v e l o p i n g e r e c t axes from c r u s t . Arrows p o i n t t o t r a n s i t i o n r e g i o n between c r u s t and e r e c t a x i s ...32 F i g u r e 2d. E r e c t axes w i t h l a t e r a l branches grown i n c u l t u r e 3 2 F i g u r e 3a. Herbarium specimen of A . j a p o n i c u s showing c l e a r e d l a t e r a l branches where p l u r a n g i a have r e l e a s e d swarmers 34 F i g u r e 3b. C r u s t s of A . j a p o n i c u s 34 F i g u r e 4a. L o n g i t u d i n a l s e c t i o n of A . j a p o n i c u s c r u s t 36 F i g u r e 4b. C r o s s - s e c t i o n a l view of c r u s t w i t h h a i r s X (h) developing beneath c o r t e x 36 F i g u r e 4c. C r o s s - s e c t i o n of o v e r l y i n g A.japonicus c r u s t showing r h i z o i d s (rh - arrows) developing from lower c o r t e x ....36 Figu r e 5a. L o n g i t u d i n a l s e c t i o n through apex of an e r e c t a x i s of A. japonicus .38 F i g u r e 5b. Apex of e r e c t a x i s of A.japonicus. Arrows point t o zone o f d i v i s i o n s at base of c o r t i c a l f i l a m e n t s .38 F i g u r e 5c. L o n g i t u d i n a l s e c t i o n through r e g i o n of t r a n s i t i o n between e r e c t a x i s and c r u s t .38 F i g u r e 5d. View of t r a n s i t i o n between c r u s t medulla (CM) and medulla of e r e c t a x i s (EM) .38 F i g u r e 5e. L o n g i t u d i n a l view of c e n t r a l region of e r e c t a x i s showing medullary c e l l s which are being s p l i t apart (arrow) as a x i s becomes hollow (he) ....38 F i g u r e 6a. L o n g i t u d i n a l s e c t i o n of a s t e r i l e e r e c t a x i s of A. j a p o n i c u s .40 F i g u r e 6b. L o n g i t u d i n a l s e c t i o n of e r e c t a x i s showing developing l a t e r a l branch. Arrows p o i n t to zone of d i v i s i o n s i n subcortex 40 F i g u r e 6c. Developing l a t e r a l branch. Arrows p o i n t to zone of d i v i s i o n s at base of c o r t i c a l f i l a m e n t s i n sub-cortex ..40 F i g u r e 7. Secondary f i l a m e n t systems i n e r e c t a x i s c f A. j a p o n i c u s .42 F i g u r e 8a.. L o n g i t u d i n a l s e c t i o n of u n i l o c u l a r l a t e r a l x i b r a n c h o f A. j a p o n i c u s 44 F i g u r e 8b. U n i l o c u l a r s p o r a n g i a .1 44 F i g u r e 8c. U n i l o c u l a r s p o r a n g i a ......................44 F i g u r e 8d. U n i l o c u l a r s p o r a n g i a i n c r o s s - s e c t i o n a l view showing l a y e r s o f s t a i n i n g 44 F i g u r e 8e. U n i l o c u l a r s p o r a n g i a i n c r o s s - s e c t i o n ..... 44 F i g u r e 8 f . U n i l o c u l a r s p o r a n g i a i n c r o s s - s e c t i o n .....44 F i g u r e 9. R e l e a s e o f s p o r e s f r o m u n i l o c u l a r s p o r a n g i a . . . . 4 6 F i g u r e 10a. L o n g i t u d i n a l s e c t i o n o f p l u r i l o c u l a r a x i s i n A. j a p o n i c u s 4 8 F i g u r e 10b. . l o n g i t u d i n a l s e c t i o n o f p l u r i l o c u l a r t i s s u e grown i n c u l t u r e , . . . . . 4 8 F i g u r e 10c.. S e c t i o n t h r o u g h l a t e r a l b r a n c h showing d e v e l o p i n g p l u r a n g i a (dp) and h a i r s (h) ...48 F i g u r e lOd. P l u r a n g i a 48 F i g u r e 11. P l u r a n g i a and s t a g e s i n t h e i r r e l e a s e ....50 F i g u r e 12. Map i n d i c a t i n g c o l l e c t i o n s i t e s i n B r i t i s h C o l u m b i a , W a s h i n g t o n , Oregon, and C a l i f o r n i a 73 F i g u r e 13. Map o f s o u t h e r n sid-e o f B a r k l e y Sound i n d i c a t i n g l o c a t i o n s o f s t u d y s i t e s n e a r t h e E a m f i e l d M a r i n e S t a t i o n .75 F i g u r e 14. Mean a x i s h e i g h t and p e r c e n t a g e o f p l u r i l o c u l a r p l a n t s i n c o l l e c t i o n s o f A . j a p o n i c u s from Bath I s l a n d between J u l y 1977 and A u g u s t 1979 77 F i g u r e 15. Mean a x i s h e i g h t and p e r c e n t a g e o f x i i p l u r i l o c u l a r p l a n t s i n c o l l e c t i o n s of A. -japonicus from A g u i l a r House Po i n t between August 1978 and p l u r i l o c u l a r p l a n t s i n c o l l e c t i o n s of A. -japonicus from Second Eeach between May 1978 and August 1979 81 F i g u r e 17. Herbarium survey r e s u l t s showing numbers of s t e r i l e , u n i l o c u l a r , and p l u r i l o c u l a r specimens examined f o r each month f o r s i x regions of A. j a p o n i c u s 1 range .83 F i g u r e 18. Summary o f e f f e c t s of v a r y i n g s a l i n i t y and temperature on growth of germlings of A. japonicus ..............106 Figu r e 19. R e l a t i v e DNA values measured by microspectrophotometry i n n u c l e i of A.japonicus ...108 F i g u r e 20. Diagrammatic r e p r e s e n t a t i o n of (a) l i f e h i s t o r y of A.japonicus i n c u l t u r e , and, (b) l i f e h i s t o r y of A. j a p o n i c u s i n nature 110 F i g u r e 21. Map of Bath I s l a n d i n the S t r a i t of Georgia, B r i t i s h Columbia 133 Figure 22. Diagram of f i e l d s i t e at Bath I s l a n d i n d i c a t i n g r e l a t i v e p o s i t i o n s i n i n t e r t i d a l r e g i o n of quadrats and t r a n s e c t used i n t h i s study 135 Figure 23. Numbers of A.japonicus p l a n t s recorded ever study p e r i o d i n QI and QII .......137 Figure 24.. Numbers of A.japonicus p l a n t s recorded over study p e r i o d i n QIV, Q U I , and QV 139 August 1979 79 Figure 16. Mean a x i s height and percentage of F i g u r e 25. Number of ft.japonicus p l a n t s recorded i n t r a n s e c t over study p e r i o d , 141 Figu r e 26.. Numbers of Ehodoaela, Sargassurn, and Cr y p t o s i p h o n i a p l a n t s recorded i n Q I - Q V and PT1- ET10 over study p e r i o d ......143 Figure 27. Numbers of i n d i v i d u a l s of M y t i l u s and M i c r c c l a d i a recorded i n Q I - Q V and PT1-PT10 over study p e r i o d 145 Figure 28. Numbers of p l a n t s with e i t h e r annual or p e r e n n i a l h a b i t recorded from Q l r Q I I # and Q I V over study p e r i o d 147 Figure 29.. Numbers of p l a n t s with e i t h e r annual or p e r e n n i a l h a b i t recorded from Q U I and QV over study p e r i o d 149 Fi g u r e 30. Mean a x i s height of A. -japonicus and numbers of A.japonicus p l a n t s recorded from Q I - Q V over study p e r i o d ....151 Figu r e 31.. Occurrence of 8 s p e c i e s of algae i n Q I - V and PT1-10 over study p e r i o d ..153 Fi g u r e 32.. Freguency d i s t r i b u t i o n of p l a n t s of fl.japonicus c o l l e c t e d from August 1978 to J u l y 1979 according to u p r i g h t a x i s height ...155 Fi g u r e 33. Type specimen of Cae_pidium antarcticum J . Ag 172 Figu r e 34. Montage of a c r o s s - s e c t i o n of c r u s t o s e t h a l l u s of C.antarcticum ..174 F i g u r e 35a. C r o s s - s e c t i o n of two l a y e r s of x i v C.antarcticum c r u s t . Arrows i n d i c a t e p e g - l i k e extensions of lower c o r t i c a l c e l l s developing i n t o r h i z o i d s 176 Fi g u r e 35b. L o n g i t u d i n a l s e c t i o n of c r u s t of £'antarcticum 176 Fi g u r e 35c. C r o s s - s e c t i o n a l view of c r u s t of C.antarcticum showing hair(h) 176 Fig u r e 35d. Medulla of c r u s t of C.antarcticum i n — r l o n g i t u d i n a l s e c t i o n . Arrow i n d i c a t e s d i r e c t i o n of growth ...176 Figu r e 36a. C r o s s - s e c t i o n of s t e r i l e bladder of C.antarcticum. Arrow p o i n t s t o w a l l breakdown and hollow c e n t r a l r e g i o n (he) ...........178 Fi g u r e 36b. S t e r i l e bladder i n c r o s s - s e c t i o n . Arrow p o i n t s to c u t i c l e - l i k e m a t e r i a l on s u r f a c e of p l a n t 178 Figure 36c. P l u r i l o c u l a r bladder. Arrows point to wall breakdown i n medulla near hollow c e n t r a l r e g i o n 178 Figure 36d. P l u r a n g i a (P) with l a r g e • p h y s o d e - c e l l ' . C u t i c l e - l i k e m a t e r i a l on t h a l l u s s u r f a c e arrowed ..178 Figure 36e. Bladder with p l u r a n g i a which have r e l e a s e d t h e i r contents ...178 Figure 36f. . Released p l u r a n g i a (RP) i n C.antarcticum bladder. Note absence of c u t i c l e - l i k e l a y e r . . Arrows point t o empty l o c u l e s of p l u r a n g i a 178 Fi g u r e 37. L o n g i t u d i n a l s e c t i o n through c r u s t at XV base of an e r e c t a x i s ............180 Figu r e 38.. L i n e drawing of a l o n g i t u d i n a l s e c t i o n a t point where e r e c t a x i s of C.antareticum grows out cf c r u s t 182 F i g u r e 39a. L o n g i t u d i n a l s e c t i o n of u p r i g h t a x i s of Cmantarcticum showing developing l a t e r a l branch (DL - curved arrow) 184 Figure 39b. . Developing l a t e r a l of C.antarcticum showing h a i r s (h) and u n i l o c u l a r sporangia (u) ....184 Figu r e 39c. L o n g i t u d i n a l s e c t i o n through apex (A) and l a t e r a l (L) of C.antarcticum 184 Figure 40a. L o n g i t u d i n a l s e c t i o n of s t e r i l e e r e c t a x i s of Cm a n t a r c t i c u m ...186 F i g u r e 40b. L o n g i t u d i n a l s e c t i o n of e r e c t a x i s of C.antarcticum showing u n i l o c u l a r sporangia ..186 F i g u r e 4 0 c . U n i l o c u l a r t i s s u e of C.antarcticum 186 x v i ^ k n o w l e d g e m en t s I wish t o e x p r e s s my a p p r e c i a t i o n f o r t h e o p p o r t u n i t i e s I h a v e e n j o y e d as a g r a d u a t e s t u d e n t i n t h e Department of B c t a n y a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , and f o r f i n a n c i a l s u p p o r t p r o v i d e d by U.B.C.. G r a d u a t e F e l l o w s h i p s and t e a c h i n g a s s i s t a n t s h i p s . I wish t o t h a n k Dr . . R.E. De Hreede f o r h i s v a l u a b l e a d v i c e and c r i t i c i s m , a s s i s t a n c e i n t h e f i e l d and on SCUBA d i v e s , and f i n a n c i a l s u p p o r t o f t h i s r e s e a r c h (NSEBC 67- 9872, UBC 2 1-9427).. I would l i k e t o a c knowledge t h e g e n e r o u s s u p p o r t o f Dr. K. M. C o l e i n a l l o w i n g me a c c e s s t o h e r l a b o r a t o r y f a c i l i t i e s and e g u i pment f o r a n a t o m i c a l , c y t o l o g i c a l and l i f e h i s t o r y s t u d i e s ( o p e r a t i n g g r a n t NSEEC 67-C645). I w i s h t o t h a n k Dr. C o l e f o r h e r e n c o u r a g e m e n t , h e l p f u l a d v i c e and c r i t i c i s m . I a l s o w i s h t o t h a n k Dr. R.F. S c a g e l f o r c o n s t r u c t i v e c r i t i c i s m and a d v i c e d u r i n g t h i s work, and i n p r e p a r a t i o n o f t h i s t h e s i s , and f o r h i s f i n a n c i a l s u p p o r t o f t h e c o l l e c t i n g t r i p t o the s o u t h e r n l i m i t o f A n a l i p u s j a p o n i c u s (NSERC A - 4 4 7 1 ) . . D r . . F . J . H a r r i s o n ' s p a r t i c i p a t i o n on my a d v i s o r y committee and h e l p f u l r e v i e w o f t h i s m a n u s c r i p t a r e g r a t e f u l l y a c k n o w l e d g e d . My r e s e a r c h was a l s o a i d e d by a G r a n t - i n - A i d o f R e s e a r c h f r c m Sigma X i . I wish t o a c k n o w l e d g e t h e use o f t h e q u a n t i t a t i v e m i c r o s c o p y f a c i l i t i e s o f Dr. J.D. B e r g e r , h i s a s s i s t a n c e and h e l p f u l s u g g e s t i o n s ^ The use o f Dr. R.E.. Foreman's x v i i u n d e r w a t e r camera and f i e l d s t a t i o n a t Bath I s l a n d a r e g r a t e f u l l y a c k n o w l e d g e d . . My t h a n k s a l s o goes t o Dr. . L . J . G o f f f o r her a s s i s t a n c e i n t h i s r e s e a r c h . Dr. G o f f l o c a t e d t h e u n i l o c u l a r p o p u l a t i o n o f A . j a p o n i c u s a t C i t y P o i n t , C a l i f o r n i a , and she and h e r s t u d e n t B r i a n G i b e s a n have p r o v i d e d me w i t h c o l l e c t i o n s and i n f o r m a t i o n f r o m t h i s s i t e . I g r e a t l y a p p r e c i a t e d Dr. G o f f ' s h e l p i n t h e f i e l d on my two c o l l e c t i n g t r i p s t o C a l i f o r n i a , h e r h o s p i t a l i t y and p r o v i s i o n o f l a b o r a t o r y f a c i l i t i e s a t t h e U n i v e r s i t y o f C a l i f o r n i a , S a n t a C r u z . The use o f s p e c i m e n s borrowed from t h e f o l l o w i n g h e r b a r i a i s g r a t e f u l l y a c k n o w l e d g e d : - B r i t i s h Museum o f N a t u r a l H i s t o r y , London; U n i v e r s i t y o f C a l i f o r n i a , B e r k e l e y ; B o t a n y D i v i s i o n , D.S.I.R., New Z e a l a n d ; T r i n i t y C o l l e g e , D u b l i n ; F a r l o w H e r b a r i u m , H a r v a r d U n i v e r s i t y ; G i l b e r t Morgan S m i t h H e r b a r i u m , H o p k i n s M a r i n e S t a t i o n ; L und U n i v e r s i t y , L u n d ; U n i v e r s i t y o f M i c h i g a n , Ann A r b o r ; U n i v e r s i t y o f W a s h i n g t o n , S e a t t l e ; and t h e p e r s o n a l c o l l e c t i o n s of Dr. M.J. Wynne and Dr. R.B. . S e a r l e s . _ I would l i k e t o thank Dr. P.C. S i l v a f o r t a x o n o m i c a d v i c e , and J u l i e C e l e s t i n o O l i v i e r a f o r a s s i s t a n c e i n t h e use o f t h e U. B.C. h e r b a r i u m . F o r a s s i s t a n c e i n t h e f i e l d and f o r c o l l e c t i o n s I would l i k e t o t h a n k t h e f o l l o w i n g p e o p l e : - Deb D o n a l d s o n , Mary C h a m b e r l a i n , L a r r y G o l d e n , S h e i l a Humphrey, Bev Hymes, S u s i e S p e n c e r , Herb V a n d e r m e u l e n , and D r s . D a v i d G a r b a r y , G a y l e Hansen, E r i c Henry, D a v i d P h i l l i p s , and C u r t x v i i i P u e s c h e l . F o r v a l u a b l e d i s c u s s i o n s and a d v i c e on a s p e c t s o f t h i s work I would l i k e t o t h ank D r s . Mary-Jo Duncan, D a v i d G a r b a r y , G a y l e Hansen, L e s l i e L o v e t t - D o u s t , C u r t P u e s c h e l and D a v i d W a l k e r , as w e l l as L i n d s a y C l a r k , S h e i l a Humphrey, and E e n i t o T a n . I would l i k e t o t h a n k Ms V. Kohn f o r t y p i n g T a b l e I I , and D a v i d Z i t t i n f o r a s s i s t a n c e w i t h t h e use o f t h e computer. 1 GENERAL INTRODUCTION 2 The brown a l g a A n a l i p u s j a p o n i c u s (Harv.) Wynne i s f o u n d i n t h e i n t e r t i d a l r e g i o n s o f t h e n o r t h e r n P a c i f i c . D e s p i t e i t s c o n s i d e r a b l e g e o g r a p h i c r a n g e t h e r e i s l i t t l e p u b l i s h e d i n f o r m a t i o n on t h i s p l a n t . P r i o r t o t h i s r e s e a r c h , a number o f i m p o r t a n t q u e s t i o n s a b o u t t h e b i o l o g y of A. -japonicus were o u t s t a n d i n g : - - t h e l i f e h i s t o r y , w hich had been r e p o r t e d t o be an a l t e r n a t i o n o f i s o m o r p h i c g e n e r a t i o n s (Abe, 1935a,b, 1936), needed v e r i f i c a t i o n ( B o l d and Wynne, 1978). t h i s p l a n t had not been grown i n c u l t u r e t o r e p r o d u c t i v e m a t u r i t y and t h e r e were no s t u d i e s o f i t s d e v e l o p m e n t beyond t h e e a r l y s t a g e s o f d i s c f o r m a t i o n . t h e r e were no s t u d i e s o f t h e e c o l o g y o r p h e n o l o g y o f A. -japonicus i n t h e e a s t e r n P a c i f i c . - t h e s i m i l a r i t i e s between A . j a p o n i c u s and t h e s u b a n t a r c t i c a l g a C a e p i d i u m a n t a r c t i c u m had been n o t e d i n t h e l i t e r a t u r e by a number o f w o r k e r s ( e . g . . S k o t t s b e r g , 1921; K y l i n , 194C), b ut no c o m p a r a t i v e s t u d i e s had been p e r f o r m e d * . T h e most d e t a i l e d work on C a e p i d i u m a n t a r c t i c u m was t h a t o f S k o t t s b e r g (1907, 1 9 2 1 ) . . t h e t a x c n o m i c p o s i t i o n of A . j a p o n i c u s was u n c l e a r . A l t h o u g h i t had been p l a c e d i n t h e R a l f s i a l e s by Nakamura (1 9 7 2 ) , t h i s c r d e r has n o t been w i d e l y a c c e p t e d and t h e p l a c e m e n t o f A . j a p o n i c u s v a r i e d between a u t h o r s ( e . g . A b b o t t and H c l l e n b e r g , 1976; L i n d s t r o m , 1977; B o l d and Wynne, 1978). Thus, t h e o b j e c t i v e o f t h i s s t u d y was t o a t t e m p t t o 3 answer some f u n d a m e n t a l q u e s t i o n s a b o u t t h e b i o l o g y c f J • j a p o n i c u s . V a r i o u s a p p r o a c h e s were t a k e n i n t h i s r e s e a r c h . A f i e l d s t u d y i n B r i t i s h C o l u m b i a a t s i t e s i n t h e S t r a i t c f G e o r g i a and on t h e west c o a s t o f V a n c o u v e r I s l a n d p r o v i d e d i n f o r m a t i o n a b o u t t h e s e a s o n a l i t y , p o p u l a t i o n d y n a m i c s , and a u t e c o l o g y o f t h i s p l a n t * C o l l e c t i o n s s o u t h o f B r i t i s h C o l u m b i a t o P o i n t C o n c e p t i o n , C a l i f o r n i a , t h e s o u t h e r n l i m i t o f t h i s p l a n t , r e s u l t e d i n i n f o r m a t i o n on t h e p h e n o l o g y o f A . j a p o n i c u s i n o t h e r p a r t s o f i t s r a n g e and f u r t h e r o b s e r v a t i o n s o f i t s h a b i t a t and g r o w t h . . An h e r b a r i u m s u r v e y was made o f ft.japonicus p l a n t s f r o m t h r o u g h o u t i t s r a n g e t o o b t a i n p h e n o l o g i c a l d a t a . C u l t u r e s t u d i e s were c a r r i e d o u t t o s t u d y (1) t h e l i f e h i s t o r y o f t h i s p l a n t , and (2) t h e e f f e c t o f s e l e c t e d e n v i r o n m e n t a l f a c t o r s on s p o r e g e r m i n a t i o n and d e v e l o p m e n t . A c y t o l o g i c a l s t u d y was p e r f o r m e d t o o b t a i n i n f o r m a t i o n on t h e p l o i d y l e v e l s o f A . j a p o n i c u s p l a n t s , and t o a t t e m p t t o i d e n t i f y t h e s i t e o f m e i o s i s and t h e o c c u r r e n c e o f f e r t i l i s a t i o n . D e velopment c f A . j a p o n i c u s and f o r m a t i o n o f r e p r o d u c t i v e s t r u c t u r e s were f o l l o w e d by a l i g h t m i c r o s c o p e s t u d y . A c o m p a r a t i v e s t u d y was made o f t h e morphology and anatomy o f A . j a p o n i c u s and C a e p i d i u m a n t a r c t i c u m . The t a x o n o m i c p o s i t i o n o f A . j a p o n i c u s was e v a l u a t e d p a r t i c u l a r l y w i t h r e s p e c t t o t h e E a l f s i a l e s and R a l f s i a c e a e where i t h a s been p l a c e d most r e c e n t l y ( B o l d and Wynne, 1978). T h i s r e s e a r c h i s p r e s e n t e d i n s i x i n t e r r e l a t e d 4 c h a p t e r s i n t h e f o l l o w i n g s e q u e n c e : - (1) t h e d e v e l o p m e n t and morphology o f t h e p l a n t , (2) t h e p h e n o l o g y and d i s t r i b u t i o n o f A . j a p o n i c u s i n t h e e a s t e r n N o r t h P a c i f i c , (3) i t s l i f e h i s t o r y and c y t o l o g y , (4) t h e a u t e c o l o g y o f A . j a p o n i c u s a t E a t h I s l a n d , (5) a c o m p a r a t i v e s t u d y o f A . j a p o n i c u s and C a e p i d i u m , and (6) an e v a l u a t i o n o f t h e E a l f s i a l e s , R a l f s i a c e a e and t h e t a x o n o m i c p o s i t i o n o f A . j a p o n i c u s . 5 T e r m i n o l o g y I n A n a l i p u s j a p o n i c u s , swarmers f r o m t h e p l u r i l o c u l a r r e p r o d u c t i v e o r g a n s p r e s u m a b l y can f u n c t i o n e i t h e r as s p o r e s o r g a m e t e s . I t i s not a p p r o p r i a t e t o a s s i g n t h e t e r m s s p o r a n g i a o r g a m e t a n g i a t o t h e s e r e p r o d u c t i v e o r g a n s u n t i l one knows t h e f a t e o f t h e i r p r o d u c t s . F o r t h i s r e a s o n , t h e t e r m p l u r a n g i u m ( p i . p l u r a n g i a ) has been employed t o r e f e r t o a p l u r i l o c u l a r r e p r o d u c t i v e s t r u c t u r e t h a t may b e a r e i t h e r s p o r e s o r gametes (Abb o t t and H o l l e n b e r g , 1S76). Where t h e f a t e o f t h e r e p r o d u c t i v e swarmers i s Jcnown, t h e terms p l u r i s p o r e s o r gametes a r e use d { f o r a s e x u a l o r s e x u a l swarmers r e s p e c t i v e l y ) . The p l u r i l o c u l a r s t r u c t u r e s o f C a e p i d i u m a n t a r c t i c u m a l s o a r e r e f e r r e d t o as p l u r a n g i a as t h e i r f u n c t i o n and p l c i d y l e v e l r e m a i n unknown. Development, m o r p h o l o g y and anatcmy o f A n a l i p u s j a p o n i c u s (Harv.) Wynne 7 I n t r o d u c t i o n Members o f t h e brown a l g a l genus A n a l i p u s K j e l l m a n p o s s e s s a d i s t i n c t i v e , p e r e n n i a l , p r o f u s e l y - l o b e d c r u s t f r c m which a r i s e e r e c t , p e r c u r r e n t a x e s . The e r e c t a x e s b e a r e i t h e r p l u r i l o c u l a r o r u n i l o c u l a r r e p r o d u c t i v e o r g a n s on s e p a r a t e p l a n t s * T h e r e a r e two s p e c i e s i n t h i s g e n u s ; A» - f i l i f o r m i s (Eupr.) P a p e n f u s s has s i m p l e a x e s and A . j a p o n i c u s p o s s e s s e s u n b r a n c h e d l a t e r a l s which c l o t h e t h e u p r i g h t a x e s . Amongst t h e few o t h e r g e n e r a i n t h e P h a e o p h y t a which show an e q u i v a l e n t e l a b o r a t i o n o f c r u s t o s e and e r e c t s y s t e m s i n t h e same g e n e r a t i o n o f t h e l i f e h i s t o r y a r e t h e m o n o t y p i c C a e p i d i u m a n t a r c t i c u m J.Ag.. (see C h a p t e r 5) and S t e r e o c l a d o n r u q u l o s u s (Bory) H a r i o t ( S k o t t s b e r g , 1907, 1921). E a r l y t a x o n o m i c t r e a t m e n t s o f A. •japonicus (as C h c r d a r i a a b i e t i n a E u p r . ex F a r l o w ) d e a l t e x c l u s i v e l y w i t h c h a r a c t e r i s t i c s o f t h e e r e c t a x e s and p l a c e d t h i s p l a n t i n t h e C h o r d a r i a l e s . L a t e r e t h e presumed l i f e h i s t o r y o f t h i s p l a n t r e s u l t e d i n i t b e i n g p l a c e d i n t o t h e g e nus H e t e r o c h o r d a r i a and t h e m o n o t y p i c H e t e r o c h o r d a r i a c e a e . Seme r e c e n t work ha s s e e n A . j a p o n i c u s p l a c e d i n t h e R a l f s i a c e a e and t h e n e w l y - e s t a b l i s h e d R a l f s i a l e s , on t h e b a s i s o f i t s g e r m i n a t i o n p a t t e r n , c h l o r o p l a s t f e a t u r e s and l i f e h i s t o r y . The t a x o n o m i c p o s i t i o n o f t h i s p l a n t i s d i s c u s s e d i n d e t a i l i n C h a p t e r 6. The most c o m p l e t e d i s c u s s i o n o f t h e d e v e l o p m e n t and 8 morphology of A.japonicus has been t h a t of Inagaki (1958) i n h i s systematic study of the C h o r d a r i a l e s i n Japan. Abe (1935a) d e s c r i b e d germination and e a r l y development c f swarmers from u n i - and p l u r i l o c u l a r s t r u c t u r e s . In a paper i n which H e t e r o c h o r d a r i a was i n c l u d e d i n Analipus and the new combination of A.japonicus r e c o g n i s e d , Wynne (1971) compared the two s p e c i e s of A n a l i p u s , p a r t i c u l a r l y t h e i r r e p r o d u c t i v e organs and c r u s t s t r u c t u r e . Wynne (1972) i l l u s t r a t e d germlings of A.japonicus up t o a six-week stage i n c u l t u r e , d e s c r i b i n g germination of p l u r i s p o r e s and formation of the i n i t i a l l y d i s c o i d a l c r u s t . T h i s m orphological and anatomical study has been undertaken to examine the development and re p r o d u c t i o n of A.japonicus. I t i s the f i r s t p art of a - c o n s i d e r a t i o n of the b i o l o g y of t h i s p l a n t and i t s s y s t e m a t i c a f f i n i t i e s w i t h i n th€ Phaeophyta. 9 M a t e r i a l s and Methods Specimens used i n t h i s s t u d y were c o l l e c t e d f r o m J u n e 1977 t o September 1979 a l o n g t h e P a c i f i c c o a s t from B r i t i s h C o l u m b i a t o c e n t r a l C a l i f o r n i a . M o n t h l y o b s e r v a t i o n s and c o l l e c t i o n s were made a t B a t h I s l a n d ( S t r a i t o f G e o r g i a , B r i t i s h C o l u m b i a - 49°09'N, 123°40'W ) and i n B a r k l e y Sound n e a r t h e B a m f i e i d M a r i n e s t a t i o n (west c o a s t o f V a n c o u v e r I s l a n d , B r i t i s h C o l u m b i a - 4 8 ° 5 0 , N , 125°09'W ) . The p l a n t s b e a r i n g u n i l o c u l a r s p o r a n g i a were c o l l e c t e d a t C i t y P o i n t ( M i s s i o n P o i n t ) M o n t e r e y C o u n t y , C a l i f o r n i a ( 3 6 ° 3 3 * N , 123°56'W ) . C o l l e c t i o n s o f p l a n t s were made e i t h e r a t low t i d e o r by use o f SCUBA. ( R e f e r t o Ap p e n d i x I f o r c o l l e c t i o n s i t e s and d a t e s . ) P l a n t m a t e r i a l was e i t h e r f i x e d i n t h e f i e l d o r f i x e d w i t h i n 48 h o u r s o f r e t u r n i n g t o t h e l a b o r a t o r y , h a v i n g been t r a n s p o r t e d f r o m t h e f i e l d wrapped i n damp newspaper i n an i c e - c o o l e d c h e s t . F i x a t i o n was i n e i t h e r 2-5% f o r m a l d e h y d e / s e a w a t e r o r 4-5% g l u t a r a l d e h y d e / s e a w a t e r . M a t e r i a l was embedded i n g l y c o l m e t h a c r y l a t e u s i n g t h e P o l y s c i e n c e s JB4 k i t , and s e c t i o n e d a t 2-5um on a S o r v a l JB4 m i c r o t o m e . S e c t i o n s mounted on g l a s s s l i d e s were s t a i n e d w i t h : - (1) t o l u i d i n e b l u e 0 (TBO) i n b e n z o a t e b u f f e r ( F e d e r and O ' E r i e n , 1968) f o l l o w i n g t h e p r o c e d u r e of M c C u l l y , G o f f and Adshead ( 1 9 8 0 ) . - (2) p e r i o d i c a c i d - S c h i f f ' s (PAS) ( J e n s e n , 1962) s o m e t i m e s 10 c o u n t e r s t a i n i n g t h i s w i t h a n i l i n e b l u e (PAS/AN). Dimedone b l o c k i n g o f b a c k g r o u n d a l d e h y d e s p r e c e d e d t h e PAS p r o c e d u r e ( F e d e r and O ' B r i e n , 1 9 6 8 ) . . - (3) a l c i a n b l u e (AB) a t pH 0.5 ( P a r k e r and D i b o l l , 1966). C u l t u r e s were s t a r t e d f r o m f e r t i l e p l u r i l o c u l a r and u n i l o c u l a r m a t e r i a l u s i n g t h e h a n g i n g d r o p t e c h n i g u e (Wynne, 1969) and were grown i n P r o v a s o l i ' s e n r i c h e d s e a w a t e r medium. ( R e f e r t o C h a p t e r 3 f o r d e t a i l s o f c u l t u r e methods.) R e s u l t s and D i s c u s s i o n G e r m i n a t i o n o f s p o r e s and growth i n c u l t u r e B o t h p l u r i s p o r e s ( F i g u r e l a ) and u n i s p o r e s ( F i g u r e 1b) o f A* j a p o n i c u s f o l l o w e d a s i m i l a r g e r m i n a t i o n s e q u e n c e . A f t e r swimming i n c u l t u r e medium f o r 1/2-4 h o u r s , t h e s p o r e s r o u n d e d up on t h e c o v e r s l i p o r d i s h , h a v i n g become p r o g r e s s i v e l y more s l u g g i s h i n t h e i r movements and e v e n t u a l l y l o s i n g t h e i r f l a g e l l a . The s p o r e s soon showed s i g n s of g e r m i n a t i o n . . A g e r m i n a t i o n t u b e was p r o d u c e d f i r s t , g i v i n g t h e s p o r e s a p e a r - l i k e shape ( F i g u r e 1 c ) . As growth p r o c e e d e d a s h o r t f i l a m e n t was p r o d u c e d , and t h e s p o r e was u s u a l l y e v a c u a t e d r e s u l t i n g i n an empty s p o r e w a l l and a s h o r t t u b e b e f o r e t h e f i r s t new w a l l was l a i d down. The c y t o p l a s m d i d n o t , however, a l w a y s l e a v e t h e germ t u b e ( F i g u r e 1 d). A f t e r 11 f o r m i n g a f i l a m e n t o f 1-3 new c e l l s beyond t h e g e r m i n a t i o n t u b e , l o n g i t u d i n a l and o b l i g u e d i v i s i o n s o c c u r r e d g i v i n g t h e g e r m l i n g s a c l u b - l i k e a p p e a r a n c e . As growth and d i v i s i o n s c o n t i n u e d , t h e d i s c t o o k on a r o s e t t e - l i k e a p p e a r a n c e ( F i g u r e 1 e ) . By a b o u t t h e 1 2 - c e l l s t a g e , t h e o r i g i n a l s p o r e and g e r m i n a t i o n t u b e were o b s c u r e d f r o m view by new growth ( i n c u l t u r e a t about day 15 a t 1 0 ° C ) . . At t h i s s t a g e i n d e v e l o p m e n t , t h e c e l l s began t o d i v i d e i n t h e p l a n e p a r a l l e l t o t h e i r a t t a c h m e n t , f o r m i n g a m u l t i l a y e r e d c r u s t . C o l o u r l e s s , u n i s e r i a t e h a i r s were p r o d u c e d by t h e c r u s t a s i t d e v e l o p e d ( F i g u r e 1e and f } . I f t h e young g e r m l i n g s were d i s l o d g e d from t h e i r s u b s t r a t e t h e b a s a l c e l l s o f t h e c r u s t became v e r y e l o n g a t e d s h o w i n g t h e i r r h i z o i d a l n a t u r e ( F i g u r e 1 f ) . . Wynne (1972 - f i g u r e 16) i l l u s t r a t e s a young c r u s t s h o w i n g t h i s c o n d i t i o n . . As t h e c r u s t s d e v e l o p e d i n c u l t u r e , t h e y o f t e n p r o d u c e d f i l a m e n t s ( F i g u r e 1f and 2 a ) . I n i t i a l l y t h e s e f i l a m e n t s were u n i s e r i a t e b u t t h e y c o u l d be d i s t i n g u i s h e d f r o m h a i r s a s t h e y p o s s e s s e d pigment, and a l s o , as t h e y d e v e l o p e d some became b i - o r m u l t i s e r i a t e . On c o n t a c t w i t h t h e s u b s t r a t e t h e s e f i l a m e n t s f o r m e d a pad o r d i s c o f c e l l s - a s e c o n d a r y d i s c - w h i c h i n t u r n p r o d u c e d h a i r s and c o u l d a l s o d e v e l o p f i l a m e n t s ; . F i g u r e s 2a and b show s u c h a s i t u a t i o n w i t h a number o f s e c o n d a r y o r s u b s i d i a r y d i s c s r e s u l t i n g from one i n i t i a l c r u s t . A l t h o u g h I was n o t a b l e t o o b s e r v e t h e e a r l y d e v e l o p m e n t o f c r u s t s i n t h e f i e l d (due t o f a i l u r e o f s e t t l e m e n t on p l a t e s - C h a p t e r 4 ) , t h i s 12 growth p a t t e r n may be a means o f v e g e t a t i v e g r o w t h , e n a b l i n g t h e c r u s t t o c o l o n i s e t h e s u b s t r a t e more r a p i d l y . I t would h e l p t o e x p l a i n t h e s p r e a d and growth of t h e c r u s t i n t h e f i e l d a t t i m e s when r e c r u i t m e n t f r o m s p o r e s i s n o t o c c u r r i n g . As t h e c r u s t s grew i n c u l t u r e t h e y e v e n t u a l l y became d e t a c h e d from t h e i r s u b s t r a t e - e i t h e r c o v e r g l a s s , g l a s s s l i d e o r base o f t h e c u l t u r e d i s h . The c r u s t s f r o m c u l t u r e had a l o o s e r and l e s s compact a p p e a r a n c e t h a n t h o s e f r o m t h e f i e l d (ccmpare F i g u r e 2c w i t h 3b) . I t h i n k t h a t t h i s c a n be a t t r i b u t e d t o t h e l a c k o f f i r m a t t a c h m e n t i n c u l t u r e c o n d i t i o n s and p e r h a p s a l s o t h e l a c k o f w a ter movement. The c r u s t s i n c u l t u r e c o n s i s t e d o f an a l m o s t c y l i n d r i c a l b r a n c h e d t h a l l u s and p r o d u c e d e r e c t a x e s f r o m t h e t i p s o f t h e i r b r a n c h e s ( F i g u r e 2 c ) . The u p r i g h t a x e s i n c u l t u r e d e v e l o p e d l a t e r a l b r a n c h e s i n some c a s e s , but g e n e r a l l y r e m a i n e d s h o r t e r and p o s s e s s e d f e w e r l a t e r a l s t h a n t h e i r f i e l d c o u n t e r p a r t s (compare F i g u r e 2d w i t h 3 a ) . R e p r o d u c t i v e m a t u r i t y o f c u l t u r e d e r e c t a x e s o c c u r r e d when t h e p l a n t s were c o n s i d e r a b l y s m a l l e r t h a n f i e l d c o l l e c t e d m a t e r i a l . M o r p h o l o g y and d e v e l o p m e n t o f t h e c r u s t The c r u s t of. A. j a p o n i c u s ( F i g u r e 3b) h a s been c l e a r l y i l l u s t r a t e d by S e t c h e l l and G a r d n e r (1903 - p i . 1 8 , f i g u r e 1 6 , 17), Okamura (1913 - p i . CXIV, f i g u r e 6) (who d e s c r i b e d i t a s " t u b e r o u s and l o b e d " ) , and most r e c e n t l y by Wynne 13 (1971 - f i g u r e 4 ) . The c r u s t s have a m u l t i l a y e r e d a p p e a r a n c e i n t h e f i e l d as l o b e s o f t h e f l a t t e n e d , b r a n c h i n g a x e s o f t e n grow o v e r e a c h o t h e r ( F i g u r e 4 c ) . . I n a g a k i (1958) d e s c r i b e d t h e d i s c o i d b a s e o f A . j a p o n i c u s as h a v i n g a m e d u l l a r y l a y e r of " p a r e n c h y m a t o u s c e l l s which a r e i r r e g u l a r l y p o l y g o n a l ' 1 , a c o r t e x c o n s i s t i n g o f "2-3 l a y e r s o f c u b i c c e l l s " , and r h i z o i d s " i s s u i n g f r o m t h e m a r g i n a l p o r t i o n o f t h e p r i m a r y d i s c o i d b a s e " . S e t c h e i l and G a r d n e r (1903) a l s o r e f e r t o t h e c r u s t a s b e i n g p a r e n c h y m a t o u s i n c o n s t r u c t i o n . Nakamura (1972) s t a t e s t h a t t h e g e r m l i n g s d e v e l o p i n t o m i nute p a r e n c h y m a t o u s d i s c s by s u c c e s s i v e c e l l d i v i s i o n s . . W y n n e (1972), however, d e s c r i b e s t h e manner o f g r o w t h o f t h e p r o s t r a t e s y s t e m as b e i n g "a b r o a d m u l t i - a x i a l ' f r o n t ' o f a s s u r g e n t f i l a m e n t s " . The c r u s t has a c l e a r l y f i l a m e n t o u s c o n s t r u c t i o n when s e e n i n l o n g i t u d i n a l s e c t i o n w i t h p r o m i n e n t a s s u r g e n t (or o b l i g u e l y c u r v i n g ) f i l a m e n t s ( F i g u r e 4 a ) . The c r u s t has a c o r t e x o f 3-4 c e l l l a y e r s on t h e uppermost s i d e and a l e s s d i s t i n c t c o r t e x o f 2-3 c e l l l a y e r s on t h e l o w e r s i d e ( F i g u r e 4a and c ) . F r e q u e n t l y t h e l o w e r m a r g i n a l c e l l s e x t e n d f o r m i n g r h i z o i d a l a t t a c h m e n t w i t h t h e s u b s t r a t e o r c r u s t l a y e r b e n e a t h ( F i g u r e 4 c ) . Growth o f t h e c r u s t o c c u r s a t t h e m a r g i n s w i t h d i v i s i o n s o c c u r r i n g a r o u n d the t i p o f e a c h l o b e ( F i g u r e 4 a ) . C e l l s i n t e r n a l t o t h e m e r i s t e m a t i c zone e l o n g a t e and e xpand. H a i r s o c c u r i n t h e c r u s t a l t h o u g h i n mature f i e l d c o l l e c t e d m a t e r i a l t h e y a r e i n f r e q u e n t l y e n c o u n t e r e d . . T h e s e a r e c o l o u r l e s s and a p p e a r t o a r i s e a t 14 t h e b a s e o f t h e c o r t i c a l l a y e r ( F i g u r e 4 b ) . The u n s t a i n e d c e l l s o f t h e c r u s t s p o s s e s s g o l d e n brown b e a d - o r d r o p l e t - l i k e c o n t e n t s . When s t a i n e d w i t h TBO t h e s e b o d i e s s t a i n b r i g h t g r e e n . T h i s m a t e r i a l i s s p r e a d t h r o u g h o u t t h e c e l l s o f t h e c r u s t c o r t e x and m e d u l l a , s t a i n i n g p a r t i c u l a r l y d e n s e l y i n t h e c e l l s of t h e u p p e r c o r t e x as s e e n i n F i g u r e s 4a and 4b. The g r e e n , t u r q u o i s e c o l o u r r e a c t i o n w i t h TBO i n d i c a t e s t h e p r e s e n c e o f p h e n o l i c m a t e r i a l s . The f u n c t i o n and o r i g i n o f t h i s m a t e r i a l i s u n c l e a r . The s t a i n i n g r e a c t i o n i s v e r y s i m i l a r t o t h a t o f t h e m a t e r i a l f o u n d i n t h e t e r m i n a l c o r t i c a l c e l l s of t h e e r e c t a x e s i n fl. -japo n i c u s . These c e l l u l a r b o d i e s have been r e c o r d e d i n brown a l g a e by many w o r k e r s from t h e t i m e o f N a e g e l i ( 1 8 4 7 ) , and a v a r i e t y o f names have been a p p l i e d t o them i n c l u d i n g t h e t e r m s p h y s o d e s and f u c o s a n v e s i c l e s . I n a r e v i e w o f t h e s e s t r u c t u r e s , Ragan (1976) a r g u e s t h a t t h e term f u c o s a n v e s i c l e s s h o u l d no l o n g e r be u s e d as t h i s i m p l i e s t h a t t h e c o n t e n t s a r e c a r b o h y d r a t e s , which has n o t been e s t a b l i s h e d . A l t h o u g h t h e r e have been a number o f s t u d i e s on t h e s e s t r u c t u r e s , and t h e i r n a t u r e has been d e b a t e d f o r o v e r a c e n t u r y , many g u e s t i o n s r e m a i n c o n c e r n i n g t h e i r o r i g i n , f u n c t i o n , and c o m p o s i t i o n (Ragan, 1976). I n a s t u d y o f t h e p h y s o d e s o f D i c t y o t a , Evans and H o l l i g a n (197 2b) c o n c l u d e t h a t t h e s e s u b c e l l u l a r b o d i e s o r i g i n a t e i n t h e p l a s t i d s , s u g g e s t i n g t h a t t h e y might be t h e p r o d u c t s o f p h o t o s y n t h e s i s . T h e i r h i s t o c h e m i c a l s t u d i e s showed t h e 15 p h y s o d e s t o be p h e n o l i c or t a n n i n - l i k e i n n a t u r e . M c C u l l y (1966) s t a t e s t h a t t h e p r e s e n c e of t h e s e b o d i e s i n t h e t h a l l u s o f F u c u s "seems c o r r e l a t e d w i t h a p p a r e n t p h o t o s y n t h e t i c a c t i v i t y , i . e . , t h e y a r e m a i n l y i n t h e e p i d e r m i s and o u t e r l a y e r s o f t h e c o r t e x . . But t h i s c o r r e l a t i o n d o e s not a p p l y f o r h o l d f a s t f i b r e s where t h e y a r e a l s o p l e n t i f u l and where p h o t o s y n t h e s i s i s p r o b a b l y m i n i m a l * " . Of t h e number o f f u n c t i o n s which have been s u g g e s t e d f o r physode m a t e r i a l i n brown a l g a e (Ragan, 1976), t h e most p l a u s i b l e f o r t h e m a t e r i a l p r e s e n t i n t h e c r u s t o f A. - j a p o n i c u s would a p p e a r t o be a m e t a b o l i c r e s e r v e s u b s t a n c e , o r as an a n t i - g r a z i n g compound. Development o f e r e c t a x e s and l a t e r a l b r a n c h e s The e r e c t axes a r i s e o u t o f t h e c r u s t f r o m t h e t i p s o f t h e l o b e s ( F i g u r e 2 c ) . I n i t i a l l y t h e u p r i g h t a x i s has a n a r r o w e r d i a m e t e r t h a n t h e l o b e from which i t a r i s e s t h u s a p p e a r i n g t o come from a d e p r e s s i o n i n t h e c r u s t t i p . . The c o l l a r o f c r u s t t i s s u e a t t h e base o f t h e u p r i g h t a x i s becomes l e s s o b v i o u s a s t h e a x i s i n c r e a s e s i n s i z e . . O p r i g h t a x e s a r e a t f i r s t s i m p l e b u t soon become c o v e r e d i n s h o r t l a t e r a l b r a n c h e s . R a r e l y t h e main a x e s a r e b r a n c h e d and even l e s s f r e q u e n t l y l a t e r a l b r a n c h e s a r e f o r k e d . I n t h e c a s e o f t h e main a x i s , i t a p p e a r s t h a t b r a n c h i n g o c c u r s a f t e r damage o r i n j u r y t o t h e p l a n t , p r e s u m a b l y a f f e c t i n g t h e a p i c a l m e r i s t e m and r e s u l t i n g i n a l a t e r a l b r a n c h becoming d o m i n a n t . 16 I n t h e t r a n s i t i o n zone between t h e c r u s t and t h e u p r i g h t a x i s , m e d u l l a r y c e l l s become e l o n g a t e d and, i n s t e a d o f t e r m i n a t i n g i n c o r t i c a l c e l l s o f t h e c r u s t , r i s e i n t o c l o s e l y p a c k e d p a r a l l e l f i l a m e n t s ( F i g u r e 5 c ) . T h e r e i s a t r a n s i t i o n between t h e r o u n d e r a l m o s t i s o d i a m e t r i c c e l l s of t h e c r u s t m e d u l l a and t h e r e c t a n g u l a r and e l o n g a t e d c e l l s of t h e u p r i g h t a x i s ( F i g u r e 5 d ) . At t h e b o r d e r s o f t h e m e d u l l a t h e c e l l s o f t h e f i l a m e n t s d i v i d e o u t w a r d s g i v i n g r i s e t o s h o r t c o r t i c a l f i l a m e n t s t e r m i n a t e d by c l u b - s h a p e d c e l l s ( F i g u r e 5c) . Rt t h e apex o f t h e e r e c t a x i s of A , j a p o n i c u s t h e r e i s a f a n o f c o r t i c a l o r a s s i m i l a t i n g f i l a m e n t s t e r m i n a t i n g t h e c l o s e l y p a c k e d m e d u l l a r y f i l a m e n t s . T h i s c a n be s e e n most c l e a r l y i n l o n g i t u d i n a l s e c t i o n s o f young m a t e r i a l ( F i g u r e 5a and b ) . The r e g i o n o f d i v i s i o n and e l o n g a t i o n o c c u r s p r i m a r i l y a t t h e b a s e o f c o r t i c a l f i l a m e n t s a t t h e apex ( a r r o w s i n F i g u r e 5 b ) . However, t h e p r o c e s s o f e l o n g a t i o n a f f e c t s t h e c e l l s below t h i s p o i n t . As t h e p l a n t grows, m e d u l l a r y c e l l s a r e s t r e t c h e d a p a r t , l e a v i n g a h o l l o w c e n t r a l r e g i o n t r a v e r s e d by o n l y a few v e r y e l o n g a t e d m e d u l l a r y c e l l s ( F i g u r e 5a and e ) . The c o r t e x and m e d u l l a i n A . j a p o n i c u s a r e w e l l d e f i n e d b u t t h e s u b c o r t e x i s l e s s c l e a r l y d i s t i n g u i s h a b l e . I have i n t e r p r e t e d t h e 1-3 l a y e r s o f r o u n d e d c e l l s b o r n e on m e d u l l a r y f i l a m e n t s and which g i v e r i s e t o c o r t i c a l f i l a m e n t s t o c o n s t i t u t e t h e s u b - c o r t e x ( F i g u r e 6 a ) . I n h e r s t u d y o f t h e M e s o g l o i a c e a e , P a r k e (1933) u s e s 17 two models o f t h a l l u s c o n s t r u c t i o n - t h e c e n t r a l t h r e a d t y p e [e.g. . M e s c g l p i a , M y r i o c l a d i a , A c r p t h r i x ) , and t h e m u l t i p l e s t r a n d t h a l l u s ( e . g . Eudesme, C a s t a g n e a , M y r i p g l o e a , C l a d p s i p h o n ) . A n a l i p u s i s most l i k e C h o r d a r i a i n t h i s t r e a t m e n t and f i t s most c l o s e l y i n t o t h e m u l t i p l e s t r a n d model. . P a r k e (1933) c o n s i d e r s t h a t t h e f i v e g e n e r a i n t h e M e s o g l o i a c e a e (which i n c l u d e s C h o r d a r i a ) a l l p o s s e s s t h e t h r e e t i s s u e t y p e s , a l t h o u g h i n d e f i n i n g t h e s u b - c o r t e x s t a t e s t h a t i t i s "an i n t e r m e d i a t e and v a r i a b l e zone l y i n g between t h e m e d u l l a and c o r t e x " . A c c o r d i n g t o F r i t s c h ( 1 945), however, C h o r d a r i a does n o t p o s s e s s a s u b - c o r t i c a l l a y e r . P a r k e (1933) a l s o d e s c r i b e s s e c o n d a r y f i l a m e n t s y s t e m s i n t h i s f a m i l y which c a n a r i s e f r o m t h e c o r t e x , s u b - c o r t e x and m e d u l l a . F r i t s c h (1945) uses t h e t e r m hyphae f o r t h e s e c o n d a r y f i l a m e n t s and i n t h e c a s e o f C h o r d a r i a d e s c r i b e s them as i s s u i n g f r o m m e d u l l a r y c e l l s and g r o w i n g b o t h upwards and downwards i n t h e t h a l l u s . I n t h e most d e t a i l e d a n a t o m i c a l s t u d y o f A. j a p o n i c u s to, d a t e ( I n a g a k i , 1958) t h e r e i s no m e n t i o n o f s e c o n d a r y f i l a m e n t s . From my e x a m i n a t i o n o f s e r i a l s e c t i o n s and o f s q u a s h e s c f m a t e r i a l p r e v i o u s l y s o a k e d i n sodium c a r b o n a t e , i t a p p e a r s t h a t s e c o n d a r y f i l a m e n t s do o c c u r i n A . j a p o n i c u s , a r i s i n g f r o m m e d u l l a r y c e l l s ( F i g u r e 7 a - d ) . I have n o t been a b l e t o i d e n t i f y any s e c o n d a r y f i l a m e n t s y s t e m s i n t h e c o r t e x o r s u b - c o r t e x . I t i s v e r y d i f f i c u l t t o t r a c e s e c o n d a r y f i l a m e n t s y s t e m s from t h e i r p o i n t s of o r i g i n i n 18 Ai - j a p o n i c u s . The m e d u l l a o f t h i s p l a n t i s t i g h t l y bound and when m a t e r i a l i s s q u a s h e d t h e c o r t i c a l f i l a m e n t s a r e r e l a t i v e l y e a s i l y d i s l o d g e d w h i l e t h e m e d u l l a and s u b - c o r t e x r e m a i n f i r m l y bound t o g e t h e r . L o n g i t u d i n a l s e c t i o n s o f e r e c t a x e s i n d i c a t e t h a t f o r m a t i o n o f l a t e r a l s i s i n i t i a t e d i n t h e s u b - c o r t e x of t h e e r e c t a x i s * F i g u r e 6a shows a l o n g i t u d i n a l s e c t i o n o f a s t e r i l e e r e c t a x i s , w hich c a n be compared w i t h F i g u r e 6b, where the f i r s t s t a g e s o f l a t e r a l f o r m a t i o n c a n be s e e n . D i v i s i o n s o c c u r i n t h e s u b c o r t i c a l zone o f t h e e r e c t a x i s c u t t i n g o f f c e l l s w h ich e i t h e r p r o d u c e c o r t i c a l f i l a m e n t s t o t h e s u r f a c e , o r i f i n t e r n a l l y l o c a t e d become e l o n g a t e d and form t h e m e d u l l a o f t h e b r a n c h ( F i g u r e 6 c ) . . A s th e l a t e r a l grows, j u s t a s i n t h e g r o w t h o f t h e main a x i s , t h e m e d u l l a r y t i s s u e g e t s p u l l e d a p a r t , and t h e b r a n c h becomes h o l l o w . Once i n i t i a t e d , growth o f t h e l a t e r a l f o l l o w s e s s e n t i a l l y t h e same p a t t e r n as t h a t o f t h e main a x i s w i t h t h e z one o f d i v i s i o n s b e i n g a t t h e apex o f t h e b r a n c h i n t h e »fan' o f a c t i v e l y d i v i d i n g c e l l s . The d i v i s i o n s o c c u r a t t h e b a s e o f t h e a s s i m i l a t o r y f i l a m e n t s i n t h e s u b - c o r t i c a l r e g i o n ( a r r o w s i n F i g u r e 6c) . As t h e main a x i s and l a t e r a l b r a n c h e s d e v e l o p , t e r m i n a l c e l l s of t h e c o r t i c a l f i l a m e n t s become e n l a r g e d and o f t e n c l u b - s h a p e d ( F i g u r e s 6 a - c ) . The f i l a m e n t s a r e 3-5 c e l l s i n l e n g t h and i n t e r s p e r s e d w i t h them a r e h a i r s which a r i s e i n t h e s u b - c o r t e x ( F i g u r e 1 0c)..The t e r m i n a l c e l l s o f c o r t i c a l f i l a m e n t s a r e f i l l e d w i t h g o l d e n - b r o w n c o n t e n t s 19 which s t a i n an i n t e n s e g r e e n w i t h TBO, i n d i c a t i n g t h e p r e s e n c e o f p h e n o l i c compounds. R e p r o d u c t i v e O r g a n s (a) U n i l o c u l a r s p o r a n g i a U n i l o c u l a r s p o r a n g i a a r e borne p r i m a r i l y on t h e l a t e r a l b r a n c h e s , a l t h o u g h t h e y can a l s o be f o u n d on t h e main a x i s o f t h e u p r i g h t s ( F i g u r e 8a) . The s p o r a n g i u m d e v e l o p s f r c m a c e l l c u t o f f by a s u b - c o r t i c a l c e l l and i s s u r r o u n d e d by s t e r i l e c o r t i c a l f i l a m e n t s 3-6 c e l l s i n l e n g t h ( F i g u r e s 8b and c ) . The s p o r a n g i u m i s immersed amongst t h e s e f i l a m e n t s u n t i l m a t u r i t y . In f r e s h l y c o l l e c t e d f e r t i l e m a t e r i a l u n i l o c u l a r s p o r a n g i a were e a s i l y d e t a c h e d from t h e s u b c o r t e x . When viewed w i t h a d i s s e c t i n g m i c r o s c o p e , d i s l o d g e d r i p e s p o r a n g i a and e m p t i e d s p o r a n g i a were seen on t h e s u r f a c e o f t h e t h a l l u s . No empty s p o r a n g i a were s e e n i n s e c t i o n e d m a t e r i a l s u g g e s t i n g t h a t t h i s d i s l o d g e m e n t may t a k e p l a c e p r i o r t o r e l e a s e • and a i d i n d i s p e r s a l o f t h e s p o r e s . S t a i n i n g w i t h TBO, AB, and PAS shows d i s t i n c t z o n e s i n t h e s p o r a n g i a . I n m a t e r i a l s t a i n e d w i t h TBO ( F i g u r e s 8b and d) t h e c e n t r a l p a r t o f t h e s p o r a n g i u m , which l a t e r r o u n d s up t o form t h e s p o r e s , s t a i n s a s a s p e c k l e d p i n k / p u r p l e z o n e. S u r r o u n d i n g t h i s l i g h t l y s t a i n e d c e n t r a l r e g i o n i s a d a r k l y s t a i n e d p u r p l i s h band. Between t h i s d a r k l y s t a i n i n g band and t h e s p c r a n g i a l w a l l i s a t h i n l i g h t l y s t a i n e d p i n k 20 l a y e r . T h i s l a y e r i s n o t v e r y n o t i c e a b l e i n y o u n g e r s p o r a n g i a b u t , as t h e y mature, i t becomes more p r o m i n e n t p a r t i c u l a r l y a r o u n d t h e apex o f t h e s p o r a n g i u m where i t f o r m s a 'dome' or 'cap'. The s p o r a n g i a l w a l l s t a i n s l i g h t - p u r p l e . Once t h e s p o r e s have r o u n d e d up, TBO s t a i n i n g r e s u l t s i n l i g h t p i n k / p u r p l e s p o r e s s u r r o u n d e d by a more d a r k l y s t a i n i n g p u r p l e b o r d e r . When u n i l o c u l a r m a t e r i a l i s s t a i n e d w i t h AB t h e p a t t e r n i s v e r y s i m i l a r ( F i g u r e 8 e ) . E ach s p o r e i s o n l y l i g h t l y s t a i n e d t u r g u o i s e b l u e and i s s u r r o u n d e d by a more d a r k l y s t a i n i n g b o r d e r . Around t h e s p o r e s i n t h e c e n t r e o f t h e s p o r a n g i u m i s a l a y e r o f b l u e - s t a i n i n g m a t e r i a l . The s p o r a n g i a l w a l l s t a i n s q u i t e d e n s e l y w i t h AB.. Between t h e w a l l and t h e l a y e r s u r r o u n d i n g t h e s p o r e s i s a t h i n l a y e r ( n e t seen i n F i g u r e 8e) which as w i t h TBO i s l i g h t l y s t a i n e d . T h i s i s a g a i n t h e l a y e r which f o r m s t h e cap a t t h e t i p o f t h e s p o r a n g i u m as i t m a t u r e s . The PAS s t a i n e d m a t e r i a l p o s s e s s e d s i m i l a r l a y e r s ( F i g u r e s 8a, c and f ) . The c e n t r a l c y t o p l a s m , and l a t e r t h e s p o r e s , s t a i n e d a dense p i n k / m a g e n t a and were o u t l i n e d w i t h a more d a r k l y s t a i n i n g b o r d e r . I m m e d i a t e l y s u r r o u n d i n g t h e s p o r e mass t h e l a y e r w hich was d a r k l y s t a i n e d w i t h TBO and AB was v i r t u a l l y u n s t a i n e d . T h i s r e g i o n was o c c a s i o n a l l y s p e c k l e d w i t h p a t c h e s o f p a l e p i n k . Between t h i s e s s e n t i a l l y u n s t a i n e d l a y e r and t h e s p o r a n g i a l w a l l ( t h i n b u t b r i g h t l y s t a i n i n g ) was a b r i g h t p i n k l a y e r and i t i s t h i s which f o r m s t h e cap a t t h e t i p o f t h e s p o r a n g i u m . . 21 T o l u i d i n e b l u e i s a t h i a z i n dye and s t a i n s m e t a c h r o m a t i c a l l y h i g h m o l e c u l a r weight compounds w i t h f r e e c a r b o x y l , s u l p h a t e or phosphate groups (Barka and Anderson, 1963; M c C u l l y , 1966). . P a r t i c u l a r l y , t h e p i n k / p u r p l e s t a i n w i t h TBO i s r e g a r d e d t o be c h a r a c t e r i s t i c of s u l p h a t e d p o l y s a c c h a r i d e s o f t e n r e f e r r e d t o as f u c o i d i n i n brown a l g a e . P e r c i v a l (1979) p r e f e r s , f o r s i m p l i c i t y , t o c a l l t h e s u l p h a t e d p o l y s a c c h a r i d e s o f the Phaeophyta • f u c a n s ' . . She n o t e s t h a t v a r i o u s names have been a p p l i e d t o t h e s e compounds and t h a t i n f a c t one i s d e a l i n g w i t h "a f a m i l y of p o l y d i s p e r s e h e t e r o m o l e c u l e s c o n t a i n i n g i n a d d i t i o n t o f u c o s e , v a r y i n g p r o p o r t i o n s o f g a l a c t o s e , mannose, x y l o s e and g l u c u r o n i c a c i d " ( P e r c i v a l , 1978). These compounds have a s t r o n g a f f i n i t y f o r Mg i o n s , and as t h e s e i o n s are h i g h l y h y d r a t e d , P e r c i v a l (1979) c o n s i d e r s t h a t they p l a y an i m p o r t a n t r o l e i n t h e p r o t e c t i o n o f t h e s e p l a n t s from d e s i c c a t i o n . A l c i a n dyes d i f f e r e n t i a t e and l o c a t e s u l p h a t e d and n o n - s u l p h a t e d p o l y s a c c h a r i d e s . A l c i a n b l u e a t pH 0.5 complexes e x c l u s i v e l y w i t h s u l p h a t e groups as t h e c a r b o x y l groups are not d i s s o c i a t e d below pH 1.0 ( P a r k e r and D i b o l l , 1966) . The pink/magenta r e a c t i o n o f PAS i s q u i t e s p e c i f i c f o r p o l y s a c c h a r i d e s h a v i n g f r e e h y d r o x y l groups on two v i c i n a l c arbon atoms ( H o t c h k i s s , 1948; M c C u l l y , 1 9 6 6 ) . . A l g i n i c a c i d i s a c a r b o x y l a t e d p o l y s a c c h a r i d e found i n t h e Phaeophyta which po s s e s s e s f r e e v i c i n a l h y d r o x y l groups and i s t h u s PAS p o s i t i v e . P e r c i v a l (1979) n o t e s t h a t a l g i n i c a c i d has a 22 h i g h a f f i n i t y f o r Ca i o n s and t h a t t h i s f e a t u r e g i v e s t h i s m a t e r i a l a h i g h g e l s t r e n g t h . Wasserman (1948, 1949) showed t h a t a l g i n i c a c i d can u n d e r g o a s a l t i n g o u t p r o c e s s i n which t h e p o r e s i z e o f t h e g e l i s g r e a t l y d e c r e a s e d , and t h a t i t c a n a l s o a c t as an e x c e l l e n t i o n - e x c h a n g e r e s i n * A l g i n a t e s ( s a l t s o f a l g i n i c a c i d ) have been f o u n d i n t e r c e l l u l a r l y and i n c e l l w a l l s (Evans and H o l l i g a n , 1972a) and a r e c o n s i d e r e d t o have s t r u c t u r a l as w e l l as i o n - e x c h a n g e f u n c t i o n s (Mackie and P r e s t o n , 1974). U s i n g t h e TBO metachromasy ( P e r c i v a l , 1964; M c C u l l y , 1966, 1968a; E v a n s and H o l l i g a n , 1972a; E v a n s , Simpson, and C a l l o w , 1973; T o t h , 1974, 1976) one can c o n c l u d e t h a t w i t h i n t h e u n i l o c u l a r s p o r a n g i a o f A . j a p o n i c u s l a y e r s can be i d e n t i f i e d w i t h t h e f o l l o w i n g c o m p o s i t i o n : - (1) t h e s p o r a n g i a l w a l l , p o s s e s s i n g b o t h a l g i n i c a c i d and f u c a n s (PAS, AB, and TBO p o s i t i v e ) , (2) a t h i n l a y e r i m m e d i a t e l y b e n e a t h t h e w a l l w hich e n c i r c l e s t h e s p o r a n g i u m and a l s o e x p a n d s a t t h e apex o f t h e s p o r a n g i u m t o form a dome o r cap c o n s i s t i n g of p r i m a r i l y a l g i n i c a c i d (PAS p o s i t i v e , TBO and AB l i g h t l y s t a i n e d ) , (3) a l a y e r p r i m a r i l y o f f u c a n s w i t h p o s s i b l y t r a c e s o f a l g i n i c a c i d (TBO and AB d a r k l y s t a i n e d , m a i n l y PAS n e g a t i v e b u t w i t h a few p a t c h e s o f s t a i n ) , and (4) s p o r e s s u r r o u n d e d w i t h a t h i n l a y e r c o n t a i n i n g b o t h a l g i n i c a c i d and f u c a n s (PAS, TBO and AB p o s i t i v e ) . R number c f w o r k e r s have s p e c u l a t e d on t h e e c o l o g i c a l and f u n c t i o n a l s i g n i f i c a n c e o f p o l y s a c c h a r i d e m a t e r i a l , p a r t i c u l a r l y t h e h y d r o p h i l i c a l g i n i c a c i d and f u c a n s w i t h i n 23 brown a l g a e . In p a r t i c u l a r , p r o t e c t i o n a g a i n s t d e s i c c a t i o n , and the osmotic b u f f e r i n g p r o v i d e d by t h e s e m a t e r i a l s i s s t r e s s e d ( M c C u l l y , 1966, 1968b; P e r c i v a l , 1964, 1978, 1979). These compounds have a l s o been i m p l i c a t e d i n the r e l e a s e of u n i s p o r e s . . K n i g h t (1923) suggested t h a t i n P i l a y e l l a r e l e a s e o f s p o r e s from the u n i l o c u l a r s p o r a n g i a was a f f e c t e d by a r a p i d i n t a k e of water r e s u l t i n g i n an i n c r e a s e i n t h e volume of t h e m u c i l a g i n o u s m a t e r i a l s u r r o u n d i n g t h e mass of s p o r e s . I n a study of P i l a y e l l a , Markey and Wilce (1976) suggested two f a c t o r s are i n v o l v e d i n t h e r e l e a s e p r o c e s s - both t h e s w e l l i n g of m u c i l a g i n o u s m a t e r i a l and a weakening of t h e i n n e r p a r t o f t h e s p o r a n g i a l w a l l . In a s t u d y of u n i l o c u l a r s p o r a n g i a of s p e c i e s o f E l a c h i s t a , Hecatonema and P i l a y e l l a , L o i s e a u x (1973) suggested t h a t t h e m u c i l a g e s u r r o u n d i n g the s p o r e s may a c t as an osmoticum, thus i n c r e a s i n g osmotic p r e s s u r e . In a s t u d y of propagule r e l e a s e i n s e v e r a l brown a l g a e , Toth (1976) observed spore r e l e a s e and the changes o c c u r r i n g i n c e l l w a l l s t r u c t u r e . Toth s u g g e s t s t h a t t h e p r o d u c t i o n and h y d r a t i o n o f c a r b o h y d r a t e s may produce an i n t e r n a l p r e s s u r e i n the s p o r a n g i a . Enzymatic breakdown of t h e w a l l p a r t i c u l a r l y i n t h e r e g i o n of t h e apex a l l o w s t h e s p o r e s t o be r e l e a s e d from t h i s p o i n t of weakness as s w e l l i n g o f p o l y s a c c h a r i d e s o c c u r s . Toth s u g g e s t s t h a t t i m i n g of r e l e a s e may occur w i t h r e i m m e r s i o n i n s e a w a t e r a f t e r t h e low t i d e exposure and d e s i c c a t i o n , the subsequent r a p i d h y d r a t i o n of the m u c i l a g i n o u s p o l y s a c c h a r i d e s 24 r e s u l t i n g i n t h e s p o r a n g i u m b u r s t i n g . As c a n be s e e n i n F i g u r e s 9a and 9b u n i l o c u l a r . s p o r a n g i a o f A. -japonicus d e h i s c e a t t h e s p o r a n g i a l t i p . The r e l e a s e o f s p o r e s from s p o r a n g i a o b s e r v e d i n t h e l a b o r a t o r y was r a p i d . The s p o r a n g i a l w a l l r u p t u r e s a t t h e t i p ( F i g u r e 9b shows a f l a p o f w a l l b e n t back a f t e r r e l e a s e ) and a mass of s p o r e s o o z e s out - sometimes l e a v i n g a few s p o r e s or a p p a r e n t l y u n c l e a v e d c y t o p l a s m b e h i n d ( F i g u r e 9 b ) . The mass o f s p o r e s s w e l l s a s i t l e a v e s t h e s p o r a n g i u m f o r m i n g a m u s h r o o m - l i k e c l o u d . W i t h i n a minute o r l e s s of r u p t u r i n g t h e s p o r a n g i a l w a l l , s p o r e s on t h e o u t s i d e o f t h e s p o r e mass b e g i n t o b r e a k away. E y e s p o t s i n t h e s p o r e s c a n be c l e a r l y s e e n i n l a t e r s t a g e s o f development w i t h i n t h e s p o r a n g i u m and a r e v e r y d i s t i n c t a f t e r r e l e a s e of t h e s p o r e mass. (b) P l u r a n g i a I n A . j a p o n i c u s p l u r a n g i a a r e b o r n e p r i m a r i l y on t h e l a t e r a l s a l t h o u g h t h e y can a l s o be f o u n d on t h e main a x i s o f t h e u p r i g h t p l a n t . They r e s u l t from t h e t r a n s f o r m a t i o n o f t h e s t e r i l e c o r t i c a l f i l a m e n t s by t r a n s v e r s e , l o n g i t u d i n a l and sometimes o b l i g u e d i v i s i o n s ( F i g u r e s 10 and 1 1 ) . They f o r m u s u a l l y u n i - t o b i s e r i a t e ( o c c a s i o n a l l y m u l t i s e r i a t e ) f i l a m e n t s which a r e t e r m i n a t e d by 1-3 s t e r i l e c e l l s ( F i g u r e 10a).. At m a t u r i t y t h e d e p t h o f t h e p l u r i l o c u l a r t i s s u e c a n be up t o 15 c e l l s deep a l t h o u g h more o f t e n t h e l a y e r f o r m e d i s 8-10 c e l l s deep. I n c u l t u r e d m a t e r i a l t h e p l a n t s became f e r t i l e when much s m a l l e r t h a n 25 t h e i r f i e l d c o u n t e r p a r t s and t h e c o r t i c a l f i l a m e n t s were n o t t r a n s f o r m e d i n t o such a deep f e r t i l e l a y e r , c o n s i s t i n g i n s t e a d o f a l a y e r of u s u a l l y 4-8 c e l l s i n d e p t h ( F i g u r e 10b) . When t h e p l u r a n g i a r e l e a s e t h e i r swarmers t h e l a t e r a l s where t h e y were a p p e a r w h i t e ( F i g u r e 3 a ) . T h i s i s b e c a u s e t h e e n t i r e p h o t o s y n t h e t i c t i s s u e i s s h e d f r o m t h e f e r t i l e a r e a s w i t h o n l y c o l o u r l e s s s u b - c o r t e x and m e d u l l a r e m a i n i n g * No r e g e n e r a t i o n o f t h e c o r t e x o c c u r s and t h u s t h e r e s u l t o f r e p r o d u c t i o n v i a p l u r a n g i a i s l o s s o f t h e l a t e r a l s and o f t e n o f t h e e n t i r e e r e c t a x i s . Swarmer r e l e a s e from t h e p l u r i l o c u l a r o r g a n s ( F i g u r e s 11a-d) can o c c u r v e r y r a p i d l y p a r t i c u l a r l y i n m a t e r i a l which has been d r i e d by e m e r s i o n and t h e n i s r e i m m e r s e d i n w a ter. The p r o c e s s o f r e l e a s e a p p a r e n t l y c o n s i s t s o f a t l e a s t two s t a g e s . The s t e r i l e t e r m i n a l c e l l s become d e t a c h e d from t h e p l u r a n g i a a l t h o u g h t h e y a r e s t i l l l a t e r a l l y c o n n e c t e d t o one a n o t h e r . . Thus i n s e c t i o n s o f r e l e a s i n g m a t e r i a l one c a n f i n d r i b b o n s o f s t e r i l e c e l l s ( F i g u r e 11b and c ) . T h e s e c u r l backwards as i f r e l e a s e d f r o m t e n s i o n when t h e y a r e d e t a c h e d f r o m t h e p l u r a n g i a . Once t h e swarmers have been shed empty p l u r a n g i a c a n n o t be f o u n d . The w a l l s a p p e a r t o d i s i n t e g r a t e and o n l y f r a g m e n t s r e m a i n ( F i g u r e s 11b-d). J a a s u n d (1964) s u g g e s t e d t h a t i n R a l f s i a v e r r u c o s a t h e s t e r i l e l a y e r above t h e s p o r a n g i a f a c i l i t a t e d r e l e a s e as t h e s e l a t e r a l l y c o n n e c t e d c e l l s r e m a i n e d a d j o i n e d and b r o k e 26 away from t h e r e s t of t h e t i s s u e b e n d i n g back w i t h some s p o r a n g i a . . E d e l s t e i n , Chen and M c L a c h l a n (1968) r e p o r t e d t h a t i n R a l f s i a f u n q i f o r m i s a s i m i l a r r e l e a s e p r o c e s s o c c u r s . I n t h i s s p e c i e s t h e r e a r e a l s o o v e r l y i n g s t e r i l e c e l l s which " f o r m a f i r m l a y e r which h o l d s the s p o r a n g i a t o g e t h e r " and t h a t when t h e s o r u s m atures t h i s l a y e r s p l i t s . They r e p o r t e d t h a t t h e s p o r a n g i a c o m p l e t e l y d i s i n t e g r a t e on s p o r e r e l e a s e and n o t e d t h a t t h i s i s n o t t h e c h a r a c t e r i s t i c mode o f p l u r i l o c u l a r r e l e a s e r e p o r t e d f o r t h e P h a e o p h y t a . R e l e a s e i s most o f t e n r e p o r t e d t o o c c u r t h r o u g h an a p i c a l p o r e . W ith TBO ( F i g u r e s 11a and b r 10b) and AB (not shown) t h e p l u r a n g i a s t a i n i n t h e f o l l o w i n g way - swarmers a r e s t a i n e d p i n k / p u r p l e and t u r q u o i s e b l u e r e s p e c t i v e l y , and a r e s u r r o u n d e d w i t h i n t h e p l u r a n g i a l w a l l by a more d a r k l y s t a i n i n g l a y e r . The p l u r a n g i a l w a l l a l s o s t a i n s i n t e n s e l y as does t h e m a t r i x s u r r o u n d i n g t h e p l u r a n g i a w i t h t h e s e two s t a i n s i n d i c a t i n g t h e p r e s e n c e o f s u l p h a t e d p o l y s a c c h a r i d e s . . O n s t a i n i n g w i t h PAS ( F i g u r e s 10a, c and d) t h e swarmers s t a i n a b r i g h t p i n k / m a g e n t a , b u t t h e b o r d e r between t h e swarmer and t h e w a l l i s u n s t a i n e d - i m p l y i n g t h e p r e s e n c e o f o n l y s u l p h a t e d p o l y s a c c h a r i d e s s u c h as f u c a n s , and not c a r b o x y l a t e d p o l y s a c c h a r i d e s s u c h as a l g i n i c a c i d . The p l u r a n g i a l w a l l s s t a i n w i t h PAS and t h e m a t r i x o u t s i d e t h e w a l l s s u r r o u n d i n g t h e p l u r a n g i a i s n o t s t a i n e d . Thus t h e m a t e r i a l s u r r o u n d i n g t h e swarmers w i t h i n t h e p l u r a n g i a and s u r r o u n d i n g t h e p l u r a n g i a i s s u l p h a t e d 27 p o l y s a c c h a r i d e or f u c a n . I t seems l i k e l y t h a t t h i s m a t e r i a l p l a y s an i m p o r t a n t r o l e i n t h e r e l e a s e p r o c e s s . The r a p i d i t y o f t h e r e l e a s e and t h e s h a t t e r i n g o f t h e p l u r a n g i a l w a l l s s u g g e s t s t h a t a s w e l l i n g o r p r e s s u r e b u i l d - u p due t o t h e h y d r a t i o n o f t h e s e compounds c o u l d be i m p o r t a n t * T h i s may be p a r t i c u l a r l y s o a f t e r d e s i c c a t i o n and t h e n r e i m m e r s i o n i n water, o r when an i n c r e a s e i n t h e h y d r a t i o n o f t h e s e m a t e r i a l s o c c u r s a f t e r a b r e a k a p p e a r s i n t h e p r o t e c t i v e s u r f a c e s t e r i l e l a y e r . A s w e l l i n g of t h e m a t r i x would f o r c e t h e p l u r a n g i a l w a l l s t o b r e a k down and a l s o f o r c e back t h e r o o f o f s t e r i l e c e l l s a l l o w i n g a more r a p i d r e l e a s e o f t h e swarmers i n t o t h e e n v i r o n m e n t . . I n e a c h p l u r i l o c u l a r p l a n t , v i r t u a l l y a l l t h e c o r t i c a l t i s s u e i n most l a t e r a l s i s c o n v e r t e d t o p l u r i l o c u l a r o r g a n s and t h u s huge numbers o f swarmers a r e p r o d u c e d . The l o s s o f c o r t i c a l t i s s u e on swarmer r e l e a s e makes t h e p l a n t v u l n e r a b l e t o e n t r y o f d i s e a s e a g e n t s ( f u n g i were f o u n d i n t i s s u e o f r e l e a s e d l a t e r a l s and were c a u s i n g d i s r u p t i o n o f t h e c e l l s , t h u s d e l e t e r i o u s l y a f f e c t i n g t h e p l a n t ) , and an i n c r e a s e i n e p i p h y t e s ( n o r m a l l y r a r e ) was a l s o f o u n d on t h e p a r t s o f t h e p l a n t where r e l e a s e had t a k e n p l a c e . The p o s t - r e p r o d u c t i v e phase o f p l u r i l o c u l a r p l a n t s v a r i e s i n l e n g t h . I n some p l a n t s t h e whole p l a n t i s f o r m i n g s p o r a n g i a v i r t u a l l y s y n c h r o n o u s l y . I n o t h e r i n d i v i d u a l s , e x t e n s i o n and new l a t e r a l f o r m a t i o n i s o c c u r r i n g a t t h e apex w h i l s t l a t e r a l s l o w e r on t h e a x i s a r e f e r t i l e and s h e d d i n g swarmers. 28 C o n c l u s i o n s A summary o f t h e l i f e h i s t o r y of A n a l i p u s •japonicus i s p r e s e n t e d i n F i g u r e 20.. The d e v e l o p m e n t sequence o f A n a l i p u s j a p o n i c u s which I o b s e r v e d i n c u l t u r e and i n f i e l d m a t e r i a l may be summarised as f o l l o w s . A swarmer r e l e a s e d f r o m e i t h e r a u n i l o c u l a r s p o r a n g i u m o r p l u r a n g i u m s e t t l e s and g e r m i n a t e s . W i t h i n t e n days a d i s t i n c t i v e d i s c o i d a l pad o f c e l l s has f o r m e d from an i n i t i a l l y one t o t h r e e - c e l l e d g e r m l i n g . The d i s c o f c e l l s d e v e l o p s t o f o r m a p s e u d o p a r e n c h y m a t o u s c r u s t . The a s s u r g e n t f i l a m e n t s o f t h e c r u s t r i s e t o f o r m an upper c o r t i c a l l a y e r o r grow downwards f o r m i n g a b a s a l c o r t e x from which r h i z o i d s w i l l o f t e n a r i s e . E r e c t axes a r e p r o d u c e d from t h e t i p s o f t h e c r u s t a x e s , a s t h e s e c y l i n d r i c a l and b r a n c h i n g a x e s e x t e n d a l o n g t h e s u b s t r a t e . The anatomy of t h e e r e c t a x i s o f t h i s p l a n t i s s i m i l a r t o t h a t o f C h o r d a r i a . I t d i f f e r s f r o m C h o r d a r i a , however, i n p o s s e s s i n g p l u r a n g i a as w e l l as u n i l o c u l a r s p o r a n g i a on s e p a r a t e i n d i v i d u a l s . . Swarmer r e l e a s e from t h e p l u r a n g i a i s r a p i d and r e s u l t s i n c o m p l e t e l o s s o f t h e c o r t i c a l t i s s u e from t h e f e r t i l e b r a n c h e s , as a l l o f t h i s t i s s u e e x c e p t f o r a t e r m i n a l l a y e r o f 1-3 c e l l s i n d e p t h i s c o n v e r t e d t o p l u r a n g i a . The u n i l o c u l a r s p o r a n g i a a r i s e f r o m t h e s u b c o r t e x and a r e f o u n d immersed among s t e r i l e c o r t i c a l f i l a m e n t s . I n b o t h t y p e s o f r e p r o d u c t i v e o r g a n s t h e r e i s c o n s i d e r a b l e p r o d u c t i o n o f p o l y s a c c h a r i d e m a t e r i a l . The s t u d i e s o f o t h e r w o r k e r s have 29 i m p l i c a t e d s u c h m a t e r i a l t o be i n v o l v e d i n t h e r e l e a s e p r o c e s s i n brown a l g a e . B o t h t h e t e r m i n a l c o r t i c a l c e l l s o f t h e e r e c t a x e s and t h e c e l l s o f t h e c r u s t c o r t e x and m e d u l l a a r e r i c h i n p h e n o l i c m a t e r i a l s . The f u n c t i o n of t h e s e 'physodes* i s n o t known a l t h o u g h i n o t h e r brown a l g a e w o r k e r s have p o s t u l a t e d t h e i r r o l e i n s t o r a g e and a s a n t i - g r a z i n g compounds(Ragan, 1976). . 30 Fig u r e 1a. S e t t l e d p l u r i s p o r e s of A.japonicus. (525x) F i g u r e 1b. S e t t l e d u nispores of A.japonicus. (525x) F i g u r e 1c. Germinating p l u r i s p o r e s . (600x) Fig u r e 1d. Developing d i s c s (from p l u r i s p o r e s ) with evacuated (e) and non-evacuated (n) germination tubes. (690x) Figure 1e. D i s c s developed from unispores showing empty spore wall (s - arrow) and h a i r (h) . . (540x) F i g u r e 1f. Developing d i s c detached from base of d i s h with c l u s t e r of h a i r s (h) , f i l a m e n t (f) and r h i z o i d s ( r h ) . (675x) 31 32 F i g u r e 2a. P r i m a r y d i s c (pd) w i t h f i l a m e n t s (f) and s e c o n d a r y d i s c s (sd) . (370x) F i g u r e 2b. D e v e l o p i n g c r u s t s on s u r f a c e o f c o v e r s l i p . C r u s t s w i t h s e c o n d a r y d i s c s i n d i c a t e d by a r r o w s . . (2;75x) F i g u r e 2c* D e v e l o p i n g e r e c t a x e s from c r u s t . A r r o w s p o i n t t o t r a n s i t i o n r e g i o n between c r u s t and e r e c t a x i s . (7.5x) F i g u r e 2d. E r e c t a x e s w i t h l a t e r a l b r a n c h e s grown i n c u l t u r e . . (1.5x)  34 F i g u r e 3a* H e r b a r i u m s p e c i m e n o f A . j a p o n i c u s showing c l e a r e d l a t e r a l b r a n c h e s where p l u r a n g i a have r e l e a s e d swarmers. F i g u r e 3b. C r u s t s o f A . j a p o n i c u s . , ( 2 . 7 5 x ) IN, 36 F i g u r e 4 a . L o n g i t u d i n a l s e c t i o n o f A . j a p g n i c u s c r u s t . UC = upper c o r t e x ; M = m e d u l l a ; LC = l o w e r c o r t e x . TBO s t a i n . (260x) F i g u r e 4b. C r o s s - s e c t i o n a l view o f c r u s t w i t h h a i r s (h) d e v e l o p i n g b e n e a t h c o r t e x . TBO s t a i n . (740x) F i g u r e 4 c . C r o s s - s e c t i o n o f o v e r l y i n g A . j a p o n i c u s c r u s t showing r h i z o i d s (rh - arrows) d e v e l o p i n g f r o m l o w e r c o r t e x . PAS s t a i n . (250x) 37 38 F i g u r e 5a. L o n g i t u d i n a l s e c t i o n t h r o u g h apex o f an e r e c t a x i s o f A . j a p o n i c u s . Note h o l l o w e d c e n t r a l r e g i o n ( h e ) . TBO s t a i n . (230x) F i g u r e 5b. .Apex o f e r e c t a x i s o f A . j a p o n i c u s . Arrows p o i n t t o zone o f d i v i s i o n s a t base o f c o r t i c a l f i l a m e n t s . TBO s t a i n . (785x) F i g u r e 5 c . L o n g i t u d i n a l s e c t i o n t h r o u g h r e g i o n o f t r a n s i t i o n between e r e c t a x i s and c r u s t . EM = e r e c t a x i s m e d u l l a ; CM = c r u s t m e d u l l a ; EC = e r e c t a x i s c o r t e x ; CC = c r u s t c o r t e x . TBO s t a i n (175x) F i g u r e 5d. View o f t r a n s i t i o n between c r u s t m e d u l l a (CM) and m e d u l l a o f e r e c t a x i s (EM). TBO s t a i n . (475x) F i g u r e 5e. L o n g i t u d i n a l view o f c e n t r a l r e g i o n of e r e c t a x i s showing m e d u l l a r y c e l l s which a r e b e i n g s p l i t a p a r t (arrow) as a x i s becomes h o l l o w ( h e ) . TBO s t a i n . (475x) 3 9 no F i g u r e 6a. L o n g i t u d i n a l s e c t i o n o f a s t e r i l e e r e c t a x i s of A . j a p o n i c u s . , C = c o r t e x ; SC = s u b - c o r t e x ; M = m e d u l l a . TBO s t a i n . (550x) F i g u r e 6 t . L o n g i t u d i n a l s e c t i o n o f e r e c t a x i s showing d e v e l o p i n g l a t e r a l branch. Arrows p o i n t t o zone of d i v i s i o n s i n s u b c o r t e x . TBO s t a i n . (550x) F i g u r e 6 c . D e v e l o p i n g l a t e r a l branch. Arrows p o i n t t o zone of d i v i s i o n s at base of c o r t i c a l f i l a m e n t s i n s u b - c o r t e x TEO s t a i n . (550x) 41 42 F i g u r e 7. S e c o n d a r y f i l a m e n t s y s t e m s i n e r e c t a x i s o f A . j a p o n i c u s . Arrows p o i n t t o s e c o n d a r y f i l a m e n t s ; he = h o l l o w c e n t r a l r e g i o n o f a x i s . . (550x)  44 F i g u r e 8 a . L o n g i t u d i n a l s e c t i o n o f u n i l o c u l a r l a t e r a l b r a n c h c f A . j a p o n i c u s . PAS s t a i n . . C = c o r t e x ; SC = sub- c o r t e x ; H = m e d u l l a ; 0 = u n i l o c u l a r s p o r a n g i a . . (215x) F i g u r e 8b. U n i l o c u l a r s p o r a n g i a . TBO s t a i n . SF = s t e r i l e f i l a m e n t s ; 1,2,3,4 = numbers a s s i g n e d t o l a y e r s w i t h i n t h e s p o r a n g i a . (625x) F i g u r e 8 c . U n i l o c u l a r s p o r a n g i a . PAS s t a i n , ( l a b e l s a s f o r b.) (625x) F i g u r e 8d. U n i l o c u l a r s p o r a n g i a i n c r o s s - s e c t i o n a l view showing l a y e r s o f s t a i n i n g . TBO s t a i n . (625x) F i g u r e 8e* U n i l o c u l a r s p o r a n g i a i n c r o s s - s e c t i o n . . A B s t a i n , (625x) F i g u r e 8 f . U n i l o c u l a r s p o r a n g i a i n c r o s s - s e c t i o n . .PAS s t a i n (625x) 45 46 F i g u r e 9. R e l e a s e o f s p o r e s from u n i l o c u l a r s p o r a n g i a . (b) arrow p o i n t i n g t o f l a p o f s p o r a n g i a l w a l l t o r n back d u r i n g r e l e a s e . (750x) 4 7 48 F i g u r e 10a. L o n g i t u d i n a l s e c t i o n o f p l u r i l o c u l a r a x i s i n fl.japonicus. TS = t e r m i n a l s t e r i l e c e l l s ; P = p l u r a n g i a ; SC = s u b - c o r t e x ; M = m e d u l l a ; he = h o l l o w c e n t r a l r e g i o n o f a x i s . .PAS s t a i n . (190x) F i g u r e 10b. L o n g i t u d i n a l s e c t i o n o f p l u r i l o c u l a r t i s s u e grown i n c u l t u r e . . S = s p o r e g e r m i n a t i n g ; l a r g e arrow p o i n t s t o t h i c k s u r f a c e l a y e r o f m u c i l a g e - l i k e m a t e r i a l . . T B O s t a i n . . (650x) F i g u r e 10c. S e c t i o n t h r o u g h l a t e r a l b r a n c h showing d e v e l o p i n g p l u r a n g i a (dp) and h a i r s ( h ) . PAS s t a i n . (600x) F i g u r e 10d. P l u r a n g i a . PAS s t a i n . . (900x) 49 50 F i g u r e 11. P l u r a n g i a and s t a g e s i n t h e i r r e l e a s e , (a) mature p l u r a n g i a (fc) r e l e a s e o f swarmers has begun - t e r m i n a l s t e r i l e c e l l s (TS) c u r v i n g back; h a i r (h) s h o w i n g o r i g i n i n t h e s u b - c o r t e x ; EP = r e l e a s i n g p l u r a n g i a (c) p l u r a n g i a h a v e r e l e a s e d t h e i r c o n t e n t s - some swarmers from t h e p l u r a n g i a (ps) can be s e e n amongst t h e p l u r a n g i a l w a l l f r a g m e n t s . (d) p l u r a n g i a l w a l l f r a g m e n t s (arrowed) r e m a i n i n g a f t e r r e l e a s e o f swarmers. TBO s t a i n , ( ( a ) - ( c ) 550x, (d) 750x)  52 CHAPTER 2, The p h e n o l o g y and d i s t r i b u t i o n o f A n a l i p u s j a p o n i c u s (Harv.) Wynne i n t h e e a s t e r n P a c i f i c 53 I n t r o d u c t i o n A n a l i p u s j a p o n i c u s i s w i d e l y d i s t r i b u t e d i n t h e n o r t h e r n P a c i f i c h a v i n g been r e c o r d e d f r o m Japan (the t y p e l o c a l i t y ) , t h e K u r i l e s , S a g h a l i e n , t h e J a p a n Sea c o a s t of S i b e r i a , t h e s e a o f O k h o t s k , Kamschatka, t h e B e r i n g S e a , t h e A l e u t i a n I s l a n d s , and t h e P a c i f i c c o a s t of N o r t h A m e r i c a from A l a s k a t o P o i n t C o n c e p t i o n , C a l i f o r n i a ( F a r l o w , 1875; Okamura, 1910,1933; S e t c h e l l and G a r d n e r , 1903, 1925; T o k i d a , 1954; I n a g a k i , 1 958) . D e s p i t e t h i s c o n s i d e r a b l e g e o g r a p h i c range t h e r e i s l i t t l e p u b l i s h e d on t h e p h e n o l o g y c r e c o l o g y o f t h i s p l a n t . The o b s e r v a t i o n s which h ave been p u b l i s h e d on t h e p h e n o l o g y o f A. j a p o n i c u s -* t h e g r o w t h o f t h e u p r i g h t a x e s from t h e c r u s t and t h e s e a s o n a l i t y o f r e p r o d u c t i o n T a r e c o n t r a d i c t o r y and s u g g e s t t h a t A . j a p o n i c u s e x h i b i t s d i f f e r e n t p a t t e r n s of s e a s o n a l growth i n d i f f e r e n t p a r t s o f i t s r a n g e : - J a p a n - " S p o r a n g i a o f b o t h k i n d s , March-May" (Okamura, 1910) ; " I n May o r June t h e f r o n d s u s u a l l y f a l l o f f f r o m t h e base which r e m a i n s a d h e r i n g t o t h e r o c k s and g i v e s r i s e t o new f r o n d s a g a i n i n September o r O c t o b e r . " ( I n a g a k i , 1958); " . . . d e v e l o p s i n Asamushi from w i n t e r t o s p r i n g . . . " (Abe, 1935a) ; S a g h a l i e n - " T h i s i s one o f t h e most common and u b i g u i t o u s seaweeds i n S a g h a l i e n . . I t f l o u r i s h e s from e a r l y s p r i n g t o l a t e summer.'! ( T o k i d a , 1954); C a l i f o r n i a - " . . . e r e c t a x e s d i s a p p e a r i n g i n l a t e f a l l , new a x e s a r i s i n g from p e r e n n i a l b a s e t h e f o l l o w i n g 54 spring.'.' ( A b b o t t and H o l l e n b e r g , 1976) . When I began t h i s s t u d y t h e r e was no p u b l i s h e d a c c o u n t o r o b s e r v a t i o n s on t h e p h e n o l o g y o f A . j a p o n i c u s i n B r i t i s h C o l u m b i a . D e s c r i p t i o n s o f t h e h a b i t o f A . j a p o n i c u s which have been p u b l i s h e d a l s o v a r y c o n s i d e r a b l y , d e s c r i b i n g i t t o be an a l g a which grows on r o c k s o r s t o n e s i n t h e u p p e r i n t e r t i d a l (Okamura, 1910; A b b o t t and H o l l e n b e r g , 1976)* i n t h e m i d d l e c r l o w e r i n t e r t i d a l r e g i o n ( S e t c h e l l and G a r d n e r , 1925; Kawabata, 1936; N a g a i , 1940; T o k i d a , 1954), a t o r j u s t below low water ( S e t c h e l l and G a r d n e r , 1903), o r , " g r o w i n g on r o c k s , s t o n e s , o r s e a m u s s e l s , v e r y o f t e n i n t i d e p o o l s , between t i d e marks" ( I n a g a k i , 1958). T a n i g u t i (1962, 1978) d e s c r i b e s A . j a p o n i c u s as f o r m i n g an i m p o r t a n t a s s o c i a t i o n i n t h e i n t e r t i d a l r e g i o n w i t h C h o r d a r i a f l a g e l l i f o r m i s i n n o r t h e r n H o k k a i d o i n t h e c o a s t a l a r e a s s t r o n g l y i n f l u e n c e d by d r i f t i c e . S c a g e l (1961) r e c o r d e d t h e v e r t i c a l d i s t r i b u t i o n o f A . j a p o n i c u s i n t h e i n t e r t i d a l r e g i o n from t h r e e s i t e s i n t h e v i c i n i t y o f Queen C h a r l o t t e S t r a i t - Hope I s l a n d , Deer I s l a n d , and K l u c k s i w i R i v e r . T h e s e t h r e e s i t e s r e p r e s e n t a t r a n s i t i o n f r o m h i g h t c low s a l i n i t y , and a l s o , f r o m e x p o s u r e t o s u r f a c t i o n a t Hope I s l a n d t o c a l m e r w a t e r s a t t h e o t h e r two s i t e s . S c a g e l f o u n d t h a t a t Hope I s l a n d A . j a p o n i c us e x t e n d s t h r o u g h a c o n s i d e r a b l y l a r g e r v e r t i c a l r a n g e and t o a much h i g h e r e l e v a t i o n t h a n a t e i t h e r o f t h e o t h e r two l o c a t i o n s , and a t t r i b u t e d t h e s e v e r t i c a l e l e v a t i o n d i f f e r e n c e s p r i m a r i l y t o t h e e f f e c t o f s u r f a c t i o n . From s t u d i e s o f 55 i n t e r t i d a l t r a n s e c t s w i t h d i f f e r i n g a s p e c t and wave e x p o s u r e , S a i t o e t a l . (1971) c o n c l u d e d t h a t A . j a p o n i c u s (as H e t e r o c h o r d a r i a a b i e t i n a ) i s a "sun f o r m " . . They f o u n d i t was n o t p r e s e n t a t e i t h e r t h e most e x p o s e d t r a n s e c t or t h e two most s h e l t e r e d t r a n s e c t s . From t h e i r d i s c u s s i o n i t i s n o t c l e a r why t h e y c h o s e t o use t h e t e r m " s u n f o r m " r a t h e r t h a n s t a t i n g t h a t A . j a p o n i c u s grew most a b u n d a n t l y a t t h e s i t e s o f i n t e r m e d i a t e e x p o s u r e . Atobe and S a i t o (1974) s t u d y i n g t h e e f f e c t s o f wave a c t i o n on a l g a l z o n a t i o n a t s i t e s i n H o k k a i d o , n o t e t h a t a t l e a s t 10 a l g a l s p e c i e s i n c l u d i n g A . j a p o n i c u s e x i s t a t h i g h e r e l e v a t i o n s a l o n g more e x p o s e d t r a n s e c t s and a t l o w e r e l e v a t i o n s a l o n g more s h e l t e r e d t r a n s e c t s . I n t h i s s e c t i o n o f my r e s e a r c h I a d d r e s s e d t h e f o l l o w i n g g u e s t i o n s a b o u t A . j a p o n i c u s i n B r i t i s h C o l u m b i a : - At what t i m e o f t h e y e a r a r e t h e u p r i g h t axes p r o d u c e d ? What i s t h e s e a s o n a l i t y o f r e p r o d u c t i o n ? What p r o p o r t i o n s o f p l u r i l o c u l a r and u n i l o c u l a r p l a n t s a r e f o u n d i n t h e f i e l d ? A f t e r t h e f i r s t y e a r o f f i e l d s t u d i e s i n B r i t i s h C o l u m b i a , I e x t e n d e d t h e s c o p e o f t h i s work and s o u g h t i n f o r m a t i o n a b o u t s e a s o n a l g r o w t h and r e p r o d u c t i o n o f A . j a p o n i c u s i n e t h e r p a r t s o f i t s r a n g e p a r t i c u l a r l y i n t h e e a s t e r n P a c i f i c . 56 M a t e r i a l s and Methods _1. A. C o l l e c t i o n s and O b s e r v a t i o n s i n B r i t i s h Columbia Over a 27 month p e r i o d , 85 c o l l e c t i o n s were made of ft.japonicus i n B r i t i s h Columbia. T h i s m a t e r i a l came from 18 s i t e s (shown i n F i g u r e 12 and 13) i n t h e s o u t h e r n p o r t i o n of t h e P r o v i n c e . (Refer t o Appendix I f o r a complete l i s t of c o l l e c t i o n s i t e s w i t h dates of c o l l e c t i o n s . ) R e g u l a r s a m p l i n g o c c u r r e d at t h r e e major s i t e s - Bath I s l a n d i n the S t r a i t o f G e o r g i a , and A g u i l a r House P o i n t and Second Beach, near the B a m f i e l d Marine S t a t i o n i n B a r k l e y Sound on the west c o a s t of Vancouver I s l a n d . Whenever p o s s i b l e , c o l l e c t i o n s c o n s i s t e d of more t h a n 50 e r e c t axes w i t h c r u s t m a t e r i a l . The c o l l e c t i o n s were p r e s e r v e d and s t o r e d i n 2-5% f o r m a l d e h y d e / s e a w a t e r . M a t e r i a l from t h e t h r e e major s i t e s was used f o r a d e t a i l e d p h e n o l o g i c a l s t u d y . The p l a n t c h a r a c t e r i s t i c s used i n t h i s study were measured a c c o r d i n g t o t h e f o l l o w i n g p r o c e d u r e : - From each c o l l e c t i o n , 25 u p r i g h t p l a n t s were chosen at random. ( T w e n t y - f i v e was found t o be a s u i t a b l e sample s i z e a f t e r r u n n i n g a t e s t s e r i e s i n c r e a s i n g t h e sample s i z e and measuring v a r i a t i o n i n the value of t h e mean h e i g h t of t h e main a x i s . ) 1) The number of axes per e r e c t p l a n t was r e c o r d e d . 2) Each p l a n t was s c o r e d f o r t h e presence/absence of l a t e r a l s . 57 3) The h e i g h t o f t h e main a x i s was measured f o r e a c h p l a n t . M e a s u r i n g 1/3 o f t h e main a x i s h e i g h t down from t h e apex, t h e n e x t 1 cm o f t h e main a x i s was used f o r c h a r a c t e r i s t i c s 4-6. 4) The number o f l a t e r a l s o c c u r r i n g i n t h e 1cm o f main a x i s was c o u n t e d as a measure o f l a t e r a l d e n s i t y . 5) The l e n g t h s o f t h e f i r s t f i v e l a t e r a l s e n c o u n t e r e d were measured. A mean v a l u e f o r l a t e r a l l e n g t h was c a l c u l a t e d when t h e r e were t h r e e o r more l a t e r a l s measured. 6) The w i d t h s o f t h e f i r s t f i v e l a t e r a l s e n c o u n t e r e d were measured, and a mean v a l u e c a l c u l a t e d when t h e r e were t h r e e o r more measured. 7) E i t h e r l a t e r a l s o r t h e main a x i s were s e c t i o n e d t o d e t e r m i n e t h e r e p r o d u c t i v e s t a t u s o f t h e p l a n t . A t o t a l o f 1125 p l a n t s were examined - 700 p l a n t s f r o m B a t h I s l a n d from c o l l e c t i o n s made o v e r 27 months, 225 p l a n t s f r c m Second Beach from c o l l e c t i o n s s p a n n i n g 16 months, and 200 p l a n t s from A g u i l a r House P o i n t c o l l e c t e d o v e r 13 months. J..E. C o l l e c t i o n s and O b s e r v a t i o n s o u t s i d e B r i t i s h C o l u m b i a F i g u r e 12 shows t h e l o c a t i o n s a t w h i c h A. -japonicus was c o l l e c t e d d u r i n g t h i s s t u d y . I n A p pendix I , t h e c o l l e c t i o n d a t a a r e g i v e n f o r t h e m a t e r i a l o f A. - j a p o n i c u s t h a t came from s i t e s o u t s i d e B r i t i s h C o l u m b i a . An e x t e n s i v e c o l l e c t i n g t r i p was made i n l a t e March 1979.. C o l l e c t i o n s were made o v e r an e i g h t - d a y p e r i o d f r o m s o u t h o f t h e 58 p u b l i s h e d s o u t h e r n l i m i t o f A . j a p o n i c u s ( P o i n t C o n c e p t i o n r C a l i f o r n i a ) t o Oregon. T h u s , b e g i n n i n g i n t h e s o u t h a t C a r p e n t e r i a R e e f (where no A . j a p o n i c u s was f o u n d ) , 17 s i t e s were v i s i t e d a t 15 o f wh i c h A . j a p o n i c u s was f o u n d and c o l l e c t e d . A n o t h e r c o l l e c t i n g t r i p t o C a l i f o r n i a was made i n e a r l y September 1979 when o v e r a f o u r - d a y p e r i o d t e n s i t e s were v i s i t e d from P i e d r a s B l a n c a s i n t h e s o u t h t o Moss Beach (San Mateo County) i n t h e n o r t h , and A . j a p o n i c u s was c o l l e c t e d a t a l l s i t e s . C o l l e c t i o n s were p r e s e r v e d and s t o r e d i n 2-5% f o r m a l d e h y d e / s e a w a t e r . Whenever p o s s i b l e , c o l l e c t i o n s c o n s i s t e d o f more t h a n 50 e r e c t a x e s . M a t e r i a l was s e c t i o n e d , and i t s r e p r o d u c t i v e s t a t u s r e c o r d e d . 2.. H e r b a r i u m S u r v e y H e r b a r i u m s p e c i m e n s o f A . j a p o n i c u s were o b t a i n e d f r o m as many p a r t s o f i t s g e o g r a p h i c r a n g e as p o s s i b l e . M a t e r i a l f o r t h i s s t u d y came from t h e f o l l o w i n g h e r b a r i a : - U n i v e r s i t y o f B r i t i s h C o l u m b i a (UBC); U n i v e r s i t y o f C a l i f o r n i a , B e r k e l e y (UC), i n c l u d i n g s p e c i m e n s now h o u s e d a t UC from t h e D u d l e y H e r b a r i u m , S t a n f o r d U n i v e r s i t y (DH), and from t h e M i s s o u r i B o t a n i c a l Garden (MO); G i l b e r t Morgan S m i t h H e r b a r i u m , H o p k i n s M a r i n e S t a t i o n (GMS); U n i v e r s i t y o f Washington (RTU) ; U n i v e r s i t y o f M i c h i g a n , Ann A r b o r (MICH); F a r l o w H e r b a r i u m , H a r v a r d U n i v e r s i t y (FH); and T r i n i t y C o l l e g e D u b l i n (the t y p e m a t e r i a l ) (TCD). P o r t i o n s c f l a t e r a l axes were s e c t i o n e d and t h e r e p r o d u c t i v e s t a t u s o f t h e m a t e r i a l was r e c o r d e d . A t o t a l 59 o f 397 s p e c i m e n s w i t h e r e c t axes were examined, w i t h t h e sample s i z e a c c o r d i n g t o g e o g r a p h i c l o c a t i o n b e i n g as f o l l o w s : - A l a s k a ( 9 3 ) , B r i t i s h C o l u m b i a ( 1 9 1 ) , W a s h i n g t o n ( 3 3 ) , Oregon ( 1 9 ) , C a l i f o r n i a ( 4 3 ) , J a p a n ( 1 7 ) , Kamschatka ( 1 ) . F i v e s p e c i m e n s c o n s i s t i n g o f o-nly t h e c r u s t o s e p o r t i o n of A. j a p o n i c u s were a l s o examined - J a p a n (1) , B r i t i s h C o l u m b i a ( 1 ) , C a l i f o r n i a ( 3 ) . R e s u l t s _1. A. C o l l e c t i o n s and O b s e r v a t i o n s i n B r i t i s h C o l u m b i a From o b s e r v a t i o n s made o f A . j a p o n i c u s o v e r t h e two y e a r s t u d y p e r i o d , i t was f o u n d t h a t i n t h e s o u t h e r n p a r t o f B r i t i s h C o l u m b i a t h i s p l a n t i s c h a r a c t e r i s t i c o f m o d e r a t e l y e x p o s e d t o e x p o s e d r o c k y i n t e r t i d a l s h o r e s b o t h on t h e west c o a s t o f Vancouver I s l a n d and i n the S t r a i t o f G e o r g i a . G e n e r a l l y i t was f o u n d n o t t o o c c u r on l o o s e r o c k s o r b o u l d e r s and was r a r e l y f o u n d i n t i d a l p o o l s . . The r e l a t i v e t i d a l h e i g h t and t h e v e r t i c a l e x t e n t o f t h e A . j a p o n i c u s r a n g e d i f f e r e d between s i t e s . . A t some s i t e s , f o r example Botany Beach ( 48°32«N, 124°27«W ) , A . j a p o n i c u s o c c u r r e d n e a r P o s t e l s i a p a l m a e f o r m i s Rupr. i n t h e u p p e r i n t e r t i d a l r e g i o n o f an exposed s h o r e , and a t o t h e r s , f o r i n s t a n c e B a t h I s l a n d and B r e a k w a t e r I s l a n d ( 4 9 ° 0 9 ' N , 1 2 3 ° 4 0 , W ) , i t e x t e n d e d i n t o t h e l o w e r i n t e r t i d a l / u p p e r s u b t i d a l r e g i o n and was f o u n d g r o w i n g ~ a l o n g s i d e 60 C r y p t o p l e u r a y i p l a c e a (J.Aq.) K y i . and L i t h o t h r i x a s p e r g i l l u m G ray. . I n g e n e r a l , A . j a p o n i c u s o c c u r r e d i n t h e m i d - i n t e r t i d a l r e g i o n s , f r e q u e n t l y on r o c k y h e a d l a n d s w h i c h were more e x p o s e d t h a n t h e s u r r o u n d i n g c o v e s and s h o r e l i n e . The f i e l d s i t e a t B a t h I s l a n d was e x p o s e d t o s u r f a c t i o n f r o m t h e s o u t h e a s t , t h e p r e d o m i n a n t wind d i r e c t i o n i n w i n t e r months and f o r most o f t h e y e a r . Wind and s t o r m s were p a r t i c u l a r l y s e v e r e i n F e b r u a r y o f b o t h y e a r s i n w h i c h f i e l d work was c a r r i e d o u t . D r i f t l o g s a c c u m u l a t e d i n t h e upper i n t e r t i d a l r e g i o n a t B a t h I s l a n d . These were d i s l o d g e d by h i g h t i d e s o r s t o r m waves and o f t e n c a u s e d c o n s i d e r a b l e damage by b a t t e r i n g and c r u s h i n g t h e i n t e r t i d a l and u p p e r s u b t i d a l f l o r a and f a u n a . . T h e s e e f f e c t s o f d r i f t l o g damage were n o t e d by Dayton (1971) i n t h e i n t e r t i d a l s h o r e s o f W a s h i n g t o n and the San J u a n I s l a n d s . I n t h e summer t h e f r e s h e t o f t h e F r a s e r R i v e r s t r o n g l y i n f l u e n c e s t h e s a l i n i t y o f t h e S t r a i t o f G e o r g i a c a u s i n g a marked d r o p o v e r t h e summer months i n t h e minimum s a l i n i t y v a l u e s . ( R e f e r t o A p p e n d i x I I I f o r m o n t h l y s a l i n i t y and t e m p e r a t u r e d a t a f o r t h e S t r a i t of G e o r g i a . ) Extreme low t i d e s i n t h e summer o c c u r a t Bath I s l a n d i n t h e m i d d l e o f t h e day (Anon, 1977,1978,1979), and t h e A . j a p o n i c u s p l a n t s c a n be e x p o s e d f o r up t o 5 h o u r s a t midday b e f o r e b e i n g r e i m m e r s e d . The smooth r o c k s h e l f o f t h e i n t e r t i d a l r e g i o n a t t h i s s i t e p r o v i d e d l i t t l e r e l i e f f r o m d i r e c t sun o r wave a c t i o n a s t h e r e a r e few c r a c k s , c r e v i c e s o r shade a r e a s and t h u s l i t t l e h a b i t a t v a r i a t i o n . 61 At Bath I s l a n d t h e ft.japonicus p o p u l a t i o n was d i s t r i b u t e d t h r o u g h t h e i n t e r t i d a l r e g i o n from t h e upper m i d t i d a l t o t h e upper s u b t i d a l - a v e r t i c a l r a n g e o f 2.75 m e t r e s . The f i e l d s i t e s a t A g u i l a r House P o i n t and Second Beach were e x p o s e d t o s u r f a c t i o n y e a r r o u n d a l t h o u g h t h e i n c i d e n c e o f s e v e r e s t o r m s was h i g h e r i n t h e w i n t e r months. (No m i d - w i n t e r c o l l e c t i o n s were made a t Second Beach as w i n t e r s t o r m s p r e v e n t e d b o a t l a n d i n g and a c c e s s t o t h e c o l l e c t i o n s i t e . ) B o t h s i t e s were c h a r a c t e r i s e d by c o n s i d e r a b l e t o p o g r a p h i c r e l i e f - c r e v i c e s , c r a c k s , r o c k f a c e s w i t h v a r y i n g s l o p e s , s u r f a c e s t r u c t u r e and a s p e c t . They d i f f e r e d m a r k e d l y i n t h i s f r o m t h e f i e l d s i t e a t B a t h I s l a n d . On the west c o a s t , i n summer t h e extreme low t i d e s o c c u r i n t h e e a r l y m o r n i n g . The i n t e r t i d a l r e g i o n s where A . j a p o n i c u s o c c u r s a r e r e i m m e r s e d by mi d - m o r n i n g . . I n summer and e a r l y autumn on t h e west c o a s t , m o r n i n g f o g i s a common, a l m o s t d a i l y , o c c u r r e n c e . . The f o g s e r v e s t o moderate t h e e f f e c t o f t h e u n c o v e r i n g o f t h e i n t e r t i d a l r e g i o n a t low t i d e . The s a l i n i t y and t e m p e r a t u r e o f t h e west c o a s t w a t e r s a r e not s u b j e c t t o t h e extreme s e a s o n a l f l u c t u a t i o n s t h a t t h e w a t e r s o f t h e S t r a i t o f G e o r g i a e x p e r i e n c e . . ( R e f e r t o A p p e n d i x IV f o r s a l i n i t y and t e m p e r a t u r e d a t a f o r t h e west c o a s t o f V a n c o u v e r I s l a n d . ) At b o t h A g u i l a r House P o i n t and Second B e a c h , A . j a p o n i c u s was p r e s e n t on r o c k y h e a d l a n d s , w i t h H e d p p h y l l u m s e s s i l e ( C . A g . ) S e t c h . o c c u r r i n g j u s t below t h e A . j a p o n i c u s and below t h a t , i n t h e l o w e r r e a c h e s o f t h e 62 i n t e r t i d a l , L e s s o n i p g s i s l i t t o r a l i s ( T i l d . ) R e i n k e , b o t h p l a n t s c h a r a c t e r i s t i c o f e x p o s e d h a b i t a t s . The v e r t i c a l e x t e n t o f t h e A . j a p o n i c u s zone i n b o t h west c o a s t s i t e s was more l i m i t e d t h a n t h a t a t B a t h I s l a n d , r a n g i n g n o t more t h a n 0.75 m, b e i n g f o u n d i n t h e mid-upper i n t e r t i d a l and n o t e x t e n d i n g i n t o t h e l o w e r r e a c h e s o f t h i s r e g i o n . The west c o a s t s i t e s were c h a r a c t e r i s e d by a more d i v e r s e f l o r a and f a u n a t h a n p r e s e n t i n t h e S t r a i t o f G e o r g i a . No p l a n t s c f A . j a p o n i c u s b e a r i n g u n i l o c u l a r s p o r a n g i a were f o u n d i n t h e B r i t i s h C o l u m b i a n c o l l e c t i o n s . E r e c t a x e s were p r e s e n t i n a l l months o f t h e y e a r g r o w i n g from t h e c r u s t s o f A . j a p o n i c u s a l t h o u g h n o t a t a l l s i t e s . P l a n t s b e a r i n g p l u r a n g i a were a l s o f o u n d y e a r r o u n d , a l t h o u g h not a t a l l s i t e s . I n F i g u r e s 14, 15, and 16 t h e r e s u l t s o f t h e p h e n o l o g i c a l s u r v e y f o r mean a x i s h e i g h t and f o r f e r t i l i t y a r e p r e s e n t e d f o r t h e A . j a p o n i c u s p o p u l a t i o n s f r o m B a t h I s l a n d , A g u i l a r House P o i n t and Second Beach, r e s p e c t i v e l y . The summary p r o v i d e d by T a b l e I g i v e s d e s c r i p t i v e m e a s u r e s of t h e sa m p l e s of B a t h I s l a n d , A g u i l a r House P o i n t and Second Beach, r e s p e c t i v e l y , c o l l e c t e d f r o m August 1978 t o J u l y 1979.. J . J . C o l l e c t i o n s and O b s e r v a t i o n s o u t s i d e B r i t i s h C o l u m b i a A l l p l a n t s t h a t were c o l l e c t e d o u t s i d e B r i t i s h C o l u m b i a were e i t h e r s t e r i l e o r p l u r i l o c u l a r e x c e p t f o r a s m a l l p o p u l a t i o n o f u n i l o c u l a r p l a n t s f o u n d a t C i t y P o i n t 63 C a l i f o r n i a . Dr. Lynda G o f f c o l l e c t e d an i n i t i a l s e t o f p l a n t s from C i t y P o i n t , C a l i f o r n i a f o r me, a s the o n l y two u n i l o c u l a r p l a n t s c o l l e c t e d i n N o r t h A m e r i c a t h a t I had s e e n were from t h a t s i t e . I c o l l e c t e d a t C i t y P o i n t two weeks a f t e r t h e f i r s t c o l l e c t i o n by Dr.. G o f f . The u n i l o c u l a r p o p u l a t i o n was s m a l l and v e r y r e s t r i c t e d i n i t s e x t e n t g r o w i n g on t h e f a c e s o f some l a r g e and a p p a r e n t l y unmcving b o u l d e r s . The p l u r i l o c u l a r p o p u l a t i o n was e x t e n s i v e a t C i t y P o i n t and a p p e a r e d t o be o c c u p y i n g a g r e a t e r v e r t i c a l e x t e n t a t t h i s s i t e t h a n t h e u n i l o c u l a r p l a n t s . A f e a t u r e o f C i t y P o i n t and many o f t h e s i t e s v i s i t e d i n C a l i f o r n i a and Oregon was t h e l a r g e amount o f s a n d p r e s e n t n e a r b y t h e r o c k s h e l v e s o r h e a d l a n d s on w h i c h A . j a p o n i c u s was g r o w i n g . T h i s was p a r t i c u l a r l y n o t i c e a b l e i n September i n C a l i f o r n i a when, a t a number o f s i t e s v i s i t e d , r o c k s w i t h A . j a p o n i c u s were p a r t i a l l y c o v e r e d by s a n d . In p l a c e s t h e c r u s t o s e b a s a l s y s t e m had been c o v e r e d c o m p l e t e l y w i t h o n l y t h e upper p o r t i o n o f t h e u p r i g h t a x i s and l a t e r a l s p r o t r u d i n g t h r o u g h t h e d e p o s i t e d s e d i m e n t s . . On t h e March c o l l e c t i n g t r i p e r e c t a x e s o f A . j a p o n i c u s were p r e s e n t a t v i r t u a l l y a l l s i t e s where c r u s t s o f t h e p l a n t were f o u n d . A . j a p o n i c u s i s a c h a r a c t e r i s t i c , and sometimes p r o m i n e n t , component o f t h e i n t e r t i d a l , i n g e n e r a l o c c u r r i n g i n t h e m i d d l e i n t e r t i d a l r e g i o n s . . O n t h e September c o l l e c t i n g t r i p , e r e c t axes o f A . j a p o n i c us were f o u n d a t a l l s i t e s where t h e c r u s t o s e p o r t i o n was f o u n d 64 e x c e p t f o r Moss Beach. A v e r a g e p l a n t s i z e was g r e a t e r i n September t h a n i n March. At b o t h t i m e s a l l t h e p o p u l a t i o n s sampled w i t h e r e c t a x e s p o s s e s s e d p l u r i l o c u l a r p l a n t s . 2. H e r b a r i u m S u r v e y R e s u l t s o f t h e h e r b a r i u m s u r v e y o f A . j a p o n i c u s a r e g i v e n i n F i g u r e 17 ( e x c e p t f o r t h e s i n g l e s p e c i m e n f r o m Kamschatka - which was p l u r i l o c u l a r and c o l l e c t e d i n A u g u s t ) . The r e s u l t s , p r e s e n t e d as h i s t o g r a m s by r e g i o n s i n d i c a t e t h a t t h e r e i s n o t an even d i s t r i b u t i o n o f h e r b a r i u m m a t e r i a l from t h r o u g h o u t t h e y e a r f o r a l l p a r t s of i t s r a n g e . T h i s was a p r o b l e m f o r m a t e r i a l from A l a s k a - n o t s u r p r i s i n g g i v e n t h e r i g o r o u s p h y s i c a l c l i m a t e and d i f f i c u l t y o f a c c e s s f o r c o l l e c t i n g - but a l s o f o r m a t e r i a l f r c m Oregon and W a s h i n g t o n . T h i s can be a t t r i b u t e d t o an a b s e n c e o f c o l l e c t i o n s r a t h e r t h a n t o a b s e n c e o f p l a n t m a t e r i a l i n t h e f i e l d b e c a u s e t h i s p l a n t d o e s p e r e n n a t e by means o f t h e c r u s t o s e base. The f i v e s p e c i m e n s c o n s i s t i n g o f o n l y t h e base were d i s t r i b u t e d as f o l l o w s : - J a p a n (1- September) , B r i t i s h C o l u m b i a (1-May), C a l i f o r n i a (1-March, 2-May). In s u r v e y i n g t h e s p e c i m e n s from t h r o u g h o u t i t s g e o g r a p h i c d i s t r i b u t i o n , t h e r a n g e o f p l a n t s i z e e n c o u n t e r e d was c o n s i d e r a b l e . The l a r g e s t p l a n t s s e e n were c o l l e c t e d on San J u a n I s l a n d (Washington) - t h e s e r e a c h e d a l e n g t h of 70 cm w i t h l a t e r a l s up t o 6 cm and were p l u r i l o c u l a r . The s m a l l e s t f e r t i l e p l a n t s ( p l u r i l o c u l a r ) 65 were l e s s t h a n 2 cm i n h e i g h t w i t h l a t e r a l s o f 1-2 mm* U n i l o c u l a r s p e c i m e n s f r o m N o r t h A m e r i c a were r e s t r i c t e d t o two l o c a l i t i e s , b o t h v e r y c l o s e l y s i t u a t e d on o p p o s i t e s i d e s c f a bay - P o i n t L o b o s and C i t y P o i n t . D i s c u s s i o n The p o p u l a t i o n s o f A . j a p o n i c u s t h a t were s t u d i e d i n B r i t i s h C o l u m b i a were f o u n d t o have c e r t a i n f e a t u r e s i n common. The p o p u l a t i o n s showed g r e a t v a r i a b i l i t y i n p l a n t s i z e a t v i r t u a l l y a l l t i m e s of t h e y e a r . ( W i t n e s s i n g t o t h i s a r e t h e l a r g e s t a n d a r d d e v i a t i o n s f o r t h e v a r i a b l e s mean a x i s h e i g h t , l a t e r a l l e n g t h and l a t e r a l w i d t h - F i g u r e s 14, 15, and 16). E r e c t axes were f o u n d t o be p r o d u c e d a t a l m o s t a l l s i t e s v i s i t e d t h r o u g h o u t t h e y e a r and t h u s a x i s i n i t i a t i o n i s n o t s y n c h r o n o u s o r c o r r e l a t e d w i t h p a r t i c u l a r e n v i r o n m e n t a l c u e s ( s u c h as d a y l e n g t h o r t e m p e r a t u r e ) . . The r e s u l t s o f c u l t u r e work r e p o r t e d i n C h a p t e r 3 a r e a l s o c o n s i s t e n t w i t h t h i s o b s e r v a t i o n . However, t h e e x t e n t t o which t h e e r e c t a x e s d e v e l o p a t v a r i o u s s i t e s and t h e d y n a m i c s o f t h e p o p u l a t i o n s s t u d i e d ^ d i f f e r e d m a r k e d l y . I t seems l i k e l y t h a t the v a r i a t i o n s i n p a t t e r n s o f growth and r e p r o d u c t i o n a t t h e d i f f e r e n t s i t e s a r e l a r g e l y due t o d i f f e r i n g p h y s i c a l and c h e m i c a l r e g i m e s e n c o u n t e r e d a t t h e s e s i t e s . The f o l l o w i n g i s my i n t e r p r e t a t i o n o f t h e e v e n t s a t 66 B a t h I s l a n d , r e f e r r i n g i n p a r t i c u l a r t o F i g u r e 14. The d e c r e a s e i n mean a x i s h e i g h t which o c c u r r e d i n b o t h y e a r s i n F e b r u a r y / M a r c h can be a t t r i b u t e d t o i n c r e a s e d wave a c t i o n and p o s s i b l y s t o r m damage ( l o g movement) i n t h e w i n t e r . The l a r g e s t p l a n t s i n t h e p o p u l a t i o n a r e p r o b a b l y t h e most v u l n e r a b l e t o t e a r i n g o r damage b e c a u s e o f t h e i r g r e a t e r d r a g and r e s i s t a n c e i n t h e w a t e r . I n s p r i n g and e a r l y summer t h e p l a n t s grew r a p i d l y , p r o b a b l y due t o i n c r e a s i n g l i g h t , d a y l e n g t h , and warmer t e m p e r a t u r e s . . T h e a b r u p t d e c r e a s e i n a x i s h e i g h t i n J u n e - J u l y c a n be r e l a t e d t o a c o m b i n a t i o n o f f a c t o r s . The two most i m p o r t a n t f a c t o r s a p p e a r t o be t h e l o w e r e d s a l i n i t y i n t h e S t r a i t o f G e o r g i a (due t o t h e i n f l u e n c e o f t h e F r a s e r R i v e r f r e s h e t ) , and t h e t i m i n g o f t h e e x t r e m e low water t i d e s ( l e a v i n g t h e p l a n t s o u t o f water f o r up t o 5-6 h o u r s i n t h e h o t t e s t p a r t o f t h e day e x p o s e d t o b o t h t h e sun and w i n d ) . The marked d e c l i n e i n t h e g e n e r a l c o n d i t i o n o f t h e B a t h I s l a n d A . j a p o n i c u s p o p u l a t i o n can be a t t r i b u t e d t o t h i s c o m b i n a t i o n o f s t r e s s e s . ( R e f e r t o C h a p t e r 4 f o r more d e t a i l a b o u t t h i s phase o f t h e s e a s o n a l growth.) A l t h o u g h t h e p o p u l a t i o n was c l e a r l y i n a s t a t e o f d e c l i n e i n l a t e summer, e r e c t a x e s were s t i l l b e i n g p r o d u c e d ; t h e i r d e v e l o p m e n t , however, was a p p a r e n t l y b e i n g i n h i b i t e d by t h e p r e v a i l i n g e n v i r o n m e n t a l r e g i m e . By O c t o b e r and November t h e p o p u l a t i o n had r e c o v e r e d , w i t h t h e mean a x i s h e i g h t a g a i n i n c r e a s i n g . At t h i s t i m e o f y e a r , however, a v a i l a b l e l i g h t and t e m p e r a t u r e a r e d e c r e a s i n g and s o a l t h o u g h t h e p l a n t s a r e g r o w i n g and 67 r e p r o d u c i n g t h e maximal growth r a t e s f o u n d i n s p r i n g and e a r l y summer a r e n o t e n c o u n t e r e d . At A g u i l a r House P o i n t ( F i g u r e 15), t h e p o p u l a t i o n d y n a m i c s o f A . j a p o n i c u s , a s measured by mean a x i s h e i g h t o r by t h e p e r c e n t a g e o f t h e p l a n t s b e a r i n g p l u r a n g i a , were f o u n d t o be d i f f e r e n t from t h o s e j u s t d i s c u s s e d f o r B a t h I s l a n d . T h e r e was no summer d e c l i n e i n t h e A g u i l a r House P o i n t p o p u l a t i o n s - i n f a c t , t h e p o p u l a t i o n showed an i n c r e a s e i n mean a x i s h e i g h t a t t h e same t i m e t h a t t h e B a t h I s l a n d p o p u l a t i o n was a t i t s l o w e s t ebb. At A g u i l a r House P o i n t , however, t h e r e was a d e c l i n e i n t h e mean a x i s h e i g h t o v e r t h e w i n t e r months, and I a t t r i b u t e t h i s t o t h e e f f e c t s o f t h e r i g o r o u s p h y s i c a l e n v i r o n m e n t and s t o r m and wave damage* The t i d a l c y c l e , s p e c i f i c a l l y , t h e t i m i n g o f t h e low t i d e s i n summer, and t h e t o p o g r a p h i c v a r i a t i o n a t t h e s i t e , r e s u l t e d i n much l e s s s t r e s s f u l summer c o n d i t i o n s t h a n f o u n d a t E a t h I s l a n d . T h u s , summer c o n d i t i o n s a t t h i s west c o a s t s i t e were a p p a r e n t l y f a v o u r a b l e t o growth o f A . j a p o n i c u s and a l l o w e d t h e p o p u l a t i o n t o r e a c h i t s h i g h e s t l e v e l s o f f e r t i l i t y . The s e a s o n a l p a t t e r n o c c u r r i n g a t Second Beach ( F i g u r e 16) a p p e a r s t o be v e r y s i m i l a r t o t h a t a t A g u i l a r House P o i n t - showing t h e b e s t growth i n t h e summertime. U n f o r t u n a t e l y , due t o t h e a b s e n c e o f m i d - w i n t e r d a t a f o r t h i s s i t e , a c o m p l e t e p i c t u r e o f t h e growth o f t h i s p o p u l a t i o n c a n n o t be c o n s t r u c t e d . The i n t e r p r e t a t i o n p r e s e n t e d h e r e c o r r e l a t e s w e l l w i t h t h e p o p u l a t i o n v a r i a t i o n s o b s e r v e d i n t h e c o u r s e o f t h i s 68 s t u d y . I t c o u l d a l s o h e l p t o e x p l a i n t h e a p p a r e n t l y c o n t r a d i c t o r y r e p o r t s on ft.japonicus p h e n o l o g y i n t h e l i t e r a t u r e which may be m e r e l y r e f l e c t i n g t h e a b i l i t y o f t h i s p l a n t t o grow i n v a r i e d h a b i t a t s and i t s d i f f e r i n g r e s p o n s e t o d i f f e r e n t p h y s i c a l r e g i m e s . T h i s i n t e r p r e t a t i o n c o u l d be t e s t e d by a number o f p r o c e d u r e s p o s s i b l y t h e most d i r e c t o f which would be r e c i p r o c a l t r a n s p l a n t e x p e r i m e n t s between p o p u l a t i o n s o f A . j a p o n i c u s f r o m t h e S t r a i t o f G e o r g i a and west c o a s t s i t e s . A number o f w o r k e r s have r e c o r d e d t h e v e r t i c a l e l e v a t i o n o f i n t e r t i d a l p l a n t s and a n i m a l s f r o m a l l z o n e s o f t h e s h o r e w i t h i n c r e a s i n g wave a c t i o n ( e . g . Moore, 1935; J o n e s and D e m e t r o p o u l o s , 1968; Morton and M i l l e r , 1968). A l t h o u g h a t some s i t e s t h e v e r t i c a l p o s i t i o n o f A . j a p o n i c u s a p p e a r s t o be r a i s e d by e x p o s u r e t o wave a c t i o n ( e . g . a t B o t a n y B e a c h ) , I do n o t t h i n k t h a t t h e v e r t i c a l r a n g e o f t h i s p l a n t i n t h e i n t e r t i d a l r e g i o n c a n be e n t i r e l y e x p l a i n e d by r e l a t i v e e x p o s u r e t o wave a c t i o n . The p r e s e n c e of A . j a p o n i c u s from h i g h t o low i n t e r t i d a l z o n e s shows t h i s p l a n t t o have c o n s i d e r a b l e e c o l o g i c a l f l e x i b i l i t y . . I n B r i t i s h C o l u m b i a , A . j a p o n i c u s showed t h e g r e a t e s t d i s t r i b u t i o n w i t h i n t h e i n t e r t i d a l r e g i o n i n the S t r a i t o f G e o r g i a . On the west c o a s t , t h e e x t e n t o f t h e i n t e r t i d a l d i s t r i b u t i o n o f t h i s p l a n t was r a t h e r l i m i t e d . I n t h o s e i n t e r t i d a l o r g a n i s m s where v e r t i c a l l i m i t s o f d i s t r i b u t i o n have been s t u d i e d , b o t h p h y s i c a l and b i o l o g i c a l f a c t o r s have been shown t o be i m p o r t a n t . I t i s 69 g e n e r a l l y u n d e r s t o o d t h a t t h e e f f e c t s o f t h e p h y s i c a l e n v i r o n m e n t p r e d o m i n a t e i n t h e u p p e r r e a c h e s o f t h e i n t e r t i d a l , and a s t h e p h y s i c a l e n v i r o n m e n t becomes r e l a t i v e l y l e s s h a r s h l o w e r i n t h e i n t e r t i d a l r e g i o n , b i o l o g i c a l f a c t o r s become i n c r e a s i n g l y i m p o r t a n t i n d e t e r m i n i n g t h e s p e c i e s n i c h e w i d t h ( C o n n e l l , 1972; Chapman, 1973a, 1974; L u b c h e n c o , 1980). F o r a l g a e , t h e main f a c t o r s i m p l i c a t e d i n d e t e r m i n i n g t h e l o w e r l i m i t s o f a s p e c i e s * v e r t i c a l d i s t r i b u t i o n a r e c o m p e t i t i o n and h e r b i v o r y (Dayton, 1975; Hruby, 1976; L u b c h e n c o , 1980)..One e x p l a n a t i o n f o r t h e d i f f e r e n c e s d e s c r i b e d h e r e i n t h e v e r t i c a l r a n g e o f A . j a p o n i c u s i n B r i t i s h C o l u m b i a , i s t h a t A . j a p o n i c u s i s c o m p e t i t i v e l y more s u c c e s s f u l i n o b t a i n i n g s u b s t r a t e ( s p a c e b e i n g t h e r e s o u r c e which i s s o c r i t i c a l t o t h e s u c c e s s o f s e s s i l e i n t e r t i d a l i n h a b i t a n t s ) i n t h e l e s s d i v e r s e f l o r a and f a u n a o f t h e S t r a i t o f G e o r g i a . . I n t h e S t r a i t o f G e o r g i a few r e p r e s e n t a t i v e s o f t h e o r d e r L a m i n a r i a l e s a r e f o u n d . At west c o a s t s i t e s , t h e k e l p zone which o c c u r r e d below t h a t o f A . j a p o n i c u s i s w e l l d e v e l o p e d and p r o v i d e s a canopy i n t h e l o w e r i n t e r t i d a l . P o s s i b l y t h e s h a d e c a s t by t h e s e p l a n t s , a w h i p l a s h a f f e c t o f t h e b l a d e s moving w i t h wave a c t i o n , or g r e a t e r a b i l i t y o f t h e s e o r o t h e r a l g a e o r s e s s i l e a n i m a l s t o c o l o n i s e and m a i n t a i n o c c u p a n c y o f s u b s t r a t e i n h i b i t s t h e e x t e n s i o n o f A . j a p o n i c u s i n t o t h e l o w e r r e a c h e s o f t h e i n t e r t i d a l r e g i o n a t t h e s e s i t e s . I t would be v e r y i n t e r e s t i n g t o t e s t e x p e r i m e n t a l l y t h e r o l e o f c o m p e t i t i o n i n d e t e r m i n i n g 70 A. j a p o n i c u s ' l o w e r i n t e r t i d a l l i m i t s ( e . g . by s e l e c t i v e r e m o v a l o f c e r t a i n s p e c i e s , by s u b s t r a t e c l e a r a n c e o r by t r a n s p l a n t i n g A . j a p o n i c u s from o t h e r p a r t s o f i t s d i s t r i b u t i o n and v e r t i c a l r a n g e ) . V a r y i n g c o m p e t i t i v e s u c c e s s a t d i f f e r e n t s i t e s would go some way t o e x p l a i n i n g t h e a p p a r e n t l y c o n t r a d i c t o r y comments i n t h e l i t e r a t u r e a b o u t t h e h a b i t o f t h i s p l a n t . The m o r p h o l o g y o f A . j a p o n i c u s s p e c i m e n s s u r v e y e d i n t h e h e r b a r i u m s t u d y and i n t h e m a t e r i a l which I c o l l e c t e d was f o u n d t o v a r y c o n s i d e r a b l y . When p l a n t s which a r e 70 cm t a l l , w i t h l a t e r a l s up t o 6 cm i n l e n g t h and b e a r i n g p l u r a n g i a ( e . g . f r o m San J uan I s l a n d , W a s h i n g t o n , i n June) a r e p l a c e d n e x t t o p l a n t s w i t h e r e c t a x e s l e s s t h a n 2 cm i n h e i g h t w i t h l a t e r a l s o f b a r e l y 2 mm ( e . g . . f r o m B o t a n y Beach, V a n c o u v e r I s l a n d , i n March) and a l s o b e a r i n g p l u r a n g i a , one i s c o n f r o n t e d v e r y g r a p h i c a l l y w i t h t h e e x t r e m e s o f t h i s v a r i a t i o n . The morphology a p p e a r s t o be a f f e c t e d by t h e h a b i t a t e x p e r i e n c e d and t h e t h r e e B. C. p o p u l a t i o n s which were s t u d i e d r e f l e c t a c e r t a i n amount o f t h i s v a r i a t i o n ( e . g . i n t h e maximum h e i g h t a t t a i n e d by t h e e r e c t a x e s i n t h e s ample, t h e d e n s i t y o f l a t e r a l b r a n c h i n g and t h e l e n g t h and w i d t h of l a t e r a l s T a b l e I ) . One c h a r a c t e r i s t i c which was f o u n d t o be v e r y c o n s i s t e n t was t h e a x i s number. Where b r a n c h e d axes o c c u r r e d i t a p p e a r e d t o be t h e r e s u l t o f damage t o t h e apex r e s u l t i n g i n one o r more l a t e r a l s t a k i n g o v e r t h e a p i c a l f u n c t i o n and i n t u r n d e v e l o p i n g l a t e r a l s . . 71 No d e f i n i t i v e s t a t e m e n t s c a n be made a b o u t t h e p h e n o l o g y o f A . j a p o n i c u s b a s e d s o l e l y on t h e h e r b a r i u m m a t e r i a l examined h e r e . .From t h e d a t a a v a i l a b l e , however, b o t h from my c o l l e c t i o n s and t h e h e r b a r i u m s u r v e y , i t a p p e a r s t h a t t h e p h e n o l o g i c a l p a t t e r n s a r e s i m i l a r l y v a r i e d i n e t h e r p a r t s o f A . j a p o n i c u s 1 r a n g e . The o c c u r r e n c e o f s p e c i m e n s w i t h e r e c t a x e s from C a l i f o r n i a c o l l e c t e d i n t e n months o f t h e y e a r ( w i t h p l u r i l o c u l a r p l a n t s i n a l l o f t h e s e months) shows t h a t t h e p l a n t s do n o t e x h i b i t e x c l u s i v e l y t h e p h e n o l o g i c a l p a t t e r n g i v e n i n A b b o t t and H o l l e n b e r g (1976) a l t h o u g h t h i s may be t h e p r e d o m i n a n t one. The most u n e x p e c t e d f i n d i n g o f t h i s p h e n o l o g i c a l s t u d y was t h e d i s t r i b u t i o n o f r e p r o d u c t i v e s t r u c t u r e s and t h e a b s e n c e o f u n i l o c u l a r s p o r a n g i a ( o r a t l e a s t t h e i r e x t r e m e r a r i t y ) o v e r t h e m a j o r i t y o f t h e r a n g e o f t h i s p l a n t . The work p r e s e n t e d i n C h a p t e r 3 shows t h a t t h e p l u r i l o c u l a r p l a n t s a r e h a p l o i d , b u t a p p a r e n t l y t h e swarmers f r o m t h e p l u r a n g i a a r e n o t f u n c t i o n i n g a s gametes o v e r the m a j o r i t y o f t h e r a n g e o f t h e p l a n t . F o r each o f t h e t h r e e s i t e s s t u d i e d i n B r i t i s h C o l u m b i a t h e p r o p o r t i o n o f p l a n t s b e a r i n g p l u r a n g i a i n t h e p o p u l a t i o n s a m p l e s c l o s e l y f o l l o w e d t h e t r e n d s shown by mean a x i s h e i g h t . The h i g h i n c i d e n c e o f f e r t i l e p l a n t s i n t h e f i e l d and i n t h e h e r b a r i u m m a t e r i a l examined, c o u p l e d w i t h t h e o b s e r v a t i o n ( C h a p t e r 1) t h a t each p l u r i l o c u l a r p l a n t p r o d u c e s a v e r y l a r g e number o f r e p r o d u c t i v e u n i t s , i n d i c a t e s t h a t t h e r e i s a c o n s i d e r a b l e i n v e s t m e n t o f e n e r g y 72 i n r e p r o d u c t i o n . The r a p i d d i e b a c k of t h e p o s t - r e p r o d u c t i v e p l a n t s f u r t h e r e m p h a s i s e s t h e p r i m a r y r o l e of r e p r o d u c t i o n f o r t h e e r e c t a x e s of A . j a p o n i c u s . 73 F i g u r e 12. Map i n d i c a t i n g c o l l e c t i o n s i t e s i n B r i t i s h C o l u m b i a , W a s h i n g t o n , Oregon, and C a l i f o r n i a . EEITISH COLOMEIA:- 1) Bath I s l a n d 2) B r e a k w a t e r I s l a n d 3) A g u i l a r House P o i n t 4) B r a d y ' s Beach 5) D i a n a I s l a n d 6) H a i n e s I s l a n d 7) H e l b y I s l a n d 8) Boss I s l e t s 9) S e p p i n g s I s l a n d 10) Second Beach 11) W i z a r d Bock 12) Pachena Bay 13) Botany Beach 14) Sombrio E i v e r 15) Sooke 16) V i c t o r i a B r e a k w a t e r 17) S i d n e y WASHINGTON:- 18) C a t t l e P o i n t , San Juan I s l a n d 19) Mar V i s t a , San J u a n I s l a n d 20) K a l a l o c h CEEGON: - 21) F o g a r t y C r e e k 22) B o i l e r Bay 23) M a r i n e G a r d e n s 24) D e v i l ' s Punchbowl 25) Y a g u i n a Head 26) Cape Arago 27) S u n s e t Beach 28) H a r r i s Beach CALIFORNIA:- 29) T r i n i d a d P o i n t 30) W e s t p o r t 31) P o i n t A r ena 32) Anchor Eay 33) Moss Beach 34) P e s c a d e r o Beach 3 5) P e b b l e Beach 36) P i g e o n E c i n t 37) D a v e n p o r t L a n d i n g 38) P o i n t J o e 39) P o i n t P i n e s 40) S t i l l w a t e r Cove 41) C i t y ( M i s s i o n ) P o i n t 42) P i e d r a s E l a n c a s 43) Montana de Oro  75 F i g u r e 1 3 . Map o f s o u t h e r n s i d e o f B a r k l e y Sound i n d i c a t i n g l o c a t i o n s o f s t u d y s i t e s n e a r t h e B a m f i e l d M a r i n e S t a t i o n .  77 F i g u r e 14. Mean a x i s h e i g h t and p e r c e n t a g e o f p l u r i l o c u l a r p l a n t s i n c o l l e c t i o n s o f A . j a p o n i c u s f r o m B a t h I s l a n d between J u l y 1977 and August 1979. E r r o r b ar i n d i c a t e s one s t a n d a r d d e v i a t i o n o f mean.  79 F i g u r e 15. Mean a x i s h e i g h t and p e r c e n t a g e o f p l u r i l o c u l a r p l a n t s i n c o l l e c t i o n s of A. .ja p o n i c u s from A g u i l a r House P o i n t between August 1978 and August 1 9 7 9 . . E r r o r b a r i n d i c a t e s one s t a n d a r d d e v i a t i o n of mean.  81 F i g u r e 16. Mean a x i s h e i g h t and p e r c e n t a g e o f p l u r i l o c u l a r p l a n t s i n c o l l e c t i o n s o f A . j a p o n i c u s f r o m Second Beach between May 1978 and August 1979. E r r o r b a r i n d i c a t e s one s t a n d a r d d e v i a t i o n o f mean.  83 F i g u r e 1 7 . H e r b a r i u m s u r v e y r e s u l t s s h o w i n g numbers o f s t e r i l e , u n i l o c u l a r , and p l u r i l o c u l a r s p e c i m e n s examined f o r e a c h month f o r s i x r e g i o n s o f A . j a p o n i c u s 1 r a n g e . Alaska n = 93 British Columbia n r!91 Washington n = 33 Oregon JZl n r !9 9, California n =43 - Japan n sl7 J F M A M J J A S O N D Months J F M A M J J A S O N D = sterile = plurilocular = unilocular TABLE I:- Descriptive measures of A. japonicus sampled from August 1978 to July 1979 at 3 sites in British Columbia. (Refer to the text for measurement procedures.) BATH ISLAND VARIABLE SAMPLE SIZE MIN. MAX. MEAN STANDARD DEVIATION axis:- number 250 1.0 2.0 1.03 0.176 height(cm) 250 0.1 33.9 4.70 6.18 lat e r a l : - density 250 0 36 5.66 9.14 length(cm) 84 0.1 1.86 0.53 0.39 width(cm) 83 0.05 0.11 0.061 0.014 f e r t i l i t y 250 - 63% ste r i l e , 37% plurilocular AGUILAR HOUSE POINT VARIABLE SAMPLE SIZE MIN. MAX. MEAN STANDARD DEVIATION axis:- number 170 1.0 4.0 1.06 0.35 height(cm) 170 0.1 15.0 3.63 2.98 lat e r a l : - density 151 0 75 18.55 17.58 length(cm) 104 0.1 1.10 0.41 0.24 width(cm) 104 0.05 0.14 0.071 0.025 f e r t i l i t y 170 - 42% st e r i l e , 58% plurilocular SECOND BEACH VARIABLE SAMPLE SIZE MIN. MAX. MEAN STANDARD DEVIATION axis:- number 175 1.0 2.0 1.02 0.13 height(cm) 175 0.4 22.5 4.62 3.34 lat e r a l : - density 175 0 83 22.33 19.50 length(cm) 129 0.1 1.46 0.46 0.29 width(cm) 128 0.05 0.16 0.073 0.027 f e r t i l i t y 175 - 29% st e r i l e , 71% plurilocular 86 CHAPTER 3, L i f e h i s t o r y and c y t o l o g i c a l s t u d i e s o f A n a l i p u s j a p o n i c u s (Harv.) Wynne 87 I n t r o d u c t i o n P r i o r t o t h i s r e s e a r c h , t h e l i f e h i s t o r y o f A n a l i p u s j a p o n i c u s had n o t been o b s e r v e d i n c u l t u r e , and a l t h o u g h Abe, i n r e p o r t i n g t h e r e s u l t s o f c y t o l o g i c a l s t u d i e s on A . j a p o n i c u s i n J a p a n (Abe, 1935a,b; 1936), s t a t e d t h a t i t e x h i b i t s an a l t e r n a t i o n o f i s o m o r p h i c g e n e r a t i o n s , t h i s had n o t been f u l l y a c c e p t e d . B o l d and Wynne (1978) s u g g e s t e d t h a t Abe's work needed v e r i f i c a t i o n . F u r t h e r e m p h a s i s o f t h e need f o r a l i f e h i s t o r y s t u d y o f A . j a p o n i c u s i s g i v e n by t h e r e c e n t p l a c e m e n t o f t h i s p l a n t i n t h e H a l f s i a l e s (Nakamura, 1972; B o l d and Wynne, 1978). The l i f e h i s t o r y p a t t e r n i s one o f t h r e e c r i t e r i a used t o d i s t i n g u i s h t h i s o r d e r and y e t t o d a t e a c o m p l e t e l i f e h i s t o r y i n v o l v i n g s e x u a l i t y h as n o t been shown f o r any member of t h e o r d e r . The growth o f A . j a p o n i c u s i n c u l t u r e s t u d i e s h as been r e p o r t e d on by I k a r i (1928), Abe (1935a) and Wynne ( 1 9 7 2 ) , b u t t h e s e were r e s t r i c t e d t o t h e e a r l y s t a g e s o f d i s c g r o w t h . P l a n t s w i t h e r e c t a x e s have n o t been grown i n c u l t u r e , and t h u s t h e development o f r e p r o d u c t i v e l y mature p l a n t s h as n o t been o b s e r v e d i n c u l t u r e e i t h e r . Abe p u b l i s h e d t h r e e p a p e r s on t h e c y t o l o g y o f A . j a p o n i c u s . Abe (1935a) r e p o r t e d t h a t A . j a p o n i c u s e x h i b i t s an a l t e r n a t i o n o f i s o m o r p h i c g e n e r a t i o n s , w i t h d i o e c i o u s p l u r i l o c u l a r p l a n t s p r o d u c i n g swarmers w h i c h f u n c t i o n as gametes. He r e p o r t e d t h a t t h e f e m a l e swarmers a r e l a r g e r t h a n t h e male, and o n l y t h e f e m a l e swarmers c a n d e v e l o p 88 w i t h o u t f u s i o n . P r i o r t o f u s i o n , he a l s o r e p o r t e d t h a t t h e male gametes swarm a b o u t t h e f e m a l e gametes. Abe (1935a and b) r e p o r t e d t h a t swarmers from u n i l o c u l a r s p o r a n g i a c o u l d d e v e l o p w i t h o r w i t h o u t f u s i o n and when f u s i o n o c c u r r e d i t c o u l d be between swarmers from t h e same i n d i v i d u a l o r f r o m t h e two k i n d s o f u n i l o c u l a r p l a n t s which he r e c o g n i s e d . I k a r i (1928) d i d n o t o b s e r v e f u s i o n o f swarmers f r o m u n i l o c u l a r s p o r a n g i a . . Abe (1936) r e p o r t e d t h a t t h e u n i l o c u l a r s p o r a n g i a a r e t h e s i t e o f m e i o s i s i n A. " j a p o n i c u s and t h a t t h e u n i s p o r e s and t h e p l u r i l o c u l a r p l a n t s a r e h a p l o i d w i t h a chromosome number o f n=20.. He (1936) c o n c l u d e d t h a t A . j a p o n i c u s e x h i b i t s an a l t e r n a t i o n of i s o m o r p h i c g e n e r a t i o n s but t h a t when t h e swarmers f r o m u n i l o c u l a r o r g a n s f u s e , o r swarmers from p l u r i l o c u l a r o r g a n s d e v e l o p w i t h o u t f u s i o n , t h e a l t e r n a t i o n o f g e n e r a t i o n s i s s u p p r e s s e d . . K y l i n (19 37) n o t e d t h a t i f A . j a p o n i c u s d o e s have an a l t e r n a t i o n o f i s o m o r p h i c g c e n e r a t i o n s and t h e p l u r i l o c u l a r p l a n t s f u n c t i o n as g a m e t o p h y t e s , as s u g g e s t e d by Abe, t h i s would mean t h a t i t would h a v e t o be p l a c e d i n t h e E c t o c a r p a l e s and not i n t h e C h o r d a r i a l e s . However, K y l i n (1940) c o n t i n u e d t o p l a c e A . j a p o n i c u s (as H e t e r o c h o r d a r i a ) i n t h e C h o r d a r i a l e s , s u g g e s t i n g t h a t i t i s a n a l o g o u s t o C a e p i d i um a n t a r c t i c u m . He (1940) s t a t e d t h a t t h e p o s s i b i l i t y c a n n o t be d i s m i s s e d t h a t p l u r a n g i a o f b o t h t h e s e p l a n t s a r e d i p l o i d and f u n c t i o n a s v e g e t a t i v e r e p r o d u c t i v e s t r u c t u r e s - t h a t i s , t h e same p a t t e r n 89 e x h i b i t e d by o t h e r members o f t h e C h o r d a r i a l e s . I n t h i s c h a p t e r I d i s c u s s t h e r e s u l t s o f c u l t u r i n g and c y t o l o g i c a l o b s e r v a t i o n s on A . j a p o n i c u s . I wanted t o o b s e r v e t h e l i f e h i s t o r y o f A . j a p o n i c u s i n c u l t u r e , and t o d e t e r m i n e : (1) t h e e f f e c t o f t e m p e r a t u r e and p h o t o p e r i o d on t h e growth of t h e c r u s t o s e phase and t h e p r o d u c t i o n o f e r e c t a x e s ; (2) under what c o n d i t i o n s t h e p l a n t s become f e r t i l e ; (3) t h e r e s p o n s e o f t h e p l a n t s t o v a r i o u s s a l i n i t y and t e m p e r a t u r e r e g i m e s ; and, (4) t h e e f f e c t o f l i g h t g u a l i t y on g e r m i n a t i o n and e a r l y d e v e l o p m e n t . I a l s o wanted t o f i n d o u t i f d i f f e r e n t p o p u l a t i o n s o f p l u r i l o c u l a r A . j a p o n i c u s a r e i n t e r f e r t i l e and t o t e s t Abe's c l a i m t h a t A . j a p o n i c u s i s a s t r i c t l y d i o e c i o u s p l a n t , w i t h two d i s t i n g u i s h a b l e f o r m s o f p l u r i l o c u l a r p l a n t s c o r r e s p o n d i n g t o f e m a l e and male. In a d d i t i o n , I wanted t o d e t e r m i n e i f f u s i o n s o c c u r r e d between swarmers from u n i l o c u l a r s p o r a n g i a , a c o n t r o v e r s i a l c l aim-made by Abe {1935a,b) and i f I c o u l d a l s o r e c o g n i s e t h e two t y p e s o f u n i l o c u l a r p l a n t s d i s t i n g u i s h e d by Abe. I n o r d e r t o t e s t Abe's c l a i m t h a t A . j a p o n i c u s e x h i b i t s an a l t e r n a t i o n o f i s o m o r p h i c g e n e r a t i o n s , I n e e ded t o d e t e r m i n e i f t h e r e was an a l t e r n a t i o n o f c a r y o l o g i c a l g e n e r a t i o n s between u n i l o c u l a r and p l u r i l o c u l a r i n d i v i d u a l s . 90 M a t e r i a l s and Methods A. C u l t u r e Wgrk C u l t u r e s o f A. -japonicus were e s t a b l i s h e d u s i n g f e r t i l e f i e l d m a t e r i a l w i t h e i t h e r p l u r i l o c u l a r o r u n i l o c u l a r o r g a n s . P l u r i l o c u l a r p l a n t s , used t o i n i t i a t e c u l t u r e s , came from a number o f l o c a t i o n s : - B r i t i s h C o l u m b i a - A g u i l a r House P o i n t , E a t h I s l a n d , B otany Beach, D i a n a I s l a n d , H e l b y I s l a n d , Second Beach, Sooke; W a s h i n g t o n - C a t t l e P o i n t , Mar V i s t a ; Oregon - B o i l e r Bay, Cape A r a g o , F o g a r t y C r e e k , M a r i n e G a r d e n s , Y a g u i n a Head; C a l i f o r n i a - C i t y P o i n t ( M i s s i o n P o i n t ) , P o i n t P i n o s . The f i e l d m a t e r i a l w i t h u n i l o c u l a r s p o r a n g i a came o n l y f r o m C i t y P o i n t , C a l i f o r n i a . ( E e f e r t o A p p e n d i x I f o r c o - o r d i n a t e s o f c o l l e c t i o n s i t e s , and t o F i g u r e 12 f o r a map o f t h e c o l l e c t i o n range.) The i s o l a t i o n p r o c e d u r e u s u a l l y employed was t h e h a n g i n g - d r o p t e c h n i g u e o f Wynne (1969)..On some o c c a s i o n s when f i e l d m a t e r i a l was immersed i n s e a w a t e r upon a r r i v a l a t t h e l a b o r a t o r y , swarmer r e l e a s e would be e x t r e m e l y r a p i d and dense. In t h e s e c a s e s , d r o p s o f t h e swarmer s u s p e n s i o n were p i p e t t e d o n t o e i t h e r g l a s s s l i d e s o r c o v e r s l i p s , and t h e swarmers a l l o w e d t o s e t t l e i n a m o i s t chamber c r e a t e d by damp p a p e r t o w e l s s u r r o u n d i n g t h e s l i d e s on t h e bottom of a c l o s e d t r a y . The s l i d e s were r i n s e d u s i n g a s q u e e z e b o t t l e f i l l e d w i t h f i l t e r e d s e a w a t e r , and t h e n immersed i n c u l t u r e s o l u t i o n . C u l t u r e s were i n i t i a l l y s e t up i n 91 d i s p o s a b l e p o l y s t y r e n e p e t r i p l a t e s - e i t h e r 60 x 15 mm o r 100 x 20 mm. A f t e r t h e m a t e r i a l had d e v e l o p e d f o r 6-10 weeks, t h e c o v e r s l i p s o r s l i d e s were p l a c e d i n t o l i d d e d P y r e x c u l t u r e c o n t a i n e r s (#3250) w i t h ca..200 ml o f c u l t u r e medium. Both P r o v a s o l i ' s ES medium (West's m o d i f i c a t i o n o f P r o v a s o l i ' s (1968) medium g i v e n i n M c L a c h l a n (1973) was used) and a r t i f i c i a l s e a w a t e r (a m o d i f i c a t i o n of ESAW of H a r r i s o n , W a t e r s , and T a y l o r , 1980 - r e f e r t o A p p e n d i x V f o r t h e c o m p o s i t i o n of t h e e n r i c h m e n t s o l u t i o n ) were used i n t h e c u l t u r e s t u d y . The media were f i l t e r e d w i t h 0.45 pm p o r e - s i z e f i l t e r o r by g l a s s f i b e r f i l t e r (Reeve A n g e l ) . . E S medium was u s e d a l t e r n a t e l y w i t h n o n - e n r i c h e d f i l t e r e d s e a w a t e r as t h e m a i n t e n a n c e c u l t u r e media. The n a t u r a l s e a w a t e r used f o r t h e c u l t u r e medium was t a k e n e i t h e r f r o m t h e s e a w a t e r s y s t e m a t t h e B a m f i e l d M a r i n e S t a t i o n ( s a l i n i t y o f 29 - 32 % ) o r c o l l e c t e d n e a r B a t h I s l a n d i n t h e S t r a i t o f G e o r g i a ( s a l i n i t y of 18 - 26 %> ) . . The a d d i t i o n o f 2 ml o f a s t o c k s o l u t i o n o f 250 mg/L germanium d i o x i d e t o a l i t r e of c u l t u r e medium was used t o e l i m i n a t e d i a t o m c o n t a m i n a n t s f r o m t h e newly e s t a b l i s h e d c u l t u r e s ( L e w i n , 1966). The use o f germanium d i o x i d e was d i s c o n t i n u e d when t h e d i a t o m s had been e l i m i n a t e d . C o n t r o l c u l t u r e s w i t h o u t germanium d i o x i d e were s e t up t o c h e c k i f t h e germanium d i o x i d e had any i n h i b i t o r y e f f e c t (as has been r e p o r t e d by M c L a c h l a n , Chen and E d e l s t e i n , 197 1a; T a t e w a k i and M i z u n o , 1 9 7 9 ) . . A r t i f i c i a l s e a w a t e r was used i n 92 the s a l i n i t y experiments as i t enabled s o l u t i o n s to be prepared of known and r e p e a t a b l e s a l i n i t y . Another c u l t u r i n g method t r i e d was the growth of spore suspensions on agar (ES s o l i d i f i e d with 15? agar - Chapman, 1973b) or on agar with l i g u i d ES forming a l a y e r on top. C u l t u r e s were grown at a v a r i e t y of temperature and photoperiod regimes:- 5° 8:16, 5° 12:12, 10° 12:12, 10° 16:8, 15° 12:12, and, 20° 12:12 were used, although r o u t i n e l y c u l t u r e s were maintained at the f i r s t f o u r of these c o n d i t i o n s . The l i g h t i n g c o n d i t i o n s provided by C o o l White f l u o r e s c e n t tubes v a r i e d between 44- -2 -1 110 ^E .m .sec . (This v a r i a t i o n was due to bulb aging, d i f f e r e n t p o s i t i o n s of dishes w i t h i n the chambers, and d i f f e r e n t chambers. To minimise the e f f e c t s of l i g h t v a r i a t i o n dishes were moved around w i t h i n the chambers a t each medium change.) C u l t u r e s were grown with or without a g i t a t i o n i n t h e i r e a r l y stages. Due to l a c k of space and equipment, c u l t u r e s could not be grown or maintained r o u t i n e l y i n a g i t a t e d c o n d i t i o n s . 1) S a l i n i t y / T e m p e r a t u r e experiment C o v e r s l i p s with f r e s h l y - s e t t l e d p l u r i s p o r e s were placed i n t o c u l t u r e d i s h e s with 50 ml of c u l t u r e medium. A r t i f i c i a l seawater was made up to the f o l l o w i n g s a l i n i t i e s - 12. 5, 18. 5, 25, 30, 37%o - by v a r y i n g the amount of the s a l t s o l u t i o n s and d i s t i l l e d water (Harrison e t a l . , 1980). A modified enrichment medium was used ( r e f e r Appendix V) and the same amount of the enrichment medium was used i n 93 e a c h o f t h e f i v e d i f f e r e n t s o l u t i o n s . T h r e e r e p l i c a t e s f o r e a c h s a l i n i t y t r e a t m e n t were p l a c e d i n growth chambers a t 5° , 10° , 15° , and 20° C w i t h a p h o t o p e r i o d o f 12:12..The g r c w t h o f t h e c u l t u r e s was r e c o r d e d by p h o t o g r a p h i c s a m p l i n g a t 5, 7, 10, and 14 d a y s a f t e r commencement. A camera a t t a c h e d t o an Olympus i n v e r t e d m i c r o s c o p e was u s e d t o t a k e t h r e e p h o t o g r a p h s o f each c u l t u r e . 2) E f f e c t s o f 1 ig_ht q u a l i t y on g e r m i n a t i o n and g e r m l i n g growth C o v e r s l i p s w i t h f r e s h l y s e t t l e d p l u r i s p o r e s were p l a c e d i n p e t r i d i s h e s w i t h ES medium. The p e t r i d i s h e s were p l a c e d i n b o x e s c o n s t r u c t e d from " C i n e m o i d 1 1 c o l o u r e d a c e t a t e s h e e t i n g w i t h t h e f o l l o w i n g t r a n s m i s s i o n ' p r o p e r t i e s : - red(#14) 370-4 14 nm (minor c o m p o n e n t ) , 588-700 nm; g r e e n (#24) 444-590 nm; b l u e (#32) 370-564 nm, 640-700 nm ( m i n o r c o m p o n e n t ) . D i s h e s were a l s o p l a c e d i n t h e chamber w i t h o u t c o l o u r e d f i l t e r s b u t shaded by l a y e r s o f c o t t o n g a u z e . A l l - 2 -1 d i s h e s r e c e i v e d 32-35 j i E .m . s e c . T h r e e d i s h e s were s e t up i n e a c h o f t h e f o u r t r e a t m e n t s . The c u l t u r e s were grown a t 10° C,12:12 p h o t o p e r i o d , and t h e medium was c h a n g e d e v e r y 10-14 d a y s . Growth o f t h e s e c u l t u r e s was o b s e r v e d f o r 6 weeks. I n t h e i n i t i a l s t a g e s growth was f o l l o w e d by p h o t o g r a p h i c s a m p l i n g (as i n 1 above) . 3) F u s i o n o f swarmers The d i a m e t e r o f r e c e n t l y s e t t l e d swarmers from d i f f e r e n t p o p u l a t i o n s was measured. In a d d i t i o n t o h a n g i n g d r o p s w i t h t i s s u e from s i n g l e p l a n t s , s m a l l p i e c e s o f t i s s u e f r o m 94 l a t e r a l s o f two p l u r i l o c u l a r p l a n t s were used f o r a s i n g l e h a n g i n g d r o p . The p l a n t s used f o r t h e s e t w o - p l a n t h a n g i n g d r o p s were e i t h e r b o t h from t h e same s i t e o r f r o m d i f f e r e n t s i t e s . M a t e r i a l from t h e f o l l o w i n g s i t e s was t r i e d i n c o m b i n a t i o n : - E a t h I s l a n d / B a t h I s l a n d , A g u i l a r House P o i n t / A g u i l a r House P o i n t , Second B e a c h / S e c o n d B e a c h , B a t h I s l a n d / A g u i l a r House P o i n t , Bath I s l a n d / S e c o n d B e a c h , Cape Arago/Cape A r a g o , B a t h I s l a n d / C a p e Arago, F o g a r t y C r e e k / F o g a r t y C r e e k , Bath I s l a n d / F o g a r t y C r e e k , F o g a r t y C r e e k / Y a q u i n a Head, F o g a r t y C r e e k / B o i l e r Bay, C i t y P o i n t / C i t y P o i n t . The r e l e a s e d swarmers were o b s e r v e d as t h e y s e t t l e d and any swarmer i n t e r a c t i o n s were n o t e d . Swarmers r e l e a s e d f r o m t h e o n l y u n i l o c u l a r m a t e r i a l a v a i l a b l e ( f r c m C i t y P o i n t , C a l i f o r n i a ) were o b s e r v e d as t h e y swam and e v e n t u a l l y s e t t l e d . B. C y t o l o q i c a l Methods M a t e r i a l t o be u s e d f o r c y t o l o g i c a l work was m o s t l y f i e l d ' c o l l e c t e d m a t e r i a l and was e i t h e r f i e l d f i x e d o r b r o u g h t b a c k t o t h e l a b o r a t o r y i n a c o o l c h e s t and f i x e d w i t h i n 24 h o u r s o f c o l l e c t i o n . The f i x a t i v e used was 3 p a r t s 95% e t h a n o l : 1 p a r t g l a c i a l a c e t i c a c i d . . T h r e e methods were t r i e d t o o b t a i n i n f o r m a t i o n a b o u t t h e c a r y o l o g i c a l p h a s e s o f t h e l i f e h i s t o r y o f A . j a p o n i c u s : - ( 1 ) chromosome s t a i n i n g w i t h (a) a c e t o c a r m i n e , and (b) a c e t o - i r o n - h a e m a t o x y l i n - c h l o r a l - h y d r a t e , and (2) q u a n t i t a t i v e DNA s t a i n i n q and measurement w i t h 95 m i c r o s p e c t r o p h o t c m e t r y . (1) (a) A c e t o c a r m i n e ( J e n s e n , 1962). The a c e t o c a r m i n e s g u a s h t e c h n i q u e was used on s m a l l p i e c e s o f p l u r i l o c u l a r t i s s u e . T i s s u e was p r e t r e a t e d by s o a k i n g i n a 6% s o l u t i o n o f sodium c a r b o n a t e t o a s s i s t i n t h e s p r e a d i n g o f t i s s u e ( N a y l o r , 1959; R o b e r t s , 1966). (Sodium c a r b o n a t e removes a l g i n i c a c i d - P a r k e r and D i b o l l , 1 9 6 6 . ) (1) (b) A c e t o - i r o n - h a e m a t o x y l i n - c h l o r a l h y d r a t e (Wittmann, 1965). T h i s s t a i n was used on b o t h s q u a s h e s o f p l u r i l o c u l a r m a t e r i a l and a l s o on c u l t u r e d g e r m l i n g s . . C o v e r s l i p s w i t h f r e s h l y - s e t t l e d p l u r i s p o r e s g r o w i n g i n e n r i c h e d s e a w a t e r a t 1 0 ° C on a 12:12 p h o t o p e r i o d were f i x e d o v e r a 24 h o u r p e r i o d a t 1 h o u r i n t e r v a l s , a f t e r 1 day i n c u l t u r e and a l s o a f t e r 3 d a y s i n c u l t u r e . No sodium c a r b o n a t e p r e t r e a t m e n t was u s e d w i t h t h i s s t a i n as t h e c h l o r a l h y d r a t e f u n c t i o n s t o s o f t e n t h e t i s s u e . (2) Q u a n t i t a t i v e DNA measurement by m i c r o s p e c t r o p h o t o m e t r y . F e u l g e n was used as a q u a n t i t a t i v e DNA s t a i n . . (The q u a n t i t a t i v e r e l a t i o n s h i p between s t a i n i n t e n s i t y and DNA c o n t e n t i s w e l l e s t a b l i s h e d e.g. L e u c h t e n b e r g e r , 1958; Wied, 1966.) The r e l a t i v e DNA c o n t e n t o f t h e n u c l e i was measured u s i n g t h e t w o - a r e a method ( G a r c i a and I o r i o , 1966) w i t h a Z e i s s p h o t o m i c r o m e t e r w i t h K o h l e r i l l u m i n a t i o n and Z e i s s f i l t e r s BG23 and GG14. M a t e r i a l was p r e p a r e d f o r m i c r o s p e c t r o p h o t o m e t r y a c c o r d i n g t o t h e f o l l o w i n g s c h e d u l e : - (a) f e r t i l e l a t e r a l b r a n c h e s were chopped with a r a z o r 96 b l a d e i n t o pieces c a . 1-2mm, and t h e n p l a c e d i n t o a 6% s odium c a r b o n a t e s o l u t i o n f o r 1-2 h o u r s . A l l t h e f o l l o w i n g s t e p s u n t i l t h e d e h y d r a t i o n s e r i e s were p e r f o r m e d i n c e n t r i f u g e t u b e s , w i t h t h e m a t e r i a l b e i n g c e n t r i f u g e d and t h e n r e s u s p e n d e d w i t h e a c h s o l u t i o n c h a n g e . (b) two r i n s e s w i t h d i s t i l l e d water (c) h y d r o l y s i s i n 3N HCI a t 60° C f o r 20 m i n u t e s (These were t h e o p t i m a l h y d r o l y s i s c o n d i t i o n s and were d e t e r m i n e d e x p e r i m e n t a l l y . ) (d) r e p e a t (b) t o s t o p h y d r o l y s i s (e) t r a n s f e r r e d t o S c h i f f ' s r e a g e n t f o r 6 h o u r s i n t h e d a r k ( - r e a g e n t made up a c c o r d i n g t o Sharma and Sharma, 1965) (f) r i n s e d i n t a p water - f o u r c h a n g e s (g) r i n s e d i n SO2 water - t h r e e c h a n g e s o f t e n m i n u t e s e a c h (h) t r a n s f e r r e d t o d i s t i l l e d w a t e r ( i ) d e h y d r a t e d i n e t h y l a l c o h o l s e r i e s - 30%, 50%, 70%, 90%, 100%, 100%, 100% - a f t e r t r a n s f e r r i n g t o g l a s s v i a l s ( j ) mounted s m a l l p i e c e s o f t i s s u e i n E u p a r a l , s g u a s h i n g f i r m l y t o s p r e a d t h e c e l l s . The r e l a t i v e DNA c o n t e n t of 40 n u c l e i was measured f o r m a t e r i a l from e a c h o f t h e t y p e s examined:- (1) B a t h I s l a n d (B.C.) - p l u r i l o c u l a r p l a n t s , (2) D a v e n p o r t L a n d i n g ( C a l i f o r n i a ) - p l u r i l o c u l a r p l a n t s , (3) C i t y P o i n t ( C a l i f o r n i a ) - p l u r i l o c u l a r p l a n t s , (4) C i t y P o i n t u n i s p o r e s i n u n i l o c u l a r s p o r a n g i a , (5) C i t y P o i n t - s t e r i l e c e l l s i n u n i l o c u l a r p l a n t s . 97 R e s u l t s A. . C u l t u r e Work The h a n g i n g d r o p t e c h n i q u e o f Wynne (1969) gave c o n s i s t e n t l y good r e s u l t s . A p a r t from a t h i c k b o r d e r o f s p o r e s a b o u t t h e r i m (a hedgerow e f f e c t ) o f t h e h a n g i n g d r o p , s p o r e s e t t l e m e n t on t h e c o v e r s l i p o r s l i d e s u r f a c e was g e n e r a l l y e v e n . When u s i n q d r o p s o f s p o r e s u s p e n s i o n f o r s e t t l e m e n t t h e r e was a t e n d e n c y f o r a l e s s e v e n d i s t r i b u t i o n o f s e t t l e d s p o r e s . Growth on a g a r , e i t h e r w i t h o r w i t h o u t a l a y e r o f l i q u i d ES medium a b o v e , was n o t good. Growth i n t h e a r t i f i c i a l s e a w a t e r was v e r y s i m i l a r t o t h a t i n t h e ES medium. T h e r e was a l s o no a p p a r e n t d i f f e r e n c e i n t h e growth o f A. -japonicus i n media w i t h o r w i t h o u t germanium d i o x i d e . The c u l t u r e s which were grown w i t h a g i t a t i o n d i d not d e v e l o p d i f f e r e n t l y t h a n t h o s e grown w i t h o u t a g i t a t i o n . D e v e l o p m e n t a l d i f f e r e n c e s between a g i t a t e d and n o n - a g i t a t e d c u l t u r e s may have a p p e a r e d i f t h e c u l t u r e s had been grown beyond t h e e a r l y c r u s t s t a g e s . The d e v e l o p m e n t and m o r p h o l o g y of c u l t u r e s grown f r o m p l u r i s p o r e s and u n i s p o r e s a r e d e s c r i b e d i n C h a p t e r 1. The c u l t u r e s i n i t i a t e d w i t h p l u r i s p o r e s p r o d u c e d a c r u s t o s e phase and e r e c t a x e s i n a l l p h o t o p e r i o d / t e m p e r a t u r e c o n d i t i o n s under which t h e y were m a i n t a i n e d , t h a t i s , 5° 8:16,, 5° 12:12, 10° 12:12, 10° 16:8, 15° 12:12.. C u l t u r e s i n i t i a t e d w i t h u n i s p o r e s a l s o p r o d u c e d b o t h c r u s t and e r e c t a x e s i n a l l t h e c o n d i t i o n s i n which t h e y were grown, t h a t 98 i s , 5 ° 8 : 1 6 , 10 ° 1 2 : 1 2 , 10 ° 1 6 : 8 . The c u l t u r e s i n i t i a t e d w i t h p l u r i s p o r e s d e v e l o p e d p l u r i l o c u l a r s p o r a n g i a u n d e r t h e c o n d i t i o n s l i s t e d a b o v e . At t h e t i m e o f w r i t i n g , t h e c u l t u r e s i n i t i a t e d w i t h u n i s p o r e s had n o t d e v e l o p e d r e p r o d u c t i v e s t r u c t u r e s . 1 ) S a l i n i t y / T e m p e r a t u r e e x p e r i m e n t The r e s u l t s o f t h i s e x p e r i m e n t a r e summarised by F i g u r e 18 i n which f i v e r e p r e s e n t a t i v e g e r m l i n g s have been f i g u r e d a t day 7 a f t e r commencement f o r e a c h o f t h e t r e a t m e n t s . . The two f a c t o r s b e i n g v a r i e d i n t h i s e x p e r i m e n t c a n be s e e n t o have i n t e r a c t i v e e f f e c t s on t h e growth of A . j a p o n i c u s . _ At 5 ° C s l o w g r o w t h o c c u r r e d a t a l l s a l i n i t i e s , b e i n g p a r t i c u l a r l y s l e w a t t h e l o w e s t s a l i n i t y 1 2 . 5 % o . At 15 ° and 2 0 ° C , t h e e f f e c t o f t h e l o w e r s a l i n i t i e s was a p p a r e n t l y s e v e r e . Thus a t 15 ° C and 12.5%o and 2 0 ° C and 1 2 . 5 and 18.5%o , t h e c e l l s have l i t t l e c y t o p l a s m and a p p a r e n t l y a r e m o s t l y v a c u o l a r s p a c e . T h e s e c e l l s a l s o have an i n f l a t e d a p p e a r a n c e b e i n g c o n s i d e r a b l y l a r g e r t h a n c e l l s a t t h e same s a l i n i t y b u t l o w e r t e m p e r a t u r e . I n g e n e r a l , t h e f i r s t o b l i q u e d i v i s i o n o f t h e d i s c o i d c r u s t o c c u r r e d when t h e g e r m l i n g s r e a c h e d a 2-3 c e l l s t a g e . I n c u l t u r e s a t 2 0 ° C between 1 2 . 5 and 30%o t h e g e r m l i n g s c o n t i n u e d t o d i v i d e t r a n s v e r s e l y f o r m i n g l o n g e r f i l a m e n t s b e f o r e t h e o b l i q u e d i v i s i o n s o c c u r r e d . . I n t h e s e c u l t u r e s a t 2 0 ° C , a number o f a b e r r a n t g e r m l i n g s were seen where t h e o b l i q u e d i v i s i o n s o c c u r r e d from s e v e r a l c e l l s i n a l o n g e r f i l a m e n t r e s u l t i n g i n a t y p i c a l and sometimes b r a n c h i n g g e r m l i n g s . . T h e s e 9 9 g e r m l i n g s a p p a r e n t l y a b o r t e d s h o r t l y a f t e r t h i s s t a g e and t h e more n o r m a l g e r m l i n g s c o n t i n u e d t o grow and d e v e l o p i n t o d i s c o i d c r u s t s . (For an example r e f e r t o F i g u r e 18 2 0 ° C a t 30% .) At 2 0 ° C and 37%o t h e g e r m l i n g s h a d a n o r m a l m o rphology and were c o n s i d e r a b l y more d e v e l o p e d t h a n t h e g e r m l i n g s a t t h e same s a l i n i t y b u t l o w e r t e m p e r a t u r e s . Growth o f c u l t u r e s beyond day 7 f o l l o w e d a s i m i l a r p a t t e r n t o t h a t i l l u s t r a t e d i n F i g u r e 1 8 . At 1 2 . 5 % o o n l y t h e c u l t u r e s a t 5 ° and 1 0 ° C s u r v i v e d beyond day 1 4 . . A t 1 8 . 5 and 25%o a l l c u l t u r e s s u r v i v e d b eyond day 2 5 e x c e p t f o r t h e s e a t 2 0 ° C . At 3 0 and 37%o a l l c u l t u r e s s u r v i v e d b e y o n d day 2 5 a l t h o u g h some g e r m l i n g s a t 3 0 % o and 2 0 ° C were a b e r r a n t i n t h e i r f o r m . 2) l i g h t Q u a l i t y Growth i n a l l t h e t r e a t m e n t s was e q u i v a l e n t w i t h no d e v e l o p m e n t a l d i f f e r e n c e s d i s t i n g u i s h a b l e up t o t h e s i x week s t a g e . 3) F u s i o n o f J warmers The s i z e o f t w e n t y swarmers from e a c h o f f o u r p o p u l a t i o n s were measured, and t h e d a t a a r e shown i n t h e f o l l o w i n g The mean d i a m e t e r s o f t h e swarmers f r o m t h e t h r e e l i s t . MATERIAL MEAN DIAMETER (jam) SD (Jim) C i t y P o i n t u n i s p o r e s 7 . 3 2 5 C i t y P o i n t p l u r i s w a r m e r s 6 . 6 7 5 Bath I s l a n d p l u r i s w a r m e r s 6 . 7 6 2 5 F c g a r t y Creek p l u r i s w a r m e r s 6 . 6 6 2 5 0 . 3 3 5 4 0. 3 8 9 8 0 . 3 0 8 6 0. 4 2 3 6 p l u r i l o c u l a r p o p u l a t i o n s were n o t f o u n d t o be s i g n i f i c a n t l y 100 d i f f e r e n t , and were a l l s i g n i f i c a n t l y s m a l l e r (0.001 c o n f i d e n c e l e v e l ) t h a n t h e mean o f t h e u n i s p o r e s measured. Thus I f o u n d no e v i d e n c e t o s u p p o r t Abe's c l a i m o f a n i s o g a m e t e s . . I d i d n o t o b s e r v e any e v i d e n c e o f p h y s i o l o g i c a l a n i s o gamy, o r swarmers swimming a r o u n d o t h e r s (as r e p o r t e d by Abe). I saw no e v i d e n c e o f f u s i o n i n t h e c o m b i n a t i o n h a n g i n g d r o p s . The u n i s p o r e s I o b s e r v e d a l s o showed no s i g n s of f u s i n g . F u r t h e r , I c o u l d not i d e n t i f y t h e two k i n d s o f u n i l o c u l a r or two k i n d s o f p l u r i l o c u l a r p l a n t s which Abe d e s c r i b e d i n J a p a n . B. C y t o l o g i c a l R e s u l t s ( 1 ) (a) A c e t o c a r m i n e . R e s u l t s o b t a i n e d w i t h t h i s method were po o r and no chromosome c o u n t s were o b t a i n e d . . The p r i m a r y p r o b l e m was o b t a i n i n g d i v i s i o n s t a g e s i n t h e t i s s u e r a t h e r t h a n f a i l u r e o f t h e s t a i n . (1)(b) A c e t o - i r o n - h a e m a t o x y l i n - c h l o r a l h y d r a t e . . T h i s t r e a t m e n t gave b e t t e r r e s u l t s t h a n t h e a c e t o c a r m i n e , but t h e s i z e o f t h e n u c l e i and c y t o p l a s m i c s t a i n i n g p r e v e n t e d c o u n t i n g chrcmcsomes. With p l u r i l o c u l a r m a t e r i a l i t was a l s o d i f f i c u l t t o f i n d n u c l e i i n d i v i s i o n s t a g e s . A 24-hour f i x a t i o n s e r i e s o f young g e r m l i n g s was c a r r i e d o u t i n t h e hope o f i d e n t i f y i n g t h e t i m e a t which n u c l e a r d i v i s i o n s were o c c u r r i n g . The g e r m l i n g s had t h e a d v a n t a g e o f b e i n g o n l y 1 c e l l - l a y e r t h i c k . With t h i s m a t e r i a l t h e c y t o p l a s m 101 was v e r y d e n s e , and o b s c u r e d t h e n u c l e i . (2) Q u a n t i t a t i v e DNA measurement. B e s u l t s o f t h e m i c r o s p e c t r o p h o t o m e t r i c t e c h n i q u e a r e g i v e n i n F i g u r e 19. F e u l g e n m i c r o s p e c t r o p h o t o m e t r y r e v e a l s t h e r e l a t i v e amount o f DNA p r e s e n t i n a c e l l . F o r a p a r t i c u l a r p l o i d y l e v e l t h i s w i l l be a r a n g e o f v a l u e s d e p e n d i n g upon t h e s t a g e o f t h e c e l l c y c l e t h a t t h e n u c l e i a r e i n . T h a t i s , i n h a p l o i d t i s s u e DNA v a l u e s w i l l r a n g e between 1x and 2x and i n d i p l o i d t i s s u e between 2x and 4 x . . A l t h o u g h a b s o l u t e DNA c o n t e n t can be c a l c u l a t e d u s i n g known s t a n d a r d s , t h i s i s n o t n e c e s s a r y h e r e as t h e p u r p o s e i s t o see i f t h e r e i s a d i f f e r e n c e i n p l o i d y l e v e l . The r a n g e o f v a l u e s o f DNA c o n t e n t f o r s t e r i l e c e l l s f r o m u n i l o c u l a r p l a n t s i s between 9 and 22, whereas t h a t o f t h e u n i s p o r e s w i t h i n t h e u n i l o c u l a r s p o r a n g i a and a l s o t h a t o f a l l t h e p l u r i l o c u l a r p l a n t s f a l l s between 3 and 12. The mean r e l a t i v e DNA c o n t e n t o f t h e p l u r i l o c u l a r and u n i s p o r e n u c l e i i s a p p r o x i m a t e l y h a l f as l a r g e as t h a t i n t h e s t e r i l e c e l l s o f t h e u n i l o c u l a r p l a n t s . 102 D i s c u s s i o n The c u l t u r e r e s u l t s i n d i c a t e d t h a t d e v e l o p m e n t o f e r e c t a x e s and p r o d u c t i o n o f p l u r i l o c u l a r r e p r o d u c t i v e o r g a n s can o c c u r i n A. -japonicus o v e r a r a n g e o f t e m p e r a t u r e and p h o t o p e r i o d c o n d i t i o n s . T h i s a g r e e s w i t h f i e l d o b s e r v a t i o n s ( r e p o r t e d i n C h a p t e r 2) where i t was f o u n d t h a t e r e c t a x e s o c c u r r e d t h r o u g h o u t t h e y e a r ( a l t h o u g h n o t n e c e s s a r i l y a t a l l s i t e s ) , and t h a t r e p r o d u c t i v e l y mature p l a n t s w i t h p l u r i l o c u l a r s p o r a n g i a were a l s o p r e s e n t y e a r r o u n d . The r e s p o n s e o f g e r m l i n g s t o v a r y i n g s a l i n i t y / t e m p e r a t u r e c o n d i t i o n s shows A . j a p o n i c u s t o be w i d e l y t o l e r a n t o f t h e s e e n v i r o n m e n t a l f a c t o r s . T h i s i s a l s o c o n s i s t e n t w i t h f i e l d o b s e r v a t i o n s a s A . j a p o n i c u s i n B r i t i s h C o l u m b i a i s f o u n d b o t h i n h i g h e r s a l i n i t y w a t e r s o f t h e open c o a s t and i n l o w e r s a l i n i t y w a t e r s o f t h e S t r a i t o f G e o r g i a . P o p u l a t i o n s o f A . j a p o n i c u s i n t h e S t r a i t o f G e o r g i a d e c l i n e i n t h e summer, u n l i k e o t h e r p o p u l a t i o n s s t u d i e d i n B.C. In summer, t h e F r a s e r R i v e r f r e s h e t r e s u l t s i n low minimum s a l i n i t y v a l u e s i n t h e w a t e r s o f t h e S t r a i t o f G e o r g i a . T h i s i s a l s o a t i m e when water and a i r t e m p e r a t u r e s a r e a t t h e i r h i g h e s t * ( R e f e r t o Appendix I I I f o r s a l i n i t y and t e m p e r a t u r e d a t a f o r t h e S t r a i t o f G e o r g i a . ) As bo r n e c u t by t h e s e c u l t u r e r e s u l t s , t h e low s a l i n i t y / h i g h t e m p e r a t u r e c o m b i n a t i o n i s p a r t i c u l a r l y s t r e s s f u l f o r t h e p l a n t s . 103 P l a n t s o f A . j a p o n i c u s a r e known t o o c c u r a t d i f f e r e n t t i d a l h e i g h t s a t d i f f e r e n t l o c a t i o n s b e i n g r e p o r t e d f r o m t h e upper i n t e r t i d a l zone t o t h e u p p e r s u b t i d a l zone ( r e f e r C h a p t e r 2 ) . The p l a n t s a t d i f f e r e n t t i d a l h e i g h t s n o t o n l y e x p e r i e n c e d i f f e r e n t amounts o f d e s i c c a t i o n , b u t a l s o e x p e r i e n c e d i f f e r e n t l i g h t g u a l i t y o v e r t h e t i d a l c y c l e due t o t h e d i f f e r e n t i a l a b s o r p t i o n o f t h e l i g h t by t h e water column. The r e s u l t s of the l i g h t g u a l i t y e x p e r i m e n t s u g g e s t t h a t A . j a p o n i c u s p o s s e s s e s a t l e a s t t h e a b i l i t y t o g e r m i n a t e and d e v e l o p under a r a n g e o f l i g h t q u a l i t y r e g i m e s . I t would be i n t e r e s t i n g t o e x t e n d t h i s work t o see whether a d u l t m o rphology and r e p r o d u c t i o n a r e a f f e c t e d by p r o l o n g e d e x p o s u r e t o r e g i m e s o f d i f f e r e n t l i g h t g u a l i t y . The use o f t h e F e u l g e n m i c r o s p e c t r o p h o t o m e t r i c method r e p o r t e d h e r e i s t h e f i r s t a c c o u n t o f t h e a p p l i c a t i o n o f t h i s t e c h n i g u e i n t h e P h a e o p h y t a a l t h o u g h i t has been u s e d cn members o f t h e C h l o r o p h y t a ( H o p k i n s and M c B r i d e , 1976; H o p k i n s , 1977; Kemp, D o y l e , and A n d e r s o n , 1979) and R hodophyta ( H u r d e l b r i n k and S c h w a n t e s , 1972; Breeman, 1979). T h i s c y t o l o g i c a l work v e r i f i e s Abe's (1936) c l a i m t h a t t h e u n i l o c u l a r s p o r a n g i a i n A . j a p o n i c u s a r e t h e s i t e o f m e i o s i s . C u l t u r e s i n i t i a t e d w i t h p l u r i s p o r e s were c a p a b l e o f a s e x u a l l y c y c l i n g t h e m s e l v e s f o l l o w i n g t h e l i f e h i s t o r y p a t t e r n o u t l i n e d i n F i g u r e 20a. A . j a p o n i c u s a p p a r e n t l y has an e x c l u s i v e l y a s e x u a l l i f e h i s t o r y f o r most o f i t s g e o g r a p h i c r a n g e ( A l a s k a , B r i t i s h C o l u m b i a , W a s h i n g t o n , 104 Oregon and much o f C a l i f o r n i a ) . The s e x u a l c y c l e and a l t e r n a t i o n o f i s o m o r p h i c g e n e r a t i o n s r e p r e s e n t e d i n F i g u r e 20fc o c c u r s i n J a p a n and i n a t l e a s t one s i t e i n C a l i f o r n i a . ( R e f e r t o C h a p t e r 2 f o r f u r t h e r d e t a i l s a b o u t the p h e n o l o g y o f A . j a p o n i c u s . ) The c y t o l o g i c a l work r e p o r t e d h e r e s h o w i n g t h e p l u r i l o c u l a r p l a n t s t o be h a p l o i d i s c o n t r a r y t o K y l i n ' s (1940) s u g g e s t i o n t h a t t h e y a r e d i p l o i d . I f Abe (1935a) was c o r r e c t i n m a i n t a i n i n g _ t h a t o n l y f e m a l e swarmers a r e c a p a b l e o f apogamous d e v e l o p m e n t t h e n t h e o b s e r v a t i o n s r e p o r t e d h e r e a r e not s u r p r i s i n g . A p p a r e n t l y f o r most o f t h e r a n g e o f A . j a p o n i c u s t h e r e a r e no s e x u a l p o p u l a t i o n s . T h u s , t h e p l u r i l o c u l a r p l a n t s which a r e f o u n d a t most s i t e s would be, a c c o r d i n g t o Abe, e x c l u s i v e l y f e m a l e and o n l y a b l e t o r e p r o d u c e a s e x u a l l y . The r e s u l t s g i v e n h e r e - t h a t t h e r e were n o t two s i z e c l a s s e s o f swarmers from t h e p l u r i l o c u l a r p l a n t s and t h a t t h e s e swarmers d i d n o t f u s e - would be c o n s i s t e n t w i t h t h i s . . O n l y where t h e r e a r e u n i l o c u l a r p l a n t s would t h e r e be t h e p o s s i b i l i t y o f male p l u r i l o c u l a r p l a n t s ( g r o w i n g f r o m u n i s p o r e s ) and t h u s t h e p o s s i b i l i t y o f s e x u a l f u s i o n . I f t h i s i n t e r p r e t a t i o n i s c o r r e c t , t h e n t h e o n l y s i t e where t h e a n i s o g a m o u s p l u r i l o c u l a r p l a n t s might be f o u n d i s C i t y P o i n t . A l t h o u g h I d i d n o t f i n d anisogamy i n t h e m a t e r i a l I e x a m i n e d , I t h i n k t h a t t h i s w a r r a n t s c l o s e r a t t e n t i o n . . I f , as a t o t h e r s i t e s , t h e a s e x u a l p o r t i o n o f t h e l i f e h i s t o r y o f A . j a p o n i c u s a t C i t y P o i n t c o n t r i b u t e s s u b s t a n t i a l l y t o t h e p o p u l a t i o n m a i n t e n a n c e and g r o w t h , t h e n one would 105 e x p e c t t h e male p l u r i l o c u l a r p l a n t s t o be r e l a t i v e l y r a r e compared t o t h e f e m a l e p l u r i l o c u l a r p l a n t s , p a r t i c u l a r l y as t h e u n i l o c u l a r p o p u l a t i o n i s so l i m i t e d i n i t s e x t e n t . . The l i f e h i s t o r y f i n d i n g s show t h a t A . j a p o n i c u s s h o u l d n o t be p l a c e d w i t h t h e C h o r d a r i a l e s w hich i s d i s t i n g u i s h e d as a d i s t i n c t c r d e r "on t h e b a s i s o f t h e h e t e r o m o r p h i c l i f e h i s t o r y i n which t h e h a p l o i d m i c r o t h a l l u s a l t e r n a t e s w i t h a d i p l o i d m a c r o t h a l l u s " ( B o l d and Wynne, 1978) . . ( B e f e r t o C h a p t e r 6 f o r a d i s c u s s i o n o f t h e t a x o n o m i c p o s i t i o n o f 106 F i g u r e 18. Summary o f e f f e c t s o f v a r y i n g s a l i n i t y and. t e m p e r a t u r e cn growth o f g e r m l i n g s o f A . j a p o n i c u s . F i v e 7 - d a y - o l d g e r m l i n g s from e a c h o f t h e t w e n t y e x p e r i m e n t a l c o n d i t i o n s a r e r e p r e s e n t e d h e r e . S c a l e t a r = 20 ûm. 107 5 ° 10° 15 ° 2 0 ° 12-5 CO n 18-5 V ft ^ 0 <̂ > 25 30 S 8 37 \ <p 20 108 F i g u r e 1 9 . . R e l a t i v e m i c r o s p e c t r o each t y p e o f DNA v a l u e s measured p h o t o m e t r y i n n u c l e i t i s s u e t h e sample s by of A . j a p o n i c u s . F o r i z e o f n u c l e i = 40.. 109 8n B a t h I s l a n d - p l u r i l o c u l a r 8- D a v e n p o r t - p l u r i l o c u l a r 8 C i t y P o i n t - p l u r i l o c u l a r 4J 3 C 8 n SI E z C i t y P o i n t - u n i s p o r e s 8- C i t y P o i n t - u n i l o c u l a r ( s t e r i l e c e l l s ) ~i 1 1 1 1 1 1 1 1 1 1 1 h- 4 8 12 16 R e l a t i v e D N A v a l u e s 20 110 F i g u r e 2 0 . D i a g r a m m a t i c r e p r e s e n t a t i o n o f (a) l i f e h i s t o r y of fl.japonicus i n c u l t u r e , and, (b) l i f e h i s t o r y o f fl.japonicus i n n a t u r e . 111 erect axes mmmm^^- plurangia t *! crusts -^FJjjjJ™™ swarmers (a) LIFE HISTORY IN CULTURE unilocular sporangia (b) LIFE HISTORY IN NATURE 1 CHAPTER 4 A u t e c o l o g y o f A n a l i p u s j a p o n i c u s a t B a t h I s l a n d , B r i t i s h C o l u m b i a 113 I n t r o d u c t i o n O v e r a two y e a r p e r i o d from J u l y 1977 t o August 1979, an a u t e c o l o g i c a l s t u d y was made o f a p o p u l a t i o n o f A n a l i p u s j a p o n i c u s a t Bath I s l a n d i n t h e S t r a i t o f G e o r g i a , B r i t i s h C o l u m b i a . When t h i s work was s t a r t e d t h e r e were no s t u d i e s on t h e e c o l o g y o f A . j a p o n i c u s i n t h e e a s t e r n P a c i f i c , and, as n o t e d i n C h a p t e r 2, t h e r e were c o n t r a d i c t o r y r e p o r t s i n t h e l i t e r a t u r e on t h e p h e n o l o g y o f t h i s p l a n t i n o t h e r p a r t s o f i t s r a n g e . I n making t h i s s t u d y i t was hoped t o d e s c r i b e d y n a m i c s o f t h e A . j a p o n i c u s p o p u l a t i o n and e l e m e n t s o f t h e community i n which i t i s fo u n d a t Ba t h I s l a n d . T h i s work s h o u l d be s e e n as h y p o t h e s i s - g e n e r a t i n g i n n a t u r e r a t h e r t h a n h y p o t h e s i s - t e s t i n g , and s e r v e a s a b a s e l i n e f o r f u t u r e e x p e r i m e n t a l e c o l o g i c a l s t u d i e s on t h i s p l a n t and t h e i n t e r t i d a l c o m m u n i t i e s f o u n d i n the S t r a i t o f G e o r g i a . The s e a s o n a l g r o w t h o f t h e A . j a p o n i c u s p o p u l a t i o n and t h e B a t h I s l a n d i n t e r t i d a l community was f o l l o w e d by permanent q u a d r a t s and t r a n s e c t s sampled e i t h e r by t h e p o i n t i n t e r s e c t method o r w i t h u n d e r w a t e r p h o t o g r a p h y * As r e p o r t e d i n C h a p t e r 2, r e g u l a r c o l l e c t i o n s were made o f A . j a p o n i c u s from t h e B a t h I s l a n d s i t e i n o r d e r t o measure v a r i o u s m o r p h o l o g i c a l c h a r a c t e r s o f t h e p l a n t s and i n o r d e r t o f o l l o w t h e p h e n o l o g y o f t h e p o p u l a t i o n . . S e t t l e m e n t p l a t e s were s e t o u t i n t h e s t u d y a r e a i n the hope o f o b s e r v i n g e a r l y s t a g e s o f c r u s t d e v e l o p m e n t and a l s o t o g e t 114 an i n d i c a t i o n o f t h e t i m e o f y e a r when most s e t t l e m e n t o c c u r s . . S u b s t r a t e w i t h i n t h e A . j a p o n i c u s zone i n t h e i n t e r t i d a l r e g i o n was c l e a r e d t o f o l l o w r e c o l o n i s a t i o n a t d i f f e r e n t t i m e s o f A. • j a p o n i c u s 1 r e p r o d u c t i v e c y c l e . M a t e r i a l s and Methods Bath I s l a n d ( 49 o09*N, 123°40'M ) i s one o f t h e F l a t Top G r o u p , a c l u s t e r o f s m a l l i s l a n d s l o c a t e d i n t h e S t r a i t o f G e o r g i a t o t h e s o u t h e a s t o f G a b r i o l a I s l a n d ( F i g u r e 2 1 ) . The f i e l d s i t e u s e d f o r t h i s s t u d y was on t h e s o u t h e a s t s i d e o f Bath I s l a n d on an e v e n l y s l o p i n g r o c k f a c e . The i s l a n d i s l o c a t e d a t t h e w i d e s t p o i n t o f the S t r a i t o f G e o r g i a w i t h t h e f i e l d s i t e f a c i n g t h e d i r e c t i o n o f t h e p r e v a i l i n g w i n d s , and an u n i n t e r r u p t e d s t r e t c h o f w a t e r t o t h e l o w e r m a i n l a n d o f B r i t i s h C o l u m b i a . Q u a d r a t s and T r a n s e c t s F i v e permanent g u a d r a t s (QI-QV, F i g u r e 22) were e s t a b l i s h e d a t t h e f i e l d s i t e a t t h e b e g i n n i n g o f J u l y 1977. An a l u m i n i u m q u a d r a t frame 0.5 m x 0.5 m was u s e d . N y l o n t h r e a d was s t r u n g t i g h t l y a c r o s s t h e f r a m e h o r i z o n t a l l y and v e r t i c a l l y , r e s u l t i n g i n i n t e r s e c t i n g l i n e s a t 5 cm i n t e r v a l s (a t o t a l o f 81 i n t e r s e c t p o i n t s w i t h i n t h e f r a m e ) . A h o l e was d r i l l e d i n e a c h o f t h e u p p e r 115 l e f t and r i g h t hand c o r n e r s o f t h e frame. The f i v e q u a d r a t s were l o c a t e d i n t h e f i e l d w i t h e x p a n s i o n b o l t s ( s e c u r e d i n t h e r o c k s u b s t r a t e a f t e r d r i l l i n q h o l e s w i t h a d r i l l powered w i t h c o m p r e s s e d a i r ) which were f l a g g e d w i t h b r i g h t p l a s t i c t a p e . The frame s l o t t e d o v e r t h e b o l t s and s a m p l i n g c o n s i s t e d of r e c o r d i n g what o c c u r r e d a t e a c h p o i n t i n t e r s e c t . The q u a d r a t s were sampled a t a p p r o x i m a t e l y m o n t h l y i n t e r v a l s f r o m J u l y 1977 t o A u g ust 1979. R e f e r t o A p p e n d i x VI f o r a l i s t o f s a m p l i n g d a t e s f o r Q u a d r a t s I-V. A permanent t r a n s e c t (PT1-10, F i g u r e 22) r u n n i n g v e r t i c a l l y t h r o u g h t h e i n t e r t i d a l r e g i o n was e s t a b l i s h e d a t t h e b e g i n n i n g o f J u l y 1977. I t was marked by two e x p a n s i o n b o l t s p l a c e d 12 m a p a r t i n t h e r o c k s u b s t r a t e and marked w i t h c o l o u r e d p l a s t i c t a p e . A c o t t o n c o r d was used as t h e t r a n s e c t l i n e and t h i s was marked a t one metre i n t e r v a l s w i t h p l a s t i c t a p e and a t 5 cm i n t e r v a l s w i t h i n d e l i b l e i n k . The c o r d was s t r e t c h e d out between t h e e n d p o i n t s and a t t a c h e d t o t h e s e by ' S' hooks a t e i t h e r end o f t h e c o r d . A 10 IE l e n g t h o f t h i s c o r d was used f o r r e c o r d i n g what o c c u r r e d a t 5 cm i n t e r v a l s t h r o u g h t h e l e n g t h o f t h e l i n e . The t r a n s e c t l i n e was sampled a t a p p r o x i m a t e l y m o n t h l y i n t e r v a l s from J u l y 1977 t o J u l y 1979. R e f e r t o A p p e n d i x Vt f o r a l i s t o f s a m p l i n g d a t e s of t h e t r a n s e c t . 116 C l e a r i n g E x p e r i m e n t As r e p r e s e n t e d i n F i g u r e 22 by T1-T12, 12 q u a d r a t s were e s t a b l i s h e d a l o n g a h o r i z o n t a l l i n e . They were marked by e x p a n s i o n b o l t s , f l a g g e d w i t h p l a s t i c t a p e and were 0.3 m x 0.3 m i n s i z e . They were s a m p l e d p h o t o g r a p h i c a l l y , f o r 19 months (frcm May 1978 t o November 1979) u s i n g an u n d e r w a t e r camera a t t a c h e d t o an a l u m i n i u m f r a m e . The r e c t a n g u l a r b a s e o f t h e frame s l o t t e d o v e r b o l t s m a r k i n g e a c h q u a d r a t . The camera and s t r o b e f l a s h l i g h t s o u r c e were s u p p o r t e d on t h e u p p e r p o r t i o n o f t h e f r a m e which was r a i s e d on l e g s a t r i g h t a n g l e s t o t h e base and a s e t d i s t a n c e f r o m t h e s u b s t r a t e b e i n g p h o t o g r a p h e d . The d i m e n s i o n s o f e a c h g u a d r a t t h a t was p h o t o g r a p h i c a l l y sampled was 0.2 m x 0.15 m. Q u a d r a t s 1,3,5,7,9,and 11 were l e f t u n d i s t u r b e d d u r i n g t h e s a m p l i n g p e r i o d . S i x g u a d r a t s were c l e a r e d a t low t i d e by s c r a p i n g and t h e n by b u r n i n g w i t h an e t h y l e n e t o r c h . Q u a d r a t s T2 and T6 were c l e a r e d i n May 1978, T4 and T8 i n June 1978, and 110 and T12 were c l e a r e d i n August 1978.. M o n t h l y C o l l e c t i o n s As d e s c r i b e d i n C h a p t e r 2, c o l l e c t i o n s were made a t Bath I s l a n d a t a p p r o x i m a t e l y m o n t h l y i n t e r v a l s and were used f o r a p h e n c l o g i c a l s t u d y . 1 1 7 P h o t o g r a p h i c Q u a d r a t s I n a d d i t i o n t o t h e g u a d r a t s T1-T12 w h i c h were o b s e r v e d p h o t o g r a p h i c a l l y a s p a r t o f t h e c l e a r i n g e x p e r i m e n t , 8 q u a d r a t s ( i n d i c a t e d i n F i g u r e 22 as Q1-Q8) were f o l l o w e d p h o t o g r a p h i c a l l y . T h e s e p h o t o g r a p h s p r o v i d e d a r e c o r d o f c r u s t and u p r i g h t a x i s g r o w t h . S e t t l e m e n t P l a t e s Frames t o h o l d g l a s s m i c r o s c o p e s l i d e s were c o n s t r u c t e d from p e r s p e x and c o n s i s t e d o f a f l a t r e c t a n g u l a r b a s e w i t h s t r i p s mounted a l o n g b o t h s i d e s t o fo r m o v e r l a p p i n g e d g e s . S i x g l a s s s l i d e s were t h u s h e l d i n a h o r i z o n t a l p o s i t i o n on t h e p e r s p e x b a s e . The g l a s s s l i d e s were g r o u n d on a r o c k p o l i s h i n g t u r n t a b l e w i t h c a r b o r u n d u m t o r o u g h e n t h e i r s u r f a c e s . The f r a m e s w i t h the s l i d e s i n p l a c e were s e c u r e d a t t h r e e v e r t i c a l h e i g h t s w i t h i n t h e ft.japonicus r a n g e by b e i n g s l o t t e d o v e r e x p a n s i o n b o l t s and were h e l d i n p l a c e by wing n u t s . The p l a t e s w i t h t h e s l i d e s were p l a c e d i n t h e f i e l d s i t e f o r v a r y i n g l e n g t h s o f t i m e and when c o l l e c t e d t h e s l i d e s were p l a c e d i n t o a 5% f o r m a l d e h y d e / s e a w a t e r s o l u t i o n . The s l i d e s were s t a i n e d w i t h l a c t o p h e n o l b l u e and o b s e r v e d m i c r o s c o p i c a l l y i n t h e l a b o r a t o r y . The f o l l o w i n g l i s t g i v e s t h e d a t e s on which s l i d e s were p l a c e d i n t h e f i e l d and t h e n c o l l e c t e d and f i x e d f o r exam i n a t i o n , : - 118 Date p l a c e d i n t h e f i e l d Date c o l l e c t e d 7 . IX • .78 7 . . X . 73 7 . X • 78 26 . XI . 78 26 . XI • 78 13 I . 79 13 . I • 79 10 . I l l . 79 10 = . I I I • 79 12 . . V . 79 12 . V • 79 10 . V I I . 79 R e s u l t s F i e l d S i t e S t o r m s and i n c r e a s e d wave a c t i o n a f f e c t e d t h e B a t h I s l a n d s t u d y s i t e i n t h e w i n t e r . D r i f t l o g s w h i c h a c c u m u l a t e i n t h e u p p e r i n t e r t i d a l r e g i o n were d i s l o d g e d by t h e h i g h e s t t i d e s o r by s t o r m waves and o f t e n c a u s e d c o n s i d e r a b l e damage by b a t t e r i n g and c r u s h i n g t h e i n t e r t i d a l and upper s u b t i d a l f l o r a and f a u n a . M o n t h l y mean, maximum and minimum s a l i n i t y and t e m p e r a t u r e v a l u e s f o r t h e s u r f a c e w a t e r s o f f E n t r a n c e I s l a n d ( 4 9 ° 1 0 « N , 124°10'W ) a r e g i v e n i n A p p e n d i x I I I . ( T h i s i s t h e c l o s e s t s i t e t o Bath I s l a n d where d a i l y s a m p l i n g o f s u r f a c e w a t e r s o c c u r s . The l i m i t e d amount o f s a l i n i t y and t e m p e r a t u r e d a t a c o l l e c t e d a t B a t h I s l a n d c l o s e l y f o l l o i i s t h e d a t a from E n t r a n c e I s l a n d . ) The f r e s h e t of t h e F r a s e r R i v e r s t r o n g l y i n f l u e n c e s s a l i n i t y i n t h e S t r a i t of G e o r g i a c a u s i n g a marked d r o p i n t h e minimum v a l u e s measured i n t h e summer months. The t i d a l r e g i m e a t B a t h I s l a n d i s a mixed s e m i d i u r n a l 119 t y p e . The e x t r e m e low t i d e s i n summertime o c c u r a r o u n d t h e m i d d l e o f t h e day, and t h e A . j a p o n i c u s p l a n t s c a n be e x p o s e d t o t h e a i r f o r up t o 5 h o u r s b e f o r e b e i n g r e - immersed by t h e i n c o m i n g t i d e (Anon, 1977,1978,1979). The i n t e r t i d a l r o c k s h e l f a t t h i s s i t e i s e v e n l y s l o p i n g w i t h l i t t l e t o p o g r a p h i c a l r e l i e f - few c r a c k s , c r e v i c e s o r shade a r e a s - and t h u s , t h e r e i s l i t t l e h a b i t a t v a r i a t i o n . A d i a g r a m m a t i c r e p r e s e n t a t i o n o f t h e f i e l d s i t e and t h e v e r t i c a l h e i g h t s above c h a r t datum, o f g u a d r a t and t r a n s e c t p o i n t s a r e g i v e n i n F i g u r e 22. The s i t e was s u r v e y e d a t e x t r e m e low water i n mid-summer t o measure t h e h e i g h t o f t h e s t u d y p o i n t s above c h a r t datum. A c c e s s t o t h e s i t e f o r most o f t h e y e a r was by d i v i n g w i t h SCUBA. From May t o August some o b s e r v a t i o n s and c o l l e c t i o n s were made a t extreme low t i d e s . . Q u a d r a t s and T r a n s e c t s A l i s t o f s p e c i e s f o u n d i n t h e q u a d r a t s and t r a n s e c t i s g i v e n i n A p p e n d i x V I I . F i g u r e 23 and 24 show numbers o f A* j a p o n i c u s p l a n t s sampled i n q u a d r a t s QI-QV o v e r t h e s t u d y p e r i o d . F i g u r e 25 shows numbers of A . j a p o n i c u s r e c o r d e d i n t h e t r a n s e c t o v e r t h e same p e r i o d w i t h t h e u p p e r e i g h t m e t r e s o f t h e t r a n s e c t h a v i n g been d i v i d e d i n t o f o u r , two- metre l e n g t h s and l a b e l l e d PTI-PTIV (from upper t o l o w e r i n t e r t i d a l r e g i o n ) r e s p e c t i v e l y . No A . j a p o n i c u s p l a n t s were r e c o r d e d i n t h e l o w e s t metre of t h e t r a n s e c t . . I n F i g u r e 26 numbers o f p l a n t s o f Sargassum muticum, Rhodomela l a r i x , 120 and C r y p t o s i p h o n i a w o o d i i s a m p l e d i n t h e t r a n s e c t and q u a d r a t s a r e shown. F i g u r e 27 shows numbers o f m u s s e l s ( M y t i l i s e d u l i s ) and M i c r o c l a d i a b o r e a l i s p l a n t s r e c o r d e d o v e r t h e s a m p l i n g p e r i o d i n b o t h t r a n s e c t and quadrats,. These- s p e c i e s were i m p o r t a n t e l e m e n t s o f t h e community (as measured by numbers r e c o r d e d i n q u a d r a t s and t r a n s e c t ) and t h e abundance o f t h e s e s p e c i e s c hanged o v e r t h e s t u d y p e r i o d . F i g u r e 31 i n d i c a t e s t h e d i s t r i b u t i o n o v e r t i m e o f e i g h t o t h e r s p e c i e s t h a t were f r e q u e n t l y r e c o r d e d i n t h e q u a d r a t s and t r a n s e c t w i t h t h e p e r i o d o f maximum abundance b e i n g i n d i c a t e d by s h a d i n g . F i g u r e s 28 and 29 show numbers o f a n n u a l and p e r e n n i a l p l a n t s sampled i n each o f t h e q u a d r a t s p l o t t e d a g a i n s t t h e t i m e o f s a m p l i n g . In F i g u r e 30 t h e t o t a l number o f A . j a p o n i c u s p l a n t s r e c o r d e d f r o m a l l q u a d r a t s and t h e mean a x i s h e i g h t o f A . j a p o n i c u s from m o n t h l y c o l l e c t i o n s from Bath I s l a n d ( as g i v e n i n F i g u r e 14) a r e p l o t t e d o v e r t h e s a m p l i n g p e r i o d . C l e a r i n g E x p e r i m e n t A d e s c r i p t i v e summary i s p r e s e n t e d i n T a b l e I I o f t h e r e s u l t s o f t h e c l e a r i n g e x p e r i m e n t . M o n t h l y C o l l e c t i o n s D e s c r i p t i v e s t a t i s t i c s and g r a p h s o f t h e mean a x i s h e i g h t and o f f e r t i l i t y o f p l a n t s c o l l e c t e d o v e r t h e s t u d y p e r i o d a r e g i v e n i n C h a p t e r 2 ( T a b l e I and F i g u r e 14). 121 F i g u r e 32 shows t h e f r e g u e n c y o f h e i g h t c l a s s e s f o r t h e A . j a p o n i c u s p l a n t s c o l l e c t e d a t B a t h I s l a n d f o r t h e p e r i o d August 1978 t c J u l y 1979. P h c t o q r a p h i c Q u a d r a t s As n o t e d i n C h a p t e r 2, e r e c t a x e s o f A . j a p o n i c u s were s e e n t o be p r o d u c e d t h r o u g h o u t t h e y e a r a l t h o u g h g r o w th o f t h e s e a x e s was g r e a t e s t i n s p r i n g and e a r l y summer. I t was o b s e r v e d t h a t A . j a p o n i c u s c r u s t s were g r o w i n g and e x t e n d i n g d u r i n g t h e w i n t e r months and i n t o t h e s p r i n g . C r u s t s as w e l l a s e r e c t a x e s were d e l e t e r i o u s l y a f f e c t e d by t h e l a t e summer c o n d i t i o n s a t t h e s i t e . E r e c t a x e s i n t h e l a t e summer were a r e d / t a n c o l o u r and e r o d e d bacJc t o s t u b b s . These o f t e n had a l g a l e p i p h y t e s on them, which were n o t o b s e r v e d a t o t h e r t i m e s o f t h e y e a r . I t was n o t p o s s i b l e t o measure growth o f c r u s t s t h r o u g h o u t t h e y e a r f r o m t h e s l i d e s as o v e r s t o r e y growth p a r t i c u l a r l y o f A . j a p o n i c u s e r e c t a x e s , and s p e c i e s o f G i g a r t i n a , R ho do me l a , Sarg_assum, and F u c u s , p r e v e n t e d a c l e a r view o f t h e c r u s t s and s u b s t r a t e . A . j a p o n i c u s was o b s e r v e d t o grow o v e r R a l f s i a c r u s t s i n t h e f i e l d - t h e r e v e r s e was n e v e r o b s e r v e d . . B o t h A . j a p o n i c u s and R a l f s i a grew e v e r P e t r p c e l i s c r u s t s and a g a i n t h e r e v e r s e s i t u a t i o n was n e v e r s e e n . * The s c u l p i n , O l i q o c g t t u s m a c u l o s u s , was o f t e n p r e s e n t i n t h e s e p h o t o g r a p h s . The c o l o u r and p a t t e r n i n g o f t h i s f i s h d i s g u i s e i t w e l l n o r m a l l y , as i t l i e s amongst t h e 122 a l g a l c o v e r c l o s e t o t h e s u b s t r a t e . S e t t l e m e n t P l a t e s Over t h e t i m e t h a t t h e p l a t e s were i n t h e f i e l d , t h r e e were l o s t a p p a r e n t l y t o s t o r m o r l o g damage, and on some o f t h e p l a t e s t h a t r e m a i n e d t h e r e was b o t h b r e a k a g e and l o s s o f i n d i v i d u a l s l i d e s . No c r u s t o s e p h a s e s o f & . j a p o n i c u s were r e c o g n i s e d on t h e ground g l a s s s l i d e s . L i m p e t s were f o u n d on a l l p l a t e s t h a t r e m a i n e d i n t h e f i e l d . G e n e r a l O b s e r v a t i o n s I n November-December 1975 and 1976, t h e i n t e r t i d a l and s u b t i d a l r e g i o n of Bath I s l a n d were g r a z e d h e a v i l y by t h e g r e e n sea u r c h i n S t r o n q y l o c e n t r o t u s d r o e b a c h i e n s i s (De Wreede, 1980). No u r c h i n s were f o u n d i n t h e s t u d y s i t e o v e r t h e c o u r s e o f t h i s work a l t h o u g h a few v e r y s m a l l u r c h i n t e s t s were f o u n d washed up i n t h e d r i f t i n 1979.. I n t h e summer o f 1977 and 1978 l a r g e numbers o f P i s a s t e r o c h r a c e u s were seen i n t h e i n t e r t i d a l r e g i o n and a p p a r e n t l y were f e e d i n g on M y t i l u s as a t r a i l o f b r o k e n s h e l l s l a y i n t h e i r wake as t h e y p a s s e d a c r o s s t h e s h o r e . B o t h M y t i l u s and P i s a s t e r were f o u n d o n l y r a r e l y i n t h e summer o f 1979. . L i m p e t s ( C o l l i s e l l a p e l t a and Notoacmea scuturn) were s e e n i n t h e g u a d r a t s and t r a n s e c t s a l t h o u g h o n l y i n f r e g u e n t l y sampled by an i n t e r s e c t p o i n t . . The i s o p o d s 123 I d o t e a w o s n e s e n s k i i and Gnorimpsphaeroma O r e g o n e n s i s were b o t h f o u n d i n f r e g u e n t l y on u p r i g h t axes o f ft.japonicus. D i s c u s s i o n The r e s u l t s p r e s e n t e d h e r e - s p e c i f i c a l l y , s h i f t s t h a t were f o u n d i n t h e s p e c i e s c o m p o s i t i o n - s u g g e s t t h a t d u r i n g 1977-79, t h e i n t e r t i d a l community was r e c o v e r i n g f r o m t h e e f f e c t s o f s e a - u r c h i n g r a z i n g . I n t h e f i r s t summer o f t h i s s t u d y M y t i l i s e d u l i s was v e r y abundant, c o v e r i n g l a r g e a r e a s o f t h e i n t e r t i d a l r e g i o n . As i l l u s t r a t e d i n F i g u r e 27, t h e numbers o f M . e d u l i s f o u n d i n t h e g u a d r a t s and t r a n s e c t d r o p p e d d r a m a t i c a l l y i n t h e s e c o n d and t h i r d summer o f t h e s t u d y . A n o t h e r p r o n o u n c e d s h i f t i n s p e c i e s abundance was t h a t o f M i c r o c l a d i a b o r e a l i s ( r e f e r F i g u r e 27) which was much r e d u c e d i n abundance i n t h e 1979 summer fr o m t h e 1978 r e c o r d s . Even more marked was t h e v i r t u a l d i s a p p e a r a n c e o f C r y p t o s i p h o n i a w g o d i i . From t h e autumn o f 1977 t h r o u g h t h e summer o f 1978, C. w o o d i i was abundant and a p p a r e n t l y an i m p o r t a n t e l e m e n t o f t h e community ( r e f e r F i g u r e 28) p a r t i c u l a r l y i n t h e upper p o r t i o n o f t h e t r a n s e c t and i n g u a d r a t s QI and Q I I . T h i s p l a n t was v i r t u a l l y a b s e n t , however, i n l a t e 1978 and i n 1979.. The p r o p o r t i o n o f s p e c i e s which e x h i b i t a n n u a l o r p e r e n n i a l l i f e p a t t e r n s c a n be u s e d as an i n d i c a t o r o f t h e s u c c e s s i o n a l s t a t e o f t h e community ( e . g . i n a s t u d y o f 124 s u c c e s s i o n o f a l g a l c o m m u n i t i e s on b r e a k w a t e r s i n J a p a n , S a i t o , S a s a k i and Watanabe (1976) n o t e d t h a t , as a r u l e , t h e dominance of most p e r e n n i a l a l g a e i n c r e a s e d as t h e s u b s t r a t a became o l d e r ; N e w e l l and Tramer (1978), i n a s t u d y o f an h e r b a c e o u s p l a n t community d u r i n g s u c c e s s i o n , f o u n d a s h i f t f r o m a n n u a l t o p e r e n n i a l l i f e s t r a t e g i e s i . e . l o n g e r l i f e s p a n s ; Sousa (1980), i n a s t u d y o f r e c o l o n i s a t i o n and s u c c e s s i o n i n t h e i n t e r t i d a l , n o t e d t h a t p e r e n n i a l a l g a e were c h a r a c t e r i s t i c o f l a t e r s u c c e s s i o n a l s t a g e s ) . When t h e numbers o f p l a n t s w i t h e i t h e r a n n u a l o r p e r e n n i a l l i f e f o r m s were g r a p h e d o v e r t h e t i m e o f t h e s t u d y p e r i o d a t B a t h I s l a n d ( F i g u r e s 28 and 29) i n t e r e s t i n g t r e n d s a r e shewn. In t h e s p r i n g and e a r l y summer o f 1978 (March t o June) o f a l l f i v e g u a d r a t s , t h e r e i s a p p a r e n t l y a s e a s o n a l peak i n abundance o f a n n u a l s p e c i e s . I n t h e same p e r i o d i n 1979, however, t h i s peak i n t h e numbers o f a n n u a l p l a n t s i s f o u n d o n l y i n Q U I and QV, t h e two l o w e s t on t h e s h o r e . In QI,QII and QIV t h e r e i s a s h i f t t o a p r e d o m i n a n c e o f p e r e n n i a l p l a n t s . F i g u r e 26 shows t h a t o v e r t h e s t u d y p e r i o d t h e a n n u a l s p e c i e s Bhodomela l a r i x and S a r g a s s u m muticum i n c r e a s e i n abundance. These p l a n t s a r e f o u n d p a r t i c u l a r l y i n Q U I and QV and i t i s t h e i n c r e a s e i n t h e s e s p e c i e s t h a t c a u s e s t h e s e a s o n a l peak i n 1979 i n t h e numbers o f a n n u a l s p e c i e s i n t h e s e two g u a d r a t s . The p h e n o l o g i c a l s t u d y ( r e p o r t e d i n C h a p t e r 2) showed t h a t t h e A . j a p o n i c u s p o p u l a t i o n a t B a t h I s l a n d e x h i b i t e d a marked s e a s o n a l i t y w i t h r e s p e c t t o t h e t i m e o f maximal 125 g r o w t h , as measured by mean a x i s h e i g h t , and t h e p r o d u c t i o n of p l u r a n g i a . The p l a n t s were a t t h e i r s m a l l e s t i n l a t e summer, and t h i s c a n be c o r r e l a t e d w i t h h i g h l y s t r e s s f u l e n v i r o n m e n t a l c o n d i t i o n s (as i n d i c a t e d by c u l t u r e e x p e r i m e n t s r e p o r t e d i n C h a p t e r 3) o f l o w e r e d s a l i n i t y , i n c r e a s e d t e m p e r a t u r e and ext r e m e low t i d e s o c c u r r i n g i n t h e m i d d l e of t h e day. The p l a n t s c o l l e c t e d between J u l y and September have an e r o d e d r e d / t a n a p p e a r a n c e and a l t h o u g h e r e c t a x e s a r e b e i n g p r o d u c e d c o n t i n u a l l y , few a t t a i n m a t u r i t y a t t h i s t i m e . F i g u r e 32 i n d i c a t e s t h a t t h e s i z e d i s t r i b u t i o n o f e r e c t a x e s measured i n t h e p h e n o l o g i c a l s t u d y i s s t r o n g l y skewed t o t h e l o w e r s i z e c l a s s e s . The t o t a l number o f A . j a p o n i c u s o c c u r r i n g i n a l l t h e g u a d r a t s ( F i g u r e 30) shows a s t e a d y and s t e e p i n c r e a s e i n t h e p o p u l a t i o n f r o m O c t o b e r t o March i n b o t h 1978 and 1979. In 1978 t h e r e i s a d e c r e a s e i n t h e numbers of £• j§J2Pjiicus r e c o r d e d between A p r i l and J u n e . In 1979 t h e r e i s a s m a l l d e c r e a s e i n t h e p l a n t numbers r e c o r d e d f o r A p r i l . As i l l u s t r a t e d i n F i g u r e 30, i n 1978 and 1979 A p r i l / M a y i s t h e t i m e t h a t t h e mean a x i s h e i g h t (and a l s o t h e f e r t i l i t y o f t h e p o p u l a t i o n ) ( F i g u r e 14) a r e a t t h e i r peak.. In J u n e - August o f 1977-1979 a s e c o n d a r y peak and d e c l i n e i s s e e n i n t h e A . j a p o n i c u s numbers. . S e v e r a l f a c t o r s a r e a p p a r e n t l y i n v o l v e d i n t h e s e p o p u l a t i o n f l u c t u a t i o n s r e p o r t e d h e r e . The d e c l i n e i n the A . j a p o n i c u s p o p u l a t i o n i n t h e s p r i n g and e a r l y summer c o i n c i d e s w i t h t h e t i m e o f maximal g r o w t h . 126 T h i s s u g g e s t s t h e i n v o l v e m e n t o f i n t r a s p e c i f i c c o m p e t i t i o n as H a r p e r (1977) s t a t e s "...one o f t h e h i g h r i s k s t a t e s i n t h e l i f e c y c l e . . . i s t h e s e a s o n a l h a z a r d when t h e s u r v i v o r s a r e g r o w i n g f a s t . The p e r i o d c f r a p i d growth i s t h e t i m e a t w hich i n d i v i d u a l s make t h e i r h e a v i e s t demands on e n v i r o n m e n t a l r e s o u r c e s and a r e most l i k e l y t o i n t e r f e r e w i t h t h e i r n e i g h b o u r ' s a c t i v i t i e s . T h i s would e x p l a i n t h e h i g h m o r t a l i t y r i s k a t t h i s t i m e and a l s o t h e f a c t t h a t t h e r i s k i s s p e c i e s s p e c i f i c . E a c h p l a n t i s most l i k e l y t o s u f f e r f r o m t h e p r e s e n c e o f a n e i g h b o u r o f i t s own s p e c i e s b e c a u s e t h e i r p h e n o l o g i e s w i l l be s y n c h r o n o u s . . I f i n t r a s p e c i f i c c o m p e t i t i o n i s i n v o l v e d i n t h i s d e c l i n e i n ft. •japonicus numbers a t a t i m e when i t s maximal growth i s b e i n g r e c o r d e d i t would be e x p e c t e d t h a t t h e r e i s a r e l a t i o n s h i p between t h e r i s k o f d e a t h and t h e d e n s i t y o f t h e p o p u l a t i o n . To t e s t t h i s h y p o t h e s i s * e x p e r i m e n t s s h o u l d be c a r r i e d out on p o p u l a t i o n s o f ft. -japonicus a t d i f f e r e n t d e n s i t i e s and d a t a g a t h e r e d on t h e l o n g e v i t y / m o r t a l i t y o f b o t h c r u s t s and u p r i g h t a x e s . I t has been r e p o r t e d t h a t p l a n t s g r o w i n g under d e n s i t y s t r e s s have a skewed d i s t r i b u t i o n o f p l a n t w e i g h t ( H a r p e r , 1 9 7 7 ) . . No measurements o f b i o m a s s were made i n t h i s s t u d y , b u t t h e skewed d i s t r i b u t i o n o f a x i s h e i g h t ( F i g u r e 32) would seem t o s u p p o r t t h e s u g g e s t i o n o f t h e r o l e o f i n t r a s p e c i f i c c o m p e t i t i o n . A n o t h e r f a c t o r t h a t may be i n v o l v e d i n t h e d e c l i n e i n ft> j a p o n i c u s numbers a t t h i s t i m e was t h e i n c r e a s e i n g r o w t h 127 and abundance o f o t h e r s p e c i e s - t h a t i s , i n t e r s p e c i f i c c o m p e t i t i o n . P a r t i c u l a r l y i n 1978, M j c r o c l a d i a was a t i t s peak abundance o v e r t h e p e r i o d o f d e c l i n e i n t h e A . j a p o n i c u s numbers. The o t h e r a n n u a l p l a n t s w h i c h i n c r e a s e d i n i m p o r t a n c e (as measured by t h e i r numbers i n t h e g u a d r a t s and t r a n s e c t ) a t t h i s t i m e were C r y p t p s i p h p n i a ( i n 1978 o n l y ) , Sargassum, Rhodomela, and F u c u s ( F i g u r e 3 1 ) . I n t e r e s t i n g l y , H a l f s i a and P e t r o c e l i s , t h e two o t h e r c r u s t s o c c u r r i n g i n t h e s t u d y s i t e were l e a s t a bundant o v e r t h e same p e r i o d o f A p r i l - J u n e . . T h i s may i n d i c a t e t h a t t h e a n n u a l p l a n t s o v e r g r o w t h e c r u s t s and t h a t t h e c r u s t s were o b s c u r e d by a t e m p o r a r y o v e r s t o r e y b u t were s t i l l p r e s e n t i n t h e community. I n t h e c a s e o f A . j a p o n i c u s t h i s i s l e s s l i k e l y as t h e p r e s e n c e o f t h e e r e c t a x e s means t h a t t h e s e c r u s t s a r e ' l e s s e a s i l y o b s c u r e d . F u r t h e r e x p e r i m e n t a t i o n i.s needed t o d e t e r m i n e t h e r e l a t i v e c o n t r i b u t i o n o f i n t r a - and i n t e r s p e c i f i c c o m p e t i t i o n t o t h e s e d y n a m i c s o f t h e A . j a p o n i c u s p o p u l a t i o n . The peak i n A . j a p o n i c u s numbers o c c u r r i n g i n l a t e summer happens 2-4 months a f t e r t h e peak i n f e r t i l i t y , and i t i s p r o b a b l y due t o s e t t l e m e n t and c o l o n i s a t i o n by s p o r e s and t h e growth o f t h e new r e s u l t a n t c r u s t s . The s h a r p d e c r e a s e i n t h i s peak (August/September 1977-1979) c o i n c i d e d w i t h a d e c r e a s e i n t h e mean a x i s h e i g h t o f t h e p o p u l a t i o n . Young p l a n t s a r e u n l i k e l y t o be a b l e t o s u r v i v e t h e e n v i r o n m e n t a l e x t r e m e s e x p e r i e n c e d a t t h i s t i m e o f t h e y e a r a t B a t h I s l a n d (- t h i s i s s u p p o r t e d by t h e work 128 r e p o r t e d i n C h a p t e r 3 ) , and t h e r e a r e few c r e v i c e s o r s haded a r e a s where new c r u s t s c o u l d e s c a p e t h e r i g o u r s o f t h e e n v i r o n m e n t . I t was hoped t h a t t h e s e t t l e m e n t p l a t e s would a l l o w s t u d y o f e a r l i e r s t a g e s o f c r u s t d e v e l o p m e n t i n t h e f i e l d and a l s o would i n d i c a t e t h e t i m e o f y e a r when maximal s e t t l e m e n t o c c u r r e d . The s e t t l e m e n t p l a t e s need t o be r e d e s i g n e d t o e x c l u d e g r a z e r s , i n t h i s c a s e l i m p e t s , w hich a p p e a r e d t o be a f f e c t i n g t h e growth o f a l g a e on t h e p l a t e s . L i m p e t s h a ve been r e p o r t e d t o c o n t r o l a l g a l g r o w t h by consuming young p l a n t s and t h e r e b y a f f e c t i n g r e c r u i t m e n t (Lodge, 1948; J o n e s , 1948; S o u t h w a r d , 1958)., O t h e r m a t e r i a l s might a l s o prove more s u c c e s s f u l s u b s t r a t e s f o r A . j a p o n i c u s p l a n t s . H a r l i n and L i n d b e r g h (1977) d e m o n s t r a t e d s u b s t r a t e s u r f a c e r e l i e f c an r e g u l a t e t h e d e v l o p m e n t of an a l g a l community: i n c r e a s e d a l g a l s e t t l e m e n t o c c u r r e d w i t h an i n c r e a s e i n p a r t i c l e s i z e . A l t h o u g h t h e s l i d e s i n my s t u d y had b een r o u g h e n e d , p o s s i b l y t h e y were s t i l l t o o smooth t o a l l o w f o r t h e d e v e l o p m e n t of t h e A . j a p o n i c u s c r u s t s . The r e s u l t s o f t h e c l e a r i n g e x p e r i m e n t ( T a b l e I I ) s u p p o r t t h e p r e v i o u s e x p l a n a t i o n o f t h e q u a d r a t r e s u l t s . The g u a d r a t s c l e a r e d i n A u g u s t had a dense c o v e r o f A . j a p o n i c u s a y e a r l a t e r . B o th p a i r s o f q u a d r a t s c l e a r e d i n May and J u n e , however, had o n l y s c a t t e r e d A . j a p o n i c u s p l a n t s o c c u r r i n g on them a f t e r a y e a r , d e s p i t e t h e f a c t t h a t t h e r e was a much h i g h e r p e r c e n t a g e o f t h e A . j a p o n i c u s p o p u l a t i o n f e r t i l e i n May and J u n e t h a n i n A u g u s t ( F i g u r e 129 1 4 ) . A p p a r e n t l y any A . j a p o n i c u s s e t t l e m e n t which d i d o c c u r on t h e s e s u b s t r a t e s i n Hay-June was u n a b l e t o s u r v i v e due t o t h e e x t r e m e s o f t h e e n v i r o n m e n t . A l s o t h e y may be u n a b l e t o compete s u c c e s s f u l l y w i t h e p h e m e r a l s p e c i e s which o c c u r r e d a t l e a s t i n t h e g u a d r a t s T2 and T6 c l e a r e d i n May. I t a p p e a r s t h a t t h e r e p r o d u c t i v e 'peak' o f A . j a p o n i c u s between A p r i l and J une a t B a t h I s l a n d makes a n e g l i g i b l e c o n t r i b u t i o n t o i t s s u c c e s s f u l s e t t l e m e n t and c o l o n i s a t i o n a t t h i s s i t e . B o t h t h e r e s u l t s o f t h e c l e a r i n g e x p e r i m e n t and t h e g u a d r a t s s u g g e s t t h a t t h e s i g n i f i c a n t r e p r o d u c t i v e p e r i o d i n t e r m s o f r e s u l t a n t r e c r u i t m e n t t o t h e p o p u l a t i o n o c c u r s i n t h e autumn, w i n t e r and e a r l y s p r i n g . I n c u l t u r e work ( d e s c r i b e d i n C h a p t e r 1) t h e c r u s t s were f o u n d t o s p r e a d by f o r m i n g " s a t e l l i t e - d i s c s " - i t would be o f c o n s i d e r a b l e v a l u e t o t h i s p l a n t i f t h i s t y p e o f v e g e t a t i v e growth can c c c u r i n t h e f i e l d and p a r t i c u l a r l y i f i t i s o p e r a t i v e i n t h e autumn p r i o r t o t h e o n s e t of i n c r e a s e d e r e c t a x i s d e v e l o p m e n t and i n c r e a s e d f e r t i l i t y i n t h e p o p u l a t i o n . The A . j a p o n i c u s p o p u l a t i o n as a whole i n c r e a s e d i n s i z e o v e r t h e s t u d y p e r i o d ( F i g u r e 30) a l t h o u g h as i l l u s t r a t e d by F i g u r e s 23, 24, and 25, t h e same t r e n d s a r e n o t o c c u r r i n g a t a l l t h e v e r t i c a l h e i g h t s s a m p l e d . An i n c r e a s e i n A« j a p o n i c u s numbers was n o t o b s e r v e d i n e i t h e r Q U I o r QV, o r i n PTIV where Sargassum and Bhodomela were becoming i n c r e a s i n g l y i m p o r t a n t ( b o t h b e i n g more numerous and d o m i n a t i n g t h e a r e a i n t e r m s o f t h e o v e r s t o r e y c o v e r 130 t h e y p r o v i d e d ) . QI, Q I I , and' QIV and t h e upper t h r e e s e c t i o n s of t h e t r a n s e c t showed an i n c r e a s e i n t h e numbers o f fl.japonicus r e c o r d e d i n t h e s e c o n d y e a r o f t h i s s t u d y . I n mid-1977 and mid-1978, fl.japonicus numbers were h i g h e s t i n QV and PTIV - a s i t u a t i o n w hich i s v i r t u a l l y r e v e r s e d by t h e f o l l o w i n g y e a r . flt o t h e r s i t e s i n B r i t i s h C o l u m b i a where I have c o l l e c t e d A . j a p o n i c u s , i t s v e r t i c a l d i s t r i b u t i o n i n t h e i n t e r t i d a l r e g i o n i s g e n e r a l l y much more r e s t r i c t e d t h a n a t B a t h I s l a n d . ( T h i s i s d i s c u s s e d i n C h a p t e r 2.) The s h i f t s e e n h e r e i n t h e v e r t i c a l p o s i t i o n o f A . j a p o n i c u s 1 maximal abundance o v e r two y e a r s may be an i n d i c a t i o n o f d i f f e r i n g c o m p e t i t i v e a b i l i t y i n A. j a p o n i c u s at d i f f e r e n t h e i g h t s * I n t h e l o w e r p o r t i o n o f i t s r a n g e A . j a p o n i c u s was c o n t e n d i n g w i t h i n c r e a s i n g c o v e r from Sargassum and Rhgdomela.. The s u c c e s s o f t h e s e two s p e c i e s and t h e c o ncommitant d e c l i n e i n A . j a p o n i c u s numbers may be r e l a t e d t o t h e o v e r s t o r e y f ormed by Sargassum and Shodomela. T h i s o v e r s t o r e y c o u l d be a f f e c t i n g t h e A . j a p o n i c u s p l a n t s by s e v e r e l y r e s t r i c t i n g t h e i n c i d e n t l i g h t . The i n c r e a s e i n Sargassum and Rhodpmela may a l s o be due t o t h e i r g e r m l i n g v i g o u r and growth b e i n g s u p e r i o r t o t h a t i n A . j a p o n i c u s . E x p e r i m e n t s , s i m i l a r t o t h o s e s u g g e s t e d i n C h a p t e r 2, t o t e s t i f t h e d e c l i n e i n A . j a p o n i c u s numbers i n t h e l o w e r i n t e r t i d a l r e g i o n i s due t o c o m p e t i t i o n w i t h t h e o v e r s t o r e y s p e c i e s c o u l d be p e r f o r m e d by r e m o v a l o f Sargassum and Rhodpmela and s u b s e g u e n t o b s e r v a t i o n o f t h e fl.japonicus p o p u l a t i o n t o s e e 131 i f i t i n c r e a s e s i n abundance or extends i t s v e r t i c a l range. Predators and h e r b i v o r e s have been shown t o play important r o l e s i n shaping community s t r u c t u r e and i n i n f l u e n c i n g d i s t r i b u t i o n and abundance of i n t e r t i d a l and s u b t i d a l p l a n t s and s e s s i l e animals (e.g. Jones and Kain, 1967; Paine and Vadas, 1969; Paine, 1971; F o s t e r , 1975; Lutchenco and Menge, 1978; Lubchenco, 1978, 1980) . Although a per i o d of heavy u r c h i n g r a z i n g had preceded t h i s study, u r c h i n s were absent from the s i t e over the study p e r i o d , and thus any e f f e c t s they had on the community were h i s t o r i c a l . . T h e dynamics of the p o p u l a t i o n s of M y t i l u s and P i s a s t e r are f a r from c l e a r . Whether the settlement of p e r e n n i a l algae i n the zones p r e v i o u s l y occupied by M y t i l u s prevented s u c c e s s f u l establishment of the M y t i l u s , or whether the g r a z i n g by P i s a s t e r r e s u l t e d i n a s m a l l p o p u l a t i o n of M y t i l u s that was unable t o r e p l a c e i t s e l f , or whether the changes i n the p o p u l a t i o n are part of a c y c l e which spans a l o n g e r p e r i o d than t h a t of t h i s study, i s not known. Whilst l i m p e t s may be grazing germlings and young c r u s t s of A. j a p o n i c u s , no h e r b i v o r e s which were a f f e c t i n g the growth of mature A.japonicus c r u s t s and e r e c t axes were seen. Thus the r o l e of both h e r b i v o r y and p r e d a t i o n i n t h i s community remains u n c e r t a i n . From t h i s study i t appears t h a t the mid- to lower- i n t e r t i d a l community at Bath I s l a n d was i n the process of r e c o v e r y i n 1977-79 from s e a - u r c h i n g r a z i n g which occ u r r e d i n 1975 and 1976. Algae dominate the space a v a i l a b l e i n the 1 3 2 i n t e r t i d a l r e g i o n and a l s o s t r u c t u r e t h e f o r m o f t h e community. M c l l u s c a n h e r b i v o r e s a p p a r e n t l y have a m i n o r e f f e c t on t h e mature s t r u c t u r a l l y d o m i n a n t p l a n t s i n t h i s community a l t h o u g h t h e y may be i n f l u e n c i n g t h e r e c r u i t m e n t o f g e r m l i n g s and a f f e c t i n g t h e g r o w t h o f e p h e m e r a l s p e c i e s . T h i s t y p e o f s i t u a t i o n has been r e p o r t e d i n t h e l o w e r i n t e r t i d a l zone o f s h o r e s o f New E n g l a n d by L u b c h e n c o and Menge (1978) and by D a y t o n (1975) on t h e Washington c o a s t . A d i s c u s s i o n c f A . j a p o n i c u s ' growth and l i f e h i s t o r y f e a t u r e s i s p r e s e n t e d i n t h e G e n e r a l D i s c u s s i o n . 133 F i g u r e 2 1. Map o f B a t h I s l a n d i n t h e S t r a i t o f G e o r g i a , E r i t i s h C o l u m b i a . 134 135 F i g u r e 22. D i a g r a m o f f i e l d s i t e a t B a t h I s l a n d i n d i c a t i n g r e l a t i v e p o s i t i o n s i n i n t e r t i d a l r e g i o n o f g u a d r a t s and t r a n s e c t used i n t h i s s t u d y . T r a n s e c t and q u a d r a t p c i n t s a r e f o l l o w i n q h e i g h t s ( i n m) above c h a r t datum:- ET 0 - 1.98 Q I -• 1.74 PT 1 - 1. 80 Q I I -• 1.36 ET 2 - 1.66 Q I I I -• 1.02 PT 3 - 1. 50 Q IV -- 1.14 PT n - 1.34 Q V -• 0.92 PT 5 - 1. 18 ET 6 - 0.98 PT 7 - 0.80 ET 8 - 0.74 PT 9 - 0. 56 ET 10 - 0. 48 ( h e i g h t o f t o p l e f t hand c o r n e r o f Q I-V measured) r -o o 00 0 0 •o o > 0 m 0 136 0 0 lO 137 F i g u r e 23* Numbers o f ft.japonicus p l a n t s r e c o r d e d o v e r s t u d y p e r i o d i n QI and Q I I . {Sample s i z e of 81 p o i n t s i n 0.5 m x 0.5 m q u a d r a t . )  139 F i g u r e 2 4. Numbers c f ft.japonicus p l a n t s r e c o r d e d o v e r s t u d y p e r i o d i n Q I V r Q U I , and QV. (Sample s i z e o f 81 p o i n t s i n 0.5 m x 0.5 m q u a d r a t . ) 1977 1978 1979 141 F i g u r e 2 5. Number of A . j a p o n i c u s p l a n t s r e c o r d e d i n t r a n s e c t o v e r s t u d y p e r i o d . Upper 8 m o f t r a n s e c t were d i v i d e d i n t o f o u r , 2 m l e n g t h s and l a b e l l e d P T I - P T I V , from upper t o l o w e r i n t e r t i d a l r e g i o n r e s p e c t i v e l y . (40 sample p o i n t s i n e a c h 2 m l e n g t h o f t r a n s e c t . ) Number of A. japonicus plants 143 / F i g u r e 26. Numbers o f Rhpdomela, Sargassum, and C r y p t o s i g h p n i a p l a n t s r e c o r d e d i n QI-QV and PT1-PT10 o v e r s t u d y p e r i o d .  145 F i g u r e 27* Numbers o f i n d i v i d u a l s o f M ^ t i l u s and M i c r o c l a d i a r e c o r d e d i n QI-QV and PT1-PT10 o v e r s t u d y p e r i o d . 1977 1978 1979 M o n t h s CD 147 F i g u r e 28. Numbers o f p l a n t s w i t h e i t h e r a n n u a l o r p e r e n n i a l h a b i t r e c o r d e d from QI, Q I I , and QIV o v e r s t u d y p e r i o d . Number of p l a n t s 149 F i g u r e 29. .Numbers of p l a n t s w i t h e i t h e r a n n u a l o r p e r e n n i a l h a b i t r e c o r d e d f r o m Q U I and QV o v e r s t u d y p e r i o d . . 80- 60—\ 40-H 20-J — a n n u a l — p e r e n n i a l Q III QV 60- 40-H 20-H * i—i—i—r O D 1977 J A 1978 M o n t h s i — i — i — i — i — r O D F T — i — i — r J A 1979 cn o 151 F i g u r e 30. Mean a x i s h e i g h t o f A . j a p o n i c u s and numbers o f A . j a p o n i c u s p l a n t s r e c o r d e d f r o m QI-QV o v e r s t u d y p e r i o d . M o n t h s ro 153 F i g u r e 31. O c c u r r e n c e o f 8 s p e c i e s o f a l g a e i n QI-V and PT1-10 e v e r s t u d y p e r i o d . Shaded a r e a s r e p r e s e n t t i m e o f maximal abundance. Cryptopleura Petalonia Scytosiphon Ulva Gigartina Petrocelis Fucus Ralfsia • A W77 I I I I • i — i — r F A J A 1978 MONTHS " i — i — i — i — i— r D F A I I I J A 1979 cn 155 F i g u r e 3 2. F r e g u e n c y d i s t r i b u t i o n o f p l a n t s o f ft.japonicus c o l l e c t e d from A u g u s t 1978 t o J u l y 1979 a c c o r d i n g t o u p r i g h t a x i s h e i g h t . .N=250 E V Ci d o m ci II c T Ci T oo Ci Ci T o Ci T ~r Ci 156 Ci oo Ci N Ci Ci m ci fi Ci Ci Ci O Ci oo r - 1 CO E w I O » oo N <o rt ci '3 % Summary of quadrat c l e a r i n g experiment. Macroscopic algae present in the quadrats are l i s t e d i n order of understorey to overstorey habit. QUADRAT NUMBERS AND DATE OF CLEARING OBSERVATIONS JUNE 20, 1978 JULY 18, 1978 DECEMBER 8, 1978 JULY 12, 1979 2 6 (May 24, 1978) dense green turf dense cover of Ulva sp. R a l f s i a , P e t r o c e l i s , crustose c o r a l l i n e , Petalonia, Ulva Colpomenia, Fucus, Sargassum, (mussels scattered) dense overstorey of Fucus and Sargassum (understorey obscured from view) dense green turf dense cover of Ulva sp. R a l f s i a , P e t r o c e l i s , Ulva, Petalonia, Colpomenia, Fucus, Sargassum, (mussels-scattered) P e t r o c e l i s , Ulva dense overstorey of Fucus 4 8 (June 20, 1978) no macroscopic algae - (limpets and sculpins) rock v i r t u a l l y bare - R a l f s i a , P e t r o c e l i s , Ulva, Petalonia, Colpomenia R a l f s i a , P e t r o c e l i s . Analipus, Ulva, Gigartina - no macroscopic algae - (limpets and sculpins) mostly bare rock - R a l f s i a , Petalonia, Ulva, Fucus mostly bare rock - R a l f s i a , Fucus, filamentous greens io - - Analipus (abundant), R a l f s i a , P e t r o c e l i s , Ulva Analipus, P e t r o c e l i s . Fucus. Gigartina 12 (August 15, 1978) - Analipus (very dense). P e t r o c e l i s , Colpomenia Analipus (very dense). Ulva, P e t r o c e l i s , R a l f s i a , G igartina, Fucus Controls 1.3,5 7,9,11 (not cleared) R a l f s i a , Analipus, P e t r o c e l i s , Gigartina, Ulva, Microcladia. Fucus, Rhodomela, Cryptosiphonia, S.irgassum R a l f s i a , Analipus, P e t r o c e l i s , Gigartina, Ulva, Colpomenia, Fucus, Petalonia, Sargassum Analipus, R a l f s i a , P e t r o c e l i s , Gigartina, Ulva, Microcladia, Fucus (malor overstorey plant) 158 CHAPTER 5, A comparison of Caepidium antarcticum J.Ag. and A n a l i p u s j a p o n i c u s (Harv.) Wynne - (Phaeophyta) 159 I n t r o d u c t i o n The m o n o t y p i c genus C a e p i d i u m a n t a r c t i c u m J.Ag» i s f o u n d i n s u b a n t a r c t i c w a t e r s . I t has been r e c o r d e d f r o m F u e g i a , t h e F a l k l a n d I s l a n d s ( t h e t y p e l o c a l i t y ) , S o u t h G e o r g i a , l i e s K e r g u e l e n , and C a m p b e l l I s l a n d , New Z e a l a n d * T h i s brown a l g a c o n s i s t s o f a c r u s t o s e s y s t e m f r o m w h i c h a r i s e b o t h u p r i g h t a x e s and w a r t - o r b l a d d e r - l i k e p r o l i f e r a t i o n s . The u p r i g h t C h o r d a r i a - l i k e a x e s a r e g e n e r a l l y u n b r a n c h e d and c l o t h e d w i t h s h o r t numerous l a t e r a l s which b e a r u n i l o c u l a r s p o r a n g i a . These e r e c t s h o o t s d e v e l o p f r o m c u p - l i k e d e p r e s s i o n s f o u n d a t t h e t i p s of t h e b r a n c h i n g p r o s t r a t e t h a l l u s . The b l a d d e r s , w h i c h have been d e s c r i b e d a s C o l p o m e n i a - l i k e , d e v e l o p out o f t h e c r u s t and form p l u r a n g i a i n s o r i . T hus, an i n d i v i d u a l o f t h i s s p e c i e s can p o s s e s s t h r e e m o r p h o l o g i c a l p h a s e s . T h i s d i s t i n c t i v e m o r p h o l o g y has posed a p r o b l e m i n i n t e r p r e t a t i o n , and C a e p i d i u m has been t r e a t e d by t a x o n o m i s t s i n b a s i c a l l y two ways; c l a s s i f i e d on t h e b a s i s of the u p r i g h t system i n t o t h e C h o r d a r i a c e a e ( e . g . A g a r d h , 1882; S k o t t s b e r g , 1921, K y l i n , 1933, 1940; L i n d a u e r , 1947; L i n d a u e r , Chapman and A i k e n , 196 1; P a p e n f u s s , 1964; Kim, 1971), o r p l a c e d i n t h e E n c o e l i a c e a e o r P u n c t a r i a c e a e on t h e b a s i s o f c r u s t and b l a d d e r c h a r a c t e r i s t i c s (e.g* S k o t t s b e r g , 1907; R e i n b o l d , 1908; L e v r i n g , 1945, 1960). S k o t t s b e r g (1907,1921) g i v e s a d e t a i l e d a c c o u n t o f t h e m o r phology and anatomy of C a e p i d i u m and p a r t i c u l a r l y i n t h e 160 l a t e r p a p e r d i s c u s s e s some f e a t u r e s of i t s e c o l o g y and p h e n o l o g y . C o t t o n (19 15) d i d n o t a c c e p t S k o t t b e r g ' s (1907) i n t e r p r e t a t i o n o f t h i s p l a n t a s p o s s e s s i n g c r u s t , u p r i g h t and b l a d d e r p h a s e s , and g u o t e s Dr. Yendo (who had e x a m i n e d t h e t y p e m a t e r i a l o f C a e p i d i u m ) as a l s o d i s a g r e e i n g w i t h S k o t t s b e r g . C o t t o n r e p o r t s t h a t Yendo c o n s i d e r e d C a e p i d i u m t o r e s e m b l e c l o s e l y C h o r d a r i a a b i e t i n a (now A n a l i p u s j a p o n i c u s ) p o s s e s s i n g " c r e e p i n g b r a n c h e s and u p r i g h t s h o o t s " , and t h a t t h e b l a d d e r - l i k e f r o n d s were g u i t e d i s t i n c t c o n s i s t i n g o f "young p l a n t s o f C o l p o m e n i a s i n u o s a o r an a l l i e d s p e c i e s " . A l t h o u g h C o t t o n (pp.168-169) g i v e s t h e i m p r e s s i o n o f a g r e e i n g w i t h Yendo t h a t t h e b l a d d e r i s n o t p a r t o f t h e c r u s t and e r e c t s y s t e m o f C a e p i d i u m , he s t a t e s l a t e r (under C o l p o m e n i a , p.173) " t h e f o r k e d b a s a l b r a n c h e s . . . . would a p p e a r t o be p a r t o f t h e C o l p o m e n i a p l a n t (and n o t o f t h e C h o r d a r i a ) and t h e s e a r e u n l i k e a n y t h i n g f o u n d i n t h e o r d i n a r y C o l p o m e n i a o f E u r o p e " . S k o t t s b e r g (1921) r e - i t e r a t e d t h a t t h e b l a d d e r - b e a r i n g C a e p i d i u m was g u i t e d i s t i n c t from C o l p o m e n i a , and t h a t t h e c r u s t , b l a d d e r s and u p r i g h t a x e s were a l l p a r t of t h e same s p e c i e s . S k o t t s b e r g (1921) p l a c e d C a e p i d i u m i n t h e C h c r d a r i a c e a e , c h a n g i n g h i s e a r l i e r view o f i t s c l a s s i f i c a t i o n a f t e r f i n d i n g u n i l o c u l a r s p o r a n g i a i n t h e e r e c t a x e s ( a l t h o u g h t h e s e had i n f a c t been d e s c r i b e d by Agardh ( 1 8 8 2 ) ) . S k o t t s b e r g (1921) s p e c u l a t e d on t h e r e l a t i o n s h i p between t h e l i f e h i s t o r y o f t h i s p l a n t and i t s m o r p h o l o g y , 161 s u g g e s t i n g t h a t t h e b l a d d e r s and e r e c t a x e s might be a l t e r n a t i n g g e n e r a t i o n s and o c c u r on d i f f e r e n t c r u s t s . He n o t e d t h e d i f f i c u l t y o f s e p a r a t i n g i n d i v i d u a l s due t o t h e o v e r l a p p i n g n a t u r e o f t h e c r u s t s . The u p r i g h t s h o o t s were o n l y f o u n d i n t h e summer months (November t o F e b r u a r y ) w h i l s t t h e b l a d d e r s and c r u s t s were p r e s e n t y e a r r o u n d . K y l i n (1933) i n t e r p r e t e d S k o t t s b e r g ' s (1921) work s l i g h t l y d i f f e r e n t l y . He a c c e p t e d t h a t t h e t h r e e m o r p h o l o g i c a l p h a s e s were a l l p a r t o f t h e same s p e c i e s , and c o n s i d e r e d t h a t t h e y a l l c o u l d be p r e s e n t on t h e same i n d i v i d u a l . K y l i n s u g g e s t e d t h a t t h e C h o r d a r i a - l i k e u p r i g h t s a r e d i p l o i d , u n d e r g o i n g a l t e r n a t i o n o f g e n e r a t i o n s w i t h m i c r o s c o p i c g a m e t o p h y t e s . The b l a d d e r s were a l s o i n t e r p r e t e d t o be d i p l o i d , and t h e p l u r a n g i a were s e e n as s e r v i n g as a means o f v e g e t a t i v e r e p r o d u c t i o n . . T h e r e a r e a number o f r e f e r e n c e s i n t h e l i t e r a t u r e t o t h e s i m i l a r i t i e s between C a e p i d i u m and t h e now A n a l i p u s j a p o n i c u s ( p r e v i o u s l y C h o r d a r i a a b i e t i n a ' and H e t e r g c h o r d a r i a a b i e t i n a ) . . K j e l l m a n (1889) n o t e d s i m i l a r i t i e s between A n a l i pus f u s i f o r m i s (now A. . f i j i f o r m i s ) and C a e p i d i u m as d i d K y l i n (1940) . . K y l i n ( 1 9 4 0 ) , as w e l l as Yendo i n C o t t o n (1915) and S k o t t s b e r g ( 1 9 2 1 ) , commented on t h e p o s s i b l e r e l a t i o n s h i p between A n a l i p u s j a p o n i c u s and C a e p i d i u m . . S k o t t s b e r g comments on t h e l i k e n e s s between t h e s e two p l a n t s n o t i n g t h a t b o t h p o s s e s s h a i r s o f t h e same t y p e on t h e h o r i z o n t a l t h a l l u s . T h i s m o r p h o l o g i c a l and a n a t o m i c a l s t u d y o f 162 C. a n t a r c t i c u m and c o m p a r i s o n wi t h A . j a p o n i c u s has been u n d e r t a k e n t o a p p r o a c h t h e q u e s t i o n o f t h e a f f i n i t i e s o f t h e s e g e n e r a r a i s e d i n t h e l i t e r a t u r e . M a t e r i a l s and Methods The m a t e r i a l c f C . , a n t a r c t i c u m used i n t h i s s t u d y came from a v a r i e t y c f s o u r c e s : - (1) t h e t y p e s p e c i m e n - borrowed f r o m t h e B o t a n i c a l Museum, Lund ( L D ) . T h i s i s shown i n F i g u r e 33.. (Yendo i n C o t t o n (1915) c o n s i d e r e d t h a t t h e t y p e o f t h i s s p e c i e s " r e s e m b l e s v e r y much an o l d and n e a r l y d e c a y e d p l a n t o f C h o r d a r i a a b i e t i n a " . ) (2) wet p r e s e r v e d m a t e r i a l c o l l e c t e d by S k o t t s b e r g i n So u t h G e o r g i a , t h e F a l k l a n d s , and T i e r r a d e l Fuego - l e n t by Dr E.B. S e a r l e s (3) wet p r e s e r v e d m a t e r i a l from A r g e n t i n a and C h i l e - l e n t by Dr B.B. S e a r l e s (4) wet p r e s e r v e d m a t e r i a l f r o m C a m p b e l l I s l a n d , New Z e a l a n d - borro w e d f r o m B o t a n y D i v i s i o n , D.S.I.E., New Z e a l a n d (CHE) (5) wet p r e s e r v e d m a t e r i a l f r o m C h i l e - l o d g e d i n t h e O.E.C. h e r b a r i u m (6) h e r b a r i u m s p e c i m e n s from C h i l e - U.B.C. c o l l e c t i o n s (7) two s p e c i m e n s from t h e K e r g u e l e n s - borrowed f r o m t h e B r i t i s h Museum, London (BM). 163 The m a t e r i a l o f A n a l i p u s j a p o n i c u s used i n t h i s s t u d y i s l i s t e d i n C h a p t e r 1. In a d d i t i o n t h e t y p e s p e c i m e n and two o t h e r s p e c i m e n s o f A . j a p o n i c u s b o r r o w e d from T r i n i t y C o l l e g e , D u b l i n , were examined. M a t e r i a l was e i t h e r h a n d - s e c t i o n e d o r embedded i n g l y c o l m e t h a c r y l a t e , s e c t i o n e d w i t h a S o r v a l JBU m i c r o t o m e , and mounted on g l a s s s l i d e s * S e c t i o n s were s t a i n e d w i t h t o l u i d i n e b l u e 0 (TBO) o r p e r i o d i c a c i d - S c h i f f • s (PAS) (sometimes w i t h a c o u n t e r s t a i n o f a n i l i n e b l u e - PAS/AN) f o l l o w i n g t h e methods g i v e n i n C h a p t e r 1. R e s u l t s and D i s c u s s i o n M o r p h o l o g y The m o r p h o l o g y o f t h e c r u s t and e r e c t a x i s of C . a n t a r c t i c u m , as f i g u r e d by S k o t t s b e r g (1921 - f i g u r e 1 2 a ) , i s v e r y s i m i l a r t o t h a t o f A n a l i p u s j a p o n i c u s . . B o t h p l a n t s have a b a s a l s y s t e m c o n s i s t i n g o f c y l i n d r i c a l , b r a n c h i n g axes f o r m i n g a l o b e d and m u l t i l a y e r e d c r u s t o s e s y s t e m . The e r e c t a x e s o f C . a n t a r c t i c u m a r i s e f r o m t h e t i p s o f t h e l o b e d c r u s t and, as i n A . j a p o n i c u s , a r e d e s c r i b e d as b e i n g C h o r d a r i a - l i k e (e.g. . S k o t t s b e r g , 1921; K y l i n , 1933; L i n d a u e r , 1947), c o n s i s t i n g o f c y l i n d r i c a l , p e r c u r r e n t a x e s which become c l o t h e d i n l a t e r a l s . I t i s a t t h i s p o i n t t h a t t h e m o r p h o l o g i c a l s i m i l a r i t i e s between t h e s e two s p e c i e s 164 end as C . a n t a r c t i c u m p o s s e s s e s b l a d d e r s o r w a r t - l i k e p r o l i f e r a t i o n s which a l s o a r i s e out o f t h e c r u s t . C r u s t o s e System The c r u s t o f C a e p i d i u m has a p s e u d o p a r e n c h y m a t o u s c o n s t r u c t i o n a l t h o u g h i n c r o s s - s e c t i o n ( F i g u r e 34 and 3 5 a ) , t h e a x e s o f t h e c r u s t may a p p e a r t o be p a r e n c h y m a t o u s . However, t h e f i l a m e n t o u s c o n s t r u c t i o n o f t h e c r u s t c a n be c l e a r l y seen i n l o n g i t u d i n a l s e c t i o n s o f t h e s e a x e s ( F i g u r e s 35b and d ) . The f i l a m e n t s w i t h i n t h e m e d u l l a o f t h e c r u s t c o n s i s t o f l a r g e , e l o n g a t e d c e l l s w h ich a r e l o o s e l y p a c k e d . These f i l a m e n t s c u r v e o u t o b l i q u e l y t o t h e s u r f a c e l a y e r s o f t h e c r u s t . The c o r t i c a l l a y e r s (2-4 c e l l s deep on t h e u p p e r s u r f a c e and 1-2 c e l l s deep on t h e l o w e r s u r f a c e ) c o n s i s t o f c l o s e l y a r r a n g e d s m a l l c e l l s . As p r e s e n t e d i n C h a p t e r 1, A . j a p o n i c u s a l s o p o s s e s s e s a f i l a m e n t o u s c r u s t . I t d i f f e r s f r o m C . a n t a r c t i c u m i n t h e r e l a t i v e d i m e n s i o n s o f t h e c o r t i c a l and m e d u l l a r y c e l l s , as i n A . j a p o n i c u s t h e r e i s much l e s s d i f f e r e n c e i n s i z e between t h e s e c e l l t y p e s (compare F i g u r e s 4 and 3 5 b ) . . When i n c o n t a c t w i t h a n o t h e r p a r t of t h e c r u s t o r w i t h t h e s u b s t r a t e , t h e c o r t i c a l c e l l s o f t h e l o w e r s u r f a c e can form p e g - l i k e e x t e n s i o n s ( F i g u r e 35a) which d e v e l o p i n t o e i t h e r u n i - o r m u l t i c e l l u l a r r h i z o i d s . B h i z o i d s o f A . j a p o n i c u s and C . a n t a r c t i c u m a r e v e r y s i m i l a r i n d e v e l o p m e n t and a p p e a r a n c e . H a i r s were f o u n d i n C . a n t a r c t i c u m ( F i g u r e 35c) , and as 165 d e s c r i b e d by S k c t t s b e r g (1921), t h e s e a r e v e r y s i m i l a r t o t h o s e f o u n d i n A . j a p o n i c u s . I n b o t h s p e c i e s t h e y a r i s e f r o m a p i t below t h e c o r t e x and a r e b o r n e s i n g l y or i n twos, but do not a p p e a r i n c l u s t e r s o r i n c r y p t o s t o m a t a . The g r o w t h o f t h e c r u s t , as i n A . j a p o n i c u s , r e s u l t s f r c m d i v i s i o n s of c o r t i c a l c e l l s a t t h e m a r g i n s o f t h e c r u s t , p a r t i c u l a r l y a t t h e t i p o f each l o b e . E l o n g a t i o n and e x p a n s i o n of c e l l s i n t e r n a l t o t h e m e r i s t e m a t i c zone o c c u r s , and i n C . a n t a r c t i c u m , u n l i k e A . j a p o n i c u s , one c a n f i n d h o l l o w e d p o r t i o n s where m e d u l l a r y c e l l s have been p u l l e d a p a r t by t h e e x t e n s i o n o f t h e c r u s t ( F i g u r e 3 8 ) . I n t e r s p e r s e d w i t h t h e s m a l l and c l o s e l y p a c k e d c e l l s o f t h e c o r t e x a r e c e l l s o f v a r y i n g s i z e , f r e g u e n t l y v e r y l a r g e , which a r e d i s t i n g u i s h a b l e by t h e i r c o n t e n t s . The m a t e r i a l i n t h e s e c e l l s when u n s t a i n e d i s g o l d e n - b r o w n and has a b e a d - l i k e , a l m o s t g r a n u l a r , a p p e a r a n c e . When s t a i n e d w i t h TBO t h i s m a t e r i a l becomes b r i g h t g r e e n i n d i c a t i n g t h e p r e s e n c e o f p h e n o l i c s ( r e f e r t o C h a p t e r 1 f o r a d i s c u s s i o n o f the TBO s t a i n i n g r e a c t i o n ) . I t i s u n s t a i n e d by PAS. T h e s e c e l l s were n o t e d by S k o t t s b e r g (1907, 1921), F r i t s c h (1945) and L i n d a u e r e t a l . (1961) as o c c u r r i n g w i t h t h e p l u r i l o c u l a r s p o r a n g i a on t h e b l a d d e r s . As shown i n F i g u r e s 34, 35a, 35b, 37 and 38, t h e s e s t r u c t u r e s a r e f u l l y immersed i n t h e t i s s u e and t h e y o c c u r on t h e s o l i d c r u s t s as w e l l as the s t e r i l e b l a d d e r s ( F i g u r e s 36a and b ) . . I have d e s i g n a t e d t h e s e c e l l s t o be 'physode- c e l l s ' a l t h o u g h t h e y a r e n o t t h e o n l y c e l l s i n which p h e n o l i c m a t e r i a l s a r e 166 l o c a t e d i n C . a n t a r c t i c u m (as c a n be s e e n i n F i g u r e s 34, 35 and 3 6 ) . . W i t h i n t h e c r u s t o f A . j a p o n i c u s t h e r e a l s o i s a c o n s i d e r a b l e amount o f p h e n o l i c m a t e r i a l p r e s e n t , b u t t h e r e i s no c e l l t y p e e q u i v a l e n t t o t h e s e p h y s o d e - c e l l s . E r e c t A x i s The e r e c t a x e s o f C . a n t a r c t i c u m a r i s e from t h e l o b e - t i p s o f t h e c r u s t . An a x i s i s s u r r o u n d e d by c r u s t t i s s u e i n a c u p - l i k e d e p r e s s i o n ( F i g u r e 37 and 3 8 ) . In t h e zone o f t r a n s i t i o n between t h e c r u s t and t h e u p r i g h t a x i s , t h e m e d u l l a c e l l s become more c l o s e l y p a cked and n a r r o w e r as t h e y r i s e i n f i l a m e n t s o f e l o n g a t e , c y l i n d r i c a l c e l l s ( F i g u r e 37b and 3 8 ) . The c o r t e x o f t h e c r u s t g i v e s way t o a C h c r d a r i a - l i k e c o r t e x i n t h e d i p o f t h e d e p r e s s i o n ( a r r o w e d i n F i g u r e 37a and 38), c o n s i s t i n g o f f i l a m e n t s 2-4 c e l l s i n l e n g t h t e r m i n a t e d by a l a r g e r somewhat c l u b - s h a p e d c e l l ( F i g u r e 4 0 a ) . T h e s e t e r m i n a l c e l l s a r e f i l l e d w i t h g o l d e n - brown c o n t e n t s which s t a i n g r e e n w i t h TBO i n d i c a t i n g t h e i r p h e n o l i c c o m p o s i t i o n ( F i g u r e s 4 0 a - c ) . S c a t t e r e d i n t h e c o r t e x a r e c o l o u r l e s s h a i r s a r i s i n g f r o m t h e s u b - c o r t e x ( F i g u r e 39b and 4 0 c ) . . As i n A. • j a p o n i c u s , t h e s u b - c o r t e x o f C. a n t a r c t i c u m c a n be u n d e r s t o o d as c o n s i s t i n g o f 2-j-3 c e l l l a y e r s o f r o u n d e d c e l l s , a r i s i n g f r o m t h e m e d u l l a , and which g i v e r i s e t o t h e c o r t i c a l f i l a m e n t s and t h e u n i l o c u l a r s p o r a n g i a . The growth of t h e e r e c t a x i s f o l l o w s a p a t t e r n s i m i l a r to t h a t o f A . j a p o n i c n s w i t h m e r i s t e m a t i c a c t i v i t y l o c a t e d 167 a p i c a l l y a t t h e b a s e o f a ' f a n ' o f c o r t i c a l f i l a m e n t s ( F i g u r e 3 9 c ) ; As i n A . j a p o n i c u s , t h e growth o f C . a n t a r c t i c u m c o u l d be d e s c r i b e d as f i t t i n g the m u l t i p l e s t r a n d model o f P a r k e ( 1 9 3 3 ) . The a x i s o f C . a n t a r c t i c u m becomes h o l l o w e d a s t h e p l a n t grows ( F i g u r e 3 9 a ) , i n a s i m i l a r f a s h i o n t o t h a t d e s c r i b e d f o r A . j a p o n i c u s ( C h a p t e r 1 ) . I f o u n d s e c o n d a r y f i l a m e n t s i n t h e m e d u l l a of t h e e r e c t a x i s o f C . a n t a r c t i c u m , as i n A . j a p o n i c u s , but no i n d i c a t i o n of s e c o n d a r y s y s t e m s i n t h e c o r t e x or s u b c o r t e x . The e r e c t a x i s o f C . a n t a r c t i c u m p r o d u c e s t h i n l a t e r a l b r a n c h e s s i m i l a r t o t h o s e f o r m e d i n A. -japonicus. D i v i s i o n s o c c u r r i n g i n t h e s u b - c o r t e x r e s u l t i n t h e c o r t i c a l f i l a m e n t s b e i n g pushed o u t w a r d . The m e r i s t e m a t i c a c t i v i t y c o n t i n u e s a t t h e b a s e o f t h e a p i c a l c o r t i c a l ' f a n ' ( F i g u r e s 3 9 a - c ) . As w i t h t h e main a x i s , when t h e l a t e r a l c o n t i n u e s grow, the m e d u l l a i s s p l i t a p a r t . The e r e c t a x i s o f C . a n t a r c t i c u m p r o d u c e s u n i l o c u l a r s p o r a n g i a immersed i n t h e c o r t i c a l f i l a m e n t s . . T h e s p o r a n g i a a r e c u t o f f f r c m s u b - c o r t i c a l c e l l s ( F i g u r e s 39b, 40b and c ) . A l t h o u g h t h e anatomy of t h e e r e c t a x e s o f t h e s e two s p e c i e s i s v e r y s i m i l a r , t h e s p o r a n g i a d i f f e r i n s i z e . . The u n i l o c u l a r s p o r a n g i a i n A . j a p o n i c u s a r e u s u a l l y a t m a t u r i t y 60-90 pm x 25-40 ;im, and i n C . a n t a r c t i c u m a p p a r e n t l y mature s p o r a n g i a a r e 30-45 pm x 10-15 jam. The c o r t i c a l f i l a m e n t s o f A . j a p o n i c u s s u r r o u n d i n g t h e s p o r a n g i a a r e u s u a l l y 4-7 c e l l s i n l e n g t h , whereas i n C . a n t a r c t i c u m t h e s e s t e r i l e f i l a m e n t s a r e o n l y between 2 and 3 c e l l s l o n g . 168 A l t h o u g h t h e e r e c t a x e s o f C . a n t a r c t i c u m which I have s e e n f a l l w i t h i n t h e s i z e r a n g e o f A . j a p o n i c u s p l a n t s , t h e y a r e s m a l l e r t h a n most A . j a p o n i c u s p l a n t s , w i t h t h i n n e r l a t e r a l b r a n c h e s and main a x e s . B l a d d e r s The b l a d d e r s o f C . a n t a r c t i c u m , which i n i t i a l l y a r e s m a l l , w a r t - l i k e s t r u c t u r e s , become f o l d e d and c o n v o l u t e d b l a d d e r s a s t h e y e x t e n d . They a r i s e f r o m t h e s u r f a c e o f t h e c r u s t . When t h e b l a d d e r s a r e s e c t i o n e d t h e y can be s e e n c l e a r l y t o be p a r t o f t h e b a s a l s y s t e m o f C . a n t a r c t i c u m , and a s S k o t t s b e r g (1921) o b s e r v e d , t h e r e i s no d i f f e r e n c e i n t h e anatomy o f t h e c r u s t o r t h e b l a d d e r . The b l a d d e r becomes h o l l o w e d o u t a s i t d e v e l o p s , and t h e f i l a m e n t o u s c o n s t r u c t i o n o f the t h a l l u s i s o b s c u r e d as t h e c e l l f i l e s a r e b r o k e n . The b l a d d e r has a t i g h t l y p a c k e d c o r t e x j u s t as t h e c r u s t d o e s , and below t h i s a r e l a r g e r o u n d e d c e l l s w hich c l e a r l y show s i g n s o f b e i n g p u l l e d a p a r t w i t h t h e growth of t h e p l a n t ( F i g u r e s 36a, c and e ) . A l t h o u g h t h e b l a d d e r s have been i n t e r p r e t e d as b e i n g ' C o l p o m e n i a - l i k e ' i n c o n s t r u c t i o n (e.g. C o t t o n , 1915; L e v r i n g , 1945, 1960; L i n d a u e r , 1947), t h e i r d e v e l o p m e n t i s n o t p a r e n c h y m a t o u s . T h u s , t h e p l a c e m e n t o f t h i s s p e c i e s i n t h e P u n c t a r i a c e a e ( L e v r i n g , 1945, 1960) i s i n a p p r o p r i a t e . F r i t s c h (1945) c o r r e c t l y n o t e d t h e r e s e m b l a n c e o f t h e s e v e s i c l e s t o L e a t h e s i a and makes t h e f o l l o w i n g comment - " I n view o f t h e c l o s e r e l a t i o n s h i p between M e s o g l o i a c e a e and 169 L e a t h e s i a c e a e t h e r e i s n o t h i n g v e r y s u r p r i s i n g i n t h e e r e c t g r o w t h s from t h e b a s a l s t r a t u m r e p e a t i n g t h e s t r u c t u r a l c h a r a c t e r i s t i c s o f t h e two f a m i l i e s . " . The b l a d d e r s d i f f e r f r o m L e a t h e s i a i n n o t p o s s e s s i n g a f i l a m e n t o u s c o r t e x . They a r e a l s o l e s s m u c i l a g i n o u s , and t h e c e l l s more t i g h t l y bound t h a n i n L e a t h e s i a * The b l a d d e r s grow o u t of t h e c r u s t and can form r h i z o i d s t o a t t a c h t h e m s e l v e s t o o t h e r p a r t s o f t h e c r u s t and t o t h e s u b s t r a t e . The r h i z o i d s o f t h e b l a d d e r (as w e l l as t h o s e o f t h e c r u s t ) i s s u e f r o m t h e l o w e r c o r t e x and c a n be u n i - or m u l t i c e l l u l a r . The p l u r a n g i a which o c c u r on t h e b l a d d e r s a r e f o r m e d by t r a n s f o r m a t i o n o f t h e s u r f a c e c e l l s i n t o f i l a m e n t s w i t h u s u a l l y H-6 c e l l s p e r f i l e . The s u r f a c e o f t h e t h a l l u s becomes pushed up where t h e p l u r a n g i a d e v e l o p - r a i s e d by t h e d e v e l o p m e n t o f t h e s e f i l a m e n t s . The s u r f a c e o f t h e b l a d d e r p o s s e s s e s a c u t i c l e - l i k e l a y e r ( s t a i n i n g p u r p l e w i t h TBO) and t h i s becomes more p r o n o u n c e d o v e r t h e s c r i ( F i g u r e s 3 6 a - d ) . On r e l e a s e o f t h e swarmers, t h e c u t i c l e - l i k e l a y e r i s no l o n g e r p r e s e n t , and t h e empty c e l l w a l l s o f the b a s a l p l u r a n g i a ! r e m a i n ( F i g u r e s 36e and f) . The s y s t e m a t i c a f f i n i t i e s o f C . a n t a r c t i c u m and i t s r e l a t i o n s h i p t c A. j a p o n i c u s A l t h o u g h t h e s e two s p e c i e s s h a r e a p s e u d o p a r e n c h y m a t o u s c o n s t r u c t i o n and s i m i l a r m o r p h o l o g i e s , t h i s c o m p a r a t i v e s t u d y s u p p o r t s t h e s e p a r a t i o n o f t h e s e p l a n t s i n d i s t i n c t g e n e r a . I n p a r t i c u l a r , t h e p o s s e s s i o n o f 170 b l a d d e r s b e a r i n g p l u r a n g i a and t h e c h a r a c t e r i s t i c p h y s o d e - c e l l s a r e d i s t i n g u i s h i n g f e a t u r e s o f C . a n t a r c t i c u m . C . a n t a r c t i c u m has a l s o been r e p o r t e d t o p o s s e s s p y r e n o i d s ( A s e n s i , D e l e p i n e , and G u g l i e l m i , 1977), a f e a t u r e > w h i c h a c c o r d i n g t c some w o r k e r s h a s p h y l o g e n e t i c s i g n i f i c a n c e (Evans, 1966; H o r i , 1972; Magne, 1978) and which i s not s h a r e d by A. j a p o n i c u s ( H o r i , 1972) . As o u t l i n e d i n t h e i n t r o d u c t i o n t o t h i s c h a p t e r , S k o t t s b e r g (1921) and K y l i n (1933) p u t f o r w a r d d i f f e r e n t i n t e r p r e t a t i o n s o f t h e l i f e h i s t o r y o f C * a n t a r c t i c u m u s i n g t h e i n f o r m a t i o n a v a i l a b l e f r o m t h e morphology of t h e p l a n t . T h i s s t u d y c a n n o t r e s o l v e which o f t h e s e i n t e r p r e t a t i o n s i s c o r r e c t as no l i f e h i s t o r y s t u d i e s o f C . a n t a r c t i c u m were p o s s i b l e . I f , as s u g g e s t e d by S k o t t s b e r g ( 1 9 2 1 ) , t h e b l a d d e r - b e a r i n g and e r e c t a x i s - b e a r i n g c r u s t s a r e a l t e r n a t i n g g e n e r a t i o n s , t h e p l a c e m e n t o f C . a n t a r c t i c u m w i t h i n t h e C h o r d a r i a l e s c a n n o t be c o n t i n u e d . Such an a l t e r n a t i o n o f p a r t i a l l y h e t e r o m o r p h i c g e n e r a t i o n s w i t h a p s e u d o p a r e n c h y m a t o u s c o n s t r u c t i o n would make t h i s g e n us v e r y u n u s u a l i n t h e brown a l a g e . I t c o u l d be accommodated w i t h i n t h e E c t o c a r p a l e s i n a new f a m i l y . A l t h o u g h M u l l e r (1972) r e p o r t e d t h a t t h e g a m e t o p h y t e and s p o r o p h y t e g e n e r a t i o n s o f E c t o c a r p u s s i l i c u l o s u s c o u l d be m o r p h o l o g i c a l l y d i s t i n g u i s h e d , t h e d i f f e r e n c e s between t h e s e g e n e r a t i o n s were n o t n e a r l y as s u b s t a n t i a l as t h o s e between u p r i g h t - b e a r i n g and b l a d d e r - b e a r i n g c r u s t s . 171 I f K y l i n ' s (1933) s u g g e s t i o n i s c o r r e c t , w i t h C . a n t a r c t i c u m p o s s e s s i n g a m i c r o s c o p i c gametophyte s t a g e a l t e r n a t i n g w i t h d i p l o i d b l a d d e r s , e r e c t a x e s , and c r u s t s , t h i s s p e c i e s c a n be l e f t w i t h i n t h e C h o r d a r i a l e s . I t would seem, however, t h a t a s e p a r a t e f a m i l y would be w a r r a n t e d f o r t h i s genus on t h e b a s i s o f t h e e l a b o r a t e d c r u s t o s e s y s t e m and b l a d d e r s b e a r i n g p l u r a n g i a f u n c t i o n i n g i n a s e x u a l r e p r o d u c t i o n . B e f o r e t h e a f f i n i t i e s o f C . a n t a r c t i c u m c a n be f u l l y a p p r e c i a t e d , i n f o r m a t i o n on t h e l i f e h i s t o r y and c a r y o l o g i c a l s t a t e s o f t h e m o r p h o l o g i c a l p h a s e s i s n e e ded. A m i c r o s p e c t r o p h o t o m e t r i c a n a l y s i s of DNA c o n t e n t o f c r u s t s , b l a d d e r s and e r e c t a x e s , i s now b e i n g p u r s u e d t o f u r t h e r t h i s end. 0 V 172 F i g u r e 33. Type s p e c i m e n o f C a e p i d i u m a n t a r c t i c u m J . Ag.  174 F i g u r e 34. Montage o f a c r o s s - s e c t i o n o f c r u s t o s e t h a l l u s o f C . a n t a r c t i c u m . UC = upper c o r t e x ; LC = l o w e r c o r t e x ; M = m e d u l l a ; a r r o w s p o i n t t o ' p h y s o d e - c e l l s • . (200x) 1 7 5 176 F i g u r e 35a. C r o s s - s e c t i o n o f two l a y e r s o f C . a n t a r c t i c u m c r u s t . Arrows i n d i c a t e p e g - l i k e e x t e n s i o n s of l o w e r c o r t i c a l c e l l s d e v e l o p i n g i n t o r h i z o i d s . . (165x) F i g u r e 35b* L o n g i t u d i n a l s e c t i o n o f c r u s t o f C . a n t a r c t i c u r n < (125x) F i g u r e 35c. C r o s s - s e c t i o n a l view o f c r u s t o f C . a n t a r c t i c u m showing h a i r ( h ) . . (375x) F i g u r e 35d. M e d u l l a o f c r u s t o f C . a n t a r c t i c u m i n l o n g i t u d i n a l s e c t i o n . Arrow i n d i c a t e s d i r e c t i o n o f growth. (385x) 177 178 F i g u r e 36a. C r o s s - s e c t i o n o f s t e r i l e b l a d d e r of C . a n t a r c t i c u m . Arrow p o i n t s t o w a l l breakdown and h o l l o w c e n t r a l r e g i o n ( h e ) . (200x) F i g u r e 36b. S t e r i l e b l a d d e r i n c r o s s - s e c t i o n . Arrow p o i n t s t o c u t i c l e - l i k e m a t e r i a l on s u r f a c e o f p l a n t . . (675x) F i g u r e 36c. P l u r i l o c u l a r b l a d d e r . A r rows p o i n t t o w a l l breakdown i n m e d u l l a n e a r h o l l o w c e n t r a l r e g i o n . (200x) F i g u r e 36d. P l u r a n g i a (P) w i t h l a r g e • p h y s o d e - c e l l • . C u t i c l e - l i k e m a t e r i a l on t h a l l u s s u r f a c e a r r o w e d . . (675x) F i g u r e 36e. B l a d d e r w i t h p l u r a n g i a which have r e l e a s e d t h e i r c o n t e n t s . Arrow i n d i c a t e s w a l l breakdown i n m e d u l l a . (200x) F i g u r e 3 6 f . R e l e a s e d p l u r a n g i a (RP) i n C . a n t a r c t i c u m b l a d d e r * Note a b s e n c e o f c u t i c l e - l i k e l a y e r . Arrows p c i n t t o empty l o c u l e s o f p l u r a n g i a . . ( 6 7 5 x ) 179 180 F i g u r e 37* L o n g i t u d i n a l s e c t i o n t h r o u g h c r u s t a t b a s e o f an e r e c t a x i s . (a) s e c t i o n t h r o u g h c r u s t o s e p o r t i o n i n c u p - l i k e d e p r e s s i o n . Arrows p o i n t t o zone o f t r a n s i t i o n o f c o r t e x . (t) s e c t i o n t h r o u g h b a s e o f e r e c t a x i s and c r u s t d e p r e s s i o n . CM = c r u s t m e d u l l a ; EM = e r e c t a x i s m e d u l l a (190x) 181 182 F i g u r e 38. L i n e d r a w i n g o f a l o n g i t u d i n a l s e c t i o n a t p o i n t where e r e c t a x i s o f C . a n t a r c t i c u m grows out of c r u s t . Arrows (T) i n d i c a t e p o i n t o f t r a n s i t i o n i n t h e c o r t e x c f c r u s t . Arrow (EA) i n d i c a t e s d i r e c t i o n o f growth o f e r e c t a x i s . . (140x) 2<> 18a F i g u r e 39a. L o n g i t u d i n a l s e c t i o n o f u p r i g h t a x i s o f C . a n t a r c t i c u m s h o w i n g d e v e l o p i n g l a t e r a l b r a n c h (DL - c u r v e d arrow) . C e n t r e o f a x i s i s becoming h o l l o w (he) . S o l i d arrow p o i n t s t o e n l a r g e d c e l l lumen r e s u l t i n g f r c m c r o s s w a l l breakdown. . (175x) F i g u r e 3 9b. D e v e l o p i n g l a t e r a l o f C . a n t a r c t i c u m , showing h a i r s (h) and u n i l o c u l a r s p o r a n g i a ( u ) . Growth o c c u r s a t t h e base o f t h e c o r t i c a l f i l a m e n t s . (a60x) F i g u r e 3 9 c . . L o n g i t u d i n a l s e c t i o n t h r o u g h apex (A) and l a t e r a l (I) o f C . a n t a r c t i c u m . (190x)  186 F i g u r e 40a. L o n g i t u d i n a l s e c t i o n o f s t e r i l e e r e c t a x i s o f C . a n t a r c t i c u m . C = c o r t e x ; SC = s u b - c o r t e x ; W = m e d u l l a ; a r r o w s p o i n t t o c e l l s d e v e l o p i n g f r o m s u b - c o r t e x . (675x) F i g u r e 40b. L o n g i t u d i n a l s e c t i o n o f e r e c t a x i s o f C . a n t a r c t i c u m s h o w i n g u n i l o c u l a r s p o r a n g i a , ( a r r o w s t o u = u n i l o c u l a r s p o r a n g i a ) (675x) F i g u r e 4Cc. U n i l o c u l a r t i s s u e o f C . a n t a r c t i c u m . . H = h a i r s ; u = u n i l o c u l a r s p o r a n g i a . (675x) 187 An e v a l u a t i o n o f t h e R a l f s i a l e s , R a l f s i a c e a e , and t h e t a x o n o m i c p o s i t i o n o f A n a l i p u s j a p o n i c u s (Harv.) Wynne 189 lh§ R a l f s i a l e s The o r d e r - R a l f s i a l e s was e s t a b l i s h e d by Nakamura (1972) "on t h e b a s i s o f R a l f s i a e s p e c i a l l y b e c a u s e o f p e c u l i a r i t i e s i n t h e e a r l y d e v e l o p m e n t o f t h e t h a l l u s , and i n t h e s t r u c t u r e o f c e l l s a n d r e p r o d u c t i v e o r g a n s " * Nakamura gave t h e f o l l o w i n g c r i t e r i a a s d i s t i n g u i s h i n g f e a t u r e s o f t h i s o r d e r : - (1) a s i n g l e , p a r i e t a l , p l a t e - l i k e c h l o r o p l a s t w i t h o u t p y r e n o i d per c e l l , (2) d i s c a l - t y p e o f d e v e l o p m e n t c f t h e t h a l l u s , and, (3) an E c t o c a r p u s - t y p e o f l i f e c y c l e . He r e c o g n i s e d t h r e e f a m i l i e s i n t h e o r d e r s e p a r a t i n g them e n t i r e l y on t h e b a s i s o f t h e p l a c e m e n t o f r e p r o d u c t i v e o r g a n s . T h u s , i n Nakamura's a r r a n g e m e n t , i n t h e L i t h o d e r m a t a c e a e a r e p l a c e d p l a n t s w i t h t e r m i n a l l y b o r n e u n i l o c u l a r and p l u r i l o c u l a r o r g a n s , i n t h e Nemodermataceae a r e t h o s e w i t h i n t e r c a l a r y u n i l o c u l a r and l a t e r a l p l u r i l o c u l a r o r g a n s , and i n t h e R a l f s i a c e a e a r e t h e p l a n t s w i t h l a t e r a l u n i l o c u l a r and i n t e r c a l a r y p l u r i l o c u l a r o r g a n s , a l t h o u g h r e c o g n i s i n g t h e s e f a m i l i e s , Nakamura do e s s t a t e "... t h e p o s s i b i l i t y c a n n o t be e x c l u d e d t h a t t h e s e t h r e e f a m i l i e s s h o u l d be u n i t e d i n t o a s i n g l e f a m i l y R a 1 f s i a c e a e . " . I n d e s c r i b i n g t h e R a l f s i a l e s , Nakamura r e f e r s i n p a r t i c u l a r t o R a l f s i a v e r r u c o s a and A n a l i p u s j a p o n i c u s ( H e t e r p e n o r d a r i a a b i e t i n a ) . P r i o r t o Nakamura's t r e a t m e n t , A . j a p o n i c u s had been p l a c e d i n t h e C h o r d a r i a c e a e and 190 C h o r d a r i a l e s (Wynne, 1971) (and p r e v i o u s l y as H e t e r p c h o r d a r i a a b i e t i n a i n t h e H e t e r o c h o r d a r i a c e a e , C h o r d a r i a l e s ) . Nakamura m a i n t a i n s t h a t t h e genus A n a l i p u s s h o u l d be i n c l u d e d i n t h e E a l f s i a c e a e b e c a u s e o f i t s d i s c a l - t y p e d e v e l o p m e n t and anatomy, p a r t i c u l a r l y o f i t s r e p r o d u c t i v e o r g a n s * T h i s t a x o n o m i c judgement was s u p p o r t e d by Wynne (1972) who c o n s i d e r e d t h a t t h e a f f i n i t y between R a l f s i a and A n a l i p u s s h o u l d be r e c o g n i s e d by t h e s e g e n e r a b e i n g c l a s s i f i e d i n t h e same f a m i l y . A l t h o u g h Wynne and L o i s e a u x (1976) n o t e t h a t "many p r o b l e m s r e m a i n t o be r e s o l v e d b e f o r e t h i s o r d e r c a n be m a i n t a i n e d s a f e l y " , t h e o r d e r R a l f s i a l e s as d e s c r i b e d by Nakamura (1972) h a s been a c c e p t e d by B o l d and Wynne (1978) i n t h e i r r e c e n t a l g a l t e x t b o o k , and i n a s l i g h t l y m o d i f i e d f o r m by Tanaka and C h i h a r a ( 1 9 8 0 ) . The o r d e r has n o t been used by a number o f w o r k e r s ( e . g . J o h n and Lawson, 1974; R u s s e l l and F l e t c h e r , 1975; A b b o t t and H o l l e n b e r g , 1976; P e d e r s e n , 1976; R u e n e s s , 1977) and t h e i r t r e a t m e n t s o f c r u s t o s e brown a l g a e h a v e v a r i e d . T h e r e a r e f u n d a m e n t a l o b j e c t i o n s t o t h i s o r d e r e s t a b l i s h e d by Nakamura as c a n be r e a l i s e d when one ex a m i n e s t h e t h r e e d i s t i n g u i s h i n g c r i t e r i a o f t h e o r d e r : - (1)a s i n g l e , p a r i e t a l , p l a t e - l i k e c h i o r o p l a s t , w i t h o u t p y r e n p i d , p e r c e l l I f t h e c r i t e r i o n o f one c h l o r o p l a s t per c e l l i s e n f o r c e d , a number o f c r u s t o s e brown a l g a e , c l a s s i f i e d i n 191 e i t h e r t h e R a l f s i a c e a e o r L i t h o d e r m a t a c e a e a r e e x c l u d e d f r o m t h e o r d e r e . g . . B a s i s p o r a a f r i c a n a , B . . s a x i g e n a (John and Lawson, 1974); D i p l u r a s i a u l a n s , E n d o p l u r a a u r e a ( H o l l e n b e r g , 1969); L i t h o d e r m a f a t i s c e n s (Waern, 1949); P s e u d o l i t h o d e r ma p a r a d o x u s ( S e a r s and K i l c e , 1973) P. , r o s c o f f e n s i s ( L o i s e a u x , 1968) , P. r o s e n v i n q j i (Lund, 1959, as d e s c r i b e d i n Waern, 1949). P y r e n o i d s have a l s o been r e p o r t e d i n a number o f t h e s e c r u s t o s e p l a n t s e.g.. R a l f s i a b o r n e t i and R. c l a y a t a ( E d e l s t e i n , Chen and M c L a c h l a n , 1970), R. c l a y a t a ( L c i s e a u x , 1968) , R. v e r r u c o s a ( F l e t c h e r , 1978) , R a l f s i a and P e t r o d e r m a ( R u s s e l l and F l e t c h e r , 1975); S p r a p i o n k j e l l m a n i i , Symphypcarpus l o n g i s e t u s , S. s t r a n g u l a n s , (Lund, 1959). ( A l t h o u g h Tanaka and C h i h a r a (1980) a c c e p t t h e o r d e r R a l f s i a l e s , t h e y do n o t c o n s i d e r c h l o r o p l a s t f e a t u r e s - e . g . number, shape o r a b s e n c e o f p y r e n o i d s - t o be one o f t h e c r i t e r i a f o r membership i n t h i s o r d e r . ) ( 2 ) d i s c a l - t y p e c f d e v e l o p m e n t Nakamura (1972) d i s c u s s e d t h e e a r l y d e v e l o p m e n t o f brown a l g a l t h a l l i and d i s t i n g u i s h e d t h r e e d e v e l o p m e n t a l p a t t e r n s : - (a) c r e e p i n g - t y p e ( e x h i b i t e d by members o f t h e E c t o c a r p a c e a e and M y r i o n e m a t a c e a e ) , (b) e r e c t - t y p e (as f o u n d , f o r example, i n t h e D i c t y o t a l e s and F u c a l e s ) , and (c) d i s c a l - t y p e . In d e s c r i b i n g t h e d i s c a l t y p e , Nakamura us e d as examples R a l f s i a v e r r u c o s a and A n a l i p u s - j a p o n i c u s . He s t a t e d " T h i s d i s c a l - t y p e o f d e v e l o p m e n t has been d e s c r i b e d by Kuckuck (1897) i n Nemoderma t i n q i t a n a , by 192 K y l i n (1934) and L o i s e a u x (1968) i n R a l f s i a c l a v a t a and by Sauvageau (1907) i n members o f t h e S p h a c e l a r i a l e s . " . I n f a c t , Kuckuck (1897) as c i t e d by Nakamura, does n o t r e f e r t o Nemoderma, a l t h o u g h Kuckuck (1912) does d e s c r i b e and f i g u r e a d i s c a l m o r p h o g e n e s i s f o r Nemoderma S c h o u s b e . .While K y l i n (1934) shows d i s c a l d e v e l o p m e n t f o r s p o r e s f r o m u n i l o c u l a r s p o r a n g i a o f R a l f s i a c l a v a t a , t h e r e i s a c o n s i d e r a b l e amount o f work c o n t r a d i c t i n g t h i s . L o i s e a u x (1 968), E d e l s t e i n e t a l . . (1970) , and F l e t c h e r (1978) have d e s c r i b e d and i l l u s t r a t e d t h e u n i s p o r e s o f R. c l a v a t a d e v e l o p i n g i n i t i a l l y i n t o a u n i s e r i a t e f i l a m e n t and l a t e r f o r m i n g a d i s c s t r u c t u r e . I t would a p p e a r t h a t t h e non- d i s c o i d a l e a r l y d e v e l o p m e n t o f R. c l a v a t a u n i s p o r e s i s w e l l documented, and t h a t t h e m a t e r i a l c u l t u r e d by K y l i n was u n l i k e l y t o have been t h e same e n t i t y . O t h e r d e v e l o p m e n t a l s t u d i e s on c r u s t o s e brown a l g a e have shown f i l a m e n t o u s d e v e l o p m e n t ( e . g . . P e t r o d e r m a M c u l i f o r m e , (Wynne, 1969; W i l c e , Webber and S e a r s , 1970), P s e u d o l i t h d e r m a n i g r a ( H o l l e n b e r g , 1969), P..paradoxum ( S e a r s and W i l c e , 1973), P. r o s c o f f e n s i s ( L o i s e a u x , 1 968; F l e t c h e r , 1 9 7 8 ) ; R a l f s i a b o r n e t i ( E d e l s t e i n e t a l . , 1970), !• c o n f u s a , R . . i n t e q r a ( H o l l e n b e r g , 1 9 6 9 ) ) , and f e w e r w i t h d i s c o i d a l d e v e l o p m e n t ( e . g . . R a l f s i a p a c i f i c a (Wynne, 1972), R . . v e r r u c o s a ( L o i s e a u x , 1968; E d e l s t e i n , Chen and M c L a c h l a n , 1971: F l e t c h e r , 1978); A n a l i p u s j a p o n i c u s (Wynne, 1972; C h a p t e r 1 ( i n t h e c a s e o f t h e l a t t e r two s p e c i e s , c o n f i r m i n g t h e work of Nakamura, 1972).. A l t h o u g h 193 t h e p a t t e r n e x h i b i t e d by a s p e c i e s seems t o be a c o n s t a n t f e a t u r e , a t l e a s t w i t h i n one genus, namely B a l f s i a on which t h i s o r d e r i s b a s e d , b o t h t y p e s o f g e r m i n a t i o n - d e v e l o p m e n t p a t t e r n s o c c u r . T h i s may be a r e f l e c t i o n of t h e s t a t e o f R a l f s i a taxonomy r a t h e r t h a n o f t h e v a l u e o f t h i s c h a r a c t e r a t t h e g e n e r i c l e v e l . I n my o p i n i o n , h o w e v e r , t h e d e v e l o p m e n t a l p a t t e r n o r t y p e , as o u t l i n e d by Nakamura, does n o t c o n s t i t u t e a good o r d i n a l c r i t e r i o n f o r t h i s g r o u p of c r u s t o s e brown a l g a e . ( 3 ) E c t o c a r p u s ^ t y p e l i f e c y c l e Nakamura was r e f e r r i n g h e r e t o t h e c y c l e c l a s s i c a l l y u n d e r s t o o d as t h e E c t p e a r p u s - t y p e w i t h a a l t e r n a t i o n o f i s c m o r p h i c g e n e r a t i o n s . In t h e i r r e v i e w o f l i f e h i s t o r y s t u d i e s i n t h e P h a e o p h y t a , Wynne and L o i s e a u x (1976) n o t e t h a t t h e c l a s s i c a l i n t e r p r e t a t i o n of E c t o c a r p a l e a n l i f e h i s t o r i e s h a s been s u b j e c t e d t o m o d i f i c a t i o n s o v e r t h e p a s t t w e n t y y e a r s . The work o f M i i l l e r (1967, 1972) on E c t o c a r p u s s i l i c u l o s u s r e v e a l s a complex l i f e h i s t o r y which i n v o l v e s d i f f e r e n t g r o w t h f o r m s f o r t h e g a m e t o p h y t e and s p o r o p h y t e g e n e r a t i o n s a s w e l l as a number o f p o s s i b l e c y t o l o g i c a l p athways. I t would a p p e a r t h a t t h e d e s c r i p t i o n o f a l i f e h i s t o r y p a t t e r n by t h e term ' E c t o c a r p u s - t y p e 1 i s n o t p a r t i c u l a r l y c l e a r , g i v e n t h e v a r i a t i o n now known t o o c c u r i n t h i s genus. A l t h o u g h l i f e h i s t o r y s t u d i e s have been c a r r i e d o u t on a number o f members o f t h i s g r o u p o f brown a l g a e ( e . g . L o i s e a u x , 1968; Wynne, 1969, 1972; E d e l s t e i n e t a l . , 194 1970,1971; F l e t c h e r , 1974, 1978), o n l y " d i r e c t 1 o r a p o m e i o t i c l i f e h i s t o r i e s ( e . g . P e t r o d e r m a m a c u l i f o r m e , Wynne, 1972), o r l i f e h i s t o r i e s i n v o l v i n g t h e c r u s t s i n t h e l i f e h i s t o r i e s o f members o f t h e S c y t o s i p h o n a l e s ( e . g . E a I f g i g c l a v a t a and B. . b o r n e t i , E d e l s t e i n e t a l . , 1970) have been r e p o r t e d , and no c o m p l e t e d l i f e h i s t o r y w i t h e v i d e n c e o f s e x u a l i t y has been shown. Kuckuck (19 12) g i v e s a c o n v i n c i n g d e s c r i p t i o n o f anisogamy i n Nemoderma t i n g i t a n a and t h e f u s i o n o f s e t t l e d f e m a l e and swimming male gametes. A r e p r o d u c t i v e l y mature p l a n t was n o t grown from t h e r e s u l t i n g z y g o t e s , however, and no c y t o l o g i c a l e v i d e n c e was g i v e n . I t would a p p e a r t h a t t h r o u g h o u t t h i s g r o u p t h e r e has o c c u r r e d a r e d u c t i o n i n l i f e h i s t o r y s t a g e s and p o s s i b l y i n some, a l o s s o f s e x u a l i t y . The l i f e h i s t o r i e s o f t h e s e o r g a n i s m s need f u r t h e r work, however, i n b o t h c u l t u r e and, p a r t i c u l a r l y , i n c y t o l o g i c a l s t u d i e s , b e f o r e u s e f u l g e n e r a l i s a t i o n s c a n be made a b o u t them.. I t i s my o p i n i o n t h a t t h e R a l f s i a l e s c a n n o t be r e c o g n i s e d f o r t h e r e a s o n s o u t l i n e d above; t o c o n t i n u e t o use t h i s t a x o n w i l l o n l y f u r t h e r c o n f u s e t h e a l r e a d y t r o u b l e d s t a t e o f t h e taxonomy o f t h e s o - c a l l e d ' l o w e r ' brown a l g a e . 195 The R a l f s i a c e a e As a l r e a d y m e n t i o n e d , Nakamura (1972) r e c o g n i s e d t h r e e f a m i l i e s i n h i s t r e a t m e n t o f t h e c r u s t o s e brown a l g a e , and t h e s e f a m i l i e s were s e p a r a t e d by him e n t i r e l y on t h e b a s i s o f t h e p l a c e m e n t o f t h e r e p r o d u c t i v e s t r u c t u r e s . T h i s t r e a t m e n t was a l s o f o l l o w e d by B o l d and Wynne (19 7 8 ) . A l t h o u g h b o t h H c l l e n b e r g (1969) and F l e t c h e r (1 978) make p a s s i n g r e f e r e n c e t o c h l o r o p l a s t numbers b e i n g used g e n e r a l l y a s a c h a r a c t e r t o d i s t i n g u i s h t h e R a l f s i a c e a e and t h e L i t h o d e r m a t a c e a e , i n f a c t i n t h o s e works where f a m i l y d e s c r i p t i o n s a r e p r o v i d e d ( e . g . T a y l o r , 1957, 1960) c h l o r o p l a s t c h a r a c t e r s a r e n o t used b u t r a t h e r t h e d e s c r i p t i o n s a r e b a s e d on t h e p l a c e m e n t o f r e p r o d u c t i v e s t r u c t u r e s - i n k e e p i n g w i t h t h e o r i g i n a l d e s c r i p t i o n s o f t h e s e f a m i l i e s . Tanaka and C h i h a r a (1980) c o r r e c t l y o b s e r v e t h a t c h l o r o p l a s t number c a n n o t be used as a c r i t e r i o n t o d i s t i n g u i s h t h e s e f a m i l i e s . Feldmann (1937) e s t a b l i s h e d t h e m o n o t y p i c f a m i l y , t h e Nemodermataceae, f o r Nemoderma t i n q i t a n a S c h o u s b e . Members o f t h i s genus p o s s e s s i n t e r c a l a r y u n i l o c u l a r s p o r a n g i a , and l a t e r a l p l u r i l o c u l a r o r g a n s w h i c h a r e c l e a r l y d i s t i n g u i s h a b l e by s i z e as o o g o n i a l or a n t h e r i d i a l i n n a t u r e , and p r o d u c e a n i s o g a m e t e s . Feldmann c o n s i d e r e d t h a t t h e s e r e p r o d u c t i v e s t r u c t u r e s were s u f f i c i e n t l y d i s t i n c t f r c m t h o s e f o u n d i n t h e R a l f s i a c e a e (which he used i n i t s w i d e r s e n s e t o i n c l u d e L i t h o d e r m a t a c e a e ) t o w a r r a n t 196 f a m i l i a l s t a t u s . Nemoderma i s r e s t r i c t e d i n i t s d i s t r i b u t i o n t c t h e C a n a r y I s l a n d s , M o r o c c o , A l g e r i a , and T u n i s i a , , and f o r t h i s r e a s o n i s o f t e n n o t d i s c u s s e d o r d e a l t w i t h i n d e s c r i p t i o n s o f c r u s t o s e brown a l g a e . P a p e n f u s s (1951, 1955) i n c l u d e s Nemoderma i n t h e R a l f s i a c e a e i n h i s t a x o n o m i c t r e a t m e n t s o f t h e P h a e o p h y t a . F l e t c h e r (1978) i n s p e c u l a t i n g on t h e p h y l o g e n y o f c r u s t s m e n t i o n s Nemoderma a s a p o s s i b l e example o f an a d v a n c e d p l a n t i n an e v o l u t i o n a r y l i n e o f s o f t c r u s t s s t a r t i n g f r o m a P e t r o d e r m a - l i k e a n c e s t o r . As t h e o n l y c r u s t o s e brown a l g a w i t h c l e a r l y o o g o n i a l and a n t h e r i d i a l p l u r i l o c u l a r s t r u c t u r e s which a r e l a t e r a l l y b o r n e and a l s o the o n l y one w i t h i n t e r c a l a r y u n i l o c u l a r o r g a n s , Nemoderma i s v e r y d i s t i n c t i v e . I n my o p i n i o n , t h e c o n t i n u e d use o f t h i s f a m i l y i s j u s t i f i e d . The R a l f s i a c e a e was d e s c r i b e d by Hauck (1885) and was a l s o u s e d , but i n a somewhat d i f f e r e n t s e n s e , by K j e l l m a n (1891-93).. The L i t h o d e r m a t a c e a e was f i r s t s u g g e s t e d by K j e l l m a n ( 1 8 8 3 ) , d e s c r i b e d by Hauck (1885), and a g a i n by K j e l l m a n ( 1 891-93).. As u n d e r s t o o d by t h e s e two w o r k e r s , t h e p r i m a r y d i f f e r e n c e between t h e members o f t h e s e two f a m i l i e s was t h e p o s i t i o n i n which s p o r a n g i a were p r o d u c e d . T h u s , t h e L i t h o d e r m a t a c e a e were s e e n as b e i n g d i s t i n c t i v e b e c a u s e t h e u n i l o c u l a r s p o r a n g i a were f o r m e d by t r a n s f o r m a t i o n o f s u r f a c e c e l l s r a t h e r t h a n b e i n g f o u n d immersed among s t e r i l e c o r t i c a l f i l a m e n t s , a s i n t h e R a l f s i a c e a e * The p l u r i l o c u l a r o r g a n s i n t h e 197 L i t h o d e r m a t a c e a e were d e s c r i b e d by K j e l l m a n (1883) as • a r r a n g e d i n s p e c i f i c s t a n d s i s s u i n g f r o m t h e s u r f a c e - c e l l s o f t h e f r o n d ' , and by K j e l l m a n (1891-93) as b e i n g l a t e r a l o u t g r o w t h s from s u r f a c e c e l l s . In t h e R a l f s i a c e a e , t h e p l u r i l o c u l a r s p o r a n g i a were d e s c r i b e d as b e i n g formed by t h e t r a n s f o r m a t i o n o f t h e c o r t i c a l f i l a m e n t s (Hauck, 1885). When t h e s e f a m i l i e s a r e r e c o g n i s e d as s e p a r a t e e n t i t i e s t o d a y , t h e u n i l o c u l a r s p o r a n g i a a r e u n d e r s t o o d as f i r s t r e p r e s e n t e d by Hauck and K j e l l m a n , a l t h o u g h t h e u n d e r s t a n d i n g c f t h e p l u r i l o c u l a r o r g a n s i s somewhat d i f f e r e n t , w i t h t h o s e i n t h e R a l f s i a c e a e b e i n g d e s c r i b e d as i n t e r c a l a r y and t h o s e i n t h e L i t h o d e r m a t a c e a e a s t e r m i n a l i n o r i g i n ( e .g. B o l d and Wynne, 1978). I n t h e l i t e r a t u r e t h e r e has been c o n s i d e r a b l e v a r i a t i o n i n t h e t r e a t m e n t o f c r u s t o s e P h a e o p h y t a , b o t h i n t e r m s o f how many f a m i l i e s a r e r e c o g n i s e d and, a t a h i g h e r t a x c n o m i c l e v e l , w i t h which o r d e r t h e y a r e t o be p l a c e d . Some w o r k e r s do n o t c o n s i d e r t h e L i t h o d e r m a t a c e a e t o be a d i s t i n c t f a m i l y r e c o g n i s i n g o n l y t h e R a l f s i a c e a e ( e . g . S e t c h e l l and G a r d n e r , 1925; Feldmann, 1937; P a p e n f u s s , 1951, 1955; Lund, 1959; H o l l e n b e r g , 1969; John and Lawson, 1974; R u s s e l l and F l e t c h e r , 1975; L i n d s t r o m , 1977; E u e n e s s , 1977). O t h e r w o r k e r s r e c o g n i s e b o t h f a m i l i e s ( e . g . T a y l o r , 1957, 1960; W i l c e , 1959; B o l d and Wynne, 1978; T a n a k a and C h i h a r a , 1980) , and r a r e l y , o n l y t h e L i t h o d e r m a t a c e a e i s r e c o g n i s e d ( e . g . T o k i d a , 1954).A number o f workers ( e . g . S e t c h e l l and G a r d n e r , 1925; R u s s e l l and F l e t c h e r , 1975; 198 F l e t c h e r , 1978) have d i s c u s s e d t h e s i m i l a r i t i e s between t h e M y r i o n e m a t c e a e and t h e R a l f s i a c e a e / L i t h o d e r m a t a c e a e (Nemodermataceae i s sometimes a l s o mentioned) and t h e r e a r e t h o s e who have c a r r i e d t h i s u n d e r s t a n d i n g f u r t h e r and c l a s s i f i e d t h e s e p l a n t s t o g e t h e r i n one f a m i l y , t h e M y r i o n e m a t a c e a e ( e . g . . S k o t t s b e r g , 1921; B o r g e s e n , 1926; Hamel, 1939; L o i s e a u x , 1967a and b, 1 9 6 8 ) . . Sometimes t h e M y r i o n e m a t a c e a e and t h e o t h e r f a m i l i e s of c r u s t o s e brown a l g a e a r e a l l c l a s s i f i e d i n t h e same o r d e r , t h i s b e i n g e i t h e r t h e E c t o c a r p a l e s ( e . g . S e t c h e l l and G a r d n e r , 1925; Feldmann, 1937; R u s s e l l and F l e t c h e r , 1975; Kornmann and S a h l i n g , 1977) or t h e C h o r d a r i a l e s ( e . g . . K y l i n , 1947; L i n d a u e r , Chapman and A i k e n , 1961; T a y l o r , 1957, 1960; A b b o t t and H o l l e n b e r g , 1976; L i n d s t r o m , 1977). O t h e r w o r k e r s , however, have p l a c e d t h e M y r i o n e m a t a c e a e i n t h e C h o r d a r i a l e s and t h e R a l f s i a c e a e / L i t h o d e r m a t a c e a e i n t o t h e E c t o c a r p a l e s ( e . g . . K y l i n , 1933; P a p e n f u s s , 1951, 1955), s e p a r a t i n g a t an o r d i n a l l e v e l what o t h e r w o r k e r s c o n s i d e r t o be v e r y c l o s e l y r e l a t e d p l a n t s . P e d e r s e n (1976) w h i l e r e c o g n i s i n g b o t h t h e M y r i o n e m a t a c e a e and the-. L i t h o d e r m a t a c e a e i n t h e E c t o c a r p a l e s , has p l a c e d t h e genus R a l f s i a i n t o t h e S c y t o s i p h o n a l e s - n o t o n l y t h o s e s p e c i e s °f R a l f s i a w hich have been shown t o be i n v o l v e d i n t h e l i f e h i s t o r i e s of members o f t h e S c y t o s i p h o n a l e s b u t a l s o t h o s e s p e c i e s which i n c u l t u r e s t u d i e s have shown no c o n n e c t i o n as y e t t o s c y t o s i p h o n a l e a n l i f e h i s t o r i e s . The c r u s t s v a r y a n a t o m i c a l l y , i n p a r t i c u l a r , i n t h e 199 o r i g i n , d e v e l o p m e n t , and d e g r e e o f c o h e s i o n of t h e e r e c t f i l a m e n t s e.g. t i g h t l y packed a s s u r g e n t and downwardly c u r v i n g c e l l rows from a c e n t r a l l a y e r i n R a l f s i a f u n g i f o r m i s c f . l o o s e l y a s s o c i a t e d e r e c t f i l a m e n t s a r i s i n g f r o m 1 - s e v e r a l l a y e r s o f p r o s t r a t e f i l a m e n t s i n B a s i s p o r a a f r i c a n a . O t h e r v e g e t a t i v e f e a t u r e s s u c h a s a c u t i c l e l a y e r , h a i r s , and r h i z o i d s , may be a b s e n t o r p r e s e n t * As n o t e d e a r l i e r , g e r m i n a t i o n and e a r l y d e v e l o p m e n t i n t h e s e p l a n t s t a k e s p l a c e i n a t l e a s t two d i f f e r e n t ways - e i t h e r i n a d i s c o i d a l f a s h i o n e.g. R a l f s i a p a c i f i c a , fi. v e r r u c o s a , o r f i l a m e n t o u s l y e.g. P e t r o d e r m a m a c u l i f o r m e , P s e u d p l i t h o d e r m a n i g r a . C h l o r o p l a s t c h a r a c t e r s s u c h as number, shape and t h e p r e s e n c e o r a b s e n c e o f p y r e n o i d s v a r y c o n s i d e r a b l y amongst t h e s e p l a n t s . The c l a s s i c a l c h a r a c t e r used t o d i f f e r e n t i a t e t h e s e p l a n t s - namely, t h e p l a c e m e n t o f r e p r o d u c t i v e s t r u c t u r e s - c a u s e s some p r o b l e m s e*g. R a l f s i a l u c i d a p o s s e s s e s u n i l o c u l a r s p o r a n g i a w hich may be i n a t e r m i n a l , l a t e r a l , or i n t e r c a l a r y p o s i t i o n (Lund, 1967); E d e l s t e i n , Chen and M c l a c h l a n (1968) n o t e t h a t i n t h e e a r l y s t a g e s of d e v e l o p m e n t o f R a l f s i a f u n g i f o r m i s t h e u n i l o c u l a r s p o r a n g i a a r e t e r m i n a l on e r e c t f i l a m e n t s and i t i s o n l y once a p a r a p h y s i s d e v e l o p s on t h e same f i l a m e n t t h a t t h e u n i l o c u l a r o r g a n can be s a i d t o be l a t e r a l l y p o s i t i o n e d ; F l e t c h e r (1978) r e c o r d s t h e p r o d u c t i o n o f t e r m i n a l u n i l o c u l a r s p o r a n g i a which a r e u n a c c o m p a n i e d by p a r a p h y s e s i n c u l t u r e s o f R. s p o n q i o c a r p a ; L i t h o d e r m a f a t i s c e n g 200 A r e s c h . emend. Waern (the t y p e s p e c i e s o f a f a m i l y now u n d e r s t o o d t o h a v e t e r m i n a l p l u r i l o c u l a r s t r u c t u r e s ) p o s s e s s e s l a t e r a l l y i n s e r t e d p l u r i l o c u l a r s p o r a n g i a (Waern, 1949) . As p r e v i o u s l y n o t e d , t h e l i f e h i s t o r i e s o f R a l f s i a c e a e / L i t h o d e r m a t a c e a e which have been examined t o d a t e show two t a s i c p a t t e r n s - e i t h e r a d i r e c t t y p e w i t h o u t e v i d e n c e o f s e x u a l i t y , o r one w h i c h shows t h e c r u s t s t o be s t a g e s i n t h e l i f e h i s t o r y o f p l a n t s i n t h e S c y t o s i p h o n a l e s . A number of w o r k e r s c o n s i d e r t h a t t h e R a l f s i a c r u s t s i n v o l v e d i n s c y t o s i p h o n a l e a n l i f e h i s t o r i e s b e l o n g to t h e s u b g e n u s S t r a g u l a r i a (e.g. Wynne, 1969; E d e l s t e i n e t a l . , 1970; J R o e l e v e l d , D u i s t e r h o f and Vroman, 1974). S t r a g u l a r i a was f i r s t d e s c r i b e d as a genus by S t r o m f e l t (1886) and l a t e r g i v e n s u b - g e n e r i c r a n k w i t h E u r a l f s i a by B a t t e r s (1890). S t r a g u l a r i a was d i s t i n g u i s h e d f r c m E u r a l f s i a t y h a v i n g a l o o s e r v e g e t a t i v e s t r u c t u r e and d i f f u s e s p o r a n g i a o r i n t h e words o f B a t t e r s (1890) "The s p e c i e s b e l o n g i n g t o t h i s s u b genus r e s e m b l e M yrionemata i n f r u c t i f i c a t i o n and L i t h o d e r m a t a i n s t r u c t u r e . . Wynne (1969) v o i c e s t h e o p i n i o n t h a t S t r a g u l a r i a i s g u i t e d i s t i n c t f r o m R a l f s i a s e n s u s t r i c t o and much o f t h e c u l t u r e e v i d e n c e a v a i l a b l e s u p p o r t s t h i s v i e w . . T h e r e a r e , however, r e p o r t s which show S t r a g u l a r i a - t y p e R a l f s i a s p e c i e s t o have d i r e c t l i f e h i s t o r i e s i n c u l t u r e e.g. R. c l a y a t a ( L o i s e a u x ; 1968), R. c o n f u s a ( H o l l e n b e r g , 1969), R. s p o n g i o c a r p a ( F l e t c h e r , 1S78) . As Wynne (1969) n o t e s , e v e n S t r a g u l a r i a 201 may i n f a c t be a h e t e r o g e n e o u s c o l l e c t i o n o f s p e c i e s and w a r r a n t f u r t h e r s u b d i v i s i o n . F l e t c h e r (1978) c o n s i d e r s t h a t a s u b d i v i s i o n o f t h e E u r a l f s i a s p e c i e s may be j u s t i f i e d . The c o m p l e x i t y o f t h e s i t u a t i o n i s f u r t h e r shown by c u l t u r e s t u d i e s of members of t h e S c y t o s i p h o n a l e s w hich showed no c r u s t o s e phase i n t h e i r l i f e h i s t o r i e s (e.g. F l e t c h e r , 1974; S auvageau, 1927), and s t u d i e s o f o t h e r members of t h e S c y t o s i p h o n a l e s which showed t h e i n v o l v e m e n t o f m y r i o n e m a t o i d s t a g e s i n t h e i r l i f e h i s t o r i e s ( e . g . L o i s e a u x , 1970; M c L a c h l a n , Chen and E d e l s t e i n , 1971b; F l e t c h e r , 1978). I t can be s e e n from t h e a f o r e g o i n g t h a t t h e r e i s c u r r e n t l y no c o n s e n s u s on t h e t a x o n o m i c a f f i n i t i e s e i t h e r w i t h i n t h e c r u s t o s e brown a l g a e , o r between t h e s e p l a n t s and o t h e r members of t h e P h a e o p h y t a . A l t h o u g h t h e s e c r u s t s s h a r e a s i m i l a r h a b i t and a p s e u d o p a r e n c h y m a t c u s c o n s t r u c t i o n , t h e many d i f f e r e n c e s which e x i s t s u g g e s t t h a t t h e y do n o t f o r m a n a t u r a l a s s e m b l a g e . At p r e s e n t , however, t h e r e i s no c l e a r g r o u p i n g o f c h a r a c t e r s which a l l o w s f o r t a x o n o m i c s e g r e g a t i o n above t h e l e v e l o f genus. Even a t t h e g e n e r i c l e v e l t h e r e have been some l o n g s t a n d i n g and f r e q u e n t l y d i s c u s s e d p r o b l e m s i n t h e l i t e r a t u r e , s u c h a s , f o r e x a mple, t h e c i r c u m s c r i p t i o n o f t h e g e n e r a L i t h p d e r m a ( e . g . . L u n d , 1938; Waern, 1949) and R a l f s i a . . (In t h e c a s e o f t h e l a t t e r t h i s s t i l l i s u n r e s o l v e d . ) N e i t h e r t h e d i v i s i o n o f t h e c r u s t o s e brown a l g a e i n t o t h e two f a m i l i e s R a l f s i a c e a e and L i t h o d e r m a t a c e a e , n o r t h e g r o u p i n g o f t h e s e 202 p l a n t s w i t h t h e M y r i o n e m a t a c e a e , i n my o p i n i o n , seems t o r e f l e c t a n a t u r a l g r o u p i n g , n o r does i t a p p e a r t o h e l p us more e a s i l y c o n c e p t u a l i s e t h e r e l a t i o n s h i p s and a f f i n i t i e s o f t h e s e p l a n t s . T h u s , I f a v o u r t h e r e c o g n i t i o n o f t h e M y r i o n e m a t a c e a e , t h e R a l f s i a c e a e ( t o i n c l u d e t h e L i t h o d e r m a t a c e a e ) and a l s o t h e Nemodermataceae, and c o n s i d e r t h a t a t p r e s e n t t h e y a r e most a p p r o p r i a t e l y p l a c e d i n t h e E c t o c a r p a l e s . A s more i s l e a r n t a b o u t t h e members o f t h e R a l f s i a c e a e t h i s f a m i l y w i l l p r o b a b l y be d i v i d e d i n t o g r o u p i n g s b a s e d more on r e l a t i o n s h i p s and a f f i n i t i e s and l e s s on t h e c u r r e n t b a s i s o f h a b i t and p s e u d o p a r e n c h y m a t o u s c o n s t r u c t i o n . T h f t a x o n o m i c p o s i t i o n o f A n a l i p u s j a p o n i c u s (Harv.) Wynne E a r l y t a x o n o m i c t r e a t m e n t s o f A n a l i p u s j a p o n i c u s (Harv.) Wynne (as C h o r d a r i a a b i e t i n a Rupr. ex F a r l o w and l a t e r a s H e t e r c c h o r d a r j a a b i e t i n a (Rupr. ex F a r l o w ) S. and G. ) d e a l t e x c l u s i v e l y w i t h c h a r a c t e r i s t i s t i c s o f t h e e r e c t a x e s and p l a c e d t h i s p l a n t e i t h e r i n t h e C h o r d a r i a c e a e ( e . g i F a r l o w , 1875; Okamura, 1910, 1933; K y l i n , 1940; S c a g e l , 1957; I n a g a k i , 1958) o r t h e H e t e r o c h o r d a r i a c e a e (a m o n o t y p i c f a m i l y e r e c t e d by S e t c h e l l and G a r d n e r , 1925) (e . g . N a g a i , 1940; S m i t h , 1944). Nakamura ( 1972) p l a c e d A . j a p o n i c u s i n t h e R a l f s i a c e a e and t h i s t r e a t m e n t h a s been f o l l o w e d by Wynne (1972) and B o l d and Wynne (1978) a l t h o u g h 203 A b b o t t and H o l l e n b e r g (1976) c l a s s i f y t h i s p l a n t i n t h e Ch c r d a r i a c e a e . As d i s c u s s e d a t t h e b e g i n n i n g o f t h i s c h a p t e r , Nakamura's j u s t i f i c a t i o n f o r t h e p l a c e m e n t o f A n a l i p u s w i t h t h e R a l f s i a c e a e was i t s d i s c o i d a l e a r l y d e v e l o p m e n t , and anatomy, p a r t i c u l a r l y t h a t o f i t s r e p r o d u c t i v e s t r u c t u r e s . The r e v i e w o f t h e l i t e r a t u r e h e r e has shown t h a t t h e s e c r i t e r i a a r e n o t s h a r e d by many o t h e r members of t h e g r o u p o f p l a n t s g e n e r a l l y known a s t h e R a l f s i a c e a e . A l t h o u g h t h e anatomy o f t h e A . j a p o n i c u s c r u s t i s s i m i l a r t o t h a t o f R a l f s i a v e r r u c o s a , i t i s v e r y d i f f e r e n t f r o m t h a t o f some o t h e r R a l f s i a s p e c i e s and o t h e r members o f t h e R a l f s i a c e a e (e.g. P e t r o d e r m a , Symphyocarpus, B a s i s p o r a , R. c l a y a t a , J» s p o n q i o c a r p a ) . A l t h o u g h t h e u n i l o c u l a r s p o r a n g i a i n A n a l i p u s 'are t e r m i n a l / l a t e r a l and immersed i n p a r a p h y s e s , and t h e p l u r i l o c u l a r o r g a n s a r e i n t e r c a l a r y , t h e y a r e b o r n e on the u p r i g h t t h a l l u s s y s t e m and t h e c r u s t i s n e v e r f e r t i l e . I c o n s i d e r t h a t A n a l i p u s i s g u i t e d i s t i n c t f r o m t h e m a j o r i t y o f c r u s t s now r e c o g n i s e d i n t h e R a l f s i a c e a e . The genus A n a l i p u s does n o t seem t o be a p p r o p r i a t e l y p l a c e d i n t h e C h o r d a r i a c e a e . At t h e moment t h e r e a r e a number o f d i f f e r e n t u n d e r s t a n d i n g s o f t h e o r d e r C h o r d a r i a l e s . Some t r e a t m e n t s o f t h i s o r d e r i n c l u d e t h e C h o r d a r i a l e s w i t h i n t h e E c t o c a r p a l e s ( e.g. R u s s e l l , 1964; P a r k e and D i x o n , 1964) as o r i g i n a l l y p r o p o s e d by O l t m a n n s (1922) and f o l l o w e d by F r i t s c h (1945). B o l d and Wynne 204 (1S78) c o n s i d e r , however, t h a t t h e C h o r d a r i a l e s s h o u l d be m a i n t a i n e d as a d i s t i n c t o r d e r "on t h e b a s i s o f t h e h e t e r o m o r p h i c l i f e h i s t o r y i n which t h e h a p l c i d m i c r o t h a l l u s a l t e r n a t e s w i t h a d i p l o i d m a c r o t h a l l u s " . . I f t h e s e p a r a t i o n o f t h e E c t o c a r p a l e s and C h o r d a r i a l e s i s u n d e r s t o o d t o be p r i m a r i l y on l i f e h i s t o r y c h a r a c t e r i s t i c s , A n a l i p u s s h o u l d n o t be p l a c e d i n t h e C h o r d a r i a l e s as i t has been shown t c p o s s e s s an a l t e r n a t i o n o f i s o m o r p h i c g e n e r a t i o n s . I n my o p i n i o n t h e genus A n a l i p u s s h o u l d n o t be p l a c e d i n e i t h e r t h e R a l f s i a c e a e o r C h o r d a r i a c e a e but w a r r a n t s p l a c e m e n t i n a s e p a r a t e f a m i l y . The m o n o t y p i c f a m i l y H e t e r o c h o r d a r i a c e a e was e r e c t e d by S e t c h e l l and G a r d n e r i n 1925 f o r H e t e r o c h p r d a r i a a b i e t i n a (Rupr. §x F a r l o w ) S. and G. . H e t e r o c h o r d a r i a was merged w i t h A n a l i p u s by Wynne (1 9 7 1 ) . The f a m i l y i s l e g i t i m a t e as i t was b a s e d on a l e g i t i m a t e genus H e t e r o c h o r d a r i a S. and G..1924 and "B e c a u s e i t s merger w i t h A n a l i p u s i s a m a t t e r o f t a x o n o m i c o p i n i o n , i t was not n o m e n c l a t u r a l l y s u p e r f l u o u s when p u b l i s h e d . " (Dr P.C. S i l v a - p e r s . c o m m . ) . . S e t c h e l l and G a r d n e r ' s (1925) d e s c r i p t i o n o f t h e H e t e r o c h o r d a r i a c e a e i n c l u d e d 1) v e g e t a t i v e c h a r a c t e r s o f t h e u p r i g h t a x e s , and 2) t h e o c c u r r e n c e o f i s o m o r p h i c p l a n t s b e a r i n g e i t h e r u n i l o c u l a r o r p l u r i l o c u l a r s t r u c t u r e s . I t was t h i s s e c o n d f e a t u r e which was t h e p r i m a r y b a s i s f o r s e p a r a t i o n o f t h i s genus f r o m t h e C h o r d a r i a c e a e . . W h e n S m i t h (1944) u s e d t h e H e t e r o c h o r d a r i a c e a e ( p l a c e d i n t h e E c t o c a r p a l e s ) t h e f a m i l y 205 c h a r a c t e r i s t i c s g i v e n i n c l u d e d m e n t i o n o f the c r u s t o s e s y s t e m , and t h e p o s i t i o n o f t h e u n i l o c u l a r and p l u r i l o c u l a r s p o r a n g i a . I n o r d e r t o accommodate b o t h A . , f i l i f o r m i s (Rupr.) P a p e n f u s s and A . j a p o n i c u s i n t h i s f a m i l y t h e o r i g i n a l d e s c r i p t i o n n e e d s t o be emended.. Two o t h e r s p e c i e s p o s s i b l y f a l l w i t h i n t h i s f a m i l y but r e g u i r e c l o s e r e x a m i n a t i o n b e f o r e t h i s c a n be e f f e c t e d . . T h e s p e c i e s C h o r d a r i a g u n j i i Yendo (Yendo, 1913) (or H e t e r o c h o r d a r i a g u n j i i (Yendo) T o k i d a - T o k i d a , 1938; N a g a i , 1940) was d i s c u s s e d by Wynne (1971). I have examined t h e t y p e m a t e r i a l o f t h i s s p e c i e s and on t h e b a s i s of t h e anatomy o f t h e e r e c t s y s t e m ( o f b o t h p l u r i l o c u l a r and u n i l o c u l a r p l a n t s ) i t would a p p e a r t o f a l l w i t h i n t h e genus A n a l i p u s . As I have o n l y s e e n t h e e r e c t a x e s o f t h i s s p e c i e s and none o f t h e c r u s t o s e p o r t i o n which h a s been r e p o r t e d , I am n o t a b l e t o make any judgement on i t s t a x o n o m i c p o s i t i o n . I t d e s e r v e s c l o s e a t t e n t i o n , however, as a l i k e l y t h i r d s p e c i e s i n t h i s genus. The o t h e r s p e c i e s o f i n t e r e s t h e r e i s t h e s u b - a n t a r c t i c p l a n t S t e r e o c l a d p n r u g u l p s u m ( S k o t t s b e r g , 1921; P a p e n f u s s , 1964) w h i c h p o s s e s s e s a c r u s t o s e r a l f s i o i d s y s t e m from which a r i s e e r e c t a x e s w i t h a " C h o r d a r i a " - l i k e c o n s t r u c t i o n . T h i s p l a n t h a s n o t r e c e i v e d a t t e n t i o n s i n c e S k c t t s b e r g ' s t r e a t m e n t and need s f u r t h e r e x a m i n a t i o n t o d e t e r m i n e i t s t a x o n o m i c a f f i n i t i e s . 206 GENERAL DISCUSSION 207 A v a r i e t y o f a p p r o a c h e s have been used i n t h i s s t u d y of t h e b i o l o g y o f A n a l i p u s j a p o n i c u s . D a t a o b t a i n e d f r o m l i f e h i s t o r y , a n a t o m i c a l and c y t o l o g i c a l s t u d i e s have e n a b l e d a r e - e v a l u a t i o n of t h e t a x o n o m i c a f f i n i t i e s o f t h i s a l g a . A m o r p h o l o g i c a l and a n a t o m i c a l c o m p a r i s o n o f A . j a p o n i c u s and C a e p i d i u m a n t a r c t i c u m was made b e c a u s e o f r e p o r t s i n t h e l i t e r a t u r e o f t h e i r s i m i l a r i t y . W h i l s t c e r t a i n d i f f e r e n c e s and s i m i l a r i t i e s o f t h e s e two were d i s t i n g u i s h e d , l i f e h i s t o r y and c y t o l o g i c a l d a t a a r e n e e ded f o r C . a n t a r c t i c u m b e f o r e i t s a f f i n i t i e s c an be f u l l y u n d e r s t o o d . T h i s g e n e r a l d i s c u s s i o n i s an a t t e m p t t o i n t e g r a t e t h e d a t a d e r i v e d from t h e v a r i o u s a p p r o a c h e s I e m p l o y e d , and t o p o i n t t o g u e s t i o n s which r e m a i n o u t s t a n d i n g . The c u l t u r e and c y t o l o g i c a l s t u d i e s have c o n f i r m e d an a l t e r n a t i o n o f i s o m o r p h i c g e n e r a t i o n s f o r A. j a p o n i c u s . However, t h e p h e n o l o g i c a l and e c o l o g i c a l s t u d i e s show t h a t f o r A . j a p o n i c u s , i n t h e e a s t e r n P a c i f i c , t h e h a p l c i d p l u r i l o c u l a r phase p r e d o m i n a t e s and s e x u a l r e p r o d u c t i o n and u n i l o c u l a r p l a n t s a r e r a r e . I n a r e v i e w o f t h e P h a e o p h y t a , R u s s e l l (1973) o b s e r v e d t h a t i n t h o s e brown a l g a e w h i c h have been s t u d i e d , a number have been shown t o p o s s e s s s o l e l y a s e x u a l l i f e h i s t o r i e s , r e p r o d u c i n g e i t h e r by p l u r i s p o r e s o r u n i s p o r e s . I t has been s u g g e s t e d t h a t i n some o f t h e s e s p e c i e s t h e r e p o r t s may be a r t i f a c t s o f c u l t u r e c o n d i t i o n s ( E d e l s t e i n , Chen and M c L a c h l a n , 197 1 ) . I t has a l s o been f o u n d t h a t d i f f e r e n t p o p u l a t i o n s o r 208 i s o l a t e s o f t h e same s p e c i e s e x h i b i t d i f f e r e n t l i f e h i s t o r i e s ( e . g . Hecatpnema m a c u l a n s - L o i s e a u x , 1969; E d e l s t e i n e t a l . , 1971). B u s s e l l (1973) s t a t e s , however, " t h e i n c i d e n c e o f g e n u i n e l y a s e x u a l l i f e h i s t o r i e s , p a r t i c u l a r l y i n t h e E c t o c a r p a l e s , seems l i k e l y t o be h i g h " . I n t h i s s t u d y , p o t e n t i a l l y s e x u a l p o p u l a t i o n s o f A . j a p o n i c u s were i d e n t i f i e d o n l y i n J a p a n and i n one s i t e i n C a l i f o r n i a . The g e n e r a l l y a c c e p t e d wisdom i n b i o l o g y i s t h a t s e x u a l r e p r o d u c t i o n i s a p o s i t i v e f a c t o r i n t h e e v o l u t i o n a r y p o t e n t i a l o f a p o p u l a t i o n and i n i t s s u r v i v a l (Mayr, 1963; W i l l i a m s , 1975). As W i l l i a m s (1975) n o t e s , i f t h e s e b e n e f i t s a r e i m p o r t a n t , t h e r e l a t i v e i n f e r i o r i t y o f a s e x u a l f o r m s s h o u l d be d e m o n s t r a b l e i n t e r m s o f e c o l o g i c a l s u c c e s s and g e n e t i c p o t e n t i a l . T h e r e i s l i t t l e e v i d e n c e , however, t o s u p p o r t t h i s i d e a . Amongst weeds b o t h a s e x u a l i t y and s e l f i n g a r e common ( A l l a f d , 1965; B a k e r , 1 9 7 4 ) . . W i l l i a m s (1975) o b s e r v e s t h a t t h e s u c c e s s o f a s e x u a l o r s e l f e d weeds may be due t o a number o f p o s s i b l e r e a s o n s " t h e a b i l i t y t o e s t a b l i s h t h e m s e l v e s w i t h s i n g l e p r o p a g u l e s , t h e i r l a c k o f r e p r o d u c t i v e l y w a s t e f u l male f u n c t i o n s , t h e i r f r e g u e n t p o l y p l o i d y , o r f r e e d o m f r o m r e c o m b i n a t i o n a l l o a d " . A p p a r e n t l y a s e x u a l i t y i n A . j a p o n i c u s i s a d e r i v e d f e a t u r e , w i t h a t l e a s t one remnant s e x u a l p o p u l a t i o n r e m a i n i n g i n C a l i f o r n i a . No i n f o r m a t i o n i s a v a i l a b l e on t h e d i s t r i b u t i o n o f u n i l o c u l a r and p l u r i l o c u l a r p o p u l a t i o n s o f A . j a p o n i c u s i n t h e w e s t e r n 209 P a c i f i c . I t would be most i n t e r e s t i n g t o have more i n f o r m a t i o n on t h e r e l a t i v e c o n t r i b u t i o n s of s e x u a l and a s e x u a l r e p r o d u c t i o n t o p o p u l a t i o n m a i n t e n a n c e and v a r i a t i o n i n t h e s e r e g i o n s . A l t h o u g h t h e g e n e r a t i o n s o f A . j a p o n i c u s a r e i s o m o r p h i c , w i t h i n e a c h g e n e r a t i o n t h e t h a l l u s can be r e g a r d e d as c o n s i s t i n g o f two g u i t e d i f f e r e n t p a r t s - t h e c r u s t and t h e u p r i g h t a x e s b o r n e cn t h e c r u s t . The c r u s t i s p e r e n n i a l and a p p e a r s t o p l a y a s i g n i f i c a n t r o l e i n t h e s u r v i v a l and s u c c e s s o f t h i s s p e c i e s . I t i s t h e c o l o n i s i n g p o r t i o n o f t h e p l a n t . A l t h o u g h i t d o e s n ' t b e a r s p o r a n g i a , t h e c r u s t i s a b l e t o s p r e a d v e g e t a t i v e l y h o l d i n g o n t o s e t t l e m e n t s p a c e - t h e r e s o u r c e which i s a t a premium i n t h e i n t e r t i d a l r e g i o n . . I t a p p e a r s t o be r e s i s t a n t t o e n v i r o n m e n t a l s t r e s s and a l s o t o g r a z i n g . The e r e c t a x e s , however, a r e r e l a t i v e l y s h o r t l i v e d and a r e c o n s t a n t l y b e i n g r e p l a c e d by g r o w t h from t h e c r u s t . P a r t i c u l a r l y i n t h e p l u r i l o c u l a r p l a n t s t h e r e i s a v e r y h i g h i n v e s t m e n t i n r e p r o d u c t i v e t i s s u e . T hus, t h e two p a r t s o f t h e t h a l l u s h ave v e r y d i f f e r e n t m o r t a l i t y r a t e s and a d a p t a t i o n s t o t h e i r i n t e r t i d a l e x i s t e n c e . The l i f e h i s t o r i e s o f a number o f a l g a e i n c l u d e b o t h a c r u s t and an e r e c t phase i n s e p a r a t e g e n e r a t i o n s ( e . g i R a l f s i a and P e t a l o n i a ; P e t r o c e l i s and G i g a r t i n a ) and i t h as been c o n s i d e r e d t h a t t h e s e d i s t i n c t m o r p h o l o g i c a l p h a s e s have e v o l v e d under d i f f e r e n t s e l e c t i v e p r e s s u r e s ( L i t t l e r and L i t t l e r , 1980; L u b c h e n c o , 1980). S t u d i e s on 210 P e t r o c e l i s m i d d e n d g r f i i have shown i t t o be slow g r o w i n g , w i t h b o t h lew r e c r u i t m e n t and m o r t a l i t y r a t e s ( P a i n e , Slocum and D u g g i n s , 1979). . I t s a l t e r n a t e g e n e r a t i o n G i g a r t i n a p a p i l l a t a has a s i g n i f i c a n t l y d i f f e r e n t g r o w t h s t r a t e g y and: p r o d u c t i v i t y ( L i t t l e r and L i t t l e r , 1980) . A . j a p o n i c u s d i f f e r s from t h e s e p l a n t s , however, i n p o s s e s s i n g two m o r p h o l o g i c a l l y and f u n c t i o n a l l y d i f f e r e n t p a r t s w i t h i n t h e same c a r y o l o g i c a l phase. T h e r e a r e many examples o f a l g a e s u c h as A . j a p o n i c u s i n which an e r e c t a x i s o r b l a d e i s b o r n e on a c r u s t o s e o r p e r e n n a t i n g s t r u c t u r e <e.q.Apgphloea l y a l l i i , I r i d a e a c o r d a t a , C h o n d r u s c r i s p u s , P h a e o s t r p p h i o n i r r e q u l a r e , C a e p i d i u m a n t a r c t i c u m ) . The p l a n t s have been t e r m e d H e m i c r y p t o p h y c e a e by Feldmann (1937) on t h e b a s i s o f t h e i r " l i f e - f o r m " . T h i s i s e q u i v a l e n t t o t h e t e r m h e m i c r y p t o p h y t e u s e d i n t e r r e s t r i a l p l a n t s i n t h e t e r m i n o l o g y of B a u n k i a e r • p l a n t l i f e - f o r m s ' ( M u e l l e r - D o m b o i s and E l l e n b e r g , 1 9 7 4 ) . Frcm s t u d i e s i n B r i t t a n y , E r n s t (1968) s u g g e s t s t h a t t h e r e i s a v e r t i c a l z o n a t i o n o f l i f e - f o r m s i n marine a l g a l c o m m u n i t i e s , w i t h t h e sequence from u p p e r i n t e r t i d a l t o s u b t i d a l r e g i o n s b e i n g p h a n e r o p h y c e a e , h e m i p h a n e r o p h y c e a e , h e m i c r y p t o p h y c e a e , and chamaephyceae. I n a s t u d y o f I r i d a e a c p r d a t a (Hansen, 1977) growth o f u p r i g h t t h a l l i was f o u n d t o o c c u r m a i n l y from t h e p e r e n n a t i n g c r u s t s . Hansen l i k e n e d t h e r o l e of t h e c r u s t t o t h a t o f t h e l a t e r a l l y s p r e a d i n g r o o t s y s t e m i n some v a s c u l a r p l a n t s . The m a j o r i t y o f t h e b l a d e s d e v e l o p e d t e t r a s p o r a n g i a and t h e r e f o r e t h e 211 p o p u l a t i o n s h o u l d h a v e had a h i g h p o t e n t i a l f o r s e x u a l r e p r o d u c t i o n . However, Hansen f o u n d t h a t g a m e t a n g i a l t h a l l i were o n l y a minor component o f t h e p o p u l a t i o n and a p p a r e n t l y t h e c o n t r i b u t i o n s o f s e x u a l r e p r o d u c t i o n and s p o r e l i n g d e v e l o p m e n t were m i n i m a l t o t h e p o p u l a t i o n g r o w t h and m a i n t a n a n c e . . L u b c h e n c o (1980) f o u n d t h a t i n New E n g l a n d s h o r e s , p e r e n n a t i o n f r o m t h e h o l d f a s t / c r u s t o f C h o n d r u s c r i s p u s was a s i g n i f i c a n t f e a t u r e i n i t s e c o l o g i c a l s u c c e s s and l o n g term p e r s i s t e n c e . M a t h i e s o n (1965) c a r r i e d o u t an a u t e c o l o g i c a l and l i f e h i s t o r y s t u d y o f P h a e o s t r p p h i o n i r r e g u l a r e , a brown a l g a which l i v e s i n h a b i t a t s which a r e s e a s o n a l l y b u r i e d i n s a n d . He o b s e r v e d t h a t t h e b l a d e s a r e e p h e m e r a l and do n o t s u r v i v e t h e s e a s o n a l submergence i n s a n d . The c r u s t , however, was f o u n d t o be an i m p o r t a n t e l e m e n t i n t h e s u r v i v a l o f P h a e o s t r o p h i o n i n i t s r i g o r o u s h a b i t a t . The c r u s t i s much more r e s i s t a n t t h a n t h e b l a d e s t o s a n d - b u r i a l and i s c a p a b l e o f i n i t i a t i n g new b l a d e s once i t i s u n c o v e r e d . M a t h i e s o n r e g a r d e d t h e l i f e h i s t o r y o f P h a e o s t r o p h i o n - a d i r e c t l i f e h i s t o r y w i t h no e v i d e n c e o f s e x u a l i t y - t o have c o n s i d e r a b l e a d a p t i v e v a l u e i n t h e p l a n t ' s s u c c e s s . A l t h o u g h v e g e t a t i v e r e p r o d u c t i o n and p e r e n n a t i o n have been d i s c u s s e d ( K n i g h t and P a r k e , 1931; D i x o n , 1965) as i m p o r t a n t f e a t u r e s i n t h e growth of p e r e n n i a l a l g a e , i n t h e o p i n i o n o f Hansen (1977) g e n e r a l l y t h e s e f e a t u r e s have been u n d e r e s t i m a t e d or n o t c o n s i d e r e d . D i f f e r e n c e s i n t h e c r u s t and e r e c t p o r t i o n s o f t h e A . j a p o n i c u s t h a l l u s d e s e r v e f u r t h e r a t t e n t i o n t o u n d e r s t a n d 212 more f u l l y t h e r e l a t i o n s h i p between t h e s e p h a s e s and t h e i r r e l a t i v e c o n t r i b u t i o n s t o t h e p l a n t ' s c o m p e t i t i v e a b i l i t y . I t would be i n t e r e s t i n g t o s t u d y p h o t o s y n t h e t i c a c t i v i t i e s o f t h e c r u s t and o f e r e c t a x e s , and t h e i r r e l a t i v e i m p o r t a n c e t o growth and r e p r o d u c t i o n - t h a t i s , whether c r u s t s a r e s u b s i d i s i n g growth and r e p r o d u c t i v e a c t i v i t y o f u p r i g h t s or whether u p r i g h t s a r e e x p o r t i n g p h o t o s y n t h a t e s t o the c x u s t f o r s t o r a g e , o r growth and e x p a n s i o n o f t h e c r u s t o s e system* The s e a s o n a l p a t t e r n o f growth and a l l o c a t i o n o f p h o t o s y n t h e t i c p r o d u c t s may w e l l v a r y , w i t h t h e c r u s t c o n t r i b u t i n g s i g n i f i c a n t l y t o growth o f e r e c t a x e s a f t e r a p e r i o d o f s t r e s s o r g r a z i n g , when e r e c t a x e s have been removed o r s u b s t a n t i a l l y r e d u c e d . E v o l u t i o n o f l i f e h i s t o r y s t r a t e g i e s o r l i f e h i s t o r y t r a i t s has r e c e i v e d c o n s i d e r a b l e a t t e n t i o n i n r e c e n t y e a r s and was r e c e n t l y r e v i e w e d by S t e a r n s (1976, 1977). M o d e l s have been d e v e l o p e d t o e x p l a i n a d a p t a t i o n o f l i f e h i s t o r y t r a i t s t o s t a b l e or t o f l u c t u a t i n g e n v i r o n m e n t s . S t e a r n s (1977) r e v i e w e d d e t e r m i n i s t i c ( e . g . M a c A r t h u r and W i l s o n , 1967; P i a n k a , 1970) and s t o c h a s t i c ( e . g . .Murphy, 1968; S c h a f f e r , 1974) models o f l i f e h i s t o r y t r a i t s . He c o n c l u d e d t h a t n e i t h e r t y p e o f model i s e m p i r i c a l l y s u f f i c i e n t " t h e i r p r e d i c t i o n s a r e n o t c o n s i s t e n t w i t h much of t h e e v i d e n c e . We do n o t y e t have a g e n e r a l and r e l i a b l e t h e o r y o f l i f e h i s t o r y e v o l u t i o n . " . S e a r l e s (1980) d i s c u s s e s t h e e v o l u t i o n o f t h e t r i p h a s i c l i f e h i s t o r y o f r e d a l g a e . . He c o n s i d e r s t h a t t h i s complex l i f e h i s t o r y p a t t e r n c a n be 213 r e l a t e d t o t h e l a c k o f m o t i l i t y o f t h e gametes and t h e a p p a r e n t l y i n f r e q u e n t o c c u r r e n c e o f syngamy.. However, S e a r l e s was a d d r e s s i n g t h e g e n e r a l i s e d l i f e h i s t o r y r a t h e r t h a n t h e a d a p t a t i o n s t r a t e g y o f an i n d i v i d u a l s p e c i e s o r p o p u l a t i o n . The s t r a t e g y o f an i n d i v i d u a l a l g a i s n o t g o i n g t o be s o l e l y d e f i n e d by i t s p o t e n t i a l mechanisms f o r s e x u a l and a s e x u a l r e p r o d u c t i o n b u t i s a l s o t h e r e s u l t o f i t s morphology o r " l i f e - f o r m " and i t s p h y s i o l o g y . T h e r e i s v e r y l i t t l e i n f o r m a t i o n a v a i l a b l e f o r seaweed s p e c i e s on b a s i c p o p u l a t i o n c h a r a c t e r i s t i c s s u c h as n a t a l i t y and m o r t a l i t y , f e c u n d i t y , r e p r o d u c t i v e e f f o r t , age o f f i r s t r e p r o d u c t i o n (Chapman, 1974). B e f o r e t h e u n d e r s t a n d i n g o f a l g a l l i f e h i s t o r y s t r a t e g i e s c a n be e x t e n d e d and g e n e r a l i s a t i o n s o f any c r e a t i v e v a l u e a r r i v e d a t , more d a t a a r e needed on t h e s e b a s i c p o p u l a t i o n p r o c e s s e s . REFERENCES , 215 A b b o t t , I . A . and H o l l e n b e r g , G . J . 1976. M a r i n e A l g a e o f C a l i f o r n i a . . S t a n f o r d U n i v e r s i t y P r e s s , S t a n f o r d . . 8 2 7 pp. Abe,K. 1935a. Zur K e n n t n i s d e r E n t w i c k l u n g s g e s c h i c h t e von H e t e r o c h o r d a r i a , S c y t o s i p h o n , und S o r o c a r p u s . Tohoku I m p . U n i v . , S e n d a i , S c i . E e p . , Ser.4 ( B i o l . ) 9:329-337. Abe,K..1935b. K o p u l a t i o n d e r Schwarmer aus U n i l o k u l a r e m S p o r a ngium von H e t e r o c h o r d a r i a a b i e t i n a . Tohoku Imp.Univ. , S e n d a i , S c i . B e p . , S e r . 4 ( B i o l . ) 10: 287-290. . Abe,K. 1 9 3 6 . . K e r n p h a s e n w e c h s e l von H e t e r o c h o r d a r i a a b i e t i n a . . T o h o k u I m p . U n i v . , S e n d a i , S c i . B e p . , S e r . 4 ( B i o l . ) J J:239-241. Agardh,J.G..1882. T i l l A l g e r n e s S y s t e m a t i k . L u n ds U n i v . A r s s k r . J7: 1-134, 3 p i . A l l a r d , B . W . 1965. G e n e t i c s y s t e m s a s s o c i a t e d w i t h c o l o n i z i n g a b i l i t y i n p r e d o m i n a n t l y s e l f - p o l l i n a t i n g s p e c i e s . I n " G e n e t i c s o f C o l o n i z i n g S p e c i e s " (ed. H.G.Baker and G . L . S t e b b i n s ) pp.49-76. Academic P r e s s . Ancn..1977,1978,1979..Canadian T i d e and C u r r e n t T a b l e s - P a c i f i c C o a s t . V o l . 5 and 6. C a n a d i a n H y d r o g r a p h i c S e r v i c e . E n v i r o n m e n t Canada. A s e n s i , A . , D e l e p i n e , R . , and G u g l i e l m i , G . 1977. N o u v e l l e s o b s e r v a t i o n s s u r 1 • u l t r a s t r u c t u r e du p l a s t i d o m e d e s P h e o p h y c e e s . S o c . P h y c o l . de F r a n c e , B u l l . . 2 2 : 1 9 2 - 2 0 5 . A t o b e , S . and S a i t o , Y . 1 9 7 4 . . P h y t o s o c i o l o g i c a l s t u d y o f t h e i n t e r t i d a l m a r i n e a l a g e . I I I . E f f e c t o f wave a c t i o n on a l g a l z o n a t i o n . B u l l . F a c . F i s h . H o k k a i d o U n i v . . 2 4 : 133-138. Baker,H.G. 1974. The e v o l u t i o n o f weeds* A n n . R e v . E c o l . S y s t . 5:1-24. B a r k a , T . .and A n d e r s o n , P . J . . 1965. H i s t o c h e m i s t r y - T h e o r y , P r a c t i c e and B i b l i o g r a p h y . H a r p e r Row. 660 pp* B a t t e r s , E . A . L . . 1 8 9 0 . A l i s t o f t h e marine a l g a e o f B e r w i c k - on-Tweed. H i s t . B e r w i c k s h . N a t . C l u b . J2:221-392, p i s . 7 - 11. B old,H.C. and Wynne,M.J. 1978. I n t r o d u c t i o n t o t h e A l g a e - S t r u c t u r e and R e p r o d u c t i o n . P r e n t i c e - H a l l , I n c . , Englewood C l i f f s , New J e r s e y . 706 pp. 216 B o r g e s e n , F . . 1 9 2 6 . . M a r i n e a l g a e f r o m t h e Ca n a r y I s l a n d s . . I I . P h a e o p h y c e a e . K. DansJce V i d e n s k a b . S e l s k a b . B i o l * Meddel. 6 (2) . Breeman,A.M. 1979..The c a r y o l o g i c a l p h a s e s i n the l i f e h i s t o r y c f AcrosymphYton p u r p u r i f e r u m ( J . A q . ) S j o s t . (Rhodophyceae, C r y p t o n e m i a l e s ) . P h y c o l o g i a 18:146- 148. . Chapman,A.R.0..1973a..A c r i t i q u e o f p r e v a i l i n g a t t i t u d e s t o w a r d c o n t r o l o f seaweed z o n a t i o n on t h e s e a s h o r e * B o t a n i c a M a r i n a .16:80-82.. Chapman rA.B.0. 1973b. .Methods f o r m a c r o s c o p i c a l g a e . .In "Handbook o f P h y c o l o g i c a l Methods" (ed. . J . S t e i n ) Cambridqe U n i v e r s i t y P r e s s . .pp.87-104. . Chapman,A.R.O. 1974. The e c o l o q y o f m a c r o s c o p i c a l g a e * A n n . B e v . E c o l . S y s t . 5:65-80. C o n n e l l , J . H . . 1 9 7 2 . . C o m m u n i t y i n t e r a c t i o n s on m a r i n e r o c k y i n t e r t i d a l s h o r e s . A n n . B e v . E c o l . S y s t . 3:169-192. C o t t o n r A . 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C . 1975. Sex and E v o l u t i o n . P r i n c e t o n U n i v e r s i t y P r e s s . 20C pp. Wittmann,W..1965. A c e t o - i r o n - h a e m a t o x y l i n - c h l o r a l h y d r a t e f o r chrcmosome s t a i n i n g . S t a i n T e c h n o l . 40:161-164. Wynne,M.J. 1969. L i f e h i s t o r y and s y s t e m a t i c s t u d i e s o f some P a c i f i c N o r t h A m e r i c a n Phaeophyceae (brown a l g a e ) . . U n i v . C a l i f . P u b l . B o t . .50: 1-88. . Wynne,M.J. 1 9 7 1 . . C o n c e r n i n g t h e pha e o p h y c e a n g e n e r a A n a l i p u s and H e t e r o c h o r d a r i a . P h y c o l o g i a .10: 169-17 5; Wynne,M.J. 1972. S t u d i e s on t h e l i f e f o r m s i n n a t u r e and i n c u l t u r e o f s e l e c t e d brown a l g a e . I n " C o n t r i b u t i o n s t o t h e S y s t e m a t i c s o f B e n t h i c M a r i n e A l g a e o f t h e N o r t h P a c i f i c " . . (ed. I . A . A b b o t t and M.Kurogi) J a p a n e s e S o c . P h y c o l o g y , Kobe. pp.133-145.. Wynne,M.J. and L o i s e a u x , S . 1976. Recent a d v a n c e s i n l i f e h i s t o r y s t u d i e s o f t h e P h a e o p h y t a . P h y c o l o g i a 15:4 35- 4 52. Yendo,K. 19 13. Some new a l g a e f r o m J a p a n . N y t t mag, n a t u r v i d . . 5 1 : 2 7 5 - 2 8 8 . . 230 APPENDICES, L i s t o f c o l l e c t i o n s i t e s and d a t e s of c o l l e c t i o n s BRITISH COLOMBIA A g u i l a r House P o i n t - 48°50'N, 125°08'W 18 August 1978, 16 O c t o b e r , 1978, 4 December 1978, 25 F e b r u a r y 1979, 25 A p r i l 1979, 10 J u n e 1979, 11 J u l y 1979, 23 August 1979 B a t h I s l a n d - 49°09'N, 123°40'W 29 June 1977, 30 J u l y 1977, 8 September 1977, 21 O c t o b e r 1977, 4 November 1977, 2 December 1977, 20 J a n u a r y 1978, March 1978, 28 March 1978, 7 A p r i l 1978, 12 May 1978, 24 May 1978, 21 J u n e 1978, 18 J u l y 1978, 13 August 1978, 6 September 1978, 6 O c t o b e r 1978, 19 November 1978, 8 December 1978, 13 J a n u a r y 1979, 10 March 1979,' 10 A p r i l 1979, 12 May 1979, 11 June 1979, 10 J u l y 1979, 24 August 1979 Botany Beach - 48°32'N, 124°27'W 10 March 1978, 22 June 1978, 3 F e b r u a r y 1979, 10 March 1979, 17 March 1979, 13 May 1979, 9 J u l y 1979, 15 March 198C B r a d y ' s Beach - 4 8 ° 4 9 ' N , 125°09'W 5 F e b r u a r y 1978, 17 August 1978, 2 December 1978, 25 F e b r u a r y 1979, 26 A p r i l 1979 B r e a k w a t e r I s l a n d - 49°08'N, 123°40'W 7 A p r i l 1978, 19 J u l y 1978 D i a n a I s l a n d - 4 8 ° 5 0 ' N , 125°11«H 16 O c t o b e r 1978, 2 December 1978, 10 June 1979, 12 J u l y 1979 H a i n e s I s l a n d - 4 8 ° 5 0 ' N , 125°10'W 28 May 1979 H e l b y I s l a n d - 4 8 ° 5 1 ' N , 125°10«W 26 A p r i l 1979, 27 May 1979, 12 J u l y 1979, 23 August 1979 Pachena Bay - 4 8 ° 5 1 ' N , 125°20'W 26 A p r i l 1979 Ross I s l e t s - 48°52'N, 125°09'W 21 May 1978 232 Second Beach - 48°49'N, 125°10'W 28 May 1978, 17 August 1978, 15 O c t o b e r 1978, 2 December 1978, 27 A p r i l 1979, 27 May 1979, 9 June 1979, 12 J u l y 1979, 23 August 1979 S e p p i n g s I s l a n d - 48°51'N, 125°12'W 28 May 1979 S i d n e y - 48°38'N, 123°25'W 20 J u l y 1978 Sombrio R i v e r - 4 8 ° 3 0 ' N , 124°17'W 14 May 1979 Sooke - 48°2 1 ' N , 1 2 3 ° 4 4 , W 18 F e b r u a r y 1978, 1 May 1978, 21 June 1978, 20 J u l y 1978, 3 March 1979, 1 A p r i l 1979 V i c t o r i a B r e a k w a t e r - 48°24»N, 123°24'W 1 A p r i l 1979, 19 Au g u s t 1979 W i z a r d Rock - 4 8 ° 5 1 ' N , 125°09'W 26 A p r i l 1979, 27 May 1979, 9 J u n e 1979, 12 J u l y 1979, 23 August 1979 WASHINGTON C a t t l e P o i n t - 48°27»N, 122°58'W 6 J u l y 1978 K a l a l o c h - 4 7 ° 5 5 ' N , 124°38'W 8 September 1978 Mar V i s t a - 4 8 ° 2 8 ' N , 123°01'W 6 J u l y 1978 OREGON B o i l e r Bay - 44 ° 5 0 ' N , 124°05'W 21 June 1978, 29 March 1979 Cape Arago ( s o u t h cove) - 4 3 ° 1 9 ' N , 124°24'W 1 13 June 1979 D e v i l ' s Punchbowl - 44°47'N, 124°05'W 30 March 1979 F o g a r t y C r e e k - 4 4 ° 4 9 ' N , 124°05'W 24 June 1978 H a r r i s Beach - 43°10»N, 124°20»W 28 March 1979 M a r i n e Gardens - 4 4 0 4 8 « N , 124°05'W 20 June 1978, 30 March 1979, 7 August 1979 S u n s e t Beach - 4 3 ° 1 8 « N , 124°23«W 29 March 1979 Y a g u i n a Head - 44°40'N, 124°04»W 20 June 1978 CALIFORNIA Anchor Bay - 3 8 ° 4 0 « N , 123°30'W 27 March 1979 C i t y ( M i s s i o n ) P o i n t - 36 ° 3 3«N, 1 2 1 ° 5 5«W 22 August 1979, 6 September 1979 D a v e n p o r t L a n d i n g - 3 7 3 0 0 ^ , 122° 11 • W 25 March 1979, 8 September 1979 Montana de Oro - 3 5 0 2 0 ^ , 120°50'W 23 March 1979 Moss Beach - 3 7 0 3 0 ^ , 122°30'W 5 September 1979 P e b b l e Beach - 3 7 ° 2 0 « N , 122°30«W 5 September 1979 P e s c a d e r o Beach - 3 7°25«N, 1 2 2 ° 2 8 , W 5 September 1979 P i e d r a s E l a n c a s - 3 5 ° 3 9 « N , 121°17»W 24 March 1979, 6 September 1979 P i g e o n P o i n t - 3 7°15'N, 122°30'W 26 March 1979, 8 September 1979 P o i n t Arena - 3 8 ° 5 5 « N , 1 2 3 ° 5 0 , W 27 March 1979 P o i n t J o e - 3 6 ° 3 7 « N , 1 2 1 ° 5 7«W 7 September 1979 P o i n t P i n o s - 3 6 ° 3 7 » N , 1 2 1 ° 5 7«W 7 September 1979 S t i l l w a t e r Cove - 36°36'N, 121°56«W 24 March 1979, 7 September 1979 234 T r i n i d a d P o i n t - 4 1 ° 0 5 ' N , 124°10«W 28 March 1979 W e s t p o r t - 3 9 ° 3 8 , N , 123°46«W 28 March 1979 235 Appendix I I R e p r e s e n t a t i v e h e r b a r i u m s p e c i m e n s of A n a l i p u s j a p o n i c u s (Harv.) Wynne PLUBILOCULAR J a p a n Onagava, R i k u z e n - 37°53«N, 140°04«E - UC 315269, FEB 1926 M u r c r a n , H o k k a i d o - 42°2C«N, 140°58«E - UBC A56339, 1 JUNE 1949; UBC A45981, 9 AUG 1971 A l a s k a Rocky P o i n t s , M a s s a c r e Bay, A t t u I s l a n d - 52°49«N, 173°11'W - UC M61586, 3 JULY 1953 T r a p p e r ' s Cove, Adak - 51°48'N, 176°50'W - UBC A8421, 14 JUNE 19 60 Cape Muzon, D a l l I s l a n d - 54°4 0 ' N , 132°41«W - UBC A20564, 13 JUNE 1965 K l o k a c h e f I s l a n d - 57°25'N, 135°53«W - UBC A22598, 30 JUNE 1965 S t . M a k a r i u s Bay, Amc h i t k a I s l a n d - 5 1 ° 2 3 ' N , 179°12'E - MICH, 16 MARCH 1968 Br i t i s h C o l u m b i a G l a c i e r P o i n t , V a n c o u v e r I s l a n d - 48 024'N, 123°59«W - UBC A32537, 23 MARCH 1968 Kyuguot, Garden I s l a n d - 50°01'N, 127°21«W - UBC A4074, 27 MAY 1959 O r l e b a r P o i n t , G a b r i o l a I s l a n d - 49°1 1 « N , 123°39'W - UBC A28922, 27 APRII 1967 M u i r C r e e k - 48 ° 2 2 « N , 123°52'W - UBC A 3400, 24 JUNE 1957 K e l s e y Bay - 50 ° 2 4 « N , 125°58»W - UBC A4076, 4 JUNE 1959 A g u i l a r House Beach - 48°50 « N , 125°08»W - UBC A41766, 29 JULY 1969 N o r t h Kalanawa R i v e r , V a n c o u v e r I s l a n d - 4 8 0 4 2 ^ , 124°59»W - UEC A7400, 10 JULY 1959 S a n d s t o n e C r e e k - 48 024'N, 125°00'W - UBC A3393, 10 AUG 1957 Maude I s l a n d , S t r a i t o f G e o r g i a - 50°07»N, 125°21«W - UBC A74C1, 15 AUG 1959 T r i a n g l e I s l a n d - 5 0 0 5 2 ^ , 129°05«W - UBC A19199, 7 SEPT 1964 B r o o k s P e n i n s u l a - 50°07«N, 1 2 7 0 4 2 « W - UBC A19734, 6 OCT 1964 R o l l e r Bay, Hope I s l a n d - 50°55«N, 1 2 7 0 5 6 » W - UBC A43856, 13 SEPT 1970 P o i n t - N o - P o i n t , V a n c o u v e r I s l a n d - 48 023'N, 123°59'W - UBC A39096, 22 NOV 1968 236 Washington West B e a c h , Whidbey I s l a n d - 48°13'N, 122°46«W - WTU 244774, 21 MAY 1966 T u r n P o i n t , San J u a n I s l a n d - 48°32»N, 122°59»W - WTU 244770, 18 JUNE 1955; OEC A2379 6 JULY 1956 Mukkaw Eay - 4 8 o 2 0 « N , 124°40«W - WTU 244771, 21 AUG 1949 Waadah I s l a n d - 4 8 ° 2 3 « N , 124°36«W - UBC A2771, 15 AUG 1958 Oregon M i d d l e Bay, Cape Arago - 43°29«N, 124°23»W - UC 751284, 30 JUNE 1942 H a r r i s Beach - 43°10«N, 124°20'W - UBC A5051, 19 MAY 1958 S u n s e t Beach - 43°1 8 « N , 124°23«W - UBC A60700, 29 March 1979 C a l i f o r n i a P o i n t S a l , S a n t a B a r b a r a Co. - 3 4 ° 4 0 M 2 0 , 45°'W - UC M200208, 14 MAY 1949 Pun t a G o r d a , Humboldt Co.. - 40°15'N, 124°20»W - UC 1- 462021, 13 JUNE 1968 E a g l e G l e n Canyon, S a n t a C r u z Co. - 37°05 ' N , 122°05«W - UC M316643, 10 JULY 1964 Moss Beach, San Mateo Co.. - 37°«N, 122°30'W - UC M106575, 5 SEPT 1925 P i e d r a s B l a n c a s - 3 5 0 2 5 ^ , 121°20«W - UBC A60650, 23 MAECH 1979 Tomales Head, M a r i n Co.. - 38°14'N, 122°59'W - UBC A19985, 24 JUNE 1964 M i s s i o n P o i n t - 3 6 ° 3 2 » N , 121°56«W - GMS 2507, 26 APBIL 1967 F o r t E o s s - 3 8 ° 3 2 ' N , 123°15'W - MICH, 2 NOV 1945 P o i n t L o b o s , M c n t e r e y Co. - 3 6 0 3 2 « N , 121°55'W - MICH, 3 DEC 1965 UNILOCULAR Ja p a n Ayukawa, Honshu - 38°15'N, 141°30«E - UBC A5503, 5 FEB 1957 Mu r o r a n , H o k k a i d o - 42°20'N, 140 o58'E - UC 360340, 31 MARCH 1926; UBC A56338,6 JUNE 1951 A p p e t s u , H o k k a i d o - 42°27«N, 142°40'E - UC 360338, 4 JUNE 1925 O s h o r o , H o k k a i d o - 42°53»N, 140°35«E - UC 360339, 10 JUNE 1926 C a 1 i f c r n i a P o i n t L o b o s , M c n t e r e y Co. - 36°32«N, 121°55«W - UC 74617, 15 JULY 1892 C i t y P o i n t , C a r m e l - 36°33'N, 1 2 1 0 5 5 ^ - GMS 1694, 19 JULY 1958 237 A p p e n d i x I I I The mean monthly s a l i n i t y and t e m p e r a t u r e o f s u r f a c e w a ter a t E n t r a n c e I s l a n d ( 4 9 0 1 2 « N , 123°48»W ) f o r J u l y 1977 t o August 1979. B a r s i n d i c a t e t h e r e c o r d e d m o n t h l y maxima and minima. ( G i o v a n d c , 1977, 1978, 1979) Entrance Island o 30H 25H l E C 20H o in 15H J L J L J I L J I L J I I I I I L o 20H 16- 12H a 3 * . 0 i . 01 a E 8H 1977 1978 1979 ~ i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — i — r * A O D F A J A O D F A J A Months ^ CP 239 Appendix IV The mean monthly s a l i n i t y and t e m p e r a t u r e o f s u r f a c e w a ter a t a m p h i t r i t e P o i n t ( 4 8 ° 5 4 « N , 125°33« H ) f o r J u l y 1978 t o J u l y 1979. B a r s i n d i c a t e t h e r e c o r d e d m o n t h l y maxima and minima. ( G i o v a n d c , 1978, 1979) 240 Amphitrite Point 30H o 25H 20- 16 u 14-H 12H 4) 5 io-*- a a) 8-a E 0) , h- 6" 4H 1978 s N M M 1979 241 Appendix V C o m p o s i t i o n o f e n r i c h m e n t s o l u t i o n s added t o a r t i f i c i a l s e a w a t e r . The volume added t o 1L o f s e a w a t e r b a s e i s i n b r a c k e t s . N u t r i e n t s and T r a c e M e t a l s (10ml) NaN03 4.667 g Na2Si03.9H20 3.000 Na g l y c e r o P04 0.667 Na 2EDTA 0.553 Fe (NH4) 2 (SO4) 2.6H20 0.234 FeC13.H20 0.016 MnS04.H20 0.041 ZnS04.7H20 0.007 COSC4.7H20 0.0016 g l a s s d i s t i l l e d w a t e r 1000 ml V i t a m i n s (1ml) T h i a m i n e HCI I n i s i t o l N i c o t i n i c a c i d C a l c i u m p a n t o t h e n a t e p - a m i n o b e n z o i c a c i d V i t a m i n B12 B i c t i n F o l i c a c i d g l a s s d i s t i l l e d w a ter 0.5 g 0.5 0.03 0.03 0.003 0.002 0.001 0.0006 1000 ml Appendix VI Sample dates of quadrats (QI-V) and t r a n s e c t (PT1-10) Quadrat 1. VII.77 29. VII.77 8. IX.77 21. X.77 5. XI.77 2. XII.77 20. 1.78 2. III.78 7. IV.78 11. V.78 23. V.78 20. VI.78 19. VII.78 14.VIII.78 6. IX.7|8 6. X.78 26. XI.78 13. 1.79 9. . 1 1 1 . 7 9 11. V.79 12. VI.79 9. VII.79 24. VIII.79 Transect 2. VII.77 29. VII.77 8.. IX.77 5. XI.77 20. 1.78 3. III.78 12. . V.78 24. V.78 20.. VI.78 19. VII.78 7.. IX.78 26. XI.78 13. . 1.79 9. I I I . 7 9 9. IV.79 12. V. 79 12. VI.79 9. VII* 79 243 a p p e n d i x V I I L i s t o f s p e c i e s r e c o r d e d f r o m g u a d r a t s and t r a n s e c t s a t B a t h I s l a n d s t u d y s i t e CHIGROPHYTA D i v a s p . PHAEOPHYTA A n a l i p u s j a p o n i c u s C o l p o m e n i a p e r e q r i n a C o l p o m e n i a b u l l o s a F u c u s d i s t i c h u s P e t a l o n i a f a s c i a Sargassum muticum S c j t o s i p h o n 1 o m e n t a r i a R a l f s i a sp. RHCCOPHYTA B o s s i e l l a o r b i q n i a n a C e r a m i u n s p . Cor a l i i , na v a n c c uver i e n s i s c r u s t o s e c o r a l l i n e s C r y p t o p l e u r a y i o l a c e a C r y p t o s i p h o n i a w o o d i i G i g a r t i n a a q a r d h i i I r i d a e a sp. 244 L i t h o t h r i x a s p e r g i l l u m B i c r p c l a d i a b o r e a l i s P e t r o c e l i s f r a n c i s c a n a Plocamium t e n u e P p r p h y r a s p . R h o d o j e l a l a r j x CHBYSOPHYTA N a y i c u l a s p . VERTEBRATES O l i g p c o t t u s m a c u l u l p s u s INVERTEERATES C o l l i s e l a p e l t a Notoacmea scutum I d c t e a w o s n e s e n s k i i G n crimosphaeroma o r e g o n e n s i s P i s a s t e r o c h r a c e u s

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