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Ecological study of Laminaria sinclairii and L. longipes Markham, James W. 1969

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AN ECOLOGICAL STUDY OF LAMINARIA SINCLAIRII AND L. LONGIPES  by  JAMES W. MARKHAM A.B., S t a n f o r d U n i v e r s i t y , S t a n f o r d , C a l i f o r n i a , 1961 M.S., U n i v e r s i t y o f Washington, S e a t t l e , Washington, 1963  A THESIS SUBMITTED I N PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY  i n t h e Department of BOTANY  We a c c e p t t h i s t h e s i s as c o n f o r m i n g t o t h e required standard  THE UNIVERSITY OF BRITISH COLUMBIA JUNE, 1969  In  presenting  an  advanced  the I  Library  further  for  this  thesis  degree shall  agree  scholarly  in partial  fulfilment  of the requirements f o r  a t the U n i v e r s i t y  of British  Columbia,  make that  permission  purposes  by  his representatives.  of  this  written  thesis  i t freely  may  be g r a n t e d  gain  of  Tetany  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, C a n a d a  June 10,  1969  Columbia  copying  b y t h e Head  It i s understood  for financial  f o r reference  f o r extensive  permission.  Department  Date  available  shall  that  n o t be a l l o w e d  and  thesis  Department or  that  Study.  of this  o f my  copying  I agree  or  publication  without  my  ABSTRACT  Laminaria s i n c l a i r i i  (Harvey) F a r l o w , Anderson and E a t o n ,  found from S o u t h e r n C a l i f o r n i a t o C e n t r a l B r i t i s h  Columbia,  and L ^ l o n g i p e s Bory, found from S o u t h e a s t A l a s k a t o t h e K u r i l e I s l a n d s , d i f f e r i n s e v e r a l ways from most o t h e r k e l p plants.  T h e i r most d i s t i n c t i v e f e a t u r e i s t h e r h i z o m e - l i k e  h o l d f a s t , composed o f many h a p t e r a , from w h i c h a r i s e m u l t i p l e s t i p e s , each b e a r i n g a s i n g l e b l a d e .  The two s p e c i e s a r e  v e r y s i m i l a r t o each o t h e r and have been d i s t i n g u i s h e d i n t h e p a s t p r i m a r i l y by t h e p r e s e n c e o f m u c i l a g e d u c t s i n t h e s t i p e of L. s i n c l a i r i i and t h e absence o f t h e s e i n t h e s t i p e o f L. l o n g i p e s . are  I n o r d e r t o d e t e r m i n e whether  t h e two s p e c i e s  i n d e e d d i s t i n c t , t h e i r d i s t r i b u t i o n , e c o l o g y , growth,  and r e p r o d u c t i o n were s t u d i e d i n t h e l a b o r a t o r y and on beaches i n A l a s k a , B r i t i s h Columbia, and Oregon. The g r o s s d i s t r i b u t i o n o f b o t h s p e c i e s appears t o be c o n t r o l l e d by temperature.  T r a n s p l a n t s and l a b o r a t o r y  c u l t u r e s i n d i c a t e t h a t L. l o n g i p e s i s adapted t o lower t e m p e r a t u r e s t h a n L. s i n c l a i r i i . little  S a l i n i t y a p p a r e n t l y has  i n f l u e n c e on d i s t r i b u t i o n , as b o t h s p e c i e s t o l e r a t e  wide ranges o f s a l i n i t y . L. s i n c l a i r i i was s t u d i e d in_ s i t u on t h r e e beaches i n N o r t h e r n Oregon, where t h e p l a n t s a r e s u b j e c t e d t o heavy  surf.  The sand l e v e l on t h e beaches r i s e s through t h e summer so t h a t t h e p l a n t s a r e p a r t l y o r w h o l l y b u r i e d under sand by l a t e summer.  The f i r s t heavy storms i n t h e f a l l remove most o f t h e  sand.  Maximum growth o c c u r s i n e a r l y summer, p r i o r t o  burial.  The b l a d e s a r e l o s t i n December and new ones a r e  regenerated  i n January.  R i p e s o r i a r e produced on t h e o l d  b l a d e s j u s t b e f o r e they a r e l o s t and on t h e new b l a d e s a f t e r they appear.  There i s l i t t l e e v i d e n c e  just  from e i t h e r  field  or l a b o r a t o r y s t u d i e s t o i n d i c a t e t h a t t h e gametophytes w h i c h develop  from t h e spores i n these s o r i n o r m a l l y produce  sporophytes.  S e x u a l r e p r o d u c t i o n o f t h i s type i s d i f f i c u l t  because o f t h e s c o u r i n g a c t i o n o f t h e sand.  I n March and  A p r i l 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 new s t i p e s and b l a d e s from t h e h a p t e r a a t t h e m a r g i n s o f t h e h o l d f a s t .  This  v e g e t a t i v e p r o l i f e r a t i o n i s a p p a r e n t l y t h e normal method o f reproduction. L. l o n g i p e s was o b s e r v e d occasions. i n December.  i n s i t u i n A l a s k a on o n l y f i v e  Growth i s g r e a t e s t i n summer and s o r i a r e produced Laboratory  cultures i n d i c a t e that sexual  reproduction i s very rare i n t h i s species.  L . longipes i s  r a r e l y a s s o c i a t e d w i t h sand. T r a n s p l a n t s and l a b o r a t o r y c u l t u r e s i n d i c a t e t h a t production o f mucilage  d u c t s i n t h e s t i p e s o f t h e two s p e c i e s  i s n o t a f f e c t e d b y changes i n e n v i r o n m e n t a l c o n d i t i o n s . Comparison o f t h e two s p e c i e s shows they d i f f e r i n s e v e r a l other p o i n t s besides mucilage  ducts, i n c l u d i n g length of  s t i p e s , w i d t h o f b l a d e s , w i n t e r l o s s o f b l a d e s , morphology o f gametophytes, and h a b i t a t .  The e v i d e n c e  s h o u l d be r e t a i n e d as two s e p a r a t e s p e c i e s .  c o n f i r m s t h a t they  iv TABLE OF CONTENTS PAGE ABSTRACT  i i  LIST OF TABLES  v  LIST OF FIGURES I. II. III.  IV.  V.  i i i  x  INTRODUCTION  1  TERMINOLOGY  7  GENERAL MATERIALS AND METHODS  8  A.  G e n e r a l F i e l d Methods  8  B.  C o l l e c t i o n , Transport  C.  Tagging and Growth Measurements  D.  Laboratory  and T r a n s p l a n t i n g  culture  8 9 11  DISTRIBUTION AND AUTEOOLOGY OF LAMINARIA SINCLAIRII  13  A.  Geographical  13  B.  H a b i t a t and Autocology on Oregon beaches  15  1.  Environment  15  2.  Occurrence o f L. s i n c l a i r i i  31  3.  Associated plant species  33  4.  Seasonal c y c l e s  37  5.  Discussion  40  Distribution  DISTRIBUTION AND AUTE OOLOGY OF LAMINARIA LONGIPES  42  A.  Geographical  42  B.  H a b i t a t and Autecology Coronation I s l a n d  Distribution a t A a t s Bay,  43  1.  Environment  43  2.  Occurrence o f L. l o n g i p e s and a s s o c i a t e d species  46  3.  Growth and r e p r o d u c t i o n o f L. l o n g i p e s a t A a t s Bay  47  Discussion  48  4.  V PAGE VI.  EXPERIMENTAL ECOLOGY  50  A.  F i e l d Work  50  1.  Transplants  50  a. b. c.  50 52  d. 2.  3.  B.  A l a s k a beaches Oregon beaches B r i t i s h Columbia beaches i. Whiffen S p i t ii. R i v e r Jordan iii. S t a n l e y park Discussion  Rock c l e a r i n g  59  a. b. c. d.  59 59 59 62  Introduction Methods Results Discussion  T r a n s i t i o n zone experiments  63  a. b. c.  63 63 65  Introduction Procedures and r e s u l t s Discussion  Laboratory 1.  2.  3.  54 55 56 57  Work  66  Sporophytes  66  a. b. c. do  66 67 67 69  Photoperiod Salinity Temperature Discussion  Gametophytes  69  a. b. c. do  69 70 71 72  introduction M a t e r i a l s and Methods Results Discussion  Haptera  73  a. b. c.  73 74 78 79 80 80  Introduction M a t e r i a l s and Methods Results i. Temperature ii. Light intensity iii. i n t i a l Size  vi PAGE VII.  TAXONOMY OF L . S I N C L A I R I I AND L . L O N G I P E S  84  A.  Introduction  84  B.  L. s i n c l a i r i i  85  1.  Description  85  2.  Distribution  86  3.  Habitat  86  4.  Comments  86  C.  D. VIII. IX. X. XI. XII.  L. l o n g i p e s  87  1.  Description  88  2.  Distribution  88  3.  Habitat  88  4.  Comments  89  Comparison  of Species  89  GENERAL D I S C U S S I O N AND CONCLUSIONS  93  SUMMARY  101  BIBLIOGRAPHY  104  TABLES I - X X I I I  111  APPENDIX I .  Summary D e s c r i p t i o n Stations  of Field 151  vii  LIST OF TABLES TABLE I  II III IV V  VI VII VIII IX X XI XII XIII XXV XV XVI XVII XVIII  PAGE Seawater temperature and s a l i n i t y over d i s t r i b u t i o n ranges o f L. s i n c l a i r i i and L_. l o n g i p e s .  112  D i s t r i b u t i o n of Laminaria s i n c l a i r i i .  114  D i s t r i b u t i o n of Laminaria longipes.  116  A n a l y s i s o f sand g r a i n s i z e on Oregon beaches.  118  Temperature  and p r e c i p i t a t i o n a t A l a s k a  stations.  119  Mean temperature on Urup I s l a n d .  120  Temperature and p r e c i p i t a t i o n a t B r i t i s h Columbia s t a t i o n s . Temperature and p r e c i p i t a t i o n a t Oregon stations.  123  Long term temperature and p r e c i p i t a t i o n a t two California stations.  124  Measured v a l u e s o f seawater temperature and s a l i n i t y on Oregon beaches.  125  C a l c u l a t e d mean monthly seawater on Oregon beaches.  126  121  temperatures  Mean v a l u e s o f seawater temperature and s a l i n i t y a t A r c h Cape f o r 1960-1963.  127  Sand l e v e l a t I n d i a n Beach.  128  Sand l e v e l a t S h o r t Sand Beach.  129  Sand h e i g h t s o f beach a t A r c h Cape, Oregon.  13 0  Associated plant species.  131  S e a s o n a l d i s t r i b u t i o n o f p l a n t s p e c i e s a t Indian Beach.  136  Seasonal d i s t r i b u t i o n o f p l a n t s p e c i e s a t Short Sand Beach.  139  viii  TABLE XIX XX XXI XXII XXIII  PAGE Seasonal d i s t r i b u t i o n o f a l g a l species a t A a t s Bay, C o r o n a t i o n I s l a n d .  142  Summary o f s e a s o n a l c y c l e s on Oregon beaches.  145  Summaries o f i n s i t u growth measurements on Oregon beaches.  146  Growth o f m u l t i - p u n c h e d b l a d e o f L. s i n c l a i r i i i n s i t u a t I n d i a n Beach.  148  Dimensions o f p r e s s e d specimens o f L. l o n g i p e s from 33 A l a s k a n s i t e s . 149  ix  LIST OF FIGURES FIGURE 1 2 3 4  PAGE D i s t r i b u t i o n ranges o f Laminaria and L. l o n g i p e s .  sinclairii 2  Mean seawater t e m p e r a t u r e s over ranges o f L. l o n g i p e s and L. s i n c l a i r i i  4  Mean a i r t e m p e r a t u r e s over ranges o f L. l o n g i p e s and L. s i n c l a i r i i .  5  Emergence and submergence o f v e r t i c a l d i s t r i b u t i o n extremes o f L. s i n c l a i r i i d u r i n g p e r i o d from June 1966 t o September 1967.  14  S i t e s i n Oregon where L_. s i n c l a i r i i was c o l l e c t e d or s t u d i e d .  18  A e r i a l and g e n e r a l views o f I n d i a n Beach and S h o r t Sand Beach.  23  7  I n d i a n Beach s t u d y a r e a .  24  8  S h o r t Sand Beach s t u d y  26  9  V a r i a t i o n i n sand l e v e l s a t I n d i a n Beach i n 1967,  27  10  V a r i a t i o n i n sand l e v e l a t s t u d y r o c k s a t I n d i a n Beach i n 1967.  28  11  V a r i a t i o n i n sand and water l e v e l s a t S h o r t Sand Beach i n 1967.  30  12  L. s i n c l a i r i i and a s s o c i a t e d s p e c i e s a t I n d i a n Beach.  34  13  Submergence i n f r e s h w a t e r and a s s o c i a t i o n o f L. s i n c l a i r i i w i t h o t h e r s p e c i e s a t S h o r t Sand Beach.  36  14  A a t s Bay s t u d y a r e a .  44  15  Transplant stations.  51  16  C l e a r e d r o c k a t I n d i a n Beach.  61  17  R e s u l t s o f growth e x p e r i m e n t s on h a p t e r a o f L. s i n c l a i r i i .  77  H a b i t o f L. s i n c l a i r i i and L. l o n g i p e s .  91  5 6  18  area.  ACKNOWLEDGEMENTS  I wish to express my g r a t i t u d e and a p p r e c i a t i o n to Dr. R. F. S c a g e l f o r h i s d i r e c t i o n , a d v i c e , and c r i t i c i s m throughout  the course o f t h i s study.  I a l s o wish to expres  my a p p r e c i a t e t o Dr. G. C. Hughes, Dr. K. Cole, and Dr. G.L. p i c k a r d f o r t h e i r a d v i c e throughout  the study; to  Dr. G.H.N. Towers f o r a d v i c e on t h i s manuscript; to Mr. Stephen Borden f o r programming the computer a n a l y s e s ; to Mr. J . Thorpe f o r a s s i s t a n c e w i t h v a r i o u s k i n d s o f equipment; to Mr. W.A.  Markham f o r topographic surveys o f  the Oregon beaches; t o L t . C o l . M.R. Mrs. W.A. to  Simmonds and  Markham f o r making p r e c i p i t a t i o n data a v a i l a b l e ;  the Oregon S t a t e park Department f o r a l l o w i n g access to  I n d i a n Beach a f t e r hours;  to the Department o f Botany and  I n s t i t u t e o f Oceanography, U n i v e r s i t y o f B r i t i s h Columbia for  facilities,  equipment, and a s s i s t a n c e ; and f i n a l l y to  the many people who a s s i s t e d me on v a r i o u s beaches, e s p e c i a l l y a t n i g h t i n the w i n t e r . w i t h thanks  I wish to acknowledge  the support p r o v i d e d by f e l l o w s h i p s from the  U n i v e r s i t y o f B r i t i s h Columbia and the N a t i o n a l Research C o u n c i l o f Canada.  1  I.  INTRODUCTION  L a m i n a r i a Laraouroux, the most common k e l p genus o f n o r t h temperate w a t e r s ,  u s u a l l y has a v e r y s i m p l e morphology.  It  c o n s i s t s o f a h o l d f a s t from w h i c h a r i s e s a s i n g l e s t i p e w i t h a f l a t t e n e d l a m i n a , or b l a d e .  In t h r e e v e r y u n u s u a l  species  the h o l d f a s t , composed o f many b r a n c h e d h a p t e r a , i s expanded i n t o a r h i z o m e - l i k e o r g a n from w h i c h a r i s e many s t i p e s , each bearing a s i n g l e blade. Bornet,  One  o f these s p e c i e s , L. r o d r i g u e z i i  o c c u r s a t depths o f 100 t o 150 m i n the  and A d r i a t i c Seas (Bornet, 1 8 8 8 ) .  Mediterranean  The o t h e r two a r e f o u n d i n  the i n t e r t i d a l and s u b t i d a l zones o f the n o r t h p a c i f i c Ocean. L_. l o n g i p e s Bory i s found from the K u r i l e I s l a n d s through A l e u t i a n I s l a n d s and L. s i n c l a i r i i  the  the G u l f o f A l a s k a i n t o S o u t h e a s t A l a s k a .  (Harvey) F a r l o w , Anderson and E a t o n o c c u r s  c e n t r a l B r i t i s h Columbia t o S o u t h e r n C a l i f o r n i a no m a t e r i a l o f L_. r o d r i g u e z i i was  from  (Fig. 1).  a v a i l a b l e , the two  Since  pacific  s p e c i e s o n l y a r e the s u b j e c t o f t h i s study. There a r e s e v e r a l r e a s o n s why L. l o n g i p e s seemed w a r r a n t e d .  a s t u d y o f L. s i n c l a i r i i  and  The p r e s e n c e o f m u l t i p l e s t i p e s  s u g g e s t e d t h a t the growth o f these p l a n t s might d i f f e r somewhat from t h a t o f o t h e r k e l p s . p o r t i o n s o f L. s i n c l a i r i i  I t had been o b s e r v e d t h a t the b a s a l a r e o f t e n b u r i e d under sand; as  few  p l a n t s can w i t h s t a n d such b u r i a l , t h i s s u g g e s t e d t h a t the e c o l o g i c a l a d a p t a t i o n s o f t h i s p l a n t might be somewhat u n u s u a l . F i n a l l y , most taxonomic t r e a t m e n t s  have s e p a r a t e d these  two  s i m i l a r s p e c i e s by the p r e s e n c e o f m u c i l a g e d u c t s i n the s t i p e s  b  Figure  1  Distribution and  L.  ranges of Laminaria  longipes.  Sinclair  3  of  L. s i n c l a i r i i and t h e i r absence i n the s t i p e s o f L, l o n g i p e s .  Burrows  (1964) has shown t h a t presence o r absence o f m u c i l a g e  d u c t s i n t h e b l a d e o f L. s a c c h a r i n a (L.) Lamour. can be c o n t r o l l e d by t e m p e r a t u r e .  I n her s t u d i e s , p l a n t s grown a t  10°C  d e v e l o p e d m u c i l a g e d u c t s , whereas t h o s e grown a t 5°C d i d  not.  T h i s i s o f i n t e r e s t because the average temperature o f  the  seawater i n A l a s k a and the K u r i l e I s l a n d s i s m a r k e d l y  lower than t h a t o f the seawater i n the a r e a from B r i t i s h Columbia t o C a l i f o r n i a  (Fig. 2).  The d i f f e r e n c e i n a i r  t e m p e r a t u r e i s even g r e a t e r ( F i g . 3 ) .  In view of t h i s , i t  appeared t h a t t h e p r e s e n c e o r absence o f m u c i l a g e d u c t s i n the  two s p e c i e s m i g h t be m e r e l y a r e s p o n s e t o the  and m i g h t be a l t e r e d by c h a n g i n g t h e environment.  environment i f this  were the c a s e , t h e p r e s e n c e o r absence o f m u c i l a g e d u c t s would n o t be a s u f f i c i e n t c r i t e r i o n f o r s e p a r a t i n g the two species. m i g h t be h.-  I t seemed p o s s i b l e , t h e r e f o r e , t h a t the two s p e c i e s one. s i n c l a i r i i and L. l o n g i p e s have been d e s c r i b e d by  s e v e r a l a u t h o r s , most r e c e n t l y by D r u e h l (1968) as p a r t o f a g e n e r a l taxonomic t r e a t m e n t o f the genus. e x p e r i m e n t a l work has been done w i t h them.  However, v e r y l i t t l e Setchell  (1905)  s t u d i e d growth and r e g e n e r a t i o n i n the b l a d e and s t i p e o f L. s i n c l a i r i i and Myers (1925) c u l t u r e d the gametophytes o f L. s i n c l a i r i i . done on L.  A p p a r e n t l y no e x p e r i m e n t a l work has ever been  longipes.  This i n v e s t i g a t i o n consists of three parts.  The f i r s t i s  an e x a m i n a t i o n o f the d i s t r i b u t i o n o f the two s p e c i e s throughout  ure  2  Mean seawater temperatures over ranges o f L. l o n g i p e s and L. s i n c l a i r i i .  P o r t Hueneme i s i n Ventura County,  California,  very near the southern l i m i t o f d i s t r i b u t i o n o f L. s i n c l a i r i i .  S i t k a i s the n e a r e s t p o i n t to  C o r o n a t i o n I s l a n d f o r which there a r e long range seawater temperature d a t a .  Note l a c k o f o v e r l a p i n  w i n t e r and annual means f o r the two ranges.  20  C O L D E S T  18  W A R M E S T  MONTH  A N N U A L  M O N T H  M E A N  L. sin c. Port Huenerne  15 I  L. long. Sitkqi  4 L. s i n c.  13 I2  Port Hueneme  L. s i n e .  Short Sand Beach  Port Huenemec  Pescadero Pis Urup  10  [ope Island  9 Sitka  8  ^ ^ H o p IS.  Hope Island ^ 1  7  5 4  Attu  Urup  L. long. Sitka  m Urup  Mean S e a w a t e r of L. Ion g i p e s  Temperatures over & L. s i n c l a i r i i  Ranges  gure 3  Mean A i r t e m p e r a t u r e s over ranges o f L_. l o n g i p e and L. s i n c l a i r i i .  Oxnard i s i n V e n t u r a County, C a l i f o r n i a .  Bull  Harbour i s on Hope I s l a n d , a t t h e n o r t h end o f Vancouver overlap  I s l a n d , B r i t i s h Columbia.  i n winter,  two r a n g e s .  Note l a c k o f  summer, and a n n u a l means f o r t h e  20  °C  19  COLDEST MONTH  WARMEST MONTH L. sine.  -6  JUrup  u  Mean  Air  L. l o n g i p e s  Oxnqrd  Is.  Temperatures &  ANNUAL MEAN  L.  over  sinclairii  Ranges  of  6  t h e i r r a n g e s i n r e l a t i o n to o c e a n o g r a p h i c c o n d i t i o n s , t o g e t h e r w i t h a more d e t a i l e d p r e s e n t a t i o n o f a u t e c o l o g i c a l s t u d i e s on f o u r b e a c h e s . Coronation  L. l o n g i p e s was  studied at Aats  I s l a n d , A l a s k a d u r i n g f i v e v i s i t s i n 1965  L. s i n c l a i r i i was  and  Bay, 1966.  s t u d i e d d u r i n g r e g u l a r v i s i t s i n 1965-1967  on t h r e e beaches i n N o r t h e r n Oregon:  I n d i a n Beach, A r c h Cape,  and S h o r t Sand Beach. I n the second p a r t , growth and r e p r o d u c t i o n o f b o t h s p e c i e s under e x p e r i m e n t a l s p o r o p h y t e s was  c o n d i t i o n s are t r e a t e d .  Growth o f  f o l l o w e d i n the f i e l d and i n tanks o f seawater  i n the l a b o r a t o r y under v a r i o u s c o n d i t i o n s .  Reciprocal  t r a n s p l a n t e x p e r i m e n t s were c a r r i e d o u t i n an a t t e m p t t o  assess  the i n f l u e n c e o f environment on the morphology and anatomy o f the p l a n t s . reproduction.  Gametophytes were c u l t u r e d i n a study o f The  L. s i n c l a i r i i was  sexual  growth o f i s o l a t e d p i e c e s o f h a p t e r a  of  also investigated i n laboratory culture.  I n t h e t h i r d p a r t o f the i n v e s t i g a t i o n , the v a l i d i t y the two s p e c i e s as s e p a r a t e  e n t i t i e s i s examined i n l i g h t o f  the i n f o r m a t i o n o b t a i n e d from f i e l d o b s e r v a t i o n s and f i e l d  experiments.  of  and  laboratory  7  II.  TERMINOLOGY  Each o f t h e two s p e c i e s  s t u d i e d has a h o l d f a s t o f b r a n c h e d  h a p t e r a w h i c h b e a r s numerous s t i p e s , each s t i p e i n t u r n a blade, or lamina. confusion  bearing  Because o f t h e m u l t i p l e s t i p e s ,  may a r i s e as t o what c o n s t i t u t e s an i n d i v i d u a l p l a n t .  I n t h i s s t u d y , t h e terms " p l a n t " and "clump" a r e used t o r e f e r t o t h e h o l d f a s t w i t h a l l i t s s t i p e s and t h e i r b l a d e s .  When  an i n d i v i d u a l s t i p e o r b l a d e i s r e f e r r e d t o , i t i s d e s i g n a t e d " s t i p e " or "blade".  8  III.  GENERAL MATERIALS AND  A. The  G e n e r a l F i e l d Methods  f l u c t u a t i o n i n sand h e i g h t a t c e r t a i n l o c a t i o n s on  Oregon beaches rocky  r e f e r e n c e p o i n t s was  two beaches  two  (Indian Beach and Short Sand Beach) r e l a t i v e to measured each month by means o f a  pocket r u l e and an Abney l e v e l .  A d e t a i l e d map  o f each o f  ( F i g . 7, 8) showing the l o c a t i o n of a l l r o c k s  L. s i n c l a i r i i as w e l l as i t s upper and prepared u s i n g a surveyor's 1130  METHODS  computer was  lower l i m i t s ,  t r a n s i t and  the with  was  stadia rod.  An  IBM  used to c a l c u l a t e the t o t a l time i n any  given  month t h a t p l a n t s a t v a r i o u s h e i g h t s were out o f water or under water i n d a y l i g h t and t i d e s and  i n darkness, based on p u b l i s h e d data f o r  hours o f d a y l i g h t  B.  ( F i g . 4)  Collection, Transport,  and  Transplanting  P l a n t s were c o l l e c t e d i n the f i e l d by c u t t i n g the l o o s e from the r o c k s w i t h a k n i f e .  In most i n s t a n c e s ,  b e a r i n g more than 40 s t i p e s were separated parts. ice  The  i n t o two  p l a n t s were then t r a n s p o r t e d i n wet  holdfasts holdfasts  or more  newspaper on  i n a f r e e z e r chest to h o l d i n g tanks a t the U n i v e r s i t y o f  B r i t i s h Columbia or d i r e c t l y , when p o s s i b l e , to another beach in  t r a n s p l a n t experiments.  more than 36 hours and treatment.  The  p l a n t s were never out o f water  showed no apparent damage from t h i s  In the case o f t r a n s p l a n t s between A l a s k a  and  Vancouver the p l a n t s were k e p t i n l a r g e h o l d i n g tanks i n the shade on  the deck o f the s h i p en r o u t e .  The  water i n these  9  t a n k s was changed e v e r y two t o f i v e days and never r e a c h e d more than 10°C, due t o t h e l o w ambient a i r t e m p e r a t u r e . In t h e t r a n s p l a n t e x p e r i m e n t s , two methods were employed f o r a t t a c h i n g the p l a n t s i n the f i e l d .  The f i r s t , used i n  a r e a s w i t h l i t t l e s u r f and many l o o s e b u t s t a b l e b o u l d e r s (Volga i s l a n d , R i v e r J o r d a n , W h i f f e n S p i t , S t a n l e y  park)  ( F i g . 6) c o n s i s t e d o f p l a c i n g two l a r g e r u b b e r bands around a b o u l d e r and t h e h o l d f a s t .  I n t h e second method, employed  i n a r e a s w i t h g r e a t e r s u r f and l a c k i n g l o o s e b o u l d e r s Bay,  (Aats  I n d i a n Beach, S h o r t Sand Beach), two o r t h r e e s p i k e s were  d r i v e n i n t o t h e b e d r o c k and t h e h o l d f a s t s were a t t a c h e d t o t h e s e w i t h s e v e r a l r u b b e r bands. C.  T a g g i n g and Growth Measurements  I n t h e f i e l d and i n l a b o r a t o r y t a n k s , p l a n t s were i d e n t i f i e d b y a p i e c e o f p l a s t i c f l a g g i n g tape t i e d around a p o r t i o n o f the h o l d f a s t .  I n d i v i d u a l s t i p e s were i d e n t i f i e d by  a p i e c e o f f l a g g i n g tape t i e d around t h e s t i p e . g r o w i n g i n s i t u , e a c h s t i p e was c o n s i d e r e d  For plants  separately.  For  t r a n s p l a n t s i n the f i e l d or i n the l a b o r a t o r y , three s t i p e s , designated plant.  A, B, and C were s e l e c t e d f o r growth s t u d y on each  A was t h e s h o r t e s t s t i p e (and perhaps t h e y o u n g e s t ) ,  C was t h e l o n g e s t , and B was a s t i p e o f i n t e r m e d i a t e  length,  u s u a l l y c l o s e r to C than to A i n length. In o r d e r  t o d e t e r m i n e t h e growth more a c c u r a t e l y , a s m a l l  h o l e was punched 10 cm above t h e base o f each b l a d e measured, using a cork borer  and an a p p a r a t u s l i k e t h a t used by Sundene  10  (1964), c o n s i s t i n g o f a c e n t i m e t e r r u l e w i t h a n o t c h e d  piece  o f m e t a l f a s t e n e d t o t h e lower end and a h o l e a t t h e 10 cm mark.  The n o t c h was p l a c e d over t h e s t i p e a t t h e t r a n s i t i o n  between s t i p e and b l a d e and t h e b l a d e was punched through t h e hole i n the r u l e .  A new h o l e was punched i n t h e same manner  e v e r y time a measurement was made and t h r e e f i g u r e s were recorded:  then  l e n g t h o f s t i p e , l e n g t h o f b l a d e , and d i s t a n c e  between t h e l a s t two h o l e s .  I n l a b o r a t o r y c u l t u r e s and f i e l d  t r a n s p l a n t s , t h e w i d t h o f t h e b l a d e a t 5 cm above t h e base was a l s o r e c o r d e d .  However, i n p l a n t s growing  measurement was n o t made a f t e r i t was o b s e r v e d was r a r e l y  i n situ  this  t h a t a change  recorded.  I t was found t h a t t y i n g f l a g g i n g tape around s t i p e s i n the f i e l d sometimes i n j u r e d them, e s p e c i a l l y those i n a r e a s s u b j e c t e d t o heavy s u r f .  I n l a t e r f i e l d measurements made  i n s i t u no i d e n t i f y i n g markers were used a t a l l , a punched h o l e b e i n g t h e o n l y i n d i c a t i o n t h a t a b l a d e and i t s s t i p e had been measured p r e v i o u s l y .  Thus, s t i p e s c o u l d n o t be i d e n t i f i e d  as i n d i v i d u a l s b u t o n l y a s members o f a p a r t i c u l a r group.  How-  e v e r , t h e d i s t a n c e between punched h o l e s p r o v i d e d an a b s o l u t e measure o f g r o w t h o f one p a r t o f t h e p l a n t even i f t h e s t i p e and i t s b l a d e c o u l d n o t be i d e n t i f i e d as a p a r t i c u l a r i n d i v i d u a l i n t h e p r e v i o u s month's d a t a .  A t the beginning o f  each s e r i e s o f measurements 20 t o 25 b l a d e s were punched w i t h i n a s m a l l area  ( c a . 20x20 cm) on a r o c k and i n s u c c e e d i n g  months o n l y t h o s e f o u n d t o have h o l e s were punched and  11  measured a g a i n .  T h i s c o n t i n u e d u n t i l too few  (usually- l e s s  than 10) p r e v i o u s l y punched b l a d e s were found, a t w h i c h time new b l a d e s were punched.  I n w i n t e r , when no b l a d e s a r e  p r e s e n t , a few s t i p e s were tagged t o i n d i c a t e a d e f i n i t e a r e a on the r o c k and then 2 0 t o 25 s t i p e s were measured i n t h i s small area. D.  Laboratory Culture /<  F o r l a b o r a t o r y c u l t u r e work,two New B r u n s w i c k p s y c r o t h e r m I n c u b a t o r Shakers and t h r e e w a l k - i n c o n t r o l l e d rooms were employed.  temperature  The P s y c r o t h e r m s were k e p t a t  8°C.  V a r i o u s l i g h t p e r i o d s were employed u s i n g c o o l w h i t e f l u o r e s c e n t tubes w i t h an i n t e n s i t y o f 150 f t - c . and w i t h o u t s h a k i n g . were m a i n t a i n e d a t 5 ° ,  C u l t u r e s were grown w i t h  The t h r e e c o n t r o l l e d temperature rooms 8 ° , and 10°C.  I n each o f them two  light  i n t e n s i t i e s , a p p r o x i m a t e l y 20 f t - c ( 2 1 5 . 2 l u x ) and 150 f t - c (1614.0  l u x ) were employed.  I n each o f the t h r e e c o n t r o l l e d temperature rooms t h r e e 120  1 t a n k s o f seawater were used f o r c u l t u r i n g s p o r o p h y t e s .  Two o f t h e  t a n k s i n each room c o n t a i n e d seawater from Juan  de Fuca Auto C o u r t near O t t e r p o i n t on the west c o a s t o f Vancouver  I s l a n d , d e s i g n a t e d West Coast w a t e r .  was 30.9%o  t  0„5%„  The  salinity  depending on the month o f c o l l e c t i o n . The  t h i r d tank i n each room c o n t a i n e d water from S t a n l e y p a r k , Burrard I n l e t park water.  ( s a l i n i t y = 27.9%o  t  0.4% ) o  designated Stanley  The water was changed a t a p p r o x i m a t e l y monthly  i n t e r v a l s w i t h f r e s h l y c o l l e c t e d water.  A e r a t i o n was  12  p r o v i d e d by an aquarium b u b b l e r .  The t a n k s were i l l u m i n a t e d  by c o o l w h i t e f l u o r e s c e n t tubes w h i c h gave an i n t e n s i t y a t the s u r f a c e o f the water o f 150-200 f t - c (1614-2152 l u x ) . d e p t h o f the water was 35-40 cm.  The  p h o t o p e r i o d s r a n g i n g from  8 t o 16 hours were employed d u r i n g the c o u r s e o f the s t u d y . Gametophytes and p o r t i o n s o f h a p t e r a were c u l t u r e d i n 250 ml g l a s s c u l t u r e d i s h e s , s t a n d a r d 100 mm g l a s s P e t r i d i s h e s and 60 mm p l a s t i c P e t r i d i s h e s .  Four d i f f e r e n t seawater media  were employed, a l l b a s e d on West Coast seawater: (SW), seawater f i l t e r e d t h r o u g h c o t t o n wadding  raw seawater  (SWF),  E r d - S c h r e i b e r medium (ES) (F0yn, 1934), and e n r i c h e d E r d - S c h r e i b e r (ES+) i n w h i c h 1 ml o f "ASP  2" medium ( P r o v a s o l i , M c L a u g h l i n ,  and Droop, 1957) was added t o each l i t e r o f E r d - S c r e i b e r . a d d i t i o n , s t r a i g h t ASP cultures.  One mg/1  2 was employed i n a few  In  gametophyte  o f Ge02 was added t o each medium t o r e t a r d  the growth of diatoms.  The media were changed a t i n t e r v a l s  r a n g i n g from one day t o one month, depending on the e x p e r i m e n t .  13  IV.  D I S T R I B U T I O N A N D A U T E C O L O G Y OF L A M I N A R I A  A. L.  Geographic  sinclairii  Distribution  i s f o u n d f r o m Hope i s l a n d ,  (50 56'N, 127°58'W) t o V e n t u r a C o u n t y ,  Island  1).  The  near  Hope I s l a n d I ) .  annual seawater  31.7%  0  salinity  T h e mean a n n u a l s e a w a t e r  from n o r t h t o south, from  v a l u e s show an i n c r e a s e  temperatures range  P o r t Hueneme.  temperature  10°C a t H o p e i s l a n d  water also  Mean  t o 16.8°C between  o f L. l o n g i p e s .  As  more t h a n t h e y a r e under  ( F i g . 4), t h e a i r t e m p e r a t u r e a n d p r e c i p i t a t i o n a r e important considerations.  Tables V I I , VIII  average  temperatures and p r e c i p i t a t i o n  British  Columbia,  Oregon,  temperatures range Ventura these  range  thedifference  these conditions and those i n t h e range a r e o u t o f water  0  also  t o 13.2°C a t p o r t H u e n e m e . from  33.6%  8.6°C a t H o p e I s l a n d t o  Figure 2 illustrates  many o f t h e p l a n t s  t h e mean a t  Mean F e b r u a r y t e m p e r a t u r e s  f r o m 7.2°C a t H o p e I s l a n d  at  coast o f  a n d a t p o r t Hueneme, V e n t u r a C o u n t y ,  14.3°c a t p o r t H u e n e m e .  August  Hope  Island. mean  increases  (34°19'N,  Tofino on t h e southwest  from n o r t h t o south i n t h e plant's range, with  (Table  Columbia  I t has n o t been r e c o r d e d between  and Box I s l a n d ,  Vancouver  British  California  o  119°23.3'w) ( F i g .  SINCLAIRII  County.  conditions  from  f o r selected  and California.  a n d those i n t h e r a n g e  show  sites i n  T h e a n n u a l mean a i r  8.7°C a t H o p e I s l a n d  Figure 3 illustrates  and IX  t o 15.2°C i n  thedifferences o f L.  longipes.  between  14  Figure 4  Emergence and submergence of v e r t i c a l d i s t r i b u t i o n extremes of L. s i n c l a i r i i  during  period from June 1966 to September 1967.  Any v e r t i c a l l i n e drawn from top to bottom of each graph adds up to 100 percent of the hours i n that month.  "Dark" i s percentage of t o t a l hours  the point was i n darkness.  "Light" i s percentage  of t o t a l hours the point was i n daylight.  Line  between "wet" and "dry" divides time under water from time out of water.  6 T 8 9  10 II  LIGHT  1967  1966  12 I  I I I I 1 1 1I  2  3  I  4  I  5  6  7  6  I  I  1  I  3  DARK  Highe s t • 3.0 f t (•0.92m) D R Y  L. Sinclqirii Short  WET  Sand  Beach  Highest + 1.5 ft. D R Y (•0.46 m) WET  L. sinclairii Indian B e a c h  DRY 0 . 0 ft. (0.0 nt)  Mean  Tide  WET  DRY Lowest -1.5 ft. (-0.46 m)  L. s i n c l a i r i i WET Short  Sand  Beach  DRY Low e s t - 2 . 0 ft. (-0.6lm)  yVET  L. s i n c l a i r i i Indian  Beach  15  Table collected obtained  II lists or  observed.  University of  sinclairii  and  has  Columbia and  f o u n d on  lower  exposed  i n sand  i n the  They are  few  to  level.  intertidal  a l s o observed  in  British  Oregon are  s u r f and  have  They are  generally  the  those  a  i n more  southern  that plants than  L.  plants  zone and  a l s o l a r g e r near  I t was  own  places  the  of  limits  from  from  Oregon,  q u a n t i t a t i v e m e a s u r e m e n t s w e r e made.  H a b i t a t and  Autecology  on  Oregon  Beaches  Environment The  beaches Arch  autecology i n Northern Cape  Thus the  Short  f o r e s t and  disturbed behind an  almost  the beach.  of  L.  sinclairii  Oregon:  ( F i g . 5).  S t a t e p a r k and  is  author's  published records.  p r o d u c e much more m u c i l a g e  no  B.  the  exceptions,  fully  was  U n i v e r s i t y of  i s m u c h m o r e common i n  few  fluctuation  distribution.  although  and  been  p h y c o l o g i c a l Herbarium  Columbia;  beaches which are  California  and  British  with v e r y  exposed s i t e s .  1.  Berkeley; the  has  distribution  a v a i l a b l e i n the  Washington, but  when f o u n d  sinclairii  i n f o r m a t i o n on  been found at r e l a t i v e l y  marked seasonal  of  a t w h i c h L.  collections;  California,  larger  The  Herbarium,  observations  and  sites  from c o l l e c t i o n s  California the  the  Indian  Indian  s t u d i e d on  Beach,  Short  three  Sand  Beach i s l o c a t e d i n  Beach,  Ecola  Sand Beach i s i n Oswald West S t a t e the  land i n general  the beaches.  continuous South of  was  the  row  of  To  the  are  north  houses along  Cape t h e r e  are  very  relatively of Arch the few  land  park. little  Cape  there  bordering  houses  in  16  the  f i r s t k i l o m e t e r , p a r t l y because o f the v e r y s t e e p c l i f f s .  The f i r s t two beaches were v i s i t e d a t v a r i o u s t i m e s over the p e r i o d 1965-1967 and a t l e a s t once a month from August t h r o u g h September are  1967.  A t A r c h Cape L. s i n c l a i r i i  1966  plants  f o u n d i n an a r e a w h i c h i s v e r y d i f f i c u l t t o r e a c h e x c e p t  i n the c a l m e s t weather and a t the l o w e s t t i d e s o f the summer. The s t u d y s i t e here was v i s i t e d o n l y i n June, J u l y , August and September  1967, a l t h o u g h i t was o b s e r v e d from a d i s t a n c e on  many o t h e r o c c a s i o n s . On a l l t h r e e beaches t h e r e i s a r e g u l a r c y c l e o f r e m o v a l and d e p o s i t i o n o f sand ( F i g . . 9,10,11,16) . the  The sand l e v e l on  b e a c h r i s e s from A p r i l o r May u n t i l the f i r s t b i g storms  ( u s u a l l y i n September  o r October) remove most o f the sand i n  a v e r y s h o r t t i m e , sometimes  i n as l i t t l e as 24 h o u r s .  The  sand forms o f f s h o r e b a r s d u r i n g t h e w i n t e r and then i s t r a n s p o r t e d t o t h e b e a c h a g a i n t h e f o l l o w i n g summer.  The s u p p l y  i s c o n t i n u a l l y augmented by sand w h i c h i s b r o u g h t t o the sea by n e a r b y streams and r i v e r s . Sand samples f o r each beach were a n a l y z e d f o r g r a i n using a standard set of Endecott sieves are  v e r y s i m i l a r f o r a l l beaches.  well sorted.  (Table I V ) .  The  size, results  The sand i s v e r y c l e a n and  T h i s , and t h e s i z e d i s t r i b u t i o n r e f l e c t the  g r e a t wave a c t i o n on the beaches.  Over 97 p e r c e n t o f the sand  on each beach i s f i n e g r a i n e d (0.25 -0.1 mm diam} t o v e r y f i n e grained  (0.1 - 0.05 mm d i a m . ) .  S p e c t r o s c o p i c a n a l y s i s shows  t h a t the sand i s p r i m a r i l y composed o f q u a r t z .  Most o f i t i s  d e r i v e d from metamorphic r o c k s w i t h a s m a l l e r p e r c e n t a g e o f  17  granitic the  material.  Columbia River  Pers.  (W.R.  Danner,  (1963) n o t e d t h a t  t h e summer o f t e n  the  source of the sand i s very Dept. o f Geology,  varies  previous winter.  Oregon  beaches.  The  the supply  directly  T h i s was winter  with  sand  t h e b e a c h was  level  1967  than i n  A l l winter  on  storms the surf there  storms.  noticably  are i n f u l l y i s very  higher  of  beach both to the north  The  Arch  Cape i s a r o c k  the  During  calmest  to approach  Beach have  Oregon.  larger  Each of the  Cape i s a h e a d l a n d w i t h and  directly  the south,  a long  so t h a t  The  study  w e s t o f t h e Cape.  the other beaches are s i t u a t e d  of  the headlands.  At  I n d i a n Beach and S h o r t Sand Beach, L.  at  each end o f t h e beach.  These  of  a  E a c h i s e x p o s e d more t o t h e s o u t h than  t o waves f r o m a l l d i r e c t i o n s .  north  the  i n t h e summer  waves t e n d  on  the  and  exposed l o c a t i o n s .  i n northern  headland ( F i g .5).  the north.  Arch  the  s h o r t beach formed i n an i n d e n t a t i o n  to  exposed  (Table V I I I )  I n d i a n Beach and S h o r t Sand  i s a relatively  prominent  runoff  calm weather but from the southwest  waves t h a n a l l o t h e r beaches two  t r u e on  heavy and even on  i s some s u r f .  the northwest during  during  t h e amount o f  1966.  three beaches  summer d a y s  the beach  o f 1966-1967 had much more  than the previous winter  from  U.B.C,  o f sand on  observed to be  precipitation  of  likely  comm.). Shepard  in  The m a i n  i t i s site  The  fully  at  study  somewhat b a c k o f t h e  outermost p o i n t s were  The  stretch  sites  points  never reached.  sinclairii  areas of intensive  occurs  study  end o f I n d i a n Beach and the south end o f Short  were Sand  18  Figure 5  Sites  i n Oregon where L. s i n c l a i r i i  collected  was  or studied.  "A" and "B" near A r c h Cape a r e s i t e s where precipitation  was measured.  124°  0 0'  W  19  Beach.  Although  the Arch Indian  cape s i t e  Beach S i t e  The The  a l l t h e s i t e s must be considered  tides  tidal  and then  i n this  pattern  semidiurnal  i s slightly  with  highs of  two h i g h  a n d two l o w w a t e r s  Oregon.  for Arch  Cape.  to September mean h i g h e r water  tidal  location  data  -4.47  From t i d a l  data  (MHHW),  mean t i d e  taken  f t . (3.4m). tides  A l l heights  a s 0.0 f t .  to within  than  this  0.5 f t .  was  period  average heights  lowest  were  obtained (June 1966 obtained:  3.91 f t . (1.19m); mean  low water  are i n relation  Beach and Short  to correspond  t o these  i s  lower  low water (MLLW), t o long  Thus t h e a v e r a g e a m p l i t u d e  The extreme a m p l i t u d e  For Indian  appear  The  a r e data i s  (MLHW), 2 . 3 7 f t ( 0 . 7 2 m ) ; m e a n h i g h e r  a b o u t 8.3 f t . ( 2 . 5 m ) .  actual  there  f o r the study  the following  high water  f t (-1.36m).  successive  f o r any o f t h e beaches  f o rwhich  No i n f o r m a t i o n o t h e r  1967)  heights.  interval  There a r e  each day, and  (MHLW), - 1 . 8 1 f t . ( - 0 . 5 5 m ) ; m e a n l o w e r  l l  25 h o u r s .  an  i n winter.  The n e a r e s t  Astoria,  tide with  high  i n the daytime i n e a r l y morning i n  T h e r e a r e no p u b l i s h e d  high  between successive  o r lows a r e g e n e r a l l y o f d i f f e r e n t  studied.  type.  o f two component t i d e s , t h e  high waters o f about  summer a n d a t n i g h t  site.  a r e o f the mixed semidiurnal  an i n t e r v a l  the low waters occurs  exposed,  followed by the  Sand Beach  12% hours and the d i u r n a l  between successive usually  the Short  i s the result  tide,  waters o f about  area  more exposed,  fully  term  i s  approximately  Sand Beach t h e calculated  heights  20  T h i s a r e a i s c h a r a c t e r i z e d by r e l a t i v e l y heavy p r e c i p i t a t i o n and moderate t e m p e r a t u r e s .  The n e a r e s t p l a c e f o r w h i c h t h e r e  a r e p u b l i s h e d r e c o r d s o f p r e c i p i t a t i o n and a i r temperature Seaside, Oregon.(Pig.  5).  is  U n o f f i c i a l p r e c i p i t a t i o n records  have been k e p t by two i n d i v i d u a l s a t A r c h Cape ( F i g . 5 ) , one s i n c e 1957  (A), and one s i n c e 1965  (B).  The r a i n gauges a r e  s i t u a t e d about one k i l o m e t e r a p a r t , each atop a sea  cliff  about 20 m e a s t o f the b e a c h , and each near a house. n o r t h e r n one  (A) i s l o c a t e d a t a somewhat h i g h e r a l t i t u d e  n e a r e r t o a house  than the s o u t h e r n one  (B).  The  (1965-67) had b e l o w normal p r e c i p i t a t i o n .  and  records  f o r S e a s i d e and A r c h Cape (Table V I I I ) show t h a t the period  The more  The  study records  show more p r e c i p i t a t i o n a t s t a t i o n B than a t A and more a t each than a t S e a s i d e .  The maximum p r e c i p i t a t i o n u s u a l l y o c c u r s  i n December o r J a n u a r y and the minimum i n J u l y o r August. summer o f 1967  was p a r t i c u l a r l y d r y .  p r e c i p i t a t i o n f o r S e a s i d e i s 79.7  The mean t o t a l  The  annual  inches.  S e a s i d e i s s i t u a t e d n o r t h o f T i l l a m o o k Head, a l a r g e promontory, and I n d i a n Beach i s i n the s o u t h e r n p a r t o f T i l l a m o o k Head.  Thus, t h e r e may  be s i g n i f i c a n t d i f f e r e n c e s  i n m e t e o r o l o g i c a l c o n d i t i o n s a t the two a r e a s , p a r t i c u l a r l y i n precipitation.  However, as no o t h e r i n f o r m a t i o n i s a v a i l a b l e  from a n e a r e r l o c a t i o n , the S e a s i d e p r e c i p i t a t i o n r e c o r d s a r e p r e s e n t e d as an a p p r o x i m a t i o n o f the c o n d i t i o n s a t I n d i a n Beach. The r e c o r d s f o r S t a t i o n B a t A r c h Cape a r e p r e s e n t e d as  an  a p p r o x i m a t i o n o f p r e c i p i t a t i o n a t A r c h cape and S h o r t Sand Beach.  I n the absence o f any o t h e r r e c o r d s , the S e a s i d e a i r  21  t e m p e r a t u r e s a r e used f o r a l l t h r e e beaches.  The a n n u a l mean  a i r temperature i s 11.0°c (Table V I I I ) , r a n g i n g from a monthly mean o f 6.3°C i n J a n u a r y t o 15.7°C i n August.  The  y e a r 1967 d i f f e r s from t h e mean, e s p e c i a l l y i n t h e h i g h e r summer t e m p e r a t u r e s . There a r e p u b l i s h e d r e c o r d s o f seawater temperature and s a l i n i t y f o r A r c h cape (1949-1967).  (1960-1963) and t h e S e a s i d e Aquarium  I n a d d i t i o n , water samples were t a k e n a t I n d i a n  Beach and S h o r t Sand Beach once a month from October 1966 t h r o u g h September 1967 (Table X ) .  The samples were c o l l e c t e d  a t l o w t i d e i n t h e s u r f as near t o t h e s t u d y r o c k s as p o s s i b l e and t e m p e r a t u r e r e a d i n g s were t a k e n i m m e d i a t e l y .  The s a l i n i t y  d e t e r m i n a t i o n s were made w i t h an i n d u c t i v e s a l i n o m e t e r a t t h e I n s t i t u t e o f Oceanography,  U n i v e r s i t y o f B r i t i s h Columbia.  I n t h e A r c h cape r e c o r d s t h e r e i s c o n s i d e r a b l e v a r i a t i o n between t h e d i f f e r e n t y e a r s , p r o b a b l y p a r t l y because o f t h e g r e a t v a r i a t i o n i n number o f samples.  The mean monthly seawater  t e m p e r a t u r e r a n g e d from 7.4°c i n March 1962 t o 15.8°C i n J u l y 1963. The mean m o n t h l y s a l i n i t y r a n g e d from 26.9%„ i n A p r i l 1962 t o 32.9%  c  i n August 1961. The c o n d i t i o n s f o r 1967  a r e assumed t o be w i t h i n t h i s r a n g e .  Table X I I g i v e s the  monthly means f o r t h e p e r i o d 1960-1963. The m o n t h l y r e a d i n g s o b t a i n e d a t S h o r t Sand Beach and I n d i a n Beach were compared w i t h t h e r e c o r d s from S e a s i d e .  In  a l m o s t a l l i n s t a n c e s , when r e a d i n g s f o r s p e c i f i c d a t e s were compared, t h e S e a s i d e t e m p e r a t u r e was h i g h e r .  The mean  d i f f e r e n c e was 2.5C° a t I n d i a n Beach and 2.0C° a t S h o r t Sand  22  Beach.  These d i f f e r e n c e s were s u b t r a c t e d from t h e monthly  means f o r S e a s i d e f o r each month over t h e p e r i o d s t u d i e d t o o b t a i n an a p p r o x i m a t i o n o f t h e temperature beach (Table X I ) .  c o n d i t i o n s a t each  Assuming t h i s t o be v a l i d , a t I n d i a n Beach  the mean m o n t h l y seawater  temperature  ranged from 7.6°C i n  March t o 13.9°C i n J u l y , w i t h a y e a r l y mean o f 10.5°C, whereas a t S h o r t Sand Beach t h e range was from 7.1°C i n March t o 13.4°C i n J u l y , w i t h a y e a r l y mean o f 10.0°C. t y p i c a l t h i s y e a r may have been. temperature  I t i s n o t known how  Monthly means o f seawater  f o r S e a s i d e over t h e p e r i o d o f y e a r s  (1949-1967)  f o r w h i c h t h e r e a r e r e c o r d s a r e o f no use because t h e a n n u a l means have r i s e n over almost t h e e n t i r e p e r i o d .  T h i s may be  a t t r i b u t e d i n p a r t t o t h e s t e a d y w i d e n i n g o f t h e S e a s i d e beach due t o i n c r e a s e d sand d e p o s i t i o n w h i c h has o c c u r r e d s i n c e t h e c o n s t r u c t i o n o f t h e Columbia R i v e r j e t t y a p p r o x i m a t e l y 25 km north of Seaside. No i n t e r p o l a t i o n s a r e p o s s i b l e from a comparison  o f the  s a l i n i t y measurements a t t h e two s t u d y beaches and S e a s i d e . L o c a l freshwater r u n o f f o b v i o u s l y a f f e c t s the readings s i g n i f i c a n t l y a l t h o u g h i r r e g u l a r l y , e s p e c i a l l y a t S h o r t Sand Beach (Table X ) . The  s t u d y s i t e a t I n d i a n Beach i s an a r e a a d j a c e n t t o two  r o c k y p o i n t s a t t h e n o r t h end o f t h e beach, one a headland, the o t h e r a young s t a c k ( F i g 6, 7 ) . There a r e numerous r o c k s w h i c h e x t e n d above, t h e l e v e l o f t h e beach. is primarily olivine basalt.  Most o f t h e r o c k  A f r e s h w a t e r stream f l o w s a c r o s s  the beach j u s t s o u t h o f t h e study a r e a .  I t s course i s a l t e r e d  23  Figure 6  A e r i a l and g e n e r a l v i e w s o f I n d i a n Beach and S h o r t Sand Beach.  a.  A e r i a l v i e w o f n o r t h end o f I n d i a n Beach.  Arrow  shows a p p r o x i m a t e l i n e o f s i g h t i n f i g u r e b below. R e c t a n g l e i n d i c a t e s a r e a shown i n f i g u r e "7, b.  I n d i a n Beach a t h i g h t i d e i n A u g u s t 1967.  c.  A e r i a l v i e w o f s o u t h end o f S h o r t Sand Beach. A r r o w shows a p p r o x i m a t e l i n e o f s i g h t i n f i g u r e d below.  R e c t a n g l e i n d i c a t e s a r e a shown i n  f i g u r e 8. d.  S h o r t Sand Beach s h o r t l y a f t e r low t i d e i n March 1967.  24  Figure 7  I n d i a n Beach s t u d y a r e a .  L. s i n c l a i r i i o c c u r s on a l l r o c k s shown i n f i g u r e and a t s e v e r a l p o i n t s on the r o c k f o r m i n g the shore.  Growth measurements were made on "A" and "B".  "C" was p a r t l y c l e a r e d (see F i g . 1 6 ) . was measured a t p o i n t "F" and a t "A".  Sand h e i g h t  SCALE  25  w i t h each t i d a l c y c l e , b u t i t does n o t n o r m a l l y come i n c o n t a c t w i t h any o f t h e p l a n t s s t u d i e d .  The sand l e v e l was r e c o r d e d  a t monthly i n t e r v a l s from August 1966 t h r o u g h September 1967 a t f o u r p o i n t s on t h e b e a c h (Table X I I I ) .  The t o t a l  f l u c t u a t i o n i n sand l e v e l i s a t l e a s t 1.2 m and p r o b a b l y more on some a r e a s w h i c h were n o t measured.  On most o f I n d i a n Beach,  the sand l e v e l i s l o w e r e d i n w i n t e r b u t a sandy beach remains.  still  On t h e a r e a s t u d i e d however, a l l t h e sand i s removed  i n w i n t e r , e x p o s i n g t h e bedrock  ( F i g . 9a, 1 0 a ) .  Most o f t h e a r e a s t u d i e d a t I n d i a n Beach i s s i t u a t e d so t h a t i t i s s h i e l d e d from d i r e c t i n s o l a t i o n d u r i n g most o f t h e time t h e r o c k s a r e exposed a t l o w t i d e .  I t was o b s e r v e d  that  the r o c k s and t h e p l a n t s growing on them always r e t a i n e d enough water so as t o appear wet even i f t h e y had been exposed by t h e r e c e d i n g t i d e f o r as l o n g as 3 hours.  The o n l y  e x c e p t i o n t o t h i s o c c u r r e d d u r i n g an e x t r e m e l y d r y , h o t p e r i o d i n A u g u s t 1966 when t h e r e l a t i v e h u m i d i t y o f t h e a i r f e l l b e l o w 2 0 % and t h e a i r temperature  r e a c h e d 30°C.  The r o c k s  appeared c o m p l e t e l y d r y and some o f t h e p l a n t s were c u r l i n g , a l m o s t as i f b u r n e d . The s t u d y s i t e a t S h o r t Sand Beach i s a s e r i e s o f r o c k y o u t c r o p s a l o n g t h e n a r r o w m a r g i n between t h e s o u t h e r n end o f the sandy b e a c h and t h e s t e e p c l i f f s w h i c h form t h e s o u t h e r n boundary o f t h e cove i n w h i c h t h e b e a c h i s l o c a t e d ( F i g . 6, 8 ) . The r o c k s a r e p r i m a r i l y sandstone, a t I n d i a n Beach.  and much s o f t e r than  those  A f r e s h w a t e r stream f l o w s a c r o s s the beach  p a r a l l e l t o and near t h e o u t c r o p s .  The sand l e v e l was r e c o r d e d  lire 8  S h o r t Sand Beach s t u d y a r e a .  L. s i n c l a i r i i o c c u r s on a l l r o c k s shown and at  s e v e r a l p o i n t s on t h e r o c k f o r m i n g the s h o r e .  P a r t i c u l a r measurements o r o b s e r v a t i o n s were made on "A" and "B".  27  Figure 9  V a r i a t i o n i n sand l e v e l s a t I n d i a n Beach in  1967.  a.  April  1967  b.  May  c.  June  1967  d.  July  1967  1967  I n c and d, the l e f t b u c k e t (arrow) i s on r o c k A, the r i g h t b u c k e t  (arrow) on r o c k C.  r o c k i s the s t a c k i n d i c a t e d i n f i g u r e  The 7.  large  28  F i g u r e 10  v a r i a t i o n i n sand l e v e l a t s t u d y r o c k s a t I n d i a n Beach i n  a.  A p r i l 1967. a t low  b.  May  1967.  Note t h a t s u r f i s around r o c k s , even  tide.  1967.  Sand has advanced b u t i s n o t y e t  a r o u n d most o f the r o c k s . c.  June  1967  d.  July  1967  e.  A u g u s t 1967.  Some o f the b l a d e s o f L_. s i n c l a i r i i  have been s c o u r e d o f f r o c k A. f.  September  1967.  A r r o w i n a l l f i g u r e s i n d i c a t e s r o c k B.  I n c, d,  e, f , b u c k e t i n f o r e g r o u n d i s on r o c k A, b u c k e t i n b a c k g r o u n d i s on r o c k C (= c l e a r e d r o c k ; see F i g . 1 6 ) .  29  a t monthly i n t e r v a l s from A u g u s t 1966 t h r o u g h September 1967 at  two p o i n t s on t h e b e a c h  (Table X I V ) .  The g r e a t e s t  r e c o r d e d f l u c t u a t i o n s o f t h e sand l e v e l a t t h e s t u d y s i t e s i s 70 cm.  The f r e s h w a t e r s t r e a m w h i c h f l o w s so near t h e r o c k s  i n t e r a c t s w i t h t h e sand t o produce a somewhat d i f f e r e n t s i t u a t i o n from t h a t on I n d i a n Beach.  The c o u r s e o f t h e s t r e a m  i s s h i f t e d s l i g h t l y d u r i n g each t i d a l c y c l e .  However, most  of  t h e time t h e s t r e a m f l o w s n e x t t o t h e r o c k s , a t l e a s t i n  the  upper p a r t o f t h e beach, s e p a r a t i n g t h e r o c k s from t h e  r e s t o f the beach.  A s t h e sand l e v e l r i s e s i n summer, t h e  l e v e l o f t h e f r e s h water r i s e s n e x t t o t h e r o c k s , so t h a t b y l a t e summer as much as 30 cm o f some r o c k s i s submerged i n f r e s h water a t l o w t i d e the  ( F i g . 1 1 ) . T h i s i n c l u d e s much o f  p o r t i o n o f t h e r o c k s w h i c h i s i n h a b i t e d by L. s i n c l a i r i i .  The p r e s e n c e o f t h e s t r e a m , and i t s c o n s t a n t e r o s i o n o f a c h a n n e l , i n t u r n p r e v e n t s t h e r o c k s from b e i n g b u r i e d so d e e p l y under sand.  D u r i n g t h e l a t t e r p a r t o f t h e summer o f  1967 t h e l e v e l o f t h e sand on a l l t h e beach n o r t h o f t h e s t r e a m was a t l e a s t 1 m above t h e l e v e l o f t h e t o p s o f t h e r o c k s studied (Fig. l l f ) . the  The r o c k s had about 35-45 cm exposed above  sand i n t h e bottom o f t h e s t r e a m c h a n n e l and about 5-10  cm o f t h i s exposed above t h e f r e s h w a t e r , depending on t h e height o f the rock.  I n w i n t e r , when sand i s c o m p l e t e l y a b s e n t  n e x t t o t h e r o c k s , t h e r e i s u s u a l l y a 10-20 cm s t r i p o f r o c k at  t h e bottom, where t h e s t r e a m f l o w s , w h i c h i s c o m p l e t e l y  bare o f p l a n t s .  The r o c k above, w h i c h s u p p o r t s L. s i n c l a i r i i  and v a r i o u s o t h e r a l g a e , i s n o t t o u c h e d b y t h e s t r e a m w a t e r .  30  F i g u r e 11  V a r i a t i o n i n sand and water Sand B e a c h i n  l e v e l s at Short  1967.  a.  L e v e l o f sand and f r e s h w a t e r stream i n June  1967.  b.  L e v e l o f sand and f r e s h w a t e r stream i n J u l y  c.  L e v e l o f sand and f r e s h w a t e r stream i n August  1967. 1967.  Note t h a t sand i s h i g h e r than s t u d y r o c k s . I n a, b, and c, b u c k e t i n f o r e g r o u n d i s on r o c k B,  bucket  i n b a c k g r o u n d i s on r o c k A. d.  Study a r e a from above i n A p r i l 1967. of p i c t u r e .  Rock A i s o u t  Rocks i n background a r e c o m p l e t e l y  b u r i e d l a t e r i n summer. e.  Study a r e a from above i n J u l y  1967.  f.  Study a r e a from above i n A u g u s t 1967.  Sand i n  b a c k g r o u n d i s h i g h e r than s t u d y r o c k s and  freshwater  stream has r i s e n a l s o . I n e and f , l e f t b u c k e t i s on r o c k A, r i g h t b u c k e t i s on r o c k  B.  31  The s t u d y s i t e a t A r c h Cape i s a l a r g e b a s a l t i c d i r e c t l y west o f a l a r g e s t a c k west o f the Cape.  outcrop  The r o c k i s  about 7 m from e a s t t o west, 2 m from n o r t h t o s o u t h  and  about 1 m i n h e i g h t from the l o w e s t w i n t e r sand l e v e l .  It is  s e p a r a t e d from the s t a c k by a s t r e t c h o f beach about 20 m wide.  Somewhat n o r t h and west o f i t i s a n o t h e r , l a r g e r r o c k .  W i t h t h e e x c e p t i o n o f t h i s l a r g e r r o c k , the study r o c k r e c e i v e s a g r e a t e r f o r c e o f s u r f than any o t h e r p o i n t f o r about 5 km a l o n g the beach.  A f r e s h w a t e r stream f l o w s a c r o s s  the b e a c h about 30 m n o r t h o f the r o c k b u t does n o t u s u a l l y come i n t o c o n t a c t w i t h i t . No measurements o f sand f l u c t u a t i o n have been made near the s t u d y s i t e a t A r c h Cape.  Measurements o f sand h e i g h t  made a t a p o i n t a p p r o x i m a t e l y 1 km f a r t h e r n o r t h on the same b e a c h show a maximum f l u c t u a t i o n o f o n l y a l i t t l e over 1 m (Table XV). 2.  I t i s p r o b a b l y g r e a t e r near the r o c k .  O c c u r r e n c e o f L.  sinclairii  L. s i n c l a i r i i o c c u r s a t I n d i a n Beach on many r o c k s a t the n o r t h end o f the b e a c h ( F i g . 7 ) . r e g i o n between the +1.5  and the -2.0  t i d e l e v e l s on a l l o f the r o c k s .  The  I t i s r e s t r i c t e d t o the f t . (+0.46 and -0.61m) s i z e o f the p l a n t s  tends t o be l a r g e r i n the lower p a r t s o f the d i s t r i b u t i o n . A t S h o r t Sand Beach, L. s i n c l a i r i i o c c u r s on n e a r l y a l l r o c k s w h i c h b o r d e r the sandy beach a t the s o u t h end I t o c c u r s o n l y between the +3.0 tide levels.  and -1.5  (Fig. 8).  f t . (0.92 and -0.46m)  A t the upper l i m i t s o f i t s d i s t r i b u t i o n i t i s  32  very small.  I n June 1967,  when the sand was  d u r i n g a -6.1  s t i l l v e r y low, the lower l i m i t s o f  L. s i n c l a i r i i were d e t e r m i n e d point.  f t (-1.86m) t i d e  on r o c k s f a r o u t toward  the  Below the l o w e s t L. s i n c l a i r i i , a p a t c h o f about 15  cm o f c o m p l e t e l y b a r e r o c k was  evident.  A t A r c h Cape, the d i s t r i b u t i o n o f L. s i n c l a i r i i was mapped. Cape.  not  I t o c c u r s on s e v e r a l r o c k s a t the o u t e r p a r t o f the I t c o v e r s the e n t i r e s u r f a c e o f the r o c k on which i t  was measured.  The v e r t i c a l d i s t r i b u t i o n was n o t measured.  Based on o b s e r v a t i o n s d u r i n g v a r i o u s t i d e s , a l l the p l a n t s appear t o be c o n f i n e d t o the a r e a between MLLW (-4.5 (-1.37 m)  and LLLW ( c a . -7 f t . ) (-2.1  on the o t h e r b e a c h e s .  m).  ft.)  T h i s i s lower  than  However, the d i s t r i b u t i o n does n o t  e x t e n d i n t o the s u b t i d a l  zone.  As i s shown i n F i g u r e 4, t h e r e i s a c o n s i d e r a b l e d i f f e r e n c e i n t o t a l time o f submergence between the upper and lower l i m i t s of p l a n t d i s t r i b u t i o n .  P l a n t s growing above the 0.0  f t . tide  l e v e l a r e a c t u a l l y o u t o f water f o r a g r e a t e r p e r c e n t a g e the t o t a l time then t h e y a r e under water.  of  As the t o t a l time  o u t o f water v a r i e s and w i t h i t the t o t a l time w h i c h the p l a n t i s exposed t o s u n l i g h t w h i l e o u t o f w a t e r , d i f f e r e n c e s i n temperature,  t h e r e a r e many  l i g h t q u a l i t y and i n t e n s i t y ,  p o t e n t i a l for desiccation at d i f f e r e n t l e v e l s .  Thus i t i s to  be e x p e c t e d t h a t growth and s i z e v a r y a t the d i f f e r e n t The  and  levels.  s i t e s where growth measurements were made a r e a l l a t  different heights. Beach a r e a t the 0.0  The measured p l a n t s on r o c k A a t I n d i a n f t . tide level.  Those on r o c k B a r e a t  33  -0.5  f t (-0.15m).  The  t o t a l time w h i c h the p l a n t s on r o c k  A  a r e out o f water each month i s about 7 p e r c e n t g r e a t e r than t h a t f o r those on r o c k B.  The p l a n t s on r o c k B have a g r e a t e r  s i z e and g r e a t e r average monthly growth than those on r o c k A (Table X X I ) . about the +2.0  The measured p l a n t s a t S h o r t Sand Beach a r e a t f t . (0.61m) t i d e l e v e l  ( c a . MLHW).  These  p l a n t s a r e somewhat s m a l l e r than those a t I n d i a n Beach. 3.  Associated Plant  Species  L i s t s o f a l l the p l a n t s p e c i e s c o l l e c t e d on the  three  Oregon beaches t o g e t h e r w i t h the h e i g h t s and months o f c o l l e c t i o n are presented  i n Tables  XVI/ X V I I / X V I I I .  two c o l l e c t i o n s were made a t A r c h Cape. species present there.  Only  There a r e v e r y  On most o f the r o c k s where L.  few sinclairii  grows, no o t h e r s p e c i e s a r e found w i t h the e x c e p t i o n o f a  few  p l a n t s o f L. s e t c h e l l i i on the o u t e r and most exposed p a r t s o f the r o c k s .  A t I n d i a n Beach L. s i n c l a i r i i  i s the most  abundant p l a n t i n terms o f cover on a l l the r o c k s on w h i c h i t grows ( F i g . 11, 1 2 ) . found w i t h i t .  However, s e v e r a l o t h e r s p e c i e s are commonly  Phaeostrophion  i r r e g u l a r e o c c u r s a t the same  h e i g h t and s l i g h t l y h i g h e r than L. s i n c l a i r i i .  Several  s p e c i e s o f B o s s i e l l a a r e o f t e n found growing w i t h L. D i l s e a c a l i f o r n i c a i s common, u s u a l l y a t a s l i g h t l y l e v e l on the same r o c k s ,  v a r i o u s crustos'e c o r a l l i n e  a r e found on the same r o c k s above and b e l o w L.  sinclairii. higher algae  sinclairii.  A few p l a n t s o f L. s e t c h e l l i i a r e o f t e n a s s o c i a t e d w i t h L. s i n c l a i r i i , a l t h o u g h L. s e t c h e l l i i tends t o occur  lower i n  34  F i g u r e 12  L. s i n c l a i r i i and a s s o c i a t e d s p e c i e s a t I n d i a n Beach.  a.  A s s o c i a t i o n o f L. s i n c l a i r i i w i t h o t h e r s p e c i e s a t 1.0 f t . b e l o w mean t i d e l e v e l .  L. s e t c h e l l i i and  P h y l l o s p a d i x s c o u l e r i o c c u r w i t h L. s i n c l a i r i i . Hedophyllum s e s s i l e i s h i g h e r on r o c k . b.  June 1967.  B u r i a l under sand o f L. s i n c l a i r i i i n August 1967. Some b l a d e s have been e r o d e d away.  c.  P l a n t s on r o c k B i n June 1967. is  d.  Multipunched blade  visible.  P l a n t s on r o c k A i n June 1967. are v i s i b l e . luxuriant  Some punched b l a d e s  Note l e s s complete dominance and l e s s  growth o f L. s i n c l a i r i i as compared w i t h  t h a t on r o c k B w h i c h i s l o w e r .  35  the i n t e r t i d a l zone and i n s l i g h t l y more exposed l o c a t i o n s than L. s i n c l a i r i i .  P h y l l o s p a d i x s c o u l e r i i s common w i t h  L. s e t c h e l l i i and the lower p a r t s o f the L_. p o p u l a t i o n ( F i g . 12a).  sinclairii  Gymnogongrus l i n e a r i s and Codium  s e t c h e l l i i o c c u r from the lower l i m i t s o f the L_.  sinclairii  p o p u l a t i o n down to about 50 cm b e l o w these l i m i t s .  Both  u s u a l l y b u r i e d under sand f o r most o f the summer.  are  They a r e  a p p a r e n t l y w e l l a d a p t e d t o t h i s , as they appear h e a l t h y when the sand a g a i n r e c e d e s .  L. s i n c l a i r i i  dominance on lower r o c k s  ( F i g . 13).  A t S h o r t Sand Beach, L. s i n c l a i r i i  shows more complete  i s dominant on o n l y  a few o f the l o w e s t , most exposed r o c k s on which i t o c c u r s . On most o f the r o c k s where i t o c c u r s t h e r e a r e s e v e r a l o t h e r a l g a e i n e q u a l o r g r e a t e r abundance ( F i g . 13 d , c ) .  Dilsea  c a l i f o r n i c a and F a r l o w i a m o l l i s a r e v e r y abundant on some o f the r o c k s , u s u a l l y s l i g h t l y h i g h e r than L.  sinclairii.  B o s s i e l l a plumosa, M i c r o c l a d i a b o r e a l i s , p t i l o t a a s p l e n o i d e s , P_. f i l i c i n a , P_. p e c t i n a t a , I r i d a e a sp. and P r i o n i t i s  lyallii  a r e v e r y common a t the same h e i g h t a s , and s l i g h t l y h i g h e r than L. s i n c l a i r i i .  On many o f these r o c k s , the dominant  cover i s composed o f M i c r o c l a d i a b o r e a l i s and p t i l o t a  spp.  C r u s t o s e c o r a l l i n e a l g a e o c c u r on the same r o c k s , a b o v e , and w i t h L. s i n c l a i r i i .  below,  Gymnogongrus l i n e a r i s o c c u r s a t the  same h e i g h t and somewhat lower than L. s i n c l a i r i i .  Below the  l e v e l o f most o f the L_. s i n c l a i r i i , p h y l l o s p a d i x s c o u l e r i i s abundant and s e v e r a l p l a n t s o f L. s e t c h e l l i i a r e  found.  Gymnogongrus l i n e a r i s and P h y l l o s p a d i x s c o u l e r i a r e o f t e n n e a r l y  36  F i g u r e 13  Submergence i n f r e s h w a t e r and a s s o c i a t i o n o f L. s i n c l a i r i i w i t h o t h e r s p e c i e s a t S h o r t Sand Beach.  a.  Some o f L. s i n c l a i r i i f o r w h i c h growth was measured, hanging i n t o f r e s h w a t e r stream, June 1967.  Two  punched h o l e s a r e v i s i b l e . b.  L a t e summer submergence, August 1967. r o c k A.  Bucket i s on  P l a n t s on r o c k A a r e p a r t l y b u r i e d under  sand a s w e l l a s b e i n g submerged i n f r e s h w a t e r . c.  B u r i a l under sand i n f r e s h w a t e r stream o f L. s i n c l a i r i i , J u l y 1967.  d.  Some o f L. s i n c l a i r i i f o r w h i c h growth was measured, showing s m a l l s i z e o f clump and r e l a t i v e l a c k o f dominance compared w i t h o t h e r s p e c i e s , May 1967.  e.  S m a l l p a t c h e s o f L. s i n c l a i r i i on r o c k w i t h cover o f p t i l o t a spp and M i c r o c l a d i a b o r e a l i s .  June 1967.  37 c o m p l e t e l y b u r i e d under sand.  Most o f t h e o t h e r s p e c i e s  mentioned a r e p a r t l y b u r i e d o r a t l e a s t c o a t e d w i t h sand f o r much o f t h e summer. 4.  Seasonal  Cycles  In t h e c o u r s e o f a normal y e a r , s e a s o n a l c y c l e s o f f o u r d i f f e r e n t phenomena o c c u r i n L. s i n c l a i r i i .  The p l a n t s grow;  t h e y b e a r r i p e s o r i ; t h e y l o s e t h e i r b l a d e s ; and they a r e b u r i e d under sand. account  The o b s e r v a t i o n s on w h i c h t h e f o l l o w i n g  i s b a s e d a r e summarized i n T a b l e XX.  In e a r l y January, without blades.  the p l a n t s are normally  completely  The r o c k s on w h i c h t h e p l a n t s a r e a t t a c h e d  a r e c o m p l e t e l y uncovered and sand may be absent from t h e e n t i r e area.  Growth o f t h e p l a n t s i s v e r y s l o w .  Later i n  J a n u a r y t h e ends o f t h e s t i p e s s p l i t and from t h e m e d u l l a r y r e g i o n s new b l a d e s b e g i n t o d e v e l o p . growing  The b l a d e s  and t h e r a t e o f growth a c c e l e r a t e s .  grow, b u t much more s l o w l y than t h e b l a d e s .  continue  The s t i p e s T a b l e XXI  summarizes t h e growth measurements o f in_ s i t u p l a n t s on t h e t h r e e Oregon b e a c h e s . E a r l y i n F e b r u a r y when t h e p l a n t s a r e o n l y 2-3 cm i n l e n g t h , f e r t i l e s o r i begin t o develop a t the blade t i p s .  Ripe  s o r i a r e produced i n March and sometimes as l a t e as A p r i l . At the beginning o f t h i s reproductive p e r i o d there i s c o n s i d e r a b l e d i f f e r e n c e i n t h e time o f s o r u s development and s i z e o f s o r i developed.  L a t e r i n the p e r i o d , the sorus  p r o d u c t i o n becomes more s y n c h r o n i z e d .  I n A p r i l 1967, a t t h e  b e g i n n i n g o f a l o w t i d e s e r i e s , n e a r l y a l l the b l a d e s i n a  38  p o p u l a t i o n on one r o c k a t I n d i a n Beach had r i p e s o r i on t e r m i n a l 2 cm o f b l a d e .  the  These remained 4 days u n t i l the h i g h  t i d e b e f o r e the l a s t low t i d e w h i c h would uncover the p l a n t s for  more than an hour i n t h i s t i d e s e r i e s .  low t i d e i t was  observed  D u r i n g the  last  t h a t a l l the s o r i had been dropped,  l e a v i n g a b i t e - s h a p e d h o l e a t the end o f each b l a d e . L a t e i n March, and i n A p r i l and May o f new  the maximum i n i t i a t i o n  s t i p e s and b l a d e s o c c u r s from the h o l d f a s t s .  s t i p e s can be d i s t i n g u i s h e d from the o l d e r ones by  New color.  D u r i n g the f i r s t y e a r the s t i p e s a r e the same l i g h t brown c o l o r as the b l a d e s .  By the time the s t i p e s a r e one year o l d  and have l o s t t h e i r b l a d e s once, they a r e v e r y dark, b l a c k , and v e r y u n l i k e the b l a d e s i n c o l o r .  almost  The growth o f  the o t h e r s t i p e s and b l a d e s i s g r e a t e s t i n May  and June.  In A p r i l the sand on the beach b e g i n s to b u i l d up,  but  i s n o t y e t around the r o c k s on w h i c h L. s i n c l a i r i i grows. Sometime i n May  o r June, the low p l a c e s a r e u s u a l l y c o v e r e d  w i t h sand, l e a v i n g the r o c k s b e a r i n g L. s i n c l a i r i i s t i c k i n g o u t o f the sand ( F i g . 1 0 c ) .  still  I n y e a r s when the sand  l e v e l i s h i g h , the r o c k s as w e l l as the h o l d f a s t s and o f most o f the L. s i n c l a i r i i p l a n t s on them a r e  stipes  completely  b u r i e d by m i d - J u l y , l e a v i n g o n l y the b l a d e s o f L_. s i n c l a i r i i exposed ( F i g . l O d ) . A r c h Cape i n 1967.  Such was  the case a t I n d i a n Beach and  I t would have o c c u r r e d a t S h o r t Sand  Beach a l s o , had the stream n o t k e p t the sand away from most o f the r o c k s .  I n o t h e r y e a r s , as i n 1966,  the sand d i d n o t  r i s e as h i g h and by September o n l y the h o l d f a s t s o f the p l a n t s  39  were b u r i e d .  I n September  o r O c t o b e r , the f i r s t heavy storms  remove most o f the sand, e x p o s i n g a l l the r o c k s and p l a n t s a g a i n u n t i l the f o l l o w i n g summer. As the p l a n t s become b u r i e d , growth o f the b l a d e s d e c r e a s e s (Table X X I ) .  However, i t was n o t e d t h a t the d e c r e a s e  i n growth i s much l e s s marked i n p l a n t s w h i c h a r e growing lower i n the i n t e r t i d a l zone and i n more exposed a r e a s .  In  t h e s e a r e a s the p e r i o d o f maximum growth may be one o r two months l a t e r than i n o t h e r a r e a s .  T h i s can be seen i n  comparing r e s u l t s a t I n d i a n Beach w i t h t h o s e a t S h o r t Sand Beach, w h i c h i s s l i g h t l y l e s s exposed, and a t A r c h Cape, w h i c h i s more exposed.  I t i s a l s o e v i d e n t i n comparing the two  I n d i a n Beach s i t e s , w h i c h d i f f e r i n exposure.  The growth a t  S h o r t Sand Beach may a l s o have been i n h i b i t e d by the i n c r e a s i n g c o n t a c t w i t h the f r e s h w a t e r stream w h i c h r o s e and impinged on the  p l a n t s more as the sand l e v e l under i t r o s e .  As a  consequence o f the g r e a t e r growth, t h e p l a n t s g e n e r a l l y have l o n g e r s t i p e s and b l a d e s i n the more exposed a r e a . As t h e p l a n t s become b u r i e d , the s c o u r i n g a c t i o n o f the sand i n c r e a s e s .  I n some a r e a s , many o r a l l o f the b l a d e s may  be l o s t , p o s s i b l y due t o t h e s c o u r i n g . w i t h exposure.  However, t h i s  varies  I n the summer o f 1967, many b l a d e s were l o s t  a t S h o r t Sand Beach, fewer a t I n d i a n Beach, and none a t A r c h Cape. of  The b u r i a l was more.complete  the o t h e r s t u d y a r e a s .  a t A r c h cape than a t any  The s t r o n g f r e s h w a t e r i n f l u e n c e a t  S h o r t Sand Beach may a l s o have c o n t r i b u t e d t o t h e b l a d e l o s s .  40  The p l a n t s grow v e r y l i t t l e from September t h r o u g h December.  I n October  are developed.  the f i r s t r i p e s o r i o f the o l d b l a d e s  I n November l a r g e a r e a s o f the b l a d e s a r e  covered w i t h oblong patches of s o r i .  Experiments  show t h a t  t h e s e s o r i produce zoospores which d e v e l o p i n t o gametophytes, at  l e a s t under l a b o r a t o r y c o n d i t i o n s .  However, these  gametophytes do n o t u s u a l l y produce s p o r o p h y t e s . M e c h a n i c a l e r o s i o n o f the b l a d e t i p s o c c u r s t o some e x t e n t at  a l l times.  Under normal c o n d i t i o n s growth proceeds  faster  t h a n e r o s i o n , and the b l a d e l e n g t h e n s d u r i n g the y e a r . B e g i n n i n g i n November when growth has p r a c t i c a l l y  ceased,  endogenous d i s i n t e g r a t i o n o f the b l a d e t i p s b e g i n s t o o c c u r i n a d d i t i o n to e r o s i o n .  These p r o c e s s e s c o n t i n u e u n t i l  by  mid-December most o f the b l a d e s a r e c o m p l e t e l y gone, l e a v i n g o n l y the b a r e 5.  stipes.  Discussion L. s i n c l a i r i i was o b s e r v e d a t s e v e r a l beaches i n Washington,  Oregon, and C a l i f o r n i a  (Table I I ) .  A t a l l o f these beaches  the g e n e r a l c o n d i t i o n s a r e v e r y s i m i l a r t o those on the s t u d y beaches.  The beaches a r e f u l l y exposed  t o s u r f and t h e r e i s  e v i d e n c e o f a marked s e a s o n a l f l u c t u a t i o n i n sand l e v e l causes t h e p l a n t s t o be b u r i e d i n summer.  which  The a s s o c i a t e d p l a n t  s p e c i e s a r e a l s o s i m i l a r t o those found on the t h r e e s t u d y beaches (Table X V I ) . s e l e c t e d f o r study are  Thus i t appears t h a t the beaches typical.  The p l a n t s o f L. s i n c l a i r i i bear r i p e s o r i d u r i n g two p e r i o d s , once j u s t a f t e r the new b l a d e s b e g i n t o d e v e l o p and once j u s t  41  b e f o r e the o l d b l a d e s a r e l o s t .  I t appears e v i d e n t t h a t s o r u s  i n i t i a t i o n i s n o t a t a l l dependent upon the age o f the b l a d e tissue.  The f a c t o r s w h i c h i n d u c e t h i s i n i t i a t i o n have n o t  been d e t e r m i n e d . B o t h p e r i o d s o f s o r u s p r o d u c t i o n o c c u r d u r i n g the time when the sand i s a t i t s l o w e s t .  T h i s l e a v e s the maximum  amount o f b a r e r o c k a v a i l a b l e f o r the zoospores t o a t t a c h and produce gametophytes w h i c h then c o u l d produce s p o r o p h y t e s . However, no e v i d e n c e has been found t h a t t h i s a c t u a l l y o c c u r s (See a l s o pp 59-63 and 69-73). L. s i n c l a i r i i grows b e t t e r the lower i t i s i n the i n t e r t i d a l zone b u t Kas n o t ^ f o u n d i n the s u b t i d a l zone.  It  appears t h a t i t may r e q u i r e a s l i g h t amount o f d e s i c c a t i o n b u t cannot t o l e r a t e v e r y much d e s i c c a t i o n .  T h i s h y p o t h e s i s has  n o t been t e s t e d , a l t h o u g h a l a c k o f d e s i c c a t i o n may  partly  e x p l a i n the poor growth i n l a b o r a t o r y t a n k s (See pp 66-69) I f the p l a n t s a r e g r o w i n g i n the optimum p o s i t i o n w i t h r e g a r d t o s u r f and d e s i c c a t i o n , b u r i a l by sand appears t o cause no damage.  T h i s i n d i c a t e s t h a t the p l a n t i s w e l l adapted t o  w i t h s t a n d b u r i a l b u t i s more s e n s i t i v e t o v a r i a t i o n s i n d e s i c c a t i o n and wave exposure.  The r e s u l t s , such as l o s s o f  b l a d e s and c e s s a t i o n o f growth i n l e s s f a v o r a b l e s i t e s , n o t be d i r e c t l y due t o sand b u r i a l a t a l l .  may  42  V.  DISTRIBUTION AND AUTECOLOGY OF LAMINARIA LONGIPES  A-  Geographical  Distribution  I n t h e Western p a c i f i c , t h e s o u t h e r n  l i m i t of d i s t r i b u t i o n  o f L. l o n g i p e s i s Urup I s l a n d (46°00'N, 150°00'E) i n t h e K u r i l e Islands  (Table I I I , F i g . 1 ) . N o r t h o f t h i s , t h e p l a n t  i s f o u n d on S o u t h S a k h a l i n , on v a r i o u s o t h e r i s l a n d s i n the K u r i l e I s l a n d s , a t s e v e r a l p o i n t s a l o n g t h e e a s t c o a s t o f Kamchatka, and on B e r i n g I s l a n d .  I n the Eastern  pacific,  L. l o n g i p e s i s f o u n d on S t . P a u l I s l a n d i n t h e B e r i n g Sea, and from A t t u I s l a n d i n t h e A l e u t i a n I s l a n d s through Alaska to Coronation Alaska,  the Gulf of  I s l a n d (55°49.6'N, 134°17'W) i n S o u t h e a s t  A s u b t i d a l p o p u l a t i o n has been r e p o r t e d by D r u e h l  (1968) a t Salmon Bank Washington.  (48°26'N, 123°01'W) San Juan I s l a n d ,  The p l a n t s from Salmon Bank have b l a d e s up t o 2 0  cm b r o a d , b u t o t h e r w i s e f i t t h e d e s c r i p t i o n o f L_. l o n g i p e s . U n t i l more e x t e n s i v e s u b t i d a l c o l l e c t i o n s a r e made i n t h e N o r t h e a s t p a c i f i c , i t i s n o t p o s s i b l e t o a s s e s s the s i g n i f i c a n c e o f t h i s c o l l e c t i o n as an e x t e n s i o n o f t h e r a n g e . The a r e a i n w h i c h L. l o n g i p e s grows i s c h a r a c t e r i z e d by v e r y l o w w i n t e r seawater t e m p e r a t u r e s (Table I , F i g . 2) r a n g i n g from means o f 0.5°C a t Urup I s l a n d t o 4.4°C a t S i t k a . Summer t e m p e r a t u r e s range from a h i g h monthly mean o f 9.3°C a t p y r a m i d Cove t o 14.1°c a t S i t k a . The y e a r l y means range o o from 5.1 C a t Urup I s l a n d and A t t u I s l a n d t o 8.5 C a t S i t k a . The a i r t e m p e r a t u r e s a r e a l s o v e r y l o w (Table V, V I , F i g . 3)  43  p a r t i c u l a r l y a t Urup I s l a n d .  The s a l i n i t y w i t h i n t h e p l a n t ' s  range v a r i e s from a y e a r l y mean o f 27.7%  0  a t Sitka to  32.1%  0  a t P y r a m i d Cove. T a b l e I I I l i s t s t h e s i t e s a t w h i c h L.. l o n g i p e s has been c o l l e c t e d o r observed.  A l l i n f o r m a t i o n on d i s t r i b u t i o n i n  R u s s i a n w a t e r s has been o b t a i n e d from p u b l i s h e d r e c o r d s .  Most  o f t h e r e c o r d s o f d i s t r i b u t i o n i n American waters have been o b t a i n e d from c o l l e c t i o n s a v a i l a b l e i n t h e P h y c o l o g i c a l Herbarium, U n i v e r s i t y o f B r i t i s h Columbia.  A t most o f t h e  s i t e s o b s e r v e d by t h e a u t h o r , L_. l o n g i p e s grows on r o c k y r e e f s i n m o d e r a t e l y exposed t o m o d e r a t e l y s h e l t e r e d a r e a s .  No  p a r t i c u l a r v a r i a t i o n i n morphology was n o t e d i n p l a n t s from v a r i o u s p a r t s o f t h e range o f d i s t r i b u t i o n . B.  H a b i t a t and A u t e c o l o g y a t A a t s Bay, C o r o n a t i o n I s l a n d , Alaska  1.  Environment The s t u d y s i t e a t A a t s Bay i s a r o c k y r e e f a d j a c e n t t o a  beach composed o f g r a v e l and c o a r s e sand ( F i g . 1 4 ) . The r e e f i s composed o f a r g i l l i t e r o c k and i s c u t by s e v e r a l deep surge channels.  There i s no e v i d e n c e t o i n d i c a t e t h a t t h e r e e f i s  ever b u r i e d under sand o r t h a t t h e sand l e v e l changes.  The  beach i s i n a l o c a t i o n w h i c h i s m o d e r a t e l y s h e l t e r e d t o m o d e r a t e l y exposed.  Although there i s f r e q u e n t l y surf i n the  w i n t e r , i n summer t h e water i s o f t e n c o m p l e t e l y calm. t i d e s i n t h i s a r e a s , as i n Oregon, a r e o f t h e mixed type.  The  semidiurnal  The l o w e s t o f t h e l o w w a t e r s o c c u r s i n the daytime i n  44  F i g u r e 14  L o c a t i o n o f A a t s Bay s t u d y  Helm P o i n t i s the s o u t h e r n d i s t r i b u t i o n of  L. l o n g i p e s .  area.  l i m i t of  45  summer and a t n i g h t i n w i n t e r . Coronation  The maximum t i d a l a m p l i t u d e a t  I s l a n d i s 16.8 f t (5.12m) t h e d i u r n a l a m p l i t u d e i s  10.7 f t . (3.26m), and t h e mean t i d e l e v e l i s 8.7 f t . (2.65m) above MLLW (Anon., 1968b). T h i s a r e a i s c h a r a c t e r i z e d by r e l a t i v e l y heavyp r e c i p i t a t i o n and l o w t e m p e r a t u r e s . meteorological records f o r Coronation Decision island  There a r e no p u b l i s h e d Island.  However, Cape  (56°00'N, 134°08"W) i s o n l y 10 km from  Coronation  ( F i g . 15) and i t i s assumed t h a t t h e m e t e o r o l o g i c a l  c o n d i t i o n s a r e v e r y s i m i l a r a t t h e two p l a c e s . f o r cape D e c i s i o n  The r e c o r d s  (Table V) show t h a t t h e a n n u a l mean a i r  temperature i s 6.3°C, r a n g i n g from a monthly mean o f 0.9°c i n J a n u a r y t o 11.7°C i n August.  The mean t o t a l a n n u a l  p r e c i p i t a t i o n i s 76.12 i n c h e s , w i t h t h e g r e a t e s t amount o c c u r r i n g i n October and t h e l e a s t i n June.  A comparison o f  t h e s e r e c o r d s w i t h those f o r S i t k a (Table V) shows t h a t t h e average m o n t h l y p r e c i p i t a t i o n i s g r e a t e r a t S i t k a t h a n a t Cape D e c i s i o n f o r e v e r y month o f t h e y e a r .  A comparison o f t h e  t e m p e r a t u r e r e c o r d s shows a g r e a t e r a n n u a l temperature range a t S i t k a than a t Cape D e c i s i o n .  T h i s might be e x p e c t e d , s i n c e  S i t k a i s on a l a r g e i s l a n d and i s l a r g e l y surrounded by l a n d , whereas cape D e c i s i o n i s more s u b j e c t t o t h e m o d e r a t i n g i n f l u e n c e o f surrounding  water masses.  Maps o f the a r e a  i n d i c a t e t h e harbor a t S i t k a r e c e i v e s more f r e s h w a t e r  runoff  than does t h e s e a around c o r o n a t i o n I s l a n d and cape D e c i s i o n , although  exact f i g u r e s are l a c k i n g .  r a i n f a l l , i s important areas.  T h i s , together w i t h the  i n c o n s i d e r a t i o n s o f s a l i n i t y i n t h e two  46  There a r e no p u b l i s h e d r e c o r d s o f s a l i n i t y o r water temperature a t C o r o n a t i o n  Island.  I n t h i s study these f a c t o r s  were measured o n l y once, i n December 1966. A t t h i s time t h e temperature was 4.5°C, 1C° b e l o w t h e r e a d i n g o b t a i n e d a t V o l g a I s l a n d , S i t k a , 24 hours l a t e r .  The s a l i n i t y was 3 0 . 7 8 % , o  w h i c h was 0.5% b e l o w t h a t r e c o r d e d a t V o l g a i s l a n d . o  Long term  r e c o r d s o f t h e seawater temperature and s a l i n i t y have been published for Sitka  (Anon 1967d; see a l s o Table X X I I I ) . I f  the d i f f e r e n c e r e c o r d e d i n December i s v a l i d f o r t h e whole y e a r , t h e seawater temperature range a t C o r o r a t i o n I s l a n d s h o u l d be from a p p r o x i m a t e l y  3.4°C i n F e b r u a r y  A u g u s t , w i t h a y e a r l y mean o f 7.5°C.  t o 13.1°c i n  These may be f a i r l y c l o s e  t o t h e c o r r e c t v a l u e s , b u t adequate i n f o r m a t i o n i s l a c k i n g . Because o f t h e g r e a t e r p r e c i p i t a t i o n and r u n o f f a t S i t k a , i t seems l i k e l y t h a t t h e average s a l i n i t y i n S i t k a Harbor i s lower than t h a t a t C o r o n t a t i o n I s l a n d .  Inasmuch as t h e one s e t  o f r e c o r d e d o b s e r v a t i o n s show t h e i n v e r s e r e l a t i o n s h i p , t h e average monthly s a l i n i t i e s cannot be e s t i m a t e d from t h e available 2.  data.  O c c u r r e n c e o f L. l o n g i p e s and a s s o c i a t e d p l a n t s p e c i e s L. l o n g i p e s o c c u r s on t h e r o c k y r e e f i n t h e lower  i n t e r t i d a l and upper s u b t i d a l zones.  Most o f t h e p l a n t s a r e  uncovered by a -10.2 f t . (-3.11m) t i d e .  On t h e s e c t i o n s o f  the r e e f where i t i s found, i t i s u s u a l l y t h e dominant p l a n t i n terms o f t o t a l c o v e r , a l t h o u g h i n a few p l a c e s A l a r i a marginata  i s e q u a l l y abundant.  S l i g h t l y h i g h e r on t h e same  47  r e e f s , i n the m i d - i n t e r t i d a l p o s i t i o n , Laminaria groenlandica, Hedophyllum s e s s i l e and A l a r i a t e n u i f o l i a a r e abundant. complete  A  l i s t o f t h e s p e c i e s found on t h i s beach i s p r e s e n t e d  i n Table XIX. 3.  Growth and r e p r o d u c t i o n o f L. l o n g i p e s a t A a t s Bay Growth o f i n s i t u p l a n t s was f o l l o w e d f o r o n l y one month,  J u l y 1966. H o l e s were punched i n t h e b l a d e s 10 cm above t h e base i n t h e u s u a l manner.  Based on t h e h o l e s t h e average  growth o f t h e b l a d e s f o r J u l y was 8 cm. b u t much more s l o w l y .  The s t i p e s grew a l s o ,  I t i s assumed t h a t maximum growth  occurs  i n t h e summer months b u t d i r e c t measurements a r e l a c k i n g . The b l a d e r e a c h e s i t s maximum l e n g t h sometime d u r i n g t h e summer and then b e g i n s t o decrease i n l e n g t h due t o e r o s i o n and d i s i n t e g r a t i o n o f t h e d i s t a l end. T h i s p r o c e s s c o n t i n u e s u n t i l t h e b l a d e r e a c h e s a minimum l e n g t h sometime i n t h e w i n t e r . L a b o r a t o r y c u l t u r e s i n d i c a t e t h a t endogenously d i s i n t e g r a t i o n i s more i m p o r t a n t t h a n t h i s process.  physical abrasion i n  I n l a b o r a t o r y t a n k s , t h e same a n n u a l c y c l e  o c c u r r e d i n t h e absence o f any water Druehl  controlled  motion.  (1968) s t a t e s t h a t L. l o n g i p e s i s " p e r e n n i a l from  the s t i p e and h o l d f a s t " , i n d i c a t i n g t h a t t h e b l a d e i s l o s t c o m p l e t e l y down t o t h e s t i p e e v e r y y e a r , as i s t h e case w i t h L_. s i n c l a i r i i .  The p r e s e n t s t u d y i n d i c a t e s t h a t a l t h o u g h t h e  b l a d e s a r e much r e d u c e d i n w i n t e r , t h e y a r e never lost.  There a r e t h r e e l i n e s o f e v i d e n c e f o r t h i s :  December, when L. s i n c l a i r i i  completely (1) I n  p l a n t s are u s u a l l y t o t a l l y devoid  48  of blades,  p l a n t s o f L. l o n g i p e s a t A a t s  short blades transplants  i n 1965 a n d 1966.  (2) i n l a b o r a t o r y t a n k s  t o Oregon L. l o n g i p e s never  completely.  (3) I n h e r b a r i u m  Bay a l l possessed  lost  i t s blades  s p e c i m e n s o f L_. l o n g i p e s  f r o m a l a r g e number o f l o c a t i o n s ( T a b l e X X I I I ) often of  a remnant o f the p r e v i o u s  the current year's Plants  information cultures into  i s available  show t h a t t h e s e  ripe  f o r other sori  Little  sori  produce  i s very  a t the d i s t a l  liberate  evidence  sporophytes  i n December.  w i n t e r months. spores  was  or i n the laboratory to indicate  normally  4.  blade  there  plants  end  blade.  o f L. l o n g i p e s bear  gametophytes.  field  year's  and i n  found  Laboratory  which  develop  either  i n the  that these  (See a l s o pp  No  gametophytes  69-73).  Discussion Aats  period:  twice  (Appendix are  B a y was v i s i t e d  I ) .  somewhat  detailed  only  i n the winter Consequently  limited.  five  t i m e s i n t h e two y e a r  and three  I t would be very  i n f o r m a t i o n on s e a s o n a l  it  w o u l d be v a l u a b l e t o know i f t h e r e  from  "moderately  Exposure subject very  changes i n seawater  u s e f u l t o have  more  collected  water  well  and s a l i n i t y .  Also,  i s a change i n sand  level.  at sites  ranging  exposed"  t o mean t h e e x t e n t  t o s u r f and other  difficult  here  temperature  sheltered" to "fully  i s understood  presented  growth o f t h e p l a n t s , as  seasonal  l o n g i p e s has been  i n t h e summer  the observations  as  L.  times  study  motion.  i n exposure  (Table X X I I I ) .  to which a beach i s This  t o p u t i n q u a n t i t a t i v e terms.  terminology i s Each  collector  has  49  described  h i s c o l l e c t i n g s i t e s i n terms o f h i s own concept o f  r e l a t i v e exposure. i n summer.  Most c o l l e c t i o n s i n A l a s k a have been made  I n some p l a c e s i n A l a s k a , p a r t i c u l a r l y t h e  A l e u t i a n I s l a n d s , there i s apparently between summer and w i n t e r  a very great  conditions.  difference  I f a s i t e was v i s i t e d  o n l y once, i t i s p o s s i b l e t h a t t h e c o n d i t i o n s were v e r y a t y p i c a l on t h a t p a r t i c u l a r day, w h i c h c o u l d l e a d t o an i n c o r r e c t e v a l u a t i o n o f exposure.  F o r many s t a t i o n s i n A l a s k a n o t  v i s i t e d by the author, i t i s impossible  t o s t a t e whether t h e  e x p o s u r e , as i t has been r e c o r d e d , i s r e p r e s e n t a t i v e . Most o f t h e " f u l l y exposed" s i t e s i n A l a s k a w h i c h t h e a u t h o r has v i s i t e d a r e n o t exposed t o as s e v e r e s u r f , o r on s u c h a r e g u l a r b a s i s , as any o f t h e s i t e s on t h e c o a s t o f Washington, Oregon, o r C a l i f o r n i a .  There a r e u n d o u b t e d l y  p l a c e s i n A l a s k a w h i c h a r e as exposed as most p l a c e s on t h e c o a s t from Washington t o C a l i f o r n i a .  However, most o f  t h e s e have n o t been v i s i t e d by c o l l e c t o r s , due t o t h e d i f f i c u l t y i n l a n d i n g on such beaches from a s h i p .  Future  c o l l e c t i o n s , perhaps made w i t h t h e a i d o f a h e l i c o p t e r i n o t h e r w i s e i n a c c e s s i b l e p l a c e s , may a l t e r t h e p r e s e n t p i c t u r e o f h a b i t a t d i s t r i b u t i o n o f some s p e c i e s . On t h e b a s i s o f p r e s e n t i n f o r m a t i o n , L. l o n g i p e s i s n o r m a l l y  found on beaches w h i c h  a r e l e s s exposed than those w h i c h L. s i n c l a i r i i i n h a b i t s .  50  VI.  EXPERIMENTAL ECOLOGY  A. 1.  F i e l d Work  Transplants In o r d e r to i n v e s t i g a t e the growth and s u r v i v a l o f  L. s i n c l a i r i i and L. l o n g i p e s i n o t h e r n a t u r a l environments, to  d e t e r m i n e whether m u c i l a g e  or s u p p r e s s e d  duct development c o u l d be  induced  by d i f f e r e n t temperatures, s e v e r a l f i e l d  t r a n s p l a n t s were c a r r i e d o u t . categories:  These f a l l i n t o t h r e e main  L_. s i n c l a i r i i from Oregon was  two beaches i n A l a s k a ; L_. l o n g i p e s was  t r a n s p l a n t e d to  t r a n s p l a n t e d from A l a s k a  to  two beaches i n Oregon; and b o t h s p e c i e s were t r a n s p l a n t e d  to  t h r e e beaches i n B r i t i s h Columbia.  F i g u r e 15 shows the  l o c a t i o n of a l l s i t e s u t i l i z e d i n transplant s t u d i e s . a.  A l a s k a beaches Two  experiments.  beaches i n A l a s k a were u t i l i z e d f o r t r a n s p l a n t A a t s Bay was  chosen because i t was  i n s i t u s t u d i e s on L. l o n g i p e s . A p p e n d i x I) was  the s i t e o f  Volga I s l a n d , S i t k a  chosen because i t was  (See  assumed t o have a  s i m i l a r water t e m p e r a t u r e t o A a t s Bay and i s e a s i l y from S i t k a .  reached  A t V o l g a i s l a n d , p l a n t s were a t t a c h e d to l o o s e  r o c k s w h i c h were p l a c e d i n t i d e p o o l s where o t h e r l o o s e r o c k s were p r e s e n t .  A t A a t s Bay the p l a n t s were a t t a c h e d t o the  r e e f on b a r e s p o t s among the p l a n t s o f L_. l o n g i p e s . In  and  the f i r s t t r a n s p l a n t attempt,  f i v e p l a n t s o f L.  s i n c l a i r i i were p l a c e d a t V o l g a I s l a n d i n June 1965.  A l l of  these were s u b s e q u e n t l y  five  lost.  In the second attempt,  51  F i g u r e 15  Transplant stations  p l a n t s were p l a c e d a t V o l g a i s l a n d and f o u r p l a n t s a t A a t s Bay i n December 1965. lost.  A l l o f those p l a c e d a t A a t s Bay were  Of those p l a c e d a t V o l g a I s l a n d , two were l o s t , one  remained b u t d i e d , and one remained a l i v e t o J u l y 1966.  It  showed some growth i n l e n g t h as w e l l as p r o d u c t i o n o f new stipes.  S e c t i o n s o f t h e s e new s t i p e s d i d n o t r e v e a l any  mucilage ducts. I n t h e t h i r d a t t e m p t , i n J u l y 1966, f o u r p l a n t s were p l a c e d a t A a t s Bay and f o u r a t V o l g a i s l a n d . r e v i s i t e d one month l a t e r . remained.  A a t s Bay was  A t t h i s time t h r e e o f the p l a n t s  Two showed no change.  One had grown 8 cm, the  same growth shown by L. l o n g i p e s p l a n t s marked and measured i n situ.  I n December 1966, two p l a n t s remained a t C o r o n a t i o n  I s l a n d , b u t i n v e r y poor c o n d i t i o n . at Volga i s l a n d a t t h i s time.  Two t r a n s p l a n t s were found  B o t h had a t t a c h e d t o t h e r o c k s  on w h i c h they were p l a n t e d and appeared h e a l t h y . b.  Oregon beaches I n d i a n Beach and S h o r t Sand Beach i n Oregon were  u t i l i z e d f o r transplant studies.  On b o t h beaches t h e p l a n t s  were a t t a c h e d t o bedrock i n a r e a s where L. s i n c l a i r i i already present.  was  I n the f i r s t a t t e m p t , f i v e p l a n t s o f  L. l o n g i p e s were p l a n t e d a t I n d i a n Beach i n August 1965. A l l of  t h e s e were l o s t d u r i n g t h e f o l l o w i n g month.  I n t h e second  a t t e m p t , f i v e more were p l a n t e d a t I n d i a n Beach i n December 1965.  A g a i n , a l l were l o s t i n t h e n e x t month.  In the t h i r d  attempt, two p l a n t s were p l a n t e d a t I n d i a n Beach i n May 1966. One o f t h e s e s u r v i v e d w i t h o u t growing t o June 1966 and then d i e d  53  In August 1966,  two p l a n t s o f L. l o n g i p e s were p l a n t e d a t  I n d i a n Beach, and two a t S h o r t Sand Beach.  In a d d i t i o n ,  one  L. s i n c l a i r i i from I n d i a n Beach was p l a n t e d a t S h o r t Sand Beach, and one from S h o r t sand Beach was p l a n t e d a t I n d i a n Beach.  One o f the L. l o n g i p e s a t I n d i a n Beach d i s a p p e a r e d the  f o l l o w i n g month.  The o t h e r showed s l i g h t growth d u r i n g  September and then was n o t found a g a i n .  The L.  sinclairii  from S h o r t Sand Beach grew d u r i n g September and October, i t s b l a d e s i n December, and d i s a p p e a r e d i n J a n u a r y . Sand Beach one L. l o n g i p e s and the L. s i n c l a i r i i  l o s t i n November.  At Short  remained  t h r o u g h November and were t o r n l o o s e i n December. L. l o n g i p e s was  lost  The o t h e r  When examined i n September,  a l l t h r e e had grown, b o t h o f the L. l o n g i p e s s l i g h t l y more than the L_. s i n c l a i r i i .  T h e r e a f t e r no e v i d e n c e o f growth was  found  and the b l a d e s became p r o g r e s s i v e l y s h o r t e r u n t i l the p l a n t s were t o r n l o o s e . In December 1966,  the f i n a l t r a n s p l a n t s were s t a r t e d .  Four p l a n t s o f L. l o n g i p e s were p l a c e d a t I n d i a n Beach and f o u r a t S h o r t Sand Beach. almost immediately.  Those a t I n d i a n Beach were l o s t  A t S h o r t Sand Beach, t h r e e remained  t h r o u g h March and then d i s a p p e a r e d .  Of t h e s e , one showed some  growth d u r i n g J a n u a r y and F e b r u a r y , then d e t e r i o r a t e d u n t i l i t was  lost.  The o t h e r s remained  unchanged t h r o u g h  then d e t e r i o r a t e d u n t i l t h e y washed away.  January,  i n January, when  the L. s i n c l a i r i i p l a n t s had no b l a d e s o r were j u s t b e g i n n i n g t o d e v e l o p new ones, the L. l o n g i p e s p l a n t s r e t a i n e d the l o n g b l a d e s o f the p r e v i o u s y e a r .  54  c.  B r i t i s h Columbia  beaches  Three beaches i n B r i t i s h Columbia were u t i l i z e d : W h i f f e n S p i t and R i v e r J o r d a n on t h e west c o a s t o f Vancouver I s l a n d , and S t a n l e y P a r k , Vancouver  (see Appendix  f i r s t two were chosen because L_. s i n c l a i r i i  I ) . The  had been found  t h e r e o c c a s i o n a l l y , a l t h o u g h t h e y a r e i n a much more s h e l t e r e d a r e a than t h e normal h a b i t a t o f L. s i n c l a i r i i .  The t h i r d  s i t e , S t a n l e y p a r k , i s c o m p l e t e l y s h e l t e r e d and has a lower s a l i n i t y than t h e open c o a s t h a b i t a t s . I t was chosen i n an a t t e m p t t o determine why h. found i n such h a b i t a t s .  sinclairii  and : L . l o n g i p e s a r e n o t  A t t h e B r i t i s h Columbia  stations,  e s p e c i a l l y a t S t a n l e y p a r k , t h e range o f temperature f o r b o t h a i r and seawater i s g r e a t e r than i n Oregon o r A l a s k a . In most e x p e r i m e n t s , p l a n t s were p l a c e d a t a l l t h r e e beaches a t n e a r l y t h e same t i m e .  To a v o i d c o n f u s i o n , t h e r e s u l t s f o r  each beach a r e c o n s i d e r e d s e p a r a t e l y . i.  Whiffen S p i t - A t Whiffen S p i t , a l l t r a n s p l a n t s  were a t t a c h e d t o l o o s e b o u l d e r s w h i c h were then p l a c e d a t t h e base o f t h e landward s i d e o f a l a r g e r o c k .  The r o c k p r o v i d e d  an easy means o f f i n d i n g t h e t r a n s p l a n t s on subsequent  visits.  In t h e f i r s t a t t e m p t , f o u r p l a n t s o f L. l o n g i p e s were p l a n t e d a t W h i f f e n S p i t i n J u l y 1965. A l l o f these were l o s t b e f o r e f u r t h e r o b s e r v a t i o n s were made.  I n November, two L. s i n c l a i r i i  were p l a n t e d and t h e s e d i s a p p e a r e d a l s o . f o u r L_. l o n g i p e s were p l a n t e d .  I n J a n u a r y 1966,  A l l remained i n p l a c e and  showed a l a r g e b l a d e growth and p r o d u c t i o n o f new s t i p e s t h r o u g h March.  When t h e s i t e was l a s t v i s i t e d i n August 1966,  55  two s t i l l remained.  They appeared h e a l t h y and had grown i n  the p r e v i o u s month.  One had a t t a c h e d t o t h e r o c k on w h i c h i t  was p l a n t e d .  T h i s l a t t e r was c o l l e c t e d and s e c t i o n e d .  evidence of mucilage  d u c t s was found.  One a d d i t i o n a l p l a n t  o f L. l o n g i p e s was p l a n t e d i n March 1966. June and then was l o s t . were p l a n t e d .  No  I t grew through  I n March 1966, f o u r L. s i n c l a i r i i  One d i s a p p e a r e d v e r y soon, one d i e d i n t h e f i r s t  month, and two showed good growth t h r o u g h May.  They s u r v i v e d  t h r o u g h June and then were l o s t . The growth o f b o t h t h e L. s i n c l a i r i i t r a n s p l a n t s and t h e L. l o n g i p e s t r a n s p l a n t s was v e r y g r e a t from March through relative  May  t o t h a t a t o t h e r seasons and on o t h e r beaches.  However, t h e L. l o n g i p e s t r a n s p l a n t s showed a s l i g h t l y g r e a t e r growth.  Both s p e c i e s produced wider b l a d e s than i n t h e i r  normal h a b i t a t .  There was no t w i s t i n g and t h e b l a d e s appeared  to have d e v e l o p e d n o r m a l l y .  A l l p l a n t s which s u r v i v e d a t  W h i f f e n S p i t were shaded and a l m o s t hidden by l a r g e p l a n t s o f Hedophyllum hanging  from t h e v e r t i c a l f a c e o f the l a r g e r o c k  above. ii.  R i v e r J o r d a n - A t R i v e r J o r d a n the t r a n s p l a n t s  were a t t a c h e d t o l o o s e b o u l d e r s from v a r i o u s p a r t s o f t h e beach w h i c h were then r e - p l a c e d .  Most o f these b o u l d e r s were  hidden under l a r g e p l a n t s o f E g r e g i a m e n z i e s i i and P h y l l o s p a d i x s c o u l e r i w h i c h were a t t a c h e d nearby and l a y a c r o s s l a r g e adjacent areas. planted. this  I n November 1965, two L. s i n c l a i r i i were  These were n o t found a g a i n u n t i l March 1966.  At  time the b l a d e s were s h o r t e r o r absent and t h e r e was no  evidence  t h a t growth had o c c u r r e d .  When examined a g a i n i n May  56  t h e r e were s e v e r a l new  s t i p e s and c o n s i d e r a b l e b l a d e growth  had o c c u r r e d .  One  In March 1966,  two o t h e r _L. s i n c l a i r i i were p l a n t e d .  examined i n May  o f the p l a n t s had a t t a c h e d t o the r o c k .  t h e s e showed about the same b l a d e growth as  t r a n s p l a n t s s e t o u t i n November and s e v e r a l new present.  When  s t i p e s were  I n June one o f the p l a n t s had a t t a c h e d to the r o c k ,  the o t h e r was  dead.  In January  1966,  were t r a n s p l a n t e d to R i v e r J o r d a n . w e l l t h r o u g h June  two p l a n t s o f L_. l o n g i p e s  These remained and grew  1966.  As a t W h i f f e n S p i t , b o t h s p e c i e s produced wider  blades  a f t e r b e i n g t r a n s p l a n t e d b u t o t h e r w i s e appeared h e a l t h y and "normal".  A l l p l a n t s a t R i v e r J o r d a n w h i c h grew were  p r o t e c t e d from d i r e c t i n s o l a t i o n by l a r g e f r o n d s o f E g r e g i a m e n z i e s i i l y i n g on top o f them. iii  S t a n l e y p a r k - The  t r a n s p l a n t s a t S t a n l e y park  were a t t a c h e d t o l o o s e b o u l d e r s w h i c h were then p l a c e d near one  l a r g e r o c k f o r ease i n f i n d i n g on subsequent v i s i t s .  L. s a c c h a r i n a o c c u r s above and b e l o w the a r e a used f o r transplanting.  I n the f i r s t t r a n s p l a n t experiment,  L. s i n c l a i r i i were p l a n t e d i n November 1965. disappeared. growing  The o t h e r s u r v i v e d t o June 1966.  One  of  two these  I t continued  the whole time a c c o r d i n g to measurements between  punched h o l e s .  I t a t t a c h e d t o the r o c k on w h i c h i t was  The b l a d e s were never c o m p l e t e l y l o s t .  They d e c r e a s e d i n  l e n g t h t o March and then began i n c r e a s i n g i n l e n g t h . experiment  w i t h L_. s i n c l a i r i i was  s i x p l a n t s were t r a n s p l a n t e d .  planted.  The  second  s t a r t e d i n March 1966 when  Of t h e s e , one d i e d , one  grew  57  t h r o u g h May, and f o u r grew t h r o u g h June.  Of the l a t t e r  group,  one had a t t a c h e d t o t h e r o c k by May. F i v e p l a n t s o f L. l o n g i p e s were t r a n s p l a n t e d t o S t a n l e y P a r k i n August 1965.  A l l o f t h e s e d i e d v e r y soon.  L. l o n g i p e s were t r a n s p l a n t e d i n December 1965.  Two more  They grew  f a i r l y w e l l t h r o u g h May and then d i e d i n June. In w i n t e r t h e L. l o n g i p e s appeared t o be h e a l t h i e r the  than  L. s i n c l a i r i i , b u t as summer approched, t h e L_. s i n c l a i r i i  appeared h e a l t h i e r .  B o t h s p e c i e s produced b l a d e s c o n s i d e r a b l y  b r o a d e r than i n t h e i r normal h a b i t a t s .  They became v e r y  t w i s t e d and overgrown w i t h b r y o z o a n s , diatoms, and v a r i o u s other e p i p h y t i c algae. d.  Discussion I t was o b s e r v e d t h a t no m u c i l a g e d u c t s were p r e s e n t  i n new s t i p e s produced by L. s i n c l a i r i i p l a n t s a t V o l g a I s l a n d . T h i s would seem a t f i r s t t o c o n f i r m t h e o r i g i n a l i d e a t h a t m u c i l a g e d u c t f o r m a t i o n r e q u i r e s a temperature h i g h e r than t h a t normally encountered i n Southeast A l a s k a . new s t i p e s w h i c h were s e c t i o n e d were v e r y s m a l l .  However, t h e Subsequently,  e q u a l l y s m a l l s t i p e s from p l a n t s t a k e n d i r e c t l y from t h e f i e l d i n Oregon were s e c t i o n e d . ducts e i t h e r .  These d i d n o t show any m u c i l a g e  I t i s c o n c l u d e d t h a t t h e s t i p e s o f L. s i n c l a i r i i  must a t t a i n a c e r t a i n minimum s i z e b e f o r e m u c i l a g e d u c t s a r e produced and t h i s s i z e was n o t a t t a i n e d by any o f the new s t i p e s produced i n A l a s k a . The A l a s k a n s i t e s , w i t h one e x c e p t i o n a t A a t s Bay i n 1966, were o n l y v i s i t e d a t s i x month i n t e r v a l s .  Most o f t h e p l a n t s  58  were l o s t b e f o r e any growth:measurement c o u l d be made.  i t is  q u i t e p o s s i b l e t h a t i f t h e A l a s k a n s i t e s had been v i s i t e d as o f t e n as t h e o t h e r s a comparable number o f growth measurements m i g h t have been o b t a i n e d b e f o r e l o s s o f t h e p l a n t s . Attachment o f t h e h o l d f a s t t o t h e r o c k s on w h i c h the p l a n t s had been p l a c e d o c c u r r e d o n l y a t S t a n l e y p a r k L. s i n c l a i r i i ) R i v e r J o r d a n  (two L. s i n c l a i r i i ) ;  (three  Whiffen S p i t  (two L_. s i n c l a i r i i , one L. l o n g i p e s ) ; and V o l g a I s l a n d (one L. s i n c l a i r i i ) .  None o f t h e s e p l a c e s i s v e r y exposed, w h i c h  i s p r o b a b l y an i m p o r t a n t f a c t o r .  More i m p o r t a n t , however, i s  the method o f p l a n t i n g employed.  On a l l o f these s i t e s ,  loose  b o u l d e r s a r e p r e s e n t and t h e p l a n t s were f i x e d t o these by means o f rubber bands.  T h i s method h o l d s the b o t t o m o f the  h o l d f a s t f i r m l y a g a i n s t the rock.  I n t h e more exposed a r e a s ,  w h i c h l a c k l o o s e b o u l d e r s , the method o f f a s t e n i n g t h e p l a n t s to  s p i k e s i n t h e r o c k o n l y h e l d the o u t p l a n t s from washing  away b u t d i d n o t h o l d them i n any one p o s i t i o n f o r any l e n g t h of time.  S e v e r a l u n s u c c e s s f u l attempts  were made t o d e v i s e  a b e t t e r p l a n t i n g method. The r e s u l t s from t h e t r a n s p l a n t s t o S t a n l e y p a r k , S p i t , and R i v e r J o r d a n i n d i c a t e t h a t L. s i n c l a i r i i  i s less  a d v e r s e l y a f f e c t e d by h i g h summer seawater temperatures i s L. l o n g i p e s .  Whiffen  than  The r e s u l t s from Oregon and A l a s k a a r e n o t  as c l e a r c u t . However the summer temperatures a r e n o t as h i g h at  t h e s e s i t e s as i n Oregon o r A l a s k a , e i t h e r .  59  2.  Rock c l e a r i n g a.  Introduction A t v a r i o u s times over a p e r i o d o f one y e a r , p o r t i o n s  of  r o c k s were c l e a r e d a t I n d i a n Beach and S h o r t Sand Beach.  The o b j e c t o f t h e s e experiments  was t o determine whether t h e  a d j a c e n t L. s i n c l a i r i i p l a n t s would c o l o n i z e t h e c l e a r e d a r e a s . b.  Methods In a l l e x p e r i m e n t s , p o r t i o n s o f t h e L. s i n c l a i r i i  h o l d f a s t s were f i r s t removed w i t h a k n i f e .  The a r e a o f r o c k  thus exposed was then s c r a p e d and f i n a l l y burned t o w i t h i n 5 cm o f t h e r e m a i n i n g p l a n t s .  T h e r e a f t e r t h e c l e a r i n g s were  examined a t monthly i n t e r v a l s as f a r as p o s s i b l e . c.  Results In  t h e f i r s t experiment,  a s t r i p about 35 cm wide was  c l e a r e d between two l a r g e clumps o f L. s i n c l a i r i i a t I n d i a n Beach i n May 1966.  I n June a s l i g h t growth o f h a p t e r a  toward t h e c l e a r i n g from b o t h s i d e s was e v i d e n t . some h a p t e r a had grown 2 cm onto t h e c l e a r e d a r e a .  tips  I n August By October  the c l e a r e d a r e a had been c o m p l e t e l y c r o s s e d by h a p t e r a  from  b o t h s i d e s and some s t i p e s and b l a d e s had been produced  from  these haptera. The second c l e a r i n g a t I n d i a n Beach was made i n December 1966.  I n F e b r u a r y , t h e r e was no e v i d e n c e o f growth o f h a p t e r a  nor was t h e r e a n y t h i n g e l s e growing on t h e c l e a r e d a r e a .  In  March, two p l a n t s o f Hedophyllum s e s s i l e each 15 cm l o n g , one plant of A l a r i a  (marginata ?) 12 cm l o n g and s e v e r a l c r u s t o s e  c o r a l l i n e a l g a e were p r e s e n t .  60  The  t h i r d c l e a r i n g e x p e r i m e n t a t I n d i a n Beach was s t a r t e d  i n A p r i l 1967. A s t r i p was c l e a r e d from 10 cm above t h e sand on  one s i d e o f a r o c k , up over t h e t o p and down t o w i t h i n 10  cm o f t h e sand on t h e o t h e r s i d e ( F i g . 16) . removed from t h e r o c k .  S i x s p e c i e s were*  L_j_ s i n c l a i r i i c o v e r e d most o f t h e a r e a  w h i c h was c l e a r e d , b u t t h e r e were o c c a s i o n a l gaps i n t h e h o l d f a s t w h i c h p r o v i d e d attachment space f o r o t h e r s p e c i e s . On t o p o f t h e r o c k P h a e o s t r o p h i o n most abundant s p e c i e s .  i r r e q u l a r e was t h e second  There were a l s o s e v e r a l p l a n t s o f  B o s s i e l l a plumosa and an u n i d e n t i f i e d c r u s t o s e c o r a l l i n e a l g a on t o p .  On b o t h s i d e s H i l d e n b r a n d i a s p . and B o s s i e l l a plumosa  were found.  On one s i d e , e x t e n d i n g down t o b e l o w t h e sand  l e v e l , was an e x t e n s i v e mat o f Codium s e t c h e l l i i .  I n May,  the h a p t e r a a t t h e edges o f t h e L. s i n c l a i r i i h a d s t a r t e d g r o w i n g toward t h e c l e a r e d a r e a .  S e v e r a l patches o f Hildenbrandia  and a few t u f t s o f B o s s i e l l a were e v i d e n t  (Fig. 16a). For  the f o l l o w i n g t h r e e months t h e r o c k was c o m p l e t e l y b u r i e d under sand ( F i g . 16 b , c , d ) . the r o c k .  I n September t h e sand r e c e d e d ,  exposing  A t t h i s time t h e L. s i n c l a i r i i and Codium s e t c h e l l i i  appeared h e a l t h y and unchanged from t h e June c o n d i t i o n . Otherwise,  o n l y some d y i n g t u f t s o f B o s s i e l l a plumosa were noted.  A t S h o r t Sand Beach, t h e f i r s t c l e a r i n g experiment was s t a r t e d i n November 1966.  By March 1967, s e v e r a l h a p t e r a o f  L. s i n c l a i r i i had encroached onto t h e c l e a r e d a r e a and were p u t t i n g up s t i p e s and b l a d e s i n t h e a r e a . c o r a l l i n e a l g a e were a l s o p r e s e n t .  Several t u f t s of  61  F i g u r e 16  a.  C l e a r e d r o c k a t I n d i a n Beach  Rock C i n May 1967, one month a f t e r was c l e a r e d .  b.  Rock C i n June 1967.  c.  Rock C i n J u l y 1967.  d.  Rock C i n August 1967.  strip  62  The second experiment a t S h o r t Sand Beach was begun i n A p r i l 1967. for  A v e r t i c a l s t r i p was c l e a r e d n e x t t o the p l a n t s  w h i c h growth was b e i n g measured i n s i t u .  The r o c k a t  t h i s l o c a t i o n i s s o f t sandstone w i t h many h o l e s produced various animals.  by  The s u r f a c e was c l e a r e d i n the u s u a l manner,  b u t the h o l e s were n o t touched.  One month l a t e r numerous  tube worms were seen p r o t r u d i n g from the h o l e s i n the r o c k . By J u l y s e v e r a l e n c r u s t i n g a n i m a l s had e s t a b l i s h e d  themselves  on t h e r o c k b u t no p l a n t s were p r e s e n t . d.  Discussion S e v e r a l o f the c l e a r i n g e x p e r i m e n t s were c a r r i e d o u t  at  t i m e s when the nearby L. s i n c l a i r i i p l a n t s bore r i p e  Under of  sori.  such c i r c u m s t a n c e s i t seems l i k e l y t h a t gametophytes  L. s i n c l a i r i i s h o u l d have e s t a b l i s h e d themselves on the  cleared areas.  A l t h o u g h the gametophytes would n o t have been  v i s i b l e t h e y s h o u l d have produced s p o r o p h y t e s which would have been e a s i l y r e c o g n i z e d .  However, on a l l the c l e a r e d a r e a s  and s e v e r a l o t h e r r o c k s o b s e r v e d on these beaches over a p e r i o d o f two y e a r s , no young s p o r o p h y t e s o f L.  sinclairii  were ever o b s e r v e d w h i c h had d e f i n i t e l y a r i s e n from gametophytes.  I n e v e r y i n s t a n c e , c a r e f u l e x a m i n a t i o n showed  t h a t the young s p o r o p h y t e s had a r i s e n from h a p t e r a o f o l d e r p l a n t s and thus were outgrowths o f t h e o l d e r p l a n t s . n o t e d p r e v i o u s l y , two Hedophyllum  As  p l a n t s and an A l a r i a became  e s t a b l i s h e d on the r o c k a l t h o u g h t h e r e were r e l a t i v e l y few o f these p l a n t s very c l o s e .  This evidence i n d i c a t e s that  p r o d u c t i o n o f L. s i n c l a i r i i s p o r o p h y t e s by s e x u a l means i s r a r e .  63  On the o t h e r hand, the e x p e r i m e n t s show t h a t c o l o n i z a t i o n of  new a r e a s by o u t g r o w t h o f h a p t e r a o f L. s i n c l a i r i i  takes  p l a c e r e a d i l y , a t l e a s t d u r i n g the p e r i o d when the p l a n t s a r e n o t b u r i e d under 3.  sand.  T r a n s i t i o n zone e x p e r i m e n t s a.  Introduction In most members o f the L a m i n a r i a l e s the t r a n s i t i o n  zone between t h e s t i p e and b l a d e i s g e n e r a l l y s t a t e d t o be the p r i n c i p a l . r e g i o n o f growth. be the case i n L. s i n c l a i r i i .  I t appeared t h a t t h i s might n o t A c c o r d i n g l y , two t y p e s o f  e x p e r i m e n t s were conducted i n the f i e l d t o determine the i m p o r t a n c e o f the t r a n s i t i o n zone. of  The f i r s t o f t h e s e c o n s i s t e d  c u t t i n g o f f s t i p e s w e l l b e l o w the t r a n s i t i o n zone and  o b s e r v i n g t h e b e h a v i o r o f the r e m a i n i n g h o l d f a s t s and stubs.  stipe  T h i s was done t w i c e a t I n d i a n Beach and once a t S h o r t  Sand Beach.  The second e x p e r i m e n t c o n s i s t e d o f m a r k i n g o f f  r e g u l a r i n t e r v a l s on a p l a n t and then o b s e r v i n g where the g r e a t e s t growth o c c u r r e d . b.  T h i s was done once a t I n d i a n Beach.  p r o c e d u r e s and r e s u l t s In  the f i r s t e x p e r i m e n t s i n w h i c h t r a n s i t i o n  zones  were removed, s t i p e s were c u t a t b o t h I n d i a n Beach and S h o r t Sand Beach i n October 1966.  A t I n d i a n Beach a l l the s t i p e s  i n a l a r g e clump on top o f one r o c k were c u t o f f a t l e a s t 5 cm b e l o w the t r a n s i t i o n zone, u s i n g a l a r g e p a i r o f s h e a r s .  At  S h o r t Sand Beach t h e same p r o c e d u r e was f o l l o w e d e x c e p t t h a t the it.  clump was f i r s t i s o l a t e d by c l e a r i n g a s t r i p o f r o c k around  64  In November, the s t i p e s t u b s on b o t h beaches appeared be h e a l i n g o v e r .  to  I n a d d i t i o n , a t S h o r t Sand Beach t h e r e was  c o n s i d e r a b l e growth o f h a p t e r a a t the margins o f the clump. L a t e i n December many o f the c u t s t i p e s on b o t h beaches had produced new b l a d e s from the t i p s i n an a p p a r e n t l y normal manner.  A t t h i s t i m e , a few o f the normal uncut s t i p e s a t  I n d i a n Beach had b l a d e s w i t h r i p e s o r i , b u t the m a j o r i t y had no b l a d e s a t a l l .  A t S h o r t Sand Beach a l l the uncut  were w i t h o u t b l a d e s .  stipes  I n J a n u a r y no f u r t h e r changes i n the  c u t s t i p e s were n o t e d .  I n F e b r u a r y the b l a d e s were s l i g h t l y  l o n g e r than b e f o r e and t h e r e a f t e r no f u r t h e r changes were n o t e d at either  beach.  I n F e b r u a r y 1967, a t I n d i a n Beach.  t h e second c u t t i n g experiment was  started  T h i s time a l l the l a r g e s t i p e s i n one group  were c u t o f f w e l l b e l o w the t r a n s i t i o n zone, b u t a l l the p r e s e n t y e a r ' s s t i p e s and b l a d e s were l e f t untouched. s t i p e s o f t h e p r e s e n t y e a r a r e e a s i l y r e c o g n i z e d , as  (The previously  noted.) A f t e r one month t h e c u t s t i p e s showed no change.  The  u n c u t s m a l l b l a d e s had grown about 3 cm p a s t the c u t ends w i t h a t o t a l b l a d e l e n g t h o f 6-7  cm.  By t h e end o f A p r i l about  20 p e r c e n t o f the c u t s t i p e s had produced new b l a d e s and r e s t were unchanged.  I n June no more c u t s t i p e s had  the  produced  b l a d e s , b u t the growth o f h a p t e r a a t t h e margins o f the clump seemed t o be g r e a t e r than An experiment was  s t a r t e d i n May  normal.  t o determine the r e g i o n o f g r e a t e s t growth 1967  a t I n d i a n Beach.  I n a clump o f s t i p e s  where s e v e r a l s t i p e s and b l a d e s were b e i n g measured  65  p e r i o d i c a l l y , f o u r b l a d e s were punched w i t h many h o l e s . two o f t h e s e , the h o l e s were punched e v e r y 2 cm, 4 cm above the base o f the b l a d e .  In  beginning  I n the o t h e r two,  the h o l e s  were punched e v e r y 2 cm, b e g i n n i n g a t 2 cm above the b a s e . Both s t i p e s and b l a d e s were measured, b u t no c o n v e n i e n t  way  was  1967,  found t o mark o f f i n t e r v a l s on the s t i p e s .  o n l y one o f the f o u r p l a n t s was  In June  found, one o f those w h i c h had  been punched from 4 cm above the base ( F i g . 1 2 c ) . was  e r o d e d a t the end, undoubtedly  the t o t a l l e n g t h was  The  blade  weakened by the h o l e s , and  l e s s than a t the b e g i n n i n g .  The  total  growth o f the b l a d e as c a l c u l a t e d  from the t o t a l growth between  a l l the h o l e s was  (Table X X I I ) .  a t l e a s t 6.3  cm  i n c r e a s e d 1 cm i n the same p e r i o d .  The  stipe  The r e s u l t s show t h a t the  g r e a t e s t growth i s i n the t r a n s i t i o n zone.  The  t o t a l blade  g r o w t h i s l e s s than 50 p e r c e n t o f the average f o r o t h e r b l a d e s i n the same p l a c e a t the same time i s p r o b a b l y due  (Table X X I ) , w h i c h  t o the i n j u r i o u s e f f e c t s o f the l a r g e number  of holes. C.  Discussion Setchell  (1905), s t u d y i n g growth and r e g e n e r a t i o n i n  L. s i n c l a i r i i , n o t e d t h a t a b l a d e was produced from a wound on the s i d e o f a s t i p e . L. s i n c l a i r i i  From h i s s t u d i e s he p o s t u l a t e d t h a t  c o u l d r e g e n e r a t e from the stump o f a s t i p e .  p r e s e n t experiments  show t h a t t h i s i s c o r r e c t .  The  The  results  show t h a t a l t h o u g h growth i s g r e a t e s t i n the t r a n s i t i o n zone, i t i s n o t l i m i t e d t o t h i s zone and the zone i s not f o r the p r o d u c t i o n o f new  blades.  necessary  66  Under normal c o n d i t i o n s t h e b l a d e s a r e l o s t i n December and r e g e n e r a t e d i n J a n u a r y .  On t h e s t i p e s which were c u t i n  O c t o b e r , new b l a d e s appeared i n December, about one month e a r l i e r than n o r m a l .  However, a f t e r t h i s e a r l y b e g i n n i n g t h e  b l a d e s showed l i t t l e f u r t h e r development,  a t l e a s t i n that year.  Removal o f the t r a n s i t i o n zone a l s o seemed t o s t i m u l a t e t h e growth o f h a p t e r a .  I t i s p o s s i b l e t h a t these r e s u l t s i n d i c a t e  t h e p r e s e n c e o f an a u x i n o r o t h e r s u b s t a n c e i n the o l d b l a d e w h i c h i n h i b i t s new b l a d e f o r m a t i o n and growth o f h a p t e r a . However, such a p o s s i b i l i t y has n o t been i n v e s t i g a t e d , and a t p r e s e n t no adequate e x p l a n a t i o n o f t h e s e phenomena i s available. B. 1.  L a b o r a t o r y Work  Sporophytes Sporophytes o f L. s i n c l a i r i i and L. l o n g i p e s were c u l t u r e d  for  v a r y i n g p e r i o d s o f time i n the 120 iT'tanks o f seawater  previously described.  The c u l t u r e s were p l a n n e d t o determine  the e f f e c t s o f d i f f e r e n t t e m p e r a t u r e s , s a l i n i t i e s ,  and  p h o t o p e r i o d on the growth and r e p r o d u c t i o n o f the p l a n t s .  New  p l a n t s were c o l l e c t e d and i n t r o d u c e d i n t o t h e tanks s e v e r a l times d u r i n g a two-year p e r i o d ,  p l a n t s were removed whenever  t h e y showed no change f o r a p e r i o d o f t h r e e months o r were obviously deteriorating, a.  photoperiod P l a n t s o f b o t h s p e c i e s were grown a t p h o t o p e r i o d s  of  16 hours l i g h t / 8 hours dark, 12 hours l i g h t / 1 2 hours dark,  67  and 8 hours l i g h t / 1 6 hour d a r k ,  p l a n t s were grown i n each  p h o t o p e r i o d f o r up t o t h r e e months under each regime  (5°, 8°, and 10°C).  temperature  The p h o t o p e r i o d s were u s u a l l y s i x  months o u t o f phase w i t h t h o s e found i n n a t u r e , i . e . , s h o r t days i n summer.  The v a r i o u s p h o t o p e r i o d s d i d n o t have any  n o t i c e a b l e e f f e c t on e i t h e r growth o r r e p r o d u c t i o n o f p l a n t s of  t h e two s p e c i e s . A f t e r c o n c l u s i o n o f t h e p h o t o p e r i o d  e x p e r i m e n t s a l l c u l t u r e s were r u n a t 16 hours l i g h t / 8  hours  dark. b.  Salinity, The S t a n l e y p a r k seawater was used as low s a l i n i t y  water and t h e West Coast seawater was used as h i g h s a l i n i t y water, 10°C.  p l a n t s were grown i n each type o f water a t 5°, 8° and No d i f f e r e n c e was e v i d e n t i n growth o f t h e p l a n t s i n  t h e d i f f e r e n t s a l i n i t i e s a t t h e same t e m p e r a t u r e . c.  Temperature V a r i o u s p l a n t s were grown a t 5°, 8°, and 10°C over  a p e r i o d o f two y e a r s .  D u r i n g t h e f i r s t n i n e months t h e  p h o t o p e r i o d was changed e v e r y t h r e e months. was m a i n t a i n e d a t 16 h o u r s l i g h t / B h o u r s d a r k . o f b o t h s p e c i e s o c c u r r e d somewhat e a r l i e r than a t 8° o r 10°C.  Thereafter i t Maximum growth  i n t h e y e a r a t 5°C  The maximum growth o f L. s i n c l a i r i i was  i n March a t 5°C and i n June a t 8° and 10°c.  The maximum  growth o f L. l o n g i p e s o c c u r r e d i n J a n u a r y a t 5°C and i n F e b r u a r y a t 8° and 10°C. t e m p e r a t u r e s appeared growth o f t h e p l a n t s .  With these e x c e p t i o n s , the d i f f e r e n t  t o have v e r y l i t t l e e f f e c t on t h e  68  I n a l l p l a n t s c u l t u r e d i n t h e t a n k s , c e r t a i n common f e a t u r e s were e v i d e n t .  No v a r i a t i o n s i n t e m p e r a t u r e ,  salinity,  or p h o t o p e r i o d had any e f f e c t on p r o d u c t i o n o f s o r i o r l o s s and r e g e n e r a t i o n o f b l a d e s .  S o r i were produced on L.  s i n c l a i r i i p l a n t s c o l l e c t e d i n l a t e September a f t e r one week i n culture.  However, o t h e r p l a n t s o f t h e same p o p u l a t i o n became  r i p e a t t h e same time i n n a t u r e .  B l a d e l o s s always o c c u r r e d  a t t h e same time as i n n a t u r e u n l e s s t h e l o s s was due t o t h e plant's dying.  Likewise, blade regeneration occurred a t  t h e same time as i n n a t u r e a t a l l t e m p e r a t u r e s , a l t h o u g h t h e r a t e o f growth o f t h e new b l a d e s v a r i e d w i t h t e m p e r a t u r e . new b l a d e s were produced,  the " c o l l a r s "  After  ( S e t c h e l l , 1896)  formed by t h e f r a y e d ends o f t h e s p l i t s t i p e ends were much more o b v i o u s t h a n i n t h e f i e l d , presumably because t h e r e was no wave a c t i o n t o remove t h e e x c e s s  tissue.  G e n e r a l l y , t h e p l a n t s w h i c h grew i n t h e l a b o r a t o r y t a n k s grew w e l l and showed normal b e h a v i o u r f o r two t o f o u r months and then ceased showing b l a d e growth.  A t t h i s time some o f  them then began t o d e t e r i o r a t e ; b u t i n most, growth o f h a p t e r a and p r o d u c t i o n o f new s t i p e s and b l a d e s from t h e h o l d f a s t s c o n t i n u e d f o r up t o 15 months a f t e r b l a d e had c e a s e d .  growth  Many o f t h e s e h a p t e r a were removed and used i n  t h e e x p e r i m e n t s on h a p t e r a d e s c r i b e d below.  A l t h o u g h new  s t i p e s were produced from t h e h a p t e r a , no growth was r e c o r d e d on any o f t h e s t i p e s w h i c h were a l r e a d y p r e s e n t on p l a n t s when b r o u g h t i n t o the l a b o r a t o r y .  69  d.  Discussion The  tank r e s u l t s a r e  v a l u a b l e i n t h a t they  indicate  t h a t p r o d u c t i o n o f s o r i and b l a d e l o s s and r e g e n e r a t i o n a r e n o t c o n t r o l l e d by temperature,  s a l i n i t y , or photoperiod.  However, t h e r e s u l t s i n t h e tanks cannot g i v e a v e r y good p i c t u r e o f t h e growth o f t h e p l a n t s .  The p l a n t s a p p a r e n t l y  behaved n o r m a l l y f o r a s h o r t time a f t e r b e i n g p l a c e d i n t h e tanks.  However,  enough  t o g i v e a t r u e p i c t u r e o f growth over a whole y e a r .  The  p l a n t s were n o t p l a c e d i n the t a n k s o f t e n  t a n k s c o n t a i n e d water v e r y s i m i l a r t o t h a t i n w h i c h t h e  p l a n t s n o r m a l l y grow, b u t t h i s water was o n l y changed once a month.  T h i s a l l o w e d a much g r e a t e r b u i l d u p o f waste p r o d u c t s  t h a n o c c u r r e d i n n a t u r e and p r o b a b l y a c o n s i d e r a b l e d e p l e t i o n of various n u t r i e n t s .  T h i s , and t h e l a c k o f t i d e s and wave  a c t i o n , make t h e t a n k s v e r y poor s u b s t i t u t e s f o r the n a t u r a l conditions. 2.  Gametophytes a.  Introduction A t t h e b e g i n n i n g o f t h i s s t u d y i t appeared t h a t  L. s i n c l a i r i i and L. l o n g i p e s were p o s s i b l y ecotypes o f the same s p e c i e s . to  I n o r d e r t o t e s t t h i s h y p o t h e s i s , i t was p l a n n e d  c r o s s t h e two s p e c i e s .  To a c c o m p l i s h t h i s , i t was n e c e s s a r y  to o b t a i n s e x u a l l y mature gametophytes i n c u l t u r e . Gametophytes o f L. s i n c l a i r i i were c u l t u r e d by Myers In  (1925).  her c u l t u r e s , t h e gametophytes produced s p o r o p h y t e s ,  indicating a typical kelp l i f e history.  thus  However, she n o t e d t h a t  70  the gametophytes c o n t i n u e d t o grow v e g e t a t i v e l y and, a f t e r s i x months, were l a r g e r than any o f the sporophytes culture.  produced i n  So f a r as i s known, the gametophytes o f L.  longipes  have never been c u l t u r e d p r e v i o u s l y . b.  M a t e r i a l s and Methods S p o r o p h y t e s b e a r i n g r i p e s o r i were c o l l e c t e d a t  v a r i o u s times d u r i n g the w i n t e r s o f 1965, (L.  l o n g i p e s was  the l a b o r a t o r y .  1966  and  1967  o n l y c o l l e c t e d i n December) and b r o u g h t i n t o They were then k e p t i n the dark i n a m o i s t  p l a s t i c bag a t 5°, 8°, or 10° C f o r f o u r t o 36 hours, paper t o w e l f o r two t o e i g h t hours a t the same  then i n a  temperature.  A f t e r t h i s p e r i o d o f p a r t i a l d r y i n g the s o r i were wiped c a r e f u l l y t o remove e p i p h y t i c contaminants and then c u t i n t o small pieces with a razor blade. SWF  These p i e c e s were p l a c e d i n  or ES i n 250 ml g l a s s c u l t u r e d i s h e s or s t a n d a r d 100  P e t r i dishes.  The  d i s h e s were p l a c e d i n 5°, 8°, or 10°C  20 f t - c or 150 f t - c .  mm under  A f t e r i n t e r v a l s w h i c h rangedfrom s i x  hours t o s i x days i n v a r i o u s e x p e r i m e n t s ( a p p a r e n t l y depending upon the s t a g e o f development o f the s o r u s a t the time o f c o l l e c t i o n ) zoospores c o u l d be seen swarming from the p i e c e s of sorus.  S e v e r a l m i l l i l i t e r s o f the spore s u s p e n s i o n  p i p e t t e d i n t o each c u l t u r e d i s h . d i f f e r e n t combinations Five SWF,  were  A l a r g e number o f  o f c u l t u r e c o n d i t i o n s were t e s t e d .  d i f f e r e n t media were employed a t v a r i o u s t i m e s :  ES, ES+,  and ASP  2.  Large g l a s s c u l t u r e dishes  standard glass P e t r i dishes  (250 m l ) ,  (100mm) and s m a l l (60mm) P e t r i  d i s h e s o f b o t h g l a s s and p l a s t i c were used.  SW,  In v a r i o u s  71  e x p e r i m e n t s the medium was 10,' 14, or 28 days. year.  changed a t i n t e r v a l s o f 1, 2, 4,  Most c u l t u r e s were k e p t a t l e a s t  7,  one  A f t e r a c u l t u r e had r u n f o r two t o t h r e e months the  medium was u s u a l l y changed o n l y once a month.  The  were m a i n t a i n e d a t 5°, 8°, 10°, and 15°C  light  under  cultures  i n t e n s i t i e s o f 20 f t - c o r 150 f t - c p r o v i d e d by c o o l w h i t e fluorescent lights.  Three p h o t o p e r i o d s were t e s t e d :  8 hours  l i g h t / 1 6 hours dark, 12 hours l i g h t / 1 2 hour dark, and 16 l i g h t / 8 hours d a r k .  hours  I n one experiment gametophyes o f v a r i o u s  ages were m a i n t a i n e d f o r p e r i o d s o f one day t o two weeks i n t o t a l darkness.  C u l t u r e s were a l s o k e p t i n an e a s t - f a c i n g  window where the temperature v a r i e d from 10° to 25°C.  Some  c u l t u r e s were r u n on the shaker i n the p s y c r o t h e r m i n c u b a t o r . I n s e v e r a l i n s t a n c e s c u l t u r e s were a l s o s u b j e c t e d t o changes o f t e m p e r a t u r e , media, f r e q u e n c y o f media change, l i g h t i n t e n s i t y , p h o t o p e r i o d and v a r i o u s c o m b i n a t i o n s o f these f a c t o r s . I t was n o t p o s s i b l e t o c a r r y o u t any c r o s s i n g e x p e r i m e n t s , "for v-easot\s hottd below, c. R e s u l t s Under s e v e r a l d i f f e r e n t c o n d i t i o n s s p o r o p h y t e s were produced by L. s i n c l a i r i i gametophytes, few i n any one c u l t u r e . sporophytes t w i c e :  b u t never more than a  L. l o n g i p e s gametophytes o n l y produced  once i n 1966 a f t e r 10 weeks i n ES a t  under 20 f t - c ( f i v e s p o r o p h y t e s ) and once i n 1967 a f t e r weeks i n ES+ a t 5°C under 150 f t - c ( t h r e e s p o r o p h y t e s ) .  10°C 24 The  v a r i o u s d i f f e r e n c e s i n c u l t u r e c o n d i t i o n s seemed t o have l i t t l e e f f e c t on t h e p r o d u c t i o n o f s p o r o p h y t e s .  I n the  ES+  medium the gametophytes had l a r g e r , more d e e p l y pigmented t h a n i n the o t h e r media.  Longer and l e s s d e e p l y  pigmented  cells  72  c e l l s were p r o d u c e d under h i g h e r l i g h t i n t e n s i t i e s . the of  gametophytes had fewer and s m a l l e r c e l l s .  At  5°c  However, none  t h e s e d i f f e r e n c e s appeared t o a f f e c t t h e p r o d u c t i o n o f  o o g o n i a , a n t h e r i d i a , and u l t i m a t e l y , s p o r o p h y t e s .  Under most  c o n d i t i o n s the gametophytes c o n t i n u e d growing v e g e t a t i v e l y and appeared h e a l t h y a f t e r more than one year i n c u l t u r e . In most i n s t a n c e s t h e male and f e m a l e gametophytes o f L. s i n c l a i r i i were e a s i l y d i s t i n g u i s h e d .  As i s t y p i c a l o f k e l p  gametophytes, t h e males have s m a l l e r , more numerous c e l l s and many b r a n c h e s , whereas t h e f e m a l e s c o n s i s t o f a few l a r g e r c e l l s , and few b r a n c h e s . a n t h e r i d i a were n o t e d . situation prevails.  I n s e v e r a l i n s t a n c e s , o o g o n i a and  I n L. l o n g i p e s , however, a d i f f e r e n t  The gametophytes a r e a p p a r e n t l y a l l o f  one m o r p h o l o g i c a l type and a r e t y p i c a l o f n e i t h e r male nor f e m a l e k e l p gametophytes.  The c e l l s a r e o f i n t e r m e d i a t e s i z e  and t h e b r a n c h e s , a l t h o u g h numerous, a r e n o t as numerous as i n male gametophytes o f L. s i n c l a i r i i . o o g o n i a were e v e r n o t e d . to  No a n t h e r i d i a o r  I n L. l o n g i p e s i t was thus i m p o s s i b l e  d i s t i n g u i s h males from f e m a l e s . d.  Discussion I t i s p o s s i b l e t h a t t h e gametophytes o f L. l o n g i p e s  are  bisexual  b u t u n t i l a n t h e r i d i a and o o g o n i a a r e f o u n d ,  w i l l remain i n question.  this  W i t h t h e e x c e p t i o n o f Chorda  tomentosa (Sundene, 1963), t h e gametophytes o f a l l k e l p s reported are unisexual. The c r o s s i n g e x p e r i m e n t s were n o t c a r r i e d o u t because two e s s e n t i a l c o n d i t i o n s were l a c k i n g .  The gametophytes o f  73  11- l o n g i p e s c o u l d n o t be d i s t i n g u i s h e d from each o t h e r and thus males and f e m a l e s c o u l d n o t be i s o l a t e d .  Secondly,  no  c o n d i t i o n s were found under w h i c h e i t h e r s p e c i e s would produce sporophytes  r e g u l a r l y , w h i c h would have s e r v e d as a c o n t r o l .  In p r e v i o u s e x p e r i m e n t s t h e author has c u l t u r e d gametophytes o f L a m i n a r i a s a c c h a r i n a sessile  (L.) Lamour., Hedophyllum  (C. Ag.) S e t c h , and N e r e o c y s t i s l l u e t k e a n a (Mert.) P.  and R. under t h e same c o n d i t i o n s o f temperature, media employed i n t h e p r e s e n t s t u d y . sporophytes  were produced.  l i g h t , and  I n many i n s t a n c e s abundant  R o b i n s o n (1967) c u l t u r e d s e v e r a l  s p e c i e s o f A l a r i a under t h e same c o n d i t i o n s i n t h e same c u l t u r e rooms a t t h e same time t h a t L. s i n c l a i r i i being cultured.  One s p e c i e s  and L. l o n g i p e s were  ( A l a r i a marginata  p. and R.) was  o b t a i n e d on t h e same Oregon beaches as L. s i n c l a i r i i . o f R o b i n s o n s c u l t u r e s sporophytes 1  were produced  I n most  abundantly.  The p r e s e n t s t u d y i n d i c a t e s t h a t , r e l a t i v e t o o t h e r members of the L a m i n a r i a l e s , the s e x u a l p r o d u c t i o n o f by gametophytes o f L. s i n c l a i r i i 3.  sporophytes  and L. l o n g i p e s i s v e r y  limited.  Haptera a.  Introduction F i e l d and l a b o r a t o r y o b s e r v a t i o n s show t h a t t h e r e  i s v e g e t a t i v e p r o d u c t i o n o f new b l a d e s and s t i p e s from t h e h a p t e r a a t t h e edges o f t h e h o l d f a s t s o f L. s i n c l a i r i i .  This  phenomenon was s t u d i e d f u r t h e r u s i n g l a b o r a t o r y c u l t u r e s o f i s o l a t e d pieces o f haptera  (Markham, 1968).  74  b.  M a t e r i a l s and Methods Sporophytes o f L. s i n c l a i r i i growing i n t h e l a r g e  c u l t u r e t a n k s p r o v i d e d a s o u r c e o f h a p t e r a f o r use i n t h e experiments.  A f t e r two t o t h r e e months i n t h e t a n k s , many  h a p t e r a had grown o u t from t h e bottoms and edges o f t h e h o l d f a s t s , e s p e c i a l l y a t 8° and 10°C.  These h a p t e r a were removed from  the p a r e n t p l a n t s , washed i n f i l t e r e d seawater, and c u t t o a u n i f o r m l e n g t h o f 20 mm.  I n the experiments t e s t i n g the  e f f e c t o f i n i t i a l s i z e , p i e c e s 2.5, 5, 10, and 15 mm l o n g were used.  F o r a l l s i z e s t h e h a p t e r a e i t h e r had one growing t i p  o r were c u t o f f a t b o t h ends; i n t h e l a t t e r i n s t a n c e , t h e most d i s t a l end was a t l e a s t 20 mm back o f t h e o r i g i n a l apex o f the h a p t e r o n .  The p i e c e s w i t h one c u t end and one growing t i p  were d e s i g n a t e d "E h a p t e r a " , whereas t h o s e w i t h b o t h ends c u t o f f were d e s i g n a t e d "1J h a p t e r a " . P r e l i m i n a r y e x p e r i m e n t s were r u n t o t e s t t h e e f f e c t s o f v a r i o u s media and c u l t u r e d i s h s i z e s .  A l l o f these  e x p e r i m e n t s were r u n a t 8° and 10°C and 150 f t - c . were employed:  ES+, ES, SWF, and SW.  Four media  Two t y p e s o f c u l t u r e  d i s h e s were t e s t e d : 250 m l g l a s s c u l t u r e d i s h e s c o n t a i n i n g 200 m l o f media and 20 mm p l a s t i c P e t r i d i s h e s c o n t a i n i n g 15 ml o f media.  The media were changed a t 2-week i n t e r v a l s .  These e x p e r i m e n t s showed t h a t t h e maximum p e r c e n t a g e o f h a p t e r a produced outgrowths i n 250 m l d i s h e s c o n t a i n i n g 200 m l o f ES+. A f t e r t h e p r e l i m i n a r y r e s u l t s were o b t a i n e d , e x p e r i m e n t s were r u n t o t e s t t h e e f f e c t s o f s h a k i n g and sand s c o u r i n g .  In  75  t h e s e experiments  20 mm p i e c e s o f h a p t e r a were s e c u r e d t o s m a l l  r o c k s w i t h r u b b e r bands around t h e m i d d l e p o r t i o n s o f t h e haptera. dishes.  These r o c k s were then p l a c e d i n 250 ml c u l t u r e G e n e r a l l y t h e r o c k s were l a r g e enough so t h a t o n l y  one c o u l d f i t i n each c u l t u r e d i s h . experiments,  For the shaking  t h e d i s h e s were then f i l l e d w i t h j u s t enough ES+  to cover t h e r o c k and i t s a t t a c h e d h a p t e r a . p l a c e d i n the Psycrotherm  These were then  i n c u b a t o r s h a k e r s and t h e speed o f  s h a k i n g a d j u s t e d so t h a t t h e medium r e a c h e d t h e r i m o f t h e c u l t u r e dishes but d i d not s p i l l over. experiments  F o r the s c o u r i n g  s t e r i l i z e d beach sand from Oregon was added t o  the c u l t u r e d i s h e s u n t i l i t c o v e r e d t h e r o c k s b u t n o t t h e h a p t e r a on t o p o f t h e r o c k s .  The d i s h e s were then f i l l e d w i t h  ES+ u n t i l t h e h a p t e r a were j u s t c o v e r e d and t h e d i s h e s were p l a c e d on t h e s h a k e r s w i t h those i n t h e s h a k i n g  experiments.  The water m o t i o n caused t h e sand t o wash back and f o r t h over the h a p t e r a .  Two p i e c e s o f h a p t e r a o f L. l o n g i p e s were a l s o  c u l t u r e d on t h e s h a k e r , each on a r o c k w i t h o u t sand. i n c u b a t o r s h a k e r s t h e temperature  In the  was 8°C, and t h e l i g h t  i n t e n s i t y was 150 f t - c , w i t h a p h o t o p e r i o d o f 16 hours 8 hours d a r k . Psycrotherm  C o n t r o l experiments  light/  were r u n i n a second  incubator without shaking.  The medium was changed  e v e r y two weeks. A 3-cm p i e c e o f s t i p e was c u t o u t o f each o f two p l a n t s o f L. s i n c l a i r i i ,  h a l f w a y between t h e h o l d f a s t and b l a d e  ( a p p r o x i m a t e l y 10 cm o f s t i p e was c u t o f f each e n d ) .  These  p i e c e s were then f a s t e n e d t o r o c k s and p l a c e d i n c u l t u r e  76  d i s h e s i n the same manner as t h e h a p t e r a . sand and ES+,  the o t h e r o n l y ES+.  These were c u l t u r e d w i t h o u t  s h a k i n g i n the c o n t r o l p s y c r o t h e r m . for  One d i s h c o n t a i n e d  A l l h a p t e r a were c u l t u r e d  14 weeks and s t i p e s f o r 20 weeks. These e x p e r i m e n t s were p r e l i m i n a r y i n t h a t they were s i m p l y  d e s i g n e d t o d e t e r m i n e whether  i t was p o s s i b l e f o r h a p t e r a to  produce b l a d e s and s t i p e s a t a l l under s u c h c o n d i t i o n s .  For  t h i s r e a s o n r e s u l t s were r e c o r d e d o n l y a t the end o f the e x p e r i m e n t s and then o n l y as p r e s e n c e o r abs_ence o f b l a d e s . The f i n a l s e r i e s o f e x p e r i m e n t s t e s t e d the e f f e c t s o f t e m p e r a t u r e , l i g h t i n t e n s i t y , and i n i t i a l s i z e o f h a p t e r a . A l l t h e s e c u l t u r e e x p e r i m e n t s were c a r r i e d o u t i n 250 ml g l a s s c u l t u r e d i s h e s c o n t a i n i n g 200 ml o f ES+ medium.  The medium  was changed e v e r y two weeks e x c e p t i n the experiments  testing  very small s i z e s .  I n t h e l a t t e r i n s t a n c e i t was  e v e r y f o u r weeks.  E x p e r i m e n t s on the e f f e c t o f temperature  were r u n a t 5°, 8°, 10°, 15°, and 20°C. were r u n o n l y a t 8° and/or 10°C. (21.5 l u x ) , 150 f t - c (1614 l u x ) ,  A l l o t h e r experiments  Light i n t e n s i t i e s of 2 f t - c 250 f t - c (2690 l u x ) ,  500 f t - c (5380 l u x ) were employed i n l i g h t experiments. only.  and  intensity  A l l o t h e r e x p e r i m e n t s were r u n under 150 f t - c  The p h o t o p e r i o d f o r a l l e x p e r i m e n t s was 16 hours  l i g h t / 8 hours d a r k . weeks.  changed  A l l t h e s e e x p e r i m e n t s were r u n f o r t e n  The r e s u l t i n g d a t a were s o r t e d w i t h an IBM 113 0  computer.  I n t h e f i n a l a n a l y s i s , a l l 20 mm  h a p t e r a were  a n a l y z e d f o r t h e i r r e s p o n s e s t o l i g h t and temperature ( F i g . 17a) whereas the e f f e c t s o f i n i t i a l s i z e were a n a l y z e d s e p a r a t e l y .  77  F i g u r e 17  R e s u l t s o f growth e x p e r i m e n t s  on h a p t e r a o f  L. s i n c l a i r i i  a.  percentage  o f E and N h a p t e r a p r o d u c i n g b l a d e s ,  h a p t e r a , and growth i n l e n g t h a t v a r i o u s temperatures b.  and v a r i o u s l i g h t  intensities.  2.5 mm p i e c e s o f N. h a p t e r a , c u l t u r e d a t 10°c under 150 f t - c , showing l a t e r a l b l a d e s .  c.  20 mm p i e c e s o f E and N h a p t e r a , c u l t u r e d a t 10°C, under 150 f t - c , showing l a t e r a l and t e r m i n a l b l a d e s and one l a t e r a l  d.  hapteron.  20 mm p i e c e s o f E and N h a p t e r a , c u l t u r e d a t 10°C, under 2 f t - c , showing l a t e r a l and t e r m i n a l b l a d e s and l a t e r a l  e.  haptera.  Nonmedian s e c t i o n o f t h e s t i p e o f a b l a d e a r i s i n g a l a t e r a l outgrowth  as  from a h a p t e r o n , x 70  (Diameter o f h a p t e r o n i s 1 mm). f.  An e n l a r g e d p o r t i o n o f e, x 210. A l l r e s u l t s a t 10 weeks,  h, o r i g i n a l  p i e c e o f hapteron;  from o r i g i n a l  b, b l a d e ; co, c o r t e x , l h , l a t e r a l hapteron  h a p t e r o n ; me, meristoderm;  s t , stipe.  arising  78  c.  Results The r e s u l t s o f t h e c u l t u r e experiments were v a r i e d .  The h a p t e r a e i t h e r showed no change; grew i n l e n g t h ; l a t e r a l haptera, i . e . ,  they branched  produced  ( F i g . 17c,d); o r t h e y  produced l a t e r a l o r t e r m i n a l b l a d e s ( F i g . 1 7 b , f , c , ) .  Normally  a b l a d e o u t g r o w t h appeared f i r s t as a s m a l l b l a d e , and two to t h r e e weeks l a t e r a s m a l l s t i p e d e v e l o p e d between the b l a d e and t h e p a r e n t h a p t e r o n . S e c t i o n s i n d i c a t e t h a t o n l y the outer c o r t e x i s i n v o l v e d i n p r o d u c t i o n o f l a t e r a l o u t g r o w t h s from h a p t e r a ( F i g . 1 7 e , f ) . In t h e s h a k i n g e x p e r i m e n t s b l a d e s were produced i n b o t h the  s h a k i n g and c o n t r o l c u l t u r e s and t h e s h a k i n g appeared t o  have l i t t l e e f f e c t .  The h a p t e r a o f L. l o n g i p e s produced b l a d e s  i n t h e same manner as t h e h a p t e r a o f L. s i n c l a i r i i . The sand s c o u r i n g e x p e r i m e n t s showed t h a t b o t h branches and b l a d e s can be produced under c o n d i t i o n s o f c o n s t a n t s c o u r i n g , a l t h o u g h t h e number and s i z e o f outgrowths a r e l e s s than i n the c o n t r o l experiments.  I n two i n s t a n c e s i n t h e  s c o u r i n g c o n d i t i o n s , a h a p t e r o n produced one b r a n c h w h i c h b e n t downward and a t t a c h e d f i r m l y t o a r o c k . The two p i e c e s o f s t i p e each produced outgrowths b o t h ends.  from  A l t h o u g h somewhat deformed, t h e outgrowths were  r e c o g n i z a b l e as b l a d e s .  The s t i p e s had been h e l d h o r i z o n t a l l y  i n t h e d i s h e s , and t h e b l a d e s a l l b e n t upward.  There i s a  s t r o n g i n d i c a t i o n t h a t the s t i p e l a c k s any i n t e r n a l p o l a r i t y . However, f u r t h e r e x p e r i m e n t s on p o l a r i t y were n o t conducted.  79  In most o f t h e e x p e r i m e n t s , d i f f e r e n t r e s u l t s were o b t a i n e d f o r E h a p t e r a and N h a p t e r a ; f o r b l a d e outgrowths and h a p t e r a o u t g r o w t h s ; and f o r l a t e r a l b l a d e outgrowths and t e r m i n a l blade outgrowths. made.  A few g e n e r a l i z a t i o n s can be  The N h a p t e r a never grew i n l e n g t h under any c o n d i t i o n s .  When E h a p t e r a grew, t h e growth always o c c u r r e d a t t h e uncut end.  A g r e a t e r p e r c e n t a g e o f E h a p t e r a produced outgrowths  d i d N h a p t e r a under any g i v e n s e t o f c o n d i t i o n s .  than  The N h a p t e r a  never produced b o t h h a p t e r a and b l a d e outgrowths on t h e same h a p t e r o n , whereas E h a p t e r a o c c a s i o n a l l y d i d . For o r i g i n a l  h a p t e r a l e n g t h s o f l e s s than 15 mm,  lateral  b l a d e s were produced by N h a p t e r a , whereas t e r m i n a l b l a d e s were produced by E_ h a p t e r a . F o r t h e s t a n d a r d 20 mm h a p t e r a , N h a p t e r a g e n e r a l l y produced produced  l a t e r a l b l a d e s whereas E h a p t e r a  l a t e r a l and/or t e r m i n a l b l a d e s .  g e n e r a l l y appeared  earlier  Terminal blades  than l a t e r a l b l a d e s .  I n the f i n a l  a n a l y s e s o f t h e r e s u l t s , outgrowths were c o n s i d e r e d m e r e l y as b l a d e s o r h a p t e r a , w i t h no r e g a r d t o whether they were l a t e r a l or  terminal. i.  Temperature - A t 20°C t h e r e was a c o n s i d e r a b l e  growth o f v a r i o u s c o n t a m i n a n t s .  No  p o s i t i v e growth  response  was shown by any o f t h e h a p t e r a and s e v e r a l began t o disintegrate.  I t i s c o n c l u d e d t h a t 2 0°C i s a l e t h a l  temperature.  W i t h i n t h e l i m i t s o f 5° t o 15°C, v a r i o u s r e s u l t s were o b t a i n e d . The g r e a t e s t p e r c e n t a g e o f E h a p t e r a grew i n l e n g t h and produced  l a t e r a l h a p t e r a o r branches a t 10°C ( F i g . 17a). No  growth o c c u r r e d a t 15°C.  The g r e a t e s t p e r c e n t a g e o f N h a p t e r a  80  produced  l a t e r a l h a p t e r a a t 8 C.  The p e r c e n t a g e o f E h a p t e r a  p r o d u c i n g b l a d e s r o s e w i t h i n c r e a s i n g temperature, w i t h t h e maximum o f 100 p e r c e n t a t 15°C.  The maximum p e r c e n t a g e o f  N h a p t e r a produced b l a d e s a t 5°C.  The average number o f  b l a d e s p e r E h a p t e r o n w h i c h produced b l a d e s was h i g h e s t a t 8°c, or  i n t e r m e d i a t e t e m p e r a t u r e , whereas f o r N h a p t e r a i t was  h i g h e s t a t t h e extremes o f t e m p e r a t u r e , 5° and 15°C ( F i g . 17a). ii.  L i g h t I n t e n s i t y - The g r e a t e s t p e r c e n t a g e o f  E h a p t e r a grew i n l e n g t h and produced ft-c  l a t e r a l h a p t e r a a t 150  ( F i g . 1 7 a ) . No growth o c c u r r e d a t 500 f t - c .  p e r c e n t a g e o f N h a p t e r a produced  The g r e a t e s t  l a t e r a l haptera a t 2 f t - c .  The p e r c e n t a g e o f E_ h a p t e r a p r o d u c i n g b l a d e s r o s e w i t h i n c r e a s i n g l i g h t i n t e n s i t y t o a maximum o f 100 p e r c e n t a t 500 f t - c .  The  maximum p e r c e n t a g e o f N h a p t e r a produced b l a d e s a t 250 and 500 f t - c .  The average number o f b l a d e s p e r E_ h a p t e r o n  produced b l a d e s r o s e s t e a d i l y w i t h i n c r e a s i n g l i g h t to a maximum a t 500 f t - c . ( F i g .  which  intensity  1 7 a ) . F o r N h a p t e r a t h e number  o f b l a d e s was h i g h e s t a t 2 f t - c , b u t t h e n e x t h i g h e s t o c c u r r e d a t t h e o t h e r extreme,  500 f t - c .  H a p t e r a tended t o a r i s e a t lower l i g h t i n t e n s i t i e s and i n i n s t a n c e s where b o t h h a p t e r a and b l a d e s appeared a t l o w l i g h t intensities, iii.  t h e h a p t e r a g e n e r a l l y appeared  first.  I n i t i a l S i z e - The s i z e experiments showed t h a t  even p i e c e s o f h a p t e r a o n l y 2.5 mm l o n g can produce  outgrowths  of normal appearance ( F i g . 1 7 b ) . However, two e f f e c t s o f t h e s m a l l s i z e were n o t e d ,  p i e c e s o f h a p t e r a s h o r t e r than 15 mm  81  produced b l a d e s , b u t never h a p t e r a .  S e c o n d l y , the s i z e o f the  o u t g r o w t h appears t o be c o r r e l a t e d w i t h the s i z e o f the o r i g i n a l h a p t e r o n ; l a r g e r p i e c e s produce l a r g e r o u t g r o w t h s . d.  Discussion The d i f f e r e n c e s between the E h a p t e r a w i t h one  growing t i p and the N h a p t e r a w i t h no i n t a c t t i p s can be i n t e r p r e t e d i n a number o f ways.  The e f f e c t may be  chiefly  due t o i n j u r y caused i n removing the ends, as i n d i c a t e d by the  s c a r c i t y o f t e r m i n a l outgrowths from the N h a p t e r a .  It  may be due t o the r e l a t i v e age o f the p i e c e s o f h a p t e r a , s i n c e N h a p t e r a a r e f a r t h e r from the growing t i p and thus o l d e r .  It  c o u l d be e x p e c t e d from t h i s t h a t the N h a p t e r a would have l e s s p o t e n t i a l f o r r e d i f f e r e n t i a t i o n and would thus produce outgrowths.  fewer  Such was o b s e r v e d t o be the case i n t h i s s t u d y .  However, f i e l d s t u d i e s a l s o showed t h a t an e n t i r e p l a n t can be c u t i n h a l f and new h a p t e r a w i l l grow out from the c e n t e r , or  oldest, haptera.  Y e t a n o t h e r p o s s i b i l i t y , w h i c h has n o t  been i n v e s t i g a t e d , i s t h a t some s u b s t a n c e such as an a u x i n , w h i c h s t i m u l a t e s h a p t e r a growth and b l a d e p r o d u c t i o n i s produced o r s t o r e d i n the apex. the  S e v e r a l a u t h o r s have r e p o r t e d  p r e s e n c e o f a u x i n s i n various'members  o f the L a m i n a r i a l e s  (Van Overbeek, 1940a, b; W i l l i a m s , 1949; Mowat, 1965) and of the  one  them (van Overbeek, 1940b) r e p o r t e d a u x i n s p e c i f i c a l l y i n haptera of C o s t a r i a c o s t a t a . S e v e r a l a u t h o r s , most r e c e n t l y Dawson (1966), have n o t e d  t h a t L. s i n c l a i r i i r e p r o d u c e s v e g e t a t i v e l y from a r h i z o m e - l i k e holdfast.  The p r e s e n t s t u d y has shown t h a t the h a p t e r a need  82  n o t be a t t a c h e d t o t h e p a r e n t p l a n t f o r such r e p r o d u c t i o n t o occur.  Even v e r y s m a l l p i e c e s o f h a p t e r a can produce b l a d e s  and s t i p e s and thus u l t i m a t e l y a whole new p l a n t .  Assuming  t h a t t h i s c o u l d take p l a c e i n nature a f t e r a p o r t i o n of a h a p t e r o n i s a c c i d e n t a l l y c u t l o o s e from the p a r e n t p l a n t , the E type o f h a p t e r o n , t h a t i s , one w h i c h has o n l y been c u t once, seems much more l i k e l y . The c o n d i t i o n s under w h i c h the maximum number o f b l a d e s a r e produced per h a p t e r o n a r e n o t the same as t h o s e under which the maximum number o f h a p t e r a produce b l a d e s .  The  latter  would seem t o be the more i m p o r t a n t c o n s i d e r a t i o n , i f t h i s p r o d u c t i o n i s r e g a r d e d 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 . Each p i e c e o f h a p t e r o n , no m a t t e r how many a t t a c h e d s t i p e s b l a d e s i t has, i s s t i l l o n l y p a r t o f one  and  plant.  The e x p e r i m e n t s i n d i c a t e t h a t 20°C i s a l e t h a l temperature, b u t the h i g h e s t l i g h t i n t e n s i t y t e s t e d , 500 f t - c , i s n o t harmful.  T h i s i s t o be e x p e c t e d from o b s e r v e d n a t u r a l c o n d i t i o n s .  A water temperature o f 20°C i s v e r y u n l i k e l y on the Oregon C o a s t , whereas l i g h t i n t e n s i t i e s g r e a t e r than 500 f t - c do occur.  S e p a r a t e temperature and l i g h t i n t e n s i t y e x p e r i m e n t s  show a maximum b l a d e p r o d u c t i o n a t 15°C  and 500 f t - c .  However,  a l l temperature e x p e r i m e n t s were r u n a t 150 f t - c and a l l l i g h t i n t e n s i t y e x p e r i m e n t s a t 10°C,  so t h a t i t may  be  i n c o r r e c t t o assume t h a t a c o m b i n a t i o n o f h i g h temperature and h i g h l i g h t i n t e n s i t y would r e s u l t i n h i g h b l a d e p r o d u c t i o n .  83  N e v e r t h e l e s s , i f one makes t h i s a s s u m p t i o n , a maximum b l a d e p r o d u c t i o n s h o u l d o c c u r i n e a r l y summer when t h e water temperature i s r i s i n g t o 15°C and t h e l i g h t i n t e n s i t y i s s t i l l h i g h because t h e p l a n t s have n o t y e t been b u r i e d under sand.  F i e l d s t u d i e s i n d i c a t e t h i s i s i n d e e d the c a s e .  The s c o u r i n g e x p e r i m e n t s i n d i c a t e t h a t such p r o d u c t i o n can o c c u r even i f t h e h a p t e r a a r e p a r t l y b u r i e d i n sand, p r o v i d e d there i s s u f f i c i e n t  light.  84  VII.  TAXONOMY OF L. SINCLAIRII AND  A.  L. LONGIPES  Introduction  L. s i n c l a i r i i and L. l o n g i p e s a r e v e r y d i s t i n c t from a l l o t h e r s p e c i e s o f the genus. s p e c i e s may  The most s i m i l a r o f the o t h e r  be the M e d i t e r r a n e a n  because o f i t s m u l t i p l e s t i p e s .  p l a n t , L. r o d r i g u e z i i  Bornet,  However, i l l u s t r a t i o n s o f  L. r o d r i g u e z i i i n d i c a t e t h a t a l t h o u g h i t has m u l t i p l e s t i p e s , t h e y a r e v e r y few and w i d e l y s e p a r a t e d , u n l i k e the s i t u a t i o n i n L. s i n c l a i r i i and L. l o n g i p e s .  Furthermore,  unlikea l l  other species of Laminaria, i t i s r e s t r i c t e d to very great depths (100-150 m) (Bornet, 1888). L_. s i n c l a i r i i and L. l o n g i p e s a r e much more s i m i l a r to each o t h e r than t o any o t h e r s p e c i e s i n the genus. t h e y a r e d i s t i n c t from each o t h e r .  Nevertheless,  Previously published reports  have emphasized t h e presence o r absence o f m u c i l a g e the s t i p e f o r d i s t i n g u i s h i n g the two s p e c i e s .  The  ducts i n present  s t u d y has r e v e a l e d a t l e a s t f i v e d i f f e r e n c e s between the s p e c i e s w h i c h have n o t been p r e v i o u s l y r e p o r t e d .  These a r e  b l a d e w i d t h , s e a s o n a l l o s s o f b l a d e s , morphology o f gametophytes, and temperature  tolerances.  On the b a s i s o f  t h e s e and o t h e r d i f f e r e n c e s r e p o r t e d below, i t i s c l e a r t h a t they a r e two s e p a r a t e t a x a .  85  B.  L a m i n a r i a s i n c l a i r i i (Harvey ex Hooker f . and Harvey) F a r l o w , Anderson and E a t o n , 1877-1889. Harvey, 1852, p. 87 (as L e s s o n i a s i n c l a i r i i ) F a r l o w , Anderson and E a t o n , 1878, f a s c . 3, p. 118 Areschoug, 1883, p. 6 (as H a f q y q i a s i n c l a i r i i ) Anderson, 1891, p. 220 Howe, 1893, p. 67 De T o n i , 1895, p. 343 C o l l i n s , Holden and S e t c h e l l , 1895-1919, f a s c . 7 S e t c h e l l , 1896, pp. 44-46; 1905, pp. 139-169; 1912 pp. 131, 134, 137, 140, 141, 148, 150 Myers, 1925, pp. 114-116 S e t c h e l l and Gardner, 1925, p. 598 Okamura, 1932, p. 73 S m i t h , 1944, p. 135, p i . 31 Doty, 1947, p. 40 Sanborn and Doty, 1947, pp. 9, 13, 21, 30 Shchapova, 1948, pp. 99, 100, l l 7 , 120 S c a g e l , 1957, p. 98 S i l v a , 1957, pp. 43,44 Dawson, 1958a, p. 66; 1958b, pp. 186, 188, 201, 204; 1958c, pp. 235, 238, 242, 260; 1959, pp. 144, 161, 162; 1961, p. 396 H o l l e n b e r g and A b b o t t , 1966, p. 25 D r u e h l , 1968, p. 541 Markham, 1968, p. 125-131  1.  Description Sporophytes  up t o 3 m l o n g , p e r e n n i a l from the h o l d f a s t  and s t i p e , r e g e n e r a t i n g new b l a d e s a f t e r complete ones.  loss of o l d  H o l d f a s t an e x t e n s i v e r h i z o m e - l i k e system o f  h a p t e r a , sometimes c o v e r i n g an a r e a o f 5 m^  or more.  branched The  h a p t e r a g i v i n g r i s e t o many s t i p e s , more than 100 i n some instances.  S t i p e s f l e x i b l e , c y l i n d r i c a l , l-3mm i n d i a m e t e r ,  up t o 60 cm l o n g o r g r e a t e r , m u c i l a g e d u c t s p r e s e n t . narrowly l i n e a r ,  u s u a l l y l e s s than 3.0  Blades  cm wide, o f v a r i a b l e  l e n g t h up t o 2.5 m, e n t i r e , w i t h o u t b u l l a e , m u c i l a g e  ducts  present.  B l a d e s g e n e r a l l y l o s t i n December, r e g e n e r a t e d i n  January,  p l a n t s t e n d i n g t o be l a r g e r near the s o u t h e r n  limits  86  of  distribution.  S o r i i n patches, oblong to i r r e g u l a r i n  o u t l i n e , one t o many p e r b l a d e ,  p l a n t s produce s o r i i n  October, November, J a n u a r y , F e b r u a r y , and March.  Gametophytes  f i l a m e n t o u s , c a p a b l e o f p r o l o n g e d v e g e t a t i v e growth, u n i s e x u a l , w i t h males and females d i s t i n c t l y d i f f e r e n t , r a r e l y p r o d u c i n g sporophytes. 2.  Distribution Hope I s l a n d  (50°55'N, 127°58'W) B r i t i s h Columbia t o  V e n t u r a County, (34°19'N, 119°23.3'W) C a l i f o r n i a . 3.  Habitat Growing on r o c k s i n t h e lower i n t e r t i d a l r e g i o n ,  u s u a l l y p a r t l y b u r i e d under sand i n summer, u s u a l l y i n f u l l y exposed a r e a s , b u t o c c a s i o n a l l y i n m o d e r a t e l y s h e l t e r e d t o m o d e r a t e l y exposed 4.  areas.  Comments The mean l e n g t h o f t h e l o n g e s t s t i p e s from each o f  135 p l a n t s c o l l e c t e d i n Oregon i s 26.3 cm.  The l o n g e s t  measured s t i p e i n t h i s group was 52 cm and t h e s h o r t e s t was 11 cm.  P l a n t s lower i n t h e i n t e r t i d a l r e g i o n a r e g e n e r a l l y  l o n g e r b u t were r a r e l y c o l l e c t e d because o f d i f f i c u l t conditions.  Consequently,  surf  t h e mean l e n g t h o f a l l s t i p e s on  the Oregon beaches i s p r o b a b l y g r e a t e r than 27 cm.  plants of  L_. s i n c l a i r i i i n C a l i f o r n i a a r e known t o be g e n e r a l l y l a r g e r than those i n Oregon b u t no comprehensive  data are a v a i l a b l e .  The mean w i d t h o f t h e b l a d e s on t h e l a r g e s t s t i p e s o f 54  87  p r e s s e d specimens from Oregon i s 1.75  cm.  From measurements  o f b o t h f r e s h and p r e s s e d specimens from I n d i a n Beach, the shrinkage a f t e r p r e s s i n g i s c a l c u l a t e d this correction  Applying  f a c t o r , the mean b l a d e w i d t h . f o r a l l specimens  examined i s a p p r o x i m a t e l y  C.  t o be 20%.  2.2  cm.  L a m i n a r i a l o n g i p e s Bory,  1826  Agardh, C , 1820, p 133 Bory, 1826, v o l . 9, p 189 P o s t e l s and R u p r e c h t , 1840, p. 10 (as L. s a c c h a r i n a f . angustifolia) K i i t z i n g , 1849, p. 574 R u p r e c h t , 1851, p. 232, 350 (as L e s s o n i a repens) Le J o l i s , 1855, p. 307-308,311 (as L. r u p r e c h t i a n a ) Agardh, J . , 1867, p. 26 (as Arthrothamnus ? l o n g i p e s ) A r e s c h o u g , 1883, p. 15 K j e l l m a n , 1889, pp. 7, 9, 17, 43 De T o n i , 1895, p. 370 (as Ar t h r o thamnus ? l o n g i p e s ) S e t c h e l l , 1899, pp. 591, 592, p i . 95 S e t c h e l l and Gardner, 1903, p. 260 Yendo, 1909, p. 215; 1910, p. 295 S e t c h e l l , 1912, pp. 136, 148, 150 Okamura, 1916, p. 172 S e t c h e l l and Gardner, 1925, p. 597 Okamura, 1928, p. 53, p i s . 13-15 A r w i d s s o n , 1932, p. 153 Miyabe and N a g a i , 1932, pp. 196, 197; 1933, pp. 86, 87 Okamura, 1932, p. 72; 1933, pp. 88, 95 Z i n o v a , E., 1933, pp. 24, 25, f i g s . 8-10 Okada, 1934, p. 46, p i . 43 Yamada, 1935, pp. 1, 6, 7, 18, p i . 6 (as L. l o n g i p e s var. l a t i f o l i a ) Okamura, 1936, p. 251, f i g . 139 N a g a i , 1940, pp. 67-70; 1941, p. 262 (as L. l o n g i p e s f . a n g u s t i f o l i a , f. l i n e a r i s , & f. l a t i f o l i a ) Shchapova, 1948, pp. 93, 98, 100, 120, 127, 130, 131 T o k i d a , 1954, pp. 30, 114, 115 ( i n c l . L. l o n g i p e s f . t y p i Z i n o v a , E., 1954, p. 378 Z i n o v a , A., 1959, p. 153 Dawson, 1961, p. 396 V o z z h i n s k a y a , 1964, p. 425 D r u e h l , 1968, p. 541 0  0  88  1.  Description Sporophytes  up t o 1 m l o n g , p e r e n n i a l from the h o l d f a s t  and s t i p e , r e g e n e r a t i n g new b l a d e s w h i l e remnants o f the o l d ones remain. branched  H o l d f a s t an e x t e n s i v e r h i z o m e - l i k e system  of  h a p t e r a , sometimes c o v e r i n g an a r e a up to 5 m2.  The  h a p t e r a g i v i n g r i s e t o many s t i p e s , more than 100 i n some instances.  S t i p e s f l e x i b l e , c y l i n d r i c a l , 1-3 mm  i n diameter,  up t o 40 cm l o n g , u s u a l l y n o t e x c e e d i n g 20 cm i n l e n g t h , mucilage ducts absent. than 5.0  Blades narrowly l i n e a r , u s u a l l y l e s s  cm wide, o f v a r i a b l e l e n g t h up t o 80 cm,  without b u l l a e , mucilage ducts present.  entire,  Blades g e n e r a l l y  s h o r t e s t i n December o r J a n u a r y , b u t n o t l o s t c o m p l e t e l y . S o r i i n p a t c h e s , round t o i r r e g u l a r i n o u t l i n e , one per b l a d e ,  t o many  p l a n t s produce s o r i i n December; no i n f o r m a t i o n i s  a v a i l a b l e f o r o t h e r w i n t e r months.  Gametophytes f i l a m e n t o u s ,  c a p a b l e o f p r o l o n g e d v e g e t a t i v e growth, a p p a r e n t l y a l l o f type, very r a r e l y producing 2.  one  sporophytes.  Distribution Urup I s l a n d  (46°00'N, 150°00'E) K u r i l e I s l a n d s around o o p a c i f i c Rim t o C o r o n a t i o n I s l a n d (55 49.6'N, 134 17'W) A l a s k a . 3.  Habitat Growing on r o c k s i n the lower i n t e r t i d a l and upper  s u b t i d a l r e g i o n s , u s u a l l y i n moderately  exposed o r  moderately  s h e l t e r e d a r e a s , b u t o c c a s i o n a l l y i n f u l l y exposed a r e a s .  89  4.  Comments , The mean l e n g t h o f t h e s t i p e s measured on herbarium  specimens from 33 l o c a t i o n s i s 8.4 cm.  The average s t i p e  l e n g t h i s g r e a t e r on p l a n t s from more exposed s i t e s XXIII).  (Table  However, t h e l o n g e s t s t i p e r e c o r d e d , 3 5 cm, i s on a  p l a n t from an a r e a which i s o n l y "moderately  exposed".  The  mean w i d t h o f t h e b l a d e s o f p r e s s e d specimens from 33 l o c a t i o n s i s 1.97 cm.  From measurements on b o t h f r e s h and p r e s s e d  specimens from A a t s Bay, t h e s h r i n k a g e a f t e r p r e s s i n g i s c a l c u l a t e d t o be 37%. T h i s i s g r e a t e r than t h a t f o r L. sinclairii.  However, e x a m i n a t i o n o f o t h e r s p e c i e s from A l a s k a  i n d i c a t e s a g r e a t e r s h r i n k a g e i n them a l s o .  I t i s believed  t h a t t h e c o n c e n t r a t i o n o f f o r m a l i n used t o p r e s e r v e t h e p l a n t s was n o t t h e same f o r Oregon and A l a s k a .  Applying this  c o r r e c t i o n f a c t o r , t h e mean b l a d e w i d t h f o r a l l specimens examined i s a p p r o x i m a t e l y 3.1 cm.  A c c o r d i n g t o some a u t h o r s ,  (e.g., yamada, 1935) some p l a n t s i n t h e K u r i l e I s l a n d s have b l a d e s c o n s i d e r a b l y b r o a d e r than  D.  this.  Comparisons o f S p e c i e s  L. s i n c l a i r i i and_L_. l o n g i p e s a r e two d i s t i n c t s p e c i e s . The p r e s e n t s t u d y has shown t h a t t h e r e a r e a t l e a s t 10 d i f f e r e n c e s between t h e two s p e c i e s , as f o l l o w s :  90 L.  sinclarii  L.  longipes  1.  Mucilage ducts present i n stipe.  1,  M u c i l a g e d u c t s a b s e n t from stipe.  2.  S t i p e i n mature p l a n t u s u a l l y over 20 cm l o n g .  2,  S t i p e i n mature p l a n t u s u a l l y l e s s than 20 cm l o n g .  3.  B l a d e o f mature p l a n t 3, u s u a l l y l e s s t h a n 3 cm wide.  B l a d e o f mature p l a n t u s u a l l y more then 3 cm wide.  4.  E n t i r e b l a d e l o s t and then new one r e g e n e r a t e d .  4,  P r o x i m a l remnant o f b l a d e r e t a i n e d and new one produced w h i l e remnant s t i l l present.  5.  Male and female gametophytes i n c u l t u r e morphologically different.  5.  Gametophyes i n c u l t u r e morphologically i n d i s t i n g u i s h a b l e as t o s e x .  6.  Occurs from Southern California to Central B r i t i s h Columbia.  6.  Occurs from S o u t h e a s t A l a s k a throughout the A l e u t i a n Islands to the K u r i l e Islands.  7.  O p t i m a l growth a t mean temperature higher than 8°C.  7.  O p t i m a l growth a t mean temperature lower t h a n 8 c  8.  Grows i n lower i n t e r t i d a l region.  8.  Grows i n lower i n t e r t i d a l and upper s u b t i d a l r e g i o n s .  9.  U s u a l l y o c c u r s on f u l l y exposed s i t e s .  9.  U s u a l l y o c c u r s on m o d e r a t e l y exposed o r m o d e r a t e l y sheltered sites.  10.  Usually associated with sand and b u r i e d f o r p a r t of year.  10.  Rarely associated with  sand.  F i g u r e 18 i l l u s t r a t e s t h e d i f f e r e n c e s i n b l a d e w i d t h and stipe length. the p o i n t . f i g u r e 18.  Extreme examples were chosen t o i l l u s t r a t e  The d i f f e r e n c e i s u s u a l l y n o t as g r e a t as i n As n o t e d above, p o i n t s 3, 4, 5, 7, and 9 have n o t  been r e p o r t e d b e f o r e ; t h e o t h e r s have been mentioned,  usually  91  F i g u r e 18  a.  H a b i t o f L. s i n c l a i r i i and L.  L. s i n c l a i r i i from P e s c a d e r o P o i n t , California, x  b.  longipes  1/3.  L. l o n g i p e s from cape Spencer, A l a s k a , x  1/3.  92  without  emphasis, by v a r i o u s a u t h o r s .  Except f o r d i f f e r e n c e s  i n gametophytes and m u c i l a g e d u c t s , none o f these p o i n t s a l o n e would seem s u f f i c i e n t as a b a s i s f o r s e p a r a t i n g t h e two s p e c i e s .  However, a l l o f t h e s e d i f f e r e n c e s c o l l e c t i v e l y  c o n f i r m t h a t L. s i n c l a i r i i and L. l o n g i p e s s h o u l d be r e t a i n e d as two s e p a r a t e  species.  definitely  93  VIII.  GENERAL DISCUSSION AND CONCLUSIONS  The two s p e c i e s s t u d i e d , L. s i n c l a i r i i and L. l o n g i p e s , a r e v e r y d i s t i n c t from a l l o t h e r s p e c i e s o f t h e genus w i t h t h e p o s s i b l e e x c e p t i o n o f L. r o d i g u e z i i as n o t e d .  They a r e a l s o  d i f f e r e n t from most o t h e r members o f t h e o r d e r L a m i n a r i a l e s . There a r e t h r e e f e a t u r e s o f these two s p e c i e s w h i c h a r e v e r y distinctive. reproduction.  There i s an apparent  suppression of sexual  They have a g r e a t p o t e n t i a l f o r  d e d i f f e r e n t i a t i o n o f s u p p o s e d l y mature t i s s u e s and r e l a t e d t o t h i s , a more d i f f u s e m e r i s t e m a t i c a r e a w h i c h a l l o w s c o n s i d e r a b l e growth i n r e g i o n s o t h e r than t h e t r a n s i t i o n zone, e s p e c i a l l y i n the haptera. A few o t h e r k e l p s e x h i b i t c o n s i d e r a b l e growth i n r e g i o n s o t h e r than t h e t r a n s i t i o n zone.  E g r e g i a spp. develops a v e r y  complex t h a l l u s p r i m a r i l y by l a t e r a l outgrowths from t h e f l a t t e n e d s t i p e , w i t h v e r y l i t t l e growth o c c u r r i n g i n t h e t r a n s i t i o n zone.  I n Dictyoneurum c a l i f o r n i c u m Rupr. t h e  s t i p e becomes p r o s t r a t e and forms a r h i z o m e - l i k e s t r u c t u r e which attaches t o the s u b s t r a t e by l a t e r a l haptera. s p l i t s i n t h e b l a d e cause segmentation  Repeated  and t h e f o r m a t i o n o f  many b l a d e s , each w i t h a s t i p e t h a t becomes p r o s t r a t e so t h a t a clump i s formed ( S e t c h e l l and Gardner, 1925). Arthrothamnus b i f i d u s  In  (Gmel.) J . Ag. t h e b a s a l margins o f t h e  b l a d e s become m e r i s t e m a t i c and numerous secondary  blades  arise.  The p r o c e s s i s r e p e a t e d many t i m e s , f o r m i n g a r h i z o m e - l i k e s t r u c t u r e b e a r i n g many b l a d e s  (Yamada, 1938).  The most  94  s i m i l a r development t o t h a t o f L. s i n c l a i r i i i s found i n t h e Japanese s p e c i e s E c k l o n i a s t o l o n i f e r a Okam.  The h a p t e r a grow  i n t o s t o l o n i f e r o u s s t r u c t u r e s and produce new b l a d e s a t t h e tips.  The b l a d e s a r e deciduous  (Okamura, 1915).  Okamura  s t a t e s a l s o t h a t w i t h t h i s v e g e t a t i v e r e p r o d u c t i o n , the f o r m a t i o n o f z o o s p o r a n g i a l s o r i seems t o be s u p p r e s s e d .  However,  E c k l o n i a d i f f e r s from L a m i n a r i a i n h a v i n g l a t e r a l outgrowths from t h e b l a d e s . With the p o s s i b l e exception o f E c k l o n i a s t o l o n i f e r a , the s u p p r e s s i o n o f s e x u a l r e p r o d u c t i o n i n t h e two s p e c i e s s t u d i e d appears t o be unique i n t h e k e l p s . suppressed  Sorus p r o d u c t i o n i s n o t  i n these two s p e c i e s b u t t h e r e i s l i t t l e  t h a t t h e r e s u l t i n g gametophytes n o r m a l l y produce  evidence  sporophytes  i n any number. In v i e w o f t h e d i f f e r e n c e s between L. s i n c l a i r i i and L. l o n g i p e s and o t h e r s p e c i e s o f t h e genus, i t i s p o s s i b l e t h a t they s h o u l d be removed from t h e genus L a m i n a r i a . c h i e f reasons  One o f t h e  f o r removing them from t h e genus i s t h e  m e r i s t e m a t i c a c t i v i t y o f the .haptera w h i c h r e s u l t s i n m u l t i p l e stipes.  On t h i s b a s i s , however, t h e two s p e c i e s would have t o  be removed n o t o n l y from t h e genus L a m i n a r i a , b u t a l s o from the f a m i l y L a m i n a r i a c e a e ,  as t h e f a m i l i 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 b a s e d on t h e type o f growth i n t h e meristematic area.  Thus, b e f o r e such a taxonomic r e v i s i o n  c o u l d be made, a comprehensive r e v i e w o f t h e e n t i r e o r d e r would have t o be c a r r i e d o u t .  F o r t h i s r e a s o n , no changes o f taxonomy  are proposed a t the present  time.  95  D e s p i t e the a l m o s t i d e n t i c a l e x t e r n a l morphology  of  L. s i n c l a i r i i and L. l o n g i p e s , and the many common f e a t u r e s by w h i c h t h e y d i f f e r from o t h e r k e l p s , the two s p e c i e s a r e a l s o d i s t i n c t from each o t h e r , as has been demonstrated. of  these d i f f e r e n c e s may be the r e s u l t o f a d a p t a t i o n t o d i f f e r e n t  habitats. of the  Many  The most s t r i k i n g d i f f e r e n c e between the h a b i t a t s  the two s p e c i e s i s i n the temperature o f the seawater air.  and  The mean t e m p e r a t u r e s f o r the c o l d e s t w i n t e r months  and the a n n u a l means show no o v e r l a p p i n g v a l u e s a t a l l (Table I , F i g . 2, 3 ) .  I n summer t h e d i f f e r e n c e i s n o t as g r e a t , b u t  t h e r e i s o n l y a s l i g h t o v e r l a p f o r seawater temperature and none f o r a i r t e m p e r a t u r e .  The e v i d e n c e i n d i c a t e 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 o f v e r y l i t t l e i m p o r t a n c e t o these s p e c i e s . N e v e r t h e l e s s , i t may be s i g n i f i c a n t t h a t the most marked d i f f e r e n c e i n temperature between the a r e a s where the two s p e c i e s a r e f o u n d o c c u r s a t the time o f year when b o t h s p e c i e s bear r i p e  sori.  I t was o b s e r v e d t h a t p l a n t s o f L. s i n c l a i r i i c o l l e c t e d i n C a l i f o r n i a containedmuch more m u c i l a g e and became v e r y s l i m y soon a f t e r c o l l e c t i o n . n o t become s l i m y a t a l l .  The p l a n t s c o l l e c t e d i n Oregon d i d S e v e r a l a u t h o r s have s t a t e d t h a t  k e l p s growing i n the i n t e r t i d a l zone a r e p r o t e c t e d from r a p i d d e s i c c a t i o n by the p r e s e n c e o f m u c i l a g e .  The a i r temperature  and degree o f i n s o l a t i o n i s much g r e a t e r on C a l i f o r n i a than on Oregon beaches.  beaches  Hence the danger o f d e s i c c a t i o n i s  much g r e a t e r i n C a l i f o r n i a .  The g r e a t e r p r o d u c t i o n o f m u c i l a g e  i n the C a l i f o r n i a p l a n t s i s p r o b a b l y an a d a p t a t i o n i n r e s p o n s e  96  to air  this.  I n A l a s k a , where L. l o n g i p e s i s found, t h e average  temperature and t h e danger o f d e s i c c a t i o n a r e  c o n s i d e r a b l y l e s s than i n Oregon.  L. l o n g i p e s does n o t  produce m u c i l a g e t o any n o t i c e a b l e degree. lacks mucilage ducts i n the s t i p e .  Furthermore, i t  No d e f i n i t e  correlation  has been demonstrated between t h e p r e s e n c e o f m u c i l a g e d u c t s and t h e p r o d u c t i o n o f m u c i l a g e .  Nevertheless, c o n s i d e r i n g the  value o f mucilage i n p r o t e c t i o n a g a i n s t the e f f e c t s o f high t e m p e r a t u r e s and Burrows'  (1964) d e m o n s t r a t i o n t h a t m u c i l a g e  d u c t s t e n d t o be produced o n l y a t h i g h e r t e m p e r a t u r e s , i t i s q u i t e p o s s i b l e t h a t such a c o r r e l a t i o n may e x i s t .  Thus t h e  d i f f e r e n c e i n m u c i l a g e d u c t s i n t h e two s p e c i e s may have a r i s e n o r i g i n a l l y as a r e s p o n s e t o d i f f e r e n t t e m p e r a t u r e s , even though i t c a n a p p a r e n t l y no l o n g e r be a l t e r e d by changing the  temperature. Temperature may be t h e f a c t o r w h i c h l i m i t s t h e d i s t r i b u t i o n  of  L. s i n c l a i r i i s o u t h o f V e n t u r a County, C a l i f o r n i a .  At  S a n t a M o n i c a , j u s t s o u t h o f t h e s o u t h e r n l i m i t s o f L. s i n c l a i r i i , b o t h t h e summer mean t e m p e r a t u r e s and t h e y e a r l y mean t e m p e r a t u r e s a r e more t h a n 1C°  g r e a t e r than i n V e n t u r a  County.  A t S h o r t Sand Beach t h e p l a n t s o f L. s i n c l a i r i i were r e g u l a r l y immersed f o r s e v e r a l hours i n a l m o s t f r e s h water d u r i n g t h e l a t t e r p a r t o f t h e summer.  Many o f t h e r e d a l g a e  growing on t h e same r o c k s d i e d i n l a r g e numbers d u r i n g t h i s time.  L. s i n c l a i r i i showed r e d u c e d growth and c o n s i d e r a b l e  l o s s o f b l a d e s b u t was a p p a r e n t l y n o t i r r e p a r a b l y damaged.  97  These o b s e r v a t i o n s , t o g e t h e r w i t h t h e r e s u l t s i n laboratoryc u l t u r e and t h e g r e a t v a r i a t i o n i n s a l i n i t y w i t h i n t h e ranges of b o t h s p e c i e s i n d i c a t e t h a t s a l i n i t y i s o f l i t t l e importance i n c o n t r o l l i n g the d i s t r i b u t i o n of e i t h e r s p e c i e s . L. l o n g i p e s i s n o t n o r m a l l y found a s s o c i a t e d w i t h sand, a l t h o u g h t h e r e i s no e v i d e n c e from t h i s s t u d y t o i n d i c a t e t h a t i t s h o u l d be any l e s s w e l l adapted t o sand t h a n i s L. s i n c l a i r i i . The c h i e f r e a s o n may be s i m p l y t h a t w i t h i n t h e a r e a t o w h i c h L. l o n g i p e s i s adapted by i t s temperature r e q u i r e m e n t s t h e r e are  v e r y few sandy beaches o f t h e t y p e found i n Washington,  Oregon, and C a l i f o r n i a . The c h i e f f a c t o r c o n t r o l l i n g t h e d i s t r i b u t i o n o f L. s i n c l a i r i i w i t h i n i t s temperature range appears t o be t h e p r e s e n c e o f sand.  I t i s n e a r l y always r e s t r i c t e d t o beaches  w h i c h have a l a r g e s e a s o n a l f l u c t u a t i o n o f sand and where i t i s periodically buried. environment.  I t i s w e l l adapted to t h i s very harsh  I n such an a r e a , s e x u a l r e p r o d u c t i o n by  gametophytes i s d i f f i c u l t because o f t h e danger o f sand scouring.  F u r t h e r , i n such a w e l l - a d a p t e d p l a n t a n o n s e x u a l  means o f r e p r o d u c t i o n would ensure s u c c e s s i v e g e n e r a t i o n s o f g e n e t i c a l l y s i m i l a r , well-adapted plants.  Three o t h e r s p e c i e s  w h i c h a r e s i m i l a r l y w e l l - a d a p t e d t o t h i s type o f environment apparently lack sexual reproduction altogether: linearis  Gymnogongrus  (Smith, 1944), Ahnf e l t d a c o n c i n n a (Smith, 1944), and  Phaeostrophion i r r e g u l a r e  ( M a t h i e s o n , 1967).  Laboratory  e x p e r i m e n t s show t h a t s e x u a l r e p r o d u c t i o n i n L. s i n c l a i r i i i s s u p p r e s s e d even i n environments where sand s c o u r i n g i s a b s e n t .  98  The need f o r an a l t e r n a t i v e , or a c c e s s o r y , n o n s e x u a l means of  r e p r o d u c t i o n i n t h i s p l a n t i s t h e r e f o r e apparent.  This  s t u d y has shown t h a t such a means e x i s t s i n the p r o d u c t i o n o f new  s t i p e s and b l a d e s from h a p t e r a and can o p e r a t e even i f  the h a p t e r a a r e detached from the p a r e n t p l a n t . The g e n e r a l i z e d m e r i s t e m a t i c a c t i v i t y and the c o n s i d e r a b l e regeneration i n t h i s p l a n t serve a f u r t h e r adaptive f u n c t i o n i n e n a b l i n g the p l a n t t o r e c o v e r a f t e r v a r i o u s p a r t s have been accidentailyremoved.  T h i s i s a g r e a t advantage i n an  environment where sand s c o u r i n g and the f o r c e o f the s u r f a r e often very great. As L_. s i n c l a i r i i i s r a r e l y found where i t does n o t undergo p e r i o d i c b u r i a l under sand, i t would appear t h a t t h i s b u r i a l c o n f e r s some advantage on the p l a n t .  i t s unique methods o f  growth and r e p r o d u c t i o n e n a b l e i t t o s u r v i v e i n t h i s v e r y h o s t i l e environment which e x c l u d e s most o t h e r a l g a e . C o l o n i z a t i o n by outgrowths  from h a p t e r a i s v e r y s u c c e s s f u l i n  a r e a s w h i c h a r e p e r i o d i c a l l y b u r i e d under sand because spores or  gametophytes would be s c o u r e d away.  i n a r e a s which a r e not  b u r i e d under sand, c o l o n i z a t i o n by gametophytes or spores i s more s u c c e s s f u l because a much l a r g e r a r e a can be c o l o n i z e d i n a s i n g l e season.  S i n c e L. s i n c l a i r i i a p p a r e n t l y  reproduces  by gametophytes v e r y r a r e l y , i t i s e x c l u d e d from a r e a s where such r e p r o d u c t i o n i s an advantage. C o l o n i z a t i o n by h a p t e r a can o c c u r e a s i l y o n l y on of  areas  r o c k a d j a c e n t t o the a r e a where the p l a n t i s a l r e a d y  growing.  C o l o n i z a t i o n o f new  and i s o l a t e d r o c k s i s a more  99  d i f f i c u l t problem.  I t i s p o s s i b l e t h a t new  p l a n t s of  L. s i n c l a i r i i a r e i n i t i a t e d from gametophytes on such a r e a s . The  r e l e a s e o f s p o r e s a t a time when l i t t l e o t h e r  growth i s o c c u r r i n g and sand i s m o s t l y a b s e n t , and  algal a  subsequent r a p i d growth o f gametophytes and p r o d u c t i o n  of  young s p o r o p h y t e s , might e n a b l e the p l a n t s t o become e s t a b l i s h e d before  c o m p e t i t i o n and/or sand s c o u r i n g e l i m i n a t e d them.  It  i s assumed t h a t young s p o r o p h y t e s a r e much more r e s i s t a n t to sand s c o u r i n g than a r e gametophytes. I t was  n o t e d t h a t ; L . s i n c l a i r i i appears to be adapted to  exposure to s e v e r e s u r f .  i n the a r e a s o f g r e a t e s t exposure,  the p l a n t s are l a r g e r and c o n t i n u e b u r i a l by sand.  growing l o n g e r  after  A l s o , i n t h e s e a r e a s L. s i n c l a i r i i shows more  complete dominance on the r o c k s on w h i c h i t grows.  Thus, a t  A r c h Cape, on the e n t i r e r o c k where L. s i n c l a i r i i was o n l y one o t h e r s p e c i e s was  found.  L. s i n c l a i r i i can  studied, withstand  b u r i a l b e t t e r than many s m a l l e r p l a n t s because i t i s l o n g enough by l a t e summer so t h a t a p o r t i o n o f the b l a d e s protrudes any,  from the sand.  conduction  the p l a n t .  occurs  usually  I t i s n o t known to what e x t e n t , i f  from the exposed t o the b u r i e d p a r t s o f  When the p l a n t i s b u r i e d , g r e a t e r exposure to  s u r f m i g h t be an advantage i n t h a t g r e a t e r s u r f a c t i o n would cause g r e a t e r s t i r r i n g o f the sand and a t the same time a l l o w g r e a t e r water m o t i o n and a e r a t i o n around the b u r i e d p a r t s o f the p l a n t .  100  L. s i n c l a i r i i has been shown t o be u n i q u e l y adapted to a v e r y h a r s h environment. g e n e r a l l y l a c k s sand.  The environment o f L. l o n g i p e s However, i t too i s a v e r y h a r s h  environment, i n p a r t because o f the low seawater temperatures and even lower a i r t e m p e r a t u r e s t o w h i c h the p l a n t i s frequently subjected.  F u r t h e r s t u d i e s o f the type c a r r i e d  o u t on L. s i n c l a i r i i a r e needed t o determine e x a c t l y what e n v i r o n m e n t a l f a c t o r s a r e most s i g n i f i c a n t i n c o n t r o l l i n g L. l o n g i p e s and the ways i n w h i c h the p l a n t i s adapted t o i t s environment. the  Such i n f o r m a t i o n s h o u l d a l s o c l a r i f y  further  r e l a t i o n s h i p o f L. l o n g i p e s t o L. s i n c l a i r i i and the  r e l a t i o n s h i p o f t h e s e two unique s p e c i e s t o o t h e r k e l p s .  101  IX.  The Laminaria  SUMMARY  d i s t r i b u t i o n , ecology,  growth, and r e p r o d u c t i o n o f  s i n c l a i r i i and L. l o n g i p e s were s t u d i e d i n t h e  l a b o r a t o r y and on beaches i n A l a s k a , B r i t i s h Columbia, and Oregon over a two-year p e r i o d .  The two s p e c i e s d i f f e r  from  most o t h e r k e l p s i n h a v i n g m u l t i p l e s t i p e s a r i s i n g from an e n l a r g e d r h i z o m e - l i k e h o l d f a s t , composed o f many h a p t e r a . Each o f t h e s t i p e s b e a r s a b l a d e .  The two s p e c i e s a r e a l m o s t  i d e n t i c a l i n e x t e r n a l morphology,  p r e v i o u s workers have  d i s t i n g u i s h e d t h e two s p e c i e s p r i m a r i l y on t h e i n t e r n a l anatomy o f t h e s t i p e :  t h e s t i p e o f L. s i n c l a i r i i has  m u c i l a g e d u c t s whereas t h a t o f L. l o n g i p e s does n o t .  L.  l o n g i p e s o c c u r s from t h e K u r i l e I s l a n d s t h r o u g h t h e A l e u t i a n I s l a n d s and i n t o S o u t h e a s t A l a s k a . Northern  L. s i n c l a i r i i o c c u r s  B r i t i s h Columbia t o S o u t h e r n C a l i f o r n i a .  from  The g r o s s  d i s t r i b u t i o n o f b o t h s p e c i e s appears t o be c o n t r o l l e d b y seawater t e m p e r a t u r e .  S a l i n i t y appears t o have l i t t l e i n f l u e n c e  on d i s t r i b u t i o n , as b o t h s p e c i e s o c c u r  t h r o u g h o u t wide  r a n g e s o f s a l i n i t y and L. s i n c l a i r i i i s a b l e t o w i t h s t a n d  great  fluctuations i n salinity at a single site. S e a s o n a l c y c l e s o f growth and r e p r o d u c t i o n i n L. s i n c l a i r i i were s t u d i e d on t h r e e beaches i n N o r t h e r n p l a n t s a r e s u b j e c t e d t o heavy s u r f . o c c u r s i n e a r l y summer.  Oregon where t h e  The g r e a t e s t growth  As summer advances, t h e p l a n t s a r e  g r a d u a l l y b u r i e d under sand u n t i l o n l y t h e ends o f t h e b l a d e s protrude.  Growth d e c l i n e s d u r i n g t h i s p e r i o d and i s v e r y  slow  102  i n l a t e summer.  The f i r s t heavy storms i n f a l l remove t h e  sand and expose t h e p l a n t s . the p l a n t s b e a r r i p e s o r i .  I n November and e a r l y December L a t e r i n December t h e b l a d e s a r e  l o s t , l e a v i n g only the bare s t i p e s . are r e g e n e r a t e d  I n J a n u a r y new b l a d e s  and when they a r e o n l y 2-3 cm i n l e n g t h new  s o r i a r e produced.  S o r i a r e produced i n t o March and  t h e r e a f t e r o n l y v e g e t a t i v e growth o c c u r s .  I n March and A p r i l  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 new s t i p e s and b l a d e s from t h e h a p t e r a a t t h e m a r g i n s o f t h e h o l d f a s t . The  d i s t r i b u t i o n o f L. s i n c l a i r i i w i t h i n i t s temperature  r a n g e i s p r i m a r i l y c o n t r o l l e d b y t h e p r e s e n c e o f sand.  It is  a d a p t e d t o t h i s h a r s h environment b y i t s growth and reproduction.  I t p o s s e s s e s g r e a t powers o f r e g e n e r a t i o n and  i s p o t e n t i a l l y meristematic  i n a l m o s t any r e g i o n .  Although  i t r e g u l a r l y produces s o r i , t h e r e i s l i t t l e e v i d e n c e from e i t h e r f i e l d or l a b o r a t o r y studies to i n d i c a t e that the gametophytes w h i c h d e v e l o p from t h e s p o r e s i n these n o r m a l l y produce s p o r o p h y t e s .  sori  Sexual reproduction of t h i s  type i s d i f f i c u l t because o f t h e s c o u r i n g a c t i o n o f t h e sand. The normal method o f r e p r o d u c t i o n i s a p p a r e n t l y  vegetative  p r o l i f e r a t i o n from t h e h a p t e r a a t t h e margins o f t h e h o l d f a s t s . L. l o n g i p e s was o b s e r v e d i n t h e f i e l d i n A l a s k a on o n l y three occasions  i n summer and t w i c e i n w i n t e r .  g r e a t e s t d u r i n g t h e summer months. i n December.  Laboratory  Growth i s  The p l a n t s bear r i p e s o r i  cultures i n d i c a t e that sexual  reproduction i s very rare i n t h i s species. reduced i n winter but are not completely  The b l a d e s a r e  lost.  L_. l o n g i p e s i s  103  n o t n o r m a l l y a s s o c i a t e d w i t h sand. T r a n s p l a n t e x p e r i m e n t s showed t h a t each o f the two  species  can s u r v i v e f o r a time i n the h a b i t a t o f the o t h e r b u t cannot s u r v i v e a whole y e a r .  T r a n s p l a n t s t o s i t e s i n B r i t i s h Columbia  where the t e m p e r a t u r e range i s g r e a t e r than i n Oregon or A l a s k a showed t h a t L.. s i n c l a i r i i i s more a d v e r s e l y a f f e c t e d by low w i n t e r t e m p e r a t u r e s ,  whereas L. l o n g i p e s i s more a d v e r s e l y  a f f e c t e d by h i g h summer t e m p e r a t u r e s .  Transplanting to higher  o r lower t e m p e r a t u r e s d i d not a f f e c t the p r o d u c t i o n o f mucilage  ducts.  Comparison o f the two s p e c i e s shows they d i f f e r i n s e v e r a l p o i n t s besides mucilage  ducts, i n c l u d i n g length of s t i p e s ,  w i d t h o f b l a d e s , w i n t e r l o s s o f b l a d e s , morphology o f gametophytes and h a b i t a t .  The  evidence  c o n f i r m s t h a t they  s h o u l d be r e t a i n e d as two s e p a r a t e s p e c i e s .  104  X. A g a r d h , C. seen).  BIBLIOGRAPHY  1820. Species algarum  Agardh, J.G. 4:1-36.  1 8 6 7 . De L a m i n a r i e i s . Lund.  . . . . V o l . 1:1-168 ( N o t  Lunds Univ.  Anderson, C L . 1891. L i s t o f C a l i f o r n i a notes. z o e 2:217-225. Anon.  marine  Arsskr.  algae,  with  1965a. Ambient seawater temperature and s a l i n i t y . B i o l o g i s t ' s and Engineer's Daily Report. Vancouver Public Aquarium. 1965b. Record.  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K a i s o - z u h u ( I l l u s t r a t i o n s o f M a r i n e A l g a e ) (Jap. E x p l a n . ) (Not s e e n ) . Okamura, K. 1915. I c o n e s o f Japanese A l g a e I I I . [3] + 218 pp + [8] .  + 3 pis  . 1916. Enumerations o f Japanese A l g a e , 2nd E d i t i o n . (Not s e e n ) . Wks.  1928. A l g a e from Kamschatka. j a p a n . 1:52-55.  Rec.  Oceanogr.  1932. The d i s t r i b u t i o n o f marine a l g a e i n p a c i f i c w a t e r s . Rec. Oceanogr. Wks Japan 4:30-150.  108 Okamura, K. 1933. Y. K o b a y a s h i . (in  On the a l g a e from A l a s k a c o l l e c t e d by Rec. Oceanogr. Wks. j a p a n 5 ( l ) : 8 5 - 9 7 , 2 p i s .  1936. Nihon K a i s o s i Japanese).  (Marine A l g a e o f japan)  O l i p h a n t , M., Wyatt, B. and K u j a l a , N. 1962. Surface temperature and s a l i n i t y o b s e r v a t i o n s a t shore s t a t i o n s on the Oregon Coast f o r 1961. Ore. S t . U n i v . Dept. Ocean. ONR Data Rept. 8. P o s t e l s , A. and R u p r e c h t , F. 1840. . . i v + 22 pp + [ 2 ] , 40 p i s .  I l l u s t r a t i o n e s algarum St. Petersburg.  . .  P r o v a s o l i , L., M c L a u g h l i n , J.J.A., and Droop, M.R. 1957. The development o f a r t i f i c i a l c u l t u r e media f o r marine a l g a e . A r c h . M i k r o b i o l . 25:392-428. Robinson,, G.G.C. 1967. C y t o l o g i c a l i n v e s t i g a t i o n s o f the genus A l a r i a G r e v i l l e , as i t o c c u r s on the West c o a s t o f N o r t h America. Ph.D. T h e s i s , U n i v . B r i t i s h Columbia, x i + 136 pp. R u p r e c h t , F . J . 1851. Tange des o c h o t s k i s c h e n Meeres. I n M i d d e n d o r f f , A.T. von, R e i s e i n den a l i s s e r s t e n Norden und Osten S i b i r i e n s wahrend der J a h r e 1843 und 1844. B o t a n i k 1 ( 2 ) : 191-435, p i s . 9-18. St. Petersburg. Sanborn and Doty. 1947. The marine a l g a e o f the Coos Bay-Cape Arago R e g i o n o f Oregon. Ore. S t . Monogr. S t u d . Bot. No. 8, 66 pp, 4 p i , 1 map. S c a g e l , R.F. 1957. An a n n o t a t e d l i s t o f the marine a l g a e o f B r i t i s h Columbia and n o r t h e r n Washington. Nat. Mus. can. B u l l . 1 5 0 . v i + 289 pp. S e t c h e l l , W.A.  1896.  Notes on k e l p s .  E r y t h e a 4:41-48, p l . l .  1899. A l g a e o f the P r i b i l o f I s l a n d s . I n D. S. J o r d a n , Fur S e a l s and Fur S e a l I s l a n d s o f the N o r t h P a c i f i c Ocean, v o l 3:589-596, Washington. 1905. R e g e n e r a t i o n among k e l p s . P u b l . B o t . 2:139-169, p i s 15-17.  Univ. C a l .  1912. The k e l p s o f the U n i t e d S t a t e s and A l a s k a . Appendix K. I n Cameron, F.K. A preliminary r e p o r t on the f e r t i l i z e r r e s o u r c e s o f the U n i t e d S t a t e s . Appendix K. U.S. Senate Document No. 190, pp. 130-178. S e t c h e l l , W.A. America.  and Gardner, N.L. 1903. A l g a e o f N o r t h w e s t e r n U n i v . C a l . p u b l . B o t . 1 ( 3 ) 1 6 5 - 4 1 8 . p i s 17-27. :  109 S e t c h e l l , W.A. and Gardner, N.L. 1925. The marine algae of the p a c i f i c Coast o f N o r t h A m e r i c a . P t . I I I . Melanophyceae. U n i v . C a l . p u b l . B o t . 8(3):383-898. Shchapova, T.F. 1948. Geografiicheskoye rasprostaneniye p r e d s t a v i t e l e y poryadka L a m i n a r i a l e s v severnoy C h a s t i t i k h o g o Okeana ( G e o g r a p h i c a l d i s t r i b u t i o n o f r e p r e s e n t a t i v e s o f the Order o f L a m i n a r i a l e s i n the N o r t h e r n p a r t o f the p a c i f i c Ocean.)(In Russian) Trudy I n s t . Okeanol. 2:89-138. Shepard, F.P. 1963. Submarine Geology 1 map. Harper and Row, New Y o r k . S i l v a , p.c. 1957. 14(2):41-51.  (2nd Ed.)  Notes on p a c i f i c marine a l g a e .  x v i i i + 557  pp.  Madrono  S m i t h , G.M. 1944. M a r i n e A l g a e o f the Monterey p e n i n s u l a , C a l i f o r n i a . S t a n f o r d U n i v . p r e s s , i x + 622 pp. S t i l l , R., Wyatt, B. and K u j a l a , N. 1963. S u r f a c e temperature and s a l i n i t y o b s e r v a t i o n s a t shore s t a t i o n s on the Oregon Coast f o r 1962. Ore. S t . U n i v . Dept. Ocean. ONR Data Rept. 11. Sundene, O. 1963. R e p r o d u c t i o n and e c o l o g y o f Chorda N y t t Mag. B o t . 10:159-167.  tomentosa.  1964. The e c o l o g y o f L a m i n a r i a d i g i t a t a i n Norway i n v i e w o f t r a n s p l a n t e x p e r i m e n t s . N y t t Mag. Bot. 11:83-107. T a t e w a k i , M. 1931. The p r i m a r y s u r v e y o f the v e g e t a t i o n o f the M i d d l e K u r i l e s . J . F a c . A g r i c . Hokkaido U n i v . 2 9 ( 4 ) 1 2 7 - 1 9 0 , p i s I-X. :  T o k i d a , J . 1954. The marine a l g a e o f s o u t h e r n S a g h a l i e n . F a c . F i s h . Hokkaido U n i v . 2 ( l ) : l - 2 6 4 , p i s . I-XV. Van Overbeek, J . 15:291-299. B o t . Gaz.  1940a.  A u x i n i n marine a l g a e .  Mem.  P i . Physiol.  1940b. A u x i n i n marine p l a n t s . I I . 101:940-947.  V o z z h i n s k a y a , V.B. 1964. The bottom f l o r a o f S a k h a l i n (In R u s s i a n ) Trudy I n s t . Okeanol. 69:33 0-440. Widdowson, T.B. 1965. A s u r v e y o f the d i s t r i b u t i o n o f i n t e r t i d a l algae along a coast t r a n s i t i o n a l i n r e s p e c t to s a l i n i t y and t i d a l f a c t o r s . J . F i s h . Res. Bd. Can. 22(6):1425-1454. W i l l i a m s , L.G. agardhii.  1949. G r o w t h - r e g u l a t i n g substances i n L a m i n a r i a S c i e n c e 110:169.  110 Wyatt, B. and G i l b e r t / W. s a l i n i t y observations s t a t i o n s f o r 1965 and Ocean. ONR Data Rept.  1967. S u r f a c e temperature and a t p a c i f i c Northwest shore 1966. Ore. S t . U n i v . Dept. 25.  Wyatt, B., S t i l l R., and Haag, C. 1965. S u r f a c e temperature and s a l i n i t y o b s e r v a t i o n s a t p a c i f i c Northwest shore s t a t i o n s f o r 1963 and 1964. Ore. S t . U n i v . Dept. Ocean. ONR Data Rept. 21. Yamada, Y. 1935. The marine a l g a e from Urup, the m i d d l e K u r i l e s , e s p e c i a l l y from t h e v i c i n i t y o f Iema Bay. S c i . pap. I n s t . A l g o l . Res. Hokkaido U n i v . l ( l ) : l - 2 6 , p i s . I-X. Ag.  1938. O b s e r v a t i o n s on Arthrothamnus b i f i d u s J . S c i . Pap. I n s t . A l g o l . Res. Hokkaido U n i v . 2:113-118.  Yendo, K. 1909. Notes on a l g a e new t o Japan. Tokyo. 23(270):117-133. (Not  1910. seen)  K a i s a n Shokubutsu-gaku.  Bot.  Mag.,  (Marine Botany)  Z i n o v a , A.D. 1959. ( L i s t o f marine a l g a e o f S o u t h e r n S a k h a l i n and t h e S o u t h e r n K u r i l e I s l a n d s . ) ( I n R u s s i a n ) I s s l e d . D a l ' N e v o s t . Morea SSSR. 6:146-161. Z i n o v a , E.S. 1933. L e s a l g u e s de Kamtschatka ( i n R u s s i a n w i t h F r e n c h summary). I n s t . H y d r o l . E x p l o r . d. Mers d'URSS. F a s c . 17:7-42. 1954. (Marine a l g a e o f e a s t e r n Kamschatka). ( i n R u s s i a n ) Komarovskie C h t e n i y a B o t . I n s t . Akad. Nauk. SSSR. 2(9):365-400.  Ill  X I . TABLES I - X X I I I .  Table I  Seawater Temperature and s a l i n i t y over D i s t r i b u t i o n ranges o f L_. s i n c l a i r i i and L. l o n g i p e s .  LOCATION  MEAN FEB. (or c o l d e s t month) T°C S%  YEARS COVERED  0  [~~ Urup I s l a n d  0o5  1949-1962  (Mar.)  MEAN AUG„ (or warmest month) T°C S% 11.6  YEARLY MEAN T°C 5.1  SOURCE  S%,  Eber, Saur & S e t t e , 1968  Attu Island: Murder P o i n t P y r a m i d Cove  1959 1946- 1964  1.6 2.1  31. 9  32. 9  11. 3 9. 3  31. 0 31. 5  5. 4 5. 1  31o8  32.1  Anon. 1967d Anon. 1967d  *Sitka  1943-  1964  4.4  30o 4  14. 1  25. 5  8. 5  27.7  Anon. 1967d  Coronation Island  In Dec. 1966, 1 C° below S i t k a , 0.5% below S i t k a . 0  Hope I s l a n d , B.C» (Pine I s l a n d 1940-1965 records) R i v e r Jordan Whiffen S p i t •Stanley park L a push  Hollister 1966 Widdowson 1965 Widdowson 1965  31.3  10.0  31.8  1957  8.0  30o7  15.0  30.5  1957  8.0  30.7  15.0  30.5  1965-1967  7.1  27.9  12.1  26.8  9.4 27.6  Anon. 1965a, 1966a,1967a  7.4  30 3  11.6 32.3  9.8 31.2  Anon. 1967c  (Mar.) 7.6 27.7  (Jul.) 13.9 32.3  10.5 ?30.9?  (Neah Bay r e c o r d s ) 1936-1964 I n d i a n Beach (Seaside records c o n v e r t e d f o r T°)  31.7  7.2  .  1 9 6 6  "  .  1 9 6 7  o  8.6  Gilbert & Wyatt, 1968  Table I  i—i  Continued  H  ^|  9.7 31.4 (Mar.)  13.2 32.2 (Jul.)  1966-1967  (Mar.) 7.1 21.5  (Jul.) 13.4 33.5  pescadero P o i n t ( p a c i f i c Grove records)  1920-1964  11.9  33.2  p o r t Hueneme  1920-1963  13.2  **Santa Monica  1946-1964  13o6  A r c h Cape  1960-1963  S h o r t Sand Beach (Seaside r e c o r d s c o n v e r t e d f o r T°)  * Transplant station only.  11.3  30.9  Kujala, Wyatt, e t a l , 1961-1965  10.0  Gilbert & Wyatt, 1968  14.2 33.7 (Sep.)  12.9  33.5 . Anon. 1967d  33„3  16.8  33.6  14.8  33.6  Anon. 1967d  33.5  19.7  33.7  16.2  33.7  Anon. 1967d  No L. s i n c l a i r i i o r L. l o n g i p e s p r e s e n t  ** L o c a t e d j u s t s o u t h o f s o u t h e r n l i m i t e s o f d i s t r i b u t i o n o f L. s i n c l a i r i i (Fig.l).  114 Table I I D i s t r i b u t i o n o f Laminaria SITE B r i t i s h Columbia P l o v e r i s . , Hope I s l a n d Box I s l a n d , W. Coast Vancouver I s . * * R i v e r J o r d a n , W.C. Vancouver I s . * * W h i f f e n S p i t , W.C. Vancouver I s .  sinclairii ACC. NO.  50°56 'N, 127°58'W  SOURCE * UBC  49°04 'N, 125°47'W  UBC  17012  48°25 .4'N, 124°04'W  JWM+  48°21 •N, 123°43'W  JWM  POSITION  Washington Salmon Bank, San Juan I s . 48°26'N, 123°01'W San Juan Co. 48°23'N, 124°43.5'W Cape F l a t t e r y , C l a l l a m Co. p a r t r i d g e Bank, San Juan Co. 48 16'N, 122 51'W Agate Beach, C l a l l a m Co. **La push, C l a l l a m Co. Oregon **Indian Point, Indian Beach, C l a t s o p Co. * * B a l d P o i n t , I n d i a n Beach, C l a t s o p Co. * * E c o l a p o i n t , C l a t s o p Co. * * A r c h Cape, C l a t s o p Co. **Cape F a l c o n , T i l l a m o o k Co. * * S h o r t Sand Beach, N. end, T i l l a m o o k Co. * * S h o r t Sand Beach, S. end, T i l l a m o o k Co. Cape Kiawanda, T i l l a m o o k Co. Y a q u i n a Bay Mouth, L i n c o l n Co. Coos Bay-Cape A r a g o R e g i o n , Coos Co. (Bassendorf Beach, L i g h t h o u s e Beach, Squaw I s l a n d , N o r t h Bay, Coos Bay, cape Arago) California T r i n i d a d , Humboldt Co. S h e l l Beach, Sonoma Co. Second S l e d Road, D i l l o n Beach, M a r i n Co. Duxbury Reef, M a r i n Co. San F r a n c i s c o , San F r a n c i s c o Co. * * P e s c a d e r o P o i n t , San Mateo Co.  411  48 °10«N, 124°43.8'W 47 53.9'N, 124 37.6'W  Druehi, 1965 UC++ Druehl, 1965 17023 UBC 24574 UBC  45°55.9'N, 123°58.8'W  UBC  24673  123 58.6 'W 123°58.5 'W 12.3°58.0'W 123 58.5'W  JWM JWM JWM UBC  30535  45°45.8'N, 123°58.2'W  JWM  45";45.5 N, 123 58'W 45 13.2'N, 123°9.6'W  UBC UC  44°36.8'N, 124°4.1'W  Kjeldsen, 1967  43°20'N, 124°23'W  Sanborn & Doty, 1947  41°03.8«N, 124 09'W approx. 38 30'N, 123 25'W  UC  38°15.3'N, 122°25.2'W 37°53.3'N, 122°42'W  UBC UC  37°47'N, 122°30.8'W  UC  37°14.5'N, 122°25.2'W  UBC  45^55.6'N, 45 55.2'N, 45°48.2'N, 45 46.4'N,  ,  24933  UC  24464  115  Table I I - Continued  SITE  Cruz Co. S a n t a C r u z , S a n t a Cruz Co. A s i l o m a r p o i n t , Monterey Co. P o i n t Lobos, Monterey Co. L u c i a , Monterey Co. p i e d r a s B l a n c a s P o i n t , San L u i s Obispo Co. E s t e r o Bay, San L u i s Obispo Co. Morro Bay, San L u i s Obispo Co. Pismo Beach, San L u i s Obispo Co. p o i n t S a l , S a n t a B a r b a r a Co. P o i n t p e d e r n a l e s , Santa B a r b a r a Co. G a v i o t a , S a n t a B a r b a r a Co. P o i n t Conception, Santa B a r b a r a Co. C a r p i n t e r i a Beach S t a t e P a r k , V e n t u r a Co. Two M i . N.W. o f V e n t u r a , V e n t u r a Co. M u s s e l S h o a l s , V e n t u r a Co.  * ** + ++  POSITION  SOURCE  approx. 37°10' N, 122°20*W 36°57' N, 122°01.8'W 36°37. 5'N, 121°56. 5'w 36°31. 2'N, 121°57. 2 w 36°01. 3 N, 121°33 W  UC UC Smith,1944 Smith,1944 UC  35°40' N, 121 17'W 35°27« N, 120°57 «W 35°22'N, 120°51.3'W  UC UC UC  1  1  1  u  35°08. 9 N, 120°38. 8 w UC 34°54. 2 N, 120°40. 4' vr UC 34°36. 1 N, 120°38. 5'w 34°28. 3 N, 120°12. 3 'w  UC UC  34°26. 8 N, 120°28. 2 'w  UC Dawson, 19 58a Dawson 1958a Dawson 1958a  34°23. 1 N, 119°30'W 34°22. 5 N, 119°28'W 34°19'N, 119°23.3'w  UBC = Specimen i s i n P h y c o l o g i c a l Herbarium, U.B.C. Seen b y a u t h o r a t t h i s s i t e JWM = Seen by a u t h o r a t t h i s s i t e b u t no c o l l e c t i o n s made. UC = Specimen i s i n Herbarium, U n i v . o f C a l i f o r n i a , B e r k e l e y .  11.6 Table I I I D i s t r i b u t i o n o f Laminaria longipes  UoS.S.R.  SITE  Kurile Islands* Urup i s l a n d (Uruppu) Shimushir i s . (Simusiru)  POSITION  SOURCE  ACC. NO.  Okamura, 1928; N a g a i , 1940 152°uO'E Okamura, 1928 N a g a i , 1940 152°28'E N a g a i , 1940 152°49'E N a g a i , 1940 154°31'E N a g a i , 1940 153°51'E N a g a i , 1940 154°31'E N a g a i , 1940 154°45 «E N a g a i , 1940 155°50'E~ Miyabe & 156°20'E -Nagai, 1932; 157°27*E N a g a i , 1940 143°00'E Miyabe & Nagai,1932; V o z z h i n s k a y a , 1964 166 15'E Miyabe & Nagai 1932 158°36'E Z i n o v a , 1933 p Z i n o v a , 1933 159°57"E Z i n o v a , 1954 160°36'E Z i n o v a , 19 54 162°24'E Okamura, 1928 p Okamura, 1928 p Okamura, 1928 p Z i n o v a , 1954  46°00'N, 150°00'E 47°00»N,  Ketoy I s . (Ketoi) Yankicha I s l s . (Usisiru) Matua i s . (Matuwa) Lovushki I s . (Musisiru) K h a r i m k o t a n I s . (Harumkotan) Onekotan I s . (Onnekotan) paramushir I s . (paramusiru) Shumushu I s . (Simusiyu) Atlasova I s . (Alaid, Araid) S. S a k h a l i n  47°20»N, 47°31'N, 48°05«N, 48°32'N, 49 °07 'N/ 49°25'N, 50°25'N, 50°45'N, 50°53'N, 48°00«N,  Bering Island Kamschatka A v a c h i n s k a y a Bay Kuimska Bay Morzhovaya Bay K r o n o t s k y Bay pankara Drankinsky Barankorfa Zavodsk Cape  55 00 'N, 52°56 'N, p 53°16 'N, 54°12 •N, 58°37 'N, p p p  Alaska St. Paul Island  57°10 'N, 170°15'W  Aleutian Islands Murder p o i n t , A t t u I s . Casco Bay, A t t u I s . Chichagof P t , A t t u I s . Agattu i s .  Setchell & Gardner, 1903  52°48 •N, 52°48 'N, 52°57 'N, 52°55 'N,  UBC+ UBC UBC Setchell & Gardner,1903 Setchell & Gardner,1903 UBC  8145 8309 8381  UBC UBC UBC  8391 8150 8361  UBC  13593  UBC  27796  Kiska I s . T r a p p e r ' s Cove, Adak I s . N o r t h I s , Bay o f I s l a n d s Adak I s . Cape Agagdak, Adak I s . Zeto P o i n t , Adak I s . Bugle P o i n t , Great Sitkin I s . **Ram P t . Beach, C h e r n o f s k i Hbr, U n a l a s k a i s .  /  173°09'E 173°10'E 173°15'E 173°10'E  52°00 *N, 177°30'E 51°48 'N, 176°50'W 51°50 •N, 176°48'W 52°00 'N, 176°37"W 51°55 'N, 176°34'W 52°02 .3' N, 175°58.8'W 53°24 .6'N. 167 31.6'W  9631  117  Table I I I - Continued SITE  POSITION  **Ram P t . , C h e r n o f s k i Hbr, Unalaska I s . 53°25 'N, 167 31 .5'W **Cape A i a k - L a n c e P t . , Unalaska I s . 53°19..1'N, o 25. 9 •w * * S t a r a y a Bay, U n a l a s k a I s . 53°37..4-N, 165°30. 6 W Cape S a r i c h e f I , Unimak I s . 54 35 'N, 164 56 •w Cape S a r i c h e f I I , Unimak Is. 54°35 'N, 164^56 •w **Raven P o i n t , Cape S a r i c h e f , Unimak I s . 54°38 'N, 164^51 'W **E. Anchor Cove, I k a t a n Pen* Unimak I s . 54°41..6'N, 163°03. 2 W Gulf o f Alaska * * E a g l e Rock, N.E. Harbor Sanak I s . .6'N, o o 35. 4 •w **Nagai I s l a n d 55°12..6'N, 159°55. 2 'W **paul i s . 55°48 .7 'N, o 21 W * * C h i g n i k Bay, Nakchamik I s . 56°21 .3 • N, o 48. 2 •w **Aghiyuk I s . , S e m i d i i s l s 56°13..7'N, o 47 W **Chirikof Is. 55 49..5'N, 155°33. 5 •w **Gurney Bay, Cape I k o l i k Kodiak I s . 56°T7..7 'N, 154°44. 9 •w pasagshak p o i n t , K o d i a k I s .57 25 'N, 152 29 'W **Cape C h i n i a k , K o d i a k I s . 57°37..3-N, o 09. 5 •w **Chiniak I s . 57°36..6"N, 152° 09..6•w * * P e r i l Cape, Afognak I s . 58°07..5-N, 1 5 2 ° 16 W **Kayak I s . 59°51 »N, 144 33 •w **Wingham I s . 60 03 'N, 144°24 •w Southeast A l a s k a **Cape Spencer 58°14 'N, 136 35 •w **Cape Ommaney, Baranof I s . 56°10 'N, 134 40 •w * * A a t s Bay, C o r o n a t i o n I s . 55"53..7'N, o 16 W **Helm p o i n t . C o r o n a t i o n I s . 55 49..6'N, 134°17 W Washington ++Salmon Bank, San Juan I s . 48° 2 6 'N, 123 01 •w 1 6 7  5 4  2 6  1 6 2  1 5 9  1 5 7 1 5 6  1 5 2  1 3 4  *  SOURCE  ACC.NO.  UBC  27797  UBC UBC UBC  27724 27793 8338  UBC  8214  UBC  27790  UBC  27736  UBC UBC UBC UBC UBC UBC  13620 27715 26878 27836 27831 27672  UBC UBC UBC UBC UBC UBC UBC  27071 8162 8161 27682 25644 20935 22765  UBC UBC UBC UBC  20865 20384 20336 20319  Druehi, 1968  Ceded b y j a p a n t o U.S.S.R. i n 1945; names i n p a r e n t h e s e s a r e former Japanese names. ** L. l o n g i p e s seen b y a u t h o r a t t h i s s i t e + UBC = Specimen i s i n P h y c o l o g i c a l Herbarium, U.B.C. ++ S u b t i d a l Only  118  T a b l e IV  A n a l y s i s o f Sand G r a i n S i z e on Oregon Beaches  SIZE 2 nun +  INDIAN BEACH %  ARCH CAPE %  SHORT SAND BEACH %  0.01  0.04  0.01  0.06  0.15  0.02  0.61  2.14  0.27  67.85  74.99  75.22  31.41  22.64  24.43  0.05 - 0.1 mm  0.05  0.01  0.04  L e s s t h a n 0.05 mm  0.01  0.03  0.01  Very coarse  1 - 2 mm Coarse  0.5 - 1 mm Medium  0.25 - 0.5 mm Fine  0.1 - 0.25 mm Very f i n e  Total sieved:  100.00%  100.00%  100.00%  2575.5 g  2427.2 g  2415.0 g  119  T a b l e V.  Temperature and p r e c i p i t a t i o n a t A l a s k a s t a t i o n s  1965  1966  1967  30-Year Mean  Ppt Ppt T°C T°C T°C (xn.) Ltka ( S i t k a M a g n e t i c ) (Anon. , 1966b, 1967b, 1968a) T°C  Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. m.  Ppt.  -1.8 0.1 2.4 4.6 5.8 8.9 12.3 12.4 11.0 7.1 1.8 -0.3  12.81 7.31 2.81 5.42 6.55 8.60 2.42 6.90 6.02 18.66 5.05 11.51  -3.9 1.2 2.1 4.1 6.4 10.8 12.7 12.1 10.8 5.0 0.4 0.9  5.61 6.99 7.68 5.49 9.78 1.11 4.59 8.63 11.13 20.49 8.72 4.96  -0.8 1.3 -0.7 4.2 8.1 10.9 12.1 13.9 10.9 7.2 2.5 0.7  7.91 8.97 3.44 2.87 4.46 2.60 4.73 7.43 16.03 14.07 12.16 6.26  3.2 4.7 5.9 9.0 12.2 14.9 16.3 16.7 14.9 10.8 6.9 3.8  7.77 6.38 6.95 5.35 4.66 3.46 5.20 7.86 11.49 15.27 12.01 10.17  5.3  93.88  5.2  95.18  5.9  90.93  9.9  96.57  C o r o n a t i o n I s l a n d (Cape D e c i s i o n ) ( A n o n . , 1966b, 1967b, 1968a ) Jan.  Feb.  Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Ann.  0.5 2.2 3.9 6.0 6.1 9.4 11.1 11.6 10.3 8.6 4.4 1.8  12.09 8.64 1.66 6.07 5.30 3.83 2.58 1.53 2.63 18.43 4.22 8.81  -1.5 2.8 3.5 4.9 6.4 10.3 11.6 11.4 10.5 6.4 2.7 2.5  5.66 7.23 10.48 3.03 7.65 1.48 2.67 4.42 10.65 11.17 3.86 7.39  1.1 3.1 1.0 5.2 8.7 9.5 11.2 13.6 11.8 8.4 5.2 2.8  6.84 9.55 1.08 1.77 4.07 2.19 3.94 6.07 10.65 11.30 10.97 9.53  0.9 1.9 2.6 4.8 7.3 9.7 11.4 11.7 10.4 7.9 4.7 2.9  6.19 5.54 5.35 4.85 4.20 2.73 3.64 5.05 7.56 12.25 9.68 9.08  6.3  75.77  6.0  75.69  6.8  77.96  6.3  76.12  120 Table V - Continued 1965  1966  1967  T°C P p t . T°C P p t . T°C A t t u I s l a n d (Anon., 1966b, 1967b, 1968a) Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Ann.  -0.7 0.4 0.5 2.7 4.8 6.7 9.1 10.4 8.5 5.0 1.9 0.4  5.47 15.45 17.80 16.60 2.59 1.26 5.36 3.96 3.93 6.72 3.93 5.33  -0.8 -2.1 -0.5 0.9 3.8 5.8 8.6 10.6 9.7 5.4 3.4 0.4  5.14 4.90 2.62 3.48 7.81 2.18 2.90 4.39 11.52 5.02 9.50 3„75  4.3  88.40  3.8  63.21  Table V I  -1.2 -1.7 0.3 1.4 4.9 7.2 9.0 10.9 8.9 5.8 1.9 0.4  15-Year Mean  Ppt.  T°C  6.71 5.87 0.65 0.02 0.01 0.04 1.20 3„74 5.87 7.81 1.95 0.78  -0.6 -0.7 -0.1 1.7 4.0 6.6 9.0 10.5 8.9 5.3 2.1 0.7  4.0 34.65  Ppt.  4.18 4.14 3.86 4.68 3.85 3.19 4.75 5.64 6.04 6.68 4.79 4.35  4.0 56.15  Mean Temperature on Urup I s l a n d ( T a t e w a k i , 1931)  Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.  T°C -1 t o -8 -3 t o -8.5 -4 t o -8 1 to 2 2.5 t o 4 5 t o 10 11 10 t o 15 9.5 t o 11 2 to 3 No D a t a 0 t o -1.5  Ann.  2.5 t o 4.0  121  T a b l e V I I Temperature and p r e c i p i t a t i o n a t B r i t i s h Columbia S t a t i o n s B u l l Harbour ( f o r Hope I s l a n d )  Jan. Feb. Mar. Apr. May jun. Jul. Aug. Sep. Oct. Nov. Dec. ANNUAL  Long term Mean Ppt T° 7.64 3.8 4.4 5.71 5.9 5.96 4.27 7.6 10.0 3.06 12.0 1.95 13.5 2.59 2.94 13.7 12.2 4.73 9.7 9.12 6.6 9.72 4.9 10.37 8.7 68.06  Sooke (E. Sooke - A n d e r s o n Cove) (Anon., 1965b, 1966c, 1967c) 1965 T  Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.  °  c  * * * * * * * * * * * *  1966  «?« \ fc  * * * *  **  * * * * * *  T  °  * * * *  c  10.7 13.2 15.0 15.8 14.7 9.4 6.9 6.1  p  P  1967 f c  * * * *  1.02 0.80 1.07 0.55 1.43 6.03 5.98 9.96  T  °  c  p  P  Long Term Mean  f c  T  5.0 5.7 5.1 7.0 11.4 15.4  13.88 8.18 3.76 2.62 1.07 0.25  18.0 15.4 10.7 7.0 4.1  0.00 2.91 11.58 4.32 7.46  *  *  °  c  p  P -  * * * * * * * * * * * *  fc  * * * * * * * * * * * *  122  Table V I I  Continued  R i v e r J o r d a n (Anon., 1965b, 1966c, 1967c)  Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. ANNUAL  1965 T°C P p t . (in.) 4.0 12.57 5.3 12.69 6.3 1.58 8.2 4.06 9.6 3.80 12.6 0 66 0.84 * 2.28 * 12.2 3.47 8.26 * 8.5 11.26 9.9 10.35 o  *  71.85  Vancouver ( K i t s i l a n o ) 1965 T C Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.  1966 T°C P p t . 4.9 5.8 6.0 8.2 9.8 12.5 14.1 14.8 13.8  *  11.16 6.63 9.14 2.65 3.01 2.81 1.49 1.99 2.97 10.55 12.55 18.83  *  83.98  *  7.4  5.8 5.4 7.0 10.4 14.3  *  15.9  * *  7.8 4.1  *  1966  3.7 8.85 4.8 3.51 6.9 4.20 9.2 1.24 12.7 2.43 15.2 2.54 17.1 2.84 17.8 1.67 15.8 2.85 10.0 7.09 7.1 9.45 6.1 14.62  10.38 50.11  10.53 61.29  * = No Data  *  Long Term Mean Ppt. T°C  22.28 17.24 8.58 3.48 1.50 0.29 0.80 0.28 5.67 22.71 8.53 15.71  4.0 4.1 6.5 9.0 11.2 13.0 15.0 15.1 14.3 11.0 8.4 5.3  10.37 8.59 6.80 4.52 2.74 1.90 1 15 1.58 3.47 8.50 10.61 13.19  107.07  9.7  73.42  0  (Anon. , 1965b, 1966c, 1967c)  Ppt. (in.) 7.35 9.63 2.26 2.45 2.15 0.57 0.35 3.11 0.58 8.20 5.68 7.78  2.8 4.6 5.7 9.9 11.6 16.1 18.8 18.0 13.8 11.7 7.8 3.8  1967 Ppt. T°C  T°C  Ppt.  1967 T°C  Ppt.  4.8 5.5 5.7 8.0 13.1 18.3 18.8 20.3  11.87 5.53 5.72 3.21 2.15 0.55 1.07 0.23  * * * * *  * * * * *  Long Term Mean (HMCS D i s c o v e r y ) T°C Ppt. 5.7 5.9  8.59 6.41  12.6 17.4  2.67 2.67 1.70 1.64 3.38 6.84 8.14 9.95  * * *  19.4 16.4 11.0 7.3 4.2  *  * *  61.75  123  Table V I I I  Temperature and P r e c i p i t a t i o n a t Oregon S t a t i o n s  S e a s i d e (Anon., 1966b, 1967b, 1968a) ~ 1965 T C  Un.)  „ 1966 Ppt. T"C  Long Term Mean Ppt. T°C  1967 Ppt. T C  Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.  6.4 7.4 9.1 9.8 10.6 12.8 15.1 16.4 14.6 13.6 10.8 6.1  19.11 6.42 1.22 4.49 2.91 1.18 0.41 1.85 0.49 3.75 12.53 12.89  6.8 7.1 7.3 10.4 10.5 14.1 15.2 15.2 15.8 12.2 9.9 8.7  10.42 6.81 9.82 2.92 1.95 1.64 0.81 0.76 1.98 6.18 9.17 15.63  7.7 7.4 6.9 7.5 11.3 14.4 16.0 16.4 16.8 13.4 10.3 6.3  16.85 6.86 8.75 5.48 1.00 1.33 0.23 0.09 2.46 10.55 7.47 11.59  6.3 7.2 7.9 9.7 11.8 14.0 15.3 15.7 14.9 12.5 9.1 7.4  11.90 9.84 9.27 5.41 3.31 3.12 1.23 1.53 2.94 7.57 10.40 13.18  Ann.  11.1  67.25  11.1  68.09  11.2  72.66  11.0  79.70  A r c h Cape S t a t i o n s A & B 1965  P r e c i p i t a t i o n only (in.)  1966  1967 Ten-year ( S t a t i o n Mean A  B 22.60 7.62 1.50 7.59 2.94 1.57 1.24 2 62 1.17 4.49 11.83 16.76  A 8.98 6.06 9.97 2.13 2.39 1.90 1.36 1.36 3.34 6.76 10.96 14.70  B 11.14 6.68 12.37 2.71 2.69 2.15 1.46 1.85 3.43 8.05 9.27 20.81  A  B  Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.  A 17.99 6.34 1.68 6.41 2.41 1.54 1.26 2.37 1.02 3„78 11.42 14.53  17.14 5.42 10.78 5.16 2.12 1.46 0.47 0.20 4.16 9.00 6.16 12„49  21.79 6.73 13.05 5.75 2.10 1.65 0.51 0.20 4.51 12.59 7.20 15.74  12.71 9.77 8.82 5.60 3.29 3.11 1.26 2.19 3.32 7.26 12.09 11.46  Ann.  70o35  81.93  69.89  82.61  74.54  91.82  81.45  0  124  Table IX  Long Term Temperature and P r e c i p i t a t i o n a t two C a l i f o r n i a S t a t i o n s .  S a n t a C r u z , S a n t a Cruz Co. (Anon., 1968a) T°C  Ppt.(in.)  Oxnard, V e n t u r a Co. (Anon., 1968a) T°C  Ppt.  Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.  9.4 10.4 11.7 13.0 14.7 16.9 17.2 17.2 17.4 15.4 12.6 10.3  6.84 5.81 4.15 2.11 1.01 0.21 0.03 0.06 0.27 1.38 9.58 7.12  11.9 12.2 12.9 14.0 15.2 16.4 18.2 18.4 18.1 16.8 14.8 12.9  3.33  Ann  13.9  31.25  15.2  14.75  2o99  2.27 1.13 0.13 0.05 0.00 0.03 0.08 0.40 1.14 3.20  T a b l e X Measured V a l u e s o f Seawater Temperature and S a l i n i t y on Oregon Beaches  INDIAN BEACH Date  S%o  MONTHLY MEAN SEASIDE  SHORT SAND BEACH T°C  Date  T°C  Date  7.8  31.6  14.8  10.0  Sep. '66 Oct. '66  31.8  13.4  T°C  17 Aug.  •66  33.843  13 Sep.  •66  32.058  8.0  14 Sep.  •66  15 Oct.  •66  30.898  9.0  16 Oct.  •66  11 Nov.  •66  30.840  10o0  12 Nov.  '66  8.876  10.0  Nov.  '66  31.6  11.8  27 Dec.  •66  25.210  9 2  28 Dec.  •66  23.974  9.0  Dec.  •66  30.1  11.4  24 Jan.  '67  29.689  8.5  25 J a n .  •67  16.222  8.5  27.9  10.6  23 Feb.  '67  27.735  9.0  24 Feb.  '67  21.479  8.5  Jan. '67 Feb. '67  26.3  9.9  29 Mar.  '67  28.608  7o2  28 Mar.  •67  17.797  7.2  Mar.  •67  27.4  9.6  24 A p r .  '67  29.252  9.0  26 A p r .  •67  21.103  9.5  •67  26.2  11.0  23 May  •67  25.821  9.5  22 May  •67  24.194  10.0  Apr. May  •67  27.5  12.8  20 Jun.  •67  28.988  13.0  22 J u n .  •67  8.521  11.0  J u n . •67  27.9  13.9  19 J u l .  •67  23.827  15.7  20 J U l o  '67  5.357  14.5  Jul.  '67  29.8  15.9  17 Aug.  '67  32.295  12.0  18 Aug.  •67  33.523  9.0  Aug.  •67  29.7  14.5  2 Sep.  •67  32.190  14.0  3 Sep.  '67  31.115  13.0  29.0  15.8  0  30.833  -  Sep. •67  * Wyatt and G i l b e r t , 1967; G i l b e r t and Wyatt, 1968.  126 Table X I  1966  1967  C a l c u l a t e d Mean M o n t h l y Seawater * on Oregon Beaches .  Temperatures  INDIAN BEACH  SHORT SAND BEACH  ( S e a s i d e - 2.0 C°)  ( S e a s i d e - 2.5 C°)  Oct.  11.4^  10.9  Nov.  9.8  9.3  Dec.  9.4  8.9  Jan.  8.6  8.1  Feb.  7.9  7.4  Mar.  7.6  7.1  Apr.  9.0  8.5  May  10.8  10.3  Jun.  11.9  11.4  Jul.  13.9  13.4  Aug.  12.5  12.0  Sep,  13.8  13.3  10.55°C  10.05°C  Mean:  *  C a l c u l a t e d b y s u b t r a c t i n g mean d i f f e r e n c e from S e a s i d e monthly means (Wyatt and G i l b e r t , 1967; G i l b e r t and Wyatt, 1968) f o r October 1966 t o September 1967.  127  T a b l e X I I Mean v a l u e s o f Seawater Temperature and „ *  S a l i n i t y a t A r c h Cape f o r 1960-1963. MONTH  T°C  S%»  January  9.03  30.84  February  9.70  31.36  March  9.01  31.03  April  10.88  28.66  May  12.36  29.53  June  12.61  30.91  July  13.13  31.62  August  13.22  32o24  September  12.46  32.12  October  12.41  32.00  November  11.15  30.96  December  9.80  30.45  11.31  30.97  MEAN:  *  K u j a l a and Wyatt, 1961; O l i p h a n t , Wyatt and K u j a l a , 1962; S t i l l , Wyatt and K u j a l a , 1963;  Wyatt, S t i l l ,  and Haag, 1965.  128  Table X I I I  Sand L e v e l a t I n d i a n Beach. p o i n t s shown on F i g u r e 7.  L o c a t i o n of  A l l h e i g h t s are  cm down from s t a n d a r d r e f e r e n c e p o i n t . SAND by A: DATE  ROCK A  X (Inner)  •66:17/8  146  208  183  193  257  213  13/9  146  157  163  193  246  *  15/10  146  246  218  193  320  254  11/11  146  246  218  193  234  *  27/12  146  246  218  193  259  236  '67:24/1  146  246  218  193  259  236  23/2  146  246  218  193  259  236  29/3  146  246  218  193  259  236  25/4  146  246  218  193  259  236  25/5  146  246  218  193 J  191  185  21/6  146  218  *  193  198  *  19/7  146 i  112  112  193 .'  137  137  17/8  146 J  130  130  193 I  137  137  146  246  218  193  208  *  2/9  Y (Outer)  PT. F  SAND b y F: E D (Inner) (Outei  * := No D a t a 1 •  - Rock i s B u r i e d  -  Number u n d e r l i n e d = under water  Table XIV  Sand L e v e l a t S h o r t Sand Beach. p o i n t s shown on F i g u r e 8.  Location of  A l l heights are  cm down from s t a n d a r d r e f e r e n c e p o i n t .  ROCK B 213  SAND by B 267  239 269 277 320  213 213 213  252 282 279  213  287  320 320 320  213  287  24/2 28/3  211 211 211  213 213  287 287  26/4  211  213  *  24/5  211  213  22/6 20/7  211 ^ 211  297 320 274 302  213 213  287 259  18/8  2ll  *  3/9  211  249  DATE •66:18/8 14/9 16/10 11/11 28/12 •67:25/1  ROCK A 211 211 211 211 211  SAND by A 269  BEACH NORTH OF STREAM N. o f A N. o f B  * * * * * * * * * *  287  198 198  213  *  213  254  * = No Data  * * * * * * * * * * *  WATER-LEVEL DOWN FROM TOP OF ROCK A ROCK  * * 81  * * * * * * *  * * * * 94  * * * 135  * 64  207  64 33  168  *  18  13  226  *  28  *  25  130  T a b l e XV  Sand H e i g h t s o f Beach a t A r c h Cape, Oregon (Markham, 1967 unpubl.)  Date Distance 0+00 +50 1+00 +50 2+00 +50 3+00 +50 4+00 +50 5+00 +50 6+00 +50 Edge o f cobbles  31 J u l 66 50.00 40.00 37.18 36.03 35.60 33.56 31.83 31.10 29.99  Date  24 Feb 67  Mar 67  50.00 39.83 37.93 35.93 34.37 33.05 31.87 30.77 29.82 28.87 28.02 27.12 26.27  50.00 39.96 37.26 35.39 33.82 32.29 31.02 29.89 28.94 28.16 27.59 26.84  Distance 0+00 +50 1+00 +50 2+00 +50 3+00 +50 4+00 +50 5+00 +50 6+00 +50 7+00 +50 8+00  Deep pool  5 Sep 66 50.00 40.53 39.72 38.86 38.16 37.06 35.98 34.28 32.72 31.70 31.06 30.48  2 Oct 66 50.00 40.18 38.13 36.50 35.16 33.71 32.21 31.01 30.01 29.16 28.31  2 Nov 66 50.00 40.30 38.20 36.66 34.89 33.34 31.95 31.39 30.09  27 Nov 66 50.00 40.31 37.98 36.03 35.31 32.93 31.63 30.44 29.41 28.58 27.80 27.18 26.68 26.28  31 Dec 66 50.00 39.85 37.82 36.22 34.77 33.42 32.22 31.12 30.32 29.57 28.62  0+50.4  30 A p r 67 50.00 39.93 37.52 35.57 3.3.65 32.11 30.88 29.98 28.42 28.62 28.42 28.17 27.72 27.27 28.82 25.87 25.07  24 May 67  1Jul 67  6 Aug 67  50.00 40.07 37.57 35.84 34.08 32.72 31.35 30.26 29.51 29.02 28.45 29.02 27.16  50.00 40.09 37.63 36.49 34.58 32.84 31.45 30.63 29.59 29.59 26.87 27.27 27.07  50.00 40.10 37.60 36.50 35.00 32.66 30.59 29.80 29.23 28.40 28.00 27.84 26.64 26.12 25.67  26 J a n 67 50 00 39.60 37.42 35.72 34.25 33.14 31.78 30.78 29.81 28.85 28.14 27.60 26.74 o  0+48.4  Edge o f 0+47.0 0+47.0 0+44 0+40 Cobbles N o t e s : D i s t a n c e 0+00 i s a f i x e d p o i n t from w h i c h a l l o t h e r p o i n t s a r e measured and e s t a b l i s h e d a t 5 0 - f o o t i n t e r v a l s ; i t has a r b i t r a r i l y been a s s i g n e d a h e i g h t o f +50 f e e t and a l l o t h e r h e i g h t s have been c o r r e c t e d t o t h i s r e f e r e n c e . Edge o f c o b b l e s i s t h e d i s t a n c e west o f 0+00 a t w h i c h c o b b l e s and sand meet. A l l h e i g h t s and d i s t a n c e s a r e i n f e e t .  ro H  Table XVI  RHODOPHYTA A h n f e l t i a c o n c i n n a J.Ag. A h n f e l t i a p l i c a t a (Huds.) F r i e s A n t i t h a m n i o n sp. Bonnemaisonia nootkana (Esp.) S i l v a B o s s i e l l a c o r y m b i f e r a (Manza) S i l v a B o s s i e l l a dichotoma (Manza) S i l v a B o s s i e l l a plumosa (Manza) S i l v a B o s s i e l l a sp. C a l l i a r t h r o n sp. C a l l i t h a m n i o n pikeanum Harv. C a l l o p h y l l i s sp. Ceramium w a s h i n g t o n i e n s e K y i i n Constantinea simplex Setch Constantinea s u b u l i f e r a Setch. C o r a l l i n a o f f i c i n a l i s L. C o r a l l i n a V a n c o u v e r i e n s i s Yendo C o r a l l i n a sp. c o r a l l i n e (crustose) coralline C r y p t o p l e u r a sp. C r y p t o s i p h o n i a w o o d i i J . Ag. Cumagloia a n d e r s o n i i ( F a r l . ) S. & G. Dermatolithon dispar (Fosl.) F o s l . D i l s e a c a l i f o r n i c a ( J . Ag) 0. Kuntze E n d o c l a d i a m u r i c a t a (Harv.) J . Ag. E r y t h r o p h y l l u m d e l e s s e r i o d e s J . Ag. E u t h o r a c r i s t a t a (L.) J . Ag. F a r l o w i a m o l l i s (Harv. & B a i l . ) F a r l Fauchea sp. G i g a r t i n a sp.  Associated P l a n t Species Aats La Indian Arch Bay Push Beach Cape Heights:S L M H L M H L M H L M H *  *  X  X  Cape Short Pesc. F a l c o n Sand P o i n t L M H L M H L M H *  *  X  X  X X  X  X X  * *  X  X  x  x  X  X  X  X  X  X  X X  X  x X  X  X  X X X  X  X  X x  X  X  x  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X X  X  X  X  X  X X  X X  X  *  X  X  X  *  X X  X  X  X  X  X  X X  X  X  X  X  X  X  X  X  X  X  X  X  & Setch.  X X  X  X  X  X  X  X  X  X X  X  X  X X  X  X X  X  X  X  X  X X X  X  Con t i n ued Aats Bay S L M H G l o i o p e l t i s f u r c a t a (p. & R.) J . Ag. Gloiosiphonia v e r t i c i l l a r i s F a r l . G r a c i l a r i a v e r r u c o s a (Huds.) papenf. G r a t e l o u p i a sp. Gymnocfongrus l i n e a r i s (Turn,) J.Ag. H a l o s a c c i o n g l a n d i f o r m e (Gmel.) Rupr. Halymenia c a l i f o r n i c a S m i t h & H o l l e n b . H i l d e n b r a n d i a sp. I r i d a e a sp. Kallymenia sp. L a u r e n c i a s p e c t a b a l i s P. & R. L i t h o t h a m n i o n sp. L i t h o t h r i x a s p e r g i l l u m J . E. Gray M e l o b e s i a sp. Membranoptera sp. M i c r o c l a d i a b o r e a l i s Rupir. M i c r o c l a d i a sp. O d o n t h a l i a k a m t s c h a t i k a (Rupr.) J . Ag. Odonthalia sp. Opuntiella c a l i f o r n i c a (Farl.) K y l i n P e t r o c e l i s f r a n c i s c a n a S & G. P e t r o c e l i s m i d d e n d o r f i i (Rupt.) K j e l l . Peyssonelia p a c i f i c a Kylin Phycodrys s p . Plocamium oregonum Doty Plocamium p a c i f i c u m K y l i n Plocamium v i o l a c e u m F a r l . Plocamium sp. P o l y n e u r a l a t i s s i m a (Harv.) K y l i n P o l y p o r o l i t h o n sp. P o l y s i p h o n i a sp. P o r p h y r a sp„  La Push L M H  Indian Arch Cape Beach L M H L M H  Short Cape F a l c o n Sand L M H L M H  x x  *  x X  X  X  *  X  x x  X  X  X  *  *  X  X X X  X  *  X  X  x  X  x  X  x  X  X  X X X  X  X  X X  X X X X X  *  X  X  X  X  X  X  X  *  X  X  X X X  X  X  *  X X X  X  X  X  X  X  X X  X  X X X  X  * *  X  X  *  X X  *  X  X  X  * *  X  X  X X X  X X  X  X  X  X X X X  X X X  x  X  *  X  X  X  X  X  X  Table oo ro  Continued  Aats Bay Heights: S L M H  H  P o r p h y r e l l a g a r d n e r i Smith & Hollenb. P r i o n i t i s l a n c e o l a t a Harvey Prionitis linearis Kylin p r i o n i t i s l y a l l i i Harv. P r i o n i t i s sp. P t e r o s i p h o n i a b i p i n n a t a (p. & R.) F a l k e n b . Pterosiphonia sp. P t i l o t a a s p l e n o i d e s (Esp.) c Ag. P t i l o t a f i l i c i n a ( F a r l . ) J.Ag. P t i l o t a p e c t i n a t a (Gunn.) K j e l l . P t i l o t a tenuis Kylin P t i l o t a sp. Rhodoglossum s p . Rhodomela l a r i x (Turn.) C. Ag. Rhodymenia palmata (L.) Grev. Rhodymenia s p . PHAEOPHYTA A l a r i a f i s t u l o s a P. & R. A l a r i a m a r g i n a t a p. & R. A l a r i a nana Schrader A l a r i a t e n u i f o l i a Setch. C o s t a r i a c o s t a t a (Burn.) Saund. Cymathere t r i p l i c a t a (p. & R.) J . Ag. C y s t o s e i r a geminata c. Ag. C y s t o s e i r a osmundaceae (Menz.) c. Ag. D e s m a r e s t i a munda S. & G. D e s m a r e s t i a v i r i d i s ( M u l l . ) Lamour. Demarestia sp. Ectocarpus sp. E g r e g i a m e n z i e s i i (Turn.) A r e s c h .  La Push L M H  Indian Arch Beach Cape L M H L M H  Cape S h o r t F a l c o n Sand L M H L M H x  x x x  x  X  x x  X  X X  X  X  X  X  X  X  X  X  X  X X  X  x x x  X  X X  X  X  X X  * *  X  X X  X X  X X  X  *  X X  X  *  X  x x x x x x x* x x  *  X  X  X  X X  X X  X  X  X X  X  X  X  *  X  X X X  T a b l e XVI Cont Aats Bay Heights: S L M x Fucus s p . Haplogloia andersonii (Farl.) Lev. X X Hedophyllum s e s s i l e (C. Ag.) S e t c h . X H e t e r o c h o r d a r i a a b i e t i n a (Rupr.) S. & G. L a m i n a r i a g r o e n l a n d i c a Rosenv. xxx L a m i n a r i a l o n g i p e s Bory X Laminaria s e t c h e l l i i S i l v a L a m i n a r i a s i n c l a i r i i (Harv.) F a r . , And. & E a t . X L e a t h e s i a d i f f o r m i s (L.) A r e s c h . L e s s o n i o p s i s l i t t o r a l i s ( F a r l . & Setch.) R e i n k e Macrocystis i h t e g r i f o l i a Bory Myelophycus i n t e s t i n a l e Saund. X N e r e o c y s t x s l u e t k e a n a (Mert.) P. & R. P e l v e t i o p s i s l i m i t a t a (Setch.) Gard. P h a e o s t r o p h i o n i r r e g u l a r e S. & G. P l e u r o p h y c u s g a r d n e r i S e t c h . & Saund. X P o s t e l s i a p a l m a e f o r m i s Rupr. Punctaria sp. X P y l a i e l l a sp. R a l f s i a sp. Sargassum muticum (Yendo) Fens. X S c y t o s i p h o n l o m e n t a r i a (Lyng.) J . Ag. X S o r a n t h e r a u l v o i d e a p. & R. X X CHLOROPHYTA Cladophora s p . Codium s e t c h e l l i i Gard. Enteromorpha i n t e s t i n a l i s (L.) L i n k Enteromorpha l i n z a (L.) J . Ag. Enteromorpha s p .  d La Push L M H  Indian Arch Beach Cape L M H L M H  x X X  X  x  x  X  x  X  X  X  *  *  X  X  X  X  * x x  X  * X  X X  X  X  *  X  X  X  X  X X X  X  X X  *  X  X  *  X  X  X  * *  X  X  X  X  X  X  X X  Cape S h o r t F a l c o n Sand L M H L M H  X  X  X  X  X  X  T a b l e XVI  in n  Continued Aats Bay L M H  rH  Rhizoclonium Spongomorpha Spongomorpha U l o t h r i x sp. Ulva lactuca Ulva sp.  sp. c o a l i t a (Rupr.) sp.  X  * ab^ove  Cape S h o r t F a l c o n Sand L M H L M H X  X  X  X  *  L.  X X  X  X  X X X  X  X  * * X  X  X  X  X  X  intertidal  X  X  * *  **  s u b t i d a l ; L, lower i n t e r t i d a l ; M, m i d i n t e r t i d a l ; H, h i g h = a s s o c i a t e d w i t h sand.  Pesc. Point L M H  X  X  X  ANTHOPHYTA P h y l l o s p a d i x s c o u l e r i Hook.  #  Indian Arch Beach Cape L M H i L M H  Coll.  CHRYSOPHATA C o l o n i a l diatoms  S  La Push L M H  X  X  Table XVII  Seasonal  D i s t r i b u t i o n of P l a n t S p e c i e s a t I n d i a n Beach  cn  Height:  H  RHODOPHTYA A h n f e l t i a c o n c i n n a J . Ag. A h n f e l t i a p l i c a t a (Huds.) F r i e s Bonnemaisonia nootkana (Esp.) S i l v a B o s s i e l l a dichotoma (Manza) S i v a B o s s i e l l a plumosa (Manza) S i l v a B o s s i e l l a sp. C a l l i t h a m n i o n pikeanum Harv. Constantinea simplex Setch. C o r a l l i n a o f f i c i n a l i s L. C o r a l l i n a V a n c o u v e r i e n s i s Yendo C o r a l l i n a sp. c o r a l l i n e (crustose C r y p t o p l e u r a sp. C r y p t o s i p h o n i a w o o d i i J . Ag. D i l s e a c a l i f o r n i c a ( J . Ag.) 0 Kuntze E n d o c l a d i a m u r i c a t a (Harv.) J . Ag. G i g a r t i n a sp. Gymnogongrus l i n e a r i s (Turn.) J . Ag. H i l d e n b r a n d i a sp. I r i d a e a sp. K a l l y m e n i a sp. L a u r e n c i a s p e c t a b a l i s P. & R. M e l o b e s i a sp. M i c r o c l a d i a b o r e a l i s Rupr. O d o n t h a l i a sp. P e t r o c e l i s f r a n c i s c a n a S. & G. Peyssonelia p a c i f i c a Kylin Plocamium - oregonum Doty Plocamium p a c i f i c u m K y l i n Polocamium v i o l a c e u m F a r l . Plocamium sp.  May67 L M H  Jun67 L M H  * x  *  * x  x  *  Jun 6 6 L M H  X  Apr 6 7 L M H  x X  x  X  X  X  X  X  Jul67 L M H  Aug67 L M H  Sep67 L M H  x  *  X  X  X  x  X  X  X  X X X X  X  X  *  X  X  X  X X  X  X X  X  X  X  X  X  X  X  X  X  X  X  X  X  X X X  X X  X  X  *  X X  X X  X X  X  X  * *  X  X  X  X  *  X  X  X  * X  X  X  X  X  *  X  * *  X  X  X  X  X  X X  X  Table XVII  Cont i n u e d  Jun 6 6 Apr 6 7 May 6 7 Jun67 Height: L M H L M H L M H L M H  H  P o l y s i p h o n i a sp. P o r p h y r a sp. P o r p h y r e l l a g a r d n e r i Smith & H o l l e n b . P r i o n i t i s l y a l l i i Harv. P r i o n i t i s sp. P t e r o s i p h o n i a b i p i n n a t a (p & R.) F a l k e n b . P t e r o s i p h o n i a sp. P t i l o t a tenuis Kylin P t i l o t a sp.  X X  X X  X  X X X X  X X  * X X  X  X  *  X  X  X  X X  X  X  X X X X X X X X X  X  X X X X X  X X X X  X X X X X  X X X X  X X X  X X X  X X X X X  X X  X  Rhodomela l a r i x (Turn.) C. Ag.  X  X X  X  X X  X  c  PHAEOPHYTA A l a r i a m a r g i n a t a P. & R. A l a r i a nana S c h r a d . D e s m a r e s t i a munda S. & G. Desmarestia sp. H a p l o g l o i a a n d e r s o n i i ( F a r l . ) Lev. Hedophyllum s e s s i l e (C. Ag.) S e t c h . Laminaria s e t c h e l l i i S i l v a L a m i n a r i a s i n c l a i r i i (Harv.) F a r l . And. & E a t . L e s s o n i o p s i s l i t t o r a l i s ( F a r l . & Setch.) Reink P h a e o s t r o p h i o n i r r e g u l a r e S. & G. R a l f s i a sp. CHLOROPHYTA Codium s e t c h e l l i i Gard. Enteromorpha l i n z a (L.) J . Ag. Enteromorpha s p . U l v a sp.  X  * X  J u l 6 7 Aug 6 7 L M H L M H  X X  X  x  X X  Buried . . .  *  Buried  X X  Table XVII  Continued  Jun66 Height: L M H  Apr 6 7 May 6 7 Jun 6 7 JU167 L M H L M H L M H L M H  CHRYS OP HYTA C o l o n i a l diatoms  *  * above  *  X  ANTHOPHYTA p h y l l o s p a d i x s c o u l e r i Hook.  x = associated with  X  sand.  * X  *  X  Aug 6 7 Sep67 L M H L M H X  * X  X X  * X  Table X V I I I  Seasonal  D i s t r i b u t i o n o f P l a n t S p e c i e s a t S h o r t Sand Beach  Height: RHODOP HYTA A h n f e l t i a c o n c i n n a j . Ag. A h n f e l t i a p l i c a t a (Huds.) F r i e s B o s s i e l l a c o r y m b i f e r a (Manza) S i l v a B o s s i e l l a plumosa (Manza) S i l v a B o s s i e l l a sp. C a l l i t h a m n i o n pikeanum Harv. Constantinea simplex Setch. C o r a l l i n a V a n c o u v e r i e n s i s Yendo c o r a l l i n e (crustose) coralline C r y p t o p l e u r a sp. C r y p t o s i p h o n i a w o o d i i J . Ag. Cumagloia a n d e r s o n i i ( F a r l . ) S. & G. Dermatolithon dispar (Fosl.) F o s l . D i l s e a c a l i f o r n i c a ( J . Ag.) 0. Kvintze E n d o c l a d i a m u r i c a t a (Harv.) J . Ag. E r v t h r o p h y l l u m d e l e s s e r i o d e s J . Ag. F a r l o w i a m o l l i s (Harv. & B a i l ) F a r l . & S e t c h . G i g a r t i n a sp. Gloisiphonia v e r t i c i l l a r i s F a r l . G r a t e l o u p i a sp. Gymnogonqrus l i n e a r i s (Turn.) j Ag. Halvmenia c a l i f o r n i c a S m i t h & H o l l e n b . H i 1 d e n b r a n d i a sp. I r i d a e a sp. K a l l y m e n i a sp. M i c r o c l a d i a b o r e a l i s Rupr. O d o n t h a l i a sp. P e t r o c e l i s f r a n c i s c a n a S. & G. Plocamium oregonum Doty  Jun. 66 L M H  Apr. 67 L M H  *  * x  x  May 67 L M H  Jul. 67 L M H  *  Aug. 67 L M H * x  x  x x  X X X X  X  X  X X X  X  X X  X  X X  x  X  X X  *  X  X  X  X  X  X  X X  0  0  Jun. 67 L M H  X  X  X  X  X  *  *  X  X  *  X  X X  X  X X X  X  X  X  X X  X  *  X  *  X  X X  X  *  X  X  X  x  X  X  X  X  X  X  X  X  X X  X  X  Table o  I Continued  Jun. 66 Height: L M H  r-i  p o l y n e u r a l a t i s s i m a (Harv.) K y l i n P o l y s i p h o n i a sp. Porphyra sp. p o r p h y r e l l a g a r d n e r i Smith & H o l l e n b . p r i o n i t i s l a n c e o l a t a Harv. prionitis linearis Kylin p r i o n i t i s l y a l l i i Harv. P r i o n i t i s sp. P t e r o s i p h o n i a b i p i n n a t a (p. & R.) F a l k e n b . Pterosiphonia sp. P t i l o t a a s p l e n o i d e s (Esp.) C. Ag. p t i l o t a f i l i c i n a ( E a r l . ) J . Ag. P t i l o t a p e c t i n a t a (Gunn.) . K j e l l . P t i l o t a sp. Rhodomela l a r i x  May 67 L M H  x  x  x X  X  Jun. 67 L M H  Jul. 67 L M H  Aug. 67 L M H  Sep. 67 L M H  X  x  X X  x  X X X X  X  X  *  * *  X  X  X  X  X  X  X  * X X  *  X X  X X  X  X  X  x  x  x  x *  x  (Turn.) C. Ag.  PHAEOPHYTA A l a r i a m a r g i n a t a p. & R. A l a r i a sp. D e s m a r e s t i a munda S. & G. Desmarestia sp. Ectocarpus sp. Fucus s p . Hedophyllum s e s s i l e (C. Ag.) S e t c h . Laminaria s e t c h e l l i i S i l v a L a m i n a r i a s i n c l a i r i i (Harv.) F a r l . And. & E a t . L e a t h e s i a d i f f o r m i s (L.) A r e s c h . L e s s o n i o p s i s l i t t o r a l i s ( F a r l . & Setch.) Reinke P e l v e t i o p s i s l i m i t a t a (Setch.) Gard.  Apr. 67 L M H  X  X  X  X  *  X  X  X  X  X X X X  X  X  X  X  X  *  X  X  X X X  *  X X  X  *  *  X  X  X  X  X  *  X  Table XVIII  Height:  Continued Jun. 66 L M H  P h a e o s t r o p h i o n i r r e g u l a r e S. & G. p i l a y e l l a sp. S o r a n t h e r a u l v o i d e a P. & R. CHLOROPHYTA Cladophora s p . Enteromorpha i n t e s t i n a l i s (L.) L i n k Enteromorpha s p . Rhizoclonium sp. Spongomorpha c o a l i t a (Rupr.) C o l l . U l v a l a c t u c a L. Ulva sp.  Apr. 67 L M H  May 67 L M H  Jun. 67 L M H  Jul. 67 L M H  X X  X  X  X  X  X  X X  * *  X  X  X  X  X  X X X X  X  X  X  X  X  X  * above x = a s s o c i a t e d w i t h sand  Sep. 67 L M H  X  CHRYSOPHYTA C o l o n i a l diatoms ANTHOPHYTA P h y l l o s p a d i x s c o u l e r i Hook.  Aug. 67 L M H  * X  *  X  X  *  X  *  X  * X  *  X  X  Table XIX  Seasonal D i s t r i b u t i o n of A l g a l Height:  RHODOPHYTA A n t i t h a m n i o n sp. B o s s i e l l a sp. C a l l i a r t h r o n sp. C a l l o p h y l l i s sp. Constantinea s u b u l i f e r a Setch. C o r a l l i n a V a n c o u v e r i e n s i s Yendo C o r a l l i n a sp. c o r a l l i n e (crustose) C r y p t o p l e u r a sp. C r y p t o s i p h o n i a w o o d i i J . Ag. D i l s e a c a l i f o r n i c a ( j T A g . ) 0. Kuntze E n d o c l a d i a m u r i c a t a (Harv.) J . Ag. E u t h o r a c r i s t a t a (L.) J . Ag. F a r l o w i a m o l l i s (Harv. & B a i l . ) F a r l . & S e t c h . Fauchea sp. G i g a r t i n a sp. G l o i o p e l t i s f u r c a t a (p. & R.) J„ Ag. G r a t e l o u p i a sp. H a l o s a c c i o n g l a n d i f o r m e (Gmel.) Rupr. I r i d a e a sp. K a l l y m e n i a sp. L i t h o t h a m n i o n sp. L i t h o t h r i x a s p e r g i l l u m J„E. Gray Membr anop t e r a sp. M i c r o c l a d i a sp. O d o n t h a l i a k a m t s c h a t i k a (Rupr.) J . Ag. O d o n a t h a l i a sp. Opuntiella c a l i f o r n i c a (Farl.) K y l i n P e t r o c e l i s m i d d e n d o r f i i (Rupr ) K j e l l . Plocamium sp. Polyporolithon sp.  Spe  es a t A a t s Bay, Jun. 67 S L M H  C o r o n a t i o n Is l a n d  Dec. 65 S L M H x  x  Jun. 66 S L M H  X  x  X  X  X  X X X X  X  Dec. 66 S L M H  X X  X  X X  X  X  X X X  X  X  X  X X X  X  X X  X X X X  X X X X X  X X  x x x  0  X X  X  Jun. 65 S L M H P o l y s i p h o n i a sp. P o r p h y r a sp. P t e r o s i p h o n i a sp. Rhodoglossum s p . Rhodomela l a r i x (Turn.) c. Ag. Rhodvmenia palmata (L.) Grev. Rhodymenia sp. PHAEOPHYTA A l a r i a f i s t u l o s a p. & R. A l a r i a m a r g i n a t a P. & R. A l a r i a t e n u i f o l i a Setch. C o s t a r i a c o s t a t a (Turn.) Saund. Cymathere t r i p l i c a t a (p. & R.) j . A g . C V s t o s e i r a osmundaceae (Menz.) C. Ag. D e s m a r e s t i a v i r i d i s ( M u l l . ) Lamour. D e s m a r e s t i a sp. Fucus sp. Hedophvllum s e s s i l e (C. Ag.) S e t c h . H e t e r o c h o r d a r i a a b i e t i n a (Rupr.) S. & L a m i n a r i a g r o e n l a n d i c a Rosenv. L a m i n a r i a l o n g i p e s Bory L e a t h e s i a d i f f o r m i s (L.) A r e s c h . Mvelophvcus i n t e s t i n a l e Saund. N e r e o c v s t i s l u e t k e a n a (Mert.) p. &.R. P l e u r o p h y c u s g a r d n e r i S e t c h . & Saund. P l a n e t a r i a sp. Sargassum muticum (Yendo) Fens. S c v t o s i n h o n l o m e n t a r i a (Lyngb.) J . Ag. S o r a n t h e r a u l v o i d e a p. & R.  Dec. 65 S L M H  J u n . 66 S L M H X  X  Dec. 66 S L M H  X X  X  X  x x  X  X X  X  X  X X  x  X  X  X X X X  X  X  X X  X X  X  X X  X X  X  X X  X  X  X  X  X  X  X  X  X X  X  X  X X  X X  X X  X  X  X  X X X  X X X X  X  X  Table XIX  Continued Jun. 65 S L M H  CHLOROP HYTA Cladophora sp. Codium s e t c h e l l i i Gard. Enteromorpha s p . Monostroma sp. R h i z o c I o n i u m sp. Spongomorpha s p . Ulothrix Ulva  x  x  Dec. 65 S L M H  Jun. 66 S L M H x  x x X  X X X X X X X X X X  Dec. 66 S L M H  145 T a b l e XX  Summary o f S e a s o n a l C y c l e s on Oregon Beaches  INDIAN BEACH  SHORT SAND BEACH  8/66 Sand h i g h . Many b l a d e s m i s s i n g  Sand h i g h . F r e s h water manyplants  9/66 Sand h i g h e r . Holdfasts buried.  Sand h i g h e r .  ARCH CAPE  touches  10/66 Sand a l l gone. Most p l a n t s r i p e .  Sand lower Many p l a n t s r i p e .  11/66 P l a n t s r i p e .  Sand lower y e t . plants ripe.  12/66 Some p l a n t s r i p e , most l a c k b l a d e s .  Sand a l l gone. A l l blades missing.  1/67  New b l a d e s (1 cm)  present  New b l a d e s (1 cm)  present  2/67  New b l a d e s  (3 cm)  New b l a d e s  (3 cm)  3/67  Small terminal s o r i . Small terminal s o r i .  4/67  Sand coming b a c k . Small terminal s o r i - a l l dropped a t l a s t t i d e . Many new s t i p e s and blades.  Sand coming back. Many new s t i p e s and b l a d e s .  5/67  Sand h i g h e r - n o t yet to rocks bearing plants.  Sand h i g h e r - r o c k s s t i l l exposed.  6/67  Sand a r o u n d r o c k bases.  Sand up, stream flooding  7/67  Sand h i g h e r , has Most p l a n t s b u r i e d , r a i s e d stream l e v e l , plants partly immersed i n f r e s h water Most p l a n t s b u r i e d . Sand down s l i g h t l y . Sand v e r y h i g h A l l blades present. I n s h o r e , many b l a d e s Many b l a d e s missing; farther out, missing, plants are intact.  8/67  9/67  F i r s t plants measured.  Sand c o v e r s a l l rocks, holdfasts, s t i p e s ; only blades protruding.  Sand a l l gone. Most p l a n t s healthy.  Sand unchanged near Sand has r e c e d e d 50 r o c k s . Few b l a d e s cm. P l a n t s appear p r e s e n t e x c e p t on healthy. r o c k t o p s untouched by f r e s h w a t e r .  146  Table XXI  Summaries o f i n s i t u growth measurements on Oregon Beaches.  DATE  NUMBER MEASURED  MEAN STIPE LENGTH (cm)  0 5  28.1 30.4  41.9 46.5  0 0 0  11.7 14.0 17.9  0  18.3  0 6 26 9  *  16.3 19.8 21.8 Buried Buried  17.0 13.2 2.7 1.0 6.1 13.0 18.8 29.9 40.1 42.0 Buried  0 10  35.9 37.8  52.1 66.4  I n d i a n Beach (A Rock) 5 6 7 8 9 10 11 67/ 121 2 3 4 5 6 7 8  5 5 0 0 0 25 7 7 Est. 15 Est. 23 29 26 9 0  MEAN DISTANCE BLADE BETWEEN LENGTH HOLES (cm) (cm)  NUMBER PREVIOUS HOLES  * * *  *  *  * * *  *  *  * * *  -7.4 * * *  -10.8  9.2 4.6  *  I n d i a n Beach (B Rock) (more exposed) '67/ 5 6  20 10  15.7  147  T a b l e XXI DATE  NUMBER MEASURED  Continued  NUMBER MEAN PREVIOUS STIPE HOLES LENGTH (cm)  MEAN BLADE LENGTH (cm)  DISTANCE BETWEEN HOLES (cm)  S h o r t Sand Beach •66/ 8 9 10 11 12 '67/ 1 2 3 4 5 6 7 8 9  2 2 25 22 0 Est. Est. Est. 27 25 24 20 0 0  0 2 0 5 * * * * 0 8 24 19 0 0  •67/ 6 7 8 9  23 8 7 8  0 7 5 6  14.5. 14.5 13.1 12.6  16.8 17.7 19.9 17.6  9,.0 12,.9 15,.1 14.0  3.0 5.0 12.7 20.2 10.6, 30.2 10.8 33.3 3.5 No b l a d e s l e f t i n measurement area  * * * *  * *  *1.0  -  1.5  -  1.1  *  -  -  A r c h Cape  N.B.  17.8 Buried Buried 22.6  26.5 36.1 34.3 36.3  6.0 6.3 1.6  * = No D a t a I n some i n s t a n c e s where number measured i s g r e a t e r t h a n number o f h o l e s , h o l e s were p r e s e n t b u t o b v i o u s l y o l d e r t h a n one month.  148  Table XXII  Growth o f M u l t i - p u n c h e d B l a d e o f L.  sinclairii  i n s i t u a t I n d i a n Beach.  D i s t a n c e (cm) a t time of punching holes 23 May B l a d e base t o A  4.0  '67  D i s t a n c e (cm) a f t e r one Month 20 Jun 8.0  Growth per Month  »67 4.0 (2 cm/2  A to B  2.0  3.5  B to C  2.0  2.5  C to D  2.0  2.2  D to E  2.0  2.1  E to F  2.0  2.0  F to G  2.0  2.0  G to H  2.0  2.0  H to I  2.0  2.0  1.5 0.5 0.2 0.1 0.0 0.0 0.0 0.0  cm)  149  Table  XXIII  Dimensions L.  o f p r e s s e d Specimens o f  l o n g i p e s from  SITE:  33  Alaskan  B L A D E W I D T H (cm) M i n . Max. Mean  Moderately sheltered Chichagof P t . , A t t u I s . T r a p p e r ' s Cove, Adak I s . E a g l e R o c k , NE H b r . , Sanak I s . Gurney Bay, K o d i a k I s . Aats Bay, Coronation I s . TOTALS  S T I P E LENGTH(cm) M i n . Max. Mean  0. 5 1. 5  2. 0 3. 5  1. 2 2. 6  2. 0 11. 0 3. 0 8. 0  7. 2 4. 8  3. 5 0. 5 1. 0  3. 5 4. 5 4. 0  3. 5 1. 2 1. 7  4. 0 7. 0 5. 0 15. 0 4. 0 9. 0  5. 5 6. 7 7. 3  0. 5  4. 5  2. 0  2. 0 15. 0  6. 3  3. 1. 3. 1. 2.  2. 1. 2. 1. 1.  Moderately exposed Casco Bay, A t t u I s . 2. 0 0. 5 Cape A g a g d a k , Adak i s . Ram P t . B e a c h , U n a l a s k a I s . 1. 5 S t a r a y a Bay, Unalaska I s . 1. 0 C a p e S a r i c h e f I , U n a l a s k a I s . 1. 5 Cape S a r i c h e f I I , U n a l a s k a 2. 0 Is. 2. 0 E. A n c h o r Cove, Unimak I s . E a g l e R o c k , NE H b r . , Sanak I s . 1. 0 Nagai I s . 1. 5 0. 5 Paul I s . 1. 5 Chirikof I s . pasagshak P t . , Kodiak I s . 1. 5 2. 0 Cape C h i n i a k , K o d i a k I s . 1. 0 Chiniak I s . P e r i l cape, Afognak I s . 1. 5 2. 5 E n g l i s h Bay 2. 0 Wingham I s . Cape Spencer 1. 0 Helm P t . , C o r o n a t i o n I s . 0. 5 TOTALS  sites.  0.5  0 5 0 0 0  3 0 0 0 6  4. 0 2. 5  3. 0 2. 2  2. 1. 1. 2. 3. 5. 3. 2. 3. 2. 4. 3.  1. 1. 0. 1. 3. 3. 2. 1. 2. 2. 2. 1.  0 5 0 0 5 0 0 0 0 5 0 0  5.0  5 5 7 7 0 3 8 6 8 2 7 4  2.0  3. 7. 1. llo 7.  0 0 0 0 0  12. 20. 10. 11. 15.  0 7. 5 0 11. 1 0 8. 5 0 11. 0 0 11. 2  4. 0 3. 0 5. 0 9 . 0 2 5 . 0 17. 3 5. 7. 6. 8. 2. 5. 8. 2. 9. 4. 6. 4.  0 0 0 0 0 0 0 0 0 0 0 0  1.0  20. 9. 6. 9. 7. 9. 21. 6. 11. 6. 15. 16. 35.0  0 7. 0 0 8. 0 0 6. 0 0 8. 5 0 3. 0 0 8. 1 0 16. 3 0 4. 2 0 10. 0 0 5. 5 0 8. 3 0 8. 4 8.6  150  Table X X I I I  SITE:  Continued  BLADE WIDTH (CM) M i n . Max. Mean  F u l l y exposed Murder P t . , A t t u I s . N o r t h I s . , Adak I s . Zeto P t . , Adak I s . Cape A i a k - L a n c e P t . , Unalaska I s . Raven P t . , Unimak I s . C h i g n i k Bay, Nakchamik I s . Aghiyuk I s . , Semidi i s l a n d s Kayak I s . Cape Ommaney, B a r a n o f I s . TOTALS  EXTREME MINIMA & MAXIMA & TOTAL MEANS:  STIPE LENGTH(CM) M i n . Max. Mean  1.5 2.5 0.5  3.5 3.5 2.0  2.6 2.9 1.1  5. 0 14.0 9.1 6. 0 12.0 7.8 10. 0 15.0 12.7  1.0 2.5 0.5 0.5 2.5 1.0  2.5 3.0 1.0 2.0 3.5 2.0  1.6 2.8 0.7 1.2 2.8 1.5  9. 0 7. 0 4. 0 3. 0 7. 0 5. 0  0.5  3.5  1.9  3.0 15.0  9.1  0.5  5.0  2.0  1.0 35.0  8.4  12.0 10.6 14.0 9.2 5.0 4.5 15.0 9.9 13.0 9.5 11.0 9.0  151  XII:  APPENDIX I  Summary D e s c r i p t i o n s o f F i e l d S t a t i o n s V o l g a I s l a n d , S i t k a , A l a s k a . (57°02.5'N, 135°20.8'W) Rocky r e e f w i t h many l o o s e r o c k s l y i n g i n t i d e p o o l s and surge c h a n n e l s . M o d e r a t e l y t o f u l l y exposed t o s u r f . Mean a n n u a l seawater temperature = 8.5 C; s a l i n i t y = 27.7% Mean a n n u a l a i r t e m p e r a t u r e = 6.3 C; p r e c i p i t a t i o n = 96.57 i n c h e s . N e i t h e r L. s i n c l a i r i i nor L. l o n g i p e s p r e s e n t . S i t e used f o r t r a n s p l a n t s t u d i e s o n l y . O b s e r v a t i o n s made: 1965: June, December 1966: J u l y , December c  A a t s Bay, C o r o n a t i o n I s l a n d , Alaska«, (55°52.7'N, 134°16'W) Rocky r e e f ( a r g i l l i t e ) w i t h many deep surge c h a n n e l s , a d j a c e n t t o beach o f g r a v e l and c o a r s e sand. No d a t a on mean a n n u a l seawater temperature and s a l i n i t y . Mean a n n u a l a i r t e m p e r a t u r e — 6.3°C; P r e c i p i t a t i o n = 76.12 i n c h e s . L. l o n g i p e s p r e s e n t i n abundance„ S i t e used f o r i n s i t u s t u d i e s o f L. l o n g i p e s , t r a n s p l a n t s t u d i e s , and as a s o u r c e f o r a l l L. l o n g i p e s used i n t r a n s p l a n t s and l a b o r a t o r y e x p e r i m e n t s . O b s e r v a t i o n s made: 1965: June, December 1966: June, J u l y , December. R i v e r J o r d a n , Vancouver  I s l a n d , B r i t i s h Columbia.  (48°25.4'N 124 04-W)  Loose r o c k s i n sandy mud bottom. M o d e r a t e l y s h e l t e r e d t o m o d e r a t e l y exposed t o s u r f . No d a t a on mean a n n u a l seawater temperature and s a l i n i t y . Mean a n n u a l a i r t e m p e r a t u r e = 9.7°C; P r e c i p i t a t i o n = 73.42 i n c h e s . L. s i n c l a i r i i p r e s e n t as s m a l l p l a n t s i n v e r y s m a l l q u a n t i t i e s , h i d d e n under E g r e g i a , Hedophyllum, and P h y l l o s p a d i x . S i t e used f o r t r a n s p l a n t s t u d i e s o n l y . O b s e r v a t i o n s made: 1965: August, November 1966: J a n u a r y , March, May, June, August.  152  Sooke Harbour, W. o f W h i f f e n S p i t , Vancouver I s l a n d , Columbia. (48021.2'N, 123°44'W)  British  Loose r o c k s and l a r g e f l a t o u t c r o p s w i t h much sandy mud. A l l p l a n t s f r e q u e n t l y covered w i t h s i l t l a y e r . M o d e r a t e l y s h e l t e r e d from s u r f . No d a t a on mean a n n u a l seawater temperature and s a l i n i t y or a i r temperature o r p r e c i p i t a t i o n . L. s i n c l a i r i i p r e s e n t as s m a l l p l a n t s i n s m a l l q u a n t i t i e s , o n l y o c c a s i o n a l l y f o u n d . S i t e used f o r t r a n s p l a n t s t u d i e s only. O b s e r v a t i o n s made: 1965: J u l y , August, November. 1966: J a n u a r y , F e b r u a r y , March, May, June, August. S t a n l e y P a r k , Vancouver,  B r i t i s h Columbia.  (48°18'N, 123°06'W)  Loose r o c k s , g r a v e l and mud, w i t h s c a t t e r e d l a r g e b o u l d e r s . F u l l y sheltered. No d a t a on mean a n n u a l a i r t e m p e r a t u r e ; p r e c i p i t a t i o n = 61.75 i n c h e s Mean a n n u a l seawater temperature = 9.4°C: s a l i n i t y = 27.6% . N e i t h e r L. s i n c l a i r i i nor L_. l o n g i p e s p r e s e n t . S i t e used f o r t r a n s p l a n t s t u d i e s o n l y . O b s e r v a t i o n s made: 1965: August, November, December 1966: J a n u a r y , F e b r u a r y , March, May, June. 1967: J a n u a r y , May, J u l y . 0  I n d i a n P o i n t , I n d i a n Beach, C l a t s o p County, Oregon. (45°55.9'N, 123°58.8'W) Sandy b e a c h w i t h r o c k y o u t c r o p s ( b a s a l t ) . Sand l e v e l f l u c t u a t e s 1-2 m t h r o u g h y e a r . F u l l y exposed t o heavy s u r f . Mean a n n u a l seawater t e m p e r a t u r e = 10.5 C; s a l i n i t y = approx. 30.97%a. Mean a n n u a l a i r temperature = 11.0 C; P r e c i p i t a t i o n = 79.7 i n c h e s . L. s i n c l a i r i i p r e s e n t i n abundance as dominant p l a n t i n terms of c o v e r . S i t e used f o r i r i s i t u s t u d i e s o f L. s i n c l a i r i i , t r a n s p l a n t s t u d i e s , and as a s o u r c e f o r L. s i n c l a i r i i used i n t r a n s p l a n t s and l a b o r a t o r y e x p e r i m e n t s . O b s e r v a t i o n s made: 1965: F e b r u a r y , J u l y , August, December. 1966: One t o t h r e e times e v e r y month e x c e p t J a n u a r y , March, A p r i l , J u l y . 1967: One t o f i v e times on one l o w t i d e s e r i e s e v e r y month, J a n u a r y through September. 1968: March. Q  153  A r c h cape, C l a t s o p County, Oregon. ( 4 5 4 8 . 2 N , 123°58.2'W) 0  !  Sandy beach w i t h l a r g e r o c k y o u t c r o p s . Sand l e v e l f l u c t u a t e s 1 - 2 m through year. F u l l y exposed t o heavy s u r f . Mean a n n u a l seawater temperature = 11.3 ; s a l i n i t y = 30.97%.. Mean a n n u a l a i r temperature = 11.0°C; P r e c i p i t a t i o n = 81.4 i n c h e s . L. s i n c l a i r i i p r e s e n t i n abundance, b u t o n l y a c c e s s i b l e a t l o w e s t summer t i d e s . S i t e used f o r i n s i t u s t u d i e s o f L. s i n c l a i r i i . O b s e r v a t i o n s made: 1967: May, June, J u l y , August, September. S h o r t Sand Beach, T i l l a m o o k County, Oregon. (45 45.5'N, 123°58'W) 0  Sandy beach w i t h r o c k y o u t c r o p s ( s a n d s t o n e ) . Sand l e v e l f l u c t u a t e s 1 - 2 m t h r o u g h y e a r . F r e s h w a t e r stream f l o w s a c r o s s beach a t l o w t i d e , o f t e n c o v e r i n g some o f p l a n t s studied. F u l l y exposed t o heavy s u r f . Mean a n n u a l seawater temperature = 10.0 C; i n s u f f i c i e n t d a t a on s a l i n i t y . Mean a n n u a l a i r t e m p e r a t u r e = 11.0°C; p r e c i p i t a t i o n = approx. 81.4 i n c h e s . L. s i n c l a i r i i p r e s e n t i n abundance. S i t e used f o r i n s i t u s t u d i e s o f L. s i n c l a i r i i , t r a n s p l a n t e x p e r i m e n t s , and as a s o u r c e f o r L. s i n c l a i r i i used i n t r a n s p l a n t s and l a b o r a t o r y e x p e r i m e n t s . O b s e r v a t i o n s made: 1966: June, A u g u s t , September, October, November, December. 1967: Once o r t w i c e on one l o w t i d e s e r i e s e v e r y month, J a n u a r y t h r o u g h September. 1968: March.  

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