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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., St 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 of Washington, S e a t t l e , Washington, 1963 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n the Department of BOTANY We accept t h i s t h e s i s as conforming to the r e q u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA JUNE, 1969 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r an a d v a n c e d d e g r e e a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e a n d S t u d y . I f u r t h e r a g r e e t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d b y t h e Head o f my D e p a r t m e n t o r b y h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . D e p a r t m e n t o f T e t a n y  The U n i v e r s i t y o f B r i t i s h C o l u m b i a V a n c o u v e r 8, C a n a d a Date June 10, 1969 ABSTRACT Laminaria s i n c l a i r i i (Harvey) Farlow, Anderson and Eaton, found from Southern C a l i f o r n i a to C e n t r a l B r i t i s h Columbia, and L^ longipes Bory, found from Southeast A l a s k a to the 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 other k e l p p l a n t s . Their most d i s t i n c t i v e f e a t u r e i s the r h i z o m e - l i k e h o l d f a s t , composed of many haptera, from which 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 blade. The two species are very s i m i l a r to each other and have been d i s t i n g u i s h e d i n the past p r i m a r i l y by the presence of mucilage ducts i n the s t i p e of L. s i n c l a i r i i and the absence of these i n the s t i p e of L. l o n g i p e s . In order to determine whether the two species are indeed 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 , ecology, growth, and r e p r o d u c t i o n were s t u d i e d i n the 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 gross d i s t r i b u t i o n of both species appears to be c o n t r o l l e d by temperature. Transplants 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. longipes i s adapted to lower temperatures than L. s i n c l a i r i i . S a l i n i t y apparently has 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 both species t o l e r a t e wide ranges of 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 three beaches i n Northern Oregon, where the p l a n t s are subjected to heavy s u r f . The sand l e v e l on the beaches r i s e s through the summer so that the p l a n t s are p a r t l y or wholly b u r i e d under sand by l a t e summer. The f i r s t heavy storms i n the f a l l remove most of the sand. Maximum growth occurs i n e a r l y summer, p r i o r to b u r i a l . The blades are l o s t i n December and new ones are regenerated i n January. Ripe s o r i are produced on the o l d blades j u s t before they are l o s t and on the new blades j u s t a f t e r they appear. There i s l i t t l e evidence from e i t h e r f i e l d or l a b o r a t o r y s t u d i e s to i n d i c a t e t h a t the gametophytes which develop from the spores i n these s o r i normally produce sporophytes. Sexual r e p r o d u c t i o n of t h i s type i s d i f f i c u l t because of the sco u r i n g a c t i o n of the sand. In March and A p r i l there i s co n s i d e r a b l e p r o d u c t i o n of new s t i p e s and blades from the haptera a t the margins of the 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 apparently the normal method of re p r o d u c t i o n . L. longipes was observed i n s i t u i n Al a s k a on only f i v e o c casions. Growth i s g r e a t e s t i n summer and s o r i are produced i n December. Laboratory c u l t u r e s i n d i c a t e t h a t sexual r e p r o d u c t i o n i s very r a r e i n t h i s s p e c i e s . 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. Transplants 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 that p r o d u c t i o n of mucilage ducts i n the s t i p e s of the two species i s not a f f e c t e d by changes i n environmental c o n d i t i o n s . Comparison of the two species 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 l e n g t h of s t i p e s , width of blades, winter l o s s of blades, morphology of gametophytes, and h a b i t a t . The evidence confirms t h a t they should be r e t a i n e d as two separate s p e c i e s . i v TABLE OF CONTENTS PAGE ABSTRACT i i LIST OF TABLES v i i LIST OF FIGURES i x I. INTRODUCTION 1 I I . TERMINOLOGY 7 I I I . GENERAL MATERIALS AND METHODS 8 A. General F i e l d Methods 8 B. C o l l e c t i o n , Transport and Transplanting 8 C. Tagging and Growth Measurements 9 D. Laboratory culture 11 IV. DISTRIBUTION AND AUTEOOLOGY OF LAMINARIA SINCLAIRII 13 A. Geographical D i s t r i b u t i o n 13 B. Habitat and Autocology on Oregon beaches 15 1. Environment 15 2. Occurrence of L. s i n c l a i r i i 31 3. Associated plant species 33 4. Seasonal cycles 37 5. Discussion 40 V. DISTRIBUTION AND AUTE OOLOGY OF LAMINARIA LONGIPES 42 A. Geographical D i s t r i b u t i o n 42 B. Habitat and Autecology at Aats Bay, Coronation Island 43 1. Environment 43 2. Occurrence of L. longipes and associated species 46 3. Growth and reproduction of L. longipes at Aats Bay 47 4. Discussion 48 V PAGE VI. EXPERIMENTAL ECOLOGY 50 A. F i e l d Work 50 1. Transplants 50 a. Alaska beaches 50 b. Oregon beaches 52 c. B r i t i s h Columbia beaches i . Whiffen Spi t 54 i i . River Jordan 55 i i i . Stanley park 56 d. Discussion 57 2. Rock cle a r i n g 59 a. Introduction 59 b. Methods 59 c. Results 59 d. Discussion 62 3. T r a n s i t i o n zone experiments 63 a. Introduction 63 b. Procedures and r e s u l t s 63 c. Discussion 65 B. Laboratory Work 66 1. Sporophytes 66 a. Photoperiod 66 b. S a l i n i t y 67 c. Temperature 67 do Discussion 69 2. Gametophytes 69 a. introduction 69 b. Materials and Methods 70 c. Results 71 do Discussion 72 3. Haptera 73 a. Introduction 73 b. Materials and Methods 74 c. Results 78 i . Temperature 79 i i . Light i n t e n s i t y 80 i i i . i n t i a l Size 80 v i PAGE V I I . TAXONOMY OF L . S I N C L A I R I I AND L . LONGIPES 84 A. I n t r o d u c t i o n 84 B. L . s i n c l a i r i i 85 1. D e s c r i p t i o n 85 2. D i s t r i b u t i o n 86 3. H a b i t a t 86 4. Comments 86 C. L. l o n g i p e s 87 1. D e s c r i p t i o n 88 2. D i s t r i b u t i o n 88 3. H a b i t a t 88 4. Comments 89 D. C o m p a r i s o n o f S p e c i e s 89 V I I I . GENERAL DISCUSSION AND CONCLUSIONS 93 I X . SUMMARY 101 X. BIBLIOGRAPHY 104 X I . TABLES I - X X I I I 111 X I I . A P P E N D I X I . Summary D e s c r i p t i o n o f F i e l d S t a t i o n s 151 v i i LIST OF TABLES TABLE PAGE 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 of L. s i n c l a i r i i and L_. longipes. 112 II 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 III D i s t r i b u t i o n of Laminaria longipes. 116 IV Analysis of sand grain s i z e on Oregon beaches. 118 V Temperature and p r e c i p i t a t i o n at Alaska sta t i o n s . 119 VI Mean temperature on Urup Island. 120 VII Temperature and p r e c i p i t a t i o n at B r i t i s h Columbia stat i o n s . 121 VIII Temperature and p r e c i p i t a t i o n at Oregon sta t i o n s . 123 IX Long term temperature and p r e c i p i t a t i o n at two C a l i f o r n i a stations. 124 X Measured values of seawater temperature and s a l i n i t y on Oregon beaches. 125 XI Calculated mean monthly seawater temperatures on Oregon beaches. 126 XII Mean values of seawater temperature and s a l i n i t y at Arch Cape for 1960-1963. 127 XIII Sand l e v e l at Indian Beach. 128 XXV Sand l e v e l at Short Sand Beach. 129 XV Sand heights of beach at Arch Cape, Oregon. 13 0 XVI Associated plant species. 131 XVII Seasonal d i s t r i b u t i o n of plant species at Indian Beach. 136 XVIII Seasonal d i s t r i b u t i o n of plant species at Short Sand Beach. 139 v i i i TABLE PAGE XIX Seasonal d i s t r i b u t i o n of a l g a l species a t Aats Bay, Coronation I s l a n d . XX Summary of seasonal c y c l e s on Oregon beaches. XXI Summaries of i n s i t u growth measurements on Oregon beaches. XXII Growth of multi-punched blade of L. s i n c l a i r i i i n s i t u a t Indian Beach. 142 145 146 148 X X I I I Dimensions of pressed specimens of L. longipes from 33 Alaskan s i t e s . 149 i x LIST OF FIGURES FIGURE PAGE 1 D i s t r i b u t i o n ranges of Laminaria s i n c l a i r i i and L. l o n g i p e s . 2 2 Mean seawater temperatures over ranges of L. longipes and L. s i n c l a i r i i 4 3 Mean a i r temperatures over ranges of L. longipes and L. s i n c l a i r i i . 5 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 p e r i o d from June 1966 to September 1967. 14 5 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 6 A e r i a l and general views of Indian Beach and Short Sand Beach. 23 7 Indian Beach study area. 24 8 Short Sand Beach study area. 9 V a r i a t i o n i n sand l e v e l s a t Indian Beach i n 1967, 26 27 10 V a r i a t i o n i n sand l e v e l a t study rocks a t Indian 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 Short 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 species at Indian Beach. 34 13 Submergence i n freshwater and a s s o c i a t i o n of L. s i n c l a i r i i w i t h other species a t Short Sand Beach. 36 14 Aats Bay study area. 44 15 Transplant s t a t i o n s . 51 16 Cleared rock at Indian Beach. 61 17 R e s u l t s of growth experiments on haptera of L. s i n c l a i r i i . 77 18 Habit of L. s i n c l a i r i i and L. lo n g i p e s . 91 ACKNOWLEDGEMENTS I wish to express my gratitude and appreciation to Dr. R. F. Scagel for his d i r e c t i o n , advice, and c r i t i c i s m throughout the course of this study. I also wish to expres my appreciate to Dr. G. C. Hughes, Dr. K. Cole, and Dr. G.L. pickard for their advice throughout the study; to Dr. G.H.N. Towers for advice on t h i s manuscript; to Mr. Stephen Borden for programming the computer analyses; to Mr. J . Thorpe for assistance with various kinds of equipment; to Mr. W.A. Markham for topographic surveys of the Oregon beaches; to L t . Col. M.R. Simmonds and Mrs. W.A. Markham for making p r e c i p i t a t i o n data available; to the Oregon State park Department for allowing access to Indian Beach a f t e r hours; to the Department of Botany and I n s t i t u t e of Oceanography, University of B r i t i s h Columbia for f a c i l i t i e s , equipment, and assistance; and f i n a l l y to the many people who ass i s t e d me on various beaches, e s p e c i a l l y at night i n the winter. I wish to acknowledge with thanks the support provided by fellowships from the Univer s i t y of B r i t i s h Columbia and the National Research Council of Canada. 1 I . INTRODUCTION Laminaria Laraouroux, the most common k e l p genus of n o r t h temperate waters, u s u a l l y has a very simple morphology. I t c o n s i s t s of a h o l d f a s t from which 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 lamina, or blade. In three very unusual species the h o l d f a s t , composed of many branched haptera, i s expanded i n t o a r h i z o m e - l i k e organ from which a r i s e many s t i p e s , each be a r i n g a s i n g l e blade. One of these species, L. r o d r i g u e z i i Bornet, occurs a t depths of 100 to 150 m i n the Mediterranean and A d r i a t i c Seas (Bornet, 1888). The other two are found i n the i n t e r t i d a l and s u b t i d a l zones of the n o r t h p a c i f i c Ocean. L_. longipes Bory i s found from the K u r i l e Islands through the A l e u t i a n I s l a n d s and the Gulf of Alaska i n t o Southeast A l a s k a . L. s i n c l a i r i i (Harvey) Farlow, Anderson and Eaton occurs from c e n t r a l B r i t i s h Columbia to Southern C a l i f o r n i a ( F i g . 1 ) . Since no m a t e r i a l of L_. r o d r i g u e z i i was a v a i l a b l e , the two p a c i f i c species o n l y are the s u b j e c t of t h i s study. There are s e v e r a l reasons why a study of L. s i n c l a i r i i and L. longipes seemed warranted. The presence of m u l t i p l e s t i p e s suggested that the growth of these p l a n t s might d i f f e r somewhat from that of other k e l p s . I t had been observed that the b a s a l p o r t i o n s of L. s i n c l a i r i i are o f t e n b u r i e d under sand; as few p l a n t s can withstand such b u r i a l , t h i s suggested t h a t the e c o l o g i c a l adaptations of t h i s p l a n t might be somewhat unusual. F i n a l l y , most taxonomic treatments have separated these two s i m i l a r species by the presence of mucilage ducts i n the s t i p e s b F i g u r e 1 D i s t r i b u t i o n ranges o f L a m i n a r i a S i n c l a i r and L. l o n g i p e s . 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 of L, longipes. Burrows (1964) has shown that presence or absence of mucilage ducts i n the blade of L. saccharina (L.) Lamour. can be c o n t r o l l e d by temperature. In her s t u d i e s , p l a n t s grown a t 10°C developed mucilage ducts, whereas those grown a t 5°C d i d not. This i s of i n t e r e s t because the average temperature of the seawater i n A l a s k a and the K u r i l e Islands i s markedly lower than t h a t of the seawater i n the area from B r i t i s h Columbia to C a l i f o r n i a ( F i g . 2). The d i f f e r e n c e i n a i r temperature i s even greater ( F i g . 3 ) . In view of t h i s , i t appeared t h a t the presence or absence of mucilage ducts i n the two s pecies might be merely a response to the environment and might be a l t e r e d by changing the environment. i f t h i s were the case, the presence or absence of mucilage ducts would not 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 sp e c i e s . 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 species might be one. h.- s i n c l a i r i i and L. longipes have been de s c r i b e d by s e v e r a l authors, most r e c e n t l y by Druehl (1968) as p a r t of a general taxonomic treatment of the genus. However, very l i t t l e e xperimental work has been done w i t h them. S e t c h e l l (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 blade and s t i p e of L. s i n c l a i r i i and Myers (1925) c u l t u r e d the gametophytes of L. s i n c l a i r i i . Apparently no experimental work has ever been done on L. l o n g i p e s . This i n v e s t i g a t i o n c o n s i s t s of three p a r t s . The f i r s t i s an examination of the d i s t r i b u t i o n of the two species throughout ure 2 Mean seawater temperatures over ranges of L. longipes and L. s i n c l a i r i i . Port Hueneme i s i n Ventura County, C a l i f o r n i a , very near the southern l i m i t of d i s t r i b u t i o n of L. s i n c l a i r i i . Sitka i s the nearest point to Coronation Island for which there are long range seawater temperature data. Note lack of overlap i n winter and annual means for the two ranges. 20 18 15 I 4 13 I 2 10 9 8 7 5 4 C O L D E S T M O N T H W A R M E S T M O N T H L. sin c. A N N U A L M E A N Port Huenerne Sitkqi L. long. Port Huenemec L. sin c. Pescadero Pis Urup Port Hueneme L. s i n e . S h o r t Sand B e a c h [ope I s l a n d Sitka ^ ^ H o p Hope Island 1^ Attu IS. L. long. Urup Sitka m Urup Mean S e a w a t e r T e m p e r a t u r e s o v e r R a n g e s of L. Ion g ipe s & L. s i n c l a i r i i gure 3 Mean A i r temperatures over ranges of L_. longipe and L. s i n c l a i r i i . Oxnard i s i n Ventura County, C a l i f o r n i a . B u l l Harbour i s on Hope I s l a n d , a t the n o r t h end of Vancouver I s l a n d , B r i t i s h Columbia. Note l a c k of ove r l a p i n w i n t e r , summer, and annual means f o r the two ranges. 2 0 19 °C -6 u C O L D E S T M O N T H W A R M E S T M O N T H L. sine. A N N U A L M E A N Oxnqrd JUrup Is. M e a n A i r T e m p e r a t u r e s o v e r R a n g e s o f L. l o n g i p e s & L. s i n c l a i r i i 6 t h e i r ranges i n r e l a t i o n to oceanographic c o n d i t i o n s , together 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 of a u t e c o l o g i c a l s t u d i e s on four beaches. L. longipes was s t u d i e d a t Aats Bay, Coronation I s l a n d , A l a s k a during f i v e v i s i t s i n 1965 and 1966. L. s i n c l a i r i i was s t u d i e d during r e g u l a r v i s i t s i n 1965-1967 on three beaches i n Northern Oregon: Indian Beach, Arch Cape, and Short Sand Beach. In the second p a r t , growth and r e p r o d u c t i o n of both species under experimental c o n d i t i o n s are t r e a t e d . Growth of sporophytes was f o l l o w e d i n the f i e l d and i n tanks of seawater i n the l a b o r a t o r y under various c o n d i t i o n s . R e c i p r o c a l t r a n s p l a n t experiments were c a r r i e d out i n an attempt to assess the i n f l u e n c e of environment on the morphology and anatomy of the p l a n t s . Gametophytes were c u l t u r e d i n a study of sexual r e p r o d u c t i o n . The growth of i s o l a t e d pieces of haptera of L. s i n c l a i r i i was a l s o i n v e s t i g a t e d i n l a b o r a t o r y c u l t u r e . In the t h i r d p a r t of the i n v e s t i g a t i o n , the v a l i d i t y of the two species as separate e n t i t i e s i s examined i n l i g h t of the i n f o r m a t i o n obtained from f i e l d observations and l a b o r a t o r y and f i e l d experiments. 7 I I . TERMINOLOGY Each of the two species s t u d i e d has a h o l d f a s t of branched haptera which bears numerous s t i p e s , each s t i p e i n turn b e a r i n g a blade, or lamina. Because of the m u l t i p l e s t i p e s , confusion may a r i s e as to 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 . In t h i s study, the terms " p l a n t " and "clump" are used to r e f e r to the 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 blades. When an i n d i v i d u a l s t i p e or blade i s r e f e r r e d t o , i t i s designated " s t i p e " or "blade". 8 I I I . GENERAL MATERIALS AND METHODS A. General F i e l d Methods The f l u c t u a t i o n i n sand height at c e r t a i n locations on two Oregon beaches (Indian Beach and Short Sand Beach) r e l a t i v e to rocky reference points was measured each month by means of a pocket r u l e and an Abney l e v e l . A d e t a i l e d map of each of the two beaches (Fig. 7, 8) showing the location of a l l rocks with L. s i n c l a i r i i as well as i t s upper and lower l i m i t s , was prepared using a surveyor's t r a n s i t and stadia rod. An IBM 1130 computer was used to calculate the t o t a l time i n any given month that plants at various heights were out of water or under water i n daylight and i n darkness, based on published data for tides and hours of daylight (Fig. 4) B. Collection, Transport, and Transplanting Plants were c o l l e c t e d i n the f i e l d by cutting the holdfasts loose from the rocks with a k n i f e . In most instances, holdfasts bearing more than 40 stipes were separated into two or more parts. The plants were then transported i n wet newspaper on ic e i n a freezer chest to holding tanks at the University of B r i t i s h Columbia or d i r e c t l y , when possible, to another beach i n transplant experiments. The plants were never out of water more than 36 hours and showed no apparent damage from t h i s treatment. In the case of transplants between Alaska and Vancouver the plants were kept i n large holding tanks i n the shade on the deck of the ship en route. The water i n these 9 tanks was changed every two to f i v e days and never reached more than 10°C, due to the low ambient a i r temperature. In the t r a n s p l a n t experiments, 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 areas w i t h l i t t l e s u r f and many loose but s t a b l e boulders (Volga i s l a n d , R i v e r Jordan, Whiffen 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 of p l a c i n g two l a r g e rubber bands around a boulder and the h o l d f a s t . In the second method, employed i n areas w i t h greater s u r f and l a c k i n g loose boulders (Aats Bay, Indian Beach, Short Sand Beach), two or three spikes were d r i v e n i n t o the bedrock and the h o l d f a s t s were attached to these w i t h s e v e r a l rubber bands. C. Tagging and Growth Measurements In the f i e l d and i n l a b o r a t o r y tanks, p l a n t s were i d e n t i f i e d by a p i e c e of 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 of 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 of f l a g g i n g tape t i e d around the s t i p e . For p l a n t s growing i n s i t u , each s t i p e was considered s e p a r a t e l y . 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 A, B, and C were s e l e c t e d f o r growth study on each p l a n t . A was the s h o r t e s t s t i p e (and perhaps the youngest), C was the lon g e s t , and B was a s t i p e of intermediate length, u s u a l l y c l o s e r to C than to A i n le n g t h . In order to determine the growth more a c c u r a t e l y , a sm a l l hole was punched 10 cm above the base of each blade measured, using a cork borer and an apparatus l i k e t h a t used by Sundene 10 (1964), c o n s i s t i n g of a centimeter r u l e w i t h a notched piece of metal fastened to the lower end and a hole a t the 10 cm mark. The notch was pl a c e d over the s t i p e a t the t r a n s i t i o n between s t i p e and blade and the blade was punched through the hole i n the r u l e . A new hole was punched i n the same manner every time a measurement was made and three f i g u r e s were then recorded: l e n g t h o f s t i p e , l e n g t h of blade, and dis t a n c e between the l a s t two holes. In 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 , the width of the blade a t 5 cm above the base was a l s o recorded. However, i n p l a n t s growing i n s i t u t h i s measurement was not made a f t e r i t was observed t h a t a change was r a r e l y 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 areas subjected to heavy s u r f . In 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 hole being the o n l y i n d i c a t i o n that a blade 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 ould not be i d e n t i f i e d as i n d i v i d u a l s but o n l y as members of a p a r t i c u l a r group. How-ever, the d i s t a n c e between punched holes provided an absolute measure of growth of one p a r t of the p l a n t even i f the s t i p e and i t s blade c o u l d not 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 the previous month's data. At the beginning of each s e r i e s of measurements 20 to 25 blades were punched w i t h i n a s m a l l area (ca. 20x20 cm) on a rock and i n succeeding months only those found to have holes were punched and 11 measured again. This continued 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 blades were found, a t which time new blades were punched. In wi n t e r , when no blades are present, a few s t i p e s were tagged to i n d i c a t e a d e f i n i t e area on the rock and then 2 0 to 25 s t i p e s were measured i n t h i s s m a l l area. D. Laboratory C u l t u r e /< For l a b o r a t o r y c u l t u r e work,two New Brunswick psycrotherm Incubator Shakers and three w a l k - i n c o n t r o l l e d temperature rooms were employed. The Psycrotherms were kept at 8°C. Various l i g h t p e riods were employed u s i n g c o o l white 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 of 150 f t - c . Cultures were grown w i t h and without shaking. The three c o n t r o l l e d temperature rooms were maintained a t 5 ° , 8 ° , and 10°C. In each of them two l i g h t i n t e n s i t i e s , approximately 20 f t - c (215.2 lux) and 150 f t - c (1614.0 lux) were employed. In each of the three c o n t r o l l e d temperature rooms three 120 1 tanks of seawater were used f o r c u l t u r i n g sporophytes. Two of the tanks i n each room contained seawater from Juan de Fuca Auto Court near Otter p o i n t on the west coast of Vancouver I s l a n d , designated West Coast water. The s a l i n i t y was 30.9%o t 0„5%„ depending on the month of c o l l e c t i o n . The t h i r d tank i n each room contained water from S t a n l e y park, Bur r a r d I n l e t ( s a l i n i t y = 27.9%o t 0.4%o) designated Stanley park water. The water was changed a t approximately 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 provided by an aquarium bubbler. The tanks were i l l u m i n a t e d by c o o l white f l u o r e s c e n t tubes which gave an i n t e n s i t y a t the surface of the water of 150-200 f t - c (1614-2152 l u x ) . The depth of the water was 35-40 cm. photoperiods ranging from 8 to 16 hours were employed during the course of the study. Gametophytes and p o r t i o n s of haptera 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 dishes, standard 100 mm g l a s s P e t r i dishes and 60 mm p l a s t i c P e t r i dishes. Four d i f f e r e n t seawater media were employed, a l l based on West Coast seawater: raw seawater (SW), seawater f i l t e r e d through cotton wadding (SWF), Erd-Schreiber medium (ES) (F0yn, 1934), and enriched Erd-Schreiber (ES+) i n which 1 ml of "ASP 2" medium ( P r o v a s o l i , McLaughlin, and Droop, 1957) was added to each l i t e r of E r d - S c r e i b e r . In a d d i t i o n , s t r a i g h t ASP 2 was employed i n a few gametophyte c u l t u r e s . One mg/1 of Ge02 was added to each medium to 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 ranging from one day to one month, depending on the experiment. 13 I V . D I S T R I B U T I O N AND AUTECOLOGY OF LAMINARIA S I N C L A I R I I A. G e o g r a p h i c D i s t r i b u t i o n L. s i n c l a i r i i i s f o u n d f r o m Hope i s l a n d , B r i t i s h C o l u m b i a (50 o56'N, 127°58'W) t o V e n t u r a C o u n t y , C a l i f o r n i a (34°19'N, 119°23.3'w) ( F i g . 1). I t h a s n o t b e e n r e c o r d e d b e t w e e n Hope I s l a n d a n d B o x I s l a n d , n e a r T o f i n o o n t h e s o u t h w e s t c o a s t o f V a n c o u v e r I s l a n d . The mean a n n u a l s e a w a t e r s a l i n i t y v a l u e s show a n i n c r e a s e f r o m n o r t h t o s o u t h i n t h e p l a n t ' s r a n g e , w i t h t h e mean a t Hope I s l a n d 31.7% 0 a n d a t p o r t Hueneme, V e n t u r a C o u n t y , 33.6%0 ( T a b l e I ) . The mean a n n u a l s e a w a t e r t e m p e r a t u r e a l s o i n c r e a s e s f r o m n o r t h t o s o u t h , f r o m 8.6°C a t Hope I s l a n d t o 14.3°c a t p o r t Hueneme. Mean F e b r u a r y t e m p e r a t u r e s r a n g e f r o m 7.2°C a t Hope I s l a n d t o 13.2°C a t p o r t Hueneme. Mean A u g u s t t e m p e r a t u r e s r a n g e f r o m 10°C a t Hope i s l a n d t o 16.8°C a t P o r t Hueneme. F i g u r e 2 i l l u s t r a t e s t h e d i f f e r e n c e b e t w e e n t h e s e c o n d i t i o n s a n d t h o s e i n t h e r a n g e o f L. l o n g i p e s . A s many o f t h e p l a n t s a r e o u t o f w a t e r more t h a n t h e y a r e u n d e r w a t e r ( 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 a l s o i m p o r t a n t c o n s i d e r a t i o n s . T a b l e s V I I , V I I I a n d I X show a v e r a g e t e m p e r a t u r e s a n d p r e c i p i t a t i o n f o r s e l e c t e d s i t e s i n B r i t i s h C o l u m b i a , O r e g o n , a n d C a l i f o r n i a . The a n n u a l mean a i r t e m p e r a t u r e s r a n g e f r o m 8.7°C a t Hope I s l a n d t o 15.2°C i n V e n t u r a C o u n t y . F i g u r e 3 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 b e t w e e n t h e s e c o n d i t i o n s a n d those i n t h e r a n g e o f L. l o n g i p e s . 14 Figure 4 Emergence and submergence of vertical distribution extremes of L. s i n c l a i r i i during period from June 1966 to September 1967. Any vertical line drawn from top to bottom of each graph adds up to 100 percent of the hours in that month. "Dark" i s percentage of total hours the point was in darkness. "Light" i s percentage of total hours the point was in daylight. Line between "wet" and "dry" divides time under water from time out of water. 1 9 6 6 1 9 6 7 6 T 8 9 10 II 12 I 2 3 4 5 6 7 6 3 I I I I 1 1 1 I I I I I 1 I • 3 .0 f t (•0.92m) DRY W E T + 1.5 ft. DRY (•0.46 m) W E T 0 . 0 ft. (0.0 nt) DRY W E T DRY -1.5 ft. (-0.46 m) W E T DRY -2 .0 ft. (-0.6lm) yVET L IGHT D A R K Highe s t L. Sinclqirii Shor t S a n d B e a c h H i g h e s t L . s i n c l a i r i i Indian B e a c h Mean T i d e Lowest L. s i n c l a i r i i S h o r t Sand B e a c h Low e s t L. s i n c l a i r i i Indian B e a c h 15 T a b l e I I l i s t s t h e s i t e s a t w h i c h L . s i n c l a i r i i h a s b e e n c o l l e c t e d o r o b s e r v e d . The i n f o r m a t i o n o n d i s t r i b u t i o n was o b t a i n e d f r o m 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 U n i v e r s i t y o f C a l i f o r n i a H e r b a r i u m , B e r k e l e y ; t h e p h y c o l o g i c a l H e r b a r i u m o f t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a ; t h e a u t h o r ' s own o b s e r v a t i o n s a n d c o l l e c t i o n s ; a n d p u b l i s h e d r e c o r d s . L. s i n c l a i r i i h a s b e e n f o u n d a t r e l a t i v e l y f e w p l a c e s i n B r i t i s h C o l u m b i a a n d W a s h i n g t o n , b u t i s much more common i n O r e g o n a n d C a l i f o r n i a , with v e r y f e w e x c e p t i o n s , t h e p l a n t s a r e f o u n d o n b e a c h e s w h i c h a r e f u l l y e x p o s e d t o s u r f a n d h a v e a m a r k e d s e a s o n a l f l u c t u a t i o n i n s a n d l e v e l . T h e y a r e g e n e r a l l y l a r g e r when f o u n d l o w e r i n t h e i n t e r t i d a l z o n e a n d i n more e x p o s e d s i t e s . T h e y a r e a l s o l a r g e r n e a r t h e s o u t h e r n l i m i t s o f d i s t r i b u t i o n . I t was a l s o o b s e r v e d t h a t p l a n t s f r o m C a l i f o r n i a p r o d u c e much more m u c i l a g e t h a n t h o s e f r o m O r e g o n , a l t h o u g h n o 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. B. H a b i t a t a n d A u t e c o l o g y o n O r e g o n B e a c h e s 1. E n v i r o n m e n t The a u t e c o l o g y o f L. s i n c l a i r i i was s t u d i e d o n t h r e e b e a c h e s i n N o r t h e r n O r e g o n : I n d i a n B e a c h , S h o r t S a n d B e a c h , a n d A r c h Cape ( F i g . 5). I n d i a n B e a c h i s l o c a t e d i n E c o l a S t a t e p a r k a n d S h o r t S a n d B e a c h i s i n O s w a l d West S t a t e p a r k . T h u s t h e f o r e s t a n d t h e l a n d i n g e n e r a l a r e r e l a t i v e l y l i t t l e d i s t u r b e d b e h i n d t h e b e a c h e s . To t h e n o r t h o f A r c h Cape t h e r e i s a n a l m o s t c o n t i n u o u s r o w o f h o u s e s a l o n g t h e l a n d b o r d e r i n g t h e b e a c h . S o u t h o f t h e Cape t h e r e a r e v e r y f e w h o u s e s i n 16 the f i r s t k i l o m e t e r , p a r t l y because of the very steep c l i f f s . The f i r s t two beaches were v i s i t e d a t various times over the p e r i o d 1965-1967 and a t l e a s t once a month from August 1966 through September 1967. At Arch Cape L. s i n c l a i r i i p l a n t s are found i n an area which i s very d i f f i c u l t to reach except i n the calmest weather and a t the lowest t i d e s of the summer. The study 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, although i t was observed from a distance on many other occasions. On a l l three beaches there i s a r e g u l a r c y c l e of removal and d e p o s i t i o n of sand (Fig.. 9,10,11,16) . The sand l e v e l on the beach r i s e s from A p r i l or 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 or October) remove most of the sand i n a very s h o r t time, sometimes i n as l i t t l e as 24 hours. The sand forms o f f s h o r e bars during the winter and then i s t r a n s p o r t e d to the beach again the f o l l o w i n g summer. The supply i s c o n t i n u a l l y augmented by sand which i s brought to the sea by nearby streams and r i v e r s . Sand samples f o r each beach were analyzed f o r g r a i n s i z e , u s i n g a standard s e t of Endecott si e v e s (Table I V ) . The r e s u l t s are very s i m i l a r f o r a l l beaches. The sand i s very clean and w e l l s o r t e d . T h i s , and the s i z e d i s t r i b u t i o n r e f l e c t the great wave a c t i o n on the beaches. Over 97 percent of the sand on each beach i s f i n e g r a i n e d (0.25 -0.1 mm diam} to very f i n e g r a i n e d (0.1 - 0.05 mm diam.). Spectroscopic a n a l y s i s shows tha t the sand i s p r i m a r i l y composed of quartz. Most of i t i s d e r i v e d from metamorphic rocks w i t h a smaller percentage of 17 g r a n i t i c m a t e r i a l . The m a i n s o u r c e o f t h e s a n d i s v e r y l i k e l y t h e C o l u m b i a R i v e r (W.R. D a n n e r , D e p t . o f G e o l o g y , U . B . C , P e r s . comm.). S h e p a r d (1963) n o t e d t h a t t h e s u p p l y o f s a n d o n t h e b e a c h i n t h e summer o f t e n v a r i e s d i r e c t l y w i t h t h e amount o f r u n o f f t h e p r e v i o u s w i n t e r . T h i s was o b s e r v e d t o b e t r u e o n t h e O r e g o n b e a c h e s . The w i n t e r o f 1966-1967 h a d much more p r e c i p i t a t i o n t h a n t h e p r e v i o u s w i n t e r ( T a b l e V I I I ) a n d t h e s a n d l e v e l o n t h e b e a c h was n o t i c a b l y h i g h e r i n t h e summer o f 1967 t h a n i n 1 9 6 6 . A l l t h r e e b e a c h e s a r e i n f u l l y e x p o s e d l o c a t i o n s . D u r i n g w i n t e r s t o r m s t h e s u r f i s v e r y h e a v y a n d e v e n o n t h e c a l m e s t summer d a y s t h e r e i s some s u r f . The waves t e n d t o a p p r o a c h f r o m t h e n o r t h w e s t d u r i n g c a l m w e a t h e r b u t f r o m t h e s o u t h w e s t d u r i n g s t o r m s . I n d i a n B e a c h a n d S h o r t S a n d B e a c h h a v e l a r g e r w a v e s t h a n a l l o t h e r b e a c h e s i n n o r t h e r n O r e g o n . E a c h o f t h e two i s a r e l a t i v e l y s h o r t b e a c h f o r m e d i n a n i n d e n t a t i o n o f a p r o m i n e n t h e a d l a n d ( F i g . 5 ) . 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 t h e n o r t h . A r c h Cape i s a h e a d l a n d w i t h a l o n g s t r e t c h o f b e a c h b o t h t o t h e n o r t h a n d t h e s o u t h , s o t h a t i t i s f u l l y e x p o s e d t o w a v e s f r o m a l l d i r e c t i o n s . The s t u d y s i t e a t A r c h Cape i s a r o c k d i r e c t l y w e s t o f t h e Cape. The s t u d y s i t e s o n t h e o t h e r b e a c h e s a r e s i t u a t e d somewhat b a c k o f t h e p o i n t s o f t h e h e a d l a n d s . T h e s e o u t e r m o s t p o i n t s w e r e n e v e r r e a c h e d . A t I n d i a n B e a c h a n d S h o r t S a n d B e a c h , L . s i n c l a i r i i o c c u r s a t e a c h e n d o f t h e b e a c h . The a r e a s o f i n t e n s i v e s t u d y w e r e t h e n o r t h e n d o f I n d i a n B e a c h a n d t h e s o u t h e n d o f S h o r t S a n d 18 F i g u r e 5 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 . "A" and "B" near Arch Cape are s i t e s where p r e c i p i t a t i o n was measured. 124° 0 0 ' W 19 B e a c h . A l t h o u g h a l l t h e s i t e s m u s t b e c o n s i d e r e d f u l l y e x p o s e d , t h e A r c h c a p e s i t e i s s l i g h t l y more e x p o s e d , f o l l o w e d b y t h e I n d i a n B e a c h S i t e a n d t h e n t h e S h o r t S a n d B e a c h s i t e . The t i d e s i n t h i s a r e a a r e o f t h e m i x e d s e m i d i u r n a l t y p e . The t i d a l p a t t e r n i s t h e r e s u l t o f two component t i d e s , t h e s e m i d i u r n a l t i d e , w i t h a n i n t e r v a l b e t w e e n s u c c e s s i v e h i g h w a t e r s o f a b o u t 1 2 % h o u r s a n d t h e d i u r n a l t i d e w i t h a n i n t e r v a l b e t w e e n s u c c e s s i v e h i g h w a t e r s o f a b o u t 25 h o u r s . T h e r e a r e u s u a l l y two h i g h a n d two l o w w a t e r s e a c h d a y , a n d s u c c e s s i v e h i g h s o r l o w s a r e g e n e r a l l y o f d i f f e r e n t h e i g h t s . 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 t h e d a y t i m e i n e a r l y m o r n i n g i n summer a n d a t n i g h t i n w i n t e r . T h e r e a r e no p u b l i s h e d t i d a l d a t a f o r a n y o f t h e b e a c h e s s t u d i e d . The n e a r e s t l o c a t i o n f o r w h i c h t h e r e a r e d a t a i s A s t o r i a , O r e g o n . No i n f o r m a t i o n o t h e r t h a n t h i s was o b t a i n e d f o r A r c h C a p e . F r o m t i d a l d a t a f o r t h e s t u d y p e r i o d ( J u n e 1966 t o S e p t e m b e r 1967) t h e f o l l o w i n g a v e r a g e h e i g h t s w e r e o b t a i n e d : mean h i g h e r h i g h w a t e r (MHHW), 3.91 f t . (1.19m); mean l o w e r h i g h w a t e r (MLHW), 2.37 f t (0.72m); mean h i g h e r l o w w a t e r (MHLW), - 1 . 8 1 f t . (-0.55m); mean l o w e r l o w w a t e r (MLLW), -4.47 f t (-1.36m). A l l h e i g h t s a r e i n r e l a t i o n t o l o n g t e r m mean t i d e t a k e n a s 0.0 f t . Thus t h e a v e r a g e a m p l i t u d e i s a b o u t 8.3 f t . (2.5m). The e x t r e m e a m p l i t u d e i s a p p r o x i m a t e l y l l f t . (3.4m). F o r I n d i a n B e a c h a n d S h o r t S a n d B e a c h t h e a c t u a l t i d e s a p p e a r t o c o r r e s p o n d t o t h e s e c a l c u l a t e d h e i g h t s t o w i t h i n 0.5 f t . 20 This area 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 temperatures. The nearest place f o r which there are p u b l i s h e d records of p r e c i p i t a t i o n and a i r temperature i s Seaside, Oregon.(Pig. 5). 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 kept by two i n d i v i d u a l s a t Arch 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 are s i t u a t e d about one k i l o m e t e r apart, each atop a sea c l i f f about 20 m east of the beach, and each near a house. The more northern one (A) i s l o c a t e d a t a somewhat higher a l t i t u d e and nearer to a house than the southern one (B). The records f o r Seaside and Arch Cape (Table V I I I ) show that the study p e r i o d (1965-67) had below normal p r e c i p i t a t i o n . The 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 Seaside. The maximum p r e c i p i t a t i o n u s u a l l y occurs i n December or January and the minimum i n J u l y or August. The summer of 1967 was p a r t i c u l a r l y dry. The mean t o t a l annual p r e c i p i t a t i o n f o r Seaside i s 79.7 inches. Seaside i s s i t u a t e d n o r t h of Tillamook Head, a l a r g e promontory, and Indian Beach i s i n the southern p a r t of Tillamook Head. Thus, there 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 areas, p a r t i c u l a r l y i n p r e c i p i t a t i o n . However, as no other i n f o r m a t i o n i s a v a i l a b l e from a nearer l o c a t i o n , the Seaside p r e c i p i t a t i o n records are presented as an approximation of the c o n d i t i o n s a t Indian Beach. The records f o r S t a t i o n B a t Arch Cape are presented as an approximation of p r e c i p i t a t i o n a t Arch cape and Short Sand Beach. In the absence of any other records, the Seaside a i r 21 temperatures are used f o r a l l three beaches. The annual mean a i r temperature i s 11.0°c (Table V I I I ) , ranging from a monthly mean of 6.3°C i n January to 15.7°C i n August. The year 1967 d i f f e r s from the mean, e s p e c i a l l y i n the higher summer temperatures. There are p u b l i s h e d records of seawater temperature and s a l i n i t y f o r Arch cape (1960-1963) and the Seaside Aquarium (1949-1967). In a d d i t i o n , water samples were taken a t Indian Beach and Short Sand Beach once a month from October 1966 through September 1967 (Table X ) . The samples were c o l l e c t e d a t low t i d e i n the s u r f as near to the study rocks as p o s s i b l e and temperature readings were taken immediately. The s a l i n i t y determinations were made w i t h an i n d u c t i v e salinometer a t the I n s t i t u t e of Oceanography, U n i v e r s i t y of B r i t i s h Columbia. In the Arch cape records there i s con s i d e r a b l e v a r i a t i o n between the d i f f e r e n t years, probably p a r t l y because of the great v a r i a t i o n i n number of samples. The mean monthly seawater temperature ranged from 7.4°c i n March 1962 to 15.8°C i n J u l y 1963. The mean monthly s a l i n i t y ranged from 26.9%„ i n A p r i l 1962 to 32.9% c i n August 1961. The c o n d i t i o n s f o r 1967 are assumed to be w i t h i n t h i s range. Table X I I gives the monthly means f o r the p e r i o d 1960-1963. The monthly readings obtained a t Short Sand Beach and Indian Beach were compared w i t h the records from Seaside. In almost a l l i n s t a n c e s , when readings f o r s p e c i f i c dates were compared, the Seaside temperature was higher. The mean d i f f e r e n c e was 2.5C° a t Indian Beach and 2.0C° a t Short Sand 22 Beach. These d i f f e r e n c e s were su b t r a c t e d from the monthly means f o r Seaside f o r each month over the p e r i o d s t u d i e d to o b t a i n an approximation of the temperature c o n d i t i o n s a t each beach (Table X I ) . Assuming t h i s to be v a l i d , a t Indian Beach the mean monthly seawater temperature ranged from 7.6°C i n March to 13.9°C i n J u l y , w i t h a y e a r l y mean of 10.5°C, whereas at Short Sand Beach the range was from 7.1°C i n March to 13.4°C i n J u l y , w i t h a y e a r l y mean of 10.0°C. I t i s not known how t y p i c a l t h i s year may have been. Monthly means of seawater temperature f o r Seaside over the p e r i o d of years (1949-1967) f o r which there are records are of no use because the annual means have r i s e n over almost the e n t i r e p e r i o d . This may be a t t r i b u t e d i n p a r t to the steady widening of the Seaside beach due to i n c r e a s e d sand d e p o s i t i o n which has occurred s i n c e the c o n s t r u c t i o n of the Columbia R i v e r j e t t y approximately 25 km n o r t h of Seaside. No i n t e r p o l a t i o n s are p o s s i b l e from a comparison of the s a l i n i t y measurements a t the two study beaches and Seaside. 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 although i r r e g u l a r l y , e s p e c i a l l y a t Short Sand Beach (Table X ) . The study s i t e a t Indian Beach i s an area adjacent to two rocky p o i n t s a t the north end of the beach, one a headland, the other a young stack (Fig 6, 7). There are numerous rocks which extend above, the l e v e l of the beach. Most of the rock i s p r i m a r i l y o l i v i n e b a s a l t . A freshwater stream flows across the beach j u s t south of the study area. I t s course i s a l t e r e d 23 F i g u r e 6 A e r i a l and general views of Indian Beach and Short Sand Beach. a. A e r i a l view of n o r t h end of Indian Beach. Arrow shows approximate l i n e of s i g h t i n f i g u r e b below. Rectangle i n d i c a t e s area shown i n f i g u r e "7, b. Indian Beach a t high t i d e i n August 1967. c. A e r i a l view of south end of Short Sand Beach. Arrow shows approximate l i n e of s i g h t i n f i g u r e d below. Rectangle i n d i c a t e s area shown i n f i g u r e 8. d. Short Sand Beach s h o r t l y a f t e r low t i d e i n March 1967. 24 F i g u r e 7 Indian Beach study area. L. s i n c l a i r i i occurs on a l l rocks 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 rock forming 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 . 16). Sand height was measured a t p o i n t "F" and a t "A". S C A L E 25 w i t h each t i d a l c y c l e , but i t does not normally come i n contact w i t h any of the p l a n t s s t u d i e d . The sand l e v e l was recorded at monthly i n t e r v a l s from August 1966 through September 1967 a t four p o i n t s on the beach (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 probably more on some areas which were not measured. On most of Indian Beach, the sand l e v e l i s lowered i n winter but a sandy beach s t i l l remains. On the area s t u d i e d however, a l l the sand i s removed i n w i n t e r , exposing the bedrock ( F i g . 9a, 10a). Most of the area s t u d i e d a t Indian Beach i s s i t u a t e d so that 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 during most of the time the rocks are exposed a t low t i d e . I t was observed that the rocks and the p l a n t s growing on them always r e t a i n e d enough water so as to appear wet even i f they had been exposed by the recedin g t i d e f o r as long as 3 hours. The only exception to t h i s occurred d u r i n g an extremely dry, hot p e r i o d i n August 1966 when the r e l a t i v e humidity of the a i r f e l l below 20% and the a i r temperature reached 30°C. The rocks appeared completely dry and some of the p l a n t s were c u r l i n g , almost as i f burned. The study s i t e a t Short Sand Beach i s a s e r i e s of rocky outcrops along the narrow margin between the southern end of the sandy beach and the steep c l i f f s which form the southern boundary of the cove i n which the beach i s l o c a t e d ( F i g . 6, 8 ) . The rocks are p r i m a r i l y sandstone, and much s o f t e r than those a t Indian Beach. A freshwater stream flows across the beach p a r a l l e l to and near the outcrops. The sand l e v e l was recorded l i r e 8 Short Sand Beach study area. L. s i n c l a i r i i occurs on a l l rocks shown and at s e v e r a l p o i n t s on the rock forming the shore. P a r t i c u l a r measurements or observations were made on "A" and "B". 27 F i g u r e 9 V a r i a t i o n i n sand l e v e l s a t Indian Beach i n 1967. a. A p r i l 1967 b. May 1967 c. June 1967 d. J u l y 1967 In c and d, the l e f t bucket (arrow) i s on rock A, the r i g h t bucket (arrow) on rock C. The l a r g e rock i s the stack i n d i c a t e d i n f i g u r e 7. 2 8 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 study rocks a t Indian Beach i n 1967. a. A p r i l 1967. Note t h a t s u r f i s around r o c k s , even a t low t i d e . b. May 1967. Sand has advanced but i s not yet around most of the r o c k s . c. June 1967 d. J u l y 1967 e. August 1967. Some of the blades of L_. s i n c l a i r i i have been scoured o f f rock A. f. September 1967. Arrow i n a l l f i g u r e s i n d i c a t e s rock B. In c, d, e, f, bucket i n foreground i s on rock A, bucket i n background i s on rock C (= c l e a r e d rock; see F i g . 16). 29 at monthly i n t e r v a l s from August 1966 through September 1967 at two p o i n t s on the beach (Table XIV). The g r e a t e s t recorded f l u c t u a t i o n s of the sand l e v e l a t the study s i t e s i s 70 cm. The freshwater stream which flows so near the rocks i n t e r a c t s w i t h the sand to 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 Indian Beach. The course of the stream i s s h i f t e d s l i g h t l y during each t i d a l c y c l e . However, most of the time the stream flows next to the ro c k s , a t l e a s t i n the upper p a r t of the beach, s e p a r a t i n g the rocks from the r e s t of the beach. As the sand l e v e l r i s e s i n summer, the l e v e l of the f r e s h water r i s e s next to the rocks, so t h a t by l a t e summer as much as 30 cm of some rocks i s submerged i n f r e s h water a t low t i d e ( F i g . 11). This i n c l u d e s much of the p o r t i o n of the rocks which i s i n h a b i t e d by L. s i n c l a i r i i . The presence o f the stream, and i t s constant e r o s i o n of a channel, i n t u r n prevents the rocks from being b u r i e d so deeply under sand. During the l a t t e r p a r t of the summer of 1967 the l e v e l of the sand on a l l the beach n o r t h of the stream was a t l e a s t 1 m above the l e v e l of the tops of the rocks s t u d i e d ( F i g . l l f ) . The rocks had about 35-45 cm exposed above the sand i n the bottom of the stream channel and about 5-10 cm of t h i s exposed above the freshwater, depending on the height of the rock. In w i n t e r , when sand i s completely absent next to the r o c k s , there i s u s u a l l y a 10-20 cm s t r i p of rock at the bottom, where the stream flo w s , which i s completely bare of p l a n t s . The rock above, which supports L. s i n c l a i r i i and v a r i o u s other algae, i s not touched by the stream water. 30 F i g u r e 11 V a r i a t i o n i n sand and water l e v e l s at Short Sand Beachin 1967. a. L e v e l of sand and freshwater stream i n June 1967. b. L e v e l of sand and freshwater stream i n J u l y 1967. c. L e v e l of sand and freshwater stream i n August 1967. Note t h a t sand i s higher than study r o c k s . In a, b, and c, bucket i n foreground i s on rock B, bucket i n background i s on rock A. d. Study area from above i n A p r i l 1967. Rock A i s out of p i c t u r e . Rocks i n background are completely b u r i e d l a t e r i n summer. e. Study area from above i n J u l y 1967. f. Study area from above i n August 1967. Sand i n background i s higher than study rocks and freshwater stream has r i s e n a l s o . In e and f, l e f t bucket i s on rock A, r i g h t bucket i s on rock B. 31 The study s i t e a t Arch Cape i s a l a r g e b a s a l t i c outcrop d i r e c t l y west of a l a r g e stack west of the Cape. The rock i s about 7 m from east to west, 2 m from north to south and about 1 m i n height from the lowest winter sand l e v e l . I t i s separated from the stack by a s t r e t c h of beach about 20 m wide. Somewhat n o r t h and west of i t i s another, l a r g e r rock. With the exception of t h i s l a r g e r rock, the study rock r e c e i v e s a greater f o r c e of s u r f than any other p o i n t f o r about 5 km along the beach. A freshwater stream flows across the beach about 30 m n o r t h of the rock but does not u s u a l l y come i n t o contact w i t h i t . No measurements of sand f l u c t u a t i o n have been made near the study s i t e a t Arch Cape. Measurements of sand height made at a p o i n t approximately 1 km f a r t h e r n o r t h on the same beach show a maximum f l u c t u a t i o n of o n l y a l i t t l e over 1 m (Table XV). I t i s probably greater near the rock. 2. Occurrence of L. s i n c l a i r i i L. s i n c l a i r i i occurs a t Indian Beach on many rocks a t the n o r t h end of the beach ( F i g . 7 ) . I t i s r e s t r i c t e d to the r e g i o n between the +1.5 and the -2.0 f t . (+0.46 and -0.61m) t i d e l e v e l s on a l l of the r o c k s . The s i z e of the p l a n t s tends to be l a r g e r i n the lower p a r t s of the d i s t r i b u t i o n . At Short Sand Beach, L. s i n c l a i r i i occurs on n e a r l y a l l rocks which border the sandy beach a t the south end ( F i g . 8 ) . I t occurs o n l y between the +3.0 and -1.5 f t . (0.92 and -0.46m) t i d e l e v e l s . At the upper l i m i t s of i t s d i s t r i b u t i o n i t i s 32 very s m a l l . In June 1967, during a -6.1 f t (-1.86m) t i d e when the sand was s t i l l very low, the lower l i m i t s of L. s i n c l a i r i i were determined on rocks f a r out toward the p o i n t . Below the lowest L. s i n c l a i r i i , a patch of about 15 cm of completely bare rock was e v i d e n t . A t Arch Cape, the d i s t r i b u t i o n of L. s i n c l a i r i i was not mapped. I t occurs on s e v e r a l rocks a t the outer p a r t of the Cape. I t covers the e n t i r e surface of the rock 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 not measured. Based on observations during v a r i o u s t i d e s , a l l the p l a n t s appear to be confined to the area between MLLW (-4.5 f t . ) (-1.37 m) and LLLW (ca. -7 f t . ) (-2.1 m). This i s lower than on the other beaches. However, the d i s t r i b u t i o n does not extend 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, there i s a considerable d i f f e r e n c e i n t o t a l time of 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 . t i d e l e v e l are a c t u a l l y out of water f o r a greater percentage of the t o t a l time then they are under water. As the t o t a l time out of water v a r i e s and w i t h i t the t o t a l time which the p l a n t i s exposed to s u n l i g h t w h i l e out of water, there are many d i f f e r e n c e s i n temperature, l i g h t q u a l i t y and i n t e n s i t y , and p o t e n t i a l f o r d e s i c c a t i o n at d i f f e r e n t l e v e l s . Thus i t i s to be expected t h a t growth and s i z e vary a t the d i f f e r e n t l e v e l s . The s i t e s where growth measurements were made are a l l a t d i f f e r e n t h e i g h t s . The measured p l a n t s on rock A at Indian Beach are a t the 0.0 f t . t i d e l e v e l . Those on rock B are at 33 -0.5 f t (-0.15m). The t o t a l time which the p l a n t s on rock A are out of water each month i s about 7 percent greater than t h a t f o r those on rock B. The p l a n t s on rock B have a greater s i z e and greater average monthly growth than those on rock A (Table XXI). The measured p l a n t s a t Short Sand Beach are a t about the +2.0 f t . (0.61m) t i d e l e v e l (ca. MLHW). These p l a n t s are somewhat smaller than those a t Indian Beach. 3. A s s o c i a t e d P l a n t Species L i s t s of a l l the p l a n t species c o l l e c t e d on the three Oregon beaches together w i t h the heights and months of c o l l e c t i o n are presented i n Tables XVI/ XVII/ X V I I I . Only two c o l l e c t i o n s were made at Arch Cape. There are very few species present there. On most of the rocks where L. s i n c l a i r i i grows, no other species are found w i t h the exception of a few p l a n t s o f L. s e t c h e l l i i on the outer and most exposed p a r t s of the r o c k s . A t Indian 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 of cover on a l l the rocks on which i t grows ( F i g . 11, 12). However, s e v e r a l other species are commonly found w i t h i t . Phaeostrophion i r r e g u l a r e occurs a t the same height and s l i g h t l y higher than L. s i n c l a i r i i . S e v e r a l species of B o s s i e l l a are o f t e n found growing w i t h L. s i n c l a i r i i . 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 higher 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 algae are found on the same rocks above and below L. s i n c l a i r i i . A few p l a n t s of L. s e t c h e l l i i are 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 , although L. s e t c h e l l i i tends to occur lower i n 34 Fi 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 species at Indian Beach. a. A s s o c i a t i o n of L. s i n c l a i r i i w i t h other species a t 1.0 f t . below 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 occur 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 higher on rock. June 1967. b. B u r i a l under sand of L. s i n c l a i r i i i n August 1967. Some blades have been eroded away. c. P l a n t s on rock B i n June 1967. Multipunched blade i s v i s i b l e . d. P l a n t s on rock A i n June 1967. Some punched blades are v i s i b l e . Note l e s s complete dominance and l e s s l u x u r i a n t growth of L. s i n c l a i r i i as compared w i t h th a t on rock B which i s lower. 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 of the L_. s i n c l a i r i i p o p u l a t i o n ( F i g . 12a). Gymnogongrus l i n e a r i s and Codium  s e t c h e l l i i occur from the lower l i m i t s of the L_. s i n c l a i r i i p o p u l a t i o n down to about 50 cm below these l i m i t s . Both are u s u a l l y b u r i e d under sand f o r most of the summer. They are apparently w e l l adapted to t h i s , as they appear healthy when the sand again recedes. L. s i n c l a i r i i shows more complete dominance on lower rocks ( F i g . 13). At Short Sand Beach, L. s i n c l a i r i i i s dominant on only a few of the lowest, most exposed rocks on which i t occurs. On most of the rocks where i t occurs there are s e v e r a l other algae i n equal or greater abundance ( F i g . 13 d,c). D i l s e a  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 are very abundant on some of the r o c k s , u s u a l l y s l i g h t l y higher than L. s i n c l a i r i i . 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 asplenoides, 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 l y a l l i i are very common a t the same height as, and s l i g h t l y higher than L. s i n c l a i r i i . On many of these r o c k s , the dominant cover i s composed of 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. Crustose c o r a l l i n e algae occur on the same rocks,above, below, and w i t h L. s i n c l a i r i i . Gymnogongrus l i n e a r i s occurs a t the same height and somewhat lower than L. s i n c l a i r i i . Below the l e v e l of most of 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 of L. s e t c h e l l i i are 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 are o f t e n n e a r l y 36 F i g u r e 13 Submergence i n freshwater and a s s o c i a t i o n of L. s i n c l a i r i i w i t h other species a t Short Sand Beach. a. Some of L. s i n c l a i r i i f o r which growth was measured, hanging i n t o freshwater stream, June 1967. Two punched holes are v i s i b l e . b. Late summer submergence, August 1967. Bucket i s on rock A. P l a n t s on rock A are p a r t l y b u r i e d under sand as w e l l as being submerged i n freshwater. c. B u r i a l under sand i n freshwater stream of L. s i n c l a i r i i , J u l y 1967. d. Some of L. s i n c l a i r i i f o r which growth was measured, showing s m a l l s i z e of clump and r e l a t i v e l a c k of dominance compared w i t h other s p e c i e s , May 1967. e. Small patches of L. s i n c l a i r i i on rock w i t h cover of 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 completely b u r i e d under sand. Most of the other species mentioned are p a r t l y b u r i e d or a t l e a s t coated w i t h sand f o r much of the summer. 4. Seasonal Cycles In the course of a normal year, seasonal c y c l e s of four d i f f e r e n t phenomena occur i n L. s i n c l a i r i i . The p l a n t s grow; they bear r i p e s o r i ; they l o s e t h e i r blades; and they are b u r i e d under sand. The observations on which the f o l l o w i n g account i s based are summarized i n Table XX. In e a r l y January, the p l a n t s are normally completely without b l a d e s . The rocks on which the p l a n t s are attached are completely uncovered and sand may be absent from the e n t i r e area. Growth of the p l a n t s i s very slow. L a t e r i n January the ends of the s t i p e s s p l i t and from the medullary r e g i o n s new blades begin to develop. The blades continue growing and the r a t e of growth a c c e l e r a t e s . The s t i p e s grow, but much more s l o w l y than the blades. Table XXI summarizes the growth measurements of in_ s i t u p l a n t s on the three Oregon beaches. E a r l y i n February when the p l a n t s are only 2-3 cm i n len g t h , f e r t i l e s o r i begin to develop a t the blade t i p s . Ripe s o r i are produced i n March and sometimes as l a t e as A p r i l . At the beginning of t h i s r e p r o d u c t i v e p e r i o d there i s con s i d e r a b l e d i f f e r e n c e i n the time of sorus development and s i z e of 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 synchronized. In A p r i l 1967, a t the beginning of a low t i d e s e r i e s , n e a r l y a l l the blades i n a 38 p o p u l a t i o n on one rock a t Indian Beach had r i p e s o r i on the t e r m i n a l 2 cm of blade. These remained 4 days u n t i l the high t i d e before the l a s t low t i d e which would uncover the p l a n t s f o r more than an hour i n t h i s t i d e s e r i e s . During the l a s t low t i d e i t was observed t h a t a l l the s o r i had been dropped, l e a v i n g a bite-shaped hole a t the end of each blade. Late i n March, and i n A p r i l and May the maximum i n i t i a t i o n of new s t i p e s and blades occurs from the h o l d f a s t s . New 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 c o l o r . During the f i r s t year the s t i p e s are the same l i g h t brown c o l o r as the blades. By the time the s t i p e s are one year o l d and have l o s t t h e i r blades once, they are very dark, almost b l a c k , and very u n l i k e the blades i n c o l o r . The growth of the other s t i p e s and blades 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 begins to b u i l d up, but i s not yet around the rocks on which L. s i n c l a i r i i grows. Sometime i n May or June, the low places are u s u a l l y covered w i t h sand, l e a v i n g the rocks b e a r i n g L. s i n c l a i r i i s t i l l s t i c k i n g out of the sand ( F i g . 10c). In years when the sand l e v e l i s high, the rocks as w e l l as the h o l d f a s t s and s t i p e s of most of the L. s i n c l a i r i i p l a n t s on them are completely b u r i e d by mid-July, l e a v i n g o n l y the blades of L_. s i n c l a i r i i exposed ( F i g . lOd). Such was the case a t Indian Beach and Arch Cape i n 1967. I t would have occurred at Short Sand Beach a l s o , had the stream not kept the sand away from most of the r o c k s . In other years, as i n 1966, the sand d i d not r i s e as high and by September only the h o l d f a s t s of the p l a n t s 39 were b u r i e d . In September or October, the f i r s t heavy storms remove most of the sand, exposing a l l the rocks and p l a n t s again 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 of the blades decreases (Table X X I ) . However, i t was noted that the decrease i n growth i s much l e s s marked i n p l a n t s which are growing lower i n the i n t e r t i d a l zone and i n more exposed areas. In these areas the p e r i o d of maximum growth may be one or two months l a t e r than i n other areas. This can be seen i n comparing r e s u l t s a t Indian Beach w i t h those a t Short Sand Beach, which i s s l i g h t l y l e s s exposed, and a t Arch Cape, which i s more exposed. I t i s a l s o evident i n comparing the two Indian Beach s i t e s , which d i f f e r i n exposure. The growth at Short 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 contact w i t h the freshwater stream which rose and impinged on the p l a n t s more as the sand l e v e l under i t rose. As a consequence o f the greater growth, the p l a n t s g e n e r a l l y have longer s t i p e s and blades i n the more exposed area. As the p l a n t s become b u r i e d , the scouring a c t i o n of the sand i n c r e a s e s . In some areas, many or a l l of the blades may be l o s t , p o s s i b l y due to the s c o u r i n g . However, t h i s v a r i e s w i t h exposure. In the summer of 1967, many blades were l o s t a t Short Sand Beach, fewer at Indian Beach, and none a t Arch Cape. The b u r i a l was more.complete a t Arch cape than a t any of the other study areas. The strong freshwater i n f l u e n c e a t Short Sand Beach may a l s o have c o n t r i b u t e d to the blade l o s s . 40 The p l a n t s grow very l i t t l e from September through December. In October the f i r s t r i p e s o r i of the o l d blades are developed. In November l a r g e areas of the blades are covered w i t h oblong patches of s o r i . Experiments show that these s o r i produce zoospores which develop 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 not u s u a l l y produce sporophytes. Mechanical e r o s i o n of the blade t i p s occurs to some extent a t a l l times. Under normal c o n d i t i o n s growth proceeds f a s t e r than e r o s i o n , and the blade lengthens during the year. Beginning 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 of the blade t i p s begins to occur i n a d d i t i o n to e r o s i o n . These processes continue u n t i l by mid-December most of the blades are completely gone, l e a v i n g o n l y the bare s t i p e s . 5. D i s c u s s i o n L. s i n c l a i r i i was observed 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 ) . At a l l of these beaches the general c o n d i t i o n s are very s i m i l a r to those on the study beaches. The beaches are f u l l y exposed to s u r f and there i s evidence of a marked seasonal f l u c t u a t i o n i n sand l e v e l which causes the p l a n t s to be b u r i e d i n summer. The a s s o c i a t e d p l a n t species are a l s o s i m i l a r to those found on the three study beaches (Table XVI). Thus i t appears t h a t the beaches s e l e c t e d f o r study are t y p i c a l . The p l a n t s of L. s i n c l a i r i i bear r i p e s o r i during two p e r i o d s , once j u s t a f t e r the new blades begin to develop and once j u s t 41 before the o l d blades are l o s t . I t appears evident that sorus i n i t i a t i o n i s not a t a l l dependent upon the age of the blade t i s s u e . The f a c t o r s which induce t h i s i n i t i a t i o n have not been determined. Both periods of sorus production occur during the time when the sand i s a t i t s lowest. This leaves the maximum amount of bare rock a v a i l a b l e f o r the zoospores to a t t a c h and produce gametophytes which then could produce sporophytes. However, no evidence has been found t h a t t h i s a c t u a l l y occurs (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 but Kas not^found i n the s u b t i d a l zone. I t appears t h a t i t may r e q u i r e a s l i g h t amount of d e s i c c a t i o n but cannot t o l e r a t e very much d e s i c c a t i o n . This hypothesis has not been t e s t e d , although a lack of d e s i c c a t i o n may p a r t l y e x p l a i n the poor growth i n l a b o r a t o r y tanks (See pp 66-69) I f the p l a n t s are growing i n the optimum p o s i t i o n w i t h regard to 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 to cause no damage. This 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 to withstand b u r i a l but i s more s e n s i t i v e to 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 of blades and c e s s a t i o n of growth i n l e s s f a vorable s i t e s , may not be d i r e c t l y due to sand b u r i a l at a l l . 42 V. DISTRIBUTION AND AUTECOLOGY OF LAMINARIA LONGIPES A- Geographical D i s t r i b u t i o n In the Western p a c i f i c , the southern l i m i t of d i s t r i b u t i o n of L. longipes i s Urup I s l a n d (46°00'N, 150°00'E) i n the K u r i l e I s l a n d s (Table I I I , F i g . 1 ) . North of t h i s , the p l a n t i s found on South S a k h a l i n , on v a r i o u s other 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 along the east coast of Kamchatka, and on Bering I s l a n d . In the Eastern p a c i f i c , L. longipes i s found on St. P a u l I s l a n d i n the Bering Sea, and from A t t u I s l a n d i n the A l e u t i a n I s l a n d s through the Gulf of Al a s k a to Coronation I s l a n d (55°49.6'N, 134°17'W) i n Southeast A l a s k a , A s u b t i d a l p o p u l a t i o n has been re p o r t e d by Druehl (1968) a t Salmon Bank (48°26'N, 123°01'W) San Juan I s l a n d , Washington. The p l a n t s from Salmon Bank have blades up to 2 0 cm broad, but otherwise f i t the d e s c r i p t i o n of L_. longipes. U n t i l more extensive s u b t i d a l c o l l e c t i o n s are made i n the Northeast p a c i f i c , i t i s not p o s s i b l e to assess the s i g n i f i c a n c e of t h i s c o l l e c t i o n as an extension of the range. The area i n which L. longipes grows i s c h a r a c t e r i z e d by very low winter seawater temperatures (Table I , F i g . 2) ranging from means of 0.5°C at Urup I s l a n d to 4.4°C at S i t k a . Summer temperatures range from a high monthly mean of 9.3°C at pyramid Cove to 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 to 8.5 C a t S i t k a . The a i r temperatures are a l s o very low (Table V, VI, 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 the p l a n t ' s range v a r i e s from a y e a r l y mean of 27.7% 0 a t S i t k a to 32.1%0 a t Pyramid Cove. Table I I I l i s t s the s i t e s at which L.. longipes has been c o l l e c t e d or 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 Russian waters has been obtained from published records. Most of the records of d i s t r i b u t i o n i n American waters have been obtained from c o l l e c t i o n s a v a i l a b l e i n the P h y c o l o g i c a l Herbarium, U n i v e r s i t y of B r i t i s h Columbia. At most of the s i t e s observed by the author, L_. longipes grows on rocky r e e f s i n moderately exposed to moderately s h e l t e r e d areas. No p a r t i c u l a r v a r i a t i o n i n morphology was noted i n p l a n t s from v a r i o u s p a r t s of the range of d i s t r i b u t i o n . B. H a b i t a t and Autecology at Aats Bay, Coronation I s l a n d , A l a s k a 1. Environment The study s i t e a t Aats Bay i s a rocky r e e f adjacent to a beach composed of g r a v e l and coarse sand ( F i g . 14). The r e e f i s composed of a r g i l l i t e rock and i s cut by s e v e r a l deep surge channels. There i s no evidence to i n d i c a t e t h a t the re e f i s ever b u r i e d under sand or that the sand l e v e l changes. The beach i s i n a l o c a t i o n which i s moderately s h e l t e r e d to moderately exposed. Although there i s f r e q u e n t l y s u r f i n the wi n t e r , i n summer the water i s o f t e n completely calm. The t i d e s i n t h i s areas, as i n Oregon, are of the mixed semidiurnal type. The lowest of the low waters occurs i n the daytime i n 44 F i g u r e 14 L o c a t i o n of Helm P o i n t i s d i s t r i b u t i o n of L. Aats Bay study area. the southern l i m i t of l o n g i p e s . 45 summer and a t n i g h t i n w i n t e r . The maximum t i d a l amplitude at Coronation I s l a n d i s 16.8 f t (5.12m) the d i u r n a l amplitude i s 10.7 f t . (3.26m), and the mean t i d e l e v e l i s 8.7 f t . (2.65m) above MLLW (Anon., 1968b). This area 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 low temperatures. There are no pub l i s h e d m e t e o r o l o g i c a l records f o r Coronation I s l a n d . However, Cape D e c i s i o n (56°00'N, 134°08"W) i s only 10 km from Coronation i s l a n d ( F i g . 15) and i t i s assumed th a t the m e t e o r o l o g i c a l c o n d i t i o n s are very s i m i l a r at the two p l a c e s . The records f o r cape D e c i s i o n (Table V) show th a t the annual mean a i r temperature i s 6.3°C, ranging from a monthly mean of 0.9°c i n January to 11.7°C i n August. The mean t o t a l annual p r e c i p i t a t i o n i s 76.12 inches, w i t h the g r e a t e s t amount o c c u r r i n g i n October and the l e a s t i n June. A comparison of these records w i t h those f o r S i t k a (Table V) shows that the average monthly p r e c i p i t a t i o n i s greater a t S i t k a than a t Cape D e c i s i o n f o r every month of the year. A comparison of the temperature records shows a greater annual temperature range at S i t k a than a t Cape D e c i s i o n . This might be expected, 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 land, whereas cape D e c i s i o n i s more su b j e c t to the moderating i n f l u e n c e of surrounding water masses. Maps of the area i n d i c a t e the harbor a t S i t k a r e c e i v e s more freshwater r u n o f f than does the sea around coronation 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 . T h i s , together w i t h the r a i n f a l l , i s important i n c o n s i d e r a t i o n s of s a l i n i t y i n the two areas. 46 There are no p u b l i s h e d records of s a l i n i t y or water temperature at Coronation I s l a n d . In t h i s study these f a c t o r s were measured o n l y once, i n December 1966. At t h i s time the temperature was 4.5°C, 1C° below the reading obtained a t Volga 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 30.78% o, which was 0.5% o below th a t recorded a t Volga i s l a n d . Long term records of the seawater temperature and s a l i n i t y have been p u b l i s h e d f o r S i t k a (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 recorded i n December i s v a l i d f o r the whole year, the seawater temperature range a t Cororation I s l a n d should be from approximately 3.4°C i n February to 13.1°c i n August, w i t h a y e a r l y mean of 7.5°C. These may be f a i r l y c l o s e to the c o r r e c t values, but adequate i n f o r m a t i o n i s l a c k i n g . Because of the greater p r e c i p i t a t i o n and r u n o f f at S i t k a , i t seems l i k e l y t h a t the 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 Corontation I s l a n d . Inasmuch as the one set of recorded observations show the in v e r s e r e l a t i o n s h i p , the average monthly s a l i n i t i e s cannot be estimated from the a v a i l a b l e data. 2. Occurrence of L. longipes and a s s o c i a t e d p l a n t species L. longipes occurs on the rocky r e e f i n the lower i n t e r t i d a l and upper s u b t i d a l zones. Most of the p l a n t s are uncovered by a -10.2 f t . (-3.11m) t i d e . On the s e c t i o n s of the r e e f where i t i s found, i t i s u s u a l l y the dominant p l a n t i n terms of t o t a l cover, although i n a few places 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 higher on the 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 are abundant. A complete l i s t of the species found on t h i s beach i s presented i n Table XIX. 3. Growth and re p r o d u c t i o n of L. longipes at Aats Bay Growth of 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. Holes were punched i n the blades 10 cm above the base i n the usual manner. Based on the holes the average growth of the blades f o r J u l y was 8 cm. The s t i p e s grew a l s o , but much more s l o w l y . I t i s assumed th a t maximum growth occurs i n the summer months but d i r e c t measurements are l a c k i n g . The blade reaches i t s maximum leng t h sometime during the summer and then begins to decrease i n length due to e r o s i o n and d i s i n t e g r a t i o n of the d i s t a l end. This process continues u n t i l the blade reaches a minimum leng t h sometime i n the w i n t e r . Laboratory c u l t u r e s i n d i c a t e t h a t endogenously c o n t r o l l e d d i s i n t e g r a t i o n i s more important than p h y s i c a l abrasion i n t h i s process. In l a b o r a t o r y tanks, the same annual c y c l e occurred i n the absence of any water motion. Druehl (1968) s t a t e s that L. longipes i s "perennial 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 the blade i s l o s t completely down to the s t i p e every year, as i s the case w i t h L_. s i n c l a i r i i . The present study i n d i c a t e s that although the blades are much reduced i n win t e r , they are never completely l o s t . There are three l i n e s of evidence f o r t h i s : (1) In December, when L. s i n c l a i r i i p l a n t s are u s u a l l y t o t a l l y devoid 48 o f b l a d e s , p l a n t s o f L. l o n g i p e s a t A a t s B a y a l l p o s s e s s e d s h o r t b l a d e s 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 a n d i n t r a n s p l a n t s t o O r e g o n L. l o n g i p e s n e v e r l o s t i t s b l a d e s c o m p l e t e l y . (3) I n h e r b a r i u m s p e c i m e n s o f L_. l o n g i p e s p l a n t 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 ) t h e r e i s v e r y o f t e n a r e m n a n t o f t h e p r e v i o u s y e a r ' s b l a d e a t t h e d i s t a l e n d o f t h e c u r r e n t y e a r ' s b l a d e . P l a n t s o f L . l o n g i p e s b e a r r i p e 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 m o n t h s . L a b o r a t o r y c u l t u r e s show t h a t t h e s e s o r i l i b e r a t e s p o r e s w h i c h d e v e l o p i n t o g a m e t o p h y t e s . L i t t l e e v i d e n c e was f o u n d e i t h e r i n t h e f i e l d o r i n t h e l a b o r a t o r y t o i n d i c a t e t h a t t h e s e g a m e t o p h y t e s n o r m a l l y p r o d u c e s p o r o p h y t e s ( See a l s o pp 6 9 - 7 3 ) . 4. D i s c u s s i o n A a t s B a y was v i s i t e d o n l y f i v e t i m e s i n t h e two y e a r s t u d y p e r i o d : t w i c e i n t h e w i n t e r a n d t h r e e t i m e s i n t h e summer ( A p p e n d i x I ) . C o n s e q u e n t l y t h e o b s e r v a t i o n s p r e s e n t e d h e r e a r e somewhat l i m i t e d . I t w o u l d b e v e r y u s e f u l t o h a v e more d e t a i l e d i n f o r m a t i o n o n s e a s o n a l g r o w t h o f t h e p l a n t s , a s w e l l a s s e a s o n a l c h a n g e s i n s e a w a t e r t e m p e r a t u r e a n d s a l i n i t y . A l s o , i t w o u l d b e v a l u a b l e t o know i f t h e r e i s a c h a n g e i n s a n d l e v e l . L . l o n g i p e s h a s b e e n c o l l e c t e d a t s i t e s r a n g i n g i n e x p o s u r e f r o m " m o d e r a t e l y s h e l t e r e d " t o " f u l l y e x p o s e d " ( T a b l e X X I I I ) . E x p o s u r e i s u n d e r s t o o d t o mean t h e e x t e n t t o w h i c h a b e a c h i s s u b j e c t t o s u r f a n d o t h e r w a t e r m o t i o n . T h i s t e r m i n o l o g y i s v e r y d i f f i c u l t t o p u t i n q u a n t i t a t i v e t e r m s . E a c h c o l l e c t o r h a s 49 d e s c r i b e d h i s c o l l e c t i n g s i t e s i n terms of h i s own concept of r e l a t i v e exposure. 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 summer. In some places i n A l a s k a , p a r t i c u l a r l y the A l e u t i a n I s l a n d s , there i s apparently a very great d i f f e r e n c e between summer and winter c o n d i t i o n s . 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 that the c o n d i t i o n s were very a t y p i c a l on t h a t p a r t i c u l a r day, which c o u l d l e a d to an i n c o r r e c t e v a l u a t i o n of exposure. For many s t a t i o n s i n A l a s k a not v i s i t e d by the author, i t i s impossible to s t a t e whether the exposure, as i t has been recorded, i s r e p r e s e n t a t i v e . Most of the " f u l l y exposed" s i t e s i n Al a s k a which the author has v i s i t e d are not exposed to as severe s u r f , or on such a r e g u l a r b a s i s , as any of the s i t e s on the coast of Washington, Oregon, or C a l i f o r n i a . There are undoubtedly places i n A l a s k a which are as exposed as most places on the coast from Washington to C a l i f o r n i a . However, most of these have not been v i s i t e d by c o l l e c t o r s , due to the 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 the a i d of a h e l i c o p t e r i n otherwise i n a c c e s s i b l e p l a c e s , may a l t e r the present p i c t u r e of h a b i t a t d i s t r i b u t i o n of some sp e c i e s . On the b a s i s of present i n f o r m a t i o n , L. longipes i s normally found on beaches which are l e s s exposed than those which L. s i n c l a i r i i i n h a b i t s . 50 V I . EXPERIMENTAL ECOLOGY A. F i e l d Work 1. Transplants In order to i n v e s t i g a t e the growth and s u r v i v a l of L. s i n c l a i r i i and L. longipes i n other n a t u r a l environments, and to determine whether mucilage duct development could be induced or suppressed 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 out. These f a l l i n t o three main c a t e g o r i e s : L_. s i n c l a i r i i from Oregon was t r a n s p l a n t e d to two beaches i n A l a s k a ; L_. longipes was t r a n s p l a n t e d from Alaska to two beaches i n Oregon; and both species were t r a n s p l a n t e d to three beaches i n B r i t i s h Columbia. Fi 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 t r a n s p l a n t s t u d i e s . a. A l a s k a beaches Two beaches i n Alaska were u t i l i z e d f o r t r a n s p l a n t experiments. Aats Bay was chosen because i t was the s i t e of i n s i t u s t u d i e s on L. longipes. Volga I s l a n d , S i t k a (See Appendix I) was chosen because i t was assumed to have a s i m i l a r water temperature to Aats Bay and i s e a s i l y reached from S i t k a . At Volga i s l a n d , p l a n t s were attached to loose rocks which were placed i n t i d e pools where other loose rocks were present. At Aats Bay the p l a n t s were attached to the r e e f on bare spots among the p l a n t s of L_. longipes. In 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 of L. s i n c l a i r i i were placed a t Volga I s l a n d i n June 1965. A l l of these were subsequently l o s t . In the second attempt, f i v e 51 F i g u r e 15 Transplant s t a t i o n s p l a n t s were placed at Volga i s l a n d and four p l a n t s at Aats Bay i n December 1965. A l l of those placed a t Aats Bay were l o s t . Of those placed a t Volga I s l a n d , two were l o s t , one remained but di e d , and one remained a l i v e to J u l y 1966. I t showed some growth i n length as w e l l as production of new s t i p e s . S e ctions of these new s t i p e s d i d not r e v e a l any mucilage ducts. In the t h i r d attempt, i n J u l y 1966, four p l a n t s were placed a t A a t s Bay and four a t Volga i s l a n d . Aats Bay was r e v i s i t e d one month l a t e r . At t h i s time three of the p l a n t s remained. Two showed no change. One had grown 8 cm, the same growth shown by L. longipes p l a n t s marked and measured i n s i t u . In December 1966, two p l a n t s remained a t Coronation I s l a n d , but i n very poor c o n d i t i o n . Two t r a n s p l a n t s were found a t Volga i s l a n d a t t h i s time. Both had attached to the rocks on which they were p l a n t e d and appeared healthy. b. Oregon beaches Indian Beach and Short Sand Beach i n Oregon were u t i l i z e d f o r t r a n s p l a n t s t u d i e s . On both beaches the p l a n t s were attached to bedrock i n areas where L. s i n c l a i r i i was alr e a d y present. In the f i r s t attempt, f i v e p l a n t s of L. longipes were p l a n t e d a t Indian Beach i n August 1965. A l l of these were l o s t during the f o l l o w i n g month. In the second attempt, f i v e more were p l a n t e d a t Indian Beach i n December 1965. Again, a l l were l o s t i n the next 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 Indian Beach i n May 1966. One of these s u r v i v e d without growing to June 1966 and then di e d 53 In August 1966, two p l a n t s of L. longipes were planted a t Indian Beach, and two at Short Sand Beach. In a d d i t i o n , one L. s i n c l a i r i i from Indian Beach was p l a n t e d a t Short Sand Beach, and one from Short sand Beach was p l a n t e d at Indian Beach. One of the L. longipes a t Indian Beach disappeared the f o l l o w i n g month. The other showed s l i g h t growth during September and then was not found again. The L. s i n c l a i r i i from Short Sand Beach grew during September and October, l o s t i t s blades i n December, and disappeared i n January. At Short Sand Beach one L. longipes and the L. s i n c l a i r i i remained through November and were torn loose i n December. The other L. longipes was l o s t i n November. When examined i n September, a l l three had grown, both of the L. longipes s l i g h t l y more than the L_. s i n c l a i r i i . Thereafter no evidence of growth was found and the blades became p r o g r e s s i v e l y shorter u n t i l the p l a n t s were t o r n loose. 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 of L. longipes were placed a t Indian Beach and four a t Short Sand Beach. Those a t Indian Beach were l o s t almost immediately. At Short Sand Beach, three remained through March and then disappeared. Of these, one showed some growth during January and February, then d e t e r i o r a t e d u n t i l i t was l o s t . The others remained unchanged through January, then d e t e r i o r a t e d u n t i l they washed away. i n January, when the L. s i n c l a i r i i p l a n t s had no blades or were j u s t beginning to develop new ones, the L. longipes p l a n t s r e t a i n e d the long blades of the previous year. 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 : Whiffen S p i t and Ri v e r Jordan on the west coast of Vancouver I s l a n d , and Stanl e y Park, Vancouver (see Appendix I ) . The f i r s t two were chosen because L_. s i n c l a i r i i had been found there o c c a s i o n a l l y , although they are i n a much more s h e l t e r e d area than the normal h a b i t a t of 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 park, i s completely s h e l t e r e d and has a lower s a l i n i t y than the open coast h a b i t a t s . I t was chosen i n an attempt to determine why h. s i n c l a i r i i and : L . longipes are not found i n such h a b i t a t s . At the B r i t i s h Columbia s t a t i o n s , e s p e c i a l l y a t Stanl e y park, the range of temperature f o r both a i r and seawater i s greater than i n Oregon or A l a s k a . In most experiments, p l a n t s were placed a t a l l three beaches at n e a r l y the same time. To a v o i d confusion, the r e s u l t s f o r each beach are considered s e p a r a t e l y . i . Whiffen S p i t - At Whiffen S p i t , a l l t r a n s p l a n t s were attached to loose boulders which were then p l a c e d a t the base of the landward s i d e of a l a r g e rock. The rock provided an easy means of f i n d i n g the t r a n s p l a n t s on subsequent v i s i t s . In the f i r s t attempt, four p l a n t s of L. longipes were p l a n t e d at Whiffen S p i t i n J u l y 1965. A l l of these were l o s t before f u r t h e r observations were made. In November, two L. s i n c l a i r i i were p l a n t e d and these disappeared a l s o . In January 1966, four L_. longipes were p l a n t e d . A l l remained i n place and showed a l a r g e blade growth and production of new s t i p e s through March. When the 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 healthy and had grown i n the previous month. One had attached to the rock on which i t was pl a n t e d . This l a t t e r was c o l l e c t e d and sectioned. No evidence of mucilage ducts was found. One a d d i t i o n a l p l a n t of L. longipes was p l a n t e d i n March 1966. I t grew through June and then was l o s t . In March 1966, four L. s i n c l a i r i i were p l a n t e d . One disappeared very soon, one die d i n the f i r s t month, and two showed good growth through May. They s u r v i v e d through June and then were l o s t . The growth of both the L. s i n c l a i r i i t r a n s p l a n t s and the L. longipes t r a n s p l a n t s was very great from March through May r e l a t i v e to that a t other seasons and on other beaches. However, the L. longipes t r a n s p l a n t s showed a s l i g h t l y greater growth. Both species produced wider blades 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 the blades appeared to have developed normally. A l l p l a n t s which s u r v i v e d a t Whiffen S p i t were shaded and almost hidden by l a r g e p l a n t s of Hedophyllum hanging from the v e r t i c a l face of the l a r g e rock above. i i . R i ver Jordan - At River Jordan the t r a n s p l a n t s were attached to loose boulders from various p a r t s of the beach which were then r e - p l a c e d . Most of these boulders were hidden under l a r g e p l a n t s of Egregia 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 which were attached nearby and l a y across l a r g e adjacent areas. In November 1965, two L. s i n c l a i r i i were planted. These were not found again u n t i l March 1966. At t h i s time the blades were shorter or absent and there was no evidence t h a t growth had occurred. When examined again i n May 56 there were s e v e r a l new s t i p e s and considerable blade growth had occurred. One of the p l a n t s had attached to the rock. In March 1966, two other _L. s i n c l a i r i i were planted. When examined i n May these showed about the same blade growth as t r a n s p l a n t s set out i n November and s e v e r a l new s t i p e s were present. In June one of the p l a n t s had attached to the rock, the other was dead. In January 1966, two p l a n t s of L_. longipes were t r a n s p l a n t e d to R i v e r Jordan. These remained and grew w e l l through June 1966. As a t Whiffen S p i t , both species produced wider blades a f t e r being t r a n s p l a n t e d but otherwise appeared healthy and "normal". A l l p l a n t s at River Jordan which 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 fronds of Egregia  menziesii l y i n g on top of them. i i i S tanley park - The t r a n s p l a n t s at Stanley park were attached to loose boulders which were then p l a c e d near one l a r g e rock f o r ease i n f i n d i n g on subsequent v i s i t s . L. saccharina occurs above and below the area used f o r t r a n s p l a n t i n g . In the f i r s t t r a n s p l a n t experiment, two L. s i n c l a i r i i were p l a n t e d i n November 1965. One of these disappeared. The other s u r v i v e d to June 1966. I t continued growing the whole time according to measurements between punched holes. I t attached to the rock on which i t was planted. The blades were never completely l o s t . They decreased i n l e n g t h to March and then began i n c r e a s i n g i n length. The second experiment w i t h L_. s i n c l a i r i i was s t a r t e d i n March 1966 when s i x p l a n t s were t r a n s p l a n t e d . Of these, one d i e d , one grew 57 through May, and four grew through June. Of the l a t t e r group, one had attached to the rock by May. F i v e p l a n t s of L. longipes were t r a n s p l a n t e d to Stanley Park i n August 1965. A l l of these d i e d very soon. Two more L. longipes were t r a n s p l a n t e d i n December 1965. They grew f a i r l y w e l l through May and then d i e d i n June. In winter the L. longipes appeared to be h e a l t h i e r than the L. s i n c l a i r i i , but as summer approched, the L_. s i n c l a i r i i appeared h e a l t h i e r . Both species produced blades co n s i d e r a b l y broader than i n t h e i r normal h a b i t a t s . They became very t w i s t e d and overgrown w i t h bryozoans, diatoms, and various other e p i p h y t i c algae. d. D i s c u s s i o n I t was observed that no mucilage ducts were present 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 Volga I s l a n d . This would seem a t f i r s t to confirm the o r i g i n a l idea that mucilage duct formation r e q u i r e s a temperature higher than t h a t normally encountered i n Southeast A l a s k a . However, the new s t i p e s which were s e c t i o n e d were very s m a l l . 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 taken d i r e c t l y from the f i e l d i n Oregon were sec t i o n e d . These d i d not show any mucilage ducts e i t h e r . I t i s concluded t h a t the s t i p e s of 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 before mucilage ducts are produced and t h i s s i z e was not a t t a i n e d by any of the new s t i p e s produced i n A l a s k a . The Alaskan s i t e s , w i t h one exception a t Aats 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 of the p l a n t s 58 were l o s t before any growth:measurement could be made. i t i s q u i t e p o s s i b l e t h a t i f the Alaskan s i t e s had been v i s i t e d as o f t e n as the others a comparable number of growth measurements might have been obtained before l o s s of the p l a n t s . Attachment of the h o l d f a s t to the rocks on which the p l a n t s had been placed occurred o n l y a t Stanley park (three L. s i n c l a i r i i ) R iver Jordan (two L. s i n c l a i r i i ) ; Whiffen S p i t (two L_. s i n c l a i r i i , one L. longipes) ; and Volga I s l a n d (one L. s i n c l a i r i i ) . None of these places i s very exposed, which i s probably an important f a c t o r . More important, however, i s the method of p l a n t i n g employed. On a l l of these s i t e s , loose boulders are present and the p l a n t s were f i x e d to these by means of rubber bands. This method holds the bottom of the h o l d f a s t f i r m l y a g a i n s t the rock. In the more exposed areas, which l a c k loose boulders, the method of f a s t e n i n g the p l a n t s to spikes i n the rock o n l y held the o u t p l a n t s from washing away but d i d not hold them i n any one p o s i t i o n f o r any length of time. Several unsuccessful attempts were made to devise 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 the t r a n s p l a n t s to Stanley park, Whiffen S p i t , and Ri v e r Jordan i n d i c a t e that L. s i n c l a i r i i i s l e s s a d v e r s e l y a f f e c t e d by high summer seawater temperatures than i s L. lo n g i p e s . The r e s u l t s from Oregon and Alaska are not as c l e a r c ut. However the summer temperatures are not as high at these s i t e s as i n Oregon or A l a s k a , e i t h e r . 59 2. Rock c l e a r i n g a. I n t r o d u c t i o n At v a r i o u s times over a p e r i o d of one year, p o r t i o n s of rocks were c l e a r e d a t Indian Beach and Short Sand Beach. The o b j e c t of these experiments was to determine whether the adjacent 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 the c l e a r e d areas. b. Methods In a l l experiments, p o r t i o n s of the 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 area of rock thus exposed was then scraped and f i n a l l y burned to w i t h i n 5 cm of the remaining p l a n t s . Thereafter the 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. R e s u l t s In the 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 of L. s i n c l a i r i i a t Indian Beach i n May 1966. In June a s l i g h t growth of haptera t i p s toward the c l e a r i n g from both s i d e s was evident. In August some haptera had grown 2 cm onto the c l e a r e d area. By October the c l e a r e d area had been completely crossed by haptera from both s i d e s and some s t i p e s and blades had been produced from these haptera. The second c l e a r i n g at Indian Beach was made i n December 1966. In February, there was no evidence of growth of haptera nor was there anything e l s e growing on the c l e a r e d area. In March, two p l a n t s of Hedophyllum s e s s i l e each 15 cm long, one p l a n t of A l a r i a (marginata ?) 12 cm long and s e v e r a l crustose c o r a l l i n e algae were present. 60 The t h i r d c l e a r i n g experiment a t Indian 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 the sand on one s i d e of a rock, up over the top and down to w i t h i n 10 cm of the sand on the other s i d e ( F i g . 16) . S i x species were* removed from the rock. L_j_ s i n c l a i r i i covered most of the area which was c l e a r e d , but there were o c c a s i o n a l gaps i n the h o l d f a s t which provided attachment space f o r other s p e c i e s . On top of the rock Phaeostrophion i r r e q u l a r e was the second most abundant spec i e s . There were a l s o s e v e r a l p l a n t s of B o s s i e l l a plumosa and an u n i d e n t i f i e d crustose c o r a l l i n e a l g a on top. On both sides Hildenbrandia sp. and B o s s i e l l a plumosa were found. On one s i d e , extending down to below the sand l e v e l , was an extensive mat of Codium s e t c h e l l i i . In May, the haptera at the edges of the L. s i n c l a i r i i had s t a r t e d growing toward the c l e a r e d area. S e v e r a l patches of Hildenbrandia and a few t u f t s of B o s s i e l l a were evident ( F i g . 16a). For the f o l l o w i n g three months the rock was completely b u r i e d under sand ( F i g . 16 b,c,d). In September the sand receded, exposing the rock. At t h i s time the L. s i n c l a i r i i and Codium s e t c h e l l i i appeared healthy and unchanged from the June c o n d i t i o n . Otherwise, only some dying t u f t s of B o s s i e l l a plumosa were noted. At Short Sand Beach, the 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 haptera of L. s i n c l a i r i i had encroached onto the c l e a r e d area and were p u t t i n g up s t i p e s and blades i n the area. Several t u f t s of c o r a l l i n e algae were a l s o present. 61 F i g u r e 16 Cleared rock a t Indian Beach a. Rock C i n May 1967, one month a f t e r s t r i p 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. 62 The second experiment at Short Sand Beach was begun i n A p r i l 1967. A v e r t i c a l s t r i p was c l e a r e d next to the p l a n t s f o r which growth was being measured i n s i t u . The rock 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 holes produced by v a r i o u s animals. The surface was c l e a r e d i n the usual manner, but the holes were not 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 holes i n the rock. By J u l y s e v e r a l e n c r u s t i n g animals had e s t a b l i s h e d themselves on the rock but no p l a n t s were present. d. D i s c u s s i o n S e v e r a l of the c l e a r i n g experiments were c a r r i e d out a t times 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 s o r i . Under such circumstances i t seems l i k e l y t hat gametophytes of L. s i n c l a i r i i should have e s t a b l i s h e d themselves on the c l e a r e d areas. Although the gametophytes would not have been v i s i b l e they should have produced sporophytes which would have been e a s i l y recognized. However, on a l l the c l e a r e d areas and s e v e r a l other rocks observed on these beaches over a p e r i o d of two years, no young sporophytes of L. s i n c l a i r i i were ever observed which had d e f i n i t e l y a r i s e n from gametophytes. In every i n s t a n c e , c a r e f u l examination showed that the young sporophytes had a r i s e n from haptera of older p l a n t s and thus were outgrowths of the o l d e r p l a n t s . As noted p r e v i o u s l y , two Hedophyllum 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 rock although there were r e l a t i v e l y few of 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 roduction of L. s i n c l a i r i i sporophytes by sexual means i s r a r e . 63 On the other hand, the experiments show that c o l o n i z a t i o n of new areas by outgrowth of haptera of L. s i n c l a i r i i takes place r e a d i l y , a t l e a s t during the p e r i o d when the p l a n t s are not b u r i e d under sand. 3. T r a n s i t i o n zone experiments a. I n t r o d u c t i o n In most members of 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 the s t i p e and blade i s g e n e r a l l y s t a t e d to be the p r i n c i p a l . r e g i o n of growth. I t appeared that t h i s might not be the case i n L. s i n c l a i r i i . A c c o r d i n g l y , two types of experiments were conducted i n the f i e l d to determine the importance o f the t r a n s i t i o n zone. The f i r s t of these c o n s i s t e d of c u t t i n g o f f s t i p e s w e l l below the t r a n s i t i o n zone and observing the behavior of the remaining h o l d f a s t s and s t i p e stubs. This was done twice a t Indian Beach and once a t Short Sand Beach. The second experiment c o n s i s t e d of marking 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 observing where the g r e a t e s t growth occurred. This was done once a t Indian Beach. b. procedures and r e s u l t s In the f i r s t experiments i n which t r a n s i t i o n zones were removed, s t i p e s were cut a t both Indian Beach and Short Sand Beach i n October 1966. A t Indian Beach a l l the s t i p e s i n a l a r g e clump on top of one rock were cut o f f a t l e a s t 5 cm below the t r a n s i t i o n zone, using a l a r g e p a i r of shears. At Short Sand Beach the same procedure was f o l l o w e d except that the 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 of rock around i t . 6 4 In November, the s t i p e stubs on both beaches appeared to be h e a l i n g over. In a d d i t i o n , a t Short Sand Beach there was considerable growth of haptera a t the margins of the clump. Late i n December many of the cut s t i p e s on both beaches had produced new blades from the t i p s i n an apparently normal manner. At t h i s time, a few of the normal uncut s t i p e s a t Indian Beach had blades w i t h r i p e s o r i , but the m a j o r i t y had no blades a t a l l . A t Short Sand Beach a l l the uncut s t i p e s were without blades. In January no f u r t h e r changes i n the cut s t i p e s were noted. In February the blades were s l i g h t l y longer than before and t h e r e a f t e r no f u r t h e r changes were noted a t e i t h e r beach. In February 1967, the second c u t t i n g experiment was s t a r t e d a t Indian Beach. This time a l l the l a r g e s t i p e s i n one group were cut o f f w e l l below the t r a n s i t i o n zone, but a l l the present year's s t i p e s and blades were l e f t untouched. (The s t i p e s of the present year are e a s i l y recognized, as p r e v i o u s l y noted.) A f t e r one month the cut s t i p e s showed no change. The uncut s m a l l blades had grown about 3 cm past the cut ends w i t h a t o t a l blade l e n g t h of 6-7 cm. By the end of A p r i l about 20 percent of the cut s t i p e s had produced new blades and the r e s t were unchanged. In June no more cut s t i p e s had produced blades, but the growth of haptera a t the margins of the clump seemed to be greater than normal. An experiment to determine the r e g i o n of g r e a t e s t growth was s t a r t e d i n May 1967 a t Indian Beach. In a clump of s t i p e s where s e v e r a l s t i p e s and blades were being measured 65 p e r i o d i c a l l y , four blades were punched w i t h many holes. In two of these, the holes were punched every 2 cm, beginning 4 cm above the base of the blade. In the other two, the holes were punched every 2 cm, beginning a t 2 cm above the base. Both s t i p e s and blades were measured, but no convenient way was found to mark o f f i n t e r v a l s on the s t i p e s . In June 1967, o n l y one of the four p l a n t s was found, one of those which had been punched from 4 cm above the base ( F i g . 12c). The blade was eroded a t the end, undoubtedly weakened by the holes, and the t o t a l l e n g t h was l e s s than a t the beginning. The t o t a l growth of the blade as c a l c u l a t e d from the t o t a l growth between a l l the holes was a t l e a s t 6.3 cm (Table X X I I ) . The s t i p e i n c r e a s e d 1 cm i n the same p e r i o d . 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 growth i s l e s s than 50 percent of the average f o r other blades i n the same place a t the same time (Table XXI), which i s probably due to the i n j u r i o u s e f f e c t s of the l a r g e number of h o l e s . C. D i s c u s s i o n S e t c h e l l (1905), studying growth and regeneration i n L. s i n c l a i r i i , noted t h a t a blade was produced from a wound on the s i d e o f a s t i p e . 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 L. s i n c l a i r i i could regenerate from the stump of a s t i p e . The present experiments show that t h i s i s c o r r e c t . The r e s u l t s show t h a t although 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 not l i m i t e d to t h i s zone and the zone i s not necessary f o r the production of new blades. 66 Under normal c o n d i t i o n s the blades are l o s t i n December and regenerated i n January. On the s t i p e s which were cut i n October, new blades appeared i n December, about one month e a r l i e r than normal. However, a f t e r t h i s e a r l y beginning the blades showed l i t t l e f u r t h e r development, a t l e a s t i n t h a t year. Removal of the t r a n s i t i o n zone a l s o seemed to s t i m u l a t e the growth of haptera. I t i s p o s s i b l e that these r e s u l t s i n d i c a t e the presence of an auxin or other substance i n the o l d blade which i n h i b i t s new blade formation and growth of haptera. However, such a p o s s i b i l i t y has not been i n v e s t i g a t e d , and at present no adequate e x p l a n a t i o n of these phenomena i s a v a i l a b l e . B. Laboratory Work 1. Sporophytes Sporophytes of L. s i n c l a i r i i and L. longipes were c u l t u r e d f o r v a r y i n g periods of time i n the 120 iT'tanks of seawater p r e v i o u s l y d e s c r i b e d . The c u l t u r e s were planned to determine the e f f e c t s of d i f f e r e n t temperatures, s a l i n i t i e s , and photoperiod on the growth and re p r o d u c t i o n of 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 intr o d u c e d i n t o the tanks s e v e r a l times during a two-year p e r i o d , p l a n t s were removed whenever they showed no change f o r a p e r i o d of three months or were o b v i o u s l y d e t e r i o r a t i n g , a. photoperiod P l a n t s of both species were grown a t photoperiods 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 dark, p l a n t s were grown i n each photoperiod f o r up to three months under each temperature regime (5°, 8°, and 10°C). The photoperiods were u s u a l l y s i x months out of phase w i t h those found i n nature, i . e . , short days i n summer. The va r i o u s photoperiods d i d not 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 or rep r o d u c t i o n of p l a n t s of the two s p e c i e s . A f t e r c o n c l u s i o n of the photoperiod experiments a l l c u l t u r e s were run a t 16 hours l i g h t / 8 hours dark. b. S a l i n i t y , The Stanley park seawater was used as low s a l i n i t y water and the West Coast seawater was used as high s a l i n i t y water, p l a n t s were grown i n each type of water at 5°, 8° and 10°C. No d i f f e r e n c e was evident i n growth of the p l a n t s i n the d i f f e r e n t s a l i n i t i e s a t the same temperature. c. Temperature Various p l a n t s were grown a t 5°, 8°, and 10°C over a p e r i o d of two years. During the f i r s t nine months the photoperiod was changed every three months. Thereafter i t was maintained at 16 h o u r s l i g h t / B h o u r s dark. Maximum growth of both species occurred somewhat e a r l i e r i n the year a t 5°C than at 8° or 10°C. The maximum growth of 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 of L. longipes occurred i n January a t 5°C and i n February a t 8° and 10°C. With these exceptions, the d i f f e r e n t temperatures appeared to have very l i t t l e e f f e c t on the growth of the p l a n t s . 68 In a l l p l a n t s c u l t u r e d i n the tanks, c e r t a i n common featu r e s were evident. No v a r i a t i o n s i n temperature, s a l i n i t y , or photoperiod had any e f f e c t on production of s o r i or l o s s and r e g e n e r a t i o n of blades. 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 c u l t u r e . However, other p l a n t s of the same p o p u l a t i o n became r i p e a t the same time i n nature. Blade l o s s always occurred a t the same time as i n nature unless the l o s s was due to the p l a n t ' s dying. L i k e w i s e , blade r e g e n e r a t i o n occurred a t the same time as i n nature a t a l l temperatures, although the r a t e of growth of the new blades v a r i e d w i t h temperature. A f t e r new blades were produced, the " c o l l a r s " ( S e t c h e l l , 1896) formed by the f r a y e d ends of the s p l i t s t i p e ends were much more obvious than i n the f i e l d , presumably because there was no wave a c t i o n to remove the excess t i s s u e . G e n e r a l l y , the p l a n t s which grew i n the l a b o r a t o r y tanks grew w e l l and showed normal behaviour f o r two to four months and then ceased showing blade growth. At t h i s time some of them then began to d e t e r i o r a t e ; but i n most, growth of haptera and prod u c t i o n of new s t i p e s and blades from the h o l d f a s t s continued f o r up to 15 months a f t e r blade growth had ceased. Many of these haptera were removed and used i n the experiments on haptera d e s c r i b e d below. Although new s t i p e s were produced from the haptera, no growth was recorded on any of the s t i p e s which were a l r e a d y present on p l a n t s when brought i n t o the laboratory. 69 d. D i s c u s s i o n The tank r e s u l t s are valuable i n that they i n d i c a t e t h a t production of s o r i and blade l o s s and regeneration are not c o n t r o l l e d by temperature, s a l i n i t y , or photoperiod. However, the r e s u l t s i n the tanks cannot g i v e a very good p i c t u r e of the growth of the p l a n t s . The p l a n t s apparently behaved normally f o r a short time a f t e r being placed i n the tanks. However, p l a n t s were not pl a c e d i n the tanks o f t e n enough to gi v e a true p i c t u r e of growth over a whole year. The tanks contained water very s i m i l a r to th a t i n which the p l a n t s normally grow, but t h i s water was only changed once a month. This allowed a much greater b u i l d u p of waste products than occurred i n nature and probably a considerable d e p l e t i o n of v a r i o u s n u t r i e n t s . T h i s , and the l a c k of t i d e s and wave a c t i o n , make the tanks very poor s u b s t i t u t e s f o r the n a t u r a l c o n d i t i o n s . 2. Gametophytes a. I n t r o d u c t i o n At the beginning of t h i s study i t appeared that L. s i n c l a i r i i and L. longipes were p o s s i b l y ecotypes of the same s p e c i e s . In order to t e s t t h i s hypothesis, i t was planned to cross the two s p e c i e s . To accomplish t h i s , i t was necessary 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 (1925). In her c u l t u r e s , the gametophytes produced sporophytes, thus i n d i c a t i n g a t y p i c a l k e l p l i f e h i s t o r y . However, she noted that 70 the gametophytes continued to 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 of the sporophytes produced i n c u l t u r e . So f a r as i s known, the gametophytes of 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 Sporophytes bear 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 during the winters of 1965, 1966 and 1967 (L. longipes was only c o l l e c t e d i n December) and brought i n t o the l a b o r a t o r y . They were then kept i n the dark i n a moist p l a s t i c bag a t 5°, 8°, or 10° C f o r four to 36 hours, then i n a paper towel f o r two to e i g h t hours a t the same temperature. A f t e r t h i s p e r i o d of 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 to remove e p i p h y t i c contaminants and then cut i n t o s m a l l pieces w i t h a r a z o r blade. These pieces were placed i n SWF or ES i n 250 ml g l a s s c u l t u r e dishes or standard 100 mm P e t r i d i s hes. The dishes were placed i n 5°, 8°, or 10°C under 20 f t - c or 150 f t - c . A f t e r i n t e r v a l s which rangedfrom s i x hours to s i x days i n v a r i o u s experiments (apparently depending upon the stage of development of the sorus a t the time of c o l l e c t i o n ) zoospores could be seen swarming from the pieces of sorus. S e v e r a l m i l l i l i t e r s of the spore suspension were p i p e t t e d i n t o each c u l t u r e d i s h . A l a r g e number of d i f f e r e n t combinations of c u l t u r e c o n d i t i o n s were t e s t e d . F i v e d i f f e r e n t media were employed a t v a r i o u s times: SW, SWF, ES, ES+, and ASP 2. Large g l a s s c u l t u r e dishes (250 ml), standard g l a s s P e t r i dishes (100mm) and s m a l l (60mm) P e t r i dishes of both g l a s s and p l a s t i c were used. In v a r i o u s 71 experiments the medium was changed a t i n t e r v a l s of 1, 2, 4, 7, 10,' 14, or 28 days. Most c u l t u r e s were kept at l e a s t one year. A f t e r a c u l t u r e had run f o r two to three months the medium was u s u a l l y changed onl y once a month. The c u l t u r e s were maintained a t 5°, 8°, 10°, and 15°C under l i g h t i n t e n s i t i e s of 20 f t - c or 150 f t - c provided by c o o l white f l u o r e s c e n t l i g h t s . Three photoperiods 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 hours l i g h t / 8 hours dark. In one experiment gametophyes of various ages were maintained f o r periods of one day to two weeks i n t o t a l darkness. Cultures were a l s o kept 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 run on the shaker i n the psycrotherm i n c u b a t o r . In 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 subjected to changes of temperature, media, frequency of media change, l i g h t i n t e n s i t y , photoperiod and various combinations of these f a c t o r s . I t was not p o s s i b l e to c a r r y out any c r o s s i n g experiments, "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 sporophytes were produced by L. s i n c l a i r i i gametophytes, but never more than a few i n any one c u l t u r e . L. longipes gametophytes onl y produced sporophytes twice: once i n 1966 a f t e r 10 weeks i n ES a t 10°C under 20 f t - c ( f i v e sporophytes) and once i n 1967 a f t e r 24 weeks i n ES+ a t 5°C under 150 f t - c (three sporophytes). 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 to have l i t t l e e f f e c t on the production of sporophytes. In the ES+ medium the gametophytes had l a r g e r , more deeply pigmented c e l l s than i n the other media. Longer and l e s s deeply pigmented 72 c e l l s were produced under higher l i g h t i n t e n s i t i e s . At 5°c the gametophytes had fewer and smaller c e l l s . However, none of these d i f f e r e n c e s appeared to a f f e c t the production of oogonia, a n t h e r i d i a , and u l t i m a t e l y , sporophytes. Under most c o n d i t i o n s the gametophytes continued growing v e g e t a t i v e l y and appeared healthy a f t e r more than one year i n c u l t u r e . In most ins t a n c e s the male and female gametophytes of 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 of k e l p gametophytes, the males have s m a l l e r , more numerous c e l l s and many branches, whereas the females c o n s i s t of a few l a r g e r c e l l s , and few branches. In s e v e r a l i n s t a n c e s , oogonia and a n t h e r i d i a were noted. In L. l o n g i p e s , however, a d i f f e r e n t s i t u a t i o n p r e v a i l s . The gametophytes are apparently a l l of one morphological type and are t y p i c a l of n e i t h e r male nor female k e l p gametophytes. The c e l l s are of intermediate s i z e and the branches, although numerous, are not as numerous as i n male gametophytes of L. s i n c l a i r i i . No a n t h e r i d i a or oogonia were ever noted. In L. longipes i t was thus impossible to d i s t i n g u i s h males from females. d. D i s c u s s i o n I t i s p o s s i b l e that the gametophytes of L. longipes are b i s e x u a l but u n t i l a n t h e r i d i a and oogonia arefound, t h i s w i l l remain i n question. With the exception of Chorda  tomentosa (Sundene, 1963), the gametophytes of a l l k e l p s r e p o r t e d are u n i s e x u a l . The c r o s s i n g experiments were not c a r r i e d out 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 of 73 11- longipes c o u l d not be d i s t i n g u i s h e d from each other and thus males and females could not be i s o l a t e d . Secondly, no c o n d i t i o n s were found under which e i t h e r species would produce sporophytes r e g u l a r l y , which would have served as a c o n t r o l . In previous experiments the author has c u l t u r e d gametophytes of Laminaria saccharina (L.) Lamour., Hedophyllum  s e s s i l e (C. Ag.) Setch, and Ne r e o c y s t i s l l u e t k e a n a (Mert.) P. and R. under the same c o n d i t i o n s of temperature, l i g h t , and media employed i n the present study. In many instances abundant sporophytes were produced. Robinson (1967) c u l t u r e d s e v e r a l species of A l a r i a under the same c o n d i t i o n s i n the same c u l t u r e rooms a t the same time t h a t L. s i n c l a i r i i and L. longipes were being c u l t u r e d . One species ( A l a r i a marginata p. and R.) was obtained on the same Oregon beaches as L. s i n c l a i r i i . In most of Robinson 1s c u l t u r e s sporophytes were produced abundantly. The present study i n d i c a t e s t h a t , r e l a t i v e to other members of the L a m i n a r i a l e s , the sexual production of sporophytes by gametophytes of L. s i n c l a i r i i and L. longipes i s very l i m i t e d . 3. Haptera a. I n t r o d u c t i o n F i e l d and l a b o r a t o r y observations show that there i s v e g e t a t i v e production of new blades and s t i p e s from the haptera a t the edges of the h o l d f a s t s of 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 using l a b o r a t o r y c u l t u r e s of i s o l a t e d pieces of haptera (Markham, 1968). 74 b. M a t e r i a l s and Methods Sporophytes of L. s i n c l a i r i i growing i n the la r g e c u l t u r e tanks provided a source of haptera f o r use i n the experiments. A f t e r two to three months i n the tanks, many haptera had grown out from the bottoms and edges of the 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 haptera were removed from the parent p l a n t s , washed i n f i l t e r e d seawater, and cut to a uniform l e n g t h of 20 mm. In the experiments t e s t i n g the e f f e c t of i n i t i a l s i z e , pieces 2.5, 5, 10, and 15 mm long were used. For a l l s i z e s the haptera e i t h e r had one growing t i p or were cut o f f a t both ends; i n the l a t t e r i n s t a n c e , the most d i s t a l end was a t l e a s t 20 mm back of the o r i g i n a l apex of the hapteron. The pieces w i t h one cut end and one growing t i p were designated "E haptera", whereas those w i t h both ends cut o f f were designated "1J haptera". P r e l i m i n a r y experiments were run to t e s t the e f f e c t s of var 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 of these experiments were run a t 8° and 10°C and 150 f t - c . Four media were employed: ES+, ES, SWF, and SW. Two types of c u l t u r e dishes were t e s t e d : 250 ml g l a s s c u l t u r e dishes c o n t a i n i n g 200 ml of media and 20 mm p l a s t i c P e t r i dishes c o n t a i n i n g 15 ml of media. The media were changed a t 2-week i n t e r v a l s . These experiments showed that the maximum percentage of haptera produced outgrowths i n 250 ml dishes c o n t a i n i n g 200 ml of ES+. A f t e r the p r e l i m i n a r y r e s u l t s were obtained, experiments were run to t e s t the e f f e c t s of shaking and sand sco u r i n g . In 75 these experiments 20 mm pieces of haptera were secured to small rocks w i t h rubber bands around the middle p o r t i o n s of the haptera. These rocks were then p l a c e d i n 250 ml c u l t u r e dishes. G e n e r a l l y the rocks were l a r g e enough so that o n l y one could f i t i n each c u l t u r e d i s h . For the shaking experiments, the dishes were then f i l l e d w i t h j u s t enough ES+ to cover the rock and i t s attached haptera. These were then p l a c e d i n the Psycrotherm incubator shakers and the speed of shaking adjusted so t h a t the medium reached the r i m of the c u l t u r e dishes but d i d not s p i l l over. For the scouring experiments s t e r i l i z e d beach sand from Oregon was added to the c u l t u r e dishes u n t i l i t covered the rocks but not the haptera on top of the r o c k s . The dishes were then f i l l e d w i t h ES+ u n t i l the haptera were j u s t covered and the dishes were p l a c e d on the shakers w i t h those i n the shaking experiments. The water motion caused the sand to wash back and f o r t h over the haptera. Two pieces of haptera of L. longipes were a l s o c u l t u r e d on the shaker, each on a rock without sand. In the incubator shakers the temperature was 8°C, and the l i g h t i n t e n s i t y was 150 f t - c , w i t h a photoperiod of 16 hours l i g h t / 8 hours dark. C o n t r o l experiments were run i n a second Psycrotherm incubator without shaking. The medium was changed every two weeks. A 3-cm pi e c e of s t i p e was cut out of each of two p l a n t s of L. s i n c l a i r i i , halfway between the h o l d f a s t and blade (approximately 10 cm of s t i p e was cut o f f each end). These pieces were then fastened to rocks and placed i n c u l t u r e 76 dishes i n the same manner as the haptera. One d i s h contained sand and ES+, the other o n l y ES+. These were c u l t u r e d without shaking i n the c o n t r o l psycrotherm. A l l haptera were c u l t u r e d f o r 14 weeks and s t i p e s f o r 20 weeks. These experiments were p r e l i m i n a r y i n that they were simply designed to determine whether i t was p o s s i b l e f o r haptera to produce blades and s t i p e s a t a l l under such c o n d i t i o n s . For t h i s reason r e s u l t s were recorded o n l y a t the end of the experiments and then o n l y as presence or abs_ence of blades. The f i n a l s e r i e s of experiments t e s t e d the e f f e c t s of temperature, 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 of haptera. A l l these c u l t u r e experiments were c a r r i e d out i n 250 ml g l a s s c u l t u r e dishes c o n t a i n i n g 200 ml of ES+ medium. The medium was changed every two weeks except i n the experiments t e s t i n g very s m a l l s i z e s . In the l a t t e r i n s t a n c e i t was changed every four weeks. Experiments on the e f f e c t of temperature were run a t 5°, 8°, 10°, 15°, and 20°C. A l l other experiments were run o n l y a t 8° and/or 10°C. L i g h t i n t e n s i t i e s of 2 f t - c (21.5 l u x ) , 150 f t - c (1614 l u x ) , 250 f t - c (2690 l u x ) , and 500 f t - c (5380 lux) were employed i n l i g h t i n t e n s i t y experiments. A l l other experiments were run under 150 f t - c onl y . The photoperiod f o r a l l experiments was 16 hours l i g h t / 8 hours dark. A l l these experiments were run f o r ten weeks. The r e s u l t i n g data were s o r t e d w i t h an IBM 113 0 computer. In the f i n a l a n a l y s i s , a l l 20 mm haptera were analyzed f o r t h e i r responses to l i g h t and temperature ( F i g . 17a) whereas the e f f e c t s of i n i t i a l s i z e were analyzed 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 of growth experiments on haptera of L. s i n c l a i r i i a. percentage of E and N haptera producing blades, haptera, and growth i n len g t h a t various temperatures and various l i g h t i n t e n s i t i e s . b. 2.5 mm pieces of N. haptera, 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 blades. c. 20 mm pieces of E and N haptera, 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 blades and one l a t e r a l hapteron. d. 20 mm pieces of E and N haptera, 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 blades and l a t e r a l haptera. e. Nonmedian s e c t i o n of the s t i p e of a blade a r i s i n g as a l a t e r a l outgrowth from a hapteron, x 70 (Diameter of hapteron i s 1 mm). f. An enlarged p o r t i o n of e, x 210. A l l r e s u l t s a t 10 weeks, b, blade; co, cortex, h, o r i g i n a l p i ece of hapteron; l h , l a t e r a l hapteron a r i s i n g from o r i g i n a l hapteron; me, meristoderm; s t , s t i p e . 78 c. R e s u l t s The r e s u l t s of the c u l t u r e experiments were v a r i e d . The haptera e i t h e r showed no change; grew i n length; produced l a t e r a l haptera, i . e . , they branched ( F i g . 17c,d); or they produced l a t e r a l or t e r m i n a l blades ( F i g . 17b,f,c,). Normally a blade outgrowth appeared f i r s t as a s m a l l blade, and two to three weeks l a t e r a s m a l l s t i p e developed between the blade and the parent hapteron. Sections i n d i c a t e t h a t o n l y the outer corte x i s i n v o l v e d i n p r oduction of l a t e r a l outgrowths from haptera ( F i g . 17 e , f ) . In the shaking experiments blades were produced i n both the shaking and c o n t r o l c u l t u r e s and the shaking appeared to have l i t t l e e f f e c t . The haptera of L. longipes produced blades i n the same manner as the haptera of L. s i n c l a i r i i . The sand scouring experiments showed that both branches and blades can be produced under c o n d i t i o n s of constant sc o u r i n g , although the number and s i z e of outgrowths are l e s s than i n the c o n t r o l experiments. In two instances i n the scouring c o n d i t i o n s , a hapteron produced one branch which bent downward and attached f i r m l y to a rock. The two pieces of s t i p e each produced outgrowths from both ends. Although somewhat deformed, the outgrowths were r e c o g n i z a b l e as blades. The s t i p e s had been held h o r i z o n t a l l y i n the dishes, and the blades a l l bent upward. There i s a strong i n d i c a t i o n that 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 experiments on p o l a r i t y were not conducted. 79 In most of the experiments, d i f f e r e n t r e s u l t s were obtained f o r E haptera and N haptera; f o r blade outgrowths and haptera outgrowths; and f o r l a t e r a l blade outgrowths and te r m i n a l blade outgrowths. A few g e n e r a l i z a t i o n s can be made. The N haptera never grew i n leng t h under any c o n d i t i o n s . When E haptera grew, the growth always occurred at the uncut end. A greater percentage of E haptera produced outgrowths than d i d N haptera under any given s e t o f c o n d i t i o n s . The N haptera never produced both haptera and blade outgrowths on the same hapteron, whereas E haptera o c c a s i o n a l l y d i d . For o r i g i n a l haptera lengths of l e s s than 15 mm, l a t e r a l blades were produced by N haptera, whereas t e r m i n a l blades were produced by E_ haptera. For the standard 20 mm haptera, N haptera g e n e r a l l y produced l a t e r a l blades whereas E haptera produced l a t e r a l and/or t e r m i n a l blades. Terminal blades g e n e r a l l y appeared e a r l i e r than l a t e r a l blades. In the f i n a l analyses of the r e s u l t s , outgrowths were considered merely as blades or haptera, w i t h no regard to whether they were l a t e r a l or t e r m i n a l . i . Temperature - At 20°C there was a considerable growth of va r i o u s contaminants. No p o s i t i v e growth response was shown by any of the haptera and s e v e r a l began to d i s i n t e g r a t e . I t i s concluded that 2 0°C i s a l e t h a l temperature. Within the l i m i t s of 5° to 15°C, va r i o u s r e s u l t s were obtained. The g r e a t e s t percentage of E haptera grew i n length and produced l a t e r a l haptera or branches a t 10°C ( F i g . 17a). No growth occurred a t 15°C. The g r e a t e s t percentage of N haptera 80 produced l a t e r a l haptera a t 8 C. The percentage of E haptera producing blades rose w i t h i n c r e a s i n g temperature, w i t h the maximum of 100 percent a t 15°C. The maximum percentage of N haptera produced blades a t 5°C. The average number of blades per E hapteron which produced blades was highest at 8°c, or i ntermediate temperature, whereas f o r N haptera i t was highest a t the extremes of temperature, 5° and 15°C ( F i g . 17a). i i . L i g h t I n t e n s i t y - The g r e a t e s t percentage of E haptera grew i n l e n g t h and produced l a t e r a l haptera at 150 f t - c ( F i g . 17a). No growth occurred a t 500 f t - c . The g r e a t e s t percentage of N haptera produced l a t e r a l haptera a t 2 f t - c . The percentage of E_ haptera producing blades rose 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 to a maximum of 100 percent a t 500 f t - c . The maximum percentage of N haptera produced blades a t 250 and 500 f t - c . The average number of blades per E_ hapteron which produced blades rose 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 i n t e n s i t y to a maximum a t 500 f t - c . ( F i g . 17a). For N haptera the number of blades was highest a t 2 f t - c , but the next highest occurred at the other extreme, 500 f t - c . Haptera tended to 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 instances where both haptera and blades appeared a t low l i g h t i n t e n s i t i e s , the haptera g e n e r a l l y appeared f i r s t . i i i . I n i t i a l S i z e - The s i z e experiments showed that even pieces of haptera o n l y 2.5 mm long can produce outgrowths of normal appearance ( F i g . 17b). However, two e f f e c t s of the small s i z e were noted, pieces of haptera s h o r t e r than 15 mm 81 produced blades, but never haptera. Secondly, the s i z e of the outgrowth appears to be c o r r e l a t e d w i t h the s i z e of the o r i g i n a l hapteron; l a r g e r pieces produce l a r g e r outgrowths. d. D i s c u s s i o n The d i f f e r e n c e s between the E haptera w i t h one growing t i p and the N haptera 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 of ways. The e f f e c t may be c h i e f l y due to 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 of t e r m i n a l outgrowths from the N haptera. I t may be due to the r e l a t i v e age of the pieces of haptera, since N haptera are f a r t h e r from the growing t i p and thus o l d e r . I t could be expected from t h i s t h a t the N haptera 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 fewer outgrowths. Such was observed to be the case i n t h i s study. However, f i e l d s t u d i e s a l s o showed that an e n t i r e p l a n t can be cut i n h a l f and new haptera w i l l grow out from the center, or o l d e s t , haptera. Yet another p o s s i b i l i t y , which has not been i n v e s t i g a t e d , i s that some substance such as an auxin, which s t i m u l a t e s haptera growth and blade production i s produced or stored i n the apex. S e v e r a l authors have report e d the presence of auxins i n various'members of 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 one of them (van Overbeek, 1940b) r e p o r t e d auxin s p e c i f i c a l l y i n the haptera of C o s t a r i a c o s t a t a . S e v e r a l authors, most r e c e n t l y Dawson (1966), have noted that L. s i n c l a i r i i reproduces 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 h o l d f a s t . The present study has shown that the haptera need 82 not be attached to the parent p l a n t f o r such reproduction to occur. Even very s m a l l pieces of haptera can produce blades 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 tha t t h i s c ould take p l a c e i n nature a f t e r a p o r t i o n of a hapteron i s a c c i d e n t a l l y cut loose from the parent p l a n t , the E type of hapteron, t h a t i s , one which has o n l y been cut once, seems much more l i k e l y . The c o n d i t i o n s under which the maximum number of blades are produced per hapteron are not the same as those under which the maximum number of haptera produce blades. The l a t t e r would seem to be the more important c o n s i d e r a t i o n , i f t h i s p roduction i s regarded as a means of v e g e t a t i v e r e p r o d u c t i o n . Each piece of hapteron, no matter how many attached s t i p e s and blades i t has, i s s t i l l o n l y p a r t of one p l a n t . The experiments i n d i c a t e t h a t 20°C i s a l e t h a l temperature, but the highest 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 not harmful. This i s to be expected from observed n a t u r a l c o n d i t i o n s . A water temperature of 20°C i s very u n l i k e l y on the Oregon Coast, whereas l i g h t i n t e n s i t i e s greater than 500 f t - c do occur. Separate temperature and l i g h t i n t e n s i t y experiments show a maximum blade production a t 15°C and 500 f t - c . However, a l l temperature experiments were run a t 150 f t - c and a l l l i g h t i n t e n s i t y experiments a t 10°C, so t h a t i t may be i n c o r r e c t to assume th a t a combination of high temperature and high l i g h t i n t e n s i t y would r e s u l t i n high blade production. 83 Nevertheless, i f one makes t h i s assumption, a maximum blade production should occur i n e a r l y summer when the water temperature i s r i s i n g to 15°C and the l i g h t i n t e n s i t y i s s t i l l high because the p l a n t s have not yet 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 indeed the case. The s c o u r i n g experiments i n d i c a t e t h a t such production can occur even i f the haptera are p a r t l y b u r i e d i n sand, provided there i s s u f f i c i e n t l i g h t . 84 V I I . TAXONOMY OF L. SINCLAIRII AND L. LONGIPES A. I n t r o d u c t i o n L. s i n c l a i r i i and L. longipes are very d i s t i n c t from a l l other species of the genus. The most s i m i l a r of the other species may be the Mediterranean p l a n t , L. r o d r i g u e z i i Bornet, because of i t s m u l t i p l e s t i p e s . However, i l l u s t r a t i o n s of L. r o d r i g u e z i i i n d i c a t e t h a t although i t has m u l t i p l e s t i p e s , they are very few and w i d e l y separated, 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, u n l i k e a 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. longipes are much more s i m i l a r to each other than to any other species i n the genus. Nevertheless, they are d i s t i n c t from each other. P r e v i o u s l y p u b l i s h e d r e p o r t s have emphasized the presence or absence of mucilage ducts i n 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 present study 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 species which have not been p r e v i o u s l y reported. These are blade width, seasonal l o s s of blades, morphology o f gametophytes, and temperature t o l e r a n c e s . On the b a s i s of these and other 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 that they are two separate taxa. 85 B. Laminaria s i n c l a i r i i (Harvey ex Hooker f. and Harvey) Farlow, Anderson and Eaton, 1877-1889. Harvey, 1852, p. 87 (as Lessonia s i n c l a i r i i ) Farlow, Anderson and Eaton, 1878, f a s c . 3, p. 118 Areschoug, 1883, p. 6 (as Hafqyqia s i n c l a i r i i ) Anderson, 1891, p. 220 Howe, 1893, p. 67 De Toni, 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 Smith, 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 Scagel, 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 Hollenberg and Abbott, 1966, p. 25 Druehl, 1968, p. 541 Markham, 1968, p. 125-131 1. D e s c r i p t i o n Sporophytes up to 3 m long, 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 blades a f t e r complete l o s s of o l d ones. H o l d f a s t an extensive r h i z o m e - l i k e system of branched haptera, sometimes covering an area of 5 m^  or more. The haptera g i v i n g r i s e to many s t i p e s , more than 100 i n some in s t a n c e s . 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 diameter, up to 60 cm long or g r e a t e r , mucilage ducts present. Blades narrowly l i n e a r , u s u a l l y l e s s than 3.0 cm wide, of v a r i a b l e l e n g t h up to 2.5 m, e n t i r e , without b u l l a e , mucilage ducts present. Blades g e n e r a l l y l o s t i n December, regenerated i n January, p l a n t s tending to be l a r g e r near the southern l i m i t s 86 of d i s t r i b u t i o n . 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 to many per blade, p l a n t s produce s o r i i n October, November, January, February, and March. Gametophytes filamentous, capable of prolonged 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 producing sporophytes. 2. D i s t r i b u t i o n Hope I s l a n d (50°55'N, 127°58'W) B r i t i s h Columbia to Ventura County, (34°19'N, 119°23.3'W) C a l i f o r n i a . 3. H a b i t a t Growing on rocks i n the 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 areas, but o c c a s i o n a l l y i n moderately s h e l t e r e d to moderately exposed areas. 4. Comments The mean l e n g t h of the longest s t i p e s from each of 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 longest measured s t i p e i n t h i s group was 52 cm and the s h o r t e s t was 11 cm. P l a n t s lower i n the i n t e r t i d a l r e g i o n are g e n e r a l l y longer but were r a r e l y c o l l e c t e d because of d i f f i c u l t s u r f c o n d i t i o n s . Consequently, the mean l e n g t h of a l l s t i p e s on the Oregon beaches i s probably greater than 27 cm. p l a n t s of L_. s i n c l a i r i i i n C a l i f o r n i a are known to be g e n e r a l l y l a r g e r than those i n Oregon but no comprehensive data are a v a i l a b l e . The mean width of the blades on the l a r g e s t s t i p e s of 54 87 pressed specimens from Oregon i s 1.75 cm. From measurements of both f r e s h and pressed specimens from Indian 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 to be 20%. Applying t h i s c o r r e c t i o n f a c t o r , the mean blade w i d t h . f o r a l l specimens examined i s approximately 2.2 cm. C. Laminaria longipes Bory, 1826 Agardh, C , 1820, p 0 133 Bory, 1826, v o l . 9, p 0 189 P o s t e l s and Ruprecht, 1840, p. 10 (as L. saccharina f. a n g u s t i f o l i a ) K i i t z i n g , 1849, p. 574 Ruprecht, 1851, p. 232, 350 (as Lessonia repens) Le J o l i s , 1855, p. 307-308,311 (as L. ruprechtiana) Agardh, J . , 1867, p. 26 (as Arthrothamnus ? longipes) Areschoug, 1883, p. 15 K j e l l m a n , 1889, pp. 7, 9, 17, 43 De Toni, 1895, p. 370 (as Ar thro thamnus ? longipes) 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 Arwidsson, 1932, p. 153 Miyabe and Nagai, 1932, pp. 196, 197; 1933, pp. 86, 87 Okamura, 1932, p. 72; 1933, pp. 88, 95 Zinova, 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. longipes v a r . l a t i f o l i a ) Okamura, 1936, p. 251, f i g . 139 Nagai, 1940, pp. 67-70; 1941, p. 262 (as L. longipes 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 Tokida, 1954, pp. 30, 114, 115 ( i n c l . L. longipes f . t y p i Zinova, E., 1954, p. 378 Zinova, A., 1959, p. 153 Dawson, 1961, p. 396 Vozzhinskaya, 1964, p. 425 Druehl, 1968, p. 541 88 1. D e s c r i p t i o n Sporophytes up to 1 m long, 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 blades w h i l e remnants of the o l d ones remain. H o l d f a s t an extensive r h i z o m e - l i k e system of branched haptera, sometimes covering an area up to 5 m2. The haptera g i v i n g r i s e to many s t i p e s , more than 100 i n some in s t a n c e s . 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 to 40 cm long, u s u a l l y not exceeding 20 cm i n length, mucilage ducts absent. Blades narrowly l i n e a r , u s u a l l y l e s s than 5.0 cm wide, of v a r i a b l e l e n g t h up to 80 cm, e n t i r e , without b u l l a e , mucilage ducts present. Blades g e n e r a l l y s h o r t e s t i n December or January, but not l o s t completely. S o r i i n patches, round to i r r e g u l a r i n o u t l i n e , one to many per blade, 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 other winter months. Gametophytes filamentous, capable of prolonged v e g e t a t i v e growth, apparently a l l of one type, very r a r e l y producing sporophytes. 2. D i s t r i b u t i o n Urup I s l a n d (46°00'N, 150°00'E) K u r i l e Islands around o o p a c i f i c Rim to Coronation I s l a n d (55 49.6'N, 134 17'W) A l a s k a . 3. H a b i t a t Growing on rocks 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 or moderately s h e l t e r e d areas, but o c c a s i o n a l l y i n f u l l y exposed areas. 89 4. Comments , The mean le n g t h of the 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 greater on p l a n t s from more exposed s i t e s (Table X X I I I ) . However, the longest s t i p e recorded, 3 5 cm, i s on a p l a n t from an area which i s o n l y "moderately exposed". The mean width of the blades of pressed specimens from 33 l o c a t i o n s i s 1.97 cm. From measurements on both f r e s h and pressed specimens from Aats Bay, 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 to be 37%. This i s greater than that f o r L. s i n c l a i r i i . However, examination of other species from Alaska i n d i c a t e s a greater shrinkage i n them a l s o . I t i s b e l i e v e d that the co n c e n t r a t i o n of f o r m a l i n used to preserve the p l a n t s was not the same f o r Oregon and A l a s k a . Applying t h i s c o r r e c t i o n f a c t o r , the mean blade width f o r a l l specimens examined i s approximately 3.1 cm. According to some authors, (e.g., yamada, 1935) some p l a n t s i n the K u r i l e I slands have blades c o n s i d e r a b l y broader than t h i s . D. Comparisons of Species L. s i n c l a i r i i and_L_. longipes are two d i s t i n c t s p e c i e s . The present study has shown th a t there are a t l e a s t 10 d i f f e r e n c e s between the two s p e c i e s , as f o l l o w s : 90 L. s i n c l a r i i L. longipes 1. Mucilage ducts present i n 1, s t i p e . 2. S t i p e i n mature p l a n t 2, u s u a l l y over 20 cm long. 3. Blade of mature p l a n t 3, u s u a l l y l e s s than 3 cm wide. 4. E n t i r e blade l o s t and then 4, new one regenerated. 5. Male and female gameto- 5. phytes i n c u l t u r e m o r p h o l o g i c a l l y d i f f e r e n t . 6. Occurs from Southern 6. C a l i f o r n i a to C e n t r a l B r i t i s h Columbia. 7. Optimal growth a t mean 7. temperature higher than 8°C. 8. Grows i n lower i n t e r t i d a l 8. r e g i o n . 9. U s u a l l y occurs on f u l l y 9. exposed s i t e s . 10. U s u a l l y a s s o c i a t e d w i t h 10. sand and b u r i e d f o r p a r t of year. Mucilage ducts absent from s t i p e . 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 long. Blade of mature p l a n t u s u a l l y more then 3 cm wide. Proximal remnant of blade r e t a i n e d and new one produced while remnant s t i l l present. Gametophyes i n c u l t u r e m o r p h o l o g i c a l l y i n d i s t i n g -u i s h a b l e as to sex. Occurs from Southeast A l a s k a throughout the A l e u t i a n I s l a n d s to the K u r i l e I s l a n d s . Optimal growth a t mean temperature lower than 8 c 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 . U s u a l l y occurs on moderately exposed or moderately s h e l t e r e d s i t e s . R a r e l y a s s o c i a t e d w i t h sand. F i g u r e 18 i l l u s t r a t e s the d i f f e r e n c e s i n blade width and s t i p e l e n g t h . Extreme examples were chosen to i l l u s t r a t e the p o i n t . The d i f f e r e n c e i s u s u a l l y not as great as i n figure 18. As noted above, p o i n t s 3, 4, 5, 7, and 9 have not been r e p o r t e d before; the others have been mentioned, u s u a l l y 91 F i g u r e 18 Habit of L. s i n c l a i r i i and L. longipes a. L. s i n c l a i r i i from Pescadero P o i n t , C a l i f o r n i a , x 1/3. b. L. longipes from cape Spencer, A l a s k a , x 1/3. 92 without emphasis, by various authors. Except f o r d i f f e r e n c e s i n gametophytes and mucilage ducts, none of these p o i n t s alone would seem s u f f i c i e n t as a b a s i s f o r separating the two sp e c i e s . However, a l l of these d i f f e r e n c e s c o l l e c t i v e l y confirm that L. s i n c l a i r i i and L. longipes should d e f i n i t e l y be r e t a i n e d as two separate s p e c i e s . 93 V I I I . GENERAL DISCUSSION AND CONCLUSIONS The two species s t u d i e d , L. s i n c l a i r i i and L. longipes, are very d i s t i n c t from a l l other species of the genus w i t h the p o s s i b l e exception of L. r o d i g u e z i i as noted. They are a l s o d i f f e r e n t from most other members of the order L a m i n a r i a l e s . There are three f e a t u r e s of these two species which are very d i s t i n c t i v e . There i s an apparent suppression of sexual r e p r o d u c t i o n . They have a great 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 of supposedly mature t i s s u e s and r e l a t e d to t h i s , a more d i f f u s e meristematic area which allows considerable growth i n regions other than the 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 other k e l p s e x h i b i t c o nsiderable growth i n regions other than the t r a n s i t i o n zone. Egregia spp. develops a very 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 the f l a t t e n e d s t i p e , w i t h very l i t t l e growth o c c u r r i n g i n the t r a n s i t i o n zone. In Dictyoneurum c a l i f o r n i c u m Rupr. the 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 to the su b s t r a t e by l a t e r a l haptera. Repeated s p l i t s i n the blade cause segmentation and the formation of many blades, each w i t h a s t i p e that becomes p r o s t r a t e so that a clump i s formed ( S e t c h e l l and Gardner, 1925). In Arthrothamnus b i f i d u s (Gmel.) J . Ag. the b a s a l margins of the blades become meristematic and numerous secondary blades a r i s e . The process i s repeated many times, forming 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 blades (Yamada, 1938). The most 94 s i m i l a r development to tha t of L. s i n c l a i r i i i s found i n the Japanese species E c k l o n i a s t o l o n i f e r a Okam. The haptera 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 blades a t the t i p s . The blades are 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 formation of zoosporangial s o r i seems to be suppressed. However, E c k l o n i a d i f f e r s from Laminaria i n having l a t e r a l outgrowths from the bla d e s . With the p o s s i b l e exception of E c k l o n i a s t o l o n i f e r a , the suppression of sexual r e p r o d u c t i o n i n the two species s t u d i e d appears to be unique i n the k e l p s . Sorus production i s not suppressed i n these two species but there i s l i t t l e evidence t h a t the r e s u l t i n g gametophytes normally produce sporophytes i n any number. In view of the 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. longipes and other species of the genus, i t i s p o s s i b l e that they should be removed from the genus Laminaria. One of the c h i e f reasons f o r removing them from the genus i s the meristematic a c t i v i t y of the .haptera which r e s u l t s i n m u l t i p l e s t i p e s . On t h i s b a s i s , however, the two species would have to be removed not o n l y from the genus Laminaria, but a l s o from the f a m i l y Laminariaceae, as the f a m i l i e s of the order L a m i n a r i a l e s are based on the type of growth i n the meristematic area. Thus, before such a taxonomic r e v i s i o n could be made, a comprehensive review of the e n t i r e order would have to be c a r r i e d out. For t h i s reason, no changes of taxonomy are proposed a t the present time. 95 Despite the almost 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 features by which they d i f f e r from other k e l p s , the two species are a l s o d i s t i n c t from each other, as has been demonstrated. Many of these d i f f e r e n c e s may be the r e s u l t of adaptation to d i f f e r e n t h a b i t a t s . 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 of the two species i s i n the temperature of the seawater and the a i r . The mean temperatures f o r the c o l d e s t winter months and the annual means show no o v e r l a p p i n g values at a l l (Table I , F i g . 2, 3 ) . In summer the d i f f e r e n c e i s not as great, but there i s o n l y a s l i g h t o v e r l ap f o r seawater temperature and none f o r a i r temperature. The evidence i n d i c a t e s that sexual r e p r o d u c t i o n i s of very l i t t l e importance to these s p e c i e s . Nevertheless, 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 areas where the two species are found occurs a t the time of year when both species bear r i p e s o r i . I t was observed 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 mucilage and became very sli m y soon a f t e r c o l l e c t i o n . The p l a n t s c o l l e c t e d i n Oregon d i d not become s l i m y a t a l l . Several authors 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 are 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 presence of mucilage. The a i r temperature and degree of i n s o l a t i o n i s much greater on C a l i f o r n i a beaches than on Oregon beaches. Hence the danger of d e s i c c a t i o n i s much greater i n C a l i f o r n i a . The greater production of mucilage i n the C a l i f o r n i a p l a n t s i s probably an adaptation i n response 96 to t h i s . In A l a s k a , where L. longipes i s found, the average a i r temperature and the danger of d e s i c c a t i o n are con s i d e r a b l y l e s s than i n Oregon. L. longipes does not produce mucilage to any n o t i c e a b l e degree. Furthermore, i t l a c k s mucilage ducts i n the s t i p e . No d e f i n i t e c o r r e l a t i o n has been demonstrated between the presence of mucilage ducts and the production of mucilage. Nevertheless, c o n s i d e r i n g the value of 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 of high temperatures and Burrows' (1964) demonstration that mucilage ducts tend to be produced o n l y a t higher temperatures, 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 the d i f f e r e n c e i n mucilage ducts i n the two species may have a r i s e n o r i g i n a l l y as a response to d i f f e r e n t temperatures, even though i t can apparently no longer be a l t e r e d by changing the temperature. Temperature may be the f a c t o r which l i m i t s the d i s t r i b u t i o n of L. s i n c l a i r i i south of Ventura County, C a l i f o r n i a . At Santa Monica, j u s t south of the southern l i m i t s of L. s i n c l a i r i i , both the summer mean temperatures and the y e a r l y mean temperatures are more than 1C° greater than i n Ventura County. A t Short Sand Beach the p l a n t s of 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 almost f r e s h water during the l a t t e r p a r t of the summer. Many of the r e d algae growing on the same rocks d i e d i n l a r g e numbers during t h i s time. L. s i n c l a i r i i showed reduced growth and considerable l o s s of blades but was apparently not i r r e p a r a b l y damaged. 97 These observations, together w i t h the r e s u l t s i n laboratory-c u l t u r e and the great 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 the ranges of both species i n d i c a t e t h a t s a l i n i t y i s of 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. longipes i s not normally found a s s o c i a t e d w i t h sand, although there i s no evidence from t h i s study to i n d i c a t e that i t should be any l e s s w e l l adapted to sand than i s L. s i n c l a i r i i . The c h i e f reason may be simply that w i t h i n the area to which L. longipes i s adapted by i t s temperature requirements there are very few sandy beaches of the type 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 the d i s t r i b u t i o n of L. s i n c l a i r i i w i t h i n i t s temperature range appears to be the presence of sand. I t i s n e a r l y always r e s t r i c t e d to beaches which have a l a r g e seasonal f l u c t u a t i o n of sand and where i t i s p e r i o d i c a l l y b u r i e d . I t i s w e l l adapted to t h i s very harsh environment. In such an area, sexual reproduction by gametophytes i s d i f f i c u l t because of the danger of sand scou r i n g . F u r t h e r , i n such a well-adapted p l a n t a nonsexual means of re p r o d u c t i o n would ensure successive generations of g e n e t i c a l l y s i m i l a r , well-adapted p l a n t s . Three other species which are s i m i l a r l y well-adapted to t h i s type of environment apparently l a c k sexual r e p r o d u c t i o n a l t o g e t h e r : Gymnogongrus  l i n e a r i s (Smith, 1944), Ahnf eltda concinna (Smith, 1944), and Phaeostrophion i r r e g u l a r e (Mathieson, 1967). Laboratory experiments show th a t sexual 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 suppressed even i n environments where sand scouring i s absent. 98 The need f o r an a l t e r n a t i v e , or accessory, nonsexual 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 study has shown th a t such a means e x i s t s i n the production of new s t i p e s and blades from haptera and can operate even i f the haptera are detached from the parent p l a n t . The g e n e r a l i z e d meristematic a c t i v i t y and the considerable r e g e n e r a t i o n 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 to recover a f t e r various p a r t s have been accidentailyremoved. This i s a great advantage i n an environment where sand scouring and the f o r c e of the s u r f are o f t e n 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 not undergo p e r i o d i c b u r i a l under sand, i t would appear that t h i s b u r i a l confers some advantage on the p l a n t . i t s unique methods of growth and r e p r o d u c t i o n enable i t to s u r v i v e i n t h i s very h o s t i l e environment which excludes most other algae. C o l o n i z a t i o n by outgrowths from haptera i s very s u c c e s s f u l i n areas which are 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 scoured away. i n areas which are 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 area can be c o l o n i z e d i n a s i n g l e season. Since L. s i n c l a i r i i apparently reproduces by gametophytes very r a r e l y , i t i s excluded from areas 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 haptera can occur e a s i l y only on areas of rock adjacent to the area where the p l a n t i s already growing. C o l o n i z a t i o n of new and i s o l a t e d rocks 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 that new p l a n t s of L. s i n c l a i r i i are i n i t i a t e d from gametophytes on such areas. The r e l e a s e o f spores a t a time when l i t t l e other a l g a l growth i s o c c u r r i n g and sand i s mostly absent, and a subsequent r a p i d growth of gametophytes and production of young sporophytes, might enable the p l a n t s to become e s t a b l i s h e d before competition and/or sand scouring e l i m i n a t e d them. I t i s assumed th a t young sporophytes are much more r e s i s t a n t to sand s c o u r i n g than are gametophytes. I t was noted that ; L . s i n c l a i r i i appears to be adapted to exposure to severe s u r f . i n the areas of g r e a t e s t exposure, the p l a n t s are l a r g e r and continue growing longer a f t e r b u r i a l by sand. A l s o , i n these areas L. s i n c l a i r i i shows more complete dominance on the rocks on which i t grows. Thus, a t Arch Cape, on the e n t i r e rock where L. s i n c l a i r i i was s t u d i e d , o n l y one other species was found. L. s i n c l a i r i i can withstand b u r i a l b e t t e r than many smaller p l a n t s because i t i s long enough by l a t e summer so that a p o r t i o n of the blades u s u a l l y protrudes from the sand. I t i s not known to what extent, i f any, conduction occurs from the exposed to the b u r i e d p a r t s of the p l a n t . When the p l a n t i s b u r i e d , greater exposure to s u r f might be an advantage i n t h a t greater s u r f a c t i o n would cause greater s t i r r i n g of the sand and a t the same time a l l o w greater water motion and a e r a t i o n around the b u r i e d p a r t s of the p l a n t . 100 L. s i n c l a i r i i has been shown to be uniquely adapted to a very harsh environment. The environment of L. longipes g e n e r a l l y l a c k s sand. However, i t too i s a very harsh environment, i n p a r t because of the low seawater temperatures and even lower a i r temperatures to which the p l a n t i s f r e q u e n t l y subjected. Further s t u d i e s of the type c a r r i e d out on L. s i n c l a i r i i are needed to determine e x a c t l y what environmental f a c t o r s are 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. longipes and the ways i n which the p l a n t i s adapted to i t s environment. Such i n f o r m a t i o n should a l s o c l a r i f y f u r t h e r the r e l a t i o n s h i p of L. longipes to 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 of these two unique species to other k e l p s . 101 IX. SUMMARY The d i s t r i b u t i o n , ecology, growth, and rep r o d u c t i o n of Laminaria s i n c l a i r i i and L. longipes were s t u d i e d i n the 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 species d i f f e r from most other k e l p s i n having m u l t i p l e s t i p e s a r i s i n g from an enlarged r h i z o m e - l i k e h o l d f a s t , composed of many haptera. Each of the s t i p e s bears a blade. The two species are almost i d e n t i c a l i n e x t e r n a l morphology, previous workers have d i s t i n g u i s h e d the two species p r i m a r i l y on the i n t e r n a l anatomy o f the s t i p e : the s t i p e of L. s i n c l a i r i i has mucilage ducts whereas th a t of L. longipes does not. L. longipes occurs from the K u r i l e I s l a n d s through the A l e u t i a n I s l a n d s and i n t o Southeast A l a s k a . L. s i n c l a i r i i occurs from Northern B r i t i s h Columbia to Southern C a l i f o r n i a . The gross d i s t r i b u t i o n of both species appears to be c o n t r o l l e d by seawater temperature. S a l i n i t y appears to 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 both species occur throughout wide ranges of s a l i n i t y and L. s i n c l a i r i i i s able to withstand great f l u c t u a t i o n s i n s a l i n i t y a t a s i n g l e s i t e . Seasonal c y c l e s of growth and rep 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 three beaches i n Northern Oregon where the p l a n t s are subjected to heavy s u r f . The g r e a t e s t growth occurs i n e a r l y summer. As summer advances, the p l a n t s are 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 the ends of the blades protrude. Growth d e c l i n e s during t h i s p e r i o d and i s very slow 102 i n l a t e summer. The f i r s t heavy storms i n f a l l remove the sand and expose the p l a n t s . In November and e a r l y December the p l a n t s bear r i p e s o r i . L a t e r i n December the blades are l o s t , l e a v i n g o n l y the bare s t i p e s . In January new blades are regenerated and when they are o n l y 2-3 cm i n length new s o r i are produced. S o r i are 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 occurs. In March and A p r i l there i s con s i d e r a b l e p r o d u c t i o n of new s t i p e s and blades from the haptera a t the margins of the h o l d f a s t . The d i s t r i b u t i o n of L. s i n c l a i r i i w i t h i n i t s temperature range i s p r i m a r i l y c o n t r o l l e d by the presence of sand. I t i s adapted to t h i s harsh environment by i t s growth and re p r o d u c t i o n . I t possesses great powers of regeneration and i s p o t e n t i a l l y m eristematic i n almost any r e g i o n . Although i t r e g u l a r l y produces s o r i , there i s l i t t l e evidence from e i t h e r f i e l d or l a b o r a t o r y s t u d i e s to i n d i c a t e t h a t the gametophytes which develop from the spores i n these s o r i normally produce sporophytes. Sexual r e p r o d u c t i o n of t h i s type i s d i f f i c u l t because o f the scouring a c t i o n of the sand. The normal method of r e p r o d u c t i o n i s apparently v e g e t a t i v e p r o l i f e r a t i o n from the haptera a t the margins of the h o l d f a s t s . L. longipes was observed i n the f i e l d i n A l a s k a on only three occasions i n summer and twice i n w i n t e r . Growth i s g r e a t e s t during the summer months. The p l a n t s bear r i p e s o r i i n December. Laboratory c u l t u r e s i n d i c a t e t h a t sexual r e p r o d u c t i o n i s very r a r e i n t h i s s p e c i e s . The blades are reduced i n winter but are not completely l o s t . L_. longipes i s 103 not normally a s s o c i a t e d w i t h sand. Transplant experiments showed that each of 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 of the other but cannot s u r v i v e a whole year. Transplants to s i t e s i n B r i t i s h Columbia where the temperature range i s greater than i n Oregon or A l a s k a showed th a t L.. s i n c l a i r i i i s more adversely a f f e c t e d by low winter temperatures, whereas L. longipes i s more adversely a f f e c t e d by high summer temperatures. T r a n s p l a n t i n g to higher or lower temperatures d i d not a f f e c t the production of mucilage ducts. Comparison of the two species 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 l e n g t h of s t i p e s , width of b l a d e s , winter l o s s of blades, morphology of gametophytes and h a b i t a t . The evidence confirms that they should be r e t a i n e d as two separate s p e c i e s . 104 X. BIBLIOGRAPHY A g a r d h , C. 1 8 2 0 . S p e c i e s a l g a r u m . . . . V o l . 1:1-168 ( N o t s e e n ) . A g a r d h , J.G. 1 8 6 7 . De L a m i n a r i e i s . L u n d s U n i v . A r s s k r . 4:1-36. L u n d . 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The development of a r t i f i c i a l c u l t u r e media f o r marine algae. Arch. 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 of the genus A l a r i a G r e v i l l e , as i t occurs on the West coast of North America. Ph.D. Thesis, Univ. B r i t i s h Columbia, x i + 136 pp. Ruprecht, F . J . 1851. Tange des ochotskischen Meeres. In Middendorff, A.T. von, Reise i n den al i s s e r s t e n Norden und Osten S i b i r i e n s wahrend der Jahre 1843 und 1844. Botanik 1(2): 191-435, p i s . 9-18. St. Petersburg. Sanborn and Doty. 1947. The marine algae of the Coos Bay-Cape Arago Region of Oregon. Ore. S t . Monogr. Stud. Bot. No. 8, 66 pp, 4 p i , 1 map. Scagel, R.F. 1957. An annotated l i s t of the marine algae of B r i t i s h Columbia and northern Washington. Nat. Mus. can. B u l l . 150.vi + 289 pp. S e t c h e l l , W.A. 1896. Notes on k e l p s . Erythea 4:41-48, p l . l . 1899. Algae of the P r i b i l o f I s l a n d s . In D. S. Jordan, Fur Seals and Fur S e a l I s l a n d s of the North P a c i f i c Ocean, v o l 3:589-596, Washington. 1905. Regeneration among k e l p s . Univ. C a l . P u b l . Bot. 2:139-169, p i s 15-17. 1912. The k e l p s of the U n i t e d S t a t e s and A l a s k a . Appendix K. In Cameron, F.K. A p r e l i m i n a r y r e p o r t on the f e r t i l i z e r resources of 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. and Gardner, N.L. 1903. Algae of Northwestern America. Univ. C a l . p u b l . Bot. 1(3) :165-418. 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 of North America. P t . I I I . Melanophyceae. Univ. C a l . p u b l . Bot. 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 Chastitikhogo Okeana (Geographical d i s t r i b u t i o n of r e p r e s e n t a t i v e s of the Order of L a m i n a r i a l e s i n the Northern p a r t of 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 (2nd Ed.) x v i i i + 557 pp. 1 map. Harper and Row, New York. S i l v a , p.c. 1957. Notes on p a c i f i c marine algae. Madrono 14(2):41-51. Smith, G.M. 1944. Marine Algae of 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 Univ. 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. Surface temperature and s a l i n i t y observations a t shore s t a t i o n s on the Oregon Coast f o r 1962. Ore. S t . Univ. Dept. Ocean. ONR Data Rept. 11. Sundene, O. 1963. Reproduction and ecology of Chorda tomentosa. Ny t t Mag. Bot. 10:159-167. 1964. The ecology of Laminaria d i g i t a t a i n Norway i n view of t r a n s p l a n t experiments. Nytt Mag. Bot. 11:83-107. Tatewaki, M. 1931. The primary survey of the vegetation of the Middle K u r i l e s . J . Fac. A g r i c . Hokkaido Univ. 29(4) :127-190, p i s I-X. Tokida, J . 1954. The marine algae of southern Saghalien. Mem. Fac. F i s h . Hokkaido Univ. 2 ( l ) : l - 2 6 4 , p i s . I-XV. Van Overbeek, J . 1940a. Auxin i n marine algae. P i . P h y s i o l . 15:291-299. 1940b. Auxin i n marine p l a n t s . I I . Bot. Gaz. 101:940-947. Vozzhinskaya, V.B. 1964. The bottom f l o r a of S a k h a l i n (In Russian) Trudy I n s t . Okeanol. 69:33 0-440. Widdowson, T.B. 1965. A survey of the d i s t r i b u t i o n of 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 respect 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. 1949. Growth-regulating substances i n Laminaria  a g a r d h i i . Science 110:169. 110 Wyatt, B. and G i l b e r t / W. 1967. Surface temperature and s a l i n i t y observations a t p a c i f i c Northwest shore s t a t i o n s f o r 1965 and 1966. Ore. S t . Univ. Dept. Ocean. ONR Data Rept. 25. Wyatt, B., S t i l l R., and Haag, C. 1965. Surface temperature and s a l i n i t y observations 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 . Univ. Dept. Ocean. ONR Data Rept. 21. Yamada, Y. 1935. The marine algae from Urup, the middle K u r i l e s , e s p e c i a l l y from the v i c i n i t y of Iema Bay. S c i . pap. I n s t . A l g o l . Res. Hokkaido Univ. l ( l ) : l - 2 6 , p i s . I-X. 1938. Observations on Arthrothamnus b i f i d u s J . Ag. S c i . Pap. I n s t . A l g o l . Res. Hokkaido Univ. 2:113-118. Yendo, K. 1909. Notes on algae new to Japan. Bot. Mag., Tokyo. 23(270):117-133. 1910. Kaisan Shokubutsu-gaku. (Marine Botany) (Not seen) Zinova, A.D. 1959. ( L i s t of marine algae of Southern S a k h a l i n and the Southern K u r i l e I s l a n d s . ) ( I n Russian) I s s l e d . Dal'Nevost. Morea SSSR. 6:146-161. Zinova, E.S. 1933. Les algues de Kamtschatka ( i n Russian w i t h French summary). I n s t . H y d r o l . E x p l o r . d. Mers d'URSS. Fasc. 17:7-42. 1954. (Marine algae of eastern Kamschatka). ( i n Russian) Komarovskie Chteniya Bot. I n s t . Akad. Nauk. SSSR. 2(9):365-400. I l l XI. 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 of L_. s i n c l a i r i i and L. longipes. LOCATION [~~ Urup I s l a n d A t t u I s l a n d : YEARS COVERED 1949-1962 MEAN FEB. MEAN AUG„ (or c o l d e s t (or warmest month) month) YEARLY MEAN SOURCE T°C S%0 0o5 (Mar.) T°C 11.6 S% T°C 5.1 S%, Eber, Saur & Sette, 1968 Murder P o i n t 1959 1.6 31. 9 11. 3 31. 0 5. 4 31o8 Anon. 1967d Pyramid Cove 1946- 1964 2.1 32. 9 9. 3 31. 5 5. 1 32.1 Anon. 1967d * S i t k a 1943- 1964 4.4 30o 4 14. 1 25. 5 8. 5 27.7 Anon. 1967d Coronation I s l a n d Hope I s l a n d , B.C» (Pine I s l a n d records) R i v e r Jordan Whiffen S p i t •Stanley park In Dec. 1966, 1 C° below S i t k a , 0.5%0 below S i t k a . 1940-1965 1957 1957 1965-1967 7.2 31.3 8.0 30o7 8.0 30.7 7.1 27.9 10.0 31.8 15.0 30.5 15.0 30.5 12.1 26.8 8.6 31.7 La push (Neah Bay records) 1936-1964 Indian Beach (Seaside records . . converted f o r T°) 1 9 6 6 " 1 9 6 7 7.4 30 o3 11.6 32.3 (Mar.) 7.6 27.7 (Jul. ) 13.9 32.3 9.4 27.6 9.8 31.2 10.5 ?30.9? H o l l i s t e r 1966 Widdowson 1965 Widdowson 1965 Anon. 1965a, 1966a,1967a Anon. 1967c G i l b e r t & Wyatt, 1968 i—i H Table I Continued Arch Cape Short Sand Beach (Seaside records converted f o r T°) pescadero P o i n t ( p a c i f i c Grove records) ^| p o r t Hueneme **Santa Monica 1960-1963 1966-1967 1920-1964 1920-1963 1946-1964 9.7 31.4 (Mar.) (Mar.) 7.1 21.5 11.9 33.2 13.2 33„3 13o6 33.5 13.2 32.2 (Jul.) ( J u l . ) 13.4 33.5 14.2 33.7 (Sep.) 16.8 33.6 19.7 33.7 11.3 30.9 10.0 K u j a l a , Wyatt, et a l , 1961-1965 G i l b e r t & Wyatt, 1968 12.9 33.5 . Anon. 1967d 14.8 33.6 Anon. 1967d 16.2 33.7 Anon. 1967d * Transplant s t a t i o n o n l y . No L. s i n c l a i r i i or L. longipes present ** Located j u s t south of southern l i m i t e s of d i s t r i b u t i o n of L. s i n c l a i r i i ( F i g . l ) . 114 Table I I D i s t r i b u t i o n of Laminaria s i n c l a i r i i SITE B r i t i s h Columbia Plover 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 Jordan, W.C. Vancouver I s . **Whiffen S p i t , W.C. Vancouver I s . POSITION SOURCE 50°56 'N, 127°58'W * UBC 49°04 'N, 125°47'W UBC 48°25 .4'N, 124°04'W JWM+ 48°21 •N, 123°43'W JWM ACC. NO. 411 17012 Washington Salmon Bank, San Juan I s . San Juan Co. 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. Agate Beach, C l a l l a m Co. **La push, C l a l l a m Co. 48°26'N, 123°01'W 48°23'N, 124°43.5'W 48 16'N, 122 51'W 48 47 °10«N, 124°43.8'W 53.9'N, 124 37.6' Druehi, 1965 UC++ Druehl, 1965 UBC W UBC 17023 24574 Oregon **In d i a n P o i n t , Indian Beach, Clatsop Co. ** B a l d P o i n t , Indian Beach, Clatsop Co. **E c o l a p o i n t , Clatsop Co. **Arch Cape, Clatsop Co. **Cape F a l c o n , Tillamook Co. **Short Sand Beach, N. end, Tillamook Co. **Short Sand Beach, S. end, Tillamook Co. Cape Kiawanda, Tillamook Co. Yaquina Bay Mouth, L i n c o l n Co. Coos Bay-Cape Arago Region, Coos Co. (Bassendorf Beach, Lighthouse Beach, Squaw I s l a n d , North Bay, Coos Bay, cape Arago) 45°55.9'N, 123°58.8'W 45^55.6'N, 45 55.2'N, 45°48.2'N, 45 46.4'N, 123 58.6 'W 123°58.5 'W 12.3°58.0'W 123 58.5'W 45°45.8'N, 123°58.2'W 45";45.5,N, 45 13.2'N, 123 58'W 123°9.6'W 44°36.8'N, 124°4.1'W 43°20'N, 124°23'W UBC JWM JWM JWM UBC JWM UBC UC 24673 30535 24933 Kj e l d s e n , 1967 Sanborn & Doty, 1947 C a l i f o r n i a 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, Marin Co. Duxbury Reef, Marin Co. San F r a n c i s c o , San F r a n c i s c o Co. **Pescadero P o i n t , San Mateo Co. 41°03.8«N, 124 09'W UC approx. 38 30'N, 123 25'W UC 38°15.3'N, 122°25.2'W UBC 37°53.3'N, 122°42'W UC 37°47'N, 122°30.8'W UC 37°14.5'N, 122°25.2'W UBC 24464 115 Table I I - Continued SITE POSITION SOURCE approx. Cruz Co. 37°10' N, 122°20*W UC Santa Cruz, Santa Cruz Co. 36°57' N, 122°01.8' W UC Asilomar p o i n t , Monterey Co. 36°37. 5' N, 121°56. 5' w Smith,1944 P o i n t Lobos, Monterey Co. 36°31. 2' N, 121°57. 2 1 w Smith,1944 L u c i a , Monterey Co. 36°01. 3 1 N, 121°331 W UC pi e d r a s Blancas P o i n t , San L u i s Obispo Co. 35°40' N, 121u17'W UC Estero Bay, San L u i s Obispo Co. 35°27« N, 120°57 «W UC Morro Bay, San L u i s Obispo Co. 35°22' N, 120°51.3' W UC Pismo Beach, San L u i s Obispo Co. 35°08. 9 N, 120°38. 8 w UC p o i n t S a l , Santa Barbara Co. 34°54. 2 N, 120°40. 4' vr UC P o i n t pedernales, Santa Barbara Co. 34°36. 1 N, 120°38. 5' w UC Gaviota, Santa Barbara Co. 34°28. 3 N, 120°12. 3 ' w UC P o i n t Conception, Santa Barbara Co. 34°26. 8 N, 120°28. 2 ' w UC C a r p i n t e r i a Beach S t a t e Park, 34°23. Dawson, Ventura Co. 1 N, 119°30' W 19 58a Two M i . N.W. of Ventura, Dawson Ventura Co. 34°22. 5 N, 119°28' W 1958a Mussel Shoals, Ventura Co. 34°19' N, 119°23.3' w Dawson 1958a * UBC = Specimen i s i n P h y c o l o g i c a l Herbarium, U.B.C. ** Seen by author a t t h i s s i t e + JWM = Seen by author a t t h i s s i t e but no c o l l e c t i o n s made. ++ UC = Specimen i s i n Herbarium, Univ. of C a l i f o r n i a , Berkeley. 11.6 Table I I I D i s t r i b u t i o n of Laminaria longipes SITE UoS.S.R. K u r i l e I s l a n d s * Urup i s l a n d (Uruppu) Shimushir i s . (Simusiru) Ketoy I s . (Ketoi) Yankicha I s l s . ( U s i s i r u ) Matua i s . (Matuwa) Lovushki I s . (Musisiru) Kharimkotan I s . (Harumkotan) Onekotan I s . (Onnekotan) paramushir I s . (paramusiru) Shumushu I s . (Simusiyu) A t l a s o v a I s . ( A l a i d , A r a i d ) S. S a k h a l i n POSITION 46°00'N, 47°00»N, 47°20»N, 150°00'E 152°uO'E 152°28'E 47°31'N, 152°49'E 154°31'E 153°51'E 154°31'E 154°45 «E 155°50'E~ 156°20'E -157°27*E 143°00'E 48°05«N, 48°32'N, 49 °07 'N/ 49°25'N, 50°25'N, 50°45'N, 50°53'N, 48°00«N, SOURCE Okamura, 1928; Nagai, 1940 Okamura, 1928 Nagai, 1940 Nagai, 1940 Nagai, 1940 Nagai, 1940 Nagai, 1940 Nagai, 1940 Nagai, 1940 Miyabe & Nagai, 1932; Nagai, 1940 Miyabe & Nagai,1932; ACC. NO. Vozzhinskaya, 1964 Bering I s l a n d 55 00 'N, 166 15'E Miyabe & Nagai Kamschatka 1932 Avachinskaya Bay 52°56 'N, 158°36'E Zinova, 1933 Kuimska Bay p p Zinova, 1933 Morzhovaya Bay 53°16 'N, 159°57"E Zinova, 1954 Kronotsky Bay 54°12 •N, 160°36'E Zinova, 19 54 pankara 58°37 'N, 162°24'E Okamura, 1928 Drankinsky p p Okamura, 1928 Barankorfa p p Okamura, 1928 Zavodsk Cape p p Zinova, 1954 A l a s k a S t . P a u l I s l a n d 57°10 'N, 170°15'W S e t c h e l l & / Gardner, 1903 A l e u t i a n I s l a n d s Murder p o i n t , A t t u I s . 52°48 •N, 173°09'E UBC+ 8145 Casco Bay, A t t u I s . 52°48 'N, 173°10'E UBC 8309 Chichagof P t , A t t u I s . 52°57 'N, 173°15'E UBC 8381 Agattu i s . 52°55 'N, 173°10'E S e t c h e l l & Gardner,1903 K i s k a I s . 52°00 *N, 177°30'E S e t c h e l l & Gardner,1903 Trapper's Cove, Adak I s . 51°48 'N, 176°50'W UBC 9631 North I s , Bay of Isl a n d s Adak I s . 51°50 •N, 176°48'W UBC 8391 Cape Agagdak, Adak I s . 52°00 'N, 176°37"W UBC 8150 Zeto P o i n t , Adak I s . 51°55 'N, 176°34'W UBC 8361 Bugle P o i n t , Great 52°02 .3' N, S i t k i n I s . 175°58.8'W UBC 13593 **Ram P t . Beach, Chernofski 53°24 .6' N. Hbr, Unalaska i s . 167 31.6'W UBC 27796 117 Table I I I - Continued SITE POSITION SOURCE ACC.NO. **Ram P t . , Chernofski Hbr, Unalaska I s . 53°25 'N, 167 31 .5' W UBC 27797 **Cape Aiak-Lance P t . , Unalaska I s . 53°19. .1' N, 1 6 7 o 25. 9 •w UBC 27724 **Staraya Bay, Unalaska I s . 53°37. .4-N, 165° 30. 6 W UBC 27793 Cape S a r i c h e f I , Unimak I s . 54 35 'N, 164 56 •w UBC 8338 Cape S a r i c h e f I I , Unimak I s . 54°35 'N, 164^ 56 •w UBC 8214 **Raven P o i n t , Cape S a r i c h e f , Unimak I s . 54°38 'N, 164^51 'W UBC 27790 **E. Anchor Cove, Ikatan Pen* Unimak I s . 54°41. .6' N, 163° 03. 2 W UBC 27736 G u l f of A l a s k a **Eagle Rock, N.E. Harbor Sanak I s . 5 4 o 2 6 .6' N, 1 6 2 o 35. 4 •w UBC 13620 **Nagai I s l a n d 55°12. .6' N, 159° 55. 2 'W UBC 27715 **paul i s . 55°48 .7 ' N, 1 5 9 o 21 W UBC 26878 **Chignik Bay, Nakchamik I s . 56°21 .3 • N, 1 5 7 o 48. 2 •w UBC 27836 **Aghiyuk I s . , Semidi i s l s 56°13. .7' N, 1 5 6 o 47 W UBC 27831 * * C h i r i k o f I s . 55 49. .5' N, 155° 33. 5 •w UBC 27672 **Gurney Bay, Cape I k o l i k Kodiak I s . 56°T7. .7 ' N, 154° 44. 9 •w UBC 27071 pasagshak p o i n t , Kodiak I s . 57 25 'N, 152 29 'W UBC 8162 **Cape Chiniak, Kodiak I s . 57°37. .3-N, 1 5 2 o 09. 5 •w UBC 8161 * * C h i n i a k I s . 57°36. .6" N, 152° 09. .6 •w UBC 27682 * * P e r i l Cape, Afognak I s . 58°07. .5-N, 1 5 2 ° 16 W UBC 25644 **Kayak I s . 59°51 »N, 144 33 •w UBC 20935 **Wingham I s . 60 03 'N, 144°24 •w UBC 22765 Southeast A l a s k a **Cape Spencer 58°14 'N, 136 35 •w UBC 20865 **Cape Ommaney, Baranof I s . 56°10 'N, 134 40 •w UBC 20384 **Aats Bay, Coronation I s . 55"53. .7' N, 1 3 4 o 16 W UBC 20336 **Helm p o i n t . Coronation I s . 55 49. .6' N, 134° 17 W UBC 20319 Washington ++Salmon Bank, San Juan I s . 48° 2 6 'N, 123 01 •w Druehi, 1968 * Ceded by japan to U.S.S.R. i n 1945; names i n parentheses are former Japanese names. ** L. longipes seen by author 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 Table IV A n a l y s i s of Sand Grain S i z e on Oregon Beaches INDIAN BEACH ARCH CAPE SHORT SAND BEACH SIZE % % % 2 nun + 0.01 0.04 Very coarse 0.01 1 - 2 mm 0.06 0.15 0.02 Coarse 0.5 - 1 mm 0.61 2.14 0.27 Medium 0.25 - 0.5 mm 67.85 74.99 75.22 F i n e 0.1 - 0.25 mm 31.41 22.64 24.43 Very f i n e 0.05 - 0.1 mm 0.05 0.01 0.04 Less than 0.05 mm 0.01 0.03 0.01 100.00% 100.00% 100.00% T o t a l s i e v e d : 2575.5 g 2427.2 g 2415.0 g 119 Table 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 T°C T°C Ppt T°C Ppt T°C Ppt. (xn.) Ltka ( S i t k a Magnetic) (Anon. , 1966b, 1967b, 1968a) Jan. -1.8 12.81 -3.9 5.61 -0.8 7.91 3.2 7.77 Feb. 0.1 7.31 1.2 6.99 1.3 8.97 4.7 6.38 Mar. 2.4 2.81 2.1 7.68 -0.7 3.44 5.9 6.95 Apr. 4.6 5.42 4.1 5.49 4.2 2.87 9.0 5.35 May 5.8 6.55 6.4 9.78 8.1 4.46 12.2 4.66 Jun. 8.9 8.60 10.8 1.11 10.9 2.60 14.9 3.46 J u l . 12.3 2.42 12.7 4.59 12.1 4.73 16.3 5.20 Aug. 12.4 6.90 12.1 8.63 13.9 7.43 16.7 7.86 Sep. 11.0 6.02 10.8 11.13 10.9 16.03 14.9 11.49 Oct. 7.1 18.66 5.0 20.49 7.2 14.07 10.8 15.27 Nov. 1.8 5.05 0.4 8.72 2.5 12.16 6.9 12.01 Dec. -0.3 11.51 0.9 4.96 0.7 6.26 3.8 10.17 m. 5.3 93.88 5.2 95.18 5.9 90.93 9.9 96.57 Coronation I s l a n d (Cape Decision)(Anon., 1966b, 1967b, 1968a ) Jan. 0.5 12.09 -1.5 5.66 1.1 6.84 0.9 6.19 Feb. 2.2 8.64 2.8 7.23 3.1 9.55 1.9 5.54 Mar. 3.9 1.66 3.5 10.48 1.0 1.08 2.6 5.35 Apr. 6.0 6.07 4.9 3.03 5.2 1.77 4.8 4.85 May 6.1 5.30 6.4 7.65 8.7 4.07 7.3 4.20 Jun. 9.4 3.83 10.3 1.48 9.5 2.19 9.7 2.73 J u l . 11.1 2.58 11.6 2.67 11.2 3.94 11.4 3.64 Aug. 11.6 1.53 11.4 4.42 13.6 6.07 11.7 5.05 Sep. 10.3 2.63 10.5 10.65 11.8 10.65 10.4 7.56 Oct. 8.6 18.43 6.4 11.17 8.4 11.30 7.9 12.25 Nov. 4.4 4.22 2.7 3.86 5.2 10.97 4.7 9.68 Dec. 1.8 8.81 2.5 7.39 2.8 9.53 2.9 9.08 Ann. 6.3 75.77 6.0 75.69 6.8 77.96 6.3 76.12 120 Table V - Continued 1965 1966 1967 15-Year Mean T°C Ppt. T°C Ppt. T°C Ppt. T°C Ppt. A t t u I s l a n d (Anon., 1966b, 1967b, 1968a) Jan. -0.7 5.47 -0.8 5.14 -1.2 6.71 -0.6 4.18 Feb. 0.4 15.45 -2.1 4.90 -1.7 5.87 -0.7 4.14 Mar. 0.5 17.80 -0.5 2.62 0.3 0.65 -0.1 3.86 Apr. 2.7 16.60 0.9 3.48 1.4 0.02 1.7 4.68 May 4.8 2.59 3.8 7.81 4.9 0.01 4.0 3.85 Jun. 6.7 1.26 5.8 2.18 7.2 0.04 6.6 3.19 J u l . 9.1 5.36 8.6 2.90 9.0 1.20 9.0 4.75 Aug. 10.4 3.96 10.6 4.39 10.9 3„74 10.5 5.64 Sep. 8.5 3.93 9.7 11.52 8.9 5.87 8.9 6.04 Oct. 5.0 6.72 5.4 5.02 5.8 7.81 5.3 6.68 Nov. 1.9 3.93 3.4 9.50 1.9 1.95 2.1 4.79 Dec. 0.4 5.33 0.4 3„75 0.4 0.78 0.7 4.35 Ann. 4.3 88.40 3.8 63.21 4.0 34.65 4.0 56.15 Table VI Mean Temperature on Urup I s l a n d (Tatewaki, 1931) T°C Jan. -1 to -8 Feb. -3 to -8.5 Mar. -4 to -8 Apr. 1 to 2 May 2.5 to 4 Jun. 5 to 10 J u l . 11 Aug. 10 to 15 Sep. 9.5 to 11 Oct. 2 to 3 Nov. No Data Dec. 0 to -1.5 Ann. 2.5 to 4.0 121 Table 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 (for Hope Island) Long term Mean T° Ppt Jan. 3.8 7.64 Feb. 4.4 5.71 Mar. 5.9 5.96 Apr. 7.6 4.27 May 10.0 3.06 j u n . 12.0 1.95 J u l . 13.5 2.59 Aug. 13.7 2.94 Sep. 12.2 4.73 Oct. 9.7 9.12 Nov. 6.6 9.72 Dec. 4.9 10.37 ANNUAL 8.7 68.06 Sooke (E. Sooke - Anderson Cove) (Anon., 1965b, 1966c, 1967c) 1965 1966 1967 Long Term Mean T ° c «?«fc\ T ° c p P f c T ° c p P f c T ° c pP f c-Jan. * * * * 5.0 13.88 * * Feb. * * * * 5.7 8.18 * * Mar. * * * * 5.1 3.76 * * Apr. * * * * 7.0 2.62 * * May * * 10.7 1.02 11.4 1.07 * * Jun. * * 13.2 0.80 15.4 0.25 * * J u l . * * 15.0 1.07 * * * * Aug. * * 15.8 0.55 18.0 0.00 * * Sep. * * 14.7 1.43 15.4 2.91 * * Oct. * * 9.4 6.03 10.7 11.58 * * Nov. * * 6.9 5.98 7.0 4.32 * * Dec. * * 6.1 9.96 4.1 7.46 * * 122 Table V I I Continued River Jordan (Anon., 1965b, 1966c, 1967c) 1965 1966 1967 Long Term Mean T°C Ppt. T°C Ppt. T°C Ppt. T°C Ppt. (in.) Jan. 4.0 12.57 4.9 11.16 * 22.28 4.0 10.37 Feb. 5.3 12.69 5.8 6.63 5.8 17.24 4.1 8.59 Mar. 6.3 1.58 6.0 9.14 5.4 8.58 6.5 6.80 Apr. 8.2 4.06 8.2 2.65 7.0 3.48 9.0 4.52 May 9.6 3.80 9.8 3.01 10.4 1.50 11.2 2.74 Jun. 12.6 0 o66 12.5 2.81 14.3 0.29 13.0 1.90 J u l . * 0.84 14.1 1.49 * 0.80 15.0 1 015 Aug. * 2.28 14.8 1.99 15.9 0.28 15.1 1.58 Sep. 12.2 3.47 13.8 2.97 * 5.67 14.3 3.47 Oct. * 8.26 * 10.55 * 22.71 11.0 8.50 Nov. 8.5 11.26 7.4 12.55 7.8 8.53 8.4 10.61 Dec. 9.9 10.35 * 18.83 4.1 15.71 5.3 13.19 ANNUAL * 71.85 * 83.98 * 107.07 9.7 73.42 Vancouver ( K i t s i l a n o ) (Anon. , 1965b, 1966c, 1967c) 1965 1966 1967 Long Term Mean (HMCS Discovery) T C Ppt. T°C Ppt. T°C Ppt. T°C Ppt. (in.) Jan. 2.8 7.35 3.7 8.85 4.8 11.87 5.7 8.59 Feb. 4.6 9.63 4.8 3.51 5.5 5.53 5.9 6.41 Mar. 5.7 2.26 6.9 4.20 5.7 5.72 * * Apr. 9.9 2.45 9.2 1.24 8.0 3.21 * * May 11.6 2.15 12.7 2.43 13.1 2.15 12.6 2.67 Jun. 16.1 0.57 15.2 2.54 18.3 0.55 17.4 2.67 J u l . 18.8 0.35 17.1 2.84 18.8 1.07 * 1.70 Aug. 18.0 3.11 17.8 1.67 20.3 0.23 19.4 1.64 Sep. 13.8 0.58 15.8 2.85 * * 16.4 3.38 Oct. 11.7 8.20 10.0 7.09 * * 11.0 6.84 Nov. 7.8 5.68 7.1 9.45 * * 7.3 8.14 Dec. 3.8 7.78 6.1 14.62 * * 4.2 9.95 10.38 50.11 10.53 61.29 * * * 61.75 * = No Data 123 Table V I I I Temperature and P r e c i p i t a t i o n at Oregon S t a t i o n s Seaside (Anon., 1966b, 1967b, 1968a) ~ 1965 „ 1966 1967 Long Term Mean T C T"C Ppt. T C Ppt. T°C Ppt. U n . ) Jan. 6.4 19.11 6.8 10.42 7.7 16.85 6.3 11.90 Feb. 7.4 6.42 7.1 6.81 7.4 6.86 7.2 9.84 Mar. 9.1 1.22 7.3 9.82 6.9 8.75 7.9 9.27 Apr. 9.8 4.49 10.4 2.92 7.5 5.48 9.7 5.41 May 10.6 2.91 10.5 1.95 11.3 1.00 11.8 3.31 Jun. 12.8 1.18 14.1 1.64 14.4 1.33 14.0 3.12 J u l . 15.1 0.41 15.2 0.81 16.0 0.23 15.3 1.23 Aug. 16.4 1.85 15.2 0.76 16.4 0.09 15.7 1.53 Sep. 14.6 0.49 15.8 1.98 16.8 2.46 14.9 2.94 Oct. 13.6 3.75 12.2 6.18 13.4 10.55 12.5 7.57 Nov. 10.8 12.53 9.9 9.17 10.3 7.47 9.1 10.40 Dec. 6.1 12.89 8.7 15.63 6.3 11.59 7.4 13.18 Ann. 11.1 67.25 11.1 68.09 11.2 72.66 11.0 79.70 Arch Cape S t a t i o n s A & B P r e c i p i t a t i o n only (in.) 1965 1966 1967 Ten-year Mean A B A B A B ( S t a t i o n A Jan. 17.99 22.60 8.98 11.14 17.14 21.79 12.71 Feb. 6.34 7.62 6.06 6.68 5.42 6.73 9.77 Mar. 1.68 1.50 9.97 12.37 10.78 13.05 8.82 Apr. 6.41 7.59 2.13 2.71 5.16 5.75 5.60 May 2.41 2.94 2.39 2.69 2.12 2.10 3.29 Jun. 1.54 1.57 1.90 2.15 1.46 1.65 3.11 J u l . 1.26 1.24 1.36 1.46 0.47 0.51 1.26 Aug. 2.37 2 062 1.36 1.85 0.20 0.20 2.19 Sep. 1.02 1.17 3.34 3.43 4.16 4.51 3.32 Oct. 3„78 4.49 6.76 8.05 9.00 12.59 7.26 Nov. 11.42 11.83 10.96 9.27 6.16 7.20 12.09 Dec. 14.53 16.76 14.70 20.81 12„49 15.74 11.46 Ann. 70o35 81.93 69.89 82.61 74.54 91.82 81.45 124 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 . Santa Cruz, Santa Cruz Co. Oxnard, Ventura Co. (Anon., 1968a) (Anon., 1968a) T°C P p t . ( i n . ) T°C Ppt. Jan. 9.4 6.84 11.9 3.33 Feb. 10.4 5.81 12.2 2o99 Mar. 11.7 4.15 12.9 2.27 Apr. 13.0 2.11 14.0 1.13 May 14.7 1.01 15.2 0.13 Jun. 16.9 0.21 16.4 0.05 J u l . 17.2 0.03 18.2 0.00 Aug. 17.2 0.06 18.4 0.03 Sep. 17.4 0.27 18.1 0.08 Oct. 15.4 1.38 16.8 0.40 Nov. 12.6 9.58 14.8 1.14 Dec. 10.3 7.12 12.9 3.20 Ann 13.9 31.25 15.2 14.75 Table X Measured Values of Seawater Temperature and S a l i n i t y on Oregon Beaches MONTHLY MEAN INDIAN BEACH SHORT SAND BEACH SEASIDE Date S%o T°C Date T°C Date T°C 17 Aug. •66 33.843 13 Sep. •66 32.058 8.0 14 Sep. •66 30.833 7.8 Sep. '66 31.6 14.8 15 Oct. •66 30.898 9.0 16 Oct. •66 - 10.0 Oct. '66 31.8 13.4 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 02 28 Dec. •66 23.974 9.0 Dec. •66 30.1 11.4 24 Jan. '67 29.689 8.5 25 Jan. •67 16.222 8.5 Jan. '67 27.9 10.6 23 Feb. '67 27.735 9.0 24 Feb. '67 21.479 8.5 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 Apr. '67 29.252 9.0 26 Apr. •67 21.103 9.5 Apr. •67 26.2 11.0 23 May •67 25.821 9.5 22 May •67 24.194 10.0 May •67 27.5 12.8 20 Jun. •67 28.988 13.0 22 Jun. •67 8.521 11.0 Jun. •67 27.9 13.9 19 J u l . •67 23.827 15.7 20 JUlo '67 5.357 14.5 J u l . '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 Sep. •67 29.0 15.8 * Wyatt and G i l b e r t , 1967; G i l b e r t and Wyatt, 1968. 126 Table XI C a l c u l a t e d Mean Monthly Seawater Temperatures * on Oregon Beaches . INDIAN BEACH SHORT SAND BEACH (Seaside - 2.0 C°) (Seaside - 2.5 C°) 1966 Oct. Nov. Dec. 1967 Jan. Feb. Mar. Apr. May Jun. J u l . Aug. Sep, Mean: 11.4^ 9.8 9.4 8.6 7.9 7.6 9.0 10.8 11.9 13.9 12.5 13.8 10.55°C 10.9 9.3 8.9 8.1 7.4 7.1 8.5 10.3 11.4 13.4 12.0 13.3 10.05°C * C a l c u l a t e d by s u b t r a c t i n g mean d i f f e r e n c e from Seaside 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 to September 1967. 127 Table X I I Mean values of Seawater Temperature and „ * S a l i n i t y a t Arch 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 A p r i l 10.88 28.66 May 12.36 29.53 June 12.61 30.91 J u l y 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 MEAN: 11.31 30.97 * K u j a l a and Wyatt, 1961; Oliphant, 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 Indian Beach. L o c a t i o n of points shown on Fi g u r e 7. A l l heights are cm down from standard reference p o i n t . SAND by A: SAND by F: DATE ROCK A X Y PT. F E D (Inner) (Outer) (Inner) (Outei •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 2/9 146 246 218 193 208 * * : = No Data 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 Short Sand Beach. L o c a t i o n of po i n t s shown on Figu r e 8. A l l heights are cm down from standard reference p o i n t . BEACH NORTH WATER-LEVEL DOWN SAND SAND OF STREAM FROM TOP OF DATE ROCK A by A ROCK B by B N. of A N. of B ROCK A ROCK •66:18/8 211 269 213 267 * * * * 14/9 211 239 213 252 * * * * 16/10 211 269 213 282 * * 81 * 11/11 211 277 213 279 * * * * 28/12 211 320 213 287 * * * 94 •67:25/1 211 320 213 287 * * * * 24/2 211 320 213 287 * * * * 28/3 211 320 213 287 * * * * 26/4 211 297 213 * * * * 135 24/5 211 320 213 287 * * * * 22/6 211 274 213 259 198 * 64 64 20/7 ^ 211 302 213 287 198 207 33 25 18/8 2 l l * 213 * 168 * 18 13 3/9 211 249 213 254 226 * 28 * * = No Data 130 Table XV Sand Heights of Beach a t Arch Cape, Oregon (Markham, 1967 unpubl.) Date 31 J u l 5 Sep 2 Oct 2 Nov 27 Nov 31 Dec 26 Jan Distance 66 66 66 66 66 66 67 0+00 50.00 50.00 50.00 50.00 50.00 50.00 50 o00 +50 40.00 40.53 40.18 40.30 40.31 39.85 39.60 1+00 37.18 39.72 38.13 38.20 37.98 37.82 37.42 +50 36.03 38.86 36.50 36.66 36.03 36.22 35.72 2+00 35.60 38.16 35.16 34.89 35.31 34.77 34.25 +50 33.56 37.06 33.71 33.34 32.93 33.42 33.14 3+00 31.83 35.98 32.21 31.95 31.63 32.22 31.78 +50 31.10 34.28 31.01 31.39 30.44 31.12 30.78 4+00 29.99 32.72 30.01 30.09 29.41 30.32 29.81 +50 31.70 29.16 28.58 29.57 28.85 5+00 Deep 31.06 28.31 27.80 28.62 28.14 +50 pool 30.48 27.18 27.60 6+00 26.68 26.74 +50 26.28 Edge o f cobbles 0+50.4 0+48.4 Date 24 Feb Mar 30 Apr 24 May 1 J u l 6 Aug 67 67 67 67 67 67 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 Edge o f Cobbles 50.00 50.00 50.00 50.00 50.00 50.00 39.83 39.96 39.93 40.07 40.09 40.10 37.93 37.26 37.52 37.57 37.63 37.60 35.93 35.39 35.57 35.84 36.49 36.50 34.37 33.82 3.3.65 34.08 34.58 35.00 33.05 32.29 32.11 32.72 32.84 32.66 31.87 31.02 30.88 31.35 31.45 30.59 30.77 29.89 29.98 30.26 30.63 29.80 29.82 28.94 28.42 29.51 29.59 29.23 28.87 28.16 28.62 29.02 29.59 28.40 28.02 27.59 28.42 28.45 26.87 28.00 27.12 26.84 28.17 29.02 27.27 27.84 26.27 27.72 27.16 27.07 26.64 27.27 26.12 28.82 25.67 25.87 25.07 0+47.0 0+47.0 0+44 0+40 Notes: Distance 0+00 i s a f i x e d p o i n t from which a l l other p o i n t s are measured and e s t a b l i s h e d a t 50-foot i n t e r v a l s ; i t has a r b i t r a r i l y been assigned a height of +50 f e e t and a l l other heights have been c o r r e c t e d to t h i s r e f e r e n c e . Edge of cobbles i s the d i s t a n c e west of 0+00 a t which cobbles and sand meet. A l l heights and dis t a n c e s are i n f e e t . ro H Table XVI A s s o c i a t e d P l a n t Species Heights:S RHODOPHYTA A h n f e l t i a concinna J.Ag. A h n f e l t i a p l i c a t a (Huds.) F r i e s Antithamnion sp. Bonnemaisonia nootkana (Esp.) S i l v a B o s s i e l l a corymbifera (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 washingtoniense 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 Vancouveriensis Yendo C o r a l l i n a sp. c o r a l l i n e (crustose) c o r a l l i n e Cryptopleura sp. Cryptosiphonia woodii J . Ag. Cumagloia a n d e r s o n i i ( F a r l . ) S. & G. Dermatolithon d i s p a r (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 Endocladia muricata (Harv.) J . Ag. Erythrophyllum d e l e s s e r i o d e s J . Ag. Euthora 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. & Setch. x Aats La Indian Arch Cape Short Pesc. Bay Push Beach Cape Falcon Sand P o i n t L M H L M H * L M H * L M H 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 X X X X X X X X X X X X X X X X X X X X X X X X X X X G l o i o p e l t i s f u r c a t a (p. & R.) J . Ag. G l o i o s i p h o n i a 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 verrucosa (Huds.) papenf. Gra t e l o u p i a sp. Gymnocfongrus l i n e a r i s (Turn,) J.Ag. Halosaccion glandiforme (Gmel.) Rupr. Halymenia c a l i f o r n i c a Smith & Hollenb. Hildenbrandia sp. I r i d a e a sp. Kallymenia sp. Laurencia s p e c t a b a l i s P. & R. Lithothamnion 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 Melobesia 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. Odonthalia kamtschatika (Rupr.) J . Ag. Odonthalia sp. O p u n t i e l l a c a l i f o r n i c a ( F a r l . ) 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 . P e y s s o n e l i a p a c i f i c a K y l i n Phycodrys sp. Plocamium oregonum Doty Plocamium p a c i f i c u m K y l i n Plocamium violaceum F a r l . Plocamium sp. Polyneura 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. Porphyra sp„ Con t i n ued Aats Bay 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 La Push L M H * X X * X X X * X X X X X X X X Indian Beach 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 Arch Cape L M H * X X X X X * X X x * X X Cape Falcon L M H x X X X X X Short Sand 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 Table oo ro H Heights: 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 P r i o n i t i s l i n e a r i s K y l i n 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. Pt e r o s i p h o n i a b i p i n n a t a (p. & R.) Falkenb. P t e r o s i p h o n i a sp. P t i l o t a asplenoides (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 t e n u i s K y l i n P t i l o t a sp. Rhodoglossum sp. Rhodomela l a r i x (Turn.) C. Ag. Rhodymenia 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 marginata 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. Desmarestia munda S. & G. Desmarestia v i r i d i s (Mull.) Lamour. Demarestia sp. Ectocarpus sp. Egregia m e n z i e s i i (Turn.) Aresch. Continued Aats Bay 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 La Push L M H x X X X X X Indian Beach L M H x x X X X X X X X X X X X * X X X Arch Cape L M H x X X Cape Falcon L M H * X X * X Short Sand 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 Table XVI Cont Heights: Fucus sp. Hap 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.) Setch. Heterochordaria a b i e t i n a (Rupr.) S. & G. Laminaria groenlandica Rosenv. Laminaria longipes Bory Laminaria s e t c h e l l i i S i l v a Laminaria s i n c l a i r i i (Harv.) F a r ., And. & Eat. L e a t h e s i a d i f f o r m i s (L.) Aresch. 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 M a c r o c y s t i s 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. Nereocystxs luetkeana (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. Phaeostrophion i r r e g u l a r e S. & G. Pleurophycus g a r d n e r i Setch. & Saund. P o s t e l s i a palmaeformis Rupr. P u n c t a r i a sp. P y l a i e l l a sp. R a l f s i a sp. Sargassum muticum (Yendo) Fens. Scytosiphon lomentaria (Lyng.) J . Ag. Soranthera u l v o i d e a p. & R. CHLOROPHYTA Cladophora sp. 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 sp. Aats Bay S L M x X X X x x x X X X X X X X X X X d La Push L M H x X X * X X X X X X X X Indian Beach L M H x X X X X X * X X X * X X X X X Arch Cape L M H x X * X X * X X X Cape Falcon L M H x X Short Sand L M H * x x X * X X X X X X X X * * X X X X X X in n rH Table XVI Continued Rhizoclonium sp. Spongomorpha c o a l i t a (Rupr.) C o l l . Spongomorpha sp. U l o t h r i x sp. Ulva l a c t u c a L. Ulva sp. CHRYSOPHATA 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. Aats Bay L M H La Push L M H Indian Beach L M H i Arch Cape L M H Cape Falcon L M H Short Sand L M H Pesc. P o i n t 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 S # s u b t i d a l ; L, lower i n t e r t i d a l ; M, mid i n t e r t i d a l ; H, hig h i n t e r t i d a l * ab^ove = a s s o c i a t e d w i t h sand. Table XVII Seasonal D i s t r i b u t i o n cn H Height: RHODOPHTYA Ah 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 Vancouveriensis Yendo C o r a l l i n a sp. c o r a l l i n e (crustose Cryptopleura sp. Cryptosiphonia woodii J . Ag. D i l s e a c a l i f o r n i c a ( J . Ag.) 0 Kuntze Endocladia muricata (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. Hildenbrandia sp. I r i d a e a sp. Kallymenia sp. Laurencia s p e c t a b a l i s P. & R. Melobesia sp. M i c r o c l a d i a b o r e a l i s Rupr. Odonthalia sp. 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 y s s o n e l i a p a c i f i c a K y l i n Plocamium - oregonum Doty Plocamium p a c i f i c u m K y l i n Polocamium violaceum F a r l . Plocamium sp. of P l a n t Species a t Indian Beach Jun 6 6 L M H * x x X X X X X X X X X X X X X X X Apr 6 7 L M H x X X X X X X X May67 L M H * x * X X X X X * X Jun67 L M H * x X X X X X X X X X X X X X X X X X Jul67 L M H X X * X X * X X X X X X * X X X X Aug67 L M H * x * X X X X X X * X X X Sep67 L M H x X X X X * * X X * X X Table XVII Cont inued H Height: 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 gardneri Smith & Hollenb. 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. Pte r o s i p h o n i a b i p i n n a t a (pc & R.) Falkenb. P t e r o s i p h o n i a sp. P t i l o t a tenuis K y l i n P t i l o t a sp. Rhodomela l a r i x (Turn.) C. Ag. PHAEOPHYTA A l a r i a marginata P. & R. A l a r i a nana Schrad. Desmarestia munda S. & G. Desmarestia sp. Hap 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.) Setch. Laminaria s e t c h e l l i i S i l v a Laminaria s i n c l a i r i i (Harv.) F a r l . And. & Eat. Le 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 Phaeostrophion 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 sp. Ulva sp. Jun 6 6 Apr 6 7 May 6 7 Jun67 J u l 6 7 Aug 6 7 L M H L M H L M H 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 Buried . . . X X X * X Buried X X X X Table XVII Continued Height: CHRYS OP HYTA 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. Jun66 Apr 6 7 May 6 7 Jun 6 7 JU167 Aug 6 7 Sep67 L M H L M H L M H L M H L M H L M H L M H * * 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. Table XVIII Seasonal D i s t r i b u t i o n Height: RHODOP HYTA A h n f e l t i a concinna 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 corymbifera (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 Vancouveriensis Yendo c o r a l l i n e (crustose) c o r a l l i n e Cryptopleura sp. Cryptosiphonia woodii 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 Endocladia muricata (Harv.) J . Ag. Ervthrophyllum delesseriodes J . Ag. F a r l o w i a m o l l i s (Harv. & B a i l 0 ) F a r l . & Setch. G i g a r t i n a sp. G l o i s i p h o n i a 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 0 Ag. Halvmenia c a l i f o r n i c a Smith & Hollenb. Hi1denbrandia sp. I r i d a e a sp. Kallymenia sp. M i c r o c l a d i a b o r e a l i s Rupr. Odonthalia 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 of P l a n t Species at S hort Sand Beach Jun. 66 L M H * x 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 X X X X X X X * X X May 67 L M H x X * X X X X Jun. 67 L M H * x X X X X X X * X J u l . 67 L M H x X X X X X X X * X X X X X X Aug. 67 L M H * x x * X X X Table o r-i Height: polyneura 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 & Hollenb. p r i o n i t i s l a n c e o l a t a Harv. p r i o n i t i s l i n e a r i s K y l i n 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. Pt e r o s i p h o n i a b i p i n n a t a (p. & R.) Falkenb. P t e r o s i p h o n i a sp. P t i l o t a asplenoides (Esp.) C. Ag. p t i l o t a f i l i c i n a ( E arl.) 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 (Turn.) C. Ag. PHAEOPHYTA A l a r i a marginata p. & R. A l a r i a sp. Desmarestia munda S. & G. Desmarestia sp. Ectocarpus sp. Fucus sp. Hedophyllum s e s s i l e (C. Ag.) Setch. Laminaria s e t c h e l l i i S i l v a Laminaria s i n c l a i r i i (Harv.) F a r l . And. & Eat. L e a t h e s i a d i f f o r m i s (L.) Aresch. 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. I Continued Jun. 66 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 Apr. 67 L M H x X X X * * X X X X May 67 L M H x x X X X Jun. 67 L M H * x x X X X * X J u l . 67 L M H x x * X X X * X * X * X 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 Table XVIII Continued Height: Phaeostrophion i r r e g u l a r e S. & G. p i l a y e l l a sp. Soranthera u l v o i d e a P. & R. CHLOROPHYTA Cladophora sp. Enteromorpha i n t e s t i n a l i s (L.) L i n k Enteromorpha sp. Rhizoclonium sp. Spongomorpha c o a l i t a (Rupr.) C o l l . Ulva l a c t u c a L. Ulva sp. 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. Jun. Apr. May Jun. J u l . Aug. Sep. 66 67 67 67 67 67 67 L M H L M H L M H L M H 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 * above x = a s s o c i a t e d w i t h sand Table XIX Seasonal D i s t r i b u t i o n of A l g a l Spe Height: RHODOPHYTA Antithamnion 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 Vancouveriensis Yendo C o r a l l i n a sp. c o r a l l i n e (crustose) Cryptopleura sp. Cryptosiphonia woodii 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 Endocladia muricata (Harv.) J . Ag. Euthora 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 . & Setch. 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. Grateloupia sp. Halosaccion glandiforme (Gmel.) Rupr. I r i d a e a sp. Kallymenia sp. Lithothamnion 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 ter a sp. M i c r o c l a d i a sp. Odonthalia kamtschatika (Rupr.) J . Ag. Odonathalia sp. O p u n t i e l l a c a l i f o r n i c a ( F a r l . ) 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 0) K j e l l . Plocamium sp. P o l y p o r o l i t h o n sp. es at Aats Bay, Coronation I sland Jun. 67 S L M H Dec. 65 S L M H x x X X X X X X X X X Jun. 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 Dec. 66 S L M H X X X X X X X X X X X X X P o l y s i p h o n i a sp. Porphyra sp. P t e r o s i p h o n i a sp. Rhodoglossum sp. 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 marginata 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.Ag. CV s t o s e i r a osmundaceae (Menz.) C. Ag. Desmarestia v i r i d i s (Mull.) Lamour. Desmarestia sp. Fucus sp. Hedophvllum s e s s i l e (C. Ag.) Setch. Heterochordaria a b i e t i n a (Rupr.) S. & Laminaria groenlandica Rosenv. Laminaria longipes Bory L e a t h e s i a d i f f o r m i s (L.) Aresch. Mvelophvcus i n t e s t i n a l e Saund. Nereocvstis luetkeana (Mert.) p. &.R. Pleurophycus gardneri Setch. & Saund. Planetaria sp. Sargassum muticum (Yendo) Fens. Scvtosinhon lomentaria (Lyngb.) J . Ag. Soranthera u l v o i d e a p. & R. Jun. 65 S L M H x X X X X X X X X X X X X Dec. 65 S L M H X X X X X X X X X X X X Jun. 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 Dec. 66 S L M H x x X X X X X X X X X X X Table XIX Continued CHLOROP HYTA Cladophora sp. Codium s e t c h e l l i i Gard. Enteromorpha sp. Monostroma sp. RhizocIonium sp. Spongomorpha sp. U l o t h r i x U l v a Jun. 65 S L M H x x Dec. 65 S L M H x x X Jun. 66 S L M H x X X X X X X X X X X Dec. 66 S L M H 145 Table XX Summary of Seasonal Cycles on Oregon Beaches INDIAN BEACH 8/66 Sand high. Many blades m i s s i n g 9/66 Sand higher. H o l d f a s t s b u r i e d . 10/66 Sand a l l gone. Most p l a n t s r i p e . 11/66 P l a n t s r i p e . 12/66 Some p l a n t s r i p e , most l a c k b l a d e s . 1/67 2/67 3/67 4/67 5/67 6/67 7/67 8/67 9/67 New blades present (1 cm) New blades (3 cm) Small t e r m i n a l s o r i . Sand coming back. Small t e r m i n a l 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 blade s . Sand higher - not ye t to rocks b e a r i n g p l a n t s . Sand around rock bases. Sand covers a l l rocks, h o l d f a s t s , s t i p e s ; o n l y blades p r o t r u d i n g . SHORT SAND BEACH Sand high. Fresh water touches manyplants Sand higher. Sand lower Many p l a n t s r i p e . Sand lower y e t . p l a n t s r i p e . Sand a l l gone. A l l blades m i s s i n g . New blades present (1 cm) New blades (3 cm) Small t e r m i n a l s o r i . Sand coming back. Many new s t i p e s and b l a d e s . ARCH CAPE Sand higher - rocks s t i l l exposed. Sand up, stream f l o o d i n g F i r s t p l a n t s measured. Sand higher, 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 , p l a n t s p a r t l y immersed i n f r e s h -water Most p l a n t s b u r i e d . A l l blades present. Sand down s l i g h t l y . Sand very high Inshore, many blades Many blades m i s s i n g ; f a r t h e r out, m i s s i n g , p l a n t s are i n t a c t . Sand a l l gone. Sand unchanged near Sand has receded 50 Most p l a n t s r o c k s . Few blades cm. P l a n t s appear he a l t h y . present except on healthy. rock tops untouched by freshwater. 146 Table XXI Summaries of i n s i t u growth measurements on Oregon Beaches. Indian Beach (A Rock) DATE NUMBER NUMBER MEAN MEASURED PREVIOUS STIPE HOLES LENGTH (cm) 67/ MEAN DISTANCE BLADE BETWEEN LENGTH HOLES (cm) (cm) 5 5 0 28.1 41.9 -6 5 5 30.4 46.5 7.4 7 0 * * * * 8 0 * * * * 9 0 * * * * 10 25 0 11.7 17.0 -11 7 0 14.0 13.2 -12 7 0 17.9 2.7 -1 E s t . * * 1.0 -2 15 0 18.3 6.1 -3 E s t . * * 13.0 -4 23 0 16.3 18.8 -5 29 6 19.8 29.9 10.8 6 26 26 21.8 40.1 9.2 7 9 9 Buried 42.0 4.6 8 0 * B u r i e d B u r i e d * Indian Beach (B Rock) (more exposed) '67/ 5 6 20 10 0 10 35.9 37.8 52.1 66.4 15.7 147 Table XXI Continued DATE NUMBER NUMBER MEAN MEAN DISTANCE MEASURED PREVIOUS STIPE BLADE BETWEEN HOLES LENGTH LENGTH HOLES (cm) (cm) (cm) Short Sand Beach •66/ 8 9 10 11 12 '67/ 1 2 3 4 5 6 7 8 9 2 2 25 22 0 Est. E s t . E s t . 27 25 24 20 0 0 0 2 0 5 * * * * 0 8 24 19 0 0 14.5. 14.5 13.1 12.6 * * * * 9, 12, 15, 14.0 * * .0 .9 .1 16.8 -17.7 1.5 19.9 -17.6 1.1 * * 1.0 -3.0 -5.0 -12.7 -20.2 10.6, 30.2 10.8 33.3 3.5 No blades l e f t i n measurement area Arch Cape •67/ 6 7 8 9 23 8 7 8 0 7 5 6 17.8 B u r i e d B u r i e d 22.6 26.5 36.1 34.3 36.3 6.0 6.3 1.6 * = No Data N.B. In some ins t a n c e s where number measured i s greater than number of ho l e s , holes were present but o b v i o u s l y o l d e r than one month. 148 Table XXII Growth of Multi-punched Blade of L. s i n c l a i r i i i n s i t u a t Indian Beach. Blade base to A A to B B to C C to D D to E E to F F to G G to H H to I Distance (cm) at time of punching holes 23 May '67 4.0 2.0 2.0 2.0 2.0 2.0 2.0 2.0 2.0 Distance (cm) a f t e r one Month 20 Jun »67 8.0 3.5 2.5 2.2 2.1 2.0 2.0 2.0 2.0 Growth per Month 4.0 (2 cm/2 cm) 1.5 0.5 0.2 0.1 0.0 0.0 0.0 0.0 149 T a b l e X X I I I D i m e n s i o n s o f p r e s s e d S p e c i m e n s o f L . l o n g i p e s f r o m 33 A l a s k a n s i t e s . S I T E : BLADE WIDTH (cm) S T I P E LENGTH(cm) M i n . Max. Mean M i n . Max. Mean M o d e r a t e l y s h e l t e r e d C h i c h a g o f P t . , A t t u I s . 0. 5 2. 0 1. 2 2. 0 11. 0 7. 2 T r a p p e r ' s C o v e , A d a k I s . 1. 5 3 . 5 2. 6 3 . 0 8. 0 4. 8 E a g l e R o c k , NE H b r . , S a n a k I s . 3 . 5 3 . 5 3 . 5 4. 0 7. 0 5. 5 G u r n e y Bay, K o d i a k I s . 0. 5 4 . 5 1. 2 5. 0 15. 0 6. 7 A a t s B a y , C o r o n a t i o n I s . 1. 0 4. 0 1. 7 4. 0 9 . 0 7. 3 TOTALS 0. 5 4 . 5 2. 0 2. 0 15. 0 6. 3 M o d e r a t e l y e x p o s e d 0 C a s c o B a y , A t t u I s . 2. 0 3 . 0 2. 3 3 . 0 12. 7. 5 Cape A g a g d a k , A d a k i s . 0. 5 1. 5 1. 0 7. 0 20. 0 11 . 1 Ram P t . B e a c h , U n a l a s k a I s . 1. 5 3 . 0 2. 0 1. 0 10. 0 8. 5 S t a r a y a B a y , U n a l a s k a I s . 1. 0 1. 0 1. 0 l l o 0 11 . 0 11. 0 Cape S a r i c h e f I , U n a l a s k a I s . 1. 5 2. 0 1. 6 7. 0 15. 0 11. 2 Cape S a r i c h e f I I , U n a l a s k a I s . 2. 0 4. 0 3 . 0 3 . 0 5 . 0 4. 0 E . A n c h o r C o v e , U n i m a k I s . 2. 0 2 . 5 2. 2 9 . 0 25. 0 17. 3 E a g l e R o c k , NE H b r . , S a n a k I s . 1. 0 2. 0 1. 5 5. 0 20. 0 7. 0 N a g a i I s . 1. 5 1. 5 1. 5 7. 0 9 . 0 8. 0 P a u l I s . 0. 5 1. 0 0. 7 6. 0 6. 0 6. 0 C h i r i k o f I s . 1. 5 2. 0 1. 7 8. 0 9 . 0 8. 5 p a s a g s h a k P t . , K o d i a k I s . 1. 5 3 . 5 3 . 0 2. 0 7. 0 3 . 0 Cape C h i n i a k , K o d i a k I s . 2. 0 5. 0 3 . 3 5. 0 9 . 0 8. 1 C h i n i a k I s . 1. 0 3 . 0 2. 8 8. 0 21. 0 16. 3 P e r i l c a p e , A f o g n a k I s . 1. 5 2. 0 1. 6 2. 0 6. 0 4. 2 E n g l i s h B a y 2. 5 3 . 0 2. 8 9 . 0 11 . 0 10. 0 Wingham I s . 2. 0 2. 5 2. 2 4. 0 6. 0 5. 5 Cape S p e n c e r 1. 0 4 . 0 2. 7 6. 0 15. 0 8. 3 H e l m P t . , C o r o n a t i o n I s . 0. 5 3 . 0 1. 4 4. 0 16. 0 8. 4 TOTALS 0.5 5 .0 2 .0 1.0 35 .0 8.6 150 Table X X I I I Continued SITE: BLADE WIDTH (CM) STIPE LENGTH(CM) Min. Max. Mean Min. Max. Mean F u l l y exposed Murder P t . , A t t u I s . North I s . , Adak I s . Zeto P t . , Adak I s . Cape Aiak-Lance P t . , Unalaska I s . Raven P t . , Unimak I s . Chignik Bay, Nakchamik I s . Aghiyuk I s . , Semidi i s l a n d s Kayak I s . Cape Ommaney, Baranof I s . 1.5 3.5 2.6 5. 0 14.0 9.1 2.5 3.5 2.9 6. 0 12.0 7.8 0.5 2.0 1.1 10. 0 15.0 12.7 1.0 2.5 1.6 9. 0 12.0 10.6 2.5 3.0 2.8 7. 0 14.0 9.2 0.5 1.0 0.7 4. 0 5.0 4.5 0.5 2.0 1.2 3. 0 15.0 9.9 2.5 3.5 2.8 7. 0 13.0 9.5 1.0 2.0 1.5 5. 0 11.0 9.0 TOTALS 0.5 3.5 1.9 3.0 15.0 9.1 EXTREME MINIMA & MAXIMA & TOTAL MEANS: 0.5 5.0 2.0 1.0 35.0 8.4 151 X I I : APPENDIX I Summary D e s c r i p t i o n s of F i e l d S t a t i o n s Volga 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 loose rocks l y i n g i n t i d e pools and surge channels. Moderately to f u l l y exposed to s u r f . Mean annual seawater temperature = 8.5 C; s a l i n i t y = 27.7% c Mean annual a i r temperature = 6.3 C; p r e c i p i t a t i o n = 96.57 inches. Neither L. s i n c l a i r i i nor L. longipes present. 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 . Observations made: 1965: June, December 1966: J u l y , December Aats Bay, Coronation 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 channels, adjacent to beach of g r a v e l and coarse sand. No data on mean annual seawater temperature and s a l i n i t y . Mean annual 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 inches. L. longipes present i n abundance„ S i t e used f o r i n s i t u s t u d i e s of L. longipes, t r a n s p l a n t s t u d i e s , and as a source f o r a l l L. longipes 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 experiments. Observations made: 1965: June, December 1966: June, J u l y , December. R i v e r Jordan, Vancouver I s l a n d , B r i t i s h Columbia. (48°25.4'N 124 04-W) Loose rocks i n sandy mud bottom. Moderately s h e l t e r e d to moderately exposed to s u r f . No data on mean annual seawater temperature and s a l i n i t y . Mean annual a i r temperature = 9.7°C; P r e c i p i t a t i o n = 73.42 inches. L. s i n c l a i r i i present as s m a l l p l a n t s i n very s m a l l q u a n t i t i e s , hidden 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 . Observations made: 1965: August, November 1966: January, March, May, June, August. 152 Sooke Harbour, W. of Whiffen S p i t , Vancouver I s l a n d , B r i t i s h  Columbia. (48021.2'N, 123°44'W) Loose rocks and l a r g e f l a t outcrops 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 . Moderately s h e l t e r e d from s u r f . No data on mean annual seawater temperature and s a l i n i t y or a i r temperature or p r e c i p i t a t i o n . L. s i n c l a i r i i present as s m a l l p l a n t s i n small 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 found. 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 . Observations made: 1965: J u l y , August, November. 1966: January, February, March, May, June, August. Stanley Park, 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 boulders. F u l l y s h e l t e r e d . No data on mean annual a i r temperature; p r e c i p i t a t i o n = 61.75 inches Mean annual seawater temperature = 9.4°C: s a l i n i t y = 27.6% 0. Nei t h e r L. s i n c l a i r i i nor L_. longipes present. 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 . Observations made: 1965: August, November, December 1966: January, February, March, May, June. 1967: January, May, J u l y . Indian P o i n t , Indian Beach, Clatsop County, Oregon. (45°55.9'N, 123°58.8'W) Sandy beach w i t h rocky outcrops ( 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 through year. F u l l y exposed to heavy s u r f . Mean annual seawater temperature = 10.5 C; s a l i n i t y Q = approx. 30.97%a. Mean annual a i r temperature = 11.0 C; P r e c i p i t a t i o n = 79.7 inches. L. s i n c l a i r i i present i n abundance as dominant p l a n t i n terms of cover. 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 source 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 experiments. Observations made: 1965: February, J u l y , August, December. 1966: One to three times every month except January, March, A p r i l , J u l y . 1967: One to f i v e times on one low t i d e s e r i e s every month, January through September. 1968: March. 153 Arch cape, Clatsop County, Oregon. (45 048.2 !N, 123°58.2'W) Sandy beach w i t h l a r g e rocky outcrops. 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 to heavy s u r f . Mean annual seawater temperature = 11.3 ; s a l i n i t y = 30.97%.. Mean annual a i r temperature = 11.0°C; P r e c i p i t a t i o n = 81.4 inches. L. s i n c l a i r i i present i n abundance, but o n l y a c c e s s i b l e a t lowest 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 . Observations made: 1967: May, June, J u l y , August, September. Short Sand Beach, Tillamook County, Oregon. (45 045.5'N, 123°58'W) Sandy beach w i t h rocky outcrops (sandstone). Sand l e v e l f l u c t u a t e s 1 - 2 m through year. Freshwater stream flows across beach a t low t i d e , o f t e n c overing some of p l a n t s s t u d i e d . F u l l y exposed to heavy s u r f . Mean annual seawater temperature = 10.0 C; i n s u f f i c i e n t data on s a l i n i t y . Mean annual a i r temperature = 11.0°C; p r e c i p i t a t i o n = approx. 81.4 inches. L. s i n c l a i r i i present 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 experiments, and as a source 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 experiments. Observations made: 1966: June, August, September, October, November, December. 1967: Once or twice on one low t i d e s e r i e s every month, January through September. 1968: March. 

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