@prefix vivo: . @prefix edm: . @prefix ns0: . @prefix dcterms: . @prefix skos: . vivo:departmentOrSchool "Science, Faculty of"@en, "Botany, Department of"@en ; edm:dataProvider "DSpace"@en ; ns0:degreeCampus "UBCV"@en ; dcterms:creator "Druehl, Louis D."@en ; dcterms:issued "2011-10-28T19:01:37Z"@en, "1965"@en ; vivo:relatedDegree "Doctor of Philosophy - PhD"@en ; ns0:degreeGrantor "University of British Columbia"@en ; dcterms:description """This study of the brown algal genus Laminaria Lamour consisted of (1) a critical review of the taxonomy and distribution of taxa of Laminaria occurring in the northeast Pacific, (2) a description of the life histories and growth patterns of long and short stipe forms of L. groenlandica and L. saccharina, (3) an evaluation of the roles of temperature, salinity, and water motion as possible determinants of local distributions of long and short stipe forms of L. groenlandica and L. saccharina, and (4) an evaluation of the roles of temperature, salinity, exposure, and submarine illumination in determining the vertical distribution of L. saccharina. These studies were made from 1961 to 1965. Ten species of Laminaria are recognized for the northeast Pacific: L. groenlandica Rosenv.; L. farlowii Setchell; L. saccharina (L.) Lamour.; L. setchellii Silva; L. dentigera Kjellman; L. longlpes Bory; L. sinclairii (Harvey ex Hooker f. et Harvey) Farlow, Anderson et_ Eaton; L. ephemera Setchell; L. yezoensis Miyabe; and L. complanata (Setchell et Gardner) Setchell. Laminaria cordata Dawson is considered conspecific with L. saccharina, L. personata Setchell and Gardner is regarded conspecific with L. yezoensis, and L. platymeris De la Pyl. (sensu Setchell and Gardner) is considered conspecific with L. groenlandica. Pour forms of L. groenlandica are recognized for the northeast Pacific. These forms are not considered as legitimate taxonomic entities but are distinguished merely to provide a means of facilitating discussion. The known habitat requirements for all ten species were broadened and the known distributions of all species, excepting L. groenlandica, were extended. Laminaria saccharina and L. groenlandica produced sori in the late spring and winter. New sporophytes of L. groenlandica appeared throughout the year, whereas those of L. saccharina appeared in late winter and early fall. Depending upon culture conditions, two morphologically distinct forms of gametophytes were produced by both species: large gametophytes were produced in conditions of high temperature and low salinity; and small gametophytes in conditions of low temperature and high salinity. Abnormal sporophytes were observed under conditions conducive to formation of large gametophytes. Patterns of growth for the blades of the two species were essentially the same.. The growth rate decreased with increase in distance from the blade base, and the position of greatest longitudinal growth coincided with the position of greatest lateral growth. The distributions of L. saccharina and the two forms of L. groenlandica about Vancouver Island were correlated with temperature, salinity, and water motion. The two forms of L. groenlandica were absent from areas of high temperature and low salinity; L. saccharina was absent from areas subjected to surf. These field conclusions were subjected to laboratory and field tests involving gametophytes and sporophytes of both species. The distributions of the two forms of L. groenlandica can be explained on the basis of temperature and salinity distributions. Both forms require low temperature and high salinity for survival. Laminaria saccharina has a wide range of tolerance to temperature and salinity. Surf appears to be the agent controlling the distribution of this species. The upper limits of L. saccharina, as observed in Burrard Inlet, are thought to-be directly related, to air temperature and insolation and indirectly related to tidal characteristics. The lower limits appear to reflect the compensation depth of this species."""@en ; edm:aggregatedCHO "https://circle.library.ubc.ca/rest/handle/2429/38378?expand=metadata"@en ; skos:note "The U n i v e r s i t y of B r i t i s h Columbia FACULTY OF GRADUATE STUDIES PROGRAMME OF THE FINAL ORAL EXAMINATION FOR THE DEGREE OF DOCTOR OF PHILOSOPHY of LOUIS DIX DRUEHL B.S., Washington State U n i v e r s i t y , 1959 M.S., Univ e r s i t y of Washington, 1961 WEDNESDAY, SEPTEMBER 29, 1965, AT 3:00 P.M. ROOM 3332, BIOLOGICAL SCIENCES BUILDING COMMITTEE IN CHARGE Chairman: I. McT. Cowan 1. M. Cole R. F. Scagel G. L. Pickard T. M. C. Taylor G. E. Rouse G. H. N. Towers E. B. Tregunna External Examiner: M. Neushul Univ e r s i t y of C a l i f o r n i a , Santa Barbara, C a l i f o r n i a , ABSTRACT ON THE TAXONOMY, DISTRIBUTION., AND ECOLOGY OF THE BROWN ALGAL GENUS LAMINARIA IN THE NORTHEAST PACIFIC This study of the brown a l g a l genus Laminaria Lamour. consisted of (1) a c r i t i c a l review of the. taxonomy and d i s t r i b u t i o n of taxa of Laminaria occurring i n the northeast Pacific,, (2) a d e s c r i p t i o n of the l i f e h i s t o r i e s and growth patterns of long and short stipe forms of L, groenlandica and L. saccharina,, (3) an evaluation of the roles of temperature, s a l i n i t y , and water motion as possible determinants of l o c a l d i s t r i b u t i o n s of long and short stipe forms of L„ groenlandica and L saccharina ; and (4) an evaluation of the roles of temperature 3 salinity;, exposure 3 and submarine i l l u m i n a t i o n i n determining the v e r t i c a l d i s t r i b u t i o n of L. saccharina, These studies were made from 1961 to 1965, Ten species of Laminaria are recognized for the northeast P a c i f i c : L. groenlandica Rosenv.; L. f a r l o w i i S e t c h e l l ; L, saccharina (L„) Lamour.; L s e t c h e l l i i S i l v a ; L. dentigera Kjellman; L. Longipes Bory; L„ s i n c l a i r i i (Harvey ex Hooker f„ et Harvey) Farlow, Anderson et Eaton; L. ephemera S e t c h e l l ; L, yezoens is Miyabe; and L, compJanata (Setchell et Gardner) S e t c h e l l . Laminaria cordata Dawson i s considered conspecific with L, saccharina„ _L_. personata Se t c h e l l and Gardner i s regarded conspecific with L yezoensis, and L. platymeris De l a Pyl. (sensu S e t c h e l l and Gardner) i s considered conspecific with L. groenlandica. Four forms of L. groenlandica are recognized for the northeast P a c i f i c . These forms are not considered as legitimate taxonomic e n t i t i e s but are distinguished merely to provide a means of f a c i l i t a t i n g discussion. The known habitat requirements for a l l ten species were broadened and the known d i s t r i b u t i o n s of a l l species, excepting L. groenlandica., were extended. Laminaria saccharina and L. groenlandica produced s o r i i n the late spring and winter. New sporophytes of L, groenlandica appeared throughout the year, whereas those of L, saccharina appeared i n late winter and early f a l l . Depending upon culture i conditions, two morphologically d i s t i n c t forms of gametophytes were produced by both species: large gametophytes were produced in conditions of high temperature and low s a l i n i t y j and small gametophytes i n conditions of low temperature and high salinity„ Abnormal sporophytes were observed under conditions conducive to formation of large gametophytes. Patterns of growth for the blades of the two species were e s s e n t i a l l y the same. The growth rate decreased with increase i n distance from the blade base, and the p o s i t i o n of greatest l o n g i t u d i n a l growth coincided with the p o s i t i o n of greatest l a t e r a l growth, The d i s t r i b u t i o n s of L„ saccharina and the two forms of L groenlandica about Vancouver Island were correlated with temperature, s a l i n i t y , and water motion. The two forms of L, groenlandica were absent from areas of high temperature and low s a l i n i t y ; L. saccharina was absent from areas subjected to surf. These f i e l d conclusions were subjected to laboratory and f i e l d tests involving gametophytes and sporophytes of both species. The d i s t r i b u t i o n s of the two forms of L. groenlandica can be explained on the basis of temperature and s a l i n i t y d i s t r i b u t i o n s . Both forms require low temperature and high s a l i n i t y for s u r v i v a l . Laminaria saccharina has a wide range of tolerance to temperature arid s a l i n i t y . Surf appears to be the agent 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 t h i s species., The upper l i m i t s of L, saccharina, as observed i n Burrard I n l e t , are. thought to be. d i r e c t l y r e l a t e d to a i r temperature and i n s o l a t i o n and i n d i r e c t l y r e l a t e d to t i d a l c h a r a c t e r i s t i c s . The lower l i m i t s appear to r e f l e c t the compensation depth of t h i s species. GRADUATE STUDIES Marine Phytoplankton Synoptic Oceanography Dynamic Oceanography Chemical Oceanography Advanced Phycology R. F. Scagel R, F. Scagel G. ]. Pickard R. W. Burling P. W. Williams ON THE TAXONOMY, DISTRIBUTION, AND ECOLOGY OP THE BROWN ALGAL GENUS LAMINARIA IN THE NORTHEAST PACIFIC by LOUIS D, DRUEHL B.S. Washington State University, Pullman, Washington, 1959 M.S. University of Washington, Seattle, Washington, 196l A' THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in the Department of BIOLOGY AND BOTANY We accept t h i s thesis as conforming.to the required standard THE UNIVERSITY OF BRITISH COLUMBIA August, 1965 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 advanced degree 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 and 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 by t h e Head o f my Department o r by h i s r e p r e s e n t a t i v e s . . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i -c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Department o f Biology & Botany The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada Date September 30, 1965 i i ABSTRACT This study of the brown a l g a l genus Laminaria Lamour. consisted of ( l ) a c r i t i c a l review of the taxonomy and d i s t r i b u t i o n of taxa of Laminaria occurring i n the north-east P a c i f i c , (2) a description of the l i f e h i s t o r i e s and growth patterns of long and short stipe forms of L. groen- landica and L. saccharina, (3) an evaluation of the roles of temperature, s a l i n i t y , and water motion as possible determinants of l o c a l d i s t r i b u t i o n s of.long and short stipe forms of L_. groenlandica and L„ saccharina, and (4) an evaluation of the roles of temperature, s a l i n i t y , exposure, and submarine illumination in determining the v e r t i c a l d i s t r i b u t i o n of L. saccharina. These studies were made from 1961 to , 1965. Ten species of Laminaria are recognized for the north-east P a c i f i c : L. groenlandica Rosenv.; L. f a r l o w i i Setchell; L. saccharina (L.). Lamour.; L. s e t c h e l l i i S i l v a ; L. dentigera Kjellman; L. longlpes Bory; L. s i n c l a i r i i (Harvey ex Hooker f. et_ Harvey) Parlow, Anderson et_ Eaton; L. ephemera Setchell; L. yezoensis Miyabe; and L. complanata (Setchell et Gardner) Set c h e l l . Laminaria cordata,Dawson i s considered conspecific with L. saccharina, L. personata Setchell and Gardner i s regarded conspecific with L. yezoensis, and L. platymeris De l a Pyl. (sensu Setchell and Gardner) i s considered con-s p e c i f i c with L. groenlandica. Pour forms of L. groe;nlandica are recognized for the northeast P a c i f i c . These forms are i i i not considered as legitimate taxonomic e n t i t i e s but are dis-tinguished merely to provide a means of f a c i l i t a t i n g discussion. The known habitat requirements for a l l ten species were broad-ened and the known d i s t r i b u t i o n s of a l l species, excepting L. groenlandica, were extended. Laminaria saccharina and L. groenlandica produced s o r i in the late spring and winter. New sporophytes of L. groenlandica appeared throughout the year, whereas those of L. saccharina appeared i n . l a t e winter and early f a l l . Depending upon culture conditions, two morphologically d i s t i n c t forms of gametophytes were produced by both species: large gametophytes were produced in conditions of high temperature and low s a l i n i t y ; and small gametophytes i n conditions of low temperature and high s a l i n i t y . Abnormal sporophytes were observed under conditions conducive to formation of large gametophytes. Patterns of growth f o r the blades of the two species were e s s e n t i a l l y the same.. The growth rate decreased with increase in distance from the blade base, and the position.of greatest longitudinal growth coincided with the position of greatest l a t e r a l growth. The d i s t r i b u t i o n s of L. saccharina and the two forms of L. groenlandica about Vancouver. Island were correlated with temperature, s a l i n i t y , and water motion. The two forms of L..,groenlandica were absent from areas of high temperature and low s a l i n i t y ; L. saccharina was absent from areas sub-jected to surf. These f i e l d conclusions were subjected to laboratory and f i e l d , t e s t s involving gametophytes i v and sporophytes of both species. The d i s t r i b u t i o n s of the two forms of L. groenlandica can be explained on the basis of temperature and s a l i n i t y . d i s t r i b u t i o n s . Both forms require low temperature and high s a l i n i t y for s u r v i v a l . Laminaria saccharina has a wide range of tolerance to temperature and s a l i n i t y . Surf appears to be the agent c o n t r o l l i n g the d i s t r i b u t i o n . o f t h i s species. The upper l i m i t s of L. saccharina, as observed in Burrard Inlet, are thought to-be d i r e c t l y related, to a i r temperature and i n s o l a t i o n and i n d i r e c t l y related to t i d a l c h a r a c t e r i s t i c s . The lower l i m i t s appear to r e f l e c t the com-pensation depth of t h i s species. V TABLE OP CONTENTS ' Page ABSTRACT i i LIST OP TABLES .. . . . .. . . ..... v i i l LIST OP FIGURES ix ACKNOWLEDGEMENTS . . . xvi INTRODUCTION '.. .. 1 GENERAL METHODS AND MATERIALS 4 F i e l d Methods and Materials . 4 Culture F a c i l i t i e s and Methods . 5 Evaluation of Plant Response 7 Oceanographic and Meteorologic Data 9 TAXONOMY AND.DISTRIBUTION OF NORTHEAST PACIFIC SPECIES OF LAMINARIA 12 C l a s s i f i c a t i o n . ... . . . . . . 13 Generic Diagnosis. 13 Key to the Species of Laminaria of the Northeast Pa c i f i c . . . . 13 Description of Species .14 Laminaria longipes . . . • . . . . . . . . .... . . 14 Laminaria s i n c l a i r i i .... .... 15 Laminaria ephemera 16 Laminaria yezoensis . . . 17 Laminaria groenlandica : 18 Laminaria saccharina . . . . 22 Laminaria f a r l o w i i 24 Laminaria complanata . 25 Laminaria s e t c h e l l i i . 26 y i Laminaria dentlgera 2 7 LIFE HISTORIES AND GROWTH PATTERNS OF LAMINARIA GROENLANDICA LONG AND SHORT STIPE FORMS AND \" LAMINARIA SACCHARINA 2 9 L i f e H istories of. Laminaria groenlandica. Long and Stipe Forms 2 9 Gametophyte and Microscopic Sporophyte Phases of • Laminaria groenlandica Long Stipe Form . . . . . . . . 3 1 . Growth Patterns of Laminaria groenlandica Long and Short S t i p e Forms 3 2 In S i t u Growth Rates.of Laminaria groenlandica Long and Short Stipe Forms . . 3 3 Transplant Studies of. Laminaria groenlandica Long and Short Stipe Forms 3 4 L i f e History of Laminaria saccharina . . . . . . . . . 3 6 Gametophyte and Microscopic Sporophyte Phases of Laminaria saccharina 3 6 Growth Pattern of Laminaria saccharina 3 8 Growth Rates of In Situ Laminaria' saccharina . . . . . . 3 8 LOCAL DIST IBUTIONS OF LAMINARIA SACCHARINA ANDLAMINARIA GROENLANDICA LONG AND SHORT STIPE FORMS AS RELATED TO TEMPERATURE/ SALINITY, AND WATER MOTION . . 40 EXPERIMENTAL EVALUATION OF THE EFFECTS OF VARIOUS OCEANOGRAPHIC FACTORS ON LAMINARIA SACCHARINA AND ; TWO FORMS OF'LAMINARIA GROENLANDICA . . . 4 4 Whole Plant Response of Laminaria saccharina Sporo-phytes to Various Temperature and S a l i n i t y Situations . . 4 4 Whole Plant Response of Laminaria saccharina Sporophytes to Various Temperatures at Constant S a l i n i t y and to Various S a l i n i t i e s at-Constant Temperature 4 5 Photosynthetic and Respiratory Response of Laminaria saccharina and Laminaria groenlandica Long Stipe Form Sporophytes to Various Temperature and S a l i n i t y Situations . . . . 4 6 v i i Responses of Laminaria saccharina and Laminaria groenlandica Long Stipe Form Gametophytes to Various Temperature and S a l i n i t y Situations . . . . 4.7 Photosynthetic and Respiratory Response of Laminaria saccharina and of Laminaria groenlandica Long .Stipe Form Sporophytes to Seawater of Different Origins . . 49 Responses of Laminaria saccharina and Laminaria groenlandica Long Stipe Form Gametophytes to Seawater of Different Origins . . . . . 49 Transplantation Studies 50 STUDIES. ON THE VERTICAL ' DISTRIBUTION OF.. LAMINARIA SACCHARINA 52 The V e r t i c a l D i s t r i b u t i o n of Laminaria saccharina as Related to Possible Determining Factors . . . . 52 Experimental Evaluation of some Factors as Possible Determinants of D i s t r i b u t i o n of Laminaria saccharina ... . . . 53 Temperature and S a l i n i t y . . . . . . 53 Submarine Illumination 54 DISCUSSION AND CONCLUSIONS 56 Taxonomy and Dis t r i b u t i o n s 56 L i f e History and Morphology . 59 Local D i s t r i b u t i o n in Relation to Oceanographic Factors 63 V e r t i c a l D i s t r i b u t i o n of ! Laminaria saccharina . . . . . . . 67 SUMMARY . 70 BIBLIOGRAPHY . 74 APPENDIX I. Voucher specimens of Laminaria 80 APPENDIX I I . Observed, specimens of Particulartaxonomic' Significance 89 v i i i LIST GP TABLES Table Page •I. Longitudinal growth rates in s i t u of Laminaria groenlandica .. „ . . . . 91 I I . Stipe length/blade width as determined i n s i t u for Laminaria groenlandica \". . \"'92 I I I . Stipe length/blade width as determined from transplants of Laminaria groenlandica 93 IV. Values of temperature and s a l i n i t y at three depths i n Howe Sound, near Keats Island . . . . . . . 94 V> I n i t i a l tests on the s a l i n i t y and temperature tolerance of Laminaria saccharina 95 VI. Net photosynthesis, r e s p i r a t i o n , and net photo-synthesis/respiration r a t i o s of Laminaria saccharina and Laminaria groenlandica i n various temperature-salinity, situations . . . 96 VII. Sporophyte production by gametophytes of Laminaria saccharina and Laminaria groenlandica in various temperature-salinity situations 97 ix LIST OP FIGURES Figure Page 1. Spectral curves of Westinghouse F 4 0 Blue, Westinghouse F40HE37 Green and one Westinghouse Blue to two Westinghouse Green fluorescent tubes 98 2. Spectral curve of Sylvania Cool White fluorescent tubes .99 3 . Percent transparency of stained glass used in f i l t e r i n g l i g h t and percent transparency per meter f o r coastal waters 99 4. Relative s e n s i t i v i t y of the Photovolt Electronic Photometer Model. 501-M 99 5. Habit of Laminaria longipes . 100 6. Habit of -Laminaria S i n c l a i r ! ! / 1 exposed form . . . 100 7. Habit of Laminaria s i n c l a i r i i , sheltered form .. . 100 8. Habit of Laminaria ephemera, exposed form ... . . 100 9. Habit of Laminaria yezoensis . . . . 101 10. Discoid holdfast of Laminaria yezoensis . . . . . 101 11. Habit of Laminaria groenlandica shade form .. ... 101 12. Regeneration of Laminaria groenlandica shade form 101 13. Habit of Laminaria groenlandica short stipe form, bulla t e blade . . . . . . . . . . . . . . . . . . 102 14. Habit of Laminaria groenlandica. long stipe form . 102 15. Habit of - Laminaria groenlandica short stipe form, abullate blade. . 102 16. Habit of Laminaria groenlandica f l a t stipe form . 102 17. Habit of Laminaria saccharina, bullate blade . . 103 18. Habit of Laminaria saccharina, abullate blade . . 103 1.9. Habit of Laminaria f a r l o w i i 103 20. Habit of Laminaria complanata . . ....... . . . . 103 X Figure Page 21. Habit of Laminaria s e t c h e l l i i , Glacier Point form 104 22. Habit of Laminaria s e t c h e l l i i , Botany Beach form . 104 23 . Regeneration of Laminaria s e t c h e l l i i . . .... . . 104 24. Habit of Laminaria dentigera 104 25. Laminaria longipes, cross section of blade ...... . 105 26. Laminaria s i n c l a i r i i , cross section of blade . . . 105 27. Laminaria ephemera, cross section of blade . . . . 105 28. Laminaria yezoensis, cross section of blade .... 105 29. Laminaria groenlandica shade form, cross section of blade .......... . . . . . . . . 105 '•'30. Laminaria groenlandica long stipe form, cross section of blade 105 31 . Laminaria groenlandica f l a t stipe form, cross section of blade 105 32. Laminaria saccharina, cross section of blade .. . . 105 33. Laminaria f a r l o w i i , cross section of blade ... . . 105 34. Laminaria complanata, cross section of blade . . . 105 35. Laminaria s e t c h e l l i i , cross section of blade ... . 105 36. Laminaria dentigera, cross section of blade . . . 105 37. Laminaria longipes, cross section of stipe . . . 106 38. Laminaria s i n c l a i r i i , cross section of stipe . . . 106 39. Laminaria ephemera, cross section of stipe . . . . . 106 40. Laminaria yezoensis, cross section of stipe . . . 106 41. Laminaria groenlandica shade form, cross section of stipe 106 42. Laminaria. groenlandica short stipe form, cross section of stipe 106 4 3 . Laminaria groenlandica long stipe form, cross section of stipe 106 . x i Figure Page 44. Laminaria groenlandica f l a t stipe form, cross section of stipe 107 45. Laminaria saccharina, cross section of stipe . . . 107 46. Laminaria f a r l o w l i , cross section of stipe . . . . 107 47. Laminaria complanata, cross section of stipe . . . 107 48. Laminaria s e t c h e l l i i , cross section of stipe . .. . 107 49. Laminaria dentigera, cross section of stipe . . . . 107 50. Northeast P a c i f i c d i s t r i b u t i o n of Laminaria longipes and Laminaria dentigera .... . . . . . . . 108 •51. Northeast P a c i f i c , d i s t r i b u t i o n , north of 48° N l a t i -tude, of Laminaria s i n c l a i r i i , Laminaria ephemera, Laminaria f a r l o w i i , and Laminaria complanata . .. . 109 52. Northeast P a c i f i c d i s t r i b u t i o n , north of 48° N l a t i -tude, of Laminaria s e t c h e l l i i and Laminaria yezoensis 110 , 53._ Northeast P a c i f i c d i s t r i b u t i o n , north of 48° N l a t i -tude, of Laminaria saccharina and Laminaria groenlandica . . . . . . . . . . . I l l 54. L i f e h i s t o r y of Laminaria groenlandica . . . . . . 112 55. Gametophyte and.microscopic sporophyte phases of Laminaria groenlandica . 113 56. D i s t r i b u t i o n of longitudinal growth, i n culture, of the blade of Laminaria'groenlandica short stipe form . . . . . .. . . . . . . . . . .\"'.. . .... 114 57. D i s t r i b u t i o n of; longitudinal growth, i n culture, of the blade of'Laminaria groenlandica long stipe form . . . . . . . 114 58. D i s t r i b u t i o n of•longitudinal growth, in s i t u , of Laminaria groenlandica long and short stipe form . 114 59. • D i s t r i b u t i o n of l a t e r a l growth, i n culture, of the blade of Laminaria groenlandica short stipe , form 115 60. D i s t r i b u t i o n of l a t e r a l growth, i n culture, of the blade of Laminaria groenlandica. long stipe form . . 115 61 . D i s t r i b u t i o n of- l a t e r a l growth, in s i t u , of the blades of Laminaria groenlandica long.and short stipe forms . . . . . . . . 115 x l i Figure Page 62. L i f e history of Laminaria saccharina ..'. . . . ... 116 6 3 . Gametophyte and microscopic sporophyte phasis of Laminaria saccharina ....... 117 64. D i s t r i b u t i o n of longitudinal growth, in culture, of the blade of Laminaria saccharina .' . 118 6 5 . D i s t r i b u t i o n of, longitudinal growth, i n s i t u , of the.blade of Laminaria saccharina ..... . . . . . 118 66. D i s t r i b u t i o n of l a t e r a l growth, in culture, of the blade of Laminaria saccharina . . . . . . . . 119 67. D i s t r i b u t i o n of l a t e r a l growth, in s i t u , of the blade of Laminaria saccharina . . . . . . 119 68. Longitudinal growth rates and theore t i c a l lengths of the blades of Laminaria saccharina 120 69. Seasonal v a r i a t i o n in temperature and s a l i n i t y at three depths for Sheringham Point and Gordon Head 121 70. Seasonal v a r i a t i o n in temperature and s a l i n i t y at three depths for Burrard Inlet near Stanley Park 122 .'71. Seasonal v a r i a t i o n in surface temperature and s a l i n i t y for several stations about Vancouver Island . . . . . . . . . . ... . 123 72. Total range of seasonal s a l i n i t y and temperature v a r i a t i o n summarized from Figure '71 '. . 124 73. Percent increase in blade surface per day for Laminaria saccharina af t e r 15 days at various temperatures . . 125 74. Percent increase in blade surface per day f o r Laminaria saccharina a f t e r 15 days at-various s a l i n i t i e s . . . . . . . . . . . . . . . . 125 75. Sporophyte growth of Laminaria saccharina and Laminaria .groenlandica in'seawaters from d i f f e r e n t areas .....' 126 76. Sporophyte growth of Laminaria saccharina in seawaters d i f f e r e n t ' o r i g i n s and of d i f f e r e n t s a l i n i t i e s . . . . . . . . . . . . . . . . . . . . . . 126 x i i i Figure Page 77. Upper v e r t i c a l l i m i t s of Laminaria saccharina ... . 127 78. Some t i d a l features for Burrard Inlet . . . . . . . 128 79. A i r temperature for K i t s i l a n o , Vancouver . . . . . 129 80. Daylength as calculated f o r 49°N . . 130 81. Hours bright sunshine observed f o r the Univ. B r i t i s h Columbia, Vancouver.. . . . . . . . . . . . 130 82. Approximate mean d a i l y i n s o l a t i o n f or l i g h t i n the v i s i b l e range as observed for the Univ. B r i t i s h Columbia 130 83. Approximate submarine l i g h t i n t e n s i t y at three depths i n Burrard Inlet near Stanley Park ... . . . 131 84. Apparent photosynthesis of Laminaria saccharina at d i f f e r e n t l i g h t i n t e n s i t i e s . 132 85. Whole plant.response of Laminaria'saccharina to different, l i g h t i n t e n s i t i e s . ........ . . . . . 132 86. Mean monthly seawater temperature and s a l i n i t y at Departure Bay and Pine Island ... 133 xiv ACKNOWLEDGEMENTS I sincerely wish to express my gratitude to Dr. R.P. Scagel for h i s d i r e c t i o n , advice and f i n a n c i a l support, provided by a research grant from the Defence Research Board of Canada (DRB 9520-14); to Dr. K.M. Cole, Dr. G.L. Pickard and Dr. T.M.C. Taylor for t h e i r assistance throughout the course of t h i s study; to Dr. G.E. Rouse and Dr. E.B. Tregunna for t h e i r assistance i n preparation of t h i s manuscript; to Dr. M.A. Newman, and his s t a f f at the Vancouver Public Aquarium for making available research space and f a c i l i t i e s ; to Mr. J. Green and Mr. R. D r i s k i l l for making subtidal c o l l e c t i o n s ; and to the s t a f f of the following herbaria: Allan Hancock Foundation, Friday Harbor Laboratories, and the University of C a l i f o r n i a , Berkeley, for making available specimens. I wish to record my indebtedness to the Department of Biology and Botany and the Institute of Oceanography, University of B r i t i s h Columbia for continued help throughout a l l phases of t h i s study; to the National Research Council of Canada and the University of B r i t i s h Columbia for scholarships for the years 1962-1965. F i n a l l y , I would l i k e to thank Mrs. Louis D. Druehl f o r her support and assistance i n t h i s study. \"Not everyone has peered Into clear rock pools at the hour of dawn, when the tide i s at i t s lowest ebb. and recognized in the dusky shadowy forms of the young kelp, l i v i n g creatures belonging to the fa r distant past. Only the fortunate can-know the true meaning of the Greek work phaios and f u l l y appreciate i t s beauty.\" Josephine Tilden, 1935 INTRODUCTION The brown a l g a l genus, Laminaria Lamouroux, i s the most frequently encountered member of the Laminarlal.es i n north-ern temperate coastal waters. Along much of the northeast P a c i f i c coast, p a r t i c u l a r l y north of 48°N, l a t i t u d e , i t i s the dominant lower i n t e r t i d a l plant. Setchell and Gardner (1925) recognized 11 species and 7 forms of Laminaria i n the northeast P a c i f i c , and t h e i r t r e a t i s e i s .the most recent com-prehensive taxonomic treatment of t h i s genus i n t h i s area. Since then S i l v a (1957) and Widdowson (1959) have contributed to an understanding of the taxonomy of i n d i v i d u a l species recognized by Setchell and Gardner; Dawson (1950) has recog-nized a new species, L. cordata Dawson. B i o l o g i c a l studies on Laminaria \"in the northeast P a c i f i c have been c h i e f l y morphological (Setchell, 1905; Griggs, 1906; P a l l i s , 1916; and Myers, 1925). The present investigation consists of two parts. The f i r s t , a c r i t i c a l evaluation of the taxonomy and species d i s t r i b u t i o n of Laminaria i n the northeast P a c i f i c , was based on extensive c o l l e c t i o n s available for the coasts of B r i t i s h Columbia and Alaska (Scagel, i n herb., UBC). These were supplemented.by c o l l e c t i o n s • o f the author from northern Washington to the southern extreme of Alaska. For the remainder of the .northeast P a c i f i c , , the study was based on herbarium material provided by the University of Washington (WTU), University of C a l i f o r n i a , Berkeley (UC), and the 2 Allan Hancock Foundation (AHFA). The second part i s an eval-uation of the e f f e c t s of ( l ) temperature, s a l i n i t y , and water motion In determining the horizontal d i s t r i b u t i o n s of Laminaria saccharina (L.) Lamour. and two forms of L. groenlandica Rosenv. about Vancouver Island, B r i t i s h Columbia; and ( 2 ) temperature, s a l i n i t y , exposure, and sub-marine illumination i n determining the v e r t i c a l d i s t r i b u t i o n of L. saccharina i n Burrard Inlet, near Vancouver, B r i t i s h Columbia. To- accomplish the second part, the patterns of growth, growth rates, and, l i f e h i s t o r i e s of L. saccharina and L. groenlandica were determined. An understanding of these b i o l o g i c a l aspects of the plants was necessary to evaluate plant response to various environmental conditions. The d i s t r i b u t i o n s of these two species were subsequently correlated.with the above mentioned environmental parameters. Where possible the established correlations were subjected to laboratory and.field tests. Several features make species of - Laminaria ideal for an ecological study of the type described above. The ver-t i c a l d i s t r i b u t i o n s of the species extend from the subtidal to the lower i n t e r t i d a l regions. This v e r t i c a l positioning minimizes the influence of environmental factors operative during periods of emergence, and supports the hypothesis presented here that horizontal d i s t r i b u t i o n of Laminaria i s controlled primarily by ©ceanographic factors rather than meteorological conditions. Since the species studied are perennial, t h e i r absence from any l o c a l i t y r e f l e c t s the unfavorableness of the environmental conditions in that area for the establishment and maintenance of a population. Laminaria saccharina and the two forms of L. groenlandica studied are e a s i l y recognized i n the f i e l d . The ease with which the sporophytes and gametophytes of these two species can be cultured makes them id e a l for _in v i t r o experimentation. The taxonomy and d i s t r i b u t i o n of Laminaria was studied throughout i t s known northeast.Pacific range--California to Alaska. The d i s t r i b u t i o n s of Laminaria saccharina and the two forms of L. groenlandica were correlated with oceanographic factors at several stations about Vancouver Island. JCn s i t u studies on L. saccharina were made i n Burrard Inlet (49°l8'9\"N 123°7'30\"w). near Stanley Park, Vancouver, B r i t i s h Columbia; on L. groenlandica long stipe form at Glacier Point ( 4 8 ° 2 3 ' W N 1 2 3 0 5 9 ' l o \" w ), Vancouver Island, B r i t i s h Columbia; and on L. groenlandica short stipe form at the Ogden Breakwater (48 ° 2 4 ' 5O\"N 123°23'3o\"w), V i c t o r i a , Vancouver Island, B r i t i s h Columbia. 4 GENERAL METHODS AND MATERIALS The methods and materials described below apply generally to a l l aspects of thi s study. Sp e c i f i c information describ-ing experimental conditions i s discussed along;with the res u l t s of the i n d i v i d u a l experiments. F i e l d Methods To f a c i l i t a t e r e l o c a t i n g plants employed i n f i e l d studies two types of plant markers were used. When i t was necessary to relocate a s p e c i f i c plant, the plant was tagged with a coded p l a s t i c l a b e l secured to the stipe with a nylon l i n e . When i t was not necessary to Identify i n d i v i d u a l s s p e c i f i c a l l y , the plants were tagged with red or blue nylon tape t i e d to the stip e . Transport of l i v i n g material was accomplished by wrapping the plants in newspaper saturated in seawater and storing i n an iced chest. The period from c o l l e c t i o n to establishment i n a new environment never exceeded 24 hours. Plants transported i n t h i s manner showed no signs of injury. When placed i n the new environment, the transplanted plants were attached to rocks by rubber bands overlapping the haptera. The v e r t i c a l l i m i t s of d i s t r i b u t i o n for L. saccharina were determined by measuring the v e r t i c a l distance from the plants to the water surface and from the water surface to a bench, mark. The lower l i m i t s were measured with a metered l i n e , oriented by a SCUBA diver, and the upper l i m i t s 5 with a meter r u l e . The position of the bench mark i n r e l a t i o n to various t i d a l heights was found by measuring the v e r t i c a l distance between the bench mark and the water surface and then r e l a t i n g t h i s measurement to the t i d a l height predicted for the time of measurement (Anon., 1962). Culture F a c i l i t i e s and Methods Four d i f f e r e n t f a c i l i t i e s were employed i n culturing Laminaria. (1) P l a s t i c ( l u c i t e ) aquaria. Five p l a s t i c aquaria were employed as culture chambers f o r large plants and as incubators for small sporophyte and gametophyte cultures. These aquaria measured 45 cm,long x 25 cm wide x 60 cm high. When culturing large plants d i r e c t l y in these aquaria, 40 to 50 1 of seawater were used. When incubating culture dishes the water l e v e l was maintained immediately below the l i p of the dishes. The. culture dishes used for small sporophytes were 1 l i t r e glass covered dishes. Those used f o r the gametophytes were 250 ml glass covered dishes. The aquaria were kept i n a 7°C constant temperature room. Temperatures higher than 7°C were maintained by small aquarium heaters equipped with thermostats. (2) Constant temperature water bath table. This table consisted of a 20 cm deep galvanized iron tray measuring 1.1 m x 2.2 m. Temperature was controlled by means of a r e f r i g e r a t i o n c o l l f l u s h ..with the bottom of the tray but external to the water bath. This table was used as an incubator for BOD (B i o l o g i c a l Oxygen Demand) bottles employed 6 i n p h ysiological studies. (3) Culture tank. A large tank ( 3 . 8 m. long x 0 .8 m wide x 1.0 m high) was provided .by the Vancouver Public Aquarium at Stanley Park. This culture tank was connected to a c i r c u l a t i n g seawater system which was augmented with fresh seawater every day. The replacement time of seawater in the culture tank was approximately 7 hours. Temperature was controlled by passing the incoming water through a refrige r a t e d c o i l . (4) Constant temperature rooms. Constant temperature rooms of 5, 10, 15, and 20°C were employed. These are main-tained by the Department of Botany, University of B r i t i s h Columbia. A l l experiments employing, culture f a c i l i t i e s ( l ) , (2), and (3). were illuminated with two green fluorescent tubes (Westinghouse F40HE 37) to each blue .fluorescent tube (Westinghouse F40 Blue). The spectrum of l i g h t r e s u l t i n g from the above combination i s shown i n Figure 1. The l i g h t transmission curve as observed i n inshore waters (after Jerlov, 1951) i s shown i n Figure 3b. Experiments conducted i n the constant temperature rooms were illuminated with Sylvania fluorescent, tubes (FT12/CW) f i l t e r e d through stained glass (Fig. 2, 3 a ) . A l l l i g h t measurements were made with a Photovolt Elect r o n i c Photometer (Photovolt Corporation, New York 16, N.Y., Model 501-M). The r e l a t i v e s e n s i t i v i t y of t h i s l i g h t meter at various wave lengths i s i l l u s t r a t e d i n Figure 4.. The seawater employed i n culture studies was usually obtained from the same area from which the plants were c o l -lected. One exception was a study on the response of plants to waters collected from d i f f e r e n t regions. No nutrients were added to the seawater in any of the culture experiments. F i l t r a t i o n of the seawater was effected by passing the water through a column of packed glass wool 2 to 3 times for sporophyte culture, and 6 to 7 times for gametophyte culture. The f i l t e r i n g of seawater for gametophyte culture usually resulted i n u n i a l g a l cultures. Contaminated cultures were discarded. S a l i n i t y adjustment was achieved by heating seawater at 30 to 35°C u n t i l a concentration of 35 to 50$» was reached. Subsequently, concentrated seawater was added to a larger volume of raw seawater and the s a l i n i t y was determined using an inductively coupled salinometer (Model 601, MK III, Auto-Lab Industries, Sydney, A u s t r a l i a ) . Lower s a l i n i t i e s were obtained by d i l u t i n g t h i s seawater mixture with d i s -t i l l e d water. Evaluation of Plant Response (l) Responses of entire sporophytes to conditions of l i g h t , temperature, and s a l i n i t y were determined by measuring the surface area of plants at the beginning, during, and at the end of the experiment. The surface area was determined by measuring the outline of the blade with.a compensating polar planimeter (Keuffel &•. Esser Co., Germany, Model 4236). The blade outline was obtained by either a paper tracing or 8 a photograph of the blade against a metered g r i d . (2) Responses of discs of sporophyte blades to condi-tions of temperature, l i g h t , s a l i n i t y , and.different water origins were determined by measuring apparent photosynthesis and r e s p i r a t i o n , and by cal c u l a t i n g the net photosynthesis and the r a t i o of net photosynthesis to r e s p i r a t i o n i n the following manner. (a) Net photosynthesis was determined by the equation: Net PhotosynthesiSQ^ = Apparent Photosynthesis,^ + RespirationQ where Apparent Photosynthesis,-^ i s determined by' subtracting the i n i t i a l 0 2 content of the culture medium from the 0 2 content present a f t e r a period of exposure t o . l i g h t . RespirationQ^ i s the difference between the i n i t i a l 0 2 content and the 0 2 content a f t e r a period of darkness. The units employed were ynl 0 2 evolved or utilized/cm /hr. (b) . The Net Photosynthesis to Respiration r a t i o was determined by d i v i d i n g the rate of Net Photosynthesis by the re s p i r a t i o n rate.. The above measurements were made on discs having an area of 33 square cm. These were cut d i s t a l l y to the basal.15 cm of the blade i n areas of uniform color and free from apparent injury. After cutting, the discs employed in temperature and s a l i n i t y studies were preconditioned for f i v e days i n the situa t i o n to be tested. In studies on l i g h t the discs were not preconditioned but kept i n darkness f o r 24 hours p r i o r to t e s t i n g . At the time of measurement, one disc was placed i n each 300 ml BOD b o t t l e . Five discs were tested in each temperature and s a l i n i t y s i t u a t i o n , and six discs 9 were employed -in each l i g h t i n t e n s i t y tested. The BOD bottles were then f i l l e d with f i l t e r e d seawater of a known oxygen concentration ,and of the desired temperature and s a l i n i t y . These bottles were then placed i n l i g h t for 2 hours. At the end of the period.150 ml aliquots of the seawater were siphoned from the 300 ml BOD bottles to . 150 ml BOD bottles and the oxygen content.was determined by the Winkler Method as described by Strickland and Parsons ( i 9 6 0 ). The seawater remaining with the plant discs in the 300 ml BOD bottles was then replaced with new seawater of known oxygen content and the bottles were placed i n darkness for 4 hours. At the end of the dark period the oxygen content was determined. Control BOD bottles were employed i n each experiment to detect changes in oxygen concentration due to microbial a c t i v i t y . The con-t r o l bottles d i f f e r e d from the experimental bottles only i n that they did not contain plant.discs. (3) . Responses of gametophytes to various temperature, s a l i n i t y , and l i g h t situations were determined by.the a b i l i t y of the plants to produce sporophytes. (4) !Responses of gametophytes to samples of seawater from different.areas were determined by measuring the length of the r e s u l t i n g sporophytes a f t e r a period of growth. Measurements were made with a calibrated ocular micrometer. Oceanographic and Meteorologic Data With the exception of oceanographic data from Burrard Inlet a l l data employed in t h i s study were available i n published form. Where pertinent references to these data 10 are provided i n the text. Monthly determinations of the temperature and s a l i n i t y structure of the water column i n Burrard Inlet were made from June, 1963 to May, 1964. An inductive salinometer (Model R S 5 2 , Industrial Instruments Inc., New Jersey) was used. Approximate submarine l i g h t i n t e n s i t i e s for three depths in Burrard Inlet were determined using the following expres-sion ( a f t e r Strickland, 1 9 5 7 ) : I z = ant i l o g ( l o g 1 0 I Q - Kz), where I i s the l i g h t of the v i s i b l e range ( 3 8 0 0 - 7 2 0 0 8) entering the water, K the extinction c o e f f i c i e n t of the water and z the depth investigated. The manner in,which the values for these three terms was determined i s as follows: ( I Q ) ' I D = O .96 ( l ± - 0 . 5 ) , where I ^ . i s the monthly mean incident l i g h t as recorded from 1 2 : 0 0 noon to. 1 :00 PMby an Eppley Pyrheliometer at the o University of B r i t i s h Columbia for the wave band 3 0 0 0 - 5 0 , 0 0 0 A (Anon., 1 9 6 4 ) . The value 0 . 5 i s a conversion factor for determination of the percentage of 1^ i n the v i s i b l e range. This value assumes clear sky and unobscured sun,.with sky l i g h t contributing one t h i r d of the t o t a l r a d i a t i o n (Strickland, 1957).. The value O .96 corrects f o r 4$ r e f l e c t i o n of the incident l i g h t from the water surface (Holmes, 1 9 5 7 ) . (K) .• The extinction c o e f f i c i e n t (K) i s derived from the equation: where D i s the secchi disc depth expressed i n meters (Strickland, 1957). The mean secchi disc depth f o r Burrard Inlet as deter-mined from monthly readings (June, 1963 to May, 1964) taken at high tide was 3-2 m. This value was used i n the l a s t equation, (z) To rel a t e the distance that, l i g h t must pass from the water surface to the three depths Investigated, i t was necessary to compensate f o r t i d a l f l u c t u a t i o n s . This compensation was achieved by employing the monthly mean t i d a l height as observed at 12:30 PM. The resultant values for submarine illumination,are approximations and are employed here to i l l u s t r a t e the seasonal trend (Fig. 83). 12 TAXONOMY AND DISTRIBUTION OP NORTHEAST PACIFIC SPECIES OF. LAMINARIA The following taxonomic and d i s t r i b u t i o n a l treatment of taxa of Laminaria occurring i n the northeast P a c i f i c consti-tutes the f i r s t c r i t i c a l review of t h i s large and important genus since Setchel and Gardner's study i n 1925• Extensive i n t e r t i d a l c o l l e c t i o n s and a few subtidal c o l l e c t i o n s i n B r i t i s h Columbia and Alaska have altered previously reported d i s t r i b u t i o n patterns. A l l d i s t r i b u t i o n a l records north of 48°N la t i t u d e have been taken from c o l l e c t i o n s available i n the Phycological Herbarium, University of B r i t i s h Columbia, and the Friday Harbor Laboratory Herbarium, University of Washington. D i s t r i b u t i o n a l data for c o l l e c t i o n s made north of 48°N.lati-tude are presented i n Figures 50 to 53. Further d i s t r i b u t -ional data are provided i n Appendix I. The l i m i t s of Laminaria, south of 48°N l a t i t u d e , are taken from f l o r i s t i c studies of Smith ( l 9 4 4),Doty (1947), and Dawson (1961) . A, l i s t i n g of some of the plants studied for t h e i r p a r t i c u l a r taxonomic significance i s given i n Appendix I I . The descriptions and phenological data of taxa of north-east P a c i f i c Laminaria were derived from plants c o l l e c t e d i n northern Washington, B r i t i s h Columbia,. and Alaska. Where possible, species are distinguished on the basis of q u a l i t a t i v e characters. The use of size measurements has been, minimized because of the considerable v a r i a t i o n encoun-tered. C l a s s i f i c a t i o n D i v i s i o n : Phaeophyta Glass: Phaeophyceae Order: Laminariales Family: Laminariaceae Genus: Laminaria Lamouroux, 1813 Generic Diagnosis Holdfast of branched haptera or a continuous disc. Stipe simple, s o l i d or hollow, complanate or terete, with or without mucilage ducts. Transition region between stipe and blade undivided. Blade entire or l o n g i t u d i n a l l y dissected, with or without bullae, with or without mucilage ducts. Sori occurring on blade as continuous or iso l a t e d patches. Key to the Species of Laminaria of the Northeast P a c i f i c 1. Holdfast d i s c o i d 2 1. Holdfast of d i s t i n c t , branched haptera 3 2. Mucilage ducts absent from blade.. L. ephemera, p. 16. 2 . Mucilage ducts present i n blade... L. yezoensis, p. IJ. 3. Numerous stipes, each terminated by a single blade a r i s i n g from an extensive holdfast 4 3. One stipe, terminated by a single blade a r i s i n g from.a holdfast 5 4. Mucilage ducts present i n s t i p e . . . L. s i n c l a i r i i , p. 15 4. Mucilage ducts absent from sti p e . . L. longipes, p. 14. . 14 5. Mucilage ducts absent from stipe 6 5. Mucilage ducts present i n stipe , 8 6. Stipe complanate, usually longer than 30 cm . . L. complanata, p. 25 . 6. Stipe terete, usually shorter than 20 cm. 7 7. Bullae, when present, i n two rows oriented p a r a l l e l . t o the margin of. the blade. L. saccharina, p. 22. 7. Bullae covering entire blade........ L. f a r l o w i i , p. 24. 8. Stipe terete throughout; blade deeply dissected i n t o many narrow, uniform segments. 9 8. Stipe often s l i g h t l y to extensively complanate; blade, when dissected, of few segments varying in width and: length . . L. groenlandica, p.. 18. 9. Mucilage ducts positioned near surface of stipe • L. dentigera, p. 27. 9. Mucilage ducts positioned i n the. mid-cortical region of the stipe L. s e t c h e l l i i , p. 26. Description of Species Laminaria longipes Bory Setchell and Gardner, 1925, p. 597. Dawson, 1 9 6 l , p. 396. Description. Pig. 5, 25, 37. Holdfast an extensive system of branched haptera, the haptera giving r i s e to.many stipes, each terminated by a single blade. Stipe usually less than 20 cm;long, mucilage ducts absent. Blade l i n e a r , cuneate, usually less than 5 cm wide, of variable length, mucilage ducts present. No plants observed with s o r i . 15 Northeast P a c i f i c D i s t r i b u t i o n . Pig. 50; App. I, Sect. A. Attu Island to Kenai Peninsula, Alaska. Known e a r l i e r only from the Bering Sea. Habitat. Growing on rocks i n the lower i n t e r t i d a l region, often i n muddy or sandy areas. Comments. Only herbarium specimens observed. Laminaria s i n c l a i r i i (Harvey ex Hooker f. _et Harvey), Parlow, Anderson et Eaton Setchell, 1905, P. 139-Setchell and Gardner, 1925, p. 598. Smith, 1944, .p. 135. Scagel, 1957, P. 98. S i l v a , 1957, P. 43 . Dawson, 1961, p. 396. Description. Pig. 6, 7, 26, 38. Sporophyte perennial from the stipe, of variable length and width. Holdfast composed of extensive, branched haptera, the haptera giving r i s e to, many stipes, each terminated by a single.blade. Stipe usually not exceeding 20 cm i n length, mucilage ducts present. Blade of i n t e r t i d a l plants of variable length, usually less than 5 cm wide, .mucilage ducts pre-sent. Blade of subtidal plants of variable length, up to 20 cm wide, with mucilage ducts. Plants with s o r i i n February. Northeast P a c i f i c D i s t r i b u t i o n . F i g . 51; App. I, Sect. B. Hope Island, B r i t i s h Columbia to Ventura County, C a l i f o r n i a . E a r l i e r known from southern B r i t i s h Columbia to 16 Ventura County, C a l i f o r n i a . Habitat. Growing on rocks in the lower i n t e r t i d a l region in areas of surf. Known only from the subtidal region i n sheltered areas. Laminaria ephemera Setchell Setchell, 1901, p. 121. Griggs, 1906, p. 247 (as Renfrewia parvula). Setchell and Gardner, 1925, p. 603. Smith, 1944, p. 136. Doty, 1947, P . 40. Scagel, 1957, p.- 95. Dawson, 1961, p. 396. Description. F i g . 8, 27, 39. Sporophytes annual, mucilage ducts wanting. Holdfast a small disc. Stipe terete, 1 5 + cm long in exposed i n t e r t i d a l regions, much shorter for subtidal plants. Blade l i n e a r , 6 + cm wide, of variable length, infrequently dissected. Plants with s o r i i n May and June. Northeast P a c i f i c D i s t r i b u t i o n . F i g . 51; App. I, Sect. C. Amphitrite Point, Vancouver Island, B r i t i s h Columbia to Monterey Peninsula, C a l i f o r n i a . E a r l i e r known from Port Renfrew, B r i t i s h Columbia to Monterey Peninsula, C a l i f o r n i a . Habitat. On rocks i n the lower i n t e r t i d a l and upper subtidal regions i n areas subjected to surf; r e s t r i c t e d to the subtidal region i n sheltered areas. 17 Laminaria yezoensis Miyabe Saunders, 1901, p. 429 (as L..solidungula), see comments on page 18. Miyabe, 1902 (English edition, 1957, p. 2 3 ) . Setchell and Gardner, 1924, p. 10 (as L. personata). Setchell and Gardner, 1925? p. 599 (as L. personata). Dawson, 1961, p. 396 (as L. personata). Description. Pig. 9, 10, 28, 40. Sporophytes up to 1 nr. long. Holdfast a scutate disc, often very expansive and united with other discs. Stipe terete, up to 40 cm.long, mucilage ducts absent. Blade of variable length, entire or lo n g i t u d i n a l l y dissected, mucilage ducts present. One plant with sorus collected from Queen Charlotte Islands i n July. Northeast P a c i f i c D i s t r i b u t i o n . Pig. 52; App. I, Sect. D. Adak Island, Alaska to Hope Island, B r i t i s h Columbia. Heretofore not known from the northeast P a c i f i c . Habitat. Growing on rocks i n exposed areas i n the lower i n t e r t i d a l region. No.subtidal data available. Comments. The plants observed displayed much varia-b i l i t y but were i n complete accord with Miyabe 1s description (1902). Setchell and Gardner, i n t h e i r o r i g i n a l description of L. personata S. & G. (1924) and i n a l a t e r discussion (1925) of t h i s species, make no mention of L. yezoensis. However, i t must be assumed that Setchell was aware of t h i s species since he discussed i t i n an e a r l i e r paper (1908). Prom comparison of Setchell and Gardner's description (1924) of L. personata and Miyabe's description (1902) of 18 L. yezoensis i t i s apparent that the two authors are describing the same species. The type specimen of L. personata (N.L. Gardner #3951,• UC#26649l) appears to be a juvenile form of L. yezoensis and. has a l l the diagnostic c h a r a c t e r i s t i c s of that species. The species L. personata S. & G. i s here reduced to synonomy under the older name L. yezoensis Miyabe. Saunders (1901) described L. solidungula J. Ag. from Yakutat Bay,•Kukak Bay, and Popof Island, Alaska. Laminaria solidungula, common to the northwest A t l a n t i c , i s distinguished from L. yezoensis by the presence of mucilage ducts i n i t s sti p e . Subsequent c o l l e c t i o n s (Scagel, i n herb.) made i n the Yakutat region s p e c i f i c a l l y and Alaska. generally have f a i l e d to rediscover L. solidungula. Possibly the ent i t y referred to L. solidungula by Saunders was Cymathere t r i p l i c a t a (Post, and Rupr.) J. Ag.. Cymathere t r i p l i c a t a i s found throughout the range collected by Saunders and i s similar to L. solidungula i n that it.has mucilage ducts i n the stipe and a well developed discoid holdfast. Laminaria groenlandica Rosenvinge Setchell and Gardner, 1925, p. 600 (as L. cuneifolia) and p. 605 (as L. platymeris). Doty, 1947, p. 39 (as L. c u n e i f o l i a ) . Scagel, 1957, P. 93 (as L. cuneif.olia) and p. 95 (as L. platymeris). Widdowson, 1959, P. 56 (as L. c u n e i f o l i a ) . Wilce, 1959,-P. 158 (as L. c u n e i f o l i a ) . 19 Wllce, I960,.p. 203. Dawson, 1961, p. 396 (as L. cun e l f o l i a and L. platymeris). Description. . Pig. 11, 12, 13, 14, 15, 16, 29, 30, 31, 4 l , 42, 43, 44. Sporophyte perennial from the stipe. Holdfast of many; branched haptera. Stipe varying in length from 1 to 60 cm, terete to complanate, mucilage ducts pre-sent. A l l plants producing s o r i i n winter; plants older than one year often producing s o r i i n early summer. Northeast P a c i f i c D i s t r i b u t i o n . ..Fig. 53; App. I, Sect. E, F, G. Attu Island, Alaska to Cape Blanco, Oregon. . Habitat. Growing on rocks i n the lower i n t e r t i d a l region and upper subtidal region i n areas exposed to consider-able water motion. Restricted to the subtidal region in sheltered, estuarine waters. Comments. Wilce (1959) did. not r e t a i n i n L. cu n e i f o l i a (= L. groenlandica) those plants from the northeast P a c i f i c referred to t h i s species by Setchell and Gardner (1925). Wilce's decision to remove the northeast P a c i f i c plants from L. c u n e i f o l i a was based on a comparative study of the color and texture of the plants, and of the size, shape, and arrangements of the surface c e l l s and the i r protoplasts. However, Wilce.did agree that the general shape of the north-east P a c i f i c plants and the d i s t r i b u t i o n of t h e i r mucilage ducts conforms with the A t l a n t i c L. groenlandica. I cannot agree with Wilce's choice of c h a r a c t e r i s t i c s ' fo r distinguishing the northeast P a c i f i c plants from L. groenlandica. The surface c e l l s , because of the meristematic nature, are prone to be variable. Inherent 2 0 and environmental factors affecting.the rate and mode of growth would influence the character of the meristematic c e l l s . Burrows ( 1 9 6 4 ) i n a study of L. saccharina related v a r i a b i l i t y of c e l l size, blade thickness, and other blade c h a r a c t e r i s t i c s to temperature. In•my study, those plants conforming to Setchell and Gardner's . ( 1 9 2 5 ) L. cuneif o l i a are considered to be L. groenlandica. Setchell and Gardner ( 1 9 2 5 ) distinguished L. platymeris De l a Pyl. from L. c u n e i f o l i a on the basis of the abullate blade and long, often complanate stipe of the former i n contrast to the bulla t e blade and short, often terete stipe of the l a t t e r . Widdowson ( 1 9 5 9 ) suggested that Setchell and Gardner misinterpreted De l a Pylaie's description of L. platymeris. Setchell and Gardner characterize L. platymeris as having a stipe up to 1 m long and compressed from just above the holdfast. De.la Pylaie ( 1 8 2 9 , . p . 5 2 ) states: \"L. stipete brevi, t e r e t i , . mihuto....\" and ;below he says \"Le stipe est cylindrique et egal dans toute sa longeur...seulement long de 9 a. 1 2 centimetres.... Results from a series of in s i t u studies on L. groenlandica at Glacier Point indicate that the form referred to 'as L. platymeris by Setchell and Gardner i s a form of L. 1 groenlandica at least one year old. These studies are discussed i n d e t a i l on page 3 0 . The above-mentioned f i e l d studies combined with Setchell and Gardner's apparent misunderstanding of De l a Pylaie's description of L.. platymeris lead me to place plants f i t t i n g t h i s species description (sensu.Setchell and Gardner, 1925) i n L. groenlandica. During,the course of my study four forms of L. groenlandica were recognized for the northeast P a c i f i c . These forms are not considered by me at present as legitimate taxonomic e n t i t i e s hut are distinguished merely to provide a means of f a c i l i t a t i n g d i s c u s s i o n . u n t i l such time as a c r i t i -cal monographic study for the entire genus i s made. Laminaria. groenlandica f l a t stipe form. F i g . . l 6 , 31, 44, 53; App. I, Sect. E. This form i s t y p i f i e d by having an abullate blade and complanate stipe usually exceeding 40 cm.in ;length. Plants f i t t i n g .this description.have been collected only from exposed areas in the lower i n t e r t i d a l region .about north Graham Island, Queen Charlotte Islands, B r i t i s h Columbia. Laminaria groenlandica short.stipe form. Pig. 13, • 15, 53; App. I, Sect. P. Blade with or without bullae. Stipe usually less than ,10 cm long, terete to s l i g h t l y complanate. Two patterns of b u l l a t i o n have been observed for t h i s form. The f i r s t pattern .consists of two longitud-i n a l rows of bullae running the length of the blade.. This form clo s e l y f i t s Setchell and Gardner's (1925) I±' cuneif o l i a f. subsimplex. The second pattern i s charac-te r i z e d by having the' entire blade i r r e g u l a r l y b u l l a t e . This form f i t s Setchell/and Gardner's (1925) L. cu n e i f o l i a f. amplissima. The'short stipe form occurs i n t e r t i d a l l y and s u b t i d a l l y in regions of. l i t t l e surf action from northern Washington to Attu Island, Alaska. 22 Laminaria groenlandica long stipe form. F i g . , l 4 , 30, 43, 53; App. I, Sect. G. Blade with or without bullae. Stipe s l i g h t l y flattened or terete, usually. 10 to 30 cm..long. This form i s s i m i l a r to Setchell and Gardner's (1925) L. c u n e i f o l i a f. c u n e i f o l l a and encompasses at least i n part t h e i r L. platymeris. The long stipe form ; i s common in regions of heavy surf from southern Vancouver Island, B r i t i s h Columbia to Attu Island, Alaska. To understand better the r e l a t i o n s h i p between the long and short stipe forms, a series of transplantations were per-formed . involving, these two forms. Complete d e t a i l of these studies i s presented on page 34. Laminaria groenlandica shade form. F i g . 11, 12, 29, 4 l . This form i s the same as Setchell and Gardner's (1925) L. c u n e i f o l i a f. angusta. This i s a shade form of the long and short stipe forms. The habitat.and. d i s t r i b u t i o n , of the shade form i s thought.to coincide w i t h t h a t of the long,and short stipe forms. The plants are small, usually less than 30 cm long.and 5 cm wide. The stipe i s usually terete and less than 5 cm long. For a complete discussion of the shade form and. i t s r e l a t i o n s h i p to the long.and short stipe forms see page 29. Laminaria saccharina (L.) Lamouroux Setchell and Gardner, 1925, p. 595. Doty, 1947, P. 39. Scagel, 1957, P. 96. Dawson, 1950, p. 153 (as L. cordata). 23 Dawson,. 1961, p. 396 (as. L. cordata and L. saccharina) . Description. Pig. 1J,,18, 32, 45. Sporophyte perennial from the stipe. Holdfast of many branched haptera. Stipe of varying length, terete to s l i g h t l y complanate, mucilage ducts absent. Blade usually entire, cuneate to cordate, of varying length and width, with or without bullae, mucilage ducts present. A l l plants producing s o r i i n winter; plants older than 9 months often producing s o r i i n late spring. Northeast P a c i f i c D i s t r i b u t i o n . F i g . 53; App. I , Sect. H. Prom Kenai Peninsula, Alaska to Coos Bay, Oregon, with an i s o l a t e d population on Santa Catalina Island, C a l i f o r n i a . E a r l i e r known from the Alaskan Peninsula to Coos Bay, Oregon. Habitat. Growing on s h e l l , , wood, and rock i n sheltered waters from the lower i n t e r t i d a l region to the upper subtidal region. Known only in. the subtidal region, in areas exposed to.surf. Comments. Setchell and Gardner (1925) recognized three forms of L. saccharina in.the northeast P a c i f i c . The form saccharina i s most often encountered. This form f i t s the description given above. Form l i n e a r i s d i f f e r s from form saccharina i n that i n the former the haptera extend up the stipe whereas in the l a t t e r the haptera are r e s t r i c t e d to the lower end of the stipe. Setchell and Gardner (1925) describe the d i s t r i b u t i o n of form l i n e a r i s as extending from Unga, Alaska to northern Washington, growing.on rocks i n the upper subtidal region.. In t h i s study form, l i n e a r i s has been observed only as d r i f t . Form membranacea i s the 24 t h i r d form recognized by Setchell and Gardner. This plant i s characterized by having an ample and membranous blade. Plants f i t t i n g the description of form membranacea have been collected i n the lower i n t e r t i d a l region at Sechelt Inlet, S t r a i t of Georgia, and B u l l Harbour, Hope Island, B r i t i s h Columbia. Setchell and Gardner describe i t s d i s t r i b u t i o n as extending from Alaska to Coos Bay, Oregon. Dawson .(195©) j u s t i f i e d describing L. cordata Dawson as a new species from Santa Catalina Island, C a l i f o r n i a on the grounds that \" . . . t h i s species i s unlike any previously described from the north P a c i f i c by manner of i t s short-s t i p i t a t e , non-bullate, entire but terminally eroded, broad-cordate blade, and holdfast of branched haptera.\" This species described by Dawson has the same mucilage duct dis-t r i b u t i o n as does L. saccharina. Further, the c h a r a c t e r i s t i c s employed i n setting L. cordata apart from other north P a c i f i c Laminaria are a l l shared by the variable species L. saccharina. Examination.of the type specimen (Dawson #56001, AHFH #36922) revealed that t h i s plant i s similar i n a l l regards except the s l i g h t l y coarser haptera to the Laminaria saccharina observed by me i n the northeast P a c i f i c . These observations lead me to reduce L. cordata Dawson to synonomy with the older species, L. saccharina (L.) Lamour. f. saccharina. Laminaria f a r l o w i i Setchell. Setchell,. 1891, p. 220. Setchell, 1905, p. 139. Setchell and Gardner, 1925, p. 599. Smith, 1944, p. 136. Dawson,. 1961, P. • 396. Description.. F i g . 1 9 * ' 3 3 , 46. Sporophyte 4 o + cm long. Holdfast of many, branched haptera. Stipe 4 cm long, terete, without mucilage ducts. Blade entire, cuneate, entire surface i r r e g u l a r l y bullate, mucilage ducts present. Northeast P a c i f i c D i s t r i b u t i o n . . F i g . 51; App. I, Sect. I. Central to southern C a l i f o r n i a and one isol a t e d c o l l e c t i o n at Comox, B r i t i s h Columbia. E a r l i e r known from Santa Cruz, C a l i f o r n i a to Bahia del Rosario, Baja C a l i f o r n i a . Habitat. The Comox specimens were collected, i n sheltered waters. Comments. The above description i s based on two plants (V 001484) collected by John Macoun,. 1915, at Comox, B r i t i s h Columbia. Laminaria complanata (Setchell and Gardner) Setchell Setchell and Gardner, 1903, p. 262. (as L. saccharina f. complanata). Setchell,, 1 9 1 2 , p. 149. Setchell and Gardner, 1925, p. 596. Scagel, 1957, P. 93 . Dawson, 1961, p. 396. Description. F i g . 2 0 , . 3 4 , 47. Holdfast of stout, branched haptera. Stipe less than 50 cm long, of variable width, being terete below and complanate near the blade base, mucilage ducts absent. Blade of variable size, trun-cate to cordate, mucilage ducts present. No plants with 26 s o r i observed. Northeast P a c i f i c D i s t r i b u t i o n . . Pig. 51; App. I, Sect. J. Known.only from northern Washington, James Bank, B r i t i s h Columbia, and Graham Island, Queen Charlotte Islands, B r i t i s h Columbia. Earlier ;known only from Friday Harbor, Washington. Habitat. Growing on rocks i n the subtidal region i n sheltered areas. The Queen Charlotte c o l l e c t i o n was made from an exposed.area i n the i n t e r t i d a l region. Comments. The species referred to here as L. complanata S. closely f i t s Kjellman's (1877) L. d i g i t a t a f. complanata and may well be an i s o l a t e d population of that form. Laminaria s e t c h e l l i i S i l v a Setchell, 1905, p. 139 (as L. .andersonii). Setchell and Gardner, 1925> P. 605 (as L. andersonii). Smith, 1944, p. 137 (as L. andersonii). Scagel, 1957, P. 97. S i l v a , 1957, P. 42. Dawson, 1961, p. 396. Description. F i g . 21 , 22, 23, 35, 48. Sporophyte perennial from the stipe, up to 1.5 m long. Holdfast of many stout, branched haptera. Stipe terete, r i g i d , up to 50 cm.long, usually greater than 2 cm thick at the base, mucilage ducts present, in the mid-cortical region. Blade of variable length, cuneate, deeply dissected into many segments, mucilage ducts conspicuous. Plants producing 27 s o r l i n late winter and early summer. Northeast P a c i f i c D i s t r i b u t i o n . F i g . 52j App. I, Sect. K. From Yakutat, Alaska to southern C a l i f o r n i a . E a r l i e r known from northern B r i t i s h Columbia to southern C a l i f o r n i a . Habitat. Growing on rocks i n the lower i n t e r t i d a l region and upper subtidal region i n areas of heavy surf. Restricted to the subtidal region in.sheltered areas. Comments. Laminaria s e t c h e l l i i displays considerable morphological v a r i a t i o n characterized by the blade becoming broader and the degree of dissection decreasing, with a t r a n s i t i o n from exposed to sheltered water (Fig. 21, 2 2 ) . Plants with newly rejuvenated blades are e a s i l y recog-nized as belonging.to L. s e t c h e l l i i by t h e i r heart shaped blades (Fig. 2 3 ) . A strong a f f i n i t y between L. s e t c h e l l i i and L. s i n c l a i r i i i s suggested on the basis of the blade and stipe anatomy. Both species have d i s t i n c t medullary regions i n the blade and deep seated mucilage ducts which are positioned.in the mid-cortical region i n both the blade and stipe. Laminaria dentigera Kjellman Setchell and Gardner, 1925, p. 604. Dawson, 1961, p. 396. Description. F i g . 24, 36, 49. Holdfast of stout, branched haptera. Stipe terete, usually more than 2 cm thick, u s u a l l y - l e s s than 40 cm. long, mucilage ducts pre-sent. Blade dissected into many narrow segments of variable width and length, mucilage ducts present but sparse. 28 Northeast P a c i f i c D i s t r i b u t i o n . Pig. 51; App. I, Sect. L. Prom Attu Island, Alaska to Yakutat, Alaska. E a r l i e r reported (Setchell and Gardner, 1925) as extending from the Aleutian Islands to the Bering S t r a i t . Habitat. Growing on rocks i n the lower i n t e r t i d a l region i n exposed areas. Comments. Only herbarium specimens studied. This species i s e a s i l y distinguished from L. s e t c h e l l i i on the basis of the mucilage duct positioning. In L. s e t c h e l l i i the ducts are deep seated while i n L. dentigera they are positioned near the periphery of the blade and sti p e . In the field.these two species are d i f f i c u l t to dis-tinguish. One c h a r a c t e r i s t i c which i s of assistance i n distinguishing these species i s the depth-to which the blades are dissected. In L. s e t c h e l l i i the blades are dissected almost to the blade base, whereas i n L. dentigera the d i s -section ceases some distance above the base. Yakutat, Alaska appears to be the northern extent of L. s e t c h e l l i i and the southern extent of L. dentigera. 29 LIFE HISTORIES AND GROWTH PATTERNS OF LAMINARIA GROENLANDICA LONG AND SHORT STIPE FORMS AND LAMINARIA SACCHARINA Studies on the l i f e h i s t o r i e s and growth, patterns were made to increase our understanding of the plants, to apply t h i s to subsequent studies on di s t r i b u t i o n , , and to develop a basis for evaluation of plant response to environmental factors. Further, i t was intended that the following studies on the two forms of L. groenlandica might lead to a better understanding of the taxonomic l i m i t s of t h i s species. L i f e H i s t o r i e s of Laminaria groenlandica Long and Short Stipe Forms Studies on the l i f e h i s t o r i e s of i n t e r t i d a l l y occurring L. groenlandica long, and short stipe forms were made during the years 1962-1964, at Glacier Point and Ogden Breakwater. These studies consisted of observations on populations and tagged.individuals. The l i f e h i s t o r i e s of both forms are e s s e n t i a l l y the same. With both forms, the sporophytic generation f i r s t became conspicuous i n February (Fig. 5 4 ) . By A p r i l two types of plants were apparent: large plants t y p i c a l of both forms, and.small plants t y p i c a l of the shade form (= L. c u n e i f o l i a f. angusta Setchell and Gardner, 1925). From observational studies, the small form i s thought.to r e s u l t from shading i n areas heavily populated by-larger seaweeds. In June, 1962 and 1964, many of the larger plants became f e r t i l e , whereas i n June, 1963, the larger plants remained, in the vegetative state. F a i l u r e of the June, 1963 plants to produce, s o r i may have resulted from the severe sunburn received by them during the spring tides of that, period. In June, 1962 and 1964, there was no evidence of severe sunburn. The small plants (shade form) were not observed with s o r i i n June. The large plants generally disappeared af t e r releasing t h e i r meiospores. Natural removal of some of the larger plants i n June and July exposed many of the small plants. These newly exposed plants grew rapidly, whereas those s t i l l shaded remained e s s e n t i a l l y the same siz e . During the November to January period there was no mea-surable vegetative growth and a l l observed L. groenlandica produced s o r i . Following release of meiospores, most of the blade was l o s t . In February regeneration of a new blade was i n i t i a t e d , and by June i t was not possible to d i s t i n g u i s h between the shade form and the long and short stipe forms. In June most of the two-year plants became f e r t i l e except those severely sunburnt. Laminaria groenlandica long stipe form, p e r s i s t i n g into the second year, closely f i t s the description of L. platymeris (sensu Setchell and Gardner, 1925). The stipe length of the former exceeded the greatest length attributed by Setchell and Gardner (1925) to L. c u n e i f o l i a (= L. groenlandica). The stipe was usually flattened and the blades, as observed i n June, were free of b u l l a t i o n s and 31 i r r e g u l a r l y dissected. A large population of new sporophytes became conspicuous in February and was augmented throughout the year. The February plants apparently had t h e i r o r i g i n from gametophytes a r i s i n g from meiospores released l n the November to January period. Sporophytes a r i s i n g at times other than February may have t h e i r o r i g i n from meiospores released from i n t e r -t i d a l plants i n June or from subtidal plants. However, the phenology of the subtidal plants i s unknown. Gametophyte and Microscopic Sporophyte Phases of Laminaria groenlandica Long Stipe Form One of two forms of gametophyte was produced under cul-ture conditions. Under unfavorable conditions of s a l i n i t y and temperature (p. 4 7 ) the u n i c e l l u l a r female gametophytes produced larger oogonia (Fig. 55b) than were produced under favorable conditions ( Fig. 5 5 c ) . The male gametophytes grown under favorable conditions were small, consisting of a few c e l l s most of which became antheridia ( Fig. 55d). When cul-tured under unfavorable conditions, the male gametophytes were quite large, consisting of many c e l l s few of which became antheridia (Fig. 55a) . Sporophytes grown under favorable culture conditions were at f i r s t . u n i s e r i a t e filaments which l a t e r became broader by a series of p e r i c l i n a l d i v i s i o n s (Fig. 55e, 55f) . When grown under-adverse conditions, structures which are thought to be abnormal sporophytes developed; these were ir r e g u l a r , consisting of a globular mass of large c e l l s 32 (Pig. 55g) . Growth Patterns of Laminaria groenlandica Long and Short Stipe Forms ? To define the growth patterns i n the blades of these two forms, the growth of the plants was followed i n both culture and i n s i t u . This was accomplished by punching a series of holes at close i n t e r v a l s along the longitudinal axis of the blade. Both the distances between holes and the blade width at each hole were recorded at the beginning of the experiment and again, a f t e r 30 days. In s i t u growth studies were made on three plants of each form ranging from 40-60 cm.in length from July 20 to August 19, 1962. Similar studies were made under culture conditions from September 23 to October 23, on f i v e plants of each form ranging i n length from 20-30 cm. These plants were cultured in p l a s t i c aquaria at 10°C, 28-31$* s a l i n i t y , and 4-50 f t - c illumination, with a regime of 12 hours l i g h t and 12 hours dark. D i s t r i b u t i o n of meristematic a c t i v i t y in the blades of the long and.short stipe forms was sim i l a r . In a l l cultured plants 90$ of the measurable longitudinal growth was i n the basal.7 cm of the blade (Fig. 56, 5 7 ) . No longitudinal growth was recorded more than 11 cm d i s t a l to the blade base. In a l l of the i n s i t u plants,. 90$ of the longitudinal growth was observed i n the basal 17 cm of the blade (Fig. 5 8 ) . Longi-tudinal growth was r e s t r i c t e d to the basal 22 cm of the blade. 33 Measurable l a t e r a l growth i n the cultured plants was limit e d to the basal 12 cm.of the blade (Pig. 59, 6 0 ) , but in plants studied, i n s i t u , l a t e r a l growth extended 20 cm d i s t a l to the blade base (Pig. 6 l ) . The greatest per cent increase'in width for the cultured plants was 80$ and f o r the i n s i t u plants, 450$. The gross morphology of L. groenlandica long and short stipe forms i s i l l u s t r a t e d i n Figures 13, 14, and 15. Large plants of the short stipe form are characterized by a broad blade with.a truncate-cordate base and a short st i p e . The long stipe form has a l i n e a r blade usually with a cuneate base and a.long,stipe. Young plants of both forms, when subjected to shading,.produce a growth form t y p i c a l of Setchell and Gardner's (1925) L. cu n e i f o l i a f. angusta. This growth form i s distinguishable by i t s small size. In Situ Growth Rates of Laminaria groenlandica Long and Short Stipe Forms The longitudinal growth rates for small groups of long and short stipe forms of L. groenlandica were determined at di f f e r e n t periods of the year (Table I ) . This was. accom-plished by punching holes at short i n t e r v a l s along the length of the blade and observing.the d i s t r i b u t i o n s of holes a f t e r a period of time. The greatest growth rates were observed p r i o r to July; the slowest rates were recorded i n the f a l l , p r i o r to sorus development. 34 Transplant Studies on Laminaria groenlandica Long and Short Stipe Forms As was mentioned above, two of the c h a r a c t e r i s t i c s dis-tinguishing long and short,stipe forms are blade width and stipe length. To quantify these c h a r a c t e r i s t i c s the stipe length to blade width (St/W) r a t i o of several populations was determined at d i f f e r e n t times of the year (Table I I ) . Measurements were made on tagged populations several times and on untagged populations once. The grand mean St/W r a t i o for a l l measurements of long stipe populations was 0.52 and the means of the various populations measured varied from 0 . 2 9 - 0 . 8 5 . The grand mean St/W r a t i o for a l l short stipe populations measured was 0.16 while the means of the i n d i v i d u a l populations varied from 0 . 1 2 - 0 . 2 2 . The general trend.was fo r the young plants of both forms to have a St/W r a t i o of. 0.20-0.40. As these plants increased i n . s i z e , the long stipe St/W r a t i o increased and.the short stipe St/W r a t i o decreased. Between February 22 and A p r i l 28, 1963, twenty-five long,stipe plants from Glacier Point and 20 short stipe plants from the Ogden Breakwater were transplanted to a sheltered area i n Burrard Inlet near Stanley Park, Vancouver. The plants selected f o r transplantation were the smallest that could be handled conveniently. None of these plants exceeded 45 cm i n length. The St/W r a t i o was determined at the time of trans-plantation and f o r t n i g h t l y as long as the plants remained in Burrard I n l e t . It was possible to-follow change i n the St/W r a t i o f or only 12 long stipe and 14 short stipe plants. Of the 19 plants l o s t early, i n the study most were thought-l e s s l y destroyed by picnickers. Remaining transplants were l a s t observed on June 10, 1963. Subsequent loss of these plants was thought to be the re s u l t of unfavorable oceano-graphic conditions. The f i n a l mean St/W r a t i o was determined by averaging the l a s t i n d i v i d u a l measurements regardless of when they were made. The I n i t i a l mean St/W r a t i o for the long stipe plants was 0 . 2 7 ; the f i n a l mean r a t i o was 0.40. The St/W r a t i o f o r the short stipe plants changed from.an i n i t i a l mean, of 0 .20 to a f i n a l mean of 0.17 (Table I I I ) . Five short stipe plants were transplanted from Ogden Breakwater to Glacier Point on May 22, 1963. The St/W r a t i o of these plants changed from an i n i t i a l mean.of 0 .23 to a f i n a l mean.of 0 .34 as observed October 6, 1963 (Table I I I ) . Results of the transplantations of long and short stipe plants to Burrard Inlet indicate that the differences i n stipe length and blade width between these forms i s geneti-c a l l y controlled and i s not the r e s u l t of environmentally induced phenotypic p l a s t i c i t y . However, response of short stipe plants placed.in the long stipe plants' environment contradicts t h i s hypothesis. The r e s u l t s of both .transplant experiments are considered.inconclusive because of the small number of plants tested and the p o s s i b i l i t y of precondition-ing of the plants p r i o r to transplantation. 3 6 L i f e History of Laminaria Saccharina The l i f e h istory of L. saccharina was followed i n Burrard Inlet near Stanley Park, Vancouver from. 1962 to 1965. This study consisted of general observations of the entire popula-t i o n and s p e c i f i c observations of tagged plants. The sporophytes f i r s t became evident i n the February-March, period. Measurable growth of these plants continued u n t i l mid-November, by which time a l l plants had produced s o r i (Fig. 6 2 ) . After release of the meiospores a large portion of the blade was l o s t . Those plants surviving the remainder of the winter i n i t i a t e d blade regeneration i n February. A l l plants of t h i s second year group became f e r t i l e i n June, a f t e r which the i n t e r t i d a l plants were l o s t . Young sporophytes, presumably r e s u l t i n g from meiospores released in. June, became apparent i n September. These plants, through a rapid growth phase, became f e r t i l e i n October-November. Unlike L. groenlandica, new L..saccharina sporo-phytes became apparent only i n two b r i e f periods, February-March and September, and the population.was not augmented throughout the year. Gametophyte and Microscopic Sporophyte Phases of Laminaria saccharina The gametophytes took on one of two forms, depending upon culture conditions. In adverse conditions (p. 47) .the male and female gametophytes were large, filamentous, and consisted of many c e l l s (Fig. 63a, 63b) . Under favor-able conditions the female gametophytes were u n i c e l l u l a r , 37 the single c e l l a r i s i n g at the end of a germination tube through which the protoplasm passed from the meiospore (Pig. 6 3 c ) . This c e l l functioned as the oogonium. Under favorable conditions the male gametophyte was greatly reduced in size, consisting of many c e l l s most of which became antheridia .(Fig. 63d) . Young sporophytes a r i s i n g from the fusion of egg and sperm t y p i c a l l y underwent a series of a n t i c l i n a l d i v i s i o n s r e s u l t i n g in a uniseriate filament. Later, the sporophyte tha l l u s became wider as a r e s u l t of p e r i c l i n a l d i v i s i o n s (Fig. 63e, 6 3 f ). Under unfavorable conditions plants thought to be abnormal sporophytes were produced; these were ir r e g u l a r , globular masses of c e l l s (Fig. 63g). The o r i g i n of these structures i s questionable, as gametes were not observed i n adverse culture conditions. In October, 1963 and. 1964, glass s l i d e s were inoculated with meiospores and placed in Burrard Inlet near Stanley Park. Inoculation was achieved by placing s l i d e s in dishes of meiospore-suspension f o r 24 hours. Then the. s l i d e s with the attached meiospores were placed i n shallow staining dishes and established in Burrard Inlet at the one foot tide l e v e l . A few s l i d e s were removed each fortnight for microscopic examination. Sporophytes were produced by December of both years. In a l l cases only 6-12 sporophytes were produced on each s l i d e . The sporophytes and remnants of the female gametophytes were of the type shown i n Figures 63e, 63f. No male or u n f e r t i -l i z e d female gametophytes were observed. Growth Pattern of Laminaria saccharina The d i s t r i b u t i o n of l a t e r a l and.longitudinal growth was determined i n 1962 for 6 plants i n culture and 3 plants i n s i t u . The cultured plants varied i n length from.14-19 cm at the s t a r t , and t h e i r growth.was followed from June 10-23. The i n s i t u plants were 26-35 cm long at the beginning, and were studied from August,1-13. The method for culturing and for growth.study of these plants was the same as described f o r L. groenlandica (p. 3 2 ) . A l l measurable longitudinal growth .in the cultured plants was i n the basal 5 cm of the blade, 90$. of the growth being in the basal 4 .5 cm (Pig. 64). Observed, longitudinal growth i n s i t u was i n the basal 14 cm of the blade; 90$ of . the growth was r e s t r i c t e d to the basal 8 cm (Pig. 6 5 ) . Measurable l a t e r a l growth i n both cultured and i n s i t u plants was r e s t r i c t e d to the basal.15 cm of the blade. The i n s i t u plants increased.in width a maximum of 100$, and the cultured plants a maximum of 30$ during the 13 day period of study. ' Growth Rates of In Situ Laminaria saccharina In May, 1962, twenty-five plants were punched in.the manner described on page 33 for L. groenlandica. . The longitudinal growth of these plants was followed u n t i l December, 1962. Figure 68 depicts the th e o r e t i c a l length of the population and the growth rate throughout the period of measurement. The t h e o r e t i c a l length was determined by adding the increase i n length by the in t e r c a l a r y meristem 39 to the i n i t i a l length of the plants as observed i n May. This measurement did not take into account, loss of the blade through erosion at the d i s t a l end. The mean th e o r e t i c a l length of the population at the end of the growth period was 195 em, wliereas the mean actual length of the population was approximately 30 cm. The longitudinal growth rate of t h i s population shows a normal d i s t r i b u t i o n from August to December (.Pig. 6 8 ) . Prior to August the growth rate was i r r e g u l a r . This can be accounted for i n part by the severe sunburn these plants received during the spring tides i n June. The greatest mean growth rate was approximately 1.4 cm per day for the August to September period. ho LOCAL DISTRIBUTIONS OP LAMINARIA SACCHARINA AND OF LAMINARIA GROENLANDICA LONG AND SHORT STIPE FORMS.AS RELATED TO TEMPERATURE, SALINITY, AND WATER MOTION This study was confined to the coastal region from north Vancouver Island to the southern extent of the San Juan Archipelago because of the abundance of oceanographic and d i s t r i b u t i o n a l data available for t h i s region. The d i s t r i b u t i o n s of L. saccharina and the two forms of L. groenlandica about Vancouver Island and adjacent waters are i l l u s t r a t e d in Figure 53-The i n t e r t i d a l d i s t r i b u t i o n of L. saccharina on the west coast of Vancouver Island i s r e s t r i c t e d to sheltered.bays and i n l e t s , and on the east coast i t i s abundant from Hope Island (50O58'N 127°55'W), at the north, to the San Juan Archipelago in the south. Laminaria saccharina i s known, in the subtidal region only, at Amphritrite Point (48°56'N 125°33'W), on the west coast of Vancouver Island, and at Salmon Bank (48°26'N 1 2 3 ° 0 l ' w ) , i n the San Juan Archipelago. Laminaria groenlandica long stipe form i s found pre-dominantly i n exposed regions along the north and west coast of Vancouver Island, the southern extent of i t s continuous west coast d i s t r i b u t i o n being Sooke (48°2 I 'N 1 2 3 ° 4 3 'W ) , with a population., occurring on the southwest coast of San Juan Island. The d i s t r i b u t i o n . o f L. groenlandica short, stipe form i s intermediate between that of L. groenlandica long stipe form and that of L. saccharina, often overlapping the d i s t r i b u t i o n 41 of the l a t t e r . The short stipe form i s encountered i n t e r -t i d a l l y at the mouths of i n l e t s along the west coast of Vancouver Island. This form i s abundant from Sooke to Jan Juan Island and along the east coast of Vancouver Island from the south side of Hope Island to the southern end of Johnstone S t r a i t ( 5 0 ° 2O'N) and i s infrequently encountered from t h i s point to the southern end of Texada I s l a n d ( 4 9 ° 3 0 ,N). One subtidal c o l l e c t i o n of the short stipe form has been made, near Keats Island, Howe Sound (49°24*N 123°28 'w) . The r e l a t i o n s h i p s between the horizontal d i s t r i b u t i o n s of L. saccharina and the two forms of L. groenlandica and temperature and s a l i n i t y were established by comparing the species' d i s t r i b u t i o n s i n selected areas with the seasonal v a r i a t i o n of surface temperature and s a l i n i t y . This seasonal v a r i a t i o n was characterized by p l o t t i n g mean monthly tempera-tures against mean monthly s a l i n i t i e s in.a T-S diagram. The r e l a t i o n s h i p of seasonal v a r i a t i o n f o r surface temperature and s a l i n i t y as related to the v a r i a t i o n in subsurface waters i s i l l u s t r a t e d in Figures 69 and 70. Variation of temperature and s a l i n i t y i n the upper 6.m i s n e g l i g i b l e at Sheringham Point, a region inhabited, by L. groenlandica long stipe form, and at Gordon Head, a region inhabited by both L. saccharina and L. groenlandica short stipe form (Fig. 6 9 ) . In Burrard Inlet, a region occupied by L. saccharina, considerable v a r i a t i o n of tempera-ture and.salinity i s encountered i n the upper 5 m (Fig. 7 0 ) . Laminaria groenlandica long stipe form occurs i n areas with cold, high s a l i n i t y water having l i t t l e seasonal v a r i a t i o n and exposed to heavy surf (Fig. 71, 7 2 ) . Pine Island and Kains Island t y p i f y t h i s habitat (Fig. 71) . This form i s absent from areas with considerable temperature and s a l i n i t y variations and from.areas where surf action i s n e g l i g i b l e , e.g., Friday Harbor (Fig. 71) . Laminaria groenlandica short stipe form i s absent from areas having extreme seasonal v a r i a t i o n of temperature and s a l i n i t y , e.g., Deep Cove and Departure Bay, and from areas subjected to heavy surf (Fig. 71, 7 2 ) . Laminaria saccharina inhabits temperature and s a l i n i t y situations t y p i c a l of those encountered by the two forms of L. groenlandica (Fig. 71, 72) as well as situations of extreme temperature and s a l i n i t y v a r i a t i o n . Laminaria saccharina i s absent from i n t e r t i d a l areas subjected to surf. Available subtidal d i s t r i b u t i o n a l data suggest that L. groenlandica short stipe form and L. saccharina may succeed.in geographical areas where surface water conditions are p r o h i b i t i v e , providing they grow at.a depth below the influence of these conditions. This hypothesis i s supported by subtidal c o l l e c t i o n s of L. saccharina from Salmon Bank and Amphritrite Point,- places where i t i s not known i n t e r -d i a l l y . Both of these areas are subjected.to heavy surf. :Laminaria groenlandica short stipe form i s known sub t i d a l l y only from one c o l l e c t i o n , that i n 35 to 40 feet of water near Keats Island, Howe Sound. Here, surface temperature and s a l i n i t y conditions (Table IV) appear unfavorable for th i s form on the basis of the data presented in Figures 71 and 72. However, temperature and s a l i n i t y conditions near Keats Island, at the depth inhabited by the short stipe form, are much, less variable (Table IV). Prom the above analysis of f i e l d data the following correlations have been made: Laminaria saccharina d i s t r i b u t i o n i s independent of l o c a l temperature and s a l i n i t y conditions and dependent upon the presence of calm water. Laminaria groenlandica short stipe form d i s t r i b u t i o n i s r e s t r i c t e d by temperature and/or s a l i n i t y only when any mean monthly value of temperature exceeds 1-8°C and/or when s a l i n i t y i s lower than 23$* (Fig. 7 2 ). Further, t h i s form i s excluded from.areas of heavy surf. Laminaria groenlandica long stipe form d i s t r i b u t i o n i s re s t r i c t e d . t o areas of heavy surf. These areas are characterized by low temperature and high s a l i n i t y . 44 EXPERIMENTAL EVALUATION OF THE EFFECTS OF VARIOUS OCEANOGRAPHIC FACTORS ON LAMINARIA SACCHARINA AND TWO FORMS OF LAMINARIA GROENLANDICA The following experiments were designed to test the responses of vegetative and reproductive phases of the two species to various temperature, s a l i n i t y , and water motion regimes. Culture studies were r e s t r i c t e d to L. saccharina and L. groenlandica long stipe .form; transplant studies were conducted on two forms of L. groenlandica. Also, a preliminary study was conducted to evaluate the possible e f f e c t s on the plants of seawater from d i f f e r e n t o r i g i n s . Whole Plant Response of Laminaria saccharina Sporophytes to Various Temperature and S a l i n i t y Situations Two plants were placed i n each of a variety of temperature-salinity situations on May 16, 1963. These situations consisted of 10, 20, 30, 40, and 50$p s a l i n i t i e s , at 5> 10, 15,. and 20°C. The plants were cultured i n 1 l i t r e dishes i n constant temperature rooms, under 330-345 f t - c l i g h t i n t e n s i t y , 12 hours l i g h t alternating with.12 hours dark. The culture medium was changed weekly. The increase i n surface area of the plants' blades was determined every 7 days for 4 weeks afte r which time the experiment was terminated. Injury to or death of the plants was recorded. Injured plants were characterized by a green-white color and were considered dead upon disintegration of the t h a l l u s . A l l plants maintained at 10 and 50$» s a l i n i t i e s died, death being e a r l i e r at the higher temperatures (Table V ) . A l l plants cultured at 20°C died by the end of the t h i r d week. Plants maintained at 5 to 15°C, i n 20 to 4o$? s a l i n i t y , survived the f u l l four weeks. Optimal conditions for growth were 10 to 15°C, in 20 to 30$0, s a l i n i t y . Whole Plant Response of Laminaria saccharina Sporophytes ' to Various Temperatures at Constant S a l i n i t y and to Various S a l i n i t i e s at Constant Temperature In the spring of 1964 f i v e plants were subjected to each of 5 s a l i n i t i e s ranging from 16 to 31$« > at a constant temperature of 10°C, and to 4 temperatures ranging from 7 to l6°C, at a constant s a l i n i t y of 2&%> . These plants were cultured i n 1 l i t r e dishes using.plastic aquaria as incubat-ors. The culture medium was changed every 5 days. Light conditions were 425 f t - c , 12 hours on and 12 hours o f f . The mean increase i n blade surface area per day was determined a f t e r 13 days for the plants subjected to various s a l i n i t i e s and a f t e r 15 days for the plants subjected to various temperatures. The optimal temperature tested.was 10°C (Fig. 7 3 ) . However, with the exception of one plant at. 10°C, plants grown at 7.5 and 12.5°C responded s i m i l a r l y . Plants grew most rapidl y at the higher s a l i n i t i e s (Fig. 7 4 ) . 46 Photosynthetic and Respiratory Responses of Laminaria saccharina and Laminaria groenlandica Long Stipe Form Sporophytes to Various Temperature and S a l i n i t y Situations This study employed discs cut from the blades of L. saccharina collected May 10, 1964 and L. groenlandica collected June 5, 1964. One disc from each of f i v e plants was placed.in a 1 l i t r e culture dish f o r each of the following temperature-salinity conditions: s a l i n i t i e s of 20, 24, 26, 28, 30, . and 32$*, at 7, 10, 13, 15, and l 8°C. The discs were'preconditioned to these temperature-salinity situations for 5 days. Light conditions were 425 f t - c , 12 hours l i g h t a l t e rnating with 12 hours dark. The sea-water was changed on the second and fourth days of pre-conditioning. On the morning of the sixth day the photo-synthesis and r e s p i r a t i o n rates were determined. The l i g h t period during which.photosynthesis was measured lasted 2 hours at an i n t e n s i t y of 650 f t - c and the dark period for r e s p i r a t i o n , 4 hours. In general, net photosynthesis and r e s p i r a t i o n i n L..saccharina discs increased with increase i n temperature but were r e l a t i v e l y uniform at a l l s a l i n i t i e s (Table VI). The resultant net photosynthesis/respiration r a t i o remained f a i r l y constant at a l l temperatures and. s a l i n i t i e s . For L. groenlandica long stipe form discs, the rate of net photosynthesis and the net photosynthesis/respiration r a t i o decreased with increasing temperature but increased with increasing s a l i n i t y (Table VI). There was no measurable net photosynthesis at s a l i n i t i e s lower than 2&%> , for temper-atures greater than 10°C, or i n any s a l i n i t y at. l 8°C. The rate of r e s p i r a t i o n increased with increasing temperature to 15°C, above which the rate decreased. The r e s p i r a t i o n rate was nearly constant at a l l s a l i n i t i e s . Measurable respiration.occurred i n a l l temperature-salinity situations tested. Responses of Laminaria saccharina and Laminaria groenlandica Long Stipe Form Gametophytes to Various Temperature and S a l i n i t y Situations Meiospores of these two species, attached to glass s l i d e s were placed i n 250 ml dishes, incubated i n p l a s t i c aquaria. Duplicate cultures were established for s a l i n i t i e s of 17, 20 , 23 , 26, 29, and 32$* , at 7, 10, 13, and ' l 6°C. Light i n t e n s i t y was approximately 100 f t - c ; the regime was 12 hours on and 12 hours o f f . The culture medium was changed weekly. Laminaria saccharina gametophytes were maintained under the above conditions from October 23 to December 15, 1964, and L. groenlandica long stipe form gametophytes, from-December 20, 1964 to January 25, 1965. S u i t a b i l i t y of the gametophytes' environment was determined by the plants' a b i l i t y to produce normal sporo-phytes (Table VII). Laminaria saccharina gametophytes produced normal sporophytes under most of the tested temperature-salinity conditions (Fig. 63e, 6 3 f). Plants thought to be abnormal sporophytes, consisting of i r r e g u l a r , globular masses of 48 c e l l s ( F i g . 63g) were produced at 17, 20, and 23$* s a l i n i t y and 16°C. Gametophytes of L. s a c c h a r i n a produced i n h i g h tempera-t u r e , low s a l i n i t y s i t u a t i o n s were l a r g e and f i l a m e n t o u s ( F i g . 63a, 63h) . S i m i l a r gametophytes were produced i n crowded c u l t u r e s . Gametophytes produced under uncrowded c o n d i t i o n s , a t low temperatures and h i g h s a l i n i t i e s , were, very s m a l l , the m a j o r i t y of t h e i r c e l l s f u n c t i o n i n g as gametangia ( F i g . 63c, 6 3 d ) . Gametophytes of L. g r o e n l a n d i c a long s t i p e form pro-duced normal sporophytes ( F i g . 55e, 55f) i n a l l s a l i n i t i e s , at 7 and.10°C, and only i n 29 and 3 2 $ * s a l i n i t i e s , at 13°C (Table V I I ) . P l a n t s thought t o be abnormal sporophytes ( F i g . 55g) were observed In. 17$* s a l i n i t y , at 10°C, and In 20-32$* s a l i n i t i e s , at 1 3°C S t r u c t u r e s resembling sporo-phytes were not observed In. 17$* s a l i n i t y , at.13°C, o r - i n any s a l i n i t y , at l 6°C. Gametophytes of L. g r o e n l a n d i c a long s t i p e form, grown under c o n d i t i o n s conducive to normal sporophyte p r o d u c t i o n , were s m a l l e r than those grown under c o n d i t i o n s g i v i n g r i s e to abnormal sporophytes. Development of what were thought t o be gametophytes, i n a l l t e s t e d s a l i n i t i e s , at l 6°C, was r e s t r i c t e d t o a few c e l l u l a r d i v i s i o n s r e s u l t i n g i n c o c c o i d t h a l l i . I t was. not p o s s i b l e t o d i s t i n g u i s h male from female gametophytes. 49 Photosynthetic and Respiratory Responses of Laminaria saccharina and of Laminaria groenlandica Long Stipe Form Sporophytes to Seawater of Different Origins The experimental procedure employed here was i d e n t i c a l to that described on page 46. Discs from f i v e plants of each species were placed in three water types: Glacier Point seawater, Burrard Inlet seawater, and Glacier Point seawater dil u t e d with Capilano River water. (This r i v e r enters the north side of Burrard Inlet.) S a l i n i t y of the three types of water was adjusted to 2&% and the cultures were maintained at 7 ° C After 5 days preconditioning, r e s p i r a t i o n and net photosynthesis of the discs were determined. No s i g n i f i c a n t differences i n response of the two species to the three water types was observed. Responses of Laminaria saccharina and Laminaria groenlandica Long Stipe Form Gametophytes and Microscopic Sporophytes to Seawater of Different Origins Responses of L. saccharina and L. groenlandica long stipe form to seawater of d i f f e r e n t origins were observed in November-December, 1963, and, for L. saccharina, again in June-July, 1964. The method of culture was as described on page 47. In the' November-December tests water from Burrard Inlet, Glacier Point, and the Ogden Breakwater was adjusted to 2&% s a l i n i t y , a n d maintained at 7°C. In the June-July tests water from Burrard Inlet and Glacier Point was adjusted to 28, 30, and 32$* s a l i n i t i e s and maintained at 7°C . 50 Both the November-December and June-July cultures were maintained f o r 35 days a f t e r inoculation with meiospores. At the end of t h i s time 25 sporophytes, randomly selected, were measured with an ocular micrometer. In both experiments the longest L. saccharina sporophytes developed in the Burrard Inlet water (Fig. 75, 7 6 ) . The longest L_. groenlandica sporophytes were produced i n Glacier Point and the Ogden Breakwater seawater. Transplantation Studies A series of transplantations was made to test the ef f e c t of oceanographic conditions encountered i n Burrard Inlet and Departure Bay on two forms of Laminaria groenlandica. These areas are inhabited by L. saccharina but not by L. groenlandica. Further, they are characterized by having considerable temperature and s a l i n i t y variations and are not exposed to any surf (Fig. 70, 8 6 ) . Twenty young sporophytes of L. groenlandica long stipe form and 25 of L. groenlandica short stipe form were transplanted to Burrard Inlet between February 22 and A p r i l 28, , 1963. Plants of both forms survived u n t i l June 10, 1963. On August 5, 1963, twenty plants of both forms were transplanted to Departure Bay. Only remnants of these plants were observed at the next, low tide series, 14 days l a t e r . Ten plants of both forms were transplanted to Burrard Inlet on January 13, 1964. A l l of these plants had i n i t i a t e d soral production by the time of transplantation. The majority of these plants survived i n Burrard Inlet u n t i l May 28, 1964. On December 20, 1964, ten plants of the long stipe form were transplanted to Burrard Inlet. Microscopic examination of representative plants disclosed that apparent soral production had not yet been i n i t i a t e d . A l l plants had produced .sori by mid-January. Fragments of these s o r i , when placed under culture conditions, released meiospores. One set of gametophytes of L . groenlandica long stipe form were placed i n Burrard Inlet on glass s l i d e s January 20, 1965. Subsequent observations of the glass s l i d e s revealed no laminarialean gametophytes, and the s l i d e s were overgrown with diatoms. 52 STUDIES O N THE VERTICAL DISTRIBUTION OP LAMINARIA, SACCHARINA The V e r t i c a l D i s t r i b u t i o n of Laminaria saccharina as Related to Possible Determining Factors The upper l i m i t s of L. saccharina d i s t r i b u t i o n were observed at i n t e r v a l s from November, 1962 to December, 1963 (Fig. 7 7 ) . Three positions for the upper l i m i t s were observed, the f i r s t near Point Atkinson in Burrard Inlet (49°20'N :-123°l6' W), an area with tide pools,, the remaining two-in Burrard Inlet, near Stanley Park. Position 1. Plants occupied tide pools situated above the mean,low water (MLW) throughout the year. Position 2. Plants were found above MLW i n areas free of tide pools only when the spring low tides occurred at night (September-April). Position 3- The upper-most extent of L_. saccharina was below MLW when the spring low tides occurred during the day (April-September). The lower l i m i t s of .L..saccharina were found.on two occasions in June, 1963 to be approximately 6 .5 m below MLW. The occurrence of L. saccharina throughout the year in tide pools situated above MLW suggests that factors operative during emergence are responsible for determining the upper l i m i t s of t h i s species. F i r s t , only those plants which are constantly submerged may remain.above MLW throughout the year. Second, water temperature encountered i n the tide pools would approximate or exceed extreme values encountered i n the i n l e t . The upper v e r t i c a l l i m i t s of L. saccharina are lowered from 20 cm.above MLW to 40 cm below MLW.with a s h i f t from night to day of the spring low tides (Fig. 77) . Prom September to A p r i l the plants above MLW are subjected to greater frequency and accumulative duration of emergence and longer i n d i v i d u a l emergences than are plants situated below MLW from A p r i l to September (Pig. 7 8 ) . This suggests that the factors determining the upper l i m i t s are more severe during the day than during the night. A i r temperature and i n s o l a t i o n are thought to-be agents responsible in determining the upper l i m i t s of L.. saccharina ('Pig. 79-82) . Plants exposed by low spring tides occurring during the day are subjected to more in s o l a t i o n and higher a i r temperatures than.are plants exposed by low spring tides occurring during the night. The lower i n s o l a t i o n values encountered by the f i r s t group r e s u l t from lower incident radiation and a greater mean water depth through which the l i g h t must pass. The lower l i m i t s of L. saccharina are thought to r e f l e c t the compensation depth of t h i s species. The compensation depth Is mainly a function of submarine illumination and temperature (Pig. 70, 8 3 ) . Experimental Evaluation of Some Factors as Possible Determinants of D i s t r i b u t i o n of Laminaria saccharina Temperature and S a l i n i t y . The i n v i t r o response of 54 L. saccharina sporophytes i l l u s t r a t e s the a b i l i t y of t h i s species to thrive i n conditions of temperature and s a l i n i t y exceeding extremes encountered i n Burrard Inlet (Fig. 70j Tables V, VI). Cultured gametophytes of L. saccharina grew under temperature and s a l i n i t y conditions representative of extreme values f o r Burrard Inlet (Table VII). However, production of normal sporophytes was r e s t r i c t e d to tempera-tures below and s a l i n i t i e s above the extremes encountered during summer months in the Inl e t . These summer extremes were l8°C and. 15$* s a l i n i t y . Gametophytes grown under con-d i t i o n s p r o h i b i t i v e to formation of normal sporophytes may produce normal sporophytes when subsequently introduced to favorable conditions. If t h i s i s the case, then extreme temperatures and s a l i n i t i e s encountered i n Burrard Inlet would not determine the upper l i m i t s of L. saccharina but would only delay normal sporophyte production. Submarine Illumination. Several young plants were cultured under 6 d i f f e r e n t . l i g h t i n t e n s i t i e s at the Van-couver Public Aquarium In the spring of 1964. The water temperature was,10°C and the l i g h t regime was 12 hours l i g h t , 12 hours dark. The increase i n blade surface area was determined a f t e r 22 days (March 2 1-April 13)• The great-est growth occurred at 700 f t - c (Fig. 8 5 ) . Blades of plants cultured under 850 f t - c were a yellow-white color. The apparent photosynthesis of blade discs was deter-mined at 10 l i g h t i n t e n s i t i e s ranging from 20 to 900 f t - c at 10 C. The discs were cut at the time of c o l l e c t i o n (June 1, 1964) from injury free blades and placed i n darkness for 24 hours at 10°C. Then, the apparent photosynthesis of each disc was determined i n the manner described on page 46. The l i g h t compensation point was found to be between 20 to 40 f t - c (Fig. 84). The rate of apparent photosynthesis increased with increasing, l i g h t i n t e n s i t y to 500 f t - c , above which i t was constant. The a b i l i t y of L_. saccharina gametophytes to produce sporophytes under various l i g h t i n t e n s i t i e s was tested i n June, 1964. Meiospores attached to glass s l i d e s were placed i n eight l i g h t i n t e n s i t i e s ranging from 25 to 710 f t - c . They were maintained i n open dishes submerged i n the culture tank at the Vancouver Public Aquarium. The water temperature was 10°C, l i g h t regime 12 hours l i g h t and 12 hours dark. Gametophytes produced normal sporophytes within four weeks under a l l l i g h t i n t e n s i t i e s tested. 56 DISCUSSION; AND CONCLUSIONS Taxonomy and D i s t r i b u t i o n Ten morphologically d i s t i n c t species of Laminaria are recognized by me for the northeast P a c i f i c . In many instances these species form morphologically s i m i l a r p a i r s . The members of a pa i r are usually distinguishable by one charac-t e r i s t i c and often occupy similar ecological niches separated geographically. These species pairs are described below. Laminaria yezoensis--Laminaria solidungula. ' These two species can be distinguished on the basis of the d i s t r i -bution, of mucilage ducts. Laminaria solidungula has mucilage ducts i n the stipe, L. yezoensis does not. Laminaria yezoensis i s found i n the northern P a c i f i c (Miyabe, 1902) and L. solidungula i n both the .northern A t l a n t i c (Taylor, 1957) and A r c t i c Oceans (Kjellman, 1883). Laminaria saccharina--Laminaria f a r l o w i i . Laminaria f a r l o w i i can be distinguished from L. saccharina by having the entire blade b u l l a t e ; in. L. saccharina the bullae, when present, are r e s t r i c t e d to two rows p a r a l l e l to the blade margin. Laminaria saccharina i s known i n the north A t l a n t i c (Taylor, 1957), the A r c t i c (Kjellman, 1883), the northwest P a c i f i c (Tokida, 1945) , and the northeast P a c i f i c south to Oregon, with an is o l a t e d population at Santa Catalina Island, C a l i f o r n i a . Laminaria f a r l o w i i i s known from several locations i n C a l i f o r n i a and one i n B r i t i s h Columbia. Laminaria.saccharina—-Laminaria agardhil. . The members of t h i s p a i r can be separated from each other on the basis of the d i s t r i b u t i o n of mucilage ducts; L. saccharina has ducts i n the blade, L. agardhli does not. Laminaria agardhil Is known from the north A t l a n t i c (Taylor, 1957) and the A r c t i c (Kjellman, • 1883). The d i s t r i b u t i o n of L. saccharina i s given above. Laminaria longipes--Laminaria s i n c l a i r i i . Laminaria s i n c l a i r i i can be distinguished from L. longipes by the presence of mucilage ducts i n i t s stipe. Laminaria s i n c l a i r i i i s known from C a l i f o r n i a to central B r i t i s h Columbia, L. longipes, from central Alaska to Japan (Tokida, 1954). Laminaria s e t c h e l l i i- - L a m i n a r i a dentigera. These two species can be separated from each other by the positioning of the mucilage ducts. In L. s e t c h e l l i i the ducts are deep seated; i n L. dentigera they are situated near the periphery. Laminaria s e t c h e l l i i i s known from C a l i f o r n i a to southern Alaska and L. dentigera from central Alaska to Japan (Tokida, 1954). Recent f i e l d and culture studies have demonstrated considerable morphological p l a s t i c i t y of many characteris-t i c s used for s p e c i f i c d i s t i n c t i o n i n Laminaria. Burrows (1964, 1964a) studied the eff e c t of temperature on the blade morphology and- anatomy of L. .saccharina and L. agardhil, Results from t h i s study indicate that temperature can. modify such c h a r a c t e r i s t i c s as b u l l a t i o n , mucilage duct production, thickness of blade, and.cell s i z e . Sundene (1962, 1964, 1964a) observed considerable morphological p l a s t i c i t y in.two. forms of L. d i g i t a t a subjected to transplantation. He noted that plants from exposed regions when grown i n sheltered areas developed a broader blade and became more bullate, thus resembling the form native to the sheltered areas. Evidence of morphological p l a s t i c i t y presented by Burrows and Sundene suggests a closer r e l a t i o n s h i p between the members of the above l i s t e d pairs than accorded them at present. C l a r i f i c a t i o n of the taxonomy of, Laminaria depends upon (l ) d e f i n i t i o n of phenotypic p l a s t i c i t y f o r those character-is t i c s ' employed i n delineating species and (2) an understand-ing of., the genetic relationships between species-. The f i r s t can be accomplished by culture and transplant studies similar to those of Sundene (1962, 1962a, 1964) and Burrows (1964, 1964a). The second may be accomplished by i n t e r f e r t i l i t y studies comparable to those of Sundene (1958) and Tokida and Yabu (1962), and by c y t o l o g i c a l studies similar to those of Kemp and Cole (1961), Evans (1964), and others. The geographical ranges of a l l the species studied, with the exception of L. groenlandica, have been extended i n t h i s study. Further, the habitats to which some species were previously thought to be r e s t r i c t e d have been enlarged. Laminaria saccharina and L. complanata were thought to be limited to sheltered waters; both have been revealed, i n exposed regions, L. saccharina subt i d a l l y and L. complanata ' I n t e r t i d a l l y . Laminaria s e t c h e l l i i , L. ephemera, and L. s i n c l a i r i i were previously thought to be r e s t r i c t e d to exposed areas. They have recently been disclosed s u b t i d a l l y i n sheltered areas. . These r e s u l t s emphasize the importance of an integrated submarine and i n t e r t i d a l exploration program for studying d i s t r i b u t i o n s of benthic organisms which occur both.inter-t i d a l l y and s u b t i d a l l y . This approach has not been exten-s i v e l y employed i n the past. L i f e History and Morphology Both i n t e r t i d a l l y occurring L. groenlandica long and short stipe forms and L..saccharina :produced s o r i i n October-January, and again i n June. Parke (1948) observed s o r i of L..saccharina In Great B r i t a i n throughout the year. She noted the greatest numbers of f e r t i l e plants between October and March. Her investigation encompassed i n t e r t i d a l and subtidal plants. F a i l u r e of i n t e r t i d a l L. saccharina i n British.Columbia to produce s o r i throughout the year may r e s u l t from damage to p o t e n t i a l l y reproductive.tissues during low daytime spring tides. The reproductive cycle of subtidal plants i n B r i t i s h Columbia i s unknown. Parke (1.948) noted that sporophytic tissue must be at least 6 months old for soral i n i t i a t i o n i n L. saccharina, whereas Kireeva and Schapova (1938) stated that t h i s species does not reach maturity u n t i l the t h i r d year of growth. In t h i s study, a l l observed L. saccharina produced s o r i i n the October-November period, including plants which f i r s t became conspicuous i n February and September of the same year. Plants which produced s o r i i n June were at, l e a s t 9 months old. It would.appear from my study that soral production by B r i t i s h Columbia plants i n the winter i s independent of 6o the age of sporophytic tissue, but i n June may be dependent in.part upon the age of t h i s tissue. Much morphological v a r i a b i l i t y of gametophytes under culture conditions was observed for L. saccharina and to a much lesser degree f o r L. groenlandica (Fig. 55, 6 3 ) . Laminaria saccharina meiospores produced filamentous gameto-phytes under crowded culture conditions and i n -situations of high,temperature and. low s a l i n i t y . Under conditions of low temperature and. high s a l i n i t y and i n uncrowded culture conditions the meiospores produced female gametophytes of a single c e l l , which functioned as the oogonium, and male plants of several c e l l s , most of which functioned as antheridia. In s i t u observations of L. saccharina gameto-phytes i n the winter revealed u n i c e l l u l a r female plants but no male plants. Filamentous gametophytes have been observed by Kemp and Cole (1961) for Nereocystis luetkeana under conditions of high temperature; by Sundene (1962) for L. d i g i t a t a under conditions of feeble day l i g h t , low temperature, and unchanged media; and by Kain (1964) for L. hyperborea under conditions of high temperature, low illumination, and unchanged media. In many species' of Laminariales vegetative growth of gametophytes can occur at temperatures above those conducive to gamete production (Myers, 1928; Schreiber, 1930; Hollenberg, 1939; Salt©, 1956, 1956a; Kemp and Cole, 1 9 6 l ; Sundene, 1963). 61 From the above studies it.would appear that Laminaria can produce one of two forms of gametophytes depending upon environmental conditions: a small gametophyte consisting of few c e l l s most of which function as gametangia, and a large filamentous gametophyte which may or may not produce gametangia while subjected to the environmental conditions under which the gametophyte developed. The small form appears adapted for immediate sporophyte production. However, f e r t i l i z a t i o n l i m i t s the reproductive pot e n t i a l of the female plant. The large form may remain in.the vegetative state u n t i l conditions become favorable for sporophyte production. Consideration of re s u l t s obtained from culture studies, here and elsewhere, and of the l i f e history of L. saccharina as observed i n B r i t i s h Columbia have led.me to the following opinion. Gametophytes developing from meiospores released in June are of the large and vegetative type. These plants remain i n the vegetative condition through most of the summer while conditions of high temperature and low s a l i n i t y per-s i s t . In the late summer when the temperature i s lower and s a l i n i t y higher, gametes are produced and f e r t i l i z a t i o n occurs. The resultant sporophytes are first |c o n s p i c u o u s in early f a l l . Gametophytes r e s u l t i n g from the f a l l and winter crops of meiospores develop into small gametophytes and are almost immediately f e r t i l e . The resultant sporophytes f i r s t become conspicuous i n February. Plants thought to be abnormal sporophytes are produced by L. saccharina and L. groenlandica long stipe form under culture conditions of high temperature and low s a l i n i t y '(Fig.. 55, 63) . Similar plants have been described from culture by Schreiber (1930), Segi and Kida (1958), Kemp and Cole (1961), and Tokida and Yabu (1962). These plants have been referred to as parthenogenetic sporophytes. Only Kemp and Cole (1961) were able to demonstrate that these plants had the haploid number of chromosomes. Intercalary growth observed i n both L. groenlandica long and short stipe forms and L. saccharina i s greatest i n the lower-most portion of the blade, and decreases d i s t a l l y . The region of greatest.longitudinal growth coincides with the region of greatest l a t e r a l growth (Fig. 5 8 - 6 l , 64 -67) . Similar d i s t r i b u t i o n s of i n t e r c a l a r y growth have been observed by F a l l i s .(1916) for L. .saccharina i n Washington, by Parke (1948) for L. saccharina i n Great B r i t a i n , and by Hasegawa (1962) for L. angustata i n Japan. Sundene (1964) noted considerable v a r i a t i o n in. the d i s t r i b u t i o n of i n t e r -calary growth i n L. d i g i t a t a . During June he observed some plants In which the greatest increments of growth occurred d i s t a l to the basal 5 cm of the blade base. There i s a seasonal v a r i a t i o n i n the growth rates of both L. saccharina and L. groenlandica long and short stipe forms as observed i n t h i s study (Fig. 68; Table I ) . In both forms of L. groenlandica vigorous growth commences in February when blade regeneration i s i n i t i a t e d . Rapid growth-continues u n t i l May-June. The growth rate then diminishes u n t i l November when sorus development i s i n i t i a t e d . Growth i s n e g l i g i b l e from November u n t i l February. In L. saccharina the growth rate increases throughout the spring and summer 63 u n t i l September afte r which time It., decreases. No measurable growth.was observed i n December and January. Seasonal v a r i a t i o n of growth for L. groenlandica i s i n close accord with that reported by others in that the most active growth occurs i n . l a t e winter and i n spring (Printz, 1926, for L. hyperborea; Parke, 1948, for L. saccharina; Tseng, Wu, and Sun, 1957, for L. japonica; Hasegawa, 1962, for L. angustata; Sundene, 1964, for L. d i g i t a t a ) . The period of active growth i n L. saccharina as observed i n B r i t i s h Columbia d i f f e r s from the above species i n that i t extends from late winter u n t i l September. Local D i s t r i b u t i o n i n Relation to Oceanographic Factors The approach used i n t h i s study i s an extension of a d i s c i p l i n e i n i t i a t e d on the west coast of North America by the late William Albert Setchell at the turn of the century. Setchell (1893, 1917, 1935) attempted to explain marine a l g a l d i s t r i b u t i o n s on the basis of temperature d i s t r i b u t i o n at a time when the marine environment was poorly defined. Following his studies l i t t l e was done i n t h i s f i e l d u n t i l the recent studies of Scagel (1961, 1 9 6 l a , 1962, 1963, 1963a). Scagel related the d i s t r i b u t i o n s of marine algae in.the northeast P a c i f i c to various oceanographic and meteorologic conditions. . He has pointed out that one can suggest the role of p a r t i c u l a r environmental factors i n governing a l g a l d i s t r i b u t i o n s by r e l a t i n g environmental conditions t o . a l g a l d i s t r i b u t i o n s , and that the ultimate determination of the e f f e c t of any factor on.algal 64 d i s t r i b u t i o n s may be established by both experimental f i e l d and laboratory studies (Scagel, 1962). In the present study, the d i s t r i b u t i o n s of L.'. saccharina and L . groenlandica are related to factors that are con-spicuously d i f f e r e n t between the habitats of these species: v i z . , water motion, temperature, and s a l i n i t y . Conditions of l i g h t and substratum were considered i n s i g n i f i c a n t because the species are known to have approximately the same v e r t i -c a l and l a t i t u d i n a l d i s t r i b u t i o n s and substratum requirements. The significance of b i o l o g i c a l and chemical factors other than those related to s a l i n i t y were only investigated i n a super-f i c i a l fashion, and should be followed up in future studies. D i s t r i b u t i o n a l studies on the two forms of L, groenlandica and on L. saccharina indicate that the former are excluded from areas having high temperature and low s a l i n i t y , and the l a t t e r i s excluded from areas of surf action (Fig. 71) . Further, L. groenlandica long stipe form i s found only i n areas subjected to considerable surf, and i s replaced by the short stipe form in.quieter waters. Evaluation of responses of L_. groenlandica long stipe form and L. saccharina sporophytes and gametophytes to several temperature and s a l i n i t y situations demonstrates that the range of tolerance to these factors i s narrow i n the former plants but wide i n the l a t t e r (Fig. 73, 74; Tables V, VI, VII). Transplant studies on L. groenlandica long and short stipe forms i l l u s t r a t e that these plants are r e s t r i c t e d to narrow ranges of both temperature and s a l i n i t y . They are able to grow well i n Burrard Inlet, an area where they do not occur naturally, only from December to June, the period when temperature i s low and s a l i n i t y high (Fig. 8 6 ) . In the long stipe form vegetative growth and soral production proceed in the sheltered transplant area, and are not dependent upon surf action. Responses of gametophytes of L. groenlandica long stipe form.and L. saccharina to water of d i f f e r e n t origins indicate that conditions most favorable for growth are encountered in water from the plant's natural habitat (Fig. 75, 7 6 ) . The d i f f e r e n t i a l responses may r e s u l t from differences in. Ionic r a t i o s of the seawater constituents, or from the presence and/or absence of organic substances. Physiological res-ponse of sporophytes of both species was similar to. a l l water types tested. From the above culture and transplant studies the following conclusions are drawn: (1) The l o c a l d i s t r i b u t i o n of L. groenlandica i s independent.,of surf conditions. This, i n d i r e c t l y , suggests that the two forms.are eco l o g i c a l adaptations of one geno-type. (2) Temperature and.salinity acting upon the sporo-phyte of the two forms of L. groenlandica determine t h e i r d i s t r i b u t i o n s about Vancouver Island. Considering the temperature and s a l i n i t y conditions, the gametophytes of these forms could produce normal sporophytes during the winter and early s p r i n g . i n . a l l l o c a l environments inhabited by L. saccharina. In the summer sporophyte production 66 would be prohibited. No .attempt has been,made to separate the e f f e c t s of temperature and s a l i n i t y on the physiology and growth of L. groenlandica. The culture experiments indicate that the best s i t u a t i o n for growth i s low temperature i n combination with high s a l i n i t y . The detrimental e f f e c t s of high tempera-ture may be of f s e t by/high s a l i n i t y , and the harmful e f f e c t s of low s a l i n i t y , by low temperature. E a r l i e r culture studies have i l l u s t r a t e d the importance of temperature i n l i m i t i n g growth and reproduction in Laminaria (Burrows, 1961, in L. saccharina; Tokida and Yabu, 1962, i n L. r e l i g i o s a ; Kain, 1964, i n L. hyperborea; Sundene, 1964, i n L. saccharina, L. hyperborea, L. d i g i t a t a ) . Transplant studies on L. japonica (Tseng, Wu, and Sun, 1957) and on A l a r i a esculenta and L. d i g i t a t a (Sundene, 1962, 1962a, 1964) have yielded r e s u l t s similar to those recorded here. These authors found that plants transplanted from a colder to a warmer region were able to grow successfully through the winter and spring months but;were severely injured or k i l l e d by summer temperatures. Sundene 1s (1964a). studies on L. saccharina, L. hyperborea, and L. d i g i t a t a i n Norway indicate that s a l i n i t y i s a minor factor in determining regional d i s t r i b u t i o n s . (3) The l o c a l d i s t r i b u t i o n of L. saccharina i s indepen-dent of. temperature and s a l i n i t y with the possible exception of extreme estuarine conditions. Its d i s t r i b u t i o n i s res-t r i c t e d by surf action since (a) the plant was observed i n areas subjected to surf only when:positioned below.the dire c t influence of t h i s factor and (b) the delicate nature of the plant, i t s thin blade and stipe, and f i n e holdfast do not equip i t f o r survival i n areas subjected-to surf. (4) Competition between L. saccharina and L. groenlandica may be an important factor influencing the d i s t r i b u t i o n s of these species i n areas not subjected to surf. This i s i n d i -cated by the responses of gametophytes to water of d i f f e r e n t o r i g i n s . V e r t i c a l D i s t r i b u t i o n of Laminaria saccharina\\ The upper l i m i t s of L. saccharina as observed i n Burrard Inlet are thought to be determined by a i r temperature and i n s o l a t i o n . ( F i g . 7 7 ) . The tide i s considered an i n d i r e c t agent Influencing the upper l i m i t s , as i t determines the time and duration of exposure to a i r and dir e c t i n s o l a t i o n . Culture studies show that prolonged exposure .to l i g h t of an i n t e n s i t y above the saturation l e v e l (5.00 to 700 f t - c ) r e s u l t s i n a growth decrease and i n tissue injury (Fig. - 8 5 ) . B r i e f exposures to these i n t e n s i t i e s do not appear to be detrimental (Fig. 84). These studies suggest that submarine l i g h t , i n t e n s i t i e s may play.'a s i g n i f i c a n t role i n determining the upper l i m i t s . However, because of the greater Intensity and spectral d i s t r i b u t i o n of l i g h t s t r i k i n g emerged plants, d i r e c t i n s o l a t i o n would be more harmful. Other workers have recorded considerably/lower satur-ation, l e v e l s . Segi and Kida (1958) found the best growth of Fndaria undariodes occurred between. 1500 to 2000 lux 68 (ca. 140 to. 180 f t - c ) ; at 4800 lux (ca. 440 f t - c ) growth was i n h i b i t e d . Kain (1964, 1965) reports a saturation value of 1000 lux (ca. 93 f t - c ) at 10°C for sporophytes and 350 lux (ca. 33 f t - c ) f o r gametophytes of L. hyperborea. These workers employed ligh t , sources of d i f f e r e n t spectral q u a l i t i e s than those used.in t h i s study, thus a comparison of r e s u l t s i s not meaningful. The lower l i m i t s of L. saccharina as observed i n Burrard Inlet are thought to r e f l e c t the compensation depth of t h i s species. There was no evidence of competition with- other algae and the substratum was uniform from above the upper l i m i t s to below the lower l i m i t s of v e r t i c a l d i s t r i b u t i o n . The culture studies on sporophytes of L. saccharina have shown the l i g h t compensation.point.to be between 20 to 40 f t - c at,;10°C (Pig. 84). Increase i n blade sur-face was approximately Vfo per day at 180 f t - c and 10°C (Pig. 8 5 ) . Gametophytes cultured under 20 f t - c illumination at 10°C produced normal sporophytes. Lower i n t e n s i t i e s were not tested. Kain (1964, 1965) found 20 lux. (ca. 1.9 f t - c ) at 10°C to be the, minimal l i g h t i n t e n s i t y for development of gametophytes and young sporophytes of L. hyperborea. The lowest values of submarine illumination, i n Burrard Inlet are encountered.during, the winter, months. Laminaria saccharina gametophytes resulting.from the June crop of meiospores should be able to e s t a b l i s h themselves at, a greater depth than.the winter population because of the greater l i g h t , i n t e n s i t y i n June. The resultant sporophyte 69 generation could be eliminated, as the l i g h t i n t e n s i t y may f a l l below the compensation point i n the winter months (Fig. 8 3 ) . During t h i s period gametophytes would not succeed to the depth populated by the summer ones. I n s u f f i c i e n t data on fluctuations of the. lower l i m i t s and on the l i f e h i story of the subtidal plants precludes precise d e f i n i t i o n of agents responsible for the lower l i m i t s . 70 SUMMARY Ten species of Laminaria are recognized by me for the northeast P a c i f i c : L. groenlandica Rosenv.; L. f a r l o w i i Setchell; L. saccharina (L.)>Lamour.; L. complanata (Setchell et Gardner) Set c h e l l ; L. s e t c h e l l i i S i l v a ; L. dentigera Kjellman; L. longipes Bory; L. s i n c l a i r i i (Harvey ex Hooker f. et Harvey), Farlow, Anderson et Eaton; L. ephemera Setchell; and L... yezoensis Miyabe. Laminaria cordata Dawson i s considered conspecific with L. saccharina, L. personata Setchell and Gardner i s regarded as conspecific with L. yezoensis, and L. platymeris De l a Pyl. (sensu Setchell and Gardner, 1925) i s considered conspecific with L. groenlandica. Pour forms of L. groenlandica are recognized for the northeast P a c i f i c : long stipe form, short stipe form, f l a t stipe form and shade form. These forms are not considered by me at present as legitimate taxonomic e n t i t i e s but are distinguished merely to provide a means of f a c i l i t a t i n g discussion u n t i l such time as a c r i t i c a l monographic study f o r the entire genus i s made. The geographical ranges of a l l the species studied, with the exception of _L. groenlandica, have been extended i n t h i s study. Further, the habitats to which a l l ten species were previously/thought to be r e s t r i c t e d have been enlarged. LatiyLnaria saccharina and L. complanata were thought to be limited to sheltered waters: both have been revealed in exposed regions. Laminaria s e t c h e l l i i , L. ephemera, and L. s i n c l a i r i i were previously thought to be r e s t r i c t e d to 71 exposed areas. They have recently been disclosed s u b t i d a l l y in sheltered areas. Laminaria saccharina and L. groenlandica produced s o r i i n the l a t e spring.and winter. Sorus production by L. saccharina i n the winter i s independent of the age of the sporophyte tissue but in spring may be dependent i n part upon the age of t h i s t i s s u e . New sporophytes of L. groenlandica appeared throughout the year, whereas those of L. saccharina appeared i n ;late winter and early f a l l . Depending upon culture conditions, two morphologically d i s t i n c t forms of gametophytes were produced by both species: large gameto-phytes i n conditions of high temperature and low s a l i n i t y ; and small gametophytes i n .conditions.of low temperature and high s a l i n i t y . ' Abnormal sporophytes were observed under conditions conducive to formation of large gametophytes. These may be parthenosporophytes. Patterns of growth for the blades of the two species were e s s e n t i a l l y the same. The growth rate decreased with increase i n distance from the blade base, and the p o s i t i o n of greatest longitudinal growth coincided with the position of greatest l a t e r a l growth. The d i s t r i b u t i o n s of L. saccharina and the two forms of L. groenlandica about.Vancouver Island were correlated with temperature, s a l i n i t y , and water motion. The two forms 72 of L. groenlandica were absent from areas of high temperature and low s a l i n i t y , L. saccharina was absent from areas sub-jected to surf. These f i e l d conclusions were subjected to laboratory and f i e l d tests involving gametophytes and sporophytes of both species. The d i s t r i b u t i o n s of the two forms of L. groenlandica can be explained on the basis of temperature and s a l i n i t y d i s t r i b u t i o n s . Both forms require low temperature and high s a l i n i t y for s u r v i v a l . Laminaria saccharina has a wide range of tolerance to temperature and s a l i n i t y . Surf appears to be the agent 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 t h i s species. The upper l i m i t s of L. saccharina, as observed i n Burrard Inlet, are thought to be d i r e c t l y related to a i r temperature and i n s o l a t i o n and i n d i r e c t l y related to t i d a l c h a r a c t e r i s t i c s . The lower l i m i t s appear to r e f l e c t the compensation depth of t h i s species. The above studies i l l u s t r a t e the p o t e n t i a l value of combined f i e l d and laboratory studies i n solving problems of a l g a l taxonomy, morphology, physiology, and ecology. In.the laboratory i t i s possible to define and control environmental parameters and to monitor the organisms studied. However, culture conditions as used at present are poor approximations of the natural environment. In the f i e l d one deals with a natural environment, however, in t h i s case the environment i s usually poorly defined and the specimens are not read i l y accessible for monitoring. Employ-ment of modern oceanographic instrumentation and SCUBA gear a s s i s t s In overcoming these d i f f i c u l t i e s . 74 BIBLIOGRAPHY Anon., 1948. Observations of seawater temperature, s a l i n i t y , and density on the P a c i f i c coast of Canada. Fish. Res. Bd. Can., Nanaimo. Pacif. Oceanogr. Group. Vol. 4, 1940, 1941. 131 pp. 1952. Observations of seawater temperature and s a l i n i t y on the P a c i f i c coast.of Canada. Fish'. Re_s. Bd. Can., Nanaimo. Pacif. Oceanogr. Group. Vol. ' 1 1 , 1951. 62 pp. 1955. Physical and chemical data record. Juan de Fuca S t r a i t Project 1951-52. Fish. Res. Bd. Can., Nanaimo. Pacif. Oceanogr. Group. 182 pp., 2 f i g s . . ; 1957. Observations of seawater temperature and s a l i n i t y on the P a c i f i c coast of Canada. Fish. Res, Bd. 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Renfrewla paryula, a new kelp from Vancouver Island. Postelsia .(1906) :245-274, pi's. 16-19. Hasegawa, Y. 1962. An ecological study of Laminaria angustata Kjellman on the coast of Hldaka Prov,, Hokkaido. B u l l . Hokkaido Reg. Fish. Res. Lab. 24:116-138. ,(Not seen). 76 Hollenberg, G.J. 1939. Culture studies of marine algae. I. Eisenia arborea. Am. Jour. Bot. 26:34-41, 3 f i g s . Holmes, R.W. 1957. Chapter 6. Solar radiation, submarine daylight, and photosynthesis. _In: Treatise on marine ecology and paleoecology. Vol. I, Ecology. Geol. Soc. Am. Mem. 67. Pp. 109-128, 1 f i g s . , 6 tables. Jerlov,. N. L. 1951. Optical studies of ocean waters. Reports of the Swedish Deep-Sea Expedition. Vol. I l l , Physics and Chemistry. No. 1, 1-59, 51 f i g s , 33 tables. Kain, J.M. 1964. Aspects of the biology of Laminaria .hyperborea. I I I . Survival and growth of gametophytes. • J. Mar. Bi o l . Ass. U.K. 44:415-433, 10 f i g s . 1965. Aspects of the biology of Laminaria hyperborea IV. Growth of early sporophytes. J. Mar. B i o l . Ass. U.K. 45:129-143, 8 f i g s . Kemp, L. and K. Cole. 196l. Chromosomal alternation of generations in Nereocystis luetkeana (Mertens) Postels and Ruprecht. Can. J. Bot. 39:1711-1724, 2 p i s . , 1 table. Kireeva, M.S. and T.P. Schapova. 1938. Rates of growth, age and sporebearing•of Laminaria saccharina and _L. d i g i t a t a i n Kola Fjord. Trans. Inst. Mar. Fi s h . 0ceanogr„ Moscow 7:29-58. (English summary). (Not seen). Kjellman, F.R. 1877. Bidrag t i l l Kannedomen af Kariska hafvets Algvegetation. Ofver. af K. Vet.-Akad. Forhand., No. 2, 3-30, 1 table. 1883. The algae of the A r c t i c Sea. K. Sv. Vet.-Akad. Handl. 20:1-350, 31 p i s . Miyabe, K. 1902. On the Laminariaceae of Hokkaido. (English e d i t i o n by J. Tokida, 1957) Jour. Sapp. Agr. C o l l . , Fac. Agr. Hokkaido Univ. 1:1-50, 29 p i s . Myers, M.E. 1925. Contributions towards a knowledge of the l i f e - h i s t o r i e s of the Melanophyceae. Univ. C a l i f . Publ. Bot. 13:109-124, 3 p i s . 1928. The l i f e - h i s t o r y of the brown alga Egregia menziesii. Univ. C a l i f . Publ. Bot. 14:225-245, p i s . 49-52, 1 table. Parke, M. 1948. Studies on the B r i t i s h Laminariaceae. I. Growth in Laminaria saccharina (L.) Lamour. J. Mar. B i o l . Ass. U.K. 27:651-709, 10 f i g s . , p i s . 5-12, 8 tables. 77 Printz, H. 1926. Die Algenvegetation des Trondhjemsfjorden. Skr. norske Vedensk-Akad., No. 5, 1-273. (Not seen). Saito, Y. 1956. An ecological study of Undaria pinnatifIda Sur. I. On the Influence of environmental factors upon the development of gametophytes. B u l l . Jap. Soc. S c i . Pish. 22:229-34, 6 f i g s . , 1 p i . 1956a. An ecological study of Undaria p i n n a t i f i d a Sur. I I . On the influence of environmental factors upon maturity of gametophytes and early development of sporophytes. B u l l . Jap. Soc. S c i . Pish. 22:235-239, 5 f i g s . Saunders, de A. 1901. Papers from the Harriman Alaska Expedition. XXV. The algae of the expedition. Proc. Wash. Acad. Sciences 3:391-486, p i s . 10-26. Scagel, R.P. 1957. An annotated l i s t of the marine algae of B r i t i s h Columbia and northern Washington (including keys to genera). B u l l . Nat. Mus. Can. No. 152, 286 pp., 1 f i g . 1 9 6 l . The d i s t r i b u t i o n of certain benthonic algae in Queen Charlotte S t r a i t , B r i t i s h Columbia, in r e l a t i o n to some environmental f a c t o r s . P a c i f i c Science 14: 494-539, 51 f i g s . 1 9 6 l a . A synthetic approach to some problems in marine a l g a l ecology. _n: Recent Advances i n Botany, pp. 175-180, 1 f i g . Univ. of Toronto Press. 1962. Benthic a l g a l productivity in the north P a c i f i c with p a r t i c u l a r reference to the coast of B r i t i s h Columbia. Proc. Ninth P a c i f i c S c i . Congress, 1957. 4:181-187. 1963. Some problems in a l g a l d i s t r i b u t i o n i n the north P a c i f i c . Proc. 4 t h International Seaweed Symposium, B i a r r i t z , , pp. 259-264. 1963a. D i s t r i b u t i o n s of attached marine algae in r e l a t i o n to oceanographic conditions i n the northeast P a c i f i c . __: M.J. Dunbar (Ed.), Marine D i s t r i b u t i o n s , Univ. of Toronto Press. Roy. Soc. Canada Spec. Publ. No. 5, 37-50, 11 f i g s . Schreiber, E. 1930. Untersuchungen uber Parthenogenesis, Geschlechtsbestimmung und Bastardierungsvermogen bei Laminarien. Planta 12:331-353. (Not seen). Segi, T. and .W. Kida. 1958. Studies on the development of Undaria undarioides (Yendo) Okamura. I I . On the development of the sporophytes and influence of l i g h t on i t . Rep. Pac. Pish. Pref. Univ. Mie 3:236-246, p i s . 6 - 8 , 7 f i g s , , 1 table. 78 Setchell, W.A. 1891. In: Anderson, C.L. L i s t of C a l i f o r n i a marine algae with notes. Zoe 2:217-225. m 1893. On the c l a s s i f i c a t i o n and geographical d i s t r i -bution of the Laminariaceae. Trans. Conn. Acad. Arts S c i . 9 :333-375. 1901. Notes on Algae. I. Zoe 5 1 2 1 - 1 2 9 . 1 _____ 1905. Regeneration among kelps. Univ. C a l i f . Publ. Bot. 2:139-169, P i s . 15-17. 1908,: C r i t i c a l notes on the Laminariaceae. La Nuova Notarisia 19:90-101. 1912. The kelps of the United States and Alaska. Appendix K. In: Cameron, F.K. A preliminary'report on the f e r t i l i z e r resources of the United States. Appendix K. United States Senate Document No. 190, 130-178. 1917. Geographical d i s t r i b u t i o n of marine algae. Science 45:197-204. . 1935. Geographical elements of the marine f l o r a of the North P a c i f i c Ocean. Amer. Nat. 69:560-577, 12 f i g s . S etchell, W.A. and ;N.L. Gardner. 1903. Algae of North-western America. Univ. C a l i f . Publ. Bot. I : l 6 5 - 4 l 8 , p i s . 17-27. , 1924. Phycological contributions, VII. Univ. C a l i f . Publ. Bot. 13:1-13. 192.5. The marine algae of the P a c i f i c Coast of North America. Part I I I . Melanophyceae. Univ. C a l i f . Publ. Bot. 8 :383-898, p i s . 34-107. S i l v a , P.O. 1957. Notes on P a c i f i c marine algae. Madrona, 14:41-80. Smith, G.M. 1944. Marine algae of Monterey Peninsula, C a l i f o r n i a . Stanford Univ-. Press, ix + 622 pp., 98 p i s , Strickland, J.D.H. 1957. Solar .radiation penetrating the ocean. F i s h . Res. Bd. Can., Pacif. Oceanogr. Group. Nanaimo. 46 pp., 9 f i g s . , 4 tables. Strickland, J.D.H. and T.R. Parsons, i 9 6 0 . A manual of sea water analysis. (With special reference to the more common micronutrients and to p a r t i c u l a t e organic material.) . F i s h . Res. Bd. B u l l . No. 125, v l + 185 pp., . 13 tables. 79 Sundene, 0. 1958. I n t e r f e r t i l i t y between forms of Laminaria d i g i t a t a . Nytt Mag. Bot. 6:121-128, 3 f i g s . , 4 p i s . , 3 tables. 1962. Growth in the sea of Laminaria d i g i t a t a sporophytes from culture. Nytt Mag. Bot. 9 :5-24 , 18 f i g s . , 3 p i s . , 1 table. 1962a. The implications of transplant and culture experiments on the growth and d i s t r i b u t i o n of A l a r i a esculenta, Nytt Mag. Bot. 9:155-174, 8 f i g s . , 6 p i s . , 1 table. 1964. The ecology of Laminaria d i g i t a t a in Norway in view of transplant experiments. Nytt Mag. Bot. 11:83-107, 20 f i g s . i t9 -o 124° 125° 1 2 7 o 127° 127° 1 2 ? o 126° 128° 1 2 7 o 128° 1 2 7 o 128° 127° 127 126° 127° 125° 125° 3 2 1 w 4518 RFS UBC 9610 52; w 4419 RFS UBC 8181 24; w 4613 RFS UBC 8218 02 w 926 RFS UBC 2107 10 w 7035 RFS UBC 16936 Ol! w UBC 2485 34 w 7370 RFS UBC 16940 32 w 670 LD UBC 18057 26! w 170 LD UBC .14754 47 w 500 LD UBC 17006 46 w 8173 RFS UBC 17859 32, w 8081 RFS UBC 17846 55, w 8264 RFS UBC. 17865 36 w 7974 RFS UBC 17870 34 w 2839 RFS UBC 7391 25, w 8410 RFS UBC 19339 21 w 2419 RFS UBC 4111 2 5 , w 8341 RFS UBC 19336 29 w 7922 RFS UBC 17893 02 w 2633 RFS UBC 4097 27, w 7760 RFS UBC 17868 37, w 8150 RFS UBC 17861 4o ' w 2202 RFS UBC 4106 2 6 ' w 3450 RFS UBC 7409 11J w 3085 RFS -UBC 7380 03 w 3184 RFS UBC 7199 Appendix I, Section R, cont. Voucher Specimens of Laminaria s e t c h e l l i i Area Location Approximate Latitude Position Longitude C o l l No. Acc. No. II I Ogden Point Breakwater 48°. 24* N 123° 123° 23 w UBC 12677 I I I Otter Point 48° 49° 49° 21 N 49 w 3851 RFS UBC 12173 I I I Perez Rocks 25, N 126° 126° 37, w 2886 RFS UBC 7418 I I I Sharp Point 21 N 16 w 2986 RFS UBC 7379 III •• Sheringham Point 48° 5^0 5°o 22 N 123° 55, w 191 TBW UBC 3448 I I I Solander Island 07, N 127° 127° 56 w 8293 RFS UBC 17851 I I I Spring Island 0 0 , N 25, w UBC 7 4 l l I I I Tofino 49° 48° 09, N 125° 129° 125° 55, w 569 LD UBC 18055 III Triangle Island 55, N 0 5 , w 8436 RFS UBC 19340 III -Wouwer Island N 22 w 2028, RFS UBC 4109 IV Cape Sutile 5°o 53 N 128° 0 3 ! w 6409 RFS UBC 16998 IV Roller Bay, Hope Island 5°o 5°o 48° 48° 5 6 N 127 56 w 6323 RFS UBC 16993 TV Rosebush Island 14 N 1 2 5 ° 1 2 4 o 09 w 3626 RFS UBC 6947 V Neah Bay 22 N 37, w UBC 399 V-(D) Salmon Bank 33 N 123 10 w 1125 RFS UBC- 2363 Appendix I, Section L. Voucher Specimens of Laminaria dentigera I Cape Agagdak, Adak I Cape Sarichef, Unimak I Chicagof Point, Attu I Murder Point, Attu I Ocean Cape, Yakutat I Pasagshak Point, Kodiak 5 1 o 540 5 2 o 52° K 57 60, N 35, N 27, N 48 N 31, N 25 N .0 W • 4070 RFS UBC 8277 w 4204 RFS UBC 8278 E 4o49 RFS UBC 8248 E 4003 RFS UBC 8258 W 4419 RFS • UBC 8315 W 4377 RFS UBC 8175 Appendix I I . Observed Specimens of Particular Taxonomic Significance Legend. Co l l e c t o r s . (EYD) E. Y. Dawson (MP) M, Poslie (NLG) N, L. Gardner (WAS) W. A. Setchell (S&G) W. A. Setchell :(JT) J. Tokida (JMW) J. M. Weeks Herbaria. (uc) University of (AHPH). Allan Hancock University of Southern C a l i f o r n i a . Appendix I I . Observed Specimens of Particular Taxonomic Significance Specimen Location Collector No. Date Herb No. L. b u l l a t a f. amplissima (= L. groenlandica short stipe) L. bu l l a t a f. angusta (= L. groenlandica shade form) L. b u l l a t a f . subsimplex TYPE (= L. groenlandica short stipe) L. cordata HOLOTYPE (= L. saccharina) L. ephemera TYPE L. personata TYPE .(= yezoensis) L_. platymeris (= L. groenlandica long stipe) L_. platymeris (= L. groenlandica.long stipe) L, solidungula L. yezoensis Esquimalt, B.C., Canada Whidbey Island, Wash., U.S.A. Whidbey Island, Wash., U.S.A. Santa Catalina Island, C a l i f o r n i a , U.S.A. Carmel Bay, Ca l i f o r n i a , U.S.A. Sitka, Alaska, U.S.A. Sooke Harbour, Vancouver Island, Canada San Juan Island, Wash., U.S.A. Dlevie Bay, Spitsbergen, Norway Nishiwada, Nemuro, Hokkaida, Japan S&G, 6 8 0 a May, 1 9 0 1 S&G 1 0 9 S&G 157 EYD•56OO Dec. 1 , 1 9 4 8 JMW A p r i l 29, I 8 9 8 NLG 3 9 5 1 July,. 1 9 1 7 WAS June 3 0 , . 1 9 3 0 NLG M P JT 3 5 0 Sept. 2 , . 1 9 2 5 uc 9 6 9 2 6 uc 9 6 9 3 2 uc 9 6 9 2 0 AHFH 3 6 9 2 2 uc 9 6 9 6 5 UC 2 6 6 4 9 1 UC 4 6 3 9 9 6 uc 3 9 6 7 0 1 uc 9 7 1 3 2 UC 5 4 3 9 2 91 Table I. Longitudinal Growth Rates i n Situ of Laminaria groenlandica No. PI. I n i t i a l Date Mean I n i t i a l P i n a l Date Mean Increase Length Length/Day Laminaria groenlandica short stipe form 5 July 20, 1962 12.4 cm July 30, 1962 0.59 cm/day 4 July 30, 1962 18.8 cm Aug. 14, 1962 O.36 cm/day 4 Aug. 14, 1962 31.8 cm Sept, .12, 1962 0.38 cm/day 8 Apr. 26, 1963 90.3 cm May 22, 1963 0.64 cm/day 4 May 22, 1963 . 106.4 cm June. 9, 1963 0.71 cm/day jaminaria groenlandica long stipe form 8 July 30, 1962 21.3 cm Aug. 14, 1962 o.4o cm/day 3 Aug. 14, 1962 26.9 cm Sept, .12, 1962 0.15 cm/day 3 Sept, ,12, 1962 31.3 cm Nov. 15, 1962 0.05 cm/day 5 Apr. 26, 1963 72.4 cm May 22, 1963 1,68 cm/day 3 May 22, 1963 .118.2 cm June 9, 1963 0.26 cm/day 92 Table I I . Stipe Length/Blade Width as Determined.in Situ f o r Laminaria groenlandica -Date No. PI. Mean Stipe Mean Blade Mean Length Width • St/W Laminaria groenlandicalong stipe form 1. July 2 0 , 1962 8 3.1 cm 4.2 cm 0.73 2. Aug. 14, 1962 8 3.2 cm 6.9 cm : 0.46 3 . Apr. 26, 1963 5 4 . 7 cm. 16.0 cm :0.29 4. May 22,. 1963 5 7.0 cm 18.2 cm 0.38 5. Apr. 26, 1963 10 4.9 cm 14.9 cm 0.32 6. Aug. 4, 1963 8 10.3 cm 14.3 cm 0.72 7. Aug. 4, 1963 17 4 . 9 cm 5.7 cm 0.85 8. Sept. 15, 1962 9 1.3 cm 3.1 cm o . 4 i Laminaria groenlandica short stipe form 9. July 20, 1962 5 1.3 cm 5.8 cm 0.22 10. Aug. 14, 1962 - 5 2.0 cm 14.8 cm 0.13 11. Apr. 26, 1963 8 2.4 cm 12.6 cm 0.19 12. May 22, 1963 8 3.0 cm 14.5 cm 0.20 13. Apr. 26, 1963 9 2.7 cm 14.7 cm 0.18 14. Aug. 4, 1963 8 3.9 cm 25.7 'cm . 0.15 15. Aug. 4, 1963 17 1.3 cm 7.8 cm 0.16 16. Sept. 15, 1962 10 0 . 7 cm 5.6 cm 0.12 93 Table I I I . Stipe Length/Blade Width as Determined from Transplants of Laminaria groenlandica Date No. PI. Mean Stipe Mean Blade Mean Length Width St/W Laminaria groenlandica long stipe form l a Feb. 22-Apr. 28, 1963 12 2.1 cm 7.6 cm 0.27 lb Apr. 28-June 10, 1963 . 12 3.2 cm 8.0 cm 0.40 Laminaria groenlandica short stipe form 2a Feb. 22-Apr. 28, 1963 14 1.6 cm 8.0 cm 0.20 2b Apr. 28-June.lO, 1963 . 14 2.7 cm 15.3 cm 0.17 3a May 22, 1963 5 2.3 cm 9.9 cm 0.23 3b Oct. 6, 1963 5 3.0 cm 8.7 cm 0.34 Laminaria groenlandica Long and Short Stipe Forms as Observed at the Beginning (a) and End (b) of Transplant Studies. Numbers 1 and 2 were transplanted to Burrard Inlet, number 3 to Glacier Point. 94 T a b l e I V . Values o f T e m p e r a t u r e a n d S a l i n i t y a t T h r e e D e p t h s i n H o w e S o u n d , n e a r K e a t s I s l a n d (49°24'N 1 2 3 ° 1 7 'W) o m 6 m 10. m D a t e T ° C S $ o T ° C T ° C S$o J u n e , 1957 16.9 9.3 11.4 22.1 9.2 27.1 J u l y , 1957 18.3 15.0 12.6 25 .9 11.2 26 .9 S e p t . , 1957 15.2 12.1 14.1 25 .9 12.2 27.0 N o v . , 1957 7.8 27.4 9.0 2 8 . 0 9.1 28.4 A n o n . , 1958. S t a t i o n 2 . 95 Table V. I n i t i a l Tests on the S a l i n i t y and Temperature Tolerance of Laminaria saccharina (L.) Lamour. May 16 to June 11, 1963 Temperature S a l i n i t y May 21 May 28 June 4 June 11 10$, I I I. I I I D\" D 20$* 7$ 28$ 23$ 35^ 32$ 47$ 34$ 5 C 12$ 21$ 20$ 51$ 23$ 62$ 32$ 69$ 40$, — 20$ 33$ 53$ 50% I I I I D D 10$, I I I I D D 10°C 20$, 24$ 16$ 43$ 28$ 49$ 37$ 64$ 40$ 30$* 9$ 17$ 22$ 34$ 31$ 37$ 41 40$„ 13$ . 17$ 19$ 33$ 30$ 45$ 34$ 53$ 50$, I I I I D D 10$, 6$ 15$ I I D D 15°C 20foo 15$ 18$ 27$ 38$ 42$ 47$ 46$ 52$ 30$, 2«$ 27$ 41$ 52$ 42$ 67$ 51$ 74$ 40$, 20$ 15$ 34$ 22$ 31$ 45$ 36$ 50$, I I D D 10$, D D 20°C 20$, 20$ 7$ I I D D 30$, b$ 7$ I I D D 40$, 1$ 4$ I I D D 50$, D D Two plants were placed in each temperature-salinity s i t u a t i o n . (D) Death; (I) Injury; ($) increase in area of the blade from May 16, 1963. To face page 96 Table VI. Net photosynthesis, r e s p i r a t i o n , and net photo-synthesis/respiration r a t i o s of Laminaria. saccharina and Laminaria groenlandica -.long stipe form in various temperature-salinity situations. Units: /(lOg/cmS/hour LAMINARIA SACCHARINA PHOTOSYNTHESIS SALINITY' '%. 20 24 26 28 30 32 m 7 6.1 8.3 6.8 6 .8 7.0 7.5 7.1 TO 8.1 8.0 8.1 7.9 8.0 8 .3 8 J 13 7.8 8.5 8.6 8.2 8.9 8.6 8.4 15 10 .0 11 .0 10.5 9.2 10.6 9 .3 10.1 18 10 .8 11.9 10.4 8.3 12.8 12.1 11.1 ni 8.6 9.5 8.9 8.1 9.5 9.2 RESPIRATION 7 0 .5 0.5 0.6 0 .7 0.5 0 .7 0 .6 TO 1.0 0 .5 0.6 0 .4 0 .8 0 .5 0.6 13 0 .7 0 .7 0.6 0.7 0.6 0 .7 0 .7 15 0 .7 1.0 0.7 0 .8 0.9 0 .7 0 .8 18 0.6 0 .8 0 . 7 0 .7 0.8 0 .9 0 .8 rfl 0 .7 0 .7 0.6 0 .7 0 .7 0 .7 PHOTOSYNTHESIS / RESPIRATION 7, 12 .2 16.6 11.3 9.7 14 .0 10.7 12.4 10 8.1 16 .0 13.5 19 .8 10.0 16.6 14.0 13 11.1 12.T 14.3 11.7 14.8 12 .3 12 .7 15 14 .3 11 .0 15.0 11.5 11.8 13 .3 12.8 18 18 .0 14 .9 14.9 11.9 16.0 13 .4 14.9 rfi 12 .7 14.1 13.8 12.9 13.3 13 .3 LAMINARIA GROENLANDICA PHOTOSYNTHESIS SALINITY 20 24 26 28 30 32 ffT 7 5.7 5.7 6 .0 5.6 5.7 7.0 6 . 0 10 2.4 3.7 2,5 4 .2 5,9 5,0 4 , 0 13 0 0 0 0 2 .7 2 .7 0 .9 15 0 0 0 2.9 1.4 0 0 .7 18 0 0 0 0 0 0 0 rR 1.6 1.9 1.7 2 .5 3.1 2.9 RESPIRATION C O 7 1,6 1.2 1.2 T.T T.O 0 .7 1.1 o TO 1.8 1.8 0 .8 1.9 l . T 1.2 1.4 L L J cr: T3 2.2 2 .3 2.2 1.6 2 .4 1.8 2.1 ro I— 15 2.2 2.2 2.5 2 .6 2 ,8 2.4 2 .5 cr: U J n 18 1.2 0 .5 1.5 1.6 1.8 2.1 K 5 rs L U 1— ffl 1.8 1.6 1.6 1.7 1.8 PHOTOSYNTHESIS / RESPIRATION 1.6 7 3.6 4 .8 5.0 5,1 5.7 10 .0 5,7 10 1.3 2.1 3.1 2.2 5.4 4 .2 3.1 13 - - - - 1.1 1.5 0 .4 15 - - - 1.1 0 .5 - 0 .3 18 — _ — — — — m 1,0 1.4 1.6 1.7 2.5 3.1 97 LAMINARIA SACCHARINA SALINITY %. 17 20 23 26 29 32 7 N N N N N N TO N N N N N N 13 N N N N N N 16 A A A N N N LAMINARIA GROENLANDICA SALINITY\" 17 20 23 26 29 32 7 N N N N N N 10 NA N N N N N 13 A A A NA NA N = NORMAL SPOROPHYTE A = ABNORMAL SPOROPHYTE Table VII. Sporophyte production by gametophytes of Laminaria saccharina and Laminaria groenlandica long stipe form i n various temperature-salinity sit u a t i o n s . 40 00 5000 o 6000 7000 WAVE LENGTH (A) Figure 1. Spectral curves of (a) Westinghouse F 4 0 Blue, ( b ) Westinghouse F40HE37 Green, and (c) one Westinghouse Blue to two Westinghouse Green fluorescent tubes. (Westinghouse,.personal communication.) To face page 99 Figure 2 . Spectral curve of Sylvania Cool White (F48T 12-CW) fluorescent tubes. Figure 3. (a) Percent transparency of stained glass used in f i l t e r i n g , l i g h t in constant temperature control rooms. (b) Percent transparency per meter for coastal waters with sun at an a l t i t u d e of 45° (After J e r l o v / 1951, water type no. 7.) Figure 4. Relative s e n s i t i v i t y of the Photovolt Elec t r o n i c Photometer Model 501-M. 99 3 5 0 0 4 5 0 0 5 5 0 0 6 5 0 0 3 0 0 0 4 0 0 0 5 0 0 0 6 0 0 0 W A V E L E N G T H (A ) To face page 1 0 0 Figure 5. Habit of Laminaria longipes. Figure 6. Habit of Laminaria s i n c l a i r i i , exposed form. Figure 7. Habit of Laminaria s i n c l a i r i i , sheltered form. Figure 8. Habit of Laminaria ephemera, exposed form. 100 To face page 101 Figure 9. Habit of Laminaria yezoensis. Figure 1 0 . Discoid holdfast of Laminaria yezoensis. Figure 11 . Habit of Laminaria groenlandica shade form. Figure 1 2 . Regeneration of Laminaria groenlandica shade form. Figure 13. Habit of Laminaria bu l l a t e . Figure 14. Habit of Laminaria Figure 15. Habit of Laminaria abullate blade. Figure 16. Habit of Laminaria To face page 102 groenlandica short stipe form, groenlandica long stipe form. groenlandica short stipe form, groenlandica f l a t stipe form. 102 13 To face page 103 Figure 17. Habit of Laminaria saccharina, bul l a t e blade. Figure 18. Habit of Laminaria saccharina, abullate blade. Figure 19. Habit of Laminaria f a r l o w i i . Figure 20. Habit of Laminaria complanta. 103 To face page 104 Figure 21. Habit of Laminaria s e t c h e l l i i , Glacier Point form. Figure 22. Habit of Laminaria s e t c h e l l i i , Botany Beach form. Figure 23. Regeneration of Laminaria s e t c h e l l i i . Figure 24. Habit of Laminaria dentigera. To face page 105 Figure 25 . Laminaria longipes, cross section of blade. Figure 2 6 . Laminaria s i n c l a i r i i , cross section of blade. Figure 27. Laminaria ephemera, cross section of blade. Figure 28 . Laminaria yezoensis, cross section of blade. Figure 29 . Laminaria groenlandica shade form, cross section of blade. Figure 30. Laminaria groenlandica long stipe form, cross section of blade. Figure 31 . Laminaria groenlandica f l a t stipe form, cross section of blade. Figure 3 2 . Laminaria saccharina, cross section of blade. Figure 33 . Laminaria f a r l o w i i , cross section of blade. Figure 34. Laminaria complanata, cross section of blade. Figure 35 . Laminaria s e t c h e l l i i , cross section of blade. Figure 36. Laminaria dentigera, cross section of blade. ( < ) indicates the position of mucilage ducts. To face page 106 Figure 37. Laminaria longipes, cross section of stipe. Figure 38. Laminaria s i n c l a i r i i , cross section of st i p e . Figure 39 . Laminaria ephemera, cross section of stipe. Figure 40. Laminaria yezoensis, cross section of stipe. Figure 4 l . Laminaria groenlandica shade form, cross section of stipe. Figure 42. Laminaria groenlandica short stipe form, cross section of stipe. Figure 43 . Laminaria groenlandica long stipe form, cross section of stipe. ( ) indicates the position of mucilage ducts. The l i n e drawings indicate the r e l a t i v e shape and size (2X) of the stipes as observed in cross section. The area of these drawings darkened represents the position of the section represented in the accompanying photograph. To face page 1 0 7 Figure 4 4 . Laminaria groenlandica f l a t stipe form, cross section of stipe. Figure 4 5 . Laminaria saccharina, cross'section of st i p e . Figure 46. Laminaria f a r l o w i i , cross section of stipe. Figure 4 7 . Laminaria complanata, cross section of st i p e . Figure 48. Laminaria s e t c h e l l i i , cross section of st i p e . Figure 4 9 . Laminaria dentigera, cross section of stipe. ( — — — ) indicates the position of mucilage ducts. The l i n e drawings indicate the r e l a t i v e shape and size (2X) of the stipes as observed in cross section. The area of these drawings darkened represents the position of the section in the accompanying photograph. 107 6 0 ° N 5 5 ' c7* 0 5 0 * N ^ I 8 0 ° W I 7 0 ° W + I 5 0 ° W L. LONGIPES A L. DENTIGERA • Figure 50. Northeast P a c i f i c d i s t r i b u t i o n of Laminaria longipes and Laminaria dentigera. ' — 1 o CO 109 Figure 51. Northeast P a c i f i c d i s t r i b u t i o n , north of 48° N la t i t u d e , of Laminaria s i n c l a i r i i , Laminaria ephemera, Laminaria f a r l o w i i , and Laminaria complanata\". I80°W I60°W 140° W Figure 52 60°N 55°N L. S E T C H E L L I I # L. Y E Z O E N S I S • SUBTIDAL O N L Y Q Northeast P a c i f i c d i s t r i b u t i o n , north of 48°N l a t i t u d e , of Laminaria s e t c h e l l i i and Laminaria yezoensis-. ~ o I 70°E I80°W I70°W I60°W I50°W I40°W I30°W 120 \" 1 1 1 I I I _ J Figure 53. Northeast P a c i f i c d i s t r i b u t i o n , north of 48°N l a t i t u d e , of Laminaria groenlandica and Laminaria saccharina. 1 1 2 FEBRUARY SPOROPHYTE GENERATION FIRST CONSPICUOUS*-APRIL SHADE FORM JUNE SHADE FORM NOVEMBER-JANUARY SHADE FORM FEBRUARY JUNE V LONG & SHORT STIPE FORMS I M DEAT SORUS PRODUCTION*? t ATH SORUS PRODUCTION ATTRITION OF BLADE DEATH REGENERATION OF BLADE SORUS PRODUCTION DEATH Figure 54. L i f e h i s t o r i e s of Laminaria groenlandica long stipe form, short stipe form, and shade form-as observed at Glacier Point and Ogden Breakwater, Vancouver Island. Large arrows denote phenology of the sporophyte generation, small arrows, the gameto-phyte generation. 113 Figure 55. Gametophyte and microscopic sporophyte phases of Laminaria groenlandica long and short stipe forms: [a). male gametophyte developed i n adverse culture conditions, (b) female gametophyte developed i n adverse culture conditions, (c) female gametophyte developed in favourable culture conditions, (d) male gametophyte developed in favourable culture conditions, (e, f) sporophytes developed in favourable culture conditions, (g) plants, thought to be sporophytes, developed in.adverse culture con-d i t i o n s . To face page 114 Figure 56. D i s t r i b u t i o n of longitudinal growth, in culture, of the blade of Laminaria groenlandica short stipe form. Figure 57. D i s t r i b u t i o n of•longitudinal growth, in culture, of the blade of Laminaria groenlandica long stipe form. Figure 58. D i s t r i b u t i o n of longitudinal growth, i _ s i t u , of the blades of Laminaria groenlandica long stipe form (&,) and short stipe form (o). 114 D I S T A N C E FROM B L A D E B A S E (CM) To face page 115 Figure 59. D i s t r i b u t i o n of l a t e r a l growth, in culture, of the blade of Laminaria groenlandica short stipe form. Figure 60. D i s t r i b u t i o n of. l a t e r a l growth, i n culture, of the blade of Laminaria groenlandica long stipe form. Figure 6 l . D i s t r i b u t i o n of. l a t e r a l growth, i n s i t u , of the blades of Laminaria groenlandica long stipe form (A) and short stipe form (o) ™ 115 116 FEBRUARY-MARCH OCTOBER-NOVEMBER SPOROPHYTE GENERATION FIRST CONSPICUOUS SORUS PRODUCTION-FEBRUARY JUNE SEPTEMBER OCTOBER-NOVEMBER FEBRUARY Figure 62. ATTRITION OF BLADE REGENERATION OF BLADE SORUS PRODUCTION DEATH SPOROPHYTE GENERATION FIRST CONSPICUOUS SORUS PRODUCTION -t ATTRITION OF BLADE \\ REGENERATION OF BLADE L i f e history of Laminaria saccharina as observed in Burrard Inlet. Large arrows denote phenology of sporophyte generation, small arrows, gametophyte generation. 117 Figure 63. Gametophyte and microscopic sporophyte phases of Laminaria saccharina: (a) Female gametophyte developed in adverse culture conditions, (b). male gametophyte developed i n adverse culture condi-tions, (c) female gametophyte developed .in favor-able culture conditions, (d)male gametophyte developed in favorable culture conditions, (e. f) sporophytes developed in favorable conditions, (g) plants, thought to be sporo-phytes, developed in adverse culture conditions. 118 D I S T A N C E F R O M B L A D E B A S E (CM) Figure 64. D i s t r i b u t i o n of longitudinal growth, in culture, of the blade of Laminaria saccharina. Figure 65. D i s t r i b u t i o n of longitudinal growth, in s i t u , of the blade of.Laminaria saccharina. 119 96 I N C REASE IN WIDTH A F T E R 13 DAYS Figure 66. D i s t r i b u t i o n of l a t e r a l growth, In culture, of the blade of.Laminaria saccharina. Figure 67. D i s t r i b u t i o n of l a t e r a l growth, _in s i t u , of the blade of Laminaria saccharina. MAY JUNE JULY AUG. SEPT. OCT. NOV. Figure 68. Longitudinal growth rates (A) and theoretical lengths (o) of the blades of Laminaria saccharina, as observed in 1962, in Burrard Inlet. Numbers accompanying each growth rate value denote number of plants measured. The v e r t i c a l bars represent extent of the extreme values. 121 S%o 28 30 32 28 30 32 0 m j — i — ' — 1 — 1 — i — j I — i — i — i — i — 1 ~ Figure 69. Seasonal v a r i a t i o n in temperature and s a l i n i t y at three depths for Sheringham Point (48° 23 N 123° 59 ' W ) , on the r i g h t , and Gordon Head (48° 30*N .: 1 2 3 ° . 17 'w); on the l e f t . Anon., 1955, stations 3a and 10a. 122 Figure. 7 0 . Seasonal v a r i a t i o n in temperature and s a l i n i t y at three depths for Burrard Inlet near Stanley Park. To face page 123 Figure JI. Seasonal v a r i a t i o n i n surface temperature and s a l i n i t y at l o c a l i t i e s inhabited by Laminaria groenlandica long stipe form ( l ) , Laminaria saccharina (2), and Laminaria groenlandica short stipe form and Laminaria saccharina ( 3 ) , as observed at Amphitr.ite Point, Kains Island, Race Rocks, Departure Bay, Cape Mudge, Pine Island, f o r 1 9 4 8 - 5 7 , ' ( A n o n . , 1958); Beaver Point 1954-57, (Anon.,. 1958); Pulteney Point, 1955-57, (Anon., 1958); Nootka, 1942-51 (Anon., 1952); Ladysmith Harbour, 1950-56, (Anon., 1957); Texada Mines, 1954-56, (Anon., 1957); William Head, 1921-38, (Anon., 1948); Friday Harbor, 1935-52, (Anon.,.1962a); Neah Bay, 1936-60, (Anon., 1962a); Deep Cove, 1963, (Anon., 1964). PINE I. PULTENEY PT. CAPE MUDGE TEXADA I. r o o o <0 O 0/ (0 8 > < Q \\ UJ o < u. cc CO UJ Q < -I CD UJ CO < UI oc o I 8 r 125 12 -6 7 3 I8r 12 6 l 1 » 1 1 1 t I I I 1 I I I ! ! l _ 8 9 10 || 12 13 14 15 16 TEMPERATURE °C 74 A _ l L A A k A J L j I A A A J I A A A A A I J L 15 17 19 21 23 25 SALINITY %o A A A A A 27 29 31 Figure 73. Percent increase in blade surface per day for Laminaria saccharina a f t e r 15 days culture at 7 . 5 , 10, 12 .5 , and 15.5°C, at 2 8 ^ s a l i n i t y . Figure 7h. Percent increase i n blade surface per day for Laminaria saccharina after- 1 5 days culture at 16, 19 .5 , 2 3 . 5 , 27, and 31$* s a l i n i t y , at 1 0 ° C . 126 120 -8 0 x o z UJ UJ r ->-X 0-o ct o 0_ CO 4 0 75 B.I. V. G.R B.I. V. G.R L. GROENLANDICA 76 BURRARD IN. % 0 32 30 28 L. SACCHARINA G L A C I E R PT, % 0 32 30 28 8 0 4 0 t 4 + + L. SACCHARINA Figure 75. Sporophyte growth of Laminaria saccharina and Laminaria groenlandica long stipe form in sea-waters from d i f f e r e n t areas: Burrard Inlet .(B.I.), V i c t o r i a (V.), Glacier Point (G.P.). The ho r i -zontal l i n e represents the mean length; the rectangle, the 95$ confidence\"limits; the v e r t i c a l . l i n e , the extent of v a r i a t i o n . Figure 76. Sporophyte growth of Laminaria saccharina in seawater from Burrard Inlet and Glacier Point at three s a l i n i t i e s . 1 9 6 3 N . D i J . F . M . A . M . J . J . A . S . O . B , gure 77. Upper v e r t i c a l l i m i t s of L. saccharina as observed i n Burrard Inlet near Stanley Park. 0,4,, 0, represent, d i f f e r e n t year classes. To face page 128 Figure 78, Some t i d a l features for Burrard Inlet (after Anon.. 1962). The three v e r t i c a l bars under each month.represent three i n t e r t i d a l positions. Reading from r i g h t to l e f t : 30 cm above MLT, MLT, and 30 cm below MLT. D U R A T I O N OF L O N G E S T E X P O S U R E H O U R S E X P O S E D / MONTH 6 4 2 0 2 4 6 i — l — i 1 — i — i — | 1 — i — i — i — i — i DAY . NIGHT 100 60 20 0 20 60 100 I — l — l 1 — l 1 — I — l — i — I — i DAY ^NIQHT F 3 \" =3 3 F F E N U M B E R C X P 0 8 U R E S / MONTH 20 12 4 0 4 i — r ~ i — i — i — r 12 20 - i — i — I D A Y , NIGHT = 3 3 > 2 X CO I J F M A M J J A S O N D MONTHS Figure 79. A i r temperature for Kit s i l a n o , Vancouver, 1963 (Anon. 1964a). (c) the monthly mean temperature range mid-point, (£) monthly M mean maximum and (P) monthly mean minimum. ro To face page 130 Figure 80. Day length as calculated for 49°N (Anon. 1961.) . Figure 8 l . Hours bright sunshine observed for the Univ. B r i t i s h Columbia, Vancouver, 1963 (Anon. 1964a). Figure 82. Approximate monthly mean d a i l y insolation for l i g h t in the v i s i b l e range (3800-7200 A) as observed for the Univ. B r i t i s h Columbia,. 1963. (After Anon. 1964b). INSOLATION ( L Y / DAY) — ro OJ o o o o o o i 1 1 1 1 BRIGHT S U N ( H O U R S / M O N T H ) O O o o o o T T DAY L E N G T H (HOURS) 00 O IU i 0) I I I I I I T \\ >-_J 7500 H5000 _ o X H2500 H m 250 _2 -< i o oo \"H 75 50 25 0 Figure 83. MONTHS Approximate submarine l i g h t i ntensity at ( ).6.5 meters below MLT, ( ) 30 cm below MLT, and ( ) 30 cm above MLT between 12:00 AM and 1:00 PM. 0 0 CO CO UJ CO o 12 8 5 ^ LU OC < CL CL < ^ 4 0 -4 0 8 4 i A & A A A 2 I 0 -I -2 _L 2 0 0 4 0 0 6 0 0 8 0 0 A A 20 40 60 _ l I I UJ Q < _J CO UJ CO < UJ u: or cc O 3 ? CO < a o < 8 0 8 5 A A A A A A A A Figure 84. Figure 85. 2 0 0 4 0 0 6 0 0 8 0 0 LIGHT INTENSITY (ft-c) Apparent photosynthesis of L. saccharina sporo-phytes at d i f f e r e n t l i g h t i n t e n s i t i e s . Whole plant response of:L. saccharina to di f f e r e n t l i g h t i n t e n s i t i e s . SALINITY 22 24 26 28 (96o) 3 0 3 2 3 4 O o LU < cr 8 10 12 a. 14 UJ 16 18 1 ' 1 r - p r 12 T r i r II DEPARTURE BAY 10 6 9 2 3 U 12 II 10 6 9 7 8 PINE ISLAND - — l 1 8 i 1 1 1 i i i • » Figure 86. Mean, monthly seawater temperature and s a l i n i t y at Departure Bay and Pine Island, 1931-1958 (Anon. 1959). 00 "@en ; edm:hasType "Thesis/Dissertation"@en ; edm:isShownAt "10.14288/1.0105409"@en ; dcterms:language "eng"@en ; ns0:degreeDiscipline "Botany"@en ; edm:provider "Vancouver : University of British Columbia Library"@en ; dcterms:publisher "University of British Columbia"@en ; dcterms:rights "For non-commercial purposes only, such as research, private study and education. Additional conditions apply, see Terms of Use https://open.library.ubc.ca/terms_of_use."@en ; ns0:scholarLevel "Graduate"@en ; dcterms:title "On the taxonomy, distribution, and ecology of the brown algal genus Laminaria in the Northeast Pacific"@en ; dcterms:type "Text"@en ; ns0:identifierURI "http://hdl.handle.net/2429/38378"@en .