Science, Faculty of
Botany, Department of
Zoology, Department of
DSpace
UBCV
Scagel, Robert Francis
Master of Arts - MA
University of British Columbia
Only one species, Macrocvatls integrifolia Bory, is present in British Columbia coastal waters. It is found distributed along the whole coast in regions approaching open ocean conditions, but protected against the full ocean force, where the seawater has a specific gravity of over 1.018 at 15 degrees Centigrade and where a suitable rocky bottom, between three feet above to twenty feet below zero tide level, is available. The method of growth of the sporophyte, which is partly dichotomous and partly unilateral, is fully described from the time it is first discernible to the naked eye up to maturity. Dichotomous and subdichotomous divisions of the basal growth region are responsible for the extensive development and perennial habit of the plant. Active zoospore liberation from mature plants and young sporophytes (single, undivided blade stage) were found at least from July 1 to September but the limits were not determined. The gametophytic stages were not followed. Maximum growth (elongation) of Macrocystis recorded was 3.10" per day for a period of 29 days at the north-east of Vancouver Island. No regeneration from the holdfast or at the cut surfaces of stipes were shown as a result of cutting. Stipes of Macro cystis may continue to grow after being severed from their holdfasts. Plants removed to deeper water are retarded in rate of growth. Various practical aspects of this economically important plant are considered. It is believed two harvests per year are possible if proper precautions are taken with respect to the relative position of the terminal laminae of the plants when harvesting. A method of estimating quantities using the diameter of the stipe as an index is suggested.
^^3 _£y> SA < 3 Co/> . I THE LIFE HISTORY AMD DISTRIBUTION OF M&CROCYSTIS IN BRITISH COLUMBIA COASTAL W&TERS fey Robert Francis; Scagel, B.A. -o-A Thesis submitted i n Partial Fulfilment of The Requirements for the Degree of MASTER OF ARTS In the Department of BIOLOGY AND BOTANY -o-The University of British Columbia October, 194# ABSTRACT: Only one species, Macrocvatls in t e g r i f o l i a Boiy, i s present i n Bri t i s h Columbia coastal waters. I t i s found distributed along the whole coast i n regions approaching open ocean conditions, but protected against the f u l l ocean force, where the seawater has a specific gravity of over 1.018 at 15 degrees Centigrade and where a suitable rocky bottom, between three feet above to twenty feet below zero tide le v e l , i s available. The method of growth of the sporophyte, which i s partly dichotomous and partly unilateral, i s ful l y described from the time i t i s f i r s t discern-ible to the naked eye up to maturity. Dichotomous and subdichotomous div-isions of the basal growth region are responsible for the extensive develop-ment and perennial habit of the plant. Active zoospore liberation from mature plants and young sporophytes (single, undivided blade stage) were found at least from July 1 to September but the limits were not determined. The gametophytic stages were not followed. Maximum growth (elongation) of Macrocystis recorded was 3.10" per day for a period of 29 days at the north-east of Vancouver Island. No regeneration from the holdfast or at the cut.surfaces of stipes were shown as a result of cutting. Stipes of Macro cystis may continue to grow after being severed from their holdfasts. Plants removed to deeper water are retarded i n rate of growth. Various practical aspects of this economically important plant are considered. It i s believed two harvests per year are possible i f proper precautions are taken with respect to the relative position of the term-inal laminae of the plants when harvesting. A method of estimating quan-t i t i e s using the diameter of the stipe as an index i s suggested. - i -TABLE 03? COHEENTS I a g e T a b l e o f Contents i L i s t o f C h a r t s • i i L i s t o f F i g u r e s . . . , i i i L i s t o f Tables v L i s t o f P l a t e s v i . , A b s t r a c t • v i - i - i , ^«<4w3 ffflkr/ja^JL I . INTRODUCTION 1 I I . HISTORICAL BACK&ROUM) AMD REVIEW OF L I T E R A T U R E . . . . . . . . 2 I I I . DETERMINATION OF SPECIES 8 I V . GEOGRAPHICAL DISTRIBUTION. 11 1 . R e g i o n Surveyed 11 2 . E x p l a n a t i o n o f C h a r t s 13 3 . Method of E s t i m a t i n g 15 4 . Equipment Used 20 V . FACTORS AFFECTING DISTRIBUTION 22 1 . G e n e r a l . • 22 2 . Temperature.. 24 3 . L i g h t v 24 4 . S a l i n i t y 25 5 . C u r r e n t s . . . 26 V I . ECOLOGICAL ASSOCIATIONS 26 1 . Other P l a n t s 26 2 . A n i m a l s 27 . V I I . L I F E HISTORY 28 1 . A l t e r n a t i o n o f G e n e r a t i o n s . . . 28 2 . E a r l y Development o f the Sporophyte 30 3 . L a t e r Development o f t h e Sporophyte 32 4 . Growth R a t e and P r o p o r t i o n s 41 V I I I . SUMMARY 53 I X . ACKNDWIEDGaffiOTS 56 X . BIBLIOGRAPHY 57 APPENDICES i x -- i i -LIST OF CHARTS I . D i s t r i b u t i o n o f Macro c y s t i s . A key t o Char ts I I t o V I i n c l u s i v e , showing a r e a s f o r w h i c h l a r g e s c a l e d e t a i l s are g i v e n . I I . D i s t r i b u t i o n o f Macro c y s t i s . Large s c a l e C h a r t showing d i s t r i b u t i o n a t the n o r t h e r n I n t e r n a t i o n a l Boundary i n the v i c i n i t y o f Dundas I . and a d j a c e n t i s l a n d s . I l l . D i s t r i b u t i o n o f M a c r o c y s t i s . Large s c a l e c h a r t showing d i s t r i b u t i o n a r o u n d P o r c h e r I . , Stephens I . , and a d j a c e n t i s l a n d s . I V . D i s t r i b u t i o n o f Macro c y s t i s . Large s c a l e c h a r t showing d i s t r i b u t i o n around Banks I . -V . D i s t r i b u t i o n o f M a c r o c y s t i s . Large s c a l e c h a r t showing d i s t r i b u t i o n a r o u n d A r i s t a z a b a l I . V I . D i s t r i b u t i o n o f Macro c y s t i s . A key t o C h a r t s V I I t o X I i n c l u s i v e , showing a r e a s f o r which l a r g e s c a l e d e t a i l s a r e shown. V I I . D i s t r i b u t i o n o f Macro c y s t i s . Large s c a l e c h a r t showing d i s t r i b u t i o n i n S h e l t e r Bay and v i c i n i t y . . . V I I I . D i s t r i b u t i o n o f M a c r o c y s t i s . Large s c a l e c h a r t showing d i s t r i b u t i o n a t Hope I . and v i c i n i t y . I X . D i s t r i b u t i o n o f Macro c y s t i s . Large s c a l e c h a r t showing d i s t r i b u t i o n i n Hardy Bay and G o l e t a s C h a n n e l . X . D i s t r i b u t i o n o f Macro c y s t i s . L a r g e ' s c a l e c h a r t o f d i s t r i b u t i o n f r o m D i l l o n P o i n t t o K l u c k s i w i R i v e r j key a l s o t o C h a r t X I (Deer I s l a n d ) . X I . D i s t r i b u t i o n o f M a c r o c y s t i s . Large s c a l e c h a r t o f d i s t r i b u t i o n around Deer I s l a n d . X I I . D i s t r i b u t i o n o f M a c r o c y s t i s . Large s c a l e c h a r t o f d i s t r i b u t i o n around M a l c o l m I . l i i -LIST OF FIGURES 1. Diagrammatic representation of the Life-Cycle of Macrocystis int e g r i f o l i a Bory. 2. Diagrammatic representation of the origin of the " s p l i t s " and development of the various laminae produced i n the young sporophyte of Macrocystis integ r i f o l i a Bory. , 3. Diagrammatic representation of the development of a young sporophyte of Macrocystis in t e g r i f o l i a Bory. 4. Diagrammatic representation of several stipes rising from the same holdfast of Macrocystis i n t e g r i f o l i a Bory showing general position taken by the stipes i n the water at four different stages of the tide. 5. Drawings of early development of the thallus of the sporophyte of Macrocystis i n t e g r i f o l i a Bory. 6. Drawing of a young sporophyte of Macrocystis in t e g r i f o l i a Bory. 7. Drawing of a young sporophyte of Macrocystis in t e g r i f o l i a Bory. 8. Drawing of a young sporophyte of Macrocystis i n t e g r i f o l i a Bory showing early " s p l i t s " i n thallus. 9. Drawing of & young sporophyte of Macrocystis in t e g r i f o l i a Bory showing early '"splits" in thallus. 10. Drawing of a. young sporophyte of Macrocystis in t e g r i f o l i a Bory showing early separations in laminae of the thallus. 11. Drawing of a young sporophyte of Macrocystis in t e g r i f o l i a Bory showing early separations i n laminae of the thallus. 12. Drawing showing formation of a " s p l i t " in the differentiation of the thallus of the sporophyte of Macrocystis i n t e g r i f o l i a Bory. 13. Drawing showing somewhat of an abnormal type of "sp l i t t i n g " i n the thallus of the sporophyte of Macrocystis i n t e g r i f o l i a Bory. 14.. Drawing of upper portion of a young stipe of Macrocystis i n t e g r i f o l i a Bory (see Fig. 15 for lower portion of the same stipe). 15. Drawing of lower portion of a young stipe and the growing "point" of a sporophyte of Macrocystis integ r i f o l i a Bory (see Fig. 14 for upper portion of the same stipe). 16. Drawing of ;a sporophyll of Macrocystis i n t e g r i f o l i a Bory in a comparatively early stage of division. LIST OF FIGURES (CONCLUDED) 17. Drawing of a sporophyll of Macrocystis i n t e g r i f o l i a Bcay at .a later stage of division. 18. Drawing of a sporophyll of Macrocystis in t e g r i f o l i a Bory without a pneumatocyst. 19. Drawing of a sterile lamina of Macrocystis i n t e g r i f o l i a Bory. 20. Drawing of a terminal lamina oi. a young (short) stipe of Macrocystis in t e g r i f o l i a Bory. 21. Drawing of a terminal lamina of a stipe of Macrocystis in t e g r i f o l i a •Bory. 22. Drawing of a terminal lamina of a stipe of Macrocystis in t e g r i f o l i a Bory. . 23. Drawing of a terminal lamina of a stipe of Macrocystis- integ r i f o l i a Bory (older). 24. Drawing of a young holdfast of Macrocystis i n t e g r i f o l i a Bory showing • i n i t i a l stages of "rhizome11 formation. 25. Drawing of a young holdfast of Macrocystis i n t e g r i f o l i a Bory showing i n i t i a l stages of "rhizome" formation. 26. Drawing of a portion of a mature holdfast of Macrocystis i n t e g r i f o l i a Bory showing origin of stipes of various ages. 27. Drawing of an extensive hapteron from a holdfast of Macrocystis integr i f o l i a Bory. 28. Diagrammatic representation of the method of branching found i n the holdfast of Macrocystis in t e g r i f o l i a Bory. LIST OF TABLES I. Distribution and Abundance of Macrocystis i n t e g r i f o l i a Bory. II. Eate of Growth and Division of Very Small Plants of Macrocystis integr i f o l i a Bory located on a large rock at northwest of Deer I. A l l plants between 1 and 2 feet above zero tide l e v e l . III. Bate of Growth of Stipes of Small Complete Plants of Macrocystis integrifolia Bory attached to holdfasts at -10 f t . for a period of 29 days, then moved to -14 f t . for a period of 17 days. IV. Bate of Growth (Elongation) of Stipes of Macrocystis i n t e g r i f o l i a Bory severed from holdfasts and anchored. V. Bate of Growth (Elongation) of Stipes of Macrocystis i n t e g r i f o l i a Bory fastened to holdfast. VI. Size of Stipes of Macrocystis in t e g r i f o l i a Bory and Distribution of Weight. VII. Elongation of Stipes of Macrocystis integ r i f o l i a Bory from which the terminal laminae, including the upper .2-foot portion, have been severed. VIII. Density and Temperature Eecordings on Sestwater Samples. LIST OF PLATES I.Photograph of holdfast of Macrocystis in t e g r i f o l i a Bory at 1.5 f t . tide l e v e l , 8 a.m. Aug. 27, 1946, Welcome Harbour (Porcher I.) II. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory, 8 a.m. Aug. 27, 1946, Welcome Harbour (Porcher I.) I l l . Photograph of same holdfast of Macrocystis i n t e g r i f o l i a Bory as shown i n Plate II from a different position and closer. Taken at 1.5 f t . tide level, 8 a.m. Aug. 27, 1946, Welcome Harbour (Porcher I.) IV. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at Aristazabal I. at 8 f t . tide, 13:10 Aug. 18, 1946. V. Photograph of bed of Nereocystis ltletkeana (Mert.) P. & R. outside bed of Macrocystis i n t e g r i f o l i a Bory at Melville I. at 10 f t . tide, 12:26, Aug. 23, 1946. VI. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory -at N. Banks I. (near Deadman I.) at 9 f t . tide, 13:30 Aug. 19, 1946. VII. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at N. Banks I. (near Deadman I.) at 8 f t . tide, 13:10, Aug. 19, 1946. VIII. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at N. Banks I. just at Deadman I. at 8 f t . tide 13:10, Aug. 19, 1946. IX. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at approaches to Norway Inlet at 10 f t . tide, Aug. 19, 1946. X. Photograph of bed of Macrocystis i n t e g r i f o l i a Boiy at 15 f t . tide a t entrance to Norway Inlet, 8:00, Aug. 20, 1946. (See Plate XI for the same bed at lower tide). XI. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at 10 f t . tide at entrance to Norway Inlet, 15:40, Aug. 19, 1946. XII. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at 11 f t . tide at E. Hope I., 12:30, Aug. 16, 1946. XIII. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at 5 f t . tide at Deer Island, 10:00 Aug. 15, 1946. XIV. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at 5 f t . tide at Deer Island, 10:00 Aug. 15, 1946. XV. Photograph of three terminal laminae of Macrocystis i n t e g r i f o l i a Bory showing several laminae on each i n progressive stages of separation. - v i i -LIST OF PLATES (CONCLUDED) XVI. Photograph of terminal lamina of Macrocystis i n t e g r i f o l i a Bory showing great number of laminae i n progressive stages of separation. XVII. Photograph of a young stipe of Macrocystis i n t e g r i f o l i a Bory broken off with a small section of the holdfast at the growing "point." XVIII. Photograph of the upper portion of a major stipe of Macrocystis in t e g r i f o l i a Bory. XIX. Photograph of an abnormal development of two laminae from one point on the stipe of Macrocystis in t e g r i f o l i a Bory. XX. Photograph of an abnormal development i n the stipe of Macrocystis in t e g r i f o l i a Bory. XXI. Photograph of a holdfast of Macrocystis integ r i f o l i a Bory showing numerous stipes from the same holdfast. XXII. Photograph of a holdfast of Macrocystis in t e g r i f o l i a Bory showing characteristics of a mature holdfast. XXIII. Photograph of plants of Macrocystis in t e g r i f o l i a Bory torn loose and washed high on beach. XXIV. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at northeast of Deer Island. XXV. Photograph of bed of Macrocystis i n t e g r i f o l i a Bory at northwest of Deer 'Island. XXVI. Photograph of numerous plants of Macrocystis i n t e g r i f o l i a Bory growing on a large rock exposed at zero tide, north side of Deer Island. XXVII. Photograph of plants of Laminaria saccharina (L.) Lamour. growing on rocks, mixed with small plants of Macrocystis integ r i f o l i a Bory, at about aero tide l e v e l on reef at north of Deer Island. THE LIFE HISTORY AMD DISTRIBUTION UF MICROCYSTIS IN BRITISH COLUMBIA COASTAL WATERS I. INTRODUCTION: This paper presents an account of the lif e - h i s t o r y and distribution of Macrocystis as encountered i n British Columbia coastal waters, and i s the result of the f i e l d work done during two summers, plus post-laboratory work done during the two subsequent f a l l and spring sessions at the University of B r i t i s h Columbia i n the Depart-ment of Biology and Botany, through the courtesy and assistance of Dr. A. H. Hutchinson, Head of the Department. The general distribution of Macrocystis i n B r i t i s h Columbia coastal waters was studied by the writer (69) during a period of employment i n the summer of 1946 with the Fisheries Research Board of Canada, under the direction of Dr. R. E. Foerster, Director, Pacific Biological Station, Departure Bay, B. C , and i n conjunction with a survey sponsored by the B r i t i s h Columbia Research Council, University of British Columbia (12 and 13). The more detailed study of the distribution i n the vic i n i t y of Hardy Bay, British Columbia, together with the c r i t i c a l study of the plant, i t s method of growth, and other aspects, was accomplished during a period of employment i n July, August, and September in the summer of 1947 (70) with the Provincial Fisheries Department under the direction of Mr. George J . Alexander, Deputy Minister. The main endeavour during this l a t t e r investigation was directed towards a more thorough understanding of the practical aspects of commercially import-ant species of algae and related problems. A l l work on the latter i n -vestigation was conducted from a station at Deer Island (Charts X and XI) and detailed work was restricted to the northern portion of Vancouver Island i n the immediate vi c i n i t y of the Port Hardy airport (Chart X) and Deer Island (Charts X and XI). The more detailed investigations on distribution have been supplemented by casual observations made during the summer of 1948 while engaged i n other work for the Fisheries Research Board. The study, i n so far as detailed distribution i s concerned, could not be extended to the west coast of Vancouver Island, the Queen Charlotte Islands, nor to a few minor, rather inaccessible points on the coast during the period of the investigations, although a few casual observations have since been made at several points i n these regions not previously touched. These observations indicate a general distribution a l l along the west coast of Vancouver Island and the writer f u l l y expects i t w i l l be found i n abundance along portions of the Queen Charlotte Islands, where occurrence at least has been noted during the summer of 1948 at various points along the west coast of Moresby and Graham Islands and the north coast of Graham Island. The life-history and growth of Macrocystis have been studied in d e t a i l i n so far as the sporophyte generation i s concerned, but the gametophytic phases have not been attempted i n this paper. The writer hopes to supplement this present paper with a morphological study of the microscopic phases of the life-history as well as a complete cytological study of Macrocystis. I I . HISTORICAL BACKGROUND AND REVIEW UF THE LITERATURE: The genus Macrocystis i s widely distributed i n both northern and southern hemispheres. Tilden (87) says that Macrocystis has perhaps the most extended range as regards temperature of any species (from 5°G. to 25°C. i n the southern hemisphere and 10°C. to 25°C. i n the north-ern hemisphere). This extremely wide distribution i s no doubt the chief reason for not only the very wide interest shown by world-scattered authors, but also the confusion which came about (and s t i l l exists, a l -though to a lesser degree) with regards to specific determination as opposed to geographical or seasonal variation. The determination of the species i s dealt with i n detail i n Section III of this paper under "Deter-mination of Species." A f a i r l y complete, although not exhaustive, bibliography at the end of this paper leads one to a comprehension of the differences of opinion amongst writers i n the past concerning the species of Macrocystis. In spite of the great interest at various times over the past century there are s t i l l many unanswered questions concerning the distribution and taxonomy of the genus, and our own section of the Pacific Coast i s not without exception. Many of the records, particu-l a r l y with respect to distribution along the northern part of the Pacific Coast of North America s t i l l stand uncorrected or unverified and w i l l remain so u n t i l further extensive work has been done, particu-l a r l y at the northern and southern extremes i n the northern hemisphere. To the early navigators approaching the shores of Mexico and California this plant was called "Porra" and was looked upon as an un-f a i l i n g sign of proximity to land. F i r s t mention, according to Tilden (87) goes back to the middle of the 16th century. I t was often alluded to i n the narrative of Cook's voyage. In so far as distribution of - 4 -Macrocystis on the British Columbia portion of the Pacific Coast i s concerned ve have a few records which, to date, have not been summarized and correlated. Before dealing with the references which definitely encompass the portion of the coast of Br i t i s h Columbia dealt with i n this paper i n detail, brief mention should be made of some of the early records estab-lished for the area to the Worth, some of which are questioned as to distribution and others as to speoies. I t i s most l i k e l y that these particular references are directly, or indirectly responsible for regarding the distribution of the different species i n the manner to be discussed l a t e r . Alexander Postels and Henry Mertens made a t r i p to Alaska in 1840 i n the ship Seniavin under Captain Lfltke and obtained specimens of Macrocystis near Sitka (60) . These were recorded, however, by Postels and Ruprecht (60) as Lessonla c i l i a t a . I t was also reported at this time from Unalaska. Murray (54-) records the distribution of Macrocystis up the whole coast of America to Alaska and over to the coast of Asia at Kamtschatka. There i s no corroboration, however, for i t s presence on the Asiatic coast or at Unalaska (76), and i t seems very unlikely that such i s the case unless i t may have drifted there and floating specimens were taken. Ruprecht (67) does not mention i t as occurring i n the Ochotsk. I t seems quite possible that i t may have drifted there, how-ever, since the pneumatocysts may keep portions of the plant floating for considerable time. The writer has found pieces of Macrocystis d r i f t -ing i n the north Pacific 150 miles off shore i n the Japanese Current. Saunders (68) on the Harrinan Alaska Expedition in 1901 recorded M. pyrifera (Turn.) Ag. from Juneau, Sitka and Wrangell, Alaska* Further records for Alaska were established by Johnston (43) and Setchell (93). There are some early records of specimens taken along the coast of British Columbia, but with the exception of Cameron's report (14), none attempted to enlarge on the distribution within this area. Dr. Archibald Menzies visited the northwest coast in a trading vessel somewhere between 1779 and 1786, and again in 1792, 1793 and 1794, as a member of the exploring expedition in command of Captain George Vancouver. He made a collection, which was partly figured and described by Dawson Turner (89) in Fuci, published in 1808 tonl819, including M. pyrifera (Turner) Ag. Dr. David Lyall sent a collection from the region of Esquimalt to Harvey (33) in Dublin, who described a specimen of M. pyrifera (Turn.) Ag. Tilden (86) recorded M. pyrifera (Turn.) Ag. from Fort Renfrew. Setchell and Gardner (75) recorded M. plrifera (Turner) Ag. (obviously misspelt for M. pyrifera) from Unalaska and Sitka to the • west coast of Whidbey Island, Washington, in dealing with distribution from the Strait of Juan de Fuca to Alaska. They mention itrias not being plentiful in Puget Sound. Their distribution at Unalaska, however, rested on the report of Postels and Ruprecht (60). The record from Whidbey Island was based on Gardner's specimen No. 861. Collins (18) record8 M. pyrifera (Turn.) Ag. from the west coast of Vancouver Island, and makes reference to a specimen taken by Macoun at Victoria, More exact records were established by Setchell (74-) for the west coast of Vancouver Island from specimens collected by Setchell and Parks (77) i n 1930 and 1931. Specimens identified as M. i n t e g r i f o l i a Bory are deposited i n the Herbarium of the University of California from Quatsino (1930, #445012), Clayoquot (1930, #445007 and #445013), and Banfield i n Barkley Sound (1931, #445014). It i s based on this record that Smith (83) has recorded distribution of M. in t e g r i f o l i a Bory from British Columbia (Vancouver I.) to Central California (Carmel Bay). The distribution recorded by Smith (83) for M. pyrifera (L.) Ag. i s from Alaska to Baja, California (Magdalena Bay). A more extensive survey of a portion of the coast of B r i t i s h Columbia by Cameron (14) was the f i r s t attempt to record a more detailed distribution of Macrocystis i n these waters, and the plant was referred to as M. pyrifera (Turn.) Ag. Cameron's distribution, together with that brought out by this paper, w i l l be discussed i n greater detail i n Section IV under "Geographical Distribution." The writer has confirmed most of the distribution recorded by Cameron (14)> added to the distribution within those areas surveyed by him, as well as extended the survey to areas not previously covered. While Macrocystis pyrifera has been recorded on the B r i t i s h Columbia coast by previous workers (14 and 18), this would appear to be the result of the confusion which existed u n t i l a more thorough study and separation of the two species M. i n t e g r i f o l i a and M. pyrifera. on the basis of the characteristics of the holdfast, was made by more - 7 -recent workers (74, 76, 82, and 83). For this same reason i t appears l i k e l y that many of the earlier publications (75 > 27, 43, 65, and 73), regarding investigations i n Alaska which li3ted M. pyrifera as occurring there, were actually dealing with M. i n t e g r i f o l i a . From the previous experience of the writer i t i s known that the same type, with the peculiar holdfast of "creeping rhizomes" (M. integrifolia) occurs as far north as Dundas Island and was the only type located from points a l l along the coast from Dixon Entrance south to Sooke on the southern end of Vancouver Island, This would seem to agree with Setchell (74), who considered Macrocystis integrifolia to be the only species along the whole stretch of coast-line north of Point Conception. The taxonomy of the species of Macrocystis rests primarily on a review of the works of the following: Agardh (1, 2, and 3), Areschoug (6), Bory (7, 8, and 9), Harvey (30, 31, 32, 33, and 34), Hooker (37), Howe (38), Kjellman (44), Linnaeus (47), and Skottsberg (78 and 79), The gametophytic phases of the life-history of Macrocystis have s t i l l not been f u l l y studied. Some preliminary work has been done, however, by Brandt (10), Delf and Levyns (23), Levyns (46), and Papenfuss (58), The method of growth, division of laminae, as well as development of the holdfast have been variously considered by Fritsch (26), Setchell (74), Skottsberg (79), as well as i n a recent paper by the writer (70). Brandt (10) and Moore (53) have also reported something on the growth of Macrocysti3. The general anatomy has been considered i n some detail by Fritsch (26), Rosenthal (66), Skottsberg (79), Sykes (85), and Will (92). - 8 -Reproduction has been separately discussed by Hoffman (36) and also by Smith and Whitting (80). Ill.DETERMINATION OF THE SPECIES: Linnaeus (47) listed the type of the genus Macrocystis in his diversified and all-including genus "Fucus" as Fucus pyrlferus. The specimen on which Linnaeus founded the species came from the "mere Aethiopicum" which has been surmised as being somewhere among the islands of the Antarctic (probably Kerguelen according to Setchell (74)). Throughout the reference material up until Setchell and Gardner (76) published their "Melanophyceae" in 1925 the type was referred to, how-ever, as M. pyrifera (Turn.) Ag. The genus Macrocystis was delimited by Agardh (1) in his monumental revision. The credit to Turner (89) came through his publication of "Fuci"j however, since establishment of the prior right of Linnaeus (47), as set forth by Setchell and Gardner (74), the type is now referred to as M. pyrifera (L.) Ag. At various times in the past some thirty-odd species of Macrocystis have been accorded specific rank. The early conclusion reached by Harvey (31) and Hooker (37) was that al l so-called species should be referred to the single species M. pyrifera and not to very variable species. These two workers claimed that many of the reputed species are found "growing together at times on the different parts of the same stem." The differences were attributed to the result of age, degree of denticulation, rugose character of the surfaces of the laminae, proportion of length to breadth of lamina, and holdfast characteristics. - 9 -Since the more recent review of the work by ttowe (38), Setchell and Gardner (76), and Setchell (74), the concensus of opinion i s that there are only two species; namely, M. pyrifera (L.) Ag. and M. i n t e g r i f o l i a Bory. In addition there i s some reason to believe that a third form, M. angustlfolia Bory may merit specific rank, although i t i s often included with M. i n t e g r i f o l i a Bory. According to Setchell (74) some two or even three species of reasonable cleavage might be expected within such a range of temperature. The separation of these two species i s based primarily on the characteristics of the holdfast, although secondary characteristics pertaining to the general proportion of the length of laminae to breadth, particularly the terminal lamina, are stated to be diagnostic f o r the two species i n the waters off California (83). From a l l the plants observed by the writer i n the waters along the coast of Br i t i s h Columbia, however, i t i s believed that only the holdfast characteristics may be used as a basis for separation. The position from which the laminae are obtained must be taken into consideration i f the l a t t e r method i s to be used, and even then there i s such a range in variation to be found i n these proportions that they cannot be relied upon. The proportions given by Smith (83) for the terminal blade (lamina) of M. pyrifera are length 3 to 5 times breadth, with 10 to 20 young later a l blades i n progressive stages of differentiation, and for M. jntegrifolla 6 to 10 times breadth and 3 to 6 young blades. According to the proportions given for the laminae of these two species by Smith (83) one would assign the Macrocystis found i n B r i t i s h Columbia coastal waters to the species M. pyrifera ( L . ) Ag. - 10 -These characteristics of the holdfast, however, very definitely conform to those described for M. integrifolia Borv. There is undoubtedly a considerable range in variation between plants of this species along this coast and those of the type locality (Valparaiso) originally described by Bory (7). A comparison with the photographs by Howe (38j in the "Marine Algae of Peru" indicates the extent of this variation. The basis of separation using the holdfast characteristics, therefore, as outlined by Setchell and Gardner (76), seems to be the best-and only way, from macroscopic characteristics at least, of differentiating between the two species in these waters. The basic difference in the holdfasts of these two species rests on the presence or absence of the flattened, creeping, so-called "rhizome-portion" of the holdfast (Fig. 26). The details of holdfast construction as well as other characteristics of the plant as found in these waters are considered later in this paper. Although the records for Macrocystis in British Columbia waters in the past (with the exception of those for the west coast of Vancouver Island by Setchell and farks (77)) have been for M. pyrifera. i t is the opinion of the writer that only M. integrifolia Bory is present, and that those records for Alaska will probably also be found as applying to M. integrifolia. Setchell (74) anticipated that M. integrifolia Bory would probably be the only species north of Point Conception, but his verification of this only extended to the west coast of Vancouver Island. The study of the distribution made by the writer has extended this verification as far as tfundas Island, at Dixon Entrance—the International Boundary between British Columbia and Alaska—as veil as to the vest coast of the Queen Charlotte Islands. Macrocystis is commonly known as the "giant kelp", "long bladder kelp", "kelp flag", "sea-ivy", or "devil's apron". Navigation charts indicating kelp generally include Nereocystis and Macrocystis without differentiation. The term kelp is usually limited to those members of the Phaeophyceae belonging in the families Laminariaceae, Lessoniaceae, and Alariaceae. The term kelp, however, is loosely used by many persons along this coast when referring to any seaweed, and is in some cases used only when referring to Nereocystis luetkeana. Some even apply the term to members of the Rhodophyceae. Tressler (88) says that "in Europe kelp usually means the ashes of seaweeds which are used for manufacture of potassium salts and iodine, although some persons use the term to designate a l l seaweeds from which these ashes are obtained. Along the Pacific coast of the United States the large brown seaweeds are called kelp." According to the classification used by Setchell and Gardner (76) the genus Macrocystis is included in the Tribe Macrocysteae, Family Lessoniaceae, Order Laminariales, Class Phaeophyceae. IV. GEOGRAPHICAL DISTRIBUTION: 1. Region Surveyed; The whole coast of British Columbia has been generally surveyed in the inside waters and around most of the larger islands between the Strait of Juan de Fuca and Dixon Entrance with the exception of the more exposed regions of Banks Island, Aristazabal Island, Estevan Island and the Barsdale Group, ao survey has been comprehensive of either the west coast of Vancouver Island or the Queen Charlotte Islands, although some casual observations have been made i n these regions verifying i t s general distribution. During the survey most of the areas reported by Cameron (L4) have been ve r i f i e d . Cameron (14) reported Macrocystis at Shusharties Bay, at the north end of Balaklava Island and i n Qlavdzeet Anchorage, none of -which were verified during the survey, but there i s no reason to question i t s occurrence at these points. Within the areas surveyed by Cameron (14) two beds i n particular of f a i r l y large size seem to have been missed; namely, at the east end of nope Island and on the north side of Malcolm Island. Both these areas are fringed with large beds of Nereocvstis which may have been the reason for missing them. In addition Cameron (14) reported distribution i n Dodger Cove, Banfield Creek and the neck of Useless Inlet, on the west coast of Vancouver Island. These areas have been ve r i f i e d by the writer and further evidence has been found i n Ucluelet Inlet and i n Sea Otter Cove near the entrance to San Josef Bay. Macrocystis has been reported to be present i n Blunden Harbour and a l l down the west side of Banks Island, Aristazabal Island and Estevan Island, but none of these reports has been authori-tatively made. None of these was verified other than the northern portion of the west coast of Banks Island. Macrocystis has been reported at Port Renfrew and the writer has found i t at Sooke as well as verifying i t s presence at Port Renfrew. It has also been located by the writer on the west coast of Vancouver Island i n the lee of Amphitrite Point. Macrocystis has not been found, other than as d r i f t , i n the inside waters along Vancouver Island from Ten-mile Point to Port McNeill. - 13 -In the Port McNeill region i t was not found south of a l i n e drawn from Ledge Point (near Port McNeill) to Lizard Point (north side of Malcolm Island) nor west of a l i n e from Lizard Point to Browning Islands i n the inner waters (see Chart XII). Observations made in the Queen Charlotte Islands indicate a general distribution along the north and west coasts. I t i s expected that i t w i l l be equally, i f not more prevalent on the east side (see Chart XIII). The writer has found i t a d r i f t i n Houston-Stewart Channel between Moresby and Kunghit Islands, and at the entrance to Tasu Sound on Moresby Island. I t was also found at the entrance to Port Louis and behind Frederick Island. In the l a t t e r location i t was quite abundant; one continuous bed i s present about 300 feet wide and over a mile long. Macrocystis was also found a d r i f t i n Masset Inlet and around the wharf, and a large bed about 300 feet by a mile or no re i s present along the east side of the entrance to Naden Harbour. It i s also present i n broad fringes behind Mippa Island i n Hippa Passage. C r i t i c a l examination of various plants i n the Queen Charlotte Island region indicates that the same species i s found there as at other points along the coast of British Columbia. 2. Explanation of Charts: Charts I to XIII give a complete geographical presentation of the distribution of Macrocystis along the coast of B r i t i s h Columbia i n so f a r as i t i s at present known and recorded. Chart I i s a small scale chart which presents a key to a l l other charts and indicates the extent of the survey. Charts VI and X - u -also give keys to other charts i n areas where more detailed surveys have been made i n addition to presenting the general distribution i n those areas. In a l l charts the presence of Macrocystis i s indicated by small dots. In one instance (Chart XI) the presence of Jiereocystis i s indicated by large dots. Except i n the large scale charts the areas indicating presence of Macrocystis are not to accurate scale, but are correspondingly exaggerated to show up on the charts. In a l l cases, however, where Macrocystis i s indicated i n any quantity a corresponding entry w i l l be found i n Table I indicating the estimated area or quantity present. Soundings and other navigational information have been ommitted from a l l charts i n order that the distribution would stand out, but these charts may be referred to standard navigation charts to obtain such information. In most cases where Macrocystis was found the limits of the beds or fringes were definitely established. Exceptions to this were made, however, i n the beds located along the west side of Dundas, Dunira and Melville Islands. The areas reported i n Table X were definitely established for these l o c a l i t i e s but their limits were not determined. I t i s quite definite that there i s considerable growth of Macrocystis along these shores which was not included i n this particular assessment— the f u l l extent was not v i s i b l e from the general shoreline, being either obscured by large reefs, rocks or, islands. The navigation here for a large boat, i n unfamiliar waters, was very dangerous, and without a power-driven smaller boat, i t was impossible to investigate the area f u l l y . - 15 -The Macrocystis reported at the north-west end of Banks Island (Norway Inlet) was also not completely investigated. While i t is fairly exposed here, there are many" passable channels amongst the small islands, which afford considerable protection, and the tonnage to be expected here is certainly much in excess of that reported i f the indications continue southward. 3. Method of Estimating: In the more general survey distances used for purposes of estimating were obtained, where possible, by locating landmarks and referring to the scale of the largest chart available, or by running the boat at a known speed and recording the time taken. In many cases, however, this was not possible and distances had to be estimated by comparison with the length of the boat or dinghy, training the eye to judge the distance. Density of plant growth was estimated by counting the number of plants in 100 sq. ft., and after determining the weight of an average plant, cut approximately 6 to 8 feet below estimated mean low tide level, the weight per square foot was calculated. Beds of Macrocystis are difficult to estimate due to the method of growth. Samples were weighed from time to time, and as a check against the method of estimating, a section a hundred square feet in area was blocked off with sticks and the whole area was cut to from 6 to 8 feet below the surface, collected and weighed. In all cases blank spaces and irregular beds were taken into consideration in estimating areas and an attempt was made to visualize a block size, or average width for the bed or fringe. In some cases, where only scattered plants or fringes were noted, only a very general - 16 -estimate was made, sometimes covering a considerable length of coast-li n e . In many cases, particularly where considerable growth was found, the beds have been shown i n Table I as made up of various sizes, although these separations have not always been made on the charts. Generally-these beds are more or less continuous, but due to variations i n width, and sometimes densities, i t was considered more accurate to treat them in this way as progress was made along the shore than to try to assess the whole area as one. In a l l cases the tonnages shown represent what may be considered as "standing crop" during the midsummer rather than annual potentiality. Indications of plant growth marked on the charts, for which no estimates are given i n Table I, may be considered as traces only or extremely scattered, thin fringes. Areas i n close proximity to the detailed distribution shown, having no markings, may be assumed as having no growth of Macrocystis of any consequence, or at least not evident at the time. Since observations were made at a l l stages of the tide, particularly i n the Prince Rupert area, i t should be pointed out that i t i s quite possible that beds have been missed. To estimate Macro- cystis beds one should view them at low tide, as i n many cases some of the plants, at least, are completely below the surface even at half-tide or less. Also, i n locations where there i s a strong t i d a l current the floating kelps may be completely dragged below the surface while the t i d a l current i s strong, and at slack tide be quite conspicuous. As a result of more accurate, although not extensive, checks made during the 1947 investigation on the size of beds i n the v i c i n i t y - 17 -of Deer Island, i t would appear that, due to the unfavourable tides, poor v i s i b i l i t y , and rapidity of checks made i n 1946, the widths of larger beds were underestimated by approximately one- to two-thirds. An aerial survey, combined with detailed "ground" work, would seem to be the most desirable method to determine the area of beds of Macrocystis i f i t i s considered necessary to more accurately assess the quantities and condition of these beds. Use of a transit might be of some use to calculate areas from the ground and water, but because of the irregu-l a r i t y of the shape of beds and presence of scattered blank patches throughout the beds, this would have i t s limitations. Provided adequate definition of the plant growth can be obtained by photographing from the a i r , these large blank spaces should be easily eliminated from areas. Estimation of density from the ground and water i s equally complicated by the fact that a great variation i s encountered, not only in the number of plants per unit of area, but also in the number of stipes arising from different plants. Also, the size and bulknof material necessary to harvest i n order to ascertain a typical cross-section i s beyond the f a c i l i t i e s of a small boat. It i s estimated that when there i s one average plant (or plant group)uper hundred square feet, there i s approximately 30-percent, coverage. Checks made on plots at the north of Deer Island (see Chart XI), except where blank patches occur, when viewed at 2- to 3-foot tides appear about 50 per cent, covered, but at approximately half-tide appear more like they were 30 per cent, covered. There does not seem to be any object i n calculating at extreme low or high tides. Cutting at the latter w i l l miss a l l or nearly a l l , and what more i s gained at a low tide i s either of l i t t l e consequence - 18 -or will damage too many young stipes to make cutting practical. This feature should be borne in mind if any photographs are taken from the air. If photographs are taken when the tide is extremely low, much of the plant material will be either completely exposed and lying on the rocks (and hence not show where the blank spaces occur amongst the beds) or will give a false impression of the coverage due to the great volume of plant material spread out in shallow water. The reverse is the case in photographing at high tide; that is, so much of the plant material is completely submerged that the coverage could not be calculated. It would seem that the ideal condition for photographing is a low-water, slack period, not lower than 2 to A feet above zero tide-level. It is beyond the capacity of a small boat, at least, to do any estimating which involves cutting and weighing an abundance of Macrocystis. For practical purposes, I believe that estimating of tonnage taking one pound per square foot, when coverage is approximately 30 to 50 per cent., gives a fairly approximate figure of the available tonnage from any bed. A careful measurement of the bed to the north of Deer Island indicates that approximately 568 tons could be expected from this bed at a single harvest (see Chart XI). An attempt was made to estimate plant material using a method that could be adapted to the facilities of a small boat. This was done, using a summary of the data present in 'fable VI. The basis of this calculation is the maximum diameter of the stipe; that is, by cutting a stipe off at any length, i t was found that there was a comparatively close correlation between the greatest diameter of the portion of the - 19 -stipe cut (at the cut surface) and the total weight of the upper, f u l l y laminated portion of the stipe. Thus, by counting the number of stipes and taking the diameter of each at the cut surface, one need not bother weighing the material, provided these figures are considered sufficiently accurate for the purpose required. In view of other variables to be taken into consideration i n any method of estimating quantities of this plant, this does not seem too great an assumption to make. The results of the data compiled in Table VI, and proposed for estimating quantities, are summarized below i n the form of standard weights estab-lished for computing the weight (approximate) of a f u l l y laminated stipe of Macrocystis in t e g r i f o l i a using as an index the stipe diameter at the cut surface: Diameter of Stipe Estimated Percentage Weight Used At Cut Surface Total Size Attained i n Estimating (Mm.). (Lb.). 3 - 5 25 0.50 6 - 7 50 1.50 8 - 9 75 2.50 10 - 12 100 A.50 In establishing these standards some range i n stipe diameter has been allowed because the stipe i s not always exactly circular i n cross-sectional outline, but i s inclined often to be somewhat oval. Actual weights of individual plants with the great number of stipes usually found may run up to 100 pounds or more. On the other hand, where only a few stipes are present from a holdfast, the total weight w i l l only be a few pounds. A sample plot measuring 20 by 20 feet at the north of Deer Island was cut somewhat lower than ordinary harvesting should be done, and was found to yi e l d 509 pounds, or 1.27 pounds per square foot. The - 20 -coverage at this point was estimated at about 75 per cent. About twenty plants were represented in this plot, giving an average per holdfast (or plant) of 25 pounds at the first harvest. 4. Equipment Used: The general survey was accomplished in a 46-foot tuna boat, supplemented by a 9-foot dinghy "powered" only with oars. Two dredges were used. One, constructed of cotton fish-netting protected from the bottom by a canvas apron and with a strap-iron "mouth", was used more in an endeavour to find associated small forms of algae. This one was found useless to obtain a holdfast of Macrocystis. being too heavy to operate from the dinghy and too difficult to manpeuver from the large boat by hand due to its weight and because the engine could not be idled slow enough. The second dredge was one improvised by the writer using a small bar to which a row of large fish hooks were wired. To the ends of the bar a rope was tied, making a loop, and opposite the bar a weight was tied on this loop. To operate this, as many of the stipes of a plant as possible were pulled in to the boat through the rope loop, were cut off at a convenient length to leave room in the boat to operate, and the bar was lowered by jerking another rope attached close to the weight on the loop, until i t had fallen to the bottom (this was determined by peering through the bottom of a glass-bottomed wooden bucket). The boat was then rowed around so that the rope would form a fairly tight coil around the stipes of the plant while the hooks fastened on to the basal part of the plant. With one person in the back of the boat holding tightly to the rope and as much of the upper part of the plant as possible, and the other making a supreme effort on the oars, i t was possible to raise an occasional holdfast from deep water i n this manner. Plants i n the shallower water, of fiourse, were much easier to obtain and often i t was possible to raise the whole plant, including the rock to which i t was attached. Accessory equipment included an ordinary garden rake and c u l t i -vator for reaching loose ends and a butcher knife for cutting. A beam-scale, 20-pound capacity, was used for taking weights used i n estimating. Samples were taken i n sacks when bulky and small samples were preserved in quart sealers i n a seawater-formalin-acetic acid solution for l a t e r laboratory examination. A set of hydrometers, t a l l glass cylinder and thermometer were used for determining densities i n the f i e l d and representative samples were taken i n small bottles and later tested by ti t r a t i o n method as a check against the hydrometer readings. A l l densities were corrected to 15 deg. centigrade. A colorimeter type pH k i t was used for taking pH readings on the seawater samples. During the extreme low-tide periods, much could be done while wading i n shallow water using hi-boots. Field glasses were constantly used. Without these i t would have been impossible to make any speed on the survey. With the existing colour differences between Macrocystis and flereocvstis (Macrocystis i s more brownish and Mereocystis blackish) and the general difference i n appearance of the plants at the surface of the water, i t was possible in a good l i g h t to make out a bed or fringe of Macrocystis, which would otherwise have escaped notice completely, considerable distance behind one of Nereocystls. - 22 -V. FACTORS AFFECTING DISTRIBUTION; 1. General; Macrocystis Integrifolia i s found growing in the v i c i n i t y of Deer Island, and elsewhere along the coast, attached to rocks from three feet above to twenty feet below zero tide-level, with the majority being i n less than 12 feet of water at zero tide. I t i s estimated that nearly 50 per cent, of the available Macrocystis i n the v i c i n i t y of Deer Island, at least, i s i n less than 6 feet of water and possibly 5 to 10 per cent, i s completely exposed at or near zero tides. An abundance of very small plants was found during the period of the investigation between the zero tide-level mark and up to about 3 feet above zero tide-level, but i n -dications are that only a limited number of these reach any great size in this location. Eventually they are either crowded out or torn loose and become fastened elsewhere. Transplantations of small plants by tying them with cloth strips close to rocky surfaces proved that these plants w i l l fasten themselves to new rocks i f torn loose, provided, of course, they are not washed up on the beach and desiccated for any length of time. The more securely fastened plants have their haptera closely adherent to the rough irregular portions of large rocks of sufficient weight to keep them from being washed away, but many plants reach considerable size fastened to almost f l a t rocks, provided there i s some roughness to the surface or an overgrowth of barnacles on the rocky surfaces. No plants of Macrocystis are found growing where the bottom i s inclined to be muddy or sandy, since there are insufficient large rocks to support the weight of the plants and prevent them from being - 23 -carried away. Occasionally stretches of sandy bottom appear to support growth, but on closer examination i t i s invariably found that the presence of a few scattered rocks or rocky reefs provide the attachment. In many cases, especially where plants were scattered, a l o t of sandy bottom was noticeable i n the v i c i n i t y , and in these cases almost invariably large sandy beaches were in evidence at the shore-line (Plates 7, 9, and 12). This was not always true, Welcome Harbour on Porcher Island being a decided exception, the whole area being composed of rocks with rough, and irregular surfaces (Plates 1 and 2). In many locations along the coast, where other conditions seem favorable for growth of Macrocystis. the great depth i s no doubt the deciding factor. In many cases rocks either go straight down from steep shores, or slope off so steeply that there i s l i t t l e , i f any, available bottom for more than a mere shore scattering of Nereoevstis plants. Macrocystis i n t e g r i f o l i a i s found along the outer sections of the coast of B r i t i s h Columbia i H narrow to wide fringes, not exposed to strong wave action, but usually protected by outer reefs, rocks or a heavy fringe of flereocystis. or i t may be i n a sheltered i n l e t where salinity s t i l l remains high. It i s found i n the northern region of the coast on the west Bides and slightly removed from the west sides, but s t i l l f a i r l y close to open ocean conditions, of the main outer islands (Dundas, Melville, Stephens, Porcher, Banks, and Aristazabal). The plants found i n deeper water are usually larger, with a greater number of stipes, but are much more scattered. The greatest number of plants i s found i n 12 feet or less and the plants are denser in that depth, although usually smaller individuals with fewer stipes. In the f a l l the south-east storms in the v i c i n i t y of Deer Island p i l e up vast quantities of Macrocystis on the beaches, particu-l a r l y along the shore adjacent to the Port Hardy airport. Plate 23 shows the results of a minor south-east storm on September 9, 194-7. In the f a l l of the year this plant material i s piled up on the beaches i n the v i c i n i t y from 5 to 12 feet deep. 2. Temperature: Temperature does not play as important a part, at least i n so far as local distribution on this portion of the coast i s concerned. Water temperatures, excluding occasional surface changes depending on the weather, did not vary appreciably during the period of the investi-gation. Within the limits of the calibration of the thermometer used, this may be set within the range 11 to 13 degrees centigrade. In every case where early-morning water temperatures were recorded, the temperature was 12 degrees centigrade. Occasionally during the day temperatures at the surface went as high as 16 degrees centigrade, but these occasions were rare. 3. Light: Vertical distribution seems to be a function of the l i g h t intensity. Plants i n deeper water are much darker i n colour. Plants transferred from shallow to deeper water become darker i n colour and when in water over 20 feet deep (with reference to zero tide level) seem to be retarded i n their growth rate (see Table III). 4. Salinity: Salinity appears to be the chief factor limiting local dis-tribution geographically. The whole general area of water under observation i n the v i c i n i t y of Deer Island i s of f a i r l y high s a l i n i t y . Density recordings made during the period at various locations around Deer Island show a range from 1.0223 to 1.0241 (compared at 15 degrees centigrade). In the immediate v i c i n i t y of the Keough River, depending on the stage of the tide, densities as low as 1.0129 were recorded, but at the same point at other periods ran as high as 1.0241. The average of densities recorded, however, excluding this t i d a l phenomenon i n the region of the Keough River, was 1.0229 (at 15 degrees centigrade). In no place where Macrocystis was encountered along the coast was the density found to be below 1.020 and i n general i t ranged from 1.020 to 1.024, the best beds being about 1.022. While no prevailing difference in water conditions seems to exist i n the immediate vic i n i t y of the Keough River, there i s an evident difference i n the appearance of the plants i n the v i c i n i t y of this fresh-water influx. The plants were noticeably healthier i n appearance, due probably to the variation i n sali n i t y created by the fresh-water influx preventing or retarding the growth of animal forms which were found to abound on the plants i n other l o c a l i t i e s . The plants in this v i c i n i t y were noticeably free of such animal colonies as Obelia and Foramlnlfera. although some were heavily covered with Melibe (the sea-slug). The latter, however, drops off almost as soon as the plant i s l i f t e d from the water. Obelia. particularly i n other locations, i s very plentiful on the plants and no doubt i s responsible for many of the laminae becoming b r i t t l e and breaking off. The - 26 -foraminiferans do not seem to be sufficiently abundant anywhere to cause much damage, and the only spot in which they were found in any great number was in a small patch at the north-east corner of Deer Island. The whole bed to the north of Deer Island is very heavily covered with Ubella. although confined mainly to the surfaces of the laminae, from the lower extremities of the plants in many cases up to within a few feet of the terminal laminae. Bryozoans (moss animals ) are quite frequently found on both the laminae and stipes, but are not so abundant in this region as was seen last year, for example, at Hope Island. 5. Currents: The increased tidal current at the sharp point of Deer Island at the north-east corner, at which spot the foraminiferans were found abundant and the Obelia almost lacking, may have been responsible for the differences noted. In general, however, the tidal flow in this region is not in excess of 1 to 3 knots per hour. VI. ECOLOGICAL ASSOCIATIONS: 1. Other Plants: Where growth of Macrocystis integrifolia and Nereocystis luetkeana occur together, the latter is invariably in the deeper water at the outer edge of the Macrocystis bed, with some intermixture at the extreme. Along the inner margins of the Macrocystis beds, mixed in with the plants in shallower water, Egregia Menziesii is abundant. The long strap-like, tough portion of this plant may reach a length of 15 feet or more. This plant is commonly called the "feather-boa kelp." It is also supported in the water by small pneumatocysts or bladders and produces many stipes arising from a common, large, conical holdfast. - 27 -Any of this latter plant harvested with Macrocystis would probably prove somewhat troublesome due to its extreme toughness. Other forms of algae, most of them minute, are epiphytic at least (some may be parasitic), but these are of little consequence. 2 . Animals: A wide variety of marine animals is found in close association with the beds of Macrocystis integrifolia. In most cases this association is probably just a provision of a convenient, sheltered spot in the water in which these forms live, here they are sheltered from strong wave-action amongst the laminae and stipes, or in many cases amongst the irregular, branching rhizomes and haptera close to the bottom of the sea. Many kinds of small, as well as large, fish occur amongst the plants, and some evidence of spawning on the laminae was detected. The molluscs Melibe and Tegula are found on the laminae in great abundance, the former particularly in the vicinity of the Keough River. The worst enemies of Macrocystis appear to be the many forms of crabs, large and small, as well as other crustaceans, which eat pieces out of both the laminae and stipes. Another enemy, which is more abundant here, is ubelia: these moss-like animals completely cover the surfaces of some laminae, particularly the lower ones on the stipe. Their harmful effect on the laminae seems to be a result of exclusion of light and contact with the water, resulting in the laminae becoming brittle and easily broken. Membranipora membranacea is almost equally as disastrous to the laminae as is Obelia. This animal forms an encrusting calcareous layer over the surfaces of the laminae and produces much the same results as Obelia. In one location at the north-east corner of Deer Island a foraminiferan i s quite abundant on the laminae, but i t i s quite small and does not seem to cause any noticeable damage. The Macrocystis plants at Porcher Island (Welcome Harbour) on the average vere larger, denser and less covered with marine fauna. Those i n Norway Inlet had considerable Obella on the laminae, in many cases with only a few feet at the upper end of the stipe being actually free from these animals, while those on the north and north-east ends of Banks Island were found with considerable Teeula. The Macrocystis bed at the east end of Hope Island had a great deal of Membranipora membranacea. i n some cases the large laminae being completely covered with the encrustations. VII. LIFE HISTORY: 1. Alternation of Generations: During this investigation no attempt was made to follow the microscopic stages which take place i n the life-cycle of this plant. The following brief account, some of which i s from reference material, w i l l provide a background which i s necessary i n order to understand the importance of certain phases i n the life-cycle of Macrocystis. This i s important, particularly from the standpoint of conservation. A diagrammatic representation of the life-cycle i s shown in Fig. 1. On the mature, massive plant of Macrocystis, which i s the form ordinarily seen and known as the sporophyte (Fig. 1 (1)) , there are certain basal laminae (sporophylls)(Fig. 1(2)) borne on the stipe close to the holdfast of the plant. These are specialized for reprod-uction. This large, sporophytic plant i s the asexual generation i n the - 29 -life-cycle; that is, i t produces asexual spores. These reproductive cells are formed in sac-like structures known as sporangia (Fig. 1 (3)), which are produced, together with paraphyses, on both surfaces of the sporophylls. When these reproductive cells are mature they are liberated from the sporangia as free-swimming zoospores (Fig. 1 (5)). They are microscopic, bean-shaped, pigmented, unicellular structures, each bearing two laterally affixed flagella which produce the movement. After a period of motility these zoospores settle down, lose their flagella, and develop into either female or male plants (gameto-phytes). These sexual plants are filamentous in structure, consisting of only a few cells (Fig. 1 (7) and (8)). The female and male plants produce eggs and sperms (spermatozoids) in structures known as oogonia and antheridia respectively. The motile sperms (spermatozoids), when released from the antheridia at maturity, swim about, and one fertilizes the egg (ovum) of the oogonium on the female plant, resulting in form-ation of a zygote., The zygote then undergoes repeated divisions and growth, resulting in the young, blade-like form which is the young sporophyte (Fig. 1 (14) and (15)). This in turn, by a definite method of splitting and rapid growth (Figs. 2 and 3), ultimately results in the mature sporophyte with its long stipes bearing numerous laminae and reaching to the surface of the water. No culture experiments were conducted to determine the micro-scopic phases of this cycle, but periodic examination of the sporophylls on mature sporophytic plants revealed that active zoospore production was taking place at a l l times during the period July 1 to Setpember 15. - 30 -The limits ( i f any) of this period, therefore, were not determined. Brandt (10) states that spores are produced at a l l times of the year fey Macrocystis. Observation on the abundance of young sporophytic plants, however, seems to indicate that there i s probably a period when sporulation i s not so active at least, since young sporophytic plants (single, undivided-lamina stage) were conspicuously more abundant during July than during the early part of September. This, of course, may be the result of other factors. Zoospores of M. pyrifera have been reported (4-6) to germinate in twenty-four hours i n culture. Young sporophytes have been reported to occur i n cultures varying from seventeen days old (46) to forty-four days old (58). It i s not known, however, at least for plants i n these waters, just how long this period takes for Macrocystis i n t e g r i f o l i a under natural conditions. Brandt (10) estimated that a young sporophyte of Macrocystis when f i r s t readily discernible as an undivided blade was approximately three months old. On this basis i t i s probable, therefore, that from four to five months are required from the time mature zoospores are liberated u n t i l the new sporophyte reaches this size. 2. Early Development of the Sporophyte: Table II shows the results of growth found amongst very small plants. From these results i t i s apparent that some variation exists. Much variation could be expected, particularly under natural conditions where the plants are more or less preyed upon by animals, obstructed by other plants, and influenced by differing physico-chemical conditions. In Table II, plants No. 52 to No. 56, inclusive, had no pneu-matocysts (floats or bladders) evident on August 4 , or at most a slight i. .... i - 31 -thickening, but at the end of the experiment (September 13) Nos. 53, 55, and 56 had small, but well-developed pneumatocysts. Nos. 52, 54, and 125 s t i l l had no pneumatocysts by September 13. I t i s obvious, by comparing the figures i n Table II with those shown for somewhat larger plants (Table I I I ) , that the early rate of growth, up to the time when ful l y developed stipes with separated laminae develop, i s comparatively slow. y On the basis of these comparisons, i t i s estimated that by the time a young sporophyte has developed i t s f i r s t pair of stipes to a length of 6 feet, i t i s approximately eight to ten months old; that i s , approximately three months were required to reach the undivided blade form f i r s t discernible, three additional months to reach a height of about 12 inches, two additional months to reach approximately 24 inches in height, and another two months to reach approximately 6 feet i n height. Growth rates from this point on w i l l be discussed under "Later Develop-ment of the Sporophyte." As already indicated, plants s t i l l i n the form of an undivided blade were found even by the middle of September, although not as abundant as i n the middle of July. I t i s obvious from examination of mature plants, which were found during the whole period of the investigation, that re-production asexually by means of active zoospore liberation was taking place during the whole period. Thus, during this period plants from a single undivided blade to a large mature plant, actively sporulating, were found at a l l times. The presence of a greater abundance of younger plants i n the middle of July may be an indication that the autumn and winter spores, which produce gametophytes i n the winter, have greater v i t a l i t y or develop best during conditions prevailing through that - 32 -period. These gametophytes, according to the estimates mentioned above, produce the sex c e l l s which, after fusion, result in develop-ment of a young sporophyte that reaches the size of a small, undivided blade by July. To some extent i t may also be because there are fewer plants l e f t i n the late winter and early spring on account of the severe winter storms, and hence fewer spores are available to be set free i n the water at this period. From the number of spores liberated throughout a season, un-doubtedly the mortality rate i s very high, although suitable bottom or some other environmental factor may be a limiting factor preventing the increase i n the size of the beds. I t i s to be expected also that the motile, unicellular stages i n the life-history of Macrocystis i n t e g r i f o l i a play an important part i n the phytoplankton of this area and are probably considerably reduced i n numbers by zooplankton, thus contribu-ting to the general economy of this portion of the sea. 3. Later Development of the Sporophyte; The method of division of the single blade of a young sporo-phyte, which ultimately results i n the massive complex plant with i t s many stipes, follows a definite procedure. This development has been carefully followed during the investigation and i s diagramma t i c a l l y represented i n Figs. 2, 3, and 28. The following briefly summarizes the significance of the various symbols used i n these three f i g u r e s : — "DI," "DII," and "Dili."—These are primary divisions taking place in development of the plant form. Each of these divisions results i n the formation of a pair of primary segments which behave similarly. DI i s the only true dichotomy, a l l other primary divisions being sub-dichotomous, alternating from outer to inner edge progressively i n order of development and a l l secondary divisions being unilateral. "dl."—This i s a secondary division which takes place i n a primary segment, resulting i n an outer secondary segment which develops into a stipe (with unilaterally developed laminae) ultimately reaching to the surface of the water after later divisions, and an inner secondary segment which acts as the growing "point" (actually a segment with a subterminal meristem) and which ultimately undergoes a primary division to repeat the cycle and continue growth of the system. From the above i t w i l l appear obvious, since subsequent to the f i r s t primary division a l l divisions are theoretically paired, that the f i r s t primary division results i n the formation of one pair of stipes and one pair of growing "points." The second primary division results, therefore, i n a total of four stipes (two pair) and four growing "points" (two pair), the third i n eight stipes and eight growing "points," and so on—the fourth, f i f t h , and sixth producing sixteen, thirty-two, sixty-four, etc.—theoretically ad infinitum. Thus by the time there have been, for example, four primary divisions, there w i l l be thirty stipes i n progressive stages of differentiation. " d 2 . " — T h i s i s the f i r s t division i n the outer secondary segment (the segment which ultimately produces the differentiated stipe) and separates off the basal lamina of a stipe, usually without a pneumatocyst at i t s proximal end. This basal lamina undergoes a series of dichotomous . divisions into a number of "bladelets" which become sporophylls. "d3."—This i s the second division i n the outer secondary segment and separates off a lamina which may be the same i n appearance, development and function as that resulting from a d2 division, or may have the addition of a pneumatocyst at i t s proximal end, but otherwise undergoes similar divisions into a number of "bladelets" which become sporophylls. The lamina produced by this division i s Immediately above that produced by the d2 division. "d4."—This i s the third division i n the outer secondary segment and usually separates off a st e r i l e lamina with a pneumatocyst at i t s proximal end, but i n some cases may result i n formation of sporophylls as in the case of a d3 division. "d5," wd6," "d7," etc.—These are fourth, f i f t h . sixth, etc. divisions respectively, and separate off sterile laminae, progressively larger from basal to terminal region of stipe, each with a pneumatocyst at i t s proximal end join-ing i t to the stipe. Actually, one might say that the d l division, which results in an outer secondary segment ultimately forming the stipe, and an inner secondary segment which acts as a growing "point" for the production of further stipes and growing "points," i s probably a division of the growing "point"—actually a subterminal growth region—one segment con-tinuing to produce new laminae along a stipe and the other segment con-tinuing to produce new stipes and growing "points" by additional seg-mentation. In Fig. 28 the arrangement i s somewhat modified to show the j relationship of the various stipes arising, but i n nature the branched system occurs so completely overlapped and tangled that sometimes i t i s practically impossible to determine whether one has a single or several holdfasts mixed up together before one. "S 1."—This i s the f i r s t ma.ior stipe to develop (paired). "S 2."—This i s the second major stipe to develop (paired). "S 3."—This i s the point at which the third ma.ior stipe develops (paired). "S 4," "S 5," etc.—These are fourth, f i f t h , etc., stipes (paired) i n order of magnitude. These are d i f f i c u l t to determine in proper order, but indications are that this i s the order usually followed (as i l l u s t r a t e d ) . Beyond this point, however, because of the rapidly increasing number of growing "points" ( "P") , the order of development i s obscured with respect to the earliest formed stipe at least. "X."—This i s the dying-out region of the holdfast— the earliest part formed and the f i r s t to die away under natural conditions. "Y."—This i s the portion bearing the most mature stipes reaching to and stretching along the surface of the water at times. "Z."—This i s the portion where the newest stipes - 36 -are being formed and the location of the basal growing "points." The dotted portion i n Fig. 23 indicates the approximate position and development of the broad, creeping rhizome-portion of the holdfast, from the two edges of which root-like structures known as haptera develop (see Fig. 27). At a l l times during the period of this investigation one could find a l l stages i n development of stipes from a common mature holdfast from the early, s t i l l incompletely separated stage, as shown in Fig. 5 (•&• to C), up to the f u l l y developed stipe (see Plate 18) reaching to and extending along the surface of the water. When mature, these stipes reach a length of 35 to 40 feet. Although a length was recorded by the writer of approximately 60 feet in Welcome Harbour (Porcher Island) i n 1946 this seems to be exceptional. Nothing as long as this was encountered i n the v i c i n i t y under investigation i n 1947. I t would appear, however, that plants of Macrocystis integr i f o l i a grow somewhat larger i n these waters than i n waters off California.> Setchell and Gardner give 5 to 8 meters as the length i n the latter location. (76). Lanceolate laminae (Fig. 19) up to 31.5 inches long and 6.5 inches wide are produced at f a i r l y regular intervals along the whole length of the stipe, becoming progressively smaller and closer together towards the terminal lamina, where they are differentiated (Figs. 20-23). The terminal lamina i s broadly falcate. Plate 15 and 16 show terminal laminae taken from mature stipes which reached to the surface of the water, Indicating how new laminae are produced and detached. As many - 37 -as twenty-five to thirty laminae may be i n progressive stages of separation from- this terminal lamina when i t i s situated on a mature stipe that reaches close to the surface of the water. On shorter, less mature stipes, however, the terminal laminae (Fig. 20) may have as few as three or four laminae in progressive stages of separation. Hence the proportion between the length and width of the terminal lamina, as well as the number of segments occurring in such laminae, i s extremely variable. Although this same variable characteristic was not studied in the v i c i n i t y of Deer Island during the winter, a brief v i s i t to Sooke, on the southern end of Vancouver Island, was made during late December for this purpose. At this time, at least, the more linear nature of the terminal laminae of short stipes was even more pronounced (Fig. 22). In addition, a l l other laminae higher up on major stipes s t i l l intact presented a decidedly linear appearance as did also the sporophylls (Figs. 16, 17, and 18), in comparison with those examined ea r l i e r i n the year (Figs. 21, 22, and 23) at the north-east portion of Vancouver Island. This may, however, in this instance be somewhat of a seasonal variation rather than one of location, and no attempt has been made to study the matter further. Each s t e r i l e lamina normally i s produced singly and attached at the proximal end to a solid stipe by means of a pyriform pneumatocyst (Fig. 19). In one case, however, two laminae were found developed from a common pneumatocyst (Plate 19), although each possessed i t s own intervening pneumatocyst. These pneumatocysts, which serve to keep the main portion of the plant afloat, and possibly aid i n distribution as well, start as small swellings at the base of the laminae, but as growth - 38 -continues may be up to 2 inches long and 1 inch i n diameter. The pneu-matocysts are hollow and are attached to the stipe by a short stalk. Pneumatocysts attached towards the base of the stipe, particularly those on which sporophylls are formed, are generally shorter and larger i n comparative diameter than those produced higher up on the stipe. The maximum separation of the laminae found on any stipe was 7.1 inches, which was i n the lower portion of the plant. Usually the lower portion of a mature stipe for a distance of up to 5 feet, or even as much as 15 feet i n some cases, i s completely devoid of laminae, having become either damaged by predators or wave action. Even the pneumatocysts are sometimes missing i n this same region. Table VI outlines various proportions of different sizes of stipes measured and weighed to show this relationship and i t s significance in determining the best depth at which the plants should be harvested. The diameter of the stipe varies with age or length of the stipe, attaining a maximum of about 0.5 or 0.6 inch i n diameter. The diameter i s not always uniform, very often being almost oval i n cross-sectional outline. The diameter of the stipe i s somewhat less i n the older portion of the stipe close to the holdfast (probably a mechanical feature) but increases in diameter i n the vi c i n i t y of the largest laminae and gradually tapers down again towards the terminal lamina, where f i n a l l y i t cannot be differentiated from the terminal lamina tissue. The surfaces of the laminae are deeply rugose and the edges are denticulate. The denticulations are 1 cm. or less apart and up to 9 mm. long, occasionally dichotomously branched. This characteristic - 3 9 -i s sometimes less pronounced i n newly produced laminae at the apex but becomes more pronounced with age. The sporophylls usually have coarser denticulations and are more deeply rugose, thicker, and tougher. The sporophylls may be found up to 1 9 . 5 inches or more in length—this probably should be somewhat greater, as the extremity of the bladelets becomes tattered and torn with age. Sometimes only one group of sporophylls i s formed close to the lower attachment of a stipe to i t s holdfast, but usually two or three groups are produced. The lowermost group i s usually without a common pneumatocyst, and the next superior one or two groups each bears a common pneumatocyst at the proximal end. In a l l cases these sporopnyll-ous laminae undergo successive divisions, producing ultimately as many as twelve bladelets i n a group. On both surfaces of these bladelets the sporangia are borne. In one specimen a rudimentary pneumatocyst was found bearing sporophyll blades, and only a small pore i n the centre of the pneumato-cyst was present, instead of being either lacking or quite hollow. Normally, when f u l l y developed, the pneumatocysts are a l l hollow, efficient gas chambers. Upwards of four major stipes may be found on the one holdfast, but usually not more than about thirty complete stipes are found (Fig. 21). The holdfast of the sporophyte starts as a small irregular expansion at the base of a short stipe (Fig. 2 (A)). Branched haptera (dichotomously branched for the most part) soon grow from the holdfast with sometimes as many as 8 divisions taking place, but usually not more than four successive divisions occurring. Sometimes the haptera become - AO -i quite extensive when developing from the flattened or rhizome-portion (Figs. 26 and 27) of the holdfast. As the stipe elongates, the haptera or root-like structures arise from along the lower portion of the stipe (Fig. 2A A and B). The lower portion of the stipe (and branches of the stipe) after bending over, probably partly due to the weight of the superior growth, becomes depressed, forming the characteristic flattened, subligulate, irregular, creeping, overlapping rhizome-portion of the holdfast from which d i -chotomously branched haptera arise along both edges (Fig. 26). The il l u s t r a t i o n of the plant in Fig. 2A shows both major branches bent over i n approximately the same direction. This i s not always the casej many extend each major branch out i n opposite directions (Fig. 25), or other peculiarities and tangled arrangements may result. As growth of the plant continues, new branches form, stipes arise, and the older portion of the rhizome (lower portion of the old stipe and branched haptera) dies away, although s t i l l allowing the plant to continue sending up new stipes each year because of continued pro-gress of the growing "points." The older parts of the rhizome and haptera of the holdfast, unlike the flexible newer portions, are more b r i t t l e . In older plants (after the f i r s t two or three major divisions have been completed) the growth of haptera advances up to the point of, and passes, the portion of the stipe at which the branching of the plant took place (Fig. 26). The haptera develop f i r s t as small, round protuberances. As i t dies off behind, the rhizome-portion thus splits up into several portions, so that ultimately the same plant may become - a -separate "plants" by isolation of i t s several growing "points." In fact the further decay advances from the i n i t i a l point of contact against the substratum, the more segments become free, although there i s usually so much overlapping of the rhizome and tangling of haptera that this characteristic i s not alwayB very apparent. Many holdfasts do indicate, however, where the main bulk has been torn away in the centre and where new growing "points" are s t i l l carrying on growth as indicated i n Fig. 2§. The f i n a l gross nature of the holdfast depends on the way i t spreads out. Some spread over a f l a t surface, making the whole flattened rhizome obvious, and the haptera seem to be extended more evenly from the sides, i n many cases remaining comparatively short i n length. However, sometimes one half of the main dichotomy creeps over the other and a more elevated mass results, hopelessly tangled up. The haptera s t i l l only come off at the two edges, even when the rhizome i s not closely adherent to a f l a t rocky surface, although usually they bend downwards more predominately i n the l a t t e r case. From a l l the observations made, i t would appear that a plant must be at least ten months old before i t has produced any stipes approaching harvestable size and before sporulation commences. 4. Growth Rate and Proportions: As a result of general conditions and the necessary limitation of the proposed program, i t was decided to concentrate on phases of the study which could be carried on i n the more immediate v i c i n i t y of Deer Island. Hence, the growth phase of Macrocystis i n t e g r i f o l i a . effect of cutting, etc., have received special emphasis during the latter investigation. - 42 -A check on the growth rate of new growing "points" near the holdfast i s d i f f i c u l t and not very significant within the period of the investigation. These early divisions are comparatively slow-forming, only two or three new "s p l i t s " being formed in about a month. Development i s comparatively rapid, however, once a stipe has become differentiated and begins to elongate, and lends i t s e l f more readily to measurable analysis. Tables IV and V outline i n detail the results of various experiments set up to determine growth rates, and Table VI attempts to " present a detailed analysis of the distribution of weight in the various parts of the plant. As might be expected, a great variation i s found i n growth rates, both under normal conditions of growth as well as under exper-imental conditions. Tide-action and wave-action tend to break off laminae, and when these are terminal i n position the production of new laminae ceases. These terminal laminae are also preyed upon, as i s the whole plant, by various crustaceans and molluscs. Other algae are epiphytic. The rate of growth also appears to vary with the depth at which the plants are growing, probably as a result of variation i n li g h t intensity. I t w i l l appear obvious from these conditions that the photo-synthetic area i n any group of plants w i l l d i f f e r widely. The cells of the plant capable of carrying on photosynthesis are present only i n the surface layer of cells; that i s , the pigmented cells. Hence, theoretically, the basis on which these plants should be compared to show relative rate of growth or increase i n weight in response to different conditions should - A3 -be compared to show relative rate of growth or increase in weight i n response to different conditions should be surface area. Obviously this i s impossible, or almost so, without a great many detailed measurements. The basis for comparison, therefore, has been selected as either the total length of the stipe, the laminated length of the stipe, or the diameter of the stipe, since i t i s f e l t , after observing many of these plants, that these measurements are the only ones which can be practically used. The comparison made of various plants of different stipe diameter and length i s shown in Table VI. Table VII shows the result of an experiment set up to show the effect of cutting off the terminal laminae of stipes. While some elon-gation i n the stipe takes place, i t i s comparatively small and i t i s Impossible for any further production of laminae to take place. The laminae already formed continue growing; that i s , they increase i n size, but do not give rise to additional laminae. Table II attempts to show the period required to complete the early divisions that take place i n the plant. The period required to increase the length of a small stipe appears to be greater than that required when the stipe i s older and longer. This factor should probably also be related to photosynthetic area. Tables III, IV, and V show detailed results of experiments conducted to show the growth rates of major stipes, either severed and kept anchored or l e f t naturally fastened to their holdfasts, as indicated. During the period of the experiments at least, i t would seem that there i s no significant difference i n the rate of growth - AA -(elongation) when severed or l e f t attached, provided the severed stipe i s maintained i n an equally satisfactory position. Those stipes which were severed were tied by means of strips of cotton to a rope loop, which i n turn was anchored to a large stone. This method gave easy access to the plant when another rope, to which a float was attached, was fastened to the rock. Usually about ten severed stipes were fastened to a common loop and each was individually tagged with a numbered aluminium tag. In Borne of the experiments the stipes became heavily weighed down with Obelia and settled more deeply in the water, probably thus contributing to the variation encountered. It was noted that in a l l cases whefce plants were removed to deeper water that, while growth s t i l l continued, i t was very much reduced and production of additional laminae at the apex stopped or was retarded. This usually resulted in the apex ending in a single pneumatocyst or bladder with f u l l y developed lamina formed at i t s end, macroscopically, at least, no different from the laminae farther down on the stipe. Counts of laminae show that the number of laminae s p l i t off, or indicated by a s p l i t at the base of the terminal lamina before moving to deeper water, continued to develop and reached almost normal, size, but no additional s p l i t s formed. Occasionally, instead of ending i n one pneumatocyst with a single blade, the one pneumatocyst gave rise to two laminae and very rarely one was found with two secondary pneumatocysts on the end of a primary pneumatocyst, each bearing a separate lamina. The colour of these plants when transferred to deeper water became much darker. In normal plants, whose terminal laminae reach to the surface of the water, the upper portion of the plant i s from olive-brown to almost yellow-brown, - A5 -but the laminae in deeper water are always a much darker brown. No significant differences were noted, in so far as growth rate i s concerned, in plants observed at the various locations under investigation. Weights before and after the experimental period were not taken because i t was f e l t the handling required to make the i n i t i a l weighing (which could only be total weight) would only increase the possibility of damage to the plants by breaking off additional laminae and would also necessitate keeping the plants out of water for an additional period. Examination and comparison of a l l tables giving data on growth rates of Macrocystis show considerable variation. In so far as this preliminary work has been carried out, however, i t seems f a i r l y evident that major stipes under favorable conditions grow in excess of 2 inches per day, at least during this period of the year. The maximum growth rate (elongation) recorded was an average of 3 . 1 0 inches per day for a period of twenty-nine days. I t may be found that beds of Macrocystis w i l l thicken somewhat after cutting has taken place because more li g h t w i l l penetrate when the heavy top growth i s cut away, and also there w i l l be less plant weight for the waves to strain against, resulting in fewer holdfasts being torn loose. There i s , however, no evidence of stimulation of existing or development of new stipes as a result of cutting. The growing "points" seem to continue their development independently. Several holdfasts were observed during the period, the major stipes of which had been cut off at several feet above the holdfast. In no case was there any noticeable evidence of an increase i n the growth rate or - 46 -decrease as a result of cutting. These early divisions, however, even under normal conditions, as already explained, take place comparatively slowly, so that i t i s possible there i s some effect which could not be detected. A longer period of observation on more specimens would be necessary to determine this. The stipes severed do not decay immediately, provided some laminae remain, but as soon as the laminae become negligible from breakage or being eaten off, the stipe begins to decay from the cut surface back to the rhizome. Hence, any stimulation as a result of cutting can only come from the growing "points" already established and no new growth can be directly established at the position of the cut as a result of cutting. In some cases during the period of observation, holdfasts checked for regeneration showed slight development of the already-formed new stipes by the differentiation of their basal pneumatocysts, but these did not d i f f e r noticeably from the normal development, and in many cases only the occurrence of one or two new splits at the growing "point" developed during the period. In other cases no further * "s p l i t s " had even developed in the new stipes at the growing "point" i n slightly over a month. Some of the smaller plants under observation for this phase of growth were l e f t on their original attachments and observed at the low-tide periods, but some of the larger plants were pulled loose and anchored so that they could be reached when desired. When .the stipe dies back from a cut surface, the intermediate region (cortex) becomes soft and pulpy, leaving an outer pigmented layer (the meristoderm) which separates off and an inner rather tough '•core" (the medulla). Finally a l l i s sloughed off except the central thread-like "core," which i s the last to disappear right back to the holdfast. If intensive harvesting of beds of Macrocystis i s to be carried out and the maximum tonnage realized from any bed, i t i s the opinion of the writer that definite precautions must be taken, partic-ularly with regard to the time at which the plants are cut, as well as the height from the bottom at which the plants are cut. If these two precautions are taken, there does not seem to be any reason why the same beds could not be expected to yield two good harvests. It i s also believed that i f these precautions are taken, the total tonnage obtained during the year from any bed w i l l be greatest. It would seem advisable to survey the beds to be harvested, making soundings and marking prominent rocks which obstruct otherwise clean sweeps, with a view to plotting sections and thus determining the most advantageous period and depth at which harvesting should be done. By marking off these areas and cutting at known depths, more of the short stipes may be conserved. But i f the tops of these stipes are cut off, no further laminae w i l l develop; the laminae already formed w i l l be more broken, and i f broken off completely, the stipes w i l l die back to the holdfast. The short stipes are the ones which would normally develop and be harvested at a later season, but when the terminal laminae are severed, they w i l l not continue growing into long, f u l l y developed stipes. The result of harvesting at different stages of the tide with respect to the point of attachment of a plant on the bottom and i t s t e r m i n a l laminae i s d i a g r a m m a t i c a l l y r e p r e s e n t e d i n F i g . 4. From o b s e r v a t i o n s made d u r i n g the p e r i o d o f t h i s i n v e s t i -g a t i o n , i t i s i m p o s s i b l e t o determine the age o f some o f the s t i p e s p r e s e n t . The appearance o f some o f the l o n g e s t s t i p e s , however, i n d i c a t e s t h a t they a r e probably s t i p e s which have been c a r r i e d through w i n t e r s torms, and as l o n g as the t e r m i n a l laminae were not b r o k e n , growth and the p r o d u c t i o n o f new laminae a t the apex have taken p l a c e . I t would be a d v i s a b l e t o check the amount, s i z e , and c o n d i t i o n o f p l a n t s i n evidence i n the e a r l y p a r t o f the y e a r to d i s c o v e r how much o f the p r e v i o u s y e a r ' s growth p e r s i s t s through the w i n t e r and to determine whether the f i r s t c u t t i n g s h o u l d be l e f t u n t i l l a t e r f rom the s t a n d p o i n t o f t o t a l p o t e n t i a l tonnage f o r the c u r r e n t y e a r . C e r t a i n l y a h a r v e s t made too e a r l y , i f made too deep, c o u l d v e r y g r e a t l y cut down the t o t a l tonnage t o be expected i n the e a r l y - o r mid-summer p e r i o d . I f , as suspected, a number o f these extremely l o n g s t i p e s (which are u s u a l l y devo id o f laminae and o f t e n pneumatocysts f o r as much as 15 f e e t o r more above the h o l d f a s t ) a r e c a r r i e d over through the w i n t e r , i t i s p o s s i b l e t h a t a c u t t i n g e a r l y i n the y e a r , made q u i t e s h a l l o w i n o r d e r not t o sever the t e r m i n a l laminae o f the succeeding s t i p e s formed, would be advantageous. T h i s procedure might ensure more v i g o r o u s , c l e a n e r h a r v e s t s a t subsequent p e r i o d s , s i n c e i t would p r o b a b l y hasten decay o f the o l d p o r t i o n o f the h o l d f a s t and p e r m i t a c t i v e growing " p o i n t s " o f the h o l d f a s t to spread out and thus p r e v e n t crowding and damage t o the new s m a l l s t i p e s be ing formed. I t would a l s o p e r m i t g r e a t e r c i r c u l a t i o n of water and p e n e t r a t i o n o f l i g h t , and by e l i m i n a t i o n o f the t a n g l e d masses o f o l d s t i p e s would leave l e s s b u l k o f p l a n t m a t e r i a l t o be dashed by the waves and l e s s e n the p o s s i b i l i t y o f t e a r i n g out the h o l d f a s t - A9 -during storms. Uare would have to be taken, as previously pointed out, that the cutting i s not made too deep at this period. I f the term-inal laminae are removed, although stipe elongation may take place to some extent, the possibility of many laminae remaining by the time the next harvest i s proposed w i l l be very slight, and those that do remain wil l be so badly broken or eaten that the portion of stipe obtained w i l l be poor i n condition, light i n weight, or even decaying back to the holdfast. I f such an early harvest were made, i t would probably y i e l d a minor tonnage, but might be worthwhile from the standpoint of increas-ing the volume of subsequent harvests during the current year. This point could only be determined by experimentally cutting a portion of a bed i n this manner. The f i r s t major cutting would probably be possible i n the early summer, but this harvest should not be made very deep i f a subsequent harvest i s anticipated in the same season. Provided only the major stipes are cut and the terminal laminae of the shorter stipes not damaged, a second major harvest should be attainable by the end of the summer. This l a s t harvest could be made deeper, as the probability of harvesting being practical through the winter seems remote. To regulate the cutting depth, both the tide and depth of the holdfast attachment must be taken into consideration. To cut only the upper portion of the plant, therefore, one must either cut less deep at a low tide or cut the normal depth of the cutting mechanism used at a higher tide. Whatever cutting policy i s adopted, under no circumstances should cutting be permitted lower than 3 to A feet above the holdfast on - 50 -t h e b o t t o m . The r e a s o n f o r t h i s w i l l a p p e a r o b v i o u s f r o m t h e f o r e g o i n g e x p l a n a t i o n s o n t h e p o s i t i o n a n d f u n c t i o n o f t h e s p o r o p h y l l s . I t w o u l d a p p e a r g o o d i n s u r a n c e , t h e r e f o r e , o n t h e p a r t o f a n y o p e r a t o r t o s p e n d some t i m e i n t h e o f f - s e a s o n s o u n d i n g a n d p l o t t i n g a r e a s w i t h some c a r e . I f t h i s p r a c t i c e i s a d h e r e d t o , i t w o u l d n o t seem n e c e s s a r y t o l e a v e s t r i p s u n c u t , p r o v i d e d t h e method o f c u t t i n g d o e s n o t t e a r l o o s e a l a r g e p r o p o r t i o n o f t h e h o l d f a s t s , s i n c e a l a r g e number o f t h e p l a n t s i n d e e p e r w a t e r (12 t o 15 f e e t o f w a t e r a t z e r o t i d e ) w i l l a l w a y s h a v e t h e i r r e p r o d u c t i v e p a r t s o u t o f r e a c h o f t h e c u t t i n g - b l a d e s b e c a u s e o f r o c k s a n d r e e f s . W h a t e v e r c u t t i n g a p p a r a t u s i s u s e d i n h a r v e s t i n g M a c r o c y s t i s , i t w i l l be n e c e s s a r y t o c u t i n t o t h e d i r e c t i o n o f t h e c u r r e n t , o t h e r -w i s e , i f one c u t s w i t h t h e c u r r e n t , much o f t h e m a t e r i a l w i l l b e p u s h e d u n d e r t h e c u t t i n g a p p a r a t u s a n d m i s s t h e b l a d e s , o r i f c u t t i n g i s a t t e m p t e d a c r o s s t h e c u r r e n t , t h e b o a t o r b a r g e w i l l be c a r r i e d s i d e w a y s i n t h e c u r r e n t a n d p r o b a b l y t e a r h o l d f a s t s l o o s e f r o m t h e b o t t o m . I t i s p o s s i b l e t h a t some o f t h e p l a n t s t o r n l o o s e may a t t a c h a g a i n , b u t more l i k e l y t h a t t h e p n e u m a t o c y s t s w i l l s u p p o r t t h e p l a n t i n t h e w a t e r a n d e v e n t u a l l y i t w i l l be c a r r i e d a s h o r e by t h e w a v e s a n d w a s h e d u p o n t h e b e a c h t o g e t h e r w i t h o t h e r p l a n t s o f M a c r o c y s t i s w h i c h i t w i l l c a r r y o u t w i t h i t i n a t a n g l e d mass a s i t p r o c e e d s t h r o u g h t h e b e d . P r o v i d e d t h e d e s c r i p t i o n s o f t h e v a r i o u s p a r t s o f t h e p l a n t , a s a l r e a d y o u t l i n e d , a r e b o r n e i n m i n d a n d r e c o g n i z e d , one c a n e a s i l y make a n e x a m i n a t i o n o f a n y h a r v e s t a f t e r i t h a s b e e n b r o u g h t i n a n d f i n d s u f f i c i e n t p r o o f t o d e t e r m i n e t h e a p p r d x i m a t e h e i g h t a t w h i c h t h e p l a n t s - 51 -have been cut with respect to their attachment on the bottom. I do not believe that anything w i l l be gained by attempting to cut the same areas more than twice a year. It should be pointed out, however, that these conclusions are based on observations made on beds during the period July 1 to September 15, and should be supplemented by observations made at different periods or the results of actual harvests. Although observations have not been made on large beds during the winter, examination of individual plants of this same species in the v i c i n i t y of Sooke, Bri t i s h Columbia, on the west coast of Vancouver Island, on the 29th of December, has substantiated this belief. While harvesting more often than twice a year may y i e l d a tonnage distributed over the whole year, i t i s my belief that the total tonnage realized from such a practice w i l l be less than i f the pre-cautions mentioned i n this section are, even i n part, given consideration. The nature of the bottom i n many locations w i l l necessitate the progress of a rowboat i n advance of any cutting mechanism unless some previous investigations have been made. Often the plant stipes lying along the surface of the water are only a few inches above alarge concealed rocks. As previously pointed out, tearing loose the older part of a plant w i l l often leave the growing "points" s t i l l intact. Many of the large, mature stipes reaching to the surface, however, may s t i l l be attached comparatively close to the growing "points," so that i f torn loose, instead of being cut, they w i l l often break off a portion of the rhizome carrying the growing "point" with i t , with the obvious result that further production of stipes from that point on the rhizome i s impossible. - 52 -Harvesting w i l l invariably have to be done i n comparatively calm water. Even i n the lee of Deer Island (.see rlates 24 and 25), which i s protected from the direct south-east gales, the ground-swell which circles around the corner of the island i s so great that harvesting would be impossible without great damage to the plants. On the sample plot at the north-west side of Deer island previously mentioned, several stipes cut off were noticed to aie DECK after a period o l titty-eight aays, but only part way towaras tne noiaiast when sporophylls or other laminae were s t i l l present. Exam-ination of these sporophylls showed that reproduction took place even after the stipe had been cut off above. This harvest was made to a depth of approximately 2 to 3 feet above the holdfasts. A total of 509 pounds was obtained from the plot on July 18 (1.27 pounds per square foot). Harvesting the same plot on September 14 (fifty-eight days later) gave 107 pounds (0.27 pounds per square foot) when cut at approximately the same depth. As previously indicated, i t was found that growth of the stipes continued after being severed from their holdfast, provided they were kept suitably maintained i n the water. This would seem to indicate that the plant may be stored for some time i n the sea even after cutting, and s t i l l continue growing with l i t t l e evidence of decay at (the severed end. It should be emphasized, however, that this experiment was conducted i n more or less open stretches where adequate circulation of water and exposure to light were provided. Also, no details of the algin or mineral content of the plants so grown were obtained. If plants are to be stored i n bulk i n the sea in any sort of trap, one could hardly expect equally favourable growth and lack of - 53 -decay, primarily because the plants i n the lower part of the trap would be excluded from the necessary intensity of l i g h t and the poorer circulation of water would tend to interfere with the activites of the plant. Particularly i f algin i s to be obtained from plants stored in this manner for any length of time, i t would be advantageous to determine what effect this period of storage has on i t s analysis. VIII. SUMMARY; In many respects this paper must be regarded as preliminary. The following briefly summarizes the essential points brought out during the investigation: The species encountered in the coastal waters of British Columbia i s considered to be Macrocystis integ r i f o l i a Bory. It i s distributed along the whole coast in regions approaching open ocean conditions where suitable depth (20 feet or less at zero tide level) and rocky bottom i s available, but s t i l l protected from f u l l force of the open ocean by reefs or other protection. It i s not found in the inside waters, nor i n water of low salinity (not below density of 1.019 at 15 degrees centigrade). The method of growth of Macrocystis integ r i f o l i a from the single-blade stage of the sporophyte to the f u l l y developed plant i s described and diagrammatically explained. The single-blade stage of Macrocystis integrifolia was found at a l l times from July 1 through to September 15, and the limits were not determined. A dichotomous (and sub-dichotomous) division of the basal growth region i s responsible for the extensive development and perennial habit of the plant. The growth region i s subterminal (a basal meristem) - 5 4 -and s p l i t t i n g i s mainly the result of increased activity of the meristo-derm. Active zoospore liberation was also found during the period from July 1 through to September 1 5 , and the limits were not determined. The gametophytic phases have not been followed during the investigation. The rate of growth in Macrocystis i n t e g r i f o l i a was found to vary greatly, depending on age or size at any given period, although other factors must also be taken into consideration. The maximum growth (elongation) of Macrocystis i n t e g r i f o l i a recorded during the investi-gation was an average of 3 . 1 0 inches per day for a period of twenty-nine days. Only the growth rate of the macroscopic sporophyte was followed. No regeneration was found to occur from the cut surfaces of stipes, nor did laminae tissue regenerate. New stipes are only produced by continued branching of the basal portion of the plant (from the rhizome portion of the holdfast), and new laminae on stipes are only produced when the terminal lamina i s intact. No direct evidence of stimulation to produce more stipes was found subsequent to cutting. Plants moved -to deeper water show retardation i n growth. Various practical aspects have been considered, since the species i s one of economic importance as an alginophyte: A method of estimating quantities was arrived at, using the diameter of the cut surface of the stipe of Macrocystis i n t e g r i f o l i a . Some allowances are made in this method, but i t i s believed adequate for practical purposes. The effect of harvesting Macrocystis i n t e g r i f o l i a was studied after cutting experimental plots as well as individual plants. I t i s - 55 -recommended that plants of Macrocystis i n t e g r i f o l i a be cut not lower than 3 to A feet above the holdfast i n order to leave sporophylls to continue asexual reproduction. I t i s also recommended that the f i r s t cutting of beds be delayed u n t i l well on in the year and that due regard be taken to protect the terminal laminae of "younger" stipes at the f i r s t harvest. It i s believed that by so doing, two harvests may be taken from the same bed i n the same year and y i e l d a total tonnage greater than by starting e a r l i e r i n the year and disregarding these precautions. The greatest destruction to beds of Macrocystis i n t e g r i f o l i a results from severe south-east storms, which either tear the plant out completely and wash i t ashore or damage the major stipes to such an extent that no further laminae are produced on the already differentiated stipes. Some minute algae are epiphytic, at least, on Macrocystis. but these do not seem to cause any great damage. Marine animals, however, including various Crustacea, molluscs (particularly the snail Tegula and, to a lesser extent, the nudibranch Melibe), Ubelia. and Membranipora membranacea. cause considerable damage to the plants either by way of eating portions of the laminae and stipes directly or fastening on to the plants i n such abundance that they weigh the plants down and restrict the l i g h t f a l l i n g on the photosynthetic surfaces. Storage of severed stipes of Macrocystis i n t e g r i f o l i a may be carried out in the sea, and the stipes w i l l continue to grow i f certain precautions are taken. The effect of such storage on the mineral and algin content i s not known. • ACifflOWLEDGMENTS: The writer wishes to acknowledge i n particular the genuine interest which has been shown in the subject of this thesis and the encouragement given by Dr. A. H. Hutchinson, Head of the Department of Biology and Botany, University of British Columbia, under whose guidance the work has been conducted. The assistance given during this investi-gation, as well as the personal attention, have been greatly appreciated. I wish also to express my sincere appreciation and grateful acknowledgment to the following persons, who have given assistance in various ways during the investigations or during the preparation of the manuscript: Mr. George J . Alexander, Deputy Minister, and the staff of the British Columbia Provincial Fisheries Department; Dr. V. A. Clemens, Head of the. Department of Zoology, University of British Columbia; Professor John Davidson, Department of Biology and Botany, University of British Columbia; Mr. Ben Farrar, B r i t i s h Columbia Research Council, University of British Columbia; Dr. R. E. Foerster, Director, Pacific Biological Station, Nanaimo, B. C ; Dr. S. E. Maddigan, Director, British Columbia Research Council, University of British Columbia; Dr. G. F. Papenfuss, Curator of Algal Collections, Department of Botany, University of California. Grateful acknowledgment i s also made to the Canada Kelp Company, Limited, for making accommodation available and for the co-operation of the members of the company in permitting the use of various f a c i l i t i e s at Deer Island. - 57 BIBLIOGRAPHY (1) Agardh, C. A. — 1823. Species algarum r i t e cognitae, cum synonymis, differentiis specificis et descriptionibus „ succinctis. GrypMswaldiae (Sumtibus Ernesti Mauritii). (2) Agardh, C. A. - 1829. Revision de"r Algengattung Macrocystis (Acta Ac. Leop.-Carol. Vol. 21). (Wot seen—cited by Kjellman, 1397). (3) Agardh, C. A. - 1848. Species, Genera, et Ordines Algarum, I. (not seen—cited by Setchell, 1932). (4) Alsberg, C. L. - 1912. A discussion of the probable food value of marine algae. F e r t i l i z e r Resources of the United States. U. S. 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The economic value of the coast kelps. Univ. Calif. Pub., Co l l . of Agr., Agr. Expt. Sta. Bull. No. 248. (12) B. C. Research Council « 1947. Technical Bull. No. /+. A Report on the Location of Marine Plants of Economic Importance in Br i t i s h Columbia Coastal waters. - 58 -BIBLIOGRAPHY (CONTINUED) (13) - B. C. Research Council - 194-8. Marine Plants of Economic Importance in B r i t i s h Columbia Coastal Waters (to be published in 1948). (14) Cameron, A . T. — 1916. The Commercial value of the kelp beds of the Canadian Pacific Coast—rA preliminary report and survey of the beds. Contributions to Can. Biology, Supplement to the 5th .Annual Report of the Department of Naval Service, Fisheries Branch. (15) Cameron, Frank K. — 1912. (with assistance of others) A preliminary report on the f e r t i l i z e r resources of the United States. Senate Doc. No. 190, 62nd Congress, 2nd Session. (16) Cameron, Frank K. - 1915. Pacific Kelp Beds as a Source of Potassium Salts. Potash from Kelp. 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(Not s e e n — c i t e d by F r i t s c h , 1945) . (42) J a n c z e w s k i , E d . de - 1875. Observat ions s u r l ' a c c r o i s s e m e n t du t h a l l e des Pheosporees . Mem. S o c . N a t . S c i . N a t . Cherbourg , 19s 112-113. (Not s e e n — c i t e d by S e t c h e l l , 1932) . (43) Johns ton , E . C . - 1912. B r i e f notes on K e l p s o f A l a s k a . F e r t i l i z e r Resources o f the U n i t e d S t a t e s . U . S . Senate Doc. No . 190 . Appendix 0, ¥ a s h . (44) K j e l l m a n , F . R . - 1897• Laminar iaceae i n D i e N & t u r l i c h e n P l a n z e n -f a m i l i e n I . T e i l . 2 A b t e i l u n g , E n g l e r , A . und K . P r a n t l , L e i p z i g . (45) L e v y n s , M . R. - 1924. The d i s t r i b u t i o n o f seaweeds of the Cape P e n i n s u l a . South A f r . J o u r n . S c i . 21 (Rep. South A f r . A s s o c . S c i . , Johannesburg, 2 2 ) , 2 6 5 - 9 . (Not s e e n — c i t e d by F r i t s c h , 1945) . (46) L e v y n s , M . R. - 1933. Sexua l r e p r o d u c t i o n i n M a c r o c y s t i s p y r i f e r a A g . A n n a l s Botany 47 :349-353 . (47) L i n n a e u s , C a r o l u s - 1771. M a n t i s s a p l a n t a r u m a l t e r a e t spec ierum e d i t i o n i s I I . (Not s e e n — c i t e d by S e t c h e l l , 1945) . BIBLIOGRAPHY (CONTINUED) (4-8) MacMillan, Conway - 1902. The kelps of Juan de Fuca. Postelsia, 1901j p. 218. (49) McFarland, Frank M. - 1912. Kelps of the Central Californian Coast. F e r t i l i z e r Resources of the United States. U. S. Senate Doc. No. 190, Appendix M. Washington. (50) McKay, H. H. - 1933. The Life History of Pterygophora Californica Ruprecht. Univ. C a l i f . Pub. i n Bot., Vol. 17, • No. 6, pp. 111-148. (51) Mertens, H. - 1329. Ueber verschiedene Fucus Arten, Linnaea, Vol. 4. (Not seen—cited by Setchell and Gardner, 1903). (52) Montagene, C. - 1839. Sertum Patagonicum (in Aleide D'Orbiguy, Voy. dans l'Amer. Merid.). Not seen—cited by Setchell, 1932). (53) Moore, L. B. - 1943. Observations on the Growth of Macrocystis i n New Zealand. Trans. Roy. Soc. New Zealand, 72, 333-40. (Not seen—cited by Fritsch, 1 9 4 5 ) . (54) Murray, George - 1895. An Introduction to the Study of Seaweeds. MacMillan and Co. (55) Myers, M. E. - 1928. The Life History of the Brown Alga, Egregia Menziesii. Univ. Ca l i f . Pub. i n Bot., Vol. 14, No. 6, pp. 225-t246. (56) Myers, M. E. - 1925. Contributions toward a knowledge of the l i f e ^ histories of the Melanophyceae. Univ. of C a l i f . Pub. in Bot., Vol. 13, No. 4, pp. 109-124. (57) Okamura, K. - 1932. The Distribution of Marine algae i n Pacific waters. Records of Oceanographic Works in Japan. Vol. IV. No. 1. pp. 30-150. (58) Papenfuss, G. F. - 1942. Studies of S. African Phaeophyceae. I. Ecklonia maxima, Laminaria pallida, Macro-cystis pyrifera. Am. Jour. Bot., 29sl5-&4. (59) Phyc Bor.-Amer. - (Not seen—cited by Smith, 1944). (60) Postels, A., et F. Ruprecht. - 1840. Illustrationes Algarum. (Not seen—cited by Setchell, § 9 3 2 ) . (61) Reinke, J. - 1903. Studien zur ve gleichenden Entwickelungsgeschichte der Laminariaceae (Kiel). (Not seen—quoted by Setchell, 1932). - 62 -BIBLIOGRAPHY (CONTINUED) ( 6 2 ) Reinsch, P. F. - 1890. Zur Meeresalgenflora von Slid Georgia. Die internat. Polarforschung. Die deut. Exp., 2:417, 418, p i . 16, f. 7a, 7b. (Not seen— cited by Setchell, 1932). (63) Rigg, 0. B. - 1912. Ecological and economic notes cn Puget Sound kelps. F e r t i l i z e r Resources of the United States. U. S. Senate Doe. No. 190, Appendix L., Washington. (64) Rigg, G. B. - 1915. The Kelp Beds of Puget Sound. Potash from Kelp. U. S. Dept. Agric. Report No. 100, Part III. (65) Rigg, G. B. - 1916. The Kelp Beds of Western Alaska. Potash from Kelp. U. S. Dept. Agric. Report No. 100, Part V. (66) Rosenthal, 0. - 1890. Zur Kennthis von Macrocystis und Thalassio-phyllum. Flora, 73:105-147, p i . 7, 8. (Not seen—cited by Setchell, 1932). (67) Ruprecht, F. J. - 1851. Tange des Ochotskischen Meeres, in Midden-dorff's sibirische Reise, 1:2, Botanik. (Not seen—cited by Setchell, 1932). ( 6 8 ) Saunders, DeAlton - 1901. The Algae. Papers from the Harriman Alaska Expedition, XXV, Proc. Wash. Acad. Sci. 3. p.431, p i . 60. (69) Scagel, R. F. — 1946. A report to the Fisheries Research Board of Canada of a seaweed survey of the coast of British Columbia, with 19 charts, 8 plates and 8 tables (MS unpublished). (70) Scagel, R. F. - 1948. An investigation on Marine plants near Hardy Bay, B. C. A report to the Provincial Fisheries Dept., Victoria, B. C. No. 1. King's Printer, Victoria. (71) Setchell, W. A. - 1893. On the classification and geographical d i s t r i -bution of the Laminariaceae. Trans. Connecti-cut Acad., 9:333-375. (Not seen—cited by Setchell, 1932). (72) Setchell, W. A. - I896. Notes on Kelps. Erythea, 4:44. (Not seen— cited by Setchell, 1932). (73) Setchell, W. A. - 1912. The kelps of United States and Alaska. F e r t i l i z e r Resources of the United States. 62nd Congr., 2nd Session, Senate Doc. 190, pp. 130-178. BIBLIOGRAPHY ( CONTINUED ) (74-) Setchell, ¥. A.. - 1932. Macrocystis and i t s holdfasts. Univ. C a l i f . Publ. Bot. 16:445-492. (75) Setchell, ¥. A. and N. L. Gardner - 1903. Algae of northwestern America. Univ. C a l i f . Publ. Bot. 1:165-413. (76) Setchell, ¥. A. and N. L. Gardner - 1925. The Marine Algae of the Pacific Coast of North -America, Part III, Melano-phyceae, Univ. C a l i f . Publ. Bot., Vol. 8, Part III, pp. 383-398, plates 34-107. (77) Setchell, ¥. A., and H. E. Parks - 1930 - In Herbarium Univ. C a l i f . #445012, 445007, 445013 and 445014. (78) Skottsberg, C. - 1903. Nagro ord om Macrocystis pyrifera (Turn.) Ag. Bot. Notiser, 40-4. (Not seen—cited by Fritsch, 1945). (79) Skottsberg, C. — 1907. Zur Kenntnis der subantarktischen und antark-tischen Meeresalgen. I, Phaeophyceen. Wiss. Ergebn. d. Schwed. Sudpolarexped. 1901-1903. 4, No. 6:1-172. Stockholm. (80) Smith, A. L. and Whitting, F. G. - 1895. Notes on the Sori of Macro-cystis and Postelsia. Phyc. Mem. Part 3:84-37; p i . 20. (Not seen—cited by Tilden, 1937). (81) Smith, G. M. - 1938. Cryptogamic Botany, Vol. I., Algae and Fungi, McGraw-Hill. (82) Smith, G. M. — 1942. Notes on Some Brown .Algae from the Monterey Peninsula, C a l i f . Jkm.. Jour. Bot. 29:645-653; 13 fig s . (83) Smith, G. M. - 1944. Marine Algae of the Monterey Peninsula, C a l i f . Standford Univ. Press. (34) Sverdrup, H. N., M. ¥. Johnson, and R. H. Fleming - 1946* "The Oceans." Prentice-Hall, New York. (85) Sykes, M. G. - 1908. Anatomy and histology of Macrocystis pyrifera and Laminaria saccharina. Ann. Bot. .22, 291-325. BIBLIOGRAPHY (CONCLUDED) (86) Tilden, Josephine E. - 1894-19Q2. American Algae, Centuries 1-6. (#518). (Not seen—cited by Setchell and Gardner, 1903) . (87) Tilden, Josephine E. - 1937. The Algae -and Their Life Relations, Univ. of Minnesota Press. (88) Tressler, Donald K. - 1923. Marine Products of Commerce. The Chemical Catalog Company, Incorp. (89) Turner, Dawson - 1809. Fuci, 2 : 1 0 3 , 104, p i . 110. (Not seen—cited by Setchell and Gardner, 1903) . (90) Turrentine, J . W. - 1912. The Composition of Kelps. F e r t i l i z e r Resources of the United States, U. S. Senate Doc. No. 190, Appendix P., Wash. (91) Turrentine, J . W. - 1912. The Technology of the seaweed industry. F e r t i l i z e r Resources of the United States, U. S. Senate Doc. No. 190, Appendix Q, Washington. (92) W i l l , Dr. H. - 1884. Zur .Anatomie von Macrocystis luxurians Hook. f i l . et Harv. Bot. Zeit. 42 :801-808, 825-830, p i . 1 2 . (Not seen—cited by Setchell, 1932) . - i x -APPENDICES Appendix A - Charts Appendix B - Tables Appendix C - Figures Appendix D - Plates APPENDIX A C H A R T S 1 CHRRT NOVUI APPENDIX U T A B L E S TABLE I Page 1 o f 5. . DISTRIBUTION AND ABUNDANCE OF MACROCYSTIS INTEGRIFOLIA BORY. LOCATION AND CHART REFERENCE SIZE OF BED AREA DENSITY TONNAGE (Pounds per - (Feet) (Square Feet) Square Foot) CHART I I . Dundas I s l a n d W h i t l y P t . ( N . E . end) t r a c e Brundige I n l e t t r a c e Goose Bay .20 x 200 4 ,000 1 .0 2 .0 A r n i s t o n P o i n t t r a c e ¥ e s t shore f r i n g e 128,000 1 .0 64 .0 200 x 500 100,000 1 ,0 50.0 100 x 300 30,000 264,000 0 . 5 7 .5 Chearnley I s l a n d ( N . E . ) 200 x 1320 1 .0 132.0 P r i n c e Leboo I s l a n d ( E . ) 200 x 200 40,000 256,000 1 .0 2 0 . 0 ( S . ¥ . ) 50 x 5120 0 . 5 64 .0 Baron I s l a n d (N. ) 100 x 600 60,000 1 .0 30 .0 M e l v i l l e I s l a n d S . ¥ . . a n d ¥ , shores 400 x 2000 800,000 0 . 5 200 .0 200 x 1000 200,000 0 . 5 50.0 300 x 500 150,000 1 .0 75.0 400 x 2700 1 ,080,000 ' 0 . 3 162.0 E . and S . E . shores 2 5 x 1320 33,000 264,000 1 .0 16 .5 200 x 1320 1 .0 132.0 D u n i r a I s l a n d ( ¥ . shore) f r i n g e 200 ,000 1 .0 100.0 CHART I I I . Stephens I s l a n d E . s i d e t r a c e E . end Canoe Pass 100 x 500 50,000 1.0 2 5 . 0 N.¥ . shore 50 x 80 4 ,000 1 .0 2 . 0 TABLE I I cont inued) Page 2 o f 5. DISTRIBUTION AND ABUNDANCE OF MACROCYSTIS INTEGRIFOLIA BORY. LOCATION AND CHART REFERENCE SIZE OF BED -AREA DENSITY TONNAGE (Feet) (Square Feet ) (Pounds p e r Square Foot) Stephens I s l a n d (concluded) '3.0 N.¥. shore f r i n g e s 6,000 1.0 S k i a k l Bay f r i n g e s 160,000 1.0 80.0 B u t l e r Cove t r a c e P r e s c o t t Channel f r i n g e s 400,000 0.75 150.0 Porcher I s l a n d Welcome Harbour ( i n c l u d i n g S . E . beds and Henry I.) f r i n g e s 7,000,000 1.0 3,500.0 N . and ¥ . Henry I. f r i n g e 20,000 1.0 10.0 S u r f P t . t o Pearce P t . t r a c e Refuge Bay 200 x 400 80,000 1.0 40.0 50 x 600 30,000 1.0 15.0 200 x 100 20,000 1.0 10.0 100 x 300 30,000 1.0 15.0 Creak Bay f r i n g e 20,000 0.5 5.0 Jap I n l e t f r i n g e 100,000 1.0 50.0 CHART I V . Banks I s l a n d N . end 500 x 15,840 7,920,000 2.0 7,920.0 50 x 200 10,000 1.0 5.0 f r i n g e 300,000 1.0 150.0 N . ¥ . shore and Norway I n l e t f r i n g e s 1,440,000 0.75 540.0 CHART V . A r i s t a z a b a l I s l a n d U l r i c P t . t o Baker P t . 50 x 6,600 330,000 1.0 165.0 S . o f Baker P t . 25 x 100 2,500 1.0 1.25 25 x 50 1,250 1.0 0.63 TABLE I (cont inued) Page 3 o f 5 . DISTRIBUTION AND ABUNDANCE OF MACROCYSTIS INTEGRIFOLIA BORY. LOCATION AND CHART REFERENCE SIZE OF BED AREA DENSITY '• TONNAGE (Pounds per (Feet) (Square Feet ) Square Foot) A r i s t a z a b a l I s l a n d (concluded) 2 M i l e s S . o f Baker P t . t r a c e CHART V I I . S h e l t e r Bay- 100 x 200 20 ,000 1 .0 1 0 . 0 100 x 200 20 ,000 1 .0 10 .0 25 x 400 10,000 1 .0 5 .0 50 x 300 15,000 1 .0 7 .5 20 x 50 1,000 1 .0 0 . 5 f r i n g e s 2 , 0 0 0 1 .0 1 .0 Wal lace I s l a n d t r a c e 200 CHART V I I I . Hope I s l a n d E . end 4,000 x 350 1,400,000 1 .0 700 .0 Heath P t . t o Turn P t . f r i n g e s 80,000 1 .0 40 .0 N i g e i I s l a n d Opposi te Lemon M t . t r a c e 100 CHART I X . Gole tas Channel (S . s i d e ) f r i n g e s 40,000 1 .0 2 0 . 0 Hardy Bay f r i n g e s 10,000 . 1 .0 5 .0 CHART X . P e e l I s l a n d f r i n g e s 2 ,000 1 .0 1 .0 Deer I s l a n d ) N . s i d e ) see a l s o CHART X I 200 x 2 , 0 0 0 400,000 0 . 7 140 .0 W. s i d e ) f r i n g e s 40,000 1 .0 2 0 . 0 Browing I s l a n d t r a c e TABLE I ( cont inued) Page A o f 5» DISTRIBUTION M D ABUNDANCE OF M&CROCYSTIS INTEGRIFOLIA BORY. LOCATION AND CHART REFERENCE SIZE OF BED AREA DENSITY TONNAGE (Pounds per (Feet) (Square Feet) Square Foot) Vancouver I s l a n d Thomas P t . t o F a l s e H d . 7,900 x 80 632,000 1.0 316.0 300 x 5,280 1,584,000 1.0 792.0 400 x 1,000 400,000 1.0 200.0 100 x 500 50,000 1.0 25.0 10,560 x 300 3,168,000 1.0 1,584.0 F a l s e H d . 200 x 1,000 200,000 1.0 100.0 F a l s e H d . t o Suquash 200 x 5,280 1,056,000 1.0 528.0 100 x 5,280 528,000 1.0 264.0 5 x 500 2,500 1.0 1.3 S i n g l e Tree P o i n t 25 x 100 2,500 1.0 1.2 800 x 200 160,000 1.0 80.0 50 x 200 10,000 1.0 5.0 N . K l u c k s i w i R i v e r 25 x 400 10,000 1.0 5.0 CHART X I I . Vancouver I s l a n d K l u c k s i m R . t o Ledge P t . 20 x 200 4,000 1.0 2.0 10 x 100 1,000 1.0 0.5 20 x 200 4,000 1.0 2.0 M a l c o l m I s l a n d T r i n i t y Bay 600nx 5,280 3,3&8,000 528,000 0.75 1,188.0 264.0 Boulder P o i n t 200 x 2,640 1.0 660 x 2,640 1,742,400 1.0 871.2 50 x 2,640 132,000 1.0 66.0 B l a c k B l u f f 200 x 2,640 528,000 1.0 264.0 ¥ . o f B l a c k B l u f f 250 x 3,960 990,000 1.0 495.0 200 x 400 80,000 1.0 40.0 TABLE I (concluded) Page 5 o f 5. DISTRIBUTION AND ABUNDANCE OF MACROCYSTIS INTEGRIFOLIA BORY. LOCATION AND CHART REFERENCE SIZE OF BED AREA DENSITY TONNAGE -(Feet) (Square Feet) (Pounds per Square Foot) M a l c o l m I s l a n d (concluded) Bere P o i n t 200 x 200 40,000 1 .0 2 0 . 0 200 x 1,000 200,000 1 .0 100.0 Bere P t . to S t a r t P t . 200 x 1,000 200,000 1 .0 100.0 300 x 1,000 300,000 1 .0 150.0 200 x 1,500 300,000 1 .0 150.0 N . ¥ . end o f I s l a n d 75 x 600 45,000 1 .0 2 2 . 5 f r i n g e 280 ,000 1 .0 140.0 TABliE II RATE OF GROWTH MP DIVISION OF VERY SMALL PLANTS OF MACROCYSTIS INTEGRIFOLIA BORY LOCATED ON A LARGE ROCK. N.W. OF DEER I. ALL PLANTS BETWEEN ONE AND TWO FEET ABOVE ZERO TIDE LEVEL. S, split only evident at base and not completely separated; C_, crease only indication of division; x, damaged. CODE NO. 50 51 52 53 54 55 56 125 AUGUST A STAGE OF DIVISION COMPLETE I I I I I I I I I I I I I I INDICATED I I I I I I I I I IV I I I I I I I STIPE LENGTHS 32" 20" 4*» 5t" 6" x l a 32" 12" x 6" 6" S c s x 2a 6" S X S x S C C S 3a AUGUST 17 6 ° SEPTEMBER 13 STAGE OF DIVISION COMPLETE I I I I I I I I I I I I I I I INDICATED IV IV rv IV I I TV i n I I STIPE LENGTHS 45x 2 5 " 5 i * 16" 8" x 13" 6* " la 69" 21" 5¥ x . 8" 9 R 13" 6 i " 8" 8" S 6" S x S 2a x 3x S x S s s s s c s c 3a S S c x NO. OF DAYS 40 40 40 40 40 40 40 28 CODE NO. STIPE LENGTH TO HOLDFAST AUGUST 4 SEPTEMBER 13 TOTAL INCREASE IN LENGTH DAILY AVERAGE INCREASE IN LENGTH 50 51 52 53 54 55 56 125 32" 32" 6" 12" 2 0 " 4 i n 5 i n 6" 5f 4l n AUGUST 17 6" 4 5 £ 69" 14* 21" 25fi 5a" 16" 8" 9 n 13° 13" 6*« 13" 37" 8" 9" 5" 1" 10§» 2 " 34" 8|" 8 i " 0 .33" 0 . 9 3 " 0 .20" 0 .23" 0 .13" 0 .03" 0.26" 0 .05" 0 .88" 0 . 2 1 " 0 . 2 1 " 0.02" Mil RATE OF GROWTH OF STIPES OF SMALL COMPLETE PLANTS OF MACROCYSTIS INTEGRIFOLIA BORY ATTACHED TO HOLDFASTS AT -10 FT, FOR A PERIOD OF 29 DAYS. THEN MOVED TO -14 FT. FOR PERIOD OF 17 DAYS. CODE NO. FIRST CHECK SUBSEQUENT CHECKS TOTAL IN-CREASE IN LENGTH NO. OF DAYS DAILY AVERAGE INCREASE DEPTH DATE STIPE LENGTH TOTAL T.flMTfffiF. DATE' STIPE LENGTH TOTAL LAMINAE 30 3 0 (Total) July 2 5 5 ' 4 " 16 Aug. 23 Sept. 9 1 2 ' 1 0 « 15*6" 2 5 3 6 7*6" 2»3 n 2 9 17 3 . 1 0 " 1 . 8 8 " - 1 0 ' (moved to - 1 4 ' ) 1 0'2" 4 6 3 1 3 1 (Total) July 2 5 6'9" 1 9 Aug. 23 Sept. 9 13170 16'6« 3 5 41 6'10" 2'9" 2 9 17 2 . 8 3 " 1 . 9 4 " - 1 0 « (moved to - 1 4 ' ) 91711 4 6 3 2 32 ( T o t a l ) July 2 5 6'8" 2 1 Aug. 23 Sept. 9 12'0° 1 2 ' 1 0 " 24 2 4 5 » 4 a O'lO" 2 9 17 2 . 2 1 " 0 . 5 8 " - 1 0 ' (moved to - 1 4 ' ) 6'2" 4 6 33 33 (Total) July 2 5 5'7 a 1 5 Aug. 23 Sept. 9 1 0 ' 3 n 1 1 ' 1 0 " 2 4 30 4 ' 8 " 1'7" 29 17 1 . 9 3 " 1 . 1 2 " - 1 0 ' . (moved to - 1 4 ' - ) 6'3" 4 6 3 4 July 2 5 2* 1 0 " 11 Aug. 23 4'6* 1 2 1 ' 8 " 2 9 0 . 6 9 " - 1 0 ' 38 July 2 5 l'lO" 1 3 Aug.. 23 3» 1 " 1 5 1'3" 2 9 0 . 5 2 " - 1 0 ' TABLE IV RATE OF GROWTH (ELONGATION) OF STIPES OF MACROCYSTIS INTEGRIFOLIA BORY SEVERED FROM HOLDFASTS AND ANCHORED. NOS. 108-113 LOCATED AT N.E. DEER I.; NOS. 72-73 AT MOUTH OF KEOUGH RIVER; AND BALANCE AT N.W. DEER ISLAND. FIRST CHECK SUBSEQUENT CHECKS CODE DATE STIPE TOTAL DATE STIPE TOTAL TOTAL IN- NO. OF DAILY DEPTH •NO. LENGTH LAMINAE LENGTH LAMINAE CREASE IN DAYS AVERAGE LENGTH INCREASE 1 July 14 14 '10" 48 Aug. 3 1 7 ' 8 " 50 2 ' 1 0 " 25 1 .36" - 1 2 ' 3 July 14 1 3 ' 0 » 62 Aug. 3 16 '10" 66 3 ' 1 0 " 25 1 .84" -12 * 3 . Sept. 4 1 9 ' 3 " 67 2 ' 5 " 27 1 .07" (moved to - 1 8 ' ) (Total) 6 » 3 " 52 42 July 2 5 2 2 ' 1 0 " 71 Aug. 25 2 8 ' 3 " 77 51 5« 31 .2.10" - 1 2 ' 42 Sept. 5 .30 ' 0 " 80 1 ' 9 " 11 1.91" - 1 2 ' (Total) 7 ' 2 " 42 45 July 25 1 5 ' 1 " 58 Aug. 25 20»5" 62 5 ' 4 H 31 2 .06" - 1 2 ' 48 July 25 1 3 1 3 " 54 Aug. 25 17*8" 55 4 ' 5 " 31 1 . 7 1 " - 1 2 ' 20 July 2 5 12 »0° 47 Aug. 2 5 16 '10" 57 4 ' 1 0 u ' 31 1 .87" -12» 20 Sept. 5 19'4" 61 2 ' 6 " 11 2 . 7 3 " - 1 2 ' (Total) 7 ' 4 " 42 15 July 25 27 '1" 78 Aug. 25 3 1 ' 4 " 79 4 ' 3 a 31 1 .65" - 1 2 ' 91 Aug. 7 12 '10" 42 Sept. 3 1 5 ' 2 " 47 2 ' 4 " 27 1 .04" - 1 5 ' 92 Aug. 7 7 ' 2 " 35 Sept. 3 9 ' 1 0 " 37 2«8» 27 1 .19" - 1 5 ' 93 Aug. 7 6 ' 8 ° 32 Sept. 3 9 ' 3 " 37 2'7" 27 1 .15" - 1 5 ' 94- Aug; 7 8 ' 6 " 39 Sept. 3 1 1 ' 6 " 39 ~ 3 ' 0 « 27 1 .33" - 1 5 ' 95 Aug. 7 6 ' 1 0 " 33 Sept. 3 8 ' 9 n 36 - I ' l l " 27 0 .85" - 1 5 ' 96 Aug. 7 8 « 0 U 34 Sept. 3 9 ' 5 " 36 1 ' 5 " 27 0 . 6 3 " -15" 97 Aug. 7 5 ' 7 " 33 Sept. 3 1 0 ' 1 1 " 34 5«4" 27 2 .37" -15' 108 Aug. 8 8 ' 1 1 " 43 Sept. 9 1 5 ' 1 " 51 6 ' 2 « 32 2 . 3 1 " - 1 2 ' 109 Aug. 3 1 1 ' 5 " 48 Sept. 9 1 6 ' 4 " 60 4 ' 1 1 " 32 1 .84" - 1 2 ' 110 Aug. 8 7 ' 0 " 33 Sept. 9 1 1 ' 7 " 41 4 ' 7 " 32 1 .72" - 1 2 ' 113 Aug. 3 1 3 ' 1 1 " 53 Sept. 9 1 9 ' 1 " 58 5 ' 2 " 32 1 .94" - 1 2 ' 72 Aug. 5 1 1 ' 8 " 47 Sept. 7 16 '10" 56 5 "2" 33 1 .83" -12'. 76 Aug. 5 6 ' 5 " 39 Sept. 7 1 2 ' 7 " 48 6 ' 2 " 33 2 . 2 4 " -12» 77 Aug. 5 1 2 ' 7 " 46 Sept. 7 1 6 ' 8 " 55 4 ' 1 " 33 1 .43" - 1 2 ' 78 Aug. 5 10 '10" 48 Sept. 7 15 '2" 50 4 » 4 " 33 1 .58" - 1 2 ' T A B L E V BATE OF GROWTH (ELONGATION) OF STIPES OF MACROCYSTIS INTEGRIFOLIA BORY FASTENED TO HOLDFAST. NOS. 98-103 LOCATED IN -12 FT. OF WATER AT THE MOUTH OF THE KEOUGH R.: BALANCE LOCATED IN THE SAME DEPTH AT N.W. DEER ISLAND. FIRST CHECK SECOND CHECK CODE NO. DATE LENGTH TOTAL NO. DATE LENGTH TOTAL NO. TOTAL INCREASE DAILY OF STIPE OF LAMINAE OF STIPE OF LAMINAE IN LENGTH AVERAGE INCREASE 98 Aug. 5 8'11" 27 Sept. 7 12»0" 37 3»1" 1 . 16" 101 Aug. 5 2*11* ? Sept. 7 V 8" 23 4'9n 1.78" 102 Aug. 5 14'6" 47 Sept. 7 18»10« 57 4'4" 1.63" 103 Aug. 5 0 ' 4 | « 5 Sept. 7 1'2" 9 0'9i" 0.30" 24 July 16 8'2" > Sept. 5 15'8" 31 7'6" 1.80" 26 July 16 2»6" 21 Sept. 5 10'6" 29 8'0" 1.92" 27 July 16 l'O" 15 Sept. 5 5*6" 18 4I6« 1.08" 13 July 16 2'0" 1 5 Sept. 5 4'5" > 2'5" 0.58" TARl'iT*! VI SIZE OF STIPES OF MACROCYSTIS INTEGRIFOLIA BORY AND DISTRIBUTION OF WEIGHT PAC je 1 of 2. CODE LENGTH OF NO. OF TOTAL LENGTH TOTAL WT. LENGTH OF WT. OF MAXIMUM APPROX. NO. STIPE TO LAMINAE OF STIPE OF'STIPE FULLY LAM- FULLY LAM- DIAMETER SIZE HOLDFAST COMPLETE WEIGHED (Pounds) INATED POR-. INATED POR- OF STIPE ATTAINED 2I0N TION (Mm.) % (Pounds) l a 30'0" 47 30 «0" 3.90 16'0" 3.40 8-9 75 2a ? 38 l l ' 4 n 2 .60 11'4° 2.60 11 100 • 3a 34'0" 63 34'0" •6.00 21»3" 5.30 11 100 4a 10'11" 37 10'11" 1.35 7*11" 1.15 8-9 75 5a ? . 28 6'7" 0.60 6'7" 0.60 7 50 6a 17 '0" 41 17'0" 2.60 12'7" 2.40 8 75 7a 9 36 16'2" 3.55 10'4" 2.80 6-7 50 8a 15*10" 34 15'10" 1.30 10'0" 1.15 6-7 - 50 9a 3*0" 11 3,0" 0.10 3'0" 0.10 3-4 25 10a 17'0" 38 17 f0" 1.60 10'6" 1.50 6-7 50 11a 3'7" 11 3'7" 0.10 3'7" 0.10 3-4 " -25 12a 9'11" 28 9'11" 0.95 9'11" 0.95 6-7 50 13a 7'8" 29 7'3" 0.75 7'3" 0.75 6 50 14a 2' 11" 12 . 2'11" 0.10 2'11" 0.10 5 25 15a 7'0" 20 7*0" 0.40 4'7" 0.35 5 25 16a 9 58 14'8" 3.90 14'8" 3.90 11 100 ' 17a 9 51 14«0" 3.20 14'0" 3.20 11 100 18a 34'5" 61 34'5" 6.30 25 '3" 5.30 11 100 19a 35'10" 55 35'10" 6.40 18'10" 5.50 11 100 20a 27'4" 42 27 '4" 3.10 19 '3" 2.70 6-7 50 21a 27'8" 42 271 gir 3.35 17'8" 2.85 8-9 75 22a 21'8" 45 21'8" 2.90 14'3" 2.50 10-11 100 23a 19'10" 38 19'10" 1.95 15'0" 1.85 8-9 75 24a ? 41 12'11" 2.70 12'11" 2.70 8-9 75 25a 9 38 12'3n 3.40 12 ' 3 n 3.50 9-10 100 26a 9 36 13'4" 3.70 13'4n 3.70 11 100 27a 9 38 L2'0» 2.90 12'0" 2.90 10-11 100 28a 9 • 23 17'D" 0.90 S ' i l " 0.70 3-4 25 TABiiE V I (concluded) SIZE OF STIPES OF MACROCYSTIS INTEGRIFOLIA BORY AND DISTRIBUTION OF WEIGHT Page 2 of 2 . CODE LENGTH OF NO. OF TOTAL LENGTH TOTAL WT. LENGTH OF W. OF MAXIMUM APPROX. NO. STIPE TO LAMINAE OF STIPE OF STIPE FULLY LAM- FULLY L A M - DIAMETER SIZE HOLDFAST COMPLETE WEIGHED (Pounds) INATED POR^ INATED POR- OF STIPE ATTAINED TION •*. TION (Mm.) % (Pounds) 29a 19 i i i t 2 9 1 9 ' 1 " 1 .80 13 '5" ' 1 .50 6-7 50 30a 38 2 2 ' 9 " 3 .50 1 4 ' 2 " 3 .05 8-9 75 31a 56 1 8 ' 7 " 4 . 8 0 1 4 ' 2 " 4 .40 12 100 32A 2 6 ' 0 » 44 2 6 ' 0 « 4 . 1 5 1 7 f 0 " 3 .60 11 100 33a 2 0 12»6» 0 .80 6 ' 7 " 0 .70 3-4 25 34a 35*6" 68 3 5 ' 6 " . 6 .40 2 4 ' 7 " 5.80 11 100 35a 1 2 ' 1 1 " 23 1 2 ' 1 1 " 1.20. 8 ' 1 0 " 1.05 8 75 36a 3 2 ' 2 " 61 3 2 ' 2 " 5.30 2 2 » 3 D 4 .70 11 100 37a 36 «0" 68 3 6 ' 0 " $ .60 2 7 ' 0 " 7.00 11 100 38a 3 0 ' 2 " 53 3 0 ' 2 " 4 .60 2 1 ' 1 0 " 4 .10 10 100 PERCENTAGE SIZE ATTAINED 25 50 75 100 TOTAL WEIGHT OF STIPE (LAM-INATED PORTION) Maximum 0.70 2 . 8 0 3 .40 7.00 Average 0.34 1.49 2 . 3 1 4 .28 Minimum 0.10 0.60 1.05 2 . 5 0 T A B L E VII ELONGATION OF STIPES OF MACROCYSTIS INTEGRIFOLIA BORY FROM M I C H THE TERMINAL LAMINAE. INCLUDING THE UPPER .2-FOOT PORTION. HAVE BEEN SEVERED. PLANTS WERE FOLLOWED FOR 31-DAY PERIOD AT DEER L , AUGUST 4 SEPTEM. BER 3 TOTAL INCREASE DAILY CODE NO. LENGTH NUMBER OF LENGTH . NUMBER OF IN LENGTH (AVEBAGE) LAMINAE LAMINAE INCREASE 57 5«6« 13 5 '10" 13 4 " .129" 58 6 « 7 " 18 6*9» 18 2 " .065" 59 5%P 17 5*11" 17 7" .226" 60 8»10« 29 9»0" 29 2 " .065" 62 8 ' 1 « 16 8»2« 16 1" .032" 63 7 » 1 " 17 7*1* 17 0 ° .000" 64 6 ' 8 « 19 6 ' 10" 19 2 " .065" 65 7 ' 2 " 27 7*2* .27 0" .000" 66 6 ' 3 " 2 1 6*5* 2 1 2 " .065" 67 6 ' 5 " 19 7*1" 19 8" .258" 68 5 ' l l n 16 5 ' 1 1 » 16 0" .000" 69 8 ' 2 » 2 1 8 ' 2 ° 21 0" .000" 70 4 ' 8 " 12 4 ' 8 " 12 0" .000" T A B L E VIII DENSITY AND TEMPERATURE RECORDINGS ON SEAWATER SAMPLES DATE TIMEg LOCATION TEMPERATURE DENSITY °F ( a t 1 5 ° C . ) A I R WATER 25/8/46 13:30 P o r t Simpson 69 60 1 .0202 25/8/46 23:30 P e a r l Harbour 59 55 1 .0197 25/8/46 7:00 B i g Bay 54 54 1.0197 23/8/46 2 0 : 0 0 Goose Bay 56 54 1.0219 23/8/46 12:15 S.W. M e l v i l l e I . 58 54 1 .0210 24/8/46 13:15 S . E . M e l v i l l e I . 59 54 1 .0204 26/8/46 14:00 P r e s c o t t Channel 68 56 1 .0215 27/8/46 9:00 Welcome Harbour ( P o r c h e r I . ) 58 54 1 .0224 21/8/46 12 :00 Creak Bay 64 54 1.0226 21/8/46 10:30 Jap I n l e t 62 54 1.0214 19 /8/46 17 :00 Norway I n l e t 58 52 1 .0235 19/8/46 13 :00 Deadman I . 56 52 1 .0233 28/8 /46 18:30 G r e n v i l l e Channel ( a t Lowe I n l e t ) 62 49 1 .0230 18/8/46 14 :45 N . E . A r i s t a z a b a l I . 68 55 1 .0224 29/8 /46 16:30 Meyers Narrows 72 52 1 .0233 17/8/46 16 :00 Promise P t . ( N . C e c e l i a I . ) 68 56 1 .0199 30/8/46 10:30 N . D u f f e r i n I . 62 58 1 .0195 30/8/46 11:00 Dundivan I n l e t 64 60 1 .0187 30/8/46 13:30 Georg ie P t . 59 59 1 .0124 31/8/46 8:45 Welcome Harbour ( C a l v e r t I . ) 58 53 1 .0202 31/8/46 10:20 C a l v e r t I . (oppos i te Addenbrooke) 59 57 1 .0182 31/8/46 12 :00 Open Bay 63 54 1 .0181 31/8/46 13 :00 K e l p Head 62 54 1 .0218 1/9/46 11:00 Southgate Group 54 49 1 .0235 1/9/46 12 :00 S h e l t e r Bay 62 54 1 .0226 16/8/46 12:30 Hope I . (South o f C . James) 58 50 1 .0240 16/8/46 9 :30 N i g e i I. (oppos i te Lemon M t . ) 58 52 1 .0235 15/8/46 20 :30 Hardy Bay 52 54 1 .0228 15 /8/46 10:30 Deer I . 60 53 1 .0223 2/9 /46 9 :30 Bere P t . (Malcolm I . ) 58 49 1 .0235 4/9 /46 8 :00 N.W. Pearse I . 58 56 1 .0241 1/9/46 16:00 T r a c y Bay (entrance) 64 54 1.0214 4/9/46 11:00 F i f e Sound (entrance) 58 54: 1 .0184 4/9 /46 12:30 Tracy I . 64 54 1 .0195 5/9/46 9 :15 P o r t Harvey 55 54 1 .0210 5/9/46 12:00 B l i n k i n s o p Bay 62 52 1 .0229 5/9/46 14:30 J a c k s o n Bay 65 56 1 .0227 7/9/46 8:30 Shark S p i t 61 60 1 .0195 7/9/46 11:00 Rebecca S p i t 64 61 1 .0163 8/9/46 10:15 Goose S p i t (Comox) 62 60 1 .0166 10/9/46 16:30 S i d n e y S p i t (S idney I . ) 64 52 1 .0222 X A 1 1 t imes P a c i f i c Standard Time APPENDIX G F I G U R E S F i g . 1 .—Diagrammatic r e p r e s e n t a t i o n o f t h e l i f e - c y c l e o f M a c r o c y s t i s . Stages 3 t o 13 a r e a l l m i c r o s c o p i c i n s i z e . The mature sporophyte (1) has b a s a l s p o r o p h y l l s (2) on b o t h s u r f a c e s o f w h i c h a r e produced u n i l o c u l a r sporangia (3) w h i c h , when mature , r u p t u r e (4) and r e l e a s e m o t i l e zoospores ( 5 ) . Zoospores e v e n t u a l l y s e t t l e down, l o s e t h e i r f l a g e l l a ( 6 ) , and develop i n t o a female gametophyte (7) o r a male gametophyte ( 8 ) . The a n t h e r i d i u m (9) produced on t h e male gametophyte r e l e a s e s a spermatozoid (10) which f e r t i l i z e s t h e oogonium (11) produced on the female gametophyte, r e s u l t i n g i n the f o r m a t i o n of a zygote (12) . C e l l d i v i s i o n of the zygote (13) b e g i n s t h e development o f the embryo ;vhich u l t i m a t e l y grows i n t o a young sporophyte (14) w i t h a n u n d i v i d e d b l a d e . T h i s s i n g l e b l a d e undergoes a s e r i e s o f d i v i s i o n s (15) which e v e n t u a l l y become d i f f e r e n t i a t e d i n t o t h e v a r i o u s p a r t s o f t h e mature p l a n t ( 1 ) . M a g n i f i c a t i o n s i n d i c a t e d i n diagram a r e approximate , i n d i c a t i n g r e l a t i v e s i z e o f v a r i o u s p a r t s ( r e d u c t i o n a p p r o x i m a t e l y o n e - h a l f o r i g i n a l ) . F i g . 2 .—Diagrammatic r e p r e s e n t a t i o n of o r i g i n o f " s p l i t s " and development o f t h e v a r i o u s laminae produced i n t h e young sporogahyte of M a c r o c y s t i s i n t e g r i f o l i a ' B o r y . i ' o r l a t e r s t a g e s , see F i g . 3. F o r e x p l a n a t i o n o f symbols , c o n s u l t c o n t e x t . (Note .—Drawings E , FV a n d G represent one h a l f o f the p l a n t ; the p o r t i o n ommitted i s a counterpar t of t h a t i l l u s t r a t e d . ) F i g . 3.—Diagrammatic r e p r e s e n t a t i o n of development of a young sporophyte of M a c r o c y s t i s i n t e g r i f o l i a B o r y . i ' o r e a r l i e r s t a g e s , see F i g . 2 . F o r complete e x p l a n a t i o n of symbols," c o n s u l t c o n t e x t ^ ( N o t e . — T h i s diagram r e p r e s e n t s o n l y one h a l f o f the p l a n t ; t h a t i s , e v e r y t h i n g shown on t h e upper s i d e o f the l i n e DI s h o u l d be repeated on the lox^er s i d e . ) t , t e r m i n a l l a m i n a ; 1, s t e r j l l e l a m i n a ; p , pneumatocysts ; s p , s p o r o p h y l l ; s t , s t i p e ; r , f l a t t e n e d , r h i z o m e - p o r t i o n o f h o l d f a s t ; h , h a p t e r a . F i g . 4 .—Diagrammat ic r e p r e s e n t a t i o n of s e v e r a l s t i p e s r i s i n g f rom t h e same h o l d f a s t of M a c r o c y s t i s i n t e g r i f o l i a B o r y , showing g e n e r a l p o s i t i o n t a k e n "by t h e s t i p e s i n t h e water at f o u r d i f f e r e n t s tages of t h e t i d e . Dot ted l i n e s i n d i c a t e t h e p o s i t i o n of t h e s u r f a c e of the w a t e r . w i t h respec t t o t h e p o s i t i o n of t h e p l a n t . S o l i d l i n e s i n d i c a t e the swath which would he removed by c u t t i n g mechanism w h i c h cuts 6 f e e t be low t h e s u r f a c e o f t h e water a t t h e v a r i o u s t i d e - l e v e l s i n d i c a t e d . F i g u r e s on t h e l e f t i n d i c a t e t i d e - l e v e l s . E n -l a r g e d t i p s on t h e s t i p e s i n d i c a t e t h e approximate p o s i t i o n of t h e t e r m i n a l l a m i n a e . (See F i g . 25 f o r c h a r a c t e r i s t i c s ) . F i g . 5,-r-Small sporophytes o f M a c r o c y s t i s i n t e g r i f o l i a Bory shoiving v a r i o u s stages i n development of the h o l d f a s t , D shows a p l a n t w i t h f i r s t d i v i s i o n t a k i n g p l a c e . ' • F i g . 6 .—Young sporophyte o f M a c r o c y s t i s i n t e g r i f o l i a B o r y which has a t t a i n e d c o n s i d e r a b l e s i z e b e f o r e f i r s t d i v i s i o n has reached any s i z e . ) F i g . 7 .—Young sporophyte of M a c r o c y s t i s i n t e g r i f o l i a B o r y showing e a r l y d i v i s i o n s i n development o f the p l a n t at the base and i n d i c a t i o n of segmentat ion a l s o t a k i n g p l a c e a t t h e apex o f t h e t h a l l u s . F i g . 8 .—Young sporophyte o f M a c r o c y s t i s i n t e g r i f o l i a B o r y shoxiring e a r l y d i v i s i o n s i n t h e t h a l l u s . F i g . 9.—A young sporophyte o f M a c r o c y s t i s i n t e g r i f o l i a B o r y showing o r i g i n of the v a r i o u s s t i p e s and growing " p o i n t s " , s t i l l t r a c e a b l e a t t h i s p o i n t t o t h e o r i g i n a l s i n g l e - b l a d e f o r m . 0 F i g . 1 0 . — A young sporophyte o f M a c r o c y s t i s i n t e g r i f o l i a B o r y showing f i r s t p a i r of s t i p e s c o m p l e t e l y s e p a r a t e d . F i g . 11.—A young sporophyte of M a c r o c y s t i s i n t e g r i f o l i a Bory showing f i r s t p a i r of s t i p e s comple te ly s e p a r a t e d . F i g . 1 2 . — P o r t i o n s o f young sporophytes of M a c r o e y s t i s i n t e g r i f o l i a B o r y showing v a r i o u s degrees i n c o m p l e t i o n of d i v i s i o n . F i g . 1 3 . — P o r t i o n of a young p l a n t of M a c r o c y s t i s i n t e g r i f o l i a B o r y showing v a r i a t i o n i n r a t e of s p l i t t i n g . F i g . 1 4 . — A c o m p a r a t i v e l y young s t i p e of M a c r o c y s t i s i n t e g r i f o l i a Bory showing p r o p o r t i o n s of t e r m i n a l l a m i n a and b a s a l l a m i n a s p l i t t i n g up i n t o s p o r o p h y l l b l a d e l e t s . (See F i g . 15 f o r l o w e r p o r t i o n of same p l a n t ) . 1 5 . — A c o m p a r a t i v e l y young sporophyte of M a c r o c y s t i s i n t e g r i f o l i a B o r y ( h a l f o n l y shown) i n d i c a t i n g e a r l y d i v i s i o n s i n f o r m -a t i o n o f t h e t h a l l u s . (See F i g . 14 f o r upper p o r t i o n o f same p l a n t . ) F i g . 1 6 , - - S p o r o p h y l l f r o m M a c r o c y s t i s i n t e g r i f o l i a Bory showing lamina d i v i d i n g i n t o b l a d e l e t s . 17.--Sporophyll o f M a c r o c y s t i s i n t e g r i f o l i a Bory showing lamina d i v i d i n g i n t o b l a d e l e t s . -/ F i g . 1 8 . - - S p o r o p h y l l o f M a c r o c y s t i s i n t e g r i f o l i a B o r y showing d i v i s i o n of l a m i n a i n t o b l a d e l e t s . Note t h e s l i g h t s w e l l i n g a t n b " i n s t e a d of a prominent pneumatocyst. ' 1 9 . - - S t e r i l e l a m i n a o f M a c r o c y s t i s i n t e g r i f o l i a B o r y . F i g * 1 2 0 . — T e r m i n a l lamina of M a c r o c y s t i s i n t e g r i f o l i a Bory f rom a young s t i p e . Note the shape, p r o p o r t i o n , and number o f b lades i n p r o g r e s s i v e s tages of d i f f e r e n t i a t i o n a s compared w i t h F i g . 2 3 . F i g . 2 1 # - ~ T e x m i B a l l a m i n a o f M a c r o c y s t i s i n t e g r i f o l i a B o r y * o F i g ; 2 2 . - - T e r m i n a l lamina of M a c r o c y s t i s i n t e g r i f o l i a B o r y . F i g . 2 3 . — T e r m i n a l l amina of M a c r o c y s t i s i n t e g r i f o l i a B o r y . F i g * ' 34.~ H o l d f a s t o f a young p l a n t of M a c r o c y s t i s i n t e g r i f o l i a B o r y showing how the h a p t e r a form a l o n g t h e s t i p e as d i v i s i o n s . c o n t i n u e and how t h e lower p o r t i o n o f t h e l a t t e r assumes a c r e e p i n g p o s i t i o n p r i o r to expanding i n t o t h e c r e e p i n g r h i z o m e . A i s a v i e w l o o k i n g down on the h o l d f a s t and B i s a v i e w of the same h o l d f a s t f rom t h e s i d e . ' F i g . 2 5 . — H o l d f a s t of a young p l a n t of M a c r o c y s t i s i n t e g r i f o l i a B o r y showing development of h a p t e r a . 6 F i g . 26.!—Growing " p o i n t " o f a p o r t i o n o f a more mature h o l d f a s t of M a c r o c y s t i s i n t e g r i f o l i a B o r y . a i s t h e point of a t t a c h -ment of a w e l l - d e v e l o p e d s t i p e t o t h e rh izome; b i s a lowermost s p o r o p h y l l on t h e s t i p e a l r e a d y shoxring f u r t h e r d i v i s i o n ; c i s the c r e e p i n g r h i z o m e ; d i s one of t h e b r a n c h i n g h a p t e r a a r i s i n g from the edge of the c r e e p i n g r h i z o m e ; f i s t h e next l a r g e s t s t i p e t o deve lop a t t h i s " p o i n t " ; g i s the lowermost s p o r o p h y l l on t h e s t i p e f ; e i s a p r i m a r y d i v i s i o n ; h and i a r e t h e i n n e r segments produced b y t h e d i v i s i o n ; k i s the p o i n t a t which t h e t h i r d l a r g e s t s t i p e w i l l d e v e l o p ; j i s the l o w e r m o s t - s p o r o p h y l l o f t h e s t i p e d e v e l o p i n g at k . 1 F i g . 27 .—Drawing of a p o r t i o n o f a more e x t e n s i v e hapteron showing method of b r a n c h i n g (b) i n H a c r o c y s t i s i n t e g r i f o l i a B o r y . The p o i n t o f at tachment t o t h e rh izome i s shown and (a) i s a c r o s s - s e c t i o n t h r o u g h the r h i z o m e . F i g . 28.- . -Diagrammatic r e p r e s e n t a t i o n o f t h e method of b r a n c h i n g found i n the h o l d f a s t of M a c r o c y s t i s i n t e g r i f o l i a Bory shaving p o s i t i o n of s t i p e s w i t h respec t t o a g e . See context f o r complete e x p l a n a t i o n o f symbols . The d o t t e d r e g i o n i n d i c a t e s t h e p o r t i o n o f t h e s t i p e which develops i n t o the broad c r e e p i n g rhizome from w h i c h b r a n c h i n g haptera extend a l o n g b o t h m a r g i n s . ( N o t e . — T h i s diagram o n l y r e p r e s e n t s one h a l f o f the system d e v e l o p i n g from one p l a n t , and i t s c o u n t e r p a r t s h o u l d be repeated on the o p p o s i t e s i d e i n order t o complete t h e t h e o r e t i c a l form a t t a i n a b l e ) . APPENDIX D P L A T E S P l a t e 1 . —Photograph of complete p l a n t o f M a c r o c y s t i s i n t e g r i f o l i a B o r y a t 1.5 f t . t i d e , 8 a . . m . A u g . 27, 1946 i n Welcome Harbour (Porch e r I s l a n d ) . P l a t e 2 . — P h o t o g r a p h of beds o f M a c r o c y s t i s i n t e g r i f o l i a Bory near t h e entrance t o Welcome Harbour (Porcher I s l a n d ) at 1 .5 f t . t i d e , 8 a . m . , A u g . 27, 1946. P l a t e 3 . — C l o s e - u p photograph of p l a n t o f M a c r o c y s t i s i n t e g r i f o l i a Bory at 1 .5 f t . t i d e , 8 a . m . A u g . 27, 1946 i n Welcome Harbour (Porcher I s l a n d ) . P l a t e 4 . — P h o t o g r a p h of bed o f M a c r o c y s t i s i n t e g r i f o l i a Bory a t t h e n o r t h e a s t end o f A r i s t a z a b a l I s l a n d a t an 8 f t . t i d e , 1;10 p . m . , August 18 , 1946. P l a t e 5. .—Photograph of a bed of M a c r o c y s t i s i n t e g r i f o l i a Bory behind a f r i n g e o f N e r e o c y s t i s l u e t k e a n a at M e l v i l l e I s l a n d on a 10 f t . t i d e , 12:26 p . m . , August 23 , 1946. P l a t e 5 P l a t e 6 . —Photograph of a bed o f M a c r o c y s t i s i n t e g r i f o l i a B o r y t a k e n at t h e n o r t h of Banks I s l a n d on a 9 f t . t i d e , 1:30 p . m . , August 19, 1946. P l a t e 7 .—Photograph o f a bed o f M a c r o c y s t i s i n t e g r i f o l i a B o r y t a k e n a t t h e n o r t h o f Banks I s l a n d near Deadman I s l a n d , ' 1:10 p . m . , August 19, 1946 on an 8 f t . t i d e . Note sandy beach i n background. P l a t e 8 ,—Photograph of bed o f M a c r o c y s t i s i n t e g r i f o l i a B o r y t a k e n a t t h e n o r t h o f Banks I s l a n d near Deadman I s l a n d on a n 8 f t . t i d e , 1:10 p . m . , . August 19 , 1946. P l a t e 9 .—Photograph o f bed of M a c r o c y s t i s i n t e g r i f o l i a Bory w i t h some s c a t t e r e d p l a n t s of N e r e o c y s t i s l u e t k e a n a around t h e r o c k s at t h e approaches t o Norway I n l e t on a 10 f t . " t i d e , August 19, 1946. P l a t e 1 0 .—P h o t o g r a p h o f bed o f M a c r o c y s t i s i n t e g r i f o l i a Bory around r o c k s a t entrance t o Norway I n l e t on 15 f t . t i d e , 8 a . m , , August SO, 1946. Note P l a t e 11 which i s t a k e n i n t h e same l o c a t i o n a t a d i f f e r e n t s tage o f t h e t i d e . P l a t e 1 1 .—B e d o f M a c r o c y s t i s i n t e g r i f o l i a Bory around r o c k s a t entrance t o Norway I n l e t on 10 f t . t i d e , 3 :40 p . m . , August 20, 1946. Note P l a t e 10 w h i c h i s t a k e n i n t h e same l o c a t i o n a t a d i f f e r e n t s tage o f t h e t i d e . ' P l a t e 11 P l a t e 12 .—Photograph o f bed of M a c r o c y s t i s i n t e g r i f o l i a Bory a t the east end of Hope I s l a n d (note sand beach i n back-ground) on an 11 f t . t i d e , 12:30 p . m . August 16 , 1946, P l a t e 1 3 . — P h o t o g r a p h of bed of M a c r o c y s t i s i n t e g r i f o l i a Bory a t the n o r t h o f Deer i s l a n d on a 5 f t . t i d e , 10:00 a . m . August 15 , 1946. F o r t Rupert i n background. P l a t e 14, —Photograph o f bed o f M a c r o c y s t i s i n t e g r i f o l i a B o r y a t t h e n o r t h of Deer I s l a n d on a 5 f t . t i d e , 10 :00 a . m . , August 15 , 1946, E a g l e I s l a n d i n background. P l a t e 12 P l a t e 14 P l a t e 15 - - P h o t o g r a p h o f t e r m i n a l laminae o f M a c r o c y s t i s i n t e g r i -f o l i a B o r y showing range i n s i z e , p r o p o r t i o n and number of laminae i n v a r i o u s degrees of s e p a r a t i o n . P l a t e 16 .—Photograph o f t e r m i n a l l a m i n a o f M a c r o c y s t i s i n t e g r i f o l i a Bory showing p r o g r e s s i v e s tages i n development o f t h e u n i l a t e r a l l y produced l a m i n a e . Note t h e attachment o f t h e laminae t o t h e s t i p e by means o f pneumatocysts . P l a t e 1 7 . — A c o m p a r a t i v e l y young s t i p e of ^Macrocys t i s i n t e g r i f o l i a Bory broken o f f w i t h a s m a l l s e c t i o n of t h e h o l d f a s t a t t h e growing " p o i n t . " A n o t h e r s m a l l e r s t i p e i s p a r t l y shown i n t h e l o w e r r i g h t c o r n e r . The growing " p o i n t " i s between. Note t h e two lowermost b l a d e s on t h e l a r g e r s t i p e : t h e lowermost i s an unbulbed s p o r o p h y l l jus t b e g i n n i n g t o s p l i t up i n t o b l a d e l e t s . The next above i s a b u l b e d s p o r o p h y l l w h i c h i s a l s o j u s t b e g i n -n i n g t o s p l i t up i n t o b l a d e l e t s . Plate 15 Plate 17 P l a t e 1 8 . — Photograph o f t h e upper p o r t i o n of a major s t i p e of M a c r o c y s t i s i n t e g r i f o l i a B o r y . The t e r m i n a l lamina i s a t t h e extreme l e f t . Note the t a t t e r e d , n a t u r e o f t h e more b a s a l laminae (at extreme r i g h t ) and t h e heavy growth o f O b e l i a . P l a t e 1 9 . — P h o t o g r a p h o f p o r t i o n cut f rom s t i p e o f a p l a n t o f M a c r o c y s t i s i n t e g r i f o l i a B o r y showing how two laminae each p o s s e s s i n g i t s own pneumatocyst a r e j o i n e d by a common pneumatocyst t o the s t i p e . P l a t e SO,—Photograph o f a p o r t i o n o f a p l a n t o f M a c r o c y s t i s i n t e g r i f o l i a Bory s k w i n g i r r e g u l a r i t y i n s t i p e f o r m -a t i o n due t o incomple te segmenta t ion . Plate 18 Plate 20 P l a t e 2 1 . — H o l d f a s t of a p l a n t of M a c r o c y s t i s i n t e g r i f o l i a Bory spread out on a l o g . The upper p o r t i o n of most s t i p e s has been cut away. Note t h e s p o r o p h y l l s on t h e s t i p e s c l o s e t o t h e h o l d f a s t and t h e number o f s t i p e s a r i s i n g from the one h o l d f a s t . P l a t e 2 2 . — C l o s e - u p photograph o f a h o l d f a s t o f M a c r o c y s t i s i n t e g r i f o l i a Bory showing a p o r t i o n of t h e broad " c r e e p i n g r h i z o m e . " P l a t e 23 .—Photograph o f M a c r o c y s t i s i n t e g r i f o l i a Bory and a few p l a n t s of N e r e o c y s t i s lue tkeana ( M e r t . ) P . & R . washed up on the beach i n t h e v i c i n i t y o f the Keough R i v e r a f t e r a southeas t s t o r m . P l a t e 23 P l a t e 24 .—Photograph o f bed o f M a c r o c y s t i s i n t e g r i f o l i a Bory a t the n o r t h of Deer I s l a n d at about 2 f t . t i d e . T w i n Rocks i n background. P l a t e 2 5 . - - P h o t o g r a p h o f bed o f M a c r o c y s t i s i n t e g r i f o l i a B o r y a t the n o r t h of Deer I s l a n d a t about 2 f t . t i d e . Pla te 25 P l a t e 26.—Photograph o f l a r g e rock at t h e n o r t h o f Deer I s l a n d a t a b o u t . z e r o t i d e l e v e l covered w i t h s m a l l p l a n t s o f M a c r o c y s t i s i n t e g r i f o l i a Bory mixed w i t h s m a l l p l a n t s of L a m i n a r i a s a c c h a r i n a (L . )Lamour . P l a t e 27.—Laminar i a s a c c h a r i n a ( L . ) L a m o u r . , growing on r o c k s , mixed w i t h s m a l l p l a n t s o f M a c r o c y s t i s i n t e g r i f o l i a B o r y , a t about z e r o t i d e - l e v e l , on r e e f a t the n o r t h of Deer I s l a n d . P l a t e 26 P l a t e 27
Thesis/Dissertation
eng
Biology
Vancouver : University of British Columbia Library
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The life history and distribution of Macrocystis in British Columbia coastal waters
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