The Distribution of Certain Benthonic Algae in Queen Charlotte Strait, British Columbia, in Relation to Some Environmental Factors
ROBERT F. SCAGEL 1
IN COMPARISON with the progress in our knowl cially of members of the Laminariales on the edge of most groups of plants-especially con Pacific Coast of North America, on the basis of cerning their life histories and distributions latitudinal and seasonal temperature distribu the advances made in marine phycology and tions. The physical data available during this marine ecology have been relatively slow. The early period were limited, but many of the prin limited access to living material or to the facil ciples set forth by Setchell concerning the dis ities to maintain the larger marine algae in the tributions of marine algae are as sound now as living condition for a prolonged period of time, when they were first proposed. Except for more the difficulties of collection-particularly in the precise knowledge of the physical and chemical subtidal zone-and the lack of any extensive factors of the environment and the distribu direct economic importance until recent years tions of the algae concerned, much of Setchell's have all contributed to this slow progress. How ecological work can still be used as a good ever, in spite of these difficulties there has been foundation for further study. Although it was a considerable amount of interest in the marine largely a two-dimensional approach to the algae, including a number of studies of their marine environment, Setchell's work made a ecology. Although this interest has been fairly significant contributiOn to the development of widespread in a number of countries, until re marine algal ecology. cently there has been little activity in the field Lamouroux (1825, 1826) had suggested the of marine ecology relating to the benthonic possibility that temperature stratification in the algae on the Pacific Coast of North America sea might account for the vertical distribution and nothing of a comprehensive nature has been of the marine algae and had considered the published for this area. It is an anachronism effect of tides on intertidal zonations, but this that this should be so in a region which received trend to analyze the vertical distribution of the such prominence some 50 years ago through marine algae was not generally taken up in de the efforts of a pioneer in the field, the late tail until much later. Coleman (1933) was one William Albert Setchell (1893, 1917, 1935). of the first to emphasize the use of tide levels Knowledge of the effect of temperature on to account for the vertical distribution of the the world-wide distribution of plants both hor marine algae in the intertidal zone. In a study izontally and vertically had developed gradually in Oregon, on the Pacific Coast of the United over a period of many years. However, it was States, Doty (1946) has given further evidence only during the last hundred years that the at for the relationship between the vertical dis tention of phycologists was brought to a con tributions of marine algae and critical tide levels. sideration of the reasons for the observed dis A number of lists of marine algae have been tributions of the marine algae. The historical published and attempts have been made not development of this trend of thought and in only to relate the floras of one area to another, vestigation has been reviewed by Setchell such as that by Okamura (1926, 19'32) in the (1917). Starting over 50 years ago, through a North Pacific, but also to account in a general series of papers from 1893 to 1935, Setchell way for distributions on the basis of ocean cur made a noteworthy atte mp t to explain the rents, such as that by Isaac (1935) in the area world-wide distribution of marine algae, espe- around South Africa and by Tokida (1954) in the region of northern Japan. However, there 1 Department of Biology and Botany, and Institute soon followed a decided shift to intertidal studies of Oceanography, University of British Columbia, Vancouver 8, Canada. Manuscript received March 31, of regional areas, such as that by Feldmann 1960. (1937) in the Mediterranean and Chapman
494
I I ...... n&...u..m.....'...... II1lM.M,liJiJUl...a:.nQj..._ Benthonic Algae-ScAGEL 495
125°W.
DIXON ENTRANCE
LANGARA I. tit 54· N. 54· N .
00: GRAHAM 1 ~ 1.- q. .BRITiSH COLUMBIA. ~t MORESBY 1...' 0", MAINLAND ~ "0 (~ ~ U'
QUEEN CHARLOTTE SOUND
NORTH PACIFIC OCEAN 49° N. 49°N,
CAPE
1300 W. 125°W.
FIG. 1. Map of geographical features of the coast of British Columbia. 496 PACIFIC SCIENCE, Vol. XV, OctOber 1961
(1950) and his students in New Zealand. Some without simplification or clarification. There attempts have also been made to describe uni have been a number of recent comprehensive versal features of intertidal zonation through papers dealing with various aspects of marine out the world (Stephenson and Stephenson, ecology which make it unnecessary to dwell at 1949). At the same time there has been atend length on a review of the trends that have been ency to place greater emphasis on the inter followed more recently in marine algal ecology relationships between the various organisms. and the results that have been attained (Gislen, Many of these intertidal studies have been 1929, 1930; Feldmann, 1937, 1951; Fischer of great value as an initial descriptive stage of Piette, 1940; Chapman, 1946, 1957; Dory, 1957; investigation, and there is a need for further HartOg, 19'59). descriptive studies of this type in new and un Although the shift in emphasis to the inter described regions. However, the variety of sys relationship of organisms was an important one, tems of nomenclature and terms that have been in some instances this approach has been re proposed by marine ecologists to describe zona sponsible for excluding adequate concurrent tion, associations, and other ecological concepts studies of the physical and chemical aspects of has frequently only complicated the descriptive the environment. It is for this reason that a case study rather than succeeded in explaining the may be made for reassessing the status of marine observed phenomena. This has led to some con algal ecology, and a critical evaluation of the fusion in terminology. It is a debatable point steps to be taken to further its progress is timely. whether there can be such a thing as a universal Perhaps what may be called a three-dimensional system of classification beyond a generalized or an oceanographic approach can be used to scheme, such as that proposed by Ekman (19'35), analyze more precisely various factOrs in the and it is questionable whether some of the sys marine environment and the relationship of tems proposed can contribute further to progress these factOrs to the benthonic algae. Steps in in marine algal ecology even in regional studies this direction have been made more recently
50"45'
VANCOUVER ISLAND
DEPTH (fATHOMS)
• OVER 100 OJ] 'JvER 50 50'30' ~ OVER to 128' D LESS THAN 10 , 127"30' 127"
FIG. 2. Map of Queen Charlotte Strait showing depth contours. Benthonic Algae-SCAGEL 497
SALI NITY %. 26 27 28 29 30 31 32 33 34 13r-----::..'------~---~---~---~------=:;.'------...::.:=------..::z..---=---___,
-- QUEEN CHARLOTTE STRAtT 12 ---- QUEEN CHARLOTTE SOUND _._._. JOHNSTONE STRAIT ...... KNIGHT INLET
2 T-S II DIAGRAMS JUNE 1953.
10 2_ -.-._._ ,I -'- -'- -'!--
QUEEN CHARLOTTE SOUND
7
FIG. 3. T-S diagrams for stations in Queen Charlotte Strait and adjacent areas in June, 1953. with some measure of success by Dawson (1945, algal ecology have been descriptive rather than 1951) in Baja California, Doty (1946) in Ore functional in nature. As in most oceanographic gon, and Womersley (1956) in Australia. work there is value in an approach from the The physical environment of the sea imposes grosser aspects to the particular. To the ocean problems of considerable magnitude which, in ographer the most complicated physical or chem contrast to many land environments, presents ical situation to explain may be the smallest formidable obstacles such as complex tidal and unit of the environment with which he is faced. circulation patterns. In an effort to manipulate This is partly a problem of instrumentation. or simulate some aspects of this environment, However, itjs usually much easier to recognize both in the field as well as in the laboratory, the significant dIscontinuities in properties, such as more generally available resources are soon taxed. temperature, salinity, and even plankton dis Thus the methods that have been used in marine tributions, over extensive areas of the ocean algal ecology have, in many instances of neces than in restricted or local regions. It is also sity, been crudely quantitative, less than ideal easier to use such information in describing experimentally, frequently encumbered by ter dynamic processes. Hence, it is suggested that minology as a result of limited concurrent phys more attention should be given to studies of the ical and chemical data, and sometimes they have general distribution of various physical and failed to establish clearly the objectives being chemical properties in the marine environment sought. As a result most of the efforts in marine in an attempt to set up some workable hypo- 498 PACIFIC SCIENCE, Vol. XV, Ocrober 1961 theses to account for observed distributions of way in which these are considered may be some marine algae. In this way we may hope to ex what a matter of interpretation. Salinity, for plain and account for biological phenomena example, may be considered directly, from a rather than be satisfied by a description of the chemical standpoint, or indirectly as a physical phenomena or by devising terms to describe factor responsible for changes in density and them which do nothing more than give names thus contributing to the pattern of circulation. to dynamic aspects of marine ecology much in Likewise, the nature of the substratum may be need of logical explanation. With the recent considered indirectly as a geological factor or increased activity in oceanography in the Pacific directly as a physical or mechanical factor' re we may now hope for more abundant and usable stricting or permitting establishment of ben data on some of the more general oceanographic thonic organisms because of particle size. There properties of the North Pacific. In specific cases, has been much written on some of these aspects particularly in more restricted areas, the ecol of ecological study in special cases, but it is ogist will be forced to turn more attention to suggested that, in a general over-all reassess obtaining in situ physical and chemical data ment of the environment, an attempt be made before further progress can be made. to proceed from this more general position to One can arbitrarily start by summarizing all the particular. This approach may initially lead the factors in the marine environment as geo only to the erection of further hypotheses, since logical, physical, chemical, and biological. The the indirect or direct nature of the action of
SALINITY '4. 28 29 30 31 32 33 10 39
___ QUEEN CHARLOTTE STRAIT T-S _____QUEEN CHARLOTTE SOUND DIAGRAMS a GOLETAS {;HANNEL MAY 1956 9 _._._. JOHNSTONE STRAIT ...... FIFE SOUND a WELLS PASS. 2\ P 21.,~ ...8 II: "':':':":\.~ ::;) .... 29 t ol( ....:::;. ...II: A. ...2 t- 7
QUEEN CHARLOTTE SOUND 56 6 55!
FIG. 4. T-S diagrams for stations in Queen Charlotte Strait and adjacent areas in May, 1956.
c:. == "'''''Il...... 4i...... iI4LL,;;:;;sw:..,.....mLWLAAl::n:>iihAl1JJlliMiiiID11W1lJlOOlt. Benthonic Algae-SCAGEL 499
128'
QUEEN CHARLOTTE SOUNO
+16 51' .PINE I.
50'45'
VANCOUVER ISLAND
STATION POSITIONS 50'30' CRUISE 53/1 128' 127'
FIG. 5. Map of station positions in Queen Charlotte Strait in June, 1953.
128'
QUEEN CHARLOTTE SOUND
51'
50"45'
VANCOUVER ISLAND
STATION POSITIONS 50'30' CRUISE 56/1 128" 127"
FIG. 6. Map of station positions in Queen Charlotte Strait in 'May, 1956. 500 PACIFIC SCIENCE, Vol. XV, October 1961
128·
QUEEN CHARLOTTE SOUND 5'·
50·45'
VANCOUVER ISLAND
~ JUNE 1953 Swfou _ ,~-- 50·30' 128· 127·
FIG. 7. Salinity distribution in Queen Charlotte Strait in June, 1953, at the surface and at a depth of 3 m.
QUEEN CHARLOTTE
SOUND
51·
50·45 '
VANCOUVER ISLAND
SALINITY t°4.) JUNE-1953 IOm. _ 20~ __ 50·30' 128· .127·
FIG. 8. Salinity distribution in Queen Charlotte Strait in June, 1953, at depths of 10 and 20 m. Benthonic Algae-SCAGEL 501
128·
QUEEN CHARLOTTE SOUND
51·
VANCOUVER ISLAND SALINITY ('Y••) MAY 1956 SURFACE _ 3 METERS __ 50·30' 128· 127·30' 127·
FIG. 9. Salinity distribution in Queen Charlotte Strait in May, 1956, at the surface and at a depth of 3 ffi.
128·
QUEEN CHARLOTTE SOUND
51·
VANCOUVER ISLAND
SALINITY (%o) MAY 1956 10 METERS _ 20NETERS__ !50METERS .. 50'30' 128' '127'
FIG. 10. Salinity distribution in Queen Charlotte Strait in May, 1956, at depths of 10, 20, and 50 ffi. 502 PACIFIC SCIENCE, Vol. XV, October 1961 any particular factor in the environment may the direction of increasing our knowledge of ultimately be established only by experimental the entities which comprise the tools of the work either in the field or in the laboratory. In algal ecologist and of completing this descrip some regions where the algal flora is well known tive phase of the study of the marine benthonic taxonomically the descriptive aspect can and algae, an annotated check list has been com has been undertaken and the time has come pleted for the coast of British Columbia and when further progress in such an area can only northern Washington (Scagel, 1957). Based on be expected by undertaking experimental field this list, further studies are now in progress to and laboratory studies. In some regions, even augment the existing data on the marine flora where the flora may be well known, inadequate of British Columbia, not only on distributions physical and chemical data militate against fur but also on life histories, growth, reproduction, ther progress and even the advancement of and seasonal aspects. These fundamental studies tenable hypotheses. Only when such hypotheses are basic to all other aspects of ecological re are put forward, based on a correlation of the search, especially when an attempt is made to observational data, can one anticipate and justify use specific organisms as indicators of ocean embarking on an experimental field and lab ographic conditions. oratory approach in an attempt to solve prob Not only does this descriptive phase require lems relative to the distribution of marine algae. an adequate consideration of the taxonomic as This is the approach that is outlined here in pects, but also a complete description of the studies on marine organisms and, particularly other factors in the environment is needed. With on the marine algae, being carried out on the increased activity recently and currently in gen coast of British Columbia, some aspects of which eral oceanographic studies of the North Pacific will be considered here in detail. As a step in by a munber of organizations on the Pacific
MONTHS
J F M A M J J A S o N o 34 ...... - ...... ----...... ------...... ---"--.....- --''--_...0.-..,
33 / ...... / ...... __.- . ._._._.-._./ ,-_._. ----.-. . . ._.- ...... -_.- 32 -
0 ~ ...------...... ------...... - .,.,...,..,...... 1. /1(" ...... -.. __--...--- ~_",_JI: >- 31 -~ /+~" I- / --", -'-JlC_x :+0-/ \+ - "'---..~ Z " + / \ /+ '- / '--~ \ ~/...... J 30 ...... 29 SAL I NIT Y (%0) PINE I. 1942-1952 (MONTHLY) ._._.- MAXIMA •...... •...... MEAN MAXIMA ---- GRAND MEAN ---- MEAN MINIMA l<-l<-MINIMA FIG. 11. Monthly salinities at Pine Island for the petiod 1942-52. Benthonic Algae-SCAGEL 503 MONTHS J F M A M JJ A S o N D 33 32 ._._._._._._._._._._._._._._._. .- - -:..-:- -:-.~:::::7" .... /"/ ...... 1(__ ":~:':':'::~:._._._._ /' ~..,. "'--lI(-x-Je._ _--...... 31 _._._'/' ...... ,. """__Il Jit__Jil:-x_J("...... , .. ~/"".,. ""...... '-, ~ 30 ./ \ '-'-, > I- ./ \ ----- z 29 oJ « \ Ul +-JiC-Jit 28 SAL I NIT Y (%.l MALCOLM 1. (PULTENEY POINT) AUG. 1954 - DEC. 1955 (MONTHLY) MAXIMA MEAN MAXIMA GRAND MEAN MEAN MINIMA ..-~- MINIMA FIG. 12. Monthly salinities at Pulteney Point, Malcolm Island, for the period 1954-55. Coast of Canada, in Japan, and in the United with large marine algae present unique culture States, there has resulted a considerable body problems both in the field as well as in the lab of scientific knowledge but it is still far from oratory. Some studies of growth and reproduc adequate for the ecologist, particularly for one tion, particularly of some of the larger Lami interested in coastal dynamics. Circulation pat nariales, have been done in this region both in terns, temperature, salinity, oxygen, and in some the field (Scagel, 1948) as well as in the lab areas phosphate, silicate, and nitrate distribu oratory. Although the size of many~f"dle cold tions are fairly well known in a broad and gen water marine algae adds special problems, at eral sense, but at the present time these ocean least some of the stages can be carried Out to ographic data are known in sufficient detail for the point where transplant experiments can be few restricted areas. Knowledge of the distribu made from the laboratory into the sea for fur tion of other chemical constituents and to a ther study. Transplant experiments of natural large extent even of the plankton composition, populations of juvenile stages, at least of these distribution, and activity is almost completely larger marine algae, even in the case of Ma lacking. Much more information is needed in crocystis, are quite feasible. order to tackle many problems relating to spe The successful use of the experimental ap cific distributions and to set up field and lab proach in the laboratory is primarily dependent oratory studies to test hypotheses. on having facilities for maintaining tempera It is already apparent that much can be done ture and light control, although the size of experimentally both in the field and in the plants may again present certain special prob laboratory with the benthonic algae. Many of the lems. Cultures of Laminariales have been main problems encountered by the ecologist dealing tained in controlled-environment tanks at the WJiJlOU):uiMt:teAllEta....~a"""...... ".&a:&.iiJIL\UllXt:w. 504 PACIFIC SCIENCE, Vol. XV, OctOber 1961 University of British Columbia for as long as 1949). These studies have dealt largely with a year, during which the complete sexual gen harvestable quantities and distributions, and erations were grown and the young sporophytes have contributed little to an evaluation or an reached a length of 14 in., well past the stage explanation, in terms of oceanographic factors, where secondary morphological characteristics of the causes for this production. had developed to a point permitting positive Obviously the ideal of a functional interpre identification. These studies have permitted in tation in the ecology of marine benthonic algae disputable identification of the sporophytes to is dependent on an adequate and balanced knowl genus, and in some cases to species. The study edge of all of the foregoing aspects-of the of cultures in this group suggests that much of qualitative and quantitative features of both the the early work on gametOphytes in the Lami organisms and the environment. Many, much nariales and in fact even on the early sporo needed data are still lacking. An attempt to fol phytes may be in some question. In most of the low this line of investigation has been pursued early studies reported in the literature, plants on the coast of British Columbia and in a some were not grown long enough to establish beyond what more restricted area at the north end of doubt the characteristic secondary morphological Vancouver 1. in Queen Charlotte Strait. Fur features of the sporophytes of the genera from ther detailed work is in progress in the Strait of which zoospores were intially obtained. In the Juan de Fuca at the south end of Vancouver 1. presence of contaminating zoospores of other between Vancouver 1. and northern Washing species which can soon supplant the original tOn. The study in Queen Charlotte Strait forms species under study, there is no other way of the major part of this paper. establishing that the same species or even the same genus in the Laminariales was obtained in the sporophyte generation succeeding the JUNE 16 HOUR OF DAY (PST) gametOphyte generations in culture. 12 ...... ~0~60~0~08:;:0~0~10:::00:c-1:=J20~0---..:1~40::::0~16:.::::00::...... '::::.:80::::0--=:20::.::0::::0---, It would be remiss not to mention much of E the worthwhile physiological work that has been w !5... done on marine algae lind other organisms. « 10 However, there is a need for a great deal more '"w.. physiological work, particularly of the type done ...w" by Gail (1918, 1919, 1922) in an attempt to .... !! 32 8 relate physiological processes more specifically ~ z and directly to the environment and ecological :::; « problems encountered in the field. In physio '" 3\ logical studies there is frequently a tendency to proceed more and more deeply into special as pects of the physiological behaviour or the bio chemistry of an organism under artificial condi 10 tions. Although this information is very often ~, of great value there is a very real need to project . ,,:;;-;:.,..-'~ // / back to the field and attempt to explain be 'x..,'-.../' haviour under the conditions existing in the natural enwonment. The statistical approach, STATION 19 (1953) as illustrated by Berquist's (1959) revealing QUEEN CHARLOTTE STRAIT -_ OmUets study of Hormosira, has also been little used :3 metU$ as yet. ----- 10 meters l(-l(-l( 11-14.5 meters Almost all of the quantitative aspects of the productivity of the benthonic algae in this area FIG. 13. Fluctuations in temperature, salinity, and have related to species of economic interest oxygen at various depths near Malcolm Island at sta (anon., 1947, 1948a; Scagel, 1948; Hutchinson, tion 19 (1953). 506 PACIFIC SCIENCE, Vol. XV, October 1961 ical (anon., 1954; Pickard and McLeod, 1953) characteristic of the coast provide an ideal area and geological data (Dawson, 1880, 1881a, in which to study the distribution of marine ben 1881b, 1888, 189'7; Bostock, 1948) have been thonic organisms particularly in relation to referred to in the literature. Earlier biological salinity over a rather extensive geographic area. observations and collections, which form part of the background for this paper, were made in GEOLOGICAL CHARACTERISTICS the area in 1946 (anon., 194'7, 1948a) and 1947 (Scagel, 1948). The results of a study of the General Coast Features phytOplanktOn and zooplanktOn collections The Coast Mountains of British Columbia, which were also made during the same cruises which run along the whole length of the prov- will be presented in a subsequent paper. HOUR OF DAY (PST) GENERAL ECOLOGICAL CHARACTERISTICS JUNE 18 JUNE 19 ::'i"..,--",,,--,,,"'-...:=-~'''~.~'~OOO~2..~.-,,22,,,,..,--·:c:"..",--".",...,--..~..~..~..,--.=OOO::o....:"",-=:, OF COASTAL REGION The Pacific Coast of Canada is ideally suited to a study of benthonic organisms and the effect of oceanographic factors on their distribution 32 both in the intertidal and the subtidal zones. Although the coast of British Columbia (Fig. 1) is only about 600 mi. long, proceeding di rectly from the Strait of Juan de Fuca to Dixon Entrance, if all its various ramifications are in cluded there is a coastline estimated at about 16,900 mi. in length. The tidal amplitude in this region is great, ranging from about 11 ft. at the southern boundary to nearly 26 ft. at the northern boundary. As a result of thorough mix ing in the coastal region the upper zone in this area, except fora few local anomalies, is char acteristically rather uniform in temperature at anyone period and fluctuations occur within narrow limits. The annual range in temperature of the seawater near the surface is from about 6° to 18°C. On the other hand, because of the runoff from large rivers, especially through the 15 long mainland inlets, there are conditions rang ing from practically fresh water at one extreme to full ocean salinity of about 34 %0 at the other. Throughout the coast the physical nature of the substratum, ranging from mud and sand at one extreme to solid rock at the other, deter mines to a large extent the organisms which are found in a specific area. However, a com STATION 21 QU((Irf QWIIlOTTE STRAIT parison of the flora and fauna on various types -.~ ...... ) . of bottom is possible in a number of regions ----·10 .... which are otherwise oceanographically rather ·---25-30111. similar. This permits a correlation of the dis tribution of a wide variety of plants and animals FIG. 15. Fluctuations in temperature, salinity, and with other physical and chemical factOrs of the oxygen at various depths near the Klucksiwi River at environment. The oceanographic conditions station 21 (1953). Benthonic AIgae-SCAGEL 507 HOUR OF DAY (PST) JUNE 19 JUNE 20 12 r-_.:.:18:..0=-0=--~2000.:.;..=--..:2::;2..:.0.:.0--=2:...:4.:0.:0--.:0~2:...:0::0:...... :0.;.4.:0:..0....:0.::.60:..0::.....:::.0:::.80:..0::...... :1:..0:::0:..0---:.12~0.::0::-..::14~0~0:...... :1~6.:.::0:..0_..::18?10~0~..=2;;:0;.0;.0_.., II o ~ 10 0..: ~ W ~ 8 32 >- 31 ~ z ~ 15 14 13 12 ~ ~ "- II 0' E ----'z 10 w (9 >- :1 X 0 8 7 STATION 22 (1953) QUEEN CHARLOTTE STRAIT o m. 3 m. 10 m ,,-x-x 14-16.5m. FIG. 16. Fluctuations in temperature, salinity, and oxygen at various depths near the Klucksiwi River at station 22 (1953). HOUR OF DAY (PST) JUNE 20 JUNE 21 JUNE 22 12 2200 2400 0200 0400 0600 0800 1000 1200 1400 1600 1800 2000 2200 2400 0200 0400 u o ...... II w cr ...... ::> I- 10 32 .. ------~ lt~- -Jr_Il-_~~~~~-=::~~~~~==~~~~=l=·= ... - ....-I'.-Il-..-,,-·--.. .. >- f:::: ------...... _------_.------z 31 ...... -.J 30 16 ...... 15 ... ~ . 14 .-, 13 -.J '- d> .s 12 z w (!) II >- x 0 10 9 8 7 6 STATION 23 (1953) QUEEN CHARLOTTE STRAIT o motors ~ motors 10 motors x--.<-x 20 motors 25.5 -~O motors ·FIG. 17. Fluctuations in temperature, salinity, and oxygen at various depths near Deer Island at station 23 (1953). Benthonic Algae-SCAGEL 509 ince with an average width of about 100 mi., (Fig. 1). These sediment-filled systems of valleys constitute the mainland coast. Although not as with steep slopes and numerous deltas can be rugged as elsewhere in the Cordillera, the west traced in some places even through the coastal ern side of the Coast Mts. rises from the sea archipelago, which represents a partly sub precipitOusly to summits in places exceeding merged margin of the Coast Mountains. West 8,000 or 9,000 ft. Several rivers, which rise in of the Coast Mountains, and in a partly sub the plateau country to the eastward, flow com merged condition, lies another chain, the In pletely across this range to the Pacific, where sular Mountains, of which Vancouver I. and the lower parts of their valleys, as well as those the Queen Charlotte Islands are projecting of many streams originating in the mountains ranges. This outer chain stands on the edge of themselves, continue in the extensive system of the continental platform with the great depths fjords along the mainland of British Columbia of the Pacific seaward from it. Between these MONTHS J F M A M JJ A S o N o j ...... / "--'-"",. / ""'._._\ / ...... \ / ...... //... . \ . / ..... / ------_...... // "./' ------" ...... ,/. / /'-"-~...... ' ..... / .. "-.. " .,-// ~-~_./ ' ...... ,- / '"+ , /"'/" /' " / . '"" ------_// / / ,+" ",/. TEMPERATURE (oG) ",, / PINE I. x-t-x_-. 1942 -1952 (MONTHLY) -._.- MAXIMA ...... MEAN MAXIMA GRAND MEAN ------MEAN MINIMA X-K--lt MINIMA FIG. 18. Monthly temperatures of seawater at Pine Island for the period 1942-52. 510 PACIFIC SCIENCE, Vol. XV,Ocrober 1961 MONTHS 12 .-- J F ...... M__"""-A M JJ.a.- A --a.S__o~....;... N o.., II ,,/...... //.". /' / ". 10 /// ...... \""":::::::-:>---. (} L 9 w ./" /./~ ~ :..::;-;~, ...... 0:: ./'" / ~ f- FIG. 19. Monthly temperatures of seawater at Pulteney Point, Malcolm Island, for the period 1954-55. twO ranges lies the Coastal Trough, part of a TEMPERATURE (CO) great depression extending intermittently north 10 10 10 10 10 10 10 westward from the Gulf of California through the Puget Sound-Williamette lowland and on into Alaska. In British Columbia this great val ley is largely submerged and comprises the ex tensive areas of the Strait of Georgia, Queen Charlotte Strait, Queen Charlotte Sound, and -~ 10 Hecate Strait (Fig. 1). The Coast Mountains consist largely of Mes I ozoic rocks ranging from Triassic ro early Ter f a. o tiary and are composed of principally granitic w g~ ~ ~ o HOUR o it 2 o OF DAY rocks with some included masses of Mesozoic o Q= ! ~ ~ N and Palaeozoic strata. The rocks of the Van STATION 19 couver 1. Ranges are composed conspicuously 16 JUNE 1953 of masses of Triassic and Jurassic lava and vol 9 10 II 12 SCALE (CO) ? ~ • , ! • canic products, lesser contemporaneous sedi· ments with subordinate amounts of later granitic FIG. 20. Temperature profiles at station 19 (1953) rocks, and marine and continental Cretaceous near Malcolm Island from bathythermograph traces. Benthonic Algae-SCAGEL 511 TEMPERATURE (QC) 10 10 10 10 10 10 10 10 10 10 10 10 I, o I. I I. I. I. I• I. I. I. I• I I I I I I I I I I I ') 10 · · - 20 - I l Q.. ·11 W o 30· j 0 0 0 0 0 0 0 HOUR 0 0 It) 0 0 0 0 0 0 0 0 0 CD 0 N - - 0 N ~ CD CD 0 N ~ CD- CD 0 OF DAY 0 - - - - - N 0 0 0 0 0 STATION 20 16 JUNE- 17 J UN E 1953 SCALE (CO) 7 8 9 Ip II 12 • • • , • FIG. 21. Temperature profiles ar sration 20 (1953) near Malcolm Island from barhythermograph rraces. sediments which have participated in its fold chiefly quartz diorite. Parts of the island groups ing. Horizontal Miocene beds occur along some near the entrance to the Strait, including Hope I. parts of the shore. and Nigei I., include some intrusive rocks sim ilar to those on the mainland side, but the major General Features of Queen Charlotte Strait part of the north end of Vancouver I. is com Along the mainland portion of Queen Char posed of Triassic to Jurassic rocks. In this latter lotte Strait (Figs. 1, 3) the rocks are chiefly region argillites, lavas, tuffs, breccia, sandstones, Jurassic and intrusive, composed of granodiorite, and limy siltstones are common. A smaller por- 512 PACIFIC SCIENCE, Vol. XV, October 1961 tion of the east coast of Vancouver 1. from 1., and Malcolm 1. are also examples of these Hardy Bay southward almost to Johnstone Strait deposits. In some places cliffs of these deposits is Cretaceous (chiefly Upper Cretaceous), with border the shoreline, with extensive accumula shales and sandstones. tions of boulders at the base. In such regions The rocks are heavily glaciated throughout the boulders, which occur in great abundance almost the whole coastal area. The Queen Char along the beaches, are probably erratics derived lotte Strait area was heavily glaciated during the from morainic material. Pleistocene, which fact is particularly evident in Although the detailed geology of Queen Char the vicinity of Deer 1., where northwestern lotte Strait region is not well known supratidally, slopes are comparatively rough in contrast to it is even less well known subtidally. The gen the grooved and polished vertical or near ver eral features described, however, indicate the tical faces on southeastern parts. Well-stratified wide variety of substrata available for the at deposits of clays, silts, and sands occur, partic tachment of benthonic organisms, particularly ularly toward the east end of the Strait and along in the intertidal and shallower subtidal zones. the Vancouver 1. side. Cormorant 1., Harwood This variety in the physical nature of the sub- TEMPERATURE 10 10 10 10 10 10 10 10 10 10 10 10 ~ 20 I ~ ~ W Cl 30 40 0 0 0 0 0 0 0 0 0 0 0 0 0 0 HOUR .,..""" 0 0 0 0 0 0 0 0 0 0 0 0 0 ~ N 0 N 0 N .. 0 - ..- ! • til til 0 0 0 0'" •0 ~ OF DAY STATION 21 /8 JUNE - 19 JUNE (1953) 7 8 9 II 12 SCALE (COl If fIG. 22. Temperature profiles at station 21 (1953) near Malcolm Island from bathythermograph traces. TEMPERATURE (DC) 10 10 10 10 10 10 10 10 10 10 10 1010 10 10 10 10 10 tJ:j ~ ." o , I ii' ii' II' j I 'i I Ii' I') I"I' j I' ) I' )I ',I J I I J I I j' I > . ~ ;:;.o :> ...... ~ 10 ~ 1 I ~ § Cl. t"' UJ 0 20i 0 0 0 It) o 0 0 0 0 o 0 0 0 0 0 0 0 HOUR 0 It) 0 r FIG. 23. Temperature profiles at station 22 (1953) near Malcolm Island from bathythermograph traces. TEMPERATURE (Oc.) o I J)I! ) I' ", )I I) I' 71 I 71' } I I J" I I' j I I } I I " I JI I "I I I I) II I, 10 ~ 20 :I: I- a.. w Cl 30 , I 0 0 0 0 0 0 0 0 o o 0 0 o o o o .. o HOUR· 0 0 0 o o ,., o ,., o o '"0 N ,., on CD 0 ,., o o '" ..'" '".. N on CD OF OAY '" 0 0 0 0 0 0 0 '" .. ... N '" N '"N '"o 40J STATION 23 20 JUNE - 21 JUNE (1953) VI 7 8 9 10 II 12 ...... SCALE (CO) • t! \j.l FIG. 24. Temperature profiles at station 23 (1953) near Deer Island from bathythermograph traces. 514 PACIFIC SCIENCE, Vol. XV, October 1961 PELAGIC DIVISION EPIPELAGIC ZONE .r z LIGHTED o . ~ (f) > • MESOPE~AGIC·. " •• • ZONE' • Cl ~* ..... "... ,... '. 0° •• o .~ • -IOOOm. Z ° •• or . .. 0 o ... ' . I '~~'. .~~RK r ARCHIBENTHONIC ": :.': ::•••••••: ':.'•• : •:: :.':..: ::.' Z ZONE 0° •••••• 0°. ° 0 ,•' •• ° "0. 0" W 0 en o .'.: -: •• : ":' • ,.0: ::::: DEEP-SEA '"o (f) SYSTEM o rI Z enW ______-----:~::-----f------"~~~~ii~ FIG. 25. Diagrammatic representation of regions of marine habitat (modified after Ekman, 1935). stratum is apparent not only geographically in geological features of the area indicates the the area but also vertically. In the deeper water present status of published knowledge (Bo soft mud bottoms predominate but sand and stock, 1948; Dawson, 1880, 1881a, 1881b, 1888, gravel are also found, and in the shallower areas 1897) concerning the Queen Charlotte Strait sand, mud, gravel, pebbles, boulders, and solid regIOn. rock are all found to varying extent, particularly along the Vancouver I. side of the Strait. In Bottom Topography of Queen Charlotte Strait the shallower regions, however, a solid rock or Although soundings are still incomplete for bouldery substratum predominates. The nature the area, a study of the bottom topography (Fig. of this rocky substratum in Queen Charlotte 2) in Queen Charlotte Strait indicates exten Strait is equally varied. There are igneous as sive shallows, particularly along the Vancouver well as sedimentary and metamorphosed sub I. side of the Strait and around Malcolm I. strata. There are basalts, dolerites, trachytic In this region an abundant and varied inter rocks, hard sandstones or quartzites, shales, con tidal and subtidal flora and fauna are supported. glomerates, argillites, fine-grained to crystalline, In the central part of the Strait and between commonly cherty limestones mixed with feld Nigei and Vancouver islands (Fig. 2), there spathic rocks, and dioritic and granitic frag are deeper channels exceeding 100 fathoms. The ments are also common. water in these deeper channels is not continu The preceding brief summary of the general ous, however, with the deep waters of the main- Benthonic Algae-SCAGEL SIS land inlets and Johnstone Strait, and exhibits Charlotte Sound with the deeper more saline physical and chemical properties quite distinct water below. The more saline water from the from the latter (Figs. 3,4). open ocean and Queen Charlotte Sound moves into the Strait centrally as well as along the Van PHYSICAL CHARACTERISTICS couver I: side of the Strait and along the north side of Malcolm I. The intrusion of high salin Hydrographic Conditions in ity water along the deep channels in the central Queen Charlotte Strait part of the Strait is also apparent. This general The salinity distribution near the surface pattern of salinity distribution, with fluctuations (Figs. 7-10) in Queen Charlotte Strait suggests to varying degrees near the surface, is pro a general circulation in a counter-clockwise fash nounced in the upper zone to a depth of at ion. The runoff from the mainland inlets along least 20 m. (Figs. 7-10). the north shore and at the east end of the Strait, Although strong winds may assist in the particularly from Knight Inlet at the east end, movement of water near the surface there is contributes large volumes of fresh water which clear evidence at times of the movement of tends to move seaward at the surface, mixing as water against the wind and there are strong it progresses along the north shore into Queen tidal currents throughout the region. The cur- MONTHS J F M A M: J. J A S o N D 30 25 20 ...... •••••••••••••••• 0° ...... <.) ° 0 .... -', -'. L ...... • .....-::._._.-. ". w 15 ... ~.,.;..~---...... :::-...... a:: " '-'-;;;"- -~'_." .."... "" . ::> ...... ,,' /. ll--«_IC...... "...... I- 10 ,,' /' ...... '" ...... ". .. -10 MAXIMA -l(-X MEAN MINIMA MEAN MAXIMA -0-0 MINIMA -15 MEAN NORMAL MEAN (w. COAST VANCOUVER I.) FIG. 26. Monthly air temperatures at Bull Harbour, B. C, for 1953. 516 PACIFIC SCIENCE, Vol. XV, October 1961 MONTHS J F M A M J J A S o N o 30 25 ...... 20 ...... 0 ~ ...... ~::::::...-=-.::-. w 15 .' "7" ... -. a::: .... /._•..",t.'...... ", ::> ..... ,,/ // ',". ". f- lO .' ". /. lI.-IC-X...... :::-...... « ...... /...... ",,""'" "'...... a::: w ._.-./ .,."...- /0-0 - 0 "'-11_" , ...... £L ._.- / ...... "/ 0 ...... 0 "'...... "'~ . 5 ~~ ~ _"./.,. / ...... 0_ ""'~_ W _- ".- 0 0__.., f- -::....x-lI.-X-IC- 0"" 0.... 0 • /0...... '0, ...... 0 0-0-0 0 /-0_0_0- AIR TEMPERATURES (OC) -5 / HARDY BAY - 1953 o -10 0/ MAXIMA IC-lt-X MEAN MINIMA MEAN MAXIMA 0-0-0 MINIMA MEAN -15 NORMAL MEAN (E. COAST VANCOUVER I.) FIG. 27. Monthly air temperatures at Hardy Bay, B. C, for 1953. rent velocities involved vary considerably but able from Pulteney Point on Malcolm I. to at times may reach at least 80 cm/sec from near analyze the seasonal fluctuations more precisely the surface to a depth of 20 m., and in deeper in the central region of the Strait, but for the portions of the Strait, although generally much period available (Fig. 12) a range of about less, they may attain as much as 57 cm/sec at 28 %0 to 32 %0 with an annual mean of 21.50 %0 a depth of 125 m. is indicated. An analysis of surface salinity data over a Seasonal data indicate that near the surface 10-year period (Fig. 11) from Pine I. Light the distribution of salinity throughout the year house, which is near the entrance to Queen follows the same general pattern, decreasing to Charlotte Strait, indicates a salinity maximum ward the mainland and being higher along the of about 33 %0 and a minimum of about 30 %0, Vancouver I. side of the Strait. At anyone with an annual mean of 31.75 %0. Although point, however, there is a general decrease in this station is not characteristic of the Strait it salinity in time from the maximum in April to self, the data available probably give a reason a minimum in midsummer, when the maximum able approximation of the annual salinity fluc runoff from the mainland inlets occurs. The tuations for the Strait. Insufficient data are avail- wimer salinity may be somewhat modified near Benthonic Algae-SCAGEL 517 the surface during the period of maximum pre Sound. The T-S diagrams for Queen Charlotte cipitation, which may reduce salinity near the Strait indicate a characteristically intermediate surface. condition between these extremes in properties The minor extent of fluctuations that occur of temperature and salinity for the greater part in the salinity distribution in the upper 20 m. of the Strait. is indicated by data taken at a number of an An analysis of surface temperature data over chor stations (Figs. 13-17). These fluctuations a lO-year period (Fig. 18) from Pine I. Light are greatest at or near the surface and, in cer house, which is near the entrance to Queen Char tain instances, as near the Klucksiwi River (Fig. lotte Strait, indicates a temperature maximum 15), show the influence of fresh-water inflow of about 12°C. and a minimum of about 5°C., of a more localized nature. with an annual mean of 8.6°C. Although this A comparison of the temperature-salinity station, as already indicated, is not characteris characteristics of various parts of the Strait and tic of the Strait itself in all respects, it prob the connecting bodies of water by means of T-S ably gives a reasonable approximation of the diagrams (Figs. 3, 4) indicates the discreteness annual temperature fluctuations. Insufficient of the water masses typical of Johnstone Strait, data are available from Pulteney Point on Mal mouth of Knight Inlet, and Queen Charlotte colm I. to analyze more precisely the seasonal MONTHS J F M A M J J A S o N D 30 25 ...... 20 ...... u /./._._.""'.- ...•...... •.. -° 15 ...•... .,..".--".",---' ----,...... " ... / ~ w .' . .,., , .. 0:: IC ~ 10 ...... /././ / _"' " II- -: _"'" ; . I- et .-.-' 11-" 0_0-- -"'~ .... 0:: .'" / .,...... 0...... 0.... "'....._ ~ "'• W 5 "'~. a.. .- - ---...... "._.... 0-0-0...... '0 - ~~ ~-)l_""'" -0_ 0 ~ __ / -0 w /0 -0-0_0 I- 0 0/°...... 0_0_°-° AIR TEMPERATURES (oG) ./ ALERT BAY - 1953 -5 MAXIMA -10 MEAN MAXIMA MEAN 1(-1(- MEAN MINIMA -15 o-e-O MINIMA NORMAL MEAN IE. COAST VANCOUVER I.) FIG. 28. Monthly air temperatures at Alert Bay, B. c., for 1953. 518 PACIFIC SCIENCE, Vol. XV, October 1961 J F M A 95 90 ~ ..--.".,....-.-.-. !:! "' >- ' t- -- '\. 0 85 '\...... '\. :iE ' ~ ...... I 80 '..... w ...... > t- « MEAN RELATIVE HUMIDITY (%l ..J 75 W BULL HARBOUR (HOPE .t.l- 1953 a::: (elev. 15') 70 0130 hours 0730 1330 1930 65 FIG. 29. Monthly relative humidity at Bull Harbour, B. c., for 1953. MONTHS J FMAM JJ A SON 100r-"O""'------""O':"--A..---.:A.:--...... -----''---...... :;;.--'':O::'''''---...:;:..--~-~~ 95 90 ..' .. , >- 85 o'= ::E ~ I80 w ...... > ...... i= « ..J75 w a::: 70 -0: MEAN RELATIVE HUMIDITY (%l PORT HARDY (elev. 74') - 1953 65 0130 hours 0730 1330 1930 FIG. 30. Monthly relative humidity at Port Hardy, B. c., for 1953. Benthonic Algae-ScAGEL 519 MONTHS J F M A M JJ A S o N D 100r-'''------'''''---~--"'''''--...... ~-~--''''-----''---..a....----''-- ...... 95 .... ~ 90 ~ ~ >- t- 85 0 ::i: ::> :I: 80 UJ > t- FIG. 31. Monthly relative humidity at Alert Bay, B. c., for 1953. MONTHS J F M A M J J A S o N o 15 PRECI PITATION (inches) ~ en BULL HARBOUR (HOPE I.) Q) ~ () -- 1953 c 10 ...... AVERAGE z .. -.' o ..../ ..... l e::{ I- '. a...... ~ 5 a::: a...... o FIG. 32. Monthly precipitation at Bull Harbour, B. c., for 1953. 520 PACIFIC SCIENCE, Vol. XV, October 1961 fluctuations in the central region of the Strait, face at most times of the year, with the water but for the period available (Fig, 19) a range along the Vancouver I. side being warmer than of from about 5° to 11 0e. with an annual mean that along the mainland side of the Strait. This of 8.25°e. is indicated. is a result of the inflow of colder, less saline The temperature distribution with depth, as water from the mainland inlets. shown by bathythermograph records taken at Seasonal data indicate that in general during shallow anchor stations, indicates a well-mixed the summer months the temperature at the sur region near the surface along Vancouver I. and face is seldom above lO°e. and at a depth of Malcolm I. (Figs. 20-24). Except for slight 20 m. is seldom above 9°e. During the winter anomalies the water is generally almost isother months it is seldom below 7.5°e. at the surface mal, with temperatures during this period of ob bu't may be about re. at a depth of 20 m. This servation seldom above 11°e. at the surface and picture of the vertical temperature distribution falling to not less than 8°e. at a depth of 20 m. is somewhat modified during the winter as a As indicated from the anchor stations, there are result of surface cooling of the water down to no marked changes with time in this general a depth of about 20 m. and a temperature in picture of the vertical distribution of tempera version has been observed in January which dis ture, and at anyone depth the fluctuations in appears by April. During this early period the dicated were less than 2 e.0 (Figs. 13-17). water at the depth of 20 m. and, in places, even However, there is a slight difference (usually to a depth of 400 m., may be warmer than that not more than 1 e. 0) in temperature at the sur- in the upper 20 m. by as much as 1 e. ° and MONTHS J F M A M J J A S o N o 15 o FIG. 33. Monthly precipitation at Hardy Bay, B. c., for 1953. Benthonic AIgae-SCAGEL 521 MONTHS J F M A M JJ A S o N D 15 PRECIPITATION (inches) lJ) Q) .c: ALERT BAY u c 10 --1953 z ...... AVERAGE 0 l- e::{ I- - a... - 0 ...... W 5 0:: a...... O...... ------J FIG. 34. Monthly precipitation at Alert Bay, B. c., for 1953. MONTHS J F M A M JJ A S o N D 100 90 ?J7-·.' ,/ 80 \. ." ,/..-:...:-/ ...... "," ./"" / ::-i? 0 \ " ...... ~ 0: 70 .,." w \ / > "-. I 0 \/ 0 60 "" / o ::J o <3 50 MEAN CLOUD COVER (%l BULL HARBOUR (HOPE I.) -1953 40 0130 hours 0730 1330 30 1930 FIG. 35. Monthly cloud cover at Bull Harbour, B. c., for 1953. 522 PACIFIC SCIENCE, Vol. XV, October 1961 MONTHS JFMA M J JA SON D 1001"'"------"'"------...... ------'------.,;,~'---.....;..---.:.'---.....:....., 90 ~ 80 a: lIJ > 70 0 (,) 0 :::> 60 0 -l MEAN CLOUD COVER (%) (,) HARDY BAY - 1953 50 _._.- 0130 hour. 0730 .. ----- 1330 .. 40 ...... 1930 .. FIG. 36. Monthly cloud cover at Hardy Bay, B. c., for 1953. MONTHS 100 ...... JFMAMJJ'--_....:...__.A:.-_--'-__"""'-_--' A ...... S __...... o _--'N D -; .... ~ -- 90 /! ------II II ---- ~ ...... "",..,.,.,,,,,----- ...... I :' ~ 80 ,;:..--".... ~ I. a: '...... 1 .: lIJ I > ...... 070 .... (,) ...... o ..' g 60 -l (,) MEAN CLOUD COVER (%) 50 ALERT BAY - 1953 -- 0730 houri ___ 1330 ...... 1930 • FIG. 37. Monthly cloud cover at Alert Bay, B. c., for 1953. CMU i...... MN."'.=.S\i&...,,::;3L.j.2Jillmj.li:w.uilzauSlr.tJ.1l'IW0''''~...lSiJ.t,saaaA_ Benthonic Algae-SCAGEL 523 then fall to a lower temperature again, beneath of organisms in this region. A comparison of this warmer upper layer, down to between 6° the meteorological data (air temperatures, pre and 7°C. in the deeper regions. cipitation, mean relative humidities, and mean Thus the environment presented to the inter cloud cover) available for the coast of British tidal and immediate subtidal zones (see Fig. 25 Columbia, particularly from Bull Harbour for terminology) appears to be a relatively (Hope I.), Hardy Bay, and Alert Bay, gives stable one as far as the temperature of the sea some picture of the meteorological conditions water is concerned. at the northeast end of Vancouver I. (Figs. 26-37). Meteorological Conditions Along the coast, air from the maritime Pacific The intertidal region, however, during periods Ocean is usually present and results in mild of exposure, is subjected to a varying degree to winters and cool summers. Holding a high mois meteorological conditions, particularly fluctua ture content, this air does not become extremely tions in temperature and precipitation, which hot or cold. However, occasional outbreaks of must be considered in assessing the environment continental air (polar) from the interior of the 1953 JAN FEB MAR APR MAY JUNE 18 16 14 12 10 8 ~. l 1\ , \ . 1\11 \1/ i 0 I- w w 1953 1L. JULY AUG SEPT OCT NOV DEC 18 16 FIG. 38. Monthly summary of tidal features at Hope Island for 1953 (see Figure 39 for significance of lines). 524 PACIFIC SCIENCE, Vol. XV, October 1961 1953 (JAN.-DEC) the inner coast and the lee side of Vancouver 1., where there is some protection from the maritime influence, precipitation and cloudiness 18 HHHW are somewhat reduced and ranges in tempera HLHW ture are somewhat increased, as at Alert Bay 16 (Figs. MHHW 28, 31, 34, 37). 14 The number of frost-free days in the whole MLHW Queen Charlotte Strait region averages 200-250 12 LHHW in a year. In both ourer and inner regions there LLHW are no months with all temperatures ranging 1-10 W below O°e. In the ourer coastal region 4 to 5 w HHl.W ll..8 \ months have temperatures above 100e., and in to \\ HlLW the inner coastal region 5 6 months have 6 \'. MHL'N temperatures above 100e. In the ourer coastal region, depending upon the locality, the mean 4 \ MLLW monthly air temperatures for January are 1.r to 4.4°e., and for July 13.3°e., and the mean 2 \ LHLW daily temperatures are -1.1° to 1.7°e. and LLLW 0 15.6° to 18.9°e. for the same months, respec tively. In the inner coastal region, depending upon the locality, the mean monthly air tem FIG. 39. Summary of tidal features for 1953. peratures for January are 1.7° to 3.3°e., and for July 15.6° to 18.3°e., and the mean daily . teIJ:lperatures are -1.1° to O°e. and 21.1 ° to continent bring cold periods during the winter, 23.9°e., respectively, for the same months. although generally the Coast and Cascade moun Although some restricted regions of British tains provide considerable protection. Along the Columbia on Vancouver 1. average as much as outer coast, the maritime conditions, which are 264 in. of rain, in the Queen Charlotte Strait present almost continually, result in high pre region the annual rainfall averages between 40 cipitation, prolonged cloudiness, and small and 60 in. at the inner end (including Alert ranges in temperature-the typical conditions Bay and Malcolm 1.), with between 5-10 per prevailing in the vicinity of Hope I. (Figs. 26, cent as snow, and between 60 and 100 in. at 29, 32, 35) and Hardy Bay (Figs. 27, 30, 33, the outer end (including Hope 1. and Port 36) , despite the fact that these points are some Hardy), with less than 5 per cent as snow. The what on the lee side of Vancouver I. Along period of minimum rainfall is in the summer, 1953 JUNE AUGUST M ,. " 2. '0 27 HHHW ,.1------HLHW 1----- A 1 A I /' A /1 A I I A II/ \/ \I \I 1/ 1/ y \ 1/ If f-----I-..- ~e M'MldnlQht PACIFIC STANDARD TIME FIG. 40. Tidal features for the periods of greatest exposure during summer months of 1953. Benthonic Algae-SCAGEL 525 1953 NOVEMBER M 19 M 20 M 21 M 22 M 23 M 18 HHHW HLHW 16 I MHHW 14 n 1\ MLHW r- 12 I \ LLI LHHW LLI l!.. I \ -10 f LLHW LLIo \ HHLW r- 8 \ l!.. o \ \ HLLW r- 6 \ :I: y MHLW Cl J V ~4 MLLW 2 LHLW o LLLW M ~ Midnight PACIFIC STANDARD TIME FIG. 41. Tidal features for period of greatest exposure during winter months of 1953. with between 2-3 in. on the average falling the period mid-May to mid-September the west during August in the Strait, and the maximum erly winds predominate and blow at speeds up rainfall occurs in the winter, with between to 25 m.p.h. Particularly during this latter 9'-13 in. on the average in December. period, however, there may be considerable calm Near the entrance of the Strait, especially periods, especially during the morning hours, along the north side of Hope I., conditions of followed by strong seas which reach a peak heavy surf generally prevail. Even when there about 1600 hrs. and then drop rapidly to rela is no sea a heavy swell is common. Farther tive calm by 2000 hrs. down in the Strait to the south and east there is an area protected by scattered islands and Tidal Characteristics reefs near the entrance, from the heavy swell The tidal amplitude in the vicinity of Hope from the open ocean, but this whole area is sub I. is usually about 17 ft. and the highest on ject to strong westerly winds in the summer record (37 years) was almost 19 ft. (December, months and to even stronger southeasterly winds 1941). An extensive intertidal flora and fauna during the winter months, so that the Strait is is exposed during low tide periods in this zone. generally subjected to strong wave action and Continuous tidal data are recorded since 1949 wind mixing. During the period mid-Septem at Alert Bay, near station 8 (Fig. 6). At the ber to mid-May the southeasterly winds are north end of Vancouver I. the tides are semi predominent and blow at speeds frequently up diurnal and only moderately declinational, and to 40 m.p.h. and occasionally higher. During thus springs and neaps are distinguishable (Figs. 526 PACIFIC SOENCE, Vol. XV, October 1961 1953 (MONTHLY) M A M JJ A 5 o N o o '" on ON (f) TIDAL REGION (FEET) o'" ~ ~ -'" ----- MLHW-MHHW (13.1-14,51 I.LJ ••••••• . . MHHW-HLHW 114.5-16.41 o 1953 (MONTHLY) M A M JJ A 5 o N o TIDAL REGION (FEET! MHLW-HLLW (5.&-6.41 MLLW- HLHW (3.Z-S.6) HLLW-HHLW (6.4-e.2) LH L W-MLLW(L7-3.21 -~-)t-lC- LLL W- LHLW 10.3-1.7) -0-0- H HLW- L LHW (8.2 -10.4) FIG. 42. Diagram showing number of times each month during 1953 varIOus levels 10 the intertidal zone were exposed. 38-41). Twice a month there are two tides a during the winter months (Fig. 41) the lowest day which are about equal, and in the intervals low waters occur late at night or early in the between there is a much greater inequality in morning. the height of any twO successive low waters than The number of times during the year when between the two high waters of the same day. each of the various levels recognized (Fig. 39) During the summer months (Fig. 40) the low are exposed is indicated in Figure 42, and the est low waters occur during daylight hours and number of days of continuous exposure of the Benthonic Algae-SCAGEL 527 upper portions of the intertidal zone are shown BIOLOGICAL CHARACTERISTICS in Figure 43. The number of times (and per cent) these levels are exposed (Fig. 44) and Biological observations, extending over the submerged (Fig. 45) are also presented in an length of the coast, indicate that there is a high attempt to indicate possible critical levels. The degree of uniformity in the populations of many tide levels are referred to in feet above or below benthonic plants and animals extending from the datum (zero point) which, for the coast of the Strait of Juan de Fuca to Dixon Entrance. British Columbia, is the level of lowest normal This would be anticipated under such relatively tides. uniform conditions of temperature. In attempt ing to correlate the distribution of some of these Chemical Characteristics organisms with salinity characteristics, as well During the summer months, when plant pro as other oceanographic factors, there are several duction is at its peak in Queen Charlotte Strait, areas on the coast which could be used for pur the surface zone may be supersaturated with poses of this study. Although some supporting observations have been made in the Strait of oxygen; values in excess of 15 mg/l are fre quently encountered. The maximum values gen Juan de Fuca and Dixon Entrance, this paper erally prevailing near the surface and to a depth is restricted largely to a consideration of the; vicinity of Queen Charlotte Strait near the north of 20 m. are between 7 and 10 mg/I. During end of Vancouver I. the summer months the values are somewhat higher than in the winter. The maximum con Horizontal Distribution of Organisms in centration of oxygen occurs at a depth between Queen Charlotte Strait 2 and 5 m. (Figs. 46, 47), rather than right at the surface during the summer, and is related Biological observations have been made to the region of maximum phytoplankton activ throughout the area although a more intensive ity. The water throughout most of the Strait study and collection has been undertaken at has a higher oxygen content near the surface Hope I., Deer I., and in the vicinity of the (Figs. 46,47) than in Queen Charlotte Sound Keogh River, the Klucksiwi River, and Malcolm or in the adjacent connecting mainland chan I. These areas present a transition from Hope nels. Although marked fluctuations occur locally I., where the highest salinities are encountered, in the upper 10 m., the general picture is more to the north and east sides of the Strait, where stable (Figs. 13-17) at greater depth. lowest salinities are found, with Deer I. and Phosphate concentrations (POrP) are not Malcolm I. being intermediate between these available for the winter months, but for the extremes. The distributions in the Strait of the summer, during which minimum amounts are more conspicuous organisms (Table 1) observed probably reached, the values present (Fig. 48) during this study are illustrated in Figure 49. were between 0.5 and 2.0 mg.-at. per liter in Although both the marine algae al'id the in the upper 20 m. The minima were generally vertebrate animals have been observed, the em in the upper 10 m. and most of the minima phasis in this study is on the more conspicuous for the Strait and Queen Charlotte Sound were marine algae. between 1.0 and 1.5 mg.-at. per liter. Some organisms in the area are more cos- NUM8ER OF 0.4T5 CONTINUOUS [)I'POSURE (1953) ""tTl FIG. 43. Diagram showing number of days various levels in the intertidal zone were subjected to conr'in uous exposure. 528 PACIFIC SCIENCE, Vol. XV, October 1961 TABLE LIST OF THE MORE CONSPICUOUS SPECIES OF MARINE BENTHONIC ORGANISMS IN QUEEN CHARLOTTE STRAIT 1. Polyneura latissima (Harvey) Kylin 50. Pleurophycus gardneri Setchell et Saunders 2. Chthamalus dalli Pilsbry 51. Agarum fimbriatum Harvey 3. Tegula funebralis (Adams) 52. Agarum cribrosum Bory 4. Acmaea instabilis (Gould) 53. Hedophyllum sessile (c. Agardh) Setchell 5. Balanus cariosus (Pallas) a. smooth form 6. Endocladia muricata (Harvey) J. Agardh b. bullate form 7. G!oiopeltis furcata (Postels et Ruprecht) J. 54. Postelsia palmaeformis Ruprecht Agardh 55. Lessoniopsis littoralis (Farlow et Setchell) Reinke 8. Prionitis lanceolata Harvey 56. Pterygophora californica Ruprecht 9. Prionitis lyallii Harvey 57. Egregia menziesii (Turner) Areschoug subsp. 10. Erythrophyllum delesserioides J. Agardh menziesii 11. Petrocelis franciscana Setchell et Gardner 58. Fuctts evanescens C. Agardh f. evanescens 12. H ildenbrandia occidentalis Setchell 59. Fucus gardneri Silva f. gardneri 13. Opuntiella californica (Farlow) Kylin 60. Pelvetiopsis limitata (Setchell) Gardner f. 14. Plocamium pacificum Kylin limitata 15. Gigartina papillata (c. Agardh) J. G. Agardh 61. Cystoseira geminata C. Agardh 16. Gigartina sitchensis Ruprecht 62. Smithora naiadum (Anderson) Hollenberg 17. Iridaea cordata (Turner) Bory 63. Porphyra perforata J. Agardh f. perforata 18. Iridaea heterocarpa Postels et Ruprecht 64. Farlowia mollis (Harvey et Bailey) Farlowet 19. Halosaccion glandiforme (Gmelin) Ruprecht Setchell 20. Rhodymenia palmata (Linnaeus) Greville f. 65. Dilsea californica (J. Agardh) O. Kuntze palmata 66. Gloiosiphonia californica (Farlow) J. Agardh 21. Gastroclonium coulteri (Harvey) Kylin 67. Nereocystis luetkeana (Mertens) Postels et 22. j'yficrocladia borealis Ruprecht Ruprecht 23. Ptilota asplenioides (Esper) C. Agardh 68. Macrocystis integrifolia Bory 24. Ptilota californica Ruprecht 69. Alaria nana Schrader 25. Ptilo'ta hypnoides Harvey 70. Alaria marginata Postels et Ruprecht 26. Hymenena setchellii Gardner 71. Alaria tenuifolia Setchell f. tenuifolia 27. Hymenena flabelligera (J. Agardh) Kylin 72. Alaria valida Kjellman et Setchell f. valida 28. Cryptopleura ruprechtiana (J. Agardh) Kylin 73. Bangia fuscopurpttrea (Dillwyn) Lyngbye 29. Polysiphonia collinsii Hollenberg var. collinsii 74. Bossiella plumosa (Manza) Silva 30. Pterosiphonia bipinnata (Postels et Ruprecht) 75. Bossiella californica (Decaisne) Silva Falkenberg vat. bipinnata 76. Calliarthron regenerans Manza 31. Laurencia spectabilis Postels et Ruprecht 77. Calliarthron schmittii Manza 32. Rhodomela larix (Turner) C. Agardh 78. Callithamnion pikeanum Harvey var. pikeanum 79. Callophyllis edentata Kylin 33. Odonthalia floccosa (Esper) Falkenberg 34. Odonthalia washingtoniensis Kylin 80. Callophyllis firma Kylin 81. Cladophora trichotoma (c. Agardh) Kiitzing 35. Prasiola meridionalis Setchell et Gardner 82. Spongomorpha coalita (Ruprecht) Collins 36. Agardhiella coulte1·i (Harvey) Setchell 83. Codium fragile (Suringar) Haribt 37. Saundersella simplex (Saunders) Kylin 84. Codium setchellii Gardner 38. Heterochordaria abietina (Ruprecht) Setchell et 85. Coilodesme californica (Ruprecht) Kjellman Gardner 86. Constantinea simplex Setchell 39. Desmarestia intermedia Postels et Ruprecht 87. Constantinea subulifera Setchell 40. Desmarestia herbacea Lamouro~x 88. Corallina officinalis var. chilensis (Harvey) 41. Desmarestia media val. tenuis Setchell et Gardner Kiitzing 42. Desmarestia munda Setchell et Gardner 89. Costaria costata (Turner) Saunders 43. Soranthera ulvoidea Postels et Ruprecht f. 90. Costaria mertensii J. Agardh ulvoidea 91. Cryptosiphonia woodii J. Agardh 44. Myelophycus intestinale Saunders 92. Cttmagloia andersonii (Farlow) Setchell et Gard 45. Scytosiphon lomentaria (Lyngbye) J. Agardh f. ner lomentaria 93. Cymathere triplicata (Postels et Ruprecht) J. 46. Coilodesme bulligera Stroemfelt Agardh 47. Laminaria cuneifolia J. Agardh f. cuneifolia 94. Halicystis ovalis (Lyngbye) Areschoug 48. Laminaria saccharina (Linnaeus) Lamouroux f. 95. Pylaiella litoralis (Lyngbye) Kjellman saccharina 96. Leathesia difformis u. ~ i::J&.,'D.l3SAil.DA,.• a:·~,.G&.....m...&,i:l'a:a:&;;J.ll&il!C;",.,.mNl.\$lJ/i.d1\i:ltlRiGRiiUi&ii Benthonic Algae-SCAGEL 529 TABLE 1 (continued) 98. Ahnfeltia concinna J. Agardh 113. Strongyloce·ntrotus franciscanus (Agassiz) 99. Rhodoglossum latissimum ]. Agardh 114. Mytilus edulis Linnaeus 100. Iridaea lineare (Setchell et Gardner) Kylin 115. Littorina planaxis Phillippi 101. Ahnfeltia plicata (Hudson) Fries 116. Dictyoneurum californicum Ruprecht 102. Schizymenia pacifica Kylin 117. Phyllospadix scouleri Hooker 103. Rhodoglossum affine (Harvey) Kylin 118. Zostera marina 1. var. marina 104. Flustrella corniculata (Smitt) 119. Balanus nubilus Darwin 105. Mytilus californianus Conrad 120. Vlva latissima 1. 106. Mitella polymerus (Sowerby) 121. Rhizoclonium ripariftm (Roth) Harvey 107. Pisaster ochraceus (Brandt) 122. Porphyrella gardneri Smith et Hollenberg 108. Balanus glandulus (Darwin) 123. Desmarestia ligulata (Lightfoot) Lamouroux 109. Styela montereyensis (Dall) 124. Amplisiphonia pacifica Hollenberg 110. Haliotis kamtschatkana Jones 125. Rhodymenia pertusa (Postels et Ruprecht) J. Ill. Strongylocentrotus drobachiensis (Muller) Agardh 112. Strongylocentrot1ts'purpuratus (Stimpson) 126. Pterochondria woodii (Harvey) Hollenberg mopolitan in their distribution, particularly in phyra perforata J. Ag. f. perfarata, Rhodomela their tolerance to extreme dilution. Extending larix (Turn.) C. Ag., Odonthalia floccosa throughout the Strait (Fig. 49) are forms such (Esper) Falk., Mytilus edulis Linnaeus, Lit as Alaria tenuifolia Setchell f. tenuifolia, Cym torina plimaxis Philippi, and Strongylocentrotus athere triplicata (P. and R.) J. Ag., Costaria drobachiensis (Muller). costata (Turn.) Saunders, C. mertensii J. Ag., Restricted to the region of highest salinity, Laminaria saccharina (1.) Lamour. f. saccharina, as at Hope I., are Postelsia palmaefarmis Rupr., Nereocystis luetkeana (Mert.) P. and R., Por- Lessoniops: r littoralis (Farl. and Setch.) Reinke, Laminaria setchellii Silva, Pelvetiopsis limitata (Setchell) Gardner f. limitata, Dilsea califor NUMBER OF TIMES EXPOSED (1953) nica '0. Ag.) O. Kuntze, Erythraphyllum deles 2.00 400 600 BOO 1000 1200 1400 serioides J. Ag., Iridaea lineare (S. and G.) Kylin, Hymenena setchellii Gardner, Ptilota 18 asplenioides (Esper) C. Ag., P. californica Rupr., HHHW P. hypnoides Harvey, Mitella polymerus (Sow HLHW 16 erby), and Flustrella corniculata (Smitt). MHHW A smooth form of Hedophyllum sessile (c. 14 MLHW Ag.) Setch., Pleurophycus gardneri Setch. and I- Gardner, and Styela montereyensis Dall are also ~ 12. LHHW ~ present in regions of highest salinity, but are LLHW w 10 somewhat less restricted in their distribution o l- and extend into the Strait as far as Deer I. LL 8 HHLW o A few organisms extend still farther into the I- Strait but only slightly beyond Deer I. Among ~ HLLW 6 MHLW w these are Alaria nana Schrader and A. marginata I 4 P. and ~. Still others extend from Hope I. MLLW throughout the Deer I. region and as far as 2. LHLW Malcolm I., but not as far as the east and north LLLW sides of the Strait. Among these are MytilttS 2.0 30 40 50 60 70 BO 90 100 californianus Conrad, Strongylocentrotus pur NUMBER OF TIMES EXPOSED (%) puratus (Stimpson), Macrocystis integrifolia Bory, Egregia menziesii (Turner) Aresch. subsp. FIG. 44. Diagram showing number (and per cent) of times tidal condition caused exposure of various menziesii, the typical bullate form of Hedophyl regions in the intertidal zone. lum sessile (c. Ag.) Setch., Alaria valida 530 PAOFIC SOENCE, Vol. XV, October 1961 NUMBER TIMES SUBMERGED (1953) conspicuous organisms at Hope 1. (Fig. 50). o 100 200 300 Comparisons are made also for some of these 18 ( Fig. 51) at Hope 1., Deer 1., and near the HHHW Klucksiwi River, to indicate the effect exposure 17 HLHW to surf has on the vertical distribution of some 16 organisms. 15 MHHW By comparing the vertical distributions of 14 organisms in Queen Charlotte Strait (Fig. 50) f- 13 MLHW w with tidal data presented with respect to emer ~12 LHHW gence and submergence (Figs. 42--45), a num II w ber of limiting levels are fairly apparent. Near ~IO LLHW f- the top of the intertidal zone is a region u.. 9 o (HHHW to HLHW) which is rarely submerged f.. 8 HHLW (Fig. 43) for more than a few hours on each of I a few days at any period of the year and which '"w 7 HLLW 6 MHLW OXYGEN (mg/L) MLLW 20 3 2 LHLW o LLLW o 5 10 15 20 50 NUMBER TIME S SUBMERGED (')I.) FIG. 45. Diagram showing number (and per cent) I of times tidal condition caused submergence of vari ous regions in the intertidal zone. 100 I ! -211 , w (Kjellm. and Setch.) f. valida, Constantinea sim .. I OXYGEN (mg/L.) ~ 150 plex Setch., and Haliotis kamtschatkana Jones. I JUNE - 1953 I ! t I _·_·_·-JOHNSTONE STRAIT Isolated populations of the abalone (Haliotis eL -----ClUEEN CHARLOTTE SOUND W • i kamtchatkana) , which have been noted farther o i --QUEEN CHARLOTTE STRAIT ! e~stward in Johnstone Strait and which may I 200 I be related to local oceanographic features, pre I I sent something of an anomaly in the general i I distribution. An exception to the general dis i I tribution described in this group is that of 250 i I Macrocystis integrifolia. Although Macrocystis i occurs in regions of high salinity and extends i i as far down the Strait as the north side of Mal i i colm 1. and the south side of Numas 1., it does 'DO i i not occur in the most exposed areas where i i Postelsia and Mitella are encountered. i i Vertical Distribution of Organisms 350 ~ The vertical distributions with reference to FIG. 46. Distribution of oxygen with depth at vari tide levels, based on data obtained using an ous stations in Queen Charlotte Strait and adjacent Abney level, are presented for some of the more regions in June, 1953. = 'DUth.. lWIUMI...i2ilWI,lWLllliiiJUiJlS:;....iJJi2LlINiliiiZli..MilJllJAiaMkUIm:m Benthonic Algae-ScAGEL 531 0, (mg./L) is submerged at least once a day. Although the upper limit of a number of the organisms (Fig. o 15 50) in this region of the intertidal zone between 13.1 and 17.4 ft. varies to some extent with the organism, it is apparent that in this general /, ,"'I'. region an upper boundary is probably deter Ii'"1/ mined directly or indirectly by the degree of exposure to climatic conditions. The precise tide 50 " ) level, if there is one, at which this boundary ,,1 occurs is not clear from the data available. The /I: effect of surf in the exposed environment may / /'1'I / A also cause some variation in the upper limit of 100 ( 2lQlz the vertical dist ributi 0 n. Since the greatest V> I I ~ I change in degree of exposure within this upper I E'" I I region (13.1-17.4 ft.) occurs between MLHW I I 1 and MHHW, a critical level is suggested at this f- I "- I W ISO I point for a considerable number of conspicuous 0 I : organisms. .1 1 '16 ) All portions of the intertidal zone below J / I / MLHW are submerged at least once a day for I / ,,I various periods. The greatest change in condi 200 1~2 I I tions of submergence (Fig. 45) in the region I OXYGEN (mg/U . 1 MAY - 1956 of the middle intertidal zone (8.2-13.1 ft.) 1 I --OUEEN CHARLOTTE STRAIT occurs between HHLWand LLHW, and it is i ------QUEEN CHARLOTTE SOUND I I 8 GOLETAS CHANNEL at this point where another critical level is sug 1 _·_·_·--JOHNSTONE STRAIT 250 .. I , ...········-FIFE SOUND a WE LLS PASS gested, in some instances as the upper limit and 1 1 in others as near the lower limit of the vertical :I I distribution of certain organisms (Fig. 50). I I In the lower intertidal zone (0-8.2 ft.) the I 1 300 "0 most extreme change in conditions of exposure (Fig. 44) and submergence (Fig. 45) occurs between MLLW and MHLW, and this is again FIG. 47. Distribution of oxygen with depth at vari ous stations in Queen Charlotte Strait and adjacent reflected both at the upper and lower limits in regions in May, 1956. the vertical distribution of certain organisms (Fig. 50). Another critical level (Fig. 50) occurs in the region between LLLWand LHLW, may be continuously exposed for as long as 4 where another region of marked change (Fig. months during the summer period. Those or 45) exists in conditions of submergence. ganisms that can tolerate such conditions are rare Although the upper limits in most instances and in this zone one finds chiefly Littorina, which appear to be relatively sharp, there is less con is capable of moving sufficiently to extend into lower less extreme zones when necessary. In the sistency in this respect concerning lower limits. region below (HLHW to MHHW), continuous This suggests that other factors, perhaps com petition for space or predation, may be responsi exposure to the air may last for periods ranging ble for limiting distribution, particularly down from a few days to almost 2 weeks, and these ward in some instances. periods occur at least twice a month, but the rest of the time this zone is submerged at least once a day. In the region below (MHHW to DISCUSSION MLHW), continuous exposure to the air is Knowing as we do from experimental work rarely for more than a few days at a time, and the responses of some organisms to environ for a few months during the winter the region mental factors, it seems reasonable to anticipate _!1tiUbl,J"ih•.&,t1J1JUJlG'MlI'tIItLWJM.L,'"UIilG lOW ••..., 532 PACIFIC SCIENCE, Vol. XV, October 1961 50 100 .-- \ U1.... \ Q) \ +0- Q) \ E \ \ \ I \ ~ 0... 150 31 \\ W \ \ 0 \ \ \ \ \ \ \ \ PHOSPHATE- PHOSPHORUS (mg.at./U \ \ MAY - 1956 \ \ \ \ ---QUEEN CHARLOTTE STRAIT \ \ \ \ 200 \ 52 ------QUEEN CHARLOTTE SOUND \ a GOLETAS CHANNEL \ \ \ _.__.- JOHNSTONE STRAIT \ I .... · ...... FIFE SOUND a WELLS PASS I I \ 250 \ il \ I I \, \ \ \ \ 300L.- ---=.:~_J5068 FIG. 48. Distribution with depth of phosphate-phosphorous at various stations in Queen Charlotte Strait and adjacent regions in May, 1956. Benthonic Algae-SCAGEL 533 NUMBERS REFER TO SPECIES LISTED IN HORIZONTAL DISTRIBUTION TABLE I kLUCKSIWI MALCOLM JoPPR. klUCkSIW, MAlCOLM SPECIES HOPE I. DEER I. KEOGH R SPECIES HOPE I. DEER KEOGH R APPR, . RIVER ISLAND INLETS I. . RIV!ft ISLAND INL!T, 116 91 10 ~ 97 24 I-- 102 23 I-- 78 60 I-- 110 55 I-- 42 - 54 I-- 46 65 I-- 57 : 94 I-- 61 126 I-- 113 122 I-- 47 106 I--- 68 104 I-- 45 100 ~ 70 99 ~ 64 103 ~ 4 81 ~ 15 49 17 - 26 ~ 36 8~ 22 50 f--- 84 25 I--- 89 56 I--- 90 109 I--- 93 98 95 101 107 124 121 77 21 III 530 123 117 83 112 87 105 96 76 43 74 44 86 62 13 73 92 82 69 I 66 5 75 6 35 9 80 28 72 29 3 32 125 33 119 39 I 34 40 37 41 38 30 51 2 , 52 16 53b 20 7 63 II 71 12 48 14 58 18 59 I 19 108 27 118 31 120 85 115 88 114 79 67 FIG. 49. Horizontal distribution of marine benthonic organisms in Queen Charlotte Strait. that the distribution of marine organisms in such a many-sided and complicated study would time and space can be explained on the basis of be an avoidance of reality. However, in such a oceanographic features-provided the geologi study one expects to make assumptions based cal, physical, chemical, and biological factors of on the knowledge available, in some instances the environment and their interaction can be of necessity by extrapolation, and to pass adequately assessed. Any attempt to oversimplify through a descriptive phase in an attempt to 534 PACIFIC SCIENCE, Vol. XV, October 1961 correlate observed distributions on the basis of There have been many attempts to describe and in relation to other factors in the environ the zonation of marine organisms on the shore, ment. These comparisons may be largely qualita and a great jumble of appalling confusion in tive in the first instance, but the intent is that ecological terminology has evolved to the point they be not only qualitative but also quantitative where some magic significance is sometimes in the final analysis and, as in all branches of associated with the terms and units involved. science, eventually permit predictions. At best, Although the need for this descriptive phase of however, only a descriptive treatment can be at the study (with a minimum of terminology) is tempted at this point and can only point the way recognized at an early stage of study or in a to further studies and hypotheses. new area under investigation, once it has served VERTICAL DISTRIBUTION TIDE LEVEL r r 'il: 'il::I: r :I: :r r :I: r :I: r r r· :I: r rr :I: :I: :I: LIST OF SPECIES ~ ~ :E :E:E :E :E :E LEVEL ("u) 51 116 ------56 79 93 89 109 - f-- Z 104 - t-- c 70- 'il: 50 til 68 /T1 67 :0 83 en 119 - 86 - :0 9 - _. 1'1 55 - ." 117 1'1 49 :0 25 -- 24 -- -t -t 10 - J> 0 8 - lD 100 r 45 1'1 en 57 -- 65 - ... "1'1 107 t-t- 0 22 - t--- 1'1 88 - en 69 18 ~ .8 ~ C 74 en 15 1- -t 53 1'1 2€ 0 29 I 38 54 Z 17 t-- 30 t-- 106 105 32 6 108 5 63 60 7 16 60 II 73 '- 115 -t-- FIG. 50. Vertical distribution of some marine benthonic organisms at Hope Island. Benthonic Algae-SCAGEL 535 HOPE I. DEER I. ------VERTICAL DISTRIBUTION KLUCKSIWI R...... TIDE LEVEL r r 3: 3:I I r r 3: 3: II r I r Ir I r I r I rI r r r rr r I I I I II LIST 0 F SPECIE S ::E ::E ::E ::E::E ::E ::E ::E ::E ::E ::E~ LEVEL (FEn) I , ..'..0 - N OJ .j:> U10"l --.J OJ lD N OJ .j:> (ji U1 .j:> OJ N 0 - O"l--.J OJ lD ENDOCLADIA MURICATA GLOIOPELTIS -'- FURCATUS HETEROCHORDARIA ABIETINA PETROCELIS FRANCISCANA PORPHYRA PERFORATA FIG. 51. Comparison of vertical distribution of some benthonic organisms at several points in Queen Char lotte Strait. its initial purpose little refinement in this on a broad scale are frequently less apparent method of approach seems conducive to an ex and confused in a local area. The distribution planation of the causal factors. From this point of the genus Macrocystis is an interesting case it becomes necessary to look at the problem in point. Although the global pattern of dis from a new perspective involving a detailed tribution of this genus is rather clearly estab study of the organisms concerned-an under lished (Setchell, 1932; Womersley, 1954) on standing of their life histories, rate of growth, the basis of temperature distributions and hence reproduction, and various physiological require follows the pattern of distribution of some of ments and tolerances in relation to the environ the cold water currents of both southern and ment. These problems may be and, as already northern hemispheres, the distribution of M. indicated, have been approached to some extent integrifolia in British Columbia follows a dis by actual field studies and experiments as well tinct salinity distribution. As yet, however, it as by laboratory studies under controlled condi cannot be conclusively stated in the latter in tions. stance that salinity itself is the causal factor. It is with this philosophy in mind that this It still remains to be established whether salin study has been approached to the extent possible ity in terms of an osmotic relationship, or some from the existing data, initially from the stand parallel factor associated with open ocean water point of the oceanographer with an analysis of of high salinity, provides a causal mechanism the environmental factors and their relation to for the distribution of M. integrifolia on this: the organisms. The success of these preliminary coast. efforts both in the field as well as in the labora It is clear that there is a horizontal distribu tory supports the conviction that the approach tion of organisms, including Macrocystis integri is a useful and instructive one. Differences folia, in Queen Charlotte Strait which follows which sometimes appear striking or significant closely the pattern of salinity distribution in the 536 PACIFIC SCIENCE, Vol. XV, October 1961 Strait. In turn this reflects the circulation within SUMMARY the area. It would be premature to say that A detailed study of the horizontal and ver salinity is directly responsible for the observed tical distributions of marine benthonic organ distributions of all the organisms encountered. isms in Queen Charlotte Strait has been limited But one may say that the distribution reflects the so far to the more conspicuous algae and in dependence on high salinity water which is vertebrates encountered. The relationship of characteristic of the open ocean and exposure. these distributions to the salinity distribution In some instances it may be salinity that is a indicates that more intensive study of the flora direct causal factor. On the other hand, the open and fauna in this area, as well as elsewhere on coast has organisms associated witl). surf condi the coast, will provide further supporting evi tions. It has been suggested that the high dence indicating not only the effect of ocean oxygen requirement of certain organisms is met ographic variables on the distribution and pro only in such an exposed environment. However, duction of marine benthonic organisms but also the distributions and concentrations of oxygen the possibility of using such organisms as in in the sea in this area do not directly support dicators of oceanographic conditions both in this argument. The oxygen content of the waters time and space. The relationship of the vertical within the sheltered Strait is as high or higher distribution of some of the organisms to certain than in the surf in the exposed regions. This is tide levels indicates the response of the different particularly true in the central part of the Strait organisms to varying degrees of exposure and when there is a heavy bloom of phytoplankton, submergence. at which time the water may be supersaturated This oceanographic approach, both qualita with oxygen to as much as 175 per cent. Like tively and quantitatively, has given a broad wise, altQOlJgh it is known that many marine understanding of some of the possible factors algae have a high inorganic phosphate require which are likely to be responsible for the ob ment, there is no evidence that this nutrient is served distributions. However, before there can ever limiting in this area within the zone occu be a clear understanding and explanation of the pied by the benthonic algae. There is a great fundamental relationships between the organ need for further knowledge of the presence, dis isms and their environment in this region, as tribution, amounts, and availability of many well as an understanding of the interrelation more dissolved inorganic as well as organic sub ships and interaction among the organisms stances and perhaps even of growth substances. themselves, additional field observations, field experiments, and laboratory experiments must There is also need for a study of the quantitative be undertaken. aspects of removal, the rate of removal of such substances, and precise requirements for growth ACKNOWLEDGMENTS and reproduction in the micro-environment of the marine benthonic algae. The restriction of Financial assistance provided by the Defence certain organisms to surf conditions suggests Research Board of Canada in support of this that constant movement of water is required research (DRB 9520-14) over a period of sev to provide nutrients and gases which may be eral years is gratefully acknowledged. The author rapidly exhausted from the immediate or micro would also like to thank the Defence Research environment of the individual fixed alga, or in Board for its assistance in providing ship trans the case of the sessile marine invertebrates, such portation through the Pacific Naval Laboratory, as Mitella polymerus, to provide particulate food. Esquimalt, and the Royal Canadian Navy for It may be that lowering the concentration or the purpose of collecting data on several cruises. removal by dilution or by water movement of In addition, assistance provided by the Na some toxic substances which may accumulate tional Research Council, the Joint Committee on above a certain concentration in the micro Oceanography, the Fisheries Research Board of environment is just as significant as the avail Canada, and the University of British Columbia ability of others. is gratefully acknowledged. Benthonic Algae-SCAGEL 537 --- 1953b. Monthly Record Meteorological REFERENCES Observations in Canada, January-December, 1953. Dept. Transport Meteorological Div., ANON. 1944. Observations of Sea Water Tem Ottawa. perature and Salinity on the Pacific Coast of Canada. Fish. Res. Bd. Can., Nanaimo. Pacif. --- 1954. British Columbia Inlet cruises, Oceanogr. Group. Vol. 5, 1942, 1943. 1953. Inst. Oceanogr. Univ. Brit. 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