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Size-fractionated chlorophyll and primary productivity and nutrient distributions off the west coast.. Harris, Shannon Lee 2001

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SIZE-FRACTIONATED C H L O R O P H Y L L AND P R I M A R Y PRODUCTIVITY AND NUTRIENT DISTRIBUTIONS O F F T H E WEST COAST O F V A N C O U V E R ISLAND  by  S H A N N O N L E E HARRIS B.Sc, University of Manitoba, 1997  A THESIS SUBMITTED IN PARTIAL F U L F I L L M E N T OF THE REQUIREMENTS FOR THE D E G R E E OF  M A S T E R OF SCIENCE in THE F A C U L T Y OF G R A D U A T E STUDIES (Department of Earth and Ocean Sciences)  We accept this thesis as conforming to the r^qujrcd standard  THE UNIVERSITY OF BRITISH C O L U M B I A December 2001 © Shannon Lee Harris, 2001  In  presenting  degree  this  at the  thesis  in  University of  partial  fulfilment  of  of  department  this or  publication of  thesis for by  his  or  of  her  representatives.  ' E - a x V r v V OCL^C  DE-6 (2/88)  ^ D ^ C ^~e*TD\  for  an advanced  Library shall make  it  agree that permission for extensive  It  this thesis for financial gain shall not  The University of British Columbia Vancouver, Canada  Date  that the  scholarly purposes may be  permission.  Department  requirements  British Columbia, I agree  freely available for reference and study. I further copying  the  is  granted  by the  understood  that  head of copying  my or  be allowed without my written  ABSTRACT  Spatial and temporal variability of nutrients, chlorophyll and primary productivity off the west coast of Vancouver Island are not well studied. This study examined how dissolved nutrients and size-fractionated biomass and primary productivity vary in time and space and evaluated the relative contribution of >5 um size fraction of phytoplankton to total biomass and primary productivity. Size-fractionated  primary productivity,  and  physical,  chemical,  and  biological  characteristics were studied during spring, summer and fall cruises for 1997 and 1998. Studies were conducted at four transects extending across the continental shelf, perpendicular to the west coast of Vancouver Island.  Transects were over the La Perouse Bank, over Barkley  Canyon, off Estevan Point and off Brooks Peninsula. Physical, chemical and biological characteristics of shelf regions were distinct from beyond shelf regions and showed a strong cross-shelf gradient. The shelf region was characterized by lower temperature and lower salinity.  In addition, higher nitrate and silicic  acid, and higher chlorophyll and primary productivity were observed in the shelf region compared to the beyond shelf region. Variability was very high off the west coast of Vancouver Island; often the mean and the standard deviation were similar. This study was conducted during a strong E l Nino (1997/98) and La Nina (1998) event and interannual variation was evident. The mixed layer depth was shallower, nitrate, silicic acid and primary productivity were higher during E l Nino.  In contrast, phytoplankton biomass,  diatom abundance and the relative contribution of >5 um sized phytoplankton were higher during La Nifia. The >5 um sized phytoplankton were dominated by the diatoms Chaetoceros spp. and Leptocylindrus danicus and contributed substantially to the biomass (62%)  and primary  productivity (57%) off the west coast of Vancouver Island. The relative contribution was higher in shelf regions than in beyond shelf regions. This study clearly showed that the contribution of the >5 um size fraction was greatest at high biomass concentrations and high productivity rates supporting the idea that in order to reach high biomass and productivity, large cells are required.  ii  TABLE OF CONTENTS Abstract  ii  Table of Contents  iii  List of Tables  v  List of Figures  x  Acknowledgements  xx  G E N E R A L INTRODUCTION  1  Coastal Upwelling The importance of cell size ENSO-E1 Nino Southern Oscillation The west coast of Vancouver Island Physical oceanography Ecological dynamics Global Ocean Ecosystem Dynamics Program (GLOBEC) Thesis goals Thesis organization  2 5 7 8 8 15 19 19 20  C H A P T E R 1: Variability of physical, chemical and biological parameters off the west coast of Vancouver Island. Introduction Materials and Methods Physical measurements Chemical and biological measurements Statistical analysis of physical, chemical and biological data Results Physical characteristics Incident Irradiance Mixed layer parameters (Temperature, salinity and rj ) Chemical parameters Dissolved nutrient concentrations Biological parameters Total Chlorophyll Phytoplankton assemblages Discussion West coast of Vancouver Island Comparison with previous studies off the west coast of Vancouver Island Comparison with other upwelling regions Summary t  21 21 22 22 24 26 27 27 27 27 35 36 43 51 63 63 65 65 66 67  iii  CHAPTER 2:  Size-fractionated biomass and primary productivity off the west coast of Vancouver Island Introduction Materials and Methods Chemical and biological measurements Statistical analysis of chemical and biological data Results : Biological parameters Size-fractionated chlorophyll Total primary productivity Size-fractionated primary productivity Carbon assimilation rates Discussion West coast of Vancouver Island Comparison with previous studies off the west coast of Vancouver Island Comparison with other upwelling regions Summary  68 68 69 70 74 74 74 74 84 87 97 101 101 106 108 109  GENERAL DISCUSSION  110  FUTURE STUDIES  114  LITERATURE CITED  115  APPENDICES  125  A: B: C: D: E: F: G:  Station details 1999 raw data for disso lved nutrients 1999 raw data for chlorophyll 1999 raw data for primary productivity Sampling stations Cell volumes for conversion of cells L" to carbon L" Incident surface irradiance during primary productivity measurements 1  125 126 142 150 151 161 164  1  H:  Vertical profiles of temperature, salinity and or  170  I: J: K: L:  Vertical profiles of dissolved nutrients Chlorophyll values Vertical profiles of size-fractionated chlorophyll Depth profiles of the contribution of >5 um fraction to chlorophyll and productivity Primary productivity data  177 183 184  M:  186 194  iv  LIST O F T A B L E S  Table 1.1  Table 1.2  Mixed layer parameters for stations occupied during 1997 cruises. Monthly mean, yearly mean ± 1 S.D. and yearly coefficient of variation, (C.V.,%) are given for the shelf and beyond shelf region. Temperature (°C), salinity, and density for the mixed layer were calculated as the mean value from the surface to the calculated mixed layer depth. Dashed line indicates that data are not available  33  Mixed layer parameters for stations occupied during 1998 cruises. Monthly mean, yearly mean ± 1 S.D. and yearly coefficient of variation (C.V., %) are given for the shelf and beyond shelf region. Temperature (°C), salinity, and density for the mixed layer were calculated as the mean value from the surface to the calculated mixed layer depth. 34  Table 1.3  Table 1.4.  Table 1.5  Table 1.6  Table 1.7  Mean nutrient concentrations (0-10m) (pM) for WCVI during each cruise in 1997 and 1998. Values are the mean of all stations during each cruise. Yearly mean ±1 S.D. and coefficient of variation (C.V., %) are included  36  Mean surface (0-10 m) nitrate, phosphate and silicic acid concentrations (pM) in 1997 and 1998 for shelf and beyond shelf stations of La Perouse Bank, Barkley Canyon, Estevan Point, Brooks Peninsula off the west coast of Vancouver Island. The mean ± 1S;D. and coefficient of variation for each year are given. The number of samples (n) for each transect is given. N D indicates nutrient concentration was not detectable. (-) indicates information not available. * indicates a significant difference between shelf and beyond shelf regions was found at p<0.05 level and a indicates a significant difference between region was found at p<0.01 level  42  Mean chlorophyll (100-1% surface light) ± 1 S.D. (mg chl m") for WCVI during each cruise in 1997 and 1998. Values are the mean of all stations during each cruise. Yearly mean ±1 S.D. and coefficient of variation (CV, %) are included  43  2  Mean integrated chlorophyll ± 1 S.D. (Chl; mg chl m") and coefficient of variation (CV; %) for 1997 and 1998 for shelf and beyond shelf stations along L a Perouse Bank, Barkley Canyon, Estevan Point and Brooks Peninsula off the west coast of Vancouver Island. Mean for each cruise and each year are given 2  List of diatoms identified from samples collected off the west coast of Vancouver Island from May 1997 to October 1998. X in 1997/1998 column signifies the diatom was observed in either April/May, July or October  50  58  v  List of autotrophic flagellates identified from samples collected off the west coast of Vancouver Island from May 1997 to October 1998. X signifies species was observed in either April/May, July or October. ...  59  Abundance of diatoms, nanoflagellates, autotrophic and heterotrophic dinoflagellates during April, July and October 1997 off the west coast of Vancouver Island. LP=La Perouse Bank BC=Barkley Canyon, EP=Estevan Point, BP=Brooks Peninsula. See Figure 1.1 for location of transects. (—) indicates no sample taken, * indicates the most abundant group  60  Abundance of diatoms, nanoflagellates, autotrophic dinoflagellates and heterotrophic dinoflagellates at 55% surface light depth during May, July and October 1998 off the west coast of Vancouver Island. LP=La Perouse Bank BC=Barkley Canyon, EP=Estevan Point, BP=Brooks Peninsula. See Figure 1.1 for location of transects. * = most abundant group  61  Summary of characteristics of shelf and beyond shelf regions off the west coast of Vancouver Island. Values are for 1997 and 1998. Units for parameters below are: M L , meters; temperature, °C, nitrate, uM; chlorophyll, mg chl i n ; C V , %)  62  Summary of characteristics of shelf and beyond shelf regions off the west coast of Vancouver Island. Values are for 1997 and 1998. Units for parameters below are: M L , meters; temperature, °C, nitrate, uM; chlorophyll, mg m" ; PP, mg C m" d" ; carbon assimilation rates, mgCmgchr h ;CV,%)  100  Cruise dates and season for 3 cruises during 1997, 1998 and 1999. Transition date is the date the prevailing wind shifted for the season and it was calculated using Thomson & Ware's wind velocity index (R. Thomson pers. comm.). Seasonal classification is based on the prevailing winds. (-) denotes information is not available  125  Table B. 1 N0 ", HP0 " and Si(OH) in May 1999 off the west coast of Vancouver Island. Dashed line (-) indicates that data point is not available. (-) indicates information is not available; b-10 indicates water sample was collected at 10 m off the bottom  126  Table 1.8  Table 1.9  Table 1.10  Table 1.11  2  Table 2.1  2  1  Table A . l  3  Table B.2  4  2  1  _,  4  N 0 \ HP0 " and Si(OH) in July 1999 off the west coast of Vancouver Island. A l l samples were collected and analyzed as outlined in methods section of Chapter 1. (-) indicates information is not available. 3  4  4  132 Table B.3  N0 ", HP0 " and Si(OH) for October 1999 on the west coast of Vancouver Island. All samples were collected and analyzed as outlined in methods section of Chapter 1. Dashed line (-) indicates that information is not available 2  3  4  4  136  vi  Chlorophyll a (mg m") in May 1999 off the west coast of Vancouver Island. A l l samples were filtered onto 0.7 pm glass fiber filters unless otherwise indicated by * which were filtered onto 5.0 pm polycarbonate filters. (-) indicates data not available  141  Table C.2  Chlorophyll a (mg m") in July 1999 off the west coast of Vancouver Island. All samples were filtered onto 0.7 pm glass fiber filters unless otherwise indicated by ^ which were filtered onto a 5.0 pm polycarbonate filters  145  Table C.3  Chlorophyll a (mg m") in July 1999 off the west coast of Vancouver Island. A l l samples were filtered onto 0.7 pm glass fiber filters unless otherwise indicated by * which were filtered onto a 5.0 pm polycarbonate filters  147  Integrated (100-1% surface light) daily primary productivity (g C m" d") in 1999 for July and October off the west coast of Vancouver Island  150  Location and water depth of stations occupied during 08-24 April 1997 (Cruise ID#9707) off the west coast of Vancouver Island. Under the station column, B indicates L a Perouse Bank, C indicates Barkley Canyon, D indicates D Line, G indicates Estevan Point, BP indicates Brooks Peninsula and CS indicates Cape Scott transects. See Figure 1.1 for location of transects  152  Location and water depth of stations occupied during 14-28 July 1997 (Cruise ID#9713) off the west coast of Vancouver Island. Under the station column, A indicates Juan de Fuca canyon, B indicates La Perouse Bank, C indicates Barkley Canyon, G indicates Estevan Point and BP indicates Brooks Peninsula transect. See Figure 1.1 for location of transects  153  Tabled  Table D . l  3  3  3  2  Table E . l  Table E.2  1  Table E.3. Location and water depth of stations occupied during 20-27 October 1997 (Cruise ID#9737) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, D indicates Line D, G indicates Estevan Point and BP indicates Brooks Peninsula. See Figure 1.1 for location of transects. ...  154  Location and water depth of stations occupied during 11-25 May 1998 cruise (Cruise ID#9810) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, G indicates Estevan Point, H indicated H Line, J indicates J Line, BP indicates Brooks Peninsula and CS indicates Cape Scott transect. See Figure 1.1 for location of transects  155  Table E.4  vii  Table E.5  Table E.6  Table E.7  Table E.8  Table E.9  Table F.l  Table F.2  Table J.l  Location and water depth of stations occupied during 14-26 July 1998 (Cruise ID#9823) off the west coast of Vancouver Island. Under the station column, A indicates Juan de Fuca canyon, B indicates La Perouse Bank, C indicates Barkley Canyon, D indicates D Line, G indicates Estevan Point, BP indicates Brooks Peninsula and E R indicated Endeavor Ridge transect. See Figure 1.1 for location of transects  156  Table E.6 Location and water depth of stations occupied during the 0516 October 1998 cruise (Cruise ID#9836) off the west coast of Vancouver Island. Under the station column, indicates La Perouse Bank, C indicates Barkley Canyon, G indicates Estevan Point, BP indicates Brooks Peninsula and CS indicated Cape Scott. See Figure 1.1 for location of transects  157  Location and water depth of stations occupied during May 1999 (Cruise ID#9911) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, D indicates D Line, G indicates Estevan Point, J indicates J Line, BP indicates Brooks Peninsula and CS indicates Cape Scott. See Figure 1.1 for location of transects  158  Location and water depth of stations occupied during July 1999 (Cruise ID#9928) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, G indicates Estevan Point and BP indicates Brooks Peninsula. See Figure 1.1 for location of transects  159  Location and water depth of stations occupied during October 1999 (Cruise ID#9935) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, D indicates D Line, G indicates Estevan Point, J indicates J Line, BP indicates Brooks Peninsula and CS indicates Cape Scott transect. See Figure 1.1 for location of transects  160  Cell volume and carbon per cell pf the diatom species observed off the west coast of Vancouver Island during 1997 and 1998. P indicates pennate diatoms and C indicates centric diatoms  162  Cell volume and carbon per cell for other phytoplankton species except diatoms) observed off the west coast of Vancouver Island during 1997 and 1998. P indicates autotrophic and H indicates heterotrophic nutrition  153  Size-fractionated chlorophyll (surface to 1% light depth +1 SD) and the relative contribution of the >5um fraction to total chlorophyll for: the west coast of Vancouver Island, the shelf regions and the beyond shelf regions off the west coast of Vancouver Island during 1997 and 1998. Means for the 1997, 1998 and all cruises are given along with the minimum and the maximum  183 viii  Table M . l  Table M.2  Total primary productivity ±1 S.D. (surface to 1% light depth) for the west coast of Vancouver Island (WCVI) and the shelf and beyond shelf region in 1997 and 1998. Means for 1997, 1998, and for all cruises are given along with the minimum and maximum for all cruises. Numbers in brackets are the number of stations sampled  194  Size-fractioned primary productivity ±1 S.D. and relative contribution of >5 pm fraction to total productivity off the west coast of Vancouver Island, the shelf and beyond shelf regions in 1997 and 1998. Means for the 1997, 1998, and all cruises are given with the minimum and the maximum  195  ix  LIST O F F I G U R E S  Figure 1  Figure 2 Figure 1.1  Figure 1.2  Figure 1.3  Figure 1.4  Idealized cycle of nutrient and carbon phytoplankton processes in an upwelling region (adapted from Wilkerson and Dugdale, 1987). Solid arrows represent nutrient rich upwelling water mass and dashed arrows represent a nutrient deplete upwelling plume water mass. pN represents nitrate uptake rate and pC represents carbon uptake rates. Day # on top of each zone estimates the days since upwelling was initiated. The sun represents solar heating necessary for stabilization of water column Surface circulation off the west coast of Vancouver Island in summer (Thomson et al., 1989). Dashed line marks the 200 m contour Location of transects off the west coast of Vancouver Island. Dashed line delineates the 200 m contour. A=Juan de fuca Canyon (Line A), B=La Perouse Bank (Line B), C=Barkley Canyon (Line C), D=D line, G=Estevan Point (Line G), H=H Line, J=J line, BP=Brooks Peninsula (BP Line) and CS=Cape Scott (CS Line). Transects with * will be discussed in this chapter Incident surface irradiance for A) April 1997, B) July 1997, C) October 1997, D) May 1998 and E) July 1998. Station labels are placed on peak to day of primary productivity measurements. See Appendix E for location of stations. Irradiance data not available for October 1998 Surface (0-10m) nitrate concentration for all cruises in 1997 and 1998 along transects on La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Shaded area represents the shelf region; solid lines and closed symbols are for 1997 and dashed lines and open symbols are for 1998. No data are available for La Perouse Bank in 1998. The distance offshore increases as the station number increases Surface (0-10m) phosphate concentration for all cruises in 1997 and 1998 along transects on La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Shaded area represents the shelf region; solid lines and closed symbols are for 1997 and dashed lines and open symbols are for 1998. No data are available for La Perouse Bank in 1998. The distance offshore increases as the station number increases  4 ^  23  29  38  39  x  Figure 1.5  Figure 1.6  Figure 1.7  Figure 1.8  Figure 1.9  Figure 1.10  Figure 1.11  Surface (0-10m) phosphate concentration for all cruises in 1997 and 1998 along transects on La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Shaded area represents the shelf region; solid lines and closed symbols are for 1997 and dashed lines and open symbols are for 1998. No data are available for La Perouse Bank in 1998. The distance offshore increases as the station number increases  40  Total chlorophyll ±1S.D. off the west coast of Vancouver Island in 1997 and 1998. Values are the mean of shelf and beyond shelf stations during each cruise  44  Interannual mean and annual means of total chlorophyll ±1 S.D. of the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the mean of all shelf and beyond shelf stations sampled. Numbers above each bar are the total number of stations for each mean  45  Seasonal variability of total chlorophyll ± 1 S.D. of the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the mean of all shelf and beyond shelf stations during each cruise. Numbers above each bar=the total number of stations  ^6  Integrated chlorophyll (mg chl m") for all cruises in 1997 and 1998 along transects on La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) on the west coast of Vancouver Island. Shaded area represents shelf region; solid lines and closed symbols are for 1997 and dashed lines and open symbols are for 1998. No data are available for La Perouse Bank in 1998. The distance offshore increases as the station number increases.  47  Total chlorophyll of the shelf and the beyond shelf regions of the La Perouse Bank, Barkley Canyon, Estevan Point and Brooks Peninsula transect off the west coast of Vancouver Island in 1997 and 1998. A April, M=May, J=July and 0=October  49  Interannual mean and annual means of total cell abundance and phytoplankton biomass ± 1 S.D. for the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf values are the mean of all shelf stations sampled and beyond shelf values are the mean of all beyond shelf stations  52  2  xi  Figure 1.12  Figure 1.13  Seasonal variability of total cell abundance and phytoplankton biomass ± 1 S.D. for the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf means are the average of all shelf stations sampled during each cruise and beyond shelf values are the man of all beyond shelf stations during each cruise  53  Contribution of each phytoplankton group to A) total cell abundance, and B) total phytoplankton biomass (\xg L )of the shelf and beyond shelf regions off the west coast of Vancouver Island for 1997, 1998 and the 2 yr mean  54  Contribution of each phytoplankton group to total cell abundance and biomass of: A) shelf and B) beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Groups contributing <2% were not included  56  Location of study area off the west coast of Vancouver Island. Dashed line is the 200 m contour. The box delineates the study area. CI, LC4, LG3, BP2 are shelf stations and LB16, LC9, LG9, BP7 are beyond shelf stations. CI and L B 16 =La Perouse Bank transect, LC4 and LC9 = Barkley Canyon transect, LG3 and LG6 = Estevan Point transect and BP2 and BP7 = Brooks Peninsula transect  70  Interannual mean and annual means of size-fractionated chlorophyll ± 1 S.D. for the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf values are the mean of all shelf and beyond shelf stations, respectively. Numbers of the top of each panel represent the percentage of the total chlorophyll that was accounted for by the > 5 um fraction  76  Cruise means of <5 um and > 5 um chlorophyll ± 1 S.D. for the west coast of Vancouver Island in 1997 and 1998. Values are the mean of shelf and beyond shelf stations during each cruise. Numbers on the top of each panel represent the percentage of the total chlorophyll that was accounted for by the >5 um fraction  77  Interannual mean and annual means of size-fractionated chlorophyll ± 1 S.D. for the shelf and beyond shelf region off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the mean of all shelf and beyond shelf stations sampled. Numbers on the top of each panel represent the percentage of the total chlorophyll that was accounted for by the >5 um fraction  78  _1  Figure 1.14  Figure 2.0  Figure 2.1  Figure 2.2  Figure 2.3  xii  Figure 2.4  Figure 2.5  Figure 2.6  Figure 2.7  Figure 2.8  Figure 2.9  Figure 2.10  Mean of size-fractionated chlorophyll ± 1 S.D. for the shelf and the beyond shelf region off the west coast of Vancouver Island in April (n=2), July (n=3) and October 1997 (n=4) and May (n=4), July (n=4) and October (n=4) 1998. Shelf and beyond shelf values are the mean of all shelf and beyond shelf stations. Numbers on the top of each panel represents the percentage of the total chlorophyll that was accounted for by the >5 pm fraction  80  Relative contribution of < 5.0 pm size fraction and > 5.0 pm size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during July 1997. Relative contribution of > 5.0 pm fraction to depth integrated chlorophyll is in right hand corner of each graph. Brooks Peninsula shelf and Barkley Canyon beyond shelf stations were not sampled  82  Relative contribution of < 5.0 pm size fraction and > 5.0 pm size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during July 1998. Relative contribution of > 5.0 pm fraction to depth integrated chlorophyll is in right hand corner of each graph. Brooks Peninsula shelf and Barkley Canyon beyond shelf stations were not sampled  83  Daily mean primary productivity ± 1 S.D. off the west coast of Vancouver Island for April, July and October 1997 and May, July and October 1998. Values are the mean of all stations sampled during each cruise  84  Mean primary productivity ± 1 S.D. (g C m" d") of the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the average of all shelf and beyond shelf stations sampled  85  Total mean primary productivity of shelf and beyond shelf regions of the La Perouse Bank, Barkley Canyon, Estevan Point and Brooks Peninsula transect off the west coast of Vancouver Island. One station was sampled for each of the shelf and the beyond shelf region of each transect. A/M=April and May, J=July and 0=October  87  Interannual mean and annual means of size-fractionated primary productivity ± 1 S.D. for the west coast of Vancouver Island in 1997 and 1998. Number on the top of each panel represent the percentage of the total primary productivity that was accounted for by the >5 pm fraction  88  2  1  xiii  Figure 2.11  Figure 2.12  Figure 2.13  Figure 2.14  Figure 2.15  Figure 2.16  Figure 2.17  Cruise means of <5 jam and > 5 um primary productivity ± 1 S.D. for the west coast of Vancouver Island in 1997 and 1998. Values are the mean of shelf and beyond shelf stations during each cruise. Numbers on the top of each panel represent thepercentage of the total primary productivity that was accounted for by the >5 um fraction  89  Interannual mean and annual means of size-fractionated primary productivity ± 1 S.D. for the shelf and beyond shelf region off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the mean of all shelf and beyond shelf stations sampled. Numbers on the top of each panel represent the percentage of the total primary productivity that was accounted for by the >5 um fraction  91  Mean of size-fractionated primary productivity ± 1 S.D. for the shelf and the beyond shelf region off the west coast of Vancouver Island in April (n=2), July (n=3) and October 1997 (n=4) and May (n=4), July (n=4) and October (n=4) 1998. Shelf and beyond shelf values are the mean of all shelf and beyond shelf stations. Numbers on the top of each panel represents the percentage of the total primary productivity that was accounted for by the >5 um fraction  93  Percent contribution of >5 um size fraction relative to: A) total chlorophyll and B) total primary productivity off the west coast of Vancouver Island. Values are for all stations in 1997 and 1998 in the shelf and the beyond shelf region  94  Relative contribution of < 5 um size fraction and > 5 um size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during July 1997. Relative contribution of > 5 um fraction to depth integrated primary productivity is in the right hand corner of each graph. Brooks Peninsula shelf and Barkley Canyon beyond shelf stations were not sampled  95  Relative contribution of < 5 um size fraction and > 5 um size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during July 1998. Relative contribution of > 5 um fraction to depth integrated primary productivity is in the right hand corner of each graph. Brooks Peninsula shelf and Barkley Canyon beyond shelf stations were not sampled  96  Carbon assimilation rates ± 1 S.D. off the west coast of Vancouver Island in 1997 and 1998. Values are the mean of all stations sampled during each cruise  97  Figure 2.18  Figure 2.19  Figure G. 1  Figure G.2  Figure G.3  Figure G.4  Carbon assimilation rates ± 1 S.D. of the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf means are the average of all shelf stations sampled during each cruise and beyond shelf values are the mean of all beyond shelf stations during each cruise  98  Carbon assimilation rates of the shelf and the beyond shelf region of La Perouse Bank, Barkley Canyon, Estevan Point and Brooks Peninsula transect off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond means are the average of all shelf and beyond shelf stations, respectively  99  Incident surface irradiance for primary productivity measurements during April 1997 off the west coast of Vancouver Island. Incubation period was 24 hours. Primary productivity was not measured at the beyond shelf station of the Brooks Peninsula transect. See Figure 2.1 for location of transects  ^5  Incident surface irradiance during primary productivity measurements in July 1997 off the west coast of Vancouver Island. The shaded region demarcates the incubation period. Primary productivity was not measured at the shelf station of the Brooks Peninsula transect or the beyond shelf station of the Barkley Canyon transect. Number in the right hand corner of each plot is the percentage that the incubation period represented of the total daily irradiance. See Figure 2.1 for location of transects  166  Incident surface irradiance during primary productivity measurements in October 1997 off the west coast of Vancouver Island. The shaded region demarcates the incubation period. Primary productivity was not measured in the beyond shelf region of the Estevan Point and La Perouse Bank transects. Number in right hand corner of each plot is the percentage that the incubation period represented of the total daily irradiance. Note scale change relative to Figure G . l and G.2. See Figure 2.1 for location of transects Incident surface irradiance during primary productivity measurements in May 1998 off the west coast of Vancouver Island. The shaded region demarcates the incubation period. Number in the right hand corner of each plot is the percentage that the incubation period represented of the total daily irradiance. See Figure 2.1 for location of transects  167  168  xv  Figure G.5  Figure H . l  Figure H.2  Figure H.3  Figure H.4  Figure H.5  Incident surface irradiance during primary productivity measurements during July 1998 off the west coast of Vancouver Island. The shaded region demarcates the incubation period. Number in the right hand corner of each plot is the percentage that the light during the incubation period represented of the total daily irradiance. See Figure 2.1 for location of transects Vertical profiles of density, salinity and temperature in April 1997 at a shelf and beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP) and Brooks Peninsula (BP) off the west coast of Vancouver Island. Data for deep stations only shown for 0 - 200 m depth. See Figure 1.1 for location of transects. Data not available for beyond shelf station on the Brooks Peninsula transect. B indicates beyond shelf region Vertical profiles of density, salinity and temperature in July 1997 at a shelf and beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP) and Brooks Peninsula (BP) off the west coast of Vancouver Island. Data set for deep stations only shown for 0-200 m depth. Data set not available for beyond shelf station on the Barkley Canyon transect. See Figure 1.1 for location of transects. B indicates beyond shelf region Vertical profiles of density, salinity and temperature in October 1997 at a shelf and a beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Esteven Point (EP) and Brooks Peninsula (BP) off the west coast of Vancouver Island. Data for deep stations only shown for 0-200 m depth. See Figure 1.1 for location of transects. Data not available for the beyond shelf station of the La Perouse Bank transect. B indicates beyond shelf region Vertical profiles of density, salinity and temperature in May 1998 at a shelf and a beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP) and Brooks Peninsula (BP) off the west coast of Vancouver Island. Data for deep stations only shown for 0-200 m. Beyond shelf stations for the Estevan Point and Brooks Peninsula transect only sampled to 100 m depth. See Figure 1.1 for location of transects. B indicates beyond shelf region Vertical profiles of density, salinity and temperature in July 1998 at a shelf and beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP), and Brook Peninsula (BP) off the west coast of Vancouver Island. Data for deep stations shown for 0200 m depth. The beyond shelf station of La Perouse Bank was only sampled to 100 m depth. See Figure 1.1 for location of transects. B indicates beyond shelf region  ^  171  172  173  174  175  xvi  Figure H.6  Figure 1.1  Figure 1.2  Figure 1.3  Figure 1.4  Figure 1.5  Figure 1.6  Vertical profiles of density, salinity and temperature in October 1998 at a shelf and a beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP) and Brooks Peninsula (BP) on the west coast of Vancouver Island. Data set for deep stations shown for 0-200 m depth. See Figure 1.1 for location of transects. B indicates beyond shelf region  ^  Vertical profiles of nitrate, phosphate and silicic acid (uM) in April 1997 at all stations along transects of La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Station name is located at bottom of each graph  1^7  Vertical profiles of nitrate, phosphate, and silicic acid (uM) in July 1997 at all stations along transects of La Perouse Bank (Line B), Barkley Canyon, (Line C), Estevan Point (Line G), and Brooks Peninsula (Line BP) on the west coast of Vancouver Island. Station name is located at the bottom of each graph  1  /  0  Vertical profiles of nitrate, phosphate and silicic acid (uM) in October 1997 at all stations along transects of La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) on the west coast of Vancouver Island. Station name is located at the bottom of each graph  1^9  Vertical profiles of nitrate, phosphate and silicic acid (uM) in May 1998 at all stations along transects of Barkley Canyon (Line C), BEstevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Station name is located at the bottom of each graph. No data are available for the La Perouse Bank transect  180  Vertical profiles of nitrate, phosphate and silicic acid (uM) in July 1998 at all stations along transects of Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Station name is located at the bottom of each graph. No data are available for the La Perouse Bank transect  181  Vertical profiles of nitrate, phosphate and silicic acid (uM) in October 1998 at all stations along transects of Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) on the west coast of Vancouver Island. Station name is located at the bottom of each graph. No data are available for the La Perouse Bank transect  182  xvii  Figure K. 1  Figure K.2  Figure L . l  Figure L.2  Figure L.3  Figure L.4  Figure L.5  Vertical profiles of size-fractionated chlorophyll a for April, July, and October 1997 at shelf and beyond shelf stations along transects of La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP off the west coast of Vancouver Island. Data are not available for the > 5.0 pm fraction for station C I , BP2, B16 and BP7. Samples taken down to 1% surface irradiation. See Figure 2.1 for location of stations  184  Vertical profiles of size-fractionated chlorophyll a May, July, and October 1998 at shelf and beyond shelf stations along transects of La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Samples taken down to 1% light depth. See Figure 2.1 for location of stations  185  Relative contribution of <5 pm size fraction and >5 pm size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during April 1997. The relative contribution of > 5 pm fraction to depth integrated chlorophyll is given in top right hand corner of each plot  186  Relative contribution of <5 pm size fraction and >5 pm size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during Oct. 1997.Relative contribution of >5 pm fraction of depth integrated chlorophyll in right hand corner of each graph  187  Relative contribution of <5 pm size fraction and >5 pm size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during May 1998. Relative contribution of >5 pm fraction depth integrated chlorophyll in right hand corner of each graph  188  Relative contribution of <5 pm size fraction and >5 pm size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during Oct. 1998. Relative contribution of >5 pm fraction depth integrated chlorophyll in right hand corner of each graph. ns=no sample  189  Relative contribution of <5 pm size fraction and >5 pm size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during April 1997. Relative contribution of >5 pm fraction to depth integrated primary productivity is in the right hand corner of each graph  190  xviii  Figure L.6  Figure L.7  Figure L.8  Relative contribution of < 5.0 um size fraction and > 5.0 um size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during Oct. 1997. Relative contribution of > 5.0 um fraction to depth integrated primary productivity in right hand corner of each graph  191  Relative contribution of < 5 um size fraction and > 5 um size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during May 1998. Relative contribution of depth integrated primary productivity in right hand corner of each graph  1^  Relative contribution of < 5 um size fraction and > 5 um size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during Oct. 1998. Relative contribution of > 5 um fraction to depth integrated primary productivity in right hand corner of each graph  1^3  xix  ACKNOWLEDGEMENTS  I am indebted to my research supervisor, Dr. Paul J. Harrison for many many things, too many to mention specifically. He was always supportive and incredibly positive at all stages of my degree. I especially value his assistance with pulling this thesis together. It was truly an honor to work with him. Thanks to Dr. Lewis for sharing your knowledge and enthusiasm of zooplankton ecology but also for always providing advice when needed. I thank my thesis committee of Dr. Lewis and Dr. Taylor for always being helpful and approachable. Many thanks to Dr. Susan Allen for your enthusiasm and support at U B C and at sea. My thanks to Dr. Ian Perry for his assistance in the field, for his encouragement at all stages of this thesis and his commitment to the G L O B E C program. I am indebted to Dr. Roger Pieters, without his incredible flexibility and I know this thesis would have never been completed. Without the field assistance of many people, this research would not have been possible. My thanks are extended to B. Beaith, E . Bornhold, B. Ferris, M . Henry, L . Lee, J. McKay, J. Needoba and Carine Vmdeirinho for your 'around the clock commitment'. My deepest gratitude to Hugh McLean who never missed a Globec adventure. The Oceanography department was lucky to have such a committed and happy sea-going technician-we will miss you. I owe my gratitude to many staff at the Institute of Ocean Sciences including B. Minkley, S. Romaine, D. Moore, D. Thule and D. Yelland for their assistance in the field. Special thanks to Doug Yelland for providing me with endless streams of data. My appreciation is extend to Janet Bardwell-Clarke who provided helpful advice and assistance concerning our troublesome technicon. Without Janet's assistance and wisdom, I am not sure if our technicon would ever have run again. Many thanks are extended to all the officers and crew of the C.S.S. "John P. TullyT. Members of the Paul Harrison Lab have been very helpful. Thanks to Ming Xin Guo and H. Toews for their help with analyzing the nutrient samples. Many thanks to Lauren Ross for her many encouraging e-mails. Special thanks to Robert Strzepek for the late night coffee and ice cream breaks. His company when the hallway was empty and the sky was dark was always appreciated and will be missed dearly. Many thanks are extended to D. Varela who provided helpful advice in the field and the lab. My gratitude is extended to Ramzi Mirshak, who not only was a kind friend but help me with G M T . I greatly appreciate the professionalism of Rowan Haigh who did the phytoplankton taxonomy analysis. Unfortunately, I do not have the skill and knowledge necessary for the fine art of taxonomic analysis. Rowan kindly analyzed all the taxonomy samples and provided the information necessary for data processing. Thanks for his patience with my never-ending questions. Financial support was provided by the Natural Sciences and Engineering Research Council (through the Canadian Global Ocean Ecosystem Dynamics Program) and by Fisheries and Oceans Canada. Thanks to Tracy Laval for your friendship in this new city. Special thanks are extended to Russell Hobbs for your love and support. Your guidance and encouragement is always valued. Although my father has passed away, the lessons he taught me proved invaluable at every stage of this work. My love for you is forever stored deeply in my heart. I miss you dearly and I wish you could share this accomplishment of mine. Finally, I can not find the words to express my respect for my mother, Marg. Her strength, support, love, encouragement and advice were always a phone call away. Only 5 years ago I witnessed my Mom triumph when faced with adversity and what I learned from watching her is what allowed me to complete this thesis. She is incredibly strong and caring in the same breath, a quality I am striving for. I hope you are as proud of me as I am of you. xx  GENERAL INTRODUCTION  Traditionally the growth rate of phytoplankton in the ocean has been regarded to be limited by nitrogen, but this view has recently been challenged by evidence that indicates during particular periods and in certain regions other nutrients or trace metals may also limit growth. For example, the equatorial Pacific may be phosphorus limited (Smith, 1984; Harrison et al., 1990; Karl et al., 1997). New evidence suggests that silicate may regulate primary production in upwelling regions of the equatorial Pacific (Dugdale and Wilkerson, 1998), the Southern Ocean (Nelson and Truguer, 1992) and in the Gulf Stream (Nelson and Brzezninski, 1990). The growth of phytoplankton in certain areas called high nitrate, low chlorophyll (HNCL) regions are now known to be limited by the availability of iron. Oceanic regions such as the equatorial Pacific (Cole et al,  1996), the Southern Ocean (Boyd et al, 2001), the N E subarctic Pacific  (Martin and Fitzwater, 1988; Boyd et al, 1996) and California coastal upwelling regions such as Big Sur (Hutchins and Bruland, 1998) are now known to be iron limited. Carbon limitation of photosynthesis has been considered rare because of the high availability of inorganic carbon in the oceans, but some researchers suggest that carbon may control phytoplankton growth and phytoplankton community structure (Tortell, 2000). Vertical mixing of phytoplankton in the water column is very important as mixing controls the light and nutrient regimes to which the phytoplankton are exposed. Lack of mixing may cause photoinhibition of phytoplankton if the cells are exposed to very high irradiances (Neale, 1987). Conversely, phytoplankton may suffer an 'energy crisis' when mixed from high to low light (Falkowski and LaRoche, 1991). During this crisis the cell shifts its focus from the synthesis of molecules necessary for growth to the production of light harvesting complexes that allow for adaptation and growth to low light environments. This shift is called photoacclimation  1  and it causes a temporary decrease in phytoplankton growth rate. It is important that the depth of the mixed layer is no deeper than the critical depth. The critical depth is defined as the depth above which photosynthetic production for the water column equals the respiration of the water column per unit surface area.  If mixing is deeper than the critical depth, phytoplankton  respiration will exceed photosynthesis and there will be no growth. Deep mixed layers can cause a depression in photosynthesis (Huntsman and Barber, 1977) and nitrate uptake rates (Zimmerman et al., 1987) because the phytoplankton are maintained in light-limited conditions.  Coastal Upwelling The aquatic environment accounts for approximately 40% of the total photosynthesis on earth (Falkowski, 1994) and much of this photosynthesis occurs in coastal upwelling regions which occupy only 0.1% of the total ocean area. 95% percent of the world's fisheries occurs within 320 km of the shore (Thurman and Trujillo, 1999). They are fertile regions relative to oceanic regions and they are characterized by relatively high levels of biomass and phytoplankton (Barber and Smith, 1981).  Primary productivity rates >300 g C m" y" are 2  1  common in upwelling regions compared to <50 g C mf y" in oceanic regions (Ryther, 1969). 2  1  Coastal upwelling occurs along the western margins of continents in both the southern and northern hemispheres.  When northwest winds transport water offshore in the Northern  Hemisphere, cold nutrient-rich subsurface water is upwelled to the surface to replace the water advected offshore.  In the absence of upwelling, nutrient-rich subsurface water is vertically  separated from the nutrient-poor surface waters by a density gradient that prevents mixing. Upwelling is a physical process that provides an injection of new nutrients and a "seed" phytoplankton population to the euphotic zone and in the presence of light, rapid growth can  2  occur.  Following an upwelling event, phytoplankton respond to high nutrients and light  availability by a series of specific physiological transition stages that occur along the axis of the upwelling plume.  Jones et al., (1983) characterized a series of 4 idealized zones that are  summarized by Wilkerson and Dugdale (1987) as a 'conveyor phytoplankton processes in an upwelling zone.  belt' of nutrient and carbon  These zones are summarized in Figure 1. In  Zone 1, upwelling occurs and phytoplankton nutrient and carbon uptake rates are "shifted down" because the cells are taking up nutrients and growing considerably slower than their maximal rates. This lag is due to cells becoming acclimated to both high nutrient concentrations and near-surface light intensities and/or a period of biological conditioning may be required because of inherent toxic factors (Barber and Huntsman, 1977). Zone 2 is downstream of the upwelling plume where cells have acclimated to the new conditions and uptake rates increase or are "shifted up". Initially, nitrogen uptake rates are elevated, but nitrogen uptake is quickly followed by increases in carbon uptake and growth rate.  In Zone 3, phytoplankton rate  processes are functioning at their maximal rates and nutrients will quickly be depleted and phytoplankton biomass will increase.  Finally in Zone 4, nutrient concentrations are exhausted  by the fast-growing "shifted-up" phytoplankton cells and consequently cells undergo a "shift down" in rate processes. Following nutrient exhaustion, phytoplankton rapidly sink out of the photic zone (Bienfang and Ziemann, 1992). The time and space domain where these sequences of physiological changes take place is only 8-10 days within 30-60 km of the coast (Maclsaac et al., 1985; Zimmerman et al., 1987).  3  Figure 1 Idealized cycle of nutrient and carbon phytoplankton processes in an upwelling region (adapted from Wilkerson and Dugdale, 1987). Solid arrows represent a nutrient rich upwelling water mass and all dashed arrows represent the nutrient deplete upwelled plume. pN represents nitrate uptake rate and pC represents carbon uptake rates. Day # on top of each zone estimates the days since upwelling was initiated. The sun represents solar heating necessary for stabilization of water column. 4  Globally, picoplankton (0.2-2.0 pm) account for more than 80% of primary productivity in oligotrophic waters (Stockner and Antia, 1986), but in upwelling regions Malone (1980) has shown that phytoplankton biomass is characterized by episodic production of chain-forming diatoms or large-celled diatoms or dinoflagellates.  There is increasing evidence from N tracer 1 5  experiments that show that the larger size fractions primarily use new nitrogen sources (NO3") and nanoplankton and picoplankton primarily use regenerated sources such as ammonium and urea (Probyn, 1985; Probyn et al, 1990). Often diatoms develop in part due to the rapid uptake of nutrients and large vacuoles that allow for storage of nutrients (Turpin and Harrison, 1979)  The importance of cell size The size of organisms at any trophic level can be a determining factor in the length of the food chain, the ecological efficiency of energy transfer, and the yield of organisms living at the highest trophic level (Ryther, 1969). Ryther (1969) showed that the yield of fish from a marine ecosystem dominated by large-celled phytoplankton was greater than from areas dominated by small-celled phytoplankton. Cushing (1989) suggested that food chains based on large phytoplankton are ecologically more efficient than those based on small phytoplankton because small phytoplankton can not be directly grazed by copepods.  As a result, a greater  proportion of the carbon fixed in small cells is lost through respiration and excretion because of the additional trophic level introduced by microzooplankton grazing on small phytoplankton, before being consumed by mesozooplankton. The size structure of phytoplankton assemblage affects the vertical flux of organic carbon in the oceans; only the production of large, rapidly sinking particles can result in a  5  significant transfer of anthropogenic carbon dioxide into the deep ocean (Joint et al., 1993). This flux of organic carbon is often called export production (Berger et al., 1989) and in nutrient replete regions it can account for up to 50% of the total carbon fixation (Bienfang and Ziemann, 1992). Export production derived from the uptake of inorganic carbon by phytoplankton results in the transfer of carbon out of the upper ocean to the deep ocean.  Phytoplankton are an  important component of this biological pump. Parsons and Takahashi (1973) suggested that phytoplankton cell size selectivity is determined on the basis of ecological and species specific physiological data such as: 1) the rate of nitrate or ammonium uptake by the cell, 2) the extinction coefficient of the water, 3) the mixed layer depth, 4) the light intensity, 5) the sinking rate of phytoplankton, and 6) the upwelling velocity of water.  They suggested the physiological differences between Ditylum  brightwelli (30 um) and Emiliania huxleyi (5 um) can account for different growth rates in different environments. Only in regions of high light intensity and high nutrient concentrations, is it possible for large cells to grow faster than small cells. Michaelis-Menten uptake kinetics where Vma  X  The uptake of nitrogen follows  is the maximum uptake rate and K is the s  concentration supporting half the maximum rate of uptake (half-saturation constant) (Dugdale, 1967). Eppley et al. (1969) have shown that the large variation in Vmax and Ks of phytoplankton may account for competitive selection. Small cells with high surface area to volume ratios have a competitive advantage over large cells with respect to the acquisition of nutrients. Generally, small cells have a lower K and consequently in a low nutrient environment, they should s  dominate (Dugdale, 1967).  There is a general negative relationship between the size of  phytoplankton cells and their ability to take up nutrients (Eppley et al., 1969). Conversely, large cells usually have a large storage capacity for nutrients and consequently in environments where  6  nutrients are delivered in pulses, large cells often rapidly acquire and store nutrients and sustain growth for longer periods than smaller cells (Turpin and Harrison, 1979).  Often the  composition and size structure of the autotrophic assemblage is a major determinant of the quantity of production that can occur.  Upwelling regions generally support relatively large  phytoplankton stocks typically dominated by large phytoplankton composed mainly of diatoms.  E N S O - E l Nino Southern Oscillation Marine ecosystems undergo large interannual to decadal fluctuations that are beyond those attributable to direct harvest effects.  There is increasing evidence that persistent and  synchronous ecological changes have occurred that are linked to the variation in interdecadal climate (e.g. Venrick et al., 1987; Beamish, 1993; Beamish et al., 1999). The E l Nino Southern Oscillation (ENSO) is a natural climatic process that causes significant variability in living resources. ENSO is an interannual climatic condition that results in sea-surface warming, and a deeper thermocline and nutricline in eastern boundary currents (Philander, 1983).  A deep  thermocline is usually correlated with a deeper mixed layer and hence a reduction in productivity due to a decrease in the average light. Because light decreases exponentially as a function of depth, the depth of the surface mixed layer in which phytoplankton are homogeneously distributed, determines the quantity and quality of light that can be harvested by the phytoplankton (Sverdrup, 1953). Often coastal winds do not weaken and in fact, they may intensify during an E l Nino because of increased thermal differences between land and sea (Enfield, 1981).  Coastal upwelling may still occur, but the thermocline and nutricline are  depressed below the depth of entrainment (40-80 m). This effect was seen during the 1982-83  7  El Nino off the west coast of South Africa where coastal winds and locally forced winds continued through March 1983, but after November 1982, the surface water had significantly reduced nutrients and increased temperatures (Barber and Chavez, 1986) The effects of ENSO events such as those of 1982/1983 and 1991/1992 which enhanced the poleward movement of warmer water in winter and reduced summer upwelling along much of the west coast of North America, appears to have been responsible for altering the distribution of Pacific hake and salmon and for changing the predator-prey balance for many coastal species (Ware and Thomson, 1983). ENSO events may have a global geochemical impact also, since algal photosynthesis removes upwelled CO2 and reduces the pC02 gradient between the sea surface and the atmosphere. Gammon et al. (1985) found that the rate of increase of CO2 (pC02 anomaly) in the atmosphere fell to zero for a period during the 1982-1983 E l Nino. The ENSO event of 1997/8 showed unual development, both in terms of scale and mode of development.  Post 1976 ENSO events start to develop in approximately November to  December with the largest sea surface temperature anomalies recorded January to March of the preceding year. The 1997/8 E l Nino was different in that the first anomalies were seen along the equator in March and by June the anomalies had penetrated through the entire northeastern Pacific.  This thesis will examine physical, chemical, and biological parameters during the  1997/8 ENSO event.  The west coast of Vancouver Island Physical Oceanography Vancouver Island lies on the west coast of British Columbia (Canada) between approximately 48 to 51°N and 123 to 128.5°W. The western coastline of Vancouver Island has  8  numerous sounds and inlets that vary considerably in their dimensions (Pickard, 1963). Total precipitation varies along Vancouver Island, but generally a maximum occurs in December and a minimum in July (Dodimead, 1967). The river outflow volumes follow the annual cycle of precipitation except for the Gold and Stamp Rivers, which receive snow melt resulting in a second maximum outflow in May.  Shelf width and bathymetric profiles vary considerably along the western margin of Vancouver Island. The continental margin gradually narrows northward of La Perouse Bank from 65 km wide to just 5 km in width off Brooks Peninsula.  Near the southwest tip of  Vancouver Island, westward of Juan de Fuca Strait, the continental margin is cut by a deep (>250 m) narrow ( « 7 km wide) submarine canyon called the Juan de Fuca Canyon that extends seaward from the mouth of Juan de Fuca Strait. Northwest of this canyon is a series of isolated banks (z = 40-80 m), two additional submarine canyons, Nitinat and Barkley, and semi-enclosed basins (z >120 m).  Bottom slopes along banks and basins are steep. The shelf break is  approximately delineated at the 200 m isobath (Dodimead, 1985).  The Juan de Fuca Strait shows typical estuarine circulation that has a direct forcing effect on the southern margin of Vancouver Island. Brackish water flows out of the strait at the surface and denser water flows into the Strait near the bottom.  The strength of the flow is  mainly controlled by runoff from the Fraser River. The flow out of the strait is maximum in early summer when discharge from the Fraser River is maximal.  The Aleutian Low and the North Pacific High are two large scale pressure systems that govern the oceanic wind regimes off the coast of Vancouver Island (Favorite et al., 1976; Thomson, 1981).  The location and intensities of these two pressure systems control the  9  prevailing wind patterns along the west coast. Generally, from August to December (Northern Hemisphere fall/early winter) the Aleutian Low intensifies and shifts southward from the Bering Sea to the Gulf of Alaska.  Southeasterly to southwesterly winds persist from late August to  early spring as air flows counterclockwise around the dominant Aleutian Low. The Aleutian Low then progressively weakens until it is no longer evident in July as the North Pacific High intensifies until it covers the entire Gulf of Alaska during May to August.  The resulting  pressure pattern of the intensified North Pacific High and the diminished Aleutian Low during May through September, results in northwesterly winds as the air flows clockwise around the dominant North Pacific High pressure cell.  The study area for this thesis falls into the coastal upwelling domain described by Dodimead et al. (1963) and is subject to dynamic forcing from wind-induced upwelling. Continental shelf and slope waters off the west coast of Vancouver Island are at the northern end of an extensive Eastern Boundary Current system called the California Current that stretches from Baja California (25°N) to the northern tip of Vancouver Island (50.5° N) (Ware and McFarlane, 1989). Upwelling favorable winds are not as strong as those off the coast of California (Nelson, 1977) but are strong enough to induce classical wind-induced upwelling (Thomson, 1981). Conditions are favorable for upwelling in the summer, roughly from late March to the end of September, followed by a relatively abrupt reversal of the prevailing alongshore winds that produce winter downwelling favorable conditions.  Summer wind-  induced upwelling occurs during periods of persistent northwesterly (equatorward) coastal winds associated with the establishment of high pressure systems along the coast, while winter downwelling occurs during times of southeasterly (pole-directed) coastal winds associated with the passage of winter lows (Thomson et al., 1989).  10  There is considerable variability in the timing, duration and intensity of the summer upwelling and winter downwelling season. The summer season starts between early April and late June and lasts 146.4 ± 35.2 days, roughly three times longer than the winter downwelling season which starts in late September and runs until late November and lasts 53.7 ± 28.1 days (Thomson and Ware, 1996). A spring and a fall transition season separate the summer and the winter seasons. There is considerable interannual variability in the timing and duration of these two transitions. The spring transition can occur as early as January during 1995, or as late as mid-April during 1987, and usually lasts 79.2 ± 38.6 days. This variability in timing and duration is caused by changes in the prevailing winds, coastal runoff, alongshore pressure gradients, and other forcing mechanisms (Thomson and Ware, 1996). The fall transition can begin as early as August (1987) or as late as November and lasts 64.8 ± 27.4 days (Thomson and Ware, 1996). The fall transition variability is related to the degree of storm activity in the North Pacific and the positions of storm tracks relative to the coast of North America (Thomson and Ware, 1988).  Wind-induced upwelling is dynamic but variable along the west coast of  Vancouver Island.  Thomson and Ware (1996) report dramatic interannual variations in the  intensity of upwelling; for example the current velocity index (a measure of upwelling intensity) shows upwelling intensity during 1993 was approximately three times higher than the upwelling intensity during 1992.  Examination of a 32 yr record (1965-1997) of the Bakun Upwelling  Index, a measure of upwelling intensity, shows on average that coastal upwelling peaked in 1995 and 1996 and then steady declined to the lowest measurement recorded in 1997 (Robinson and Ware, 1999). Clearly, the relative forcing of wind-induced upwelling on environmental conditions off the west coast of Vancouver Island varies considerably.  11  The west coast of Vancouver Island is also subject to forcing from the Pacific Ocean. The surface circulation of the N E Pacific is dominated by the Subarctic Current that originates from the mixing of the Kuroshio and Oyashio currents (Dodimead et al., 1963; Tabata, 1975). The Subarctic Current flows eastward and divides into two streams about 300 km offshore at the latitude of Vancouver Island to form the northward-flowing Alaska Current and the southward flowing California Current (Dodimead et al., 1963; Tabata, 1975). In the winter, the northward flowing Alaskan Current intensifies and in response to the prevailing southeasterly winds, the surface flow along the shelf break is northward parallel to the shore (Manner, 1926; Freeland et al., 1984). This appears to be the northward extension of the Davidson Current that originates off California (Thomas and Emery, 1986). In the winter, the bottom current (up to 1 m s' ) 1  flows to the northwest, and is strongest in the late autumn or winter. In the summer, the surface flow along the shelf break is reversed in response to the shift in the California Current towards Vancouver Island and a reversal in the prevailing winds. Persistent northwesterly winds cause southward surface flow along the shelf break (Figure 2) and surface current speeds are often >40 cm s" to depths of 50 m (Mackas, 1992). Beneath the shelf break current, the California 1  Undercurrent flows northwest at 5-10 cm s' along the continental slope below 200 m and is 1  characterized by high salinity (34) and low dissolved oxygen (0.5-2.0 ml l" ) (Hickey, 1979; 1  Thomson, 1981). There is some evidence that suggests that the California Undercurrent extends as far north as Estevan Point (Freeland et al., 1984). The current pattern along the inner part of the shelf is different from that along the shelf break by the persistent northward flowing Vancouver Island Coastal Current (VICC) (See Figure 2). The VICC is a poleward flowing, buoyancy-driven surface current off the west coast of Vancouver Island (LeBlond et al. 1986; Thomson et al. 1989). It is a permanent feature of  12  the surface circulation that flows year-round from the entrance of Juan de Fuca Strait and can be found beyond Brooks Peninsula. It is generally confined to the innermost part of the shelf within about 25 km of the coast, but occasionally meanders seaward across the shelf (Freeland et al. 1984; Thomson et al. 1989). This current is driven by the flux of low density water onto the shelf from Juan de Fuca Strait (influenced by Fraser River discharge) andfromfreshwater runoff from coastal streams along the west coast of Vancouver Island.  These two sources of  freshwater differ in the timing of peak discharge and spatial distribution of discharge. Direct discharge from coastal streams peaks during the fall winter rainy season, while discharge from the Fraser River peaks in June. This current is characterized by low surface salinity (30-31.5). Maximal near surface speeds can reach 50 cm s" within the core of the current, but average 1  longshore flow from October - March is « 25 cm s" and from April - September is « 10 cm s~\ 1  Current speeds are reduced in the summer because strong northwesterly winds tend to retard the flow while during the winter southeasterly winds augment the northward flow. Summer and autumn circulation at the southern end of Vancouver Island is often dominated by the Tully eddy or alternatively called the Juan de Fuca eddy (Figure 2)  This was  first observed by Tully (1942) and it is situated over the southwest margin of Vancouver Island. Freeland and Denman (1982) found that this cyclonic eddy is controlled by the interaction between bathymetry and the local flows off the southwest margin of Vancouver Island.  A  shallow region called Finger Bank, deflects currents to the west and starts the eddy on its course (Freeland and Denman, 1982). The average residence time of water in the upper 40 m is 4.2 d, with a range of 1.2 to 17 d and water exits either north onto La Perouse Bank or south to Washington. (Freeland, 1988).  Although this eddy is a local effect, it does dominate the  productivity in this region during the summer months (Mackas and Sefton, 1982).  13  129°  128°  127°  126°  125°  124°W  Figure 2 Surface circulation off the west coast of Vancouver Island in summer (redrawn from Thomson et al., 1989). Dashed line marks the 200 m contour. Vancouver Island coastal current =VICC.  Long-term coastal temperature variability (Brainard and McLain, 1985) and long term variability in coastal sea levels (Enfield and Allen, 1980) has been studied for many years. Thomson et al. (1984) examined a 40 year sea-surface temperature data set of and found decadal scale cycles in the water temperature along the west coast.  Current observations  suggest the ocean climate has recently undergone a change during the 1990s. Sea-surface temperatures measured at Amphitrite Point lighthouse on the west coast of Vancouver Island show two large temperature anomalies during the 1990's, the 1992 and the 1997 E l Niftos (Robinson and Ware, 1999). Relative to sea-surface temperatures dating back to 1000's, the  14  1990's were an unusually warm decade, which ended abruptly in 1999 with conditions typical of the 1960's and early 1970's (Mann et al, 1999).  Ecological Dynamics The physical oceanography of the west coast of Vancouver Island has been studied much more extensively than the biological oceanography and generally studies of ecological dynamics have been restricted to the southern margin of Vancouver Island. Historically, several large research programs have been completed in this area including Coastal Oceanic Dynamic Experiment (CODE), La Perouse, and Marine Survival of Salmon Species (MASS) program. C O D E was initiated to study the general circulation, the dominant physical mechanisms, and the resulting planktonic ecosystem dynamics on the continental shelf. The La Perouse project was initiated in 1985 following the major 1982/83 E l Nino event in the Pacific Ocean to investigate the causes of annual and interannual recruitment variability in herring, sablefish, Dungeness crab and Pacific cod. The MASS program was initiated to investigate the interrelationships between biophysical events and salmon distribution and survival on an annual and interannual time scale.  The surface waters off the west coast of Vancouver Island are characterized by high levels of dissolved nutrients (Mackas et al., 1980; Denman et al., 1981, 1982) supplied by wind mixing, episodic wind-driven upwelling, topographically controlled upwelling  (Freeland and  Denman, 1982) and the outflow from Juan de Fuca Strait where deep nutrient-rich water is advected to the surface by tidal mixing and estuarine circulation (Mackas et al., 1980). The outflow from Juan de Fuca is a potential source of nutrients for phytoplankton for both the shelf and offshelf regions due to cross-shelf transport of water to the outer shelf (Mackas and  15  Yelland, 1999; Crawford, 1989). Mackas et al, (1980) estimated the flux of nitrogen (nitrate + nitrite) out of Juan de Fuca Strait to be 30 kg s" depending on the rate of outflow of water from 1  Juan de Fuca and the nitrogen concentration of this outflow water and estimates that windinduced upwelling accounts for on average -20% of the estimated contribution from estuarine flow.  The relative contribution of upwelling may increase 2-3 fold during periods of active  upwelling, but on average the relative contribution of upwelling is less than Juan de Fuca. Crawford and Dewey (1989) estimated nitrogen flux rates for wind-driven upwelling at 15 kg s" wind mixing in the upper 20 m at 5 kg s" and turbulent mixing less than 2 kg s". 1  In  1  summary, the nutrient rich surface waters off the west coast of Vancouver Island are mainly derived from estuarine flow out of the Juan de Fuca, followed by upwelling, and tidal mixing (Crawford and Dewey, 1989). Surface nitrate is rarely depleted to undetectable levels. At the southern margin of Vancouver Island, shoreward of the shelf break current and extending into the Juan de Fuca Strait, near surface nitrate in excess of 20 u M is common. On average, the nitrate concentration of the inner part of the shelf is >10 uM, and at beyond shelf regions, nitrate concentrations are lower but generally between 1-5 uM. The shelf region off Vancouver Island is characterized by relatively high phytoplankton biomass.  Maximum chlorophyll concentrations between 10-50 mg chl m" arefrequentlyfound 3  (Mackas and Sefton, 1982), and generally there is a cross-shelf gradient. On average, nearshore surface layer chl a concentrations >5 mg chl m" are common compared to 1-3 mg m" for 3  offshore regions (Mackas, 1992). al  3  Phytoplankton biomass is patchy in distribution. Denman et  (1981) found persistent regions of high surface chlorophyll (20 mg chl m") along a 3  nearshore 20 km wide band and a zone of high biomass centered over the 80 m bathymetric contour. The seasonal cycles of the shelf and the beyond shelf are different. Phytoplankton  16  biomass on the inner shelf is persistently high in the spring, summer and fall; on average biomass peaks in August/September.  In the offshore regions, biomass blooms in March/April,  decreases in May and June coupled with depletion of nutrients and increases throughout the season as nutrients increase until October/November when phytoplankton are likely limited by light. The studies by Denman et al. (1981), Mackas and Sefton (1982), Forbes and Denman (1991), and Taylor and Haigh (1996) are among the few to analyze phytoplankton species composition off the west coast of Vancouver Island. Most studies were limited to the southwest margin of Vancouver Island and limited to identification of the dominant species, except for the investigation  of Taylor and Haigh (1996) who completed  microplankton community in Barkley Sound.  Taylor and Haigh (1996) found and identified  potentially harrnful phytoplankon species in B.C. coastal waters. focussed  exclusively on the distribution of  pungens),  Forbes and Denman (1991)  Pseudo-nitzschia pungens  (formally  a diatom known to produce domoic acid and shellfish poisoning (Subba Rao  1988). Denman  Nitzchia  a systematic study of the  et al.  (1981) found the diatoms,  Rhizosolenia setegera  Nitzschia et al.,  (-300 um long) and  spp. (-40 um long) dominated the phytoplankton assemblage when biomass was high.  There have been no detailed taxonomic studies off the west coast of Vancouver Island that extend to the northern reaches of Vancouver Island, although Taylor and Waters (1982) have studied the subarctic Pacific beyond the continental shelf. Large diatom populations typically develop during late July and August (Mackas et al,  1980; Denman et al., 1981; Mackas and  Sefton, 1982). Detailed studies examining temporal and spatial variability of primary productivity are limited on the west coast of Vancouver Island. Most of the study spatially focussed on the  17  southern margin of Vancouver Island during short-time scales. Persistent zones of high primary productivity that last for several months have been measured on the southwestern coast of Vancouver Island (Denman et al., 1981).  Primary productivity at the surface as high as  136 mg C m" h" was measured during May of 1981. 3  During July and August, maximum  1  primary productivity was 49.1 and 61.4 mg C m" h" respectively. 3  1  The assimilation numbers  measured were 3.92 and 3.75 mg C mg chl" h" , while Forbes and Denman (1991) found 1  1  assimilation numbers ranging from 3.5-17.5 mg C mg chl" h" . 1  1  The La Perouse Bank is one of the most productive fishing areas in the Northern Hemisphere and it generates a landed value to the British Columbia economy in excess of $40 million annually (Ware and Thomson, 1991).  The west coast of Vancouver Island is an  important feeding and breeding region for many pelagic fish species including Walleye pollock, Pacific cod, Pacific halibut, Pacific hake, Pacific sardine, Pacific herring, Pacific mackerel and northern anchovy. It is a migration corridor for returning salmon. When phytoplankton stocks are abundant, Pacific hake, sardine and mackerel migrate into Canadian waters in the summer to feed and return to southern Baja California where they spawn in the winter and spring (Ware and McFarlane, 1988). Recent changes in the pelagic fish community of species abundance and production dynamics have been observed during the 1990's off the west coast of Vancouver Island.  Increased biomass of mackerel, a reappearance of Pacific sardine (Hargreaves et al.  1994), more abundant but smaller Pacific hake (Ware and McFarlane, 1995) and poorer growth of Pacific herring have been observed (Tanasichuk, 1997).  18  Global Ocean Ecosystem Dynamics Program ( G L O B E C ) In 1997 the Canadian G L O B E C (Global Ocean Ecosystem Dynamics) program set out to understand how living marine resources are affected by variability in their physical environment. Previous work by the La Perouse Project, MASS (Marine Survival of Salmon), COPRA and salmon-index streams programs, focussed exclusively on the southern margin of the west coast of Vancouver Island and found strong upstream influences on water properties (Thomson et al., 1989). One objective of G L O B E C was to extend the study area to the northern region of Vancouver Island to provide good spatial and temporal coverage for the west coast of Vancouver Island. Despite extensive studies on the southern margin of Vancouver Island, little information is available for primary productivity. Previous studies have focussed extensively on nutrient and chlorophyll distributions and there has been limited study of primary productivity on isolated cruises covering small spatial and temporal scales. Very little is known about seasonal and  interannual variability of primary productivity and the  importance of  different  phytoplankton size fractions have not been investigated for the west coast of Vancouver Island.  Thesis Goals Physical, chemical and biological parameters were investigated to document spatial and temporal variability in transects crossing the continental margin on the west coast of Vancouver Island. Specifically, during 1997 and 1998 the variability of physical, chemical and biological parameters was studied: a) for the shelf and beyond the shelf regions, b) for 4 distinct geographic regions of the west coast of Vancouver Island such as a region cut by a underwater  19  canyon, or strongly affected by strong estuarine flow, and c) over the spring, summer and fall growing seasons.  This thesis also evaluates the size structure of primary productivity and  phytoplankton biomass.  Primary productivity was examined in the upper water column on  horizontal transects across the continental margin to document the spatial and temporal variability in the size distribution of phytoplankton. Three specific questions were addressed by this research: (1) Do dissolved nutrients, chlorophyll concentrations and primary productivity vary seasonally, interannually and spatially? (2) Does the large size fraction (>5.0 pm) contribute substantially to the phytoplankton biomass and primary productivity during the upwelling season? (3) Was the primary productivity impacted during the strong ENSO event of 1997 and 1998? Field studies were conducted as part of the Canadian G L O B E C program aboard the C.S.S "John P. Tully".  Cruises off the west coast of Vancouver Island were undertaken  seasonally over a period of two years from 1997-1999. The data for the 1999 cruises are contained in Appendices B (chlorophyll), C (nutrients), and D (primary productivity). Thesis Organization Chapter one of this thesis examines seasonal variability of physical, chemical and biological parameters along the west coast of Vancouver Island. Four transects perpendicular to the coast were studied in detail and the results will be discussed in Chapter one.  The four  transects were: over La Perouse Bank, over Barkley Canyon, off Estevan Point, and off Brooks Peninsula.  Chapter two examines size-fractionated biomass and primary productivity of the  four study transects.  At each transect, sampling stations were chosen in order to have one  station on and one station off of the continental shelf.  20  CHAPTER 1  VARIABILITY OF PHYSICAL, C H E M I C A L AND BIOLOGICAL PARAMETERS O F F T H E WEST COAST OF VANCOUVER ISLAND  INTRODUCTION  The goal of G L O B E C was to determine how and why marine ecosystems change in response to variations in the physical oceanic environment.  To quantify and interpret these  changes on the continental margin of British Columbia, it is essential to expand upon existing time series observations, both temporally and spatially.  Long-term time series data sets are  necessary to achieve this goal because many of these changes are known to occur on interannual or decadal time scales (Dickson et al., 1988; Beamish and Bouillan, 1993; Steele, 1998) which can only be identified and understood using multi-year data sets. It is essential to expand the time base of existing time series by incorporating new observations to allow identification of low frequency variability important to both physical and biological processes. The southern margin of the west coast of Vancouver Island has been the focus of extensive studies but there has been little study of the northern coast of the Island.  It is  important to expand studies to the northern regions of Vancouver Island because previous studies have found strong upstream influences on water properties off the southwest coast of Vancouver Island (Thomson et al., 1989). In addition to the expansion of the time-series observations (this chapter), and process studies (Chapter 2), numerical ecosystem models are being developed that require boundary conditions and validation data and expansion of historical data sets will greatly increase confidence in the results of model predictions (Ianson et al., submitted).  21  The main objective of this chapter was to show the seasonal, interannual and spatial variability of physical, chemical and biological parameters in transects that cross the continental margin off the west coast of Vancouver Island.  MATERIALS AND METHODS  Six cruises aboard the C.S.S. John P. Tully were undertaken on the west coast of Vancouver Island as part of the Canadian Global Ocean Ecosystem Dynamics program (GLOBEC).  Studies were conducted during 3 cruises in 1997 and three in 1998, which  correspond to the annual spring transition, the summer upwelling season and fall transition period. Cruise details are available in Table A. 1. Studies were also conducted during 3 cruises in 1999 and these data are in Appendix B (dissolved nutrients), Appendix C (chlorophyll), and Appendix D (primary productivity). Studies were conducted at several stations along transects extending perpendicular to the west coast of Vancouver Island from the southern to the northern tip of Vancouver Island (Figure 1.1).  These transects bisected the continental shelf, and the  shelf break and ended in deep water beyond the shelf. A n attempt was made to occupy similar stations during each cruise, but sampling logistics and weather conditions prohibited complete replication. During the spring transition cruise in April 1997, several stations were sampled again one week after the initial occupation. Station details such as latitude, longitude and water depth for all stations sampled during each of the 3 cruises in 1997, 3 cruises in 1998 and 3 cruises in 1999 are presented in Appendices E . Physical Measurements Incident surface solar irradiance (photosynthetically active radiation - I PAR) was 0  continuously measured with a Licor Quantum Sensor Model LI-190SB calibrated for use in air  22  Figure 1.1 Location of transects off the west coast of Vancouver Island. Dashed line delineates the 200 m contour. A=Juan de Fuca Canyon (Line A), B=La Perouse Bank (Line B), C=Barkley Canyon (Line C), D=D Line, G=Estevan Point (Line G), H=H Line, J=J Line, BP=Brooks Peninsula (BP Line) and CS=Cape Scott (CS Line). Transects with * will be discussed in this chapter.  23  and mounted in a shade-free area on the after-deck of the ship. The measurements were logged as 10 min averages using a Licor datalogger Model LI-1000. Light data were obtained for all cruises except the October 1998 cruise due to a datalogger failure; therefore for October 1998, primary productivity calculations the light data from October 1997 was used. In May 1998, the datalogger did not log continuously, and therefore for the days without continuous data, the data for a similar day was used. Vertical profiles of underwater irradiance were measured with a Biospherical QSP-200 L4S 4495 PAR sensor calibrated for use in water. When light profiles could not be measured due to weather conditions, pre-dawn sampling, or unfavorable sampling logistics, estimates based on previous casts were used. For these stations, actual and estimated light depths may have been slightly different. Due to a processing error, underwater irradiance is not available for most stations at the time of writing. Vertical profiles of conductivity, temperature, pressure, and chlorophyll fluorescence were obtained using a Seabird® Model SBE 911 C T D Serial #0437 mounted with a SeaTech +  fluorometer.  Personnel from the Institute of Ocean Sciences (Sidney, B.C.) provided all  physical data. The raw data are stored at the Institute of Ocean Sciences. Vertical profiles of cr were derived from temperature and salinity data using the t  expression given by Millero and Poisson (1981). The mixed layer depth was identified as the depth where a 0.125 change in a was first observed relative to a surface reference value (after t  Levitus, 1982). Chemical and Biological Measurements The time of the day when stations were occupied was variable among stations and between cruises and was dependent upon the time of arrival on station. Seawater was collected using acid-cleaned 10-L P V C Niskin bottles equipped with Teflon® coated springs and fittings 24  and silicone tubing mounted on an instrumented rosette sampler. Seawater samples for vertical profiles of chemical and biological parameters were taken at 0, 10, 20, 30, 50, 75, 100, 125, 150, 175, 200, 250, 300, 500, 600, 700, 800, 1000, 1200, 1400, 1500, 1600, 1700 m depending on the bottom depth. After collection of dissolved oxygen, water samples were immediately collected for dissolved nutrients, chlorophyll a, phytoplankton identification and primary productivity measurements (Chapter 2).  Seawater was also collected for particulate nitrogen and carbon,  ammonium and urea concentrations and nitrogen uptake rate experiments (Varela et al., in prep.). Water samples for nitrate + nitrite (NCV + N t V ) , soluble reactive phosphate (HPO4 "), 2  and silicic acid (Si(OH) ) were filtered directly out of the Niskin bottles using an acid-cleaned 4  60 ml Nalgene® syringe fitted with 25 mm Millipore Swinnex® filter holder and a combusted (460°C for 4.5 h) 25 mm glass-fibre filter.  In 1997, seawater was filtered through 25 mm  Whatman™ GF/F glass-fibre filters (nominal pore size 0.7 (am) and in 1998, 25 mm A M D ™ GF75 glass-fibre filters (nominal pore size 0.75 um) were used. Seawater was gently filtered into acid-cleaned Nalgene® bottles and stored at -20°C until analysis ashore. Gloves were used for all nutrient sampling to avoid contamination. All dissolved nutrients were processed using a Technicon® Autoanalyzer® II. Nitrate plus nitrite, soluble reactive phosphate and silicic acid were determined using the procedures of Wood et al. (1967), Hager et al. (1968), Armstrong et al. (1967), respectively.  Combined nitrate and nitrite concentrations are reported as nitrate.  Shelf region stations were east of the shelf break, as defined by the 200 m contour, and the beyond shelf stations were those beyond the shelf break. The number of stations varied at each transect and each cruise and are summarized in Appendix E . Chlorophyll corrected for phaeopigments was determined at 0, 10, 20, 30, and 50 m water depth by in vitro fluorometry (Yentsch and Menzel, 1963). A 500 ml water sample was 25  filtered onto 25 mm diameter glass-fibre filters using a vacuum pressure differential of  <100  mm of Hg and stored at - 2 0 ° C in a dark desiccator until analysis ashore. In 1997, seawater was filtered through 25 mm Whatman™ GF/F glass-fibre filters and in 1998, 25 mm A M D ™ GF75 glass-fibre filters were used. A l l samples were analyzed within two weeks of collection. Chl a was extracted in 10 ml of 90% acetone by sonication in an ice bath for 10 min and then subsequently stored in the dark for 20-24 h at - 2 0 ° C . The fluorescence of the acetone extract was measured before and after the addition of three drops of 10% HC1 to estimate phaeopigments in a Turner Designs™ Model 10-AU fluorometer calibrated with a solution of commercially available chlorophyll a obtained from Sigma Chemical Company. Chlorophyll a calculated using the equation of Parsons et al. (1984). Samples for phytoplankton identification were fixed with acidic Lugol's iodine solution during the 1997 sampling season and neutral Lugol's iodine solution during the 1998 sampling season (Throndsen, 1978; Parsons et al., 1984).  The samples were stored in the dark until  identification and enumeration was performed using inverted microscopy following Utermohl (1958) procedures. Depending on the biomass, 10 or 25 ml was settled in a counting chamber for at least 12 h. This methodology underestimated the heterotrophic crytomonads that would best be distinguished using an epifluorescence technique (Geider, 1988). The counts were then converted into cells l" . Cell carbon was calculated according to the equations of Strathman 1  (1967). Cell volumes specific to each species were required for the conversion of cells l" to 1  carbon. Cell volume calculated using measurements of representative cells and equations for simple geometric shapes spheroids, cylinders, boxes, and cones were supplied by R. Haigh (unpubl. data). The cell volume for each species is given in Appendix F.  26  Statistical analysis of chemical and biological data Replicate casts were not completed due to time constraints in the cruise schedule and the labor-intensive nature of this study. Routinely a single water sample was collected from each depth for analysis of chemical and biological parameters. One factor analysis of variance (ANOVA) and a Tukey test were used to examine spatial and temporal variation. For analysis of temporal variation, both interannual and seasonal, the physical, chemical and biological data were grouped according to mean values for the west coast of Vancouver Island (WCVI) and for shelf and beyond shelf regions.  For analysis of  spatial variation, the cross shelf and along shore direction, the data were grouped according to cruise and year.  Contour plots were created using Wavemetric Igor Pro (v. 4.0) using 64x triangulation.  RESULTS  I. Physical Parameters A)Incident Irradiance The continuous recordings of incident surface irradiance are shown for April, July and October of 1997 and May and July 1998 in Figure 1.2.  Irradiance data are not available for  October 1998 due to a datalogger malfunction. For the two study years, the highest values were observed in July 1997 (Figure 1.2b) and the lowest values in October 1997 (Figure 1.2c). For July 1997, surface irradiance was consistently high, and during October 1997, surface irradiance was consistently low for the duration of the cruise.  There was considerable variability in  surface irradiance during May and July of 1998 (Figure 1.2 D & E) while incident surface irradiance in April and July of 1997 was more consistent.  27  The incident solar irradiance on the day of the primary productivity experiments (Chapter 2) is included in Appendix G.  Included in the plots is the percentage that the  incubation period represented of the total daily irradiance. B) Mixed Layer Parameters (Depth, temperature, salinity and a ) t  Very high variability of mixed layer parameters was measured off the west coast of Vancouver Island, and frequently the standard deviation and mean were similar. Trends and patterns will be discussed, but the differences between regions, cruises or years are not statistically significant unless otherwise noted. Mixed layer (ML) depth and mixed layer parameters are shown in Table 1.1 for April, July and October 1997 and Table 1.2 for May, July and October 1998 for the primary productivity stations. BI) MIXED LAYER (ML) DEPTH The M L was consistently deeper at beyond shelf regions than those over the shelf. On average over the 2 years, the M L was 23 ± 1 2 m in the beyond shelf region and 12 ± 6 m in the shelf region. The M L was deeper in 1998 compared to 1997 for both the shelf and the beyond shelf region. The mean M L in the shelf region was 11 ± 7 m in 1997 and was 13 ±5 m in 1998 and for the beyond shelf region was 22 ± 1 9 m in 1997 and 24 ± 6 m in 1998. There was a consistent seasonal pattern for the shelf region in 1997 and 1998. The M L was deepest in October and shallowest during July, the month of continuously high surface irradiance. There was no consistent seasonal pattern for the beyond shelf region in 1997 and 1998. In 1997 the deepest M L was in April, and in 1998 was in October.  28  2500  April 1997  July 1997  16  2500  CD o -r^ CD CO T3 CM  g g  500  20  21  22  23  24  25 26  October 1997  0 2500  o ci  19  1500 -E 1000  SS 3  18  2000 -E  'E  g  17  BP2 BP7  G3  C9  C1 C4  22  23  24  25  26  27 G7  2000 1500  May 1998  1000 500 -_ 0  BP2 C1 C4 14 15  BP7 16  17  18 19  20  21  22  23  24  2500 2000 -_  B16  1500  July 1998  1000 500 0  h/UJUAf 14  15  16  17 18  19 20 21 22 23 24  25  26  Day of the Month Figure 1.2 Incident surface irradiance for A) April 1997, B) July 1997, C) October 1997, D) May 1998 and E) July 1998. Station labels on top of peaks indicate day primary productivity was measured. See Appendix E for location of stations. Irradiance data are not available for October 1998.  No consistent south-north trend was observed in mixed layer depth at shelf or beyond shelf stations. Vertical profiles of temperature, salinity and a for the primary productivity t  stations are provided in Appendix H . Variability was higher in 1997 than in 1998 for the shelf and the beyond shelf region (see coefficient of variation in Tables 1.2 and 1.3).  Seasonal  variability was highest in July for both the shelf and the beyond shelf region. B 2 ) ML TEMPERATURES M L temperatures were on average warmer (~ 1°C) at the beyond shelf region than at the shelf regions. On average over the 2 years, the M L temperature was 12.8 ±2.1°C in the beyond shelf region and 11.9 ±1.7°C in the shelf region. The M L temperature in the shelf region were similar in 1997 and in 1998, while in the beyond shelf region M L temperatures were on average warmer in 1998 than in 1997. In 1997, the spring cruise was approximately one month earlier than the spring cruise in 1998, this might explain the lower temperature measured in 1997. If spring cruise is not include in the seasonal mean, the is no significant difference between 1997 (14.1°C)and 1998 (13.9°C). There was a consistent seasonal pattern for the shelf region and the beyond shelf region in 1997 and 1998. The M L temperatures in the shelf region were warmest in July arid coolest in April. The water column was generally weakly stratified during April 1997 but was stratified by July.  In July, the thermocline was -10 m, but by October it was depressed to ~40 m  (Appendices, H1-H3).  In 1998, the water column was generally strongly stratified for all  cruises; the thermocline was -20-30 m in the beyond shelf region and -10-20 m for shelf regions (Appendices, H4-H6). In 1997, the seasonal variability in the M L temperature in the shelf region was significant (p<0.01), while in 1998 no significant seasonal variability was  30  found.  The seasonal variability in the M L temperature of the beyond shelf region was  significant during both study years (p<0.01). A consistent alongshore trend in M L temperature was not observed for the shelf region while a consistent trend was observed for the beyond shelf region. Beyond shelf stations of the southern transects L a Perouse Bank or Barkley Canyon tended to be warmer than the northern transects, Estevan Point and Brooks Peninsula. The northern transects have shorter continental shelves so upwelled water has less distance and less time to warm before it is transported to the beyond shelf region. In contrast, by the time cold upwelled water arrives at the beyond shelf stations of the southern transects, solar irradiation has warmed it. Low M L temperatures corresponded to high salinity measurements, consistent with characteristics of an upwelling region. Similarly, at stations with high M L temperature, the lowest salinity was noted. Variability was higher in 1997 than in 1998 for the shelf and the beyond shelf region (see coefficient of variation in Tables 1.2 and 1.3). Variability was highest in October for both the shelf and the beyond shelf region in 1997 and 1998. M L temperatures ranged from 9.017.5°C during 1997 and 10.3-14.6°C during 1998. B3) M L SALINITY A N D D E N S I T Y  The M L salinity and density were on average lower in shelf regions than at the beyond shelf regions. On average over the 2 years, the M L salinity was 31.5 ±0.7 in the beyond shelf region and 31.1 ±0.9 in the shelf region. The M L salinity and density in the shelf region and beyond shelf region were lower in 1997 than in 1998. The lowest M L salinity and density was measured in the shelf region in 1997. There was no consistent seasonal trend in M L salinity and density in the shelf region. Salinity and density tended to be lowest in April suggesting the V I C C lowered the salinity, but 31  by July and October the salinity increased perhaps due to upwelling. There was no consistent seasonal trend for the beyond shelf regions. No consistent south-north (latitudinal) trend was observed at either shelf or beyond shelf stations during either 1997 or 1998. The transect that showed the highest and the lowest salinity and density, varied during each cruise. Variability in salinity and density was low during 1997 and 1998, but was higher in 1997 than in 1998 for both shelf and the beyond shelf regions (see coefficient of variation in Tables 1.2 and 1.3). Variability was similar during each cruise for both the shelf and the beyond shelf regions in 1997 and 1998. M L salinity ranged from 29.18-32.38 during 1997 and 30.99 - 32.35 during 1998. B4) SUMMARY OF PHYSICAL PARAMETERS In summary, a strong seasonal cycle was noted for solar radiation, the highest flux was measured in July and the lowest during October. A shoaling in the mixed layer depth was noted in both regions in 1997 relative to 1998.  Mixed layer temperature was similar in both years.  Salinity and density were lower in 1997 than in 1998 for both regions.  A strong cross-shelf  gradient was observed in 1997 and 1998. The mixed layer depth was consistently deeper and temperature, salinity and density were all higher in the beyond shelf regions than in the shelf region.  32  Table 1.1 Mixed layer parameters for stations occupied during 1997 cruises off the west coast of Vancouver Island (WCVI). Monthly mean, yearly mean ± 1 S.D. and yearly coefficient of variation, (C.V.,%) are given for the shelf and beyond shelf region. Temperature (°C), salinity, and density for the mixed layer were calculated as the mean value from the surface to the calculated mixed layer depth. Dashed line indicates that data are not available. Transect  Shelf La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula WCVI WCVI WCVI WCVI SD CV Beyond Shelf La Perouse Bank  Date  April July Oct. April July Oct. April July Oct. April July Oct. April July Oct. 1997 1997 1997  Mixed Layer (m)  Temp.  3 3 25 17 10 14 14 9 16 5 6 7 10 7 16 11 6.6 62  9.16 11.6 10.9 9.70 14.3 14.0 9.16 15.1 14.0 8.97 11.9 13.5 9.2 13.2 13.1 11.9 2.3 19  Salinity  Cc)  Density (<*<)  29.39 31.70 31.45 29.18 30.81 31.13 30.40 30.16 30.59 29.46 31.41 . 29.98 29.6 31.0 30.8 30.47 0.9 3  April 67 8.67 32.38 30.78 July 13 17.5 Oct. April 11 9.23 31.28 Barkley Canyon July Oct. 19 13.5 31.01 Estevan Point April 15 9.06 30.15 July 9 14.9 30.20 Oct. 38 12.9 31.92 April Brooks Peninsula July 14 13.0 31.10 Oct. 12 13.2 30.36 WCVI April 31 9 31.3 WCVI July 12 15.0 30.7 WCVI Oct. 23 13.2 31.1 WCVI 1997 22 12.4 31.0 1997 19.0 2.9 0.8 SD CV 1997 86 24 2 Latitude, longitude and bottom depth are found in Appendix E.  22.73 24.17 24.07 22.48 22.92 23.23 23.52 22.26 22.84 22.51 23.84 22.44 22.8 23.3 22.9 23.00 0.7 3 25.15 22.20 -  24.20 -  23.24 23.34 22.33 24.04 23.40 22.80 24.2 22.6 23.4 23.4 0.9 4  Table 1.2 Mixed layer parameters for stations occupied during 1998 cruises off the west coast of Vancouver Island (WCVI). Monthly mean, yearly mean ± 1 S.D. and yearly coefficient of variation (C.V., %) are given for the shelf and beyond shelf region. Temperature (°C), salinity, and density for the mixed layer were calculated as the mean value from the surface to the calculated mixed layer depth. Transect  Shelf  La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula WCVI WCVI WCVI WCVI SD CV  Date  April July Oct. April July Oct. April July Oct. April July Oct. April July Oct. 1997 1997 1997  Mixed Layer (m)  Temp.  19 8 11 10 9 19 19 15 16 8 11 9 15 11 14 13 5 35  10.3 11.9 12.2 11.9 12.1 11.5 12.4 13.4 11.8 11.0 12.9 11.4 11.4 12.6 11.7 11.9 0.8 7  Salinity  Density  CQ 30.99 32.27 " 32.10 31.16 31.89 32.30 31.49 31.27 31.97 31.79 31.44 31.79 31.4 31.7 32.0 31.7 0.4 1  23.81 24.51 24.34 23.66 24.18 24.61 23.83 23.46 24.30 24.31 23.68 24.23 23.9 24.0 24.4 24.1 0.4 2  Beyond Shelf  La Perouse Bank  April 23 11.6 31.91 July 26 14.6 32.35 Oct. 37 13.8 32.13 Barkley Canyon April 16 11.9 31.64 July 23 13.9 32.10 Oct. 31 14.5 32.22 Estevan Point April 22 11.4 31.97 July 31 14.4 32.05 Oct. 23 13.6 32.15 April 21 10.6 31.90 Brooks Peninsula July 25 13.6 31.94 Oct. 15 13.0 32.19 WCVI April 21 11.4 31.9 WCVI July 26 14.1 32.1 WCVI Oct. 27 13.7 32.2 WCVI 1997 24 13.1 32.0 1.3 0.2 1997 6.3 SD 1997 26 10 1 CV Latitude, longitude and bottom depth are found in Appendix E.  24.34 24.04 24.04 24.02 23.98 23.96 24.37 23.86 24.08 24.45 23.93 24.24 24.3 24.0 24.1 24.1 0.2 1  II. Chemical Parameters The following sections will examine the distribution of surface (0-10 m) nutrients in space and time. A l l values that will be discussed are 0-10 m surface concentrations. Vertical profiles of all three nutrients are provided in Appendix I. For the nutrient data, three spatial scales were examined: the mean of all stations sampled, including all shelf and the beyond shelf stations, will be referred to as the mean for the WCVI; a cross-shelf gradient (east/west) and for the nutrient data is referred to as N03~ hdf, s  HP0 - eif, Si(OH) helfOr NCV beyond? HPO4 beyond or Si(OFf)4beyond for each transect; and in an 2  4  sh  4s  alongshore gradient (latitudinal gradient) from the southern La Perouse transect to the northern Brooks Peninsula transect for the shelf and beyond shelf region.  Three time scales will be  examined for each spatial scale: a 2-yr mean (1997 and 1998); annual mean for 1997 and 1998; and season means. Very high variability was measured off the west coast of Vancouver Island, and frequently the standard deviation and mean were similar. Trends and patterns will be discussed, but the differences between regions, cruises or years are not statistically significant unless otherwise noted. A) DISSOL VED NUTRIENT CONCENTRA TIONS  A l ) MEAN VALUES FOR THE WCVI The average W C V ^ ^ , WCVI osphate, WCVI ii ph  mean WCVI  nitra  S  icic a c i d  are shown in Table 1.3. The 2-yr  te was 4.1 u M , WCVI h s hate was 0.59 and WCVI H p  0  P  si  cic  aci  d was 10.5 uM.  WCVI itrate in 1997 was 5.2 u M and in 1998 it was 2.9 uM, which was significantly lower than n  in 1997 (p<0.05). The WVCIhosphate in 1997 was 0.55 u M and in 1998 it was 0.63 uM. The p  WCVIsiHcic acid concentration in 1997 was 12.9 u M and in 1998 was 8.12 u M , which were significantly lower than in 1997 (p<0.01).  35  A seasonal trend was evident in 1997 for WCVInj^te and WCVI jij i S  C C  jd but not for  ac  WVCIphosphate. On average in 1997, WCVI trate and WCVI | i id concentrations were highest in ni  si  ic  C ac  April and decreased as the season progressed to the lowest during October. In 1998, nutrients were consistently high in October and there was a significant difference in nitrate (p<0.05), phosphate (p<0.05), and silicic acid (p<0.01) concentration between cruises in 1998. In April 1997, surface (0-10 m) N0 "ranged from 2.5-13.5 uM, HP0 "ranged from 0.32  3  4  1.0 uM, and Si(OH) ranged from 11.2-29.2 uM. Variability was consistently higher in 1998 4  than in 1997 for nitrate, phosphate and silicic acid.  Table 1.3 Mean nutrient concentrations (0-10m) (uM) for WCVI during each cruise in 1997 and 1998. Values are the mean of all stations during each cruise. Yearly mean ±1 S.D. N0  HPO "  Si(OH)  6.6 ±2.1 6.4 ±5.4  0.5±0.1 0.6 ±0.4  15 ±3.4 13 ±9.8  3.3 ±2.3  0.6 ±0.2  11 ± 3 . 4  1997 Mean 1997 C V  5.4 ±4.0 47  0.6 ±0.3 28  13 ±6.8 28  1998 May July Oct.  0.9 ±1.1 3.4 ±3.6 5.1 ±2.8  0.3 ±0.1 0.8 ±0.3 0.8 ±0.4  2.7 ±1.5 10 ±5.5 9.8 ±6.0  1998 Mean 1998 C V  3.1 ±3.3 91  0.6 ±0.4 40  7.0 ±5.9 52  2 year 2 year C V  4.4 ±3.8 86  0.6 ±0.3 56  11 ±7.0 65  1997 April July Oct.  2  3  4  4  36  A2) CROSS-SHELF GRADIENT  The mean nutrient concentrations are shown for each cruise in 1997 and 1998 in Table 1.4. Note that during 1998 there was no data available for La Perouse Bank. On average nutrient concentrations were all higher in the shelf region than in the beyond shelf region. In 1997, nutrient concentrations in the shelf region were all significantly higher than in the beyond shelf region. In 1998, phosphate and silicic acid were significantly higher, while nitrate was not (p>0.05). Nutrient concentrations were higher in 1997 than in 1998, but the differences were significant only for nitrate and silicic acid (p<0.05). Generally as the distance from shore increased, the nutrient concentrations decreased (Figure 1.3-1.5) and in many instances the decreasing concentrations are observed after the shelf break. In April 1997, nutient concentrations did not reach detection limits, while in July and October nutrient concentrations were observed at detection limits in the beyond shelf region. The most obvious feature in 1998 was that  NO3" and HPO4 " 2  concentrations at many transects  were generally at or below detection limits in the shelf and beyond shelf regions during the May cruise and these were the lowest concentrations that were measured for all three nutrients during the two study years.  In July 1998, surface nutrients in the shelf region were higher than in  May, but nutrient concentrations decreased to detection limits as the distance from shore increased. The same pattern was observed in October 1998. There was no consistent seasonal trend for nitrate, phosphate or silicic acid for shelf or beyond shelf regions in 1997 or 1998. The month during which maximum concentrations were measured, varied between region and between years.  One consistent trend was clear; on  average, the lowest nutrient concentration over the two years was in May 1998.  37  April/May  1  20  2  3  4  5  July  6  7  Brooks Peninsula  1  2  3  4  October  5  6  7  1  2  3  4  5  6  7  6  7  Brooks Peninsula  Brooks Peninsula • 4  f  1  2  3  4  5  6  7  1  2  3  4  5  Station Number - A - April 1997 • A - May 1998  July 1997 • G - July 1998  October 1997 October 1998  Figure 1.3 Surface (0-10m) nitrate concentration for all cruises in 1997 and 1998 along transects on La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Shaded area represents the shelf region; solid lines and closed symbols are for 1997 and dashed lines and open symbols are for 1998. No data are available for La Perouse Bank in 1998. The distance offshore increases as the station number increases.  38  July  April/May 3.0  I  .La.P6rpu«JB.ank...  October  !  _  ^..Perouse Bank  I  La Perouse Bank  t  ]  "1 2.0  -4 -  1.5  \  1.0 -I  \  0.5-| 2  0.0  c o  2  c fl) o c o o  £  "] 1.0 —I 0.5-|  8  2  9 10 11 12 14 16  8  9 10 11 12 14 16  Barkley Canyon  B £  2.0  A 12  0.0 3.0  (0  o a.  Barkley Canyon  2  1.5  2.5  sz  9 10 11 12 14 16  -I  2.0  «J a  8  H  3 4 7 8 9101112  Estevan Point  C  1  2  3  4 7  8  9  1011  Estevan Point  1 2 3 4 7 8 9 1011C12  Q  1.51.0  -  \  0.50.0 3.0 2.5-  1  2  3  4  5  6  Brooks Peninsula  7  Q  1  L., 2  3  4  5  6  Brooks Peninsula  7  H  L  Brooks Peninsula i  I  2.0-  I  I  1.5-  !  El  1.0-  m i \  0.5 0.0 1  2  3  4  5  6  7  1  2  3  4  5  6  7  1  2  3  4  5  6  7  Station Number April 1997 May 1998  -Q-  July 1997 July 1998  -EH-  October 1997 October 1998  Figure 1.4 Surface (0-1 Om) phosphate concentration for all cruises in 1997 and 1998 along transects on La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Shaded area represents the shelf region; solid lines and closed symbols are for 1997 and dashed lines and open symbols are for 1998. No data are available for La Perouse Bank in 1998. The distance offshore increases as the station number increases.  39  July  April/May  October  La Perouse Bank  30-  La Perouse Bank  2010 0 2  8  9 10 11 12 14  Barkley Canyon  16  £  2  Estevan Point  20  9  10 11 12 14  16  M  1 2 3 4 7 8 9 101112 30-  8  Q  2  8  9  10 11 12 14  | Barkley.Canyon  1  2  3  4  7  8  9 10 11  M  16  J  1 2 3 4 7 8 9 1011C12  Estevan Point  j£  -  10 0 2 30-|  3  4  5  Brooks Peninsula  6  7  Q  1  2  3  4  5  6  7  1  2  3  4  5  6  7  Brooks |_| PenJnsuia  M  20 10  Q.-O 0 1  2  3  4  5  6  7  Station Number -A— April 1997 • A - May 1998  July 1997 •o- July 1998  October 1997 ••»- October 1998  Figure 1.5 Surface (0-10m) silicic acid concentration for all cruises in 1997 and 1998 along transects on La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Shaded area represents the shelf region; solid lines and closed symbols are for 1997 and dashed lines and open symbols are for 1998. No data are available for La Perouse Bank in 1998. The distance offshore increases as the station number increases.  40  Variability was high for all nutrient concentrations in both the shelf and beyond shelf region. In 1997, variability was highest in the beyond shelf region for all nutrients. In 1998 the region with the highest variability depended on the kind of nutrient. Nitrate was more variable in the beyond shelf region, while phosphate and silicic acid were more variable in the shelf region (Table 1.4). In general, variability was higher in 1998 than in 1997. A3) NORTH- SOUTH GRADIENT There was no consistent north/south gradient in nutrient concentrations in the shelf region or in the beyond shelf region for 1997 or for 1998.  For shelf regions, the highest  concentrations were generally measured over La Perouse Bank in 1997 while in 1998, the location of peak concentrations was variable for each cruise and for each nutrient. For 1997, there was a weak trend for nitrate to decrease north from La Perouse Bank to Estevan Point, then increase again at Brooks Peninsula.  A4) SUMMARY OF NUTRIENT CONCENTRATIONS In summary, significantly higher nitrate and silicic acid concentrations were measured in 1997 than in 1998, but phosphate concentrations were similar for both years. Nutrient concentrations were higher in shelf regions than beyond shelf regions for both years. Generally, the highest nutrient concentrations were measured on La Perouse Bank but on occasion, higher concentrations were measured at Brooks Peninsula.  In May and July 1998, surface nitrate  concentrations were frequently at or near detection limits suggesting either a lower nutrient supply rate or greater nitrate utilization in 1998 compared to 1997.  41  Table 1.4. Mean surface (0-10 m) nitrate, phosphate and silicic acid concentrations (pM) in 1997 and 1998 for shelf and beyond shelf stations of La Perouse Bank, Barkley Canyon, Estevan Point, Brooks Peninsula off the west coast of Vancouver Island. The mean ± 1 S.D. and coefficient of variation (CV, %) for each year and for the 2 year average are given. The number of samples (n) for each transect is given. N D indicates nutrient concentration was not detectable. (-) indicates information not available. * indicates a significant difference between shelf and beyond shelf regions was found at p<0.05 level and w indicates a significant difference between region was found at p<0.01 level. Region Shelf  Nitrate Beyond  Phosphate Shelf Beyond  Silicic Acid Shelf Beyond  n Shelf  Beyond  April 1997  La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula April WCVI mean  9.5 6.1 5.4 8.8 7.5  5.7 6.5 3.3 4.1 4.9  0.8 0.4 0.5 0.5 0.6  0.7 0.6 0.3 0.5 0.5  22 18 15 14 18  14 14 16 11 13.0  3 3 3 2 111  4 3 3 4 114  14.7 6.9 5.0 14.2 10.2  0.3 0.5 1.6 7.8 2.6  1.1 0.9 0.5 1.3 0.6  0.1 0.1 0.2 0.6 0.3  31 18 9 23 20.2  3 3 4 45 6.0  3 3 2 3 111  4 4 2 4 114  3 2 3 3 111 33  2 4 4 3 113 41  4 3 1  4 4 6 114  July 1997  La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula July WCVI mean October 1997  La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula Oct. WCVI mean 1997 Mean 1997 CV  7.1 5.3 2.5 4.2 4.8 n7.5±3.8 50  ND 3.4 0.1 3.8 1.8 H3.1 ±2.7 56  0.7 0.7 0.5 0.6 0.6 n0.7 ±0.3 39  0.7 15 0.5 13 0.2 10 0.5 14 0.5 13.1 n0.4 ±0.2 H17±6.2 54 36  12 10 3.0 12 9.2 n9.4 ±4.8 51  May 1998  Barkley Canyon Estevan Point Brooks Peninsula WCVI mean  3.2 ND 0.3 1.2  0.8 1.1 0.05 0.7  0.3 0.1 0.5 0.2  0.4 0.4 0.3 0.3  5.08 0.27 2.6 2.7  3.4 2.0 3.0 2.8  10 2.3 5.2 6.0  0.1 0.1 2.2 0.8  1.3 0.8 1.1 1.1  0.5 0.6 0.5 0.5  18 12 16 15  2.8 5.6 7.5 5.3  1.1 1.5 1.0 1.2 *0.9±0.5 55 0.8 ±0.3 47  0.4 0.2 0.6 0.4 *0.4±0.1 31 0.4 ±0.1 22  16 16 *10±7.4 72 14 ±5.5 39  3.8 3.8 *4.0±1.9 47 6.6 ±3.5 52  July 1998  Barkley Canyon Estevan Point Brooks Peninsula WCVI mean October 1998  Barkley Canyon Estevan Point Brooks Peninsula WCVI mean 1998 Mean 1998 CV 2 year Mean 2 year CV  3.3 0.8 9.5 5.1 7.1 4.5 6.6 3.5 4.6±3.8 1.6+1.9 82 117 *6.1±2.7 *2.4±1.5 45 62  18 4 3 3  no 4 2 3  19 27  4 4 4 112 4 3 4 111 37  42  III. A)  Biological Parameters (Chlorophyll and phytoplankton species composition) CHLOROPHYLL  For all cruises in 1998, no data are available for the La Perouse transect due to time constraints and sampling logistics.  A l ) MEAN VALUE FOR WEST COAST OF VANCOUVER ISLAND The 2 yr average off the west coast of Vancouver Island was 77.2 ±55.4 mg chl m"  2  (Table 1.5).  On average, biomass was significantly higher in 1998 than in 1997 (p<0.05). In  1998, biomass was 1.6-fold higher than biomass measured for 1997. There was a consistent seasonal peak in July in 1997 and 1998 but the increase in July was more obvious in 1998 than in 1997 (Figure 1.6). Variability was high off the west coast of Vancouver Island but was lower for 1997 than for 1998 (see coefficient of variation in Table 1.5); during both years the highest variability was measured in July. Table 1.5 Mean chlorophyll (100-1% surface light) ±1 S.D. (mg chl m" ) for WCVI during each cruise in 1997 and 1998. Values are the mean of all stations during each cruise. Yearly mean ±1 S.D. and coefficient of variation (CV, %) are included. 2  Chlorophyll 1997 April July Oct. Mean CV  56.3 ±26.4 (23) 77.1+55.2(24) 47.4 ±17.7(26) 60.2 ±37.4 (73) 62  May July Oct. 1998 Mean 1998 C V 97/98 Mean  67.6 ±32.8 (24) 149 ±87.1 (19) 82.9 ±48.9 (21) 99.9 ±67.6 (65) 68 77.2 ±55.4 72  1997 1997 1998  43  250  n=19  200 H  150  ^  H  n=24 n=24  100 H n=26 50 H  April 1997  Julv 1997  October 1997  May 1998  July 1998  October 1998  Figure 1.6 Total chlorophyll ± 1 S.D. off the west coast of Vancouver Island in 1997 and 1998. Values are the mean of shelf and beyond shelf stations during each cruise.  A2) CROSS-SHELF GRADIENT 2 yr and annual means The biomass in shelf regions was on average 2-fold higher than in the beyond shelf regions (Figure 1.7). These plots clearly show higher chlorophyll in the shelf region. The 2 yr average chlorophyll concentration was 101 ±62.9 and 54.0 ±34.7 mg chl m' for shelf and 2  beyond shelf regions, respectively (Table 1.6). On average, chlorophyll was significantly higher in the shelf than in the beyond shelf region in 1997 (p<0.05) and in 1998 (p<0.05). The biomass in the shelf region was significantly higher in 1998 compared to the biomass in the shelf region in 1997 (p<0.05). In the beyond shelf region there was little difference between years.  44  n=33  200 H  E o  O)  E  n=74 150  H n=32  1997 mean  2-yr mean  \Z2 Shelf  m  1998 mean  Beyond Shelf  Figure 1.7 Interannual mean and annual means of total chlorophyll ± 1 S.D. of the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the mean of all shelf and beyond shelf stations sampled. Numbers above each bar are the total number of stations for each mean.  Seasonal variation  Generally, a consistent seasonal pattern was observed in the shelf regions in 1997 and 1998 (Figure 1.8). Biomass was consistently high in July during each year, but the increase in 1998 (2.0-fold) was larger than in 1997 (1.4-fold). Very high increases in biomass in the shelf region from April to July were observed.  For example, at one station in 1998 biomass at  Barkley Canyon increased almost 500% from April to July. In 1998, the biomass in July was significantly higher than May and October (p<0.05), but there was no significant difference between May and October (Tukey test, p>0.05). The seasonal dynamics of biomass in the beyond shelf region were different relative to the shelf region. Biomass in the beyond shelf region showed little seasonal change during 45  cruises in 1997, while in 1998 considerable seasonal variation was observed. In 1998, biomass peaked in July. ;  300 - ,  1997  1997  1997 122  Shelf  —i  1998  1998  1998  B e y o n d Shelf  Figure 1.8 Seasonal variability of total chlorophyll ± 1 S.D. of the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the mean of all shelf and beyond shelf station during each cruise. Numbers above each bar = the total number of station.  Transects Figure 1.9 shows how integrated chlorophyll varies as distance from shore increases. Generally, biomass decreases as the distance from shore increases.  A striking feature  noteworthy in Figure 1.9 was that peaks in biomass were generally observed in 1998 rather than in 1997, specifically for July 1998. In addition, the maximum biomass measured in 1998 was higher than in 1997. For 1998 the maximum Chl a was 428 mg chl m' in the shelf region of 2  Estevan Point (Figure 1.9 G), whereas in 1997, the maximum biomass was 380 mg chl m" for 2  the shelf region of La Perouse Bank (Figure 1.9 E). It should be noted that the July 1998 cruise  46  400-  _ La Perouse Bank  La.P.erouse.Bank  A  A  200-  E  '•  /\  300 •  La Perouse. Bank.  •  10002 400  October  July  April/May  8  9 10 11 12 14  16  Barkley Canyon  2  8  9 10 11 12 14  16  2  8  9 10 11 12 14  16  Barkley Canyon  Barkley Canyon  B  J  F  300  o  200  E  100-  O)  * !»_.  Q  a ®  r-,.J3 • *.  \ G  012  Q.  O vO SZ  400  O  300  iS o  200  3  4  7  8  9  1011  Estevan Point  12  3  4  7  8  9  1011  |- « Estevan Point  c  |  ' . \ • . • j- -"V ''.V :  1  ' ' * . -  »  . b.  a. i  i  1  3 4  2  7  8  9  1011  Estevan Point  G  K  Q *  b  100 0 1 400  2  3  4  5  6  7  Brooks Peninsula  D  1 2  3  4  5  6  7  Brooks Peninsula  300  I  i  200  I  o I  1  a  3  4  5  6  Brooks Peninsula  7  L  " H \  3-0.  100  2  1  3  fa'  0 1  2  3  4  5  6  7  1  2  3  4  5  6  7  1  2  3  4  5  6  7  Station Number —A— April 1997 July 1997 -m—A— May 1998 - o - July 1998  October 1997 October 1998  Figure 1.9 Integrated chlorophyll (mg chl m" ) for all cruises in 1997 and 1998 along transects on La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) on the west coast of Vancouver Island. Shaded area represents shelf region; solid lines and closed symbols are for 1997 and dashed lines and open symbols are for 1998. No data are available for La Perouse Bank in 1998. The distance offshore increases as the station number increases. 2  47  was the only cruise in the two year sampling program that had consistent northwesterly winds (R. Thomson, pers. comm.). This may explain the elevated biomass during the July 1998 cruise. Both regions showed high variability but on average, variability was higher in the shelf region than the beyond shelf region. A3) ALONG-SHORE GRADIENT There was no consistent alongshore trend for the shelf or beyond shelf region in 1997 or 1998 (Figure 1.10). The transect where the highest biomass was measured was different during each cruise. In fact, during one cruise the maximum may be at one particular transect whereas for the next cruise the minimum may be at the same transect. For example, a north-south trend in the shelf region was observed during April 1997 (Figure 1.1 OA). Biomass tended to increase north of La Perouse Bank but then for July 1997, biomass decreased from the southern transect to the northern transect. For October 1997, the difference in biomass between Estevan Point and Barkley Canyon in the shelf region was signifiantly different (Tukey test, p<0.05). The result of an A N O V A test indicates highly significant differences between the transects during July 1998 (pO.Ol).  A  Tukey test showed a significant difference between Estevan Point and both Barkley Canyon and Brooks Peninsula (p<0.05), but no significant difference between Barkley Canyon and Brook Peninsula (p>0.05). A4) SUMMARY OF CHLORPHYLL CONCENTRATIONS In summary, chlorophyll was significantly higher in 1998 compared to 1997. A crossshelf gradient was observed where chlorophyll was significantly higher in the shelf regions than in the beyond the shelf regions. There was no north-south gradient in the shelf or the beyond shelf region for 1997 or 1998. Chlorophyll was consistently higher in July relative to April/May or October for both study regions. Maximum biomass measured for the 2 study years was <20.0  48  mg chl m" , or 428 mg chl m" . During this study period, the highest biomass was measured in 3  2  July 1998, which was the only cruise during the two year study that had conditions favorable for upwelling.  300  beyond Shelf 1997  H  MM  o £  O o ta J?  Beyond Shelf 1998  o £  A/M  LZ]  La Perouse Bank E 3  Barkley Canyon  Z3  Jy  Estevan Point  •  Brooks Peninsula  Figure 1.10 Total chlorophyll of the shelf and the beyond shelf region of the La Perouse Bank, Barkley Canyon, Estevan Point and Brooks Peninsula transect off the west coast of Vancouver Island in 1997 and 1998. A=April, M=May, J=July and 0=October.  49  Table 1.6 Mean integrated chlorophyll ± 1 S.D. (mg chl m") and coefficient of variation (CV; %) for 1997 and 1998 for shelf and beyond shelf stations along La Perouse Bank, Barkley Canyon, Estevan Point and Brooks Peninsula off the west coast of Vancouver Island. Mean for each cruise and each year are given. 2  Date  April 1997  July 1997  Oct. 1997  1997 May 1998  July 1998  Oct. 1998  1998 Mean 2 yr Mean  Region  Chlorophyll Shelf Beyond  La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula WCVI Mean La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula WCVI Mean La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula W C V I Mean WCVI Mean  38.8 ±0.80 61.0 ±13.5 90.4+61.3 93.9 +63.6 71.0 ±34.8 187+122 130+60.5 80.4 ±25.7 38.8 ±7.3 109+53.9 54.4 ±24.2 29.6 ±6.0 78.8+13.3 35.1 ±8.3 49.5 ±13.0 76.5 ±45.8  32.2+12.3 18.9+11.3 59.6 ±22.6 55.2 ±20.3 41.4 ±16.6 36.6 ±15.8 68.4 ±22.9 30.1 45.2 ±38.2 45.1 ±28.2 27.7 38.3+6.80 58.8+47.1 56.6+36.8 45.4+22.7 43.9+15.5  La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula WCVI Mean La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula WCVI Mean La Perouse Bank Barkley Canyon Estevan Point Brooks Peninsula WCVI Mean W C V I Mean WCVI  _  _  Shelf  2.2 22 67 68 38 65 46 32 19 41 45 20 17 24 26 60  CV Beyond  38 60 38 37 43 43 33  85 40  18 80 65 54 35  n Shelf  Beyond  2 4 4 2 112 5 3 3 3 114 5 3 4 3 115 41  3 3 3 2 111 2 4 1 3 110 1 3 3 4 111 32  _  _  _  _  81 36  6 4 1  4 3 6 113  39  37 35 23 31  -  -  -  -  -  -  94.1 ±28.4 306 ±88.0 147 ±66.6 182+61.0  54.6 ±11.1 172 ±103.8 121 ±49.0 116 ±54.6  30 29 45 36  20 60 40 40  4 3 3 110  4 3 4 19  113 +91.5 61.5+22.3 95.4 89.8 ±37.9  29.8 34.8 71.2 45.3  ±11.0 ±12.1 ±16.0 ±13.0  -  111  -  -  -  -  -  -  103 ±53.8 120+64.2 152+62.5 125.0 ±60.2 132+70.7 101 ±62.9  37.2 +14.7 48.1 ±10.4 37.0 ±6.80 40.8+10.6 67.4 ±48.3 54 ±34.7  55 54 41 50 53 63  40 22 18 27 39 64  4 3 3 111 33 74  4 2 4 HO 32 64  50  B) PHYTOPLANKTON  ASSEMBLAGES  Phytoplankton taxonomic analysis was completed at the same stations that primary productivity were measured. At a single station in each of the shelf and beyond shelf regions was two samples collected, one at each of the 55% and 1% surface light level.  A list of all  species found during 1997 and 1998 are listed in Tables 1.7 and 1.8. The category 'diatoms' includes all species listed in Table 1.7 and photosynthetic flagellates refers to all species listed in Table 1.8.  Abundances of diatoms, nanoflagellates, and dinoflagellates for 1997 and 1998  are included in Table 1.9 and 1.20.  Bl) TOTAL ABUNDANCE AND BIOMASS On average, the total cell abundance was higher in 1997 (6.1 xlO cells L" ) compared to 6  1  1998 (4.7 xlO cells L" ), in contrast to higher biomass in 1998 compared to 1997 (Figure 1.11). 6  1  The decrease in cell abundance in 1998 was due to fewer nanoflagellates and the increase in biomass in 1998 was due to increased presence of diatoms. Total cell abundance and total biomass were consistently higher in shelf regions than beyond shelf regions (Figure 1.11). For 1998, the difference in cell abundance and biomass between the two regions was significantly different (p<0.05). A seasonal trend was evident for total cell abundance for 1997 and 1998 (Figure 1.12). Generally total abundance peaked in July. For total biomass, a consistent seasonal trend was not observed for 1997 and 1998. In 1997, biomass peaked in July while in 1998 biomass peaked in May.  51  8-1  ¥2  Shelf  m  Beyond Shelf  Figure 1.11 Interannual mean and annual means of total cell abundance and phytoplankton biomass ± 1 S.D. for the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf values are the mean of all shelf stations sampled and beyond shelf values are the mean of all beyond shelf stations.  52  n=4  12 H 0>  o ^ c at  <° =  o  O  10 H n=4 8H  6H 4 H  2H  April a) (/>  1400  E  1200  n  o m c  n=3 n=3  ^  800  2  600  Q. O  '  Oct.  '  May  '  July  Oct  July 1998  Oct 1998  1000  S O "  c  July  400  H  n=4  n=3 n=3  200  April 1997  July 1997 22  n=4 n=2  Oct 1997 Shelf  May 1998  Beyond Shelf  Figure 1.12 Seasonal variability of total cell abundance and phytoplankton biomass ± 1 S.D. for the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf meansare the average of all shelf stations sampled during each cruise and beyond shelf values are the mean of all beyond shelf stations during each cruise. B2) COMMUNITY STRUCTURE The phytoplankton community of shelf and beyond shelf regions were generally numerically dominated by nanoflagellates during the study period (Figure 1.13A). Generally nanoplankton were mainly composed of unidentified miscellaneous flagellates, but peaks of  53  Mantoniella  squamata, Micromonas  pusilla  and  coccolithophores  were  observed.  Chrysochromulina spp. and Crypotomonas spp. were observed but they never dominated the nanoflagellate assemblage.  B) Total Phytoplankton Biomass  ES)  Heterotrophic Dinoflagellates  •  Photosynthetic Dinoflagellates  ZZi  Diatoms  E3  Nanoflagellates  Figure 1.13 Contribution of each phytoplankton group to: A) total cell abundance, and B) total phytoplankton biomass (ug L" ) of the shelf and beyond shelf region off the west coast of Vancouver Island for 1997, 1998 and the 2 yr mean. 1  54  Despite their numerical dominance, they contributed substantially less to total biomass due to their small size (Figure 1.13B). For 1998, a change in community structure was observed. The contribution of diatoms to total abundance and biomass increased significantly in 1998 from 1997 (p<0.05) (Figure 1.13). The increase was particularly notable for May and July 1998 (Figure 1.14).  The diatom assemblage was frequently composed of Pseudo-nitzschia spp  Chaetoceros spp.,  Skeletonema costatum, Leptocylindrus danicus, Detonula pumila,  Asterionella glacialis. The contribution of diatoms to abundance and biomass was less in the beyond shelf region. The relative contribution of diatom to total biomass for the October 1998 cruise was low, which was expected since the cruise occurred after the fall transition had occurred. Three obvious features of the relative contribution of the different phytoplankton groups are: 1) diatoms tend to account for a high proportion of total biomass in April/May and July, particularly for the shelf region, 2) the contribution by photosynthetic dinoflagellates, dominated by Gymnodinium spp. tends to increase in the fall (Figure 1.14)  and 3) heterotrophic  dinoflagellates dominated by Gyrodinium spp., contributed the least to total cell abundance during the study period. The variability in cell abundance was higher for diatoms than for nanoflagellate during both study years.  The variability of autotrophic and heterotrophic dinoflagellates abundance  was high. The variability in biomass was higher in diatoms than nanoflagellates during 1997, while for 1998 the variability was higher for nano flagellates. Mesodinium rubrum is a ciliate that contains chloroplasts (Taylor et al, commonly blooms in upwelling regions.  1971) and  Generally the abundance was higher in the shelf  region compared to the beyond shelf region; 4% of the total biomass was accounted for by Mesodinium rubrum in the shelf region compared to 1 % in the beyond shelf region. Abundance 55  and biomass were higher in 1997 relative to 1998. On average, they accounted for 5% of the biomass for 1997 and 1% for 1998.  Generally, the contribution of Mesodinium rubrum to  abundance and biomass was low during the study period.  A) Shelf  1997  B) Beyond Shelf  1998  1997  1998  EiSl  Heterotrophic Dinoflagellates Photosynthetic Dinoflagellates Diatoms t-V'-Vl Nanoflagellates Hi  Figure 1.14 Contribution of each phytoplankton group to total cell abundance and biomass of: A) shelf and B) beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Groups contributing <2% were not included.  56  B3) SUMMARY OF PHYTOPLANKTON ASSEMBLAGES In summary, phytoplankton biomass was higher in 1998 due to increased abundance of diatoms.  Diatom blooms in 1998 were mainly composed of Chaetoceros debilis and  Leptocylindrus danicus. The phytoplankton biomass was higher in shelf regions than beyond shelf regions for all cruises and all transects. The biomass of diatom species dominated the shelf regions during the spring and summer cruises and autotrophic flagellates dominated the biomass during October. The contribution of autotrophic and heterotrophic dinoflagellates was generally low, but particularly during the fall, the relative contribution to abundance and biomass increased. A change in community structure was observed in 1998, with blooms of diatoms that were not observed during the 1997 season.  57  Table 1.7 List of diatoms identified from samples collected off the west coast of Vancouver Island from May 1997 to October 1998. X in 1997/1998 column signifies the diatom was observed in either April/May, July or October. 1998 1997 Bacillariophyceae Species X X Chaetoceros spp.* Centric Diatoms X X Chaetoceros compressus Chaetoceros convolutus X X X X Chaetoceros debilis X Chaetoceros eibenii X X Chaetoceros radicans X Chaetoceros socialis X Detonula pumila Leptocylindrus danicus X X Leptocylindrus minimus X X Proboscia alata (formally Rhizosolenia alata) X X Dactyliosolen fragilissimus (formally X X Rhizosolenia fragilissima) Rhizosolenia setigera X Guinardia striata X X (formally Rhizosolenia stolterfothii) Asterionella glacialis Cylindrotheca closterium Fragilaria spp. Navicula spp. Nitzschia spp. Pseudo-nitzschia spp.* Pseudo-nitzschia delicatissima * Pseudo-nitzschia pungens Thalassionema nitzschoides Thalassiosira aestivalis Thalassiosira spp. * Thalassiosira rotula Thalassiosira nordenskioeldii Skeletonema costatum* Synedra spp. * Most abundant species  Pennate Diatoms  X X X X X X X X X X X X  X X X X X X X X X X X X X  Table 1.8 List of autotrophic flagellates identified from samples collected off the west coast of Vancouver Island from May 1997 to October 1998. X signifies species was observed in either April/May, July or October. 1997 1998 Species X Alexandrium tamarense Dinophyceae X Ceratium kofoidii X X Gymnodinium spp. X Gyrodinium fusiforme X X Gyrodinium spp. X X Katodinium rotundatum X Prorocentrum balticum X X Prorocentrum gracile X Protoperidinium rhomboidalis X Protoperidinium spp. X X Chrysochromulina spp.* Prymnesiophyceae X X Coccol ithophores X Dictyocha speculum Chrysophyceae X X Prasinophysceae Micromonas pusilla X X Mantoniella squamata X Cryptophyceae Leucocryptos marina X Cryptomonad spp. X X Mesodinium rubrum Ciliate * Most abundant species  Table 1.9. Abundance of diatoms, nanoflagellates, autotrophic and heterotrophic dinoflagellates during April, July and October 1997 off the west coast of Vancouver Island. LP=La Perouse Bank BC=Barkley Canyon, EP=Estevan Point, BP=Brooks Peninsula. See Figure 1.1 for location of transects. (—) indicates no sample taken, * indicates the most abundant group. Region  Nanoflagellates  Diatoms  Autotrophic Dinoflagellates 10 Cells r  Heterotrophic Dinoflagellates 10 Cells r  10 Cells 1  10 Cells I  0.96 0.23 1.52 0.02 14.2 2.67 13.4 5.13 7.5 2.0  46.1 * 12.2 * 96.6* 85.4* 33.3 * 75.4* 67.1 * 24.6* 60.8 * 49.4*  0.16 0.08 0.76 0.02 0.25 0.49 0.17 0.78 0.34 0.34  0.08 0.09 0.41 0.14 0.16 0.24 0.43 0.15 0.27 0.16  52.8* 0.01 15.7 8.36 0.38 2.38 25.6 0.92  37.2 4.58* 86.0* 33.2 * 81.4* 40.4* 52.2* 42.1 *  0.23 0.02 0.22 0.52 0.10 0.34 0.32 0.15  0.19 0.02 0.16 0.51 0.13 0.28 0.29 0.14  1.80 0.72 0.31 10.2 0.32 1.68 3.26 1.0 12.1 1.3  28.4* 7.67* 16.4* 35.8 * 64.3 * 47.8* 34.1 * 32.1 * 49.0* 41.0*  0.20 0.05 0.06 0.49 0.32 0.21 0.26 0.13 0.31 0.20  0.19 0.12 0.11 0.34 0.34 0.77 0.24 0.43 0.27 0.25  s  1  s  1  s  1  s  1  April 1997  LP-Shelf LP-Beyond Shelf BC-Shelf BC-Beyond Shelf EP-Shelf EP-Beyond Shelf BP-Shelf BP-Beyond Shelf Mean-Shelf Mean-beyond July 1997  LP-Shelf LP-Beyond Shelf BC-Shelf BC-Beyond Shelf EP-Shelf EP-Beyond Shelf BP-Shelf BP-Beyond Shelf Mean-Shelf Mean-beyond October 1997  LP-Shelf LP-Beyond Shelf BC-Shelf BC-Beyond Shelf EP-Shelf EP-Beyond Shelf BP-Shelf BP-Beyond Shelf Mean-Shelf Mean-Beyond 1997 Mean-Shelf 1997 Mean-Beyond  Table 1.10 Abundance of diatoms, nano flagellates, autotrophic dinoflagellates and heterotrophic dinoflagellates at 55% surface light depth during May, July and October 1998 off the west coast of Vancouver Island. LP=La Perouse Bank BC=Barkley Canyon, EP=Estevan Point, BP=Brooks Peninsula. See Figure 1.1 for location of transects. * = most abundant group. Region  Nanoflagellates  Diatoms 10 Cells r 5  1  10 Cells I s  1  Autotrophic Dinoflagellates 10 Cells r s  1  Heterotrophic Dinoflagellates 10 Cells r s  1  May 1998  LP-Shelf LP-Beyond Shelf BC-Shelf BC-Beyond Shelf EP-Shelf EP-Beyond Shelf BP-Shelf BP-Beyond Shelf Cruise Mean-shelf Cruise Mean-beyond  46.8* 0.01 6.99 0.003 1.20 74.4* 15.2 8.55 17.5 * 20.7*  3.32 43.6* 8.34 * 6.21* 30.3 * 34.7 15.6* 11.5 * 11.7 10.0  0.52 0.13 0.04 0.01 0.07 1.62 0.02 0.02 0.16 0.41  0.44 0.06 0.03 0.02 0.64 1.69 0.06 0.12 0.29 0.47  8.19* 0.02 102 * 0.01 74.4 0.01 81.2 7.44 66.4* 1.86  3.32 2.81 * 43.9 0.21 26.9* 8.29* 12.5 33.0 * 21.7 9.06*  0.04 0.01 0.05 0.02 1.62 0.02 0.91 0.15 0.78 0.05  0.10 0.18 0.58 0.04 1.68 0.04 1.08 0.23 0.86 0.12  0.55 0.39 19.0* 0.09 1.91 0.18 1.18 0.34 5.66 0.24 29.9* 7.6  4.55 * 37.7* 15.0 1.18* 25.4* 17.0 9.15* 3.54 * 13.5 * 14.9* 15.6 11.3 *  1.45 0.09 0.12 0.01 5.87 0.48 5.46 0.06 3.22 0.16 1.40 0.20  0.34 0.28 0.35 0.05 0.57 0.17 0.38 0.05 0.41 0.13 0.52 0.24  July 1998  LP-Shelf LP-Beyond Shelf BC-Shelf BC-Beyond Shelf EP-Shelf EP-Beyond Shelf BP-Shelf BP-Beyond Shelf Cruise mean-shelf Cruise mean-beyond October 1998  LP-Shelf LP-Beyond Shelf BC-Shelf BC-Beyond Shelf EP-Shelf EP-Beyond Shelf BP-Shelf BP-Beyond Shelf Cruise mean-shelf Cruise mean-beyond 1998 mean-shelf 1998 mean-beyond  61  Table 1.11 Siurirnary of characteristics of shelf and beyond shelf regions off the west coast of Vancouver Island. Values are for 1997 and 1998. Units for parameters below are: M L , meters; temperature, °C, nitrate, uM; chlorophyll, mg chl m" ; C V , %) Shelf Beyond 2  Mixed layer depth Mixed layer temperature Mixed layer salinity 1997 N0 1998 2 yr mean HP04 1997 1998 2 yr mean 1997 Si(OH) 1998 2 yr mean Total Chl surface range 1997 1998 2 yr mean CV % 1997 1998 2 yr mean CV 1997 % Diatoms 1998 2 yr mean CV 3  4  11.8 11.9 31.1 7.5 4.6 6.1 0.7 0.9 0.8 16.9 10.3 13.8 0.21-19.7 76.5 132 104 63 33 2 18 75 38 65 52 8  23.2 12.8 31.5 3.1 1.6 2.4 0.4 0.4 0.4 9.4 4.0 6.6 0.12-12.7 43.9 67.4 55.7 64 41 32 37 14 24 40 32 7  DISCUSSION West coast of Vancouver Island Variability was very high off the west coast of Vancouver Island, and often the standard deviation was similar to the mean.  This makes it difficult of determine the significance of  differences between regions and between years and therefore this thesis also contains comments on trends or patterns.  Landry et al. (1989) have also shown that means of temperatures,  salinities, winds, currents, nitrate and chlorophyll are only known within large limits of uncertainty. They also pointed out that the variability of these parameters was an important ecological parameter in its own right. Nutrients and biomass off the west coast of Vancouver Island were generally high but on several occasions and at several locations nutrient concentrations were at or near detection limits.  Surface nitrate up to 14-15 p M was observed but on average nitrate, phosphate and  silicic acid concentrations for the west coast of Vancouver Island were 4.1, 0.7 and 12 p M respectively for the study period. A maximum of 306 mg chl m" was observed and on average 2  chlorophyll was 77.2 mg chl m" off the west coast of Vancouver Island. 2  Generally, the mixed layer depth was deeper and more saline in 1998 for both the shelf and the beyond shelf regions.  This is reasonable considering more intense upwelling was  observed in 1998, which would inject high salinity water into the surface layers.  The mixed  layer temperatures were on average similar for both years. On average, biomass was lower in 1997 than in 1998 while on average nutrients were higher in 1997 than in 1998.  It is likely that  the high phytoplankton biomass observed in 1998 was responsible for drawing down nutrient concentrations. This scenario is even more plausible considering the high intensity of upwelling and high flux of nutrients to the surface.  Upwelling as a nutrient source should be higher in  1998 than in 1997. Depletion of surface NO3" in 1998 suggests primary productivity may have 63  been nutrient limited during periods. The seasonality of nutrient concentrations varied between years. In 1997, nitrate and silicic acid were highest in April and lowest in October, whereas in 1998 concentrations were the lowest in April and the highest in October. A distinct change in phytoplankton community structure occurred in 1998.  During the 1997 cruises, the relative  contribution of diatoms was low, but during the 1998 season the diatoms returned and contributed significantly to total biomass.  This suggests that during periods of intense  upwelling the fast growing diatoms outcompete other species. The various physical, chemical and biological parameters off the west coast of Vancouver Island varied considerably in the cross-shelf direction (E/W). shelf and beyond shelf parameters see Table 1.11.  For a summary of  The mean mixed layer depth was deeper,  warmer and more saline beyond the shelf than in the shelf region. Nutrient concentrations and biomass were higher in the shelf region than in the beyond shelf region.  The phytoplankton  community structure of shelf and beyond shelf regions was distinct from each another. The diatoms had a greater contribution to total biomass in the shelf region than in the beyond shelf region. Generally the diatom assemblage was composed of Pseudonitzschia spp., Chaetoceros debilis, Skeletonema costatum, Asterionella glacialis, Dactyliosolen fragillissimus, but was dominated by Chaetoceros debtiis and Leptocylindrus danicus. The phytoplankton assemblage in the beyond shelf regions were composed of Chrysochromulina spp., and cryptomonads, but it was generally dominated by miscellaneous unidentified flagellates. In addition to cross shelf differences, physical, chemical and biological parameters varied considerably in the along shore directions (N/S). There was no consistent trend along the length of Vancouver Island for the shelf region or the beyond shelf region. The high variability in the cross-shore direction and along shore direction suggests environmental conditions vary considerably off the west coast of Vancouver Island. 64  The distribution of nutrients and phytoplankton biomass off the west coast of Vancouver Island showed considerable variation in time. Seasonality of biomass was similar in 1997 and 1998. The seasonal peak in biomass and diatom abundance for each year was seen in July for 1997 and 1998, while the highest biomass during the study period were measured in July 1998. During May and July 1998 the relative contribution of the diatoms reached 91 and 86% of total cell biomass. It is interesting to note that during the entire study, the July cruise was the only cruise that occurred during upwelling favorable conditions.  Comparison with previous studies off the west coast of Vancouver Island A distinct cross-shelf gradient was observed during this study.  This finding confirms  previous results (Mackas, 1992) which showed a general cross-shelf gradient in physical, chemical and biological properties. The shelf was characterized by cooler temperature, lower salinity, higher dissolved nutrients, and higher biomass than the beyond shelf region. The lower salinity reflects the strong regional influence of the Vancouver Island Coastal Current (Freeland et al. 1984; Thomson et al. 1989). The results of this study are consistent with those reported by Mackas (1992). Denman et al. (1989) found that continental shelf waters are generally retained along the continental margin which may in part, explain the distinct characteristics of shelf regions and beyond shelf regions observed during this study. Nitrate concentrations reported by Mackas (1992) were similar to those found in this study.  One different feature observed during this study was for May and July 1998 nitrate  concentrations were frequently at or near detection limits, whereas Mackas (1992) found surface nitrate was very rarely depleted to undetectable levels. This difference may be explained by low upwelling during the strong E l Nino. Chlorophyll concentrations reported by this study were similar to those reported by Mackas (1992), both in mean concentration and seasonality. 65  Comparison of the taxonomy data with other studies is limited since there have been no detailed taxonomic studies off the west coast of Vancouver Island, although Taylor and Haigh (1996) have examined the microplankton community structure of Barkley Sound. They found that the summer community was dominated by diatoms, typical of coastal waters. These observations are similar to results presented in this thesis and similar to results reported by Mackas et al. 1980; Perry et al. 1999. Taylor and Haigh (1996) reported over 14 harmful phytoplankton species in Barkley Sound. This study did not examine the distribution of harmful algal species specifically, but Alexandrium tamarense, Chaetoceros convolutus, and  Pseudo-nitzschia delicatissima,  Gymnodinium auratum, Pseudo-nitzschia pungens were all observed during the study period. In addition, Taylor and Haigh (1996) suggested that strong dinoflagellate blooms are more common in the fall.  This study has shown that the contribution of dinoflagellates was the  highest in the fall of 1998.  Comparison with other upwelling regions The nutrient and biomass distribution off the west coast of Vancouver Island are similar to the coastal upwelling areas off Washington and Oregon (Landry et al. 1989) and off California (Wilkerson et al. 2000). In general, nutrient and biomass concentrations were high (>5 uM) and a sharp cross-shelf distribution of nutrients and biomass were observed. Small and Menzies (1981) have shown that the highest biomass developed within 20 km of the shore. This is consistent with the results of this study. Landry et al. (1989) reported that concentrations of nitrate are replenished in the summer after the seasonal low in April caused by the spring bloom. During 1998, the year of intensive upwelling, this study has shown that nitrate was replenished after low concentrations found in May. 66  Low or undetectable levels of nutrients were found during 1998 which differs from previous studies (e.g. Mackas, 1992) that found detection limits were rarely observed off the west coast of Vancouver Island.  Lower nutrient concentrations during the E l Nino year  compared to non-El Nino years are reported by Barber and Chavez, 1983; Barber and Chavez, 1986; and Wilkerson et al. 1987). It is possible that the strong E l Nino during 1997/98 may have reduced the winter and summer nutrient supply to the shelf region allowing for rapid depletion of a smaller nitrate pool resulting in undetectable nitrate concentrations. Off the coast of Washington and Oregon depletion of nitrate was common, particularly after the spring bloom in April (Landry et al. 1989). The higher contribution of diatoms to the abundance and biomass during May and July 1998 is consistent with results found by Chavez (1996). Generally during upwelling conditions, he found high diatom abundance that dominated the phytoplankton biomass.  SUMMARY OF CHAPTER ONE In general, the physical, chemical and biological parameters showed interannual variability.  Differences between the two sampling years include: shallower mixed layer in  1997; lower salinity and density in 1997; higher nitrate and silicic acid concentration in 1997; lower chlorophyll in 1997; higher phytoplankton abundance in 1997, lower phytoplankton biomass and lower diatom abundance and biomass in 1997. There was no difference between mixed layer temperature between 1997 and 1998.  During 1997 nanoplankton dominated the  community structure while in 1998, diatoms dominated. This suggests that the 1997 oceanic environment was not favorable for large cells, and was perhaps due to El Nino conditions.  67  CHAPTER 2 SIZE-FRACTIONATED BIOMASS AND PRIMARY PRODUCTIVITY OFF THE WEST COAST OF VANCOUVER ISLAND  Introduction There is increasing evidence of synchronous changes in the climate and the ocean environment in the Pacific Ocean (Beamish et al., 1999). Large fluctuations of Pacific salmon stocks, shifts in mesozooplankton abundance (Brodeur and Ware, 1992; Sugimoto and Tadokora, 1997), chlorophyll (Venrick et al. 1987) and nutrient distribution (Whitney et al. 1998) in this century are linked to changed in the climate and to physical changes in the ocean. It is reasonable to assume that if trends in climate match trends in fish, zooplankton and chlorophyll, then primary productivity may be impacted as well.  A persistent trend in  climate/ocean conditions is called a regime (Beamish et al. 1999b). It is important to recognize that physical and biological mechanisms may change when regimes shift. The regime shifts are important in fisheries management because the natural shifts may be large and sudden, requiring that these natural impacts be incorporated into management plans. In order to assess the possible climatic impact on ecosystem productivity, long term data bases are necessary because it is difficult to assess decadal trends when the database is less than 20 years. Often observations fail to examine critically important environmental disturbances such as ENSO (El Nino-Southern Oscillation) events. The measurement of size-fractionated biomass and primary productivity is important to understand the response of the phytoplankton assemblage to fluctuating environmental conditions.  A better understanding of the community response  understanding of the functioning of the pelagic foodweb.  will further aid our  This is the first study of size-  68  fractionated biomass and primary productivity off the west coast of Vancouver Island.  In  addition, this study examines a large fluctuation in the ocean regime and its impact on phytoplankton communities during 1997 (strong El Nino) and 1998 (strong La Nina).  The goals of this study were to: (1) investigate the size structure of the phytoplankton biomass and primary productivity along the west coast of Vancouver Island, (2) determine if increases in biomass and primary productivity are due to the larger size fraction, and (3) determine if the size structure or total primary productivity was impacted during the strong ENSO event of 1997 and 1998.  MATERIALS AND METHODS Total and size-fractionated phytoplankton biomass and primary productivity in the euphotic zone were investigated on 4 transects over the continental margin off the west coast of Vancouver Island. At each of the four transects, sampling stations were chosen in order to have one station on and one station off the continental margin, making a total of 8 stations (Figure 2.1).  The transects were off La Perouse Bank, Barkley Canyon, Estevan Point and  Brooks Peninsula.  Size-fractionated chlorophyll and size-fractionated primary productivity  were measured on water samples collected at 6 depths corresponding to 100, 55, 30, 10, 3.5, and 1% of the surface light intensity (I ). 0  69  Chemical and biological measurements Water samples for nitrate + nitrite (NO3" + NCV), soluble reactive phosphate (HPO4 '), and silicic acid (Si(OFf)4) were collected at 6 depths in the euphotic zone and processed as outlined and discussed in Chapter 1.  Figure 2.0 Location of study area off the west coast of Vancouver Island. Dashed line is the 200 m contour. The box delineates the study area. C I , LC4, LG3, BP2 are shelf stations and L B 16, LC9, LG9, BP7 are beyond shelf stations. CI and L B 16 =La Perouse Bank transect, LC4 and LC9 = Barkley Canyon transect, LG3 and LG6 = Estevan Point transect and BP2 and BP7 = Brooks Peninsula transect.  70  Duplicate samples (0.5 L) for size-fractionated chlorophyll concentration (corrected for phaeopigments) were collected at 6 water depths in the euphotic zone and processed using parallel filtration. Water samples were filtered through 25 mm 0.7 pm glass-fiber filters (Whatman GF/F or A M D GF75) for total chlorophyll and through a 47 mm Nuclepore® 5 pm polycarbonate filter for the >5 pm fraction. Samples were extracted for chlorophyll and calculations were made according to the methods outlined in Chapter 1.  The <5 pm size  fraction was calculated as the difference between total chlorophyll and the >5 pm chlorophyll sample. To evaluate the contribution of the >5 pm sized cells to the phytoplankton biomass, the >5 pm size fraction values were calculated as percentages of the total phytoplankton biomass.  Primary productivity During each cruise, an attempt was made to occupy process stations before dawn but sampling logistics and weather conditions sometimes prohibited this plan. Water was sampled using 10 L acid-cleaned P V C Niskin bottles fitted with silicone O-rings and Teflon-covered closure springs mounted on an instrumented rosette equipped with a Biospherical Instruments 4 7t light sensor. The PAR sensor trace was examined on the downcast to identify the six water depths corresponding to 100, 55, 30, 10, 3, 1% of incident surface photosynthetic photon flux density (PPFD).  Samples were transferred directly from the Niskin bottle to 70 ml acid-  cleaned polycarbonate (Nalgene®) incubation bottles without using a siphon tube to prevent contamination (Price et al. 1986). The samples were maintained under low light conditions during all manipulations until the start of the incubation, usually within 1 h of sampling. Triplicate samples were taken for all depths except for the April 1997 samples where duplicates  71  were only taken at two depths.  Triplicate dark bottles were collected at the surface and  incubated with the 100% I samples. A time zero sample was collected from the surface and 0  filtered immediately after inoculation with NaH CC»3 onto each of a 0.7 um and 5.0 um filter. 14  The N a H C 0 1 4  3  stock was stored refrigerated (5°C) but allowed to come to ambient  temperature before inoculation of the samples. Samples were inoculated with 0.37 M B q (10 uCi) of N a H C 0 1 4  3  New England Nuclear (NEC-086H) and incubated under natural light  conditions in on-deck Plexiglas® incubators using neutral density screening to simulate the light levels from which the water was taken. The 100% I incubation bottles were placed in 0  clear polycarbonate bags. No attempt was made to correct for spectral properties of incident light. Recirculating surface water controlled temperature within ±2°C. All incubations were ~6 h in duration except during May 1997 when incubations were 24 h in length. A 100 ul sample of isotope stock was taken and added directly to scintillation vials containing 100 ul of ethanoalamine (Sigma Chemical Co.) which prevents H C 0 1 4  escaping to the atmosphere.  3  from  This subsample was used to determine the total activity of  1 4  C  added (DPM t). The incubations were terminated by gravity filtration through a cascade of a to  47 mm, 5 um Nuclepore® polycarbonate filter and then filtered through a 25 mm, Whatman® GF/F 0.7 um filter using <100 mm Hg vacuum differential (Joint and Pomroy, 1983). The filters were gently sucked dry, rinsed with 15 ml of filtered seawater, folded and placed in a 20 ml scintillation vial.  250 ul of 0.5 N HC1 was added to each vial to eliminate the  unincorporated inorganic N a H C 0 and the vials were placed uncapped in the fumehood until 1 4  3  the filters were dry (approx. 24 h).  The samples were stored in the dark until processing  ashore.  72  At the onshore laboratory, 10 ml of Readysafe containing the filters.  scintillation fluor was added to the vials  The vials were capped and stored in the dark for >24 h before the  samples were counted on a Beckman® LS 6000 series liquid scintillation counterl Primary productivity was determined from the amount of  1 4  C incorporated into i  particulate organic carbon and retained on a filter (Steemann-Nielsen,  1952; Parsons et al.  1984). Rates were calculated according to Parsons et al. (1984) to obtain mg C m" h" . A time 3  zero blank correction was used for all calculations to correct for  1 4  1  C adsorption to the filters.  Hourly primary productivity rates were converted to daily productivity by dividing the primary productivity by the percentage that the incubation period represented of the total daily irradiance.  Light data were not available in October 1998 due to a datalogger failure. For  conversion of October 1998 hourly primary productivity to daily primary productivity, the light data from October 1997 were used. Vertically integrated primary productivity was calculated by averaging the measured productivity between two depths and multiplying by the depth difference (Ichimura et al. 1980). The sum of these measurements yielded the hourly production integrated over depth and expressed as mg C m" h" . To evaluate the contribution of the >5 pm 2  1  cells to the primary productivity, the >5 pm size fraction values were calculated as percentages  i of the total primary productivity. Dark bottles productivity rates were calculated but no dark bottle corrections were applied to the productivity values reported (Banse, 1993). The carbon assimilation rate is the photosynthetic rate per unit of chlorophyll and was calculated by dividing the hourly productivity rates (mg C m" h") by the chlorophyll 3  concentration (mg chl m" ). 3  1  The integrated assimilation number was calculated by dividing  integrated hourly productivity (mg C m" h") by integrated chl concentration (mg chl m" ). 2  1  2  73  Statistical analysis of chemical and biological data  j  Replicate casts were not completed due to time constraints in the cruise schedule and the labor-intensive nature of this study.  Replicate (and sometimes triplicate)!  s a m  pl  e s  w  e  r  e  routinely collected from each depth (i.e. from the same water sample) for analysis of sizefractionated chlorophyll and primary productivity. Replicate samples were oniy collected on glass fiber filters during April and July 1997.  For the remainder of the cruises, replicate  samples were collected on glass fiber filters and polycarbonate filters. The mean of the replicates is reported in this chapter.  j  i One factor analysis of variance (ANOVA) and a Tukey test were used to examine !  spatial and temporal variation. For analysis of temporal variation, interannual and seasonal, the physical, chemical and biological data were grouped according to mean values} for the WCVI i and shelf and beyond shelf regions. For analysis of spatial variation, (i.e. the cross shelf and along shore direction), the data were grouped according to cruise and year.  i j I  RESULTS  |  The following sections will examine the distribution of biomass and productivity in  j space and time. Three spatial scales will be examined: the mean of all stations will be referred to as the mean for the west coast of Vancouver Island (WCVI); how biomass and productivity  | varies in a cross-shelf manner (West/East fashion) and how shelf and beyond shelf stations  j  vary in an alongshore manner (from the southern La Perouse transect to the northern Brooks i Peninsula transect). Three time scales will also be examined for each spatial scale: a 2-yr mean (1997 and 1998); annual mean for 1997 and 1998; and a cruise mean.  I  Very high variability was observed for chlorophyll and primary productivity off the west coast of Vancouver Island.  Trends and patterns will be discussed but the differences 74  between regions, cruises, or years were not statistically significant unless otherwise noted.  A) Size-fractionated chlorophyll Size-fractionated chlorophyll analysis allows one to determine which size fraction of  i phytoplankton was responsible for the measured changes in total chlorophyll. Integrated data ±1 S.D. from all stations sampled during 1997 and 1998 for size-fractionated chlorophyll were averaged for each cruise and for all 6 cruises and are given in Appendix J. The data are also stored in the Institute of Ocean Sciences Sidney, B C database.  Vertical profiles of size-  fractionated biomass are included in Appendix K.  \  A1) MEAN VALUES FOR WCVI Annual and interannual means (WCVI) The >5.0 pm size fraction contributed substantially to biomass on the WCVI (Figure 2.1). On average over the two year study period, the >5 pmfractionaccounted i for 46% of the  I 2i total chlorophyll. The 2-yr mean biomass for all cruises was 31.0 mg chl m jfor the <5 pm i fraction and 39.8 mg chl m" for the >5 pm fraction. 2  In 1997, biomass was higher in the <5 pmfractionthan the >5 pmfraction(Figure 2.1). The mean chlorophyll concentration for 1997 was 34.9 ±19.9 for the <5 pm fraction and was 30.6 ±35.7 mg chl m" for the >5 pm fraction. The relative contribution of the >5 pm fraction 2  was greater in 1998 than in 1997.  The mean chlorophyll concentration for 1998 was 27.1  i ±15.7 in the <5 pm fraction relative to the 48.9 ±58.8 mg chl m for the >5 pm fraction. In •  i  1997, the >5 pm fraction accounted for 41% of the total chlorophyll, while in j 1998 the large cells accounted for 50% of the total chlorophyll. ji  75  140 H 120 H  100 80 60 H 40 20 H  2-yr mean  "  E2  1997 mean  <5um  "  1998 mean  E53 > 5 um  Figure 2.1 Interannual mean and annual means of size-fractionated chlorophyll ± 1 S.D. for the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf values are the mean of all shelf and beyond shelf stations, respectively. Numbers on the top of each panel represent the percentage of the total chlorophyll that was accounted for by the >5 um fraction.  76  Seasonal means (WCVI) The contribution of the >5 umfractionwas highest in April for 1997, while for 1998 the contribution of the >5 (amfractionwas highest in July (Figure 2.2). The increase contribution seen in July 1998 was due to an increase in phytoplankton >5 um since there was no seasonal change in biomass in the <5 (am size fraction.  44%  48%  ,  32%  ,  41%  „  61% n=8  160 £  48%  140  o.  S O O  120 p  •o _  Si -=  100  «  o  |  f> 80  n=6  n=8  n=4  n=8  o  2  T  n=4  60  n=8  n=6  n=8 n=8  a> N  W  20  n=8  n=8  40  H April 1997  1  'A Oct.  July 1997  1997  £2  < 5 fim  May 1998  July 1998  Oct. 1998  HSI < 5 um  Figure 2 2 Cruise means of <5 um and > 5 Mm chlorophyll ± 1 S.D. for the west coast of Vancouver Island in 1997 and 1998. Values are the mean of shelf and beyond shelf stations during each cruise. Numbers on the top of each panel represent the percentage of the total chlorophyll that was accounted for by the >5 um fraction.  77  A2) CROSS-SHELF GRADIENT Annual and 2 yr means-cross-shelf The size structure and the relative contribution of the >5 pm fraction was different in the shelf and beyond shelf regions. Phytoplankton biomass in the shelf region was dominated by the >5 pm fraction, whereas at the beyond shelf region, the <5 pm fraction accounted for the highest proportion of biomass (Figure 2.3). For the 2 year study period, large cells accounted for 62% (range between 23 and 82%) of total chlorophyll of the shelf regions while in the beyond shelf region they account for only 29% (range between 5 and 71%).  The observed  difference in the relative contribution of the >5 pm fraction between the two regions was significant (p<0.01).  140 H  Q-  120  o  100 H  " g E +J — CO .C C CJ O c,  o E  ro —  £z  0) N  55  80 60 40  20  o-^ [23 < 5 pm  m  >5um  Figure 2.3 Interannual mean and annual means of size-fractionated chlorophyll ± 1 S.D. for the shelf and beyond shelf region off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the mean of all shelfand betond shelf stations sampled. Numbers on the top of each panel represent the percentage of the total chlorophyll that was accounted for by the >5 pm fraction.  78  In the shelf region, the difference between the two size fractions was not significant whereas in the beyond shelf region, biomass was significantly higher in the <5 pm fraction (p<0.01). The biomass in the <5 pm fraction showed little cross shelf variation while the >5 pm fraction showed significantly higher biomass in the shelf region than in the beyond shelf region (p<0.01). Seasonal means-cross-shelf In 1998, in the shelf region, the contribution of the >5 pm fraction increased relative to 1997. In 1998, chlorophyll in >5 pm fraction increased 1.7-fold relative to 1997, whereas the chlorophyll in <5 pm fraction remained the same (Figure 2.4). The general pattern of higher biomass of the >5 pm fraction in shelf regions was evident during each cruise with the exception of the October 1997 cruise, when the small cells accounted for over 60% of the total chlorophyll (Figure 2.4). During all other cruises in 1997 and 1998, the >5 pm fraction accounted for the majority of the total biomass of shelf region whereas the <5 pm accounted for the majority of total biomass in beyond shelf regions. There was no consistent seasonal pattern for the relative contribution of the >5 pm fraction. For 1997, the relative contribution of the >5 pm fraction was similar in April and July, while for 1998 the relative contribution was highest in July . Seasonal changes in biomass and the lack of a consistent seasonal pattern in the relative contribution of the >5 pm fraction, suggest variable environmental conditions over the course of the season. The highest contribution of the >5 pmfractionduring the two study years was in July 1998.  79  Percentage accounted for by > 5 um fraction  £3  < 5 um  E7H > 5 um  Figure 2.4 Mean of size-fractionated chlorophyll ± 1 S.D. for the shelf and the beyond shelf region off the west coast of Vancouver Island in April (n=2), July (n=3) and October 1997 (n=4) and May (n=4), July (n=4) and October (n=4) 1998. Shelf and beyond shelf valuesare the mean of all shelf and beyond shelf stations. Numbers on the top of each pannel represents the precentage of the total chlorophyll that was accounted for by the >5 um fraction.  80  Transects-cross shelf The highest biomass for the >5 um fraction was measured in July 1997 at Barkley Canyon.  In 1998 the highest biomass was measured in July at Brooks Peninsula but high  biomass was also observed at La Perouse Bank and Estevan Point (Appendix K). In 1998, high biomass was measured in the shelf region of most transects suggesting all transects have the potential of favorable conditions for phytoplankton growth.  The minimum for both size  fractions was observed in 1997 whereas the maximum concentration for both fractions was observed in 1998. This fact coupled with a slightly higher contribution of large cells suggests that 1998 was a better growing year than 1997.  Variability Size-fractionated biomass on the WCVI showed high variability on average, with a range of 8.7-80.0 mg chl m' for the <5 pm fraction and 1.8-213.1 mg chl m" for the >5 pm 2  2  fraction. The relative contribution of the large size fraction also varied widely from a low of 5% to a high of 82% of the total chlorophyll.  A3) DEPTH PROFILES OF SIZE-FRACTIONATED CHLOROPHYLL The relative contribution of the >5 pmfractionvaried with depth in the water column in both the shelf and the beyond shelf regions (Figure 2.5 and 2.6). Only the values for July 1997 and 1998 are shown; the values for the remaining cruises are shown in Appendix L . The trends were clearer in 1998 than in 1997, but they showed that generally the contribution of the >5 pm fraction decreases with depth.  81  120 100 -  La Perouse Bank Shelf  57%  La Perouse Bank Beyond Shelf  Barkley Canyon Shelf  82%  Barkley Canyon Beyond Shelf  25%  80 60 -_ 40  -j  20 0 120 100 80 -  no data are available  60 -  Q.  P  4020  5 Q C  0)  120 100  o  80 -  Q.  60  49%  Estevan Point Shelf  Estevan Point Beyond Shelf  28%  Brooks Peninsula Beyond Shelf  24%  40 ~_ 200 120 100 80  Brooks Peninsula Shelf  no data are available  6040 20 - I 100 ' 55 ' 30 ' 10 ' 3 ' 1 '  100  55  30  10  3  1  Percent of Surface Irradiance  123  < 5 urn fraction  EZ3  > 5 Lim fraction  Figure 2.5 Relative contribution of < 5.0 um size fraction and > 5.0 um size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during July 1997. Relative contribution of > 5.0 um fraction to depth integrated chlorophyll is in right hand comer of each graph. Brooks Peninsula shelf and Barkley Canyon beyond shelf stations were not sampled. 82  120 100 -  La Perouse Bank Shelf  70%  La Perouse Bank Beyond Shelf  23%  m  80 60 40 20 0• 120 100  Barkley Canyon Shelf  62%  Barkley Canyon Beyond Shelf  71%  80 60  I  1  Ifv"  40 20 0 120  i Estevan Point Shelf  85%  Estevan Point Beyond Shelf  r 32%  m  100 80 60 40-  ill  20 0• 120 100  i  i—r Brooks Peninsula Shelf  95%  Brooks Peninsula Beyond Shelf  r 26%  (if*  80  ML  WW  60 40 20 0  100  55  30  10  3 ' 1 '  100  55  30  10  Percent of Surface Irradiance E3  < 5 um fraction n  > 5.0 um fraction  Figure 2.6 Relative contribution of <5 um size fraction and >5 um size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during July 1998. Relative contribution of >5 um fraction to depth integrated chlorophyll is in the right hand comer of each graph. 83  B) Total Primary Productivity Integrated primary productivity data for all stations during 1997 and 1998 averaged for each cruise, each year and for all 6 cruises (2 yr mean) are given in Appendix M . For this section, total and size-fractionated primary productivity will be discussed separately. Bl) WCVI-TOTAL PRODUCTIVITY The 2 yr mean primary productivity for the WCVI was 4.0 ±0.8 g C m" d" (range 2  1  between 0.8 and 5.7 g C m" d* ). In 1997, primary productivity was on average higher 2  1  (4.3 g C m" d") than in 1998 (3.4 g C m" d" ). 2  1  2  1  There was no consistent seasonal pattern in primary productivity during the two years except during both study years the lowest productivity was measured always in October (Figure 2.7).  In 1997, primary productivity was highest in April, while in 1998 primary  productivity was highest in July.  A/M 1997  A/M 1998  J 1998  1  0 ~ ^ 1998  Figure 2.7 Daily mean primary productivity ± 1 S.D. off the west coast of Vancouver Island for April, July and October 1997 and May, July and October 1998. Values are the mean of all stations sampled during each cruise.  84  B2) CROSS-SHELF GRADIENT  2 yr mean and annual means Primary productivity was generally higher in the shelf than the beyond shelf region (Figure 2.8). The 2 yr mean primary productivity was 5.1 ±2.7 and 2.0 ±0.1 g C m" d" in the 2  1  shelf and beyond shelf region. On average over the study period, primary productivity was 2.6-fold higher in shelf regions than beyond shelf regions.  During 1997, the difference  between shelf and beyond shelf regions was less clear than in 1998. Primary productivity was on average 1.8-fold higher in shelf regions in 1997, whereas in 1998 primary productivity was 4.0-fold higher.  25n=4  > u  20  8 xi 15 £ E  i «  n=4  ,  J . S 10-I  re  5  n=3  n=4  n=4  n=4 n=1  n=4  n=3  Apr 1997  July  May 1998  Oct. 1997  1997  [22  Shelf  HI  ii  n=4  July  1998  n=4  n  =  4  Oct 1998  Beyond Shelf  Figure 2.8 Mean primary productivity ± 1 S.D. (g C m" d" )ofthe shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the average of all shelf and beyond shelf stations sampled. 2  1  85  Seasonal means The general trend of higher primary productivity in the shelf region was not observed during October 1997 or July 1998 when the primary productivity was slightly higher in the beyond shelf region of the Brooks Peninsula transect (Figure 2.9). The maximum difference between the two regions occurred in July 1998 when productivity was 6.5-fold higher in the shelf.  No consistent seasonal pattern was observed in either the shelf or the beyond shelf  region.  For 1997, primary productivity in the shelf region was highest during April, while in  1998 the seasonal peak occurred in July. Variability Both regions showed high variability on average, with a range between 0.6 and 26.1 g C mf h" for the shelf region and 0.3 and 6.3 g C m" h" for the beyond shelf region. The 2  1  2  1  variability in primary productivity during both years was similar in the shelf and the beyond shelf regions. B3) ALONGSHORE DIRECTION There was no consistent pattern along the length of Vancouver Island in shelf or beyond shelf regions for 1997 or for 1998 (Figure 2.9).  The transect where biomass was the highest  varied with each cruise. No alongshore gradient in primary productivity was observed for 1997 and 1998.  86  28 26 H 24  La Perouse Ba nk  Barkley Canyon  Estevan Point  Brooks Peninsula  o 3  10CM  >  8 -  D_  43 O I-  >  6 4-  s  2 -  0  0  ' A/M '  A/M  EZ3  Shelf 1997  ESI  in Hk A/M  Beyond 1997  [31  II  Shelf 1998  A/M  •  Beyond 1998  Figure 2.9 Total mean primary productivity of shelf and beyond shelf regions of the La Perouse Bank, Barkley Canyon, Estevan Point and Brooks Peninsula transect off the west coast of Vancouver Island. One station wassampled for each of the shelf and the beyond shelf region of each transect. A/M=April and May, J=July and O O c t o b e r .  C) Size-fractionated primary productivity Integrated data ±1 S.D. from all stations sampled during 1997 and 1998 for sizefractionated primary productivity were averaged for each cruise and for all 6 cruises and are given in Appendix M . CI) WCVI-SIZE-FRACTIONATED  2 year means Productivity was consistently higher in the >5 um fraction than in the <5 pm fraction for 1997 and 1998 (Figure 2.10).  The 2 yr mean primary productivity was 1.1 ± 0 . 6 and  2.8 ±0.7 g C m" d" for the <5.0 pm and >5.0 pm fraction, respectively. On average, primary 2  1  productivity in the >5.0 pm fraction was 2.5-fold higher than productivity in the <5.0 pm fraction. On average, the >5 pm fraction accounted for 54% of the total primary productivity  87  of the W C V I .  Primary productivity o f both size fractions was on average higher in 1997 than  in 1998 (Figure 2.10). Size-fractionated  primary productivity on the W C V I  showed high variability on  average, with a range o f 0.3-1.8 g C m" d" for the <5.0 pm fraction and 0.4-5.4 g C m" d" for 2  1  2  the >5.0 p m fraction. Variability was higher in 1998 than 1997 for both size  1  fractions  (coefficient o f variation = 26 and 73% for <5 pm fraction for 1997 and 1998, respectively and 26 and 93% for the >5 p m fraction for 1997 and 1998, respectively). The relative contribution o f the >5 p m fraction varied widely from a l o w o f 44% to a high o f 62% o f the total primary productivity. Clearly the >5.0 p m fraction contributed substantially to primary productivity on the W C V I .  S  <5|jm ^  >5(jm  Figure 2.10 I n t e r a n n u a l m e a n a n d a n n u a l m e a n s o f size-fractionated p r i m a r y p r o d u c t i v i t y ± 1 S.D. for t h e w e s t coast o f V a n c o u v e r Island i n 1997 a n d 1998. N u m b e r o n t h e t o p o f e a c h p a n e l represent t h e p e r c e n t a g e o f the t o t a l p r i m a r y p r o d u c t i v i t y t h a t w a s a c c o u n t e d f o r b y t h e >5 p m f r a c t i o n .  88  Seasonal means  There was no consistent seasonal trend in primary productivity for either fraction. For 1997, productivity was highest in April and for 1998, it was the highest in July (Figure 2.11). The highest productivity of the two years was measured in the >5 pm fraction in July 1998. There was no consistent seasonal pattern of the relative contribution of the >5.0 pm size fraction during 1997 and 1998. The relative contribution of the > 5 pm fraction was similar in 1997 and 1998, but was more variable in 1998 than 1997 (coefficient of variation in 1997 was 3% and in 1998 was 18%).  55%  55%  53%  62%  61%  14 H > o 3  n  44%  n=8  12 H  •o  s  10 -\  Q.  8H  £•  I "° & O c  -2  n=8 6H  0)  n=5  n=8  4H  o  I  n=6  n=8  n=8  n=5  n=6  2H  n=8  N  V) April 1997  Oct. 1997  July 1997  IZ3  <5pm  May 1998  m  July 1998  n=8  n=8  Oct. 1998  <5um  Figure 2.11 Cruise means of <5 pm and > 5 pm primary productivity ± 1 S.D. for the west coast of Vancouver Island in 1997 and 1998. Values are the mean of shelf and beyond shelf stations during each cruise. Numbers on the top of each panel represent the percentage of the total primary productivity that was accounted for by the >5 pm fraction.  89  C2) CROSS-SHELF GRADIENT 2 year means  Generally, the size structure of primary productivity of the two regions was different. In the shelf regions, primary productivity was dominated by >5 pm phytoplankton while at the beyond shelf region <5 pm phytoplankton accounted for the highest proportion of productivity (Figure 2.12). The >5 pm phytoplankton accounted for on average 72% (range between 67 and 86%) of total primary productivity of the shelf regions and only 36% (range between 23 and 75%) for the beyond shelf regions. Shelf region primary productivity of the >5 pm fraction was significantly greater than the productivity of the <5 pm fraction (p<0.05). Productivity in the >5 pm fraction was significantly higher in the shelf than in the beyond shelf region (Figure 2.12) There was no consistent trend in productivity for the <5 pm fraction and on average productivity was similar in both regions. Variability  The variability in the shelf region was higher in the <5 pm fraction than the >5 pm fraction, in contrast to higher variability in the >5 um fraction than the <5 pm fraction in beyond shelf regions (see coefficient of variation in Appendix M) The variability of the >5 pm fraction was similar in both regions. For the <5 pm fraction variability was higher in the shelf than beyond the shelf. This implies that primary productivity in the small cells in the beyond shelf region is static while in the shelf region it undergoes changes and is dynamic. On average, variability was higher in 1998 than 1997 for bothfractionsand both regions.  90  12-T  72%  JL  38%  69%  36%  1998 mean  1997 mean  2-yr mean  34%  78%  >  n=12  O 3 TJ  10  s  Q.  |?  8H  £ E  6H  I-  c«  n=11 n=21  •o O CB  c o  4H  u  n=15  2  T  CD N  2A  n=11 n=21  n=19  tf)  04  *  Shelf  1  ~T  = n=19 n  Beyond  11  Shelf  1  n=15  mm  Beyond  n=12  n=12 n=12  Shelf  V2ZZML Beyond  E2 <5um E 3 >5 um Figure 2.12 Interannual mean and annual means o f size-fractionated primary productivity ± 1 S.D. for the shelf and beyond shelf region off the west coast o f Vancouver Island in 1997 and 1998. Shelf and beyond shelf means are the mean o f all shelfand betond shelf stations sampled. Numbers on the top o f each panel represent the percentage o f the total primary productivity that w a s accounted for by the >5 um fraction.  Seasonal Means The general pattern o f higher contribution o f the >5 u m fraction relative to the <5 um fraction was not always observed; in October 1997, the <5.0 u m fraction accounted for 59% o f the primary productivity at L a Perouse Bank. The relative contribution o f the >5.0 u m fraction in the shelf region was lowest in October in 1997 and 1998. This suggests that environmental conditions were not favorable for >5 um phytoplankton in the fall.  There was no consistent  seasonal trend in the beyond shelf region o f the relative contribution o f the >5 u m  fraction  during 1997 and 1998 (Figure 2.13). A trend may be obscured by the fact that in October 1997 only one beyond shelf station (Brooks Peninsula) was sampled and during May 1998 the  91  phytoplankton community at Brooks Peninsula beyond the shelf was more typical of a shelf region. The highest productivity of the two years was measured in the shelf region in the >5 pm fraction in July 1998. Variability In 1997, the variability was greater in the >5 pm fraction than the <5- pm fraction in contrast to higher variability in the <5 pm fraction in 1998. In 1997, the coefficient of variation was 79% for the >5 pm fraction and 66% for the <5 pm fraction. In 1998, the coefficient of variation was 138% for the >5 pm fraction and 201% for the <5 pm fraction. The pooled data clearly show that the contribution of the >5 pm size fraction was greater at high biomass concentrations and high productivity rates, supporting the idea that in order to reach high biomass and productivity, large cells are required (Figure 2.14). The lack of data points in the bottom right hand corner show that high rates of primary productivity were not measured if large phytoplankton were not abundant. The pooled data also clearly shows the distinct characteristics of shelf and the beyond shelf regions. Generally, beyond shelf stations are grouped at the bottom left hand quadrant and are characterized by low total chlorophyll composed mainly of small cells and by lower productivity, also accounted for by the smaller cells. During the two years of this study, beyond shelf stations were never found in the top right hand quadrant. C3) DEPTH PROFILES OF THE CONTRIBUTION OF >5 pm FRACTION In the shelf region, the relative contribution of the >5 pm fraction was highest at the top of the euphotic zone and then decreased as the depth increased (Figure 2.15 and 2.16). The relative contribution of the <5 pm fraction generally increased as depth. At the beyond shelf region, the <5 pm fraction was generally high at all depths with a slight increase at the bottom of the euphotic zone. 9 2  Percentage accounted for by > 5 um fraction  20 - |  73%  1997  JL  73%  54%  Shelf  43%  38%  48%  1997 Beyond Shelf  15  10 H  5H  0  wAh  J ^  f T i ^ r ^  vzmm;  20 - |  15 H  10 H  5H  May  Oct.  July  E2  <5um  May  July  Oct  E 3 >5um  Figure 2.13 Mean o f size-fractionated p r i m a r y productivity ± 1 S.D. for the shelf a n d the beyond shelf region off the w e s t coast o f Vancouver Island in April (n=2), J u l y (n=3) a n d October 1997 (n=4) a n d May (n=4), J u l y (n=4) a n d October (n=4) 1998. S h e l f a n d beyond shelf v a l u e s are the mean o f all shelf and beyond shelf stations Numbers on the top o f each pannel represents the precentage o f the total p r i m a r y productivity that w a s accounted for by the >5 pm fraction.  93  100 -f  o 0 • 0  2 o  sz u  60  o o  o %o  is  p c o  o  o o ,  •  H  40  o  o  80 H  a  A  0  o co m  20 H  m  m A  m m  m  o-l  50  100  150  200  250  300  Total Chlorophyll (mg chl m" ) 2  100 >  o o  3  •o o  80  H o  £•  E  60  B Q  40  H  C  o u CO  m A  20  0°  0  Q. CO  B  €  o  o  °o° o  0 0  °  o o  "  * • ••  o-l  12 14 10 6 2 Total Primary Productivity (g C m ' d' )  o  •  Shelf Beyond  Figure 2.14 Percent contribution of >5 um size fraction relative to: A) total chlorophyll and B) total primary productivity off the west coast of Vancouver Island. Values are for all stations in 1997 and 1998 in the shelf and the beyond shelf region.  94  > o 3  •a o  Q.  CO  E o c CD  H Q.  100 ' 55 ' 30 ' 10 ' 3 ' 1 ' 1 0 0 ' 55 ' 30 ' 10 ' 3 ' 1 Percent o f Surface I r r a d i a n c e  E3  < 5 um fraction  El  > 5 pm fraction  Figure 2.15 Relative c o n t r i b u t i o n o f < 5 u m size f r a c t i o n a n d > 5 u m size fraction to p r i m a r y p r o d u c t i v i t y at e a c h l i g h t d e p t h f o r t h e shelf a n d t h e b e y o n d shelf station o f e a c h t r a n s e c t during J u l y 1997. Relative c o n t r i b u t i o n o f > 5 p m fraction t o d e p t h i n t e g r a t e d p r i m a r y p r o d u c t i v i t y is inthe right h a n d c o r n e r o f e a c h g r a p h . Brooks Peninsula shelf a n d Barkley C a n y o n b e y o n d shelf stations w e r e n o t s a m p l e d .  95  100  55  30  10  3 ' 1  100  55  30  10  Percent of Surface Irradiance E3 < 5 Mm fraction H) > 5.0 Mm fraction Figure 2.16 Relative contribution of < 5 Mm size fraction and > 5 Mm size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during July 1998. Relative contribution of > 5 Mm fraction to depth integrated primary productivity is in the right hand corner of each graph.  96  D) CARBON ASSIMILATION RATES Dl) MEAN VALUES FOR WCVI The 2 yr mean carbon assimilation rates for the WCVI were 4.0 ±0.8 g C mg chl" d" 1  (range between 1.8 and 4.9).  1  Carbon assimilation rates were on average similar in 1997 (4.4 g  C mg chl" d") and 1998 (3.8 g C mg chl" d" ). There was no consistent seasonal pattern in 1  1  1  1  carbon assimilation rates during the two years except during both study years the highest rates were measured in July (Figure 2.17). Variability was higher in 1998 than in 1997 (coefficient of variation in 1998=46% and in 1997=6%).  (A CD  co i_  c o re  o £ O) w E CA CO  O  ° c € S CO  O  October 1998  Figure 2.17 Carbon assimilation rates ± 1 S.D. off the west coast of Vancouver Island in 1997 and 1998. Values are the mean of all stations sampled during each cruise.  97  D2) C R O S S - S H E L F G R A D I E N T  Carbon assimilation rates were on average higher in the shelf region than beyond the shelf (Figure 2.18). The 2 yr assimilation rates were 4.1 +0.8 and 3.8 ±1.1 g C mg chl" d" in 1  1  the shelf and beyond shelf regions, respectively. No consistent seasonal pattern was observed for the shelf or the beyond shelf regions.  Carbon assimilation rates in the shelf region were  higher in 1997 than in 1998, while in the beyond shelf region rates were similar in both years. The differences between values in the shelf region were not significantly different from each other. During 1997 and 1998, the variability was higher in the beyond shelf region than the shelf region.  April  1997  July  ~"  1997  Oct  "~  1997  E3  Shelf  May  1998  ESI  July  1998  Oct.  1998  Beyond Shelf  Figure 2.18 Carbon assimilation r a t e s ! 1 S.D. of the shelf and the beyond shelf regions off the west coast of Vancouver Island in 1997 and 1998. Shelf means are the average o f all shelf stations sampled during each cruise and beyond shelf values are the mean of all beyond shelf stations during each cruise.  98  D3) ALONGSHORE GRADIENT There was no trend along the length of Vancouver Island for the shelf region for the 2 yr mean. Carbon assimilation rates were on average highest at Barkley Canyon and lowest at Estevan Point, but the differences between transects were not significant. The location of the peak carbon assimilation rate during each of the three cruises in 1997 and 1998 did not show a consistent pattern (Figure 2.19) and varied during each cruise and each year. In the beyond shelf region, there was a trend for assimilation rates to increase progressively from La Perouse Bank to Brooks Peninsula during each cruise in 1997 and 1998.  14 H </>  12 H  CU  10 -\ =2 £  E °  •5) g> <n c co  8H  0  O CD  6H  co O  4H 2H  PZl  Shelf 1997 ES  Beyond Shelf 1997  •  Shelf 1998  E3  Beyond Shelf 1998  Figure 2.19 Carbon assimilation rates o f the shelf and the beyond shelf region o f La Perouse Bank, Barkley Canyon, Estevan Point a n d Brooks Peninsula transect off the west coast o f Vancouver Island in 1997 and 1998. Shelf and beyond m e a n s a r e the average o f all shelf and beyond shelf stations, respectively.  99  Table 2.1 Summary of characteristics of shelf and beyond shelf regions off the west coast of Vancouver Island. Values are for 1997 and 1998. Units for parameters below are: M L , meters; temperature, °C, nitrate, uM; chlorophyll, mg m" ; PP, mg C m" d" ; carbon assimilation rates, mgCmgchr h^CV/Zo) Beyond Shelf Shelf 2  2  1  1  11.8 11.85  Mixed Layer depth Mixed layer temperature Mixed layer salinity N0  31.1 6.1  3  23.2 12.75 31.5 2.4  range 97/98 Mean Median <5 um chlorophyll range 97/98 Mean >5 um chlorophyll range 97/98 Mean range % contribution 97/98 Mean range Total PP 97/98 Mean Median CV. range <5 um PP 97/98 Mean range >5 urn PP 97/98 Mean range % contribution 97/98 Mean  24.3-226.7 95.6 84.1 10.4-73.9  11.9-147.8  32.2 12.3-213.1 59.8  26.7 3.0-44.5 13.5  23-92 62  5-71 29  0.3-26.1 5.1 2.8 52  0.3-5.1 2.0  0.1-3.2 1.0 0.2-22.9 *  0.2-3.8 1.1 0.1-4.6  4.1  0.9  41-93 72  14-91 36  range 97/98 Mean  1.0-10.7 4.2  0.2-13.8 3.8  Total Chlorophyll  AN  * 0-9.7 when one station was excluded  49 36.7 8.7-48.8  0.9 4  DISCUSSION Distribution of biomass and primary productivity Total phytoplankton biomass and primary productivity off the west coast of Vancouver were high. Surface biomass reached 13.4 mg chl m" and mean integrated chlorophyll biomass 3  up to 227 mg chl m' was measured.  Total chlorophyll greater than 100 mg chl m" and  2  2  primary productivity >5 g C m' d" was frequently measured during all cruises. During most 2  1  cruises primary productivity > 3 g C m" d" was common. On average, primary productivity 2  1  was 4.0 g C m" d' off the west coast of Vancouver Island. 2  1  On average, biomass was higher in 1998 than in 1997, while on average primary productivity was lower in 1998 than in 1997, mainly due to differences in the beyond shelf region.  Whitney et al. (1998) have shown a shoaling and warming of the mixed layer depth in  the 1990's and consequently reduced NO3" concentrations in the surface layer. Strong vertical stratification of the water column would effectively block the vertical flux of nutrients to the surface.  Chapter one showed surface depletion of NO3" and a deeper mixed in 1998 layer  suggesting nutrient availability may have accounted for some of the difference in primary productivity between the two years. Alternately, the role of solar radiation can not be ruled out since we do not have solar radiation for the full growing season.  Solar radiation was only  collected during the duration of the cruise and consequently no corrections for mean monthly solar radiation was made. Total solar radiation has a strong seasonal signal at mid-latitudes. Phytoplankton at mid-latitudes show a strong relationship to solar radiation (Perry et al. 1989) and therefore measurement of primary productivity during periods of cloud cover (below average daily solar radiation) will cause a reduction in primary productivity.  101  The distribution of phytoplankton biomass and productivity off the west coast of Vancouver Island varied considerably in the cross-shelf gradient (E-W direction).  During the  two study years, phytoplankton biomass showed a 13-fold variability, ranging between 17.3 and 227 mg chl m' . The surface biomass in the shelf region, ranged from 1.48 -13.4 mg chl m' 2  3  while in the beyond shelf region, chlorophyll ranged between 0.17-6.35 mg chl m' . Similarly, 3  primary productivity varied considerably, ranging from 0.3-26.1 g C m" d" . 2  1  Biomass and  productivity were generally higher in the shelf region, with the exception of Brooks Peninsula. It is important to note that in regions of generally high productivity, low rates are also measured, highlighting the variability in the region. Primary productivity variability was greater in the shelf region than the beyond shelf region. In addition to cross-shelf patchiness, the distribution of phytoplankton biomass and productivity in the long shore directions (N/S) was patchy and varied considerably.  The  location along Vancouver Island (i.e. L a Perouse Bank, Barkley Canyon, Estevan Point and Brooks Peninsula) where biomass was the highest varied during each cruise.  At least once  during the two year study period, the highest biomass was measured at each of the four transects.  A weak trend of higher biomass at the northern end of Vancouver Island was  observed for both the shelf and the beyond shelf regions. The high variability in the cross-shelf and alongshore directions suggests a wide range of environmental conditions exist off the west coast of Vancouver Island. It is likely that the alongshore and the cross-shelf currents play an important role in maintaining the patchy distribution of biomass and productivity. The distribution of phytoplankton biomass and productivity showed considerable variation in time.  In April 1997, we had the opportunity to resample 4 stations within one  week. During the second week, the chlorophyll biomass was often higher and N C V was lower  102  than the previous week.  During the second week, nutrient depletion of the surface water was  also evident to greater depths. No opportunity was available to resample on a shorter time scale.  Seasonal variability was observed for biomass and productivity, but the seasonal  patterns were not similar in 1997 and 1998. The seasonal peak was seen in April 1997 and in July 1998 for both biomass and primary productivity. Over the two year study period, the highest biomass and primary productivity were measured in July 1998.  As was noted earlier,  the July cruise was during a period of upwelling favorable winds. The July cruise was the only cruise during the study period that occurred during upwelling favorable conditions.  The  intensity of upwelling was higher in 1998 than 1997 (R. Thomson, pers. comm.) which may explain high biomass and productivity in July 1998. community was composed of  In July 1998, the phytoplankton  Pseud-nitzschia delicatissima, Chaetoceros debilis, Skeletone  costatum, Asterionella glacialis, and Dactyliosolen fragillissimus, but was it dominated Chaetoceros spp. and Leptocylindrus danicus. Generally coastal environments are characterized by episodic pulses in biomass caused by chain forming diatoms and occasionally dinoflagellates  (Malone, 1980).  The phytoplankton assemblage  off the west coast of  Vancouver Island was consistent with this generalization. The high supply of nutrients during the spring/summer when solar radiation was high was responsible for the persistently high biomass and primary productivity that are characteristic of this upwelling region.  Lower  productivity measured during October of 1997 and 1998 was likely due to light limitation. Results from chapter one show light availability was lower during October relative to July and April. Nitrate concentrations were surprisingly low in October, with on average concentrations of 3.7 u M in 1997 and 4.7 uM in October 1998. Therefore, nutrient limitation can not account for the low October productivity rates. Variability was generally the highest during July (high  103  C V . relative to other cruises.) It is likely that the variance is due to the intermittent nature of upwelling during the summer. This thesis has shown the importance and substantial contribution of the >5 pm phytoplankton to biomass and primary productivity.  The relative contribution of the >5 pm  sized phytoplankton tended to increase as biomass and primary productivity increased. For example, during July 1998 when high biomass and productivity was measured, 73% of the phytoplankton community was  composed  of diatoms,  mainly Chaetoceros spp. and  Leptocylindrus danicus. We have also shown that in order to obtain high biomass and primary productivity large cells are required. This is consistent with modeling results obtained by Tremblay and Legendre (1994). The dynamics of phytoplankton biomass and primary productivity of shelf and beyond shelf regions were distinct in regard to community size structure and seasonality.  The shelf  regions tend to be dominated by >5 pm sized phytoplankton while the beyond shelf region was dominated by the <5 pm fraction, mainly various flagellates.  This is consistent with previous  observations that large phytoplankton cells tend to dominate in nutrient-rich coastal waters while small cells are characteristic of nutrient-poor waters (Malone, 1971).  Primary  productivity in the shelf region was higher than primary productivity in the beyond shelf region. The higher rates measured in the shelf region were due to >5 pm sized phytoplankton since the rates measured for the <5 pm fraction were similar in both regions.  Beyond shelf  regions are generally characterized by the dominance of the smaller fraction as shown in Figure 2.14 where beyond shelf stations are generally clustered in the bottom left hand corner of the plot.  Frequently, the beyond shelf region of Brooks Peninsula showed characteristics more  similar to the shelf region. This is not surprising considering the narrow width of the shelf at  104  Brooks Peninsula. There is a high probability that freshly upwelled water and biomass can quickly be transported beyond the 200 m contour.  In addition, wind-driven upwelling  filaments (jets) recurrently develop off Brooks Peninsula, and transport a substantial proportion of nutrients, phytoplankton and zooplankton biomass from the shelf to the deep ocean (Forbes etal. 1991).  105  Comparison with previous studies off the west coast of Vancouver Island Previous studies off the west coast of Vancouver Island by Denman et al. (1981) have focused on the southern margin of Vancouver Island.  Their region of study overlaps with my  measurements on the La Perouse Bank and Barkley Canyon. Denman et al. (1981) found 2 persistent areas of high biomass and high productivity located parallel to the coast, one near the outer edge of the shelf region and the other <20 km offshore.  During July 1998, this study  found two regions of high biomass that closely match the area described by Denman et al. (1981). On average (mean of maximum values for the three cruises), the maximum biomass and productivity found by Denman et al. (1981) were higher than those reported in this thesis. Their maximum biomass reached 38.5 mg chl m' , while the maximum measured during this 3  thesis was 20.4 mg m' . Lower measurements of biomass is reasonable considering Venrick et 3  al. (1987) has shown chlorophyll was higher and sea surface temperature was lower in the early 1980's, the period of Denman et al.'s (1981) study, compared to the early 1970's. Considering that above average sea surface temperatures were observed during the 1990's, leads one to hypothesis that chlorophyll levels would be lower in the 1990's compared to the 1980's. Denman et al.  (1981) also  found that on average  maximum productivity reached  82 mg C m' h" whereas in this study on average, the maximum was 40.4 mg C m' h" . The 3  1  3  1  carbon assimilation numbers that were reported by Denman et al. (1981) were similar. The primary productivity rates measured in this study are similar to those found by Whitney et al. (1999). They reported primary productivity values of 2.4 g C m" d" , which is 2  1  lower than values found in this thesis, but considering the more offshore nature of their station (P4) relative to the regions in this thesis, lower values for their station would be expected.  106  The high spatial and temporal heterogeneity is in close agreement with Boyd et al., (1999).  Boyd et al. (1999) reported marked seasonal and interannual variability of  phytoplankton biomass and primary productivity at an offshore P4 station (depth 1300 m). The present study found beyond shelf regions were dominated by <5 pm sized phytoplankton which is similar to the community structure found by Boyd et al. (1999). A general cross-shelf gradient in phytoplankton biomass has been reported by Mackas, (1992) and Perry et al. (1999).  Generally, the shelf region was characterized by  higher chlorophyll concentrations than beyond the shelf region. Mackas (1992) found that for zooplankton biomass the outer shelf had the least seasonal variability, which is similar to the low variability I found for chlorophyll and primary productivity in the beyond shelf region. The elevated biomass and primary productivity and the carbon assimilation rates for the beyond shelf region off Brooks Peninsula are consistent with results by Forbes et al. (1986). A comparison of size-fractionated biomass and primary productivity with previous studies in the area is not possible since this is the first study to examine size-fractionated biomass and primary productivity. However, a comparison with results of a one-dimensional simulation model of plankton and fish production was made. Robinson and Ware (1999) found annual diatom production was lower in 1997 than 1998 which agrees with observations from this study that the >5 pm biomass and productivity were lower in 1997 than in 1998 and the relative contribution to biomass by the large fraction was lower in 1997. Primary productivity for the southern margin was 3.5 mg C m' d" in 1997, lower than 4.0 mg C m" d" in 1998. 2  1  2  1  The results of this study agree well with model simulation results. Robinson and Ware (1998) found diatom production has steadily declined since the mid-1980s, largely due to a decrease in  107  upwelling intensity.  This observation  may help explain lower biomass and primary  productivity in this study compared to values found by Denman et al. (1981).  Comparison with other upwelling regions Phytoplankton dynamics found for coastal upwelling regions off Washington are similar to results obtained in this thesis off the west coast of Vancouver Island. Coastal Zone Color Scanner images off the Washington coast showed patchy distribution of biomass and features such as jets were common (Perry et al. 1989).  High phytoplankton biomass and primary  productivity in the coastal upwelling areas off Washington were reported by Perry et al. (1989). In the shelf region, biomass ranged from 1-11.0 mg chl m" , while the range for the beyond 3  shelf region was 0.3-8.5 mg chl m' . Perry et al. (1981) found a general cross-shelf gradient in 3  biomass and productivity, which agrees with the results of this study. Generally, the primary productivity off Washington was 4 g C m" d" in the shelf region, higher than 1.5 g C m' d" in 2  1  2  1  the beyond shelf region. These rates are close to those found in this study, which was 5.1 g C m" d" in the shelf region and 2.0 g C m' d" in the beyond shelf region. Generally, primary 2  1  2  1  productivity off the coast of Washington was highest in the spring, but Perry et al. (1989) also found equally high rates in the summer. This is consistent with the results of this study that found the highest primary productivity in the spring of 1997 and in the summer of 1998. Phytoplankton biomass and primary productivity was on average higher off the west coast of Vancouver Island than found for Monterey Bay, CA. (Wilkerson et al. 1999), but the results of this study agree closely with the relative contribution by the >5 um size fraction reported by Wilkerson et al. (1999).  Wilkerson et al. (1999) found that the larger sized  phytoplankton (>5 um fraction) contributed significantly to biomass and productivity in  108  Monterey Bay. On average, the >5 um fraction accounted for 50% of the total chlorophyll and 56% of the primary productivity (Wilkerson et al. 1999) compared to 49% of total chlorophyll and 57% of primary productivity found off the west coast of Vancouver Island.  SUMMARY Primary productivity and biomass were high off the west coast of Vancouver Island. The >5 pm phytoplankton contributed significantly to biomass and primary productivity off the west coast of Vancouver Island and the relative contribution of the >5 pm cells increased as the phytoplankton biomass or productivity increased.  This study  has  shown  that high  phytoplankton biomass and primary productivity develop due to the presence of large cells, which were predominately diatoms. Phytoplankton in the beyond shelf region tended to be dominated by the <5 pm size fraction, while in the shelf region the >5 pm size fraction dominated.  The phytoplankton  assemblage in the beyond shelf region was composed of various flagellates, while diatoms dominated by Chaetoceros spp. and Leptocylindrus danicus, were common in the shelf region. On average, the relative contribution of the >5 pm size to total chlorophyll was 62% in the shelf region compared to 29% in the beyond shelf region. Similarly, the relative contribution of the >5 pm size fraction to primary productivity was 72% in the shelf region, while only 36% in the beyond shelf region.  Clearly, the size structure of biomass and primary productivity  were different for the shelf and the beyond shelf regions.  109  G E N E R A L DISCUSSION This study was conducted during an extreme ENSO event. One of the strongest ever recorded E l Nino event occurred during 1997/98, the first year of this sampling program. Conversely, a strong L a Nina event was recorded during 1998/99, the second year of this sampling program. The 1997/98 El Nino event peaked in February 1998 and ended abruptly in May of 1998 but sea surface temperatures in the coastal regions remained slightly higher than average until December 1998 (Freeland, 1998).  The April 1997 cruise occurred before  temperature anomalies were observed off the west coat of Vancouver Island, while the July and October 1997 cruises occurred during the 1997/98 El Nino event. The May, July and October 1998 cruises occurred during the 1998-99 La Nina event. One would expect interannual variability in the ocean climate given the magnitude of this most recent climatic oscillation. This study has shown the mixed layer was shallower and less saline in 1997. This suggests that the level of upwelling was lower in 1997 than 1998. Examination of a 30 year record of upwelling intensity revealed that the lowest index of upwelling occurred during 1997 (I. Perry, pers. comm.).  Despite lower upwelling in 1997 relative to 1998, nitrate and silicic acid  concentrations were higher in 1997, the year of less intense upwelling. Nutrient concentration is a non-conservative tracer and is influenced by the biological community. Lower nutrient concentrations in 1998 are not surprising considering phytoplankton biomass, particularly diatom biomass was higher in 1998 which would allow for rapid depletion of the nutrient pools. In fact, nitrate concentrations near or at detection limits were frequently measured. Lower nutrient concentrations observed in this study agree with Whitney et al. (1998) who found that nitrate during 1998 was lower than the 1969-1981 average nitrate concentration.  110  The dominant time scale of variability off the west coast of the biology and physics such as ocean currents, is the annual cycle (Denman et al. 1989). On a shorter time scale, current flucuatations have time scales of the order of 10 days and winds events occur on the order of days. Turnover times for phytoplankton are typically days therefore during the course of each cruise the phytoplankton are exposed to flucuating physical conditions.  The arrival  time at stations during the cruises was random, therefore the chance to arrival at at station before or after a wind event is equal.  Denman et al. (1989) suggest that if data was collected  frequently enough, on the order of days to weeks, interannual and interdecadal anomalies can be clearly shown.  The 1997 annual mean averaged results for 2 cruises during the E l Nino  while the 1998 annual mean averages results for 3 cruises during the L a Nina.  Given the  random arrival time at the stations, the near equal time scale for biological and physical events and the equal distribution of cruises between E l Nino and L a Nina events the annual means presented in this study are valid. This study has also shown that the shelf and beyond shelf regions showed distinct physical, chemical and biological parameters.  Temperature, salinity, density, chlorophyll,  phytoplankton abundance, total phytoplankton biomass and diatom biomass all showed a strong cross-shelf gradient. Generally, the shelf region tended to be more productive than the beyond shelf region.  The cross-shelf gradient is influenced by the Vancouver Island Coastal  Current, which acts as a barrier to exchange between the shelf and the beyond shelf regions (Thomson et al. 1989). Thomson et al. (1989) also suggested that the current might act as a conduit for alongshore transport. This study did not find a consistent pattern of long shore distribution of nutrients, biomass or productivity. This does not exclude the possibility that a latitudinal gradient exists or that the VICC does not influence the distribution of chemical and  ill  biological parameters. Although this study was extensive both in time and space, the cruises were only for 2-3 weeks, three times per year and only provided a snapshot of conditions. Satellite images could expand the time series and fill in gaps between cruises. The VJ.CC is a physical feature that acts as a barrier to exchange across the shelf break, while spatially localized jet-like currents that transport shelf water offshore act as a conduit for exchange Denman et al. (1989). This study found evidence of this at the Brooks Peninsula transect, since biomass and productivity in the beyond shelf region were frequently similar to that found in the shelf region. This study has shown the substantial contribution that the >5 pm sized phytoplankton make to biomass and productivity off the west coast of Vancouver Island. importance was higher in the shelf region as was the abundance of diatoms.  The relative This study has  shown that large cells (>5 pm) are largely composed of diatoms. Frequently, Chaetoceros spp. and Leptocylindrus danicus dominated the assemblage when high chlorophyll concentrations and high primary productivity were measured. In the beyond shelf region, various flagellates dominated the assemblage, resulting in a lower contribution of >5 pm phytoplankton.  In  addition, high biomass and productivity were measured when the assemblage was dominated by large cells, not small cells. This implies that in order to obtain high biomass or productivity, large cells must be present. The dominance of large cells has important ramifications for the local food chain. Larger cells tend to support a shorter food chain leading to more efficient transfer of energy to upper trophic levels (Ryther,  1969).  The dominance of large cells  perhaps explains why the La Perouse Bank has historically supported a diverse and highly productive fishery (Ware and Thomson, 1991).  The size structure of the phytoplankton  community has important implications for the downward flux to sediments, since the  112  magnitude of the downward flux is dependent on the assemblage structure (Michaels and Silver, 1988). It is more likely that larger phytoplankton will contribute more significantly to the vertical flux than smaller sized phytoplankton.  This study has shown that there is  variability in the size structure of the phytoplankton community that must be considered in modeling efforts. This study described the physical, chemical, and biological characteristics of four transects off the west coast of Vancouver Island. This was the first study to investigate: 1) size-fractionated biomass and rates of size-fractionated primary production along transects off the west coast of Vancouver Island; b) phytoplankton species composition from the southern to the northern margin of Vancouver Island; and c) the relative contribution of small and large cells to biomass and primary productivity off the west coast of Vancouver Island.  113  FUTURE RESEARCH The  results of this study have suggested additional studies that warrant further  examination. Suggestions for future research include: a) Further studies on size-fractionated biomass and productivity in the beyond shelf region to determination the relative contribution of picoplankton, particularly during periods of surface nutrient depletion. The beyond shelf region had low biomass, low nutrients and was dominated by the <5 um size fraction and it is likely that small picoplankton cells which have favorable nutrient uptake rates, contributed to the biomass and productivity. The relative contribution of picoplankton in the shelf region is likely to be much less important. The shelf region is generally nutrient rich and therefore the competitive advantage of the small cells would be outweighed by diatoms rapid nutrient uptake rates. b) Compare rates of nitrogen uptake to rates of primary productivity to examine if uptake rates approach Redfield ratios. c) Examine primary productivity in an upwelling jet to determine physiological changes that occur as water is advected offshore.  Does the sequence of changes conform to the  "conveyor belt" proposed by Dugdale and Wilkerson (1985)? d) Determination of primary productivity on the shelf break. 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Date  Season  Transition Date  Northwesterly Winds  1997 Cruises 9707  07-14 April  Spring  Jan. 30  No  9707  21-28 April  Spring  Jan. 30  No  9713  14-28 July  Fall  May 16  No  9737  20 - 27 Oct.  Fall  June 12  No  9810  11-25 May  Spring/Summer  Feb. 25  Yes  9823  13 -27 July  Summer  May 7  Yes  9836  05-13 Oct.  Fall  Oct. 23  No  9911  04-12 May  Spring  Jan 30  No  9928  30 June-06 July  Summer  02 June  Yes  9935  23-30 Sept.  Fall  30 Aug.  No  1998 Cruises  1999 Cruises  125  APPENDIX B  1999 DISSOLVED NUTRIENT R A W DATA SET  Table B . l N0 ", HP0 " and Si(OH) in May 1999 off the west coast of Vancouver Island. Dashed line (-) indicates that data point is not available. (-) indicates information is not available; b-10 indicates water sample was collected at 10 m off the bottom. Si(OH) Depth HP<V' Niskin N0 " Cast Date Station (m) # # uM uM uM 3  4  4  4  3  B2 B2 B2 B2 B2 B4 B4 B4 B4 B4 B4 B4 B6 B6 B6 B6 B6 B6 B6 B8 B8 B8 B8 B10 B10 B10 B10 B10 B10 B10 B10 B12 B12 B12 B12 B12 B12 B12 B12 B12 B12 B12 B16 B16 B16 B16 B16 B16 B16 B16  04-May-99  04-May-99  05-May-99  05-May-99  06-May-99  06-May-99  06-May-99  4 4 4 4 4 6 6 6 6 6 6 6 8 8 8 8 8 8 8 14 14 14 14 46 46 46 46 46 46 46 46 44 44 44 44 44 44 44 44 44 44 44 43 43 43 43 43 43 43 43  5 4 3 2 1 7 6 5 4 3 2 1 7 6 5 4 3 2 1 4 3 2 1 9 8 7 6 5 4 3 2 11 10 9 8 7 6 5 4 3 2 1 8 7 6 5 4 3 2 1  0 10 20 30 50 0 10 20 30 50 75 100 0 10 20 30 50 75 100 50 75 100 130 0 10 20 30 50 75 100 125 50 75 100 125 150 175 200 250 300 400 495 40 50 75 100 125 150 175 200  18.5 19.1 22.0 23.7 14.2 1.7 12.5 12.6 10.2 19.5 13.2 32.9 0.0 1.9 11.1 8.0 16.2 12.4 25.1 15.6 16.9 22.0 33.9 0.4 0.5 4.7 4.8 5.3 7.6 23.1 33.6 4.8 10.0 24.1 28.7 33.2 35.2 17.2 36.9 35.8 36.8 28.2 8.1 4.6 12.0 30.4 32.1 27.0 34.4 35.8  1.7 1.8 2.0 2.1 1.3 0.6 1.2 1.3 0.8 1.3 0.9 2.6 0.4 0.0 1.1 0.5 1.5 1.0 2.2 1.5 1.4 1.8 3.0 0.4 0.4 0.8 0.6 0.5 0.5 2.0 2.6 0.3 1.2 2.1 2.4 2.4 2.7 1.4 2.8 2.6 2.7 2.0 1.1 0.3 1.3 2.4 2.5 2.1 2.6 2.7  29.9 29.3 33.0 34.6 29.4 19.3 25.3 25.8 19.6 27.8 22.1 50.2 12.9 7.9 20.4 14.1 29.1 22.1 44.8 23.7 24.3 60.7 82.1 12.7 11.4 14.9 13.9 12.7 13.5 30.9 47.5 9.8 15.3 29.7 37.0 41.3 47.0 29.8 54.1 58.1 55.2 37.6 15.5 10.9 29.9 82.0 80.1 78.0 83.2 47.7  APPENDIX B  Table B . l continued. Dissolved nutrients in May 1999. Depth Niskin N0 " Cast Date Station uM # m # 3  B16 B16 B16 B16 B16 B16 B16 B16 B16 C1 C1 C1 C1 C1 C1 C1 C1 C2 C2 C2 C2 C2 C2 C2 C4 C4 C4 C4 C4 C4 C4 C7 C7 C7 C7 C7 C7 C8 C8 C8 C8 C8 C8 C8 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C11 C11 C11 C11 C11 C11  05-May-99  05-May-99  05-May-99  05-May-99  05-May-99  05-May-99  05-May-99  05-May-99  42 42 42 42 42 42 42 42 42 9 9 9 9 9 9 9 9 21 21 21 21 21 21 21 23 23 23 23 23 23 23 27 27 27 27 27 27 28 28 28 28 28 28 28 30 30 30 30 30 30 30 30 30 30 30 30 30 36 36 36 36 36 36  _  8 7 6 5 4 3 2 1 7 6 5 4 3 2 1 7 6 5 4 3 2 1 6 5 4 3 2 1 8 7 5 4 3 2 1 13 12 11 10 9 8 7 6 5 4 3 2 1 13 12 11 10 9 8  500 600 700 800 1000 1200 1500 1669 1778 15 20 30 50 75 100 125 b-10 0 10 20 30 50 75 100 20 31 50 75 100 125 154 20 30 50 75 100 120 50 75 100 125 150 175 190 25 30 50 75 100 125 150 175 200 250 300 400 500 75 100 125 150 176 200  32.2 52.9 46.7 46.8 25.9 46.5 46.3 22.9 45.6 6.6 16.3 19.6 18.0 15.8 30.1 29.1 33.0 2.9 0.7 3.3 13.1 11.4 12.6 12.1 6.8 13.5 16.0 16.1 21.0 18.4 18.5 1.2 8.3 7.6 11.8 20.8 28.7 5.0 10.9 11.6 27.2 33.4 37.6 38.1 5.0 4.1 10.6 24.3 16.4 33.5 25.4 35.4 37.6 30.9 31.1 32.1 44.6 17.2 25.6 22.9 23.7 34.4 35.8  HPCV  Si(OH)  uM  uM  2.6 4.0 3.7 3.7 2.2 3.7 3.6 1.7 3.5  51.0 72.2 47.7 48.5 37.9 49.7 59.6 54.9 75.0 23.7 32.2 33.6 32.8 26.6 52.6 58.7 68.1 6.5 3.4 6.8 20.3 19.1 20.1 21.7 12.1 23.5 23.0 21.8 24.8 26.1 25.9 8.7 15.1 11.3 15.7 35.3 49.5 10.1 14.9 17.0 28.0 37.7 51.3 54.3 15.7 14.4 54.1 63.6 52.1 64.5 55.2 89.2 48.6 43.8 46.5 44.6 55.9 22.5 37.7 37.5 39.2 48.3 52.2  0.6 1.6 2.0 1.7 1.6 2.8 2.4 2.9 0.5 0.0 0.3 1.5 1.0 1.0 0.7 0.9 1.5 1.6 1.6 1.8 1.8 1.7 0.1 1.1 0.8 1.4 2.3 2.8 0.6 1.3 0.9 2.1 2.4 2.9 3.0 0.9 0.4 1.2 2.1 1.3 2.4 1.6 2.7 2.8 2.3 2.0 2.4 3.4 1.6 2.0 1.8 1.8 2.6 2.8  4  APPENDIX B  Table B . l continued. Dissolved nutrients in May 1999. Depth Niskin N0 Cast Station Date m uM # # 3  C11 C11  05-May-99  C11  C11 C11  C11 C11  C14 C14 C14 C14 C14 C14 C14 C14 C14 C14 C14 C14 C14 C14 C14 C14 C14 C14 D4 D4 D4 D4 D4 G1 G1 G1 G1 G1 G2 G2 G2 G2 G2 G2 G2 G3 G3 G3 G3 G3 G3 G3 G3 G4 G4 G4 G4 G4 G7 G7 G7 G7 G7 G7 G7 G7  06-May-99  06-May-99  07-May-99  07-May-99  07-May-99  07-May-99  06-May-99  36 36 36 36 36 36 36 40 40 40 40 40 40 40 40 40 40 40 40 40 40 40 40 40 40 50 50 50 50 50 52 52 52 52 52 52 52 52 52 52 52 52 65 65 65 65 65 65 65 65 56 56 56 56 56 59 59 59 59 59 59 59 59  7 6 5 4 3 2 1 7 6 4 3 2 5 4 3 2 1  7 6 5 4 3 2 1 8 7 6 5 4 3. 2 1 6 5 3 2 1 12 11 10 9 8 7 6 5  300 400 500 750 1000 1250 1445 0 10 20 30 50 75 100 120 150 175 200 250 300 400 500 750 1000 b-10 0 10 30 40 50 0 10 20 30 50 0 10 20 30 50 75 100 15 20 30 40 50 75 100 b-10 30 50 100 125 141 30 40 50 75 100 125 175 200  39.4 27.8 26.5 45.4 43.8 45.3 45.3 1.1 1.1 1.1 5.9 8.5 15.3 16.0 30.7 31.5 35.0 34.0 29.1 18.1 40.4 25.7 42.2 38.6 37.2 5.5 9.4 10.7 7.8 15.2 0.4 3.0 3.1 3.5 3.4 0.4 0.5 2.5 7.7 5.2 14.2 18.4 2.1 1.6 3.9 4.9 11.1 9.7 31.5 25.2 26.1 18.8 27.7 24.8 25.2 3.1 6.6 . 8.0 8.5 17.0 28.9 24.3 34.8  HPCV  Si(OH)  uM  uM  3.0 1.9 1.9 3.6 3.6 3.6 3.6 0.6 0.6 0.5 1.0 1.0 1.3 0.9 2.3 2.4 2.5 2.6 2.2 1.6 3.1 2.0 3.6 3.3 3.1 0.7 0.9 1.3 0.5 1.6 0.0 0.5 0.7 0.7 0.4 0.1 0.0 0.6 1.0 0.3 1.4 1.4 0.6 0.1 0.4 0.4 1.3 0.6 2.4 1.8 2.1 1.7 .2.0 1.3 1.5 0.7 1.0 1.1 0.9 1.5 2.2 1.9 2.7  57.0 46.4 37.8 48.0 89.4 81.7 98.7 12.1 11.4 10.5 13.5 13.6 19.3 21.6 37.8 40.7 45.2 48.2 51.7 37.1 48.9 36.0 51.9 51.8 58.8 11.0 16.0 16.3 13.7 22.9 4.3 9.2 9.0 9.2 10.3 7.4 5.2 8.0 15.4 12.0 20.1 31.8 14.3 13.0 10.7 10.3 14.3 38.2 80.3 72.1 34.5 26.2 35.0 39.3 40.8 12.6 14.2 16.8 47.3 30.5 37.8 43.2 49.0  4  APPENDIX B  Table B . l continued. Dissolved nutrients in May 1999. Station  Date  Cast #  G7 G7 G7 G7 G7 G7 G7 G7 G9 G9 G9 G9 G9 G9 G9 G9 G9 G9 G9 G9 G9 G9 G9 G9 J2 J2 J2 J2 J2 J2 J3 J3 J3 J3 J3 J3 J4 J4 J4 J4 J4 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J6 J6 J6 J6 J6  06-May-99  59 59 59 59 63 63 63 63 64 64 64 64 64 64 64 64 64 64 64 64 64 64 64 64 125 125 125 125 125 125  07-May-99  11-May-99  11-May-99  -  11-May-99  128 128 128 128 128 134 134 134 134 134 134 134 134 134 134 134 134 134 134 134 134 134 131 131 131 131 131  -  11-May-99  11-May-99  Niskin  # 4 3 2 1 12 11 9 7 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 6 5 4 3 2 1 6 5 4 3 2 1 5 4 3 2 1 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 17 16 15 14 13  Depth m 250 300 400 500 500 600 800 1000 0 10 20 30 50 75 100 125 150 175 200 300 400 500 750 1000 0 10 20 30 50 56 0 10 20 30 50 72 50 75 100 125 150 0 10 20 30 50 75 100 125 150 175 200 300 400 500 600 800 1000 20 30 50 75 100  N0  3  uM 25.8 40.4 27.2 45.1 35.9 50.4 32.1 51.8  o:8 0.5 0.2 1.1 7.7 14.2 19.7 22.2 19.1 32.7 13.2 28.5 45.1 26.0 46.7 38.6 0.7 0.9 2.4 5.7 10.8 13.8 0.6 0.6 2.9 5.4 8.2 8.8 13.4 22.1 22.4 36.9 34.8 0.5 0.3 1.4 2.6 6.3 10.7 14.1 30.5 33.6 34.9 37.6 30.7 23.8 29.5 45.3 41.6 43.6 0.5 0.6 1.2 1.3 3.4  HP<V" uM  Si(OH)  1.6 3.1 1.6 3.5 2.5 3.9 2.5 4.0 0.6 0.6 0.0 0.1 1.1 1.4 1.3 1.6 1.3 2.5 0.7 2.0 3.3 2.0 3.5 3.0 0.1 0.1 0.2 0.3 0.4 0.5 0.8 0.8 0.7 1.4 1.6 1.5 1.5 2.1 1.6 2.8 2.4 0.4 0.7 1.0 1.1 1.3 1.6 1.0 2.5 2.6 2.6 2.8 2.2 1.6 2.1 3.4 3.2 3.8 0.5 0.6 0.4 0.3 0.7  50.4 63.1 55.7 72.0 43.6 75.7 57.3 137.7 10.3 9.2 6.4 7.1 14.5 18.9 30.6 35.1 32.9 48.7 22.9 55.6 51.7 39.4 86.9 104.7  uM  -  -  2.3 2.3 2.3 4.2 9.3 8.6 22.6 34.8 36.5 49.2 41.6 2.3 2.3 2.3 2.5 4.1 7.6 25.3 44.6 47.1 50.2 55.0 50.0 42.1 44.1 58.7 66.0 144.5 2.3 2.3 2.3 2.3 2.3  4  APPENDIX B  Table B . l continued. Dissolved nutrients in May 1999. N0 Depth Niskin Cast Date Station m uM # # 3  J6 J6 J6 J6 J6 J6 J6 J6 J6 J6 J6 J6 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP8 BP8 BP8 BP8 BP8 BP8 BP8 BP8 BP8 BP8 BP8 BP8 BP8 BP8 BP8 BP8  11-May-99  10-May-99  10-May-99  10-May-99  131 131 131 131 131 131 131 131 131 131 131 131 117 117 117 117 117 117 117 117 117 117 117 117 117 117 117 117 117 123 123 123 123 123 123 123 123 123 123 123 123 123 123 123 123 110 110 110 110 110 110 110 110 110 110 110 110 110 110 110 110  12 11 10 9 8 7 6 5 4 3 2 1 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1  125 150 175 200 250 300 400 500 600 800 1000 1099 0 10 20 30 50 75 100 125 150 175 200 300 400 500 750 800 910 0 5 10 20 30 50 75 100 125 150 175 200 250 300 400 500  12.9 18.0 20.0 30.1 35.6 18.7 38.3 43.6 45.6 22.0 32.1 28.7 1.1 2.1 2.0 3.4 6.6 6.9 5.5 32.6 35.3 36.2 38.4 39.9 42.0 18.9 45.1. 45.6 24.9 2.8 2.7 3.7 3.9 6.8 7.0 10.6 12.3 25.1 27.2 28.1 29.9 42.5 35.8 39.3 42.7  21 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6  0 10 20 30 50 75 100 125 150 175 200 250 300 400 500 600  0.7 1.1 1.8 3.4 6.9 12.3 8.3 21.7 21.5 21.4 32.7 34.0 36.9 40.7 45.2 46.6  HPO/  Si(OH)  uM  uM  1.3 1.2 1.6 2.4 2.7 1.3 2.6 3.3 3.5 1.7 2.5 2.0 0.1 0.2 0.1 0.1 0.3 0.2 0.9 2.5 2.7 2.7 2.8 3.1 3.3 1.5 3.6 3.6 2.0 0.1 0.0 0.2 0.3 0.3 0.3 0.4 1.5 2.3 2.4 2.4 2.5 3.8 2.9 3.2 3.5 0.5 0.6 0.6 0.7 0.7 1.3 0.5 1.9 1.6 1.6 2.6 2.7 2.9 3.2 3.3 3.5  9.4 28.1 29.7 39.9 38.9 28.7 31.3 54.7 44.2 25.9 35.2 26.2  4  -  11.0 41.1 46.0 46.8 50.6 52.0 43.2 23.7 44.4 48.1 36.2  -  17.5 34.0 39.3 42.8 48.4 32.9 64.1 54.0 56.0 8.8 9.4 9.3 10.0 17.3 18.7 14.8 29.1 35.7 36.7 46.8 53.9 61.2 66.0 76.7 66.0  130  APPENDIX B  Table B . l continued. Dissolved nutrients in May 1999. N0 Depth Niskin Cast Date Station uM m # # 3  BP8 BP8 BP8 BP8 BP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 MP8 CPE2 CPE2 CPE2 CPE2 CPE2 CPE2 CPE2 CPE2 CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS1 CS1 CS1 CS1 CS1 CS1 CS1 CS1 CS1 CS1 CS3 CS3 CS3 CS3 CS3 CS3 CS3 CS3 CS3 CS3 CS3  10-May-99  11-May-99  09-May-99  09-May-99  08-May-99  08-May-99  110 110 110 110 110 137 137 137 137 137 137 137 137 137 137 137 137 137 137 137 137 137 107 107 107 107 107 107 107 107 95 95 95 95 95 95 95 95 95 95 95 67 67 67 67 67 67 67 67 67 67 73 73 73 73 73 73 73 73 73 73 73  5 4 3 2 1 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 8 7 6 5 4 3 2 1 11 10 9 8 7 6 5 4 3 1 1 10 9 8 7 6 5 4 3 2 1 11 10 9 8 7 6 5 4 3 2 1  800 1000 1200 1500 2000 0 10 20 30 50 75 100 125 150 175 200 300 400 500 600 800 1000 0 10 20 30 50 75 100 117 0 10 20 30 50 75 100 125 150 175 221 50 75 100 125 150 175 200 300 400 500 0 10 20 30 50 75 100 120 150 175 200  24.7 47.1 47.3 42.2 44.6 2.9 1.5 3.9 5.2 10.1 8.9 24.7 23.0 21.5 27.7 33.1 26.8 20.2 47.5 38.3 23.9 26.9 0.1 0.3 2.3 2.9 4.4 8.1 23.7 44.7 5.5 2.6 6.7 7.0 5.3 8.7 11.1 10.5 18.6 19.5 35.5 8.2 6.8 18.2 25.1 30.4 17.9 32.4 36.4 29.9 29.8 2.6 2.8 4.0 4.0 8.5 9.6 8.3 32.4 30.6 33.9 39.5  HP0 " uM z  4  2.0 3.6 3.5 3.2 3.4 1.1 0.5 1.2 0.8 1.6 1.1 2.2 1.8 1.8 2.0 2.3 2.0 1.3 3.5 3.4 2.3 2.4 0.4 0.3 1.0 0.4 1.2 0.8 2.0 3.4 0.6 0.0 0.8 0.7 0.3 1.0 1.1 1.1 1.6 1.3 2.7 1.1 0.6 1.8 2.2 2.6 1.2 2.6 3.0 2.5 2.0 0.9 1.0 1.0 0.8 1.4 1.5 0.6 2.5 2.7 2.9 3.0  Si(OH)  uM 37.7 71.9 64.1 51.6 60.1 2.3 2.3 3.0 2.8 2.3 2.3 71.8 65.6 60.0 58.7 46.2 44.2 25.3 48.7 46.3 29.2 34.1 4.7 7.5 8.6 5.9 12.3 16.7 31.1 58.3 12.0 7.6 12.5 12.9 13.1 16.0 17.3 16.5 25.4 33.7 55.8 14.9 13.0 60.5 71.3 80.8 69.4 84.2 93.2 83.4 70.1 2.3 2.3 2.3 2.3 5.7 8.5 58.0 88.0 85.8 89.6 86.6  4  APPENDIX B  Table B . l continued. Dissolved nutrients in May 1999 N0 Depth Niskin Cast Station Date uM (m) # # 3  CS5 CS5 CS5 CS7 CS7 CS7 CS7 CS7  08-May-99  CS9 CS9 CS9 CS9 CS9 CS9 CS9 CS9 CS9 CS9 CS9  08-May-99  08-May-99  77 77 77 82 82 82 82 82  3 2 1 6 5 4 3 1  88 88 88 88 88 88 88 88 88 88 88  11 9 10 8 7 6 5 4 3 2 1  30 50 57 0 10 20 30 56 0 10 20 30 50 75 100 125 150 175 179  5.6 10.4 8.0 5.6 6.0 4.6 5.6 10.8 3.6 5.3 3.4 5.8 8.8 6.0 10.2 24.7 32.1 37.1 37.7  HPCV  Si(OH)  uM  uM  0.4 1.0 0.5 0.8 0.9 0.4 0.3 1.1 0.9 1.1 0.8 1.1 1.4 0.8 0.9 2.1 2.6 2.9 2.9  12.8 17.4 16.1 13.0 13.5 10.3 12.8 20.5 2.3 2.7 2.3 3.6 2.3 2.3 60.8 35.8 47.9 50.3 53.7  4  APPENDIX B  1999 DISSOLVED NUTRIENTS R A W DATA SET  Table B.2 N 0 \ HP0 " and Si(OH) in July 1999 off the west coast of Vancouver Island. A l l samples were collected and analyzed as outlined in methods section of Chapter 1. (-) indicates information is not available. 3  Station B6 B6 B6 B6 B6 B6 B8 B8 B8 B8 B8 B8 B8 B8 B16 B16 B16 B16 B16 B16 C1 C1 C1 C1 C1 C1 C1 C2 C2 C2 C2 C2 C2 C2 C4 C4 C4 C4 C4 C4 C4 C4 C4 C4  Date 30-Jun-99  30-Jun-99  1-Jul-99  1-Jul-99  1-Jul-99  1-Jul-99  CA CA CA CA C5 C5 C5 C5 C5 C5  2-Jul-99  4  Cast # 6 6 6 6 6 6 8 8 8 8 8 8 8 8 18 18 18 18 18 18 33 33 33 33 33 33 33 34 34 34 34 34 34 34 36 36 36 36 36 36 36 36 36 36 36 36 36 36 38 38 38 38 38 38  4  Niskin #  Depth m  9 8 7 6 5 4 9 8 7 5 4 3 2 1 6 5 4 3 2 1 7 6 5 4 3 2 1 7 6 5 4 3 2 1 14 13 12 11 10 9 8 7 6 5 4 3 2 1  0 5 10 20 30 50 0 10 20 50 75 100 125 144 20 25 30 45 55 100 0 10 20 30 50 75 90 0 10 20 30 50 75 95 0 5 6 5 8 18 34 41 50 75 100 125 150 164  6 5 4 3 2 1  0 10 20 30 50 60  HPO/  Si(OH)  uM  uM  uM  4.6 7.1 15.7 32.5 22.5 29.5 2.1 19.9 15.9 28.4 22.0 24.7 34.5 20.7 0.7 2.1 6.0 10.3 9.5 20.0 1.7 8.5 33.8 36.3 38.5 41.5 42.1 3.3 14.1 25.0 29.9 33.0 27.6 35.4 12.3 4.9 12.4 13.3 11.1 32.0 22.5 12.2 26.5 26.6 20.4 27.7 21.9 28.8 3.4 7.6 35.3 36.6 40.2 40.1  0.2 0.4 1.1 1.8 1.3 1.7 0.5 1.2 1.4 1.6 1.7 2.2 3.0 1.8 0.0 0.3 0.7 0.9 0.6 1.3 0.0 0.3 2.0 2.2 2.3 2.5 2.4 0.6 1.2 2.0 2.3 2.4 2.1 2.6 1.1 0.3 1.1 1.2 0.9 2.3 2.0 1.2 2.4 2.4 1.9 2.5 2.1 2.7 0.2 0.8 2.2 2.3 2.7 2.7  8.1 9.7 16.4 53.5 44.6 49.7 9.5 17.9 18.3 11.1 21.8 46.5 63.5 45.6 26.1 28.5 30.8 30.5 25.2 31.3 6.9 13.0 37.8 42.2 44.5 46.4 70.4 15.2  N0  3  -  29.6 34.7 42.5 40.7 46.0 18.7 9.7 15.8 17.3 14.4 35.6 37.0 24.4 37.2 41.6 39.5 40.1 42.1 35.1 9.3 14.8 32.7 35.0 42.0 25.5  4  APPENDIX B Table B.2 continued. Dissolved nutrients in July 1999. Station C6 C6 C6 C6 C6 C6 C7 C7 C7 C7 C7 C7 C7 C7 C8 C8 C8 C8 C8 C8 C9 C9 C9 C9 C9 C9 C11 C11 C11  Date 2-Jul-99  Niskin  #  #  39  6 5 4 3 2 1 8 7 6 5 4 3 2 1 11 10 9 8 7 6 16 5 4 3 2 1 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 5 4 3 2 1 7 6 5 4 3 8 7 6 5 4 3 2 1  39 39 39 39 39  2-Jul-99  2-Jul-99  2-Jul-99  2-Jul-99  C11  C11 C11 C11 C11 C11 C11 C11 C11 C11 C11 C11 C11 C11 C11 C11 C11 G1 G1 G1 G1 G1 G2 G2 G2 G2 G2 G3 G3 G3 G3 G3 G3 G3 G3  Cast  3-Jul-99  3-Jul-99  3-Jul-99  40 40 40 40 40 40 40 40 41 41 41 41 41 41 42 42 42 42 42 42 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 46 61 61 61 61 61 63 63 63 63 63 65 65 65 65 65 65 65 65  Depth m 10 20  30 50 75 86 0 10 20 30 50 75 100 122 0 10 20 30 50 75 0 250 300 400 400 612 0 10 20 30 50 75 100 125 150 175 200 250 300 400 500 600 800 1000 1200 1446 0 10 20 30 50 0 10 20 30 50 0 5 10 20 30 50 75 100  N0 '  HPCV"  Si(OH)  uM  uM  uM  0.0 6.5 20.8 29.3 41.3 38.1 0.0 0.2 4.7 11.0 11.5 15.5 21.7 30.1 1.0 0.1 4.3 10.2 35.1 45.2 0.9 34.0 36.3 39.2 39.9 40.9 1.6 1.6 2.3 7.4 11.2 12.5 20.1 29.3 31.3 25.4 34.7 19.1 40.3 44.3 46.1 49.3 50.7 51.0 50.9 42.8 0.6 1.9 22.6 29.1 22.3 0.6 3.6 22.5 24.8 14.3 0.2 0.2 0.1 9.9 19.7 36.9 26.4 52.2  0.3 0.7 1.5 1.9 2.4 2.4 0.4 0.1 • 0.8 0.9 0.7 0.7 1.7 2.5 0.0 0.5 0.3 0.7 1.8 2.6 0.0 2.9 3.0 3.2 3.4 3.4 0.4 0.4 0.7 0.3 1.0 1.0 1.5 2.1 2.3 1.9 2.4 1.2 2.9 3.1 3.3 3.4 3.5 3.6 3.6 3.1 0.0 0.0 1.7 2.0 1.3 0.3 0.3 1.8 1.9 1.0 0.0 0.3 0.4 1.2 1.6 2.2 1.6 3.0  3  3.6 8.1 20.0 26.3 38.1 51.4 5.5 4.7 12.4 13.9 12.2 13.1 32.4 42.9 28.8 30.1 30.1 31.3 50.1 62.7 17.1 61.4 66.8 75.7 87.7 96.6 5.3 5.0 7.5 5.5 10.7 10.3 23.4 33.8 39.9 40.4 46.9 34.2 60.2 74.3 84.9 96.1 121.3 134.1 130.0 121.4 6.7 7.1 29.2 37.3 25.8 10.1 10.3 33.3 34.3 17.4 7.0 9.0 8.4 16.9 24.0 41.8 32.0 65.9  4  APPENDIX B  Table B.2 continued. Dissolved nutrients in July 1999. Station G6 G6 G6 G6 G6 G6 G6 G6 G6 G6 G6 G6 G6 G6 G6 G6 G6 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 BP2 BP2 BP2 BP2 BP2 BP2 BP2 BP2 BP2 BP3 BP3 BP3 BP3 BP3 BP3 BP3 BP3 BP3 BP3 BP4 BP4 BP4 BP4 BP4  Date  Cast  #  #  Depth m  3-Jul-99  70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 71 71 71 71 71 71 71 71 71 71 71 71 71 71 71 71 71 71 71 71 71 71 94 94 94 94 94 94 94 94 94 91 91 91 91 91 91 91 91 91 91 90 90 90 90 90  17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 22 21 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 9 8 7 6 5 4 3 2 1 10 9 8 7 6 5 4 3 2 1 19 18 17 16 15  0 10 20 30 50 75 100 125 150 175 200 250 300 400 500 600 800 0 5 10 20 30 50 75 100 125 150 175 200 250 300 400 500 600 800 1000 1200 1500 1769 4 6 10 15 18 30 50 75 96 0 10 20 30 50 75 100 125 150 161 0 10 20 30 50  3-Jul-99  f  6-Jul-99  6-Jul-99  6-Jul-99  Niskin  N0 uM 3  1.1 0.4 3.7 15.1 12.7 29.9 18.8 29.0 24.3 32.8 33.5 26.7 37.1 38.9 33.4 28.4 42.9 0.2 0.2 0.3 0.3 1.9 8.9 16.7 22.0 32.9 34.2 30.9 16.1 39.1 40.8 19.9 45.6 22.6 47.6 47.8 47.8 28.4 45.9 1.0 0.4 0.4 1.7 4.9 20.5 21.9 17.5 31.5 0.8 0.8 4.4 14.7 26.4 35.9 26.8 26.4 18.7 32.5 3.5 0.6 10.4 20.0 15.7  HPCV  Si(OH)  uM  UM  0.4 0.4 0.8 1.4 1.2 1.9 1.4 2.3 2.2 2.8 2.8 2.4 3.2 3.3 2.9 2.6 3.6 0.0 0.2 0.3 0.4 0.1 0.9 1.5 1.7 2.4 2.5 2.3 1.0 2.8 2.9 1.5 3.3 2.2 3.5 3.5 3.5 2.2 3.4 0.1 0.1 0.4 0.4 0.4 1.7 2.0 1.4 2.5 0.3 0.2 0.0 1.1 1.9 2.0 2.4 2.3 1.5 2.8 0.1 0.3 1.2 1.7 1.3  3.4 3.7 6.7 11.4 12.8 15.6 13.0 38.1 41.5 52.5 54.2 53.1 67.7 75.2 73.2 90.1 109.3 3.9 4.8 5.4 7.9 5.7 13.3 21.8 32.3 38.9 43.1 42.6 28.6 56.0 62.5 48.2 85.5 62.9 137.7 136.9 154.7 111.5 185.9 4.5 4.1 5.1 5.3 7.4 15.5 29.4 27.6 45.5 3.4 5.9 7.3 16.6 26.4 33.3 42.7 45.2 36.8 47.4 4.4 5.1 14.7 21.8 12.4  4  APPENDIX B  Table B.2 continued. Dissolved nutrient in July 1999. Station BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP4 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP6 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7  Date 6-Jul-99  5-Jul-99  5-Jul-99  Cast  Niskin  #  #  Depth m  90 90 90 90 90 90 90 90 90 90 90 90 90 90 89 89 89 89 89 89 89 89 89 89 89 89 89 89 89 89 89 86 86 86 86 86 86 86 86 86 86 86 86 86 87 87 87 87 87 87  14 13 12 11 10 9 8 7 6 5 4 3 2 1 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 14 13 12 11 9 8 7 6 5 4 3 2 1 6 5 4 3 2 1  75 100 125 150 175 200 250 300 400 500 600 800 1000 1080 0 10 20 30 50 75 100 125 150 175 200 250 300 400 500 600 900 125 150 175 200 300 500 700 750 800 1000 1200 1500 2250 10 11 15 22 31 37  N0 uM  HP0 uM  30.3 30.7 16.8 17.5 34.1 33.4 21.3 29.2 39.9 29.3 29.8 33.1 29.7 30.9 0.4 0.4 0.4 6.6 9.5 25.8 17.2 33.4 25.8 28.7 26.2 40.2 40.9 45.0 45.7 25.5 20.6 23.3 45.0 44.7 45.5 45.4 22.0 35.2 45.6 36.5 19.3 29.4 45.8 43.5 0.4 0.7 0.9 1.7 9.4 8.1  1.8 2.6 1.6 1.4 2.8 2.8 2.0 2.3 3.3 2.4 2.7 2.7 2.7 2.4 0.6 0.6 0.2 0.8 0.9 1.6 1.0 2.3 1.7 1.9 1.7 2.9 2.9 3.2 3.1 1.7 1.5 1.7 2.6 2.7 2.9 3.1 1.7 3.1 3.6 3.1 1.7 2.2 3.5 3.4 0.2 0.5 0.5 0.6 1.3 0.9  3  4  z  Si(OH)  uM 29.9 47.5 45.4 29.1 53.6 46.4 42.6 52.2 76.4 79.4 84.3 91.5 95.7 82.1 8.1 7.4 4.2 9.4 11.2 19.7 28.6 38.3 44.9 37.0 39.6 54.8 44.3 60.5 83.1 50.0 33.2 9.8 24.7 20.7 32.1 33.6 66.3 100.9 112.9 102.5 113.4 97.5 113.4 113.4 9.2 11.5 11.6 9.5 16.7 17.7  4  APPENDIX B  1999 DISSOLVED NUTRIENT R A W DATA SET  Table B.3 N0 ", HP0 ~ and Si(OH) for October 1999 on the west coast of Vancouver Island. A l l samples were collected and analyzed as outlined in methods section of Chapter 1. Dashed line (-) indicates that informaton is not available. HP0 Niskin Depth Si(OH) Cast Date NOj" Station uM uM uM # m # 2  3  4  4  z  4  B4 B4 B4 B4 B4 B4 B4 B6 B6 B6 B6 B6 B6 B6 B7 B7 B7 B7 B7 B7 B7 B7 B7 B7  30-Sep-99  C1  30-Sep-99  1-Oct-99  27-Sep-99  C1 C1 C1 C1  C1 C1 C1 C1 C4 C4  CA CA CA CA CA CA CA CA CA CA CI C7 C7 C7 C7 C7 C7 C7 C7  29-Sep-99  132 132 132 132 132 132 132 130 130 130 130 130 130 130 78 78 78 78 78 78 78 78 78 78 119 119 119 119 119 119 119 119 119 115 115 115 115 115 115 115 115 115 115 115 115 110 110 110 110 110 110 110 110 110  7 6 5 4 3 2 1 8 7 6 5 4 3 2 10 9 8 7 6 5 4 3 2 1 14 11 10 6 5 4 3 2 1 12 11 10 9 8 7 6 5 4 3 2 1 9 8 7 6 5 4 3 2 1  0 10 20 30 50 75 105 0 10 20 30 50 75 100 250 300 400 500 600 800 1000 1200 1500 2000 0 10 15 40 50 75 100 125 150 0 2 5 10 20 30 50 75 100 125 150 161 0 10 20 30 50 75 100 125 bot  20.3 11.3 32.5 27.2 33.4 39.3 37.3 9.2 11.4 8.4 19.1 19.8 34.9 30.4 11.9 29.6 32.2 12.5 34.8 39.7 41.5 40.4 47.3 47.4 3.0 25.1 26.0 26.9 28.7 23.5 27.3 25.5 30.6 17.5 16.3 17.7 19.4 13.9 24.2 24.6 22.9 30.2 19.9 32.7 28.0 9.0 21.4 100.7 31.6 28.4 60.7 27.2 22.8 38.4  2.0 1.7 2.9 3.0 3.1 3.3 2.9 1.0 1.1 1.4 1.7 1.7 2.9 2.6 0.7 2.3 2.4 1.0 2.5 2.9 3.0 2.9 3.5 3.7 0.4 1.8 1.8 2.1 1.7 1.8 1.6 1.8 1.5 1.9 1.8 1.8 2.1 1.7 2.2 2.1 2.3 2.5 1.7 2.8 2.2 0.6 0.6 2.3 2.5 2.1 4.4 2.4 1.8 3.1  35.4 29.3 40.8 40.3 41.3 46.4 47.5 25.4 24.8 28.0 31.4 31.6 47.7 45.1 15.6 32.7 37.4 19.3 38.3 36.3 40.4 27.9 48.6 49.0 8.7 37.7 36.1 33.7 37.2 31.4 39.4 38.6 48.4 31.4 32.3 29.8 34.8 28.2 34.8 37.1 38.7 49.8 40.0 55.4 44.6 17.3 20.2 61.3 55.0  78.8 46.4 37.9 53.2  4  APPENDIX B Table B.3 continued. Dissolved nutrients in October 1999. Station  Date  C8 C8 C8 C8 C8 C8 C8 C8 C8 C8 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9 C9  29-Sep-99  C11  29-Sep-99  29-Sep-99  C11 C11 C11  C11 C11  C11 C11 C11 C11  C11 C11 C11 C11 C11 C11 C11 C11 C11  D1  23-Sep-99  D1 D1 D1  D2 D2 D2 D2 D2 D4 D4 D4 D4 D4  23-Sep-99  23-Sep-99  Cast  Niskin  #  #  108 108 108 108 108 108 108 108 108 108 106 106 106 106 106 106 106 106 106 106 106 106 106 106 106 106 106 106 106 106 106 104 104 104 104 104 104 104 104 104 104 104 104 104 104 104 104 104 104 104 21 21 21 21 22 22 22 22 22 25 25 25 25 25  11  10 9 8 7 6 5 3 2 1 22 21 20 19 18 17 16 15 14 13 12 11 10 8 7 6 5 4 3 2 1 20 19 18 17 16 15 13 12 11 10 9 8 7 6 5 4 3 2 1 4 3 2 1 5 4 3 2 1 6 5 4 3 2  Depth m  NCV  uM  HP(V uM  0 10 20 30 50 75 100 150 175 188 0 2 5 10 15 20 25 30 40 50 75 100 125 175 200 250 300 400 500 600 613 0 10 20 30 50 75 125 150 175 200 250 300 400 500 600 800 1000 1200 1540 0 10 20 30 0 10 20 30 40 0 10 20 30 50  25.5 13.3 28.6 10.4 18.7 26.6 43.5 51.4 52.5 43.7 12.4 18.7 18.4 12.3 8.3 7.2 7.0 6.5 6.6 11.9 24.9 28.8 44.3 46.2 41.6 25.9 37.7 41.4 35.2 45.0 45.4 7.7 8.3 24.3 9.4 13.7 12.3 34.6 28.3 38.1 24.2 48.8 25.9 27.9 30.7 43.3 28.7 37.8 49.0 45.2 6.6 3.5 21.5 26.6 4.8 8.1 17.7 23.8 26.3 23.2 32.8 39.5 40.1 43.0  1.9 0.7 2.0 1.1 1.6 1.6 2.6 3.1 3.2 2.9 1.2 1.7 1.6 1.2 1.2 4.7 0.9 0.9 0.7 1.4 1.8 1.7 2.7 3.1 2.8 2.1 3.0 3.3 2.9 3.6 . 3.6 0.9 0.9 0.5 0.9 1.2 1.1 2.2 1.8 2.4 1.6 2.9 2.1 2.0 2.1 3.5 2.4 3.3 4.1 3.6 1.0 0.9 2.1 2.5 0.5 0.8 1.7 2.2 2.7 1.7 2.3 2.4 2.6 2.7  Si(OH)  uM 39.9 26.6 40.7 19.4 20.7 26.8 39.1 61.8 63.1 52.7 23.2 22.1 26.1 22.2 19.7 16.3 13.7 11.3 11.0 13.3 22.1 26.7 40.8 52.6 51.0 48.4 62.7 72.9 70.1 91.0 92.0 57.1 22.0 21.0 14.1 12.1 11.2 37.2 32.7 43.1 33.0 66.1 41.1 49.7 50.9 80.2 110.3 113.2 121.3 113.2 29.2 25.7 36.4 41.2 26.9 31.9 35.7 37.6 47.7  41.4  105.7  4  Table B.3 continued. Dissolved nutrients in October 1999. Depth Cast Niskin Station Date N0 " # m uM # 3  D8 D8 D8 D8 D8 D8 D8 D8 D8 D8 D8 D8 D8 D8 D8 D8 D8 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 D10 G1 G1 G1 G1 G1 G1 G3 G3 G3 G3 G3 G3 G3 G3 G3 G3 G3 G3 G3 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7  23-Sep-99  24-Sep-99  28-Sep-99  28-Sep-99  28-Sep-99  30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 30 32 32 32 32 32 32 32 32 32 32 32 32 32 32 32 32 32 32 32 32 88 88 88 88 88 88 92 92 92 92 92 92 92 92 92 92 92 92 92 98 98 98 98 98 98 98 98 98 98 98 98  17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 6 5 4 3 2 1 13 12 11 10 9 8 7 6 5 4 3 2 1 24 23 22 21 20 19 18 17 16 15 14 13  0 10 20 30 50 75 100 ,125 150 175 200 250 300 400 500 600 782 0 10 20 30 50 75 100 125 150 175 200 250 300 400 500 600 800 1000 1200 1475 0 10 20 30 50 55 0 2 5 10 15 20 25 30 40 50 75 100 125 0 5 10 15 20 25 30 40 50 75 100 150  4.0 6.2 8.3 19.4 31.3 98.8 17.6 33.1 35.2 38.5 28.5 27.9 25.5 28.8 103.7 31.9 49.4 1.1 1.3 2.7 6.5 6.4 6.4 18.1 20.7 19.7 43.5 25.0 33.0 20.5 36.3 22.6 26.9 29.0 46.0 45.9 32.7 15.7 16.1 14.8 25.3 25.1 26.3 12.8 13.7 14.1 8.7 13.1 17.5 17.7 18.3 17.6 23.8 35.8 37.5 33.1 4.2 5.0 5.6 6.0 7.4 7.5 7.9 12.1 9.7 18.9 36.1 41.7  HPO/  Si(OH)  uM  uM  0.6 0.8 1.1 1.5 2.6 6.5 0.7 2.4 2.6 2.8 2.0 1.9 1.7 1.9 6.8 2.5 3.6 0.2 0.3 0.3 0.5 0.6 0.6 1.5 1.6 1.4 2.7 2.0 2.3 1.5 3.0 1.6 2.2 2.6 3.7 3.6 2.7 1.7 1.8 1.6 2.7 2.9 2.9  26.8 24.7 13.8 20.2 37.5 91.7 13.7 34.7 42.9 50.3 45.4 48.1 45.1 52.6 154.1 66.0 107.6 16.7 17.3 9.4 10.9 10.0 9.9 18.7 20.0 23.9 44.9 27.8 40.1 38.4 72.1 57.9 77.6 99.1 113.5 113.5 113.6 30.5 30.5 25.3 38.4 37.3 38.5 30.2 31.2 30.3  1.3 1.5 1.5 0.9 1.3 1.8 1.8 1.7 1.7 2.3 3.4 3.1 2.9 0.7 0.7 0.8 0.8 0.9 1.0 1.0 1.2 1.1 1.4 2.3 2.7  28.3 40.3 23.9  24.9 30.6 57.7 52.3 47.9 15.3 15.0 16.1 16.2 15.2 15.2  13.4 23.9 19.4 33.9 45.7  4  Table B.3 continued. Dissolved nutrients in October 1999. Station G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 G7 J6 J6 J6 J6 J6 J6 J6 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 J8 BP2 BP2 BP2 BP2 BP2 BP2 BP2 BP2 BP2 BP2 BP2 BP5 BP5 BP5 BP5 BP5 BP5 BP5 BP5 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7  Date  Cast #  Niskin  28-Sep-99  98 98 98 98 98 98 98 98 98 98 98 98 84 84 84 84 84 84 84  12 11 10 9 8 7 6 5 4 3 2 1 19 18 17 16 15 14 13 20 19 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 12 11 10 8 7 6 5 4 3 2 1  27-Sept-99  28-Sep-99  27-Sep-99  27-Sep-99  27-Sep-99  81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 81 70 70 70 70 70 70 70 70 70 70 70 75 75 75 75 75 75 75 75 78 78 78 78 78 78 78 78 78 78  #  20 19 18 17 16 15 14 13 22 21 20 19 18 17 16 15 14 13  Depth m  N0 ' uM  175 200 250 300 400 500 600 800 1000 1200 1500 1800 0 10 20 30 50 75 100 0 10 30 50 75 100 125 150 175 200 250 300 400 500 600 800 1000 1200 1500 0 2 5 15 20 25 3 40 50 75 100 0 10 20 30 50 75 100 125 0 2 5 15 25 40 50 75 100 125  41.7 45.8 30.1 24.1 40.8 34.6 35.3 44.1 33.1 31.1 21.8 25.8 8.1 15.5 15.5 13.9 12.6 9.4 30.9 5.1 4.9 10.0 7.8 15.3 23.8 20.4 28.6 38.3 33.7 35.4 32.2 23.3 15.8 28.0 43.6 43.6 41.5 25.2 16.3 15.8 16.0 19.5 20.0 24.7 20.2 31.0 33.7 31.4 31.2  3  10.0 3.9 13.2 11.7 21.0 13.4 14.9 24.8 7.7 10.2 9.2 11.1 18.8 20.2 25.0 33.6 39.0 32.1  HPfV uM 2.8 2.9 1.9 1.9 3.3 2.9 2.7 3.5 2.6 2.5 1.9 1.9 0.5 1.6 0.0 1.3 1.2 0.5 1.3 1.1 1.1 1.4 1.2 1.8 1.5 1.2 1.9 2.4 2.2 2.4 2.5 1.9 1.5 2.4 3.7 3.7 3.5 2.2 1.4 1.4 1.4 1.5 1.6 2.0 1.4 2.0 2.4 2.0 2.5 1.2 0.3 1.3 1.1 1.7 1.1 1.3 2.0 0.9 1.3 1.3 1.4 2.2 2.3 2.2 2.1 2.4 2.1  Si(OH) uM 47.0 58.6 50.9 53.3 75.5 90.7 89.6 113.3 110.9 113.4 113.5 108.2 13.6 22.9 17.9 18.2 13.0 9.4 25.5 14.1 14.4 18.6 12.4 19.8 27.2 24.1 36.5 45.9 45.8 55.2 52.1 58.1 71.4 77.1 112.7 112.8 112.9 112.8 33.6 33.5 33.9 33.3  26.8 33.3 49.1 35.4 41.5 29.8 15.4 26.3 20.2 25.5 22.5 24.9 36.3 23.3 23.8 27.2 25.4 31.0 31.3 27.9 39.2 59.8 44.7  4  Table B.3 continued. Dissolved nutrients in October 1999. Depth N0 Niskin Station Date Cast m uM # # 3  BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 BP7 CPE2 CPE2 CPE2 CPE2 CPE2 CPE2 CPE2 CPE2 CPE2 CS7 CS7 CS7 CS7 CS7 CS7 CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS3B CS3B Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3 Q3  27-Sep-99  26-Sep-99  26-Sep-99  26-Sep-99  26-Sep-99  78 78 78 78 78 78 78 78 78 78 78 66 66 66 66 66 66 66 66 66 42 42 42 42 42 42 49 49 49 49 49 49 49 49 49 49 49 49 63 63 63 63 63 63 63 63 63 63 63 63 63 63 63 63 63  12 11 10 9 8 7 6 5 3 2 1 9 8 7 6 5 4 3 2 1 6 5 4 3 2 1 12 11 10 9 8 7 6 5 4 3 2 1 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1  150 175 200 250 300 400 500 600 800 1000 b-10 0 10 20 30 50 75 100 100 121 0 10 20 30 50 60 0 10 20 30 50 75 100 125 150 175 200 210 0 10 25 35 45 48 50 75 100 125 150 175 200 250 300 400 500  41.7 41.7 53.5 28.1 42.5 44.4 34.4 32.6 27.5 43.4 38.6 13.0 6.6 11.5 16.6 19.1 21.2 28.7 30.2 31.4 8.9 7.8 10.2 23.5 73.3 72.6 4.4 2.8 3.0 3.9 15.3 25.3 23.2 41.7 41.7 31.6 33.2 38.3 2.5 6.6 4.1 3.6 11.9 13.4 14.2 28.1 38.4 41.6 43.8 52.1 52.4 23.4 40.7 23.3 42.9  HPfV  Si(OH)  uM  uM  2.9 2.9 3.3 2.1 3.3 3.5 2.8 2.8 2.1 3.6 3.2 1.4 0.8 1.3 1.7 2.0 2.0 2.8 2.7 2.9 0.8 0.8 1.0 1.6 4.6 4.6 0.7 0.7 0.6 1.0 1.7 2.8 1.4 2.7 2.9 2.0 2.0 2.5 0.6 1.0 0.3 0.1 1.5 1.5 1.5 2.1 2.2 2.6 2.9 3.0 3.0 1.9 3.2 2.0 3.4  54.3 58.4 65.4 48.8 72.5 72.7 73.1 88.3 88.5 112.9 109.9 24.3 17.9 22.4 27.0 28.1 30.0 41.2 39.8 49.1 19.9 18.8 22.1 29.2 93.2 91.8 16.7 13.2 12.6 15.6 23.5 32.3 26.8 48.3 50.3 40.5 40.8 50.0 49.0 15.4 7.7 5.9 16.0 17.8 18.2 32.3 35.9 46.4 52.3 61.4 62.6 41.7 68.8 69.2 84.0  4  APPENDIX C 1999 R A W D A T A S E T  Table C.l Chlorophyll a (mg m") in May 1999 off the west coast of Vancouver Island. All samples werefilteredonto 0.7 pm glassfiberfilters unless otherwise indicated., by * which were filtered onto 5.0 pm polycarbonatefilters.(-) indicates data not available. Station Date Cast Niskin Depth Chl a # # (m) (mg m") 3  3  B2  4-May-99  4  B4  4-May-99  6  B6  5-May-99  8  B8  5-May-99  14  BIO  6-May-99  46  B12  6-May-99  44  B14  6-May-99  40  B16  6-May-99  43  B16* CI  6-May-99  43  5-May-99  9  CI* C2  5-May-99  9  5-May-99  21  5 4 3 1 7 6 5 4 7 6 5 4 3 5 4 3 2 1 9 8 7 6 5 15 14 13 11 18 17 16 15 14 12 11 10 9 8 7 12  0 10 20 50 0 10 20 30 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 50 0 10 20 30 50 0 10 20 30 40 50 0  4.4 4.9 2.1 1.0 14.5 8.2 6.0 2.6 8.3 12.9 6.3 2.5 0.6 0.9 1.3 2.6 4.7 1.7 1.0 1.1 0.6 0.6 0.5 0.6 0.7 0.5 0.8 0.5 0.6 0.7 0.6 0.7 0.4 0.4 0.5 0.4 0.5 1.1 0.7  12 11 10 8 7 12  0 2 5 15 2 0  2.9 5.5 8.3 11.5 6.1 3.7  7 6 5 4 3  0 10 20 30 50  2.6 3.2 3.3 0.5 0.3  Table C. 1 continued Station Date  Cast #  Niskin #  Depth (m)  Chl a (mg m )  11 10 9 8 7 6 11  0 5 10 15 20 30 0  2.9 1.1 0.6 1.3 3.6 0.4 0.1  10 8 6 5 4 12 11 10 9 8 16 15 14 14 13 12 18 17 16 15 14 5 4 3 2 1 7 6 5 4 3 5 4 3 2 1 7 6 5 4 3 12 11 10 9 7 6 12  0 10 20 30 50 0 10 20 30 50 0 5 10 20 25 30 0 10 20 30 50 0 10 20 30 41 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 5 10 20 30 40 0  2.0 0.6 0.9 0.7 0.7 1.3 1.2 0.6 0.6 0.8 1.0 1.0 1.2 0.4 0.3 0.6 0.5 0.5 0.9 0.8 1.4 4.6 4.3 4.6 1.2 1.6 3.5 3.8 0.3 0.2 0.3 3.8 2.1 1.7 1.8 1.2 1.2 1.6 1.9 0.3 0.2 0.8 0.8 0.9 0.9 0.9 0.6 0.1  16 15 14 13 12 11 16  0 10 20 30 50 0 5 10 20 30 40 0  0.9 1.2 1.0 0.7 0.1 0.6 0.6 0.4 0.3 0.5 0.8 0.1  C4  5-May-99  23  C4* C7  5-May-99  23  5-May-99  27  C8  5-May-99  28  C9  5-May-99  30  Cll  5-May-99  36  D2  6-May-99  48  D4  6-May-99  50  Gl  7-May-99  52  G2  7-May-99  53  G3  7-May-99  65  G3* G4  7-May-99  65  7-May-99  56  G7  6-May-99  59  G7*  6-May-99  59  3  Table C. 1 continued  Station  Date  Event #  Niskin #  Depth (m)  Chl a (mg m")  16 15 14 13 12 6 5 4 3 2 6 5 4 3 2 9 8 7 6 5 17 16 15 14 13 17 16 15 14 13 10 9 8 7 6 5 10  0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 2.5 5 10 15 20 0  0.2 0.2 0.3 0.4 2.7 8.4 8.2 7.6 1.8 0.4 7.5 7.9 2.2 0.7 0.4 3.2 3.0 2.4 0.9 0.3 2.5 2.5 1.6 0.6 0.4 3.2 3.4 1.1 0.8 0.2 3.8 4.0 3.9 4.1 3.7 4.2 2.8  17 16 15 14 13 16 15 14 13 12 16 15 14 13 12 11 8 16  0 10 20 30 50 0 10 20 30 50 0 2.5 5 10 15 20 100 0  3.7 4.3 5.7 1.6 1.0 3.8 3.3  21 20 19 18 17 15 14 13 12 11  0 10 20 30 50 0 10 20 30 50  0.6 0.6 0.4 0.5 0.3 3.3 5.1 7.3 2.7 1.1  G9  7-May-99  64  Jl  10-May-99  124  J2  10-May-99  125  J4  11-May-99  128  J6  11-May-99  131  J8  11-May-99  134  BP2  10-May-99  118  BP2*  10-May-99  118  BP4  7-May-99  117  BP6  10-May-99  114  BP7  10-May-99  123  BP7*  10-May-99  123  BP8  10-May-99  110  JI22  9-May-99  101  .  3  1.2 0.4 0.8 0.8 0.8 0.8 0.6 0.6 0.1 0.1  >  Table C. 1 continued Station Date  Event #  Niskin #  Depth (m)  Chl a (mg m~)  14 13 12 11 10 11 10 9 8 7 11 10 9 8 7 7 11  0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 50 0  8.0 7.8 8.3 6.9 0.7 3.3 3.2 1.8 9.8 0.4 1.9 1.6 1.4 0.9 0.5 0.4 1.1  6 5 4 3 2 6 5 4 3 2 11 10 9 8 7 8 7 6 5 4 6 5 4 3 2 17 16 15 14 13 16 15 14 13 12 16 15 14 13 12  0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50  5.7 5.1 5.9 4.2 5.0 5.2 5.4 5.0 4.2 2.3 5.1 6.5 6.1 5.0 2.2 5.0 5.9 2.5 0.4 0.3 5.7 4.6  CSl  8-May-99  67  CS3  8-May-99  73  CS3B  9-May-99  95  CS3B* CS5  9-May-99  95  8-May-99  77  CS7  8-May-99  82  CS9  8-May-99  88  CPE2  9-May-99  107  CS1B  9-May-99  93  MP8  12-May-99  137  P4  12-May-99  142  P6  12-May-99  142  3  5.7 2.9 2.3  0.7 0.7 0.6 0.2 0.3 0.6 0.6 0.6 0.4 0.3 0.6 0.5 0.6 0.3 0.3  APPENDIX C Table C.2 Chlorophyll a (mg m") in July 1999 off the west coast of Vancouver Island. All samples were filtered onto 0.7 um glass fiber filters unless otherwise indicated by * which were filtered onto a 5.0 um polycarbonate filters. Station Date Cast Niskin Depth Chl a # # (mg m ) (m) 3  3  B6  30-Jun-99  6  B8  30-Jun-99  8  B16  1-Jul-99  18  CI  30-Jun-99  33  C4  1-Jul-99  36  C4* C5  1-Jul-99  36  2-Jul-99  38  C6  2-Jul-99  39  C7  2-Jul-99  40  C8  2-Jul-99  41  C9  2-Jul-99  42  C9* Cll  2-Jul-99  42  2-Jul-99  46  9 8 7 6 5 4 3 9 8 7 6 5 5 4 3 2 1 7 6 5 4 3 14 12 11 9 8 7 13  0 5 10 20 30 50 75 0 10 20 30 50 0 10 20 30 40 0 10 20 30 50 0 6 5 18 34 41 5  2.50 2.45 2.27 1.80 0.94 0.34 0.17 8.32 4.81 0.94 0.79 0.35 0.85 0.87 0.90 0.42 0.05 6.47 4.20 1.02 1.06 0.19 4.16 3.77 4.27 0.54 0.18 0.22 3.45  6 5 4 3 2 7 6 5 4 3 8 7 6 5 4 11 10 9 8 7 16 15 14 13 12 16  0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0  3.47 2.93 1.22 0.70 0.32  20 19 18 17 16  0 10 20 30 50  0.43 0.29 0.34 0.38 0.74  0.70 0.74 2.07 0.98 0.23 0.56 0.74 1.46 0.83 0.16 0.44 0.41 1.08 0.970.19 0.46 0.41 0.55 0.60 0.52 0.12  Table C.2. continued  Station  Date  Cast #  Niskin #  Depth (m)  Chl a (mg m")  5 4 3 2 1 7 6 5 4 3 8 7 6 5 4 3 8  0 10 20 30 50 0 10 20 30 50 0 5 10 20 30 40 0  6.66 6.24 0.60 0.27 0.12 7.82 4.81 0.40 0.09 0.15 1.82 2.01 2.02 0.81 0.32 0.05 1.34  17 16 15 14 13 22 21 20 19 18 17 9 8 7 6 5 5 10 9 8 7 6 19 18 17 15 16 17 16 15 14 13 6 5 4 3 2 1 6  0 10 20 30 50 0 5 10 20 30 40 3.5 6 10 15 18 18 0 10 20 30 50 0 10 20 50 30 0 10 20 30 50 10 11 15 22 31 37 0  1.98 2.00 1.73 0.87 0.27 1.07 0.89 0.92 0.68 1.34 0.32 1.60 3.13 3.43 3.31 2.51 2.07 1.90 4.70 3.93 1.59 0.20 2.93 3.70 3.05 0.34 0.82 2.80 2.64 1.49 1.21 0.31 3.93 4.24 3.47 2.34 0.64 0.39 1.34  Gl  3-Jul-99  61  G2  3-Jul-99  63  G3  3-Jul-99  65  G3* G6  3-Jul-99  65  3-Jul-99  70  G7  3-Jul-99  71  BP2  6-Jul-99  94  BP3  6-Jul-99  91  BP4  6-JUI-99  90  BP6  5-Jul-99  89  BP7  5-Jul-99  86  BP7*  5-Jul-99  86  3  APPENDIX C  Table C.3. Chlorophyll a (mg m") in October 1999 off the west coast of Vancouver Island. All samples werefilteredonto 0.7 um glassfiberfiltes unless otherwise indicated by * which were filtered onto a 5.0 um polycarbonate filters. J  Station  Date  Cast  Niskin  #  #  Depth (m)  Chl a (mg m")  7 5 4 3 8 7 6 5 4 14 13 12 10 8 6 14  0 20 30 50 0 10 20 30 50 0 2 5 15 25 40 0  2.66 1.08 0.75 0.39 3.12 2.91 1.76 0.72 0.29 4.51 4.58 4.70 2.89 1.08 0.32 1.92  115  12 11 10 8 7 6 12  0 2 5 20 30 50 0  4.89 5.31 4.93 1.01 1.15 0.46 3.67  9 8 7 6 5 11 10 9 8 7 22 21 20 18 16 14 22  0 10 20 30 50 0 10 20 30 50 0 2 5 15 25 40 0  6.85 7.16 2.71 1.56 0.31 4.85 4.77 2.93 1.96 0.35 4.97 5.47 6.12 3.31 0.50 0.17 3.54  20 19 18 17 16 4 3 2 1 5 4 3 2 6 5 4 3 2  0 10 20 30 50 0 5 10 30 0 10 20 30 0 10 20 30 50  5.20 4.77 1.88 0.89 0.23 9.55 8.90 0.93 0.42 6.62 6.51 1.58 0.42 5.93 1.90 1.42 0.83 0.31  B4  30-Sep-99  131  B6  30-Sep-99  130  C1  30-Sep-99  119  C1* C4  30-Sep-99  C4* C7  30-Sep-99 29-Sep-99  110  C8  29-Sep-99  108  C9  29-Sep-99  106  30-Sep-99  C9*  29-Sep-99  106  C11  29-Sep-99  104  D1  23-Sep-99  21  D2  23-Sep-99  22  D4  23-Sep-99  25  3  Table C.3. continued Station Date  Cast #  Niskin #  Depth (m)  Chl a (mg m )  17 16 15 14 13 20 19  0 10 20 30 50 0 10 20 30 50 0 10 20 30 50 0 2 5 15 25 40 0 0 0 5 10 25 40 0 0 10 20 30 50 0 10 20 30 50 0 2 5 15 25 40 0 0 10 20 30 50 0 2 5 15 25 40 0 0 10 20 30 50  7.62 3.85 0.48 0.16 0.12 6.89 7.66 1.76 0.25 0.12 7.43 7.20 2.56 0.11  D8  23-Sep-99  30  D10  24-Sep-99  32  18 17  Gl  28-Sep-99  88  G3  28-Sep-99  92  G3* G7  28-Sep-99 28-Sep-99  98  G7*  28-Sep-99 27-Sep-99  84  J6  16 6 5 4 3 2 13 12 11 9 7 5 13 24 24 23 21 19 17 24 19 18 17 16  J8  27-Sep-99  81  BP2  26-Sep-99  70  BP2* BP5  26-Sep-99 27-Sep-99  75  15 20 19 18 17 16 12 11 10 8 6 4 12 20 19 18 17 16  BP7  27-Sep-99  BP7* CS7  27-Sep-99 25-Sep-99  78  42  22 21 20 19 18 17 22 6 5 4 3 2  3  0.13  8.16 7.66 8.74 4.69 0.40 0.13 5.35 0.78 1.90 1.94 1.72 1.62 0.54 1.50 2.76 3.00 2.53  1.72  0.13 1.79 1.94 2.69 2.83 0.28 6.08 6.08 5.97 3.89 1.47 0.37 4.66 8.74 7.28 3.20  1.32  0.13 9.40 10.01 9.70 1.90 0.32 0.26 3.24 1.95 2.30 2.09 0.25 0.05  Table C.3. continued Station Date  Cast #  Niskin #  Depth (m)  Chl a (mg m )  9 8 7 6 5 12 11 10 9 8 17 15 14 13 12 11 17  0 10 20 30 50 0 10 20 30 50 0 25 35 45 48 50 0  4.51 4.20 2.08 1.73 0.67 1.50 1.57 1.69 1.36 0.07 1.38 0.37 0.16 0.28 0.10 0.09 0.80  CPE2  26-Sep-99  66  CS3B  26-Sep-99  49  Q3  26-Sep-99  63  Q3*  26-Sep-99  63  3  APPENDIX D P R I M A R Y P R O D U C T I V I T Y F O R 1999  Table D . l Integrated (100-1% surface light) daily primary productivity (g C m" d") in 1999 for July and October off the west coast of Vancouver Island. Daily Production Date Station 2  1  29 June 30 June 07 July 01 July 03 July 05 July  CI C4 G3 C9 G7 BP7  0.2 2.0 0.8 0.4 0.9 1.9  30 Sept. 30 Sept. 38 Sept. 26 Sept. 29 Sept. 28 Sept. 27 Sept.  CI C4 G3 BP2 C9 G7 BP7  0.6 0.7 1.6 0.6 0.6 0.3 0.8  APPENDIX E  Sampling Stations In addition to specific time series and process study stations, water samples were taken at additional stations along transects off the west coast of Vancouver Island. Tables E.1-E.6 show latitude, longitude and water depth for all stations sampled during cruises in 1997 and 1998.  In this thesis data are presented for 1997 and 1998 for La  Perouse Bank, Barkley Canyon, Estevan Point and the Brooks Peninsula transects only. Physical, chemical, and biological data are presented in Chapter 1 and size-fractionated phytoplankton biomass and primary productivity in Chapter 2. All raw data for the above transects and the additional stations sampled during each cruise are archived in the Institute of Ocean Science's database. During three cruises in 1999 a reduced number of stations were sampled. Tables E.7-E.9 show latitude, longitude and water depth data of all stations sampled in 1999. Data for 1999 are presented in Appendices B, C and D and also archived in the database at the Institute of Ocean Science (Sidney, BC).  151  APPENDIX E Table E . l Location and water depth of stations occupied during 08-24 April 1997 (Cruise ID#9707) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, D indicates D Line, G indicates Estevan Point, BP indicates Brooks Peninula and CS indicates Cape Scott transects. See Figure 1.1 for location of transects. Bottom Depth Station Longitude Transect Latitude (m) deg° min' sec" W deg° min' sec" N 56 B2 La Perouse Bank 125° 02' 24" 48° 39' 00" 135 B8 125° 28' 39" 48° 25' 18" 153 B9 125° 34' 48" 48° 22' 00" 219 Bll 48° 15' 12" 125° 47' 45" 450 B12 48° 12' 55" 125° 51' 54" B14 966 48° 08' 29" 126° 00' 00" 1530 B16 126° 17' 00" 48° 00' 32" 156 Barkley Canyon CI 125° 15' 14" 48° 28' 59" C2 125° 30' 57" 111 48° 48' 41" 166 C4 48° 43' 28" 125° 40' 48" 129 C7 48° 32' 58" 126° 00' 30" 201 C8 126° 07' 06" 48° 29' 27" 659 C9 48° 25' 24" 126° 13' 42" 1115 CIO 126° 20' 12" 48° 22' 24" 1330 Cll 126° 26' 42" 48° 18' 57" 45 D2 D Line 125° 47' 03" 48° 58' 21" 60 D4 125° 57' 00" 48° 53' 10" 463 D7 126° 16' 48" 48° 42; 40" 1100 D9 126° 30' 00" 48° 35' 41" 60 Estevan Point Gl 126° 35' 00" 49° 20' 30" 126 G3 126° 43'42" 49° 15' 00" 150 G4 126° 49' 24" 49° 11' 18" 250 G5 126° 55' 18" 49° 07' 24" 972 G6 49° 03' 30" 127° 01' 12" 1750 G7 127° 07' 12" 48° 59' 24" 99 BP2 Brooks Peninsula 127° 54' 12" 50° 04' 00" 139 BP3 127° 55' 18" 50° 03' 12" 1090 BP4 50° 04'24" 127° 58' 06" 1230 BP5 128° 00' 00" 50° 00' 00" 1730 BP6 128° 05' 30" 49° 56' 12" 2200 BP7 128° 11' 12" 49° 52' 24" 1967 J023 Cape Scott 50° 39' 48" 129° 01' 54" 64 CS2B 128° 59' 52" 50° 55' 59" 158 CPE1 127° 50' 00" 51° 00' 00" 123 CPE2 50° 43' 00" 128° 40' 00"  152  APPENDIX E Table E.2 Location and water depth of stations occupied during 14-28 July 1997 (Cruise ID#9713) off the west coast of Vancouver Island. Under the station column, A indicates Juan de Fuca canyon, B indicates La Perouse Bank, C indicates Barkley Canyon, G indicates Estevan Point and BP indicates Brooks Peninula transect. See Figure 1.1 for location of transects. Transect Station Latitude Longitude Depth (m) deg° min' sec" N deg° min' sec" W Al 264 Juan de Fuca Canyon 48° 29' 14" 124°43'39" A4 175 125° 04' 07" 48° 19' 22" 112 A6 48° 12' 38" 125° 17' 12" 950 A10 125°43'24" 47° 59' 01" La Perouse Bank  Barkley Canyon  D Line  Estevan Point  Brooks Peninsula  B2 B8 B9 BIO B12 B14 B16 CI C2 C4 C7 C8 C9 CIO Cll D2 D4 D7 D9  48° 39' 00" 48° 25' 18" 48° 22' 00" 48° 18' 34" 48° 12' 55" 48° 08' 29" 48° 00' 32" 48° 28' 59" 48° 48' 41" 48° 43' 28" 48° 32' 58" 48° 29' 27" 48° 25' 24" 48° 22' 24"  Gl G3 G4  49° 20' 30" 49° 15' 00" 49° 11' 18" 49° 07' 24" 50° 04' 48" 50° 04' 00" 50° 03' 12" 50° 04' 24" 50° 00' 00" 49° 56' 12" 49° 52' 24"  G5 BP1 BP2 BP3 BP4 BP5 BP6 BP7  48° 48° 48° 48° 48°  18' 57" 58' 21" 53' 10" 42' 40" 35' 41"  125° 02' 24" 125°28'39" 125° 34' 48" 125° 41'21" 125°51'54" 126° 00' 00" 126°17' 00" 125° 15' 14" 125° 30'57" 125° 40' 48" 126°00'30" 126° 07' 06" 126° 13' 42" 126° 20' 12" 126° 26'42" 125° 47' 03" 125°57'00" 126° 16' 48" 126° 30' 00" 126° 35' 00" 126° 43' 42" 126° 49' 24" 126° 55' 18" 127° 52' 48" 127° 54' 12" 127° 55'18" 127° 58' 06" 128° 00' 00" 128° 05' 30" 128° 11' 12"  56 135 153 151 450 966 1530 156 111 166 129 201 659 1115 1330 45 60 463 1100 60 126 150 250 32 99 139 1090 1230 730 2200  153  APPENDIX E  Table E.3 Location and water depth of stations occupied during 20-27 October 1997 (Cruise ID#9737) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, D indicates Line D, G indicates Estevan Point and BP indicates Brooks Peninula. See Figure 1.1 for location of transects. Depth Longitude Station Latitude Transect deg°min' sec" W (m) deg°min' sec" N 145 B2 125° 02' 28" La Perouse Bank 48° 39' 36" 145 B8 125° 28' 43" 48° 25' 17" 151 B9 125° 34' 49" 48° 22' 00" 153 BIO 125° 41' 13" 48° 18' 35" 1185 B14 125° 00' 07" 48° 08' 27" 156 CI 125° 15' 14" Barkley Canyon 48° 28' 59" C4 166 125° 40' 48" 48° 43' 28" 129 C7 126° 00' 30" 48° 32' 58" 201 C8 126° 07' 06" 48° 29' 27" 656 C9 126° 13' 42" 48° 25' 24" 1115 CIO 126° 20' 12" 48° 22' 24" 1330 Cll 126° 26' 42" 48° 18' 57" 33 Dl 125° 43' 48" D Line 49° 00" 06" 45 D2 125° 47' 03" 48° 58' 21" 60 D4 125° 57' 00" 48° 53' 10" 137 D6 126° 10' 12" 48° 46' 11" 463 D7 126° 16' 48" 48° 42' 40" 60 Estevan Point Gl 126° 35' 00" 49° 20' 30" 102 G2 126° 38' 06" 49° 18' 42" 126 G3 126° 43'42" 49° 15' 00" 150 G4 126° 49' 24" 49° 11' 18" 250 G5 126° 55' 18" 49° 07' 24" 972 G6 127° 01' 12" 49° 03' 30" 1750 G7 127° 07' 12" 48° 59' 24" 32 BP1 127° 52' 48" Brooks Peninsula 50° 04' 48" 99 BP2 127° 54' 12" 50° 04' 00" 139 BP3 127° 55' 18" 50° 03' 12" 1090 BP4 127° 58' 06" 50° 04' 24" 1230 BP5 128° 00' 00" 50° 00' 00" 1730 BP6 128° 05' 30" 49° 56' 12" 2200 BP7 128° 11' 12" 49° 52' 24"  154  APPENDIX E Table E.4 Location and water depth of stations occupied during 11-25 May 1998 cruise (Cruise ID# 9810) off the west coast of Vancouver Island. Under the station column, B indicates L a Perouse Bank, C indicates Barkley Canyon, G indicates Estevan Point, H indicated H Line, J indicates J Line, BP indicates Brooks Peninula and CS indicates Cape Scott transect. See Figure 1.1 for location of transects. Depth Longitude Station Latitude Transect deg° min' sec" N deg° min' sec" W (m) 160 B7 La Perouse Bank 125° 28'06" 48° 25' 41" 145 B8 125° 28' 39" 48° 25' 18" 1530 B16 48° 00' 32" 126° 17' 00" 156 CI Barkley Canyon 125° 27' 44" 48° 50' 26" C2 125° 30' 57" 111 48° 48' 41" 120 C3 125° 34' 14" 48° 46' 57" 129 C7 126° 00' 30" 48° 32' 58" 201 C8 126° 07' 06" 48° 29' 27" 659 C9 126° 13' 42" 48° 25' 24" 1330 Cll 126° 26' 42" 48° 18' 57" 1000 C12 126° 40' 00" 48° 15' 00" 60 Estevan Point Gl 126° 35' 00" 49° 20' 30" 105 G2 126° 38' 06" 49° 18' 42" 126 G3 126°43'42" 49° 15' 00" 972 G6 127° 01' 12" 49° 03' 30" 1750 G7 127° 07' 12" 48° 59' 24" 2081 G9 127° 19' 21" 48° 51' 10" 41 H2 49° 32' 15" H Line 126° 47' 17" 97 49° 28' 37" H3 126° 52' 54" 151 49° 21' 16" H5 126° 04' 45" 1001 49° 13' 38" H7 126° 16' 07" 2079 49° 05' 31" H9 126° 28' 44" 150 49° 42' 40" J2 J Line 126° 05' 18" 150 14 49° 35' 25" 126° 16' 46" 1001 49° 27' 29 " J6 126° 28' 35" 99 BP2 Brooks Peninsula 127° 54' 12" 50° 04' 00" 1090 BP4 127° 58' 06" 50° 04' 24" 1230 BP5 128° 00' 00" 50° 00' 00" 1730 BP6 128° 05' 30" 49° 56' 12" 2200 BP7 128° 11' 12" 49° 52' 24" >2200 BP8 128° 16' 48" 49° 48' 36" 49° 44' 47" >2200 BP9 128° 22' 48" 150 CS3B Cape Scott 128° 50' 48" 51° 00' 00"  APPENDIX E Table E.5 Location and water depth of stations occupied during 14-26 July 1998 (Cruise ID#9823) off the west coast of Vancouver Island. Under the station column, A indicates Juan de Fuca canyon, B indicates La Perouse Bank, C indicates Barkley Canyon, D indicates D Line, G indicates Estevan Point, BP indicates Brooks Peninula and E R indicated Endeavor Ridge transect. See Figure 1.1 for location of transects. Transect Station Longitude Depth Latitude deg° min' sec" N deg° min' sec" W (m) 264 Al Juan de Fuca Canyon 124° 43' 39" 48° 29' 14" A2 312 124° 51" 19" 48° 26' 15" A4 175 48° 19' 22" 125° 04' 07" 112 A6 125° 17' 12" 48° 12' 38" A8 142 48° 05' 47" 125° 30' 24" 156 CI La Perouse Bank 48° 28' 59" 125° 15' 14" 1530 B16 48° 00' 32" 126° 17' 00" 94 Barkley Canyon CI 125° 27' 44" 48° 50" 26" C2 111 125° 30' 57" 48° 48' 41" 129 C7 48° 32' 58" 126° 00' 30" 201 C8 126° 07' 06" 48° 29' 27" 659 C9 48° 25' 24" 126° 13' 42" 1115 CIO 126° 20' 12" 48° 22' 24" 1330 Cll 126° 26' 42" 48° 18' 57" 33 D Line Dl 48° 49' 06" 125° 43" 48" D2 45 48° 58' 21" 125° 47' 03" D4 60 48° 53' 10" 125° 57' 00" D7 463 48° 42' 40" 126° 16' 48" D9 1100 126° 30' 00" 48° 35' 41" 60 Estevan Point Gl 49° 20' 30" 126° 35' 00" G2 105 126° 38' 06" 49° 18' 42" G3 126 49° 15' 00" 126° 43'42" G4 150 49° 11' 18" 126° 49' 24" G5 258 49° 07' 24" 126° 55' 18" G7 1800 48° 59' 24" 127° 07' 12" 32 Brooks Peninsula BP1 50° 04' 48" 127° 52' 48" BP2 99 50° 04' 00" 127° 54' 12" 139 BP3 127° 55' 18" 50° 03' 12" BP4 1090 50° 04' 24" 127° 58' 06" 1230 BP5 50° 00' 00" 128° 00' 00" 1730 BP6 128° 05' 30" 49° 56' 12" 2200 BP7 49° 52' 24" 128° 11' 12" Endeavor Ridge ER01 2200 47° 57' 29" 129° 05' 03" M3 1180 50° 05' 23" 128° 10' 12"  156  APPENDIX E  Table E.6 Location and water depth of stations occupied during the 05-16 October 1998 cruise (Cruise ID#9836) off the west coast of Vancouver Island. Under the station column, indicates La Perouse Bank, C indicates Barkley Canyon, G indicates Estevan Point, BP indicates Brooks Peninula and CS indicated Cape Scott. See Figure 1.1 for location of transects. Longitude Depth Station Latitude Transect deg° min' sec" N deg° min' sec" W (m) 156 CI La Perouse Bank 125° 15' 14" 48° 28' 59" 1530 B16 126° 17' 00" 48° 00' 32" 94 CI Barkley Canyon 125° 27' 44" 48° 50' 26" C2 111 125° 30' 57" 48° 48' 41" 167 C4 125° 40' 48" 48° 43' 28" 129 C7 126° 00' 30" 48° 32' 58" 201 C8 126° 07' 06" 48° 29' 27" 659 C9 126° 13' 42" 48° 25' 24" 1115 CIO 126° 20' 12" 48° 22' 24" 1330 Cll 48° 18' 57" 126° 26' 42" 60 Estevan Point Gl 126° 35' 00" 49° 20' 30" 126 G3 126° 43'42" 49° 15' 00" 150 G4 126° 49' 24" 49° 11' 18" 257 G5 126° 55' 18" 49° 07' 24" 1800 G7 127° 07' 12" 48° 59' 24" 32 BP1 Brooks Peninsula 127° 52' 48" 50° 04' 48" 99 BP2 127° 54' 12" 50° 04' 00" 139 BP3 127° 55' 18" 50° 03' 12" 1090 BP4 127° 58' 06" 50° 04' 24" 1230 BP5 128° 00' 00" 50° 00' 00" 1730 BP6 128° 05' 30" 49° 56' 12" 2200 BP7 128° 11' 12" 49° 52' 24" >2000 CS1 Cape Scott 129° 41' 30" 50° 34' 54" 200 CS3 129° 20' 00" 50° 45' 36" 65 CS6 51° 00' 00" 128° 50' 00" 150 CPE1 127° 50' 00" 51° 00'00" 140 CPE2 128° 40' 00" 50° 43'00"  157  APPENDIX E Table E.7 Location and water depth of stations occupied during May 1999 (Cruise ID#9911) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, D indicates D Line, G indicates Estevan Point, J indicates J Line, BP indicates Brooks Peninula and CS indicates Cape Scott. See Figure 1.1 for location of transects. Longitude Depth Latitude Station Transect deg° min' sec" W (m) deg° min' sec" N 94 125° 27' 44" CI 48° 50' 26" La Perouse Bank 56 B2 125° 02' 24" 48° 39' 00" 105 125° 08'43" B4 48° 35' 40" 110 125° 15' 31" B6 48° 32' 40" 135 125° 28' 39" B8 48° 25' 18" 151 BIO 125° 41' 21" 48° 18' 34" 450 B12 125° 51' 54" 48° 12' 55" 966 B14 126° 00' 00" 48° 08' 29" 1530 126° 17' 00" B16 48° 00' 32" 111 C2 125° 30' 57" 48° 48' 41" Barkley Canyon C4 125° 40' 48" 166 48° 43' 28" 129 C7 126° 00' 30" 48° 32' 58" 201 C8 126° 07' 06" 48° 29' 27" 659 126° 13'42" C9 48° 25' 24" 1115 Cll 126° 20' 12" 48° 22' 24" 1330 C12 126° 26' 42" 48° 18' 57" 45 D2 125° 47' 03" D Line 48° 58' 21" 60 D4 125° 57' 00" 48° 53' 10" 60 Gl 126° 35' 00" Estevan Point 49° 20' 30" 105 G2 126° 38' 06" 49° 18' 42" 126 126° 43'42" G3 49° 15' 00" 150 G4 126° 49' 24" 49° 11' 18" 250 G7 126° 55' 18" 49° 07' 24" 2081 127° 19' 21" G8 48° 51' 10" 63 49° 44' 18" 127° 02' 30" Jl J Line 79 49° 42' 40" J2 126° 05' 18" 150 49° 35' 25" 14 126° 16' 46" 1001 49° 27' 29 " 126° 28' 35" 36 >2000 49° 20' 07 " J8 126° 40' 58" 99 BP2 127° 54' 12" Brooks Peninsula 50° 04' 00" 1090 BP4 127° 58' 06" 50° 04' 24" 730 128° 05'30" BP6 49° 56' 12" >2200 BP7 128° 11' 12" 49° 52' 24" >2200 BP8 128° 16' 48" 49° 48' 36" >2200 J122 Cape Scott 129° 17' 36" 50° 39' 48" >2200 129°41'30" CS1 50° 34' 54"  158  APPENDIX E Table E.8 Location and water depth of stations occupied during July 1999 (Cruise ID#9928) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, G indicates Estevan Point and BP indicates Brooks Peninula. See Figure 1.1 for location of transects. Longitude Depth Latitude Station Transect deg° min' sec" W (m) deg° min' sec" N 110 125° 15' 31" B6 La Perouse Bank 48° 32' 40" 135 125° 28' 39" B8 48° 25' 18" 1530 126° 17' 00" B16 48° 00' 32" 90 125° 27' 44" CI 48° 50' 26" Barkley Canyon C4 48° 43' 28" 125° 40' 48" 162 90 125° 47' 24" C5 48° 39' 56" 95 125° 54' 00" C6 48° 36' 28" 129 126° 00' 30" C7 48° 32' 58" 201 126° 07' 06" C8 48° 29' 27" 659 C9 126° 13' 42" 48° 25' 24" 1115 126° 20' 12" Cll 48° 22' 24" 60 126° 35' 00" Gl Estevan Point 49° 20' 30" 105 126° 38' 06" G2 49° 18' 42" 126 126° 43'42" G3 49° 15' 00" 866 127° 07' 12" G6 49° 03' 30" 250 126° 55' 18" G7 49° 07' 24" 99 127° 54' 12" BP2 50° 04' 00" Brooks Peninsula 172 127° 55' 18" BP3 50° 03' 12" 1090 127° 58' 06" BP4 50° 04' 24" 1730 128° 05' 30" BP6 49° 56' 12" 2200 128° 11' 12" BP7 49° 52' 24"  159  APPENDIX E Table E.9 Location and water depth of stations occupied during October 1999 (Cruise ID#9935) off the west coast of Vancouver Island. Under the station column, B indicates La Perouse Bank, C indicates Barkley Canyon, D indicates D Line, G indicates Estevan Point, J indicates J Line, BP indicates Brooks Periinula and CS indicates Cape Scott transect. See Figure 1.1 for location of transects. Longitude Depth Station Latitude Transect deg° min' sec" W (m) deg° min' sec" N 156 CI 125° 15' 14" La Perouse Bank 48° 28' 59" 105 B4 125° 08' 43" 48° 35' 40" 110 B6 125° 15' 31" 48° 32' 40" 162 C4 125° 40' 48" Barkley Canyon 48° 43' 28" C7 126° 00' 30" 129 48° 32' 58" 201 126° 07' 06" C8 48° 29' 27" 659 C8 126° 13' 42" 48° 25' 24" 1330 Cll 126° 26' 42" 48° 18' 57" 33 125° 43" 48" Dl Line D 48° 49' 06" 45 D2 125° 47' 03" 48° 58' 21" 60 D4 125° 57' 00" 48° 53' 10" 760 D8 126° 23' 24" 48° 39' 10" 1475 126° 36' 36" D10 48° 32' 12" 60 Gl 126° 35' 00" Estevan Point 49° 20' 30" 126 126° 43'42" G3 49° 15' 00" 1800 G7 127° 07' 12" 48° 59' 24" 1001 49° 27' 29" J6 126° 28' 35" Line J >2000 49° 20' 07" 126° 40' 58" J8 99 BP2 127° 54' 12" Brooks Peninsula 50° 04' 00" 1230 BP5 128° 00' 00" 50° 00' 00" 2200 BP7 128° 11' 12" 49° 52' 24" 65 CS7 128° 44' 00" Cape Scott 51° 04' 30" 140 CPE2 128° 40' 00" 50° 43' 00" CS3B 129° 27' 00" 51° 00' 00" 750 Q3 129° 01' 54" 50° 39' 48"  160  APPENDIX F  CONVERSION OF C E L L S I /  1  TO CARBON L  For the analysis o f the taxonomy data conversion from cell L " to carbon L " is 1  necessary.  1  The equations given by Strathman (1967) were used for all species and are listed  below: L o g Carbon = (-0.422) + 0.758 ( L o g Volume)  Diatoms only  L o g Carbon = (-0.460) + 0.866 ( L o g Volume)  Other than Diatoms  C e l l volumes specific to each species were required for the conversion o f cells L " to 1  carbon. Cell volume calculated using measurements o f representative cells and equations for simple geometric shapes: spheroids, cylinders, boxes, and cones were supplied by R. Haigh (unpubl. data). Collection o f this information is a time consuming task and the author greatly appreciated the efforts o f R. Haigh.  Cell specific volumes for diatoms are listed in Tables F . l  and F.2 for other phytoplankton species.  161  Table F . l Cell volume and carbon per cell of the diatom species observed off the west coast of Vancouver Island during 1997 and 1998. P indicates pennate diatoms and C indicates centric diatoms.  Diatom  Type  Shape  Volume  C/cell  (10 pm ) 1.92 1.40 1.53 5.20 1.55 13.2 0.67 0.37 0.12 1.87 0.26 9.19 6.21 2.88 0.20 1.07 0.03 9.92 2.94 0.29 2.22 0.14 10.9 7.0 32.9 2.77 0.60 1.92 0.99 32.1 0.03 0.23 0.77 3.94 14.5 16.8  (10" pg/cell)  3  Asterionella glacialis Centric spp. Chaetoceros compressus Chaetoceros convolutus Chaetoceros debilis Chaetoceros eibenii Chaetoceros radicans Chaetoceros socialis Chaetoceros socialis, spore Chaetoceros spp. Cylindrotheca closterium Detonela pumila Fragilaria spp. Leptocylindrus danicus Leptocylindrus minimus Navicula spp. Nitzschia spp. Pennates (> 50 um) Pennates (25-50 um) Pseudo-nitzschia delicatissima Pseudo-nitzschia pungens Pseudo-nitzschia spp. Proboscia alata Dactyliosolen fragilissimus Rhizosolenia setigera Rhizosolenia stolterfothii Skeletonema costatum Synedra spp. Thalassionema nitzschioides Thalassiosira aestivalis Thalassiosira spp. (< 5 um) Thalassiosira spp. (< 10 um) Thalassiosira spp. Thalassiosira cotula Thalassiosira nordenskioeldii Thalassiosira rotula  P  c c c c c c c c c c c P  c c P P P P P P P  c c c c P P P  c c c c c c c  Triangle box Cylinder Cylinder Cylinder Cylinder Cylinder Cylinder Cylinder Cylinder Cylinder Ellipsoid Cylinder Rect. Box Cylinder Cylinder Cylinder Cylinder Box Box Cylinder Cylinder Cylinder Cylinder Cylinder Cylinder Cylinder Cylinder Elliptic Cylinder Rect. box Cylinder Cylinder Cylinder Cylinder Cylinder Cylinder Cylinder  3  4  1.17 0.92 0.98 2.48 0.99 5.02 0.52 0.34 0.15 1.14 0.26 3.82 2.84 1.59 0.21 0.75 0.05 4.05 1.61 0.28 1.30 0.16 4.35 3.11 10.1 1.54 0.48 1.16 0.70 9.86 0.05 0.24 0.58 2.01 0.39 6.05  162  Table F.2 Cell volume and carbon per cell for other phytoplankton species (except diatoms) observed off the west coast of Vancouver Island during 1997 and 1998. P indicates autotrophic and H indicates heterotrophic nutrition.  Type  Species  Shape  Volume (10 pm )  C/cell (10 pg/cell)  3  Dinoflagellates Alexandrium tamarense Ceratium furca Ceratium kofoidii Dinophysis spp. Alexandrium pseudogoniaulax Gymnodinium auratum  3  -4  P p P P P P P P P H H P P P H H P H  Ellipsoid Cone Ellipsoid Ellipsoid Ellipsoid EHipsoid Ellipsoid Ellipsoid Cone Ellipsoid Ellipsoid Ellipsoid Ellipsoid Ellipsoid Spherical Ellipsoid Ellipsoid Ellipsoid  26.4 96.3 8.89 23.8 47.0 1.56 1.56 26.3 9.07 0.42 17.0 0.29 0.68 3.65 149 6.53 7.09 3.33  23.4 71.8 9.12 2.36 38.6 2.02 2.02 23.3 9.28 0.64 16.0 0.48 0.99 4.22 106 6.97 7.5 3.89  Flagellates, misc Leucocryptos marina Mantoniella squamata Micromonas pusilla  P P P P P P P P P P P P  Ellipsoid Ellipsoid Ellipsoid Spheroid Spheroid Spheroid Ellipsoid Spheroid Ellipsoid Spheroid Spheroid Spheroid  0.02 0.04 0.03 0.03 0.24 0.06 0.47 4.88 0.01 0.91 0.01 0.002  0.04 0.09 0.07 0.06 0.41 0.13 0.72 5.42 0.03 1.26 0.02 0.01  Others Ciliates, misc Hetersigma carterae Mesodinium rubrum Mesodinium rubrum,  H P P P  Ellipsoid Ellipsoid Ellipsoid Ellipsoid  0.86 0.45 9.35 0.01  1.20 0.69 9.52 0.02  Gymnodinium spp. Small Gymnodinium spp. Gyrodinium fusiforme Gyrodinium spp. small Gyrodinium spp. Avg. Katodinium rotundatum Prorocentrum balticum Prorocentrum gracile Protoperidinium rhomboidalis Protoperidinium spp.  Unidentified-autotroph Unidentified-heterotroph Nanoflagellates (2-20 um)  Choanoflagellate spp. (< 5 um) Choanoflagellate spp. (<10 pm) Choanoflagellate spp. Chrysochromulina spp. (< 5 um) Chrysochromulina spp. (avg) Coccolithophores Cryptomonad spp. Dictyocha speculum-silicoflagellate  small  APPENDIX G  INCIDENT SURFACE IRRADIANCE DURING PRIMARY PRODUCTIVITY MEASUREMENTS  The  incident surface  solar irradiance during primary productivity measurements  presented in Chapter 2 is shown in Figures G1-G.6. Data are not available for October 1998 due to a datalogger malfunction. During April 1997 (Figure G.l) primary productivity incubation length was 24 hours. For the rest of the study period, primary productivity incubation length was 6 hours and the incubation period is indicated by the shaded region in each plot.  164  2 0 0 0 - La Perouse Bank Shelf  La Perouse Bank Beyond Shelf  •3?  1500 -  'A  1000-  500-  0 2000 - Barkley Canyon Shelf  Barkley Canyon Beyond Shelf  "V:  1500-  .«»•  s  1000 -  c  _ o«>wE n  500 -  0 -  t o  2 0 0 0 - Estevan Point Shelf  51  1500-  o c  1000 -  =  a.  Estevan Point Beyond Shelf  500 -  0 2000  1500  ] — ' —  1  —  1  — I  '  I  B r o o k s Peninsula Shelf  \ — 00:00  1  — i — 06:00  1  — i — 12:00  1  — r 18:00  H  1000 H 500  H  o-l 00:00  06:00  T  12:00  18:00 _  ...  Time of Day (h) Figure G.1 Incident surface irradiance for primary productivity measurements during April 1997 off the west coast of Vancouver Island. Incubation period was 24 hours. Primary productivity was not measured at the beyond shelf station of the Brooks Peninsula transect. See Figure 2.1 for location of transects.  165  2000-  La Perouse B a n k Shelf  La Perouse Bank Beyond Shelf *5| \  56%  50%  1500K:%  r  1000 -  9  1 $  500 0• 2000  1—'—i—i—r  ;  A\  i  T  Barkley Canyon Shelf  8~ S '* CS  ^  E  t  o  8«5 Sin  Estevan Point Beyond Shelf  co a . ** — t= o  5 c  ^pl  3  00:00  06:00  12:00  18:00  2000 H  B r o o k s Peninsula* Beyond Shelf  61%  1500  1000 - \  500  H  0-1 00:00  "i— —i— —r 1  06:00  1  12:00  18:00  Time of Day (h) Figure G.2 Incident surface irradiance during primary productivity measurements in July 1997 off the west coast of Vancouver Island. The shaded region demarcates the incubation period. Primary productivity was not measured at the shelf station of the Brooks Peninsula transect or the beyond shelf station of the Barkley Canyon transect Number in the right hand corner of each plot is the percentage that the incubation period represented of the total daily irradiance. See Figure 2.1 for location of transects.  166  1000 H 800  H  La Perouse Bank , Shelf  64%  200 H  04 1000 -  Barkley C a n y o n . Shelf  73%  Barkley Canyon Beyond Shelf  800600 0)  o c  ra ^ TS tfl S - E  s g  i f  Si  400200 01000 •  Estevan Point Shelf  89%  800 -  T3 E 600-  ••  400 200 0 1000  B r o o k s Peninsula Shelf  72%  800 600 400 200 O - lT 00:00  T  06:00  T  12:00  T  18:00  nr»:r»o  06:00  12:00  18:00  Time of Day (h) Figure G.3 Incident surface irradiance during primary productivity measurements in October 1997 off the west coast of Vancouver Island. The shaded region demarcates the incubation period. Primary productivity was not measured in the beyond shelf region of the Estevan Point and La Perouse Bank transects. Number in right hand corner of each plot is the percentage that the incubation period represented of the total daily irradiance. Note scale change relative to Figure G.1 and G.2. See Figure 2.1 for location of transects.  167  1000-  La Perouse B a n k ^ ^ * Shelf '  77%  La Perouse BankiV* Beyond Shelf  800600-j  •J  400 -  k  2000 -  I  1000 - \ Barkley Canyon I ^Sf Shelf ' 800 H • t  79%  X  i  • • •• ••• • ••  •i  49%  • ••  *  Barkley Canyon Beyond Shelf  ^!t*  ••  78%  • %  600  •  • r 400 -  • •*  200-j 0 1000 -  I Estevan Point Shelf '  800 -  '  I  T 72%  '  • ••  •  *  Estevan Point Beyond Shelf  72%  • t  600 -  t  400 200 0 1000 _ B k Shelf 800-j r o o  :  T s  T  T  Peninsula | •  85%  T  B r o o k s Peninsula Beyond Shelf I •  It  600-j  J  400 200 0 -  1 00:00  80%  it 1  1 06:00  1  1 12:00  1  1 18:00  1  1 00:00  1  I 06:00  1  I ' I 12:00 18:00  Time of Day (h) Figure G.4 Incident surface irradiance during primary productivity measurements in May 1998 off the west coast of Vancouver Island. The shaded region demarcates the incubation period. Number in the right hand corner of each plot is the percentage that the incubation period represented of the total daily irradiance. See Figure 2.1 for location of transects.  168  1500  H  La Perouse Bank Beyond Shelf /#  La Perouse Bank Shelf  I  *  ••  1000 H  :# •*  •**•  500  •  Jt £  TBarkley 1500  51%  H  -i—r Canyon  " Barkley Canyon - Beyond Shelf  Shelf  35%  1000 H  y  4 -  500  i i " • v . ' •"  o-l Q. E  -  Estevan Point - Shelf 1500 -  47%  * 71%  Estevan Point  Beyond Shelf  1000i  5000 -  1500  H  _  I  • I —r Brooks Peninsula  i T  1  88%  Shelf  At 64%  B r o o k s Peninsula Beyond Shelf  t  •  1000 H i  #  500  o-l 00:00  r  06:00  12:00  18:00  00:00  J  06:00  r* •f . ;  12:00  !  • • • •  18:00  Time of the Day (h) Figure C.5 Incident surface irradiance during primary productivity measurements during July 1998 off the west coast of Vancouver Island. The shaded region demarcates the incubation period. Number in the right hand corner of each plot is the percentage that the light during the incubation period represented of the total daily irradiance. See Figure 2.1 for location of transects.  169  APPENDIX H  V E R T I C A L P R O F I L E S O F T E M P E R A T U R E , SALINITY A N D C T  T  Vertical profiles of temperature, salinity, and cr for the upper 200 m at a shelf and a beyond t  shelf station of each transect of La Perouse Bank, Barkley Canyon, Estevan Point and Brooks Peninsula are presented for April 1997 (Figure H.l), July 1997 (Figure H.2), October 1997 (Figure H.3), May 1998 (Figure H.4), July 1998 (Figure H.5) and October 1998 (Figure H.6). The cruise dates when these profiles were measured are specified in Table A. 1. For the deep stations, data set only shown for 0-200 m.  170  — • — Density 22  24 ,  l  26 ,  l  2822 i  I  24  1  ,  1  26 ,  1  ,  2822  24  1  I  26 ,  2822  1  ,  I  24  I  i  I  26 i  I  28 i  Temperature (°C) 4  30  8  32  12  34  4  30  8  32  12  34  4  3 0  8  32  12  4  8  34  Salinity Figure H.1 Vertical profiles of density, salinity and temperature in April 1997 at a shelf and beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP) and Brooks Peninsula (BP) off the west coast of Vancouver Island. Data for deep stations only shown for 0 - 200 m depth. See Figure 1.1 for location of transects. Data not available for beyond shelf station on the Brooks Peninsula transect. B indicates beyond shelf region.  12  I  — • — Density  Salinity Figure H.2 Vertical profiles of density, salinity and temperature in July 1997 at a shelf and beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP) and Brooks Peninsula (BP) off the west coast of Vancouver Island. Data set for deep stations only shown for 0-200 m depth. Data set not available for beyond shelf station on the Barkley C a n y o n transect. See Figure 1.1 for location of transects. B indicates beyond shelf region.  2 D Graph 1  2 D Graph 1  — • — Density  Salinity Figure H.3 Vertical profiles of density, salinity and temperature in October 1997 at a shelf and a beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Esteven Point (EP) and Brooks Peninsula (BP) off the west coast of Vancouver Island. Data for deep stations only shown for 0-200 m depth. See Figure 1.1 for location of transects. Data not available for the beyond shelf station of the La Perouse Bank transect. B indicates beyond shelf region.  173  — • — Density  Salinity Figure H.4 Vertical profiles of density, salinity and temperature in May 1998 at a shelf and a beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP) and Brooks Peninsula (BP) off the west coast of Vancouver Island. Data for deep stations only shown for 0-200 m. Beyond shelf stations for the Estevan Point and Brooks Peninsula transect only sampled to 100 m depth. See Figure 1.1 for location of transects. B indicates beyond shelf region.  — • — Density  Salinity Figure H.5 Vertical profiles of density, salinity and temperature in July 1998 at a shelf and beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP), and Brook Peninsula (BP) off the west coast of Vancouver Island. Data for deep stations shown for 0-200 m depth. The beyond shelf station of La Perouse Bank was only sampled to 100 m depth. See Figure 1.1 for location of transects. B indicates beyond shelf region.  Density 22 i  24 .  i  26 .  i  .  2822 I  L__i  24 I  26 i  I  2822 i  I  I  24 1  1  26 1  1  2822 1  1  I  24 1  1  26 1  1  28 1  1  Salinity Figure H.6 Vertical profiles of density, salinity and temperature in October 1998 at a shelf and a beyond shelf station along La Perouse Bank (LPB), Barkley Canyon (BC), Estevan Point (EP) and Brooks Peninsula (BP) on the west coast of Vancouver Island. Data set for deep stations shown for 0-200 m depth. See Figure 1.1 for location of transects. B indicates beyond shelf region.  Silicic Acid (uM) 0  20 40  1 • I • I  0  20 40  I • I • I  0 I  20 40 i  I i I  o  -•25  20 40  I . I . I  0  0.0  1  0  20 40 I . I . I  Nitrate (uM) o  25  0  • • • I • .1  I  20 40  I . I . I  25 I I I I  I  0 I  J-  BP2MBP3 I • I  1.0  2.0 0.0  1.0  2.0 0.0  -A-  1.0  2.0 0.0  1.0  2.00.0 1.0  2.0 0.0  1.0 2.0  Phosphate (uM)  Figure 1.1 Vertical profiles of nitrate, phosphate and silicic acid (pM) in April 1997 at all stations along transects of La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Station name is located at bottom of each graph.  177  - » - Silicic Acid (pM) 0  20 40  I.I.I. 20  20 40  I . I . I .  40 0  20  it •  0  1  20  0  20 40  I . I . I .  40 0  M  20  0  20 40  0  20 40  I.I.I. I.I.I. Nitrate (pM)  40 0  20  40 0  20  40  0  20  i i i 0  40  ir  20  40  m  A i*  '  7 *  40  ill  60  \ B 2 . . B8  80 0  0  '.'V'.'i  it  4E  VI'i  •  20 40  A •  60  A  80  J I I I I  C2 C4 I I I' I ' ' ''I  1 1  I  I I I I  I  I I"  II  I I I  I  I I  0 20 40 60 80 • 1  0  1  1  1  1 •  Am  20  \  40 60 BP4  80 0.0  2.5  0.0  2.5  0.0  2.5  0.0  f>> BPS* 2.5  0.0  2.5  0.0  2.5  —A- Phosphate [\M) Figure 1.2 Vertical profiles of nitrate, phosphate, and silicic acid (uM) in July 1997 at all stations along transects of La Perouse Bank (Line B), Barkley Canyon, (Line C), Estevan Point (Line G), and Brooks Peninsula (Line BP) on the west coast of Vancouver Island. Station name is located at the bottom of ea^h graph.  178  Silicic Acid (uM) 0  20  40  I . I . I  20  0  40  20  0  40  0  20  40  0  20  40  I . I .  Nitrate (uM) 0 25 1 , . • • 1i  VA \J  0  25  1 • • • • i •  t  B9  '.V.'.'I : A  I*  25  1 1 1• • 1 •  \\  • 1  A \  o 0  B14  25 • • I  m  • i  1 5  0.0  2.5  o.O  2.5  0.0  C8  2.5  0.0  2.5  0.0  2.5  0.0  2.5  —A- Phosphate (uM) Figure 1.3 Vertical profiles of nitrate, phosphate and silicic acid (pM) in October 1997 at all stations along transects of La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) on the west coast of Vancouver Island. Station name is located at the bottom of each graph.  179  - » - Silicic Acid (pM) 0  20 40  I . I . I  0.0  2.5  0  20 40  I . I . I  0.0  2.5  0  20 40  I.I.I  0.0  2.5  0  20 40  I . I . I  00  2.5  0  20 40  I • I • I  0.0  2.5  0  20 40  I • I i I  0.0  2.5  0  20  40  ' • I • '  0.0  2.5  —A— Phosphate (\M) Figure 1.4 Vertical profiles of nitrate, phosphate and silicic acid (uM) in May 1998 at all stations along transects of Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Station name is located at the bottom of each graph. No are data available for the La Perouse Bank transect.  180  - » - Silicic Acid (\M) 0  20 40  0  20 40  0  20 40  0  20 40  0  20 40  0  20 40  0  20  —A— Phosphate (pM) Figure 1.5 Vertical profiles of nitrate, phosphate and silicic acid (uM) in July 1998 at all stations along transects of Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Station name is located at the bottom of each graph. No data are available for the La Perouse Bank transect.  181  40  * 0  20 40  I.I.I  0  20 40  l . l . l  0  20 40  Silicic Acid (pM) 0  20 40  I.I.I  L  0  20 40  0  20 40  0  20  L_i_L  - e - Nitrate (pM) 25 _LL.  0 25 I• • • • I •  0 J  -A- Phosphate (uM) Figure 1.6 Vertical profiles of nitrate, phosphate and silicic acid (uM) in October 1998 at all stations along transects of Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) on the west coast of Vancouver Island. Station name is located at the bottom of each graph. No data are available for the La Perouse Bank transect  182  40  -3  -2  ~  To > 1  £  +-» CH O S3  3. ©  o -  C  .2 3 +-• o  a  -7!  S  V  23  e8  fl  -H  S  4>  CO  ^  co  ^H  + i CN ON fl  +1 N£)  m  r-^  oo  +j  IL fl  m II fl  +i ON O  fl  ^f.  +1  lO  ON CN  II  (N II A  oo  +1  A  NO  1  5  v o  A  ON  ON  ON  WO  ^ 1 ^ fl  NO +1 CN  CO CN  fl ^  NO \J~i  S3  o «i3 g 3 ~ 2>  5 8  J-  3.  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II +1 C ON ^ NO  7  uo  OO  ^  « •c _ o  CO  oo oo oo CO 0 0  uo  c  oo  CO +1 NO  NO C N OO +1 II ^O  CO  oCO  CN  oo  fl  +1 ON  II  fl  +1 CO  II  CO  +1  fl  CN  O  OO  ON CO  CO  CO  <3  S  > 2°  uo ^o'  NO  4)  13  o w  CO  CN NO C N  =»• 6  H-*  O  71  oo  CO  *-H  3  oo CN  o  "a*  CO  h  +i2oo-i fl  "  a)  C „ CO JO  '-  ON  •sf  CO  IL  H *  +1  CN  NO CN  X  CN  oo  ©  •^31  C->  + l  CN CN  c  § '3  c o  CN  Z.  C id C  i—i  ^  CO  OH . O  >>  h  ^  fT} c o  i/o  s  E  1 ° I '3oD  H_,  fl  +i  ^ 1  + 1 ?  .8 ±21, n=4  i-i - f l on  +1  co  CN  fl  ° .2 B s > 2 •S £ j .a a fl  ~ H  CN  ,7 ±21,  O  +1 h  00  * B  c -fl £  •a  B  O >» PQ  fl , rS ^ fl  J3 t/5  o  g  +1  CN  CN  .8 ±21,  0  "a*  Ch >5.0  fl «4H 3. O  o  fl  • ' „II £  *5JD  .3  03  II  55  uirl  *  A  IK  >  +1 oo  +1 fl  so V  NO  CN NO  fl  CO  CN  CN CO  CN CO  O  ON ON  ON ON  "3  °  ON ON  ^  CN  o  h  fl°° II  •71  oo  fl  oo II C  co  CN  J" <N CO CN  O0  oo  OO  ^—i  '—i  % £  « o  NO  2^ ON  t S  C  + 1' i IIi NO A  ON ON  ON ON  ^ +1  O 00  q co  CO  <u  CO •3  B  e  kH  O  183  APPENDIX K  V E R T I C A L PROFILES OF SIZE-FRACTIONATED C H L O R O P H Y L L  Chlorophyll a (mg m" ) 0  2 2  1  I  44 6 6 8 8 I  I  10 10  I  0  I  -^afc'fj  2 J  4  6  i  8  10  0  2  4  6  8  10  0  2  4 6 8 J I I  i  10 L  3  • b  fflP * a® T/ I  m C4 Shelf  C1 Shelf  I  I  I  G3 Shelf I  BP2 Shelf  -in  J  I  L  cm a® al  B16 beyond shelf  p  -•-  C9 beyond shelf  < 5 um April  I I  G5 beyond shelf J ^  1  BP7 beyond shelf  - B - > 5 u m April  ~o~ < 5 um July  - - • - > 5 pm July - © - < 5 um October - E l - > 5 pm October  Figure K.1 Vertical profiles of size-fractionated chlorophyll a for April, July, and October 1997 at shelf and beyond shelf stations along transects of La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Data are not available for the > 5.0 um fraction for station C1, BP2, B16 and BP7. Samples taken down to 1%sutface irradiation. See Figure 2.1 for location of stations  184  Chlorophyll a (mg m" ) 3  < 5 pm May - « - > 5.0 pm May • o - < 5 p m J u l y • - 0 - - > 5.0 pm July - ® - < 5 pm Oct. - E - > 5.0 pm Oct.  Figure K.2 Vertical profiles of size-fractionated chlorophyll a May, July, and October 1998 at shelf and beyond shelf stations along transects of La Perouse Bank (Line B), Barkley Canyon (Line C), Estevan Point (Line G) and Brooks Peninsula (Line BP) off the west coast of Vancouver Island. Samples taken down to 1 % light depth. See Figure 2.1 for location of stations.  185  120 100 -  Perouse Bank \ LaBeyond Shelf  La Perouse Bank Shelf  80 no data are available  no data are available  60 40 20 0120 -  T ~ I — i — i — i — r  23%  Barkley Canyon Beyond Shelf  8%  97%  Estevan Point Beyond Shelf  55%  Barkley Canyon Shelf  100 80 • 60 40 20 0  r  120 -  r  Estevan Point Shelf  100 • 80  I  I  60 • 40 20 0  1  r—T—T—7  r  120 100 •  1  l  i  r  i  Brooks Peninsula Beyond Sheff  Brooks Peninsula Shelf  no data are available  no data are available  80 60 40 20 0  i  i  100  ,  55  i  „  30  i  „ i  10  ,  3  i  „  1  i i „„ i  100  i  55  „  30  i  i  10  „  T "  3  1  -  t  Percent of Surface Irradiance  E23  < 5 um fraction S  > 5 um fraction  Figure L.1 Relative contribution of <5 um size fraction and >5 um size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during April 1997. The relative contribution of > 5 pm fraction to depth integrated chlorophyll is given in top right hand corner of each plot  186  Estevan Point Shelf  100  29% -A'-.'il  80 60 40  -j  20 -  120 -| Brooks Peninsula 100 -  Shelf  25%  UP '  80 ~_  Brooks Peninsula Beyond Shelf  27%  60 ~_ 40 20 100  55  30  E3  3 1 100 55 30 10 Percent of Surface Irradiance  10  <5 um fraction  >5 um fraction  Figure L.2 Relative contribution of <5 um size fraction and >5 pm size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during Oct 1997. Relative contribution of >5 pm fraction of depth integrated chlorophyll in right hand corner of each graph.  187  1  1 0 0 ' 55 ' 30 ' 10  1  10  1  3  1  1 '  Percent of Surface Irradiance E3  < 5 um fraction  03  > 5.0 um fraction  Figure L.3 Relative contribution of <5 um size fraction and >5 um size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during May 1998. Relative contribution of >5 um fraction depth integrated chlorophyll in right hand corner of each graph.  188  Q. Q  2  o c u  0)  100  55 ' 30  10 ' 3  '  1  100  55 ' 30  10  Percent of Surface Irradiance E3  < 5 um fraction  HH  > 5 u m fraction  Figure L.4 Relative contribution of <5 um size fraction and >5 um size fraction to chlorophyll at each light depth for the shelf and the beyond shelf station of each transect during Oct 1998. Relative contribution of >5 pm fraction depth integrated chlorophyll in right hand corner of each graph. ns=no sample is available.  189  > o 3  •a o  o. E S o  c u  a.  Percent of Surface Irradiance  E3  < 5 um fraction  S  > 5 um fraction  Figure L.5 Relative contribution of <5 um size fraction and >5 um size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during April 1997. Relative contribution of >5 um fraction to depth integrated primary productivity is in the right hand vomer of each graph.  190  120  > o 3  •o o  l_ Q.  re  E •c a $ o  100 -  74%  Estevan Point Shelf  80 60 -  o  ot  a.  40 20  Brooks Peninsula Beyond Shelf  100 ' 55 ' 30 ' 10  3  1  100  55  30  47%  I  10  Percent of Surface Irradiance E3 < 5 um fraction •  > 5.0 um fraction  Figure L.6 Relative contribution of < 5.0 u m size fraction and > 5.0 u m size fraction to primary productivity ateach light depth for the shelf and the beyond shelf station of each transect during Oct 1997. Relative contribution of > 5.0 u m fraction to depth integrated primary productivity in right hand corner of each graph.  191  Percent o f Surface Irradiance E3  < 5 um fraction  E3  > 5.0 um fraction  Figure L.7 Relative contribution of < 5 um size fraction and > 5 um size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during May 1998. Relative contribution of depth integrated primary productivity in right hand corner o f each graph.  192  120  ioo H  > o 3 T3  O i_ Q.  co E  'C Q.  O  c (J) Q.  Brooks Peninsula Beyond Shelf  100  55  30  10  3  1  100  55  30  10  3  Percent of Surface Irradiance E3  < 5 um fraction  H  > 5.0 um fraction  Figure L8 Relative contribution of < 5 um size fraction and > 5 um size fraction to primary productivity at each light depth for the shelf and the beyond shelf station of each transect during Oct 1998. Relative contribution of > 5 um traction to depth integrated primary productivity in right hand corner of each graph.  193  Table M . l Total primary productivitiy ±1 S.D. (surface to 1% light depth) for the west coast of Vancouver Island (WCVI) and the shelf and beyond shelf region in 1997 and 1998. Means for 1997, 1998 and for all cruises are given along with the minimum and maximum for all cruises. Numbers in brackets are the number of stations sampled. Total Primary Productivity (gCm-'d ) Beyond Shelf WCVI Shelf 1  Date 1997 April July Oct. 1997 cruises 1998 May July Oct. 1998 cruises  5.2 ± 2 . 4 (8) 3.5 ±2.8 (6) 3.2 ±3.4 (5) 4.3 ±1.1 (19) 2.8 ±2.5 (8) 6.7 ±8.6 (8) 0.6 ±0.4 (8)  6.9 5.3 3.1 5.1  ±4.9 (4) ±2.9 (3) ±4.0(4) ±3.4(11)  3.4 ±2.2 (4) 1.7 ±1.2 (3) 3.3 (1) 2.8 ±1.8 (8)  3.4 ±3.0 (24)  3.9 ±2.4 (4) 11.7 ±10.2 (4) 1.1 ± 0 . 4 ( 4 ) 5.2 ±7.2 (12)  1.7 ±2.3 (4) 1.8 ±1.3 (4) 0.5 ±0.2 (4) 1.3 ± 1 . 7 ( 1 2 )  All cruises  4.0 ± 0.8 (43)  5.1 ± 2 . 7 ( 4 4 )  2.0 ± 0 . 1 (44)  Minimum Maximum  0.6 6.7  0.3 26.1  0.3 6.3  a PH  I  o  O  >  +1  O  00  fl  C N  \° o  CO  x  00 C O  CO  ON  is 13 -e PH  O N  1  02  fl  T3  e  o >> v  O N  M  V  Ii  -I  PM « PM V3  ;fl c  °:  +L  co  IL *  o  21  + 1  m  1  o  O N  C N  CO  + 1  + 1  CO  O  C N  o co  II  N O  + 1  N O  C N N °  C N  +1  O N  CO  h  1—1  * O O  + 1  4-1 NO  C  fl  0  o  NO  fl  o fl  A?  O N NO  + 1  ^-1  II C  n  o ^j-  in +1  o  + 1  + 1  + 1  CO  C N  N O  in  o  x  C N  CO  o fl  N O  00  + 1  CO  * :  o  C N  ^.  so  O N  ° :  O N ^  ^1-  ^M  r-H O N  ^M  N O  CO  +1  II  ^  o fl  ^  AT co  O TT  0  1-1  C N  A  O  N O  o  PH PH  § .2  C N  2 ^  ILfl +1  00  C N  O O  O  CO  ^M  CO  "sT  _  O N  NO  60  e  s I  _o  5' 3D Pi  _  CO  + 1  MM  00  g .s  w  CO  P*  LT)  * 00  ,7±0.  fl o o  + l  DC 0)  c  I  o  00 fl  A  o  O  O  x!  2  PH  q A  "3 X!  o  r-  IL  +L  (N co +L  «  N O  10  C N  IL  3  + 1  O  II C  C N  CO  C N  + 1  + 1  O N  C N  CO  CO  ON  CO  ON" r t  O  + 1  ^3-  II o o Hi r- e ^ o + 1  fl  m o\ +| 00  C N  ^  C N  + 1  O N  O  C O  O N  C N  + 1  To  3  PM  V  £fl »-  o  O  o „ Q* c3  >  3 Ef  "  S  3  PM PM  a I-  N J  ~  >  rl  a "oa| > <L>  O  91  < >  CO  Ii  O  A  CU  gg 2 >>  91  a "5 T3  I 2  IT)  fl  fl  + 1  H  II C  + 1 C N  PM  e q A  O  o  O  <T> 00 +L  IL  in  fl  q c j j 00 in co  O  PM PM  Ii  »  q  c> 00  N O  fl^ flL in fl  o + 1  +1  ctf  N O  ' I ST  C N  +  N O  ON  in C N  w  +1  ^10 + 1  O  c  o  o  +L  IL  <N 00  C N N O  A  C  C N  + 1  CO  + 1  q co  co  II fl  NO  I  1  h  fl ^  00  + 1  ~  2  x°  O +1  N ° Q N  O NO  C N  c> co  C N  e vo ?i  00 II  r>; 00 +L  IL  o  O  o + 1  in  m  in  C N  + 1  00  fl c r-  in  C N  O N  II  + 1  N O  + 1  C N  £  q  00  +L  IL  C N  +  ' PH  N O  r- fl o  O O  A  o + 1  co O  O O  C N  C N  CO O N O N  O N O N  '3 S-H  O  O  O  r~ O N O N  00  NO  N O  + 1  II 00 C  o  co <U  co <U O N  O N O' N  o  o  T—1  O N  H  fl  00  00 0  o & fl  £  co in  V  il  r~~ r-fl o  + 1  N O  00  IT)  C N  AT  O O  N O  T-H  CO  C N  O N  CS  o tS ^ so  N  N O  o  00  U  $  NO  NO  CO  N O  2 ^ o  C  ° S a o  q  5  00 O N O N  1—1  00  00  O N  O N  O N  O N  >.  3  o O  £  +1 ^1  CO  co  g  co  O  00 O N O N  o  O N  fl o  c o  il  > 195  i  to  

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