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Influence of hydrographic properties in Saanich Inlet on ontogenetic migration and retainment of the… French, Shirley E. 1988

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INFLUENCE OF HYDROGRAPHIC PROPERTIES IN SAANICH ON ONTOGENETIC MIGRATION AND RETAINMENT OF THE CALANOID COPEPOD NEOCALANUS PLUMCHRUS by SHIRLEY E. FRENCH B.Sc,  Brandon U n i v e r s i t y , 1984  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Department of Zoology) We accept t h i s t h e s i s as conforming t o the required standard  THE UNIVERSITY OF BRITISH COLUMBIA January 1988 ©  S h i r l e y E. French, 1988  In presenting  this thesis in partial fulfilment  of the  requirements for an advanced  degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department  or  by  his  or  her  representatives.  It  is  understood  that  copying  or  publication of this thesis for financial gain shall not be allowed without my written permission.  Department The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 .  DE-6(3/81)  i i  ABSTRACT During Saanich fjord not  Inlet  while  known b u t by  suspected Vertical Inlet 1985  the  an  to  annual  the  the  Strait  may a r i s e  of  was  Georgia  January-March  1986  to  population  to  spring  overwintering be  correlated  renewal. of  The  their  the  low  0.10-0.30 the  were  the  Fuca  ) that  exposed  to  tidal  low  is  was  abundance. from  Saanich  1985  in  (September  1986 t h e  population  transport  of  during  of  is the  copepods  to  their  few  inlet  years  adults in  contribution Decline 1986  early  in  appears  deep  water stages  influenced inlet  by  (i.e.  occurred  above  displacement  Subsequently, the  from  some  Inlet  August),  zone.  out  So  1985 a n d  bottom  due  in  an e x t e n s i v e  renewal,  oxygen  and  negligible.  of  to  i n the  introduced  potential  N. plumchrus (June  is  in Saanich  i n September  During the  the  water,  inlet.  considered  m i n i m u m (75 m) p o s s i b l y of  variation  conditions  Strait,  their  occurrence  concentrations 1  avoidance  increased  was  migration  mL L " ) .  oxygen  their  N. plumchrus  d i s t r i b u t i o n of  deep water oxygen  3  population with  this  anoxic  population  stations  (<0.20 nr  of  collected  w i t h i n the  three  the  from  low.  a n d J u a n de  the  of  absent  i n September  summer,  from r e p r o d u c t i o n  cause  in  oxygenated  samples  comparably and  are  overwinters  d i s t r i b u t i o n and  sparse;  at  the  dense,  haul  were c o l l e c t e d  the  development  their  was  they  The  overwintering  1986)  spring  plumchrus  years  1983).  a l . ,  and h o r i z o n t a l  March  In  i n other  influence  indicated to  Neocalanus  i n t r u s i o n of  N . plumchrus  the  years  ( H a r r i s o n et  followed  of  some  or  they  and perhaps  to  predation.  On t w o  occasions  in which N. plumchrus  was  present  during  the  winter  dense  i n Saanich Inlet  water  mixing  intruded,  disrupting  water  in  t h e w a t e r , below  left  the i n l e t  reproduces  sill  i n Sechelt  shallow  inlet.  sill  Sechelt  Inlet  layer  during the  though a g r e a t e r could  volume  have  remained  depth. successfully  Inlet  which  (15 m) t h a t  The f i f t h  Even  some o f t h e c o p e p o d s  Neocalanus plumchrus  in  t h e copepod  o f t h e two water m a s s e s .  of  very  (1969 & 1974), a h i g h volume o f  o v e r w i n t e r s and  i s w e l l - o x y g e n a t e d but has a  'locks'  the copepods  copepodite stages also  than i n S a a n i c h I n l e t ,  i n t o the  occupy deeper  even  depths  though the bottom  depths a r e comparable. In the  low oxygen  period  Saanich results. their  tolerance  of c a p t i v i t y ,  Inlet  versus  and the r e g i o n  the S t r a i t  could  the  loss  that  1  level  (12% m o r t a l i t y ) .  a massive d i e o f f could  o f N. p l u m c h r u s  such as  of o r i g i n ( i . e . variable  tolerable  n o t be d e t e r m i n e d , N. p l u m c h r u s  l e v e l s a s low a s 0.56 mL L ~ indicate  factors  of Georgia) caused  A l t h o u g h t h e minimum oxygen  migration  samples  e x p e r i m e n t s many  from S a a n i c h I n l e t .  during tolerated  Sediment  trap  not account f o r  iv  TABLE OF  CONTENTS  ABSTRACT  i i  L I S T OF TABLES  vi  L I S T OF FIGURES  ix  ACKNOWLEDGEMENTS  .  xi i i  1 . INTRODUCTION 1.1.  Life  1  History  of Neocalanus plumchrus  4  2 . THE STUDY AREAS  7  2.1 P h y s i c a l  7  Characteristics  2.1.1 S a a n i c h I n l e t  7  2.1.2. S t r a i t  of Georgia  10  2.1.3 S e c h e l t  Inlet  12  Selection  13  2.2.  Station  3. MATERIALS AND METHODS  16  3.1.  Hydrographic Data  16  3.2.  Zooplankton Sampling  17  3.3.  Treatment  20  3.4.  C o l l e c t i o n and M a i n t e n a n c e  3.5.  Experimental Procedures  23  3.6.  Sediment  25  3.7.  Statistical  o f Net H a u l Samples of Live  T r a p Samples Methods  Analysis  ....... 21  26  4. RESULTS 4.1.  Copepods  28 of the P h y s i c a l  Data  28  4.1.1. S a a n i c h I n l e t  28  4.1.2. S t a t i o n  G1748 & G1545  32  4.1.3. S e c h e l t  Inlet  34  4.2.  Analysis  of N e t , H a u l  Samples  35  of G e o r g i a V e r t i c a l Hauls  36  4.2.1.  Strait  4.2.2.  Satellite  4.2.3.  Saanich  Inlet  39  4.2.4.  Sechelt  Inlet  45  4.3. Analysis  of  Channel  38  Experimental Data  4.3.1.  Observations  4.3.2.  Oxygen T o l e r a n c e T e s t s  4.4.  of  46  F e d and S t a r v e d Copepods  47  S e d i m e n t T r a p Samples  49  5. DISCUSSION 5.1.  Physical  population 5.2. in 6.  52 i n f l u e n c e s on t h e N e o c a l a n u s in Saanich  A l t e r n a t i v e Causes Saanich  CONCLUSIONS  References Appendix A  46  plumchrus  Inlet for  t h e D e c l i n e o f N.  52 plumchrus 63 71 169 1 7  4  vi  L I S T OF  Table  1. S t a t i o n  Table  2. H y d r o g r a p h i c  TABLES  c o o r d i n a t e s and maximum d e p t h s and b i o l o g i c a l  stations  75  sampled i n  1985 Table  76  3. H y d r o g r a p h i c  and b i o l o g i c a l  stations  sampled i n  1986 Table  77  4. F l o w m e t e r measurements  w i t h and w i t h o u t  t h e mSCOR  net  78  Table  5. Low  Table  6. D i s s o l v e d oxygen c o n c e n t r a t i o n s a t Saa9  i n 1985.  . 81  Table  7. D i s s o l v e d oxygen c o n c e n t r a t i o n s a t Saa9  i n 1986.  . 81  Table  8. H y d r o g r a p h i c  station Table  oxygen  t o l e r a n c e e x p e r i m e n t s on CVs  data  collected  A u g u s t 5,  79  1986, a t  Saa3  81  9. D i s s o l v e d oxygen c o n c e n t r a t i o n s a t Saa0.8  i n 1985. 82  Table  10. D i s s o l v e d oxygen c o n c e n t r a t i o n s a t Saa0.8  in  1986 Table  82  11. D i s s o l v e d oxygen c o n c e n t r a t i o n s a t G1545 and  G1748 Table  83  12. D i s s o l v e d oxygen c o n c e n t r a t i o n s a t G1545 i n  1986. 83  Table  13. V e r t i c a l  Table  14. V a r i a t i o n  plankton between  hauls c o l l e c t e d replicate  i n 1985  vertical  84  haul  samples  85  Table  15. V e r t i c a l  plankton  hauls c o l l e c t e d  Table  16. H o r i z o n t a l p l a n k t o n  i n 1986  hauls c o l l e c t e d  at  station  86  vi i  Saa3 i n Table in Table  91  18. A N O V A - S t r a i t  of Table  1985 and  fall  1985 v e r s u s  Inlet  counts  f o r Saa3,  May,  July,  1986  96  Inlet  counts  from A p r i l  through 98  22. ANOVA-Saanich through August  23. ANOVA-Saanich t h r o u g h August  24. A N O V A - S a a n i c h t h r o u g h August  25. ANOVA-Saanich Georgia counts,  of  Inlet  for  Saa9 and  Saa0.8,  1986  100  Inlet  counts  for  Saa3 and  Saa0.8,  1986  102  Inlet  counts  f o r Saa9 and  Saa3,  1986  104  Inlet  counts at  Saa9 v e r s u s  Strait 105  Inlet  Georgia counts,  27. ANOVA-Saanich taken  counts  1986  26. ANOVA-Saanich  replicates Table  counts,  1986  Strait Table  versus  95  21. ANOVA-Saanich  April Table  of G e o r g i a  20. ANOVA-Saanich  April Table  fall  1986  April Table  counts,  93  19. ANOVA-Strait  August Table  of G e o r g i a  and summer  and A u g u s t , Table  collected  1986  fall Table  90  17. H o r i z o n t a l and o b l i q u e p l a n k t o n h a u l s  spring Table  1985.  counts at  versus  1986  Inlet  in J u l y  Saa0.8  counts  107 at  Saa9,  horizontal  1986  109  2 8 . Copepod f a e c a l p e l l e t p r o d u c t i o n o v e r a  12 h o u r  period. Table  2 9 . Buoyancy o f  experiments Table  110 CVs  following  t h e i r death  in  8 and 9  3 0 . C a l c u l a t e d volume o f  110 water  f i l t e r e d through  the  viii  mSCOR n e t Table  31. C o n c e n t r a t i o n s  oxygen Table  Saanich  Inlet,  dissolved  1969  11 1  Inlet  i n 1969  112  33. D i s s o l v e d oxygen c o n c e n t r a t i o n s measured i n  Saanich Table  from c e n t r a l  o f N. p l u m c h r u s and  32. D i s s o l v e d oxygen c o n c e n t r a t i o n s measured i n  Saanich Table  111  Inlet  i n 1974  34. G e l a t i n o u s z o o p l a n k t o n  1985  113 collected  i n September 114  ix  L I S T OF FIGURES  Figure  1. The s t u d y  Figure  2. L i f e  area  history  115  stages  characteristic  of Neocalanus  plumchrus  116  Figure  3. S e c h e l t  Inlet  and S t r a i t  of G e o r g i a  stations.  ...117  Figure  4. S a a n i c h  Inlet  stations  ...118  Figure  5. The m o d i f i e d  SCOR n e t  119  Figure  6. B a t h y k y m o g r a p h  Figure  7. A p p a r a t u s f o r l o w - o x y g e n t o l e r a n c e e x p e r i m e n t s . .120  Figure  8. A r e g r e s s i o n o f t h e l o g mean v e r s u s  results.  120  the l o g  variance  121  Figure  9. S a l i n i t y d i s t r i b u t i o n  for station  Figure  10. T e m p e r a t u r e d i s t r i b u t i o n  Saa9  for station  i n 1985. .122 Saa9 i n  1985  123  Figure  11. D e n s i t y  Figure  12. D i s s o l v e d  in  distribution  for station  oxygen d i s t r i b u t i o n  Saa9  i n 1985. .124  for station  Saa9  1985  125  Figure  13. S a l i n i t y d i s t r i b u t i o n  for station  Figure  14. T e m p e r a t u r e d i s t r i b u t i o n  Saa9  for station  i n 1986. 126  Saa9 i n  1986  127  Figure  15. D e n s i t y  Figure  16. D i s s o l v e d  in Figure  for station  oxygen d i s t r i b u t i o n  Saa9  i n 1986. .128  for station  Saa9  1986  129  17. Copepod c o n c e n t r a t i o n s  levels Figure  distribution  at station  Saa3, A u g u s t  18. Copepod c o n c e n t r a t i o n s  and d i s s o l v e d  oxygen  1985 and d i s s o l v e d  130 oxygen  X  levels Figure  at station  Saa3,  August  19. S a l i n i t y d i s t r i b u t i o n  1986  131  for station  Saa0.8 i n  1985. Figure  132  20. T e m p e r a t u r e  distribution  for station  Saa0.8 i n  1985 Figure  133 21. D e n s i t y d i s t r i b u t i o n  for station  Saa0.8 i n 1985. 134  Figure in  22. D i s s o l v e d  oxygen  distribution  for station  Saa0.8  1985  Figure  135  23. S a l i n i t y d i s t r i b u t i o n  for station  Saa0.8 i n  1 986 Figure  136 24. T e m p e r a t u r e  distribution  for station  Saa0.8 i n  1986 Figure  137 25. D e n s i t y  distribution  for station  Saa0.8 i n 1986. 138  Figure in Figure  26. D i s s o l v e d  oxygen  distribution  for station  Saa0.8  1986  139  27. S a l i n i t y d i s t r i b u t i o n  f o r s t a t i o n s G1545 a n d  G1748 i n 1985 Figure  28. T e m p e r a t u r e  140 distribution  f o r s t a t i o n s G1545 a n d  G1748 i n 1985 Figure  141  29. D e n s i t y d i s t r i b u t i o n  f o r s t a t i o n s G1545 a n d  G1748 i n 1985 Figure  142  30. D i s s o l v e d oxygen  distribution  f o r s t a t i o n s G1545  and G1748 i n 1985 Figure  143  31. S a l i n i t y d i s t r i b u t i o n  for station  G1545 i n 1986. 144  Figure  32. T e m p e r a t u r e  distribution  for station  G1545 i n  1986  145  Figure  33. D e n s i t y  Figure  34. D i s s o l v e d  in  oxygen  for station  distribution  G1545 i n 1986. 146  for station  G1545  1986  Figure  147  35. H y d r o g r a p h i c p a r a m e t e r s c o l l e c t e d  Sc1, Figure  distribution  November  at  station  1985  148  36. H y d r o g r a p h i c p a r a m e t e r s c o l l e c t e d  at  station  Sc1 , F e b r u a r y 1 986 Figure  37. H y d r o g r a p h i c p a r a m e t e r s c o l l e c t e d  Sc2, A u g u s t Figure  Figure  Figure  Saanich Figure  Inlet  Figure  .. .  concentrations  and S a t e l l i t e  collected  152 i n the  Channel  distribution  1 53 collected  in  i n 1985 a n d 1986. ..154  of Neocalanus plumchrus i n  Channel  43. V e r t i c a l  Saa3,  of Neocalanus plumchrus at  i n 1985 and 1986  copepod  42. V e r t i c a l  Satellite Figure  distribution  copepod c o n c e n t r a t i o n s  of G e o r g i a  41. T o t a l  station 151  G1545 and G1748.  40. T o t a l  at  1986  39. V e r t i c a l  Strait  station 150  38. H y d r o g r a p h i c p a r a m e t e r s c o l l e c t e d  stations  at  1986  Sc2a, August Figure  1 49  155 distribution  o f N. p l u m c h r u s a t  station  Saanich Inlet  44. V e r t i c a l  distribution  156 o f N. p l u m c h r u s a t  station  Saa0.8 Figure  157  45. V e r t i c a l  distribution  o f N. p l u m c h r u s a t  station  Saa9. Figure  158  46. Copepod c o n c e n t r a t i o n s  levels  at station  Saa0.8, June  and d i s s o l v e d 1986  oxygen 159  xii  Figure  47.  levels Figure  at  48.  Sc1, Figure  Copepod c o n c e n t r a t i o n s station  Vertical  Sechelt 49.  Sc2  and  d i s s o l v e d oxygen  1986  distribution  160  of N.  Inlet  Vertical  stations  Saa9, J u l y  and  plumchrus at  station  *  distribution  161  o f N.  plumchrus  at  Sc2a  Figure  50.  M o r t a l i t y of  Figure  51.  Percent  162 starved versus  m o r t a l i t y of CVs  f e d copepods  exposed to  163  different  oxygen c o n c e n t r a t i o n s Figure  52.  C a p t i v i t y p e r i o d and  subjected Figure  53.  164  t o low  Expected  percent  oxygen t o l e r a n c e results  of  low  m o r t a l i t y of  CVs  tests  oxygen  165 tolerance  experiments Figure  54.  prior Figure  The  166 hypothesized  t o , and  55.  station  during  Hydrographic sampled  the  distribution deep w a t e r  s t a t i o n s and  i n 1969  and  1974  o f N.  renewal  the  plumchrus i n 1985.  zooplankton  ...167  haul 168  xi i i  ACKNOWLEGDEMENTS I would the  C.S.S.  like  to extend  Vector  thanks t o the  for their  assistance  sampling.  I would a l s o l i k e  p e o p l e who  p a r t i c i p a t e d i n the  biological  data,  McLean and  Tony N o b l e who  captain  and  t o e x p r e s s my collection  and  crew  cooperation  during  appreciation of  of  to  hydrographic  those and  e s p e c i a l l y t o Dave T e s c h , Nancy B u t l e r , Hugh a s s i s t e d i n the  collection  of  zooplankton. I am  g r a t e f u l to Dr.  throughout advice,  the  c o u r s e of  financial  F.J.R.  development the  advice  Taylor  this  providing  helpful  and  and  Dr.  various  t o thank Dr.  unix  operating  enlightening problems.  culturing  W.E.  Neill  t h e s i s and  valuable dedicated  aspects Neill  were h e l p f u l i n  was  provide  of  this  his  perceptive  the  the  criticism  a v a i l a b l e on  study.  for reading  many  input  I would final  on  and  also  draft  and  criticism. Blachford  s y s t e m and  phytoplankton.  i n s t r u c t i o n s on  for his  r e s o l u t i o n of my  t o e x p r e s s my  The  'jellies'  for his  t o Rowan H a i g h  E l a i n e Simons f o r p r o v i d i n g  for  provided  offered valuable  c o n t r i b u t i o n s to the  H e a t h e r Dovey  supervision  e n c o u r a g e m e n t , and  W.E.  I would a l s o l i k e  Acreman and  He  John F u l t o n  Many t h a n k s t o A l i s t a i r the  study.  p r o j e c t and  t o d i s c u s s my  regarding  Lewis f o r h i s  thesis.  f i n a l manuscript.  occasions  like  of  this  support,  t i m e t o e d i t i n g my Dr.  A.G.  appreciation  h e l p f u l input  zooplankton  a l s o proved  computing to  on  samples a n a l y s e d  t o be  Judy  h e l p f u l to  by this  study. A very  s p e c i a l t h a n k s t o Tony N o b l e w i t h  whom I  discussed  xiv  my  work and  received valuable  input at a l l stages  of my  thesis.  1  1. INTRODUCTION The  oxygen  central  Peru  influence  to  1980;  0.1  0.2  species  mL  L"  California.  station  'P',  m;  C.B.  Miller,  pers.  In  coastal  regions  exchange of  interest  the  calanoid  west  coast  of  Hence,  mL  L"  minimum  and abundance  more  southern  Judkins,  above  1980  ).  copepod  450 m e t e r s  where  i n the Santa Barbara  latitudes  conditions and  a  (50°N) a t o c e a n (>  The  600  i s the e f f e c t Neocalanus  between  occurrence  Hoos,  of these  anoxic  of  the  1970; D e v o l ,  & Brinkhurst,  of the  side  characteristically  1968;  Burd  fjords  r e s t r i c t i n g the  the d i s t r i b u t i o n  (Fish,  1983;  in  on t h e s o u t h e a s t  Columbia,  conditions.  occur  sill  situated  British  Mills,  1984). Of  conditions  plumchrus. Upwelling  on  o f f the  t o reoxygenate the bottom  waters  l a t e A u g u s t and December o f e a c h  year.  o f t h e deep w a t e r  of t h i s  1967;  o f many  i s found at deeper depths  low o x y g e n  p r o v i d e s dense water  the e f f e c t  the  1967;  existed  1  higher  circulation  copepod  Saanich Inlet  (Longhurst,  in  a t or j u s t  z o o p l a n k t o n and b e n t h o s  particular  m  Longhurst,  the  bottom  and  a major  to the m i g r a t i o n  waters appears to i n f l u e n c e  Mackie  shown t o have  from  comm.).  Island,  anoxic  oxygen-poor  1981;  weak  200  of water. S a a n i c h I n l e t ,  develops  resident  In  t h e oxygen  Vancouver  been  Ocean  found dense a g g r e g a t i o n s of the  c o n c e n t r a t i o n s o f 0.2  a  to  californicus  Basin,  Pacific  e t a l . , 1984). Oxygen c o n c e n t r a t i o n s of  26°N;  e t a l . (1984)  have  has  a c t as a b a r r i e r  1  to  Calanus p a c i f i c u s  eastern  of z o o p l a n k t o n  d e p t h s o f 80  (15°S  Alldredge  which  the  California,  Alldredge  beyond  latitudes  oxygen  in  on t h e d i s t r i b u t i o n  Judkins, to  minimum  species,  r e n e w a l on t h e  i s also  of  interest.  distribution  2  N. p l u m c h r u s community  in  i s an i m p o r t a n t  terms  Georgia  (LeBrasseur  During  the spring  the  the  abundance and b i o m a s s  e t a l . , 1969; G a r d n e r ,  zooplankton  i n t h e S t r a i t of  1977; M a c k i e ,  a n d summer, when t h e c o p e p o d i t e s  and i n t o  population  in  introduced  from  perhaps  from  occurrence  1985).  a r e found i n  depth  Saanich  Inlet  Inlet  the S t r a i t  (Black,  o f G e o r g i a , Juan  r e p r o d u c t i o n of t h i s the  overwintering  e t a l . (1983),  difference  contrast,  The  Strait  i n Saanich  period  however, s t a t e d  Hoos  between t h e i n l e t  (1970) and Cowen  o f N. p l u m c h r u s ,  in fairly  summer  of copepods  de F u c a  s p e c i e s s i n c e they  overwintering population i n Saanich  depth  1984).  may t h e r e f o r e c o n s i s t  r e p r o d u c t i o n o f N. p l u m c h r u s  following  Harrison  Saanich  f o r the  reproduce a t  (Fulton,  that  and  I n l e t . The  o f an o v e r w i n t e r i n g p o p u l a t i o n i s e s s e n t i a l  successful  1969  of  t o p 80 o r 100 m o f t h e water c o l u m n , t h e y c a n be t r a n s p o r t e d  southwest  an  of  component  1973).  t h e r e was a l a c k o f  Inlet  as a r e s u l t  and the S t r a i t  of the  of G e o r g i a . In  (1982) had r e p o r t e d t h e p r e s e n c e  h i g h numbers  during  the  winter  in  and 1974. In  Saanich  distribution incomplete. species  Inlet,  the documentation  o f N. p l u m c h r u s The p u r p o s e  in relation  of t h i s  can s u c c e s s f u l l y  overwinter  generations within the i n l e t . and  chemical  N. p l u m c h r u s population The summer  from  overwintering  population 1985  Inlet in  to- the  t o t h e oxygen  regime i s  was t o d e t e r m i n e and g i v e r i s e  I t i s proposed  characteristics  i n Saanich  in  study  on t h e a b u n d a n c e and  that  of  the  and  providing  if  to  subsequent  the  fjord  physical  may a  this  hinder resident  d u r i n g some y e a r s .  Saanich fall  Inlet of  1986  was sampled in  from  early  conjunction  with  3  hydrographic influence these and  d a t a . The h y d r o g r a p h i c  the d i s t r i b u t i o n  s t a g e s a r e found  t h e r e f o r e come i n t o  Hence,  the  Strait  of  a b u n d a n t and t h e i r 1973; since  Gardner,  shallower In  1977).  sill  in  the  Inlet  contact  the  oxygen  with  samples A  were  low  examined  comparison  history  is  Sechelt  Inlet  i n depth  laboratory,  oxygen  Saanich  their  was  migration waters.  f o r t h e s e two made  with  the  to  well  documented  was sampled  Saanich  Inlet  (Fulton,  occasionally  yet  it  has  a  (15 m) and i s w e l l - o x y g e n a t e d .  oxygen was m e a s u r e d various  following  p o p u l a t i o n where t h e s p e c i e s i s known t o be life  i t i s similar  potentially  a n d abundance o f CVs and a d u l t s s i n c e  as the CIVs.  Georgia  could  a t deep depths  zooplankton  s t a g e s as w e l l  parameters  Inlet.  t o l e r a n c e of t h i s  i n an a t t e m p t  levels  trap  s p e c i e s t o low  to estimate t h e i r  and t o r e l a t e  Sediment  were examined  the  this  samples  t o depth collected  survival  at  of m i g r a t i o n in  f o r s i g n s o f d i e o f f i n N. p l u m c h r u s .  Saanich  4  1.1.  Life  Neocalanus commonly  History  plumchrus  found i n c o a s t a l  In t h e S t r a i t  of Neocalanus is  a  of G e o r g i a , N. p l u m c h r u s  bodies  populations important and  food  egg  item  copepodite  and  Mackie,  (LeBrasseur  six  of c a l a n o i d s ,  subsequent  stages  naupliar  ( C I - C V I ) ( G a r d n e r & Szabo,  (CVI)  male o r f e m a l e . The p o p u l a t i o n  while at least  Ecdysis,  adult  during moult and  commences  declining  1969;  an  Fulton,  the  figure  adult  phase  months CV  egg and  in  as t h e m u s c l e  are  bivoltine  December  through  (Fulton,  1973).  place  (Fulton,  lipid  the a d u l t  reserves  eggs p e r c l u t c h .  the females d i e at depth. A d u l t  Following  into of  females c o l l e c t e d  (1973) e a c h f e m a l e p r o d u c e d an a v e r a g e o f  56.3  at  1973).  J a n u a r y and e x t e n d s and  the  Georgia,  takes  450 m e t e r s  tissue Of  in  of  to adult  early  which i s  o n l y once a y e a r ,  of  laying  six  2 ) . The  N. p l u m c h r u s  populations  from  and  1982;  1984). In t h e S t r a i t  females are u t i l i z e d .  with  is  t o i t s abundance  (NI-NVI)  breeding  oceanic  d e p t h s between 300  by F u l t o n  clutches period  the  release  mid-April,  1968  occurs  of  univoltine,  et a l . ,  fertilization,  Egg  in  Miller  after  overwintering  the  is  some o f t h e  1973;  February,  i s t h e s e x u a l l y mature  Georgia  fertilization  into  reproductive  species  al.,  stages  either  (Fulton,  et  1).  N. p l u m c h r u s d e v e l o p s from t h e  stage  of  The  f o r many p l a n k t i v o r e s due size  (figure  transported  establish  1985).  copepod  of t h e p r e d o m i n a n t  often  where i t may  1974;  calanoid  Columbia  i s one  is  final  Strait  of  1977).  is typical  into  water  large  Gardner, As  of  (Koeller,  relatively  1973;  species  waters of B r i t i s h  components o f t h e m e s o p l a n k t o n adjacent  plumchrus  the  9.5  reproductive  males d e c r e a s e i n  number  5  prior  t o females  so p r e s u m a b l y  t h e y d i e soon a f t e r  (Gardner,  1972; F u l t o n ,  1973).  occurs As depths  nauplii from  1973).  6th  i n t h e upper  nourishment  naupliar  Fulton, do  stage  coincident Harrison  lipid  from  in  late  phytoplankton  shallow (Fulton,  nauplii into  to  that  the  phytoplankton  level  egg  some  may the  ( L e e e t a l . , 1972;  below t h e e u p h o t i c zone so  ensuring  collected  February  stages  phytoplankton  CI  copepodites  through  (Fulton,  migrators start  descent  of  they  primary  laying  of  extends  the n a u p l i i are  bloom  t o deeper  depths  between  meters  (Gardner, large  (Hoos,  (Fulton,  1973;  1977).  and dusk  indicate  i n June  i f n o t a l l CVs  200  1972; F u l t o n ,  lipid  Samples o f CVs that  they  are  i n May and b e g i n 1973). By A u g u s t  reached  and 400 m e t e r s w i t h  on  1974). The s t a g e V  (Fulton,  have  through  primarily  Gardner,  i n t h e w a t e r column  depths  progress  feeding  1970; K o e l l e r ,  overwintering  t h e m a j o r i t y below 300  1973).  reserve b u i l t  m a i n t a i n CVs and a d u l t s w h i l e  reproduce  June n a u p l i i  CIV,  1973;  t o appear  September most,  The  to early  d u r i n g dawn, m i d n i g h t  diurnal  up o v e r they  t h e summer  overwinter,  i s thought moult,  and  ( L e e e t a l . , 1972; H a r r i s o n e t a l . , 1983). D u r i n g t h e  overwintering through  released  reserves, present  response  the s p r i n g  towards  e t a l . , 1983).  copepodite  to  float  100 m e t e r s o f t h e S t r a i t  their  and  months  with  From  and  or  a t t h e s u r f a c e . The p e r i o d o f  several  their  from  reproduce  productivity  not  swim  1973). A d u l t s r e p r o d u c e  not  over  they  d e e p r e g i o n s where t h e eggs were  While  obtain  develop  fertilization  a  period, i t i s believed  reduction  in  activity  that  they  conserve  energy  b u t t h e CVs may a l s o  f e e d on  6  alternative 1983;  food  sources  Dagg & W a l s e r ,  such  but  copepodite  occupying  column a r e c o n c e n t r a t e d southern  end  of  the  stage  copepodites  fall  and  deepest northern  i s ubiquitous  varies in density across stages  p a r t of  the  extension  are  upper  in early  a d u l t s which found  Strait. of  the  the  throughout  different  i n the F r a s e r  Strait  and  winter,  (Frost  et a l . ,  1987).  Neocalanus plumchrus Georgia  as m i c r o z o o p l a n k t o n  inhabit  Strait  the  towards  (Black,  1984).  deep  individuals of G e o r g i a  of  are  waters i n the  water the Fifth  in  the  central,  located in  (Black,  of  Young  area  i n g r e a t e s t numbers Fewer  Strait  regions.  regions  plume summer  the  1984).  the  7  2. THE STUDY AREAS In t h e f o l l o w i n g  section  physical  characteristics  influence  of the t i d e s ,  variation  with  of  detailed  the  winds,  season,  physical/chemical  a  study  and  is  in  sites  in  of  the  given.  The  flow  and  relation  of t h e water. In  how t h e s e c o n d i t i o n s i n f l u e n c e d study  is  estuarine  explain  to  area  discussed  characteristics  order  description  to  the  addition,  the s e l e c t i o n  N. p l u m c h r u s  their  in  I  of s t a t i o n contrasting  environments.  2.1 P h y s i c a l  2.1.1  Saanich Saanich  Vancouver  Inlet Inlet  Island  through  a series  has  a  maximum  km)  long  the  result the  is  situated  and  is  of passages depth  on  connected  1962).  environment  Water  stagnates  Of  the  nature  c o n d i t i o n s develop  influx  (figure  of  of Georgia  1 ) . The  fjord  m i l e s (24  interest  the  throughout  much  i n t h e bottom of the f j o r d  as a  which p e r s i s t s  tidal  energy,  o f t h e 75 meter s i l l  over  side  is  water e x c h a n g e w i t h a d j a c e n t w a t e r s Anoxic  the S t r a i t  prime  o f weak e s t u a r i n e c i r c u l a t i o n ,  restrictive  southeastern  to  and c h a n n e l s  anoxic  year.  the  o f 236 m e t e r s and i s 13 n a u t i c a l  (Herlinveaux,  characteristic of  Characteristics  (Stucchi  winds,  and  which reduces  deep  & Giovando,  the winter, spring,  of o r g a n i c d e b r i s s e t t l e s  and r e d u c e s  1984).  and summer a s t h e oxygen t o  8  levels  below  1.0  mL  L"  1  (Lu e t a l . ,  The G o l d s t r e a m r i v e r an  insufficient  circulation water the  at  latter come a  rivers.  different  t h e upper  order  Saanich  Inlet,  Pacific  Winds the  Both r i v e r s  (Herlinveaux,  to  (Pickard  have  estuarine  sources  of  renewal Ocean  from the F r a s e r of  i n t h e Ekman l a y e r by  & Emery,  and  During  depth of  outer  sill  of  a  volume o f w a t e r  renewal, entering  t h e movement  the  (Pickard  of  to  water  C h a n n e l must enough f o r  the e a s t e r n  cool  water  Strait  Pacific  from  by mid  Satellite  Strait,  Channel the  coast  waters  50-300  meters  late  (Anderson  and  summer  i s moved from a over  (Stucchi fjord,  up  (Herlinveaux,  and  the  65  & Giovando, thus  from y e a r t o y e a r as w e l l  the f j o r d  cause  These  to  up  of  1982).  s a l i n i t y w a t e r moves  renewal i n the f j o r d  i n Haro  coast  & Emery,  t o veer t o the r i g h t .  varies  to  bottom waters i n  w a t e r . Water dense  1984). The volume o f water e n t e r i n g of  the  i n the winter  e v e n t t h e dense w a t e r  100 m e t e r s o r d e e p e r  run-  tend  develops o f f the east  i n t o Haro  a flushing  switch  of  1982). The d e n s e , h i g h  e n s u r i n g an a n n u a l deep water 1962).  and  t o August  upwelling,  and  maximum  through S a t e l l i t e  i n the i n l e t  from A p r i l  fresh  t h e f o r m e r i n June and  replacement  of the r e s i d e n t  J u a n de F u c a S t r a i t  extent  an  1962).  t h e water e n t e r i n g  replaced  meter  primary  influencing  blowing equatorward along  water  are  for  have a p e r i o d  direction  orientation,  the d e n s i t y  deep water  the  water  times of the y e a r ,  layers  In  a  fresh  1962). The  from a s o u t h e a s t e r l y  exceed  of  provides  i n December. D u r i n g t h e summer months t h e winds  more w e s t e r l y  in  supply  of S a a n i c h I n l e t  t h r o u g h t h e mouth of t h e f j o r d  Cowichan  off  a t t h e head  (Herlinveaux,  enter  1986).  Devol,  the  as the 1973;  9  IOUBC D a t a R e p o r t 52, The  tidal  cycle  ( G e y e r and Cannon, Ocean  Science  vertical bottom  Giovando, due  the  density  8-10  during  i s maintained  with  high  low  salinities  tidal the  days d u r i n g  &  while  (1984)  Sound.  surface  ebb  In  the f l u s h i n g  the  Sound  by  w h i c h was of  a  gravity  entrained  w a t e r above  the h i g h  w a t e r moved f u r t h e r Given  the  concurrent  tidal  the  exchange  observations  w a t e r . The h i g h  a b o v e . The  Saanich  an  last  end  of  (1982)  flow d u r i n g water  for  by in  neap  migrated  when i t came water saline  was water  jump o r t h e e n t r a i n m e n t as  (Geyer and Cannon,  be a m a j o r  a  deep  dense  1982). renewal, of  studies  and  the o c c u r r e n c e of the f l o o d  phase  As  factor  water  the  i n the extent  Inlet.  indicated,  the  density  water d e c r e a s e d  regime  cycles could  have  hydraulic  t h e sound  density  in  at  underneath the l e s s  salinity  into  tends to  event to  t h e water d e c e l e r a t e d  current  lowering  causes  and Cannon  over  moved  spring  r e n e w a l mechanism  o f low m i x i n g . When t h e dense  with the b a s i n  &  1978.  as a r e s u l t  into contact  the  Saanich Inlet,  f o u n d a maximum b a r o c l i n i c  sill  strong  (Stucchi  hence,  tide  tides  Puget  a  of  in  flow a l s o  1984).  u n d e r s t a n d t h e deep water  They  tide  flood  e x a m i n i n g a c a s e s t u d y c o m p l e t e d by G e y e r Puget  neap  inflow periods  and O c t o b e r of  can b e s t  Institute  Strait,  the  estimated  the  i s weakened d u r i n g  Giovando,  t h e t h r e e main  A u g u s t , September We  gradient  exchanged  salinities  the  d e e p e r w a t e r s . The  (Stucchi  and G i o v a n d o  at  m i x i n g i n Haro  t h e i n f l o w o f water  acceleration Stucchi  that  to strong in  by  shows  and  1984).  t h e amount o f w a t e r  1982). D a t a c o l l e c t e d  1984). The v e r t i c a l  tide  inhibit  Burd & B r i n k h u r s t ,  influences  (I.O.S.)  gradient water  1983;  various  10  during  neap t i d e  water  into  would  favour  the i n l e t .  the entrance  to  52,  1983; Burd  estuarine  upwelling  event  clear  that  general annual  The  miles  in  relatively  As  The a  may  i n the S t r a i t  of Georgia and  (IOUBC  also  be  of Georgia  Coast.  In  inlet  Data  due  to  a  or a  reduced  conclusion,  i t is  a r e c o m p l i c a t e d but i n  of the a n o x i c  depth  of the S t r a i t  (220  km)  to the P a c i f i c  the  i s a large  the mainland  the south and Johnstone  Strait  but  along the P a c i f i c  Island  Connections  water  Inlet  bottom waters  on an  of Georgia  The maximum  nautical  to  flow  cycle  i s a replacement  Strait  Vancouver  to  Saanich  basis.  2.1.2. S t r a i t  1).  in  the c o n d i t i o n s in t h i s  there  of  & B r i n k h u r s t , 1984), c o u l d be a t t r i b u t e d  the t i m i n g of the t i d a l  weakened  volumes  The d e l a y , o r r e d u c t i o n i n t h e volume o f  renewed water d u r i n g some y e a r s Report  of l a r g e r  Strait  of  Ocean  50 m s i l l greater  between  Columbia  (figure  i s 450 m e t e r s a n d i t i s  l i e through  Strait  to  Georgia  119  the is  Juan  de F u c a  north.  Most  exchanged  straits  Strait of  through  r a t h e r than  the the  Johnstone  1983).  w a t e r moves i n t h r o u g h  the S t r a i t  of B r i t i s h  located  i n l e n g t h (Schumacher e t a l . , 1978).  wide and deep s o u t h e r n  (LeBlond,  estuary  of Georgia  through  i n Rosario S t r a i t  extent  than  the southern  the  passage  Haro S t r a i t restricts  sill  at  i t i s linked  and R o s a r i o  Strait.  t h e i n f l o w o f water t o 130  m  in  Haro  Strait  (Schumacker e t a l . , 1978). T h e s e t o p o g r a p h i c  features contribute  to  Strait  the counterclockwise c i r c u l a t i o n  in  the  as  do  the  11  tides  and t h e f r e s h water On t h e w e s t e r n  generally  greater  (Schumacker with  et a l . ,  depths  difference to  be  true during The  of  fresh  inflow  tides, in  (LeBlond,  1983).  depths  salinity  de  Although  Strait  In c o m b i n a t i o n w i t h t h e side  period  tends  t h a n on t h e  the  Strait  to  be  i s primarily  i s estimated to  outflow (Waldichuck,  o f maximum the  salinity  two-layered passes  with water  water flow  Strait  a  the waters  of  the  a r o u n d May or  is  thought  to  mixed  and  hence  the gulf  and San Juan  n e a r - s u r f a c e waters  s i n k down t o  denser  late  water.  In  o f f the P a c i f i c  (Waldichuck,1957).  continual  well  1957). The  t h r o u g h J u a n de F u c a is  upwelling  in intensity  contribute  parts  freshet  summer  through  and e s t u a r i n e  is  i t varies  t e n d s t o keep  shallower,  t h e deep r e g i o n s o f t h e S t r a i t  Fuca  circulation,  in  in  the winter the cold  high  the  much  t o t h e s o u t h and c e n t r a l  of h i g h  water  Juan  water  outflow  the  g r a d u a l l y moves i n t o from  is  along the western  which  period  the  the deeper  dense,  a  water  though  as  During supply  50 m.  flow  River  fresh  reaches  even  side  d e p t h s o f 450 m  tide.  plume e x t e n d s  the  Islands  than  estuarine  total  J u n e . The f r e s h  weakened  less  by t h e F r a s e r  and  Strait  1978). The e a s t e r n  the f l o o d  water  enhance  100 m and r e a c h maximum  Schumacker e t a l . (1978) f o u n d t h e o p p o s i t e  the  Strait  than  1978).  of G e o r g i a the depths are  and m a i n t a i n e d f o r a l o n g e r  strong  influenced 80%  of the S t r a i t  i n d e p t h s , t h e ebb t i d e  side.  et a l . ,  side  usually  stronger  eastern  o u t f l o w (Schumacker  f l o w , t h e deep process  of  summer coast, Georgia  As a r e s u l t o f water  renewal  throughout the year.  and i n the depth of i n t r u s i o n i t  oxygenated  at a l l times  (LeBlond,  12  1983).  2.1.3  Sechelt  Inlet  Sechelt Columbia  Inlet  southwest  is  located  of J e r v i s  on  Inlet  the  mainland  (figure  Inlet  are connected d i r e c t l y to Sechelt  almost  perpendicular  to the l e n g t h of the f j o r d .  meets  the  region  Agamemnon C h a n n e l Sechelt  Inlet  lowest  normal  basin  of  length  of 22  tide  restricted i s only  fjord  nautical  Maximum f r e s h  by  was the  25.8  has  water  (Pickard,  corresponding  runoff  normal that  1961).  range the  Georgia between  If  we  column  or S a a n i c h I n l e t 28  and  Sechelt The anoxic  and  average  and,  on  consider  is which  into  which at The  meters  main and  average  m.  occurs during  fresher  The  1957  temperature at  11.1°C  then  and  i t is  than e i t h e r  7.43°C  31  the  apparent  the S t r a i t of  have t y p i c a l v a l u e s 30 and  the  O c t o b e r , November  t h e s e v a l u e s t o be w i t h i n Inlet,  a  1961).  s a l i n i t y in July  28.58 p p t a t 200 was  Inlet  water  1961).  (Pickard,  June,  29 p p t a t 20 m and  temperatures are s i m i l a r  flow of  the i n l e t  of s a l i n i t y f o r S e c h e l t  water  Sechelt  of G e o r g i a t h r o u g h  (Pickard,  km)  and  a t an a n g l e  a very shallow s i l l  into  1961). The  depths  Inlet  a maximum d e p t h o f 300  m i l e s (40.6  p p t a t 20 m and  (Pickard,  Strait  14 m d e e p  summer and w i n t e r months i n May and December  the  and M a l a s p i n a S t r a i t . The  is  this  of  British  1 ) . Narrows I n l e t  Salmon  central  of  of  salinity  p p t a t 200  a l t h o u g h s l i g h t l y warmer a t  200  m.  The  m  in  Inlet. bottom  waters of S e c h e l t  conditions  Inlet  a r e n o t known t o d e v e l o p  a l t h o u g h Narrows I n l e t ,  which  i s adjacent to  13  Sechelt, oxygen L"  c a n become v e r y  level  in  1  low i n o x y g e n . P i c k a r d (1961) f o u n d t h e  i n t h e b o t t o m meter o f Narrows I n l e t  July  1957.  During  minimum  in Sechelt Inlet  1.4 mL  L" .  present  i n Narrows I n l e t ,  This  1  Despite there fjord the  presence  a t 100 m e t e r s and c o n t a i n e d  low  oxygen-containing  may be i n t r o d u c e d i n t o S e c h e l t  shallow  of  waters.  sill  Inlet.  between t h e  s u p p l y o f f r e s h water  bottom  waters  indicate  a  Station  the department  sampled  the  of Oceanography  hydrographic  at  parameters  3 ) . T h i s was an i n f l u e n t i a l  research  (figure  d e c i d i n g on t h e s t a t i o n study.  from  Station  A u g u s t , September, a n d November for  the  September close  remainder  of  deepest meters copepod  the  1986, i n c l u d i n g  p r o x i m i t y o f one portion (table under  concentrations  study  G1748  study  another  to  was  factor  samples  sampled was  and  for  in July, maintained  p e r i o d from December 1985 t o  1985. B o t h and  with a depth  o v e r w i n t e r s below  stations are within  l i e within  i s an i m p o r t a n t  i n the c e n t r a l  physical  collect  1985, and G1545  October  of the S t r a i t ,  1). Depth  which  f o r both  has  at stations  biological  particular  fairly  UBC  of G e o r g i a  this  and  Selection  G1545 and G1748 i n t h e S t r a i t  in  Inlet  of water.  2.2. Over t h e y e a r s  water  a t t h e mouth o f S e c h e l t  The l a r g e  oxygenated  movement  consistently  that  mL  p e r i o d t h e oxygen  t o be a s t r o n g e s t u a r i n e c i r c u l a t i o n  and a d j o i n i n g  continuous  was l o c a t e d  implies  the very  appears  t h e same s a m p l i n g  t o be 0.15  the  central,  of approximately  characteristic  since the  200 m and r e a c h e s  region during the f a l l  400  and  maximum winter  14  months  (Fulton,  The  1972;  sill  at  exchange of water N.  plumchrus  are  below  Black,  the  mouth  thus a c t i n g  , especially 75  1984).  m.  The  of  Saanich  as a b a r r i e r  Inlet  t o sample t h e c o p e p o d s w h i c h were  from,  immigrating  characteristics Two  of the  stations  sampling  southern  station,  maximum  depth  station,  Saa9, w h i c h l i e s  of  Patricia  stations the  of  movement  from  exchanging  A  yet  it  N. was  would  fjord  Satellite of  would  receive  not a h y d r o g r a p h i c  1985  the  Physical  of  160  Inlet  (table  Reach and  t o the m  at  2).  1 ) . The  sill  location  the The  has  a  northern northwest  (figure  4).  Both  a t o p p o s i t e ends o f by  water  and  hence becomes a n o x i c  sooner  is  was Saa3  was 4).  anoxic  plumchrus with respect  4).  influenced  more  affected  and  by  therefore  water i s more  copepods.  (figure  reach  emigrating  less  (Saa3)  it  in Saanich  table  Channel  that  Also,  station  is  station  station  sill  outside waters. renewal  station  northern  parameters the  of t h e i r  o u t s i d e the  central  behind  a maximum d e p t h  to the transport  away from  just  (figure  in Squally 4;  copepods  Channel  either  of  station.  May  (figure  because  through  biological  develop  has  in  is situated m  southern  t h a n Saa9. The  exposed  216  Bay,  The  Inlet  were u s e d  period  Saa0.8,  were u s e d  inlet.  Saanich  were not m e a s u r e d a t t h i s  hydrographic  beginning  into,  when t h e  of the S a t e l l i t e  ( S a t e ) was or  the  f o r the t r a n s p o r t  a t times of the year  purpose  restricts  established The  conditions  would  copepods through probable enabling  to the d i s p l a c e d  station  location  due  that  the  the low  to l i m i t e d  to  measure  i s f a r enough most  likely  exchange deep  with water  examination oxygen z o n e .  ship  time.  of Saa3  15  Sechelt and  August  sampled  of  was  sampled  1986. D u r i n g  i n the deeper  estimate August  Inlet  the s i z e  1986, t h r e e  the population  i n November  t h e two w i n t e r  regions  of the  1985, and i n F e b r u a r y  months, s t a t i o n s S c 1 , was inlet  of the o v e r w i n t e r i n g  in  population  s t a t i o n s were u s e d t o a l l o w variability  within  the  inlet.  an  attempt  (figure  f o r an  to  3 ) . In  estimation  16  3. MATERIALS AND Field  data  fluctuations  were  Satellite  Inlet.  Hydrographic  determining  Channel,  or  of  the  oxygen  in their  series  of e x p e r i m e n t s  Inlet  natural  resulting  May  1985  hydrographic correspond Calvert  or G1748 were  of  habitats,  f o r N.  i n the p h y s i c a l  N.  chemical  c a r r i e r ! out  To  necessary  features  Hydrographic  Data  September and  sediment  to conduct  with either  (table  Saa0.8,  in  2  than  purposes  occasionally d e n s i t y and  interest. reversing  the so  &  Saanich  (table  of  3).  The  Inlet,  of D r .  S.E.  (figure  4).  station  above the  a  levels  Saanich  UBC  low  once a month  stations  Oceanography,  biological  f e a t u r e s of  1986  were measured a t e i t h e r  salinity,  from  approximately  t r a p mooring  of  our  plumchrus to o b t a i n evidence  3.1.  Saa9  in  enhance  t r a p samples  conditions.  were o n l y sampled  Temperature,  Sechelt  plumchrus to s u r v i v e  i t was  3). S t a t i o n s other  for  Saanich  importance  unfavourable  characteristics  added  in  of G e o r g i a , and  from  Department  (figure  conditions  plumchrus.  d a t a were c o l l e c t e d  the  population  i n w h i c h t h e y were s u b j e c t e d t o low  stations to  parameters  and  ability  through  of t h e  Physical  N.  of  paramount  changes of  examined  Hydrographic from  not  of  i n t h e l a b o r a t o r y . Sediment  were a l s o  mortality  analysis  Strait  were  the d i s t r i b u t i o n  oxygen  for  the  data  whether  understanding  of  collected  under v a r y i n g e n v i r o n m e n t a l  Inlet,  influenced  METHODS  G1545  mentioned  hydrographic  2 & 3).  o x y g e n were t h e  physical  T e m p e r a t u r e measurements were thermometers  or  a  Guideline  1 7  CTD  probe,  Surface  both  temperatures  thermometer Autosal from  salinometer  (1978).  0.003  used  were m e a s u r e d  to verify Density,  probe  and  using the  i n t h e 2-42 p p t s a l i n i t y  ongoing can  Seawater  range.  from  were d e t e r m i n e d  method on b o a r d  These v a l u e s  54  were  ship  with  the  temperature  with  (Carritt  a  UNESCO  and s a l i n i t y  modified  & Carpenter,  Winkler  1966). The  intervals  from  the bottom.  hydrographic  data  research project  be f o u n d  Salinity  have an a c c u r a c y o f  a s at, was c a l c u l a t e d  of  levels  s u r f a c e t o near The  Guideline  Practical  a b o v e p a r a m e t e r s were measured a t 9 t o 12 d e p t h the  the  CTD m e a s u r e m e n t s .  State  v a l u e s . Oxygen  sample w i t h a  i n obtaining c o n d u c t i v i t y values  was e s t i m a t e d  expressed  of  titration  a bucket  The s a l i n o m e t e r d e t e r m i n a t i o n s  ppt  Equation  were u s e d  o f + 0.01°C o r b e t t e r .  from  t o w i t h i n 0.1°C. The CTD  which s a l i n i t y  Scale +  o f w h i c h have an a c c u r a c y  are  being c o l l e c t e d  a s p a r t o f an  a t UBC. A d i s c u s s i o n o f t h e  i n t h e department  techniques  o f O c e a n o g r a p h y UBC d a t a r e p o r t s ,  & 55 (1985 & 1986) .  3.2. Zooplankton Satellite Sechelt hauled oxygen  samples  Channel, Inlet  Zooplankton  the  (table  were  Strait  Sampling  collected of  Georgia,  2 & 3). Vertical  a t v a r i o u s d e p t h s d e p e n d i n g on profile,  and  the  from  documented  Saanich  and  on  and h o r i z o n t a l the life  time  of  history  Inlet,  occasion, nets  were  year,  the  p a t t e r n s of  N. p l u m c h r u s . Vertical  haul  s a m p l e s were i n i t i a l l y  collected  u s i n g a SCOR  18  net  w i t h an i n s i d e  For  the  September  constructed  to  Oceanography, net  diameter  o f 57 cm a n d a 333  1985 c r u i s e ,  include  U.K.)  an  n e t t i n g . T h i s n e t was u s e d on  station,  the  vessel at 1 m sec"  m  The  1  but  was  mSCOR  sampled.  1  and r a i s e d  speed of t h e h a u l  depth  An  for the area  within  seven  of  was  5 ) . The mSCOR 293  jum  so t h a t t h e  was  nitex  hauls.  lowered  from t h e  v a r i e d somewhat w i t h  echosounding  of  to the surface at  w i n c h o p e r a t o r s . The d u r a t i o n o f t h e h a u l being  (figure  made  net  mesh.  Institute  f o r a l l subsequent v e r t i c a l  stationary sec" .  mechanism  nitex  SCOR n e t (mSCOR) was  N.I.0.(National  throttling  was o f t h e same d i a m e t e r  Once  a modified  nm  different  depended on t h e  used  vertical  0.5  depth  t o o b t a i n a maximum  hauls  could  be  kept  t o t e n m e t e r s o f t h e b o t t o m t o a v o i d damage t o t h e  net. Clarke-Bumpus n e t s , Frolander net  (1957),  was a t t a c h e d  diameter area  nets  the  (Yentsch was  metal  volume  of  housing  which  readout  & Duxbury,  roughly  towed by hand  Paquette  had  an  t o the wire  mounted a c r o s s on  the  t h e open  flowmeter  a t a time  and  were  volume  1956; Regan,  1.5 k n o t s ,  value  of  filtered 1963).  and  a 30° (  m «rev" 3  through  The  to four  f o r twenty  t o maintain  0.0043  from  opened  o f m e s s e n g e r s . C-B n e t s were towed  the  and  internal  w a t e r c o u l d be c a l c u l a t e d . T h r e e  angle. A c a l i b r a t i o n calculate  to  f o r t h e h o r i z o n t a l tows. A 500 M m  a t a s h i p speed o f about  + 5°) wire to  a  used  t o a mechanical  by a s e r i e s  minutes  used  to  were a t t a c h e d  closed  were  according  o f 12.4 cm. The i m p e l l e r b l a d e  was l i n k e d  which  modified  was  1  t h e C-B n e t s  calibration  value  t e s t e d u s i n g two o f t h e f o u r C-B n e t s . E a c h n e t was f o r a d i s t a n c e o f 15  meters  and  the  number  of  19  revolutions determined 0.0047  were  recorded  f o r net  m «rev" . 3  #23  was  This  1  for  six  0.0041  was  replications.  m «rev~ 3  considered  to  agreement w i t h t h e documented c a l i b r a t i o n replicate  tows were run  the v a r i a t i o n Paired modified  percent  of  that  vertical  haul  revolutions  of the  flowmeter  midway  average  of- 46%  calculated  between  f o r each v e r t i c a l  haul  the an  D e t e r m i n a t i o n s of  the  from  measured inside  of t h e n e t the  by  the net  (table  4 ) . As  flowmeter  correct  (volume  of  obtain  (1968),  to  of s i x  with  to  of t h e n e t and  used  intervals overlapped  G1545, 200-300 m and copepods  reported  overlap  was  the c e n t e r . copepod  filtered  i s taken  cubic  subtracted  at  with  a  the  given  meter,  data table  into  =  An  counts  7rr d«0.46, 2  each  (table  from  the  the  number  15)  1986, number  interval,  are  but  the  for  42-45, 48-49) t h e  the depth  and  number  sample v a l u e s a t t h e  concentration  collected  net  ( e . g . June  depth  39,  Both  haul  calculated  &  (figures  consideration.  the h i g h e s t  13  G1545, t h e number of c o p e p o d s from  vertical  station  for  figures  are subtracted  containing  1986,  sampled  200-395 m s a m p l e s ) . The  distribution  copepods  interval  per  i n t h e raw  vertical  June  rim  a c c e p t a n c e was  depth  occasionally  of  the  good  r = r a d i u s of the n e t , d = d i s t a n c e h a u l e d ) .  The  of  Aron  #31,  estimate  taken  order  flowmeter  i n the absence  net  series  t h e mSCOR n e t was  of the  average  in f a i r l y  an  were  among s a m p l e s . into  be  to obtain  occasions in  by G e h r i n g e r and  placed  1986  for  value. A  replicates  several  the v a r i a t i o n  the  recommended  where,  samples.  a c c e p t a n c e o f water  comparing to  between  SCOR n e t on  estimate  in July  and  1  The  of copepods  between  between  depth (i.e.  200-300 m 200-395).  was The  20  resultant and  300  calculation  m and  During tow  for  6 ) . The  between 300  t h e November  the  C-B  a  and  was  395  1986  n e t s was  instrument  obtain  i s t h e number p e r  cruise,  tested  readout.  b a t h y k y m o g r a p h d u r i n g t h e two a  maximum  of  n e t s were on The  Bioness  Patricia  of  zooplankton  sampler was  at s t a t i o n  filtered  of was  25  and  determined  acceptance  150  of  m for  five  depths  (figure  minutes  recorded  to  on  the  a variation depth  of  when t h r e e  6). the  Institute  on M a r c h 5,  Saa3. The  1986  Bioness  of 2 knots. F i v e  50  depth  calculated  tows i n d i c a t e d  owned by  utilized  200  w i t h a bathykymograph  The  C-B  (figure  15 m i n u t e s a t a s p e e d  intervals  90%  Bay  the  m e t e r s below t h e c a l c u l a t e d  the wire  at  for  10  between  m.  s e n t down t o  baseline  c u b i c meter  m  (table  f o r the  0.25  of w a t e r , a l t h o u g h  2  Science  collection  towed  obliquely  n e t s were r e l e a s e d a t  net  i t may  f o r the  was  17). m  of Ocean  be  The  volume of  water  opening  assuming  slightly  lower  a  (Dave  Mackas, p e r s . comm.).  3.3. Specimens formalin, samples N.  CV,  were  used and  nauplii being  with  examined (figure  for  2 ) . An  stages  to CIII  posed.  It  were  Haul  Samples  immediately  calcium  f o r c o u n t i n g and  CVI  of Net  were p r e s e r v e d  buffered  plumchrus  was  Treatment  one M5  after  carbonate. species  Wild Leitz  capture  Afterwards  of c a l a n o i d dissecting  species i d e n t i f i c a t i o n . examined  since  an  in  microscope  Only  the  CIV,  analysis  of  the  t o the  is  distinguish  more  difficult  to  the  copepod,  s t a g e s would c o n t r i b u t e l i t t l e also  3-5%  questions the  21  naulpii-CIII identify size it  stages  from o t h e r  t h e CIV-CVI  stages of t h i s  was  necessary  to  This  remove  species.  Szabo  Folsom  samples  splitter  samples  hundred  collected  examined  alternately  during  quantity  o f N. p l u m c h r u s  Folsom  splitter.  The l a r g e s t  averaged  11.8%  of the t o t a l  3.4. Live  (chamber  Inlet  four  stage f i v e  were  counts of a  actual  total  A ) . The maximum o v e r e s t i m a t e a v e r a g e d  and Maintenance of L i v e  c o p e p o d i t e s were  distilled  concentrated  water.  tolerance  thermos  flasks  with  a  the density  of copepods  monthly,  tests.  were  The l i v e  samples  Copepods  collected  acid  glass  containing  washed  in captivity  each  and  removed  pipette  3.5 l i t e r s  into  from  A l l equipment  s p e c i m e n s were  w a t e r . A maximum o f 55 s p e c i m e n s were p l a c e d that  three  chambers  underestimate of the  and s t o r a g e o f c o p e p o d s  so  Since  (318 CVs) were u s e d t o t e s t t h e  i n handling  into  and  used f o r S t r a i t  sub-sample  used  transferred  fourths. exceeded  was o n l y  1986, f o r oxygen  the  into  rarely  t o November  from  Gardner  (chamber C ) .  Collection  with  by  from  o f N. p l u m c h r u s a  May  rinsed  for  the species  t h e c o u n t . Ten r e p l i c a t e  known  legs  numbers  specimens, the Folsom s p l i t t e r  of G e o r g i a c o u n t s . E a c h o f t h e  of  identification.  t h e sample  i n Saanich  than average so  pair of  the  f o r species  large  was u s e d t o s p l i t  than i t i s t o  Occasionally  manuscript  reference  containing  5th  separation  taxonomic  (1982) was t h e main For  the  allowed  C a l a n u s m a r s h a l l a e . The  8.4%  copepods  o f t h e C I V a n d CV N. p l u m c h r u s was s m a l l e r  identification.  the  calanoid  and of sea  thermos  d i d n o t e x c e e d 14  22  o r g a n i s m s L~ .  The  flasks  (4°C)  until  1  refrigerator  t h e y were t r a n s p o r t e d copepods  were  subsequently For depth  um  bactericide  at  UBC.  The  experiments  #1-5  and  from  the  either  i n the S t r a i t of  remove s m a l l d e b r i s .  of time spent  measured  7°C)  The s e a w a t e r was f i l t e r e d  May  12th  with  of  was  water  t h e Accumet  model  taken  of  live  from t h e Department  within  range of t h e sea water  collected. in  No a d v e r s e e f f e c t s  the c o n t r o l While  copepod  didymus,  I_. g a l b a n a 1983)  weissflogii,  Pacific  was  s e a w a t e r were  Isochrysis  galbana,  collection  were  apparent  3 t o 5 days Chaetoceros  an  absence  be a t t r i b u t e d  by t h e  a t UBC. T. w e i s s f l o g i i a n d  a r e known t o be e a t e n by N. p l u m c h r u s  i t could  27.5 p p t ,  from which t h e copepods  from t h i s  7.4  during the  The p h y t o p l a n k t o n were s u p p l i e d  Culture  s o , i f t h e r e was  flasks,  water  t h e c o p e p o d s were f e d e v e r y  a n d C. g r a c i l i s .  North East  the  of  group.  in captivity  with T h a l a s s i s i r a  of  at the  i n sea water  o f Z o o l o g y s e a water o u t l e t  r u n s . The s a l i n i t y  the  a pH  c o p e p o d s were p l a c e d  experimental  i n the  140 pH meter  the  Some  days  2 4 t h ) . No  contained  and  water  73  July  p e r i o d . V a l u e s r a n g e d between  8  so t h a t t h e  w a t e r was  changed  the  an  The f l a s k  end o f t h e c a p t i v i t y 8.0.  with  on two o c c a s i o n s  i n the o r i g i n a l  was u s e d . The pH  was  i n the  s e a w a t e r was c o l l e c t e d  the copepods  to  collected  ship  a t 7°C.  by  Inlet.  during  t h e c o p e p o d s was c h a n g e d  water  aboard  end o f t h e c r u i s e a t w h i c h t i m e  4°C  tests,  carboys  maximum p e r i o d  flasks  at  occupied  into  containing  (the  held  or Saanich  stored  t o a c o l d r o o m (4°C o r  the laboratory  net  the  t h e y were h e l d  range  Georgia  were  of  to a lack  fecal  (Frost  pellets  o f f e e d i n g . One  et a l . , i n the thermos  23  of  copepods, c o l l e c t e d  June  1987,  comparison to  the  were of  not  the  fed  held  their  separate Water  100  in  period  of  although for  a  lowered  The  silicon  the  to  on  water  from t h e  by  a l a c k of  the  to  with  oxygen  on  water to  100  the  found at  mL  four  light.  nitrogen  for  of  oxygen  an  levels  i n the  oxygen.  control  Oxygen  overwintering  levels  The  required  c o n s i s t e n t oxygen  these  two  20  liter  to c o n t r o l the attached  test  to the  (figure  d i d not  polycarbonate  was  depths  have  any  nitrogen,  top  of  the  the  stoppers  and  funnels  separatory other  500  carboys,  of  two  c o p e p o d s were added  b e a k e r s . Once t h e  four  flow  7 ) . Two  c a r b o y and  c o n t r o l c a r b o y . Ten  from the  in  copepods.  samples the  food.  concentration.  level  s t r i p p e d of  that  funnels,  tubing  was  so  f o r e a c h e x p e r i m e n t c o n s i s t e d of  a valve  it  days  adjust  bubbled  to those  the  1987  c o p e p o d s were p l a c e d  the  nor  ensure  set-up  plastic  attached  funnel  similar  stoppers,  were  was  depended  June  Procedures  ten  reduce  The  a  copepods  8  t i m e of e x p o s u r e t o n i t r o g e n . Water  affects  r e m o v a l of  impaired  p o s s i b l e to obtain  separatory  of  for  in  enabled  s t a r v e d October  12 h o u r s t o  neither aerated  basic  glass  piece  bubbling not  for  carboy  15 h o u r s t o  Strait  deleterious  mL  test  to l e v e l s  the  Experimental  beakers  i t was  c a r b o y s was  in  mL  given  not  This  in c a p t i v i t y .  captivity  each experiment  the  a v e r a g e of  in  copepods c o l l e c t e d  food.  the  held  h e a l t h was  3.5. to  while  the  with  m o r t a l i t y r a t e of  c o p e p o d s were o n l y  Prior  provided  copepods  presumed t h a t  i n O c t o b e r and  were  a  for  funnels received to  each  in  place  24  air  b u b b l e s were removed  flushed  from  the  t h r o u g h t h e f u n n e l s . A 250  lines mL  from t h e t o p and a n a l y z e d f o r oxygen  and  water using  the  sample  water  was  was  collected  the Winkler  titration  method. Oxygen period  levels  were l o w e r e d g r a d u a l l y  of the f i r s t  hours,  water  prevent  build  volume  of  necessary  six  samples  experiments  water  water  in  beginning more  the l a s t  these  i t d i d not  appear  so t h a t  level.  Water  representation  i t was  necessary to  being tested  copepods samples  although  (table  flushed  at the  were e x p o s e d  were drawn  to  a  f o r oxygen  o f t h e t w e l v e hour  period  in  instances.  (table  v a l u e s ranged  5 ) . Oxygen  to  6.94  be  replicates  mL  L" .  but  as 0.28  collected  mL  on  it  since L~ , 1  from 0.26  tested  t h e two  t h e oxygen  e a c h was  t h e same d a t e would became  test  mL  L~  1  during  treated  that  differed  were  not  5 ) . I t had  been  oxygen  4.20  by  as  Also,  necessarily assumed  in a similar  copepods  t h r o u g h t o November d i d n o t d e m o n s t r a t e  from  as a s e p a r a t e t e s t .  funnels  r e s p o n d t o low  tests  f u n n e l s were i n t e n d e d t o  values often  (table  apparent  t o 2.24  i n the c o n t r o l s v a r i e d  i n e a c h of t h e t e s t  o f t h e CVs  However,  levels  Originally  1  the copepods  all  levels  to  large  seven e x p e r i m e n t s were o n l y  a t t h e b e g i n n i n g and end  Oxygen  much  environment,  three  relatively  perhaps a b e t t e r  i n t h e oxygen  of the experiment  c o n s t a n t oxygen  analysis  was  hour  flushed  e v e r y 3 h o u r s . In a d d i t i o n ,  the n a t u r a l  reduce the v a r i a b i l i t y  Every  the f u n n e l s  f o r t h e number o f c o p e p o d s ,  to replace  Hence,  and  5).  up o f m e t a b o l i t e s . Owing t o t h e  conditions  5).  (table  were c o l l e c t e d  a g r a d u a l d e c r e a s e i n oxygen of  over the twelve  collected  t h e same l e v e l  that  manner. from of  May  oxygen  25  tolerance. The  e x p e r i m e n t s were  durations  (table  examined  prior  categorized activity. was  to  as  control  healthy,  was  under  was  with  criteria  for classifying  a  lack  To d e t e r m i n e  which  the  as of  copepods  of  the  on  particle.  activity  60  and  t h e o r g a n i s m as  seconds  3.6. Paired at  sediment  trap  water. At s t a t i o n 135,  and  180  m.  cloudy tissues  copepods two  s t a t e . One  mid  (110 m),  out of the s i x  solution  to prevent the l o s s  the d e n s i t y  gradient  In  were p l a c e d i n s p e c i e s as  a  trap data.  Saa9 were moored (150  m)  t r a p s were d e p l o y e d a t  50,  t r a p s were a t t a c h e d  t o deployment  increasing  experiments  Samples  each depth. P r i o r of N a C l  categorized  the experiment.  samplers at s t a t i o n  one  were t h e  "dying".  t o the sediment  Trap  If  heartbeats  the copepods  Saa0.8 t h e s e d i m e n t Two  of  irregular  days a f t e r  relevant  t h r e e d e p t h s , s h a l l o w (45 m),  i t s level  observation  following  in this  Sediment  was  of  possible,  specimens,  T h i s was  were  heartbeat.  s e p a r a t e b e a k e r s t o d e t e r m i n e t h e b e h a v i o r of t h i s settling  hour  copepod  for a  Slow  for observation  dead  12  low oxygen  based  microscope  o b s e r v e d , r e c o v e r e d , 5-7  case  to  or  s i g n s of movement t h e prosome  dead.  were f o u n d  18,  Each  o r dead  showed no  i f r e c o v e r y was  " d y i n g " were h e l d  copepods  subjected  dying,  reported  21,  organisms.  a dissecting  coupled  in  24,  no a p p a r e n t h e a r t b e a t a f t e r  the copepod  as  for  Copepods  I f the copepod  examined  there  5).  run  t r a p was  and d e e p  t o t h e main c a b l e a t filled  of sediment  with  a  l a d e n water  between t h e t r a p w a t e r  and  30% by the  26  surrounding  water.  solution  o f sodium  settled  into  The  other  azide  the  trap  was  filled  (NaN ) t o p r e s e r v e  the  3  chamber  and  to  reduce  pieces  of  various  with  a  sample  the  0.5%  as  loss  of  it trap  contents. Whole o r g a n i s m s separated N.  from  plumchrus.  examined  or  the sediment Samples  from  from  March  for  in  Inlet.  A fixed  differences  Saa9  November  Statistical  plumchrus  stations  Sechelt  to  and  parts  were  t h e n examined  for  and  Saa0.8  were  1985  and  June  from  1986.  3.7.  fixed  samples  stations  through  t h r o u g h t o September  Neocalanus  trap  body  the  was  Methods  sampled  Strait  effects  on  a  basis  at  of G e o r g i a , S a a n i c h I n l e t ,  and  two-way ANOVA was  among t h e means between  to  test  s t a t i o n s and months i n  haul  through  the  GENLIN  (MTS  computing  system),  was  utilized  i n t h e 2-way ANOVA c a l c u l a t i o n s .  A l l tests  were run  at  the  level,  homogeneity tests  at  copepods station  the  vertical  Typically  the S c h e f f e ,  ANOVA was  per c u b i c  i t was  Bartlett  design  available  Chi-squared Bonferroni,  test for  and  Minimal  mean  number  i n the h o r i z o n t a l  samples  variation.  meter  Saa9 i n J u l y  ANOVA t e s t s  unbalanced  a t UBC  the  o f v a r i a n c e and  A single-factor of  program  including  f o r s o u r c e s of  The  used  the v e r t i c a l  5%  samples.  .monthly  used  t o compare t h e  collected  1986.  To meet  the  assumptions  n e c e s s a r y t o t r a n s f o r m the t o t a l  h a u l s and  data c o l l e c t e d  t h e number m~ for  3  copepod  of  the  numbers  from  from t h e h o r i z o n t a l population  studies  tows. are  27  transformed  using  N. p l u m c h r u s ,  the data  in  log(x+1) transformed  The  transformation Power  Law  log  linear  of  mean was c a l c u l a t e d  haul  samples c o l l e c t e d Sechelt The  Inlet  P  and  was  first  of  the  by T a y l o r in  which  against  the l o g  of r e p l i c a t e ,  of  vertical  of G e o r g i a ,  Saanich  line  fitted  logs /logx  this  t h e Power  from the s l o p e  Y i s the transformed  transformation  the  2  for  result  t h e l o g o f t h e mean. A  f o r t h e 22 p a i r s i n the S t r a i t  proposed  i n v e r t e b r a t e taxa  increased with  R =0.93. From t h i s  was d e t e r m i n e d  calculated  d i d not  For  Inlet,  (figure 8).  equation  y=0.07+1.60x,  1983).  from t h e mean t h e r e f o r e  r e g r e s s i o n of the l o g of the v a r i a n c e  the  X =Y,  distributed  the v a r i a n c e  Sell,  used.  transformation  (1961) f o r c o n t a g i o u s l y the  was  and  i n t h i s manner  the independence of t h e v a r i a n c e  Power Law  and  (Evans  value.  species  recommended  2  Law  Power X' 0  2  Law  which  was  transformation,  ( b ) , where b=1.60,  The is  to  p=1-b/2,  transformation  i s also  by Downing e t a l . ( 1 9 8 7 ) .  t h e power  28  4.  4.1. In t h i s  and  A n a l y s i s of  section,  are presented.  RESULTS  The  the P h y s i c a l Data  the p h y s i c a l  variations  oxygen, r e l e v a n t t o the  data  f o r the  in s a l i n i t y ,  study  of N.  the  Pacific  Georgia  and  Because  of  fjord  Saanich the  Saanich 1986.  sank and &  during  extremely  certain  shallow from  The  sill  the  showed d i s t i n c t i v e  cool,  mixed w i t h  sufficiently w a t e r s from  low  salinity  two  At  Saa0.8  cool  Oxygen August,  of  Strait the  other  study  of  year.  in Sechelt Inlet,  seasonal  this  areas.  January  and  to J u l y  31.20  levels  trends  w a t e r s c o o l e d by  ppt  than  150  ( f i g 20  m and  anoxic  (figure  waters,  19 &  mL  L~-  1  10  were  not  bottom waters  salinities  22).  began t o d e c l i n e a t Saa9 i n A p r i l 0.31  9,  months  r e p l a c e warmer  c o n d i t i o n s and  1985  winter a i r  & 2 1 ) . Hence, b o t t o m  (figures  r e a c h i n g a minimum of  in  in previous  near-surface  d e n s e t o s i n k below  31.03  times  b o t t o m w a t e r s a t Saa9 i n M a r c h  Saa0.8 were c h a r a c t e r i z e d by  between  i n f l u e n c e of  Inlet  Inlet  12).  The  n o t i c e a b l e i n the  11). T h i s water c o n t a i n e d more oxygen  (figure  at  Inlet  remains q u i t e d i f f e r e n t  4.1.1. S a a n i c h  and  especially  density,  plumchrus, are d e s c r i b e d  r e s p e c t to the p o s s i b l e c a u s a t i v e f a c t o r s . is  regions  temperature,  with  Ocean  various  i n August at  through a  to  depth  29  of  140  m.  From  May  presumably as a r e s u l t was  greater  dense  fluctuations  the  w a t e r was A supply  the  Fraser  from  than  to  in salinity  progressive  replace  decline  stagnating  August  conditions  Due  (figure  at  mL  L"  1  only  i n t o August, i n d i c a t e d  that  (figure  anoxic  17). T h i s  and t h a t  the i n l e t  was  September, d e n s e , h i g h  salinity  were  In 1985, t h e decreased  that  anoxic  onset  w a t e r was of t h i s  to  below  185  throughout  marked  displacement  of  low  situated  between  75 and  160 m d u r i n g  1  100 and  (table  160 m r a n g e d between  oxygen  September  high  L"  indicated  interest  low  of t h e w a t e r r a n g e d from 0.02  mL  was  mL  L"  1  and 0.17  mL  mL  zone  (table 9). L~  1  9 ) . In O c t o b e r oxygen v a l u e s 0.00  below  oxygen w a t e r from t h e b o t t o m  The o x y g e n c o n t e n t o f 0.26  present  water  o f t h e deep water r e n e w a l . Of p a r t i c u l a r  d e p t h s a t b o t h Saa9 and Saa0.8. At Saa0.8 t h e was  and  not  22).  parameters  suggested  75 m a t Saa9 and Saa0.8. C h a r a c t e r i s t i c s  the  1  was  (figure  1986.  of  length. In  the  L"  lower a t  circulation  1985 and mL  was  persistence  water column  0.11  perhaps  i n t h e t o p 50 m  water,  estuarine  August  and  salinity  the hydrographic  a t 165 m was  a t 185 m  salinity  I n May,  the e n t i r e  in  Small and  fresh  that  s h i p time,  Saa3  m t h e w a t e r was its  waters. depths,  lowered  added  to c i r c u l a t e  oxygen c o n c e n t r a t i o n 0.02  bottom  not  12).  Saa0.8.  suggested  to l i m i t e d  measured  the  i n oxygen  at  decreased,  Exchange o f w a t e r  of f r e s h w a t e r from G o l d s t r e a m R i v e r  to  enough  runoff.  and t e m p e r a t u r e a t d e e p e r  Saa0.8 t h a n Saa9. D e s p i t e  strong  River  salinity  below b e c a u s e t h e w a t e r was  and Cowichan R i v e r s ,  May  anoxic  near-surface  of F r a s e r  above t h e s i l l  sufficiently  the  to July  L  - 1  to  a  between .  Water  30  from  the  bottom  a v e r a g e of 0.38 At Saa9 120  m  in  40  mL  0  m  was  replaced  1  t h e low oxygen September  in  zone was  and O c t o b e r  August  oxygen  water was  Saa0.8  where  subsequently  at  being anoxic  of  17) was  the water  water  continued  Associated  were  i n oxygen  the  content  new  water x  lower  with  The  10  that  that  this  of  developed  below  low  station and  were  150 m a t Saa0.8  (between A u g u s t m  B  dense,  high  levels  were  high  density  the  o f oxygen  below  75  into  March  1986  oxygen  0.50  fell  below  0.20  A n o x i c c o n d i t i o n s were f o u n d below in contrast  t o 1985  developed  i n January ( f i g u r e  injection  of water  mL  inflow  of o l d , l e s s water  1985. Maximum during  w a t e r was  the f l u s h i n g  a  this  further  1  and  just  existed L"  1  in  190 m i n J u l y  2 2 ) . T h e r e may  event  (figure  i n which anoxic  from the s u r f a c e  A).  m.  L~  minimum zone s t i l l  the  salinity  reached  a t Saa0.8 mL  7 and  (appendix  3  by an o u t f l o w  waters introduced d u r i n g  t o J u n e , oxygen  August,  and  than  by an e x c h a n g e ,  been b a l a n c e d  and d e n s i t y  persisted  the  75  b o t h months  direction  had  t o be 2.9  d e p t h of 190 m w a t e r c o n t a i n e d  in  m  from  1983).  i n the l e v e l  Oxygenated  April  75  t o i n t r u d e a s f a r a s Saa0.8 i n December  time.  depth,  16). D u r i n g  i s not a f f e c t e d  (LeBlond,  salinity  1985,  (table  conditions  estimated  have  annual  increase  between  I t seems p r o b a b l e  of i n f l o w i n g  level  would  dense water  displaced  displaced.  and Saa9 t h e volume  Since  140 m.  forced  Based on t h e change  September  an  L~ .  2  minimum oxygen c o n c e n t r a t i o n s a t recorded  w i t h water c o n t a i n i n g  have  26). At a  above  (table  this  1 0 ) . From  deeper  depths.  and below  180  conditions been  in  m  were  a  small  or from the d i r e c t i o n  of t h e  31  sill  preventing  summer  (figure  At  Saa9 levels  month  (figure  mL  1  at  than  i n 1985  140  m  (table L"  t o 0.20  mL  increased  again  in  m  L" )  a t 170 m  1  depth  August  demonstrated  was  water had The  deep  temperature  water,  intrusion  began  contained  a t Saa3  renewal  in  w h i c h was  (figure  the  remained a n o x i c was  situated  L~  oxygen that  had by  13,  as  at  1986  as  to  (0.31  below  a s m a l l volume  intruding of  salinity  mL  this  of dense  mL  1986  high  (figure  high  low  deep  as  1.2  found  in  oxygen  1  L"  1  160 m  seen  intrusion  a t Saa0.8  but  deep  mixed  oxygen  water  i n September, minimum  not a n o x i c  a t 120-140 m  salinity  a t a maximum.  the  oxygen  low  water  the p a t t e r n  25 & 2 6 ) . Low 75 and  by  salinity,  water mL  160 m and -was L"  commenced  w a t e r was  In O c t o b e r , t h e  100 and  v a l u e s were a s low as 0.18  m  t o an  from 140  content  undoubtedly  t o d e p t h s between  between  140  p e r h a p s due  1  i n December u n l i k e  months  a t 140 m.  0.35  and J u n e , b u t  14, 23 & 2 4 ) . The h i g h  September  levels  subsequent displaced  mL  to  f r o m month t o  level  i n August  l o w e r c o n c e n t r a t i o n s of oxygen were layer  declined  i n May  oxygen  displacement  1985 when t h e h i g h  oxygen  m  August.  to decline  from  rose  0.75  indication  and  the basin  Water  to  characterized  throughout  i n December  early  of dense, oxygenated water.  in early  water  1986,  They  8 ) . The h i g h e r  further  intruded  140  t h e p r e s e n c e o f an oxygen minimum  (table  a  September  and  l e v e l s were a s low  i n J u l y . The  1  samples c o l l e c t e d  200  120 and  oxygen 7).  o f a s m a l l volume  Water  the s p r i n g  but t e n d e d t o f l u c t u a t e  1 6 ) . In A p r i l ,  decreased  intrusion  during  24).  oxygen v a l u e s between  lower  L"  complete s t a g n a t i o n  (table  zone  although  10).  32  At  Saa9,  September. levels of  b o t t o m w a t e r s were d i s p l a c e d  This  of  0.77 mL L ~  0.89 mL L "  volume  displaced  1  was l o c a t e d  of i n t r u d i n g  levels  in  November  water was c h a r a c t e r i z e d  (table  1  the  high  deep  volume  in  salinity  1986 i n d i c a t e d  the  and  incoming  of oxygen-poor  constant  water  1985  from  water p r i o r  w a t e r and  to  o f oxygen  1986  as  may  have  stagnant bottom  indicated  a t 120 and 140 m a t Saa9  low oxygen  the  intruding  smaller  t o t h e r e n e w a l . The deep  i n August  G1748 and G1545  of  displaced  in  o r from t h e p r e s e n c e o f a  water  by  the  and a l s o  zone a t Saa3.  water  and t e m p e r a t u r e c h a r a c t e r i s t i c s  indicate  a  marked  at the hydrographic s t a t i o n s  G e o r g i a and S a a n i c h I n l e t . T h i s exchange  renewal  G1748 & G1545  properties  Inlet,  oxygen  remained r e l a t i v e l y  examination of s a l i n i t y  stations  water  i n the  December,  l e v e l s of oxygen. T h i s  water,  d i s p l a c e m e n t o f a narrow  4.1.2. S t a t i o n  at  in  d e g r e e o f m i x i n g between  started  concentration  An  water  that  higher  from a g r e a t e r  r e n e w a l may have higher  minimum  a t 120 m. Owing t o t h e d e c l i n e  waters  i n 1986 c o n t a i n e d  waters  oxygen  t o December.  intrusion  resulted  by low  7 ) . In O c t o b e r , an oxygen  A c o m p a r i s o n o f t h e deep  water  t o a d e p t h o f 75 m i n  between  and t h e i n f l u e n c e fresh  can  the  be  Strait  of the P a c i f i c  Bathymetry  and  water  contributing  to differences  supply  between  W i n t e r months, r e p r e s e n t e d  similarity  i n the S t r a i t of  largely  attributed  of G e o r g i a Ocean are  in  to the  to  and S a a n i c h both  main  areas. factors  t h e two a r e a s . -  by F e b r u a r y , were  characterized  33  by  cool  temperatures  approximately replaced  the  approached, but  the  warmer  layers  fairly High  coast  1983;  Pickard  salinity  & Emery, at t h i s  the i n t r u d i n g but  high  depths at s t a t i o n were  (figure  28  The entire  similar  (figure  the  the  water  October.  column off  1962;  the  LeBlond,  at the S t r a i t  of  w a t e r s c o n t i n u e d t o move i n salinity  were  f o u n d a t much  temperatures  and  range  bottom  water  at  the  deeper  densities  t o t h o s e a t Saa9 s u g g e s t i n g t h a t the  until  t o v a l u e s a t Saa9 d u r i n g  G1545. A s s o c i a t e d  replacing  i n the  Salinity  upwelled  to a r r i v e  8°C  gradually  27).  (Herlinveaux,  waters  Strait  of  (figure  in February  G e o r g i a remained 30).  A  1985  levels  minimum  f r o m samples  present  in  t o December, c o n t a i n e d  V a l u e s much l o w e r t h a n t h i s  Saanich  depths,  presumably  started  salinity  insolation  both  Pacific stations  & 29).  Oxygen  September m.  was  year  measured 11).  was  summer  uniform within  9°C by  of  winter  As  increased  above  1985  time. Saline  also very similar  Ocean water  during  r e a c h e d a maximum v o l u m e . The  water  depth  d e c r e a s e d the s a l i n i t y  t o August  1982),  a  March.  a t deeper  throughout  waters,  April  d u r i n g December and  River  t o September  constant  from  station  renewal  in  fairly  reached l e v e l s  to  cooled  waters  remained  the F r a s e r  from May  west  of  surface  Water  o f t h e water  summer and from  September.  28).  bottom  depths  temperature  Input  remained  (figure  the  s h a l l o w d e p t h s were warmed by  rose during  Georgia  m  deeper  r a n g e . The  upper  200  from  Inlet  during  the  oxygenated of  2.77  throughout  mL  collected  a minimum o f 3.35  renewal  both  as a r e s u l t  L"  2  a t 400  t h e deep water  were f o u n d a t  0  was  1  m  (table  exchanged mL  L~  the  from  at  300  stations  in  1  of m i x i n g w i t h  34  resident  low  oxygen-containing  Similar 33).  The  winter also  s e a s o n a l t r e n d s o c c u r r e d i n 1986  water  mass  4.1.3. S e c h e l t Sechelt and  f o r the year  Inlet  February  November  was and  1985.  August  m  was  Saa9 but  o n l y by  The  2.40  mL  L"  1  where t h e y In  between fell  Sc1  Salinity  very cold  In Sc2  and  a t 300  1  figure  and  &  early  m which  was  34).  1986  Inlet  throughout  Hydrographic was  1986,  Both  293  fresher  t o be  L  - 1  remained  m  (figure the  of G e o r g i a  the  at a l l  a t Sc1  than  than  that  as a  result  were  above  t o Saa9 and  Saa0.8  levels  f e a t u r e s were measured a t  meters than  36).  above  the  Saa9, b a s e d  bottom).  on M a r c h  Temperatures  were  1986  fairly  water column w i t h t h e e x c e p t i o n o f  at the  surface below  physical  stations  1985,  .  (9  lower  Sc1  lower  in contrast  hydrographic o f 240  at s t a t i o n  somewhat c o o l e r  m  mL  were  (20.8-28.7 p p t )  and  250  m)  parameters  Saa9 i n November  3 5 ) . Oxygen  l e v e l s d i d not d e c l i n e August  t h r e e o c c a s i o n s , November  (figure  below 0.65  temperatures  Sc2a.  L"  2  or the S t r a i t  v a l u e s were s t i l l  homogeneous  oxygen  150  to a depth  for Saanich  1986.  density  Inlet  salinities  February  station  data  <1°C.  Saanich  lower  31,32,  G1545 i n f a l l  12,  on  generally tended  the  (table  0  (maximum d e p t h  Temperatures  of  mL  sampled  depths.  in  3.15  Compared t o s t a t i o n  w a t e r column a t Sc1  measured  into  (figure  Inlet  measured down t o 250 in  intruding  c o n t a i n e d a minimum o f  t h e minimum  1985,  waters.  2.75  (figure mL  L"  1  36).  Dissolved  (figure  d a t a were c o l l e c t e d  showed s e a s o n a l warming  at  36). stations  i n the  upper  35  50 m and a s t r o n g d e n s i t y  stratification  (figure  water was p r e s e n t a t t h e s u r f a c e a n d maximum 28.8  ppt  mL L "  1  at either  With Sechelt L" )  occurred at depth. station  did  August  sampling  maximum  f r e s h water  remained  lower  the S t r a i t  periods  obtained Georgia,  from  develop  areas  their  did  stations  Saa0.8  and  Saa9,  low oxygen c o n d i t i o n s (<1.0 mL  not  Analysis  vertical Inlet,  Saanich  distribution  than  either  and  t h e time of column  Saanich  Inlet  CIV, net  CV,  specimens  were  Samples  and  hauls  CVI  collected  i n the S t r a i t of  and S e c h e l t I n l e t . The main o b j e c t i v e o f  was t o compare c o n c e n t r a t i o n s o f c o p e p o d s from t o determine  factor  distribution  with  1961) y e t t h e water  of Net Haul  i f t h e r e were s i g n i f i c a n t  a b u n d a n c e . The h y d r o g r a p h i c  a possible  coincide  and d e n s i t y  plumchrus  Saanich  sampling  three  in  Inlet  of G e o r g i a .  Neocalanus  in  below 2.40  37 & 3 8 ) .  run-off (Pickard,  in salinity  4.2.  the  around  d u r i n g w i n t e r and summer months m e a s u r e d . The F e b r u a r y  1  or  (figure  not  salinities  The oxygen d i d n o t f a l l  reference to Saanich Inlet  37 & 3 8 ) . F r e s h  contributing  and  density  Inlet  were  parameters  to the  observed  o f N. p l u m c h r u s . used  of the s p e c i e s .  to  differences  were examined a s differences  Horizontal  determine  the  the  in  net hauls vertical  36  4.2.1. S t r a i t  The  of G e o r g i a V e r t i c a l  vertical  distribution  plumchrus  i s reported  Georgia.  Seasonal  changes  reported  in earlier  papers  The depth  Strait  Hauls  and c o n c e n t r a t i o n  f o r e a c h month sampled in their  of G e o r g i a  overwintering  depths.  The A u g u s t below  of  dropped  half in  sampled  copepods  significant  n e t was u s e d  few  i n September  replicates  i n August  (figure  13; f i g u r e inhabiting the presence number  to approximately  40). This  decline  1986 a n d was c o n s i d e r e d a 1 9 ) . However, t h i s may  were c o l l e c t e d ,  1985, a n d t o t a l  than  one c o p e p o d n r  3  a different  numbers o f CVs  was c a p t u r e d  i n O c t o b e r . I n November, t h e upper attempt  to  determine  the  100  m  be SCOR  increased  i n t h e upper  was  sampled  150 m in  an  d e e p e s t d e p t h a t w h i c h N. p l u m c h r u s  could  n o t be f o u n d . No c o p e p o d s  were c o l l e c t e d  0-100  m sample  0.2 n r  b u t i n December,  3  in  the  were c a p t u r e d  November  i n thetop  m. In December, t h e t o t a l  they  a  i n O c t o b e r and November. Less  100  to  3 9 ) . The  i n September  d e c r e a s e a t p=0.07 ( t a b l e  coincidental;  again  occurred  1985  already  150 m ( f i g u r e  of  1973).  (table  samples v e r i f i e d  t h e maximum A u g u s t c o n c e n t r a t i o n numbers a l s o  was  Strait  a r e evident as  in July  was f o u n d  the population  of a deep w a t e r p o p u l a t i o n overwintering  distribution  was f i r s t 3  suggested that  the  ( G a r d n e r , 1972; F u l t o n ,  o f 150 m where <1 c o p e p o d nr  39). T h i s  in  of Neocalanus  were  showing  signs  number o f c o p e p o d s of  maturation  collected  i n J a n u a r y and F e b r u a r y c o n t a i n e d  of w h i c h  were  females  visibly  collected  reproductive  i n March  contained  had d e c r e a s e d and  (figure  4 0 ) . Samples  healthy adults,  (table  1 3 ) . Some  visible  oviducts  many adult  b u t many  37  were s p e n t and p e r h a p s the  overwintering  due  to  death  p h a s e . The  obvious  of  and  and  upon c a p t u r e . The  population adults  difference  population  dead  from December  CVs  therefore  surface  and  it  juvenile  layers  occurrence  of  was  comparison  total  summer of  1986  (April  November  1985),  between s e a s o n s collected  season e f f e c t population In CIVs  The  overshadowed  1986,  broad  80%  tissue  an a d u l t (table  An occurred  was mean  80 m  samples  since  still  to  difference primarily  The  the mean  different. was  predominated  varied  haul  (August  effect.  of copepods  (table  A and  t h a n G1545 so p e r h a p s  G1545  i n the s i n g l e  the  month.  were  at  250  i n the  i n the s p r i n g  by t h e 83% c o e f f i c i e n t  female which  arrived  following  population  CVIs were f o u n d between 80 and  was  test  March  station  confidence interval  captured  to  a significant  significantly  the r i s i n g  indicated  and  plumchrus  by a  concentration  as  i n the f a l l  likely  the  fall  15).  The  reproducing  to the p r e c e d i n g f a l l  o c c u p y i n g t h e upper  copepods one  during  G1748 r a t h e r  between months was  April  replicates  and  station  the  necessary  demonstrate  18).  reproductive  at depth  February  primarily  table  the  not  (table  was  in  to July)  probably  variance.  numbers of N.  did  from  of  appeared  of  not  in  their  of  c o p e p o d i t e s most  starting  CIVs  density  overwintering  d i f f e r e n c e s w i t h an a n a l y s i s Nauplii  t o M a r c h was  as t h e y c o m p l e t e d  between t h e  the  steady d e c l i n e  (figure greatly  by 39;  among  of v a r i a t i o n  and  1 4 ) . Some C I V s ,  CVs,  m and  below.  from 250  contained  Of  m to the eggs  and  the  few  bottom, muscle  15).  unexpected i n May  1986.  decline  in  the  total  number of  copepods  E i g h t y - s e v e n p e r c e n t of t h e t o t a l  copepods  38  were c o l l e c t e d  i n t h e t o p 80 m ( f i g u r e  39). A  of  the copepods  were C I V s  the  d e e p e s t zone  t h e y were a l l CVs w i t h  adult  percentage  t o t h e p r e v i o u s month. I n the  exception  of  one  female. The  the  i n contrast  small  concentration  numbers c o l l e c t e d  samples  with  the  o f N. p l u m c h r u s  i n May. O n l y  majority  f o u n d i n June  CVs  occuping  were  present  depths  below  maximum c o n c e n t r a t i o n s o f 80.6 n r  3  situated  Concentrations  3  still  averaging  10.0 n r  surpassed in  200 m w i t h  i n t h e bottom  existed  June  95  m.  i n t h e u p p e r 80  m (figure 39). The was  a n n u a l maximum was r e a c h e d i n J u l y  virtually  absent  increased  between  averaging  125.2 i r r ,  comparable  August  declined  a v e r a g e t h e 80% c o n f i d e n c e samples  fell  number o f c o p e p o d s  4.2.2. S a t e l l i t e Satellite southeast, the  within  Channel  Stuart  Inlet  m.  N. p l u m c h r u s  Concentrations  concentrations  of  water. in the  August  to levels  apparent  decrease  interval  calculated for  the range of the J u l y (table  declined  CVs  1 4 ) . As  i n September  replicate  mentioned  the  (table 19).  Channel  south ( f i g u r e  Saanich  150  somewhat  to the previous year. Despite  samples w h i c h were much b r o a d e r total  top  were f o u n d i n d e e p  numbers  from t h e J u l y the  the  150-250 m b u t maximum 3  Copepod  from  1986.  receives  water  from Haro  Strait  to the  C h a n n e l t o t h e n o r t h w e s t , and S a a n i c h I n l e t t o 1).  Copepods  transported  must p a s s t h r o u g h t h i s  into  or  out  c h a n n e l . N. p l u m c h r u s  and CVs were c a p t u r e d a t t h e 75 m S a t e l l i t e  station  only  in  of  CIVs May  39  1985, 1985  a n d , A p r i l and May  considered assumed  captured  1986  were  these  cycle.  caught  N.  an  indication  of a higher  Inlet.  (north),  Saa3  central  station,  of  station  Saa3  Saa3,  are  1985 a n d  collected  in  To d e t e r m i n e  or l o s s  of copepods  t o sample d u r i n g the high  each  concentration  i n 1985.  samples were  Saa0.8  (south;  was  the only  Saanich  appeared  present to  (May,  July,  be h i g h e r  the  and August, (table  population  than  in  1986.  acceptance of water.passing  of  sampled  in Saa9  5 ) . The  station  t h a t was  ( f r o m May  through  station  than  during  i n 1986. The  N. p l u m c h r u s  at  1985 a n d 1986) were n o t  20). in  collected  figure  at this  and month on t h e abundance  higher  were  typically  and  a t t h e 5% l e v e l  significantly  i n May  tide.  few months o f t h e p r o g r a m  1985  (p=0.09)  increased  population  1985). The p o p u l a t i o n  of y e a r  significant  flood  tide  1985 s a m p l e s c a n n o t be c o n s i d e r e d  stations  (central),  effects  level  flood  those  a net gain  horizontal  Three  i n the f i r s t  summer  the  ebb t i d e a r e  Inlet  Saanich  the  during  i n May  Both v e r t i c a l and  August,  whereas  t h i s was n o t p o s s i b l e ,  plumchrus found  4.2.3. S a a n i c h  to  tide  i t would have been n e c e s s a r y Since  of  t h e ebb  numbers r e p r e s e n t e d  the i n l e t ,  sampled  caught d u r i n g  t o be i m m i g r a n t s . The s a m p l e s c o l l e c t e d  whether  tidal  numbers o c c u r r i n g i n  i n S a t e l l i t e Channel d u r i n g  as e m i g r a n t s and those  c o i n c i d e d with  April  to  greater  (figure 41). Copepods  1986  1986, w i t h  However,  1985 This  can may  through  at be  be  the  10%  considered due  to  an  t h e SCOR n e t u s e d  40  from  May t o A u g u s t  captured  with  The  Clarke-Bumpus n e t s  distribution  indicated  that  the  characteristic that  they  of  tended  horizontal  hauls  numbers o f  33.3  extrapolation Saa0.8 at  1985 a l t h o u g h  at  were  were  3  showing  depths.  were p r e s e n t  (table  at  150  available  m  except  In May  found  for stations  1985, maximum  16).  An  Saa9 and  of t h e water  0.0 a n d 0.70 mL L " . A t d e p t h s o f 5 1  b u t o n l y CVs were c a p t u r e d  hauls demonstrated  i n the upper  depths.  t h a t t h e oxygen c o n t e n t  between  Vertical  of Georgia  sampled,  o f t h e oxygen d a t a  140 m i n d i c a t e d  patterns  that of the depths  50 m some CIVs were p r e s e n t  these  t h e summer o f 1985  migrational  at shallower  revealed irr  in  species i n the S t r a i t  to aggregate  also  i n 1985.  N. p l u m c h r u s  CVs  this  Saa3 was p r o b a b l y  and  of  d e n s e c o n c e n t r a t i o n s were  120 m most  that both  below  C I V s a n d CVs  o f w h i c h were  CVs  (figure  43). In  July  concentrations the  haul  from  However,  1985,  the  o f 7.2 CVs n r  vertical 3  haul  i n t h e t o p 150  samples m.  the  horizontal  hauls  indicated  i n t h e 0-150 m v e r t i c a l  meters.  Samples  from  50,  that  most  haul,  around  150  contain  any CVs, b u t , c o n c e n t r a t i o n s o f 75.6 i r r  were  of  the  situated  100, a n d 120 m d i d n o t 3  were  found  at  m ( t a b l e 16). Ten  C-B n e t s irr ), 3  Unfortunately,  t h e b o t t o m o f t h e w a t e r column was n o t s u c c e s s f u l .  copepods c o l l e c t e d  150  contained  discrete  a t Saa3 i n A u g u s t  was  found  concentrations decreased  d e p t h s between  at  m  o f 0.11 mL L " 1  with  1985. The h i g h e s t d e n s i t y o f CVs (6.0  160  t o 0.02 mL L "  110 and 190 m were s a m p l e d  1  (table  16  were p r e s e n t  a t 185 m ( f i g u r e  &  figure  a t 165  m,  1 7 ) . Oxygen and  they  1 7 ) . No c o p e p o d s were  41  captured  from  170, 180, and 190  m.  S a m p l i n g a t Saa9 and Saa0.8 commenced Very  few  copepods  were  captured  September  a t any o f t h e t h r e e  stations  following  numbers  found  expected  t o have been f o u n d . Copepod c o n c e n t r a t i o n s o f  were  captured  Saa0.8  (figure  Saa9,  however,  at  not  during  renewal.  Based  a  single  o f 0.2 n r  (figure  were  3  CV o r a d u l t  w i n t e r months  Of t h e few s p e c i m e n s  Saa3 one s e x u a l l y  (table at  1 5 ) . An a d u l t  Saa3 and one a d u l t Low  at  was  IV  45; t a b l e  (figure  0.2  nr  3  of  station  occasionally total  station. until  The  January  at station  Saa0.8  each  station  f e m a l e c o p e p o d was c a p t u r e d  again  i n March  male was c a p t u r e d  found  at  at  Saa9, Saa3, and Saa0.8  the  3  13 & 1 5 ) .  at either  3  m a t u r e f e m a l e was  t o p 80 m. H o r i z o n t a l  some  irr  43 & 4 4 ) . The  (0.2 n r ) c a p t u r e d  and V c o p e p o d i t e s c o l l e c t e d  the  0.1  found a t  in April  a t Saa0.8.  numbers of C I V s and CVs began t o a p p e a r  stations  were  collected  s p e c i m e n s d i d n o t show s i g n s o f s e x u a l m a t u r a t i o n  and  numbers  Saa3 a n d Saa0.8, N. p l u m c h r u s was  number of c o p e p o d s d i d n o t e x c e e d  1986.  copepod  90 m a t Saa3 and none were f o u n d a t  to February  the  1985. Saanich  on  Saa3 i n p r e v i o u s months, h i g h e r  43 & 4 4 ) . A t o t a l  stations  captured  water  i n the bottom  Saa9 from O c t o b e r At  t h e deep  in  (figure  in April  April  41; t a b l e  1986  15). Stage  1986 were l o c a t e d  tows a t Saa0.8 i n d i c a t e d  C I V s were p r e s e n t  in  that  within  at  least  a t shallow depths of 5 m  (table  17) . Copepod numbers stations. and  lowest  increased  Concentrations at  concentrations  Saa0.8 were  i n May  remained (0.2  present  the highest  nr ; 3  at  1986 a t a l l t h r e e  figure  a t Saa9  41).  Saanich (1.4 n r ) 3  Intermediate  S a a 3 . B o t h C I V s and CVs, were  42  primarily  l o c a t e d i n the  In J u n e copepods had  1986,  (figure  the  43  upper  m  vertical  station  & 4 5 ) . At  r e g i o n of  of  the  at  of  1  0.37  mL  The located  station  a t Saa9  dissolved  m the  Despite N.  plumchrus  Inlet, i n an  the  N.  two  of  plumchrus  100  levels  obvious  to detect  in  1986  depths.  between Saa9 and  170  m  were  1.23  and  in  months, d e t e c t e d  with  20%  of  top  120 in  very  few  some  CVs  concentrations  found  i n 1986  suspected the  the  The  CVs  collected mid  0.10 mL  mL  L"  in  the  two  Saa3  depth)  where  1  (table  size  10).  the  of  and  the  Saanich  July  1986  months.  The  the primary  1985/1986.  the  at  3  Saa0.8  J u n e , and  between  x  nr ,  at  of G e o r g i a  i n May,  (station  7.1  1  between  t o be  was  L~ .  overwintering  between  25).  copepods  Strait  trends  G1545  the  (the  t o 0.07  the  negligible  difference  The  and  since  variance  (p=0.02)(table  that  above oxygen  plumchrus  difference  was  Inlet  months  was  they  were a l s o f o u n d  indicated  4 5 ) . A l l of  any  Saanich  significant  N.  r e g i o n s were compared  Georgia  i n the  where maximum c o n c e n t r a t i o n s of  population  attempt  Strait  of  oxygen d e c r e a s e d  the  s t a t i o n s but  Approximately  G1545 some CVs  m depths,  (figure  between  three  of  46).  in July  oxygen  180  110  at overwintering  were p r e s e n t  concentration  w a t e r c o l u m n . T h e r e were s t i l l  density  below 80 m  also  Below  and  (figure  highest  occurred were  L"  90  The  Saa9 were f o u n d  c o p e p o d s a t Saa0.8, h o r i z o n t a l h a u l s were p r e s e n t  43).  distribution.  Saa3 and  the  (figure  were p r e s e n t .  i n c r e a s e a t any  in their  the copepods a t m  o n l y CVs  d i d not  changed  upper 80  source  population An  month)  in  a n a l y s i s of indicated  s t a t i o n s ( p = 0 . 0 0 l ) and  homogeneous s u b s e t s  S c h e f f e , B o n f e r r o n i , and  of  among  minimal  a the the  tests  43  demonstrated July  (table  abundance steady of  similarities  i n May  abundance  single  haul  17).  significantly  The  to  v a l u e of z e r o was  (i.e.  <0.2  m;  table  3  that  therefore  concentrations  considered  significantly  this L"  1  the  17  hypothesis  means  region. although  the c o l l e c t i o n  Saa9,  August  (<0.1  taken to  equal  t h e oxygen was  1986  (<0.5  sampling 3  n r ) b u t had r e a c h e d 3  population  differences  significant  a t Saa3.  a  have c h a n g e d  May,  150  at  the and  145 m a r e above  days  and  140 m  was  prior  to  slightly.  increase  numbers were s t i l l  between  that  few c o p e p o d s were f o u n d a t  1 5 ) . The  maximum  were  results,  and  1.5  150  negligible  in dissolved  some w a t e r movement had  a t Saa9. P o p u l a t i o n  and  rejected  oxygen c o n t e n t  very  n r ; table that  was  140  measured  of the  Note  than c o n c e n t r a t i o n s  dissolved  the  i n t h e 135 and  been  at  test  depths  27).  filtered  has  copepods  greater  The  (table  a  i n J u l y at  difference  2 7 ) . B a s e d on t h e s e  are  were  26).  among  not i n c l u d e d  of water  &  of  oxygen a t 140 m s u g g e s t e d in  Saa9  in  demonstrate  used  significant  o f c o p e p o d s and may  During August station  at  (table  was  a t t h e 5% l e v e l  and  increased  not  tows was  ANOVA  June  i n J u l y . A comparison  concentrations  s e t s b e c a u s e t h e volume  mL  had  did  t h e months  a  and  t h e numbers  July)  for  m data  0.20  copepods  horizontal  mean  different  replicate  below  June,  increased  factor  samples  and  o f N. p l u m c h r u s a t f o u r d e p t h s (135,140,145,  table  one  the  between  of r e p l i c a t e  May  whereas,  (May  difference  Saa9. A  horizontal  to July,  G1545  series  station  G1545  and J u n e and t h e n  and  significant A  At  f r o m May  Saa0.8  m;  25),  between  at  Saa3.  occurred  low a t  Saa0.8  However,  the  June and A u g u s t were n o t  44  The  majority  indicated The  by  highest  of  copepods  both v e r t i c a l  concentrations  (figure  18).  The  followed  by  increase  an  The  p r e s e n c e of  that  a  small  an  at  The  amount of  for  hypothesis at  each  (table least  the  months  The  over  (table  separated  22  into  & 23).  two  reaffirming  that  significantly  lower.  The  lack  Saa9, weakened not not  (table  of  Saa0.8,  the  at  In  .each  1  depth.  indication August.  No  between  the  three Saanich  Inlet  to  August N.  The  null  p l u m c h r u s was  equal  was  1986.  rejected  analysis  Saa9 the  since  inlet.  in either and,  Each  p=0.07  between  was  was  difference difference were  the  stations  were  the at  Saa3  10%  and  as  stations  level,  Saa0.8  were  especially  between t h e  factors  the  Saa0.8  at  and,  be  and  numbers  stations  to  station  i f the  Saa3  station  it  c a s e . The  subsets  the  at  Saa0.8 were s u s p e c t e d  ANOVA t e s t s . A d i f f e r e n c e  significant difference  24).  an  L~  2  water  both comparisons  d e t e c t e d among S a a n i c h a  0  m  volume of  population  replication  mL  this  in early  to determine  homogeneous the  0.31  180  low  and  greater  For  and  43).  the  mouth of  was  similar  was  Saa3  the  Saa9 and  170  be  as  to  l e v e l s at  Saa0.8 s e p a r a t e l y  between s t a t i o n s  m may  months t e s t e d ,  from the  stations  170  m  17).  against  between  was  the  at  170 (figure  below  intruded  April  stations  station  oxygen  mean abundance of  population  s i m i l a r to  tested  of  samples  contained  m due  analysed  i n which the  21).  furthest  (table  was  station  m  in dissolved  d e n s e water  t h i s depth  below  were p r e s e n t  170  190  were  horizontal  oxygen minimum a t  variance  stations  Saa3  CVs  at  s p e c i m e n s were c o l l e c t e d a t filtered  and  of  water  at  months,  expected,  there  Saa3 and  Saa9  were  tested  for  45  h o m o g e n e i t y of null  h y p o t h e s i s was  between  stations  this  test  for  stations  at  was  present  Saa3 and  limited  the  Bartlett  at  the  Saa0.8  since  Saa9 and  at  below  100  the  5%  (table  replicate  Saa3  oxygen  At  chi-square level  23).  test.  for  test  However, t h e  exception  of  The  the  samples were not  ( w i t h the  and  this  of  the  the  use  of  available  August,  1986  Inlet  During  November,  than  Sc1  Inlet  adults  had  due  their  The among the  was  (table  15).  3  were  3  located  a l l s p e c i m e n s were p r e s e n t 33%  44).  of  low  (0.27  The  above  nr )  were  3  concentration  of  0.16  as  mL  L  "  oxygen b o t t o m water  1  during  10).  Saanich  in  throughout  decline  numbers  Inlet  contrast  to  the  2 nr  after  (figure  to  16.9  the  winter.  nr  between 80  February  population  48;  few In  and  table 3  and  inhabited  13  &  were f o u n d copepods  February, 245  the m,  1986  15).  between  found  in  number  of  presumably  reproduction.  collected  stations  3  1985  overwintering  c o n c e n t r a t i o n s of  decreased  three  i n November  a much l a r g e r  Sechelt  Saanich  nr )  m" )  Inlet  that  100-245 m a t  (<0.2  (figure  (table-  (0.2  Saa0.8  remaining  displacement  numbers c o l l e c t e d  indicated  Saa9  of  zone  renewal  4.2.4. S e c h e l t  to  Saa3 and  station,  minimum zone r a n g e d between 0 and  d e e p water  The  copepod d e n s i t y  southernmost  minimum  i n the  highest  Saa0.8, 67%  below  result  the  stations  m.  oxygen  captured  the  rejected  September  Specimens at  a  using  Saa3). During  the  variance,  i n August  sampled  (figure  1986 48  varied & 49).  considerably The  highest  46  number  of  CVs  Disregarding  (32.2  nr ) 3  the depth  meter c o l l e c t e d  range  a t Sc1a was  were  found  sampled, still  at  Sc1a ( f i g u r e 4 8 ) .  the t o t a l  much h i g h e r  number p e r c u b i c  than  either  Sc2a,  or S c 2 .  4.3. A n a l y s i s o f E x p e r i m e n t a l Live the  s p e c i m e n s o f N e o c a l a n u s p l u m c h r u s were c o l l e c t e d  intention  different observing  of examining  oxygen  a  lot  pellet  starved  copepods.  few  were  questions  tests  from May  with  starved  in  the  number o f f a e c a l two  groups  through  collections, 12  hours  a  the e x c e p t i o n of for  some  observational  by  while  conducted  on  of f e d v e r s u s  pellets  was c o u n t e d  30  copepods  were f e d e q u a l q u a n t i t i e s o f 29 f a e c a l  copepods  a n d June  f o r 8 days.  of  the  pellets.  occasion  September  1987  Ingestion  inspection  of f a e c a l  on one  from  o f 1986 were  October  purposes  an  f o r the presence  total  rate  t o November  flasks  of  arose  experimentation.  were  and t h e m o r t a l i t y  f o o d by t h e c o p e p o d s was v e r i f i e d  debris  under  c o u l d n o t be d e d i c a t e d t o a d d r e s s i n g  c o p e p o d s w h i c h were o n l y h e l d i n c a p t i v i t y of  species  with  of F e d and S t a r v e d Copepods  CVs c o l l e c t e d  fed phytoplankton  this  and d u r i n g  preliminary  production  4.3.1. O b s e r v a t i o n s  which  time  of  Many  in captivity  of  these questions, a faecal  the s u r v i v a l  concentrations.  the copepods  Although,  The  Data  in  and  The  which October  of  phytoplankton.  After  pellets  were c o l l e c t e d ,  83% o f  47  which  were f r o m The  and  t h e September  number o f c o p e p o d  starved October  months  (figure  specimens,  50).  were n o t  Each  Figure  50  copepods  fed died  shows  a  contrast,  in  their  contained  at a rate  group  followed  mortality  after  which  c o p e p o d s • w e e k " . The  number of dead  day  remained  until  (figure  of  two  a t the  i n October week . - 1  i n t h e number o f  by what a p p e a r e d  to  the  In  55th  copepods  t h e 62nd day  fed  of  2.05  were f e d d i e d  0.34  more d i e d  f o r both  30 c o p e p o d s  constant increase  specimens  unchanged  captivity  recorded over a p e r i o d  in captivity  1  28).  p e r i o d . Copepods c o l l e c t e d  fairly  the October  was  thermos  d y i n g i n the s t a r v e d  be a s h a r p r i s e  (table  deaths during  b e g i n n i n g of the r e c o r d i n g which  thermos  day.  at a rate  after  a t which  the  time a  of  12th two  50).  4.3.2. Oxygen T o l e r a n c e T e s t s Three of the i n i t i a l run  for  levels the  a  period  (0.45 mL 24  concentrations died. to  o f 24 and  L" )  period (0.83,  0.60  tolerance  21 h o u r s  i n experiment  1  hour  low oxygen  #1,  (table  5).  mL  80 and  1  At  higher 90%  S e v e n t y - t h r e e p e r c e n t of the copepods  0.91  mL  L"  1  for  21  hours.  The  5 ) . At  a l l copepods  (table L" )  experiments  number  subjected  of  i n t h e c o n t r o l s was  #4  i n which The  10% d i e d  results  (table  of the  oxygen  when  mortality  0%  mean  died  #1,  experiment  during  copepods  experiment  in  died  the  a t t h e t i m e of e x p e r i m e n t a t i o n r a n g e d days  oxygen  of  captivity  t o 45  low  were  #4  days from (table  i n a l l cases except  spent  in  12 d a y s i n 5).  The  experiment  5).  12 hour e x p e r i m e n t s were  variable.  The  48  thirteen Strait CVs  experiments  conducted  on  CVs c o l l e c t e d  o f G e o r g i a w i t h t h e e x c e p t i o n of two oxygen t e s t s  collected  low  were  from S a a n i c h I n l e t  in July  oxygen c o n c e n t r a t i o n s (0.46 t o  higher  2.11  t h a n 70% were r e c o r d e d ( f i g u r e  mortality  was  12% f o r oxygen t e s t s  L " ) and 0% f o r oxygen  levels  1  variability  i n the r e s u l t s  run  on  1986. A t b o t h h i g h and mL  L~ ),  mortalities  1  51). Likewise, the percent i n t h e lower  of 0.95 mL L "  may  i n the  range  (figure  1  be due t o t h e l e n g t h  (0.56 mL 51).  of  The  captivity  ( f i g u r e 52). Figure  52  indicates  increased  as the p e r i o d  to  days  103  experiments  in  (table  captivity  collected  8  days  collection.  that  mortalities, in  1986  collected  were  experiments yet they  t h e outcome o f t h e that  from t h e S t r a i t  used were  of days (figure  #13,  died  containing  i n water  oxygen c o n t e n t was  the  to  lower  i n experiment  were  the  3  in  experiments  from  where  within  t h e number  CVs  depth  in  3  conducted  resulted  L " , y e t , i n Saanich I n l e t , a  copepods  were r u n a t oxygen  the hypothesis that  1  2  they  1987 and were t e s t e d  tested  I t i s important t o note  copepods mL 0  influenced  of t h e  i n c r e a s e d . Copepods were  three experiments  These  supporting  captivity  51).  i n June  were p r e v i o u s l y  mortality  before  5 ) . The l a s t  specimens  levels  the  of c a p t i v i t y  on  of  that  50% o f t h e  collected  in  0.20 mL  0.47 July L"  1  L"  1  ( f i g u r e 47). The had in  Saanich Inlet  specimens  the lowest m o r t a l i t y captivity  concentrations  tested  f o r the time p e r i o d  (figure  52).  Overall,  were e x p e c t e d t o r e s u l t  with- a p r o g r e s s i v e  a t 0.56 and 0.54  increase  that  t h e y were  the  lowest  i n the h i g h e s t  i n t h e number  mL  held  oxygen  mortality,  d y i n g a s t h e p e r i o d of  49  captivity  increased  (figure  53).  differences  i n t h e t i m e of y e a r  which  the  CVs  Strait  o f G e o r g i a ) may  experiments.  In  a d u l t s and may 10  (table  i n most o f t h e control  L" .  The  1  The  ranged  Sediment and  collected  trap  from  specimens,  specimens  found  17th  samples  June  the  collected 1985.  The  N.  water  copepods  t o 57  the  t o moult  into  of e x p e r i m e n t s 8,  9 and  concentrations  Saa9.  captivity  Trap  1986.  i n June  In  1985,  were c o l l e c t e d  died these  from  135 m  and J u l y  not  140 m,  November  neither  were  3 1985.  found  next  in July  whole  samples  o f t h r e e CVs  sediment  t h r o u g h September  at a depth of  to  were r e c o v e r e d f r o m  deep water  was  had  from M a r c h t o  180 m t r a p between A u g u s t  plumchrus  three  Samples  A t Saa0.8, a t o t a l  21  of  100%  days.  plumchrus  and  prior  of  was  of the copepods  were examined  i n the samples  collected  of  of t h e c o p e p o d s  30%  of  t o September  mid  i n the  started  oxygen  rate  20 and  Sediment  s e t o u t between May  samples  dead  10,  from 31  at station in  was  survival  o r p i e c e s o f N.  collected  traps  CVs  the r e s u l t s  duration  4.4.  1985  the  t h e outcome  experiments w i t h the e x c e p t i o n  i n which  51).  experiments  influenced  v e r s u s the  52).  12 hour  funnels  (figure  have a l s o November  that  the r e g i o n i n  ( i . e . Saanich Inlet  have a f f e c t e d  mL  suggest  (May-November) and  e x p e r i m e n t s were run a t  t o 6.90  results  captured  late  5 & figure  Control 4.20  were  The  t r a p s . The collected  One 7  were  other and  two from  copepod  September  i n the sediment  trap  1986.  t o t h e d e p t h where  1985,  contained  0.06  the mL  0  2  50  L"  i n May  and  was  devoid  o f oxygen  at  the  1  depth  d e p t h oxygen due  to  water  mL  the  but  i n J u l y . In A u g u s t  1  i n September,  were <0.10  displacement  of  species  of  would  low  settle  oxygen  tissues  in  were f l o a t i n g  120 and  water d u r i n g  low  oxygen  days there  The isolated  had  was  to the  only  collected  conditions  on t h e f a t e  a small  and  of  proportion  remain t h e r e .  e x p e r i m e n t #8,  water  this of t h e  Following  t e n dead c o p e p o d s  o f 24 d a y s  were  (table 29). A f t e r  column.  In f i v e  and on t h e e i g h t h  day  a  2.5  their  gradually  days, four  the remaining four  which  observed  2 9 ) . The up  bottom  deep  were a t t h e b o t t o m of t h e b e a k e r . As  no c h a n g e ,  seven  had  m  were  copepods total  of  copepods  16  did  not  e x p e r i m e n t #9 were  also  surface.  and  increased  the  f l o a t e d to the s u r f a c e . For the f o l l o w i n g  copepods  copepods,  copepods  the  140  a l t h o u g h t h e numbers do n o t  i n t o the t r a p s  at the surface  copepods  (table  at  1  this  1  were decomposed by m i c r o - o r g a n i s m s t h e y  resuspended  float  L"  i n the l a b o r a t o r y ,  for a period  days a l l ten copepods  L " . Above  mortality.  tolerance  i n a beaker  the  the copepods  observations  mL  mL  50 m  concentration  0.41  low oxygen  a f t e r death suggests that  copepods  placed  death  the bottom  t h e oxygen  of t h e s e d i m e n t t r a p was  any a p p r e c i a b l e  A few  the  L~  2  renewal. I t i s p o s s i b l e that  indicate  six  0  concentrations  the  caused  the  0.20  had number  to  the  died  during  after  their  floated of  to  the  beaker.  at the s u r f a c e ,  Of  surface  floating  the  20  and o n l y  Lipids  forming  from  white  dead  a f t e r four at  the  five the  droplets  days  surface  day a t w h i c h t i m e a t o t a l  f l o a t e d t o the s u r f a c e of  the  copepods  eleventh  death.  of  remained specimens of  15 at had  wax-like  51  material. Based that  on  these  preliminary  60 t o 75% o f c o p e p o d s w h i c h may  conditions sediment  in  Saanich  Inlet,  t r a p s because of  percentage  of copepods  on  the  preservative.  Once  a l l of  would have been much  would  lost  relative  effectiveness  sunk  less the  likely tissues  of  not  i t i s possible  to  unfavourable  be c o l l e c t e d  after  4-11  brine  NaN  3  i n the  days.  The  would depend on  t o the depth of the the  to retain  (sodium  without  sodium  trap, azide) azide  s p e c i m e n s o f N. p l u m c h r u s •  had been decomposed,  but the l i k e l i h o o d  greater.  due  due t o r e s u s p e n s i o n  The t r a p s c o n t a i n i n g  would have been  die  resuspension  the depth a t which they d i e d and  observations,  of  the exoskeleton  redispersal  would  have  52  5. The  o b j e c t i v e of  p l u m c h r u s had  a  and  i f not,  1986,  from  plumchrus  other in and  Inlet  and  of  was  Inlet,  for  the  during  some y e a r s .  reduction  in  plumchrus  5.1.  t r a p data  and  the  in  and  distribution  hydrographic s p e c i e s was where  compared  of  to  (LeBrasseur  of N.  their  the  of G e o r g i a .  1969  of  depth On  p l u m c h r u s were  the found  (Hoos,  are  p h y s i c a l and the  1970)  of  the  possible  discussed of  low  chemical  distribution  ontogenetic loss  oxygen  and  overwintering  causes  with  and  migration  for  the  reference  to  experiments.  the Neocalanus plumchrus  population  Inlet  p l u m c h r u s , as was  d e t e r m i n e d by  compared  Saanich  Inlet.  i t s population  i t i s known t o be  assemblage  Strait  Other  hauls,  features  result  t h a t the  Saanich  of N.  h o r i z o n t a l plankton  e t a l . (1983)  of  a  in October  results  P h y s i c a l i n f l u e n c e s on  The  Harrison  species  as  eventual  population  N.  this  1985  population  influence  plumchrus d u r i n g  responsible  during  1982).  tested  Saanich  Inlet  overwintering  the  winter  i f Neocalanus  prevented  inlet.  Inlet  the  (Cowen,  hypothesis  abundance of N.  sediment  in Saanich  f a c t o r s that  fjord  during  1974  characterics  are  the  to determine  substantial populations  December The  the  Saanich  between  Saanich  was  i s a l a c k of an  in  hand,  study  e s t a b l i s h e d i n the  that there  difference  this  resident population  becoming  stated N.  and  DISCUSSION  with The  i n the  Gardner,  physical  abundance of Strait  a p r e d o m i n a n t member o f  e t a l . , 1969;  the  vertical  the  1977;  of  this  Georgia  zooplankton Harrison  et  53  al.,  1983).  The  distribution similar  An  Very  Saanich  i n Saanich  few  population  Inlet  copepods  Saa3 p r i o r  this  time  t o the onset  yet higher  of f a l l .  high concentrations  horizontal  stages  haul  150 m ( f i g u r e  4  (15.0  43 & t a b l e (Sate)  flood  tide  density  comparable  of  16).  taken  CVs  Strait  of  Georgia  sampled  t o t h e bottom; CVs  depth range detected  numbers  had been  the l a t t e r  were  maximum  of  in  the  Samples  top  15  m  3  from  being  and  o f CVs (33.3 n r ) the  d u r i n g an ebb t i d e  also  May  and CVs) were f o u n d i n  Satellite  i n May,  showed  t r a n s p o r t e d out of the i n l e t ;  at  found  in  150  at  August  figure  carried  m  in  July  overwintering (inJuly,  47 &  39).  concentrated  i t  into  is  the  (75.6 n r ) was 3  depths  be  Although,  in a very  attributed layer  o f CVs a t 160 m (table to  1)  in  Saanich narrow August  (6.0 n r ) a 3  1 6 ) . The d e c l i n e i n a  shift  of copepods, e i t h e r  i n the  distribution  of  or  o r 2) some o f t h e CVs may have d i e d due t o a d v e r s e  laterally,  the dense  in July  i n the  s t a t i o n G1748 was n o t  i n a band o v e r  concentration  c o n c e n t r a t i o n than  i n A u g u s t may  found a t  part  ( 1 50-160 m) . H o r i z o n t a l s a m p l e s c o l l e c t e d  a  much l o w e r  3  M a r c h 1986.  ( f i g u r e 42).  to densities  the  nr )  indeed  a t any o f t h e t h r e e  densities  During  t h a t t h e c o p e p o d s were  during  Inlet  is  chemical  was  1985 t h r o u g h  samples demonstrated a l a y e r  station  The  and  N. p l u m c h r u s  and 5 (CIVs  C I V s and CVs were b e i n g  inlet  of  3  fairly  probable  where t h e d e p t h  physical  (<0.4 n r ) were c a p t u r e d  copepodite  that  the  f r o m September  1985,  Channel  but  Inlet  differ.  stations during  at  were a l s o compared w i t h t h e  in Sechelt  Inlet  overwintering  lacking  results  of the s p e c i e s  to  conditions  Saanich  vertically  54  c o n d i t i o n s o r 3) The d a t a  predation.  collected  N. p l u m c h r u s was establish copepods sill in  a  present  of  i n the i n l e t  population.  In  m),  the i n l e t .  so t h a t  the l a t t e r  the copepods  completely  indicated  part  that  sufficient  to  of t h e summer t h e  depths,  well  below  were e x p e c t e d t o be  However, as i n d i c a t e d  three Saanich Inlet  1985  at d e n s i t i e s  had m i g r a t e d t o o v e r w i n t e r i n g  (75  almost  i n t h e summer  the  'locked'  by samples c o l l e c t e d  at  a l l  stations,  the p o p u l a t i o n  of N. p l u m c h r u s  had  disappeared  from t h e i n l e t  by September  oxygenated water  intruding  into  the  coincided  with  the  (table  13) . The bottom  a p p e a r a n c e of dense  w a t e r s of S a a n i c h I n l e t  observation The  oxygen  that  few CVs  profile  were l e f t  during  the  distinct  d i s p l a c e m e n t of oxygen  (figure  12 & 2 2 ) . In f a c t ,  September  I was the  station  originated  transit  suggest that  contained  1985.  The  water  p o o r water  at  t h e minimum oxygen  during  in  16), s u g g e s t i n g of s t a t i o n  the  inlet.  r e n e w a l showed a Saa9  and  Saa0.8  v a l u e s a t Saa9 i n recorded  that  oxygen  t h e water  SaaO.8 and  be  may in  inlet. at least  the e a r l y  following  data  (table  i n t h e water  N. p l u m c h r u s may supporting  deep  from t h e d i r e c t i o n  out o f t h e  inlet  to overwinter  and O c t o b e r were l o w e r t h a n p r e v i o u s l y  values at t h i s have  i n September,  is  have been  part that  o f t h e N. p l u m c h r u s was  N. p l u m c h r u s  based  on t h e d i s t r i b u t i o n  and  forced  s t a g e s o f t h e deep w a t e r a  hypothetical  transported  from t h e s t u d y . F i r s t  that  displaced  can not t o l e r a t e  case  out of the  o f N. p l u m c h r u s  levels in  out of  renewal  in  to explain  how  inlet,  of a l l , i t w i l l oxygen  population  be  < 0.20  August  using assumed mL  1985  L"  1  and  55  July  1986  species,  (figure  including  depths c o n t a i n i n g 1975; will  Raymont,  With  &  47).  calanoid < 0.20 mL  to migrate  level  reference  that  L~  150-160 m a t Saa9 a n d  displaced and  depth of migration  9).  As  low  minimum,  the  Saa3,  water  oxygen 2)  be  (figure zone  or  mixed w i t h  the incoming water.  In A u g u s t  3)  the  the  lack  that  they  were a v o i d i n g  (figure was  they  found  relocating were  within  have been c a r r i e d The c o p e p o d s winter  100-120  the  t h r o u g h o u t most  was  and between would  1)  110  avoid  t h e oxygen  water  that  they  Saa9 would have  d e p t h were an  been  indication  (0.11 mL L ~ )  at  1  of the i n l e t  t o d e p t h s between  then  75 and 100 m a t  the range of t h e s i l l  165  m  depth  Once  (75 m) t h e y  could  i n the outflowing  found  Inlet  i n Saanich  during  a residual  The  Strait  t h e CVs were Saa3.  out of t h e i n l e t  period.  1  zone o f oxygen p o o r w a t e r a t Saa0.8  o f 1985-1986 were e i t h e r  overwintering  Saa0.8  3  from t h e summer months o r were t r a n s p o r t e d the  at  been  o f CVs a t 160 m (6.0 n r )  this  t h e low oxygen  m  a d e p t h above  with  c o p e p o d s below  the  i n August  < 0.20 mL L "  copepods  copepods a t s t a t i o n  17). I f the d i s c r e t e  also  likely  of  to  (figure 54).  data c o l l e c t e d  and  1985 t h e c o n c e n t r a t i o n  and  t h e CVs  by t h e CVs w o u l d have  54), the  occupied,  Childress,  correspond  containing  along  t o be f o u n d a t  1984). S e c o n d l y ,  by m a i n t a i n i n g  displaced  crustacean  1967;  t o a d e p t h o f 75 m a t Saa9 a n d Saa0.8,  150 m a t Saa0.8  this  et a l . ,  to the hydrographic  around  &  other  (Longhurst,  1  they can t o l e r a t e  t h e maximum  6  addition,  t o depths which  1985,  (table  In  c o p e p o d s , do n o t t e n d  1983; A l l d r e d g e  be presumed  minimum oxygen  17  water.  the  fall  and  population  left  over  into  the i n l e t  during  of  Georgia  data  56  demonstrated  that  were p r e s e n t consider of  very  small  i n t h e t o p 100. m  these  the S t r a i t  transported N.  a  data  throughout  the  (<0.2 i r r ) 3  winter.  If  t o be r e p r e s e n t a t i v e o f t h e c e n t r a l  of Georgia, into  number o f c o p e p o d s  then  the i n l e t  t h e copepods  could  from t h e S t r a i t .  p l u m c h r u s may a l s o have been t r a n s p o r t e d  we  region  have  been  A few s p e c i m e n s o f  i n f r o m Juan de F u c a  Strait. The  overwintering  with  a  Bioness  1986.  A much g r e a t e r  was  possible  nets  ( t a b l e 17),  captured data  so  few  appear  was  sampled  (208 m )  was  3  sampled  yet only  one  from September  The i m p o r t a n t  specimen  to  March  i n Saanich t o March  from J a n u a r y  of encountering  there  using  probably Inlet  the  was n o t a  to  reproduce  1986. As t h e r e  t o March  was  supports the  (1985-1986)  i s that  number o f N. p l u m c h r u s  o r t h e C-B  N. p l u m c h r u s  100 a n d 150 m. T h i s  point  adults collected  of  than  were  (<0.20 n r ) , t h e 3  a copepod of t h e o p p o s i t e  sex d i d n o t  high.  N. spring  p l u m c h r u s was r e i n t r o d u c e d and  summer  consistently  higher  Saa3  spatial of  N. p l u m c h r u s  e i t h e r t h e mSCOR n e t ( t a b l e 3 0 ) ,  s u b s t a n t i a l numbers from J a n u a r y  likelihood  and  volume o f w a t e r  3  net.  of  (owned by I.O.S.) a t Saa3 on M a r c h 5,  (0.02 n r ) between  sufficient in  sampler  with  collected  mSCOR  population  than  originating stations  results  Saanich  1986. W i t h i n  concentration  a t Saa0.8  distribution  the i n l e t  of  into  (from  the  April  from t h e m a j o r i t y the i n l e t  c l o s e s t t o t h e mouth  fjord  t o August;  at  the north-south  and c o n c e n t r a t i n g Saa9).  was a Saa9  t a b l e 2 1 ) . The  of the Saanich  (Saa3 and  i n the  there  (p=0.07) o f c o p e p o d s  o f N. p l u m c h r u s a l o n g  from o u t s i d e  Inlet  The  axis  population a t t h e two fact  that  57  station an  Saa0.8 becomes more s t a g n a n t  indication  that  there  is less  of an  hence c o p e p o d s , a t  this  station.  further  that  reproduction  or  evidence  l a c k i n g i n the The  1986  captured  at  Inlet  throughout  is  (1973)  and  during  copepodites central  region  In an  be  the  the  changes  was  f r o m the  increased  f r o m May  80  and  Saa0.8  is  p l u m c h r u s was  & 41).  station  The  Strait  reduced  spring  m at  and  this  period  and  to  test  the  Saa9 d u r i n g  months. At  and  but  Saanich they 42).  plumchrus  the  the  Strait  nauplii  and  summer. Hence, some of  the  and  the  southwest  of  the  to Saanich  pattern  difference these  three  population G1545 t h e  June and  into  t h a t N.  population,  t o J u l y whereas t h e  could  (figure  May-July) could  of G e o r g i a  CVs  were  of  to demonstrate that  Strait  time  concentrations  (from  April  June because  p o r t i o n of  towards Haro S t r a i t  a t Saa9  in  and  of G e o r g i a  May,  deepest  the  Sate  CIVs  (1984) have o b s e r v e d  in high  same i n May  J u n e , and  top  d i f f e r e n c e i n the  among t h e  40  at  water,  f o l l o w i n g month, t h e y  tide.  found  used  detected  remained the  ebb  at  the  overwintering  population i n the  In  central,  dispersed  A significant  and  the  Black  c o p e p o d s a t G1545 and  (figure  tide.  i n the  in  attempt  in  numbers  of N.  months of A p r i l ,  i n the the  may  copepods are  variance  the  present  concentrated  Georgia  flood  transported  were s t i l l  low  were c o l l e c t e d  Sate d u r i n g  been  Fulton  CVs  the  have  The  exchange of  inlet.  C I V s and  during  t h a n e i t h e r Saa3 o r Saa9 i s  of be  an  increase r e l a t e d to  analysis  i n the months  numbers  population  Inlet.  number  of  (table  25).  (p=0.02)  was  had  steadily  total  numbers a t  Saa9  then  increased  in  For  b o t h s t a t i o n s , the  c o p e p o d numbers  June and  J u l y , were g r o u p e d  into  two  of  had July  in  May  homogeneous  58  subsets  (table  25). T h i s  indicated  t h a t t h e r e was  a  significant  rise  i n t h e p o p u l a t i o n a t G1545, b u t i t d i d n o t n e c e s s a r i l y l i n k  this  t o t h e i n c r e a s e i n t h e p o p u l a t i o n a t Saa9. The  to  i n c r e a s e i n numbers a t G1545 i n J u l y  the  lateral  movement  downward m i g r a t i o n Strait  (Fulton,1973;  increase  below  migration,  July  may  150 m  (figure  in July  is them  period  copepods around the J u l y were  w e l l below  about  designating this  since  replicate  On  to  July the that  as  vertical  of  barrier  i n deep  numbers  regions in  of copepods s h o r t l y  after  the  transport  of  to  the  increase  in  o f t h e CVs  the  cautious  population  As  was  related  mentioned  (table  Saa0.8  6 & 9). During  a t 150 m i n  for  July  indicates  c l o s e t o oxygen August  were above oxygen c o n c e n t r a t i o n s o f 0.11  mL  1985,  L" . 1  In  l e v e l s of  the copepods July  replicate  s a m p l e s d e m o n s t r a t e d t h a t N. p l u m c h r u s o c c u p i e d  containing  0.20  mL  0  2  L"  1  a t 140 m  and  the  1 6 ) . An e x t r a p o l a t i o n o f  and Saa9  situated  to  previously,  N. p l u m c h r u s were f o u n d  t h e CVs would have been  to  Saa9  N. p l u m c h r u s  of  between  (table  copepods  because the  a  refuge  the  were not made a t S a a 9 .  1985 and a t 160 m i n A u g u s t  1  the  as  the  at  opposed  real  hauls  concentrations  L"  to  of hand,  d e p t h . However, one has t o be  as a  distribution  0.20-0.30 mL  other  14th s a m p l i n g - d a t e when most  the s i l l  low oxygen d a t a  act  increase in  p r e s e n c e o f low oxygen b o t t o m w a t e r . maximum  region  the  from f i n d i n g  to the i n f l u x  June 5th c o l l e c t i o n  copepods p l u s the  i s not due  thought  4 7 ) . The  correspond  The  1984).  attributed  i n a deep r e g i o n o f t h e i n l e t  preventing  the i n l e t  and/or  the deep c e n t r a l  Black,  concentrated  low oxygen  water  into  i n numbers a t Saa9  becoming  of  o f CVs  of  c a n be  maximum  1986, water  concentrations  59  were  found  prior  t o the h o r i z o n t a l  CVs  at  were  1  m  ( t h e oxygen haul  level  (figure  did  170,  and  47). At  (1970)  data  Saanich  Inlet.  oxygen  The  indicate  that  with  t h e zone,  respect  The (0-0.16  CVs  below  and  150  0.36  m  L" )  between  1  120 and  coincidentally,  vertical  haul  f o u n d above  mL  located  examined  to  the  in  Saanich  was  in  this  His results  system a c t i v i t y (  <  t o 0.06  he  not  zone  low  acknowledged  that  poor  water  (table  10).  in CVs  a  the were  (figure  oxygen  44).  zone  enzyme a c t i v i t y station  which  showed t h a t within  L " ) , due  or  of the  zooplankton with  L"  to a  quantity  in (ETS was  t h e r e was  t h e oxygen  Mg-atoms 1  the  estimated.  found  zone. U n f o r t u n a t e l y , Devol  an a c c o u n t of t h e s p e c i e s but  5  mL  be  of  displaced  he  Since  oxygen  3  zones  1  a l t h o u g h a few  distribution  Inlet  equivalent  regime  the l o c a t i o n  (0.3 n r ) t h i s  of the  Saa0.8.  which  oxygen  Saa0.8  by m e a s u r i n g t h e r e s p i r a t o r y  close  zooplankton  present  at  of t h e o c c u p a n t z o o p l a n k t o n a t  approximately  provide  m  region  3  o f enzyme t r a n s p o r t  region  mL  L~ ; however,  can o n l y  N. p l u m c h r u s  (0.5 n r ) and below  Inlet  activity)  0.30  to  31).  displaced  160  from t h i s  to the l o c a t i o n  Saanich  lack  samples  (1981)  respect  the  depth  (table  content  r e n e w a l i n 1986  Perhaps  Devol  Saa3,  were shown t o o c c u p y d i s c r e t e  t o t h e oxygen  deep water mL  days  than  be g o v e r n e d by t h e low  c o n c e n t r a t i o n s of 0.10  n o t sample  the  o b l i q u e h a u l s c o v e r e d a d e p t h r a n g e o f 25 m, CVs  1.5  180 m i n A u g u s t , but  a t t h e s e d e p t h s were a l l g r e a t e r  N. p l u m c h r u s m i g r a t e s may  above  160,  figure  collected  18).  Hoos'  in  d a t a was  samples;  found at depths of  t h e oxygen L"  145  a  poor  which  is  scarcity  of  1  (1981) d i d n o t of  N. p l u m c h r u s was  zooplankton one  of the  60  common  species.  At  stations  declined renewal (0.77 may of  to  <  Saa3 and Saa9 irr  0.2  the  following  3  i n 1986. The d i s p l a c e m e n t mL  L" )  a l s o have been t r a n s p o r t e d  samples  (table  indicated  The  Cowen  between  0  for  low number o f CVs overwintering  h a s been  ( 1982)  and  hydrographic  Inlet  200  in  data c o l l e c t e d  in  Hoos's  the  showed  that  120 and  station  there  i n 1986.  i n the c e n t r a l some  175 m  (table  1974  of  (table  1969  t o w a r d s t h e mouth The  31).  indicates  A n d e r s o n and D e v o l intruding  water  differences  that  Saanich  report  (#30)  a displacement  o f low  oxygen  figure  L" ) t o 1  55).  the  However,  l o c a t e d more  south  of  t h e oxygen  centrally  o f h i g h e r c o n c e n t r a t i o n s of oxygen  80-200 m i n 1969 t h a n with  The  and  (table 32).  occurrence  mixed  irr  5.3  data  (0.0-0.17 mL  32;  years.  a t 150 m i n O c t o b e r  3  of t h e UBC  was  out  September  population  during  September  minimum d i s a p p e a r e d a t t h e s t a t i o n s  using  in  concentrations  December  (Hoos, 1970). An e x a m i n a t i o n 1969  layer t h e CVs  oxygenated water mixed w i t h the bottom waters of  water between  water  minimum  population  indicated  collected m  of t h e deep w a t e r  oxygen  f o u n d c o n c e n t r a t i o n s of 27 n r  1969 and  Inlet  7 ) . The  N. p l u m c h r u s  (Saa9) s u g g e s t s t h a t  p r e s e n c e o f an o v e r w i n t e r i n g  (1970)  dense,  of the  of  t o s h a l l o w d e p t h s and c a r r i e d  a negligible  r e g i o n s of S a a n i c h Hoos  the onset  t o 75 m i n September  1  the i n l e t  population  i n 1985  the  low  suggests that oxygen  150  in nitrate  r e n e w a l . They e s t i m a t e d  volume  bottom waters ( f i g u r e  (1973) c o i n c i d e n t l y below  a greater  between  m between  estimated July  the  55).  volume  and September  of i n t r u d i n g  w a t e r t o be  of  1969,  c o n c e n t r a t i o n s b e f o r e and a f t e r  t h e volume  of  the 4.3  3  61  x  10  m.  8  Calculations  3  i n t r u d e d and m i x e d A).  The c a l c u l a t i o n s  before  and  September  after  17  Since  in  were b a s e d the  indicated  1969  since  region  and  the  3.2  thay  the  7 t h and  oxygen  was  a g g r e g a t e d above  t h e oxygen mixed  in  1985  the o t h e r  displaced  a t Saa0.8 and Saa9.  hand,  This  to shallow  been  this  region  that  at  close the  water  south of  Saa3  10  o f w a t e r w h i c h came i n t o  not  likely  that  m  10  m lens.  out  through the e n t i r e  that  above  proportion  out of S a a n i c h  Inlet  x  10  8  m.  layer  of  sill  the  3  to  however,  is  it  left  t h e w a t e r would have the  to  i s equivalent  w a t e r would have  Therefore, of  3.0  concentrated  the i n l e t ,  a l l of the o u t f l o w i n g  p e r s . comm.).  (approximately  would be  t h e 100 m c o n t o u r . T h i s  region  located  w h i c h would need t o be l o s t  a  I t i s more p r o b a b l e  basin.  of copepods a r e  i n the p o p u l a t i o n for  have  t h e r e n e w a l e x c e p t where  majority  slightly  of  would  t o t h e s l o p e of t h e  volume  the  and  not  calculated  LeBlond,  not  w a t e r . On  the  calculations,  water  because  Saa9 and  situated  transported  concentrations  poor  volume  T h i s v a l u e was  (Paul  oxygen  t h e oxygen  f o r the d e c l i n e  copepods  3  t e n d e d t o o c c u p y d e p t h s above  assume  station  account  low  by t h e m i x i n g o f water d u r i n g  km)  m  B  150 m ( a p p e n d i x  t h e c o p e p o d s would have been d i s p l a c e d  also  we  between  of  copepods  c o p e p o d s were s i t u a t e d If  x 10  of t h e r e n e w a l on A u g u s t  zone  oxygen minimum was  depths  the  the  2.9  t o t h e r e n e w a l would have been d i s r u p t e d  that  disturbed  that  w a t e r below  on  onset  w i t h the o x y g e n - c o n t a i n i n g discrete  indicated  the r e s i d e n t  discrete  minumum p r i o r  a  1985  (appendix A ) . a  maintained  in  with  for  during  in a  flowed  ebb  tide  b a s e d on t h e s e  rough  N. p l u m c h r u s  population  c a n n o t be d e t e r m i n e d .  62  The sketchy not  oxygen but  from the  indicates that  displace  greater  data  a region  of  volume of m i x i n g  some  of  Inlet  could  have been m i x e d  and  the  therefore  CVs  deep water  low  oxygen  in  1969  located  account  data  the  as  that  UBC  report  (#37)  renewal  to shallow  (table  i n the  in  did  d e p t h s due  to a  33).  This  c e n t r a l region  retainment  of  rather  1974  to deeper depths d u r i n g  f o r the  is  suggests of  Saanich  the  some of  renewal  the  CVs  in  1974. Sechelt  Inlet  successfully central  data  demonstrate  overwinter  at  region  of  the  characteristics  of  Sechelt  Saanich  Inlet  column  due  very  to  sill  these  two  plumchrus  high  August  Inlet.  (16.9  i n the  1986  overwintering  the  water  of  Sechelt which  time  column  bottom waters are  relatively  N.  overwinter  m at  than  at  3  CVs  undisturbed deepest  water  i t has water.  in  a I of  fairly  s t a t i o n Sc 1 density Sc1  of  population  was  were  below 205 Inlet  by  the  of  the  sampled  and  features  but  was  in Saanich  the  the  the  the  Sc1a,  nr  at  that  is sufficiently  at  found  February  Inlet  physical  of  1985  t o 2.0  deep  overwintering  p l u m c h r u s was  declined  175  the  can  the  although,  exchange  eggs, c o n f i r m i n g  d e p t h s below  plumchrus w i l l  the  part  the  throughout  i n November  3  had  inlet.  during  irr )  Sc2a, a deeper m i g r a t i o n  Providing  with  circulation,  N.  latter  copepods  females contained  reproducing  The  contrast  restricting  Towards t h e  overwintering adult  m)  Georgia.  oxygenated  estuarine  in Sechelt  48).  is  plumchrus than  of  Inlet  N.  shallower  major d i f f e r e n c e s f a v o u r  concentrations  (figure  and  (15  Strait  it  strong  shallow  think N.  in that  depths  that  again  in  found  at  m at  Sc2  (figure  o x y g e n a t e d and water  possible  49). the  movements, depths.  63  5.2.  Alternative  Causes  f o r the D e c l i n e Saanich  A N.  number  plumchrus  of  during  were c o n s i d e r e d . 1) W h i l e into  regions  of l o w e r  their  renewal  containing hydrogen forced  low  t o occupy  or z o o p l a n k t o n ,  location  could  some of t h e  oxygen  sulphide.  or  3) The  could  in  the  decline  in Saanich could  CVs  perhaps  have l e f t  due  to at  c a u s e d e a t h . 2) D u r i n g  the  be  potentially  shallow  sink  conditions  may  the  in  Inlet  oxygen o r , as t h e oxygen d e c l i n e d  the  deep water  for  i n d i a p a u s e t h e CVs  of b a c t e r i a  of  causes  overwintering period  the r e s p i r a t i o n depth  plumchrus  Inlet  alternative their  of N.  mixed  toxic  depths  agents  which  the copepods  into  the  water such  CVs  as  were  more v u l n e r a b l e  to  predation. The region  notion, of  that  lower  oxygen  misconception  of the  species  evolved  insect  has  the copepods while  'diapause' many  of  d i a p a u s e , as o u t l i n e d  of dormancy, w h i c h equivalent cyclical  to  change  Elgmork  the  copepods  form  of  lipids  N.  decreased  level  Harrison  et  al.,  N.  history  are thought metabolic 1983).  definition  of  diapause,  independent  of a food s u p p l y .  a  This  features  of  (1978)  Nilssen  have an a c c u m u l a t e d  of  plumchrus.  characteristic  plumchrus in  late  c o p e p o d i t e moves t o d e e p d e p t h s p r i o r  which  from  and  life  The  in  arose  into a  (1971 ) c l a s s i f i c a t i o n  in their  fall.  diapause  sink  i n Mansingh's  diapause.  five  in  potentially  phase  copepod  stage  could  the  refer  to  as  demonstrates summer  as  a  the  t o the onset of  s u p p l y of r e s e r v e s i n the  t o be  activity According organism  sufficient (Lee to  et  to s u s t a i n al.,  Mansingh's  should  be  a  1972; (1971)  completely  64  As  the metabolic  it  enters  what  in  which  the  Nilssen, and  did  i s considered copepod  is  1978). However,  intensity  that  activities  with  were c a p t u r e d  o f t h e o r g a n i s m become  the r e f r a c t o r y stage in  this  a  deep t o r p i d  s t a t e of t o r p o r  the s p e c i e s . L i v e  of  diapause,  state  (Elgmork &  varies in duration  specimens  of  on a m o n t h l y b a s i s from May  N.  thermos  pipette.  when  The  fact  environmental refractory  t h e l i d was that  stimuli  swam  1986 about  removed and i n t h e p r e s e n c e o f a  they  were  indicated  phase of d i a p a u s e  plumchrus  t o November  n o t show s i g n s of t o r p o r . The c o p e p o d s a c t i v e l y  the  reduced  able  that  (Mansingh,  to  they  respond  were  not  to  i n the  1971; E l g m o r k & N i l s s e n ,  1978). In a d d i t i o n , when f o o d was a v a i l a b l e t h e c o p e p o d s a t e was d e m o n s t r a t e d The  feeding  environmental of  by t h e p r o d u c t i o n  stimuli  'low i n t e n s i t y  such as S a a n i c h overwintering  period,  to  such  tolerate  anaerobic  N. p l u m c h r u s would i t would  suggest  altering  environmental (figure as  this  (Mansingh,  s p e c i e s was  in  1971). In an  i t would be a d v a n t a g e o u s  to  a  state  environment during the  be  able  sink  threaten  into their  regions  o f low oxygen  that  t o the extent  survival.  to  during  be  during  able  August  depth of m i g r a t i o n ; stimuli  to  i f the organism c o u l d not  c o n d i t i o n s . I t i s t h e r e f o r e not p r o b a b l e  17 & 4 7 ) . In l i g h t  t h e oxygen  that  changes, e s p e c i a l l y  concentrations their  (table 28).  where t h e oxygen c o u l d d e c r e a s e  N. p l u m c h r u s a p p e a r e d oxygen  pellets  of N. p l u m c h r u s and i t s r e s p o n s e t o  diapause'  Inlet  respond  that  activity  of f a e c a l  as  they  to  avoid  1985  and  remained  the o v e r w i n t e r i n g  of these  observations  l e v e l s d e c l i n e due t o t h e  unfavourable July  1986 by  responsive period  i n 1986  I suggest  consumption  of  to  that, oxygen  65  through  respiration,  the copepods a d j u s t  their  position  water c o l u m n . However, a s d e n s e w a t e r  intrudes during  and  in  mixes  with  copepods c o u l d  the  the h y d r o g r a p h i c  and  in  oxygen  2) o n l y  tolerate  data  levels below  presumably  increasing  up  to  potentially  toxic  1  of  the  1986 ( s i g n i f i c a n t  oxygen  distribution  incoming water i s  be  6 & 9), able  to  may  i n September  response  the  to  bottom.  to tolerate  1985  oxygen  the  For the levels  a t Saa9 a s w e l l a s  1  i n t h e w a t e r . The v a r i a t i o n  difference  fjord  in  the  readily  apparent  The main  factor  related  t o the period  As t h e d u r a t i o n  52).  1) a s  i n August  i n the population  individual  also  be f o u n d  1985 and  mean a t f o u r  c o p e p o d s may  overwintering  in  require  copepods  in a  t o occupy a range of  (table 27).  Variation  rise  in  towards  o f oxygen, a l t h o u g h  oxygenated  minimum  because  may  of the copepods  d e p t h s a t p=0.05) s u g g e s t s t h a t  depths  depths  a t Saa0.8 and 0.62 mL L "  July  well  this  oxygen minimum zone  agents present  levels  survival  population  t h e y would have  horizontal  oxygen  and lower i n October ( t a b l e  deeper  of  the  different  to their  some o f t h e  1  the  gradient  0.71 mL L "  displaced  renewal  a s low a s 0.2 mL L " .  seek  copepods t o s u r v i v e ,  the i n l e t ,  f o r 1985 i n d i c a t e s ,  i n September  oxygen  the  be d e t r i m e n t a l  some members  Copepods would  water  be t r a p p e d below  layer. This could  low  bottom  a  i n the  i n the r e s u l t s influencing  other  of  killed  individual  copepods  was  o f t h e low oxygen e x p e r i m e n t s .  their  of time that  of c a p t i v i t y  i n the percent Many  tolerance  tolerance  appeared  t h e y were h e l d  increased  there  was  to  be  in captivity. a  consistent  i n t h e low oxygen e x p e r i m e n t s ( f i g u r e  factors  appeared  to  contribute  to  the  66  variability  i n the  experiments  could only  further  most  the S a a n i c h captivity  was  from  Inlet  elevated  and  (figure  5).  ecdysis,  of  differ  from  last  that  a  basis  preliminary  after  of  these for  0.56  mL  1967;  five  three  5). T h i s suggests to  which  were  so t h e i r  of  run tested  the  of CVs  than  to  low  rise  the  were  oxygen  first  likely  of  in  the  of  the  respiration before  and  exoskeleton  of  after  (Passano,  1976).  earlier  held  in  (3 t o 8 d a y s ) . The  that  ( i . e . in  were c o n d u c t e d  previously  been  intensified  been a t d e e p o v e r w i n t e r i n g d e p t h s  experiments  in  i n c r u s t a c e a n s have  immediately  Hagerman,  recognized in  oxygen demand would have  respiration  stages  collected  than  at  that  oxygen  after  period  lower  low  10  s p e c i m e n s w h i c h were time  that  in in  organisms  was  Georgia  c o p e p o d s was  of G e o r g i a  o r s i x months i t i s p r o b a b l e  t h e CVs  days  1  s o u r c e of v a r i a b i l i t y  shedding  a c o p e p o d has  87  L~ ) t h e p e r c e n t a g e  of S t r a i t  (table  Measurements  d u r i n g the  and  observed  t h e s e c o p e p o d s were t e s t e d  susceptibility  during  was  Georgia.  and  their  16  be more a c c l i m a t e d  of m o u l t i n g  Paranjape,  period  and  there i s a s i g n i f i c a n t  and  After  The  9  organisms  1960;  that  o f m o u l t i n g . Some of t h e CVs  process  shown t h a t  may  potential  8,  indications  tested  in captivity  the S t r a i t  experiments  these  became a p p a r e n t  s o u r c e of v a r i a b i l i t y  (0.54  g r e a t e r than  A further  the  as  5 2 ) . Even t h o u g h  levels  Saanich  those  used  copepods  (figure  same t i m e  from  be  interesting  Inlet  oxygen  survived the  so i t soon  experiments.  The  low  results  June  copepods  or  on J u n e ,  captivity  level  these  for  of m o r t a l i t y  measured, presumably  for a  July).  Strait a  of  short  i n these due  to  67  the  short  time spent  overwintering  period  obtained  oxygen  CVs  at  in Saanich  levels  of  0.20  The  range  in  mL  that  L'  experience  l e v e l s of  the  within  ability  the  inlet  water, are  ( f i g u r e 54).  through  Haro  the  Strait  A  i n the  of  Georgia  in  w o u l d be  susceptible  likely  below  the  Some  anoxic of  be  N.  to  the  most CVs  the  was  1  2  N.  low  mouth of  low  is  to  since oxygen  rule  out  the  plumchrus there oxygen and poor  to  the  t o be  is  the Of  waters, that  and  may those  into  o f N.  the  period  in  of  CVs  percentage few  copepods  based  Hence,  in  plumchrus  overwintering  a  a  contacting  transported  the  a  therefore  note  inlet  individuals.  on  this  the  is  not  mortality. s u c h as H S  gradually  2  Saanich  Inlet  (Lu  into  the  for a brief  withstand  being  to t h e i r  intruding  period  build  et a l . ,  t o have been d i s p l a c e d  mixed  remain  harmful  mortality  close  oxygen c o n d i t i o n s  expected some  their  a l s o have been t r a n s p o r t e d  m during  low  in  unexpected  not  ( S t a n i e r e t a l . , 1979). I t  be  50%  concentrations  substances  plumchrus c o u l d may  of  important  could  b o t t o m w a t e r s of HS  do  i n t o oxygen  likely  among  i t would o n l y  concentrations  the  oxygen minimum zone, o n l y  to the  toxic  readily oxidized  long  is  100  the  oxygen p o o r water and However,  It  stage  47).  survive  a m a j o r c a u s e of  Potentially in  to  ( f i g u r e 39).  variability  t o be  L"  experiments  since  top  carried  apparent  few  Strait  were p r e s e n t  &  copepods mixed  mortality.  inflowing  mL  a population  copepods s i t u a t e d c l o s e the  early  However,  0.47  ( f i g u r e 17  1  of  their  p e r c e n t a g e of  ( f i g u r e 52).  the  I n l e t were c o l l e c t e d i n , or v e r y  results  possibility  i n c a p t i v i t y and  since  i s not  s u r v i v a l . The  1986).  with  the  water. i t would  known  e x p o s e d t o H S, 2  up  or  how what  addition  of  68  HS  i n t o low  2  decrease in  the  oxygen water  (<0.15  p e r i o d of  survival  the  laboratory  (Theede,  were  low  to  be  July  small in  3,  and  September. S i n c e any  conclusions  been c a p t u r e d surface was  while  low  after  their  would  still The  days  N.  t o be,  although case  (Fulton,  plumchrus  following the  this  the  incidence  September  1985  plumchrus  in  the  17,  1973; was  of and  at  CVs  comm.). A l s o , the  dead  sediment  of  cause  i n J u l y and can  not  they  may  have  hauled  adaptation been a d e q u a t e l y et a l . ,  f o r c e d to occupy of  low  predation  on  1986  to  due  shallow  a  the  found.  decline  in  the  migration  Pacific  avoiding  In  Ocean i s  t o be  Saanich  depths of  75  to  I suggest  s p e c i e s may  of  predation  demonstrated  greater  to  they  oxygen w a t e r s . this  laboratory  c o p e p o d s but  the  1983).  the  not  deep  for  to  would  the  and  one  we  samples  The  21  i n d i c a t e s that i f  the  for  May  floated  This  trap  samples.  Saa0.8. O n l y  i n the  CVs  29).  of G e o r g i a  Harrison  displacement  two  i s predation.  Strait  never  1985,  to  substances  periods,  l a r g e r numbers t h a n were  i n p a r t , an has  two  due  trap  c a u s e of d e a t h ;  (table  alternative  of N.  plumchrus  thought  N.  have c o l l e c t e d  sediment  over  a substantial proportion  final  population  the  toxic  t r a p s were b e i n g  pers.  numerous c o p e p o d s were k i l l e d have c o l l e c t e d  to  crustaceans  Inlet  numbers were c o l l e c t e d  60-75% of  4-11  in  sediment  (Akira Taniguchi,  surface  p r e s e n c e of  captured  about  the  observed that  the  i n e i t h e r case,  such  shown  of  in Saanich  7 t o September  numbers were f o u n d  draw  it  August  been  i n some s p e c i e s  detected  However, s p e c i m e n s were o n l y to  has  1  plumchrus  oxygen c o n d i t i o n s or  expected  L" )  1973).  A m a s s i v e d i e o f f of N. either  mL  the  Inlet 100  that  have r i s e n  amount  of  m  in  light  69  available  for  the  d e t e c t i o n of p r e y and  between t h e d i s t r i b u t i o n In September predominating  1985  in  Mackie  important and M.  This it less  suggested  primarily  confused  l e a v e s A.  prey.  potential  that  N.  the  predators siphonophore  digitale  plumchrus  i s probably  i n the S t r a i t most  of  abundant  occupies  the  top  plumchrus  (although  (table  species,  50 m and t h e r e f o r e this  species  deeper  d w e l l i n g s p e c i e s ; Mackie,  digitale  as one  of the p o t e n t i a l  that  50  and  130  a non-visual  m,  is  1985).  predators since  however, I t h i n k t h a t  predator  an  Georgia  w i t h two  between  likely  the  m a t Saa9 were t h e  However,  o v e r l a p w i t h N.  resides  100  increased overlap  t h e hydromedusa A g l a n t h a  inlets.  atlantica,  easily  non-visual  for gelatinous predators  adjoining  would not  t h e two  and  (1985)  food  t h e p r e d a t o r s and  the upper  Muggiaea a t l a n t i c a 34).  of  the  exploited  the  it is  copepod  aggregation. In  the  californicus minimum  Santa  Barbara  Basin, C a l i f o r n i a ,  overwinters at  depths  zone c o n t a i n i n g 0.2  mL  0  Predators, primarily  the deep-sea  have  t o f e e d on  been  observed  'WASP' s u b m e r s i b l e sea  smelt  abundant  found  bathylagid fish  waters, found  autumn and  t h e r e f o r e may  likely  t o be  such  and as  be  hake,  f e e d i n g on N.  the  Leuroglossus  L.  oxygen  stilbius, from  the  s p e c i e s of  deep-  s c h m i d t i , i s the t h i r d  most  Strait  of G e o r g i a  w i t h t h e a d u l t s below  a p r e d a t o r of  Saanich  in  pacificus  ( A l l d r e d g e e t a l . , 1984).  1  smelt  i n the  1985). J u v e n i l e s o c c u r  Fishes  m  ( A l l d r e d g e e t a l . , 1 9 8 4 ) . The  in local  of G e o r g i a  L"  2  450  the copepod a g g r e g a t i o n  Phillips,  Strait  of  Calanus  N.  (Mason & 150  plumchrus  m in  in  the  chum salmon a r e  less  Inlet. and,  p i n k and  plumchrus  i n t h e autumn. These  fishes  70  feed  on  N. p l u m c h r u s  juveniles,  but  euphausiids,  later  1973).  Pacific  The  upon c o p e p o d s (Hart,  they  spring  shift  i n summer  in f a l l  to  and  summer  larger  (LeBrasseur  while  sized et  prey  they such  a l . , 1967;  herring,  Clupea  pallasi,  although  e u p h a u s i i d s t e n d t o be  are as  Hart,  continues t o prey preferred  1973).  In  Saanich  from p r e d a t o r s too  in  low  in  Inlet  i f they a r e capable oxygen  hydromedusae s p e c i e s (Raymont,  t h e c o p e p o d s may be r e l a t i v e l y  for their can  1983; A l l d r e d g e ,  of occupying  protected  depths that  are  p r e d a t o r s . However, some f i s h a n d  tolerate 1984).  low  oxygen  concentrations  71  6. CONCLUSIONS The  f o l l o w i n g c o n c l u s i o n s and r e c o m m e n d a t i o n s a r e c o m p i l e d  from t h e r e s u l t s c o l l e c t e d sediment  Saanich  Inlet  N. p l u m c h r u s  appeared  that  population 2. in  t h e l a b o r a t o r y and  in  had a n e g l i g i b l e 1985 (September  there  was  going  Saanich  Inlet into  during  the i n l e t  from t h e S t r a i t  3.  1985 t o March  to  N. p l u m c h r u s was p r e s e n t  probably  overwintering  be  a  i n 1986 b a s e d on t h e September  reintroduced  The  the  distributed  in fairly summer  1  and  4.  in  migration  of t h i s  Saa9 75 L"  to  1985;  they  summer  of  were 1986,  Inlet.  i s affected  The  CVs  were  t o low oxygen  that  they  were  t o shallow  able  to  detect  levels.  data  indicate  a  pronounced  deep w a t e r  (0-0.2 mL L~ ) was 1  d e p t h s (up t o 75 m) a t s t a t i o n s Saa0.8  and  i n 1985. I n 1986, t h e low oxygen zone was a l s o d i s p l a c e d t o m  1  concentrations  species  i n 1985 and 1986. The low oxygen zone  displaced  overwintering  o r above oxygen c o n c e n t r a t i o n s -of 0.1 t o 0.2 mL  Hydrographic  renewal  1986) and i t  of G e o r g i a .  i n 1985 a n d 1986 s u g g e s t i n g respond  high  i n t h e s p r i n g and  ontogenetic  at  low  population  samples.  by t h e low oxygen b o t t o m w a t e r s i n S a a n i c h  L"  the  traps:  1. of  i n the f i e l d ,  but i t c o n t a i n e d  higher  a t Saa0.8 and 0.77 mL L ~ 5.  During  have  been  the renewal located  encroaching  of  the  (0.31 mL  at Saa9).  i n September  above  d e p t h s a t o r above t h e s i l l displacement  1  c o n c e n t r a t i o n s o f oxygen  the  depth.  copepods  low o x y g e n - c o n t a i n i n g  were  presumed  oxygen minimum and hence a t  This  or  t h e CVs  resulted  their  water.  from e i t h e r t h e  avoidance  of  the  72  6.  The  coincided waters.  loss  with One  attributed  of  copepods  in  t h e a p p e a r a n c e of  primary  cause  to t h e i r  the  f o r the  transport  September dense,  1985  and  1986,  intruding  bottom  r e d u c t i o n i n number of CVs  out  of  the  inlet  during  is the  renewal. 7. N.  1974), 1985  and  occupied to  the  years  p l u m c h r u s was  and in  Of  find  in  1986.  N.  Sechelt  comparable  a very  shallow  sill  a d u l t s (2.0 the  sill  to  irr )  Saanich  of  a  with the  water  copepods  below  Inlet.  175  even  t h e e x c h a n g e of  present new  m,  However,  i t i s oxygenated  restricts  were  3  production  waters  than  depth.  i n that that  (1969  of a g r e a t e r volume  in Sechelt Inlet  depth Inlet  winter  t o have e n a b l e d  below  Saanich  p o p u l a t i o n of  in  February  generation within  Inlet.  9.  Alternative  numbers  of  overwinters to  N.  plumchrus  detect  and  The  were  'low  the  decline  examined.  intensity  a v o i d low  population  Since t h i s  diapause'  oxygen w a t e r s , is  in  not  and  species appears  changes  i n oxygen  likely  to  be  a  survival. presence  w a t e r , may  be  populations  of N.  the  of  for  at o v e r w i n t e r i n g depths  to t h e i r  10.  causes  in a state  concentration  1974),  in  during  intruding  from  indicating  threat  was  i s thought  environment  water. Reproductive  able  the  substantial  Inlet  renewal  of  plumchrus  differs  Sechelt  a  plumchrus o v e r w i n t e r s  i t has  1986,  which  in Saanich  Mixing  a suitable  regions  though  present  t h e d e e p water  by N.  8.  in  a  dense  of t o x i c  potential  substances  hazard.  p l u m c h r u s were f o u n d intruding  such as H S  However,  2  in  i n deep  years  during the winter  water which mixed w i t h the  when  (1969  &  anoxic  73  bottom waters, population. die-off  and  In  of CVs  11. with  'locked' Due  CVs  to  tolerate  oxygen  Low  were  the  region  Inlet  versus  the  The  the  Strait  the  between  may  massive  in  contact  oxygen  0.20  not  and  experiments  inlet  minimum some of  have been  0.70  mL  able  L" . 1  the  This  decline  . The to  main  be  d i d not  the  w h i c h may  source  of  the  p e r i o d of  time  have  p l u m c h r u s and  the  the  during  length  smelt N.  i n the  or  plumchrus fall  of  some of  of  the  predators. Pacific during 1985  of  month v e r s u s  Strait  increased their  of  the  and  a  safely  variability spent  in in  variation  c o p e p o d s were c o l l e c t e d  (one  provide  levels  c o p e p o d s were f o r c e d t o o c c u p y  of  a  i n oxygen t o l e r a n c e ,  i n 1985  to t e s t i n g  may  depths  below  of G e o r g i a ) ,  This  aggregations  of  entire  i n t o the  f a c t o r c o n t r i b u t i n g to  from w h i c h  prior  deep-sea  the  population  to t e s t i n g . Other p o s s i b l e causes  Inlet.  shallow  region  plumchrus  than e i t h e r  the  indication  transported  minimum oxygen  Inlet  as  be  tolerance  Saanich  between N.  of  traps.  water  o c c u r e n c e of m o u l t i n g  13.  no  variation  appeared  the  overwintering  of  N.  prior  was  loss  i n September.  estimate  experiments  the  plumchrus  levels  oxygen  captivity  and  N.  oxygen  into this  i n h a b i t e d by  the  the  have been a m a j o r  12.  be  sediment  individual  population  reliable  there  in  dense water may  i n t o low  carried  would not the  i n the  to  result  addition,  incoming  the  of  not  A p o r t i o n of  the  zone.  did  (Saanich  time five  spent  months),  experiments.  shallower Georgia  depths  in  or  Sechelt  distributional  overlap  Visual predators  h e r r i n g may their 1986.  such  have e x p l o i t e d  displacement  to  74  Recommendations: It  is  recommended  further  work  sampling  is  renewal  in  intrusions tolerance  in  required years  in  copepods  oxygen  levels,  of  Saanich  of t h i s  Georgia  of  the  provide  field  and  prior, both  Inlet.  collected to Saanich  is  still  In  from Inlet  be f o l l o w e d  laboratory.  plumchrus  unanswered.  More  and a f t e r  and the  with  extensive  deep  water  l a b o r a t o r y , t h e oxygen  regions  between S t r a i t o f  of  relatively  copepods c o l l e c t e d  information The impact  i n Saanich  up  t h e d e e p water  extensive  t r a p s e x a m i n e d on a more  more r e l i a b l e  o f N.  study  during weak  N. p l u m c h r u s w i t h i n t h e i n l e t . population  this  s p e c i e s n e e d s t o be compared  low o x y g e n . S e d i m e n t  could  that  about of  Inlet  from  high  regions  regular  basis  the m o r t a l i t y of  predators during  on  the  the renewal  Table 1: Station Coordinates and Maximum Depths. Station  Region  Latitude  Longitude  G1545 G1748  St.ofGeorgia St.ofGeorgia  49deg 15.0' 49deg 17.0'  123deg 45.0' 123deg 48.0'  406 417  Sate Saa9 Saa3 Saa0.8  Satellite Saanich Saanich Saanich  48deg 48deg 48deg 48deg  123deg 123deg 123deg 123deg  74 165 219 218  Scla Sci Sc2a  Sechelt Sechelt Sechelt Sechelt  Sc2  Ch. In. In. In. In. In. In. In.  42.1' 40.2' 36.7' 33.1'  49deg 33.5' 49deg 35.45' 49deg 38.9' 49deg 42.0'  29.1' 30.2' 30.0' 32.5'  123deg 47.25* 123deg 48.00' 123deg 50.6' 123deg 52.0'  Maximum Depth (m)  220 249 274 293  Table 2: Hydrographic and biological stations sampled in 1985. Type of Samples Horisontal Vertical Hauls Hauls  Station  Date  Hydrographic  G1545 Sate Saa9 Saa3 Saa0.8  May22 May21  V  May21 May21  V  7  -  V  v  V  -  -  —  —  V  -  May21  v  Saa3  July3  -  Saa0.8  July3  v  G1748 Sate Saa9 Saa3  Aug.8 Ang.8 Ang.7 Aug.7 Aug.7  V  Sept. 18 Sept. 18 Sept. 17 Sept. 17 Sept. 17 Oct.9 Oct.8  V  Oct.8 Oct.8 Oct.8  v  Nov.6 Nov.7 Nov.7 Nov.4 Nov.4 Nov.5 Dec. 17 Dec. 18 Dec. 16 Dec. 16 Dec. 16  Saa0.8 G1545 Sate Saa9 Saa3 Saa0.8 G1748 Sate Saa9 Saa3 Saa0.8 Sci G1545 Sate Saa9 Saa3 Saa0.8  —  July4 July3 July3  G1748 Sate Saa9 Saa3  -  V  G1748 Sate Saa9  Saa0.8  -  -  7  -  V  7  -  v  7  -  7  —  V  v  V  V  V  V V  -  -  —  -  -  v  7  v7  7  —  —  V V V V V  -  V  v  —  V  V  V V  v v  V  v v v v v  7  -  -  -  -  V  -  7  7 7  —  —  7  —  7  -  7 7 7  77  Table 3: Hydrographic and biological stations sampled in 1986. *Note: O n this occasion this was an oblique haul. Type of Samples Station G1545 Sate Saa9  Date Jan.28 Jan.29  Hydrographic  -  Saa3 Saa0.8  Jan.29 Jan.29 Jan.27  G1545 Sate Saa9  Feb. 17 Feb. 17 Feb. 17  Saa3 Saa0.8 Sci  Feb. 17 Feb. 17 Feb.20  v  Saa3  Mar.5  v  v  7  -  v  7  V V V  _  -  -  v*  V  V V  _  V  V V  v  V  V  —  v  7  v v  7  v  7  v  7  v  7  7  v v v v v  7  7  G1545 Sate Saa9 Saa3 Saa0.8  A p r . 15 Apr.14 A p r . 14 Apr.14 Apr.14  G1545  Mayl3 May 12  v  May 12 May 12 Mayl2  v  Jane3  V  June4 June5 June5 June5  Hauls  7  V V V  Mar. Mar. Mar. Mar. Mar.  G1545 Sate Saa9 Saa3 Saa0.8  Horizontal  Hauls  -  -  G1545 Sate Saa9 Saa3 Saa0.8  Sate Saa9 Saa3 Saa0.8  11 10 10 10 10  V  Vertical  V  -  7  -  V  v  7  -  V  V  v v v v  7  7 7 7 7  7 7 7 7  V  7  7  v  7  —  v  7  —  v  7  Table 3: Hydrographic and biological stations sampled in 1986, continued. Typ«: of Samples Horizontal Hydrographic Vertical Hauls Hauls  Station  Date  G1545 Sate Saa9 Saa9 Saa3  Julyl5 Julyl4 Julyl4 Julyl6 July 14  V  Saa0.8  Julyl4  V  v v  G1545 Sate Saa9 Saa3 Saa0.8 Scla Sc2a Sc2  Aug.6 Aug.5 Aug.5 Aug.5 Aug.5 Aug.7 Aug.7 Aug.7  V  v7  G1545 Sate Saa9 Saa3 Saa0.8  Sept .9 Sept.8 Sept.8  V  Sept.8 Sept.8  -  v v v  -  7  7 7  -  -  V v V 7  -  V V -  V -  v  7  v  -  7  -  7 7  -  v v v v v v v  -  7  -  7  v  7  7  -  7  7  v  7  7  7  —  v v V v  —  7 7  -  7  V  v  7  7  Table 4: Flowmeter measurements with and without the m S C O R net. flowmeter inside net without net wire 1=0 , depth of tow (0-50 m)  # of revolutions s.d. mean x=1361 x=2953  113 537  #of replicates  6 10  Table 5: Low oxygen tolerance experiments on CVs collected from the Strait of Georgia. *Note: Copepods collected from Saanich Inlet. Experiment # Funnel  Oxygen (mL L ) Mean B.d. - 1  Mortality % Ratio  1 1 1 1  Ta Tb Ca Cb  0.45 0.44 5.52 5.68  0.09 0.04 0.35 0.06  100 100 0 0  2 2 2 2  Ta Tb Ca Cb  _ 0.91  _  _  0.07  -  6.04  -  0.02  3  Ta Tb Ca Cb  0.61 0.62 6.37 6.39  Ta Tb Ca Cb  0.83  7/7  8/8  0/7 0/8  Date Collected (1985) May 12 Mayl2 Mayl2 May 12  Period of Captivity (days) 12 12 12 12  Duration of Experiment (hours) 24 24 24 24  Range of Oxygen (mL L ) 0.60-0.36 0.48-0.38 5.74-4.89 5.74-5.61  _  —  —  21  0.99-0.83  -  _ 1  73  _ 8/11  Mayl2  _ 29  0  -  0/9  -  Mayl2  -  29  21  6.06-6.00  0.35 0.38 0.03 0.03  70 83  7/10 5/6 0/9 0/6  June5 Mayl2 June5 Mayl2  15 39 15 39  12 12 12 12  1.09-0.26 1.16-0.26 6.33-6.42 6.35-6.43  0.60 6.54 6.53  0.31 0.28 0.05 0.06  80 90  June5 Mayl2 June5 May 12  21 45 21  9  8/10 9/10 1/9 1/11  45  24 24 24 24  1.23-0.51 0.97-0.32 6.62-6.50 6.48-6.58  0.56  0.26  12  Julyl4*  16  18  0.96-0.30  -  -  1/8  0  0/9  Julyl4*  16  18  7.14-6.47  5  Ta Tb Ca Cb  -  -  -  —  —  —  —  —  6 6 6 6  Ta Tb Ca Cb  0.54 0.46  0.21 0.12 0.03 0.03  4/7  7/10 2/10  Julyl4* Sept.9 Sept.9 Sept.9  87  70 0 20  31 31 31  12 12 12 12  0.84-0.33 0.64-0.38 6.94-6.85 6.94-6.86  7  Ta Tb Ca Cb  0.08 0.03 0.37 0.35  70 100 10 0  7/10 9/9 1/10 0/10  Sept.9 Sept.9 Sept.9 Sept.9  52 52 52 52  12 12 12 12  0.63-0.75 0.90-0.85 6.95-6.42 6.85-6.35  3  3 3  4 4  4  4  5 5  5  7 7 7  -  6.81  6.90 6.89 0.69  0.87 6.69 6.60  0.22  -  0 0  11  57  -  0/7  -  -  -  -  -  80  Table 5: Low oxygen tolerance experiments on CVs collected from the Strait of Georgia, continued. Duration of Experiment (hours) 12 12 12 12  Range of Oxygen (mL L" ) 1.07-1.04 0.79-0.87 6.56-6.28 6.56-6.30  8 8 8 8  Ta Tb Ca Cb  1.06 0.83 6.42 6.43  0.02 0.06 0.20 0.18  33 78 0 0  2/8 7/9 0/10 0/9  9 9 9 9  Ta Tb Ca Cb  0.92 0.71 6.90 6.84  0.04 0.04 0.06 0.05  100 100 30 0  10/10 10/10 3/10 0/10  Oct.14 Nov.12 Oct.14 Nov.12  57 28 57 28  12 12 12 12  0.94-0.89 0.73-0.68 6.94-6.85 6.87-6.80  10 10 10 10  Ta Tb Ca Cb  2.08 2.02 6.82 6.81  0.08 0.08 0.02 0.01  43 71 0 0  S/7 6/8 0/8 0/7  Nov.12 Sept.9 Nov.12 Sept.9  38 103 38 103  12 12 12 12  2.02-2.14 1.96-2.08 6.83-6.80 6.81-6.80  11 11 11 11  Ta Tb Tc Td  0.95 0.94 2.11 2.24  0.12 0.12 0.28 0.24  20 0 0 0  2/10 0/10 0/10 0/9  (1986) June16 June16 Junel6 Junel6  3 3 3 3  12 12 12 12  0.86-1.03 0.85-1.02 1.91-2.31 2.07-2.41  12 12 12 12  Ta Tb Ca Cb  1.88 2.18 4.20 4.24  0.01 0.03 0.04 0.16  0 0 0 0  0/8 0/9 0/11 0/9  Junel6 Junel6 June16 Junel6  6 6 6 6  12 12 12 12  1.87-1.89 2.16-2.20 4.17-4.22 4.13-4.35  13 13 13 13  Ta Tb Ca Cb  0.59 0.47 5.42 5.50  0.06 0.10 0.06 0.31  20 50 0 0  2/10 5/10 0/9 0/10  Junel6 Junel6 June16 Junel6  8 8 8 8  12 12 12 12  0.55-0.63 0.40-0.54 5.37-5.46 5.28-5.72  Oxygen (mL L ) Mean s.d. 1  Mortality Ratio  Period of Captivity (days) 15 15 15 15  Date Collected (1985) Nov.12 Nov.12 Nov.12 Nov.12  Experiment # Funnel  %  1  Table 6: Dissolved oxygen concentrations (mL L ) at Saa9 in 1985. 1  Depth  Jan  Feb  Mar  Apr  May  Jul  Aug  Sep  Oct  Nov  H 0 10 20 30 50 75 100 120 140  4.61 4.98 5.13 5.08 4.46 0.25 0.05 0.06  5.37 5.29 4.95 5.02 4.51 3.12 0.06 0.09  5.91 5.54 5.51 5.10 4.61 4.66 3.55 1.91  6.51 5.75 5.45 5.18 4.25 3.76 2.78 1.06  9.05 6.17 5.78 5.56 4.63 3.38 3.11 0.72  5.15 4.76 5.10 4.44 3.16 2.33 1.63 0.90  3.99 3.74 3.70 3.63 2.38 1.17 0.84 0.31  4.70 3.59 3.83 3.42 0.20 0.49 0.46 0.62  4.83 4.77 2.93 2.47 0.41 0.17 0.24 0.51  3.81 3.69 3.53 3.37 3.10 0.31 0.64 0.63  Table 7: Dissolved oxygen concentrations (mL L Depth H 0 10 20 30 50 75 100 120 140  _ 1  ) at Saa9 in 1986.  Mar  Apr  May  Jon  Jul  Aug  Sep  Oct  Nov  Dec  5.42 5.19 5.29 4.96 5.24 2.31 1.92 1.18  6.19 5.76 5.67 5.48 1.94 1.93 1.22 0.35  7.03 5.85 5.74 5.54 4.69 3.11 1.37 0.67  7.18 6.74 6.46 5.16 4.29 2.63 1.10 0.69  4.91 4.43 4.40 4.14 3.03 2.59 1.70 0.20  8.96 7.37 4.33 3.58 2.51 2.08 2.22 0.75  4.63 3.48 3.44 3.50 0.77 1.81 1.09 0.98  4.59 3.17 3.22 3.47 2.96 1.10 0.89 1.24  4.40 4.05 3.59 3.37 2.61 1.08 0.96 1.18  4.66 4.31 4.48 4.09 3.24 2.20 0.75 0.88  Table 8: Hydrographic data collected August 5, 1986, at station Saa3. Depth  Temperature (C) H 140 8.97 150 • 9.02 160 9.04 170 9.15 180 9.10 190 9.11 200 9.19  Salinity (ppt) 31.028 31.058 31.076 31.090 31.092 31.105 31.114  Density (sigma-t) 24.02 24.03 24.04 24.04 24.04 24.05 24.05  Oxygen (mL L ) 0.78 0.76 0.53 0.31 0.76 0.56 0.52 - 1  Table 9: Dissolved oxygen concentrations (mL L  )  - 1  Sep  Oct  Nov  4.70  3.08  3.95 3.56  2.89 2.57  6.56 5.38 2.67  3.75 3.60  4.24 2.49  2.77 1.13  0.74  0.29 0.06  1.92 0.19 0.25  2.04 0.51 0.16 0.00 0.07  2.33 1.59 0.68  Jan  Feb  Mar  Apr  May  Jul  Aug  0 10 20  3.69 4.09  6.09 5.22  6.77  10.08 7.04  SO  3.74  5.13 4.68 4.15  3.79 3.79  3.21 3.28  5.34 4.53 2.88  6.66  50 75 100 120  4.77 3.84 3.23 3.69  6.98 6.19 5.47  4.94 2.79  3.94  2.01 1.51  1.32 0.89  0.59 0.00 0.00 0.00  0.20 0.00 0.00 0.00  0.06 0.00  Depth  at Saa0.8 in 1985.  (m)  140 160 180 190  0.87  2.54  0.12 0.02  0.11 0.00  0.04 0.00 0.00  0.00 0.00 0.00  5.49  0.00 0.00  0.59 0.20 0.00 0.00 0.00  0.03 0.00 0.00 0.00  Table 10: Dissolved oxygen concentrations (mL L ) - 1  Depth  H  0 10 20 30 50 75 100 120 140 160 180 190  Mar  .  5.36 5.25 4.95 4.75 4.08 2.48 1.36 0.67 0.27 0.24 0.50  Apr  -  6.37 6.28 5.57 5.11 4.20 1.71 0.48 0.35 0.15 0.11 0.11  May  Jun  Jul  .  .  •  6.05 5.87 5.80 4.63 3.44 1.34 0.32 0.20 0.08 0.14 0.08  Aug  -  Sep  -  5.53 5.44 4.45  6.96 5.55 4.48  4.10 3.33 3.33  3.35 0.86 0.37 0.18 0.14 0.09  3.29 1.83 1.23 0.65 0.19 0.10 0.07  3.68 1.28 0.48 0.38 0.13 0.04 0.00  2.82 0.31 0.30 0.16 0.00 0.12 1.09  0.06  0.00  0.00  1.21  7.26 7.22 6.77 5.29  0.02 0.08 0.26 0.41 0.71  0.17 0.39 0.38  3.38  0.14 0.14 0.28 0.35 0.43  at Saa0.8 in 1986. Oct  -  Nov  -  4.34 2.95 2.64 2.24  4.48 3.23 3.07  0.65 0.26 0.18 0.18 0.33 0.68 0.65  1.44 0.20 0.17 0.18 0.36 0.38 0.42  2.56  Dec  -  3.73 3.73 3.94 3.43 2.17 1.48 0.64 0.19 0.16 0.22 0.18  Table 11: Dissolved oxygen concentrations (mL L ) at G1545 and G1748 in 1  1985. Depth  Feb  Mar  Apr  May  Jul  Aug  Sep  Oct  Nov  6.91 6.84 6.57  6.87  7.80 7.35  7.48  6.71  6.48  6.41  6.00 4.93  6.47 4.73  4.91 3.66  6.23  6.45 6.24  6.14 5.60  4.72 4.79 4.98 4.98 4.83 4.89  4.46 4.25 4.31 4.56 4.66  4.05 3.90 3.87  4.43 4.65 4.75  5.75 4.84 4.68 4.66 5.02 4.89 4.98 4.98  4.63 4.63 4.54  3.93 4.09 4.29 4.36 4.11  3.62 3.56 3.55 3.59 3.82 4.02 3.82 3.54  3.49 3.51 3.50 3.51 3.44  5.55 3.46 3.21 3.34 3.78 3.65 3.42 3.35  -  -  -  -  -  -  -  -  (m)  -  0 5 10 20  6.41 5.99 5.74 4.90 4.89  30 50 75 100 150  4.95 5.38 5.07 4.12 3.12 2.77  200 250 300 400  4.93 4.92 4.77  6.55 6.31  5.64 5.09 4.59  Table 12: Dissolved oxygen concentrations (mL L Depth  1  3.72 3.45 3.31 3.23  5.91  ) at G1545 in 1986.  Feb  Mar  Apr  May  Jun  Jul  Aug  Sep  Oct  Nov  Dec  6.87 6.94 6.47 5.95 5.57 4.97  6.39 6.18 5.45 5.32 5.27  7.34 6.39 5.89 5.52 5.28 5.06 5.20 4.76 4.31 4.04 4.02  6.81 6.36 5.53 5.44 5.02 4.90 4.97 4.72 4.43 4.28 4.05  8.35 7.55 6.42 5.14 4.76 4.71  7.54 6.16 4.97 4.90 4.53 4.48 4.46 4.40 4.19 4.04 3.99  6.85 7.68 4.73 4.51 4.38 4.14 4.09 4.09 4.05 3.92 3.87  6.96 6.52 4.23 4.09 3.96 3.96 3.67 3.59 3.62 3.67 3.51  7.63 6.75 4.54 4.08 3.69 3.44 3.44 3.40 3.52 3.34 3.33  5.68 5.62  6.31 6.15 5.92 5.58 5.43 3.67  H 0 5 10 20 30 50 75 100 150 200 250 300  4.70 4.38 4.36 4.21 4.13  4.94 4.68 4.51 4.52 4.21 3.99  5.11 4.63 4.62 4.05 4.05  5.52 5.17 3.77 3.29 3.37 3.34 3.27 3.22 3.15  3.43 3.63 3.51 3.54 3.27  Table 13: Vertical plankton hank collected in 1985. Date  Station  May21 May21  Sate Sate  Depth (m)  Sample Number  CVI  CV 12 55  0-15  17  0  0-70  18  0  CIV  Total  Number  8  20  11.56  76 26  15.03  9.42  May21  Saa3  0-15  15  0  19  21 7  May21  Saa3  0-120  16  0  47  6  53  3.83  July4 July3  G1748  0-150  31  0  1  0  1  0.06  July3  Sate Saa3  0-70 0-150  30 23  0 0  0 125  0 0  0 125  0 7.22  Ang8 Ang8 Aug8 Ang7 Ang7  G1748 G1748 Sate Saa3 Saa3  0-150 0-375 0-70 0-50 0-190  46 45 44 39 38  0 0 0 0 0  3 1566  3 1566  0.17 60.18  0 0 24  0 0 0 0 0  0 0 24  0 0 1.49  Septl8  G1748 G1748 Sate Saa9  0-200  59 60  0  3 652 0 1 2 0 1 0 0 0  0 0 0 0 0 0 0 0 0 0  3 652 0 1 2 0 1 0 0 0  0.13 31.41 0 0.09 0.29 0 0.10 0 0 0  1 724 0 0 0 0 0 1 0 0  0  1  0.06  0 0 0 0 0 0 0 0 0  724 0  29.20 0 0 0 0 0 0.12 0 0  Sept18 Septl8 Sept17 Sept17 Septl7 Sept17 Sept17 Sept17 Septl7  Saa9 Saa3 Saa3 Saa0.8 Saa0.8 Saa0.8  Oct9 Oct9 Oct8 Oct8 Oct8 Oct8 Oct8  G1545 G1545  Oct8 Oct8 Oct8  Saa3 Saa0.8 Saa0.8  Sate Saa9 Saa9 Saa3 Saa3  200-380 0-70 0-100 100-160 0-100  58 51 52 56 57 53 54 55  0 0 0 0 0 0 0 0 0  150-365 0-70 0-100  69 70 68 62  0 0 0 0  100-140 0-100 100-140 140-210 0-100 0-200  61 65 66 67 64 63  0 0 0  100-190 0-75 75-140 140-190 0-150  0 0 0  0 0 0 0 1 0 0  Table 13: Vertical plankton hauls collected in 1985, continued. Depth (m)  Sample Number  CVI  0-100 100-394 0-68  82  0  81 85 84 83  Saa0.8 Sci  0-100 100-150 0-100 100-210 0-195 100-195 0-100  0 0 0 0  74 73 71 72 76  0 0 0 0 0  Sci  100-245  75  0  Decl7 Decl7 Decl8  G1545 G1545 Sate  0-100 100-385  D12 D13  0 0  D7  Decl6 Decl6 Decl6 Decl6 Decl6 Decl6  Saa9 Saa9 Saa3 Saa3 Saa0.8 Saa0.8  0-65 0-100 100-150 0-100 100-210 0-100 100-190  0 0 0  Station  Date Nov6  G1748 G1748 Sate Saa9  Novo Nov7 Nov7 Nov7 Nov4 Nov4 Nov4 Nov4  Saa9 Saa3 Saa3 Saa0.8  Nov5 Nov5  CV  CIV  Number  Total  (m- ) 0 31.03 3  Dl D2 D3 D4 D5 D6  0 0 0 0  0 1052 0 0 0 0 0 0 0  0 0 0 0 0 0 0 0  0 282  0 0 0 0 0 0 0 0 16.86  0  2  0.17  0  320 0 0 0 0 1 0  9.74  0 0 0  0 282 2 320 0 0 0 0  0 0 0 0 0 0 0  1 0 4  0 1052 0 0 0 0 0 0 0  0 0 0 0 0.08 0 0.39  4  Table 14: Variation between replicate vertical haul samples.  Sample Number Fel8,20 A3,6 A2,5 A17.21.18 Ma5,8 Ma6,9 Mal5,20 Mal6,17 Mal8,19 Jel3,14 Jel5,16 Jel,4 Je3,6 JU,2 JU6.17 J112.13 J114.15 Au5,6 Au7,8 Aul6,17 Sel,2 Se7,8  Depth Sampled (m) 80-245 0-80 80-160 0-80 0-80 80-160 0-80 80-200 200-380 80-120 120-208 0-80 300-395 100-170 0-150 150-250 250-385 100-175 175-205 300-395 0-120 160-208  Coefficient of Variation  80% Confidence Interval Upper Lower  Station  %  Sci Saa0.8 Saa0.8 G1545 Saa0.8 Saa0.8 G1545 G1545 G1545 Saa0.8 Saa0.8 G1545 G1545  12.9 0 141.4 83.0 47.1 70.7 7.8 44.5  m" ) 1.5 0.1 0.1 37.2 0.1 0.03 24.0 0.4  58.9 141.4 141.4 34.0 27.9  0.2 0.3 0.2 4.4 24.7  0.8 0.8 33.7 3.0 3.6 7.6 7.0 20.0 84.8  Saa0.8 G1545  141.4 141.4 93.5 32.4 0 25.4 16.9 70.7 47.1  0.5 0.1 0.1  13.1 2.0 29.3  56.3 0.5 3.1 52.8 0.03  239.4 0.5 9.4 110.6  G1545 G1545 Saa3 Saa3 G1545 Saa0.8 Saa0.8  (#  3  0.1  (#  Mean  m" ) 2.7 0.1 3.8 232.7 3  1.1 0.7  (#  m~ ) 2.0 3  0.1 0.05 120.6 0.3 0.2 28.6 1.3 1.2 0.1 0.1 10.2 80.6 0.2 0.03 5.1 125.2 0.5 5.6 77.9 0.5 0.3  Table 15: Vertical plankton hauls collected in 1986. Depth (m) O-lOO  Sample Number  CVI  CV  crv  Total  J20  0  0  0  Sate Saa9  100-395 0-70 0-100  Jan29  Saa9  100-150  J21 J22 J23 J24  224 0 0 0  0 0 0 0  0 0 0 0  0 224  Jan29  Saa3 Saa3  0-100  J25 J26  0  0  1 0  1 0  0 0  1  1  0  0 0 0 0 0 0 0 0 0 1 0 0 0 0  Date  Station  Jan28 Jan28  G1545 G1545  Jan29 Jan29  Jan29 Jan27 Jan27 Feb20  Saa0.8 Saa0.8  100-210 0-100 100-210  (m"»)  G1545 G1545  80-300  G1545 Sate Saa9 Saa9 Saa3 Saa3 Saa0.8 Saa0.8 Sci Sci Sci Sci  300-385 0-65 0-80 80-150 0-80 80-205 0-80 80-192 0-80 0-80 80-245 80-245  F13 Fll F9 F10 F8 F7 F2 FI F17 F19 F18  124 81 0 0 0 0 0 0 0 0 0 35  F20  42  Marll  G1545 G1545  Mh20 Mhl9  15 136  MarlO MarlO MarlO MarlO MarlO  Sate Saa3 Saa3 SaaO.8 SaaO.8  0-200 200-395 0-70 0-80 80-215 0-80 80-200  Mhl4 Mhl2 Mhl3 Mhll MhlO  0 0 1 0 0  Aprl5 Aprl5  G1545 G1545 G1545 G1545 G1545  0-80 0-80 0-250  A17 A21 A18  80-250 80-250 250-390  A19 A22 A20 A16 A14 A15 A13 A12  0 0 0 0 2 2 0 0 0 0 0  Feb20 Feb20 Febl7 Febl7 Febl7 Febl7 Febl7 Febl7 Febl7 Feb20 Feb20 Feb20 Feb20 Mar 11  Aprl5 Aprl5 Aprl5 Aprl5 Aprl4 Aprl4 Aprl4 Aprl4 Aprl4  G1545 Sate Saa9 Saa9 Saa3 Saa3  0-80  J14 J15  0-72 0-80 80-155 0-160 160-213  F12 F14  Number  0 0 0 0 0 0 0 0  0 0 0  0 6.58 0 0 0  0  0  2 0 2  0.16 0 0.16  0 124  0 4.89  81 0 0 0 0 0 0 1  8.26 0 0 0 0 0 0 0.08 0 0 1.84 2.21  0 0 0 0 0 0 0 0  0 0 35 42  0 0 0 0 0 0 0  0 0  15 136  0 0 0 0 0  0 0 1 0 0  101 25 20  2080 420 716  2181 445 736  236.29  4 7 0 0 4 0 2 0  16 15 2 26 4 0 4 0  20 24  1.02 1.22 0.25 3.13 0.87 0 0.33 0  4 26 8 0  6 0  0.65 6.05 0 0 0.06 0 0  48.21 25.53  Table 15: Vertical plankton hauls collected in 1986, continued. Sample Number A3 A6  CVI  CV  crv  Total  Number  0 0  0 0  I I  1 1  A2 A5  0 1 0  0 0  0 0  0 1  0  0  0  0 0  0 0  0.11 0.11 0 0.11 0  0 0 0  267  n  243 20  6 3  0 0  11 14  1 0  1 0 0 0 0  33 3 17 11 1 0 0 0 1 0 0  0 5 8 5 0  71 116 27 203 1300 872 0 4 20 3 14 0  0 1 0 0 0 0 0 0  0 0 1 0 2  0 0 0 0 0  Date  Station  Depth (m)  Aprl4 Aprl4  SaaO.8 SaaO.8  0-80 0-80  Aprl4 Aprl4 Aprl4  SaaO.8 SaaO.8 SaaO.8  80-160 80-160 160-205  Aprl4  SaaO.8  160-205  Al A4  Mayl3 Mayl3 Mayl3 Mayl3  G1545 G1545 G1545 G1545  0-80 0-80 80-200 80-200  Mal5 Ma20 Mal6 Mal7  May 13 Mayl3 Mayl2 Mayl2 May 12 May 12 May 12 May 12 May 12 Mayl2 May 12 Mayl2  G1545  200-380 200-380  Mal8 Mal9 Mal4 Mal3 Mall Mal2 Ma5 Ma8 Ma6 Ma9 Ma7 MalO  Jun3 Jun3 Jun3 Jun3 Jun3 Jun3 Jun4 Jun5 Jun5 Jun5 Jun5 Jun5 Jun5 Jun5 Jun5 Jun5 Jun5  G1545 Sate Saa9 Saa3 Saa3 SaaO.8 SaaO.8 SaaO.8 SaaO.8 SaaO.8 SaaO.8 G1545 G1545 G1545 G1545 G1545 G1545 Sate Saa9 Saa9 Saa3 Saa3 SaaO.8 SaaO.8 SaaO.8 SaaO.8 SaaO.8 SaaO.8  0-72 0-155 0-80 80-214 0-80 0-80 80-160 80-160 160-200 160-210 0-80 0-80 80-200 200-300 200-395 200-395 0-70 0-120 120-157 0-120 120-215 0-80 0-80 80-120 80-120 120-208 120-208  Jel Je4 Je2 Je7 Je3 Je6 Je8 Je9  0 0 0 0 0 0  Jell Jel2 Jel3  0 0 . 0 0 0 0 0 0 0 0 0 0 0 0  Jel4 Jel5 Jel6  0 0 0  JelO Je21 Je22  2 4 3 0 0 0  0 0 0 0  0  14  30.12 26.98 1.66 0.87 0.67  34 8 25 16 1 2 4 3 1 0 0  1.64 0.96 1.40 1.73 0.07 0.22 0.43 0.33 0.11 0 0  71 117 27 203 1300 872 0 4 20 3 14 0  7.69 12.68 1.95 17.61 57.80 38.77  278 249 23 12  0 0 1 0 2  0 0.29 4.69 0.22 1.28 0 0 0 0.22 0 0.20  Table 15: Vertical plankton hauls collected in 1986, continued. Date  Station  Depth (m)  Sample Number  CVI  CV  CIV  Total  Number  Jull5 Jull5  G1545 G1545  0-150  J116 J117  0 0  1 0  0 0  1 0  0.06 0  Jull5 Jull5 Jull5  G1545 G1545  J112 J113  0 0  98 20  0 0  8.50 1.73  J114 J115  0 0  2396 1502  0  98 20 2396  Jill J110 J19 J17 J18 J13  0 0 0 0 0 0  0 0 61 0 12 0  0 0 0 0 0 0  0  0 0 7.05 0 1.10 0  0 0 0 0 0  0 0 3 0 0  0 0 0 0 0  0 0 3 0 0  0 0 0.37 0 0  0 0 0 0 0 0 0 0 0  1 576 956 752 0 0 3 0 0  0 0 0 0 0 0 0 0 0  0  4  0.04 49.96 87.23 68.61 0 0 0.49 0 0 0.46  0 0 0 0 0 0 0 0 0 0 0 0  4 16 23  0 0 0 0 0 0 0 0 0 0 0 0 0  1 576 956 752 0 0 3 0 0 4 4 16 23 0 2 0 130 0 0 1 0 35  0.46 4.62 6.65 0 0.17 0 32.18 0 0 0.14 0 3.80  Jull5 Jull4 Jull4 Jull4 Jull4 Jull4 Jull4 Jull4 Jull4 Jull4 Jull4 Jull4 Aug6 Aug6 Aug6 Aug6 Aug5 Aug5 Aug5 Aug5 Aug5 Aug5 Aug5 Aug5 Aug5 Aug5 Aug5 Aug7 Aug7 Aug7 Aug7 Aug7 Aug7 Aug7  G1545 G1545  0-150 150-250 150-250 250-385 250-385  Sate Saa9 Saa9 Saa3 Saa3  0-70 0-80 80-155 0-100 100-195  Saa0.8  0-100  Saa0.8 Saa0.8 Saa0.8 Saa0.8 Saa0.8  0-100 100-170 100-170 170-208 170-208  J14 Jll J12  G1545  0-200 200-300 300-395 300-395 0-70 0-100 100-153 0-100 0-100  Aul9 Aul8 Aul6 Aul7 Aul5 Aul4  G1545 G1545 G1545 Sate Saa9 Saa9 Saa3 Saa3 Saa3  100-175  Saa3  100-175  Saa3 Saa3 Saa0.8 Saa0.8 Scla Scla Sc2a Sc2a Sc2a Sc2 Sc2  175-205 175-205 0-100 100-203 0-175 175-210 0-175 175-205 205-265 0-205 205-285  J15 J16  Aul3 Au3 Au4 Au5 Au6 Au7 Au8 Au2 Aul Au20 Au21 Au24 Au23 Au22 Au28 Au29  0 2 0 130 0 0 1 0 35  0  1502 0 0 61 0 12  153.88 96.47  Table 15: Vertical plankton hauls collected in 1986, continued. Date Sep9 Sep9 Sep9 Sep9 Sep8 Sep8 Sep8 Sep8 Sep8 Sep8 Sep8 Sep8 Sep8 Sep8 Sep8 Sep8  Station G1545 G1545 G1545 G1545 Sate Saa9 Saa9 Saa3 Saa3 Saa0.8 Saa0.8 Saa0.8 Saa0.8 Saa0.8 Saa0.8 Saa0.8  Depth  Sample  (m) 0-120 0-120 120-250 250-372 0-72 0-100  Number  100-150 0-120 120-210 0-50 0-120 0-120 120-160 120-160 160-208 160-208  Se20 Se21 Se22  CVI  CIV  Total  Number (m- ) 3  Sel9 Sel8 Sel7 Sel6 Sel5 Se4 Sel Se2 Se5 Se6 Se7  0 0 0 0 0 0 0 0 0 0 0 0 0 0 0  Se8  0  Se23  CV  2 502 0 0  0 0 0 0 0 0  1 0 1 0 2 6 0 0 1 2  0 0 0 0 0 0 0 0 0 0  0 0  0 0 2 502 0 0 1 0 1 0 2 6 0 0 1 2  0 0 0.13 35.68 0 0 0.17 0 0.07 0 0.25 0.74 0 0 0.18 0.36  Table 16: Horizontal plankton hauls collected at station Saa3 in 1985. Date  Depth (m)  Sample Number  CVI  CV  crv  Total  Number (m- ) 3  Volume Filtered (m- ) 3  M a y 21 May 21  1 6  May21 May21  5 5 5 50  May 21 May 21 May21  50 100 100  8 3 9  May21 May21 May 21  120 120 150  4 10 5  0 0 0  July3 July3 July3 July3 July3 July3 July3 July3  5 50 100 100 120 120 150 150  26 22 21 27 20 28 19 19  0 0 0 0  Aug7 Aug7 Aug7 Aug7 Aug7 Aug7 Aug7 Aug7 Aug7 Aug7  110  37 36 50 49 48 47 43 42 41 40  130 140 145 150 155 160 170 180 190  7 2  0 0 0  3 6 10  0 0  1 1  0 0  0 0 18 1 96  2 4  5 10  3.00 7.58  4 2  14  10.69 1.01 1.71  3 1  0 0 0  0 0  0 0 0  18 1 96  0 0 5.10 1.98 33.27  0 0 0 0 0 0 317  0 0 0 0 0 0 0  0 0 0 0 0 0 317  0 0 0 0 0 0 75.61  0  0  0  0  0  0  0  0 0 0 0 0 0 0 0 0  2 3 2 0 3 25 0 0  0 0 0 0 0 0 0 0 0  0 2 3 2 0  0 0 0 0  0  3 25 0 0 0  0 0.42 0.74 1.18 0 1.49 6.04 0 0 0  1.67 1.32 1.31 2.97 0.58 3.17 0.55 3.53 0.50 2.89 2.43 4.36 5.93 4.70 4.95 6.14 4.19 0.55 4.40 4.81 4.07 1.70 1.63 2.01 4.14 4.33 4.40 4.44  91  Table 17: Horizontal and * oblique plankton hauls collected in 1986. Date  Station  Depth (m)  Sample Number  CVI  CV  crv  Total  Number (m- ) 3  Volume Filtered (m- ) 3  Mar5* Mar5* Mar5*  0-25 25-75 75-100  Mh5 Mh4 Mh3  0 0 0  0 0 0  0 0 0  100-150  Mh2  150-200  Mhl Mh9 Mh8  0  0  0 0  0 0  l 0  0 0 0  0 0 0 0  0  3  0 0 0 0.02  20.8 41.6 27.8 55.5 63.3  0  0 0 0  0 0  0 0  8.86 8.22  3  0.60  5.04 2.59 1.25 4.74  0 0 0  Mar5*  Saa3 Saa3 Saa3 Saa3  Mar5* MarlO  Saa3 Saa3  MarlO MarlO MarlO  Saa3  5 20  Saa3 Saa3  50 90  Mh7 Mh6  0 0 0  Aprl4 Aprl4 Aprl4 Aprl4  Saa0.8 Saa0.8 Saa0.8 Saa0.8  5 40 150 160  A10  0  A9 A8 A7  0 0 0  0 0 0  0 0 0  0 0 0  0 0 0  Mayl2 Mayl2  Saa0.8 Saa0.8  40 150  Ma3 Ma4  0 0  1 0  3 0  4 0  0.62 0  6.42 9.03  Jun5 Jun5 Jun5 Jun5  Saa0.8 Saa0.8 Saa0.8 Saa0.8  90 100 110 120  Je20 Jel9 Jel8 Jel7  0 0 0 0  2 0 3 0  0 0 0 0  2 0 3 0  0.34 0 0.43 0  5.41 6.43 7.03 6.44  Jull6 Jull6  Saa9 Saa9 Saa9 Saa9  120 120 130 130  J142 J137 J136 J135  0 0 0 0  2 19 0 0  0 0 0 0  2 19 0 0  0.29 3.56 0 0  6.98 5.34 5.76 4.58  Saa9 Saa9  135 135 135 135 135  J134 J121 J125 J129 J133  0 0 0 0 0  0 0 8 18 14  0 0 0 0 0  0 0 8 18 14  0 0 1.55 2.52 2.87  0.12 2.72 5.16 7.13 4.88  Saa9 Saa9 Saa9  140  J120  0  35  0  2.75  0 0  10  0  140  J124 J128  35 10  12.72  140  0  34  2.55 6.07  Saa9 Saa9  140 140  J132 J140  0 0  34 57 31  3.92 5.60  0 0  57 31  11.27 4.31  5.06 7.20  Julie Julie Julie Jul 16 Jull6 Jull6  Julie Jull6  Julie Julie Julie Jull6  Saa9 Saa9 Saa9  1 0 0  8.54 7.50  Table 17: Horizontal and * oblique plankton hauls collected in 1986, continued. Date  Station  Depth (m)  Sample Number  CVI  CV  crv  Total  Number (m- ) 3  Volume Filtered (m- ) 3  0 0 0  7 43 76  4.72 10.70 10.80  0 0  7 43 76 86 28  0 0  86 28  18.87 4.56  1.48 4.02 7.04 4.56 6.14  5 0 9 4  0 0 0  5 0 9  J130  0 0 0 0  0  4  2.28 0 1.25 2.95  2.19 0.18 7.19 1.35  160 170 180 190 170 180 230  Aul2 Aull Aul6 Au9 Au27 Au26 Au25  0 0 0 0 0 0 0  1 11 16 0 1 0 0  0 0 0 0 0 0 0  1 11 16 0 1 0 0  0.25 2.96 3.37 0 0.32 0 0  3.95 3.72 4.74 0.25 3.10 3.65 4.30  110 115 120  Sel4 Sel3 Sel2  0 0 0  0 0  0 0 0  0 0 1  0 0 0.18  3.49 7.81 5.41  Jull6 Jull6 Jull6  Saa9 Saa9 Saa9  145 145 145  J119 J123 J127  0 0 0  Jull6 Jull6  Saa9 Saa9  145 145  J131 J139  Jull6 Jull6 Jull6 Jull6  Saa9 Saa9 Saa9 Saa9  150 150 150  J118 J122 J126  150  Aug5 Aug5 Aug5 Aug5 Aug7 Aug7 Aug7  Saa3 Saa3 Saa3 Saa3 Sc2a Sc2a Sc2a  Sep8 Sep8 Sep8  SaaO.8 SaaO.8 SaaO.8  1  T a b l e 18. A N O V A - S t r a i t s p r i n g a n d summer. Transformed Fall  water  Spring/Summer(1986)  4.35637 3.65804 3.73308 4.02164  Analysis Source  Apr Apr Apr May May Jun Jun Jul Jul  4 .66318 3 .74840 3 .42746 3 .15982 3 .10584 4 .25698 3 .99294 4 .77987 4 .36907  of variance table Sum o f squares  SEASON 0.33564E-01 MONTH 2.7941 SEA*MON 1 .3480 Resid 0.30449 Total 4.4801 T e s t t e r m : RESIDUAL COUNT  versus  column t o t a l c o u n t s .  (1985)  Aug Sept Oct Nov  of G e o r g i a c o u n t s , f a l l  DF 1 3 3 4 1 1  Mean square 0.33564E-01 0.93137 0.44933 0.76123E-01  O v e r a l l mean 4.4483  F-ratio  Prob.  0.44092 12.235 5.9027  0.54300 0.01750 0.05961  Overall standard deviation 0.63819  Homogeneity o f v a r i a n c e t e s t  Factors SEASON  Bartlett Chi-square 1.2057  Probability 0.27218  DF  M u l t i p l e range tests-SEASON F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y  Size warn < 10 0.54300  Homogeneity o f v a r i a n c e t e s t  Factors MONTH  Bartlett Chi-square 0.67486  Multiple  range  Probability 0.87910  DF 3  Size warn < 10  tests-MONTHS  S c h e f f e t e s t a t 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( .2, .1; .3 ) ( . 1, .3, .4 ) B o n f e r r o n i t e s t a t 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( .2, .1, .3 )  as follows  as follows  ( .1, .3, .4 ) M i n i m a l t e s t a t 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( .2, . 1 , .3 ) ( . 1 , .3, .4 ) Homogeneity of v a r i a n c e Factors SEA,MON  Bartlett Chi-square 2.1248  test  Probability 0.54690  DF 3  M u l t i p l e r a n g e tests-SEASON,MONTH F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y  Size warn < 10 0.05961  as  follows  T a b l e 19. A N O V A - S t r a i t f a l l 1986.  of Georgia counts, f a l l  T r a n s f o r m e d w a t e r column t o t a l (1985) (1986) Aug 4.35637 Aug 4.33619 Sep 3.65804 S e p 4.21410 Sep 3.47125 Analysis Source  of v a r i a n c e t a b l e Sum o f DF squares  YEAR MONTH YR*MON Resid Total  0. 18660E-01 0.67148 0.31840E-02 0.74530E-02 0.68212 O v e r a l l mean 4.0072  COUNT Homogeneity Factors YEAR  1. 1 . 1 . 1 . 4.  counts.  Mean square 0.18660E-01 0.67148 0.31840E-02 0.74530E-02  F-ratio  Prob.  2.5036 90.095 0.42721  0.35881 0.06682 0.63145  Overall standard deviation 0.41295  of v a r i a n c e t e s t  Bartlett Chi-square Probability 0.27670E-02 0.95805  DF 1  M u l t i p l e range t e s t s - Y E A R F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y Homogeneity  1985 v e r s u s  Size warn < 10 0.35881  of v a r i a n c e t e s t  Bartlett Factors Chi-square MONTH 0.30163  Probability 0.58286  DF 1  Size warn < 10  M u l t i p l e r a n g e tests-MONTH F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y  0.06682  M u l t i p l e r a n g e tests-YEAR*MONTH F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y  0.63145  T a b l e 20. ANOVA-Saanich I n l e t and A u g u s t , 1985 a n d 1986.  counts  f o r S a a 3 , May, J u l y ,  T r a n s f o r m e d w a t e r column t o t a l c o u n t s , (1985) (1986) May 2.21236 1 .76234 May Jul 2.62653 1.64375 Jul Aug 1.88818 1.82056 Aug 1.93318 Aug Analysis  of v a r i a n c e t a b l e  Source  Sum o f squares  YEAR MONTH YR*MON Resid Total  0.31445 0.41358E-01 0.26982 0.63416E-02 0.66837  DF  Mean square  F-ratio  1 . 0.31445 49.586 2. 0.20679E-01 3.2609 2. 0.13491 21.274 1 . 0.63416E-02 6. Overall O v e r a l l mean standard deviation 1.9838 0.33376  COUNT  Prob. 0.08981 0.36462 0.15154  Homogeneity o f v a r i a n c e t e s t Factors YEAR Multiple F-ratio  Bartlett Chi-square 2.3842 range  Probability 0.12257  DF 1  Size warn < 10  tests-YEAR  i s not s i g n i f i c a n t a t p r o b a b i l i t y  0.08981  M u l t i p l e range t e s t s S c h e f f e t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 2. ) ( 1. ) B o n f e r r o n i t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 2. ) ( 1. ) M i n i m a l t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 2. ) ( 1. ) Homogeneity o f v a r i a n c e t e s t Factors MONTH  Bartlett Chi-square 4.9798  Probability 0.08292  DF 2  M u l t i p l e r a n g e tests-MONTH F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y  Size warn < 10  0.36462  as follows  as follows  as follows  M u l t i p l e range t e s t s S c h e f f e t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e i s 1 homogeneous s u b s e t w h i c h i s l i s t e d ( 2 2 , 2 1 , 2 3 , 1 3 , 1 1 , 1 2 ) B o n f e r r o n i t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e i s 1 homogeneous s u b s e t w h i c h i s l i s t e d ( 2 2 , 2 1 , 2 3 , 1 3 , 1 1 , 1 2 ) M i n i m a l t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e i s 1 homogeneous s u b s e t w h i c h i s l i s t e d ( 2 2 , 2 1 , 2 3 , 1 3 , 1 1 , 1 2 )  as  follows:  as  follows:  as  follows:  T a b l e 2 1 . ANOVA-Saanich I n l e t A u g u s t 1986.  counts  T r a n s f o r m e d w a t e r column t o t a l (Saa9) (Sa a3) 1 .43097 Apr Apr 1.51572 May  1.90365  May  1 .76234  Jun  1.88818  Jun  1 .76234  Jul  2.27544  Jul  1 .64375  Aug  1.24573  Aug Aug  1 .82056 1 .93318  Analysis Source  of variance table Sum o f Mean squares DF square  STATION MONTH STN*MON Resid . Total  3.0118 0.36343 1.2731 1.5692 6.3308  COUNT  Homogeneity o f v a r i a n c e Bartlett Chi-square 5.6840  Multiple  range  F-ratio  F-ratio  Prob.  4.7982 0.28950 0.50704  0.06868 0.87333 0.81244  19.  O v e r a l l mean 1.3865  Factors STATION  Overall standard deviation 0.57723  test  Probability 0.05831  DF 2  Size warn < 10  tests-STATION  i s not s i g n i f i c a n t a t p r o b a b i l i t y  0.06868  M u l t i p l e range t e s t s S c h e f f e t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e i s 1 homogeneous s u b s e t w h i c h i s l i s t e d ( 3., 2., 1. ) B o n f e r r o n i t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e i s 1 homogeneous s u b s e t w h i c h i s l i s t e d ( 3., 2., 1. ) M i n i m a l t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e i s 1 homogeneous s u b s e t w h i c h i s l i s t e d ( 3., 2., 1. ) Homogeneity o f v a r i a n c e t e s t Factors MONTH  through  counts. (SaaO .8) 1 .00000 Apr 1 .14870 Apr '1 .37973 May 1 .37973 May Jun 0 Jun 1 .24573 Jul 0 1 .24573 Jul 1 .14870 Aug  1.5059 0.90858E-01 0.15913 0.31385  2. 4. 6. 5.  from A p r i l  Bartlett Chi-square 7.9520  Probability 0.09335  DF 4  as  follows:  as  follows:  as  follows:  Size warn < 10  Multiple F-ratio  r a n g e tests-MONTH i s not s i g n i f i c a n t a t p r o b a b i l i t y  Homogeneity of v a r i a n c e  0.87333  test  STATION,MONTH I n c a l c u l a b l e  due t o s t a n d a r d  M u l t i p l e r a n g e tests-STATION,MONTH F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y  deviation  0.81244  of zero  Table April  22. ANOVA-Saanich I n l e t t h r o u g h A u g u s t 1986.  counts  f o r Saa9 a n d Saa0.8,  T r a n s f o r m e d water column t o t a l c o u n t s . (Saa9) (Saa0.8) 1.51572 Apr Apr 1.00000 Apr 1 . 14870 1.90365 May 1 .37973 May May 1.37973 Jun 1 .88818 0 Jun Jun 1.24573 0 2.27544 Jul Jul Jul 1.24573 1 .24573 Aug 1.14870 Aug Analysis Source STATION MONTH STN*MON Resid Total COUNT  of v a r i a n c e t a b l e Mean Sum o f DF squares square 2.1964 0.48538 1 .0092 1.5629 5.1973  1 . 4. 4. 4. 13.  O v e r a l l mean 1.2412  2.1964 0.12134 0.25229 0.39073  F- r a t i o  Prob.  5. 6214 0. 31056 0. 64570  0.07674 0.85815 0.65897  Overall standard deviation 0.63229  Homogeneity o f v a r i a n c e t e s t Factors STATION  Bartlett Chi-square 0.48875  Probability 0.48449  DF 1  Size warn < 10  M u l t i p l e range t e s t s S c h e f f e t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 2. ) ( 1. ) B o n f e r r o n i t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 2. ) ( 1. ) M i n i m a l t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 2. ) ( 1. )  as follows  as follows  as follows  Homogeneity o f v a r i a n c e t e s t Factors MONTH  Bartlett Chi-square 7.5529  Multiple  range  tests  Probability 0.10940  DF 4  Size warn < 10  F-ratio  i s not s i g n i f i c a n t a t p r o b a b i l i t y  Homogeneity o f v a r i a n c e t e s t STATION,MONTH U n c a l c u l a b l e due t o s t a n d a r d  0.85815 deviation  of zero  Table April  2 3 . ANOVA-Saanich I n l e t t h r o u g h A u g u s t 1986.  counts  f o r Saa3 a n d SaaO.8,  T r a n s f o r m e d w a t e r column t o t a l v a l u e s . (SaaO.8) (Saa3) Apr 1.43097 1.00000 Apr Apr 1.14870 1.76234 May 1.37973 May May 1.37973 Jun Jun 1.76234 0 Jun 1.24573 0 Jul 1.64375 Jul Jul 1.24573 1.82056 Aug 1.14870 Aug Aug 1.93318 Analysis  of variance  table  Source  Sum o f squares  DF  STATION MONTH STN*MON Resid Total  1 .7548 0 .66826 0 .34152 1 .5692 4 .7452  1 . 4. 4. 5. 14.  COUNT  1 .7548 0. 16707 0. 85380E-01 0. 31385  O v e r a l l mean 1.2601  Homogeneity of v a r i a n c e Factors STATION  Mean square  Bartlett Chi-square 5.6262  F-ratio  Prob.  5. 5911 0. 53231 0. 27204  0. 06439 0. 71949 0. 88436  Overall standard deviation 0.58219  test  Probability 0.01769  DF 1  Size warn < 10  M u l t i p l e range t e s t s S c h e f f e t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 2. ) ( 1. ) B o n f e r r o n i t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 2. ) ( 1. ) M i n i m a l t e s t a t 10% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 2. ) ( 1. ) Homogeneity o f v a r i a n c e Factors MONTH  Bartlett Chi-square 5.4340  test  Probability 0.24559  DF 4  Size warn < 10  as  follows  as  follows  as  follows  103  M u l t i p l e range t e s t s F - r a t i o i s not s i g n i f i c a n t at p r o b a b i l i t y Homogeneity o f v a r i a n c e t e s t STATION,MONTH I n c a l c u l a b l e due t o s t a n d a r d  0.71949 deviation  of zero.  104  Table April  24. ANOVA-Saanich I n l e t t h r o u g h A u g u s t 1986.  c o u n t s f o r Saa9 a n d Saa3,  T r a n s f o r m e d w a t e r column t o t a l (Saa9) (Saa3) Apr 1 .51572 Apr 1.43097 May 1 .90365 May 1.76234 Jun 1 .88818 Jun 1.76234 Jul 2.27544 Jul 1.64375 Aug 1.24573 Aug 1.82056 Aug 1.93318 Analysis Source STATION MONTH STN*MON Resid Total COUNT  of  variance table Sum o f DF squares  0.18923E-02 0.28402 0.48468 0.63416E-02 0.77945  1 . 4. 4. 1 . 10.  O v e r a l l mean 1.7438  Homogeneity o f v a r i a n c e Factors STATION  Bartlett Chi-square 2.6370  counts,  Mean square 0. 189E-02 0. 710E-01 0. 121 0. 634E-02  F-ratio  Prob.  0.29839 11.197 19.107  0.68171 0.22006 0.16973  Overall standard deviation 0.27919  test  Probability 0.10440  DF 1  M u l t i p l e range t e s t s F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y Homogeneity o f v a r i a n c e Factors MONTH  Bartlett Chi-square 4.1747  Size warn < 10 0.68171  test  Probability 0.38287  DF 4  Size warn < 10  M u l t i p l e r a n g e tests-MONTH F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y  0.22006  M u l t i p l e range t e s t s F - r a t i o i s not s i g n i f i c a n t at p r o b a b i l i t y  0.16973  T a b l e 25. ANOVA-Saanich I n l e t of G e o r g i a c o u n t s , 1986.  counts  T r a n s f o r m e d w a t e r column t o t a l (G1545) (Saa9) 3.15982 May May 1. 90365 May 3.10584 Jun 1. 88818 Jun 4.25698 Jun 3.99294 4.77987 Jul Jul 2. 27544 4.36907 Jul Analysis Source  of v a r i a n c e t a b l e Sum of Mean square DF squares  STATION MONTH STN*MON Resid Total  7.3856 1 .7918 0.48083 0 . 12069 9.7789  Homogeneity  Factors STATION  of  variance  Bartlett Chi-square 2.0631  Saa9 v e r s u s  F-ratio  Prob. 0.00087 0.01585 0.08988  Overall standard d e v i a t i o n 1.1056  test"  Probability 0.15090  DF 1  Size warn < 10  M u l t i p l e range t e s t s Scheffe t e s t at 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d a s ( 1. ) ( 2. ) Bonferroni t e s t at 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d a s ( 1. ) ( 2. ) Minimal t e s t at 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d a s ( 1. ) ( 2. ) Homogeneity Factors MONTH  of  variance  Bartlett Chi-square 0.70493  Strait  counts.  7.3856 183.58 0.89590 22.269 5.9758 0.24041 0 . 4 0 2 3 1 E - 01  1. 2. 2. 3. 8.  O v e r a l l mean 3.3035  COUNT  at  follows  follows  follows  test  Probability 0.70295  M u l t i p l e range tests-MONTH Scheffe t e s t at 5% p r o b a b i l i t y T h e r e a r e 2 homogeneous s u b s e t s ( . 1 , .2 ) ( . 2 , .3 )  DF 2 level which are  Size warn < 10  l i s t e d as  follows  B o n f e r r o n i t e s t a t 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( .1, .2 ) ( .2, .3 ) Minimal test at 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( .1, .2 ) ( .2, .3 ) Homogeneity of v a r i a n c e Factors STN,MON  Bartlett Chi-square 1.8838  as  follows  as  follows  test  Probability 0.38989  DF 2  M u l t i p l e range t e s t s F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y  Size warn < 10 0.08988  T a b l e 26. ANOVA-Saanich I n l e t c o u n t s a t Saa0.8 v e r s u s S t r a i t o f G e o r g i a c o u n t s , 1986. T r a n s f o r m e d w a t e r column t o t a l (Saa0.8) (G1545) May 1.37973 May 3.15982 May 1.37973 May 3.10584 Jun 0 J u n 4.25698 Jun 1.24573 J u n 3.99294 Jul 0 J u l 4.77987 Jul 1.24573 J u l 4.36907 Analysis Source STATION MONTH STN*MON Resid Total  of variance table Mean Sum o f sguares DF square 28.255 0.24211 2.6982 1.6725 32.868  1 . 2. 2. 6. 11 .  28.255 0.12105 1 .3491 0.27876  O v e r a l l mean 2.4096  COUNT Homogeneity Factors STATION  counts.  of variance  F-ratio  Prob.  101.36 0.4343 4.8396  0.00006 0.66659 0.05604  Overall standard deviation 1.7286  test  Bartlett Chi-square Probability 0.72875E-04 0.99319  DF 1  Size warn < 10  M u l t i p l e range t e s t s S c h e f f e t e s t a t 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 1. ) ( 2. ) B o n f e r r o n i t e s t a t 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 1. ) ( 2. ) M i n i m a l t e s t a t 5% p r o b a b i l i t y l e v e l T h e r e a r e 2 homogeneous s u b s e t s w h i c h a r e l i s t e d ( 1. ) ( 2. ) Homogeneity Factors MONTH  of variance  Bartlett Chi-square 1.7340  as  follows:  as  follows:  as  follows:  test  Probability 0.42022  DF 2  M u l t i p l e r a n g e tests-MONTH F - r a t i o i s not s i g n i f i c a n t a t p r o b a b i l i t y  Size warn < 10 0.66659  Homogeneity o f v a r i a n c e t e s t STATION,MONTH U n c a l c u l a b l e due t o s t a n d a r d d e v i a t i o n  of zero.  M u l t i p l e range tests-STATION,MONTH F - r a t i o i s not s i g n i f i c a n t at p r o b a b i l i t y 0.05604  T a b l e 27. ANOVA-Saanich I n l e t c o u n t s a t Saa9, r e p l i c a t e s t a k e n i n J u l y 1986. T r a n s f o r m e d w a t e r column t o t a l Depth 140 m 145 m 135 m 1.66302 0 1 .36392 1.31419 1 .60649 1.09161 1 .41136 1.20304 1.60948 1 .62325 1.23474 1 . 13336 1 .33935 1 .35454 Analysis SOURCE TOTAL GROUP ERROR  of variance Sum o f squares 101.489 0.94216 100.548  horizontal  counts, 150 m 1 . 17920 1 .04564 1 .24155  table DF  Mean square  16 3 13  6.34312 0.31405 7.73444  F-ratio 24.628  F - r a t i o = e r r MS/gr MS F  .05(2)12,3= 14.34  24.628>14.34 therefore, reject n u l l hypothesis, the mean v a l u e s a r e s i g n i f i c a n t l y d i f f e r e n t a t p=0.05.  no  Table 28: Copepod faecal pellet production over a 12 hour period. Copepod Collection Date Sept. 9 Oct. 14  Period of Captivity (dayis) 51 15  Total CVs Present  # of pellets (after 6 hours)  # of pellets (after 6 hours)  30 29  8 3  13 2  Total # of pellets (after 12 hours) 24 5  Table 29: Buoyancy of CVs following their death in experiments 8 and 9. Twenty Copepods Ten Copepods (from Experiment 9) (from Experiment 8) # of Copepods # of Copepods day bottom floating day bottom floating 7 13 0 4 2.5 10 10 10 5.5 6 5 4 6 8 12 5 5 7.5 15 5 8.5 6 11 4 4 6 10.5 6 12.5 4 6 18 4 6 24 4  Table 30: Calculated volume of water filtered through the m S C O R net (46% acceptance).  distance hauled (m) 15 25  volume distance filtered hauled (m) K)  volume filtered (m ) 3  1.73 2.88  175 180  30 37 40 50  3.46 4.27 4.61 5.77  190 195 200 205  55  6.34  208  23.99  65 70 72 75 80 90  7.50  210  24.22  8.07 8.30  215 225 240 245 250 270 285 290 295 335 340 350 365 375 380 385 390 395  24.79 25.95 27.68  8.65 9.23 10.37  95 100 110 120 125 135 140 145 150 155 160 165 170  10.96 11.53 12.69 13.84 14.42 15.57 16.15 16.72 17.30 17.87 18.45 19.03 19.61  20.18 20.76 21.91 22.49 23.06 23.64  28.25 28.83 31.34 32.87 33.44 34.02 38.63 39.21 40.36 42.09 43.25 43.82 44.40 44.98 45.55  Table 31: Concentrations of N. plumchrus ( C V ) and dissolved oxygen from central Saanich Inlet, 1969 (Hoos, 1970). Depth (m)  May 21 Oxygen (mL L ) - 1  125 150 175  0.10 0.06 0.00  Depth (m)  Jul 8 Copeods (#  125 150 175  m" ) 78.7 3  0  -  May 23 Copepods  May 28 Oxygen (mL L )  Jun 10 Oxygen (mL L " )  -  0.20 0.05 0.00  0.10 0.03 0.00  Jul 17 Oxygen (mL L " )  Sep 25 Oxygen (mL L )  Oct 8 Copepods  0.36  1.38 0.75 0.59  (# m - ) 159.2 0 3  1  0.08 0.00  - 1  - 1  1  (# m " ) 113.5 0 3  -  (# m " ) 0.12 27.0 3  -  Jun 13 Copepods  Nov24 Oxygen (mL L " ) 1  0.53 0.36 0.24  Table 32: Dissolved oxygen concentrations (mL L  - 1  ) measured in Saanich Inlet  in 1969 (see figure 55 for station location).  Depth  (m) 0  Jul 28 Oxygen  -  10 25 50 70 80 90  7.75 4.17  100 110 120  1.89  150 175 200 Depth  H  0 10 25 50 70 80 90 100  3.60  -  1.85 1.91 2.11 2.51 0.00 0.00 0.00 Jul 28 Oxygen  -  7.48 4.28 3.54 2.20  110 120 150  2.19 2.76 2.48 2.64 1.81 0.15  175 200  0.00 0.00  Depth  Jul 28 Oxygen  Station Saa3 Sep 16 Depth Oxygen (m) 0  -  Depth  Oct 28  (m)  Oxygen  -  10 25 50 70 80  4.88  0 10  3.45 2.91 1.17 0.74  25 50 70 80  3.46 3.63 2.48 2.39  90 100  0.76 1.93  1.23 0.63  110 120 150 175  1.23 0.17 0.00  90 100 110 120  0.05  150 175  200  0.85  200  Station Saa3.5 Depth Sep 16 (m) Oxygen 0 10 25 50 70 80 90 100 110 120 150 175 200  -  4.54 S.76 2.83 2.18 1.32 1.15 1.80 1.85 1.57 0.80 0.65 0.88  Depth  M  0 10 25 50 70 80 90 100 110 120 150  175 200  3.60  0.73 0.69 0.14 0.48 0.27 Oct 28 Oxygen  -  3.61 8.68 3.34 2.22 1.77 0.92 1.03 0.64 0.43 0.34 0.43 0.48  Station Saa4  (m)  -  Depth  H  Sep 16 Oxygen  -  2.45 2.74  0 10 25 SO 70 80  90 100 110 120 150  2.76 2.56 2.30 2.06 1.39  90 100 110 120 150  1.85 1.57 1.36 0.94 1.00  175 190  0.00 0.00  175 190  0.95 0.99  0 10 25 50 70 80  7.16 4.20 S.61  4.53 3.66 S.54 1.21 1.18  Depth  (m) 0 10 25 50 70 80 90 100 110 120 150 175 190  Oct 28 Oxygen  -  3.98 3.79 3.94 S.66 2.12 1.15 0.92 1.07 0.91 0.64 0.49 0.40  Table 33: Dissolved oxygen concentrations (mL L in 1974 (see figure 55 for station location).  Depth (m)  Jul 30 Oxygen  0 10 25 50 75 100 125 140 155 170 180 195  9.79 8.65 4.97 3.89 1.57 1.95 1.73 0.87 0.12 0.09 0.01 0.02  Station Saa4 Depth A u g 26 (m) Oxygen 0 10 20 30 50 75 100 150  4.05 3.51 3.47 3.46 1.05 1.56 0.91  1  ) measured in Saanich Inlet  Depth (m)  Sep 9 Oxygen  0 10 20 30 50 75 100 150 200  3.59 3.10 3.89 2.58 1.84 1.33 0.89 0.36  -  Table  34. G e l a t i n o u s  zooplankton c o l l e c t e d  GELATINOUS SPECIES 1. C l i o n e l i m a c i n a Pteropod 2. L e n s i a b a r y i Siphonophore 3. Dimophyes a r c t i c a Siphonophore 4. Nanomia c a r a Siphonophore 5. M u q q i a e a a t l a n t i c a Siphonophore 6. S a r s i a s p p . 7. L i m a c i n a h e l i c i n a . Pteropod 8. A q l a n t h a d i g i t a l e Hydromedusae 9. A e q u o r e a v i c t o r i a Hydromedusae 10. P r o b o s c i d a c t y l a f l a v i e i r r a t a Hydromedusae 11. Hybocodon p r o l i f e r Hydromedusae 12. A e q i n a c i t r e a Hydromedusae 13. P a n t a c h o g o n h a e c k e l i Hydromedusae 14. P l e u r o b r a c h i a s p . Ctenophore  Total  i n September  1985  Saa9 (0-100 m) Numbers Number m  96  8.3  16  1.4  54  4.7  4  0.3  12  1.0  4  0.3  115  Fitjure 1. The study area. Saanich I n l e t , the S t r a i t of Georgia, Sechelt I n l e t (for d e t a i l s see chart no. 3001).  and  116  Colanoid copepods  J  >  6 nouplior  5 copepodite  stoges  stoges  NI - NVI  CI - CV  v.  Adult  CVI  Figure 2. L i f e h i s t o r y stages c h a r a c t e r i s t i c of Neocalanus plumchrus and the calanoid copepods i n general (top). A l a t e r a l (Pottos l e f t ) and dorsal (bottom r i g h t ) view of the CV stage (the l a t e r a l view l a taken from Fulton, 1972).  F i g u r e 3. S e c h e l t I n l e t and S t r a i t of G e o r g i a a t a t l o n a sampled f o r z o o p l a n k t o n and h y d r o g r a p h i c p a r a m e t e r s .  118  Figure 4. Northern (Saa9). c e n t r a l ( S B B 3 J and aouthern (SaaO.8) a t a t l o n a l n Saanich I n l e t . The S a t e l l i t e Channel a t a t l o n (Sate) l a Bltuated a t the a l l l . .  Figure 5. The mSCOR net. A messenger i s used to release the b r i d l e s snabling the closure of the net canvas by the t h r o t t l i n g band.  120  a) 5  min.  b) 20  min.  c) 20  min.  F l o u r * 6. Bathykymograph r a a u l t a •) baaellna readout t o 150 a without C-B nets on tha wir« and the ship atationary, b) the f i r s t h o r i z o n t a l tow t o ISO • with three C-B nets on the wire, towed a t a speed of 1.5 knots and. c) a r e p l i c a t i o n of the f i r s t h o r i z o n t a l tow.  Figure 7. The basic apparatus f o r the low-oxygen tolerance experiments. The copepods were contained i n the t e a t funnels (Ts and Tb), and the c o n t r o l funnels (Ca and Cb) .  Log mean  F i g u r e 8. A r e g r e s s i o n of the l o g mean v e r s u s the l o g v a r i a n c e f o r 22 v e r t i c a l sample p a i r s .  haul  Figure  9.  Salinity  distribution  (ppt)  for  station  Saa9  in  19B5.  Month Jan  Figure  10.  Fab  Mar  Apr  May  Temperature d i s t r i b u t i o n  Jun  ( C) #  for  Jul  station  Aug  Saa9  in  Sap  1985.  Oct  Nov  Dec  Figure  11. D e n s i t y  distribution  (sigma-t)  for station  Saa9  i n 1985.  Figure 12. Dissolved oxygen distribution (mL L-l) for station Sea9 in 1985.  Figure  13.  Salinity  distribution  (ppt)  for  station  Saa9  in  1986.  Figure  14.  Temperature d i s t r i b u t i o n  («C)  for  station  Saa9 in  1986.  Figure  15. D e n s i t y  distribution  (sigma-t)  for station  Saa9  in  1986.  Month  Figure 16. Dissolved oxygen distribution (mL L-l) for station Sea9 in 19B6.  Oxygen (mL L-l)  1  1  1  1  1  1  1  1  1  1  1  1  1  1  1  1  1  1  r  x 800-  1  6  1 8  Copepod Concentration (# m-3) Figure 17. Copepod concentratione (aolid area) and dissolved oxygen levels (dashed line) at station S B B 3 . August 19B5. The 'x' designates the depths at which N. plumchrus was not found.  10  Oxygen (mL L-l)  i - - 1  ~T  r""I  1  1  T  1  1  1  • 1  1  1  1 - —  SO •  9  x  0  f 2  4  6  a  Copepod Concentration (# m-3) Figure 18. Copepod concentrations (eolid area) and dissolved oxygen levels (daehed line) at station Saa3. August 1986. The *x' designates the depth at which N. plumchrua was not found.  10  Month Jan  Figure  Fab  20.  Mar  Apr  Hay  Temperature d i s t r i b u t i o n  Jun  [°0  for  Jul  station  Aug  SaaO.B i n  Sap  1985.  Oct  Nov  Doc  Month  Month  Month  Feb  Mar  Apr  May  Jun  Jul  Aug  Sep  Oct  200  Figure 23. Salinity distribution (ppt) for station SaaO.8 in  1986.  Nov  Dec  Month  Figure 24. Tempereture distribution (°C) for station Seed.8 in 1986.  Figura 25. Density dletribution (eigma-t) for atetion SaaO.S in 1986.  Figure 26. Dissolved oxygen distribution (mL L-l) for stetion SaaO.8 in 1986.  Month  Figure 30. Dissolved oxygen distribution (mL L-i) for stetions 61545 and 61748 in 1985.  Month  Month  Fab  Mar  Apr  May  Jun  Jul  Aug  Sep  Figure 32. Tempereture distribution (°C) for station 61545 in 1986.  Oct  Nov  Dec  Month  Figure 34. Dissolved oxygen distribution (mL L-l) for station 61545 in 1966.  Temperature (°C)  Oxygen (mL L-l)  Figure 3 5 . Hydrographic parameters c o l l e c t e d at 1985: o x y g e n (x). d e n s i t y (O). t e m p e r a t u r e (o).  station Sci. and s a l i n i t y  November (A).  CD  Oxygen (mL L-l)  Temperature (°C) 4  0  0  -  6  -  8  10  •  «  A  ()  A  1I  A  C  A  100 -  100 -  r *> a  >  4J  a a  a  150 -  ISO -  800 -  SOO -  250 -  S50 IB  18  SO  SS  24  SS  Density (slgms t) Figure 36. Hydrographic parameters c o l l e c t e d at 1986: o x y g e n (x). d e n s i t y fo). t e m p e r a t u r e to),  <  A  (  A  <  )  <  >  station Sci. and s a l i n i t y  A  .-X..  •  24  SB  SB  February (A).  •  A  i  Ssllnlty  •  A  )  50  14  A A  ) SO -  IS  (ppt)  _1  SO  L .  38  Oxyg«n (mL L-l)  Temperature (°C) 10  0 F  7  9  11  13  15  17  19  Si  n  1  r  r  1  1  1  A  A A  50 h  50  100  100  e a a  c *> a a  150 h  150  a  a  200 h  200 r  250 h  250 h  300 r  3 0 0  13  15  17  19  21  23  A  A  Li 15  i i i i_ 17  Density (sigme t) Figure 37. Hydrographic parameters c o l l e c t e d at 1986; o x y g e n (x). d e n s i t y (O). t e m p e r a t u r e (O).  19  21  23  Salinity s t e t i o n Sc2. and s a l i n i t y  August (A).  25  27  29  (ppt)  o  Oxygen (mL L-i)  Density (sigma t) Figure 38. Hydrographic paremeters c o l l e c t e d et 1986: o x y g e n (X). d e n s i t y (O). t e m p e r e t u r e (O).  Temperature (°C)  Salinity s t a t i o n Sc2a. August end s a l i n i t y [A).  (ppt) tn ~*  cc  oo  C o p e p o d Density  i  :  i  :  i  :  i  :  r  L 0 40  0  0  <1 m~ CIV  3  OOm" •  3  (m ) - 3  C V &. CVI  Figure 39. Vertical distribution of Neocalanus plumchrus sampled from July 1985 through September 1986 (counterclockwise) at stations 61545 and 61748 (the bar perpendicular to the density line demarcates a separate vertical haul).  f  I  1  AUG  SEP  I  I  »  1  I  I  I  OCT NOV DEC JAN FEB MAR APR  I  I  MAY JUN  I  JUL  I  I  AUG SEP  Months (1985-1986) F i g u r e 4 0 . Copepod c o n c e n t r a t i o n s (CIV-CVI) c o l l e c t e d i n t h e S t r a i t o f G e o r g i a and 1 9 8 6 . The BOX c o n f i d e n c e i n t e r v a l s e r e shown f o r A p r i l t h r o u g h A u g u s t .  i n 1985  MAY  JUL  AUG  SEP  OCT  NOV  DEC  JAN  Months  FEB  MAR  APR  MAY  JUN  JUL  AUG  SEP  (1983-1966)  F i g u r e 4 1 . Copepod c o n c e n t r a t i o n s (CIV-CVI) c o l l e c t e d l n S a a n i c h I n l e t and S a t e l l i t e C h e n n e l i n 1985 e n d 1 9 8 6 . The 80% c o n f i d e n c e i n t e r v a l s e r e shown f o r r e p l i c e t e s a m p l e s c o l l e c t e d e t S a a O . 8 f r o m Mey t o S e p t e m b e r w i t h t h e e x c e p t i o n o f A u g u s t i n w h i c h t h e r e p l i c e t e s a m p l e s were c o l l e c t e d e t S a a 3 .  CD  03  Copepod Density  0 0  <1 m~ CIV  3  4  (m )  H CV  -3  & CVI  Figure 42. Vertical distribution of Neocalanus plumchrus sampled from May 1985 through September 1986 (counterclockwise) at station Sate in Satellite Channel.  156  CO  Copepod Density  4 (m )  0  0  0  <1  CIV  m  -3  -3  •  CV & CVI  Figure 43. Vertical diatribution of Neocalanua pluwchrua sampled from May 19BS through September 19B6 (counterclockwise) at atationa 8aa3 in Saanich Inlet (the bar perpendicular to the denaity line demarcates a separate vertical haul).  157  CO 00  Copepod  Density  0 4 0  £2  <1 m CIV  - 3  (m ) - 3  H CV  &  CVI  Figure 44. Vertical distribution of Neocalanus plumchrus sampled from September 1985 through September 1986 (counterclockwise) at station 8aa0.8 in Saanich Inlet.  cc  Copepod Density  0 4 0  0  <1 CIV  m  -  (m~ )  3  3  •  CV Sc CVI  Figure 45. Vertical distribution of Neocalanus plumchrus sampled from September 19B5 through September 19B6 (counterclockwiae) st station 8aa9 in Saanich Inlet.  Oxygen (mL L-l)  i  i  i  i  —  i  6  7  8  r  1  1  9 '" "  10  I"  1  9 50 •  100  150  X  " " " "  t  1 <D  1  $ 200  0  1  2  3  4  Copepod Concentrations (# m-3) Figure 46. Copepod concentratlona (aolid area) and dissolved oxygen levela (dashed line) at atatlon SaeO.8. June 1986. The ' x ' designates the depths at which N. plumchrus waa not found.  Oxygen (mL L-l)  0•  40 •  BO •  ISO  160  Copepod Concentrations (# m-3) Figure 47. Copepod concentretions (solid area) end dissolved oxygen levels (dashed line) et station Saa9. July 1966. The 'x' designetes ths depth et which N. plumchrus wss not found.  161  CO CO  Copepod  0  0  <1  CN  m  Density  - 3  •  0 4 (m ) -3  C V & CVI  Figure 48. Vertical distribution of Neocalanus plumchrus sampled from November 1985 and February 1986 (counterclockwise) at station Sci in Sechelt Inlet. In August 19B6 the copepods were collected at station 8c la.  Figure 49. Vertical distribution of Neocalanus plumchrus sampled in August 1986 at otatlona Sc2 and 9c2a in Sechelt Inlet.  •ays in Captivity  Figure  50.  Mortality  of  starved versus  fed copepods  c o l l e c t e d October  14.  1986.  2  6  4 Oxygen  (mL  L-l)  F i g u r e 5 1 . P e r c e n t m o r t a l i t y o f CVs e x p o s e d t o d i f f e r e n t o x y g e n c o n c e n t r a t i o n s . N o t e t h e v a r i a t i o n s i n t h e t i m e o f y e e r t h e c o p e p o d s were c o l l e c t e d (numbers i n d i c a t e t h e held in c a p t i v i t y p r i o r to experimentation) .  days  ^ S e ./'Oc  Se  . A Jl (Seen)  20  40  60  Number o f  Deys  80  In  •  Oxygen  +  .59-.62-.62  (mL  L-l)  .47-.48 .56-.34  X  .71-.69  O  .83-.87  V  .94-.95-.92  •  2.11-2.08-2.02  100  120  Captivity  F i g u r e 52. C a p t i v i t y p e r i o d and p e r c e n t m o r t a l i t y o f CVs s u b j e c t e d t o low o x y g e n t o l e r a n c e t e s t s . Mean o x y g e n c o n c e n t r e t i o n f o r e a c h e x p e r i m e n t i e a r r a n g e d a c c o r d i n g i t a occurrence in the legend.  to  1 6 6  o  Number of Days ln Captivity Figure 53. Expected reaults of the low oxygen tolerance experiments.  167  F i g u r e 54. T h e h y p o t h e s i z e d d i s t r i b u t i o n o f N e o c a l a n u s p l u m c h r u s p r i o r t o , a n d d u r i n g t h e d e e p M e t e r r e n e w a l i n 19B5; b a s e d o n b i o l o g i c a l d a t a and t h e o x y g e n r e g i m e o n A u g u s t 7 t h (a) a n d S e p t e m b e r 1 7 t h (c) .  Figure 55. Hydrographic s t a t i o n s and t h e z o o p l a n k t o n h a u l s t a t i o n 1974.  (Saa4. S a a 3 . 5 . a n d (Stn.E) sampled l n  Saa3) 1969 a n d  169  REFERENCES A l l d r e d g e , A . L . , B.H. R o b i s o n , A. F l e m i n g e r , J . J . T o r r e s , J.M. K i n g a n d W.M. Hammer. 1984. D i r e c t s a m p l i n g a n d i n s i t u o b s e r v a t i o n of a p e r s i s t e n t copepod a g g r e g a t i o n i n t h e m e s o p e l a g i c zone o f t h e S a n t a B a r b a r a B a s i n . Mar. B i o l . 80: 75-81. A n d e r s o n , J . J . , a n d A.H. D e v o l . 1973. 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V,= V = V =  volume volume volume  C,= C = C =  c o n c e n t r a t i o n of 0 i n t h e i n t r u d i n g water 0 c o n c e n t r a t i o n below 150 m, A u g u s t 7 0 c o n c e n t r a t i o n of mixed water below 150 m,  V = V =  unknown o f i n t e r e s t 13.0 x 1 0 m (estimate ?  2 3  2  3  2  3  o f i n t r u d i n g water o f w a t e r below 150 m o f m i x e d water 2  2  2  8  from Anderson  3  September  & Devol,  17  1973)  C,= 2.0 mL L " ( e s t i m a t e b a s e d on t h e d e c r e a s e i n 0 content renewed w a t e r i n t h e S t r a i t o f G e o r g i a ) C = 0.02 mL L " ( a p p r o x i m a t e l y 22% o f t h e water below 150 m c o n t a i n e d 0.10 mL L " ) C = 0.38 mL L " (the average 0 c o n c e n t r a t i o n below 150 m)  of  1  2  1  2  1  1  3  2  V, + V = C,V, + C + C C,V, + C c,v, - C 2  V V = C V V = C (V, + V ) V = C V, + C V Vi= C V - C V 3  2 2 2  2  3  2  3  2  3  2  3  2  v ^ c , - c )= v ( c - c ) Yi = (c - c ) v (c,- c )  2  3  3  3  3  2  V,  3  2  2  2  3  2  2  3  = 0.38 mL L ' - 0.02 mL L x 13 x 1 0 2.0 mL L " - 0.38 mL L " 1  1  1  V,  = 2.89  x 10  m  8  (volume o f i n t r u d i n g  3  m  8  3  1  water  E s t i m a t e o f volume o f i n t r u d i n g w a t e r (V^ ) ( + 0.6 x 1 0 ) - ( A n d e r s o n & D e v o l , 1973)  in  1985)  i n 1969,  4.3  x 10  8  m  3  8  C a l c u l a t i o n o f an e q u i v a l e n t balance the i n f l o w . length= width=  12 km 2.5  thickness=  km  volume  of o u t f l o w i n g  (assuming the m a j o r i t y & Saa9) (mean v a l u e )  water  to  of copepods a r e found a t  10m 10 m x 2500 m x 12,000 m = 3 x  10  8  m  3  

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