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Racial analysis of Skeena River steelhead trout (Salmo gairdneri) by scale pattern features Cox-Rogers, Steven Frank 1985

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RACIAL ANALYSIS  OF SKEENA  GAIRDNERI)  RIVER STEELHEAD  BY SCALE PATTERN  TROUT  (SALMO  FEATURES  by STEVEN FRANK COX-ROGERS B.Sc.,University  Of B r i t i s h C o l u m b i a , 1 9 8 1  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Department o f Z o o l o g y ) We  accept t h i s thesis required  as conforming standard  to the  THE UNIVERSITY OF BRITISH COLUMBIA September 1985 ©  Steven Frank  Cox-Rogers,  1985  In  presenting  degree  this thesis  in partial fulfilment  requirements  for  an  of this thesis for scholarly  department  or  by  his  or  her  I further agree that permission for  purposes  representatives.  permission.  Department of The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3  DE-6(3/81)  OCT" / o/fitf  extensive  may be granted by the head of my It  is  understood  that  publication of this thesis for financial gain shall not be allowed without  Date  advanced  at the University of British Columbia, I agree that the Library shall make it  freely available for reference and study. copying  of the  copying  or  my written  i i  ABSTRACT The scale  feasibility  patterns  to  of  using  identify  freshwater component  (Salmo g a i r d n e r i ) i n t h e  Skeena R i v e r  samples and  data  sex  originating Kispiox, different the  1984  salmon  and  from  five  years. incidental  fishery  composition,  steelhead  indicated  scale  steelhead  age  stocks.  in  scale  origin  with  accuracy  (range  Zymoetz 29%-60%, K i s p i o x  Babine  classification three  of  the  the  presence  stocks of  Skeena R i v e r  steelhead  probable  stock  the  stock  a b u n d a n c e and  general, predicted  be  62%  exist  1984 of  results  as  purposes. growth,  the  five  function  age  Skeena  analysis to c o r r e c t  average  classification  35%-60%,  Morice-Bulkley  d e p e n d i n g upon  the  stock  origin  stock support  differences the  notion  commercial  and  abundant  through  Sustut with  the  River  in that  stocks.  fishery  indicated that d i s t i n c t occur  for  Zymoetz) i n d i c a t e d  quantifiably discrete  mixed  run-timing  most  from  commercial  classified  56%-72%)  between  These  Morice-Bulkley to  and  be  of  J u v e n i l e morphometric a n a l y s i s  significant form.  steelhead  series  4  pattern  (Kispiox, Morice-Bulkley,  body  Classifying  45%  could  Sustut  model u s e d .  standardized  to  stocks  54%-64%,  Scale  (Zymoetz,  a  Area  discriminant  of  44%-76%,  over  were f o u n d between  the  between  trout  tributaries  the  that  river  steelhead  classification  in  Linear  five  year  investigated.  Sustut)  catch  differences s i z e s at  of  marine  samples were a l s o c o l l e c t e d  for p o t e n t i a l stock  and  was  River  Babine,  Adult  stocks  first  were a t t a i n e d f r o m a d u l t  Skeena  Morice-Bulkley,  Significant  River  size  and  catches peaks  of  fishery.  In  steelhead  were  run-timings  during  the  earlier River  p o r t i o n s of  steelhead  run-timings impacts  to  the  were p r e d i c t e d  through the steelhead  River  is  a  steelhead.  variance  i n the  differences.  briefly suggest  feasible Further technique  method study and  Kispiox,  t o be  fishery.  are  These o b s e r v a t i o n s patterns  fishery.  to  less  B a b i n e , and  abundant  Zymoetz  with  P o t e n t i a l commercial  later fishery  discussed. that  the  f o r stock i s required  technique  of  separation  i n Skeena  to  clarify  better e s t a b l i s h stock  scale  yearly specific  i v  TABLE OF CONTENTS  ABSTRACT  i i  L I S T OF TABLES  vi  L I S T OF FIGURES  ..  viii  ACKNOWLEDGEMENTS  xi  Introduction  1  Description Life The Materials  o f t h e Skeena R i v e r  H i s t o r y o f Skeena R i v e r Skeena R i v e r  Commercial  Drainage  5  Steelhead  8  Salmon F i s h e r y  a n d Methods  Scale  17  Data C o l l e c t i o n  Determination  and P r e p a r a t i o n  17  o f Sample S i z e s  23  Juvenile Analysis Analytical  10  24  Techniques" f o r S c a l e  Pattern  Analysis  Results  .... 25 30  Discrimination  o f Skeena R i v e r  Verification Descriptive Age  Steelhead  30  of s c a l e aging  30  statistics  30  composition  30  S i z e s a t age Scale  Pattern  35  Features  o f Skeena R i v e r  steelhead  .... 40  Scale  features  o f s m o l t age 3 a d u l t  steelhead  .... 41  Scale  features  o f s m o l t age 4 a d u l t  steelhead  .... 48  Scale  features  o f age 3.2+ a n d 4.2+ s t e e l h e a d  .... 52  Scale  pattern  variation  between y e a r s  55  Plus  growth  57  Stock D i s c r i m i n a t i o n Separate Pooled  57  freshwater  freshwater  Discrimination Commercial F i s h e r y  age d i s c r i m i n a n t  age d i s c r i m i n a n t  models  by sex Stock Composition  72  76 84  Discussion  97 Considerations  Theoretical  Applications  97  Considerations  Commercial F i s h e r y  LITERATURE CITED  58  75  Juvenile Analysis  Biological  models  105  Considerations  to Steelhead  Management  108 114 11 8  L I S T OF TABLES  Table  1.Riverine  tributaries Table for Table by Table  used  features  five  Skeena  River 9  from  the a d u l t  steelhead  scales  stock  . 22  3. Age C o m p o s i t i o n  f e a t u r e s o f Skeena R i v e r  steelhead  s e x , s m o l t a g e , a n d ocean age 4. Mean l e n g t h s  5.  One  differences steelhead  way in  34  and w e i g h t s of s t e e l h e a d  o c e a n age f o r t h e f i v e Table  the  i n the study  2. V a r i a b l e s measured each  of  stocks  ANOVA  mean  used  results  scale  zone  from t h e K i s p i o x ,  at  various  i n the study  40  f o r comparison widths  Zymoetz, a n d  at  of  age f o r  Morice-Bulkley  Rivers Table  6.  44 One  differences steelhead Table  7.  differences  Table  8.  ANOVA  way  differences  comparison  features  f o r smolt  of age 3 47  results  i n mean s c a l e zone  way  for  o c e a n age  ANOVA  by p o o l e d  One  results  i n mean s c a l e zone  by p o o l e d  One  steelhead  way  for  comparison  features  of  f o r s m o l t age 4  o c e a n age  ANOVA  51  results  i n mean s c a l e zone  for  features  comparison  of  f o r age 3.2+ a n d  4.2+ s t e e l h e a d Table  9. One way ANOVA  55 results  i n mean s c a l e zone f e a t u r e s steelhead  f o r comparison f o r Sustut  between  years  and Zymoetz  River 56  Tables  10-16.  Classification  discriminant steelhead Table  models  used  matrices to  for  classify  the  Skeena  linear River  to stock of o r i g i n  17. 1984 c o m m e r c i a l  62  fishery  steelhead catches  i n Area  4  79  Table  18.  sampled Table  for Table  from  age  Area  composition  Area  s t e e l h e a d sampled Means  comparison juvenile  sex  f o r steelhead 79  weights  of  steelhead  sampled  4 i n 1984  20. C l a s s i f i c a t i o n  21.  by  4 i n 1984  19. Mean l e n g t h s a n d  from Table  Ocean  and  80 results from the  of d i f f e r e n c e s  steelhead  to stock  Area  of o r i g i n  by week  4 i n 1984  results  81  o f one way ANOVAS f o r  i n morphological  features  in 95  vi i i  L I S T OF FIGURES  Figure  1. The Skeena R i v e r  Figure  2.  month Figure  1963  to  i n Area  Figure  4.  to  6. D i s c r i m i n a n t  Figure  7. Age c o m p o s i t i o n  the Figure the Figure  1984  i n Area  Figure  Figure  steelhead  escapement  steelhead  mean  annual  steelhead  4  scale  15 from t h e S u s t u t  River  ..... 19 28  structure  f o r the f i v e  steelhead  i n the study  stocks  used  31  o f age 3.2+  and 4.2+  stocks  used  10. Mean l e n g t h s stocks  used  11. Mean s c a l e  steelhead  from  i n the study  36  9. Mean w e i g h t s o f age 3.2+ a n d 4.2+  five  month  space  8. Mean l e n g t h s  five  by  13  5. A d u l t  five  by  4  Figure  the  harvest  13  1963  used  steelhead  4  harvest+escapement  stocks  6  1984 mean a n n u a l  3. 1963 t o 1984 mean  through Area  Figure  Drainage  steelhead  for  i n the study  36  by sex f o r age 3.2+  steelhead  from  i n the study  zone w i d t h s  38  for  steelhead  of  smolt  ages 3 and 4 Figure  12.  steelhead Figure  Figure  Yearly of smolt  13. Y e a r l y  steelead 14.  42 freshwater  scale  age 3 by p o o l e d  freshwater  scale  freshwater  i n adult  o c e a n age  zone c i r c u l i  o f s m o l t age 3 by p o o l e d Yearly  zone w i d t h s  45 counts  in  o c e a n age  scale  zone w i d t h s  45 in adult  steelhead Figure  15.  adult Figure  Yearly  16.  Yearly  18.  19.  20.  pattern Figure  21.  pattern Figure  22.  pattern  freshwater of  s c a l e zone c i r c u l i  s m o l t age  freshwater  counts  in  4  49  s c a l e zone w i d t h s  in  steelhead  freshwater  s c a l e zone w i d t h s  in  steelhead  4.2+  pattern Figure  49  53  Yearly  pattern Figure  4  3.2 +  17.  of age Figure  s m o l t age  steelhead  of age Figure  of  53  Discriminant variation  in adult  Discriminant variation  in adult  of  smolt  age  3  of  s m o l t age  ....  59  scale 4  ....  64  function analysis describing scale of  age  3.2+  67  function analysis describing  in steelhead  Discriminant variation  steelhead  in steelhead  Discriminant variation  steelhead  function analysis describing  Discriminant variation  function analysis describing scale  of  age  scale  4.2+  70  function analysis describing scale  in adult  steelhead  of p o o l e d  smolt  age 73  Figure  23.  sampled  Age  composition  i n the  1984  s t r u c t u r e by  commercial  week f o r  steelhead  fishery  77  Figure  24.  Predicted  run-timing  85  Figure  25.  Predicted  run-timing  85  Figure  26.  Predicted  run-timing  87  Figure  27.  Predicted  run-timing  Figure  28.  Predicted  run-timing  89  Figure  29.  Predicted  run-timing  89  Figure  30.  Predicted  run-timing  91  . ..  87  X  Figure Figure  31.  Predicted  32.  run-timing  Discriminant  •  function  analysis  morphological v a r i a t i o n  among  the  Morice-Bulkley  Kispiox,Zymoetz,and  91  juvenile  describing  steelhead Rivers  from 94  xi  ACKNOWLEDGEMENTS I  would  committee Walters this  like  t o t h a n k my  members  for their  thesis.  I  Drs. advice would  supervisor  Art  Tautz,  like  Parkinson,  B r u c e Ward,  and  Angelo  Fisheries  Branch  Les  Janz  Fisheries  and Oceans f o r t h e i r  discussions. Fisheries  and Oceans p r o v i d e d i s extended  Rupert D i v i s i o n  in  to  of  the  thank Mike of  Lough,  the  of Eric  Provincial  Department  of  comments t h r o u g h numerous and  financial the  development  the F e d e r a l  Branch  to  W i l i m o v s k y and  M c P h a i l , and C a r l  Fachin  helpful  The B.C F i s h e r i e s  A note of thanks Prince  and  J.D.  and a s s i s t a n c e also  Dr N . J .  the  Department  support f o r t h i s  staff  f o r t h e use of t h e i r  of  B.C.  facility  of  study. Packers  during  this  project. Finally,  I  would  like  to express sincere  w i f e M a r i e who was a l w a y s w i l l i n g and  s u p p o r t when needed  most.  to  provide  gratitude  field  t o my  assistance  1  INTRODUCTION Fisheries determining fisheries Anas  and M u r a i  populations harvesting  Conrad  of  o f one may  "stock  or  1981) i n m i x e d  populations stocks  of  a  possessing  and E v a n s , Because  genetic  productive  in  management  requires  their  a  mixed  exclusively  (Hill  salmon  their  salar)  stocks  studied  fisheries.  1962,  1978, P e l l a and 1981,  spawning  intermix,  the other  the  (McDonald,  1972, R i c k e r  The  various  spawning  (McDonald,  the  Parsons  as  received  are contributing  in  North  salmonids, 1971,  Misra  importance, well  as  America  have  although  not  and  Ni  1983).  a l l of the P a c i f i c the  considerable  species harvested  The i n c i d e n t a l  effective  time.  analyses  on  1972).  susceptible to  1981),  of which stocks  local  (Maclean  (Larkin,  are p a r t i c u l a r l y  over  1972,  to represent  be m a i n t a i n e d  change  have been n o n - t a r g e t  stock  affect  a r e taken  fishery  commercial  have  Fredin  d i f f e r e n c e s that are adaptive  (genus O n c o r h y n c h u s )  (Salmo  mixed  of  1959,  population  different  and L a r k i n  species  fisheries  concentrated  and  species  management.  knowledge  primarily  Because  (Simon  distributions  stock  mixed  in  has l e d t o e x t e n s i v e a p p l i c a t i o n s  1981) and w h i c h s h o u l d  less  Mixed  this  given  overfishing  a n d how  Whenever  several  fishery  in  1980, M a c l e a n and E v a n s  differentially  concept"  interested  1971, Cook and L o r d  1984).  single  been  1956, W o r l u n d  and Sandeman  Not s u r p r i s i n g l y ,  the  Utter  1969, B i l t o n  1981,  long  structure  and W h i t e s e l  1979, L e a r  McDonald  1981).  have  subpopulation (Clutter  Robertson  of  biologists  Atlantic  salmon  attention. incidentally  Less in  i n t e r c e p t i o n of steelhead  2  trout  (Sa-lmo  Pacific  gairdner i)  coast  of N o r t h A m e r i c a  Steelhead California  Andrews  are  the  along  Alaska  Oguss and  fisheries  and  Skeena  which  are  incidental of  steelhead  catch  steelhead  commercial each  pink  (unpublished  stock  fishing  may  range  (Oguss  1984a) w i t h to  given  to the  latter,  steelhead  commercial  BCFB  the  the  total  i s harvested  1984). the  sockeye  qorbuscha)  30%-60% of year  major Fraser  summer r u n the  and  the  (Oncorhynchus  through  affecting  Columbia,  regard  estimated  i n any  dynamics be  during  data,  In B r i t i s h  adjacent  With  salmon An  northern  Parkinson  " s t o c k s " of  harvested  the  from  their  estuaries.  return  coast  1966).  in areas  various  f o r salmon a l o n g example.  Pacific  1978,  (June-September).  Skeena R i v e r  such  throughout  River  incidentally  year  the  Evans  ( O n c o r h y n c h u s n e r k a ) and each  i s one  occurring  Skeena R i v e r h o s t s  fisheries  (Withler,  harvested  1977,  incidental River  occur  into  steelhead  i n net  Little  fishery  biological  as  i s known  nor  of  how  integrity  of  stock. Preliminary  (unpublished River  the  stock  has  each  stock  catch. specific  1982,  stocks  1984)  by  the  suggest  This,  B.C  that  ' commercial  fishery.  The  run-timing.  Without  s t e e l h e a d has  believed  to  the  such  Of  overall  c o i n c i d e with  temporal  weekly pattern  peak  Skeena  abundance  concern  knowledge t h e  been l i m i t e d ,  major  identification  c o n t r i b u t e s p r o p o r t i o n a l l y to the in turn, determines  F i s h e r i e s Branch the  show d i s t i n c t "peaks of  been, however, q u i t e d i f f i c u l t .  Skeena R i v e r stocks  data,  steelhead  through  investigations  of  each  is  how  incidental of  stock  management  of  especially  for  those  s o c k e y e and  pink  salmon  3  run-timing. stock  This  origins  s u g g e s t s t h e need  o f Skeena  River  f o r ways o f i d e n t i f y i n g  steelhead  i n the commercial  the  salmon  fishery. Several origins  of  commercial widely case  techniques are a v a i l a b l e salmonids  fisheries.  applied of  wild  problems  for  Mark  recapture  in various  studies  (Hartt  steelhead,  they  juvenile  to d i f f e r  between  populations  Electrophoretic 1977,  Chilcote  Grant  1984,), body  et  et  a l .  1980;  chum  and m e r i s t i c s  1980;  Landrum  1964 c h i n o o k s a l m o n : McGregor  composition 1983), (sockeye degrees  age  Leggett  1981; coho  structure  (Ricker  salmon: M a r g o l i s of  populations.  success  1971,  characterize  structures  such  as  Atlantic  1981;  sockeye et a l .  Smith  1969,  1962, Dark a n d s a l m o n : Amos e t  1984), Mulligan  salmon: elemental et a l .  infestations  used with different  P e r h a p s t h e most w i d e l y a p p l i e d  the use o f c a l c a r e o u s  1984b;  1972) a n d p a r a s i t i c  1958) have a l l been to  Allendorf  1984; A t l a n t i c  salmon: T a y l o r  (sockeye salmon: C a l i p r i c e  1962).  ( steelhead:  et a l .  o r known  Fournier  1924; p i n k  i n the  variation  and  sockeye salmon: Fukuhara  1963; chum s a l m o n : F o u r n i e r and  salmon:  been  recapture.  occurring  1972, T h o r p e a n d M i t c h e l l  a l .  morphology  adult  1984a,  stock  logistic  are hypothesized  a l . 1980, P a r k i n s o n  et  Riddell  and l a t e r  (steelhead: Utter  Winter  al.  substantial  (Worlund a n d F r e d i n ,  variation  s a l m o n : Nyman and P i p p y salmon:  systems t h a t  have  1962); however,  present  tagging  methods  t e c h n i q u e i s t o use n a t u r a l l y  one o r more b i o l o g i c a l  the r a c i a l  e n v i r o n m e n t s a n d i n mixed  and  both  An a l t e r n a t i v e in  in natal  for identifying  varying spawning  t e c h n i q u e h a s been  otoliths  (steelhead:  4  Mckern  et  a l .  especially and  scales  Sandeman  Clutter  1974),  (Atlantic  1980,  Anas  and  Bilton  and M e s s i n g e r  and  1956, Henry  Murai  (see Ihssen  salmon: Lear  Reddin  and W h i t e s e l  1963,  1978,  f i n rays  and  Misra 1961  et a l . Misra  1985;  ,  1975, Cook and L o r d  M c B r i d e and M a r s h a l l  1978,  Pacific  Rowland  1969, Tanaka e t a l .  1981) and Lear  salmon:  1969,  Mosher  1969, B i l t o n  1978, K r a s n o w s k i  1983, McGregor e t a l .  1971, et a l .  1983,  Conrad  1984). Scale  analysis  identification collect are  and p r e p a r e ,  and  Scales  large  1981).  have  scale  proven  Larkin  1955, Chapman Withler  Horncastle Keating  1981, 1959)  particular stock  1974,  among  and  variations  differences  between  hypothesized  to result  freshwater between used  period.  stocks  to  used  by  populations  Maher  1954,  many  studied  Maher  1966, N a r v e r  and 1969,  1958,  however,  patterns pattern  to  (Bali  characterize  analyses  zones.  rearing  in differential  them,  read  studies  Few  growth  degree  to  1978,  freshwater  determines the  separate  identification  i n t h e w i d t h s and p a t t e r n s  scale  The  easier  1977, W h a t e l y e t a l .  scale Scale  stock  o f t h e s p e c i m e n , and  1958, W i t h l e r  Whately  stocks.  yearly  1954,  other  generally  i n those  others).  have  steelhead  specific  circuli  1958, B a l i  over  s c a l e s have been  reliable  1944, S h a p o v a l o v and T a f t  and  are  stock  Steelhead  (Neave  Narver  advantages  do n o t r e q u i r e k i l l i n g  to  et a l .  authors  certain  techniques.  applicable  (Ihssen  has  of  accuracy  of  on  of s c a l e  Environmental environments  s c a l e growth scale  rely  during  are the  pattern difference statistical  o f t e n by d i s c r i m i n a n t a n a l y s i s .  models Both  5  parametric  (Anas and  Messinger  1975,  1978,  1982)  Cook  Conrad  salmonid  analysis  by  steelhead  long  time  specific  method  racially used  as  they  are  problems.  rear  Bilton  and  (Cook and  Lord  applied The  to  is particularly  to  a  p o t e n t i a l of suited  in n a t a l environments  use  stocks. the  The  pattern  steelhead  assess  of  longterm  scale pattern  differentiating  scale  separable  steelhead  have been  subject  Skeena R i v e r .  that  to  1975,  nonparametric  scale patterns  and  for  Firstly,  hypothesis  et a l .  for  watershed  regimes.  from the  fold.  and  fishery  t h e s i s examines the  practical stocks  of  growth  This  Major  1984)  t o Skeena R i v e r periods  1969,  discriminant analyses  wide r a n g e of mixed discriminant  Murai  goals  from  the  Secondly,  by  of  analysis  potential  contributions  between  steelhead  the  study  was  used  Skeena  to  two-  test  the  exist  as  analysis  was  identifying  to  a  trout  were  River  scale pattern  for  stock  a n a l y s i s as  the  weekly  commercial  salmon  approximately  30,500  the  f ishery.  Description The  of  the  Skeena  River  square  kilometers  British  Columbia  west  near  in  (figure  1).  with the  Seven  interior  tributaries,  can  Skeena be  an  the  considered  of  central  western  precipitation plateau  nearer  River  area  Climatic patterns  light  moderate temperatures  1968).  Drainage  drains  lying  direction  temperature and  Skeena R i v e r  the  and coast  tributaries, as  hosting  portion  vary and  heavy  i n an  the  east-  extremes  of  precipitation  ( L a r k i n and as  of  w e l l as major  McDonald, their  sub-  steelhead  6  Figure  1. The Skeena R i v e r D r a i n a g e . Shown a r e t h e major s t e e l h e a d t r i b u t a r i e s : the L a k e l s e , Kitsumkalum, Zymoetz, M o r i c e - B u l k l e y , K i s p i o x , B a b i n e , and S u s t u t R i v e r s ( a f t e r Whately, 1977).  7  8  stocks;  i n ascending  Lakelse,  order  Kitsumkalum,  Kispiox,  Zymoetz  Babine, and S u s t u t  various  other Khtada,  Squingula  e t c ) , smaller  known t o s u p p o r t  tributaries, large  lake  features study  rivers  Exstew,  from  t h e mouth t h e s e  (Copper),  tributaries  Gitnadoix,  are  upstream  Morice-Bulkley-Suskwa,  respectively.  (Ecstall, Kitwanga,  creeks,  steelhead  for  the  (Zymoetz,  Table five  Two  Sicintine,  of  the  Kispiox,  the  itself larger  R i v e r s , headwater i n  tributaries  Morice-Bulkley,  Eschamsiks,  and t h e m a i n s t e m Skeena  summarizes  Skeena  addition,  Kitseguecla,  production.  1  In  Khyex,  t h e B a b i n e and M o r i c e - B u l k l e y  systems.  a r e the  major  riverine  considered  Babine,  and  in this Sustut  Rivers).  Life  H i s t o r y o f Skeena R i v e r Skeena R i v e r  fishery to  Skeena  River  fourth, f i f t h ,  as a d u l t s  sixth,  life.  After overwintering  spawn  from  mid to  to late five  ocean. as  mid  April  summer w i t h  years  incidentally  (winters)  i n the commercial  run o r i g i n  which  return  from June t h r o u g h September i n  seventh,  or  eighth  plus  years  of  i n n a t a l streams t h e a d u l t s g e n e r a l l y  through June. the parr before  F r y emergence o c c u r s  remaining  smolting  i n freshwater  and  migrating  from  f o r one to  the  Not a l l a d u l t s d i e f o l l o w i n g spawning and many a r e t a k e n  kelts  migration. found  taken  a r e p r i m a r i l y o f summer and f a l l  the  their  steelhead  Steelhead  in  the  commercial  fishery  during  W i n t e r and s p r i n g r u n s t e e l h e a d  i n t h e l o w e r Skeena R i v e r  tributaries  their  seaward  (November-April) are below  Hazelton  and  Table  1 . R i v e r i n e f e a t u r e s of t h e f i v e used i n t h e s t u d y .  Skeena R i v e r  tributaries  Morice/ Sustut Ki spiox Babine Zymoetz Bulkley 128 27 W 126 43 W 1 27 40 W 1 26 42 W 127 20 W 56 00 N 55 1 5 N 55 25 N 54 24 N 54 32 N .3000 2086 6790 12300(M+B) ( s q km) 3080 1911(M) 200(B) 270 350 220 U p s t r e a m (km) 115 315(M) D i s t a n c e app. 164(M+B) 51 46 Mean Flow (m3/s) 1 38 76(M) June June June June Peak Flow June Jan. Mar. Mar . Feb. Minimum Flow Jan. 120(B) 85 65 1 37 L e n g t h (km) 80 75(M) Summer 24-52 24-33 1 5-25 1 5-24 6-15 T u r b i d i t y (JTU) 6-10 10-16 9- 1 4 11-16 6-1 1 Summer Water (C) Temperature 22-64 Water H a r d n e s s (mg/1 CaCo3) 50 t o 40 t o 40 t o 40 t o 1 00 t o Mean A n n u a l (cm) 75 75 1 00 250 350 Prec i p i t a t ion -22 -16 -15 -18 Mean J a n u a r y (C) 1 a i r temp. <1 4 <1 4 1 6 1 6 1 5 Mean J u l y ( c ) a i r temp. <50 <70 60 t o <60 t o F r o s t f r e e ( d a y s ) 60 t o 1 00 1 00 1 40 period Feature Long. Lat. Drainage  _  1  10  are  not  subject  to  any  appreciable  incidental  (commercial)  fishery. The  Morice-Bulkley  believed  to  production river The  river  support  followed  the  by  and  majority  the  Babine,  of  its  tributaries  Skeena  river  Zymoetz, S u s t u t ,  Lakelse  and  Kitsumkalum  recognized.  parr  Skeena R i v e r  i t may  have an  originating (Tredger,  1984).  previously  examined  for  River  (Whately, Taylor River  (Whately  1977),  Pinsent  and  system with  regards  to  The diverse  Commercial  commercial history  catches  1981).  the  (1968) and  Skeena R i v e r  salmon  and  of  ( L a r k i n and  Fisheries  two  statistical  1978),  Babine  River  Chudyk  production  in  the  is  i s not  rearing  less  the  productive  steelhead  the  run  have  been  i n the  Morice-  Kispiox  River  (Narver,  1969).  (1973) d e s c r i b e d  the  Both Skeena  steelhead.  1955)  fishing  on  the  characterized  Todd and effort  Area  4 adjacent increasing  estuary.  An  participate  in  fishery  Skeena R i v e r  target species,  1968,  Skeena R i v e r  the  run  features  al.  principle  of  history  fishery  McDonald  majority  1984)  Salmon F i s h e r y  (see M i l n e ,  the  The  salmon  of  Skeena R i v e r  et  data,  p r i m a r i l y winter  role  several  life  Kispiox  steelhead production  important  from  tributaries  Bulkley  are  c o n t r i b u t i o n t o summer  Mainstem  known; however, larger  their  rivers  is  steelhead  and  s y s t e m s r e s p e c t i v e l y (BCFW B r a n c h , u n p u b l i s h e d  streams although  The  system  by  1971, by  proportion they  are  a  fluctuating  gillnet  of  pink  McDonald  ( w i t h i n 25-30 km)  although  had  s o c k e y e and  Larkin  occurs  has  to  in the  seiners primarily  1  restricted species chum  taken  of  cutthroat Oguss  in  and  Evans  although  and  lengths  river  through Area  4 on  to  The  do  so.  (harvest  occur  on  days  (24  peak  Normal  fishery  the  height  of  Department f o r the  steelhead peaking just  prior  average s t e e l h e a d  been  just  over  1966  (20,000) and  harvest+escapement  for  upstream  i n 1984 the  to  McDonald, action  of  the  1963  to  i n Area  f o r a l l gear extremes  (31,000).  etc)  sockeye  1984  4 on  are  fishery generally  to four  or  more  Oceans  catch  show a v e r a g e  August  salmon  days  Seiners  f o r a l l gears  f r o m one  pass  four  effort  understood.  4  into  steelhead  three  fishing  t o peak p i n k  with  and  of F i s h e r i e s and  years  of  in area  upon t i d a l  from e a r l y t o mid  catch  13,000 p i e c e s again  take  openings last  and  catches  locations, duration  well  and  fishery.  migrating  may  small  (1977)  incidental  information,  not  can  as  b e l i e v e d to pool  fluctuating fishing  at  records  well  Andrews  depending  limited  fishery  hours/day).  s o c k e y e and  annual  of  Sunday e v e n i n g s and  test  occur  b a s i s and  is  only  tshawytscha),  3 days r e s p e c t i v e l y , Aro  a daily  abundant  (0  salmonid  ( S a l v e l i n u s malma)  the  time b e f o r e  B a s e d on  effects  July).  catches  (5 and  escapement  typically (late  of  Other  as  Oguss and  salmon a r e  rates, geartypes,  steelhead  char  commercial  v a r i a t i o n s can  flows.  salmon  reviewed  pink  4.  include chinook  Varden  (1978)  River  Area  (0 k i s u t c h )  Dolly  s o c k e y e and  Skeena  1968)  coho  Skeena R i v e r  considerable  the  fishery  of  (Salmo c l a r k i ) .  i n the  Both  regions  the  searun  trout  steelhead  and  outer  (0 k e t a ) , and  numbers  for  to the  1  just  after  harvests.  The  types  i n area  4  has  in catch occurring in The  same t i m e p e r i o d has  average been  annual  estimated  1 2  at and  37,000 p i e c e s w i t h 1984  (85,000).  distribution catches the  by  total  years  1963  of  Figures the  month.  of  1984).  native  net  sport  3 o u t l i n e the and  steelhead  test  (55,000)  general  fishery  fluctuating  temporal steelhead  nature  harvest+escapement  c a l c u l a t i o n s for steelhead  F i s h e r i e s and factors  for a ten  data,  i n 1966  for  of the  1984.  multiplication figures  occurring  4 o u t l i n e s the  River  U p s t r e a m escapement Department  2 and  commercial  Figure  Skeena to  extremes again  generated  o r more y e a r  Skeena  fisheries  fisheries  Oceans  the  estimated  are  based  indices  (BCFW B r a n c h ,  Skeena  mainstem  fishery  best  steelhead  i n much o f the  on  period  River  i n a l l of  test  are  and  escapement unpublished  also harvested  itself  and  tributaries.  on  by  by  major  13  Figure  2. 1963 t o 1984 mean a n n u a l s t e e l h e a d h a r v e s t by month i n A r e a 4. The week b e g i n n i n g c o d e s a r e Week 7 = J u l y 1, Week 8 = J u l y 8, Week 9 = J u l y l 5 , Week l O = J u l y 21 Week 11=July 29, Week 12=Aug 5, Week 13=Aug12 Week 14=Augl9 Week l5=Aug26 ( S o u r c e , u n p u b l i s h e d d a t a , BCF Branch, 1984).  Figure  3. 1963 t o 1984 mean s t e e l h e a d escapement by month t h r o u g h A r e a 4. The week b e g i n n i n g c o d e s a r e Week 7 = J u l y 1, Week 8 = J u l y 8, Week 9 = J u l y 1 5 , Week 1U=July 22 Week 11=July 29, Week 12=Aug 5, Week 13=Aug12 Week 14=Augl9 Week l5=Aug26 ( S o u r c e , u n p u b l i s h e d d a t a , BCF Branch, 1984).  • i..-o ;  i  c  B I  / I  <VT  ST  i snnnu fcT  £T  ?;T  i TI  \mc OT  6  8  /.  15  Figure  4. 1963 t o 1984 mean a n n u a l h a r v e s t + e s c a p e m e n t i n A r e a 4. d a t a , BCF B r a n c h , 1984).  steelhead ( Source, unpublished  16  * = harvest  + - escapement  17  MATERIALS AND METHODS  Scale Data C o l l e c t i o n  in  Ninety  to  the l a t e  fall  River  stocks  and P r e p a r a t i o n  one h u n d r e d a d u l t  (Kispiox,  scale pattern  major  Zymoetz, B a b i n e ,  Skeena R i v e r i n acetate  during  various  subsequent  Fisheries  samples  (n>100) sample n=l03,  n=30  1977  for  between  were  selected  s i z e s f o r each stock;  was l i m i t e d  to the  caught  previously  steelhead The  taken  m a j o r i t y of  length,  weight,  t h e l e a r n i n g samples f o r  years  for this  years  these  were:  1978 n = 9 l , S u s t u t  of y e a r l y time  comparison  was  study  that  represented  from  having  adequate  Kispiox  1975 n=30 1978 n=62, M o r i c e R i v e r  The a v a i l a b i l i t y  adequately  of  bases  projects.  scales represented  n=60, B a b i n e R i v e r  n=60.  stock.  Branch  data  s c a l e s h a d been  angler  Skeena  Morice-Bulkley)  Branch  f o r age a n d i n c l u d e d  Zymoetz R i v e r  1983  assumption  Most  major  taken  discriminant analysis.  Scale  1975  Fisheries  and r e p r e s e n t e d  These  Sustut,  Stock d e f i n i t i o n  tributaries.  s c a l e s h a d been once r e a d sex d a t a .  B.C.  analysis.  mounted  and  s c a l e samples  (1975-1983) from e a c h o f t h e f i v e  were s e l e c t e d from e x i s t i n g for  steelhead  was  the  true  series  limited. the  River  discriminant  and Kitsumkalum R i v e r s analyses  because  of  data  base  s t r u c t u r e of each for  b u t were n o t u s e d their  data  A major a p r i o r i  S i x t y s c a l e s a m p l e s were a t t a i n e d a n d a n a l y s e d  the Lakelse  1976  1977 n=30  scale  existing  population  River  likely  each  in later  winter-run  origins. Previously  prepared  s c a l e s and t h o s e  prepared  by t h e a u t h o r  18  were s a m p l e d 1956)  on  above t h e  from the  the  left  lateral  nonregenerate methods o f then  line  Chuganova  of  Major  scale  et  (1963),  and  the  first  circulus as  each  scale  identified  Given  the  the  independent a  six  last  aging for  subsample of months  nature  Narver year  along  focus  the  (Figure  and  used  annuli,  circuli  counts,  and  the  methodology  of  any  et  al  narrowing  Saltwater  region  (1969).  of  circuli  of  of  the  annuli  were  narrowing  which  spacing. of  technique  scale was  reading verified  random  sample of  one  hundred  scales  the  Whitesel  1963,  scale  a  after  the  f i r s t marine  Tanaka  growth.  in c i r c u l u s  subjective  source  by  were  following  i n c l u d i n g c u t t i n g over  c i r c u l u s in a  increases  author's  and  the  microfiche  for e s t a b l i s h i n g  followed  were  year's  3M  Chuganova  growth,  (1969),  new  a  from each s t e e l h e a d  Narver  of  the  p r e c e d e d marked  author  on  plus  Two  scales  distinguished  criteria  rows  fin.  C l u t t e r and  t h r o u g h the  s p a c e between c i r c u l i  identified  addition,  The  scale  following  assigned  1961,  scale  four  selected  1954,  Whitesel,  dorsal  under  were  axis one  to  F r e s h w a t e r and  annuli  scale  annuli  and/or  1984),  Henry  freshwater  Chuganova  (Maher  and  acetate  two  were  1972).  subsequent a n a l y s i s .  checks  the  ages  1955,  for  Freshwater  mounted  magnification  P r i n t s were made of  spawning  two  in  5).  checks,  steelhead  were  al.  posterior-anterior  (Clutter  t o the  Initial  Larkin  area  just posterior  34X  growth zones  false  each  p r e v i o u s workers  Maher and  1969,  of  (1963) and  at  reader-printer.  1956,  side  scales  projected  criteria  preferred  fifty  was  i n i t i a l reading.  (Conrad, by  scales.  reread  by  Validation  an In the of  19  Figure  5. A d u l t s t e e l h e a d s c a l e from the S u s t u t R i v e r . T o t a l age i s 3.2+. Shown i s t h e measurement a x i s u s e d f o r a g i n g and measurement o f s c a l e s i n t h i s s t u d y . E a c h a n n u l u s i s marked by t h e h o r i z o n t a l l i n e s ; a r e g i o n of s p r i n g p l u s growth p r e c e d e s o c e a n e n t r y (34X magnification).  20  21  scale  g r o w t h a t age f o r t h i s Age  The  age  of annulus  (1954).  d e p o s i t i o n was t a k e n  complete  (.) where summer  in  spring.  again  the next All  first  scale  (1)  and  after  It  spent  smolting  part  of  r e t u r n i n g i n the f a l l survived, migrated  (1) and p a r t o f t h e n e x t  zone  0.01mm  t o sea  the  next  and s p a w n i n g  back  t o s e a and  summer  again  (+) t o p o t e n t i a l l y  in  spawn  for  four  measurements  (table  zone a n d i n t h e  Measurements  were  to  The  and  Conrad  1984).  dependent  Murai  freshwater  were s e l e c t e d  1969,  Bilton  1971,  scale Lear  and from variables  freshwater  Skeena  one t o f i v e recorded  age.  Only  R i v e r age g r o u p s  studies  a n d Sandeman 1980,  years for  for analysis  pattern  As Skeena R i v e r s t e e l h e a d spend  scale  upon  study  s u c c e s s f u l use i n o t h e r  i n freshwater of  in this  on e a c h  made  h e l d t o low power under a W i l d M5 s t e r e o m i c r o s c o p e . used  Helios calipers  two  print  (Anas  using  2).  scale  and  scale  because of t h e i r  number  (1969).  31,  of l i f e .  (4) b e f o r e  i n each y e a r l y freshwater  variables  years  t o be M a r c h  year  returning i n the f a l l  scale  nearest  of Narver  spring.  counts  marine  print  before  s c a l e s were a n a l y s e d  circuli  full  winter  I t then  winter  before  plus  i n freshwater  the next  saltwater  the next  saltwater  the  winters  i t spent  (S) t h e n e x t spent  attempted.  As an example of age d e s i g n a t i o n , a s t e e l h e a d o f  4.1S1+ i s i n i t s s e v e n t h  four  was n o t  d e s i g n a t i o n s f o l l o w e d t h e methodology  time  Maher  study  from  one t o  five  i n saltwater, the  each  s t e e l h e a d was  s t e e l h e a d of  the  (3 and 4) were u s e d  dominant in this  study. One h u n d r e d  seventy  five  scale  samples  per  week  were  22  Table  2. for  V a r i a b l e s measured each s t o c k .  from  the adult  steelhead  scales  Variable Definition PG P r e s e n c e (1) a b s e n c e (2) of p l u s g r o w t h A1 D i s t a n c e t o second c i r c u l u s i n year 1 A2 Distance to fourth c i r c u l u s i n year 1 A3 Distance to sixth c i r c u l u s i n year 1 A4 T o t a l w i d t h o f s c a l e zone i n y e a r 1 A5 Number o f c i r c u l i h a l f a c r o s s y e a r 1 A6 Number o f c i r c u l i f u l l a c r o s s y e a r 1 B1 D i s t a n c e t o second c i r c u l u s i n year 2 B2 Distance to fourth c i r c u l u s i n year 2 B3 Distance to sixth c i r c u l u s i n year 2 B4 T o t a l w i d t h of s c a l e zone i n y e a r 2 B5 Number o f c i r c u l i h a l f a c r o s s y e a r 2 B6 Number o f c i r c u l i f u l l a c r o s s y e a r 2 CI D i s t a n c e t o second c i r c u l u s i n year 3 C2 Distance to fourth c i r c u l u s i n year 3 C3 Distance to sixth c i r c u l u s i n year 3 C4 T o t a l w i d t h of s c a l e zone i n y e a r 3 C5 Number o f c i r c u l i h a l f a c r o s s y e a r 3 C6 Number o f c i r c u l i f u l l a c r o s s y e a r 3 D1 D i s t a n c e t o s e c o n d c i r c u l u s i n f i r s t ocean y e a r D2 D i s t a n c e t o f o u r t h c i r c u l u s i n f i r s t ocean year D3 Distance to sixth c i r c u l u s i n f i r s t ocean y e a r D4 T o t a l w i d t h o f s c a l e zone i n f i r s t o c e a n y e a r D5 Number o f c i r c u l i h a l f a c r o s s f i r s t o c e a n y e a r D6 Number o f c i r c u l i f u l l a c r o s s f i r s t o c e a n y e a r E1 D i s t a n c e to second c i r c u l u s i n year 4 E2 D i s t a n c e to fourth c i r c u l u s i n year 4 E3 Distance to s i x t h c i r c u l u s i n year 4 E4 T o t a l w i d t h of s c a l e zone i n y e a r 4 E5 Number o f c i r c u l i h a l f a c r o s s y e a r 4 E6 Number o f c i r c u l i f u l l a c r o s s y e a r 4 Additional variables = L Length WT Weight Sex Sex FWA f r e s h w a t e r age SWA s a l t w a t e r age  23  attained 4  from  incidentally  commercial  1984.  salmon  Seiner  and  sampled a t  the  opening.  Fork  nearest  was  60%  Packers  of  the  An  limited  regenerate  by  small  sample of  salmon a s an  example  kept  true population sixty  maximum  expected  and  t o use  catch  Department  sizes data  weekly  fishery  plant  sales slips  of  randomly  (to  the  s c a l e sample.  Rupert  s c a l e data  i n the  All  of  indicted  B.C.  that  20  passes through  B.C  of  and  Fisheries  for  1984  and  and  a high  past  years  incidence  of  base.  Sizes sizes  for  and  they  this  Whitesel  showed t h a t  (95%  confidence  Previous  steelhead  t r u e mean i n any  of  area  were  0.5cm), w e i g h t  study  estimates  scale  were not  standard  s c a l e zone of one.  of  Clutter  Whitesel  the  sample means i n t h i s  study  the  sample means of  n  size  half  sockeye  variation  a circulus  with  The  a  of  sample  author  (in c i r c u l i From t h e  count)  were n o r m a l l y stock  from  should  lie  in a the  formula  assuming  distributed,  a of  used  modified  (1956, p a g e s 75-82) and  from a g i v e n  the  scale pattern variance  available.  deviation  Using  sampling  level) of  followed  (1956).  t o w i t h i n p l u s o r minus one mean  s i x week  through August  day  f o r each  Prince  sample  Clutter  scales.  Skeena R i v e r  nearest  4 incidental  steelhead  methodology  be  the  area  of Sample  Required  could  at  scales present  Determination  the  four  were r e c o r d e d  Attempts  fishery  i n the  mid-July  to  examination  total  facility.  during  two  (to  conducted  Oceans t e s t were  each  sex  steelhead  o f f l o a d s from g i l l n e t t e r s  length  Packers L i m i t e d . to  of  0.5kg) and  sampling  fishery  packer  end  caught  that  95%  of  within  24  two  standard  means t o mean,  errors  l i e within  136  scales  separation  Sample determine  from  classification  in  sockeye  methodology  I  The were  Juvenile  body  form  number of in the  then  97  required  was  the  sample  for  stock  l i e within  the  required.  stocks  involved,  and  the  the  highly  (1969) u t i l i z e d  to  diverse variable  Worlund's  f o r maximum e x p e c t e d e r r o r  favorable the  Skeena R i v e r  sample  high  estimated  seas.  error  at  sizes  Following  rate  steelhead  fishery  calculated  of  f i s h e r y were d i f f i c u l t  Murai  on  95%  a given  means t o  catch,  172)  For  of  in  between  of for  their  correct 15 and  30  samples  required  for  this  between  150  200  (90%  and  level).  Analysis 1983,  thirty  and  seine  Kispiox,  and  Morphological order  an  the  commercial  (page  weekly mixed  electroshocker  in  of  a sample of  salmon  for  In A u g u s t of  Bulkley,  90%  Anas and  chose  classification  confidence  were  stock  deducing  classifying  study  each  steelhead  (1960) p r e c i s i o n c u r v e s  percent.  c i r c u l u s of  the  fishery.  sample mean.  or minus one  s i z e s from t h e  the  given  For  b e c a u s e of  of  a  interval,  s t r u c t u r e of  nature  plus  purposes.  same c o n f i d e n c e  age  of  in  from t h e  comparisons  to assess the  steelhead lower  Zymoetz  different  rivers  Lagler  features  rivers.  following  Hubbs  (1967)  specimen.  These were head l e n g t h  (HL),  were c o l l e c t e d  r e a c h e s of  were c o n d u c t e d  morphological  and  parr  the  Ten were head  Morice-  respectively.  between t h e and  juveniles  t o compare body  overall  measurements,  attained depth  by  from (HD),  each head  25  width  (HW),  (CW),  caudal  body d e p t h  and  post  dorsal  and  size,  the  data  length  depth  (BD), body w i d t h length  (PoDL).  (base  (CD), c a u d a l  As t h e p a r r  were a d j u s t e d  regressions according  length  for  (PrDL),  length  (Thorpe,  1976).  e a c h v a r i a b l e on  to the c o r r e c t i o n  initial  was t h e a d j u s t e d  v a r i a b l e value,  regression  standard  procedure:  of each v a r i a b l e a g a i n s t  in a discriminant  (1)  value,  b was t h e r e g r e s s i o n  was t h e g r a n d mean s t a n d a r d  the  variable  length.  Linear 1981)  discriminant  was a p p l i e d  decision  quadratic  other  studies  discriminant had  features  s c a l e data  linear  steelhead  to  (Fisher  for  was  models  o f t h e two d o m i n a n t using  those  used  assess  systems.  1936, D i x o n  calculating  chosen  successfully features  freshwater scale  the  Linear  because b)  a) the  seemed t o  a n a l y s i s was r e a d i l y i m p l e m e n t a b l e .  (3 and 4) were c o n s t r u c t e d  f o r the  and l o g X'  and c l a s s i f i c a t i o n .  analysis  the  Analysis  d i s t r i b u t i o n s of s c a l e p a t t e r n  n o r m a l and c ) l i n e a r utilizing  used  coefficient  ( l o g ( y ) ) were  Pattern  separation  was  Antilogs  function analysis  to the adult  r u l e s f o r stock  versus  underlying  for Scale  Y  length,  a n a l y s i s of m o r p h o l o g i c a l  Techniques  log  standard  s e p a r a b i l i t y of j u v e n i l e s from t h e t h r e e  Analytical  ages  standardizing  log(y)=logy-b*(logX-logX')  where l o g ( y )  width  were o f v a r i o u s  were removed by  g r a n d mean s t a n d a r d  10)  peduncle  (BW), p r e d o r s a l  the e f f e c t s of a l l o m e t r y t o pooled  Log-log  peduncle  be  Models  age c l a s s e s  variables  which  26  were  both  normally  which had h i g h Univariate  F  scores  ANOVAS,  analyses performed (Dixon,  separating  and  established function  this  analyzing  way  comparisons and  analyses  ANOVAS,  of  variance.  and t h e d i s c r i m i n a n t  study  were  is  multivariate  generated  between  analysis  stems  distinguish  using  BMDP  from  (Jolicouer,  1959).  represents  an  sample)  The steelhead  array  describes space.  dispersion  Multivariate significance The  prerequisite  River  group  location  region  fora l l  analysis  of  of d i f f e r e n c e s rejection  of  fordiscriminant  statistically  univariate  methodology each  stock  (a l e a r n i n g  represented  (vector)  that  steelhead  by  a  from  each  individual  in  from t h e same  in multivariate  space  stock  defined  by  of i n d i v i d u a l s about t h e  variables variance between  group  discriminant  to  be  which  function.  of  (variance-covariance)  average  f o r using  o f s c a l e measurements  Individual  previously  within  known o r i g i n  can  for  discriminant  steelhead,  of  space which  the  o c c u p y a common  stock  Skeena  A  to  inability  normal p r o b a b i l i t y d e n s i t y  linear  multivariate  relative  groups using  established  in multivariate  multivariate  usual  known For  1979).  The r a t i o n a l e  the  in  of p observed v a r i a b l e s  variance  1936).  technique  present  (Pimental,  combination  group  between  a  differences  groups of o b j e c t s  (Fisher,  study.  in  analysis  variance  common  one  multivariate  i s the linear  maximizes  the  in  in univariate  1981) s o f t w a r e .  Discriminant  should  distributed  (the stock was  stock  equality analysis.  used  centroid).  to  test  the  centroids  in  this  between  centroids  Appendix  is a  A o u t l i n e s the  27  methodology study.  of d i s c r i m i n a n t  Figure  and d i s c r i m i n a n t stock  analysis.  analysis  6 shows t h e b a s i c  as  i t  applied  r e l a t i o n s h i p between  space f o r a h y p o t h e t i c a l  three  to  this  euclidean  variable,  three  28  Figure  6. D i s c r i m i n a n t space. Euclidean three v a r i a b l e space f o r three h y p o t h e t i c a l s t e e l h e a d s t o c k s . The m u l t i v a r i a t e swarms o f d a t a p o i n t s ( i n d i v i d u a l s ) , c o n s i d e r e d one v a r i a b l e a t a t i m e , f a i l t o t o s e p a r a t e i n e u c l i d e a n s p a c e a l o n g any s i n g l e v a r i a b l e p l a n e : wx, xy, o r y z . L i n e a r combinations of the o r i g i n a l v a r i a b l e s and p r o j e c t i o n o f t h e r e s u l t i n g c a n o n i c a l v a r i a b l e s t o two a x i s d i s c r i m i n a n t s p a c e b e s t separates the groups. The + d e n o t e s c e n t r o i d s f o r e a c h g r o u p , t h e ( + ) d e n o t e s t h e g r a n d mean c e n t r o i d w i t h a mean o f 0 and a s t a n d a r d d e v i a t i o n o f one i n d i s c r i m i n a n t s p a c e .  3 variab.le e u c l i d e a n  space  w a a + a a b b b + b b b  function  <+)  c c c + c c c  f u n c t i on  2 Two  axis discriminant  space  30  RESULTS  Discrimination  Verification The  of s c a l e  steelhead  readability.  o f Skeena R i v e r  aging s c a l e s used  of  general,  a l l scales exhibited  fifty  poor  often  age.  93%  Age  study  one  placement  scales  hundred  s i x months a f t e r  s c a l e s being  changed  and  narrow  were i n a g r e e m e n t  approximately  Descriptive  definition  annular  selected  A sample o f  nine  annular  made  randomly  source,  in this  exhibted  Some s c a l e s had t o be r e r e a d two a n d  because  which  Steelhead  scale  times  clarity.  In  growth  zones  difficult.  Still,  of the  f o r age  by  the authors'  scales  reread  the i n i t i a l  an  outside  d e s i g n a t i o n of by  the  reading  author  resulted in  f o r d e s i g n a t i o n of age.  statistics  composition Age  five  composition  s t r u c t u r e w i t h i n and between  major Skeena R i v e r  (Appendix collected scale  table  1  and  foranalysis,  growth  zones.  s t e e l h e a d s t o c k s was f o u n d figure  466 h a d For  7).  well  as s i x minor ones  each  readable  fresh  of  the  t o be d i v e r s e  Of t h e o r i g i n a l  475 s c a l e s  and  a l l s t o c k s s i x dominant  (3.1+, 3.2+, 3.3+, 4.1+, 4.2+, 4.3+) were e v i d e n t as  three  freshwater  read with  variable  saltwater  age c l a s s e s  from  the  data  (2.1+, 2.2+, 3.4+, 4.4+, 5.1+, 5.2+)  31  Figure  7. Age c o m p o s i t i o n s t r u c t u r e f o r t h e f i v e s t e e l h e a d s t o c k s used i n the s t u d y . RS d e n o t e s r e p e a t s p a w n e r s ( c o m p i l e d from a p p e n d i x t a b l e 1 ) .  KISPIOX RIVER : n=103  AGE COMPOSITION  aV  x%"  U II I I I yV  y  kS*  t.y »>* &  SUSTUT RIVER : n=90  V  JUuLu l>* Ll* U>*  y1-* ^"  &  MORICE/BULKLEY RIVER : n=89  O £ 0 . 2  %  ki* k>* AGE C L A S S  AGE CLASS  1>* V  I  yi>  &  I  1>" i%* V>* y l * y**  ACE CLASS  O* (..V »>*  I  ^  AGE C L A S S  ZYMOETZ RIVER : n=92  BABINE RIVER : n=91  2  O0 . 4  a>* i>*  ,  P._^_ -_.»-.. P  v^* y** y^* »>* kV k>* AGE C L A S S  &  aV  V  UJUL-JK yi* y V  AGE CLASS  *>*  33  and  s i x or  4.1S1+,  seven  repeat  4.2S1+,  4.1S1S1+,  t h e most common age 3.2+  (27%).  data on  base.  age  3.S1+  c l a s s e s over  Repeat Maiden  average,  spawner  etc).  spawners t o t h e  five  (45%)  freshwater  prior  to smolting  and  (15%)  in saltwater  growth  would  explain  steelhead and  rates  Taft By  the  stock,  Kispiox  River  4.2+  1966,  those  from  of  those  from  Babine River  (62%)  and  the  ages  4.2+  of a g e s 4.2+  3.3+  Table  each stock ocean had  as  age,  long  read and  ocean  3 summarizes from  90%  Babine  ocean w h i l e  only  relatively  2%  River had  spent  3+)  short  (29%),  predominantly  of  from t h e (16%), age  3.2+  composition  sex.  ocean  those  to  River  were  3.1+  or more. . The  2+  (15%);  ages  3.2+  River  were  (13%),  and  smolt  of  age,  steelhead  from the  spent  the  features  residencies  had  from  (26%)  Kispiox River  while  were  3.3+  Sustut  according  steelhead three  Shapovalov  those  and  the  River  River  (22%)  scales  r e s i d e n c i e s (32% had  the  their  (see  (25%);  4.2+  4.3+  c o n t r i b u t i o n s by  Bulkley River of  (50%),  1972)  Skeena  Morice-Bulkley  those  three  maturation  Zymoetz  o f a g e s 3.2+  were  and  in  1981).  3.2+  (43%),  and  predominantly (11%).  and  spent,  (Ricker,  of  stocks  total  years  (66%)  1, and  the had  Reduced  ages  the  the  4.1+  (26%);  two  environments  from  (35%)  predominantly  (18%),  Horncastle  were p r e d o m i n a n t l y  (25%);  (54%)  steelhead  steelhead  of a g e s 4.2+  four  freshwater  to southern  of  stocks  t o spawning.  northern  older  Withler  predominantly  and  i n harsher  compared 1954,  prior  i n 12%  table  (31%)  Skeena  and  one  3.251+,  From A p p e n d i x  spawners were a p p a r e n t  (1%),three  and  (3.1S1+,  a l l s t o c k s were 4.2+  two  years  classes  Morice-  (64% years  1+). in  the  incidence  of  34  Table  3. Age C o m p o s i t i o n f e a t u r e s o f Skeena R i v e r s t e e l h e a d by s e x , s m o l t a g e , a n d o c e a n age ( c o m p i l e d from a p p e n d i x t a b l e 1, ** d e n o t e s maiden spawners o n l y ) .  FEATURE:  PROPORTION OF STOCK Ki spiox Mor i c e Babine Sustut n=1 03 n=90 n=9l n=90  Zymoetz n = 92  1)adults of:** - s m o l t age 3 - s m o l t age 4 - o c e a n age 1+ -ocean age 2 + - o c e a n a g e 3+  46% 54% 1 4% 76% 1 0%  55% 45% 7% 61% 32%  27% 73% 64% 34% 3%  2)repeat  22%  1 4%  1 0%  2%  1 .2/1  1.1/1  1 .5/1  1 .8/1  4)females of:** - o c e a n a g e 1+ - o c e a n a g e 2+ - o c e a n a g e 3+  8% 87% 5%  1 0% 69% 3%  70% 30% 0%  7% 91% 2%  0% 85% 1 5%  5)ma1es o f - o c e a n age 1+ - o c e a n age 2+ - o c e a n a g e 3+  20% 65% 1 5%  4% 55% 41%  53% 38% 6%  9% 88% 3%  2% 49% 49%  3)sex  r a t i o (f/m)  repeat (eg  spawning  Zymoetz  Babine in  spawners  2%).  22%) This  and  stock  Maher  sample  River  and L a r k i n  3.3+  steelhead  1955).  incidence  9% 1 .5/1  f a r t h e s t away (eg kelt  survival  of  the hypothesis  of w i t h i n  stocks.  between  years  such a t r e n d Narver  (10%  (1959)  in  difficult.  in steelhead found s l i g h t  o f age 3.2+ s t e e l h e a d  steelhead  27% 73% 1% 70% 29%  c l o s e s t t o t h e ocean  i n those stocks  s i z e s made t e s t i n g  however, s u p p o r t  the proportions  1968),  least  age c l a s s h o m o g e n e i t y  studies;  i n those stocks  suggests a higher  d o w n s t r e a m Skeena Limited  in  was h i g h e s t  ' 67% 33% 8% 90% 2%  1967,  (73% i n  Several  (Maher  1954,  differences  1967,  60%  in  23% i n 1968),  and 4.2+  ( 8 % i n 1967, 11% i n 1968) i n t h e B a b i n e R i v e r  between  35  years.  62%  of  the  (1977) were of age 4.2+). in  For  this  both  3.2+  the  changed  52%).  little  Given  salmonids  (Ricker,  steelhead  may  river  S i z e s at  function  of  the  steelhead  dominance  at maturity age  selection  stocks.  saltwater  within  (ANOVA  length 8 and  o c e a n age River  and  of  the  class  study  were  age  data  used  major  age  (Morice-Bulkley  has  1976  1977  4.2+=46%  a heritable  basis in  s t r u c t u r e of  for successful  Skeena  River  reproduction  stocks  but  P<0.001, 9)  and  stock  ocean age  ocean  2+  males and  predominantly  were p r e d o m i n a n t l y Variations  2+  3.2+  in  (age  m a l e s were  the  in size  a  4.2+  steelhead  were  different  between  3.2+,  sexes  figure  were q u i t e 11%  longer  =7.9kg) t h a n  stocks  combined  10).  For  a  pronounced; (mean Morice  length= River  length=79.4cm, mean w e i g h t = 4 . 7 k g ) .  ages, K i s p i o x the  sex  be  Both  P<0.001) f o r t h e  alone  to  five  time.  and  (mean w e i g h t (mean  found  residence  significantly  sex  was  f o r the  freshwater  d i f f e r e n c e s by  heavier  Over a l l s t o c k s  not  weight  by  40%  age  and  mean s i z e s a t age  S i z e a t age  w e i g h t s of  88.3cm) and  were  26%  4.1+=43% 4.2+=21%: S u s t u t  1972)  steelhead  similar  Kispiox  River  years  2 summarizes t h e  mean l e n g t h s  given  and  in this  environments.  table  Skeena R i v e r  (figures  3.3+  used  age  Appendix  the  Sustut  between  t h a t age  reflect  specific  and  proportional  4.1+=37% , 4.2+=24% 1977 1983  steelhead  ( 1 % were of age  the M o r i c e  study  classes  Babine River  largest,  and  Sustut  River  Morice-Bulkley  River  steelhead steelhead  smallest. between Skeena  River  steelhead  stocks  36  Figure  8. Mean l e n g t h s o f age 3.2+ a n d 4.2+ s t e e l h e a d from t h e f i v e s t o c k s used i n t h e s t u d y . Shown a r e t h e means +/- one s t a n d a r d e r r o r a b o u t t h e mean, t h e 95% c o n f i d e n c e i n t e r v a l a b o u t t h e mean, and t h e sample s i z e f o r e a c h age c l a s s .  Figure  9. Mean w e i g h t s o f age 3.2+ and 4.2+ s t e e l h e a d f o r the f i v e s t o c k s used i n t h e study. Shown a r e t h e means +/- one s t a n d a r d e r r o r a b o u t t h e mean, t h e 95% c o n f i d e n c e i n t e r v a l a b o u t t h e mean, and t h e sample s i z e f o r e a c h age c l a s s .  37  -i-  12  44 I -I-  121 33  12  I •II  4.2+ IKispioxI  3.2+ 4.2+  3 . 2 + 4.2< 2+  I I 3.2+ 4.  I I 3.2+ 4.2+  IZymoetzI  IMorIce  I Bablnel  ISustut  Rivar  I  I  and A g «  27  22 42 33 12 I -I  -  + -I-  I I 3 . 2 • 4.2+  I I 3.2+ 4.2+  IKispioKl  IZymootzi  I I 3.2+ 4.2+  t i l l 3 . 2 » 4.2+ 3.2+ 4.2+  ;Moricp  I Babine!  River  I  and rtge  I bustict  I  38  Figure  10. Mean l e n g t h s by sex f o r age 3.2+ s t e e l h e a d f r o m the f i v e s t o c k s used i n the study. Shown a r e t h e means +/- one s t a n d a r d e r r o r a b o u t t h e mean, t h e 95% c o n f i d e n c e i n t e r v a l about t h e mean, and t h e s a m p l e s i z e .for e a c h s e x .  lO  9017  L E N G T H  85-! I  20  80-! (cm) •  75-I  70- !  65-1 M  F  iKispioxl  M  F  iZymoetzl  M IMorice River  I M1 1  F  and  i Sex  t  F  Babine!  ! M  i F  ISustut I  40  have  been  noted  (Whately  genetic  differences  feeding  behaviors,  in  et  a l . 1978) and p r o b a b l y r e f l e c t  ocean  or both.  growth  rates,  Table 4 reports  variable  the stock  specific  mean  l e n g t h s and w e i g h t s o f m a l e s  each  stock  Table  4. Mean L e n g t h s and W e i g h t s f o r Skeena R i v e r s t e e l h e a d o f v a r i o u s o c e a n age ( s t a n d a r d e r r o r s about t h e mean a v a i l a b l e from appendix t a b l e 2 ) . Zymoetz  WEIGHT (kg) 1)males o c e a n age 1+ o c e a n age 2+ o c e a n age 3+ 2)females o c e a n age 1+ o c e a n age 2+ o c e a n age 3+  Pattern  1981).  Kispiox  Babine  Sust  63.5 88.3 97. 1  59.0 79.4 91 .5  58.8 77.3 91 .4  55.9 84.4 94.7  64.9 75.2 84.2  57.8 80.4 87.3  56.3 72.5  60.3 76.4  63.5 77.2 87.0  2.2 7.9 8.1  2.2 7.9 9.6  1.8 4.7 7.4  2.0 4.4 7.4  1.8 6.4 8.9  2.6 4.4 5.7  2.4 5.6 7.2  1 .7 3.3  2.0 4.5  2.7 4.3 6.0  F e a t u r e s o f Skeena  to generally  RIVER Mor i c e  57.3 81.0 94.0  A n a l y s i s of the s c a l e them  age  for  use i n t h e s t u d y .  FEATURE LENGTH cm) 1)males: o c e a n age 1+ o c e a n age 2+ o c e a n age 3+ 2)females: o c e a n age 1+ o c e a n age 2+ o c e a n age 3+  Scale  a n d f e m a l e s by o c e a n  ocean  —  —  River  variables  sections  —  steelhead  used  be n o r m a l l y d i s t r i b u t e d  The f o l l o w i n g  —  summarize  in this (using only  study  revealed  BMDP7D, D i x o n , the  width  and  41  circuli  count  scale  features  the  Skeena R i v e r  steelhead  five  differences,  which  are  separately  presented The  had  adult  zone  the notion  smolt  stock  freshwater  and  age  Zymoetz  zone  classes  age 3.1+,  each stock  Morice-Bulkley  River  This  suggested  scale  age  adult  steelhead  Scale  features Appendix  scale  growth  in  adult  stocks  Significant  differences  f e a t u r e s were f o u n d . circuli  count  by p o o l e d for  Kispiox  steelhead  exhibited  yearly  freshwater  in  steelhead  Table  be  age b u t o f p o o l e d  made ocean  stocks.  steelhead  of smolt  majority  6 summarizes t h e  differences  found  (Table 5 ) .  could  between  for  age 3 from t h e f i v e  o c e a n age (3.1+, 3.2+, the  the  within  The same was was  smolt  steelhead  each  smolts of  example,  comparisons  Skeena R i v e r  i n older  age had s i m i l a r  age 4.1+ a n d 4.2+ pattern  in  steelhead  For  study  3 summarizes t h e d e s c r i p t i v e s t a t i s t i c s  Skeena R i v e r  and  in this  f o r e a c h o f t h e t h r e e age  respectively.  o f s m o l t age 3 a d u l t table  ocean  counts  counts,  age ( f i g u r e 1 1 ) ,  rates  adult  P>0.10)  of s i m i l a r  from each of t h e f i v e  smolt  3.2+, a n d 3.3+  d i f f e r e n c e s (ANOVA  that  age u s e d  growth  features.  w i d t h s and c i r c u l i  within  smolt  that  of d i f f e r e n t  for  using  found  scale pattern  River  nonsignificant scale  but  distance  summary t a b l e s .  of o l d e r  of slower  I t was a l s o  same  Intracircular  s c a l e z o n e s and more c i r c u l i  than d i d the a d u l t s  1972).  e a c h s c a l e zone f o r  b o t h zone w i d t h s and c i r c u l i  of younger  freshwater  which supports (Ricker,  stocks.  i n the appendix  steelhead  both wider  scale  reflect  found w i t h i n  of  measured  scale  the f i v e  3.3+ e t c ) .  zone stocks.  scale width Both  42  Figure  11. Mean s c a l e zone w i d t h s f o r s t e e l h e a d o f s m o l t a g e s 3 and 4. Shown a r e t h e y e a r l y s c a l e zone means (mm) f o r a l l s t o c k s combined +/- one s t a n d a r d e r r o r a b o u t t h e mean, t h e 95% c o n f i d e n c e i n t e r v a l , and t h e sample s i z e s f o r e a c h s m o l t a g e . FWA3=smolt age 3, FWA4= s m o l t age 4. The w i d t h o f t h e f i r s t o c e a n y e a r s c a l e zone i s g i v e n w i t h t h e s t a n d a r d e r r o r a b o u t t h e mean i n b r a c k e t s .  43  FRESHWATER  I OCEAN I  44  Table  5. F s t a t i s t i c s a n d a s s o c i a t e d p r o b a b i l i t i e s f o r one way ANOVAs w i t h i n s t o c k s f o r d i f f e r e n c e s i n mean y e a r l y f r e s h w a t e r s c a l e zone w i d t h s f o r v a r i o u s age c l a s s e s i n t h e K i s p i o x , Zymoetz, and M o r i c e r i v e r s . (* d e n o t e s no s i g n i f i c a n t d i f f e r e n c e a t t h e 5% l e v e l of significance).  River  Age c l a s s e s compared  Variable  Kispiox  3.1+,3.2+,3.3+  Zymoetz  3. 1 + ,3.2+,3.3 +  Morice  4.1+, 4.2+  scale  zone w i d t h s  and c i r c u l i  t h e most between t h e f i v e these  differences  Morice-Bulkley adults  from  zones.  the  first  former  scale  3  * * * * * * * * * * * * *  0 .34 2 .06 1 .46 1 .34 0 .54 1 .51 2 .20 2 .10 0 .25 1 .19 0 .89 1 .34 0 .21  counts  i n the second  stocks.  2,45 2,45 2.45 2,45 2,33 2,33 2,33 2,33 1 ,62 1 ,62 1 ,62 1 ,62 1 ,62  A d u l t s of smolt first  year  0.79 0.12 0.27 0.27 0.66 0.23 0.11 0.12 0.78 0.31 0.78 0.27 0.80  year  F i g u r e s 12 a n d  graphically.  suggests  either  year growth  13  and v i c e  after  12)  first  and  scale  age 3 from t h e  year  scale  emergence t i m e s  and/or  scale  environments  for  Interestingly,  both  zone  year  of growth.  t o show s u c h a t r e n d a n d s u g g e s t s e i t h e r  for  food  displacement  circuli  counts  i n M o r i c e - B u l k l e y R i v e r s m o l t age  stock  or  summarize  while  versa f o r the l a t t e r .  (figure  the  earlier  differed  zones  in productive rearing  13) d e c r e a s e d m a r k e d l y  adults  P  A4 B4 C4 D4 A4 B4 C4 D4 A4 B4 C4 D4 E4  R i v e r had the widest  zone w i d t h s  (figure  D.F  t h e Zymoetz R i v e r h a d t h e s m a l l e s t f i r s t  This  better  F  of  parr  into  T h i s was t h e o n l y high  competition  areas  of  less  45  Figure  12. Y e a r l y f r e s h w a t e r s c a l e zone w i d t h s i n a d u l t s t e e l h e a d o f s m o l t age 3 by p o o l e d o c e a n a g e . Shown a r e t h e means a n d t h e 95% c o n f i d e n c e i n t e r v a l a b o u t t h e means.  Figure  13. Y e a r l y f r e s h w a t e r s c a l e zone c i r c u l i c o u n t s i n s t e e l e a d o f s m o l t age 3 by p o o l e d o c e a n a g e . Shown a r e t h e means and t h e 95% c o n f i d e n c e i n t e r v a l a b o u t t h e means.  River  and Y e a r l y  Scale  Zone  47  Table  6. F s t a t i s t i c s and a s s o c i a t e d p r o b a b i l i t i e s f o r one way ANOVAS between s t o c k s f o r s m o l t age 3 s t e e l h e a d by p o o l e d o c e a n age (* no s i g n i f i c a n t d i f f e r e n c e a t t h e 5% l e v e l o f s i g n i f i c a n c e ) ) Variable A4: B4: C4: D4: A5: B5: C5: D5:  Conversely,  age  large  showed  (figures  12  productive and  13)  which  counts)  were  growth  the  stock ages. marine  parr  growth  features  similar  in highly  smolt  (figures  I n v a r i a b l y , the author  0.20,  stocks  zone  i n 3.1+ s t e e l h e a d  age 3.3+ s t e e l h e a d rates  nonsignificant  the  younger  adults  and  counts  of  narrower This  and  d i f f e r e n c e s between t h e s t o c k s  ocean  the  first  than  those  suggests  would  0.20  of within  of d i f f e r e n t  widths  t o be n o t a b l y  f r o m t h e same s t o c k .  i n the  12  f o r a d u l t s of  circuli  i n steelhead  found  age 3  was n o t s i g n i f i c a n t l y  Skeena R i v e r  f o r marine growth  widths  environments.  marine year  0.10 <P<  years  (zone  in adult  and Zymoetz r i v e r s  five  of smolt  between  T h i s may be a t t r i b u t a b l e t o a l a r g e l e v e l  variance  maturation  steelhead  growth  pattern  i n comparable  age 3 (ANOVA: w i d t h s 0.50).  scale  most  i n the f i r s t  between  adult  suggests  Scale  from t h e K i s p i o x  different  <P<  P<0. 02 P<0. 001 P<0. 005 *P>0. 20 P<0. 005 P<0. 001 P<0. 001 *P>0. 20  incremental  which suggests growth  smolt  3.80 8.18 4.50 1 .67 6.26 6.48 5.59 1 .67  Babine R i v e r  environments.  Scale  of  and  circuli  steelhead 13)  P  year 1 width. year 2 width. year 3 w i d t h . 1st ocean " . year 1 circ. year 2 circ. year 3 circ. 1st o c e a n " .  productivity. 3  DF=4,186 F  faster  result  in  when c o m b i n i n g t h e  48  ages of  in a pooled  first  marine  o c e a n age year  individuals  of  steelhead  smolt  Scale  of  Appendix scale  scale  the  same  age  f e a t u r e s of  age  in  adult  stocks  Significant  differences  f e a t u r e s were  a d u l t s of First differed adult five  than  and  circuli  of  circuli  counts  by  between in  counts)  again  a d u l t s of during  years smolt  all  smolt  (4.1+, 4.2+, of  way  4.3+  five etc).  measured  the  circuli  for  4 from t h e  F  scale  statistics  count d i f f e r e n c e s  a n a l y s i s of v a r i a n c e  w i d t h s of  stocks.  the  smolt times  stock of  From t h e  age of  3.  The  4 year  dfferences  Morice  years.  F  for  smolt  age  4 and  the River  first  steelhead.  scores, the  slower  growth  o l d s may  enhance  Figures  14  widths  pooled  (widths year  year  between  for scale  s c a l e zone g r o w t h in  fourth  4 were more d i f f e r e n t  4 showed wide  freshwater  the  scale pattern analysis.  large  for age  Interestingly,  age  summarizes  five  age  adults  Incremental  subsequent  the  residence  graphically. was  when  trend.  majority  c o u n t s and  smolt  differentiation the  used.  only  descriptive statistics of  i n a one  were a d u l t s of  longer  summarize  7  valid  4.  t h e most between  stocks  stock  stocks  age  steelhead  rates  15  year  the  the  Table  comparisons  steelhead  s c a l e zone w i d t h and  five  smolt  are  o c e a n age  for  stock  be  opposite  4 adult  pooled  found.  f o r the the  by  may  age  steelhead  Skeena R i v e r  between  ocean  4 showed t h e  smolt  Between  growth  t a b l e 4 summarizes  growth  generated  analysis.  and  and  ocean  and  and  age  circuli small  Sustut  in  River  incremental  s c a l e growth  zones  Wider  fewer c i r c u l i  were  but  49  Figure  14. Y e a r l y f r e s h w a t e r s c a l e zone w i d t h s i n a d u l t s t e e l h e a d o f s m o l t age 4. Shown a r e t h e means and t h e 95% c o n f i d e n c e i n t e r v a l a b o u t t h e means.  Figure  15. Y e a r l y f r e s h w a t e r s c a l e zone c i r c u l i c o u n t s i n a d u l t s t e e l h e a d of s m o l t age 4. Shown a r e t h e means and t h e 95% c o n f i d e n c e i n t e r v a l a b o u t t h e means.  50  n=37  n=4B  n=65  n=30  n=5<?  . 40-  W I D T H  !./!  . 30-  V  (mm)  1/  . 20-  15-I  1 2 3 4 Kispiox  1 2 3 4 Zymoetz River  I  1 2 3 4 Morice  1 2 3 4 Babine  and Y e a r l y S c a l e  n=37  n=4B  1 2 3 4 Kispiox  1 2 3 4 Zymoetz  n=65  1 2 3 4 Sustut  Zone  n=30  n=59 !  5-1 i  0  i  River  1 2 3 4 Morice  and Y e a r l y  1 2 3 4 Babine Scale  Zone  1 2 3 4 Sustut  51  Table  7. F s t a t i s t i c s a n d a s s o c i a t e d p r o b a b i l i t i e s f o r one way ANOVAS between s t o c k s f o r s m o l t age 4 s t e e l h e a d by p o o l e d o c e a n a g e (* no s i g n i f i c a n t d i f f e r e n c e a t t h e 5% s i g n i f i c a n c e l e v e l ) Variable  DF=4,234 F 8.61 7.40 7.64 4.32 1 2.82 18.45 3.40 0.93 5.79 4.99  year 1 width year 2 width year 3 width 1st ocean " year 4 width year 1 #circ year 2 #circ year 3 #circ 1st ocean " year 4 #circ  A4: B4: C4: D4: E4: A5: B5: C5: D5: E5:  apparent  i n K i s p i o x R i v e r smolt  year  (figure  15) even  width  (figure  14).  age  contrast  marine year between smolt  the  between  4  less smolt  3 adults. vs  3  year  differences. salmonid  to the r e s u l t s growth  the  first  increasing in  adults  zone  of  smolt  growth.  of the previous s e c t i o n ,  (width)  was  River  significantly  stocks f o r adult  P<0.05, c i r c u l i  count  w i t h i n stock v a r i a n c e of f i r s t age 4 a d u l t s o f d i f f e r e n t  first  different  steelhead of  P<0.05).  This  year marine  growth  o c e a n age t h a n  f o r smolt  V a r i a b l e f e e d i n g and/or m i g r a t i o n a l p a t t e r n s old  smolts  from  Healey  (1983)  notes  smolts  similarily.  p a t t e r n s of s c a l e  f i v e ' Skeena  characteristically or  z o n e s were  K i s p i o x a n d Zymoetz R i v e r  age 4 (ANOVA w i d t h  suggests  age  scale  age 4 s t e e l h e a d a f t e r  though t h e s c a l e  4 a g a i n had s i m i l a r In  P P<0.001 P<0.001 P<0.001 P<0.005 P<0.001 P<0.001 P<0.05 *P>0.50 P<0.001 P<0.002  (by  stock,  each  stock  that  size,  may  different age,  t o marine environments  etc)  by g r o w i n g  for  e x p l a i n the "types" may  of  respond  differently  52  Scale  features  of  age  Skeena R i v e r to  assess  specific remove  any  scale  i n age.  g r o w t h was  classes  (3.2+,  scale five  of  variance  in scale  as  to vary  4.2+)  noted  were  also  pattern  the  so  Age  as  to  growth a t t r i b u t a b l e previously,  two  scale  to  freshwater  in steelhead  dominant  analyzed  compare  analyzed  features.  desirable  nonsignificantly  were  to b e t t e r  tables  5 and  Figures  for  stocks.  and  8 for  17  zone w i d t h s a l o n e .  were  found  age  of  steelhead  to  satisfy  growth  in  widths  16  and  17  show t h a t  of  the  pooled  comparable However, t h e  the  age  general  the  the  the  ocean  between  and  4.2+  first  numbers of  between  stock  between  and, and  respectively.  stock  in years  were s i m i l a r  ( t a b l e 8)  12  and  first the  Figures to  those 14).  marine  result  two  first  and  in f i r s t  i n the  for  differences  steelhead.  this  are  differences  (Figures  of  analyses  features  counts  4.2+  steelhead,  circuli  way  interestingly,  differences  growth  statistics  from e a c h  scale  classes  analyses  scale  one  age  zone d i f f e r e n c e s  age  stock  of  for c i r c u l i  3.2+  nonsignificant  3.2+  steelhead  certain  steelhead  i n b o t h age  univariate  results  two the  the  4.2+  Significant  4.2+  marine year  for  in  i n a l l zones except  for  Concerning  and The  summarize  scale  three  3.2+  differences  in table  16  6 summarize  i n age  Skeena R i v e r  summarized  widths  scale  variations  Still,  features  the  and  in  age  year.  Appendix for  specific  usually  found  o c e a n age.  steelhead  is  However,  different  marine  of  identification  m e t h o d o l o g y and  4.2+  differences  possible  variations  and  steelhead  stock  stock  3.2+  year  suggests  ocean  year.  distances  to  53  Figure  16. Y e a r l y f r e s h w a t e r s c a l e zone w i d t h s i n s t e e l h e a d of age 3.2+. Shown a r e t h e means a n d t h e 95% c o n f i d e n c e i n t e r v a l a b o u t t h e means.  Figure  17. Y e a r l y f r e s h w a t e r s c a l e zone w i d t h s i n s t e e l h e a d of age 4.2+. Shown a r e t h e means and t h e 95% c o n f i d e n c e i n t e r v a l a b o u t t h e means.  54  River  and  Yearly Scale  Zone  55  Table  8. F s t a t i s t i c s and a s s o c i a t e d p r o b a b i l i t i e s f o r one way ANOVAS between s t o c k s f o r age 3.2 + and 4.2+ f r e s h w a t e r and f i r s t m a r i n e y e a r s c a l e zone w i d t h s and c i r c u l i c o u n t s (* i n d i c a t e s no s i g n i f i c a n t d i f f e r e n c e a t t h e 5% s i g n i f i c a n c e level) . Age  Variable A4: B4: C4: D4: E4: A5: B5: C5: D5: E5:  circuli all  year 1 width year 2 width year 3 width 1st o c e a n " year 5 width year 1 circ. year 2 circ. year 3 circ. 1st o c e a n " year 5 width  i n the f i r s t  significantly  are d i f f i c u l t growth  are  being  feeding  sampled Sustut  different  between t h e s t o c k s .  Either  reflected  yearly  i n the data  and  reflect  between  These  variations  results  in  ocean  base o r , a l t e r n a t i v e l y , stock  specific  genetic  years  s t o c k s between y e a r s even  ( e g 3.1+, 3.2+,3.3+  i n t h e same y e a r  exhibited  R i v e r s t e e l h e a d of smolt  revealed  ( v a r i a b l e s D1, D2, and D3) were  were made t o a s s e s s t h e d e g r e e  Zymoetz R i v e r a d u l t n=20)  zone  differences.  within ages  4. 2+ DF= 4, 185 F P 7. 41 P<0. 001 4. 75 P<0. 005 2. 93 P<0. 05 0. 1 6 *P>0. 50 1 1 .25 P<0. 001 18. 93 P<0. 001 2. 75 *P,0. 10 0. 79 *P<0. 50 4. 46 P<0. 005 5. 26 P=0. 001  Age  —  marine  are real  Comparisons  different  --  P<0. 002 P<0. 002 P<0. 001 P<0. 05  --  Scale pattern v a r i a t i o n  variation  --  5. 22 5. 15 8. 22 3. 06  to explain.  the d i f f e r e n c e s and/or  3. 2+ DF= 4, 1 36 F P 3. 97 P<0. 01 6. 64 P<0. 001 5.' 68 P<0. 001 2. 68 *P>0. 05  though  e t c ) , and thus non-significant  steelhead of brood  years,  differences.  age 4 (1977 n=24, 1983 n=38) and  s t e e l h e a d of smolt  significant  of s c a l e p a t t e r n  age  3  (1975  between y e a r s d i f f e r e n c e s  n=l9  1978  (table  9)  56  only  f o r second year  Sustut  Table  River  and  stock.  first  marine year  While  it  s c a l e growth  is . difficult  in  t o draw  the  strong  9. R e s u l t s of one way ANOVA'S f o r between y e a r s d i f f e r e n c e s i n s c a l e growth f o r Sustut R i v e r smolt age 4 and Zymoetz R i v e r s m o l t age 3 a d u l t s t e e l h e a d A = S u s t u t 1977 n=24, 1978 n-38 B=Zymoetz 1975 n=l9 1978 n=20 (* no s i g n i f i c a n t d i f f e r e n c e a t t h e 5% l e v e l )  Scale  Variable A4 A5 df 1,60 F .03 .16 (A) P .85 .69 * * df 1,39 F .95 1 .59 (B) P .33 .23 * *  conclusions  index  be  revealed  the  five  known.  this  in  For  scale pattern  than  the  stocks.  E4  E5  2.23  5.71  .04  .02  .84 *  .85 *  <. 00 1 <. 00 1  2.85  .01  .00  .17  .04  1 .62  -  -  .10 *  .92 *  .97 *  .69 *  .84 *  .21 *  -  -  results,  study  small  15.22  fairly  an  variation  between y e a r s  tributaries.  The scale  primary concern  g r o w t h between y e a r s in scale pattern  .02  within  was  stocks i f  years.  examination  p r o d u c t i v i t y and the  steelhead  s t a b l e between  stability,  . 14 *  yearly variations in scale  i n a l l Skeena R i v e r  remain  instream  changes  D5  17.68  major Skeena R i v e r on  D4  .04  considerable  fluctuations  C5  .04  environmental  data  C4  5.92  expected  conditions of  B5  4.25  from t h e s e  f e a t u r e s may rearing  B4  of  As flow  for  each  i n f l u e n c e of growth that  stocks  is  an rate of such not  any  changes  be  smaller  g r o w t h between y e a r s  between  57  Plus  growth  The the  i n c i d e n c e of  first  ocean y e a r  (35.6%) and (21.7%),  may  the  was  Sustut  differ  level  of  significant  adult  steelhead  of p l u s g r o w t h  trends  of  smolt  than  and  stocks  maturity  While  (and  reached were not age  stocks.  As  for  distributed, accepted. was  observed  by  analysis of  the The  equality  is  MANOVA  differences  separating  the m a j o r i t y  of  the  readily  still in  (Conrad,  justified  detecting  between g r o u p s  the  Babine  last  rivers smolt  size  according  freshwater  age  to  year.  a p p a r e n t , Skeena  t o show a h i g h e r  smolt  models five  of  hypothesis  workers  the  of  Kispiox  (20.6%)  between y e a r s )  River  incidence  4.  were Skeena  constructed  of  equal  steelhead  were  normally  dispersion  significant  finding  was  matrix often  However, d i s c r i m i n a n t  i n most c a s e s  (Pimental,  normality  the models, a  1984).  in  River  multivariate  r e j e c t e d f o r s e v e r a l of other  by  s c a l e v a r i a b l e s used  assumption  null  from t h e  r a p i d growth to  in  onset  Discrimination  Twelve d i s c r i m i n a n t a n a l y s i s study  to  followed  Zymoetz  3 tended  d i d a d u l t s of  Stock  this  in steelhead  growth r e f l e c t s  between  growth p r i o r  (33.3%) r i v e r s  (20.7%),  Plus  plus  highest  Morice-Bulkley  respectively. and  freshwater  because of ' and  1979).  the  power  nonsignificant  58  Separate  freshwater  Figure  18  discriminant smolt  age  and  centroids of  accounted  original  twenty  construction,  From  3 had much w i d e r  distances.The  DF=  10  indicated  The  36.3%,  highly  and of  variance  best  and S u s t u t  River  River  high  five  for steelhead  that  degree  of  age 3 a d u l t s were  most  to  the  the  B1,  years  steelhead  on t h e  two  of smolt  and age  s t e e l h e a d of  primarily  circuli  separated  on t h e b a s i s of s c a l e  of c e n t r o i d s Zymoetz)  in figure  portrays  18 the  s c a l e v a r i a b l e o v e r l a p between t h e  of smolt  the p a t t e r n s  in  by t h e i n t e r z o n e  steelhead  The p r o x i m i t y  being  steelhead  d i d Kispiox River  reflected  of  discriminant function  circulus  Sustut  River  Kispiox  relatively  Ten  selected for function  18, t h e f i r s t  second  being  A2.  canonical  discriminating variables  figure  to the  four  respectively.  s c a l e v a r i a b l e s were  Kispiox  the  stocks  17.8%, and 7.4% o f t h e  second d i s c r i m i n a n t f u n c t i o n  C1  of  scale v a r i a b l e s are  variance  178).  38.5%,  from M o r i c e - B u l k l e y  revealed  steelhead  significant  s c a l e zones than  age 3, t h i s  stocks  stock  t h a t a l l t e n c o m p a r i s o n s were  As p r e v i o u s l y n o t e d ,  (especially  Only  five  comparison  the four best  separated  variables  a  Pairwise  stock  of d i s t a n c e s  Babine  of adult  40 676 P<0.001).  for  four  C1, C2, and A2. primarily  River  ocean a g e .  3.76-7.98  between  smolt  DF=  showed  explained  three.  pooled  5.24  F=  functions  basis  Skeena  results  d i f f e r e n c e s among c e n t r o i d s f o r t h e t h e f i v e  ( a p p r o x i m a t e F=  (range  the  M u l t i v a r i a t e a n a l y s i s of  significant  of  summarizes  analysis using 3  included.  age d i s c r i m i n a n t models  age 3.  of freshwater  similar  for  the  Paitwise  comparisons  s c a l e growth i n smolt Sustut  to  Zymoetz,  59  Figure  18. Discriminant function analysis describing scale p a t t e r n v a r i a t i o n i n a d u l t s t e e l h e a d o f s m o l t age 3. The l e t t e r s i n d i c a t e t h e s t o c k c e n t r o i d s S = S u s t u t Z=Zymoetz K = K i s p i o x M=Morice B=Babine *=grand c e n t r o i d , t h e open c i r c l e s i n d i c a t e t h e 90% c o n f i d e n c e interval a b o u t e a c h c e n t r o i d ( f r o m P i m e n t a l , 1979), and t h e l i n e s p o i n t t o t h e next most s i m i l a r s t o c k i n d i s c r i m i n a n t space. The f i r s t two s t a n d a r d i z e d discriminant f u n c t i o n s a r e g i v e n below. D1  = -37.23C2 +61.33C1 +0.19A5 +0.50B4 -27.19A2 +69.28B1 +0.27PG -24.17B3 +5.80D3 +25.50A1 -0.12  D2 = 25.01C2 -58.24C1 -0.09A5 -.83B4 +29.27A2 -26.15B1 +0.28PG +0.11B3 -7.04D3 -24.49A1 -4.12  60  -2.0 -2.0  -1.5  -1.0  -0.5  0  FUNCTION 1  +0.5  +1.0  +1.5  61  Zymoetz  to  Kispiox, Kispiox  Morice-Bulkley holdout smolt  procedure,  stocks.  45.3%  Using  (range  10) by t h e c l a s s i f i c a t i o n  Figure  19  discriminant smolt  age  included. highly stocks  summarizes  analysis using 4  and  significant  between  2.04-14.87 DF=  accounted variability  the Skeena  for  the f i v e  13 2 2 2 ) .  50.5%  and  for  discriminating the f i r s t  of  smolt  of  among 52 861  two  f o r the t h i r d  4  from  the Sustut  of d i s t a n c e s t o the second c i r c u l u s  The River  distinctly  second  variables  small  steelhead  C3  and  function  from t h e o t h e r and  B3.  again  found  Pairwise  four  best  From  figure  separated  Rivers on  two and  as the  three  K i s p i o x and  t o be t h e most  comparisons  adults  primarily  similar.  separated  on  stock  scale variables  i n years  stocks  of  functions  the  primarily  four  (range  fourth  thirty  five  pairwise  between  i n the M o r i c e - B u l k l e y ) .  were a g a i n  discriminant  steelhead  different,  of  functions  and M o r i c e - B u l k l e y  basis  Zymoetz R i v e r  All  C1, B1, C3, and B3.  t h e most  i n the Sustut,  f o r the  canonical  discriminant function primarily  age  stock  indicated  significant  construction,  being  (large  again  P<0.001).  The f i r s t  five  steelhead  centroids  were  v a r i a b l e s being  origin  scale variables are  variance  stocks  function  of  a  adult  Only  T h i r t e e n of the o r i g i n a l  selected  19,  River  of  26.1% o f t h e e x p l a i n e d  (17.6% and 5.7%  respectively).  DF=  69%) o f t h e  to stock  results  ocean a g e .  7.48  (1975)  technique.  differences  ( a p p r o x i m a t e F=  Lachenbruch's  classified  Multivariate analysis  comparisons  were  pooled  t o K i s p i o x , and  Zymoetz 2 9 % - S u s t u t  age 3 a d u l t s were c o r r e c t l y  (Table  F=  to Kispiox  t o Zymoetz, B a b i n e  the  revealed  Babine  basis  of  that the  J  •s  A  62  fa  3 i-2SS32  I? UiSi  -SISS?  Id I-=sS2s  I553-S5S55 u«  ?§Ss =  Hi  s'is=i  I;  1! §1  2^ -*a5Si  1  |-  -ss?ss  !J! 1  2=SS2SS  I  33 52521 It  !l«  Si  3  V  I  SSsrs  •A  |-i!i£S  a;  i«  si I.  b  I  4-1 -V  "r^§=?  to  O >i  cn cn cu in o rtj rH u u • 4J C <0 O -ri E 4J cn C D  1!1 i.:; q  5-55=55  1?  -«Sss  5  "1  ^  1  r- - r-  *  J:  5-'.  ni  "5 SeESSSS  «  IJ  o m o •rH cn 4J 3 » M ro o u in •r-t rH Jtf in O 4J •<Hin 6 T3 in (0 rH 4-> O U C 4->  J I-5S5S5  I l-ssssi  11 i - s = = i s  K  c  *-S2s5a  -I!  ic  I  o  ri  I5  a  il.....  C  3 :::::  S3 S-5SSS5 s:l-55S5s  m>  C  E-2  u  4)  U  C  -=531=  U3  I O  i!  t>  •<H (0  E CD •H . C V4 rH U CU  cn 0) in • n 4J o> rH TJ in  Numbering e r r o r .  Text f o r l e a f 63 not  available.  64  Figure  19. Discriminant function analysis describing scale p a t t e r n v a r i a t i o n i n a d u l t s t e e l h e a d o f s m o l t age 4. The l e t t e r s i n d i c a t e t h e s t o c k c e n t r o i d s S = S u s t u t Z=Zymoetz K = K i s p i o x M=Morice B=Babine *=grand c e n t r o i d , t h e open c i r c l e s i n d i c a t e t h e 90% c o n f i d e n c e interval a b o u t e a c h c e n t r o i d ( f r o m P i m e n t a l , 1979), and t h e l i n e s p o i n t t o t h e n e x t most s i m i l a r s t o c k i n d i s c r i m i n a n t space. The f i r s t two s t a n d a r d i z e d dicriminant f u n c t i o n s a r e g i v e n below. D1 = -9.61E3 +0.15A5 +0.09D5 +8.50D3 -15.24B3 +24.92B1 -6.28E4 -22.50C3 +33.68C1 +0.13B6 +0.13E5 +1.52A4 -0.19D6 +0.89 D2 = -1.30E3 -0.05A5 +0.01D5 +0.42D3 -1.61B3 -1.00B1 -0.41E4 -2.22C3 +0.87C1 +0.01B6 -1.02E5 +0.07A4 -0.01D6 +5.64  + 1.5-  -2.0  i•  -2.0  ii  -1.5  ii  -1.0  iI  -0.5  I  i  ii  +0.5  0  FUNCTION  •  1  ii  +1.0  ii  +1.5  66  patterns most  of  freshwater  similar  Kispiox  stocks.  (range  adults  the  Using  inclusion  Lachenbruch's  size  the d i s c r i m i n a n t a n a l y s e s stock  discriminance.  stocks age.  were 57.1%  Skeena smolt and  shown (range  adults age  13)  weight  smolt  not  data age  age  (range  with  65%)  origin  of  the  the  the  smolt  Zymoetz 5 0 % - S u s t u t  with  increased five  respect to s i z e s  inclusion  of  11).  weight) in  as  to  stock  Skeena  (Table  4 Skeena  suprising  greatly  4  ( l e n g t h and  3 and  too  procedure,  age  66%)  of  at 3  of  the  (tables  12  length  and  data.  based  on  smolt  age  20  7.02  summarizes  analysis  scale  the  using  age  results  Skeena  variables alone.  3/pooled  were f o u n d  The  analysis.  40  483  P<0.001) by  accuracy  f o r t h e age  3.2+  Zymoetz  26%-Babine  71%)  scale  (range variables  variables  Zymoetz alone  in  Again,  Sustut,  five  five  stocks  stocks.  scale  conjunction 72%)  B a b i n e and  was  variables with  (Table  size 13A).  3.2+  to  the  differences (approximate  multivariate analysis  using  stock  a d u l t s of age  significant  d i s c r i m i n a n t model  38%-Babine  a  r e s u l t s were s i m i l a r  among t h e c e n t r o i d s f o r t h e  DF=  of  River  i n a l l p a i r w i s e c o m p a r i s o n s between  using  Morice-Bulkley  classified  discriminant  and  f o r smolt  58.6%  the  were  Zymoetz t o K i s p i o x ,  t o s t o c k of o r i g i n  related  differ  adults  (1975) h o l d o u t  of  Zymoetz 3 8 % - S u s t u t  and  4  4 Skeena a d u l t s were c o r r e c t l y  Figure  F=  71%)  T h i s was to  age  t o M o r i c e - B u l k l e y , and  classified of  i n smolt  t o Zymoetz,  Zymoetz 4 0 % - S u s t u t  were c o r r e c t l y  The  growth  Sustut  t o Zymoetz, B a b i n e  to Babine 58.2%  for  scale  of  variance  Classification 51.8% alone  (range and  related Using  M o r i c e - B u l k l e y R i v e r age  61% data  scale 3.2+  67  Figure  20. Discriminant function analysis describing scale p a t t e r n v a r i a t i o n i n s t e e l h e a d o f age 3.2+. The l e t t e r s i n d i c a t e t h e s t o c k c e n t r o i d s S = S u s t u t Z=Zymoetz K = K i s p i o x M=Morice B=Babine *=grand c e n t r o i d , t h e open c i r c l e s i n d i c a t e t h e 90% c o n f i d e n c e i n t e r v a l s a b o u t e a c h c e n t r o i d (from P i m e n t a l , 1979), a n d t h e l i n e s p o i n t t o the n e x t most s i m i l a r s t o c k i n d i s c r i m i n a n t s p a c e . The f i r s t two s t a n d a r d i z e d d i s c r i m i n a n t f u n c t i o n s a r e g i v e n below. D1  = +0.07C5 +0.01A5 +26.06C2 +13.90B1 -16.62A2 -0.03D5 +1.08D4 -0.08B5 +0.42B6 +5.26A1 -8.2  D2 = +0.02C5 -0.01A5 -4.28C2 +17.26B1 -10.02A2 +0.05D5 -0.35D4 -0.07B5 +0.11B6 +9.55A1 -0.4  -1.5-i -2.0  ii  -2.0  ii  -1.5  ii  -1.0  ii  -0.5  ii  i  I  0  FUNCTION  +0.5 1  I  i  +1.0  >i  +1.5  69  steelhead (figure  were 20)  separated primarily  second c a n o n i c a l basis  of  patterns most  along  B1.  of freshwater for  the  Sustut  Kispiox  stocks. 21  comparisons  to  Kispiox,  summarizes  the  Significant  d i f f e r e n c e s between s t o c k  again  Classification Zymoetz (range in  conjunction  alone,  canonical  71%)  the  the  patterns  (figure  21).  of freshwater  similar  C1  stocks.  steelhead  were  stock  of age  4.2+.  (approximate  of  model was  variance.  55.3%  (range  and  59.5%  scale variables scale  variables  steelhead  function  (figure  while  the stocks  age  the  on  comparisons  s c a l e growth i n  F=  were  age 4.2+  B1  to  five  comparisons  Using  and  Pairwise  f o r the Sustut  a  13B) u s i n g  canonical  variables  t o Zymoetz, B a b i n e  Babine  (Table  r e l a t e d data.  first  of  scale v a r i a b l e s alone  function p r i m a r i l y separated  v a r i a b l e C3  were most  size  that the  steelhead  analysis  M o r i c e and B a b i n e R i v e r  scale  of  to  with  along  p r i m a r i l y ~ by  using  44%-Sustut  the  Morice-Bulkley  pairwise  f o r t h e age 4.2+  70%)  on  revealed  centroids  multivariate  accuracy  Sustut,  separated  Kispiox  from  -40%-Sustut Zymoetz  Skeena R i v e r  P<0.001) and i n a l l  evident  and  results  analysis using  44 671  the  t o Zymoetz, Zymoetz t o K i s p i o x ,  discriminant  DF=  function  the stocks  s c a l e g r o w t h i n age 3.2+  t o Zymoetz, B a b i n e  6.91  canonical  s c a l e v a r i a b l e s C2 and A2 w h i l e  Pairwise  Kispiox  Figure  first  function p r i m a r i l y separated  variable  similar  by  the  21) second  the  revealed  4.2+  were  basis that  steelhead  t o Zymoetz, Zymoetz t o K i s p i o x ,  to Morice-Bulkley,  and  Morice-Bulkley  70  Figure  21. Discriminant function analysis describing scale p a t t e r n v a r i a t i o n i n s t e e l h e a d o f age 4.2+. The l e t t e r s i n d i c a t e t h e s t o c k c e n t r o i d s S = S u s t u t Z=Zymoetz K = K i s p i o x M=Morice B=Babine *=grand c e n t r o i d , t h e open c i r c l e s i n d i c a t e t h e 90% c o n f i d e n c e i n t e r v a l a b o u t e a c h c e n t r o i d ( f r o m P i m e n t a l , 1979), a n d t h e l i n e s p o i n t t o t h e n e x t most s i m i l a r s t o c k i n d i s c r i m i n a n t s p a c e . The f i r s t two s t a n d a r d i z e d d i s c r i m i n a n t f u n c t i o n s a r e g i v e n below. D1  = +0.21A5 -9.31E3 +9.73D3 +0.05D5 -24.11B3 +39.69B1 -3.32E4 -24.14C3 +46.52C1 +0.15A6 + 0.73  D2 = -0.09A5 -0.48E3 -0.35D3 -0.02B1 -0.30E4 +1.26C3 + 3.41  +0.01D5 -0.46B3 -0.76C1 +0.58A6  71  FUNCTION  1  72  Pooled  freshwater  Several analyses  pooled  (relative  i n age  Because  to  within  both  age  suggested  the b a s i c  freshwater  (1975) h o l d o u t pooled  smolt  conjunction  with of  Zymoetz, stocks.  age  age  4 s t e e l h e a d from scale  certain  age  i n each  analysis  model.  t h e mean  age/pooled  system.  o c e a n age  Using  72%)  using  weight.  Figure  accuracy  scale  scale  g r o w t h by  t o Zymoetz, Zymoetz,  (range  and  61.8%  summarizes  pooled  age  Morice-Bulkley  in the  f o r the a l l  revealed  Zymoetz t o B a b i n e ,  and  for  variables  Pairwise comparisons  a  52.5%  alone  22  for  Lachenbruch's  model was  variables  Table  results  classification  using scale  model.  to  stocks  (smolt)  freshwater  Kispiox  models.  ages  growth  zones  the  t h a t the v a r i o u s f r e s h w a t e r  age/ocean  Sustut  features  specific  s t o c k c e n t r o i d s i n d i s c r i m i n a n t space  pooled  f o r the  scale  Morice  l e n g t h and  p a t t e r n s of  similar  and  stock  the  50%-Sustut  Zymoetz  that large  in  (range  the  smolt  (eg s m a l l  67%)  variable  to  3 and  Zymoetz 3 5 % - S u s t u t  placement  s i z e s a t age,  discriminant  procedure  age d i s c r i m i n a n t  the assumption  to best d e s c r i b e o v e r a l l  14 summarizes  ocean  d i f f e r e n c e s ) would d i s t i n g u i s h  resorting  smolt  this  be p o o l e d  pooled  age/pooled  stock  t o grow s i m i l a r i l y  River)  the  smolt  composition,  stocks without  tended  d i s c r i m i n a n t models  were c o n s t r u c t e d under  differences  five  age  that  were most Babine to  to  Zymoetz  73  Figure  22. Discriminant function analysis describing scale p a t t e r n v a r i a t i o n i n a d u l t s t e e l h e a d of p o o l e d smolt age. The c i r c l e s i n d i c a t e t h e s t o c k c e n t r o i d s S = S u s t u t Z=Zymoetz K = K i s p i o x M=Morice B=Babine *=grand c e n t r o i d , the open c i r c l e s i n d i c a t e t h e 90% c o n f i d e n c e interval a b o u t e a c h c e n t r o i d ( f r o m P i m e n t a l , 1979), a n d t h e l i n e s p o i n t t o t h e n e x t most s i m i l a r s t o c k i n d i s c r i m i n a n t space. The f i r s t two s t a n d a r d i z e d discriminant f u n c t i o n s a r e g i v e n below. D1 = 0.45 WT -0.13 A5 -3.97 D1 +14.76 C3 -18.14 C -0.06 D5 -0.06 C5 +0.13 L -2.31 D3 +0.94 A2 +0.13 D6 +0.02 B6 -15.11 B1 +9.44 B3 -7.28 A +0.06 PG -1.81 D2 = 0.30 WT +0.15 A5 +7.85 D1 -17.25 C3 +44.83 C +0.12 D5 +0.11 C5 -0.02 L +5.66 D3 -24.54 A2 -0.13 D6 +0.26 B6 +35.63 B1 -11.13 B3 +18.53 +0.06 PG -4.47  74  + 1. 5 -  FUNCTION  1  75  Discrimination  As at  both  similar  effort  was  by  male and rates  differences  age/pooled  o c e a n age  with  52.6%  Several order.  of other  (1975) h o l d o u t  s e c o n d and  76%)  results  stock third  differentiation  of  r a n g e of v a r i a b l e  stocks,  as  i n d i c a t e d by  highly  with  the  using the  the  success must  rely  the  of  to  smolt Skeena origin (range  while stock  steelhead  lowering  Secondly,  was  had  each  r a t e s of  between model.  did  of e a c h the  to  mean  from  into parr  high  in  the v a r i a b l e s  consistently  low  of  above models a r e  growth, which suggests  fairly  male  i n a l l models than  for  on  only  65.%  Misclassifications  were  in  same m o d e l .  River  success  overlap  in  River  stock  classified  differences  significant  Skeena  ( t a b l e 15)  most p r o m i n e n t w e l l  the  were  technique  freshwater  is  little  female  to  a l l models.  grow  1959)  reason,  classified  responsible  to  were f o r a p o o l e d  Zymoetz  stocks.  of  though the  sex  a l l of  and  Thirdly,  (45%-61%) even  b a s i s of  such a model,  ( t a b l e 16)  discrimination years  Any  accuracy  concerning  three  sex.  sex  were c o r r e c t l y  generally  classification  stock  correctly  Kispiox  other  were  for  For  five  this  by  lower c l a s s i f i c a t i o n  the  the  shown  Larkin,  the  of  for  analysis.  points  consistently  chosen  age;  accuracy  Firstly,  on  constructed  steelhead  and  between basis  54%-Morice-Bulkley  have been  (Parker  stocks  at  were  Lachenbruch's  origin  the  size  steelhead  the  five  models  Skeena R i v e r  any  on  the  in  discriminant  Kispiox  steelhead  in freshwater  stocks  differentiating  using  female  made t o d i s t i n g u i s h  steelhead  River  sex  the that  stage.  between  the  classification  stock  centroids  Finally,  stock  76  discriminance ocean  was  (Sustut,  presence  of  greatest  i n those  Babine,  Morice-Bulkley)  specific  growth towards the Not the  upper  surprisingly, analyses  River  a  models.  support  the  In  results  of  incidentally  fishery  classification  of  stock  suggests  the  factors for  River  number of  the  the  selection  Skeena  i n the  general,  hypothesis  data  F i s h e r y Stock  drainage.  stocks  success  results  used for  in  all  of d i s c r i m i n a n t  discreteness in  Skeena  steelhead  of o r i g i n  was  the  s t e e l h e a d c a t c h by  Fisheries week f o r t h e  Incidental  catches  record.  shown, peak c a t c h e s  ending  J u l y 21  fishing.  in  although  Figure  compared (Table  18)  and  Only  in  females  Oceans  1984  commercial in  one  salmon year  the  the  were t h e  with  peak  of  over  the  and  4.2+  were  other  age  classes  were  m a l e s was  w h i c h were p r e d o m i n a n t l y and  weights  of  highest  on  (Weeks salmon  the  sample  s i x week  3.2+  4  fishery.  sockeye  of age  .1+  area  peak e f f o r t  t a b l e 7 summarize  collected  o c e a n age  mean l e n g t h s  salmon  of  Table  statistical  1984  occurred  appendix  steelhead  abundance o f  The  commercial  and  during  Steelhead  shifts  to  the  feasible.  steelhead  28) 23  of  1984.  proportional  of  and  composition  9-14  in  classification  (1984) were a v a i l a b l e f o r c l a s s i f i c a t i o n .  summarizes  As  Composition  scale analysis indicated that  caught  to stock  commercial  age  the  from  steelhead.  The  17  of  reduction  Commercial  of  regions  farthest which  regimes/and or  resulted in greater  discriminant analysis  growth  stocks  period  predominant found.  found  in  of o c e a n age steelhead  in  A 1984, .2+ the  77  Figure  23. Age c o m p o s i t i o n s t r u c t u r e by week f o r s t e e l h e a d s a m p l e d i n t h e 1984 c o m m e r c i a l f i s h e r y . ( c o m p i l e d from appendix t a b l e 7).  WEEK 12 1984 : N - 1 2 7  WEEK 9 1984 :n=123  3 O  %y  - -iX*. ^ 1 v i * '  ^*  k  0.2  a- * a^* y * y l * y**' %  »>*  WEEK 13 1984 :N=130  • •111  v  »i*  .III 11  T,.\* %* y\* a * y-s* %  v  I  ACE CLASS  i  .a* >* tf> k  WEEK 14 1984 :N=108  WEEK 11 1984 :N=134  *>*  t  AGC CLASS  ACE CLASS  a>* a** i>* y i *  ^  AGE CLASS  WEEK 10 1984 :N=130  I a . ^ ' a V * v a * y-i* ^  k.l* k V  %  ACE CLASS  ^  1i1 I 1.1  aN*V* b V ^ " b>* <- * ^n* AGE . a "CLASS vb* k%" a  *'  u>"  &  79  TABLE  17. S t e e l h e a d c a t c h a n d escapement s t a t i s t i c s t h r o u g h A r e a 4 f o r t h e 1984 c o m m e r c i a l salmon f i s h e r y . S.W i n d i c a t e s t h e s t a t i s t i c a l week. ( G N = g i l l n e t , S N = s e i n e ) . S o u r c e D.F.O, 1985.  Week Ending  S.W  J u l 15 J u l 21  8 9  Jul  28  1 0  Aug Aug Aug Aug  4 1 1 18 25  1 1 1 2 1 3 1 4  Sep  1  1 5  Table  weekly  stock  Catch  Days f ished  CPUE  GN GN SN GN SN GN GN GN GN SN GN  687 7021 2340 4362 4881 4269 2854 431 9 367 105 167  1 4.3 (1.3) 3 (3) 3.3) 3 5 2 (1 ) 1  2.4 10.8 13.0 7.4 23.9 8.5 9.6 17.0 4.6 3.0 3.0  TOTAL  31 372  290 649 180 591 204 504 297 252 80 35 55  H.R  Escape.  22.9  3191 5484 5750  '. 1 25 .619  3324  .735  4838 1 0090 5433 3880  .468 .221 .443 .109  — 42989  0 x=.422  18. Ocean age c o m p o s i t i o n by s e x f o r s t e e l h e a d t a k e n i n t h e 1984 Skeena R i v e r c o m m e r c i a l salmon f i s h e r y ( a s s e m b l e d from a p p e n d i x t a b l e 7 ) .  Ocean age %M .1+ %F .2+ %M %F %M .3 + %F  19),  Gear  9 .278 . 1 25 .544 .708 . 1 52 . 1 67  samples  generally  which suggests specific  1 0 .221 . 1 40 .573 .640 .208 .220  of  1 2 .464 . 1 58 .375 .631 .161 .210  1 3 .285 .171 .457 .800 .257 .028  14 .222 . 106 .587 .723 .190 . 170  increased through the f i s h e r y  a general  run-timing  Classification  Week 1 1 .333 .282 .560 .696 . 1 06 .021  shift  in  size  brought  (Table  about  by  differences. the  1984  commercial  fishery  steelhead  80  TABLE  19. Mean l e n g t h s and w e i g h t s o f s t e e l h e a d sampled i n t h e 1984 c o m m e r c i a l salmon f i s h e r y . SW i n d i c a t e s s t a t i s t i c a l week  Week Ending J u l 21 J u l 28 Aug 4 Aug 1 1 Aug 18 Aug 25  samples  SW  n  stock  discriminant these  ocean  analysis  models  previously oultlined.  age/scale  the  alone  (Model  B),  alone  low  age  class  to  relative  overall  Table commercial  specific  analyses,  assumed  occur  above f o u r m o d e l s .  the  Morice  River  ocean  smolt  ocean  age/pooled  sample five  steelhead  samples by s p e c i f i c age feasible  sizes present stocks  samples  under  because of i n the data study  in proportion  of  classifying  samples t o stock  were  to t h e i r  proportions were  with  some s t o c k s  throughout estimated  the  the  of o r i g i n  Temporal d i f f e r e n c e s i n the p o i n t  steelhead  D).  i n the f i s h e r y .  a l l f o u r models were found large  In o r d e r ,  age 4 / p o o l e d  pooled  fishery  i n t h e weekly  abundance  fishery  in  twelve  ( M o d e l C ) , a n d model C u s i n g  20 summarizes t h e r e s u l t s  present  the  i n a d d i t i o n t o t h e s c a l e v a r i a b l e s (Model  of the commercial  For a l l  of  age 3 / p o o l e d  (Model A ) , s m o l t  (eg 3.2+, 4.2+ e t c ) was n o t c o n s i d e r e d  base.  for  four  o f a d u l t s by s m o l t  variables  data  Classification class  1 .3 1 .9 1 .9 1 .9 1 .9 1 .7  utilized  v a r i a b l e s alone  related  8.2 10.5 9.2 8.3 9.5 8.2  S  origin  variables  age/scale  Weight (kg) 3.7 4.4 4.4 4.4 4.8 4.7  S  of  were c l a s s i f i c a t i o n  age/scale  size  1 32 131 127 1 22 1 33 108  9 10 1 1 1 2 1 3 1 4  to  Length (cm) 69.4 72.2 72.3 72.3 74.3 73.4  1984  by t h e  estimates  estimated  t o be  sample  period.  t o be p r e s e n t  in large  Table  20. C l a s s i f i c a t i o n r e s u l t s t o s t o c k o f o r i g i n by week f o r s t e e l h e a d sampled from A r e a 4 i n 1984. Model A: s m o l t age 3 / p o o l e d ocean a g e / s c a l e v a r i a b l e s a l o n e . M o d e l B: s m o l t age 4 / p o o l e d ocean a g e / s c a l e v a r i a b l e s alone. M o d e l C: p o o l e d s m o l t a g e / p o o l e d ocean age/scale variables alone. Model D: p o o l e d smolt a g e / p o o l e d o c e a n a g e / s c a l e v a r i a b l e s + l e n g t h and w e i g h t (+/- 95% c o n f i d e n c e l i m i t s about t h e e s t i m a t e d v a r i a n c e s in b r a c k e t s ) .  Week  P r o p o r t i o n a l Estimated Kispiox Zymoetz Sustut  Stock Composition Babine Morice  9 A B C D  065( . 181 ) 0 ( .253) 1 49(. 430) . 1 82 ( .440) 1 06(. 1 50) .038( .251 ) 0 ( .027) 0 ( .065)  A B C D  0 ( 255( 091 ( 0 (  A B C D  1 05(. 181 ) 0 ( .262) 0 ( .775) . 525(.798) 1 49(. 200) .087( .141) 052( .181) .032( .086)  .454( .298( .439( .303(  1 0  1 1  12 A B C D  .091 ) .057( .251 ) .701 ( .091 ) . 424) 0 ( .368) • 395(.121) .161) .068( . 1 73) • 496(.094) .056) 0 ( .072) . 4 1 2 .065) (  0 ( . 1 92)  1 93< .665)  1 38 (.092) 0 ( .072)  0 ( .462) .3221 .711) • 089(.313) .262( .086)  13 A B C D 14 A B C D  •  1 44 (. 181 ) 0 ( .274) 1 72 (.660) .241 ( .634) 1 72!. 101 ) • 045(.113) 144 1.065) .071 ( .086) 0 0 0 0  ( < ( (  .091 ) .097) .087) .065)  . 1 66) . 1 0 1 .214) < .590) . 2 1 6 ( .634) .101) .083( .094) .046) . 1 20 .086) (  .0261 .092) . 1 17(.086) .061 ( .042) .099( .032)  .454( .200 .254( .232 . 356(.116 .599( .072  .081 ( .094) . 1 1 6(.086) .0991 -.07 5) . 168(.056)  . 1 61 .( 1 73 .232( .112 . 244 ( . 1 04 .4 1 9(.072  • 376(.092) .223( .108) .325( .112) 0 < .030) .425( .091 ) . 1 42 .075) ( .378( .065) .093( .056)  .294( . 1 66 . 1 50(. 1 34 . 1 96(. 1 00 .443( .077  .428( .073) .307( .149) .264( .515 .290( . 1 26) . 1 93(. 1 34) 0 ( .112 • 422(.093) . 1 87 .(100) . 1 63 .094 < .336( .072) .287( .072) . 1 1 5(.072 .490( .448( . 443 .366( .681 ( .486( .639( .629(  .092) .120) ( .075) .065)  . 1 63(.079) .202( . 175 0 ( . 1 03) . 1 37 . (1 42 .2031 .095) . 1 35(.072 .21 4( .065) .204( .056  .092) 0 ( .079) .117) .054( .103) .081 ) . 1 35(.079) .072) .083< .046)  . 2 1 8 .(1 75 .243( .141 . 1 67(.101 . 1 29(.065  82  proportions  during  the  e a r l y weeks 9 t h r o u g h  steelhead  were  estimated  during  later  weeks  the  Zymoetz R i v e r proportions  steelhead during  model.  In  stocks  were  several  negative.  may  have been p r e s e n t  estimate  was  limits  were to  greater  in large Kispiox  t o be  present  point  River  estimates  Kispiox  Zymotez  stocks  v a r i e d about  and for  all  four  models,  each  success  Confidence  in  point  of  the  Morice-Bulkley  River  Zymoetz  Kispiox  and  these about  wide  analyses.  the  intervals  and  classification  upon  for  For  and  and  in variable  13 d e p e n d i n g  confidence  River  proportions  numbers.  the  than  Babine  Both  through  both the  f o r Babine, Sustut,  notably  11  However, the  discriminant  estimates  14.  the  in small  in proportion  original  week  indicated that  confidence  present  were e s t i m a t e d  instances  estimates  be  through  each  the  the  12  to  10.  the point  steelhead River  steelhead. The  point  estimates  probable  run-timing  fishery  in  steelhead used  1984.  catch  to generate  between  model  in table  curves The  of  each  20  steelhead  d i v e r s e age  suggested specific  were u s e d  t o c a l c u l a t e the  stock  through  c l a s s s t r u c t u r e of  that a l l four c l a s s i f i c a t i o n run-timing  comparison  and  a  curves.  This  more  detailed  the  the  1984  models  allowed  be for  run-timing  analysis. Calculating from  external  population by  adding  catch  the  1984  sources.  size during Department  estimates  to the  run-timing The  curves  calculation  e a c h week of  the  of  fishery  of F i s h e r i e s and weekly  first  estimated  required total was  Oceans w e e k l y steelhead  data  steelhead calculated steelhead escapement  83  past  the  test  fishery  10662  15111  12567 9107  timing curves  were  weekly  estimates  point  20) by t h e e s t i m a t e d  classification  of  calculated  by  total  weekly  steelhead 53.5%  models  (A  samples.  assumed  to  be  167 74361  the  classification  steelhead  population  the  two  and B ) ; h e r e ,  smolt  steelhead  comprised  age  class  of a d u l t s of smolt  of  smolt  for a  age  3  of  total  compositions 53.5% o f t h e  age  + escapement  given  specific  the estimates  t h e age  harvest  estimated  model ( t a b l e  size  I n week 9, f o r example,  sample was c o m p r i s e d  of the t o t a l  15 T o t a l 167 31372 198.9 42989  multiplying  for a given  r u n s i z e were s e t t o r e f l e c t  the  The g e n e r a l r u n -  12944 9752 4352  A d j u s t m e n t s were made f o r  weekly  table).  S t a t i s t i c a l Week 8 9 1011 12 13 14 687 9361 9243 4269 2854 4319 472 44.7 26.6 15.4 22.4 46.7 25.1 18.0 5484 5750 3324 4838 10090 5433 3880  Harvest Index Escapement Run S i z e  week.  (summary, n e x t  3.  Thus  i n week 9 was  adults.  Similar  a d j u s t m e n t s were made by week f o r e a c h m o d e l . Appendix predicted  weekly  classification +  table  escapement  8  stock  summarizes  the r e s u l t s  composition  estimates  from  m o d e l s ( t a b l e 20) t o t h e e s t i m a t e d run s i z e s  i n t h e 1984 c o m m e r c i a l  24 t o 31 summarize t h e e s t i m a t e d  run-timings  of a p p l y i n g the the  weekly  fishery.  four  harvest Figures  from appendix  table  8. In g e n e r a l , M o r i c e  River  and S u s t u t  River  were  found  to  84  predominate stocks  i n the  tended  run-timing  e a r l y weeks of  to predominate d u r i n g model  is  limitations  commercial  and  data  for steelhead  steelhead study  r e m a i n unknown.  The  specific  likely  within  stock  sample  sizes.  i n c r e a s e the  advantage  model  with  the  advantages  of  utilizing inclusion  (C)  plus  specific  s i z e s a t age.  fishery  selects  appreciably  The  for  morphological various  size  specific. River, a  and  similar  analysis seven  The  of  an  results  Morice-Bulkley phenomenon  of v a r i a n c e the  ten  of  may  or  if  models  adaptive  the  data  data  (D)  sizes  this  ( A and  but  may  B)  suffer  analysis  base.  but The  has  (C) has same  the  same  utilizing  stock  however,  a t age  if  the  should  change  evidence  that  stock.  provided  juvenile atlantic basis  comparing  and  is  Zymoetz  salmon  highly River,  juvenile steelhead parr  i n Skeena R i v e r significant  in  Analysis  between  River  "other"  age  age  of  unique  of  considered  suffer,  (1981)  of  a  not  added b e n e f i t of  for a given  variations  year  was  effect  smolt  size  Model D  Leggett  s t o c k s have  smolt  "best"  especially  one  1984  three  The  scale pattern variances  of  Juvenile and  only that  samples  much  the  between y e a r s  Riddell  by  pooled  other  identify,  scale pattern variances  w i t h i n stock  of  the  weeks.  In a d d i t i o n , t h e weekly  from reduced  to  acknowledgement  the  the  while  later  imposed  in  reduce  the  the  returns.  stocks  the  difficult  c o n s i d e r i n g . the  year  sampling  steelhead.  stock Kispiox  suggests  In a u n i v a r i a t e  d i f f e r e n c e s between  juvenile morphological  from  means  for  body measurements u s e d  85  Figure  24. Predicted run-timing. Estimated run-timing c o m p o s i t i o n t h r o u g h the 1984 Skeena R i v e r c o m m e r c i a l salmon f i s h e r y f o r a d u l t s t e e l h e a d of s m o l t age 3 / p o o l e d ocean age u s i n g s c a l e f e a t u r e s a l o n e .  Figure  25. Predicted run-timing. Normalized estimated runt i m i n g c o m p o s i t i o n t h r o u g h t h e 1984 S k e e n a R i v e r c o m m e r c i a l salmon f i s h e r y f o r a d u l t s t e e l h e a d of s m o l t age 3 / p o o l e d ocean age u s i n g s c a l e f e a t u r e s a l o n e .  Week E n d i n g  UV84)  87  Figure  26. Predicted run-timing. Estimated run-timing c o m p o s i t i o n t h r o u g h the 1984 Skeena R i v e r c o m m e r c i a l salmon f i s h e r y f o r a d u l t s t e e l h e a d of s m o l t age 4 / p o o l e d ocean age u s i n g s c a l e f e a t u r e s a l o n e .  Figure  27. Predicted run-timing. Normalized estimated runt i m i n g c o m p o s i t i o n t h r o u g h t h e 1984 S k e e n a R i v e r c o m m e r c i a l salmon f i s h e r y f o r a d u l t s t e e l h e a d of s m o l t age 4 / p o o l e d ocean age u s i n g s c a l e f e a t u r e s a l o n e .  88  XI00 30Key  l=Sustut 2 Mori ce 3=L<abine 4«Ki s p i ox 5=Zymoetz K  N 23-  u  m  b e r 20s  10-  I B  123456!123436!123436112345611234361123456!1234361 I I I I I I 7 10 11 1 13 14 IS J u l y 21 A u g u s t ,4 A u g u s t 18 Week E n d i n g  (1VB4)  X 100  !1234561123436112343611234361123436112343611234361 ! I I I I I ! I B V 10 11 12 13 14 13 J u l y 21 August 4 A u g u s t 18 Week E n d i n g <1VB4>  89  Figure  28. Predicted run-timing. Estimated run-timing c o m p o s i t i o n t h r o u g h t h e 1984 Skeena R i v e r c o m m e r c i a l salmon f i s h e r y f o r a d u l t s t e e l h e a d of p o o l e d s m o l t a g e / p o o l e d ocean age u s i n g s c a l e f e a t u r e s a l o n e .  Figure  29. Predicted run-timing. Normalized estimated runt i m i n g c o m p o s i t i o n t h r o u g h t h e 1984 Skeena R i v e r c o m m e r c i a l salmon f i s h e r y f o r a d u l t s t e e l h e a d of p o o l e d s m o l t a g e / p o o l e d ocean age u s i n g s c a l e f e a t u r e s a l o n e .  Weols E n d i n g  (19131)  91  Figure  30. Predicted run-timing. Estimated run-timing c o m p o s i t i o n t h r o u g h the 1984 Skeena R i v e r c o m m e r c i a l salmon f i s h e r y f o r a d u l t s t e e l h e a d of p o o l e d s m o l t a g e / p o o l e d o c e a n age u s i n g s c a l e f e a t u r e s i n c o n j u n c t i o n w i t h l e n g t h and w e i g h t .  Figure  31. Predicted run-timing. Normalized estimated runt i m i n g c o m p o s i t i o n t h r o u g h the 1984 Skeena R i v e r c o m m e r c i a l salmon f i s h e r y f o r a d u l t s t e e l h e a d of p o o l e d s m o l t a g e / p o o l e d o c e a n age u s i n g s c a l e f e a t u r e s i n c o n j u n c t i o n w i t h l e n g t h and w e i g h t .  Weel; E n d i n g  <19B4)  Week E n d i n g  U9B4)  93  in and  this  study  caudal  Table  were  found,  peduncle  especially  depth  f o r caudal  21. A d j u s t e d g e o m e t r i c means (+/- one S.D) and t h e r e s u l t s o f one way a n a l y s e s o f v a r i a n c e f o r d i f f e r e n c e s i n body m o r p h o l o g y between K i s p i o x , Zymoetz, a n d M o r i c e B u l k l e y R i v e r s t e e l h e a d p a r r . A l l measurements a r e i n cm. (* i n d i c a t e s no s i g n i f i c a n t d i f f e r e n c e a t t h e 5% l e v e l of s i g n i f icance)  The analysis  results  F=8.13 DF= between  three  selected  10 162  as  differences;  best  stock  in  Four  the  original  of  describing were,  caudal  caudal  function  (approximate  a l l pairwise ten  between  comparisons  variables stock  width, primarily  and  depth.  were  juvenile  of e n t r y , h e a d w i d t h ,  caudal  body  width.  The  separated  Kispiox  River  s t o c k s on t h e b a s i s o f c a u d a l  peduncle  f o r these  the  i n order  peduncle  from the o t h e r  mean v a l u e s  discriminant  Multivariate a n a l y s i s revealed  and  discriminant  and  32.  morphological  P<0.001)  these  depth,  juveniles  (DF=2,88) Kispiox F P n = 30 : 3.06 (0. 13) 0. 08 *>0. 05 2.11 (0. 12) 3. 1 7 <0. 05 1 .59 (0. 12) 6. 1 2 <0. 05 1.17 (0. 06) 12. 98 <0. 05 0.56 (0. 07) 14. 98 <0. 05 7. 41 2.91 (0. 22) <0. 05 1 .55 (0. 14) 9. 29 <0. 05 5.92 (0. 14) 0. 42 *>0. 05 6.14 (0. 16) 0. 40 *>0. 05  d i f f e r e n c e s between t h e s t o c k c e n t r o i d s  stocks.  peduncle  of  a r e shown i n f i g u r e  significant  width  width  (Table 21).  g r a n d mean s t a n d a r d l e n g t h = 10.29cm Variable Zymoetz Morice n = 29 n= 29 HL 3.07 (0. 13) 3. 00 (0. 21 ) HD 2. 01 (0. 24) 2.01 (0. 88) HW 1 .49 (0. 29) 1.41 (0. 06) CD 1 .1 1 (0. 07) 1 .04 (0. 04) CW 0. 48 (0. 07) 0.43 (0. 04) BD 2. 84 (0. 18) 2.70 (0. 13) BW 1 .55 (0. 17) 1 .40 (0. 07) PrDL 5. 91 (0. 20) 5.95 (0. 18) PoDL 6. 1 4 (0. 13) 6.10 (0. 19)  first  peduncle  Kispiox  peduncle  j u v e n i l e s had l a r g e  f e a t u r e s and were g e n e r a l l y more  "robust"  94  Figure  32. Discriminant function analysis describing morphological v a r i a t i o n among j u v e n i l e s t e e l h e a d from the Kispiox,Zymoetz,and M o r i c e - B u l k l e y R i v e r s . Each of t h e l e t t e r s i n d i c a t e s t h e s t o c k c e n t r o i d s Z=Zymoetz K = K i s p i o x M=Morice *=grand c e n t r o i d , t h e open c i r c l e s i n d i c a t e t h e 90% c o n f i d e n c e i n t e r v a l a b o u t e a c h c e n t r o i d ( f r o m P i m e n t a l , 1979), a n d t h e l i n e s p o i n t t o t h e n e x t most s i m i l a r s t o c k i n d i s c r i m i n a n t s p a c e . The two s t a n d a r d i z e d d i s c r i m i n a n t f u n c t i o n s a r e g i v e n below. D1 = 1.60HW +8.28CD +8.80CW -1.23BW -8.28 D2 = 0.03HW +0.84CD +1.47CW -1.68BW  -14.25  + 1.5-  +1. 0 -  -1.5-!  -2.0 -2.0  -1.5  -1.0  -0.5  0  FUNCTION  +0.5 1  +1.0  +1.5  96  in  body  shape  function stocks had  separated  mean  overall  body  stocks  were  given  Zymoetz  61.4%  correctly (1975)  48.3%-  a result  scale  j u v e n i l e s from  o f body w i d t h .  Actual  of  holdout  were  the  classified  Kispiox  evenly  features:  from  to  of  n = 30 n = 29 n = 29  divided  These  results  different.  form a n a l y s i s t o t h e a d u l t s  the  adults  Lough,  from  pers.  information  form  While  from e a c h  stock  have been n o t e d  several  Skeena  River  Comm.  1985)  and  f o r stock  (range  other  two  classification that the  Stock Z 6 6 1 4  separation  for  Skeena  extensions  s i m i l a r shape d i f f e r e n c e s c o u l d proportions  using  61.4  are  body  origin  the  M 3 19 8  steelhead  adult  three  suggest  Predicted  d i f f e r e n c e s i n j u v e n i l e body quite  the  procedure  between  K 21 4 7  70.0 65.5 48.3  juveniles  M i s c l a s s i f i c a t i o n s f o r Zymoetz  observable  that  stock  two  l e s s "robust" i n  juveniles  % correct  Stock  x=  likely  Morice  classification  70.0%).  discriminant the other  s i m i l a r t o the f i n d i n g s of a d u l t  pattern  K M Z  The s e c o n d  body w i d t h s , and were g e n e r a l l y  juveniles  stocks,  length.  the Morice River  shape.  Lachenbruch's  by  a  p r i m a r i l y on t h e b a s i s  low  River  at  be f o u n d .  body  provide  is  Observed  widely  steelhead  purposes.  of such  were n o t made, i t  to vary  may  River  between  stocks  (M.  additional  97  DISCUSSION  Biological The  Considerations  primary  separability analysis.  of  Skeena  Significant  composition, between  o b j e c t i v e of t h i s  sizes  tributaries  from  exist  for  of  the  Kispiox, each  the  major  stock  are  commercial  variability  confirms  t h e s u b d i v i s i o n o f Skeena  d i s c r e t e s t o c k s and suggests  Skeena  Zymoetz, B a b i n e ,  from  property  pattern  a g e , and j u v e n i l e body m o r p h o l o g y  catches  adaptive  scale  scale  incidental  into  racial  age  five  in  by  the  growth,  (Morice-Bulkley,  Run-timing d i f f e r e n c e s  was t o t e s t  steelhead  differences  at  steelhead  River  study  that  also  salmon  stock  River Sustut).  evident  fishery.  River  in This  steelhead  d i s c r e t e n e s s i s an  of t h e s p e c i e s t h a t has a r i s e n through n a t u r a l  selection. The River  scale  pattern  steelhead  technique  works  better  than  the  d e p e n d s upon t h e o b s e r v e d  compared stock  t o between s t o c k  definition  diverse  construction specific  used  models,  levels  which  studies,  study  are is  most  chosen  for  was n o t h i g h  for  commonly  the  stock  analysis.  The  s t e e l h e a d makes t h e  quite restricted  success  of w i t h i n  of  i n t u r n , depends upon  s t r u c t u r e o f Skeena R i v e r  mean c l a s s i f i c a t i o n in this  This,  Babine,  The s u c c e s s  o f d i s c r i m i n a t i o n models d i f f i c u l t .  discrimination The  variance.  Skeena  (Sustut,  ( K i s p i o x , Zymotez).  and t h e v a r i a b l e s  age c l a s s  differentiating  f o r some s t o c k s  Morice-Bulkley) technique  others  for  The u s e o f age  used  in  for this  classification  ( 5 0 % t o 65%: r a n g e  stock  species. models  Zymoetz 29%-  98  50%-  Sustut  55%-75%)  allocation  (20%)  stocks  involved.  (5)  and  pattern  "similarity"  T h i s may  reflect  dominant freeze  occupy  (=  also  of the  flows,  upper  occupy  sizes ages  the  age,  and  River  steelhead,  River  drainage,  a t age,  and  age),  maturity.  the  are  notably  wide  (yearly  sizes  region,  the by  at  are  smolt  by  of  older  s c a l e zones  steelhead  (=  by  s c a l e zones  (=  of  older  occupy  the  sizes  at  Skeena  smolting,  small  small  adult  Kispiox  River  Skeena  River  at  intermediate  and  which  the  the  ages  and  typified  s i z e s at age),  areas  zones  Morice-Bulkley  "inland" regions at  are  age,  steelhead,  the  which  scale  to large adult  o l d e r ages  scale  drainage,  l a r g e freshwater  mid-river  River  several  stock  freshwater  Babine River  l a r g e r s i z e s a t age,  Zymotez  to  steelhead,  ocean a g e s a t m a t u r i t y .  typified  freshwater  steelhead.  determines  River  adult  River  small  younger  occupy  intermediate at  and  large  and  ocean ages at m a t u r i t y .  (=  of  scale  drainage  River  Skeena  intermediate  typified  s c a l e zones  stocks  steelhead  smolting,  which occupy are  all  Sustut  smolting,  s i z e s at age),  freshwater  Skeena R i v e r  the  Skeena  at  l a r g e number  c o n t r i b u t e s to w i t h i n  maturity.  intermediate  freshwater  drainage  at  ages  smolt  steelhead  of  at age),  upper  large  the  separation.  o l d e r ages a t  ocean  by  i n Skeena R i v e r  regions  of  random  temperatures e t c ) .  variation  stock  by  level  than  between a l l Skeena R i v e r  low  intermediate  sizes  exists  better  c o n s i d e r i n g the  certain  peak  l a r g e smolt  older  A  f e a t u r e s of  variability  typified  acceptable  a common r e s p o n s e  Environmental  success  substantially  abiotic  up,  pattern  but  smolting, smolt  ocean lower  sizes  ages regions  at of  99  the  Skeena R i v e r  older  ages  at  intermediate age,  and  three  of  sizes  between  of  Stock  stock  between  the  populations  He  not  of  that  to evolve  observable  suggest  that  from  display Kispiox  more f u s i f o r m  significant  differences  in  discriminant  individual  dispersion).  is  between variance Sustut,  between  streams  in  each  to l o c a l  stocks  the  the  not  stock  are basis.  observed  steelhead.  each  a  environments".  c e n t r o i d s but  B a b i n e , and  into  selective  for  depends  about  British  having  between  River  stock  steelhead  i s subdivided  case  analysis  fishes  Parkinson  have a s t r o n g  Skeena  in  1972,).  species  the  Various  adaptation  populations  may  1981).  in  phenotypic  exists  adaptations  level  processes  homing  Ricker  adjacent  "this  this  variation by  of  and  differences  many  Evans,  specific  1972,  semi-isolated  discreteness  (centroid  the  genetic  and  genetic v a r i a t i o n  adaptive,  of  site  geographically  a l l  results  level  at  variability.  and  for genotypic  (Larkin  that  potential  patterns  the  to l a r g e s i z e s  River.  (Maclean  that  concluded  number  necessarily My  (=  Juveniles  than  to  s c a l e zones  maturity.  ecological  potential  in  Columbia.  While  intermediate  more " r o b u s t "  environments  showed  genetic  freshwater  morphological  notably  to such environments  large  intermediate  ( K i s p i o x , Zymoetz, M o r i c e - B u l k l e y )  have s u g g e s t e d  (1984b)  at age),  by  d i s c r e t e n e s s w i t h i n a s p e c i e s depends upon t h e  "stochastic"  provides  typified  intermediate  the M o r i c e - B u l k l e y  interaction  authors  are  ocean ages a t  stocks  j u v e n i l e s are  juveniles  of  smolt  the  and  smolting,  intermediate  significant River  drainage  Stock  only  upon  a l s o upon centroid  Morice-Bulkley  River  100  exhibit they  greater  e x h i b i t lowered  Kispiox in  separability  and  levels  Zymoetz R i v e r  discrimination  this  study  dispersive presence Sustut  are  Babine  freshwater  s i z e s a t age  size.  The  of  some  Rivers, (=  large  would  force  Miramichi  older River  energetic l o n g and to  i n New of  difficult a  steelhead the  juveniles attributed  the  of  (1975)  genetic  swimming  j u v e n i l e s (Thompson R i v e r ) endurance  (Chilliwack River) the  in  of  larger juvenile  strong  been  selective  T h o r p e and M i t c h e l l that  the  salmon  Tsuyuki  to  larger from  meet  swimming  seems  and  "downstream" type of  the  during  Williscroft  Fraser  significantly  levels  the  flows  "upstream"  treadmill  d i f f e r e n c e s to greater  larger  and  in greater  t o be  the  s e l e c t i o n has  Sustained  component.  and  and  adaptations  swimming e n d u r a n c e of  swimming  adult  Atlantic  Brunswick are  from  exhibit large  turbulent  concluded  upriver  sustained  the  selection for larger  1975,  upriver migration.  strong  (1977) f o u n d  than  ages  costs  have  Elson  represent  could  a potentially Elson  used the  s i z e s a t age)  is  levels  then  Steelhead  both  as  stock,  could  Hydrodynamic  ( S c h a f f e r and  S c h a f f e r and and  each  example,  region  several authors  in salmonids  1981). sizes  by  separability  l e a r n i n g samples  forces.  enhance  Conversely,  lower  b e c a u s e of h y d r o d y n a m i c  upstream/downstream m i g r a t i o n . suggested  the  of  smolt  beacuse  exhibit increased  stocks  for  u p p e r Skeena R i v e r  size  that  selective  s c a l e zones  adult  body  exhibit  representative  dominant  and  steelhead  Assuming  homogeneity  of  of c e n t r o i d d i s p e r s i o n .  models because they  of c e n t r o i d d i s p e r s i o n . in  i n d i s c r i m i n a t i o n models  the  River greater  Fraser  River  tests.  They  LDH-A  allele  101  in  the  Thompson  muscular  River  upstream  in  smolt to  findings  by s c a l e  (1977),  and  Whately  Kispiox,  and  respectively. same  three  food  study  of  found  that  exhibited  at  smaller the  upriver  enough t o p r o d u c e additionally  f o r age 4 s m o l t s  195mm,  and  and  203mm  Although  larger  fry,  adult  s c a l e zones river  by s c a l e sizes  (= s m a l l  stocks.  (1983),  Skeena R i v e r  l a r g e smolts a t age.  information  of s c a l e p a t t e r n  the  from  at  age y e t  smolt  sizes at  They a t t r i b u t e d t h e the  i ssufficiently  Babine  sockeye  in a  patterns,  T h i s c o n t r a s t s my f i n d i n g s a n d s u g g e s t s  benefit  187  factors control  1972).  salmon s t o c k s  had l a r g e r  Yukon  Different ' selective features  lengths  s c a l e zones t o lower p r o d u c t i v i t y i n  area.  little  163mm,  depend upon y e a r l y g r o w t h r a t e and  p r o d u c t i v i t y i n t h e upper  although  145mm,  178mm,  (Ricker,  back-  Morice-Bulkley,  McBride and M a r s h a l l  freshwater lower  the  my  Whatley  t h e mean  l a r g e r eggs a n d t h u s l a r g e r  chinook  stocks  (1969),  and environmental  smolting  River  upriver  River  were  were  genetic  availability.  Yukon  than  smaller  may  rivers  g e n e r a l l y produce  therefore  food  Rivers  distance  which c o n f i r m s  Narver  from  ability.  ascending  reported, that  smolts  s i z e s a t age i n s a l m o n i d s  sizes  Yukon  drainage,  f o r age 3  Both  eventual  age)  River  analysis.  t h e t h r e s h o l d of  swimming  with  The mean b a c k - c a l c u l a t e d  respectively.  parents  sustained  e t a l (1978),  Babine  increases  increase  Skeena  pattern  lengths  the  sizes  the  calculated  smolt  which  f a t i g u e and thus extends  Steelhead  from  stock  River  salmon  upper that high  steelhead  enhancement,  i s available. forces and l i f e  may  explain  the  history variation  observed i n Morice-  1 02  Bulkley the  River  largest  rather al.  A l t h o u g h the M o r i c e - B u l k l e y  of  Skeena R i v e r  uniform  the  over  five long,  low  (1978) a t t r i b u t e d t h e  Bulkley The  steelhead.  River  smaller  adult  Morice-Bulkley selection  and  steelhead  maturation  would  production)  in less productive  maturity  the  thus  increase  less  favorable  Bulkley  e n s u r e maximal  River  supports  the  the  The  steelhead  the  notion  distances Sustut  of  1984  steelhead  commercial  hypothesis It first  "rapid  is  instream p r o d u c t i v i t y .  suggests  that  may  Rapid  (and  the  parr  adult  to  smolt  Older  ages  c l a s s spawnings  poor p a r r  1984  of  better  in  in  fishery  this  must  quite  early,  which  makes  at  Morice-  stock;  r e l a t e d to the  steelhead  by  and  production  commercial  maturation"  River  of  a yearly basis  c l a s s spawnings.  of  maturity  strong e c o l o g i c a l  exist.  f r y seeding  risk  Morice-  long  travel.  were a l s o p r e d i c t e d t o p a s s t h r o u g h  fishery  possible  that  small  in Morice-Bulkley  genetic  (1981) documents  the  effect  from  decreasing  maturity  f o r many P a c i f i c  to  s e l e c t i o n f o r o l d e r and  size  o l d e r ages of  such  the a  tenable.  spawning  cumulative  et  y o u n g e r ages a t  i s probably  inland Morice-Bulkley  River  Whately  e a r l y p r e d i c t e d run-timing  through  however, e a r l y r u n - t i m i n g  distances.  t i m e between y e a r  biological  years.  are  e n v i r o n m e n t s on  t i m e between y e a r  would e x t e n d  i t s flows  t o low  for rapid adult maturation  minimizing  tributaries  s i z e s and  smolts  s i z e s a t age  River  is  gradient  small  steelhead  River  salmon  s i z e s a t age River  steelhead  commercial sizes stocks  and  at and  young a g e s a t represents  fishing. age  and  Ricker ages  a t t r i b u t e s the  t h u s more mature  a  at  trend  individuals in  103  commercial Bulkley as  fisheries.  River  shown by  passing 1971)  steelhead  the  from the  conclusive, effects  this  may  determining  rapids  River  Steelhead  from  similar.  range  (length  and  and  in  Zymoetz R i v e r s  overlap. the  This  two  l a r g e r and Stock  between t h e  Kispiox  patterns.  discriminating growth.  Buxton,  While  not  fishery  forces  in  spawning  of  two  only  show a  rivers  may  high  suggests  sytems are  that  somewhat  although  exhibit  separation  introduced,  stock,  two  stocks  However,  first  were n o t  first  differences reasons  This  the  models developed the  and  study.  first  i n h a b i t " c o a s t a l " type  weight) are  i n the  steelhead  commercial  at  time,  by  a  the  wider  discriminant  when a d u l t  s i z e s at  age  which, being s u b s t a n t i a l l y  implies either genetic  differences  ocean growth r a t e s and/or d i f f e r e n c e s i n ocean m i g r a t i o n  feeding  the  regimes  environments.  increases  in  Kispiox  is considerably  analysis  of  ecological  ages  scale pattern  Both stocks  of  greater  the  growth  Zymoetz R i v e r  that  than  over  (Harding  in this  Morice-  steelhead.  freshwater  environmental  frequencies  used  and  of  rather constant  suggests  age  composition  1961-1967  important  s i z e s at  Morice-Bulkley  d e g r e e of  length  1976-1977 d a t a  less  size  remained  from  evidence  be  the  has  homogenous  Moricetown  and  However, t h e  for  ocean for the  that  on  the  year  study  leads  b a s i s of  scale  even  for  (3.2+, 4.2+).  was  not  examined  separating  the  two  similarity  to the  p o t e n t i a l for pattern  ocean  o c e a n g r o w t h d i f f e r e n c e s between  pronounced  in this year  naturally  and  of  the Scale and  stocks.  freshwater  age  specific  growth  could No  after  lead  to  definitive  scale patterns  in  104  Kispiox  River  Kispiox lakes,  and  River,  being  b o g s , and  benches  which  (Whately,  Zymoetz R i v e r  creeks  steelhead  The  except from  the  moderate Zymoetz  on  a  the  environmental  The  of  results  Kispiox  River  of  heads,  Zymoetz  river  has  been  flows,  important general,  those  migration the  have et  a  al.,  ratios,  j u v e n i l e salmonid streams with  routes  may  select  (Taylor,  concentration River  is  apparently  of  heaviest because  and  of  further  comment.  e x h i b i t i n g deep In  contrast,  peduncles.  especially and  biology  juveniles,  may  be  Stream h a b i t a t etc)  is  velocity  highly  (substrate, extremely  (Northcote,  flow  the limited  maximize  Relating  steelhead in  supports  1969). and  In  longer  f o r a more f u s i f o r m body shape i n  1984).  older  the  system.  caudal  cover,  higher  to  that  a b r o a d c r o s s - s e c t i o n of  basis  1981).  noted  "fusiform", exhibiting  "thinner"  genetic  similar  close  peduncles.  in salmonids,  j u v e n i l e s to reduce drag  ability  and  be  for this  quite  and  water q u a l i t y  which thus  bear  many  hills  somewhat  "robust",  demonstrate  poolrriffle  for  quite  bodies,  low  high  a  The  i s f e d by  proximally  " t h i c k " caudal  Body s h a p e  obvious.  It should  analysis  are  juveniles  (Riddell  space,  are  j u v e n i l e s are  slender  shown t o  adaptive  and  River  b o t h body t y p e s .  has  heterogeneity  juvenile  h e a d s , deep b o d i e s ,  smaller  and  c e n t r o i d for a l l stocks  juveniles  Morice-Bulkley  flows  River  Zymoetz R i v e r  grand  notion  s e r i e s of  larger scale.  multivariate  seem  i n i t s headwaters,  s i t u a t e d in a  provide  1977).  morphology  glacial  steelhead  parr  lower  sustained  to in  the  reaches  upstream  this  swimming  study,  the  Morice-Bulkley  (Tredger,  productive  1984),  capacity.  105  Here,  the  parr  microhabitat throughout typical  Zymoetz  the  of  differences  data  and  the  be  same age  attributable growth. River  of  of  additional  rear  exhibit  the  be  hydrodynamic  less  types  important.  which  habitats.  adults  a s s u m p t i o n s of for  by  the  based the  supports  Body  form  from e a c h Skeena  River  information  for  stock  author  study.  established  diverse  precludes f o r the  I would q u e s t i o n  any  limited  was  in age  simple  utility  this  study. by  same b r o o d y e a r  would age  Any  the  Ideally, discrimination  from t h e  This  data  a l l samples  the  data  are  methods.  in  d o m i n a n t age the  Firstly,  classification  fish  differences  However, t h e  steelhead  of  stock.  analysis  i s homogenous a c r o s s  and  using  from e a c h to  present on  representativeness  discriminant  l e a r n i n g s c a l e samples.  achieved  approach except then,  the  representativeness  availability  the  to  discrimination  Secondly,  may  which  where  i n t e r p r e t a t i o n , assumed  was  misinterpretation  should  i n a wide r a n g e  provide  concern  for  body t y p e  less  parr  juveniles  swimming a b i l t y  extend  in  interpretation  used  1984)  River  and  Considerations  data, of  River  velocities  purposes.  Errors  all  growth  could  Theoretical  the  to Kispiox  Kispiox  (Taylor,  flow  j u v e n i l e s e x h i b i t b o t h body  may  and  separation  of  drainage.  River  to higher  compared  for sustained  notion  stock  subject  "refuges"  "coastal"  selection  the  are  limit  and  class age  any  and  of  variability  yearly differences in structure  of  Skeena  specific discrimination  classes of age  (3.2+, 4.2+).  specific  analyses  Even for  1 06  Skeena R i v e r found  in  steelhead.  this  different  study  total  the  potential  ages but  brood  year  B a s e d on  of  necessity  e f f e c t s of on  the  patterns  appear  have s i m i l a r p a t t e r n s against  The  to  of  the  it  is  stock.  Significant differences  samples  (and  thus brood y e a r s )  s t u d y were not  brood  years  for  evident.  clarification  regard  to d i f f e r e n t i a l  Thirdly,  the  it  Each  requirement structure  and  appears and  its  to  effect  Straying  between  which  maintain  group  should  quantify.  years  given  in scale  growth  several  of  supports  the  discrete  been  the  could  although  for  affect in  the This  substock  s u c c e s s was  i s assumed  identity  assumptions  definable.  discrimination stocks  with  growth.  developed and  in  samples.  especially  analysis  met,  used  of d i f f e r e n t  learning  v i o l a t i o n of  models  yearly  stocks  use  scale  function  be  the  specific  for a  between  i s needed,  that  on  investigated.  to  brood  e f f e c t s on  have  by  between  stock  should  growth  scale  classification  "stock"  the  freshwater  this point  is possible  argues  However,  scale harder  stock  that  This  density  This  of  appears  for linear discriminant  discrimination  study.  of  freshwater  for  constructing  Further  necessary  stable  from e a c h  models.  t h i s study are  growth  this  relatively  age  growth  steelhead  growth.  specific  differential  l i m i t e d evidence,  scale  scale  that  same s m o l t  age  r e s u l t s of  freshwater  indicate  freshwater of  of  t o be  not  minimal,  discrimination  purposes. The  assumption  discriminating violated  of  variables  because  used  tests  multivariate in for  the  normality  study  multivariate  could  for have  normality  the been were  1 07  unavailable. linear  discriminant  decision these  Multivariate  that  analysis  surfaces  surfaces  used  are  best  normality  because  to separate  actually  separate  normal,  the  "overlap"  classification comparisons  normal  stocks  to  (Pimental,  multivariate  basis  association  (ie.  for  well  (Pimental,  when  analysis the  Apparently,  frequency  that  distributions  the  Gilbert  is still  sample  sizes are large  In  other  words,  to  data are  valid  variance  structure  study.  groups  show t h e same  The  a  patterns  functions that  for classification equality  has  and of e q u a l  on  of  e f f e c t s of d i s p e r s i v e  (1969) n o t e s  type  Stock  the patterns  within  of d i s p e r s i v e  of  the  in this  should  stock).  hypothesis  can  i n reduced  multivariate  association  1979).  analysis  enough  and r e s u l t  axes and d i s c r i m i n a t i o n  the  powerful  group  estimate  i n e q u a l i t y of d i s p e r s i o n s  multivariate  each  univariate  tested  Variables  each  on c a n o n i c a l  discriminant  of  matrices describe  variable  inequality known  normal)  o f homogenous v a r i a n c e - c o v a r i a n c e  was n o t r i g o r o u s l y  linear  boundaries  1979).  variance-covariance  s p r e a d and  even  stock  does not g u a r a n t e e  assumption  specific  of  each  on  the  d i s t r i b u t i o n s a r e not  surface  relied  of  In l i n e a r a n a l y s i s ,  contours  I  nature  (multivariate  density  decision  success.  This  multivariate The  the  for  normality.  between  distribution  the  classification  ellipsoidal  I f the m u l t i v a r i a t e  randomly  of  groups.  linear  hyperspheres. then  i s e s p e c i a l l y important f o r  no  size  test  of  centroid  result  in  rejection  even  purposes  effect  on  II e r r o r s i f  (Pimental,  equality  linear  i s rejected.  real  I or type  i s not  1979).  by MANOVA i s when  slight  108  departures possible (eg  from  that  t h e use of  quadratic  assumptions  primarily  within  results.  variances  not  seem t o be t h e c a s e  study c o u l d  majority  of  1978), w h i c h make no distributions  quadratic  i n the a n a l y s i s .  separate  to error.  using  (1982) n o t e techniques  "best".  fact,  although  In  variables  that  1979), l a r g e variables variable This  no g u a r a n t e e t h a t  magnify  second  selected i s  analysis  the  "the  linear  relies  associated  and  on  (Pimental,  combinations  problems  (Johnson  stepwise  discriminant  the subset  discriminant  procedures"  was n o t f u l l y  Commercial F i s h e r y The  constructing  i n t e r c o r r e l a t i o n s between  selection  aspect  the problems of using for  the  procedures.  show some d e g r e e o f i n t e r c o r r e l a t i o n  will  with  large variable  and  is  did  the stocks i n  Johnson  There  This  I n common  analysis studies  functions.  is  d i f f e r e n c e s between  I chose t o use s t e p w i s e v a r i a b l e s e l e c t i o n  selection  have  analysis  systems,  variable  or  study.  a l s o be s u b j e c t  Wichern  analyses  and between g r o u p s , c o u l d  of which v a r i a b l e s best  discriminant  It i s  discriminant  are s i g n i f i c a n t  for this  apparent.  density  of t h e v a r i a b l e s used  choice  are  Lord,  However,  u s e f u l when t h e r e  The  and  underlying  the  this  Cook  relationships better  equality  non-parametric  analysis,  regarding  dispersive provided  dispersive  Wichern,  of with  1982).  investigated.  Considerations objective  potential  of s c a l e p a t t e r n  steelhead  stocks  caught  of  this  study  was  analysis for identifying  i n the commercial  salmon  to assess the Skeena fishery.  River As  109  previously 1984  noted,  fishery  composition learning  within varying  samples,  distinct  Although  general,  analyses  of  and  Kispiox  River  the  Sustut  less  four-model  River  nature  evidence,  five  dominance  through  4/pooled  Here, both during  same  classification  o f Skeena  River  may  ocean  not  and  numerical  River  steelhead  later  The e x c e p t i o n  age/scale stocks  variable were  time a t r e t u r n f o r s t e e l h e a d  analysis.  was  not seen  Further  study  run-  Zymoetz, and  of  only  T h i s may  of d i f f e r e n t  reduced  i n the pooled i s required  was  predicted  t h e e a r l y p a r t s of the f i s h e r y .  trend  be  classification  of B a b i n e ,  Babine and K i s p i o x  differential  The  1984.  allocation.  stock  the f i s h e r y .  o f age  in  it  f o r catch  age  i n the  patterns  was e v i d e n t  ages or e r r o r i n t h e a n a l y s i s because  sizes.  sample  smolt  to  age  clarify  point. The  weekly  point estimates  of s t o c k  sufficiently  v a r i a b l e enough t o r e s u l t  fluctuation  for  normalized this.  specific  and M o r i c e - B u l k l e y  abundant  age  t o be p r o m i n e n t  Although  The same m o d e l s p r e d i c t e d t h e  steelhead  smolt  analysis.  smolt  the  data  i n the  a r e pronounced  fishery  b a s e d on l i m i t e d  the f i s h e r y .  timings  reflect  were s e p a r a b l e  f o r 1984 p r e d i c t e d t h e e a r l y r u n - t i m i n g  dominance through  stock  run-timing  t o use age c o m p o s i t i o n  In  this  no  stems from t h e c o m p o s i t e  possible  stocks  bounds o f c o n f i d e n c e .  through the commercial  steelhead.  for  major  d i f f e r e n c e s between t h e s t o c k s  composition This  a l l five  the  run-timing  However,  run-timing may o r may  based  on  the  abundance  in considerable  estimates. not long  i n 1984 a r e  be  temporal  The a s s u m p t i o n o f  practical  term p a t t e r n s  because  of  (normal) of  1 10  steelhead that  normalized  run-timing  All  classification  several (eg  r e t u r n and e s c a p e m e n t  four negative  Zymoetz,  intervals upper  point  point  values  these  were  the  sample  estimates  reflect  the  difficulty  learning  the  1984  pattern  fishery  surprising to  be  estimates  analysis  This  were  Sustut  need  Skeena R i v e r  several  included  those  stocks  present  an with  in  the  the  negative  by s c a l e p a t t e r n  a n a l y s i s when  i s low.  River  in this  suggests other  River in  region  "stocks"  a l s o have  This  sytem a r e  stocks  the  i s not  stock.  that  stock  with  not  believed population  scale  not  patterns  considered for  fishery  samples.  Steelhead  production  defined.  (Lakelse,  scale patterns may  of  i s somewhat  previous  but  well  M o d i f i c a t i o n o f t h e method  information  confidence  contribution  investigation.  downstream  Kitwanga) c o u l d system.  levels  either  present  possibilities upper  t o be t h e S u s t u t  the  usually  stocks  estimating  success  a r e i n e r r o r or that to  the  t h e 90%  Rather,  in  study.  a n a l y s i s p r e d i c t e d t h e l a r g e s t component  as p o p u l a t i o n  high.  similar  i n low a b u n d a n c e  believe  resulted in  of  actually  period.  sample c l a s s i f i c a t i o n  Scale  However,  not  during  f o r stocks  1984  f o r some  estimates  fishery  rates  for  I t seems u n l i k e l y t h a t  estimates  I  assumption f o r t h i s  analyses  t a b l e 20).  a s s o c i a t e d with  positive limit.  negative  is a valid  estimate  Kispiox,  t o t h e Skeena R i v e r ,  In  to  the  be n e c e s s a r y  i n the  addition,  Kitsumkalum, similar  Both  Suskwa, Sustut  as f u r t h e r  becomes a v a i l a b l e .  Although discriminator  size  (length  f o r Skeena R i v e r  and  weight)  steelhead,  is  a  good  stock  i t s use f o r c o m m e r c i a l  111  fishery  classification  selectivity  by  Skeena adult  size  age  stock  slight  Of  specific  between y e a r s  applicability Stock the  be  (good) stock  low  age  will  year be  then  of  in  eventual models  commercial  variability  in  the  been p r e v i o u s l y n o t e d salmon  McDonald  (Aro  1981).  and  Temporal  shifts  s p e c i e s appears  (Larkin  McDonald,  1968)  and  For  year  pattern technique  will  fluctuate  brood i f the  in  Assuming  of  each  fewer  fishery each  abundance and  the t e c h n i q u e  to  b e c a u s e of  that  that  scale  on  the  concern.  stock  r e t u r n s to a given  the  some  effects  abundance i s of  be  between y e a r s a c c o r d i n g t o  year  s u c c e s s e s , then  present  stock  to  although  Skeena R i v e r s t e e l h e a d t h e  s u c c e s s and  for  McDonald  for these  class;  i t s proportional  adequate,  has  (high) i n a given year  brood  by  (freshwater)  run-timing  r e t u r n s by  classification. to  brood  abundance  numerical  affected  to c l a s s i f y  until  pink  1968,  of t h e s c a l e  contributing to  and  i s present.  differential  i s not  classification  used  of  classification  independent  four  s h o u l d be  run-timing  and M c D o n a l d  variability  be  size  established.  specific  Larkin  All  for  features  to  Any  b i a s the estimates  itself  scale  appear  Skeena R i v e r s o c k e y e  1968,  as  interceptions  is clearly  will  analysis  study  with c a u t i o n .  samples used  size).  this  steelhead  Stock  in  thus  in  technique  both  steelhead  (and  developed  pattern  done  fishery  fishery  selectivity  River  fishery  i n the  Scale  potential  be  the commercial  s t o c k abundance purposes.  must  stock the  a  stock  series  (more) f i s h and  for  each happen  of  poor  from  that  available  for  i s sampled a c c o r d i n g sampling  patterns should  design  is  respond  to  11 2  such  fluctuations.  development, shifts  1  year  available timing) (same  is  1 i n year 2  of  cannot  stock  i n year  curve  different  aspect  i s i t s overall  variable  enough t o r e s u l t  the  estimates  between  This the  discriminance utilizing patterns  Fournier  comprises  fish  actual changing  50%  of  the  i n week  from t h a t  stock are  abundance,  same r u n -  shifted  earlier  or b o t h .  although  further  accuracy.  or  later  F o r t h e most investigation  fishery of  gene  These  pattern  To i n c r e a s e  the  frequencies  body  etc.  (1983)  chum salmon  for have  stocks  stock used  with  with  of  scale  morphology,  The i n c l u s i o n o f their  separation such  stock  possibility  in conjunction include  is  overlap  increased  however, once e s t a b l i s h e d , t h e y  information  a l .  success  c o n t r i b u t i o n (eg  scale  success,  multivariate features pursued."  stock  be m o d i f i e d .  pattern  l i m i t s ,for some o f  s y s t e m s must be w e i g h e d a g a i n s t  valuable  distinguishing  Discrimination  level  and c l a s s i f i c a t i o n  be  of the s c a l e  i n wide c o n f i d e n c e  the  of c o l l e c t i o n ;  et  the u t i l i t y  and c a n n o t  parasites,  such c h a r a c t e r  provide  stocks  should  difficulty  and/or  run-timing),  of commercial  reflects  other  meristics,  simply  less  has  affecting  technique  Zymotez).  curve  of  required.  Another  point  stage  between  2 ( different  I have assumed t h e f o r m e r  clearly  present  distinguish  which  Either  or the run-timing abundance,  A  the  1 may have a p r e d i c t e d abundance  20%.  f o r capture  at  run-timing  F o r example,  i n week  of  part,  the technique  i n the p r e d i c t e d  abundance. catch  However,  an  favorable  could  purposes.  approach results.  for  1 13  Gear  Selectivity Ricker  (1981)  incidental  notes  salmonid  species  salmon d e p e n d s upon t h e i r taken  incidentally  the  run  at  to  be  larger  This  results  caught  and  spawn. The  average  fish  time,  fishery  (3 kg)  (L.  Janz,  and  the  extreme  estuary.  Skeena  River  differences  between  the  to  an  etc.)  unknown steelhead  "tangled" addition, of any  caught  than g i l l e d , the  given  (depth  steelhead  run  at  commercial by  that  time. fish,  fishery  size  is  to  possible.  steelhead  for smaller  is  sizes  Generally,  four year  in  tend  between t h o s e  the  o l d male  sockeye  old  sockeye  salmon  by  size  may  be  in  the  fishing effort  on  may of  the  numerical  catch.  change  Skeena  size  Although their  fishery  of  size  sizes  of  the  behavioral  susceptability  swimmimng, p r o x i m i t y  of g i l l n e t t i n g  successfully  somewhat  (1977) f o u n d t h a t mesh  1984  regardless  dense n a t u r e  incidentally  i n the  year  size  1985).  Andrews  the  average  exists.  five  steelhead  in  than t h e i r  f o r Skeena R i v e r  doubt  of  stocks  extent  salmon  steelhead  effect  incidental  chinook  selection  female  levels  Oguss and  significant  no  on  example,  some s e l e c t i o n  Comm.,  effects  sockeye  the  for larger  larger  pers.  selective  no  although  on  for  size  response  selection  fisheries  differences  genetic  selects  kg)  r e d u c e d by  size  ages a t m a t u r i t y  (2-3  have  strong  of  salmon t a k e n  which escape  similar  salmon  River  their  i n net  smaller  than  to e s t a b l i s h  Any  often  mode  For  pink  of  younger  size.  time w h i l e  those  degree  gillnet  are  the  caught  in considerable  Over  difficult and  that  that  were  to  more  shore often  (interview data). may  migrating  reduce the past  the  In  chances fishery.  11 4  This  argues  commercial  against fishing.  esapement may  Applications A  any  specific  Reductions  be more  to Steelhead  harvesting  during  the  impacts  for identifying  fishery  and  provide  specific  management  extremely  reducing  Short the  incidental solved.  appear  to  extended  sockeye  is  most  t o a wier  days  per  than  fishery  to  that  while  the  pool  and  can  be  prominent  is  not  is  of  easily harvest  steelhead  fishing  Branch,  is  unpublished  weekly h a r v e s t  rate  i n a t h r e e day  per  i n a f o u r day  per  relates  area  daily  harvested  stock  problem  continuous (BCF  the  drastically  r a t e s on  this  i n c r e a s i n g weekly h a r v e s t  or  the  i t becomes l o w e r  fishery  in  fishery  i s adverse  for steelhead  Presumably,  results  socio-economic  fishery  harvest  week  My  the  system  for steelhead  if  incidental  structuring  fleet,  the  dramatically  three  s t e e l h e a d move i n t o  specific  present  complex  In a d d i t i o n , t h e mean p e r c e n t  less  problem of  by  in  concern  for  However,  Mean w e e k l y p e r c e n t  i s higher  tend  fishery.  commercial  catches  week o r more f i s h e r y .  salmon  salmon  potential  regulated  the  increase  1983).  or  of  principle  beyond  of  steelhead i s  specific  which stocks are  resorting  steelhead  The  the  is  of  size  pressured  week  stock  stock  objectives.  dynamic and  factors.  of  commercial  a method thus  effects  Management  provide  for  overall  m a j o r management o b j e c t i v e f o r Skeena R i v e r the p o t e n t i a l  data,  in  selective  important.  to minimize  rate  size  t o the whereas  fact  sockeye  quite quickly.  rate pressure  d u r i n g peak s o c k e y e salmon  on  that  The  steelhead  run-timing  where  11 5  fishing  may  Tuesday,  actually  Wednesday,  commercial harvests  on  timing,  my  variability  an  How  The  peak  i s not  c l o s u r e s or  ensure  that than  continuous of  interspersed  fishing This  study help  by  followed  assumes,  of  reduce  best  commercial  of  steelhead the  potential  future  to  feasible.  The  "windows" on  Any  catches  should  of  cohort  (two, use,  days  area  closure  that  latter  the  four  days  An  commercial  do  of more would fishery fishery  during  of  As  any an  fishing  may  (the present  the such  etc).  f o u r d a y s of  steelhead  quite quickly.  through  days  (windows) or  long  h o u r s a day  four  d a y s of c l o s u r e  course,  during  This  during  24  or  three  basis.  on  techniques  peaks escape the  occurs  three  than  Three  i s t o make use  removal  three,  of  focus  Logistically,  weekly  of  management  stop a l l f i s h i n g  run-timing  fishing  two  peaks.  second  a  estimates  analyses.  f o r each s t o c k .  steelhead by  depend upon t h e  is  opening  two  short  incidental  entire  fishery  to c l a r i f y One  the  i s somewhat c o m p r e s s e d w i t h i n a  first  window  to  through the  is  run-  run-timing  Presently,  beneficial  would  during  portions risk  openings.  example  What  (eg Monday,  specific  for  run-timings  fishery  given  week  t h a t peak s t o c k  "short" w i l l  escapement  approach  rather  that  suggest  for reducing  apparent.  estimated  results  obtained  alternatives  are  Sunday).  strategy  composite,  time.  maximizing  days per  steelhead?  while of  for five  Saturday,  fishing  Firstly,  period  continue  be  more  continuous practice).  in fact  intensive fishery  migrate tagging  area  would  point.  problem  of  interspersed  windows  is  11 6  potentially which  apparent  is restricted  During  periods  to the  of  abundance) s e i n e r s caught  a  few  been p r e s e n t . harm than  fishing  allowing  when  fishing  to  technique  12 h o u r s p e r  the  fishing  restrict same  the  could occur  p e a k s of  steelhead  through  the  fishery  the  not  those  fleet  negotiate  taken  reopens a  few  days  later.  escapement may  be  the  anyway Under  greater  by such  using  openings. reduce  the  daily five  fishing  "nightly"  effort  salmon  closures.  Thus,  days  nightly  able  sockeye while  to  movements do,  and  fleet  continuously be  from  windows  commercial  abundance would s t i l l  (assuming  seiners  time to b u i l d ;  are  full  be  more  and  fishery  salmon  would  may  closure  f o r f o u r or stock  area.  do  seine  T h i s would c r e a t e  as  fishery  normally  pressure  movements of  extent  fleet,  such p e r i o d s  i s to simply  day.  seine  (peak s o c k e y e  mouth had  outer  steelhead  longer  the  the  that  river  fishing  the  overall  third  would not  the  to pass the  a n o r m a l p a t t e r n of  to  steelhead at  of  of  remove  during  circumstances,  24  regions  Window c l o s u r e s d u r i n g  area  The  outer  presence  seiner a c t i v i t y  managing  gillnetters  the  intense  days l a t e r  good by  steelhead  during  escape  in  fact,  occur). In summary, I b e l i e v e ' t h a t is  feasible  the  commercial  for  stocks  time.  specific  identification  salmon  statistically  mixed in  f o r the  with  the  fishery.  separating  The  technique of  stock  run-timing  the  technique  curves  can  through  be  and  used  the  patterns  steelhead  provides  m e a s u r a b l e bounds of c o n f i d e n c e  Secondly,  scale  Skeena R i v e r  technique  each  of  for at  a  means  classifying any  point  to c o n s t r u c t  fishery;  in  with  stock  further  11 7  investigation technique from  new  c a n be u s e d  various  technique  is flexible  thesis  pattern  or e f f o r t . will  of  commercial  and  does  Only  in run-timing, impacts  fishing.  and c a n t h e r e f o r e be  becomes a v a i l a b l e .  implementable  expenditure  to p r e d i c t the future  patterns  information  easily  this  to quantify yearly v a r i a b i l i t y  a n a l y s i s as a p r a c t i c a l  stock  T h i r d l y , the  easily  modified  F o u r t h l y , the technique  not  require  further extension  establish  t o any  the  the long  large  is  capital  of t h e r e s u l t s of  term u s e f u l n e s s  management  as  tool.  of s c a l e  1 18  LITERATURE CITED  Amos, M.H., R. 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L e c t u r e s in F i s h e r i e s , Univ. B r i t i s h Columbia, Canada.  of s t o c k s , Macmillan Vancouver,  128  APPENDICES  129  A p p e n d i x A:  Discriminant  Analysis  and  Classification  Discriminant analysis reduces the variable vectors for i n d i v i d u a l s and c e n t r o i d s t o s i n g l e v a l u e s ( c a n o n i c a l v a r i a b l e s , Di) by forming linear c o m b i n a t i o n s of t h e o r i g i n a l v a r i a b l e s weighted according to their contribution to between groups discriminance ( u s i n g p a r t i a l one way ANOVA v a r i a b l e F s c o r e s as e n t r y c r i t e r i a ) . The d i s c r i m i n a n t f u n c t i o n s a r e of the f o r m : D i = d Z +dZ + i 1 i1 2 i2  ....+d z p i p  where D i i s t h e d i s c r i m i n a n t s c o r e , f o r t h e i t h individual, d1, d2, ....dp a r e t h e w e i g h t i n g c o e f f i c i e n t s and z i 1 , z i 2 . . . z i p a r e s t a n d a r d i z e d v a l u e s of t h e measurements from t h e i t h i n d i v i d u a l . The weighting c o e f f i c i e n t s are c a l c u l a t e d so t h a t t h e D i a r e s t a n d a r d n o r m a l v a r i a b l e s and the g r a n d mean d i s c r i m i n a n t score is zero with a standard deviation of one. Discriminant f u n c t i o n s , t h e i r number being one less than the number of groups, are orthogonal to each other and describe group v a r i a t i o n along d i f f e r e n t d i r e c t i o n a l axes ( f i g u r e 6). The m a j o r a s s u m p t i o n s of d i s c r i m i n a n t a n a l y s i s are a) that the groups being distinguished are identifiable (b) that the v a r i a b l e s y s t e m b e i n g u s e d i s m u l t i v a r i a t e n o r m a l and ( c ) t h a t the g r o u p s a l l s h a r e a common v a r i a n c e - c o v a r i a n c e structure. Assumption b was tested as best p o s s i b l e by t h e e x a m i n a t i o n of u n i v a r i a t e frequency d i s t r i b u t i o n s . Assumption c was tested by the a p p l i c a t i o n of Box's m u l t i v a r i a t e M t e s t ( N i e , 1975). Classification matrices (confusion m a t r i c e s , J o h n s o n and W i c h e r n , 1982) a r e d e r i v e d i n d i s c r i m i n a n t a n a l y s i s t h r o u g h the use of classification functions; one for each stock. An e m p i r i c a l measure of g r o u p ( s t o c k ) s e p a r a b i l i t y i s obtained by classifying the i n d i v i d u a l s used t o c o n s t r u c t the d i s c r i m i n a n t f u n c t i o n s i n t o t h e i r most p r o b a b l e g r o u p s of origin using the classification functions. Lachenbruch's (1975) holdout c l a s s i f i c a t i o n p r o c e d u r e ( j a c k n i f i n g ) was u s e d i n t h i s s t u d y to reduce the bias i n p r e d i c t i n g c l a s s i f i c a t i o n e r r o r r a t e s when using the same individuals for both discrimination and classification. Incidentally caught steelhead from the 1984 commercial f i s h e r y p r o v i d e d t h e samples of unknown s t o c k c o m p o s i t i o n t o be classified to stock of origin. Of p r i m a r y c o n c e r n were t h e r e l a t i v e p r o p o r t i o n s of each stock predicted to be present during each week of the salmon f i s h e r y . W o r l u n d and F r e d i n (1962) first described linear equations which adjust the predicted proportional estimates from the mixed sample to a c c o u n t f o r t h e e r r o r s i n a s s i g n i n g i n d i v i d u a l s of known origin (the learning samples). Cook and Lord (1978) extended the p r o c e d u r e t o more t h a n two s t o c k s u s i n g matrix algebra. Using their m e t h o d o l o g y , t h e c l a s s i f i c a t i o n a c c u r a c y e s t i m a t e d by t h e h o l d o u t p r o c e d u r e f o r a g i v e n l e a r n i n g sample i s r e p r e s e n t e d by the s q u a r e m a t r i x C, where t h e e l e m e n t C i j i s t h e p r o p o r t i o n of t h e sample from s t o c k j t h a t i s c l a s s i f i e d as s t o c k i . L e t t i n g r be a c o l u m n v e c t o r r 1 , r 2 , r 3 , . . . . r i , where r i i s the proportion  130  of  the mixed  sample c l a s s i f i e d  as  stock  I  then:  -1 U = C  r  where e a c h e l e m e n t of t h e column v e c t o r U (U1, U 2 , . . . U i ) i s t h e e s t i m a t e of t h e p r o p o r t i o n of s t o c k I i n t h e commercial sample after correcting for t h e e r r o r s i n c l a s s i f y i n g i n d i v i d u a l s of known o r i g i n . V a r i a n c e s a b o u t t h e s e p o i n t estimates (Ui) were e s t i m a t e d u s i n g t h e f o r m u l a e of P e l l a and R o b e r t s o n (1979) and a 90% confidence interval was c a l c u l a t e d f o r e a c h e s t i m a t e . The c o r r e c t i o n p r o c e d u r e of Cook and Lord (1978) is basically a m o d i f i c a t i o n of t h e two s t o c k l e a r n i n g sample s c e n a r i o :  Actual  C l a s s i f i e d Stock A B Aa • Ab  Stock A  =C B  Ba  Bb  where Aa, Ab, Ba, and Bb a r e t h e p r o p o r t i o n s of f i s h from t h e i r r e s p e c t i v e s t o c k s c o r r e c t l y (Aa, Bb) and incorrectly (Ab, Ba) classified. Aa and Bb are the estimated probabilities of correctly classifying an unknown individual which actually belongs to one of those stocks whereas Ab and Ba a r e t h e e s t i m a t e d p r o b a b i l i t i e s of m i s c l a s s i f y i n g an i n d i v i d u a l a c t u a l l y b e l o n g i n g t o one of t h e s t o c k s as b e i n g from the other. In a mixed fishery sample, the proportions of fish a s s i g n e d by d i s c r i m i n a n t a n a l y s i s t o e a c h s t o c k (Pa, Pb) r e p r e s e n t b o t h the correctly assigned individuals plus the u n c o r r e c t l y a s s i g n e d i n d i v i d u a l s . S o l v i n g f o r Na and Nb, t h e actual proportions of each stock present in a mixture,i s by solution of two simultaneous equations: Pa Pb  = Aa = Ba  r  =  Na Na  + Ab + Bb  Nb Nb  or which reduces Lord (1978).  -1 C*U  to the m a t r i x  ..U  = C  adjustment  r procedure  of  Cook  and  The e l e m e n t s of U c a n be g r e a t e r t h a n z e r o , l e s s t h a n z e r o , or e q u a l t o z e r o d e p e n d i n g upon t h e p r o p o r t i o n of a g i v e n s t o c k a c t u a l l y i n t h e c o m m e r c i a l sample. P r o p o r t i o n a l e s t i m a t e s less than zero indicated the a b s e n c e of a p a r t i c u l a r s t o c k i n t h e s a m p l e . Any s a m p l e s r e s u l t i n g in proportional estimates less than zero in this study were reanalyzed using discriminant m o d e l s w h i c h d i d not i n c l u d e t h o s e s t o c k s .  Appendix  T . 1 . Age c o m p o s i t i o n s t r u c t u r e used i n the study.  f o r the f i v e  stocks  AGE CLASS +  RI  SEX  2. 1  1  M F  0 0  M F  3  4  5  +  +  +  +  +  3.2  3.3  4.1  4.2  4.3  -  0 0  -  4 1 .05  9 14 .25  4 1 .05  3 2 .05  13 19 .35  1 8 1 12 .02 .22  42 50 1  %  0 0  -  0 0  8 4 5 4 . 1 5 .09  1 0 .01  10 28 .43  9 10 .22  1 2 7 0 .01 .10  35 54 1  M F  %  0 1 .01  0 0  1 2 .03  12 16 .29  12 05 .16  1 1 .02  15 1 1 .25  8 5 3 10 . 1 1 .14  54 49 1  M F  0 0  0 0  2  21 35 .62  1 0 .01  1 3 .04  7 17 .26  0 1 .01  2 0 .02  34 57 1 36 54 1  -  -  1  RS  TOTA1  O.  -b  -  -  .03  M F  0 0  0 0  0 0  3 9 .13  1 6 0 4 . 1 1 .01  14 31 .50  1 1 3 . 16  1 7 .09  .05  .27  .09  .31  .06  .12  %  -  TOTALS % .002 RI  +  3.1  %  2  +  2.2  KEY :  1 2 3 4 5  = = = = =  -  -  -  ZYMOETZ, MORICE , KISPIOX, BABINE , SUSTUT ,  1974,1978 1977 1975 1978 1977,1983  . 1  1  n = 92 n = 90 n=l03 n = 9l n = 90  1  A p p e n d i x T . 2 . S i z e s a t age f o r t h e f i v e s t o c k s used i n t h e s t u d y . R e p o r t e d a r e t h e means, s t a n d a r d d e v i a t i o n s and sample s i z e s f o r t h e major age c l a s s e s . Kispiox  Age  L 3.1 +  M  X  s n F  X  s n 3.2+  M  X  s n F  X  s n 3.3 +  M  X  s n F  X  s n 4.1 +  M  X  s n X  s n 4.2+  M  X  5 n F  X  s n 4.3+  M  X  s n X  s n RS  M  X  s n F  X  s n Key  M=males  Zymoetz  WT  L  WT  Morice WT  L  Sustut  Babine L  WT  L  WT  --  -  61.1 0 1  2.3 0 1  56.7 2.63 4  1 .8 0.5 4  58.3 4.14 7  1 .8 0.5 7  57.3 1 .27 2  2.0 0 1  59.7 2.63 2  2.9 0.9 2  67.0 0 1  2.7 0 1  55.8 5.51 6  1 .6 0.5 6  60.0 0 1  2.0 0 1  86.5 5.51 10  7.7 1 .3 10  82.0 8.46 7  5.6 2.6 7  75.3 3.33 3  4.0 0.9 3  81.5 6.39 22  5.3 1 .0 22  84. 1 4.47 4  6.0 0.9 2  82.8 6.53 17  5.9 1 .8 17  75.5 3.49 15  4.1 0.6 15  73.2 5.83 5  3.3 0.4 5  78.5 4.72 20  4.6 0.9 20  77.0 4.23 8  4. 1 0.9 5  99.9 8.25 12  10.2 2.4 12  91 .4 6.59 5  7.5 1.8 5  91 .5 0 1  7.5 0 1  91.4 0 1  7.4 0 1  93.3 3.37 6  8.9 0.9 4  87.3 8.88 5  7.5 2. 1 5  88.9 0 1  6.9 0 1  -  -  -  86.4 6.58 4  6.1 1.2 3  55.9 0 1  2.0 0 1  57.8 1 .36 3  1.9 0.4 3  55.9 0 1  1 .8 0 1  62.8 2.47 2  90.0 9.45 13  8.2 2.3 13  77.9 8.35 12  -  -  -  --  --  -  -  —  59.6 4.68 10  1 .8 0.4 10  60.3 0 1  2.0 0 1  55.9 0 1  1 .8 0 1  2.4 0.7 2  56.9 3.43 30  1 .5 0.4 30  60.5 0 1  2.0 0 1  63.5 0 1  2.7 0 1  80. 1 3.91 15  5.0 0.5 15  83.5 7.77 7  5.3 1 .2 7  73. 1 8.52 5  3.4 0.9 5  84.6 7.89 13  6.8 2.4 10  5.3 1.0 12  74.9 4.72 18  4.6 1.8 18  71.9 2.58 14  3.4 0.6 14  74.3 1 .03 7  4.4 0.4 7  77.2 3.45 31  4.6 0.9 22  94.5 7.84 1 1  8.9 1.7 11  96.5 0 1  8.6 0 1  —  -  96. 1 4.02 10  8.9 1 .2 10  87.2 5.26 3  6.7 1.3 3  79.5 0 1  4.5 0 1  -—  87.6 6.0 1 .25 0 1 3  79.1 5.13 3  5.4 0.9 3  80.1 6.10 9  5.4 1 .7 9  66.5 9.19 2  3.0 1 .4 2  75.2 11.1 2  89.4 5.30 10  7.6 1.6 10  84.2 5.40 1 1  5.8 1 .2 1 1  82.6 5.37 7  5.1 1 .4 7  -  F=females  X=mean  -  -  -  s=S.D  -  -  --  n=sample  -  4.1 2.7 2  78.7 0 1  5.4 0 1  -  84.4 2.91 7  5.9 1 .1 4  -  size  133  i x T . 3 . V a r i a b l e means, s t a n d a r d d e v i a t i o n s , and one way ANOVA F s t a t i s t i c s f o r t h e f i v e s t o c k s u s e d i n t h e s t u d y by s m o l t age 3 ( l e a r n i n g s a m p l e s ) .  GROUP VARIABLE 1 PG 3 FWA 4 SWA 5 L 6 WT 7 SEX 8 A 1 9 A2 10 A4 1 1 A5 12 A6 13 B1 14 B 2 IS B 3 16 B4 17 6 5 18 B6 19 C I 2 0 C2 21 C 3 2 2 C4 23 C5 24 C 6 25 01 2 6 D2 27 0 3 28 04 29 D5 3 0 D6 COUNTS  STANDARD GROUP VARIABLE 1 PG 3 FWA 4 SWA 5 L 6 WT 7 SEX 8 A1 9 A2 10 A4 1 1 A5 12 A 6 13 B1 14 B 2 15 8 3 16 B 4 17 B 5 18 B 6 19 C1 2 0 C2 21 C 3 22 C4 23 C5 24 C6 2 5 01 26 D2 27 0 3 2B D4 2 9 D5 30 06  »  KISPIOX  COPPER  SUSTUT  BABINE  MORICE  ALL  0 . 89362 3. 00000 2 . 48936 87 . 2 3 4 0 2 7 . 27234 1 .42553 0. 09234 0 . 14362 0 . 2 1043 6 . 72340 2 . 76596 0. 04702 0 .09745 0 . 14SS3 0. 29043 1 1 .59574 5 . 7 0 2 13 0. 05128 0 . 10787 0 . 16362 0. 34234 12. 5 1 0 6 4 5 .85106 0 .08489 0 .18128 0 .28723 1 .65319 35 . 7 2 3 4 0 16 . 0 0 0 0 0  0 . 4054 1 3 OOOOO 2 . 27027 77 9 3 7 8 2 4 .. 8 8 3 7 8 1 .43243 0 .09351 0 .15568 0 .20189 5 .. 6 7 5 6 8 2.. 3 5 1 3 5 0..04865 0 .09838 0..15081 0..29459 1 1 .59459 s..56757 0..05189 0 .10622 0..16432 0 35514 12. . 6 7 5 6 8 5 .97297 0 .08811 0 .18054 0 .27784 1 .69459 35 . 1 0 8 0 9 15 . 4 5 9 4 6  0 . 26923 3 . OOOOO 2 . 73077 84 . 31 152 6 . OOOOO 1 . 38461 0. 09000 0 . 15462 0 . 23 192 6 . 8 4 6 15 2. 92308 0 . 03654 0. 09500 0 . 15038 0 . 30269 1 1 .53846 5 . 42 3 0 8 0. 04385 0 . 1 1385 0 . 17692 0 . 37654 12. 57692 5 . 8846 1 0 . 06731 0 . 16500 0 . 26885 1 .53615 32 . 1 5 3 8 4 15. 6 5 3 8 5  0. 29310 3 OOOOO 2 01724 77 . 3 3 2 7 5 4 59403 1 39655 0 .09345 0 14983 0 23017 7 32759 3 .06896 0 04931 0 .10569 0 16086 0 . 3462 1 13. . 2 2 4 1 4 6 .25862 0 .05345 0 .11586 0 17776 0 38672 14 . 0 5 1 7 2 6 .55172 0 .08552 0 .19086 0 .30655 1 .78207 34 . 6 3 7 9 2 15 . 9 1 3 7 9  0. 69565 3 . OOOOO 1 . 6 5 2 17 66 21738 2 . 80435 1 .43478 0 10087 0 16522 0 . 24783 7 . 82609 3 . 26087 0..04261 0 0 9 130 0 14 174 0 .24783 10. . 4 7 8 2 6 4 .. 6 5 2 1 7 0 .04087 0 .09174 0 14652 0 .28957 10. 9 5 6 5 2 5 .. 0 8 6 9 6 0 .08000 0 . 17478 0 28609 1 .70435 34 . 4 3 4 7 7 15 . 5 6 5 2 2  0. 50785 3 OOOOO 2 . 23560 79 4 9 7 8 9 5 28534 1 .41361 0 .09361 0 . 15194 0 22220 6 85340 2 .85864 0 .04607 0 .09906 0 .1514 1 0 .30471 11 , 9 4 7 6 4 5 .68063 0 .04979 0 .10885 0 16780 0 .35660 12 8 3 2 4 6 6 . .OOOOO 0 08272 0.,18105 0 28864 1. 6 9 0 5 7 34 .. 6 3 3 5 0 15 . 7 6 9 6 3  47.  37.  58 .  26.  GPS.  191 .  23.  DEVIATIONS KISPIOX 1 . 32261 0. 0 0 . 7481 1 9 . 74736 2. 09032 0 . 49977 0. 02098 0 . 02462 0. 05373 2. 05047 1 . 12699 0 . 00998 0. 01687 0. 02385 0. 06659 2. 14334 1 . 12123 0. 01296 0 .01488 0..02523 0 .08352 2 .91078 1 .36698 0 .01679 0 .02651 0 .03820 0 .38385 6 .33039 2 .57917  COPPER 0 . 92674 0. 0 0 . 87078 10. 74704 2. 0 3 7 9 0 0 . 50225 0. 02530 0. 02387 0. 03865 1 .27048 0. 58766 0 . 0 1 159 0. 01756 0 . 02 1 1 3 0. 06517 2. 25412 1 ,30257 0.,01221 0 .02086 0 .02882 0 .08909 2 .92550 1.32259 0 .02459 0 .04007 0 .04995 0 .30138 s .58660 2 .28028  BABINE  SUSTUT 0. 72430 0. 0 0 . 82741 7 . 26857 1 . 71277 0 . 4 9 6 14 0 . 0 2 135 0. 02970 0. 05485 1 .86959 0. 97665 0 . 01263 0. 01903 0. 02457 0..06372 2. 68672 1 .36156 0 . 01061 0 .01577 0 ,02223 0 .13323 3 .59080 1 .88312 0 .02475 0 .02470 0 .03241 0 .32883 5 .75980 2 .29682  .  0. 67560 0. 0 0 . 39698 7 . 6 1405 1 12475 0 . 49345 0. 0209 1 0 . 0 2 5 17 0 . 0 4 9 19 1 . 64783 0. 93400 0 . 01 197 0. 01836 0. 02364 0 0 9 7 13 2 . 84 106 1 39624 0 . 01 132 0 01697 0 .02527 0 .11063 2 .56441 1 .20193 0 01613 0 .03074 0 .04024 0 .26839 5 .15614 2 .30396  MORICE 0.97397 0.0 0.83168 1 1 .61193 1 .53637 0.5068 7 0.02314 0.03232 0.07580 2.62249 .48377 .01176 .02201 .02289 .05705 19233 1.07063 0.00793 0.01403 0.01774 .05653 .63702 94931 .01706 .03369 .05289 .39773 .47250 .08514  ALL  GPS.  0.95992 0.0 0.71224 9.30948 1 ..7 1 6 2 5 O 49868 O .. 0 2 2 1 6 O .. 0 2 6 3 9 O .. 0 5 3 1 8 1 ..8 6 2 1 4 1.01828 O .. 0 1 1 5 0 0 .. 0 1 8 4 2 O .. 0 2 3 2 7 o .. 0 7 5 7 7 2 .. 4 7 6 2 4 1 .. 2 7 3 3 6 O .. 0 1 1 5 0 .01685 0. 0.02485 0.09887 2.79591 1.35152 0.01958 0.03147. .04257 o .. 3 3 0 6 6 0 .. 7 8 8 9 7 5 .. 3 4 5 5 2 2 .  F TO ENTER DF = 4 186 3 350 0 .0 9 . 876 22 . 5 9 7 31 . 7 4 5 0 .069 0 .829 2 .970 3 .803 6 . 266 3 . 935 6 .677 3 315 4 . 154 8 . 178 6 . 48 1 7 .080 7 . 169 9 . 277 7 933 4 . SOO 5. 585 5 . 234 5 . 281 3 332 4 . 601 2 . 689 1 . 678 0 . 389  134  A p p e n d i x T . 4 . V a r i a b l e means, s t a n d a r d d e v i a t i o n s , a n d one way ANOVA F s t a t i s t i c s f o r t h e f i v e s t o c k s used i n t h e s t u d y by s m o l t age 4 ( l e a r n i n g s a m p l e s ) .  GROUP VARIABLE 1 PG 3 FWA 4 SWA 5 L 6 WT 7 SEX 8 A1 9 A2 10 A4 1 1 A5 12 A6 13 B1 14 B2 15 B 3 16 B 4 17 B 5 18 B 6 19 C1 2 0 C2 21 C 3 2 2 C4 23 C5 24 c e 25 01 26 02 27 0 3 28 •4 2 9 D5 30 06 31 E 1 32 E 2 33 E3 34 E4 35 E5 36 E 6 COUNTS STANDARD GROUP VARIABLE 1 PG 3 FWA 4 SWA 5 L 6 WT 7 SEX 8 A 1 9 A2 10 A4 11 A 5 12 A6 13 B1 14 B 2 15 S 3 16 B 4 17 B 5 18 B 6 19 C1 2 0 C2 2 1 C3 22 C4 23 C5 24 C 6 25 01 2 6 D2 27 0 3 2 8 D4 29 05 3 0 D6 31 E 1 32 E2 33 E 3 34 E4 35 ES 36 E6  KISPIOX  COPPER  SUSTUT  BABINE  MORICE  ALL  0 . 43243 4 . 00000 2. 64865 84 . 6 9 1 8 8 6 . 73513 1 . 51351  0. 29167 4. 00000 2. 56250 77 . 0 0 6 2 4 4. 56458 1 . 45833 0. 09687 0 . 15167 0 . 18708 5 . 3 5 4 17 2 . 12500 0. 04750 0. 09917 0 . 15021 0 . 24 146 9. 50000 4. 37500 0. 04833 0 . 10250 0 . 15833 0 . 27646 10. 4 3 7 5 0 4 . 87 5 0 0 0 . 08167 0 . 1777 1 0 . 28667 1. 7 6 4 7 9 3 6 . 16666 15 . 8 9 5 8 3 O.0 5 1 2 5 0 . 10667 0 . 16396 0 . 31292 10. 85417 4 . 89S83  0 . 33898 4 . 01695 2. 38983 8 3 . 38982 5. 94915 1 . 4067B 0 . 09220 0 . 15559 0 . 2 1 136 6 . 15254 2. 52542 0 . 04305 0 . 10220 0 . 16068 0 . 27339 9 . 89830 4 . 72881 0 . 04373 0 . 10949 0 . 17339 0 . 29797 10. 25424 4 . 94915 0. 07237 0 . 17339 0 . 27661 1. 6 3 0 3 4 32. 77965 1 5 . 6 2 7 12 0 . 04983 0 . 12220 0 . 19068 0. 37220 12 . 1 1864 5. 81356  0. 20000 4 . 00000 1 .9 0 0 0 0 72 . 9 8 6 6 6 .1 . 1 0 3 3 3 1 .3 0 0 0 0 0 . 10000 0 . 15167 0. 22900 7 . 40000 2. 93333 0 04867 0. 09933 0 . 15167 0 28167 1 1 .0 0 0 0 0 5. 20000 0 . 05067 0 . 10700 0 . 16 100 0 . 27633 10. 6 6 6 6 7 5. 03333 0. 08900 0 . 19767 0 . 3 1533 1 .79100 35 . 2 9 9 9 9 15 . 7 0 0 0 0 0 . 05267 0 . 1 1500 0 . 17633 0 . 34733 1 1 .7 0 0 0 0 5 . 56667  0 . 43077 4 .01538 1 .5846 1 6 5 . 04 3 0 8 2 . 62923 1 .3 0 7 6 9 0 . 09815 0 . 15800 0 . 22369 6. 73846 2 6 4 6 15 0 . 04092 0 . 0 9 108 0 . 13723 0 . 22985 9 . 78461 4 . 67692 0 . 04 2 0 0 0 . 09462 0 . 14338 0 . 23938 9 . 96923 4 . 6923 1 0 . 07354 0 . 1743 1 0 . 28262 1 .6663 1 3 3 . 8 1538 15. 33846 0 . 04277 0. 09569 0 . 14 8 0 0 0 . 27646 1 1 . 13846 5 . 20000  0 . 35146 4 . 00837 2 . 184 10 76. 01379 4 . 65816 1 .39330 0 . 09598 0 . 15343 0 . 20753 6 . 20502 2 . 50628 0 . 0 4 4 14 0 . 09649 0 . 14753 0 . 25121 10. 0 0 8 3 7 4 . 74059 0 . 04544 0 . 10293 0 . 15837 0 . 27 113 10. 2 5 1 0 5 4 . 89540 0. 07757 0 . 1779 1 0 . 28736 1 .72213 34 . 8 7 4 4 7 15 . 76 15 1 0 . 0489 1 0 . 10933 0 . 16837 0 . 31628 1 1 . 17 155 5 . 24686  0. 09378 0 . 14568 0 . 18216 5. 48649 2 . 37838 0 . 0 4 351 0 . 09108 0 . 13784 0 . 24135 10. 43243 4 . 97297 0 . 04622 0 . 10432 0 . 15865 0 . 27297 10. 16216 5. 08108 0 . 07838 0 . 17568 0 . 29108 1 . 85540 38. 05405 16 . 5 9 4 5 9 0. 05216 0 . 1 1 162 0 . 16784 0 . 27622 9. 70270 4 . 62162 37 .  48 .  59 .  30.  GPS.  239 .  65 .  DEVIATIONS KISPIOX 0 .95860 0.,0 0 .85687 10. . 9 0 0 8 6 2.. 1 4 5 5 6 0..50671 0..02487 0..02588 0. 04905 1 21613 0..63907 0 01060 0 .01370 0 .02175 0..05271 1 .95136 0 .95703 0 .01361 0 .01980 0 .02594 0 .06806 2 .06173 1 .78541 0 .02328 0 .01994 0 .03116 0 .38639 7 .26080 2 . 6 6 103 0 .01250 0 .02089 0 .02678 0 .05574 1 .85390 0 .95310  COPPER 0 .7 7069 0 .0 1 18333 8 .33509 1 .50325 0.,50353 0 .02085 0 .02579 0 ,03690 1 .02084 0 .53096 0 .01139 0..01699 0 .02274 0 .04807 1 .65027 0 .81541 0 .01243 0 .01781 0 .02435 0 .06406 1 .79723 0 .93683 0 .02014 0 .02860 0 . 0 4 138 0 .29673 5 .92266 2 .51158 0 .01315 0 .01849 0 . 03009 0 .07252 2 .19273 1 .01561  SUSTUT 0 .88298 0 .13019 0 .61635 8 .40368 1 ,. 8 0 4 5 3 0 49545 0..02009 0.,02430 0..04066 0 .99678 0..67864 0. 01235 0..02026 0..02399 0..05827 2 .09016 0 .94377 0 .01299 0 .02021 0 .02577 0 .06501 1 .88087 1 .02425 0 .02029 0 .03693 0 .04334 0 .28399 5 .68745 2 .37005 0 .01491 0 .02335 0 .02888 0 .09828 2 .76732 1 .80492  BABINE 0 .550B6 0 .0 0 .40258 5 .51172 0 .76089 0 .46609 0 .02133 0.,02793 0 .05215 1 8 1 184 0 . 94443 0 01224 0. 01660 0 .02183 0..08914 2 .75431 1 .37465 0 .01285 0 .02292 0 .02940 0 .05980 1 74856 0 .85029 0 .01689 0 .02788 0 03329 0 .34343 5 .79624 2 .32156 0 .01230 0 .01737 0 .02141 0 .11020 3 .97534 2 .04574  MORICE 0 .88334 0 .12403 0 .89952 10 . 8 1 0 6 3 1.50610 0 .46513 0 .02098 0 .02852 0 .05421 1 .31431 0 .75892 0 .01142 0 01724 0 .02058 0 .04185 1 .60558 0 .81216 0 .00870 0 .01370 0 . 0 2 138 0 .04596 I .63906 0 .88252 0 .02080 0 .03455 0 .04925 0 .24515 4 .98087 2 .05606 0 .00976 0 .01677 0 .02251 0 .08352 2 .68022 1 .32523  ALL  GPS.  0 83960 0 0 9 170 0. 85425 9. 24489 1 .6 3 1 3 5 0. 48720 0. 02143 0. 02650 0. 04684 1. 2 4 9 9 5 0 . 70829 0 . 01 164 0 . 01744 0. 02223 0. 05654 1. 9 6 4 9 0 0..95384 0 . 01 194 0 .01848 0 .02489 0 .06005 1 8 1472 1.10875 0 ,02051 0 .03141 0 .04195 0 .3028 1 5 .84140 2 .36067 0 .01260 0 .01963 0 .02635 0 .08568 2 .70478 1 .46743  DF=  ENTER 4 23' 0 . 539 0 .464 14 . 7 7 8 4 1 . 365 5 0 . 272 1. 565 1 .007 1 .460 8 .607 18 . 4 5 7 7 . 150 3 . 533 4 . 522 10. 842 7 . 404 3 . 401 4 . 126 3. 835 5 . 570 1 1 .3 4 8 7 .648 0 934 0 . 959 4 . 399 3 . 535 4 . 588 4 . 325 5 . 798 1 . 766 5 . 637 15 . 167 2 1 .3 0 6 12 . 8 1 9 4 . 990 4 . 938  ' 135  ix  T.5. V a r i a b l e means, s t a n d a r d d e v i a t i o n s , a n d one way ANOVA F s t a t i s t i c s f o r t h e f i v e s t o c k s u s e d i n t h e s t u d y by age 3.2 + ( l e a r n i n g s a m p l e s ) .  GROUP VARIABLE 1 PG 3 FWA 4 SWA 5 L 6 WT 7 SEX 8 Al 9 A2 10 A4 1 1 A5 12 A6 13 B1 14 B 2 15 B 3 16 B4 17 B 5 18 B 6 19 C1 2 0 C2 21 C 3 22 C 4 23 C5 24 C 6 2 5 01 26 02 27 0 3 28 04 29 D5 3 0 D6 COUNTS  STANDARD GROUP VARIABLE PG FWA SWA L WT SEX 8 A1 9 A2 10 A4 1 1 A5 12 A 6 13 B1 14 B 2 15 B 3 16 B4 17 B 5 18 B 6 19 C I 2 0 C2 21 C 3 22 C4 23 C5 24 C6 25 DI 26 D2 27 D 3 28 04 29 D5 30 06  -  KISPIOX  COPPER  SUSTUT  BABINE  MORICE  ALL  1 , OOOOO 3 . OOOOO 1 .9 2 5 9 3 83. 05554 6. 47037 1 .. 3 7 0 3 7 0..09074 0 .14407 0..21370 6. 74074 2.. 7 7 7 7 8 0 .04667 0 .09704 0 . 14519 0 28444 11 . 2 9 6 3 0 5 .. 7 0 3 7 0 O ,05148 0 .10889 0 .16630 0 .32556 11 . 8 5 1 8 5 5 .62963 0 .08296 0 .17852 0 .28630 1 .69963 37 . 2 2 2 2 1 16 . 4 4 4 4 4  0..55556 3 OOOOO 1 . 81481 74 . 4 8 5 1 7 4 . 08889 1 .44444 0..08926 0..15185 0 20000 5, 85185 2 .. 4 4 4 4 4 0. 04815 0. 09852 0 .15111 0.. 2 9 0 0 0 11 6 6 6 6 7 5. 66667 0. 05074 0..10407 0 .16037 0 .33333 12 . 2 5 9 2 6 5 .. 8 5 1 8 5 0 .08444 0 .17519 0 .27148 1 .69555 35 . 1 4 8 1 5 15 . 5 9 2 5 9  0 . 25000 3 . OOOOO 2 . OOOOO 79. 38332 4 . 90833 1 . 33333 0 . 08667 0 . 15583 0. 23167 6. 91667 2 . 83333 0 . 0 3 4 17 0 . 09167 0 . 14667 0 . 30167 1 2 . OOOOO 5. 75000 0 . 0 4 4 17 0 . 1 1000 0 . 16750 0. 30750 10. 91667 5 . 16667 0. 06583 0 . 16500 0 . 26667 1 .4 7 0 0 0 30. 75000 14 . 9 1 6 6 7  0. 29091 3 OOOOO 1 .94545 77 . 28 18 1 •4 . .5 8 7 2 7  0 60000 3 . OOOOO 1 . 40000 63 . 7 2 4 9 9 2 . 45000 1 .4 5 0 0 0 0 . 10150 0 16700 0. 25900 8. 20000 3. 45O00 0 . . 0 4 ISO 0..08950 0..14000 0 .24850 10 5 5 0 0 0 4 . 70000 0 .04000 0 .09200 0 .14700 0 .28750 10 . 9 0 0 0 0 5 .OOOOO 0 .08100 0 .17700 0 .28550 1 .67350 34 . 2 0 0 0 0 15 . 6 5 0 0 0  0 . 51773 3 . OOOOO 1 .84397 76 . 1 0 7 7 9 4 . 57660 1 . 3 9 7 16 0 . 09291 0 . 15220 0 . 22567 7 . 02837 2 . 9 3 6 17 0 . 0 4 6 17 0 . 09908 0 . 15156 0 . 30553 12 . 0 7 8 0 1 5 . 77305 0. 04979 0 . 10858 0 . 16723 0 . 34489 12 . 6 0 2 8 4 5 . 92 199 0 . 08234 0 . 18 156 0. 28943 1. 7 1 1 0 6 = 34 . 9 2 9 0 8 15. 8 7 9 4 3  27.  27.  12.  COPPER  SUSTUT  1 .3 8 1 8 2 0 .09400 0 .15018 0 23073 7 . 34545 3 .. 0 9 0 9 1 0. 04927 0 .10545 0 16018 0 .34509 13 . 2 3 6 3 6 6. 25455 0..05327 0 .11636 0 ,17836 0 .38909 14 . 1 2 7 2 7 6 .60000 0 .08509 0 . 19145 0 . 3 0 6 18 1 .79054 34 . 8 7 2 7 3 16 . 0 3 6 3 6 55.  20.  GPS.  14 1.  DEVIATIONS KISPIOX 1.38675 0.0 0.26688 9.45085 2.03334 0.49210 0.02093 0.02358 0.05197 2.04925 1.01274 0.01144 01540 02486 06606 07206 13730 01292 01340 02589 08568 .93131 .44510 .01772 .02670 0.03904 0.36378 5.04847 2.25888  1. 0 5 0 0 3 0. 0 0 . 39585 9. 55879 1 .61896 0. 50637 0. 02464 0. 02434 0 . 03772 1 . 29210 0. 64051 0,.01210 0. 01895 0. 02190 0. 06139 2 . 41788 1 38675 0..01207 0 .02080 0 .02848 0 .09004 3 .10821 1 .40613 0 .02375 0 .03817 0 .05013 0 .27301 5 .55187 2 .25762  62158 0 0 39491 01216 0 . 49237 0. 02309 0 . 03175 0 . 06043 1. 9 2 8 6 5 0 . 93744 0. 01505 0. 02368 0. 03085 0. 07371 3. 49024 1 . 71225 0 . . 0 1 164 0 .01128 0 .01485 0 ,07569 2 .90637 1 .40346 0 .01975 0 .02505 0 .03143 0 .36449 5 .97152 1 .97522  0. 0. 0. 5. t .  0. 68510 0. O 0 . 22918 7 .05696 1 .0 4 0 8 4 0 . 4903 1 0 . 02 122 0 . 02535 0. 05036 1. 6 8 0 1 5 0 . 9 4 8 15 0 01230 0. 01854 0. 0236 1 0. 09937 2 . 87365 1 ,43007 0 .01123 0 .01671 0 .02507 0 . 1 1263 2 .59667 1 .21106 0 .01620 0 .03123 0 .04039 0 .27288 5 .18563 2 .30107  0 . 94032 0. 0 0 . 50262 9 . 84280 1 .0 8 8 4 5 0 . 51042 0. 02300 0 . 03278 0. 07483 2 . 58742 1 . 50350 0. 01226 0. 02305 0 . 02406 0. 0585 1 2 . 18788 1 .0 8 0 9 3 0 .00795 0 .01473 0 .01895 0 .06034 1 .68273 0 .91766 0 .01651 0 .03278 0 .05336 0 .39001 6 .20356 2 .13431  0 . 95894 0. 0 0 . 31579 8 . 37883 1 .40357 0 . 49676 0. 02226 0. 02658 0 . OS360 1 .8 6 5 2 0 1 . 0 0 9 13 0. 01234 0. 01924 0. 02427 0. 07996 2 . 62132 1 .35178 0 . 01 139 0 01638 0 02451 0 .09467 2 .69177 1 .27927 0 .01849 0 .03167 0 .04356 0 .31798 5 .45005 2 .23700  F TO ENTER DF = 4 136 2 758 0 .0 12 . 5 4 7 16 . 5 4 3 24 . 7 5 0 0 .200 1 . 259 2 .318 3 . 978 5 . 227 3 .420 4 .605 3 .277 3 .460 6 .64 1 5 153 4 . 956 5 912 a .758 6 780 5 .685 8 . 223 7 878 2 8 18 2 602 4 . 075 2 . 678 3 . 06 1 1 . 217  136  i x T.6. V a r i a b l e means, s t a n d a r d d e v i a t i o n s , and one way ANOVA F s t a t i s t i c s f o r t h e f i v e s t o c k s u s e d i n t h e s t u d y by a g e 4.2 + ( l e a r n i n g s a m p l e s ) . MEANS GROUP VARIABLE 1 PG 3 FWA 4 SWA S L 6 WT B A1 9 A2 10 A4 1 1 A5 12 A6 13 B1 14 B 2 15 B 3 16 B4 17 B 5 18 B 6 19 C1 2 0 C2 21 C 3 22 C4 23 CS 24 C 6 25 01 26 0 2 27 0 3 28 04 29 05 3 0 D6 31 E1 3 2 E2 33 E3  34 35 36  E4 E5 E6  COUNTS  KISPIOX  COPPER  SUSTUT  BABINE  MORICE  ALL  GPS. DF =  0 .44444 4 OOOOO 2 .69444 85 . 4 9 1 6 5 6 .87222 0 .09417 0 .14583 0 .18222 5 .47222. 2 . 3 6 1 11 0 .04361 0 .09083 0 .13750 0 .24222 10. . 4 7 2 2 2 5 OOOOO 0. 04639 0 . 10417 0 . 15917 0 . 27528 10. 22222 5 . 11111 0. 07917 0 . 17556 0 . 29083 1 . 84778 37. 91666 16. 52777 0 . 05250 0 . 1 1 194 0 . 16778 0 . 27556 9. 66667 4 .61111 36.  0 .32558 4 .OOOOO 2 .74419 79 . 0 1 1 6 1 4 . 8 5 1 16 0 .09651 0 .15163 0 .18651 5 .32558 2 .09302 0 .04837 0 .10023 0 .. 15209 0 .24442 9 .. 4 4 1 8 6 4 . 39535 0. 04953 0 . 10512 0 . 16256 0 . 28116 10. 39535 4. 88372 0. 08395 0 . 181 16 0. 29070 1 . 77581 36. 06976 15. 90698 0 . 05256 0 . 10721 0 . 16442 0 . 30907 10. 72093 4 . 81395  0 .33898 4 .01695 2 .38983 83 . 3 8 9 8 2 5 . 9 4 9 15 0 .09220 0 .15559 0 .21136 6 .15254 2 .52542 0 .04 305 0 .10220 0 .16068 0 .27339 9 .89830 4 72881 0 04373 0..10949 0 . 17339 0 . 29797 10. 25424 4 . 94915 0 . 07237 0 . 17339 0 . 27661 1 . 63034 32. 77965 15. 62712 0. 04983 0 . 12220 0 . 19068 0 . 37220 12 . 1 1864 5. 81356  43.  59.  COPPER  SUSTUT  0 .23077 4 OOOOO 2 .03846 74 . 3 9 6 1 5 4 .13077 0 .10038 0 .15269 0 .22885 7 .53846 3 .OOOOO 0 .04692 0 09769 0 14885 0. 27346 10. 9 6 1 5 4 5 . 1 1538 0. 04846 0 . 10500 0 . 15846 0 . 27731 10. 7 6 9 2 3 5. 07692 0. 09000 0 . 19962 0 . 3 1846 1 . 7 9 6 15 35 6 5 3 8 4 IS. 80769 0 . 05154 0 . 1 1385 0 . 17500 0 . 34 192 1 1 .7 3 0 7 7 5 . 53846 26 .  0 .26923 4 .03846 2 .46154 76 . 2 1 9 2 2 4 .12692 0 .09654 0 .15692 0 .22731 7 .07692 2 .80769 0 .04077 0 .08923 0 .13577 0 22269 9 .80769 4 . 57692 0. 04385 0. 09538 0 . 14577 0 . 24885 9 . 88461 4 . 65385 0. 07000 0 . 16654 0 . 2 7 192 1. 7 0 3 4 6 35. 61537 15. 76923 0. 04423 0 . 09654 0 . 14923 0 . 26846 1 1 .OOOOO 5 . 15385  0 .33158 4 .01053 2 .48947 80 .58525 5 .37737 0 .09526 0 .15263 0 .20479 6 .15263 2 .50000 0 .04458 0 .09721 0 14932 0 .25400 10. . 0 3 6 8 4 4 . 73684 0 . 04621 0 . 10495 0 . 16242 0 . 28032 10. 3OOO0 4 . 94210 0. 07837 0 . 17821 0 . 28758 1. 7 3 7 16 3 5 . 27895 15. 90526 0 . 0504 2 0 . 1 122 1 0 . 17258 0 . 32126 1 1 .1 3 1 5 8 5 . 23158  26.  190.  STANDARD D E V I A T I O N S GROUP VARIABLE 1 PG 3 FWA 4 SWA 5 L 6 WT 8 A1 9 A2 10 A4 1 1 A5 12 A 6 13 B 1 14 B 2 15 8 3 16 B 4 17 B5 18 B 6 19 C1 20 C2 21 C 3 22 C4 23 C5 24 C 6 25 D I 26 0 2 27 0 3 28 D4 29 05 3 0 D6 31 E 1 32 E2 3 3 E3 34 E4 3 5 E5 36 E6  KISPIOX 0 . 96937 0. 0 0 . 821B2 9 . 89349 2 . 00494 0 . 0251 1 . 0 . 02623 0. 04975 1 .23024 0. 63932 0. 01073 0 . 01381 0 . 0 2 196 0. 05319 1 .96376 0..95618 0. 01376 0..02005 0.,02612 0 .06755 2 .05789 1 .80123 0 .02310 0 .02021 0 .03157 0 .38904 7 .31485 2 .66711 0 .01251 0 .02109 0 .02716 0 .05639 1 .86701 0 .96445  0 . 80832 0. 0 1 . 1 1468 6 . 10534 1 . 30190 0. 02080 0. 02600 0. 03810 1 .01702 0 . 52617 0 . 01111 0. 01739 0 . 0 2 3 15 0. 04896 1 .7 0 8 5 5 0. 84907 0. 01234 0. 01653 0..02128 0..06272 1 .80131 0 .93119 0 .01966 0 .02822 0 . 0 4 17 1 0 .29626 5 .96575 2 .57102 0 .01311 0 .01919 0 .03165 0 .07243 2 .11936 0 .98212  0. 88298 0 . 13019 0. 61635 8. 40368 1 .8 0 4 5 3 0. 02009 0. 02430 0. 04066 0. 99678 0. 67864 0 01235 0 02026 0. 02399 0 05827 2. 0 9 0 1 6 0 .94377 0.,01299 0 .02021 0 .02577 0 .06501 1 .88087 1 .02425 0 .02029 0 .03693 0 .04334 0 .28399 5 .68745 2 .37005 0 .01491 0 .02335 0 .02888 0 .09828 2 .76732 1 .80492  BABINE 0 . 58704 0. 0 0 . 19612 3. 76843 0 . 68397 0 . 02254 0 . 02878 0. 05450 1 .8 5 9 6 9 0 . 97979 0 . 01 192 0. 01704 0. 02142 0 0897 1 2 . 86329 1 .42343 0 . 01 156 0 .02319 0 .02880 0 .06213 1 .79572 0 .89098 0 ,01789 0 .02932 0 .03379 0 .35787 5 .95947 2 .41693 0 .01223 0 .01813 0 .02232 0 .11275 4 1042 1 2 .08290  MORICE 0 . 77757 0 . 19612 0. 85934 7. 77529 1 .29230 0 02226 0 0 2 6 19 0. 05604 1 38342 0 89529 0. 0089 1 0 . . 0 1 7 19 0 0 2 194 0 04423 1, 6 4 9 7 1 0. 94543 0..00898 0 .01421 0 .02062 0 .04366 1 . 6 8 1 12 0 .89184 0 .02040 0 .02993 0 .04656 0 .28305 5 .02056 2 .04563 0 .00902 0 .01573 0 .02331 0 .0629 1 2 .51396 1 .22286  ALL  GPS.  0. 83595 0 . 10252 0 . 7962 1 7 . 69473 1 .5 6 6 8 5 0 . 0 2 190 0 02595 0. 04628 1 .24845 0 . 72292 0 .01130 0 ,01768 0 0228 1 0.,05912 2., 0 5 8 5 4 1 ,00523 0 .01234 0 .01913 0 .02469 0 .06214 1 .86133 1 .16376 0 .02042 0 .03038 0 .04024 0 .31934 6 .04557 2 .44179 0 .01302 0 .02041 0 .02773 0 . 0 8 4 10 2 .68522 1 .48480  F TO ENTER 4 18! 0 . 296 0 .817 4 .021 12 . 3 6 6 19 . 6 2 2 0 .723 1. 0 0 4 7 .408 18 . 9 6 3 8 .044 2 . 567 3 .990 8 529 4 , 753 2 . 759 2. 945 1 ,835 2 . 47 1 6 19 1 2 934 0 . 790 0 . 703 5 . 288 4 . 728 6 . 031 3 . 182 4 . 463 0 . 807 2 . 066 8. 059 12 . 139 1 1 .2 4 9 5 . 262 4 . 983  4  Appendix  T.7. Age c o m p o s i t i o n s t r u c t u r e f i s h e r y s t e e l h e a d samples.  f o r t h e 1984  commercial  AGE CLASS + WK 9  10  SEX  +  +  +  +  +  +  2.2  3.1  3.2  3.3  4.1  4.2  4.3  %  2 0 .02  2 0 .02  10 1 .08  27 20 .35  6 4 .08  12 5 .13  14 14 .21  6 2 4 4 .08 .05  81 52 1  M F  0 0  1 0 .01  7 4 .08  28 25 .41  7 4 .08  8 3 .08  10 7 .13  7 9 7 3 . 1 1 .09  77 53 1  2 7 0 6 .01 . 10  82 52 1  M F .  %  1 1  +  2.1  -  RS  TOTAL  M F  %  2 0 .01  2 0 .01  12 5 . 13  25 25 .37  6 1 .05  1 1 8 .14  15 7 .16  M F  0 0  0 2 .02  17 6 . 18  16 25 .32  6 12 . 14  9 3 .09  5 3 9 9 0 5 . 1 1 .02 . 1 1  65 62 1  M F  %  1 0 .01  1 2 .02  17 24 2 20 . 1 5 .34  1 1 1 .09  2 4 .05  7 6 .10  7 14 0 1 1 .05 .19  84 46 1  M F  0 0 -  5 2 .06  7 3 .08  22 17 .33  7 9 7 2 . 1 4 .08  10 15 .21  3 1 .03  2 6 .07  65 53 1  TOTALS % .01  .02  .11  .36  .10  .16  .05  .10  1  WEEK KEY: 9 = 10 = 1 1 = 12= 13 = 1 4=  ending ending ending ending ending ending  12  -  %  13  14  %  •  July July Aug. Aug. Aug. Aug.  .10  21 n= 133 31 n = 1 30 7 n= 134 1 4 n= 127 21 n= 130 31 n = 1 18  13 7  138  Appendix  T.8. N u m e r i c a l r u n t i m i n g e s t i m a t e s f o r t h e f i v e s t o c k s u s e d i n t h i s s t u d y t h r o u g h t h e 1984 c o m m e r c i a l f i s h e r y . Model A: s m o l t age 3 / s c a l e v a r i a b l e s a l o n e . M o d e l B s m o l t age 4 / s c a l e v a r i a b l e s a l o n e . Model C p o o l e d s m o l t a g e / s c a l e v a r i a b l e s a l o n e . Model D p o o l e d smolt a g e / a l l v a r i a b l e s . 1984 e s t i m a t e d run t i m i n g  by week  model a 10  1  -  1 244 626 5418 44 1 -  1606 1219 2055  -  .115 . 106 .272 .632 -  9 stock 3670 mor i c e 210 babine 3670 sustut zymoetz kispiox 525 by p r o p o r t i o n .340 mor i c e .036 b a b i ne . 184 sustut zymoetz .257 kispiox model  D  9 stock 1707 mor i c e 787 babine 2003 sustut zymoetz 1223 kispiox 1001 by p r o p o r t i o n .423 morice .376 babine .252 sustut zymoetz .239 kispiox .299 model c  9  stock 5164 morice 885 babine 6368 sustut 551 zymoetz 1537 k i spiox by p r o p o r t i o n .369 morice .107 babine .224 sustut zymoetz . 1 32 .200 kispiox model d  9 stock 8689 morice 1436 babine 4395 sustut . zymoetz kispiox by p r o p o r t i o n .375 morice .136 babine  -  sustut zymoetz kispiox  .181  -  12  13  2374 2762 3850 -  1 330 1073 3225 948  -  . 149 .207 . 103 .280  .220 .469 .193 -  . 123 .182 .162  -  .463  -  .052 .087 .368 -  10  1 1  12  13  14  1064 532 1812 1 169  492  687 1 032 1 1 46 687  394 1289 693 495  375 83 751 334  4032 2089 7953 5117 3352  . 122 . 134 .336  .162  -  .349  -  .329 . 130 .224 .205  1 .00 1 .00 1 .00 1 .00 1 .00  10  1 1  12  13  14  total  3035 1232 6171 846 1 1 32  1749 1267 3793 776 1329  2067 2372 5353 1 229 1751  1 277 1920 4 1 90 426 1627  683 552 261 4 340 303  1 3975 8228 28489 4168 7679  .217 .150 .217 .203 .147  . 125 . 1 54 . 133 .186 .173  . 147 .288 .188 .294 .228  .091 .233 .147 .102 .212  .049 .067 .092 .082 .039  1 .00 1 .00 1 .00 1 .00 1 .00  10  1 1  12  13  14  total  5213 2090 5126  --  5401 1 134 4609 390 634  1459 3641 4262 3324  1850 1941 3319 644 1306  528 340 2573 491 -•  23140 10582 24284 4849 1940  .225 .198  .233 .107  .063 .344  .080 .183  .023 .032  1 .00 1.00  .211  .189 .080 .327  .176 .685  .137 .133 .673  .106 .101  1 .00 1 .00 1 .00  .263 .254 .228  -  -  -  1  -  573  -  1066 1721  -  -  -  -  .135 . 1 48  1  4  555  -  1734 257  .040 .094 .065  -  —  total 10779 5890 19952 698 2046 1 .00 1 .00 1 .00 1 .00 1 .00 total  

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