UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

Downstream migratory behavior of sockeye salmon fry, with particular reference to predation Delaney, Peter Wayne 1979

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Item Metadata

Download

Media
831-UBC_1979_A6_7 D44.pdf [ 8.21MB ]
Metadata
JSON: 831-1.0094749.json
JSON-LD: 831-1.0094749-ld.json
RDF/XML (Pretty): 831-1.0094749-rdf.xml
RDF/JSON: 831-1.0094749-rdf.json
Turtle: 831-1.0094749-turtle.txt
N-Triples: 831-1.0094749-rdf-ntriples.txt
Original Record: 831-1.0094749-source.json
Full Text
831-1.0094749-fulltext.txt
Citation
831-1.0094749.ris

Full Text

DOWNSTBEAM  MIGEATOBY BEHAVIOE  OF SOCKEYE  WITH PAETICOLAB EEFEEENCE TO  SALMON FE  PEEBATION  by PETEE WAYNE B.Sc,  University  A THESIS SUBMITTED  DELANEY  of British  Columbia,  IN PABTIAL FULFILLMENT OF  THE EEQUIEEMENTS  FOE THE DEGEEE OF  MASTEE OF  SCIENCE  in THE FACULTY  OF GEADUATE STUDIES  (Dept. o f Z o o l o g y )  We  accept t h i s to  1972  t h e s i s as conforming  the required  standard  THE UNIVEBSITY OF B E I T I S H COLUMBIA Sept,  1979  ^c) P e t e r Wayne D e l a n e y ,  1979  :-6  In p r e s e n t i n g  t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f the r e q u i r e m e n t s f o r  an advanced d e g r e e a t the U n i v e r s i t y of B r i t i s h C o l u m b i a , I a g r e e t h a t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r agree that permission  f o r extensive copying of t h i s t h e s i s  f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by the Head of my Department o r by h i s r e p r e s e n t a t i v e s .  It i s understood that copying or p u b l i c a t i o n  o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my written  permission.  Department The U n i v e r s i t y of B r i t i s h Columbia 2075 Wesbrook P l a c e V a n c o u v e r , Canada V6T 1W5  BP  7 5-5  1 IE  ABSTBACT  V a r i o u s a s p e c t s o f downstream salmon f r y ( p n c o r h y n c h u s n e r k a ) field  and  Department Fulton was  laboratory  gairdneri) with  with  and  the  on  the  emerging  and  recently  the  number  nightly the move  of  and  by  rainbow  The  to fry  migrating predators;  respond  (4)  emergent  migratory  behavior,  migrants) ,  while  migrants) ; displayed migrants;  tends  predators  (5)  and  tend  migrate  migra t o r y  with  light  behavior  patterns.  downstream  movement  behavior.  Begardless  is  are  not  all  compared  fry  similar  of the b a s i s f o r the  the  f r y tend  (3)  spec i e s  to fry  of  in  their  rapidly  (fast  similar  slower  concluded  (1)  predator-sp e c i f i c ,  (slow  rate  prior experience  and  of  modifies  period;  migrate a  :  throughout  time  newly-emerged  treatment It  showed t h a t  t h a t the  a l l  to at  behavior  different  not  fry  studies  cf predators  be  to  behavior  s u b je c t i n g  (6) of  are  s ome  generally. fry  different  p e r i o ds  similarly fry  others  to  sockeye  the  varies  presence  not  on  results  a more c o n c e n t r a t e d  over  the  (Salmo  laboratory  movement p a t t e r n o f m i g r a t i n g f r y , such downstream  at  trout  w i l l i a m s o n i ) on  downstream  (2) t h e  work  of  Columbia,  experience.  moving  series  Babine Lake, B r i t i s h  emerged f r y . The  migration period;  response  time  field  e f f e c t s of exposure to l i g h t  fry  a  sockeye  Service)  predation  various kinds of p r i o r  in  of  Marine  w h i t e f i s h (Prosopium  focused  The  (Fisheries  spawning c h a n n e l s ,  concerned  behavior  were e x a m i n e d  experiments.  of t i e Environment  Eiver  migratory  pr e d a t o r s  to  a nd  to f a s t  slow  f r y to inc reasing densities,  alters  that  fry  com mencing  in  their  mi g r a t o r y  behavioral dif ference  iii  between f a s t modifies  and  premigratory  and  slow  migrants,  e n h a n c e s downstream fry  to  behavioral interactions  light,  the  presence  of  predators  movement. F u r t h e r , s u b j e c t i n g  alters  between f r y .  downstream  movement  and  iv  TABLE OF CONTENTS  Abstract  i i  T a t l e Of C o n t e n t s  ....  .... i v  list  Of F i g u r e s  vi  List  Of T a b l e s  ix  List  Of  Appendices  acknowledgements  x ..........................................xii i  IN TIC DUCT ION SECTION  ..  A: EFFECT  OF  PBEDATOES AND  EXPERIENCE CN  MIGBATOEY  EEHAVIOB M a t e r i a l s And Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Experiment  1 - Effect  Of D i f f e r e n t  Predators  1  On  3 3  Naive  Fry  11  Methods And B e s u l t s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  11  Discussion  15  Experiment  2  -  Effect  Of  Different  Predators  Cn  Experienced  F r y .....................................  16  Methods And  Besults  16  ii  D i s c u s s i o n ............ . ... . . . . . . . . . . . . . . . . . . . . . . . . . . . Experiment Fry  3 - Comparison  Of E x p e r i e n c e d  And  22  Enumerated  ..................................................  26  Methods And B e s u l t s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  26  Biscussion  28  SECTION E: EFFECT Material  And  OF  PBEEATOF.S AND  Methods  LIGHT ON  EMEEGING FEY  ..  30 30  V  Experiment  1 - E f f e c t Of D i f f e r e n t P r e d a t o r s  Fry  ..  Methods And R e s u l t s Discussion Experiment  On,Emerging  ...........  35  .,  35  ...........................................  2  43  - E f f e c t Of D i f f e r e n t P r e d a t o r s And L i g h t  On E m e r g i n g F r y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  48  Methods And R e s u l t s '  48  Discussion  54  ..........................................  e SECTION' C: DEVELOPMENT Experiment  OF SCHOOLING BEHAVIOE  1 - F r y E e s p o n s e To L i g h t  Material  57  ...................  57  And Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  57  Results Experiment  58 2 - E f f e c t Of  Schooling  And  Fry  Densities  On  ..................................  64  And Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  64  Eesults  .............................................  67  Discussion  .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  72  Material  Behavior  Light  DISCUSSION  , . 76  A. Summary Of E e s u l t s  ..................................  76  E.  Of Sockeye F r y . . . . . . . . . . . . . . . . . . .  76  Early L i f e  History  C. S i g n i f i c a n c e E.  Migrants  Of P r e d a t i o n  ...........................  And N o n - m i g r a n t s  79 81  E. The E f f e c t Of E x p e r i e n c e . . . . . . . . . . . . . . . . . . . . . . . . . . . .  85  F. The S i g n i f i c a n c e  89  Of S c h o o l i n g  .......................  LITEEATUEE CITED  92  Appendices  96  ................................................  vi  L I S T OF  FIGDEE 1a:  Geographic Eiver.  FIGUEE  FIGURES  location  of  B a b i n e Lake  and F u l t o n  .........................................  1b:  Fulton Eiver  FIGURE 2a:  Experimental  and a d j a c e n t s p a w n i n g stream  channels.  channels;  ...  4 5  looking  downstream. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  6  FIGURE 2b:  Experimental  6  FIGUEE  Prey-trap at downstream end of experimental channels. ...................................... Migration patterns of inexperienced f r y within experimental stream channels. Calculated frcm t h e t o t a l number o f f r y c o m p l e t i n g m i g r a t i o n . .. Migration patterns of inexperienced f r y within experimental stream channels. Calculated frcm the total number of fry starting each experiment. ....................................  3:  FIGUEE 4: FIGURE 5:  stream  channels;  l o o k i n g upstream.  7 12  12  FIGURE 6:  Migration patterns of various 'experienced* f r y w i t h i n experimental stream channels; S e c t i o n A, E x p e r i m e n t 2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... 19  FIGUEE 7:  The effect of predator presence/absence on p e r c e n t m i g r a n t s f o r e a c h s p a w n i n g c h a n n e l . . . . . 19  FIGUEE 8:  Summary o f r e s u l t s f o r p e r c e n t m i g r a n t s first hour; showing response cf •experienced' f r y i n d i f f e r e n t predator ,  FIGURE 9:  FIGURE 10:  in the various regimes. 24  Migration patterns of various 'experienced' f r y within e x p e r i m e n t a l s t r e a m c h a n n e l s ; S e c t i o n A, E x p e r i m e n t 3. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Experimental  stream  channels  d u r i n g second  field  season. FIGURE 11a: P r e y - t r a p FIGUEE  24  31 a t downstream  end. . . . . . . . . . . . . . . . . . . .  11b: P r e y - t r a p i n p o s i t i o n .  FIGURE 12:  P r e y - t r a p with  FIGURE 13:  I n c l i n e d - p l a n e t r a p used fry.  32 32  f r y , on c o u n t i n g t a b l e . for collecting  ..........  33  emergent 33  vii  FIGURE  14:  Mean migration pattern f o r naive f r y within a r t i f i c i a l S e c t i o n B, E x p e r i m e n t 1.  experimental and stream c h a n n e l s ; 36  FIGURE  15:  Migration patterns of experimental f r y within artificial stream channels. Plot of 'time p e r i o d s x f r y t y p e ' i n t e r a c t i o n . A p p e n d i x V. ... 36  FIGURE  16:  Migration patterns.of experimental f r y within artificial stream channels, Plct of 'time p e r i o d s x p r e d a t o r s ' i n t e r a c t i o n . A p p e n d i x V. .. 36  FIGURE  17:  Migration patterns of (a) f a s t migrants, (t) slow migrants, and (c) e x p e r i e n c e d f r y w i t h i n a r t i f i c i a l s t r e a m c h a n n e l s . . . . . . . . . . . . . . . . . . . . . 38  FIGURE  18:  Migration patterns of experimental f r y within artificial stream channels containing (a) rainbow trout, (b) whitefish, (c) rainbow/whitefish combination, and (d) no p r e d a t o r s . .................................. . . . 4 0  FIGURE 19:  Means o f p e r c e n t migrants i n the f i r s t hour plotted for (a) fast migrants, (b) slow migrants, and (c) e x p e r i e n c e d f r y migrating w i t h i n each o f f o u r p r e d a t o r r e g i m e s . . . . . . . . . . . 41  FIGURE 20:  Means of percent migrants i n the f i r s t hour plotted f o r experimental f r y migrating within artificial stream channels containing (a) rainbow trout, (b) whitefish, (c) rainbo-w/whitef i s h combination, and (d) no p r e d a t o r s . . . . , , . . . . . , , . , . . . . . . . , . . . , , , . . . . . , . . . . . . 42  FIGURE 21:  Means o f p e r c e n t (a) m i g r a n t s , (b) n o n - m i g r a n t s , and (c) l o s t or eaten f o r experimental f r y migrating through artificial stream channels w i t h and w i t h o u t p r e d a t o r s . . . . . . . . . . . . . . . . . . . . . 44  FIGURE 22:  Mean migration pattern of experimental f r y ; S e c t i o n B, E x p e r i m e n t 2. . . . . . . . . . . . . . . . . . . . . . . . 50  FIGURE 23:  Migration patterns c f experimental f r y within artificial stream channels. Plct cf 'time p e r i o d s x f r y t y p e ' i n t e r a c t i o n , A p p e n d i x X. ... 50  FIGURE 24:  Migration patterns of experimental f r y within artificial stream channels. Plot of 'time p e r i o d s x p r e d a t o r s ' i n t e r a c t i o n , A p p e n d i x X. ... 50  FIGURE 25:  Migration patterns of (a) f a s t m i g r a n t s , (t) slow migrants, and (c) e x p e r i e n c e d fry in artificial stream channels with and without p r e d a t o r s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51  viii  FIGOEE 26:  Percent migrants w i t h i n the f i r s t , hour f o r experimental f r y , subjected to l i g h t treatments, migrating tnrough artificial stream channels w i t h and w i t h o u t p r e d a t o r s . . . . . . . . . . . . . . . . . . . . . 53  FIGURE 27:  Percent migrants f o r experimental f r y , subjected to light treatments, migrating through artificial stream channels with and without p r e d a t o r s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53  FIGURE 28:  Percent non-migrants for experimental fry, subjected to l i g h t treatments, migrating through artificial stream channels with and without p r e d a t o r s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53  FIGURE 29:  P l o t o f SWIMMING TIME f o r f r y s u b j e c t e d t o l i g h t t r e a t m e n t s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62  FIGURE 30:  P l o t o f RESTING TIME f o r f r y s u b j e c t e d t c light treatments,. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62  FIGURE 31:  P l o t o f SCHOOL LENGTH f o r f r y s u b j e c t e d t c l i g h t t r e a t m e n t s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63  FIGUEE 32:  Plot light  FIGUEE 33:  Experimental tank;  FIGURE  34:  Experimental tank containing f r y ; Section C, E x p e r i m e n t 2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66  FIGURE  35:  Distance moved by one f i s h , i n a g r c u p o f t e n f r y , d u r i n g a s e v e n h o u r ' o b s e r v a t i o n p e r i o d . . . . 68  FIGURE 36:  Movement p a t t e r n s o f d a r k - a d a p t e d f r y s u b j e c t e d t o v a r i n g t i m e p e r i o d s o f l i g h t t r e a t m e n t . ..... .71  FIGURE 37:  Summary of results l i g h t treatments.  of SCHOOL treatments.  FORMATION f o r f r y s u b j e c t e d t o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 S e c t i o n C, E x p e r i m e n t  2. . . . . 66  f o r f r y subjected to various 73  ix  L I S T OF TABLES  TABLE I :  Summary o f r e s u l t s f o r m i g r a n t s , non-migrants, e a t e n , and m i g r a n t s i n t h e f i r s t 1.5 h o u r , i n an experiment testing naive f r y migrating through artificial stream channels with and without p r e d a t o r s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14  TABLE I I :  P e r c e n t migrants i n t h e f i r s t hour f o l l o w i n g the onset o f m i g r a t i o n . Experiment t e s t e d f i v e f r y t y p e s , each from two s o u r c e s , and a l l e x p o s e d t o t h r e e p r e d a t o r c o n d i t i o n s . . . . . . . . . . . . . . . . . . . . . . 20  TAELE I I I :  Percent migrants, non-migrants, and e a t e n , of various 'experienced f r y , each frcm two s o u r c e s , and a l l s u b j e c t e d t o three different p r e d a t o r r e g i m e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 1  TAELE I V :  Description o f ' e x p e r i e n c e d ' f r y were s u b j e c t e d t o p r i o r t o e x a m i n i n g m i g r a t o r y b e h a v i o r . . . . . . . 27  TAELE V:  The p e r c e n t a g e o f through a r t i f i c i a l  TABLE V I :  Expectations and r e s u l t s f o r e x p e r i m e n t testing f a s t m i g r a n t s , slow m i g r a n t s and e x p e r i e n c e d f r y i n a r t i f i c i a l stream c h a n n e l s with and- w i t h o u t p r e d a t o r s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46  TABLE V I I :  Summary o f b e h a v i o r a l o b s e r v a t i o n s f o r f r y u n d e r s i m u l a t e d e x p e r i m e n t a l c o n d i t i o n s . . . . . . . . . . . . . . 59  'experienced' f r y migrating s t r e a m c h a n n e l s . . . . . . . . . . . . . 27  TAELE V I I I : Summary o f a n a l y s i s f o r f r y observed f o l l o w i n g v a r i a b l e l i g h t t r e a t m e n t s . . . . . . . . . . . . . . . . . . . . . . 60 TABLE I X :  E e s u l t s from a n a l y s i s o f v a r i a n c e and Duncan's New Multiple Eange T e s t f o r : (a) mean movement p e r 30 min observation period, and (b) t i n e taken to start swimming following dark r e c u p e r a t i o n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69  X  L I S T OF APPENDICES  APPENDIX I :  APPENDIX I I :  APPENDIX I I I :  Three-way a n a l y s i s of variance on a r c s i n e transformed values o f p e r c e n t migrants per sampling period; comparing differences i n migration patterns for naive fry in a r t i f i c i a l stream channels with and without p r e d a t o r s , ..................................  96  Four-way analysis of v a r i a n c e on a r c s i n e transformed values of percent migrants per sampling period; comparing differences i n migration pattern f o r various experienced fry, from two s p a w n i n g c h a n n e l s , w i t h and without p r e d a t o r s . . . . . . . . . . . . . . . . . . . . . . . . . . . .  97  Three-way a n a l y s i s of v a r i a n c e on a r c s i n e transformed values of percent migrants, n o n - m i g r a n t s and e a t e n ; c o m p a r i n g d i f f e r e n c e s in these values f o r various experienced f r y , from t h e two s p a w n i n g c h a n n e l s , i n a r t i f i c i a l s t r e a m c h a n n e l s w i t h and w i t h o u t p r e d a t o r s . .  98  APPENDIX IV:  Two-way analysis of v a r i a n c e on a r c s i n e transformed values of percent migrants per sampling period; comparing differences i n migration pattern f o r various experienced fry. ........... ...... 99  APPENDIX V:  Three-way analysis of variance cn a r c s i n e transformed values of percent migrants per sampling p e r i o d ; comparing m i g r a t i o n p a t t e r n s of various f r y types migrating through a r t i f i c i a l stream c h a n n e l s with and without p r e d a t o r s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100  APFENDIX V I :  Two-way analysis of variance on a r c s i n e transformed values of percent migrants per sampling period; examining effect cf p r e d a t o r s on m i g r a t i o n p a t t e r n o f t h e t h r e e f r y t y p e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101  APEENDIX V I I :  Two-way analysis of variance on a r c s i n e transformed values o f percent migrants per sampling period; examining the response of each f r y t y p e within the various predatcr r e g i m e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102  xi  APPENDIX V I I I : Two-way analysis of variance on arcsine transformed values o f percent migrants i n the first hour following the onset of the experiment; comparing differences i n these v a l u e s f o r each f r y type i n a r t i f i c i a l stream c h a n n e l s w i t h and w i t h o u t predators. An a posteriori Duncan's New M u l t i p l e Eange T e s t compared d i f f e r e n c e s i n t h e means f o r e a c h factor. « . . . 103 APPENDIX I X :  Two-way analysis of variance on arcsine transformed values of percent migrants, non-migrants and l o s t or eaten; comparing differences i n these values f o r the f r y types in artificial stream channels with and w i t h o u t p r e d a t o r s , . . ,.,,...................... .104  APPENDIX X:  Four-way analysis of variance on arcsine transformed values of percent migrants per sampling period; comparing differences i n migration pattern f o r the three f r y types, subjected to three light conditions, in a r t i f i c i a l stream c h a n n e l s with and without p r e d a t o r s . . .- . . . . . . . . . . . . . ................ ... 105  APPENDIX X I :  Three-way analysis of variance cn a r c s i n e transformed values of percent migrants i n the first hour, migrants and non-migrants; comparing d i f f e r e n c e s i n these v a l u e s f o r t h e fry types, subjected to three light c o n d i t i o n s , m i g r a t i n g through stream c h a n n e l s w i t h and w i t h o u t p r e d a t o r s . . . . . . . . . . . . . . . . . . 106  APEENDIX X I I :  Collection, fertilization and incubation p r o c e d u r e s f o r s a l m o n r o e and f r yu t i l i z e d i n 107 l a b o r a t o r y s t u d i e s . .............  ACKNOWLEDGEMENTS  I  would  larkin  like  for his  generous  t o e x p r e s s my s i n c e r e  cheerful  support  Service work.  helpful provided  I  would  Dickson f o r t h e i r special  encouragement, s t i m u l a t i n g  throughout a l l phases o f t h i s  Hoar, Dr. N.E. L i l e y provided  comments on t h i s support like  to  thank  assistance  mcral  support  E. , G i n e t z ,  i n the field  t o thank  assistance  my p a r e n t s f o r t h e i r  throughout  this  and  reviewed  and  t h e s i s . F i s h e r i e s and M a r i n e  Hahn who p r o v i d e d  f o r providing thank  guidance  and accommodation t h r o u g h o u t  whenever n e e d e d . I would a l s o l i k e  especially  t o D r . P.A. .  s t u d y . D r . W.S.  and A.Y. F e d o r e n k o c r i t i c a l l y  thanks t o Helen  associates  gratitude  study.  the  I . MacLean,  at  Fulton  advice  and E.  Eiver.  A  and a s s i s t a n c e  close  i n varicus  field  friends  and  ways. F i n a l l y ,  increditle  patience  I  and  1  INTRODUCTION  It  has  mortality 1953).  upon  In  Hunter  long  been known t h a t p r e d a t o r s  populations  a., t e n  year  (1959) e s t i m a t e d  study  variability  migrating  frcm  will  subjected  be  determine affect  Ginetz  grounds  and u n d e r s t a n d  predator-prey  The  on m i g r a t i n g due  Although  tc this  life  to  there  into  pink  pressure.  predation  of  i n t e r a c t i o n s during  this relatively  short  cycle.  S Larkin  was t h e o b s e r v a t i o n  (1976) t h a t  and  displayed  survival  rate  those  of  inexperienced  enumeration.  Newly  evidently  little  experiences  and  develop  e x p e r i m e n t s were d e s i g n e d development field  s u b s t a n t i a t e changes following fry  appropriate  t o f u r t h e r examine  and p h y s i c a l  emerged  fry  opportunity behavior.  various  to Thus  aspects  of  i n emergent f r y .  experiments in  an e x p e r i e n c e  l e a r n s from  an  changes  of p r e d a t i o n  had  had  f r y . These  disturbance  have  nerka)  and  1  patterns  an e x p e r i e n c e  during  'experienced  by  migration  presumably r e s u l t e d frcm  Initial  area  that  increased  behavioral  a l l salmonids  behavior  (Oncorhynchus  acguire  range  rearing  fry  fry  ('inexperienced')  to  Thus i t i s o f i n t e r e s t t o  t h e components  and G i n e t z  from  (Neave  i s a l a r g e s p a t i a l and  'enumerated' s o c k e y e s a l m o n  different  fry  and chum f r y ,  associated  impetus f o r t h e f o l l o w i n g study (1972)  salmcnid  i n t h e degree of p r e d a t i o n ,  spawning  phase o f t h e i r  emergent  mortality  between 23 and 86 p e r c e n t . temporal  of  i n f l i c t substantial  migratory  (Section  behavior  with a p r e d a t o r ,  a predator  A)  sought: of  (1)  emergent  to fry  t h a t i s , t o s e e what a  o r what i n f l u e n c e a p r e d a t c r h a s upon  2  a f r y t h a t i s not  eaten;  to  result  in  species  of p r e d a t o r s c o e x i s t i n g  thus  similar  (2)  migratory  different  behavior  behavior  predators  patterns  of  enumeration producing  procedure  affects  experiment  t c determine  spawning  two  stock,  differently  tc  Eesults  from  (1972)  provided  a  their  response  determine:  Due  a  from  a  how  the  inexperienced  fry,  a  f r y . The  prompted  a  potentially  close further  distinct  c o n d i t i o n s , respond  second  diversity  and  work  of emerging f r y  a dark  Only  a  and  i n the d i s c u s s i o n .  of l i g h t  to and on  environment. were  designed  to  f o l l o w i n g emergence,  and  of s c h o o l i n g b e h a v i c r .  o f s u b j e c t s i n v o l v e d , and work  of  study  ( S e c t i o n B)  the e f f e c t  ( S e c t i o n C)  initiation  has  brief  been p r e s e n t e d  the  from  laboratory  season  (2)  to light  experimental  has  field  of f r y e m e r g i n g  response  a re-examination  preliminary  predators;  order.  development o f presented  and  i n v o l v e d i n the  chronological  fry  behavioral characteristics  fry  the  experiments  fry,  laboratory studies  to the  writing,  naive  foregoing experiments,  for  to  (1)  (2) f a c t o r s  i f  work,  behavior  Finally,  of  to  differences in  evaluate  an e x p e r i e n c e d  r e a r e d under d i f f e r e n t  the  base  (1) t h e  migratory  and  subjected  experiences.  Ginetz's  examine:  sources  different  be c o n s i d e r a b l e ,  fry  to  naive,  t h a t resembles  of  can  examined t o d e t e c t any (3)  experiences  number c f  inexperienced  movement;  prcximity  various  i n a stream  downstream  a result  i f  p a t t e r n s . The  of  was  see  in this  associated  been  f o r ease  in  presented  in  introduction section.  to  the  A comprehensive  literature  survey  is  3  SECTION A: EFFECT OF PBEDATOHS AND EXPERIENCE ON MIGRATORY BEHAVIOR  Materials  and Methods  Field  work  Environment channels, mimic  was  (Fisheries  a t Babine  spawning  possibilities, cf  Ginetz  No.  2 (Eigure  predator-prey temporary  sock-like 0.6 the  x 0.4  troughs  and  (Figure control  channels  1a&b).  To  experimental  similar  m), c o n s t r u c t e d into  at  ±  traps  to a larger  the (Figure  to  those Channel  2  of  sections: 1 m  (1 mm  2  approximately  compensate  was  checked  6.1  for  remaining 3)  inch u> a s a  initial  1.2  consisted  mesh  size)  o f t h e t r o u g h s . The water d e p t h  cm/sec,  3/4  f y k e - t r a p c f dimensions  window s c r e e n i n g  top  2  with t h e upper  acclimatization;  spawning c h a n n e l s , the  area,  a r e a . The p r e y  remained  To  x 0.5  crosswise  interaction  a r r a n g e m e n t , and was meter.  Columbia  conditions  (7.3 x 0.6  divided  and  17  of t h e  2aSb).  m. M e t a l  averaged  Department  were p l a c e d w i t h i n l e g 1 c f Spawning  trap f i t t e d  front  the  S e r v i c e ) F u l t o n R i v e r spawning  three i d e n t i c a l stream  prey  prey-trapping  at  Lake, B r i t i s h  (1972),  was  and M a r i n e  channel  Each t r o u g h plywood,  conducted  30  cm.  adjusted  daily  using  The by a  set-up  was  of  a  0.9 x covered  velocity  mcvable-slit  Teledyne  f o r water c u r r e n t v a r i a t i o n s  an e x t e n s i v e w e i r  a  within the  flow a  m  Gurley  within the  constructed  at  upstream end. The  experimental  sockeye  salmon  f r y were c o l l e c t e d  from  FIGURE  l a : Geographic  location  of  Babine  Lake  and  Fulton  River.  FIGURE  lb: Fulton  River  and  adjacent  spawning  channels.  FIGUEE 2a:  A r t i f i c i a l stream c h a n n e l s ; l o c k i n g downstream. Each trough consisted cf twc sections: predator-prey interaction area and p r e y - t r a p p i n c a r e a . F l o w v e l o c i t y was adjusted hy movatle-slit arrangement at t h e upstream end.  FIGUEE 2h:  A r t i f i c i a l stream c h a n n e l s ; locking Erey-traps are not i n o p e r a t i o n .  upstream.  7  IIGDEE 3:  Prey-rtrap a t d o w n s t r e a m end o f e a c h t r c u g h . f r y were c o l l e c t e d i n t h e s o c k - ^ l i k e t r a p a t t a c h e d t o the l a r g e r f y k e - t r a p .  8  Spawning C h a n n e l s No. mean  lengths  (mm)  were 29.83 mm fry, yelk is,  30.32  For  mm  148,11  (mg)  the  during  these  were  upon  as  and t h e s e  2  f r y ; that  1972). two  basic  fry  migration  design,  types  hours  INEXPERIENCED  will  No.  f o r a l l experimental  downstream  defined  1f r y  channels,  (Ginetz,  the experimental  were r e e d e d ,  and f o r C h a n n e l  the  F o r both  experiments,  peak  period,  f o r C h a n n e l No.  mg r e s p e c t i v e l y .  was n e a r c o m p l e t e  use i n  Depending fry  and  5 o f development  collected hours);  and mean w e i g h t s  and 157.93 mg r e s p e c t i v e l y ;  absorption stage  1 and 2. F o r t h e e x p e r i m e n t a l  and  were  (2230-0130 ENOMERATED.  various ether types  be d e s c r i b e d  i n the  of  appropriate  sections. INEXPERIENCED were c a p t u r e d therefore  shortly  having  •experience*. collected  fry  For  had both  were assumed t o be n a i v e , t h a t i s , t h e y following  emergence  from  limited  opportunity  to  channels  during night-time  the  migration  hours  with  fences, t r a n s p o r t e d to the l a b i n buckets  darkened  10 g a l a q u a r i a fry  procedure  carried  the  were  and  fry  drained  u n t i l t h e next  were  collected  o u t by t h e F i s h e r i e s  captured  i n converging  o u t o f a 227 l i t r e  fry  dipnets  following staff.  the  For Channel  Curing  the  increases  counting  in light  which  fiberglass The b u c k e t  extended  procedure  intensity  fry  Ko.  t h r e a t t r a p s , washed  2,  into  tub i n t o a  waterfilled  was  transported  n e a r b y e n u m e r a t i o n room where t h e f r y were hand a trough  at the  counting  into  swept i n t o  were  evening.  polyethylene  and  an  and s t o r e d i n  12 l i t r e a  bucket.  gravel,  encounter  inexperienced  counting  ENUMERATED  the  back were  ( R i v e r : c a . 0.1  then  into  counted  the  subjected  to  river. rapid  f t - c ; Enumeration  9  rocm:  ca.  procedure  80 at  operation  ft-c) Channel  just  intensity  and  physical  No.  1  described,  fluctuations  handling.  varied  by  but t h e e s s e n t i a l  meter was  enumeration  comparison  plus p h y s i c a l handling  Gossen L u n a s i x 3 exposure  The  with  features of were t h e  the light  same.  used f o r a l l l i g h t  A  intensity  measurements. Eainbow 26-31  cm,  fork  length  trout  and  (Salmo  Mountain size  by  beach  downstream  of  the  for  t h r e e days  t o one  hours p r i o r  t o an  of  28-31  seining  cm,  and  box  were u s e d  angling  to  confirm  of the a r t i f i c i a l  3  fungus  Excessive within  contained  6  algal  and  after  inspection  made f o r any  counted  out  transferred light  Piver,  fish  their  were  just stored  health;  then  t r o u g h number  which,  from frcm  48  6  rainbow  to  Care  remove  1 was  void  trout,  and  was any  taken t o that  were  sediment b u i l d - u p daily  velocity  f r y that  clearing  was  might  cn t h e s c r e e n s and in  checked  be  the  early  a  close  and  present. Approximately  t o an e x p e r i m e n t , t h e number o f f r y f o r a t r i a l  intensity  setting  Both  infections.  evening,  prior  The  whitefish. and  the t r o u g h s n e c e s s i t a t e d  1 hour  as p r e d a t o r s .  of  stream c h a n n e l s at l e a s t  studies,  standardize the predator s i z e , developing  range  experiment.  predator interaction  number  size  williamsoni),  i n Fulton  p r e d a t o r s , t r o u g h number 2 c o n t a i n e d  trcugh  length  (Prosopium  spawning c h a n n e l e x i t .  in a live  transferred  For  whitefish  range  were c a u g h t  .qairdneri) , of f o r k  the the  holding lab  r e a c h e d 0.05  the s t a r t i n g  to  aquaria the f i e l d  ft-c  the  into  a  site.  As t h e  partition  bucket,  was  was and  incident lifted,  time f o r the experiment. For a p p r o x i m a t e l y  10  4 hours, the  the prey-trapping nets  number  of  f r y which had' m i g r a t e d  the f r y so c o l l e c t e d experiment,  the  n a t u r a l channel 0800  hours  were t e r m e d  following  each total then were  and  concerning  patterns period),  numbers  avenues  of  prior to data  ( i . e . t h e number and  (2)  test  was used  (Duncan's  heterogeneous,  the  released i n t o the future  use.  At  i n the trough  percentages number  of  for  were  completing  statistical  fry  migrating  relative  Multiple  processed  within  each  of  time  migrants,  analysis  of  posteriori  I f t h e v a r i a n c e s were  with  ( K r u s k a l - W a l l i s Test) .  There  (1) t h e m i g r a t i o n  parametric  Range T e s t ) .  the  migration,  analysis.  proportions  cases  either  a p p r o p r i a t e , f o l l o w e d by an a  t h e d a t a were  a n a l y s i s of variance  channel; upcn  unaccounted  interpretation:  of  the  where New  for  o f f r y m i g r a t i n g downstream  to  n o n - m i g r a n t s and e a t e n . I n b o t h variance  those  migrating, or, t o t a l  arcsine transformed two  hour  EATEN.  h a l f - h o u r , were c o n v e r t e d number  were e i t h e r  morning, f r y remaining  (NON-MIGRANTS),  data  1/2  Depending  or returned t o the aquaria f o r  assumed t o have been The  each  down t h e s t r e a m  MIGRANTS.  f r y once c o u n t e d  the  were c o u n t e d  were c h e c k e d  a  non-parametric  11  i t J, - E f f e c t  Methods  initial  inexperienced  and  Predators  further,  effect.  experiment t e s t e d whether t h e b e h a v i o r f r y was  different  whether d i f f e r e n t  For three  introduced  nightly  into  each  when p r e d a t o r s  types  trials,  percent  migrants  the time  significant,  period whereas  ( F i g u r e 4)  migration For  the  transformed  per h a l f - h o u r time  (sequence o f t h r e e effect  on  trials) was  rate,  c h a r a c t e r i z e d by  9 runs  lack  o f s i g n i f i c a n c e o f the ' n i g h t s ' f a c t o r abiotic  downstream  changes  in  and/or  duration  of the experiment  early  peak  of  measured was mean  the  by t h e ' p r e d a t o r  significant, migration  happened,  predators  x time'  and F i g u r e 4, patterns  when p r e d a t o r s  the  factors  for  timing  1.5  over the  the  were p r e s e n t  there  the  75%  of  hours.  The  that  any  the period of results. that  This  the  interaction  the transformed  three  period  of t h e m i g r a t i o n i s  interaction.  plotting  be  effect  in  than  suggested  were n o t a f f e c t i n g  on  to  cr gradual d e c l i n e .  From t h e d e s i g n o f t h e e x p e r i m e n t i t i s a p p a r e n t effect  shewed  The t i m e  within the f i r s t  biotic  I)  factor  i n the e x p e r i m e n t , more  f r y had m i g r a t e d  concerning  (Appendix  by a r a p i d  the  predators,  and t h e n i g h t s  an  were  predator-free.  pattern)  effect  fry  data  were n o t s i g n i f i c a n t .  followed either  6 of the t o t a l  periods  ( i . e . migration the predator  had t h e same  twc w i t h  and t h e t h i r d  pattern present;  inexperienced  of the t h r e e troughs;  Analysis of variance  were  of predators  75  r a i n b o w and w h i t e f i s h r e s p e c t i v e l y ,  only  on N a i v e F r y  And E e s u l t s  An of  of D i f f e r e n t  percent  n i g h t s , , shows  what  was  peak  a migration  12  3  •  •  no  •  •  rainbow t r o u t  A  A  4 0.5 h r s a m p l i n g  predators  whitefish  5 periods  FIGURE 4: M i g r a t i o n p a t t e r n s o f i n e x p e r i e n c e d f r y w i t h i n e x p e r i m e n t a l stream channels. P l o t o f 'time p e r i o d s x p r e d a t o r s +/-' i n t e r a c t i o n from a n a l y s i s o f v a r i a n c e , Appendix I . Curves c a l c u l a t e d f r o m t h e t o t a l number o f f r y c o m p l e t i n g m i g r a t i o n .  0.5 h r s a m p l i n g  periods  FIGURE 5: M i g r a t i o n p a t t e r n s o f i n e x p e r i e n c e d f r y w i t h i n e x p e r i m e n t a l stream channels. C u r v e s c a l c u l a t e d f r o m t h e t o t a l number o f f r y s t a r t i n g each experiment. Shaded a r e a r e p r e s e n t s t h e d i f f e r e n c e i n p r e d a t i o n between r a i n b o w t r o u t and w h i t e f i s h .  13  within  the  predator-free over an  the  first  hour  trough  t h e r e was  between  predator  greater first  Duncan's New  w i t h i n the  difference The  a more u n i f o r m  containing (largely  first  troughs  the  pattern  Multiple 1.5  were  eaten.  within  hours  p r e d a t o r s . Due  to  to  the  the  troughs of  not  variance of  the  migrants  e t h e r two,  be  trough  (Table I ) .  of  in  a  the  null  hypothesis  difference  heterogeneous  in  those  variance, on  trial the  non-parametric resulted  the trough  in  means f o r  M u l t i p l e Range  Test  trough  i s a reflection  p r o p o r t i o n s o f n o n - m i g r a n t s and e a t e n i s a doubtful indicator  fry  compare  between t h e w h i t e f i s h  (Table I ) . T h i s d i f f e r e n c e  the t e s t  a  Test)  that  fewer  of  to  Although  (Kruskal-Wallis  migrants,  trough  variance  done.  of  than  heterogeneity  were e q u a l , a Duncan's New  a significant  i n each  and  of the trough,  i n the c i r c u m s t a n c e s o f  variances.  Parametric difference  the  both h a v i n g  were  predator-free  three  could  there  non-predator  1), a parametric analysis  differing  pinpointed  shown t o be s i m i l a r ,  number  acceptance  T e s t , cn t h e p e r c e n t  further  Typically,  the  attributable  analysis  Range  compared t o t h e p r e d a t o r - f r e e t r o u g h  and  migrating  though  in  migration  I summarizes t h e number and p e r c e n t a g e s  ncn-migrants  the  migration;  hours.  Table  found  of  p r e d a t o r and p r e d a t o r - f r e e t r o u g h s  percentage  1.5  percent  onset  whole o f t h e n i g h t . A n a l y s i s o f v a r i a n c e , f o l l o w e d by  a posteriori  migrated  after  for  analysis the  whitefish  and  difference  related  of  variance  showed  percent  migrants  hetween the trough  rainbow,  60.0%  and  to the percent eaten  44.0 5?  a  significant  respectively;  per t r o u g h .  Plotting  with a the  TflEIE I :  Summary of results f o r migrants, ncn-aigraits, eaten, and m i g r a n t s i n t h e f i r s t 1.5 h o u r , i n an experiment testing naive fry rcigrating through artificial stream channels with and without predators.  r  -*i  T  |  |  | Beplicatesl \ — — I | #Migrants | #Non-migs | #Eaten | «Migs i n I 1.5hr  NO PBEDATGBS 1  2  3  -  | | | | |  —  T  |  BAINBOW  |  |  1  2  3  | | I | |  34 1 40 26  31 0 44 25  34 0 41 33  —+ 47 28 29  28 47 11  28 47 12  4  h  —  WHITEFISH  | +  1  2  3  J | | |  47 0 28 38  44 1 30 34  44 0 31 40  | —  +  1  | |  -i  | | | | | ^  | M i g r a n t s | 62.7 37. 3 37.3 | 45,.3 41.3 45.3 | 62.7 58.7 58.7 | | SNcn-migs j 37.3 62. 7 62.7 | 1.3 0 0 | 0 1.3 0 J | SJEaten | | 53.3 58.7 54.7 | 37.3 40.0 41.3 | | SMigs i n | 38.7 14.7 16.0 | 34.7 33.3 44.0 | 50.7 45.3 53. 3 | I 1.5hr | | | | !  j.  1  I I | Mean ^ M i g r a n t J E. N.M.E.T.  +  x  1  +  j.  J |  45.8  44.4  60.0  Mean ??Non-mig | E.N.M.B.T. I  54. 2  0.4  0.4  55.6  39.5  37. 3  49.8  Mean E E a t e n  |  —  Mean ?(Mig i n | 1.5hr I I.N.M.E.T. |  L_ ,  I,  23.1  ,  D.N.M.E.T. = Duncan*s New M u l t i p l e Bange T e s t ( U n d e r s c o r e d means a r e n o t significantly different.)  I I I  1  15  percent the  m i g r a t i n g f o r each time  difference  predator  in  troughs.  expresses  the (This  the percentages whereas  starting  the experiment).  graph)  is  rainbow t r o u t  migration graph  period,  cn  Figure  5  (55.6% and 39.5% e a t e n  differs  deals  with  for  between  frcm  that  The o b s e r v e d  being  (Figure 5 ) , c l e a r l y  patterns  of migrants  accounted  seemingly  period  the t o t a l  t h e degree  more  efficient  4  two which  i n each number  difference  by  the  Figure  migrated  shows  time  cf fry  (shaded  area  c f predation, the than  whitefish,  respectively).  Discussion  The  first  predators  in  experiment some  newly-emerged, is  the e f f e c t With  way  affect  inexperienced of d i f f e r e n t  predators  pattern  posed  was  the  questions:  downstream  predators from when  on t h e s e the  movement  was more a b r u p t  with  a  migration  period.  These  are  similar  Ginetz  (1972).  He  suggested  • a w a r e n e s s ' i n t h e f r y which may and  the  compactness  pattern. Clearly simply  of  peak  predators  the former i s a p o s s i b i l i t y ,  be d e a l t w i t h  f o r the second  migration  early  in  the the  t o f i n d i n g s by may  evoke  an  i n f l u e n c e the r a t e of m i g r a t i o n  c h a s e o r 'spook' t h e f r y downstream  This subject w i l l  (2) what  present  s c h o o l i n g ; hence t h e p e a k e d  compactness c f s c h o o l i n g remains  As  that  the  vere  downstream  results  and  of  fry?  troughs  predators  (1) do  migration  sockeye salmcn f r y ? ,  absent  dome-shaped;  two s i m p l e  a somewhat  migration  the predators  at a rapid dubious  rate.  may The  conjecture.  i n a later section.  guestion, although  the m i g r a t i o n  pattern  16  of t h e  f r y was  percent as  eaten  compared  predator-prey is  similar  i n the presence  was  significantly  to  whitefish  interactions  2 -  Methods And  :  behavior?,  (2)  the  in  (1)  In  are  trout  fry  P r e d a t o r s on  so i t  Experienced  of  designed  types  frcm  migration  Fry  two  pattern?,  factor  to  answer  the  of e x p e r i e n c e s a f f e c t f r y  the  spawning  and  (3)  fry  channels  are  i n f r y b e h a v i c r , and  experienced  was  these  guestions,  designed;  (inexperienced,  to  and  to  with  the  previously  the  enumerated,  whitefish);  exposed  with  t o rainbow  predator  i f so,  respond  similarly  x  3  do to  each 3  a  5  factors  x  5 fry  experienced  experienced  2 different  factorial  as f o l l o w s :  shocked,  t r o u t , and, from  2  by  spawning  by  previous channels;  c o n d i t i o n s (no p r e d a t o r s , w i t h  rainbow  whitefish).  In g e n e r a l , the  three  are l a c k i n g ,  predator species?  p r e v i o u s exposure  all  the  types  testing  exposure  (55.6%)  processes occurring.  was  different  an i n f l u e n c i n g  experiment types  the  of D i f f e r e n t  do  their  various  different  fully  the  B e h a v i o r a l observations of  w i t h i n the troughs  set of experiments  guestions  encounters  predators,  Eesults  A second  similar  Effect  icth  l a r g e r f o r rainhow t r o u t  (39.5%).  i m p o s s i b l e to understand  Experiment  of  outlined.  a d d i t i o n a l types  material  and  methods  were  B e s i d e s i n e x p e r i e n c e d and  followed  as  enumerated f r y ,  c f f r y - e x p e r i e n c e s were e s t a b l i s h e d .  One  17  type  was  SHOCKED; t h i s  12 l i t r e with  polyethylene  consisted of placing bucket  with  a number  perforated sides,  water a t a maximum f o r c e f o r 60 s e c o n d s .  carried  out  period, were  approximately  hours  before  and any f r y t h a t were damaged  or d i e d  removed.  PEEDATOES,  The  were  experimental  3  other  fry  two  which  stream  types  had  channel  of  cf f r y i n a and s p r a y i n g  This operation the  was  experimental  within  that  time  f r y , EXPERIENCED  previously  migrated  containing either  dc*n  TO an  rainbow t r o u t o r  whitefish. Since migrants reduced  EXPERIENCED in  the  fry  predator  t h e numbers  available  numbers r a n g i n g  were no o b v i o u s For  data  for  a  time  loss  frcm  60 t o 120 f r y p e r t r i a l .  of  migration.  the  downstream  t h e same f r y d e n s i t i e s  a l l values  period,  successful  predation  f o r subseguent experiments,  t r e n d s or e f f e c t s  analysis,  downstream an  from  the  troughs,  was n o t p o s s i b l e t o m a i n t a i n the  were  the  throughout,  However,  o f v a r y i n g t h e numbers  were e x p r e s s e d  as t h e  total  of  number  The a n a l y s i s  and i t  of f r y .  percentage  fry  o f v a r i a n c e was  there  starting  preceded  by  arcsine transformation. The  outlet the  pattern of migration; For those  end o f t h e e x p e r i m e n t a l  three  shewed  factors,  both  experience' former same  the  time  plus  trough,  time  periods  interaction  to  a l l time  interaction periods  term  periods implies  and  analysis  the  reached  the  of variance  with  'time  be s i g n i f i c a n t ,  for that  that  periods as the f o u r t h  i m p l i e s o n l y t h a t t h e number in  fry  periods  (Appendix  of f r y migrating a l l treatment  the  types  x  of  fry  I I ) . The  was n e t t h e  groups.  differences  was n o t t h e same f o r t h e v a r i o u s  variable,  between fry,  The time which  18  apparently  was  experienced  fry  experienced gradually was  migrants  the b e h a v i o r  number  constant  accepted  the  null  To  the o t h e r  significant  the  which r e a c h e d The  pattern  migration,  the  main  upon t h e the  fry  or  effects,  nightly  Not  are eaten.  over  the  migrants,  the  p r e d a t o r s t o have a s i g n i f i c a n t  whole n i g h t l y  trough; the  greater  (corresponding trout). cf  migrants,  fry  Range  Test  significantly  experience, predators  a l l o f the  III).  of  effect The  fry  had  no  f o r the  y i e l d e d the  predator  that  of had  had  fry the  the  data eaten,  variance  showed these  migrating than in  that for the  no  migrants,  consumed greatest  greatest  was  rainbow  s m a l l e s t percent  number  trough  the  w i t h i n each c f  i n the w h i t e f i s h trough  large  accomplish  percent  percent  i n t u r n g r e a t e r than  the  fry  n o n - m i g r a n t s and  the w h i t e f i s h trough  t h e no  of  migration. Analysis  having  to  Although  percent  (Appendix  latter  be  In  variance  Table I I I presents  percent  which was  experienced  p a t t e r n of m i g r a t i o n  for  predators,  of migration.  fry  absence  survival:  a percentage  significantly  percent  cutlet.  of  three categories  the  Multiple  to  there  period. This  a n a l y s i s of  a Duncan's New  then  types.  presence  effect  onset  whitefish  experienced  hours  f r y types,  showing  Although  whitefish  1.5  migration  II,  the  migrants.  hypothesis,  summarize, and  migrating  four other  Table  regimes,  whitefish  from  in  of the w h i t e f i s h  f o r the f i r s t  hour f o l l o w i n g t h e  the fewest  the  different  first  predator  had  of  i n numbers o v e r t h e  emphasized  i n the  usually  fry  remained  decre.ase  is  three  origins,  ( F i g u r e 6 ) . The  fry  a sharp  shewed  to  d e c l i n e d , whereas f o r t h e  difference  the  attributable  by  the  percent  percentage  19  50  0.5 h r s a m p l i n g  periods  FIGURE 6: M i g r a t i o n p a t t e r n s o f v a r i o u s ' e x p e r i e n c e d ' f r y w i t h i n experimental stream channels. P l o t o f 'time p e r i o d s x e x p e r i e n c e ' i n t e r a c t i o n from a n a l y s i s o f v a r i a n c e , A p p e n d i x I I . C u r v e s c a l c u l a t e d f r o m t h e t o t a l number of f r y s t a r t i n g each experiment.  FIGURE 7: The e f f e c t o f p r e d a t o r p r e s e n c e / a b s e n c e o n p e r c e n t m i g r a n t s f o r each spawning c h a n n e l . P l o t o f ' p r e d a t o r s +/- x c h a n n e l s ' i n t e r a c t i o n from a n a l y s i s o f v a r i a n c e . Appendix I I .  20  TABLE I I : P e r c e n t migrants i n t h e f i r s t hcur f o l l o w i n g the onset o f m i g r a t i o n . Experiment t e s t e d 5 f r y types (inexperienced, enumerated, shocked, e x p e r i e n c e d t o rainbow, experienced to whitefish), each from 2 sources, and a l l e x p o s e d t o t h r e e c c n d i t i c n s (no p r e d a t o r s , w i t h rainbow t r o u t , a n d w i t h w h i t e f i s h ) . T  INEXPEE.  ENUMEB,  r JO  '  — I  | SHOCKED | EAINBOW WEITE. EXPEE. . EXPEE.  +—  i  PREDATOR  Channel «•  1 2  80 68  63 37  80 43  73 50  67 20  1 2  54 50  45 80  48 48  60 72  48 40  68 73  57 73  88 88  68 83  62 53  BAINEOW Channel "  WHITEE3SH Channel "  1 2  Mean Variance  65.50  59. 17  65. 83  67.67  48.33  130.24  267.34  468.23  130.66  285.91  j. E,.N.M,R,.T.  D. N.M-E.T. = Duncan's New M u l t i p l e Range ( U n d e r s c o r e d means a r e n o t different.)  Test significantly  21  TAELE I I I :  (a)  Percent migrants, non-migrants, and e a t e n o f v a r i o u s ' e x p e r i e n c e d ' f r y , e a c h from two s o u r c e s , and a l l s u b j e c t e d t o three different predator regimes.  %MIGB ANTS NO PBEDATOB Channels  I  — .  Inexperience d Enumerated Shocked Bainbow E x p e r . W h i t e f i s h Exper.  BAINBOW —|  90.0 83.3 91.7 83. 3 91.7  7 5. 0 51.7 76.7 83.3 5 5.0  —_ _ _ _ _ _  j| 57.8 51.7 | 50.0 | 65.0 | 68.3  £5.0 86.7 75.0 80.0 63.3  I WHITEFISH I I 85. 0 80. 0 I 68. 86. 7 I 95. 03 96. I 86. 7 100. 03 I 81. 7 90. 0 I _ _ _ _ _  —  r  MEAN  78. 17  D. N.M.E.T.  **  68.28  A l lSignificantly  87. 17 Different  H  **  (b) %NON-MIGEANTS  I I I I  Inexperienced 10.0 Enumerated 8.3 Shocked 8.3 Eainbow E x p e r . 48.3 W h i t e f i s h E x p e r . | 16.7 MEAN  -+  16.7 16.7 2 5.0 23.3 45.0  |  3.9 5.0 | 13.3 | 0. 0 | 5.0  I  2. 5 0. 0 3. 3. 1. 7  |  2. 5  I 3 .3 1. 7 3 I0. 0 3| | 3.3  0. 0 3. 3 0,.0 0. 0 1. 7  L_.  21.83  3. 80  1.58  B.N.M.E.T. (C)  %EATEN  Inexperienced Enumerated Shocked Eainbow E x p e r . W h i t e f i s h Exper.  | | | | |  MEAN  |  I 38.3 12.5 | 12.5 20. 0 I 43.3 13.3 | 28.3 10.0 I 36.7 21-7 | 3.3 1.7 1 35.0 16.7 | 13.3 0i 0 1 26.7 35• 0 |15.0 £.3 27.92 11.24  D.N.M.E.T. |  Significantly  36.00 19.84 14.48 8.00  CHANNEL MEANS | D.N.M.E.T.  Different  Chan.1  fry sig.dif.  from Chan.2 f r y  D.N.M.E.T. = Duncan's New M u l t i p l e Bange T e s t (Underscored means a r e n o t s i g n i f i c a n t l y different.)  22  of  non-migrants. C h a n n e l No. 1 f r y were e a t e n  rate  compared  'predators  Channel  x channels'  significant. channels  to  This  at  No.  2  interaction  latter  approximately  fry  (Table  for  percent  difference  between  was n o t t h e same f o r t h e t h r e e  7 ) . For both migrants  predator  was  found  channels,  a  IIIc),  the  and t h e  migrants  was  t h e two spawning  predator  higher  twice  systems  value  (Figure  of  percent  f o r C h a n n e l No. 2 f r y ; whereas t h e r e v e r s e  was t r u e f o r t h e p r e d a t c r - f r e e  trough.  Discussion  Ginetz  (1972) f o u n d  displayed  behavior  inexperienced survival  results  which  experience  and  schooling  on  behavior,  f r y react i n different  was assumed a p r i o r i  produce  similar  distinct  frcm  This  differences i n  displayed not  on  set  nightly  f r y groups,  similar due  to  of  migration the  and,  predator  the  altered  1S76).  These  of, v a r i o u s  ways  i n which  regimes. experiences  but  pattern  this  would  i n patterns failed  the  various  whitefish-experienced  small  to  t o shew c l e a r c u t  among  p a t t e r n s . Assuming being  increased  and  f r y , resulting  except  effects  from  the e f f e c t s  experiments f a i l e d  all,  different  & larkin,  pattern,  migration  fry  enumeration  a l l four  the  the inexperienced  materialize.  experienced  effects  that  enumerated  tendencies,  ( G i n e t z , 1972; G i n e t z  migratory  and were  l e d t o some s p e c u l a t i o n r e g a r d i n g  experienced  were  patterns  enhanced  patterns  experiences  It  experienced  f r y . Both  rate,  migration  that  that the  fry,  results  and t h e v a r i a b i l i t y  23  being  large,  perhaps,  one  concludes  that  the  was  also  inexperienced the  first  and,  that for  expected  that  f r y would show l i t t l e  hour,  the  percent  > Enumerated > Shocked >>  would  be  i n the  results,  first  two  Since  these  summarized  expectations expectations  pattern results  f r y had as  analysis shewed  similar  effect  observed;  first,  effect hence  upon there  non-migrants different  from  the  the were in the  Whitefish (2)  migrants  experienced  fry  that  fail  partially valid  With in  than (3)  predator  the for  there systems  1972),  t o agree with confirm  and  the  the  third.  consistent  with  i t is possible that  learning histcry  percent  migrants, in  that  the all  t h a t made them a l l  predators  n c n - f f i i g r a n t s and  preceding  significant. two  Two were  experiments;  important not  phenomena  similar  f r y . Rainbow consumed s i g n i f i c a n t l y fewer the  the  fry.  the  was  =  i n t h i s experiment r e f l e c t e d  r e s u l t s as those  predator  right),  hour  Inexperienced. percent  within  the  first  hour i n t h e  seemed  some p r e v i o u s  of  cut over  anticipated  only  (Ginetz,  experienced  pre d a t o r s ,  activity  the  i n F i g u r e 8,  and  migration  respond  are  system.  experiments  the  i t was  i n the f i r s t  previous  cf  for  no  experienced  were t h a t t h e  fry. Finally,  no-predator  in  Bainbow  greater  more m i g r a n t s  The  be:  the e x p e c t a t i o n s  inexperienced wculd be  would  migrants  experienced  hour  with  ( i . e . movement more s p r e a d  crder  predators  (1)  migration  decreasing  than  fry  experienced.  It  first  inexperienced  migrants.  Secondly,  p r e d a t o r - f r e e trough  percent  eaten  was  i n the rainbow  the  not  in  eaten the were their  more f r y , percent  significantly  trough.  24  100  90 S.  80 lex. 70  cn E  S.;R.E  60  W.E. 50  lex.  40  - inexperienced  En.  - enumerated  S.  -  shocked  R.E. - r a i n b o w .W.E.  30  no p r e d a t o r s  rainbow  Experimental  stream  experienced  - whitefish  experienced  whitefish channels  FIGURE 8: Summary o f r e s u l t s f o r p e r c e n t m i g r a n t s i n t h e f i r s t h o u r ; showing response o f v a r i o u s ' e x p e r i e n c e d ' f r y i n d i f f e r e n t predator regimes.  40 V  1  2  3  4  0.5 h r s a m p l i n g  5  •  •  •  -•  inexperienced enumerated  o  o  t u b w i t h no  A  A  tub with  a  A  stirred  6  lights  lights  7  periods  FIGURE 9: M i g r a t i o n p a t t e r n s o f v a r i o u s ' e x p e r i e n c e d ' f r y w i t h i n e x p e r i m e n t a l stream channels. P l o t o f 'time p e r i o d s x e x p e r i e n c e ' i n t e r a c t i o n from a n a l y s i s o f v a r i a n c e , Appendix IV. Curves c a l c u l a t e d f r o m t h e t o t a l number o f f r y s t a r t i n g e a c h e x p e r i m e n t .  25  the  Although  the  rearing  conditions  stocks from nor  two  potentially the  two  predators  migrants percent  and  spawning c h a n n e l s a r e  an  percent  The  eaten  that  the  number  C h a n n e l No. results, of  these  showed two of  fry  fry  may  are  a  were (page 8) be  of  the  exhibiting  result  channel  different,  of  experiment, r e s u l t i n g  differences  or, frcm  methods. which m i g h t  was  (1972)  the  fry  physical  acquired  spawning this  may  slightly  Additional  of  the  fry  experience percent  results.  regime, 7).  eaten  a greater  if  the  spanning  the  For  compared  mortality. from  be  an  different may  Both  the  two  similar.  and/or  produce t h i s v a r i a t i o n  The  which  of  collection  in  for  different  artifact  disclose  to  similar  characteristics  stocks  tests  Secondly,  behavioral t r a i t s  just  was  percent  found  c h a n n e l s were e x t r e m e l y  i n n a t e or  different  conditions,  transporting  two  yet  (Figure  Ginetz  possible  of  predator  1 fry  1 fry exhibiting  o b s e r v a t i o n s are  the  neither  interesting  a  No,  IIIc).  proximity,  'channels x p r e d a t o r s '  within  Channel  (Table  C h a n n e l No.  channels the  2 fry  of  pattern  similar,  m i g r a n t s f o r e a c h c h a n n e l were d i f f e r e n t twice  and  However, a n a l y s i s  interaction  implying  different  migration  c h a n n e l s was  effect.  migrants, the  significant;  slightly  different.  spawning had  are  in close  the and  behavioral  survival.  26  Experiment  Methods  2  And  ~ Comparison  field  that  experienced  natural this  and  fry,  to  each  two-factor  interaction patterns  experiments, followed  by  interaction different  the  the  distinctly  than  resenbling  term  procedure  tested twice migrants of  to  that  different  that from  downstream o v e r  the  upon  in  separate  were  chosen  analyzing  time  IV).  periods The  the  'tub  in  the remaining  a peak,  migrating.  migration  patterns  four  a more c o n c e n t r a t e d  and  previous  number  with  per  migration  i n c r e a s e i n numbers t o the  the  ncn-migrants.  described  various types of experience note  distinct  fry,  percent  showed t h e  those  decline  into  effect  75  examine approach  (EXPERIENCES) with  the  The  p a t t e r n s . Three  (Appendix  a rapid  showed  and  to  as  v a r i a n c e , number m i g r a t i n g  significant  displaying  attempt  investigation.  behavior  of experience,  a gradual  end,  m i g h t have some  their  similar  to  Ginetz  limited  enumeration procedure  analysis  be  f o r the  interesting  migrating  n e a r i n g an  a short-term  was  type  to  were  hy  respectively  f r y was  pattern, percent  p e r i o d and  raised  of experimental  of the enumeration  migration  time  only  altering  ( T a b l e I V ) , and  point  f r y types  which i n t h e m s e l v e s  divisions  A  was  divide  possibly  a  f r y have a g r e a t e r s u r v i v a l r a t e  supply  f r y migration  processes  Enumerated | r j  inexperienced f r y i n behavioral observations i n  Since the  was  examined  f r y ; t h e s e two  q u e s t i o n was  taken  test  enumerated  non-enumerated  aguaria.  and  Results  A final (1S72),  of Experienced  The were  (Figure 9). I t  is  lights'  cur\e  was  the  fry  curves; time  period.  27  TABLE IV: D e s c r i p t i o n o f ' e x p e r i e n c e s ' f r y were s u b j e c t e d p r i o r to exaining migratory behavior.  EXPERIENCE  to  DESCRIPTION  1. I n e x p e r i e n c e d 2. E n u m e r a t e d  As I  3. T u t w i t h NO LIGHTS  previously described within  text.  As p r e v i o u s l y d e s c r i b e d w i t h i n  text.  Caught i n c o n v e r g i n g t h r o a t t r a p and emptied i n t o c o l l e c t i n g tub, fcithcut f l o o d l i g h t s cn.  |  4. Tub w i t h LIGHTS  C o l l e c t e d as 3 above, b u t with l i g h t s on. These f r y d i f f e r from enumerated f r y i n t h a t t h e y were not physically handled as during the normal enumeration procedure.  5. S t i r r e d  Collected similar tc inexperienced fry, p l a c e d i n a b u c k e t and s u b j e c t e d to continual stirring for approximately 2 minutes. This s i m u l a t e d c o n d i t i o n s f r y may e n c o u n t e r in the converging threat t r a p .  TAELE V: The p e r c e n t a g e of 'experienced' f r y migrating through artificial stream channels. Experiences as d e s c r i b e d i n T a b l e I V .  Inexper. JfMigrant  |  Hand i n tucket  Tub w i t h NO LIGHT  44.6 46. 9  45. 4 46.2  62. 5 38.5  Mean  45.75  45.80  50.50  Variance  2.65  0.32  287.99  -f  T  T  Enumer. 54. 8 49. 2 52.00 15. 68  | Tut with LIGHT 75.0 53. 1 64.05 239.87  | £.N.M.E.T. | D.N.M.R.T. = Duncan's New (Underscored different.)  M u l t i p l e Range means a r e n o t  Test significantly  28  A  similar  percent  non-migrants  heterogeneity means  parametric  cf  vas  the  Bange  Test,  Table  invalid  found  the  an a p o s t e r i o r i  test,  Duncan*s  different  V, b u t s i m i l a r  the from  'tub  with  migrants  and  due  the  to  the experience  non-parametric  test)  showed  significantly  statistically  a further  (Kruskal-Wallis  non-significant,  on t h e p e r c e n t  v a r i a n c e s a s s o c i a t e d with  (Table V ) . Although  variance  analysis  analysis  mean  values New  lights'  the three experiences  of  t o be  Multiple experience  indicated  on  t o t h a t o f t h e enumerated f r y .  Discussion  Ginetz  (1972)  someway a l t e r e d  the naive  fry; the resultant an  experience.  have an e f f e c t  found  that  the  behavior  f r y most c l o s e l y  The q u e s t i o n t h e n c e  enumeration  patterns  of  resembled  those  of  the  procedure,  newly  emerged  encountering  a r o s e , how d o e s  upon t h e f r y , and i n p a r t i c u l a r  what component  procedure i n  enumeration  a t what s t a g e , o r  has  the  most  both  enhanced  significant  effect? Enumeration tendencies survival  and an ' e x p e r i e n c e '  noted  in  i n nightly  aquarium  trough  experiments  component o f t h e e n u m e r a t i o n changing  fry  schooling-survival  procedure  behavior, and  observations,  possible  (Ginetz, proved  f r y schccling and  increased  1972).  If  one  t o be i m p o r t a n t i n i  relationships  enumeration-experience  between  could  be  postulated. Although failed  to  the experiment  show  anything  was o n l y  conclusive,  a brief  i n v e s t i g a t i o n and  i t did  result  in  an  29  interesting the  fry  p o i n t . The o n l y  was  'tub w i t h  the tub without behavior the  lights  compared  lights  are a  lights'. displayed  w h i c h had some e f f e c t  Since no  similar  upon  f r y collected in  difference  in  migratory  to the o t h e r f r y ' t y p e s ' , i t i s p o s s i b l e t h a t significant  p h r a s e d : does l i g h t , behavior  component  either  of i n e x p e r i e n c e d  factor.  artificial  fry?  So  a  new  or n a t u r a l ,  guesticn  is  i n f l u e n c e the  30  SECTION B : EFFECT OI  Material  and  The troughs  It  site  ( F i g u r e 10)  was  was  as noted  i n the  w i t h m o d i f i c a t i o n s added t o t h e whether  the  movement o f t h e  original  hindered  the  improve  upon t h e method, open b a s k e t s 11a&b) were u s e d ,  approximately baskets depth  0.33  m.  maintained  f r y i n t o the  reducing At  were removed and was  EMEBGING  FEY  foregoing section.  were c o n s t r u c t e d s i m i l a r  questioned  (Figure  LIGHT ON  Methods  field  described,  PBEBATOBS AND  each  fish  half  counted  a t 30  ± 2 cm  sockeye  fry,  and  t c these p r e v i o u s l y prey-trapping  area.  prey-trapping  design  collecting  with s l i d i n g  the  socks. front  prey-trapping hour s a m p l i n g  ( F i g u r e 12) the  Six  To  panels  area  to  period a l l  .  The  water  flew v e l o c i t y  at  14  ±  2 cm/sec. Newly emerged length: during  30.05  mm;  average  m i g r a t i o n hours  situated  in  trap  operational  was  Captured lar,  and  fry  were  between  naive  collected  fry  did  migrate migrant  downstream fry  fry, were  (average collected  inclined-plane No.  2  trap  ( F i g u r e 13).  hours  each  The  evening.  with d i p - n e t s , t r a n s p o r t e d to  the  10 g a l a q u a r i a .  which  migration period;  fast  an  0030-0130  types of experimental -  of  NAIVE  153.40 mg),  13 of Spawning C h a n n e l  nightly not  weight:  w i t h t h e use  s t o r e d i n darkened  Three MIGEANTS  leg  termed  fry had  were  utilized:  FAST  m i g r a t e d - downstream o v e r  (2) SLOW MIGEANTS d u r i n g the  which  (1)  migrated  naive  n i g h t ; and  (3)  fry  the  which  EXPEBIENCED  down a p r e d a t o r  channel.  31  IIGDEI  10:  A r t i f i c i a l stream c h a n n e l s d u r i n g second field season. Channels were similar to those previously described, except f o r prey-traps located at downstream end of troughs, l i v e - b o x e s f o r p r e d a t o r s a r e shewn a t f a r r i g h t of photo.  32  FIGUEE 11a:  Prey-trap. Wire screen prevented l e a v i n g t i e stream channels.  predators  FIGUEE 111:  Prey-trap i n p o s i t i o n . Wood panel, at r i g h t of trap, was used t o hold f r y i n the baskets and channels while sampling at each h a l f hour interval.  33  FIGUEE 1 3 :  Inclined-plane emergent f r y .  trap  used  for  collecting  34  Occasionally  a second  CHANNEL EXPERIENCED. at  the  outlet  at  least  24-30  adjacent the the  troughs,  replaced,.  Mountain w h i t e f i s h , f o r k l e n g t h  the  to  experiments,  size  half  hour  or eaten.  infections  as  a  test  Duncan's  New  trial due  to lack  held of  cf  the  percent  frcm  availability  possibly  Test,  was  of t o t a l runs  certain  fry  non-migrants,  and  by used  percent  lest  problems predator  an  a  or  with  feeding  posteriori  on a l l d a t a . A l l  starting,  t h e number  at times  of  were  normal  of  and  which  reduce  a t 60, a l t h o u g h  snout  numbers  predation;  F o r most e x p e r i m e n t a l  m o r t a l i t y was  to that d e s c r i b e d i n  periods,  followed  Range  were  o f the t r o u g h s .  of  were c o r r e c t e d t o p e r c e n t  was  predator  cf infection;  consisted  to  enclosures  infections  Analysis of the l a t t e r ,  Analysis of variance,  transformed.  large  Typically  sign  sampling  behavior.  ccunts  high  migrants,  tended  Multiple  in  was s i m i l a r  Data  size  of Spawning C h a n n e l  fungus  with t h e i n s i d e  percent  was n o t v a l i d  first  procedure  size  s t a n d a r d i z e d w i t h i n and between  developing  the  section.  the  lost  fungus  was  length  were a c c l i m a t i z e d 48 h o u r s i n  i n rainbow t r o u t .  contact  per  calculated percent  stored  Predators  fish  experimental  migrating  fork  were s e i n e d a t t h e o u t l e t  and any  foregoing  eaten,  and  f i n displayed  The  the g r a v e l  cm;  Predator  from  emerged f r c m  trout,  F o r undetermined r e a s o n s ,  resulting  large schools  (Rainbow  suffered, especially dorsal  from  termed  to capture.  to the troughs.  troughs.  used,  predators  r a n g e o f 25-31 cm) No. 2, and p r i o r  f r y was  T h e s e f r y were c o l l e c t e d  1-2 d a y s p r i o r  of  of experienced  o f C h a n n e l No. 2, h a v i n g  Experimental range  type  then of  t h e number types,  arcsine fry  was  the  per  varied numbers  35  ranging  from  45  to  150 f r y . However,  shewed no c o r r e l a t i o n the  measured  find  To  of D i f f e r e n t  i f FAST,  Predators  cn E m e r g i n g F r y  SLOW and EXPERIENCED f r y a r e d i f f e r e n t i n  migratory  behavior,  3 x 4  factorial  experiment, with  three  rainbow  of  Eesults  their  experienced  and any  results.  test  follows:  c f the data  between i n c r e a s i n g t h e numbers  Experiment J - E f f e c t  Methods  observation  fry  plus response  types  3 replications,  (fast  f r y ) t e s t e d i n each  trout,  6  whitefish,  to various predators,  migrants,  of 3  four  was  designed  slow m i g r a n t s ,  predator  rainbow/3  regimes  whitefish,  a as and (6  and no  predators). Analysis of variance data of.  (Appendix fry  V) showed  migrating  migrating  decreased  comparison,  the  on  graph;  t h e same  sampling  over  the migration  migration fewer  Significance  fry  interaction  differing migrating migrating; (Figure  from  that  i n the e a r l y and  a significant  per  evening. type'  on t h e t r a n s f o r m e d  (2)  was  period  fry  a l s o found  fry  evening,  with  periods  16), the average p a t t e r n  for  number  (Figure  percent of  fry  14).  For  f r y i s also plotted  migrated  early  in  the  f o r : (1) "time p e r i o d s x  the  other  'time  the  p a t t e r n of naive naive  pattern  d i f f e r e n c e i n the  period;  (Figure 15), cf  migration  slow types,  migrants  especially,  a , greater x the  pattern  predators'  percent  fewer not  interaction,  pre d a t or--free  trough  1  2  3  4  0.5 h r s a m p l i n g  FIGURE  6  N.M.  14: Mean m i g r a t i o n p a t t e r n f o r e x p e r i m e n t a l a n d n a i v e f r y within a r t i f i c i a l stream channels. P l o t o f 'time p e r i o d s ' f a c t o r f r o m a n a l y s i s o f v a r i a n c e , A p p e n d i x V.  1  2  3 0.S  FIGURE  5 periods  4  hr s a m p l i n g  5  6  N.M.  periods  15: M i g r a t i o n p a t t e r n s o f e x p e r i m e n t a l f r y w i t h i n a r t i f i c i a l stream channels. P l o t o f 'time p e r i o d s x f r y t y p e ' i n t e r a c t i o n f r o m a n a l y s i s o f v a r i a n c e , A p p e n d i x V.  37  differed  from  migrating  throughout  number o f A  t h a t o f the t h r e e p r e d a t o r c h a n n e l s , a g a i n the night,  and  correspondingly,  detailed  summarizes  examination  the  significant,  analysis  the  migration  period.  With  respect  the o v e r a l l  of  number  Appendices  variance  f o r each  17a-c. The  migrants  fast  (Appendix  m i g r a n t s and  greater  number c f f r y n o t  tended  to  Appendix  behave  VI  fry  significantly displayed  from  almost  the  whereas t h e s l o w  reaction  to the  the  remaining  graphs i n  yet the  pattern,  the  the  with a  experienced  fry  t h e c h a n n e l s , as n o t e d i n  term)  and  three.  patterns  migrants  in  were more  percentage  was  significantly  of  the  each o f t h e  p r e d a t o r systems.  the  -  noted  Figure  17c;  the  in the predator-free trough departing  analysis  p r e d a t o r t y p e s shows how  VII  the  the  The  fast  migrants  the  four  preeator  erratic  in  their  various predator types.  Expressing  different  within  interaction  identical  regimes;  migration  m i g r a t i n g . Only  pattern  from  was  over  d e c r e a s e i n numbers,  gradual  differently  (significant  experienced  a  effect  interaction  i s clear  VII  o f t h e f r y and  experienced f r y display  pattern of a r a p i d display  shows  and  declined  significant V)  VI  pattern  of migrants r a p i d l y  t o f r y type, the  analysis  same g e n e r a l i z e d slew  large  of the experimental f a c t o r s  p r e d a t o r t y p e s . In a l l c a s e s the m i g r a t i o n  Figure  a  non-migrants.  the reasons f o r the f o r e g o i n g r e s u l t s .  in  fewer  Except  data  fry  types  f o r the r a i n b o w  o f f r y m i g r a t i n g downstream different  significant  f o r each  interaction  in  terms react  term).  In  in  trout  the  trough,  per s a m p l i n g  of the f r y t y p e s  c f the  period  (Appendix  the t h r e e p r e d a t o r  (a) FAST MIGRANTS  0.5  hr  sampling  p e r i o d s  (C) EXPERIENCED  0.5 h r samp1ing  periods  M i g r a t i o n p a t t e r n s o f (a) FAST MIGRANT, (b) SLOW MIGRANT, and (c) EXPERIENCED f r y w i t h i n a r t i f i c i a l stream c h a n n e l s P l o t o f 'time p e r i o d s x p r e d a t o r s ' i n t e r a c t i o n from a n a l y s i s o f v a r i a n c e , Appendix V l a - c .  39  systems were  (Figures 18a-c), the experienced and  similar,  within  the  whereas  early  non-migrants.  In  the  evening the  slow  the  migration  of  a  the  types  naive  s i m i l a r to that of the f a s t Since  the  major  analysis  the  of  the  percent  substantiated  the  migration  variance  (Appendix  both the f r y  and  were  similar,  f r y . In the two  observed  in  hour  of  migrants  in  this  pattern  the the  results.  experiment,  predator  type.  A  Analysis  subseguent  Duncan's  New  compared to the f a s t migrant and experienced f r y ; and significantly  to those i n the three p r e d a t o r troughs.  showed  the  slow  migrants,  and  predator troughs f o r (Figures 19 and  than 70% o f the experienced and  predator troughs migrated the  percent  downstream  slow migrants  f a s t migrants-experienced  fry in  20),  f r y m i g r a t i n g down the  p r e d a t o r - f r e e trough to be d i f f e r e n t compared t o the  whereas  of  migrants t c be s i g n i f i c a n t l y  the percent migrants w i t h i n the f i r s t hour  Greater  further  VIII) d i s c l o s e d a s i g n i f i c a n t d i f f e r e n c e f o r  Separate a n a l y s i s of the f r y types and  again  fry  migration  period  the migrants w i t h i n the p r e d a t o r - f r e e trough t o be different  and cases  experienced  first  M u l t i p l e Bange Test showed the slow different  of  migrants and experienced f r y .  difference  p a t t e r n s occurred w i t h i n  migrants  (Figure 18d), the  examined, the m i g r a t i o n p a t t e r n of the channel was  fry  percentage  predators  fry  migrant  had fewer  greater  of  m i g r a t i o n p a t t e r n s of the three resembled  migrants  and  absence  fast  remainder.  f a s t migrant f r y i n the  within  the  first  i n a l l troughs, and  predator-free  troughs  hour; percent ranged  between 40-70%. Analysis  of the percentage  migrants, non-migrants and  lost  ( a ) RAINBOW TROUT 70  (b) WHITEFISH  I  70  1  2  4  3 0.5  h r sampling  5 p e r i o d s  6  N.M.  I»  1  2  3  4 0.5 h r s a m p l i n g  5  6  N.M.  periods  FIGURE 18: M i g r a t i o n p a t t e r n s o f e x p e r i m e n t a l f r y w i t h i n a r t i f i c i a l stream c h a n n e l s c o n t a i n i n g (a) RAINBOW TROUT, (b) WHITEFISH (c) RAINBOW/ WHITEFISH c o m b i n a t i o n , and (d) NO PREDATORS. P l o t o f 'time p e r i o d s x f r y t y p e ' i n t e r a c t i o n from a n a l y s i s o f v a r i a n c e , Appendix V l l a - d .  o  41  (a)  FAST  MIGRANTS  (b)SLOW  MIGRANTS  100  _  6  0  E 40  N.S . W  N.S. R/W  N.S.  No  R/W  No  predator s  (c)  EXPERIENCED  100  80  R  Rainbow  W  Whitefish  R/W No  60  Rainbow/Whitefish No  predators  40 **  R E J E C T Ho ** N.S.  R  FIGURE  19:  W  ,  R/W  No  Means and c o n f i d e n c e l i m i t s o f ' p e r c e n t m i g r a n t s i n t h e f i r s t h o u r ' p l o t t e d f o r (a) F A S T MIGRANT, (b) SLOW MIGRANT, and (c) EXPERIENCED f r y m i g r a t i n g w i t h i n e a c h o f f o u r predator regimes. A n a l y s i s of v a r i a n c e found only t h e experienced f r y means t o b e s i g n i f i c a n t l y different. D u n c a n ' s New M u l t i p l e R a n g e T e s t r e s u l t s a r e a l s o s h o w n on g r a p h s ; means u n d e r s c o r e d b y s o l i d l i n e a r e n o t s i g n i f icantly different.  42 (a)  RAINBOW  (b) W H I T E F I S H  100  100  80  80  60  60  40  40  s  N.S. F.M.  S.M.  fry  (c)  N.S. Ex.  F.M.  S.M.  Ex.  Ch. Ex.  (d)  PREDATORS  type  RAINBOW/WHITEFISH  100  NO  100  80  60  60  i  40  **  40  R E J E C T Ho ** N.S.  F.M.  S.M.  Ex.  F.M. F.M.  Fast  Migrants  S.M.  Slow  Migrants  Ex.  Experienced  Ch.Ex.  Channel  N.  Naive  S.M.  Ex.  Ch.  Ex.  N.  Experienced  F I G U R E 20: Means a n d c o n f i d e n c e l i m i t s o f ' p e r c e n t m i g r a n t s i n t h e f i r s t hour' p l o t t e d f o rthe experimental f r ym i g r a t i n g w i t h i n a r t i f i c i a l s t r e a m c h a n n e l s c o n t a i n i n g (a) RAINBOW TROUT, (b) W H I T E F I S H , ( c ) RAINBOW/WHITEFISH combination, and (d) NO P R E D A T O R S . A n a l y s i s o f v a r i a n c e found only t h e means f o r t h e f r y t y p e s w i t h i n t h e r a i n b o w / w h i t e f i s h c h a n n e l t o be s i g n i f i c a n t l y d i f f e r e n t . D u n c a n ' s New M u l t i p l e Range T e s t r e s u l t s a r e a l s o shown o n g r a p h s ; means u n d e r s c o r e d by s o l i d l i n e s a r e n o t s i g n i f i c a n t l y different.  43  or  eaten,  migrants  summarized and  fry  and  for  percent  percent  predator  (Figure  percent  fewer (3)  the  Summarizing,  cn t h e  following:  predators migrants fry  than  predators  noted both  other and  percent  fast  the  largest of  both  migrants  migrants  and  resultant  first  1.5  percent  again  percent  eaten  and agreed  differently f r y ; the  pattern  migrants  totally  and  patterns  pattern  fry  with  w h i t e f i s h trough.  with  more n o n - m i g r a n t s ;  of  h o u r s and  different,  non-migrants;  experienced  of  Test  significantly  migration  migration ratio  Eange  troughs,  migration  for  significant  greater  responded  correspondingly fry  was  trough  f o r the  percent  different  significantly  predator  an i n c r e a s e d p e r c e n t a g e  w i t h i n the  both  Multiple  n o n - m i g r a n t s and  slow  i n f l u e n c e d the  and as  and  New  type  a greater percent  or eaten  (1)  predator  a corresponding  d i s p l a y e d a more g r a d u a l  migrating  groups  and  analysis  migrants,  shewed  predator-free  to the  migrants  only  slow m i g r a n t s  the  lost  IX,  Duncan's  migrants  compared  percent  percent  (1)  (2)  percent  while  eaten.  found:  non-migrants; different  or  Appendix  non^migrants s i g n i f i c a n t l y  types;  lost  21)  with fewer  in  for  to  slow fewer  and  (2)  all  fry  non-migrants;  nigrating over  to  downstream  the  evening.  liscussion  The  foregoing  guestions:  (1)  migrants,  which  response alter  Are  experiment naive display  fry  designed  divided  different  to various predators?;  behavior?  was  and  to  into  fast  migration (2)  How  examine and  patterns  does  two slow and  experience  44 (a) % MIGRANTS Fry Type  Predators +/-  C ni u oo  1  F.M.  S.M.  E x .  Naive  R.  W.  R/W  No  (b) % NON-MIGRANTS Fry Type  C o c  Predators +/-  • V  F.M.  S.M.  E x .  Naive  + R.  W.  R/W  No  (c) % LOST or EATEN loo,  2  F r  y yp T  Predators +/-  e  60  o  + F.M.  FIGURE  S.M.  E x .  Naive  R.  W.  R/W  No  21: M e a n s a n d c o n f i d e n c e l i m i t s o f p e r c e n t ( a ) MIGRANTS, (b) NON-MIGRANTS, a n d ( c ) L O S T o r E A T E N f o r t h e e x p e r i m e n t a l f r y m i g r a t i n g through a r t i f i c a l stream channels with and without predators. P l o t o f ' f r ytype' and 'predator' f a c t o r s from a n a l y s i s o f v a r i a n c e , Appendix IXa-c. D u n c a n ' s New M u l t i p l e R a n g e T e s t r e s u l t s a r e s h o w n b e l o w e a c h g r a p h ; means u n d e r s c o r e d b y s o l i d l i n e s a r e n o t significantly different. N a i v e f r y were n o t i n c l u d e d i n the a n a l y s i s .  45  Initial VI.  expectations  and  E r i e f l y , i t was expected  migrant  results  that  are summarized i n Table  both  f r y would d i s p l a y a s i g n i f i c a n t  experienced  migrants/non-migrants  slow  Experience  fry.  was  ratio,  expected  migration r a t e s d i s p l a y e d by the f a s t  to  noted  (Table  VI),  migrants produced expected were not as d e f i n i t e general  behaving  the  compared t o f a s t things:  (1)  t h i s s e t of degree;  and  to  migrants,  any  finally,  parameters.  of  However,  the f a s t  experiments,  possibly  experimental  (3)  increase  f a s t and  slow  experienced  fry  migrants. This regarding  result  increased  number of migrants f o r experienced f r y ,  experiments  or  to  i n showing the expected t r e n d s ; although i n  similarly  pattern  comparison  results.  c o n f l i c t s with the f o r e g o i n g migration  compared  migrant f r y ; and  predators were expected t o i n f l u e n c e the measured As  fast  difference i n migration  p a t t e r n and percent migrant  and  since  indicating  variability; needed  the  be  cne  three  (2) the ' e x p e r i e n c e  reinforced  previous  cf  to  a  experiments  The r e s u l t s  failed  to have  migrants.  c l e a r l y i n d i c a t e t h a t sockeye f r y vary i n t h e i r  behavior with some d e s c r i b a b l e as f a s t slew  in  greater  d i s c r i m i n a t e between f a s t and slow migrants, experience may a f f e c t e d only f a s t  1  migrants  and  others  as  migrants. Both m i g r a t i o n p a t t e r n s and the r a t i o of percent  migrants  to  difference  percent  reflecting inherited  displayed  a  significant  between the f a s t and slow migrants. These f r y may  two g u i t e d i s t i n c t intergrades  non-migrants  types,  or  there  may  be  a l l degrees  between the two, the d i f f e r e n c e between  in  responsiveness  to  of  individuals  a d i f f e r e n c e i n environmental experience as differences  be  well  as  environmental  46 TABLE V I :  E x p e c t a t i o n s and r e s u l t s MIGRANTS,  f o r experiment  testing  and EXPERIENCED f r y i n a r t i f i c i a l  FAST MIGRANTS, SLOW  stream  c h a n n e l s w i t h and  without predators.  PREDATOR TROUGHS  NO PREDATOR TROUGH  MIGRATION PATTERNS Fast  Migrants (FM)  -migrate r a p i d l y  downstream  ++  - s l o w e r m i g r a t i o n compared t o ++ predator troughs, y e t f a s t e r compared t o SM i n no p r e d a t o r trough  Slow M i g r a n t s (SM)  - s l o w e r m i g r a t i o n compared t o FM ( i e . fewer m i g r a n t s w i t h i n the f i r s t hour)  ++  - s l o w e r m i g r a t i o n compared t o FM, and p o s s i b l y s l o w e r comp a r e d t o SM i n p r e d a t o r troughs  Experienced  - f a s t e r m i g r a t i o n compared t o FM ( i e . more m i g r a n t s w i t h i n the f i r s t hour)  +  - f a s t e r m i g r a t i o n compared t o FM  - g r e a t e r %'age o f m i g r a n t s compared t o SM  ++  - s m a l l e r %'age o f m i g r a n t s compared t o p r e d a t o r t r o u g h s  - s m a l l e r %'age o f compared t o SM  ++  - g r e a t e r %'age o f n o n - m i g r a n t s ++ compared t o p r e d a t o r t r o u g h s  - s m a l l e r %'age o f m i g r a n t s compared t o FM  ++  - s m a l l e r %'age o f m i g r a n t s compared t o p r e d a t o r t r o u g h s  - g r e a t e r %'age o f compared t o FM  ++  - g r e a t e r %'age o f n o n - m i g r a n t s ++ compared t o p r e d a t o r t r o u g h s  % MIGRANTS  ++  *1  &  % NON-MIGRANTS Fast  Migrants  -smaller t o SM Slow M i g r a n t s  -greater FM Experienced  non-migrants  %'age e a t e n  compared  non-migrants  %'age e a t e n  compared  —  +  - s m a l l e r %'age o f m i g r a n t s compared t o p r e d a t o r t r o u g h s  - s m a l l e r %'age o f compared t o FM  +  - g r e a t e r %'age o f n o n - m i g r a n t s ++ compared t o p r e d a t o r t r o u g h s  non-migrants  %'age e a t e n  compared  —  - a p p r o x i m a t e l y t h e same %'age o f m i g r a n t s observed i n m a j o r i t y o f experiments o b s e r v e d i n some e x p e r i m e n t s seldom o b s e r v e d never observed  ++  +  *1 - a l t h o u g h e x p e r i e n c e d f r y m i g r a t e d s l o w e r compared t o FM, C h a n n e l f r y m i g r a t e d i n a s i m i l a r p a t t e r n t o FM. *2  ++  - g r e a t e r %'age o f m i g r a n t s compared t o FM  -smaller t o FM  ++ +  *2  Experienced  i n p r e d a t o r and p r e d a t o r - f r e e t r o u g h s .  47  experience. Regardless individual different  fry,  fts  'awareness*  when  are  of p r e d a t o r s of  fry  Interpretation sockeye  the  fry rests  are  experiments  for  stream  migrating  of t h e f r y r e s p o n s e most p r o b a b l y  any  reported  that  survived exhibited  s c h o o l s . Thus t h e  also  higher  noted  which had  p r e d a t i o n , and  and  sensed  were  The  the  either  downstream  fry  array  coherent  in  tendencies  increased  are  implying  schools? but  Direct aquarium  observations),  the  presence  migration  predator  is,  present,  field in  cf fry school.  p o i n t . That  lacking,  fry  speed  presence,  behavior;  of is a  and  is  for fast  most  cases,  migrated  r a t e s and  an  in  Ginetz  (1972)  dcwn a p r e d a t o r  stream,  migrants.  were r e - r u n t h r o u g h  survival  much more q u i c k l y  schooling  last  are  to  naive  in schools.  affects  tendencies fry,  a  predators  sockeye  reflection  enhancing  in  p e r s o n a l l a b and  view t h a t  an  i n c r e a s e d r a t e compared  downstream  confirmation  to  evcke  their  disorganized  or,  with  exposure  apparently  absent.  a  rates* with  form  Experience  are  by  alters  channels, a t an  only  predators  accomplished  fry  among  cf f r y  i n some way  downstream  h e a v i l y upon t h i s  ( G i n e t z , 1972;  t o f a v o r the  groups  r e s u l t s i n terms of  increased migration fry  that  affected  The  downstream,  the  differences  f r y which  predators be  the  predators  i n the  i n the  may  of  observations  tend  differently  present.  where  of  the  clear  ' s p o o k e d ' downstream  swimming i n d i v i d u a l l y  that  also  f r y migrated  were  situations  the  for  ( G i n e t z , 1972)  migration  cf  is  p a t t e r n s . The  'chased' to  it  basis  mentioned,  predators  presence  the  experience  predators.  behavior  of  t h e same  procedure,  increased migration  speed  48  compared'  to  naive  Further,  he  suggested  (i.e.  experienced  predators (i.e.  in  naive  Experiment  fry  migrating that  f r y ) was  increasing  Effect  past  experience  more e f f e c t i v e survival  f r y i n a predator  2 -  down a p r e d a t o r - f r e e  of  than  cf only  a  trough. predator  awareness o f  of f r y i n predator  channels  trough).  Different  Predators  and  I i qht  light  enhances  on  Emerging F r y  Methods And B e s u l t s  To  test  the  hypothesis  that  a s s o c i a t i o n s o f emergent f r y ,  hence  tendencies  increased  factorial  6  slow  24  predators  and  stcred and,  and  trial,  i n covered  prior  experiment  tc  which t h e f i s h f t - c . Predator  a t normal stream was a l m o s t  types  s e t time  (fast  Each  initiation  were  at  troughs  treatments,  was p o s s i b l e . E x p e r i m e n t a l  an  (no  light of  an  ambient  contained  the  fry  any f o r e i g n  temperatures.  complete,  p e r i o d s (0,  t c two c c n d i t i c r s  the  10 g a l a q u a r i a t o r e d u c e  were m a i n t a i n e d  a 3 x 2 x 3  combination).  and 4 w h i t e f i s h , . D u r i n g  natural f r y migration the  0„01  fry  schooling  f r y ) were s u b j e c t e d t o a  a l l were e x p o s e d  hour  during  of  Three  and e x p e r i e n c e d  of  survival,  (1500 f t - c ) f o r t h r e e  one  intensity  rainbow t r o u t  designed.  rainbow/whitefish  ended  experimental  was  source  hours),  treatment  light  resulting  migrants  light  and  a  experiment  migrants, constant  and  development  mutual  2  were  stimuli;  Since  the  o n l y one r e p l i c a t e o f methods and  analysis  49  were as p r e v i o u s l y As w i t h cf  previous  f r y migrating  Analysis  of  pattern  The  downstream  variance  plus  x predators*  (Appendix  migrating percentage  evening  X) showed  of f r y migrating  x f r y type'  period.  non-migrants  22).  the o v e r a l l  migration  downstream  per  time  ( F i g u r e 23) and  'time  term  significant.  showed  that  the  hour,  compared  percent  the  greater  downstream w i t h i n t h e  increased  in t i e first  cumber  (Figure  yet a s i g n i f i c a n t l y  f r y migrated  Predators  downstream cf  interaction  were s i m i l a r ,  of experienced  sampling  the  ( F i g u r e 24) i n t e r a c t i o n s t o be  and slow m i g r a n t s  percent  during  was a d e c r e a s i n g  'time p e r i o d s x f r y t y p e '  'time p e r i o d s  fast  experiments, there  ( i . e . t h e number  periods), periods  described.  first  of  fry  resulting  i n a smaller  the  predator-free  of  the i n d i v i d u a l  to  troughs. With treatments  only  one  replicate,  and f r y t y p e s  comparison  of  the  comparison  was n o t s t a t i s t i c a l l y migration  e n l i g h t e n i n g . Ccmparing  first  patterns  the f r y types,  was a d i f f e r e n c e between t h e p r e d a t o r Fast  migrants  downstream displayed migrants. light  in  and  there  were  Experienced  were as a b r u p t  but  of  troughs,  not  f r y , (except  in their  fry  migration  light the  within the f i r s t  (Figure  hut 25)  i n most c a s e s  and p r e d a t o r - f r e e  channels  fewer  f o r experienced  effect  predatcr-free downstream  predator  a similar trend,  treatment  The  the  feasible,  migrated  a is  there  troughs. rapidly  n o n - m i g r a n t s . Slow m i g r a n t s as  distinctively  f o r cne t r i a l in  the  fast  - t h e 24 h o u r  predator  p a t t e r n as f o r f a s t  treatment  as  channel) migrants.  was n o t c l e a r c u t . F o r t h e  experienced  fry  hour f o r a l l l i g h t  migrated  rapidly  treatments.  When  50  0.5  FIGURE  22: M e a n m i g r a t i o n  0.5  FIGURE  h r sampling  h r  p e r i o d s  pattern  sampling  of experimental f r y .  •  •  f a s t  migrants  •  •  slow  migrants  *  *  e x p e r i e n c e d  p e r i o d s  23: M i g r a t i o n p a t t e r n s o f e x p e r i m e n t a l f r y within a r t i f i c i a l stream channels. P l o t o f 'time p e r i o d s x f r y t y p e ' i n t e r a c t i o n . f r o m a n a l y s i s o f v a r i a n c e , A p p e n d i x X.  —• • -•  0.5  FIGURE  h r sampling  p r e d a t o r s no  p r e d a t o r s  p e r i o d s  24: M i g r a t i o n p a t t e r n s o f e x p e r i m e n t a l f r ywithin a r t i f i c i a l stream channels. P l o t o f time p e r i o d s x p r e d a t o r s ' i n t e r a c t i o n f r o m a n a l y s i s o f v a r i a n c e . A p p e n d i x X. 1  (a)  FAST  0.5  (b)  MIGRANTS  h r sampling  p e r i o d s  FIGURE  SLOW  (c)  MIGRANTS  0.5 h r s a m p l i n g  p e r i o d s  EXPERIENCED  °-  25: M i g r a t i o n p a t t e r n s o f e x p e r i m e n t a l f r y i n a r t i f i c i a l s t r e a m c h a n n e l s with and without predators. P r i o r t o each experiment, f r y were subjected t o d i f f e r e n t l i g h t treatments.  5  h  r  sampling  p e r i o d s  52  predators early  were p r e s e n t  i n the With  slow m i g r a n t s ,  (0  twice the  The  migrants  summarized  the  0 and  of  exposure  of  fry  6 and  subjected  to  r a t e and  migrant  l i e .6 and  24  fry,  on  fast  yielded tended  especially fry  troughs.  migrants the  i n c r e a s e d , and  24 no  hour t r e a t m e n t  predator-free  predators,  percent XIa  and  the  migrants Figure  was  similar  percentage  significantly  of  fewer  first of  hcur  is  variance  showed c n l y  range t e s t s  the  fcund  6 hour t r e a t m e n t ,  a and  treatment. to  light  first  was  hour  lengthened,  became  Further,  in  the  percent  the  percent  less  affected  mcst  cases,  migrants  by  the with  within  the  a maximum.  percent  non-migrants  only the  the  Analysis  Further  a n a l y s i s of v a r i a n c e  Bange T e s t s o f t h e  26.  between t h e 0 and  6 hours of l i g h t ,  hour reached  in,  M u l t i p l e Eange T e s t  significant.  predators.  approximately  showed  with  difference  presence  percent  slow  and  evening  the  t o be  within  and  but  24 h o u r l i g h t  Finally,  but  p r e s e n t , the  at a slower  treatments  Duncan's New  migrants  first  occurred  resulted.  effect  As t h e  migrated  light,  light  i n Appendix  significant  predators  results,  predator  early  subsequent  predator  no  rate of  no  cases,  Analysis  and  of  i n the  ncr-migrants  to  i n both  effect  a l l three  migration  number o f n o n - m i g r a n t s . P r e d a t o r s  migration  subjected  were s i m i l a r  in  gave s i m i l a r  i n c r e a s e the  those  and  hour treatment)  approximately tc  major p a r t o f t h e  evening.  hour t r e a t m e n t s light  the  predator  migrants  and  New  (Appendix X l b and  (Appendix  effect  Duncan's  t o be  XIc  and  Figure  significant.  Multiple Figure 28)  Although  27)  again not  20 12 l i g h t  FIGURE  26: P e r c e n t m i g r a n t s w i t h i n t h e f i r s t h o u r f o r e x p e r i m e n t a l fry, subjected to various l i g h t treatments, migrating t h r o u g h a r t i f i c i a l stream c h a n n e l s w i t h and w i t h o u t predators.  12 l i g h t  FIGURE  ie  treatment ( h r s )  IB  treatment ( h r s )  27: P e r c e n t m i g r a n t s f o r e x p e r i m e n t a l f r y , s u b j e c t e d t o various l i g h t treatments, migrating through artificial stream c h a n n e l s w i t h and w i t h o u t p r e d a t o r s .  p r e d a t o r s no  p r e d a t o r s  f a s t  migrants  slow  migrants,  experienced  e °  40  12 l i g h t  FIGURE  18  treatment ( h r s )  28: P e r c e n t n o n - m i g r a n t s f o r e x p e r i m e n t a l f r y , s u b j e c t e d t o various l i g h t treatments, migrating through artificial stream c h a n n e l s w i t h and w i t h o u t p r e d a t o r s .  54  as  distinct  hour),  as  the  predators,  26,  basic trend  migrants,  in  percent was  (the p e r c e n t  i s evident  at the t h r e e  of  troughs  Figure  light  migrants  f o r the  treatments,  w i t h i n the  percent  migrants.  increased  the  h e n c e t h e r e were f e w e r n o n - m i g r a n t s . The lost  or eaten  between t h e  predatcr  first  and  The  number  difference  predatcr-free  negligible.  liscussion  As caused  i n foregoing  experiments,  an  i n the  increase  hour, i n c r e a s e d the t o t a l the  percent The  is  cf  percent  the  fry  i t  are  f o r the  treatment, migrants the  visual  form the  of  of  migrants  migrants,  predators  i n the and  to increased l i g h t  difficult  the  three  w i t h i n the  percentage  total  tc  first  reduced  first  reverse first  of non-migrants  (Breder  t h a t as t h e  environment.  a n a l y s i s of that  prior  With  the  s i x hour the  as  to  i n c r e a s e d . T h i s was  totally)  an  also light  percent  decreased,  of  while  increased.  1951,1967; Morrow light  treatment  observable,  associations  associations, orienting surrounding  was  hour and  the  especially  hour, i t appears  of migrants.  trend  treatments  interpret,  f r y t y p e s . From  of migrants  percent  fry-to-fry  clues  possible  with  (within the  Since  of  subjected to a longer l i g h t  the  percent  percentage  and  migrants  experiment, the noted  percentage  of f r y response  much more complex  the  presence  non-migrants.  guestion  compounding  the  treatments and  (schccls) 1948;  are  enhanced  Shaw 1 9 6 0 ) ,  i n c r e a s e , f r y may  familiarizing  With o n l y a few  by  it  is  tend  to  themselves  hours of l i g h t ,  with and  55  the  introduction  and  move  rapidly  subjected been  t o 24 h o u r s  subject  to further  the f i r s t shown  numbers  light  migrating  was  migrants,  and  subjected  to light,  light  slow  at a slower  f r y types  the  have  Since  field  p o s t u l a t i c n s are  treatment, as  f o r both  some t r e n d s  noted  migrants.  percent  migrating.  the  fry  Assuming  enhances t h e percentage  migration  pattern  cne  treatment by  fast  to i n c r e a s e  replicate  appears  o f slow  of  increasingly  initially  i t  order  experienced, were  the  happening,  were  a r e e v i d e n t . With no decreasing  f r y tended  migrants  non-migrants  previously:  As  experienced  the  downstream, s u g g e s t i n g t h a t t h e l i g h t affecting  these  may  as a c o h e r e n t  pace.  migrants/percent  o f what i s a c t u a l l y  treatment  s c h o o l s form  lacking,  be s i g n i f i c a n t ,  and no  indicative  behavior  been  study.  h o u r and p e r c e n t  percentage  schooling that  downstream  the d i f f e r e n t  to  predators,  the  are  may t e t c g r o u p  However, when t h e f r y have  reinforced,  observations  Although  the  light,  which m i g r a t e s  hehavioral  not  downstream.  sufficiently  group,  in  of a predator, the f r y reaction  that  migrants i s  a  is the  moving  component  increasing  schccling  tendencies. With light slew  predators present, experienced  migrated  first.  m i g r a n t s , , A f t e r 6 hours  experienced  f r y migrated  possibly  acting  presence  of  downstream the  downstream  on  the  predators,  of l i g h t ,  migrants  the  a t t h e same r a t e , fast  migrants  reinforcing  within  subjected  to  no  The same was t r u e f o r f a s t and  m i g r a t i o n . However a f t e r  percent  fry  migrants  the l i g h t  additively  the  24 h o u r s  the f i r s t  fast  tendency cf light  and  treatment with  the  f o r a rapid exposure,  hour f o r t h e t h r e e f r y  56  types were s i m i l a r and approximately o n e - t h i r d l e s s , compared t o the 0 and 6 hour l i g h t treatments. T h i s suggests certain  period  of l i g h t exposure,  that  extended  a  the d i f f e r e n c e s i n f r y types  become n e g l i g i b l e ; the b e h a v i o r a l i n t e r a c t i o n s o c c u r r i n g the  after  during  period of l i g h t overcome e a r l y experiences and/or  d i f f e r e n c e s i n the f r y t y p e .  57  SECTION C: DEVELOPMENT  OF SCHOOLING  BEHAVIOR  Experiment J - F r y Response t o L i g h t  Material  And Methods  Male and f e m a l e s a l m o n Collection, in  fertilization  and i n c u b a t i o n  from F u l t o n  4x11.4 The  cm)  tank  procedures are d e t a i l e d  and  were c o n d u c t e d  1.83  m section  the  in  the  e x p e r i m e n t a l chamber observed  consisted ft-c  (10x18.3  fiberglass  t h e water  by  tank. depth  dark.  under  intensity  0.1  Following f o r one  f t - c for  a  groups  f o r varying  time  from  'treatment',  hour,  then  10 m i n u t e s .  different  50 f r y  the stream"channel f o r the  the f r y  The t r e a t m e n t  of f r yto  4.0  p e r i o d s , from  -  30  12.0  seconds  24 h o u r s . Three-dimensional c o o r d i n a t e s of the f i s h  at  stream  of t a k i n g  them.in  was d a r k e n e d  of subjecting  light  cm  a t a p p r o x i m a t e l y 0.13 cm/sec,  experimental procedure c o n s i s t e d  minutes  were  gridded  o f 4.0 ± 1.0° C.  s t o r a g e t a n k s and a c c l i m a t i n g  15  a  of a l a r g e r  a t 30 cm, t i e v e l o c i t y  a temperature Tie  in  was d e s i g n e d a s a f l o w - t h r o u g h s y s t e m ,  maintained  by  Biver.  Appendix X I I . Experiments  to  r o e was c o l l e c t e d  observation  through the glass  sides,  and a m i r r c i  a p p r o x i m a t e l y 4 5 ° above t h e t a n k . T y p i c a l l y ,  observation  the  fry  were r a n d o m l y  were  distributed  determined suspended  a t the onset and s o l i t a r y  of on  58  the tank  bottom. W i t h i n a s h o r t  initiated  schooling  tendencies. Prior  were r e c o r d e d as i n d i v i d u a l s sections; recorded  however,  once  the  (time  one  fry  end  started  reached  the  ended  other end);  other  end);  (4)  time  i n a swimming,  uni-oriented  Statistically, groups hrs.), Test  (0  -  3  taken t o form  the  3+  25%  and  to test  school  was  tank. time,  initiated  movement 75%  time, or  starting  of  turning  towards  (measured  (at l e a s t  or  by  90%  the the  c f the  group). into  - 6 h r s . , 6+  w i t h a n a l y s i s o f v a r i a n c e and  applied  activity  swimming  d a t a were d i v i d e d  hrs.,  lengthwise  (1)  (3) s c h o o l l e n g t h d u r i n g swimming  and  10 the  (2) r e s t i n g  (the t i m e between r e a c h i n g an end  fry  the  and  the f r y  when g r e a t e r t h a n  tine,  grid);  activity  c f the  at l e a s t  o f t h e t a n k , and  active  length  parameters: when  of  commenced  as t h e t i m e t a k e n t o swim t h e  activity,  to t h i s  w i t h i n each  movement  a n a l y s i s examined f o u r  towards  t i m e t h e f r y became  four  treatment  - 9 h r s . , and  Duncan's New  9+  Multiple  -  >24  Bange  differences.  Besults  Preliminary ccnditions experiments generalized  o b s e r v a t i o n s o f f r y b e h a v i o r under  a r e summarized employed  in  Table  different  b e h a v i o r p a t t e r n s were  VII., although  time  variable groups  Swimming t i m e  measured. to  be  frames  (Table V i l l a )  of was  the  trial,  different;  a  four  t h e most  A n a l y s i s o f v a r i a n c e found  significantly  per  subseguent the  similar.  T a b l e V I I I summarizes t h e a n a l y s i s parameters.  experimental  measured  informative  the f i v e  Duncan's New  treatment Multiple  59  IfiELj V I I :  Summary of behavioral o b s e r v a t i o n s f o r f r y under experimental conditions discussed within text.  INTRODUCTION TO TRNK (Darkness)  IEY  i  SHOCKED  - f r y swim t o b o t t o m ; r e m a i n stationary f o r 0.5-6 min -slowly start non-directicnal movement; a p p r o x . 105? r e m a i n s t a t i o n a r y -within 15-20 min loose groups forms, swimming l e n g t h o f t a n k -movement increases; tighter grcup fcrm ( s c h o o l ) w i t h u n i d i r e c t i o n a l movement, - w i t h i n 35-45 min, movement decreases and f r y s p r e a d out cn b o t t c m - t h r o u g h o u t t h i s p e r i o d , f r y remain i n lower t h i r d o f water column - f r y r a p i d l y d i s p e r s e as a very l o o s e group -within 3-5 min g r o u p slows down and d i s t r i b u t e s over the bottcm  LIGHTS • ON *  - t i g h t g r o u p f o r m s w i t h i n 15 s e c i n f u r t h e s t c o r n e r frcm l i g h t source - w i t h i n 1-2 min, f r y r a p i d l y swimming l e n g t h o f t a n k i n a s c h o o l , and h a v e moved to. the upper t h i r d o f t a n k -within 5-10 min, the schccl slowly d i s p e r s e s and movement d e c r e a s e s  LIGHTS •OFF'  - t i g h t group f o r m s i n water column and a l l movement s t o p s ^for 1-2 min g r o u p r e m a i n s m o t i o n l e s s , t h e n s l o w l y d e s c e n d s t o l o w e r t h i r d o f tank and fry disperse - w i t h i n 5-7 min f r y s l o w l y s t a r t movement a s n o t e d above f o l l o w i n g i n t i c d u c t i c n t o t a n k  r  60  TABLE V I I I :  Summary of analysis f o r f r y observed f o l l o w i n g variable light treatments. Measured parameters described within text. r  — T  |  TREATMENT GROUPS  !  | SKCVA  1  |  0 h r s | 0+ > | 3+ > | 6+ •> 19 + > l (Fcalc) | 3hrs | 6hrs | 9hrs | >24hrs| SWIMMING TIME (sees) Mean Stan, e r r o r Sample s i z e  j  a-  I 18.69 0.37 4  24.77 1.74 14  I  35,. 85 2.36 5  31.50 2.33 2  I 27.74 1.41 3  6.62 REJECT Ho  10.18 1.08 3  I I | 1.73 | ACCEPT I Ho  3.4 3 0.46 2  0.40 ACCEPT Ho  E.N.M.R.T. b.  RESTING TIME (sees) Mean Stan, e r r o r Sample s i z e  I I I I I I 16.04 | 16.06 | 18.67 | 12. 14 0.69 0.84 I 3.44 | 1.26 | 2 4 5 I 14 I  E.N.M.R.T.  * c . SCHOOL LENGTH* Mean Stan, e r r o r Sample s i z e 1  Y  I I 3. 18 | 0.48 | 4 I  I I 3.3 0 | 0.36 | 14 I  I 3.91 | 0.51 I 5 I  I 3.99 | 0.42 | 2 I .  r  1  L  E.N.M.R.T.  __.  I  d. TIME t a k e n t o FORM SCHOOL (sec)* — — — — — Mean Stan, e r r o r Sample s i z e  38.50 4.63 4  54. 44 4.37 9  66.80 7.96 5  I I |125.00 | 45.00 I 2  23.00 4.04 3  KE0SKALWAI1IS*2 (Hcalc) 7.42 ACCEPT Ho  D.N. M.R.T.  D.N.M.E.T. = Duncan's New M u l t i p l e Range T e s t (Underscored means are not s i g n i f i c a n t l y different) ANOVA = A n a l y s i s o f v a r i a n c e t e s t i n g h y p o t h e s i s (Ho) t h a t mean v a l u e s f o r t r e a t m e n t g r o u p s a r e n o t d i f f e r e n t * i s c h o o l l e n g t h was measured cn g r i d a s number o f s g u a r e s (1 s q u a r e = 1 8 . 3 cms) * Ncn-paramentic a n a l y s i s of variance 2  61  Range T e s t showing hours  both  treatments  remaining swim  groups.  then  (0 h r s . )  significantly  and f r c m  different  Hence t h e swimming t i m e  l e n g t h o f tank)  hours,  the c o n t r o l  decreased  compared  ( i . e . time  i n c r e a s e d t o a maximum  gradually  3+  to to  over the next  24  the  taken  between f o u r hours  9  to  to s i x (Figure  29) . Analysis  of  between t r e a t m e n t school  length  appropriate, account time  showed  VIIIc).  subsequent  over  tc  is  time  Although  Duncan's  an  the  24  New  maximized  hour  increase  t i m e ) , with g r e a t e r  treatment.  analysis  significant  f o r both r e s t i n g  between  in than  the  treatments, being s i g n i f i c a n t l y cf  no  differences  ( T a b l e V I I I t ) and  statistically  Multiple  not  Eange T e s t s c a n  f o r t h e t r e n d s e v i d e n t i n F i g u r e s 30 and 31. The r e s t i n g  corresponds  length  groups  (Table  decreased  swimming  variance  However,  treatment  activity, 6  lower  since  the  to with  which  ( i . e . a decrease i n  hour  three  period,  treatments.. nine  hours  School  of l i g h t  greater or less  calculated  time  F value i n the  o f v a r i a n c e i s low compared t o t h e t a b u l a t e d v a l u e , i t  best  assumed  that  t h e mean s c h o o l l e n g t h s a r e e s s e n t i a l l y  similar. The varied  e s t i m a t e d time considerably  heterogeneity  New  is  treatment  Test) ,  rejected  o f means f o r t h e f i v e  Multiple  groups  within  not  difficult  a  of v a r i a n c e . Non-parametric  (Kruskal-Wallis sinilarity  t a k e n t o form  the  treatment  school  (Table  grcups,  resulting  analysis  cf  null  hypothesis  groups;  to finalize  different.  because  A relationship  of the large  in  variance,  a  Range T e s t showed o n l y t h e 0+ t c s i x h o u r significantly  Vllld)  of  Duncan's treatment  ( F i g u r e 32)  variance  recorded  50  40  30 r  20  10  >24 h r s  6 light  8  10  12  24  treatment (hrs)  FIGURE  2 9 : SWIMMING T I M E - t i m e t a k e n ( s e c o n d s ) t o s w i m l e n g t h o f e x p e r i m e n t a l tank f o r f r y s u b j e c t e d t o v a r i o u s l i g h t treatments. Curve f i t t e d by p o l y n o m i a l r e g r e s s i o n .  FIGURE  30: RESTING TIME- t i m e t a k e n ( s e c o n d s ) t o t u r n a t e n d o f tank f o r f r y s u b j e c t e d t o v a r i o u s l i g h t treatments. C u r v e f i t t e d by p o l y n o m i a l r e g r e s s i o n .  6  FIGURE  3 1 : SCHOOL L E N G T H - l e n g t h o f s c h o o l ( m e a s u r e d b y g r i d ) d u r i n g swimming f o r f r y s u b j e c t e d t o v a r i o u s light treatments. Curve f i t t e d by p o l y n o m i a l regression  160  '3 h r s  3+  »6 h r s  6 light FIGURE  6+  »9 h r s  8  9+  10  »>24 h r s  12  24  treatment (hrs)  3 2 : SCHOOL FORMATION- t i m e t a k e n ( s e c o n d s ) t o f o r m for f r y subjected to various l i g h t treatments. f i t t e d by p o l y n o m i a l regression.  school Curve  64  for  the s i x t o nine  Eoth  control  resulted  hour t r e a t m e n t , y e t some t r e n d s  and  greater  i n l e s s time taken  Experiment  2 - Effect  than  nine  hours  are evident.  light  treatment  t o form a s c h o o l .  o f L i g h t and F r y  Densities  on  Schooling  Behavior  Material  And Methods  Three of  components  a school  involved  (2)  communication; schooling.  some  and  The  light  (3) t i m e  following  tendencies  i n s o c k e y e f r y . The to t h i s  Similar  latter  mortality batch  of  were  ( c a . 80%) roe  Weaver C r e e k , (stage  5  of  was B.C..  storage  tubs.  to  results  foregoing  River,  with  followed  develop  provided  close  for  visual  skills  of  t o examine t h e cf schooling only  fertilization  the  collected  limited  salmon  to  were  and i n c u b a t i o n  incubation sockeye  were  Ginetz  roe  A l l procedures involved  from  a l l fry  trays  study,  as p r e v i o u s l y d e s c r i b e d .  early i n  Once  lab  Columbia.  development,  the incubation  available  in  end.  the  at Fulton  must be  factors i n the i n i t i a t i o n  U n i v e r s i t y of B r i t i s h  the  frcm  to  be  and m a i n t e n a n c e  experiments attempted  of these  information  three  must  i s required  importance  the  initiation  a r e : (1) a c e r t a i n number o f f i s h  proximity;  collected  with  phase,  with  Due  t c high  a  second  salmon  spawning i n  sufficiently  developed,  1972), t h e y  darkened  at  227  were t r a n s f e r r e d litre  fiberglass  65  Experiments s g u a r e s ; each a mirror Prior  were  square =  suspended  to  conducted  each  27 mm )  8  -  14.5°  C,  observation), ft  Treatments light  number for  -  compass d i r e c t i o n  decoded each 2  30  time  only  were  filled  a  before  subjecting  few  each  ft-c).  period,  period  water,  r a n g i n g from degrees  in  per  ppm.  the  tank  light  and  treatment.  the f r y t c a s e t period of  r a n g i n g from  period  0 t o 12  hours,  i n the dark  under  low  light  The 3 - d i m e n s i o n a l c o o r d i n a t e s and  o f t h e f r y were d e t e r m i n e d  minute  aspect.  with  a t 13 - 14  placed  f o r a 30 m i n u t e  5.0  for analysis  every  10  seconds  n o t e d on a t a p e r e c o r d e r , and  ( F i g u r e 3 4 ) . The d i s t a n c e  was c a l c u l a t e d  by t a k i n g  f r y moved  the square  later within  root  of x  2  + z ; x, y, and z b e i n g t h e 3 - d i m e n s i c n a l c o o r d i n a t e s . 2  Observing recording activity period. as  of  was  9 x 9  (Figure 3 3 ) , with  t h e f r y r e c u p e r a t e d f o r 30 m i n u t e s  (4.5  the  fry  x  3-dimensional  maintained  minutes  were t h e n o b s e r v e d  for  varying  ( c a . 15 - 30 f t - c ) ,  which  intensity  + y  20  tank  (15  experimental period  levels  of  consisted  intensity  following and  (although  set  acclimatized  the  the f u l l  and o x y g e n  gridded  a t 45° f o r a  experiment  temperature throughout  a  10 g a l a q u a r i u m  2  overhead  in  greater than t h r e e f i s h  the  10  patterns, A second  a group  second  movements,  out of t h e group  approach  throughout  presented a  problem  t h u s one f i s h ,  with  was  was t o r e c o r d  the o b s e r v a t i o n  observed  for  normal  the  the behavior of the  period.  for  time fish  66  IIGOEE 34:  Experimental tank, containing during observation period.  fry,  as  seen  67  Eesults  Typically, light  as  treatment  sinilar.  A  f r y were s u b j e c t e d  their  particular  movement  patterns  example,  the  g r o u p o f t e n was o b s e r v e d further The  relatively  gradually  s t a r t e d moving and t h e y  hour,  either  the  Subseguently,  upon t h e l e n g t h following  Once  motionless  aguaria.  of  light  light  period  period  and  a  (Figure 35).  a f t e r i n t r o d u c t i o n to the treatment  reached  treatment,  constant  essentially  began  a maximum  the  fry  withir-  the  t h e movement was m i n i m a l . Dependent  t h e dark r e c u p e r a t i o n  remained  of  movement o f one f r y i n a  the r e c u p e r a t i o n  darkened  first  were  f o r a 6 hour t r e a t m e n t  h a l f hour f o l l o w i n g  f r y remained  to i n c r e a s i n g periods  or  the  period  amount  increased,  decreased  within  cf  movement  then  activity  the  30  min  observation. To light  interpret  treatment, plus  variance for  the  (and  a Duncan's New  following blocked fry;  period,  dark  (C)  0 - 1  A n a l y s i s o f mean difference •E',  and,  between  blocks  (Table  IXb) l i k e w i s e  »C* and •D *,  time  For  taken  (Table  to  analysis fry;  both  time  ( i . e . the greater  periods).  number  min  swimming were  hr, 1 - 3  h r , 10 - 30 f r y .  shewed  were s i g n i f i c a n t l y  30  results  (B) 3 - 6  (D) 3 - 6  IXa)  start  the  of  was computed  a  significant  the four s e c t i o n s except f o r blocks  ( i . e . fewer f r y during  times  the  10 - 30 f r y ;  movement  cf f r y , analysis  o f movement p e r  (A) 0 - 1 h r , 1 - 3 hr,  the time p e r i o d o f  M u l t i p l e Bange T e s t )  t h e mean amount  recuperation.  as f o l l o w s :  of increasing  i n c r e a s i n g t h e number  t h e two s t a t i s t i c s :  observation  effect  The  different,  •A' and starting but f o r  of f r y during  both  68  FIGURE  35:  D i s t a n c e moved by one f i s h i n a g r o u p o f t e n f r y . E a c h s q u a r e i s t h e mean m o v e m e n t f o r a 10 m i n u t e o b s e r v a t i o n period. Shaded a r e a r e p r e s e n t s dark r e c u p e r a t i o n p e r i o d .  69  TAELF I X :  Eesults from a n a l y s i s o f v a r i a n c e and Duncan's Hew M u l t i p l e Eange l e s t f o r : (a) mean movement p e r 30 min o b s e r v a t i o n p e r i o d , and (b) t i m e t a k e n to start swimming f o l l o w i n g dark r e c u p e r a t i o n . Data was blocked as described within test. Underscored means are net significantly different.  (.§) Mean | Source  movement  p e r 3 0 min o b s e r v a t i o n  of V a r i a t i o n  | DF |  | Total j Among Within I  | | |  SS  period: MS  I  I  51 | 1144.15 | | 3 | 320.62 | 106.87 | 48 | 823.53 | 17.16 |  6.23  I ** | I  B  D.N.M.E.T. 9.92 4.63  Mean Variance  8.91 5.05  12.74 3. 16  5. 10 2. 36  N.S.  (b) Time t a k e n  | Source  I  to start  of V a r i a t i o n  | Total j Among | Within l _ :  swimminq:  | DF |  -+  _  | I |  l  MS  I  I  1-  V  50 | 1C06.00 | | 3 | 618.97 | 206.32 | 25.07 47 | 387.03 | 8.23 | 1  .  L  B  D.N.M.E.T. Mean Variance  SS  h  2. 87 3.72  5.38 37.98  0. 67 1, 32  1.64 2.62 N. S.  I  — ^ **  I I I  70  time p e r i o d s ) . movement  Both  these  patterns  (Figure  movement when t h e r e subjected  analyses  tended  observation  tc  were more f r y p r e s e n t ,  to greater  Interpreting  36)  and  periods  of l i g h t  the e f f e c t  it  appears t h a t g r e a t e r  of  light  in  less  treatment.  and  treatment  preceding. the  fry  apart). never cne  With 2 o r 3 f i s h start  out  greater  will  than  3 individual  within  2  maintain or  fish  squares  the  increases  by 3 s q u a r e s .  q u i c k l y reapproach  fry  I f this  the other.  fish.  the o r i g i n a l  usually  with  begins, sguares usually  d i s t a n c e i s exceeded,  Greater  numbers of f r y i n  Usually  between 2  2 - 3  fish  i n t h e same d i r e c t i o n  g r o u p may  As t h e number o f f i s h  another  join  up  will  individual cne o r  with  the  i n c r e a s e s i n the tack, the per  group;  15 f r y t h e groups, a r e u s u a l l y 2 f r y , and w i t h  3 - H f r y p e r group.  the  with the  yet i s  main d i f f e r e n c e i s an i n c r e a s e i n t h e number c f f r y such t h a t  in  movement  p a t t e r n but the i n t e r a c t i o n  orientating  hours  resulted  p a s s e s by i n t h e same h o r i z o n t a l p l a n e ,  more o f t h e f r y f r o m passing  fewer  correlate  time  t h e m s e l v e s as a g r o u p . However, i f  group c l o s e l y  cn f r y numbers,  ( c a . 0.5 - 1.0  with  i s n o t as i n t i m a t e . and  f r y were  trend.  i n t h e t a n k , once  apart  the tank d i s p l a y a s i m i l a r or  of  i n close proximity,  Their distance  when  measurements  same manner a r e needed t o s u b s t a n t i a t e t h i s observations  decreased  t h e 3 hour treatment  movement; however, f u r t h e r 6 h o u r  Behavioral  a  was c o r r e l a t e d w i t h  I n most c a s e s ,  the  treatment.  of light  movement  show  of  30,  71  FIGURE  36: Movement p a t t e r n s o f d a r k - a d a p t e d f r y , s u b j e c t e d t o v a r y i n g time p e r i o d s o f l i g h t treatment. Observation t i m e shown o n t h e a b s c i s s a , e a c h mark a 5 m i n u t e interval. O r d i n a t e shows f r y movement; a x i s d i v i d e d i n t o u n i t s o f 5. Each curve i s a separate experiment, w i t h t h e e x c e p t i o n o f 2 and 3 f r y where each graph r e p r e s e n t s a n e x p e r i m e n t w i t h movement r e c o r d e d f o r all f r y . light 0 min  treatment  30 min  3 hr  6 hr  2  fry  1  fry  : T—  1  E  *  *  * »•  S  S"  S  E  L-  ••• V  C  E  V  • I" • S  E  *  E  E  4-  fc-  -  3  fry  -  fc-  I- -  " \—  c  s  *•  *•  •  *  J  s  »  *  »  -  30  fry  20  fry  10  fry  •*-^^V  -  .  k  >  4  4V  »  72  Discussion  In  the  varying these  first  periods  f r y had  experiment, s u b j e c t i n g  of  light  intensity  been k e p t i n t h e  assumed t h a t t h e y r e s e m b l e d first  time.  occurs  with  tank,  nearly  six to  a  observation.  readily This in  a school  under new As  the  and  taken to  school  swimming  time  reduced - h a l f compared  of  when t h e  a school  prior  tendency  visual  a  light  change  under prior  similar  *searching' t c form  the  the  experience pace.  associations  stimulation,  behavior  for  light  fellows  rapid  be  observation  t o no  a  innate  gravel  f r y were h e l d  and, an  could  the  previous  maintain  Since  a noticeahle  no  treatment  their  plus,  f r y are  the  experiencing  The  increases  surroundings,  the  having  fry  had  may  be  more  s i x c r more h o u r s  a c c u s t o m t h e m s e l v e s ; hence swimming t i m e and are  both  longer.  With  i s somewhat l e s s , , b u t that  of  to  suggests that  school.  increase  'nervous'  light  with  explore  This  length  i t  to  external-conditions.  familiar  third  of  and  37,  that f r y with  reflects  absence  exploratory  Figure  Time t a k e n t c f o r m  possibly the  in  fry  behavior.  the  compared t o t h o s e s u b j e c t e d  suggesting form  their  d a r k from h a t c h i n g ,  t o swim t h e  two-fold  light  changed  f r y emerging frcm  summarized  time taken  hours of  pattern,  to  as  'newly emerged*  f r y subjected  those the  experience  h o u r s may  be  hours  light, tc  of  a reflection  between  light  and  of  structural  the  those  with array  i s much  and  only  fry greater the  a  light.  i n developing  of  the  time  s i x h c u r s of  f r y become more e f f i c i e n t time  light,  schooling  t o no  f r y subjected  difference in schooling  previous  the  24  time  with than  a no 12  school;  73  FIGURE  37:  Summary o f r e s u l t s f o r f r y s u b j e c t e d t o v a r i o u s light treatments. Shaded a r e a r e p r e s e n t s degree of u n c e r t a i n t y o n c u r v e b e t w e e n t h e two p o i n t s f o r t i m e t a k e n t o f o r m school. Source of curves f u l l y explained w i t h i n the a p p r o p r i a t e s e c t i o n o f t e x t , l a b o r a t o r y e x p e r i m e n t no. 1.  74  l e s s time fry  i s n e e d e d t o form  a coherent,  are s u b j e c t e d to longer p e r i o d s of l i g h t  cycles,  natural  schooling  rhythms  may  develop  second  experiment  duration  of l i g h t  treatment  behavior  p a t t e r n s o f t h e f r y such  was  reduced.  This  the  previous  experiment.  experimental replicates  will  showed and  studies  Keenleyside  (Healey  juvenile  maintained  the  individual  fish  o n e s ' own k i n d motor  group  fish,  school  i n the f i r s t  (Steven  a  to  balance  (2)  a  t c those  possible behavior  due t o t h e  reported  illuminated  returning,  i f  following  not  behavior  followed;  and  1966). B o t h l a b s t u d i e s h e r e i n patterns generally result  swimming  in  - fish  - fish  (Hemmings  tank,  away  reported  three  swimming  swimming (c)  of  of f i s h to  results  tendency  others  that  between t h e t e n d e n c y  (a) E x p l o r a t o r y t e n d e n c y  (b) R e t u r n i n g  difficulties  an  in  1964;  i t was f o u n d  and, t h e m u t u a l a t t r a c t i o n  1959). T h i s  patterns:  the  (1) t o o few  experiment  Generally  -  Two  and  in  tank.  1959).  by  time  the observations i n  shortcomings  treatments;  tendency  behavior  with  two  introduced  to depart,  fish  altered  1972; Hemmings 1966; Hear 1954; J o h n  Steven  the group;  fish,  movement o r swimming  o b s e r v a t i o n s were s i m i l a r  schooling  frcm  their  both the  o f f r y a r e i n c r e a s e d , so t h a t  t h a t found  1955;  of  hinder too c l o s e a comparison:  a s t h e number  not resemble  that  However,  f o r the longer l i g h t  effect  away f r c m  as  diurnal  increase  increasing  number  i n part correlates  design  Eehavioral  basic  Alsc,  resembling  which  that  the  c o n f i n e s of the smaller observation  other  group.  efficiency.  The  tank  stable  away  Following from  group  fcund  these  c f l a b work.  First,  apparent. from  this  type  75  it  i s d i f f i c u l t to  extrapolate  environment.  Fish  cf  fright  different  monotonous  i n nature are stimuli;  setting.  For  in  i t was  in  Following  i n t r o d u c t i o n , compact s c h o o l s  throughout  period the  the  changes  fish  (1972) f o u n d  behavior  and  same,  for  fish  grouped in  individuals. lab  experiment  conditions  and  a  This  group  i s in contrast  in this  study.  equipment can  t o the  However,  was  is  after  studies i s average  approximately  results  from  different results.  an  scatter  the  to  a  1S6U) .  and  compared  produce v a r y i n g  array  schooling  f o r m e d , but  although  fish  more  lab  between l a b  that  natural  (Jchn  'confidence'  individual swam  time  are  tank. Secondly, comparison  a l s o hazardous. Healey  the  shown t h a t  with  gain  the  to a c o n t i n u a l  comparison,  example,  aquarium  studies to  subjected  behavior  exploratory  the  aquarium  the  isolated the  second  experimental  76  DISCUSSION  a.  Summary, o f E e s u l t s  In  summary, t h e r e s u l t s  moving  downstream  period;  varies  showed t h a t :  throughout  (2) e m e r g e n t f r y a r e n o t  behavior, others  some t e n d  migrate  migration  to  migrate  a  slower  at  pattern,  and  rate  is  predators;  predators tend t o  fry  migration;  elicit not  the  to  be  (5)  significantly  the  presence  above b e h a v i o r  tc predators  compared  to  newly-emerged points  J-  will  jarly  life  be  Life  fast fry  to  subsequently  study  dealt  cycle  of sockeye  i n c r e a s e the  i s sufficient  migratory  behavior  (7)  of  downstream  f r y response  and  and  presence  r a t e of  the  to  tends prior  behavior subjecting  patterns.  These  discussed.  with  Fry  a comparatively  short  f r y - emergence t o downstream  which a number o f d i s c r e t e  (3)  migrants the  predator  migrants;  alters  H i s t o r y o f So c k e y e  This  by  while  (6) i n most c a s e s , f r y w i t h  slow  light  migrants),  cf  displayed different  and  migratory  migrants);  m o d i f i c a t i o n s , the  predator-specific;  experience  (slow  migration  in their  (fast  affected  o f any  t h e number o f f r y  nightly  percentage  ncE-migrants, (4)  the  a l l similar  rapidly  the  (1)  s t a n z a s can  be  defined:  phase  i n the  movement, i n  77  1,. 2. 3. 4. 5.  H a t c h i n g to i n i t i a t i o n of emergence; I n i t i a t i o n o f emergence t o emergence; Emergence f r o m g r a v e l i n t o the water c o l u m n ; Downstream (or upstream) d i s p l a c e m e n t ; and f i n a l l y , ' P o o l or stream r e s i d e n c e f o r a s h o r t period, and/or lake entry.  fit e a c h s t a g e and  a complex  internal  biotic  developmental the  fry  and  (Bams,  s e r i e s of e x t e r n a l environmental  f a c t o r s i n t e r a c t to  behavioral  a  period  of  a m t i e n t water t e m p e r a t u r e s , the  innate  gravel  negative  reserves  are  just  emergence  is  its  (Bams  McEcnald a few  1960,  below  greatest  is  influence  respect  the  During  surface  tuned  to  to  low  1969;  Heard  1965;  wave has  Hoar  and  As  low  during dawn  period the  fts the  gravel with  wave c f f r y timing high  intensity  of  water  decreases,  emerge  1953,  frcm  the  1954,1956,  Generally, dusk  darkest  approaches,  an  yelk  expected  and  fry  deeper  1958;  time, u s u a l l y a matter  emerged.  the  be  light  the  Neave 1955). I n a' s h o r t  (Hartman e t a l . 1962).  1969).  g r a v e l . The  light  upon  r e s u l t s from  (Bams,  of the  i s r e d u c e d , and  to  wiggle  emergence, e a c h  effect  confined  guickly  would  (Bams, 1 S 6 9 ) . As  peak  c l o s e l y dependent  f r y commence movement t o  time.  numbers m i g r a t i n g  intensities  with  behavior  response  the  closely  hours, the  movement  This  move i n waves, as  a t dusk  inhibitory  gravel  to  hatching  accumulates  temperatures  f r y h a t c h and  interstices.  depleted,  in  incubation,  phototactic  surface, tending variation  changes o c c u r i n g  and  1969).  Following  into  regulate  stimuli  of  downstream  t o dawn, w i t h  the  hours c f the  night  increasing  light  water  temperatures  inhibit  further  mode o f  downstream d i s p l a c e m e n t  emergence. After well  emergence, the  known: f r y a r e  swept p a s s i v e l y  downstream,  swim  is  not  actively  78  with  the  current,  movement i t was 19€5;  and  night,  the  S  intensity  aii  1958)  position On fry  in relation the  1944;  that f r y face  current  -  appeared  to  have an  downstream  migrants  process The  process  combination postulates, modulated  of  hence of  active  drift,  that  with work,  f r y ccmmence blindness.  at  i s greater  displaced  downstream  are a b l e to  maintain  pink  Hartman, e t a l . 1962,  1967)  and  cn s o c k e y e  swim  emerging  that i t  active  downstream and  are the  contact  extinction  they  and  faster  than  Erannon  mcving  response  o p p o s i t e , the  whether  fry  visual  waters  the (1972)  downstream  o r i e n t e d r h e o t a x i s , u s i n g means o t h e r  suggested  i s similar,  o b j e c t s , at  deeper  a c t i v e displacement.  fry  must he  pattern,  the  upon  and  Further, the rheotactic  u p s t r e a m . I t was behavior  1955;  downstream  sockeye  stimuli.  are  observations  Neave  t h u s t h e r e i s an that  Johnson  to reference points.  indicate  concluded  1958;  dependent  right  intensity  i s c o m p l e t e and  o t h e r hand, f i e l d  (Pritchard  visual  light  partial  r a t e of dark-adaptation..Fry  dark-adaptation  in  suggested  r a t e of ambient  until  passive  d i s p l a c e d downstream. F u r t h e r  dusk,  the  is  to the s u r f a c e , lose  migration i n a state of  than  1S54,  For  with r e s p e c t t c f i x e d  downstream the  1953,  that rheotaxis  light  h e n c e t o be  Brett  inbetween.  (Hoar 1951,  stimuli  the f r y t o r i s e  1959;  something  1S71)  mechanical  b o t t o m , and  (aii  Byrne  a reduction in  causes  do  postulated  Bams 1969;  visual  or  fry be  former  direction  actively  migration upstream  is  an  and  mcving innate  o r downstream  the  displacement. migration  is  passive displacement. most  c f upstream  than  likely  displaced  As  probably arnold  (1974)  downstream  of current determining  a  by  direction  79  cf  displacement  when  However, s i n c e t h e predation,  i t  the  fry  greatest mortality  is  inconceivable  their  movement. The  risk  of p r e d a t i o n i s s l i g h t .  are i n i t i a l l y  undoubtedly fully  number o f shaded or  or  fry  evenings,  especially hatits  through  After  1967;  McDonald  less  cover  is  ccntrcl  over when  night blindness  the  (Hoar the  schools  a  predation  stream  1953, fry  ccurse  bottom  downstream  banks,  have  1S58;  abandon  light  swimming  (Hartman e t a l . 1962;  congregated  preferring  in  in  1960;  tc light,  Hoar  a  water  McDonald  and  sclitary  1956;  exposure  and  McCart  deep  previous  (Hartman e t a l . 1962;  while  downstream;  m i g r a t i o n ever  been e x p o s e d  1960). A l s o , f o l l o w i n g  eyes  c f the l a k e  hours  1956,  the  zones.  'security'  daylight  along stream  response  areas  stage  movement. As  pursue  reach the  spending  pools  form  gravel.  intensities,  of p a r t i a l  their  upon l a k e e n t r y , t h e f r y  and  lighted  control  spread t h e i r  b u r i e d under c o v e r 1955).  f r y have no light  A state  actively  else,  holding  Neave  C.  d e t e c t and  emerged f r y e i t h e r  night,  that  v i s u a l c o n t a c t with  a rapid t r a n s i t  The  the  swept downstream. Once i n t h e c u r r e n t t h e f r y  dark-adapt,  enabling  one  can  emerge f r c m  upon f r y a t t h i s  f r y emerge a t low  does o c c u r , t h e f r y l o s e and  first  McCart  they  feeding i n  show  brightly  1967).  S i g n i f i c a n c e of Predation  Throughout the downstream (Hunter ranging  frcm  emergence  movement, p r e d a t i o n i s t h e  1959; from  s h o r t time  Neave 20 t o  1953; 80%.  Ricker  1941;  to  major s o u r c e Roos  the cf  1960),  Patterns of f r y behavior  end  of  mortality estimates  a s s o c i a t e d with  80  this  period  predation.  have  Emergence i n t o  downstream, increase  and  encounters.  Previous three  work  interesting  encounters.  et  al,  or  in  prior  Ginetz  prey  exposure  Naive f r y m i g r a t i n g  suffered  g r e a t e r m o r t a l i t i e s compared previously  nightly fry,  the  presence yet  evening, indicate  predators, predators  and  many  that naive  difference  p a t t e r n s of predators  mortality.  previous  experience  experienced to  with  Finally, f r y found and  in  was  showed predator  increased fry  stream  channels f r y , those  Secondly, and  downstream out  and  In  behavioral latter  response  fry  to  in (1)  were  the  naive  more  more  stimulus,  suffered  of  of  a  f r y , lacking  naive  compact ftlsc,  to  f r y were  susceptible of  the  migration  presence  fry  observations  a  span  the n a i v e  naive  t h a t the  to form  time  the  rapidly; over  d a n g e r o u s . The  (2)  suggested  short  the  experienced  varies.  spread  respects:  predators,  the  1976) to  predatcrs.  do  fry?  to experienced  potentially  two  This  the  predatcrs  fry)  influence  S larkin  f r y migrated  experienced  in  what  reaction  f r y became aware i n a  and  (sockeye  to  limit  non-migrants r e s u l t e d . T h i s tended  in migrating,  greater  or  migration  more  differed  predation.  the  prey  p a t t e r n of n a i v e  naive  w h i c h was  naive  somewhat s l o w e r  pricr  migration  hence  artificial  with  absence/presence  of  without  seme  interacted  downstream  in  down  a l l operate  to predators  survival.  having  do  of  transit  and  movement of  1972; of  how effect,  downstream  features  1962)  risk  rapid  schooling behavior,  (Ginetz  First,  t c reduce the  environment,  However,  occurs,  have upon t h e  developed dark  (Hartman  i f encounter  predators  a  subsequent  survival  predator-prey react  apparently  to and  schools  experienced  81  individuals fry  are  display  more  Thus  (1)  naive  predators,  and  as  to naive  compared  is  fry  fry  stimulation  and  Working  and  this  this  response  consistent (ca.  effect to  predator  an  once gained  (3)  i t  f r y changes f o l l o w i n g facilitating  migration.  of experience  the  and  (3) t h e  were  further  fry  change i n  supported  l e d t c the  relationship.  The  results  Ginetzs*  (1972)  development  presented work,  percent f r y  f r y m i g r a t i n g down p r e d a t o r percent  (2)  of  gravel.  and  to the  various  upon the f r y ;  ( S e c t i o n A)  of  of  work  of the b e h a v i o r a l composition  (1S76),  of naive egual  experimental  three general areas  p a t t e r n emerging; t h a t i s , the  approximately  encounters.  examinaticn  particular,  experiments  re-examination  10-30%)  (2)  information  e t c . ) , hence  various predators;  of Ginetz S L a r k i n  a  system, the  continued  upon emergence from  a rather interesting plus  of p r e d a t o r s .  b a s e l i n e of r e s u l t s ,  investigation  the  Preliminary results  paper  an  f r y , and,  behavior  rate  s u c c e s s f u l downstream  of f r y behavior. In  fry  m i g r a t i n g downstream a t a f a s t e r  Non-migrants  with  (1)  the  1976) of  encounters,  aspects  behavior  naive  Ginetz S l a r k i n cf  p a t t e r n of naive  in  naive  stimulated;  presence  i n subsequent  covered  covered:  aware  a predator  predator  survival  ( G i n e t z 1972;  become  by  behavior  (eg.  E. M i g r a n t s  when  f r y i n the absence  experience  t h a t the  greater  their  respond  somehow u t i l i z e d  appears  movement  active.  summarizing  results:  naive  much l e s s  non-migrants  the of  within,  shewed  a  mortality  channels  (i.e. fry  was not  82  migrating  downstream  predators.  From t h i s  ones which would  overnight) i t was  migrating  in  the absence  s p e c u l a t e d t h a t the f r y  be c a t e g o r i z e d a s  eaten  of  were  non-migrants i n p r e d a t o r - f r e e  channels. To  investigate  (Section of  E)  naive  types,  were d e s i g n e d f r y . I t was  fast  tendency fast  this  and  relationship,  composition  p o s t u l a t e d t h a t t h e r e a r e two  general f r y  slow  migrants.  Eesults  -  those  migrating  and,  downstream  over n i g h t ,  (mean p e r c e n t a g e  further  results,  migration  migrants/non-migrants individually, difference  slow  downstream  hours,  for  a  Two  guestions arise,  why  a r e t h e r e two  behavior  fixed, which  these  in  groups:  the  not  peak  migrating  = 24.3%). C o n s i s t e n c y  and  and slow  of  percent  migrants  that there i s a  tested  distinct  f r y types. origin  of t h i s  difference,  emerging? T h e . o r i g i n may  i s i n f l u e n c e d or d i r e c t e d  variables.  experience  twc  (b) e n v i r o n m e n t a l l y i n d u c e d ,  i m p l i e s t h a t the f r y l a c k as,  fast  what i s t h e  types  -  positive  during  patterns  both  between t h e s e twc  environmental  migrants  f a v o r s the i n t e r p r e t a t i o n  (a) g e n e t i c a l l y  such  showed  f o r n a i v e f r y t o be c h a r a c t e r i z e d by t h e s e  migrants  innate  experiments  t o examine t h e b e h a v i o r a l  migration  and,  succeeding  In t h i s  case,  (c)  a variety  with  an of  induction  and/or a b i o t i c  interactions  either:  or  environmental  similar biotic light,  by  be  stimuli;  other  f r y or  predators, etc.. The  e x i s t e n c e o f t h e s e twc  to i n t e r p r e t . ncr-migrants in  First, and  from  f r y types  the noted  relationship  p e r c e n t e a t e n , t h e slow  numbers s u f f i c i e n t  to  satiate  i s much mere  migrants  predators  difficult  between may  be  residing  percent present in  the  83  streams. is  Hence, a c e r t a i n  somehow more l i k e l y  patterns thus  of t h e s e  attracting  schools  while  erratic fast  p r e d a t o r s . The  migrants  be  genetically  not  fcrm  This  The  similar  experiences,  whereas slow having  between  migrants;  such  may  is  migrations  result  remain  to  behavior  lack of  migrants) i n a low  schccl  cf  have  light  more  had  regime  efficiently,  i n the e v e n i n g ,  with  other f r y .  certain  not  Another  which  differs  proportion  maternal net  may  similar  join  schooling  yelk,  (slow  or  poor  with l a r g e r sockeye  fast  fry  in  tanks.  possibility  (McCart  1967;  are  a less  noted  from  characteristic  T h e s e f r y may  to  study  to  schools  much l a t e r  example,  related  comprehensive  active  1972).  h e n c e may  emerge  s m a l l e r due  stream  Erannon  migrants  For  conditions.  A second  the  be  maintain  similar  as enough t i m e  physical  an o b s e r v a t i o n  laboratory  in  (Ginetz  associations;  fry.  may  incubation  1S72;  form  predators;  however, may  i s a simple  the  migrants)  types  of  may  t h e same o p p o r t u n i t y t o i n t e r a c t  possibility  may  cohesive  and  fry  migrants  distinct,  migrants  majority of naive f r y (fast  fry-fry  different,  a predator.  difference,  stimuli.  t o form  may  behavior  movements, when c o n f r o n t e d by  motionless i n the presence f o r experienced  f r y emergence  t o be consumed by  p r e d a t o r s . They may  contrast,  noted  c f any  m i g r a t i n g downstream, o r t h e y  swimming, w i t h In  slow  small percentage  1967,  to account  the  phencmenon  Byrne  1S72;  f o r the  1S71; Arnold  of upstream  E a l e i g h 1967, 1974;  review). It i s speculated that are  similar  reacting  as  to  upstream  normal  presence  o f two  fry  sockeye  fry  1971;  the l a t t e r the  slew  migrants;  downstream  Kelso  two  for a  migrants while  migrating  fast fry.  84  Erannon  (1972)  migrations and  are g e n e t i c a l l y  abiotic  ambient  stimuli,  light).  normally  concluded  (e.g. o l f a c t i o n ,  purpose  principally  migrants,  as  could a  a  be  certain  then e v e n t u a l l y  head  t e m p e r a t u r e and population may  of be  migrants. being  maintained  countering  I f rare conditions  downstream  fry  biotic  proportion  feature*  within  by b o t h  a spawning  between f r y emergence and l a k e maintain themselves  controlling  water  manyfold,  'safety  environmental conditions.  which  within  m a i n t a i n e d as u p s t r e a m  Their  timing  mechanisms  rased, yet influenced  Hence p o s s i b l y  downstream  genetically  that  unusual  o c c u r , such  as poor  plankton p r o d u c t i v i t y , f r y  t h e stream as a l a s t  f o r a lcnger  resort,  may  pericd, be  most  successful. Observations migrants  movement  with  Therefore  then  have  slowly  mcving  was i n i t i a l l y followed  by  much  m i g r a n t s may  an  also  b e h a v i o r c f upstream  movement  was  larger.  downstream,  t o upstream  eventual  i n fact  from  wandering  noted the existence  b e h a v i o r i n which  f r ytemporarily  Generally,  be  for a  to the  period  return  upstream.  upstream  migrants  and a r e i n i t i a l l y position  possibility  o f s t o c k which system  confined  holding  movement, k s e c o n d  migrants i n t h e Babine  eventually  the  downstream, o r m a i n t a i n i n g t h e i r  f r y result  (1S67)  f r y being  on  emerged t h e p r e v i o u s n i g h t  prior  upstream  upstream  t h e s e slow  which  those  that  the  pattern  pools,  stream,  1967)  indicated  daylight,  in  (McCart  (McCart  either in  the  i s that  normally  produce  1967).  McCart  o f an i n t e r m e d i a t e t y p e c f f r y held  a l o n g the stream edge but  move downstream.  Obviously  t h e problem  i s complex  and  further  experiments  85  a r e needed  to determine  artifact,  a  spawning fast  common  and s l o w  different  occurrence  migrants,  should  etc.),  absorption,  or  thus  differential  ?he E f f e c t  contain  two  types  are  of  be t e s t e d .  behavicrally  should  to  either  to  the  the  prey  comparing  versus  have shown d i f f e r e n t i a l Secondly,  early  may  responsible  adaptively  (size,  possible  to  speculation  increase  to the  the  yelk  compare  regarding  First,  group;  cf  enable  may  or,  actively  select  on  intrinsic innate  differences  in  consist  patterns internal repertoire  transforming f r y behavior.  The  differences.  behavior  of a  patterns  limited 1958).  external  of  1958)  vulnerability.  (Hoar  and  a  (Earns 1967). S t u d i e s o f  f r y (Earns 1967; y i b e r t  modify  two  intrinsic may  Predators  due t o  fry will  result  structural,  a  produced  for  stereotyped  or  one t c a n o t h e r  survival  hatched  be  select.,  experiences that  emergence, r e s p o n s e s rapidly  different  population.  within  natural  o f newly  may  behavioral  differentially  basis,  innate,  be  first  predation  f o r the 'abnormal'  be  also  rheotactic  Experience  differences,  hatchery  physically  both  predation,  in  predators  possibly  testing  differences  of  to  t h e s e s p e c u l a t i o n s need t h e n  It  Vulnerability  behavior  and p r e s e n t i n o t h e r  f r y a r e found  show i f t h e s e  etc.).  survival,  relative  each y e a r ,  experimental  (schooling behavior, reaction t c predators,  response,  select  f r y a r e , i f n o t an  s t o c k s . I f a l l emerging  Observations  !•  i f these  these  The  number  Following  stimuli  will  patterns;  thus  sooner  fry  acquire  86  non-instinctive  behavioral  better  chances f o r s u r v i v a l  are t h e i r  The is  importance  supported  predator-prey successful the  time  experimental  escape  o f a prey  of a defensive evidence  with  recognize  some form  (1977)  also  from  fryin Welty  a  predator-prey channel  those  encounter  containing  subseguent  significantly  migrated  It  was  the trough  very  fry  in  having  ( i . e . migrated  the  a similar survived  was s u f f i c i e n t  period. that  f a s t e r compared  t o modify  In  trained  experimental an  initial  down an a r t i f i c i a l  experience  from  fish.  and (2) i n p r e d a t o r c h a n n e l s ,  that  experienced  f r y could imitate, or  of untrained  (1972),  downstream s l i g h t l y  suggested  naive  by  behavicr.  different  p r e d a t o r s ) , had h i g h e r s u r v i v a l  encounters;  fry  within  (1)  with  found  Ginetz  found:  naive  able t o  action  avcidance  (1934) l i k e w i s e  As  individuals  evasive  time  the behavior  design  Those  short  could influence mentioned,  1966; G i n e t z 1972;  experience increases  take  that combining not  upon  Available  were t h e o r e t i c a l l y  and  f r y . This implied that experienced  1966).  predator encounters.  was  leadership experiments, fish  supposition that  found  extent  the  o f d a n g e r , and t h e r a p i d  of c o n d i t i o n e d predator  the s u r v i v a l rate  experienced  to a large  1962; Thompson  situation  with  experience,  (Thompson  encounter  survival  Dealing  learning  depend  response  initial  a dangerous  developing Patten  an  1934).  evidence. a  will  (Girsa  subseguent  survived  learn  as  1977) s u p p o r t s t h i s  survival  fry,  by  (Welty  experiences i n determining  encounters  experimental  that  of early  to external s t i m u l i , the  taken t o recognize the source  formation  Eatten  responses  to  rate  stream during  experienced naive f r y .  cf predatcr  encounter  the behavior o f naive  87  fry;  aquarium  observations substantiated behavioral  between t h e s e Part  twc f r y t y p e s .  of  the  experimental  research covered  was d e s i g n e d  to further  examine  encounters,  especially  f o l l o w i n g the d i v i s i o n  and  slow  migrants.  slightly  The r e s u l t s  differentiated  however b o t h  any  form  experience  procedure. light  A  experienced  behavior  attack,  predator suggested Ginetz in  to  or  on  the  of f r y i n t o  fast  experiment  only  and e x p e r i e n c e d frcm  migrants;  slow m i g r a n t s . I t  was n e t s u f f i c i e n t  migrants during  showed  had o b t a i n e d the  to some  experimental  f r y type  expected  were more f u l l y  with a  trends  for  discussed i n the  imply  that  encounters  innate  fry  were  situation  may  naive  fry  i s difficult  be s u f f i c i e n t behavior.  predator  (1972) f o u n d to  a  behavior  avoidance  stimulus;  f r y . Bowen  a  potentially  Actual  surrounding  to reinforce Two  modify  perceive  of  predator  fry  to  a  o r s t i m u l a t e an i n n a t e patterns  responses.  that experienced  f r y which f o r m e d  tc  to ascertain.  c o n t a c t , o r response  avoidance as  sufficient  In  f r y schooled  have  been  lab studies, prior  tc  and  with the s c h c c l s g e n e r a l l y being  compact and c o n t a i n i n g a l l t h e f r y .  the  initial  predator-prey  r e f e r e n c e s and t i e f o r e g o i n g e x p e r i m e n t s  response  naive  of  compounding  whole  i n the r e s u l t s  section.  visual  predator  fast  experiment,  p a t t e r n s . How t h e  dangerous  an  different  or,  f r y . These r e s u l t s  appropriate  predator  prior  second  effect  the experience  difference,  treatment,  Cited  of  were s i g n i f i c a n t l y  register  the  between f a s t  was s p e c u l a t e d t h a t e i t h e r  of  differences  This  loose aggregations  (1931) and W i l l i a m s  (1964)  was  in  contrast  to  seldom c o n t a i n i n g a l l suggested  that  this  88  aggregating  behavior  within  the  subject  tc greater  1S61;  confines,  Milinski  trout  within within  they  the  pursued  Ware  mcving  targets;  thus  anti-predator  prey  a  in  under  study  numerous b e i n g  with  greater  still  cf  &  that  sockeye  fry,  'bewildered  success to  itself  motionless fry  upon  (Ginetz  1  were than  be  were  verified  naive  trcut  not  Eadakov  maintaining  whereas  that  fish  schocl  f r y remained  seems  numerous  Theoretically whereas  trout  able  S to  stationary  a more  t h a t the  encounters.  experimental  any  condition  efficient  streams,  the  are for  nidwater  f r y would r e s p o n d  that  within  predator  encounter  the  the  field  two  most  surface  feeders,  being  (McCart  most  verified  with  i n one  probability  will  of  of  the  fish  was  response.  In  large  avoidance  set  conclusive  confines a  1S67).  vigorously  t c shew any the  (Dill  to  predation  failed  i f fry  whitefish,  bottcm  partially  behavior  t c detect  cohabit,  fish  a  Within  mountain  encounter a  interest  generally  remainder  surmised  troughs, to  are  T h i s was  e x p e r i m e n t s , however t h e  elicits  predators.  and  whitefish  expected  of  species  trout  ill-adapted  I t was  i t was  to various  rainbow  morphologically  natural  not  appeared  found  fry,  system  sufficient  stimulus,  (1971)  the  differentially  differences.  schccl  experienced  a  of the  lab observations a  any  (Manteifel*  what k i n d o f e x p e r i e n c e  respond  trout  periphery  individual  remaining  determining  Thus i t was  to predation;  device.  modification  to  an  were more a c t i v e and  1S76) .  feeders,  the  attacked  during  recognize  1S69).  risk  pressures  group. Secondly,  stimulation  In  cn  predation  seldom  a school  Larkin  or  the  1977aSb). P e r s o n a l  predators  invariably  reduced  be  dependent  89  upon t h e  spatial  and  temporal  distribution  c f both  the  fry  and  were  not  predators.  !•  ?he  Significance  Ginetz necessary  of Schooling  (1972) f o u n d for  modifying  b e h a v i o r a l responses designed was  responsible the  for  predator encounters  to experienced  what f e a t u r e o f t h e stimulating  fry  alone  f r y b e h a v i o r . Enumerated  similar  to determine  subjecting  that  to  a  an  short  fry.  fry displayed An  enumeration  experience  time  experiment procedure  showed  period i n the  that  light  was  importance  of  sufficient. Lab light  experiments  in  features light,  the  early  much  time  spend  a  confident  with  reflected  by  the the  lcnger  time  in  an  new  initial  with  their  structure  by  Secondly,  .  This  individuals,  increases as  with  measured by  a  is with  Finally,  g r e a t e r number o f i n d i v i d u a l s  stabilizing,  as  f r y become more  behavior.  confidence  to  forming f r y  associations  of s c h o o l i n g a  the  Three  subjection  environment.  and  fry .  activity  i n random swimming  development  the  sockeye  nervous  surroundings  period,  these  p e r i o d i n the l i g h t ,  a decrease  d e v e l o p m e n t and  in  e x p l o r i n g the  interact  of with  spent  longer  subseguent fry  is  their  t o determine  behavior  were e v i d e n t . F i r s t ,  a s s o c i a t i o n s and they  attempted  as  for a school  decrease  activity. Throughout  behavioral Although  this  response  work, enhancing  schooling serves  i t  has  fry  been survival  assumed is  that  the  schccling.  many f u r c t i o n s i n many s p e c i e s , i t has  90  been  suggested  reaction 1974;  that  f o r predator  Eadakov  Easically,  schools  or  school  aassociate  The  with pink  during  day.  group,  Yet  considered  have  in fish-to-fish  the  the  school  light  spent  development  of  move  after to  a  an  to  receptor  1964;  gravel  a  loose  moving  be  (Breder  short  initial  in  column  that  as  a  the  prime  1951,  1967;  operating  Keenleyside  in  1955;  enabling and  with  forces  fry, is first,  each  other;  interaction finally,  involved  the  some amount  secondly,  and  fine  innate  in  the of  a minimum  tuning  propensity  for  of the  schooling.  Extrapolating they  only  motivational  interacting  behavioral responses;  found  and  1S73).  prime  intensity  the  (John  development of s c h o o l i n g i n these time  water  time  interact  associated  attraction,  1960) ; y e t not  Eadakov  which  s o c k e y e f r y b u r y t h e m s e l v e s i n the  is  Shaw  and  a period cf  prior  1967). V i s i o n  maintenance of  Thus  emergence, b u t  the  1948;  structure  capture.  until  in  Morrow  1960;  schooled  1S73). of  (1956)  remained  stimulus  Moulton  not  Cullen  effect',  and  Hoar  &  chance  l e a r n to  fry  sensory  the  f r y must  once the  they  (McCart  the  ever  defense  Pitcher  the  •confusion  individuals.  Following  experience,  lessen  successful attack  were  schcoling  aggregation,  produce a  is a  (Neill  1960;  develops, g r a d u a l l y  other fry  detection  detected,  newly emerged  exposure to l i g h t .  the  of  feature  & Eiffenburgh  Once  will  behavior  1960).  adaptive  aggregations  probability  Schooling  migrating  Erock  encounters.  movement o f t h e reduces the  prime  a v o i d a n c e and  1973;  predator-prey  (Shaw  the  to the  from t h e  streambeds,  as  f r y cemmence e m e r g e n c e ,  d a r k , deep i n t e r s t i c e s  of  the  gravel to  the  91  1  surface  l a y e r s , where a m b i e n t l i g h t  (Heard  1964)..  are  formed,  decline gravel any  light  available light, initial  intensity  move downstream  subsequent  those  the  facilitating  in and  In  at  predator encounter  rapidly  deeper  i n the  learning  form  compact  gravel  would be  through  observing  to  fry-tc-fry  schooling  some  degree  associations  behavior.  With  d u s k , t h e f r y emerge f r c m  i n extremely  f r y which r e i n f o r c e d t h e i r  would  penetrates  loose  or f r i g h t  night,  p r i o r t c emergence  s c h c c l s ; whereas t h o s e  which  somewhat s l o w e r a t f o l l o w i n g  the schooling f r y .  the  aggregates.,During  s t i m u l i that  associations  a  were suit,  92  LITERATURE  ALI,  M. A. 1959. The o c u l a r structure, retincmotor and photobehavioral r e s p o n s e s o f j u v e n i l e P a c i f i c s a l m o n . Can. J . Z o o l . 37:965-996.  AENOLD, G. P, 49:515-576. BAMS,  CITED  1974. E h e o t r o p i s m  in  fishes.  Eiol.  Eev.  E . A. 1969. A d a p t a t i o n s o f s o c k e y e s a l m c n a s s o c i a t e d w i t h i n c u b a t i o n i n s t r e a m g r a v e l s , p..71-87. I n T. G, N o r t h c o t e (ed.) Symposium on Salmon and T r o u t i n Streams. Univ. E r i t i s h C o l u m b i a , V a n c o u v e r , B.C. 388p.  EOSEN, E . S. 1931. The r o l e o f s e n s e o r g a n s i n a g g r e g a t i o n s o f Ameiurus m e l a s . E c c l , Monogr. 1:3-35. EEANNON, E. L . 1967. G e n e t i c c o n t r o l o f m i g r a t i n g b e h a v i o r o f newly emerged s o c k e y e salmon f r y . P r o g r . . Eep. i n t e r n a t . P a c i f i c Salmon F i s h . Comm. ,No. 16, 3 1 p . EE ANNON, E. L. . 1972. Mechanisms c o n t r o l l i n g m i g r a t i o n o f s o c k e y e salmcn f r y . B u l l , i n t e r n a t . P a c i f i c Salmcn F i s h . Comm. No. 21, 86p. EEEDEE, C. M., J r . . 1 9 5 1 . S t u d i e s on t h e s t r u c t u r e s c h o o l . B u l l . Am. Mus. n a t . H i s t . 98:7-27. EEEDEE, C. M. , J r . ,1967. On t h e s u r v i v a l Z o o l o g i c a , N.Y. 52:25-40.  of  value o f f i s h  the  fish  schools.  EEETT, J , E . 8 M. A. A l l . 1S58. Seme observations cn t h e structure and p h o t o m e c h a n i c a l r e s p o n s e s of the P a c i f i c salmon r e t i n a . J . F i s h , E e s . Board C a n . 15:8 15-829. EEOCK, V. E. S E . H. EIFFENBUEGH. 1960. F i s h schooling: a possible f a c t o r i n r e d u c i n g p r e d a t i c n , J . Cons. perm. i n t . E x p l o r . Mer. 25:307-317. EYENE, J . E. 1971. P h o t o p e r i o d i c a c t i v i t y changes i n juvenile sockeye salmon (Oncorhynchus nerka). Can. J . Z o o l . 49:1155-1158. D H L , L. M. 1969. F u l t o n R i v e r f r y q u a l i t y and e c o l o g y program: report o f 1968 s t u d i e s . Dept. , o f F i s h e r i e s o f Canada, V a n c o u v e r , B.C., 185p. GINETZ, R. a. J . 1972, Some f a c t o r s affecting rainbow trout (Salmo g a i r d n e r i i p r e d a t i o n on m i g r a n t s o c k e y e salmcn f r y . M.Sc. T h e s i s , U n i v . B r i t i s h C o l u m b i a , V a n c o u v e r , B.C., 89p.  93  GIKETZ, R. M, & P, A. LARKIN. 1976, Factors affecting rainbow t r o u t (Salmo g a i r d n e r i ) p r e d a t i o n on m i g r a n t f r y c f s o c k e y e salmon (Oncorhynchus nerka). J . Fish. E e s . Board Can. 33:19-24. GIRSA, L . L . .1962. On t h e d e c r e a s e o f p r e d a t i o n by p r e d a t o r s on small fish with respect t o the formation i n small f i s h of c o n d i t i o n d e f e n s e mechanisms. V o p r , I k h t i o l . 2 :747-749. HARTMAN, G. F., T. G. NGRTHCOTE S C . C. LINDSEY. 1962. Comparison of inlet and o u t l e t spawning runs of rainbow t r o u t i n Loon L a k e , B r i t i s h C o l u m b i a , J . F i s h . Res. Board Can. 19:173-200. HARTMAN, W..L., C. W. STRICKLAND 6 D. T. HGCFES. 1962. S u r v i v a l and b e h a v i o r o f s o c k e y e s a l m c n f r y m i g r a t i n g into Erocks Lake, Alaska..Trans.,Amer. F i s h . Soc..91:123-139. HARTMAN, W. L., W. R. HEARD 8 B. DRUCKER. 1967, M i g r a t o r y t e h a v i o r o f s o c k e y e salmon f r y and s m e l t s . J . F i s h . Res. E o a r d Can. 24:2069-2099. HEAIY, M. C. 1972. Individual sockeye salmon (Oncorhynchus B o a r d C a n . No. 297, 16p. HEARD, W. salmcn  b e h a v i o r and group t e h a v i o r i n n e r k a ) . . T e c h . Sep. F i s h . Res.  E. .. 1964. P h o t o t a c t i c behaviour c f emerging f r y . Anim. Behav. 12:382-388.  HEMMINGS, C. C. Exp. B i o l .  1966. O l f a c t i o n 45:449-464.  and v i s i o n  in fish  sockeye  schccling.  J.  HO A E,  W. S. 1951. The b e h a v i o u r o f chum, p i n k and coho salmon i n r e l a t i o n t o t h e i r seaward m i g r a t i o n . J . F i s h . Ees. Board Can. 8: 241-263.  HOB E,  W. Eev.  S. 1953. C o n t r o l 28:437-452.  and t i m i n g  of f i s h  migration.  Biol.  HOAE, W. S. 1954. The b e h a v i o u r o f j u v e n i l e E a c i f i c s a l m o n , w i t h p a r t i c u l a r r e f e r e n c e t o the sockeye (Oncorhynchus nerka). J . F i s h . E e s . B o a r d C a n . 11:69-97. HOAE,  W. S. 1956, The behaviour o f m i g r a t i n g p i n k and chum salmon f r y . J . F i s h . E e s . ..Board Can. . 13:309^325.  HOAE, W. S. 1958, The e v o l u t i o n of migratory behaviour among juvenile salmon o f t h e genus O n c o r h y n c h u s . . J . F i s h . E e s . E o a r d Can. 15:391-428. HUNTEE, J . G. 1959. S u r v i v a l salmcn in a coastal 16:835-885.  and p r o d u c t i o n of stream. J. Fish.  pink and Res. Board  chum Can.  94  JOHN,  K. E. 1964. Illumination, Astyanax mexicanus (Fillipi)• 2l7l453-1473.  vision, and schooling of J . Fish. E e s . Board Can.  JCHNSON, W. E. 1965. On mechanisms cf self-regulation of p o p u l a t i o n abundance i n O n c p r h y n c h u s n e r k a . M i t t . i n t . V e r . L i m n o l . 13:66-87. KEENLEYSIDE, of f i s h .  M. H. A. 1955. Some a s p e c t s o f s c h o o l i n g B e h a v i o u r 8:183-247,  behaviour  KELSO, B. W. 1972., The genetic and e n v i r c r m e n t a l mechanisms controlling t h e l a k e w a r d m i g r a t i o n of young r a i n b o w t r o u t (Salmo g a i r d n e r i i from o u t l e t and inlet rearing streams. M.Sc. T h e s i s , O n i v . B r i t i s h j t o l u m b i a , V a n c o u v e r , B.C., 75p. MANTEIFEL *, B. P. 8 D. V. EADAKOV. 1961. The adaptive s i g n i f i c a n c e o f s c h o o l i n g b e h a v i o u r i n f i s h e s . R u s s i a n Bev. E i c l . 50:338-345. McCART, P. 1967. B e h a v i o u r and e c o l o g y o f s o c k e y e salmon f r y i n t h e B a b i n e R i v e r . J . F i s h . R e s . B o a r d Can. 24:375-428. MCDONALD, J . 1960. The b e h a v i o u r of P a c i f i c salmon f r y d u r i n g t h e i r downstream migration t c freshwater and saltwater n u r s e r y a r e a s . J , F i s h . Res. Board Can. 17:655-676. fclLINSKI, M. 1977a. E x p e r i m e n t s on t h e s e l e c t i o n against s p a t i a l oddity of t h e i r prey. Z. 43:311-325.  by p r e d a t o r s Tierpsychcl.  K I L O S KI, M.' 1977b. Do a l l members o f a swarm s u f f e r t h e same p r e d a t i o n ? Z. T i e r p s y c h o l . 45:389-413. MOEICW, J . E. 1948. S c h o o l i n g b e h a v i o r i n B i c l . 23:27-38.  fishes.  MCULTON, J . M. 1960. Swimming sounds f i s h e s . B i c l . „ B u l l . 119:2.10-223.  and  Quart;  Eev.  the s c h o o l i n g o f  NEAVE, F. 1953. P r i n c i p l e s a f f e c t i n g t h e s i z e o f p i n k and chum salmon p o p u l a t i o n s i n E r i t i s h C o l u m b i a . _ J . F i s h . B e s . Board Can, 9:450-491. NEAVE, F. 1955; Notes on t h e seaward m i g r a t i o n o f p i n k and chum salmon f r y . J . F i s h . Res. B o a r d Can. 12:369-374. NEILL, S. E. S t . J . & J . M. CULLEN. 1 9 7 4 . . E x p e r i m e n t s on whether s c h o o l i n g hy t h e i r p r e y a f f e c t s t h e h u n t i n g behaviour of cephalopods and fish predators. J. Zool., Lond. 172:549-569. EATTEN, E. G.' 1977. Body s i z e and l e a r n e d a v o i d a n c e as factors affecting predation on c o h o salmon, O n c c r h y n c h u s k i s u t c h , f r y by t o r r e n t s c u l p i n , C o t t u s r h o t h e u s . F i s h . B u l l . , 0. S. 75:457-459.  95  FITCHEE, T. J . 1973. The t h r e e - d i m e n s i o n a l s t r u c t u r e o f schools in t h e minnow, Phoxinus phoxinus (L.) . Anim.. Eehav. 21:673-686. FBITCHAED, A. L. 1944. P h y s i c a l c h a r a c t e r i s t i c s and b e h a v i o r o f pick salmon f r y at McClintcn C r e e k , B.C. J . F i s h . E e s . B o a r d Can. 6:217-227. EAEAKOV, D. V. 1973. S c h o o l i n g i n t h e e c o l o g y of f i s h . , Engl. Transl., I s r a e l Program S c i . T r a n s l . P u t l . , John W i l e y and S o n s , N. Y. , 173p. BAIEIGH, E . F. 1967. G e n e t i c c o n t r o l i n t h e l a k e w a r d migrations of sockeye salmcn (Oncorhynchus nerka) f r y . J . F i s h . E e s . E o a r d C a n . 24:2613-2622. EALEIGH, E. F. 1971. I n n a t e c o n t r o l o f m i g r a t i o n s o f s a l m c n and trout f r y from natal gravels t o rearing areas. Ecology 52:291-297. EICKEE, W. E. 1941. The c o n s u m p t i o n o f young sockeye f r y by p r e d a c e o u s f i s h . J . F i s h . E e s . B o a r d Can. 5:293-313. EOCS,  J . F. 1960. P r e d a t i o n c f young echo s a l m o n c n s o c k e y e salmon f r y a t C h i g n i k , Alaska. T r a n s . Am. F i s h . . Soc. 89:377-379.  SHAW,  E . 1960. The d e v e l o p m e n t P h y s i o l . Z o o l . 33:79-86.  of schooling  behavior  i n fishes.  STEVEN, D, M. 1959. S t u d i e s on t h e s h o a l i n g b e h a v i o r o f fishes. I. B e s p o n s e s o f two s p e c i e s t o c h a n g e s c f i l l u m i n a t i o n and t o o l f a c t o r y s t i m u l i . J . E x p , B i o l , 36:261-280. THCKPSON, E. B. 1966. E f f e c t s o f p r e d a t o r a v o i d a n c e c o n d i t i o n i n g on the post-release survival rate of artificially p r o p a g a t e d s a l m o n . Ph.p. T h e s i s , U n i v . W a s h i n g t o n , S e a t t l e , Wash., 155p. VIBEET, et WAEE,  E. 1958. C r i t e r e s e t t e s t s de r u s t i c a t e c h e z l e s t r u i t e s l e s saumons. V e r b , i n t . V e r , L i m n o l , 13:758-764.  D.. M. 1971. The p r e d a t o r y (Salmo g a i r d n e r i ) . Ph.D. T h e s i s , V a n c o u v e r , B.C., 156p.  WE1TY, J . C. 1934. E x p e r i m e n t s P h y s i o l . Z o o l . 7:85-128.  b e h a v i o u r o f rainbow t r o u t Univ. British Columbia,  i n group b e h a v i o r  i n fishes.  WILLIAMS, G. Cm 1 9 6 4 . „ M e a s u r e m e n t o f c o n s o c i a t i c n among fishes and comments on t h e e v o l u t i o n o f s c h o o l i n g . P u b l . Mus. M i c h i g a n S t a t e U n i v . , B i o l . . Ser. 2:349-384.  APPENDICES  APPENDIX I : Three-way analysis of variance cn a r c s i n e transformed values of percent m i g r a n t s per sampling time period. Analysis compared d i f f e r e n c e s i n migration pattern f o r naive f r y i n a r t i f i c i a l s t r e a m c h a n n e l s w i t h and w i t h o u t predators.  Source  of V a r i a t i o n  Factors: Time p e r i o d s Days P r e d a t o r s +/Interactions: Time p e r . x Days Time p e r . x P r e d s , Days x P r e d s . . Error Total  1 DF | j j I I  MS  5 I 2 I 2 1  I  F  | | 793,.56 | 38.32 | 6.88 | 603.49 | 60.35 1757.80 | 175.78 24.1 8 | 6,05 3967.82 76.65 13.76  4 I 1 20 | 1149.21  I  I  |  | | | 10 | | 10 | I  SS  53 | 7592.91  I  !  I I  13.81 0.67 | 0. 12  | | | |  1.05 3.06 0. 11  **  *  57.46  !  ;  * = s i g n i f i c a n t a t 5% l e v e l ** = s i g n i f i c a n t a t 15? l e v e l Time p e r i o d s = s a m p l i n g i n t e r v a l s (0.5 hour p e r i o d s ) Days = e x p e r i m e n t replicates P r e d a t o r s +/- = stream channels with (rainbow, w h i t e f i s h ) and w i t h o u t p r e d a t o r s  97  AEgENDix  Source  I I : Four-way analysis of variance on arcsine transformed values of percent migrants per sampling time period. Analysis compared differences i n migration pattern f o r various e x p e r i e n c e d f r y , from t h e two spawning c h a n n e l s , i n a r t i f i c i a l stream c h a n n e l s with and without predators.  of V a r i a t i o n  Eactors: Time p e r i o d s Experience Channels P r e d a t o r s +/Interactions: Time p e r . x E x p e r . Time p e r . x Chan. E x p e r . x Chan, Time p e r . x P r e d s . Exper. x Preds. Chan, x P r e d s . Error Total  I  1 DF H--  |  | |  1  I I  SS  | 4 | 20457.58 4 39.02 1 | 59.95 228.04 2  i 164 i 4 i j 88 i 2 i 96 i  | | | |  j|  |  )  1921.65 545.41 119.89 730.23 104.93 199.44 5640.67  ! 149 ! 30046.81  |  MS  I |  |  F •  * •^  J  | 5114. 39 | 9. 76 | 59. 95 | 114. 02 | | I | | | |  I -4-  120. 10 136. 35 29. 97 91. 28 13. 12 99. 72 58. 76  | 87. 0 4 | 0. 17 | 1. 02 1. 94 I | | 2. 04 | 2. 32 | 0. 51 | 1. 55 | 0. 22 | 1. 70 I I  I * = s i g n i f i c a n t a t 5% l e v e l ** = s i g n i f i c a n t a t 1% l e v e l Time p e r i o d s = s a m p l i n g i n t e r v a l s (0.5 h o u r p e r i o d s ) E x p e r i e n c e s = f r y t h a t are i n e x p e r i e n c e d , enumerated, s h o c k e d , and e x p e r i e n c e d t o p r e d a t o r s C h a n n e l s = Spawning C h a n n e l s No. 1 and 2 P r e d a t o r s +/- = s t r e a m channels with (rainbow, w h i t e f i s h ) and w i t h o u t p r e d a t o r s  *  ^  *  **  *  98  APPENDIX I I I : Three-way analysis of variance on arcsine t r a n s f o r m e d v a l u e s o f p e r c e n t m i g r a n t s , n o n - m i g r a n t s and eaten. Analysis compared differences i n these values f o r various e x p e r i e n c e d f r y , f r o m t h e two s p a w n i n g c h a n n e l s , in artificial s t r e a m c h a n n e l s w i t h and w i t h o u t p r e d a t o r s . F a c t o r s as d e s c r i b e d i n Appendix I I . (a)  % MIGSftNTS  Source  of Variation  | | | | | |  Experience Channels Predators v E x p e r . x Chans. Exper. x Preds, Chans, x P r e d s . Error Total (b) r  1  !  —  — —  r  Source  of Variation  1— —  MS  F  I  I  | 91. 37 | 2. 42 | 42. 48 | 1. 12 | 548. 28 | 14. 49 | 62. 10 | 1. 64 | 50. 11 I 1. 33 | 501. 29 | 13. 25  365.48 42.48 I 1096,55 248.41 I 4 00,90 I I 1002.58 302.67 I 1 I  !  29 | 3459.07  1  ^  ** | ** |  37. 83 |  I  -  1—  1  (C) %  ! 29  ——  1  *  T  i  MS  T  ;  |  1  F  27.36 | 0.44 | | 4.40 | 0.07 | 1289.02 | 20.61 | 7.40 | 0. 12 75.47 | 1.21 | | 81.73 | 1.31 | 62.54 I  ** |  [  | 3988.99 |  ••  T  +  |  +  1 I  I I I I  I  ^ 1 1I 1i 4 4I 1  "I  I 19 I  1  1  | DF  SS  ,_ 215.63 423.84 946,28 210.76 252.06 12.09 197.44  | +  | | | | i | |  T —  MS  1  r  | F 4— 53.91 | 1.09 423,84 | 8.59 946.28 | 19.17 52.69 j 1.07 63,01 | 1.28 0.25 12.09 | 49.36  ,  I  L__  I  A * **  2258.10 | I.  I  i 1  EATEN  | Source of V a r i a t i o n I „ . Experience Channels P r e d a t o r s */E x p e r . x Chans. Exper. x Preds. Chans, x E r e d s . Error Total  1  SS 1 DF | • I —-+ —r109.43 4 1 4.40 | 1 | | 2 I 2578. 04 29.59 | 4 I | 8 I 603.79 163.47 | 2 I 500128 8 I  Total  t  4 1 2 4 8 2 8  1  % NCN-MIGB BNTS  Experience Channels P r e d a t o r s +/Exper. x Chans. Exper. x Preds. Chans, x P r e d s . Error  r  SS  1 DF |  t  J  'APPENDIX IV:  Two-way analysis of variance on arcsine transformed values of percent migrants per sampling time period. A n a l y s i s compared differences i n migration pattern f o r various experienced f r y . ,  Source o f V a r i a t i o n  i  rI DF |  Factors: Experience Time p e r i o d s Interaction Error  4 6 24 35  MS  70.64 2516.03 1673.56 1183.24  17.66 419.33 69.73 33.81  "I"  I  0.52 12.40 2.06  ** *  I 6S | 5443,46 I  Total l  SS  J  1  J  * = s i g n i f i c a n t a t 5% l e v e l ** = s i g n i f i c a n t a t 1% l e v e l E x p e r i e n c e s = as d e s c r i b e d i n T a b l e IV Time p e r i o d s = s a m p l i n g i n t e r v a l s (0.5 h o u r  L-.  ,  periods)  .—  100  APPENDIX V: Three-way analysis of variance on arcsine transformed values of p e r c e n t migrants per time p e r i o d . A n a l y s i s examined migration p a t t e r n s of various f r y types migrating through a r t i f i c i a l s t r e a m c h a n n e l s w i t h and w i t h o u t p r e d a t o r s .  r-—  |  Source  T  of Variation  | DF  1  |  Factors: I Time p e r i o d s | 6 1 42334.89 Fry type | 170.55 2 1 P r e d a t o r s +/| 2 05.44 3 1 j Interactions: | Time p e r . x F r y | 12 | 3704.47 Time p e r , x P r e d s , | 18 | 3361.79 Fry x Preds. j 9U72 6 i Error I 252 | 1 4 0 0 1 . 5 7 Total  | 279  |  T  "i  SS'  63870.42  |  MS  |  1  •  F  I  I  I | | | j | | | |  |  7055.81 85.27 68.49 308.71 186.77 15.29 60.35  | | | j | | | |  116.9 1 1,4 1 1 . 13  **  5 . 12 3 . 10 0.25  ** **  I  ** = s i g n i f i c a n t a t ^% l e v e l Time p e r i o d s = s a m p l i n g i n t e r v a l s ( 0 . 5 h o u r p e r i o d s ) F r y t y p e = f a s t m i g r a n t s , slow m i g r a n t s and e x p e r i e n c e d P r e d a t o r s +/- = s t r e a m c h a n n e l s with (rainbow, whitefish, rainrow/whitefish combination) and w i t h o u t predators  APPENDIX V I :  (a) FAST  r  Two-way analysis of variance cn arcsine transformed values of percent m i g r a n t s per sampling p e r i o d ; examining e f f e c t c f p r e d a t o r s on m i g r a t i o n p a t t e r n o f t h e t h r e e f r y t y p e s .  MIGRANTS:  "  T  1  ——  •* l  T  | Source o f V a r i a t i o n | DF \ SS | MS t — ' — — - + * — - t — Time p e r i o d s 6 17147.06 | 2857.84 P r e d a t o r s +/3 119.31 | 39.77 Time p e r . . x Preds. 18 35.54 639.66 | Frror €3 69.33 4367.67 | I Total 90 22273.70 | (jb)  I  '  1  I  F 41.22 0.5 7 0.5 1  *  ^  SLOW MIGEANTS:  Source o f V a r i a t i o n  j DF  Time p e r i o d s P r e d a t o r s +/Time p e r . x Preds. Frror  j  SS  MS  6 | 11412.38 | 1902.06 55,. 5 8 | 18,53 3 I 18 | 77.49 1394.86 | 60. 17 77 4633.04 | I I 104 | 174S5.87 |  •1.6 1 0.3 1 1.29  r  Total  (£.)  |  EXPERIENCED:  Source of V a r i a t i o n  | DF  |  Time p e r i o d s P r e d a t o r s +/Time p e r . x P r e d s . Frror  | | | |  6 3 18 56  I 17479.91 | 2913.32 122.27 | 40.76 I | 3441.01 | 191.17 | 2887.11 I 51.56  |  83 | 23930.30 | L.  Total  .  .  * = significant  j__  a t 5% l e v e l  SS  |  MS  I  F  | 56.51 | 0.79 | 3.7 1 ]  I *  I  *  |  102  SPPENDIX VII;  (a) r-—  |  Two-way analysis of variance on arcsine transformed v a l u e s o f p e r c e n t m i g r a n t s per sampling time p e r i o d ; examining the response of each f r y t y p e w i t h i n t h e v a r i o u s p r e d a t o r regimes.  EAINEOW TJ3CDT  Source  of Variation  Time p e r i o d s Fry type Time p e r . x F r y Error Total  (t)  Source  I  Time p e r i o d s Fry type Time p e r . , x F r y Error Total  (C)  of Variation  6 | 2 | 12 | ^2 | | 62 |  I I I I  SS  MS  12690.67 104.57 1C98.84 3541.09 17435.18  2115. 11 52.29 91.57 84.31  "T——r-  I  DF |  6 | 2| 12 | 63 | | 83 | I I I I  •  SS  MS  I  15564. 90 | 49.64 | 1577.94 | 3837.77 | 21030.25 |  -T-  I  2594. 15 | 45.59 24.82 | 0.41 131.50 | 2. 16 60.92 | I  RAINBOW TROOT/WHITEFISH  r  ~T  T  |  Source  I  Time p e r i o d s Fry type Time p e r . x F r y Error Total  I  T'  | DF |  WHITEFISH  i  i  T  of V a r i a t i o n  I  DF  |  I  | 2 | | 12 | | 42 | | 62 |  SS  97S7.78 100. 96 1978.70 2193.21 14070.65  |  T-  ^  MS  |  j 1632.96 | 31,27 | 50.48 | 0.97 | 164.89 | 3. 16 | 52.22 | I |  (d) NO PREDATORS I  Source  | | I I |  Time p e r i o d s Fry type Time p e r . x F r y Error Total  I  of V a r i a t i o n  | DF |  +-,--+. I 6| I 2| I 12 | I 49 | | 69 |  SS  7643.32 7.10 1162.73 2315.75 11128.90  |  MS  |  I  F  | 1273.89 | 26.95 | 3.55 | 0.08 | 96.89 | 2.05 | 47.26 | I |  * *  103  APPENDIX V I I I :  Source  Two-way a n a l y s i s of v a r i a n c e on arcsine transformed values of p e r c e n t migrants i n t h e f i r s t hour f o l l o w i n g t h e o n s e t o f the experiment. A n a l y s i s compared d i f f e r e n c e s in these values f o r each f r y type i n a r t i f i c i a l s t r e a m c h a n n e l s w i t h and w i t h o u t predators. An a posteriori Duncan's New M u l t i p l e Eange T e s t compared d i f f e r e n c e s i n t h e means f o r e a c h f a c t o r .  of Variation  |  —H P r e d a t o r s */1 2 3 Fry type 1 Preds. x Fry type 1 6 Error | 28 Total  |  'T-  DF I  I  SS  | | | |  2279.34 3505.17 1534.76 7158.70  39 |  15477.96  MS  I  1093.1 1 1752.58 255.79 255.67  4.28 6.86 1.00  * *  * = s i g n i f i c a n t a t 5% l e v e l P r e d a t o r s +/- = s t r e a m c h a n n e l s w i t h ( r a i n b o w , whitefish, r a i n b o w / w h i t e f i s h c o m b i n a t i o n ) and w i t h o u t predators Fry t y p e = f a s t m i g r a n t s , slow m i g r a n t s and e x p e r i e n c e d  D.N.M.E.T. (a) PREDATOES +/-:  Bain, 73.87  White. 70.22  Ea,/Wh. 67.23  No p r e d . 50. 15  N.S, (b) FEY TYPE:  F a s t migs. 72.79  Experienced 72.39  N.S. N.S. = mean v a l u e s n o t s i g n i f i c a n t l y  different  Slow migs, 52.27  104  APPENDIX I X :  Two-way analysis of variance cn arcsine transformed values of percent migrants, non-migrants and l o s t or eaten. Analysis compared d i f f e r e n c e s i n t h e s e v a l u e s f o r t h e f r y t y p e s i n a r t i f i c i a l stream c h a n n e l s with and w i t h o u t p r e d a t o r s .  (a) J MIGEANTS: | Source  of V a r i a t i o n  DF |  Fry type P r e d a t o r s +/Fry type x Preds. Error I |  2 3 6 28 I  |  MS  |  F  _^ I I 764.86 | 382.43 | 4. 81 I 1019.29 | 339.77 | 4. 27 660.33 | 110.05 | 1. 38 I | 2226.15 | 7S.51 I  39 I  Total  I  SS  I  -I  *  j  4670.63 1  I  (t) % NON-MIGEANTS : -T  | Source o f V a r i a t i o n H | Fry type | P r e d a t o r s +/| F r y type x Preds, I Error |  | DF |  +  — T -  SS  3  6 28  '  |  923.57 1801.44 317.94 2530.66  2  Total  '  T  MS  |  461.79 6C0.48 52.99 90,38  «•  F  H  5. 11 6.64 0. 59  39 | 5573,61  <C) i LOST o r EATEN  | Source  of V a r i a t i o n  f  I | J I  | DF  |  P  28.88 328.29 357.12 S95.02  2 3  Fry type P r e d a t o r s +/Fry type x Preds. Error  6  28  J Total l  SS  MS  (  F  |  +  L  39 -  —  I 1709,31 | _  -^J  14.44 109.43 59.52 35.54 ,  --J.  0.41 3.08 1.68  .  * = s i g n i f i c a n t a t 5$ l e v e l ** = s i g n i f i c a n t a t 1$ l e v e l F r y t y p e = f a s t m i g r a n t s , s l o w m i g r a n t s , and e x p e r i e n c e d P r e d a t o r s +/- = s t r e a m c h a n n e l s w i t h ( r a i n b o w , w h i t e f i s h , rainbow/whitefish combination) and without predators  105  APPENDIX X:  r  | y  Source _  Four-way analysis of variance cn arcsine transformed values of percent migrants per sampling time periods. Analysis compared d i f f e r e n c e s i n m i g r a t i o n p a t t e r n f o r the t h r e e fry types, subjected to three l i g h t conditions, in a r t i f i c i a l s t r e a m c h a n n e l s w i t h and w i t h o u t predators.  of V a r i a t i o n  Factors: Time p e r i o d s Fry types Hours l i g h t P r e d a t o r s +/Interactions: Time p e r . x F r y Time p e r . x L i g h t Fry type x L i g h t Time p e r . x P r e d s . Fry type x Preds. L i g h t x Preds. Error  | DF | ,. ,_4  SS  "  T  J  T-  MS  |  +  F  +  1  ^  1  5 1 14519.55 | 2903.91 | 49.4 1 ** 2| 51.56 | 25.78 | 0.44 2 32.72 | 16.36 | 0.28 1 I . 112.85 | 112.85 | 1.92  1 1 1  10 | 10 j 4' 5| 2 2| 64 |  I  I I  1307.68 879.62 35. 85 2293.28 8.88 85^53 3760.91  | | | | | | |  130.77 87.96 8.96 458.66 4.44 42.76 58.76  | | | | | | |  —  L  2.2 2 * 1.50 0.15 7.8 1 ** 0.08 0.73  107 | 23088.42 |  Total L  r  1  ,.  L^.  1  „  L  ,  * = s i g n i f i c a n t a t 5% l e v e l ** = s i g n i f i c a n t a t 1% l e v e l Time p e r i o d s = s a m p l i n g i n t e r v a l s (0.5 hour p e r i o d s ) F r y t y p e = f a s t m i g r a n t s , slow m i g r a n t s . a n d e x p e r i e n c e d Hours l i g h t = number o f h o u r s f r y were s u b j e c t e d to light t r e a t m e n t (0, 6, 24 h o u r s ) P r e d a t o r s +/- = s t r e a m c h a n n e l s ; with (rainbow, w h i t e f i s h , r a i n b o w / w h i t e f i s h c o m b i n a t i o n ) and without predators  I  106  APPENDIX XI: Three-way analysis of variance on arcsine transformed values of percent migrants i n f i r s t hour, migrants and n o n - m i g r a n t s . A n a l y s i s compared d i f f e r e n c e s i n t h e s e v a l u e s for the f r y types, subjected to three light conditions, m i g r a t i n g t h r o u g h s t r e a m c h a n n e l s w i t h and w i t h o u t p r e d a t o r s . (a) % MIGRANTS i n FIRST HOUR r  **  I Source  T  of Variation  1  T  1  | DF |  SS  Fry t y p e Hours l i g h t P r e d a t o r s +/Fry t y p e x L i g h t Fry type x Preds, L i g h t x Preds, Error  + +1 2 | 1050.64 1 2 | 1305.99 1 1 I 2300.68 360.01 I I 760.95 I 2| I 2 | 651.27 561.07 I *» I  Total  I 17 | 6990.61  a  -  T  1  '•  |  MS  |  F  |  MS  I  F  I  4 4— | 525.32 | 3.75 i 652.99 | 4.66 | 2300.68 | 16.40 | SO.00 | 0.64 | 380.47 | 2.71 | 325.63 | 2.32 | 140.27  i' (b) % I Source  MIGRANTS  of V a r i a t i o n  I DF |  2 2 1 4 2 2 4  Fry t y p e Hours l i g h t P r e d a t o r s +/Fry type x L i g h t Fry type x Preds. L i g h t x Preds. Error  Source  | 77.78 | 38.89 0.63 | 424.30 | 212,15 j 3.45 | 1332.66 | 1232.66 | 21.66 | 241.46 | 60.36 j 0.98 | 215.51 | 1C7.76 | 1.75 | 147.72 | 73.86 | 1.20 | 246.07 | 61.52  |  **  j  NON-MIGRANTS of Variation  Fry type Hours l i g h t P r e d a t o r s +/Fry type x L i g h t Fry type x Preds. Light x Preds. Error Total l  i  17 | 2685.48  Total  (C) 2  SS  .  | DF i  HI I I I I I I  2 2 1 4 2 2 4  | | | | | | |  SS  |  MS  |  | 17 | 2965.69 |  j  I  L  I  F  (._ 437.55 | 18.77 | 0. 26 349.80 | 174.90 | 2.40 1550.50 | 1550 50 | 21.29 ** 230.41 | 57.60 | 0.79 285.31 | 142.66 | 1.96 220.87 | 110.43 | 1. 52 291.25 | 72.81 | L  I  107  APPENDIX X I I :  Collection, fertilization p r o c e d u r e s f o r s a l m o n r o e and laboratory studies.  Approximately then  divided  Concurrently, watertight securely by  20,000 eggs  evenly sperm  4  into  four  from t w e l v e  oz g l a s s  packed  were s t r i p p e d  into  jars.  a coleman  water  males  from  seven f e m a l e s ,  each and  plastic  divided  For t r a n s p o r t ,  i c e t c m a i n t a i n low t e m p e r a t u r e s ,  incubation utilized in  tight  was  cooler;  and fry  tags.  into  four  the c o n t a i n e r s container  were  surrounded  newspaper  to  absorb  of  British  shock. Fertilization Columbia nixing in  eight the  water  The  eggs  eggs  and  incubation  f o r 30  adeguate  12.5  the  oxygen. meter),  tap  (3) t h e n p l a c i n g  and  II  5  of:  (1)  (2) b a t h i n g  the  eggs  the f e r t i l i z e d  eggs  flow i n c u b a t o r ;  - 4.0  gal/ffin,  4.0°  C.  water,  A  and  the  water  dechlcrinator  while  a  water  temperature reduced  a t a p p r o x i m a t e l y 100%  through  the  incubator,  a  any  provided  (HACH k i t ' and  remained  micron  the  baffle-system  measured w e e k l y ,  YSI  saturation.  Cuno  (G78B2)  filter  effectively  removed  filters  were c h a n g e d  periodically,  particles  from  t h e w a t e r . The  depending  upon  sediment  malachite  green s o l u t i o n ,  (3 gms  cf  80 m l / t r e a t m e n t  f o r f l o w c f 3,5  water;  consisted  trays.  Oxygen was  tc circulating  Kicrc-Kleam  ~  It  seconds;  developed i n a v e r t i c a l  from in  at the U n i v e r s i t y  collection.  sperm  a p p r o x i m a t e l y 3,5  chlorine  Prior  hours a f t e r  Heath  decreasing  Oxygen  conducted  f o r 3 m i n u t e s ; and  into four  flow  was  build-up.  To p r e v e n t f u n g u s cf c r y s t a l -  infection,  dissolved 4.0  most  in 1  gal/min) ,  a  litre was  108  added  every  heating but  second  system  day  until  was a t t a c h e d t o augment  t h e f l o w was t o o g r e a t The  temperature  degree-days. dead  eggs  were  incubation  f o r i t be  was  Following  periodically  the eyes  examined  and  day,  to  laboratory  British  Columbia.  alevins  (95% s u r v i v a l  moved  to  fiberglass recycled  stage. After  a l l e g g s had h a t c h e d .  1.5° loan  Seven  from  days  tubs.  The  deg-days),  68  were  days, o f  Tc t e s t  moving  300 a l e v i n s  were  (1480 d e g - d a y s ) a l l r e m a i n i n g  stored within  at  to alevin  stage)  i n twc d a r k e n e d the  tubs  was  Biological  Station,  was a u t o m a t i c a l l y f o r 30 s e c o n d s . ,  were  227 l i t r e constantly  an a p p r o x i m a t e ' t e m p e r a t u r e  and f r y were f e d u s i n g  Nanaimo  t a c k s e v e r y hour  and  water  maintained  Oregon M o i s t Mash  later  frcm f e r t i l i z a t i o n  campus  C. The a l e v i n  into  f a c i l i t i e s a t s o u t h campus, U n i v e r s i t y o f  south  and  converted  and samples  p r o c e d u r e s and s t o r a g e t a n k s , a t 1400 d e g - d a y s , moved  temperature,  ( c a , 450 - 500  second  f o r development  (1280 d e g - d a y s ) ,  water  daily  the eyed-stage every  the  effective.  recorded  removed  were w e l l - p i g m e n t e d . ft  of 4 ±  mechanical feeders Fish Culture  released  into  the  on  Section. storage  

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
http://iiif.library.ubc.ca/presentation/dsp.831.1-0094749/manifest

Comment

Related Items