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A further investigation of the homing behaviour of the intertidal cottid, Oligocottus maculosus Girard Craik, Gwenneth Jean Steele 1978

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A FURTHER INVESTIGATION  OF THE HOMING BEHAVIOUR OF  THE INTERTIDAL COTTID, OLIGOGOTTUS MAGOLOSUS GIRARD  BY  GWENNETH JEAN STEELE CRAIK  B.Sc.(Hons.)i  Australian  National  A THESIS SUBMITTED IN PARTIAL  University,  FULFILLMENT OF  THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF  PHILOSOPHY in  THE FACULTY  OF GRADUATE  STUDIES  (Department' o f Z o o l o g y )  He a c c e p t t h i s  thesis  required  THE UNIVERSITY  as conforming t o the standard  OF BRITISH COLUMBIA  M a r c h , 1978 ((^SGwenneth J e a n  Steele  1973  Craik,  1978)  In  presenting this  thesis  an advanced degree at the I  Library shall  f u r t h e r agree  for  fulfilment of  the U n i v e r s i t y of B r i t i s h  make i t  freely available  that permission  for  the requirements f o r  Columbia,  I agree  r e f e r e n c e and  f o r e x t e n s i v e copying o f  this  that  study. thesis  s c h o l a r l y purposes may be granted by the Head of my Department or  by h i s of  in p a r t i a l  this  written  representatives. thesis  f o r f i n a n c i a l gain s h a l l  Zoology  University of B r i t i s h  2075 W e s b r o o k P l a c e V a n c o u v e r , Canada V6T 1W5  Date  is understood that copying or p u b l i c a t i o n  permission.  Department of The  It  22 inarch,  1978  Columbia  not be allowed without my  ABSTRACT The the  purposes  variability  of this  previously  observed  °J:i20£ottus m a q u l o s u s G i r a r d mechanisms olfaction was  by  which  at  British  morphological inlet in  and  12 s i t e s  Columbia.  behaviour  Consideration behaviour fish,  the  of  the  to  detail,  sensory  nature  of  analysis.  At any  the  population  and/or  results one with  clues,  could  were  involved  the use  and  examined. in  homing  and i n t r o d u c e d  of  the  home  paired  c h e m o s e n s o r y c l u e s and i n g r e a t e s t  and o l f a c t i o n  from  age  and  i n homing  behaviour.  year-class  vertebrae  decreasing differed  D i f f e r e n c e s were f o u n d which  and  differences  was  and  differences,  developed  be r e l a t e d  numbers  and  length-freguency  t i m e t h e r e a r e t h r e e major age  Age-length r e l a t i o n s h i p s  areas,  area  determination using o t o l i t h s  with  Vancouver  tidepool  length  between r e s i d e n t  Prior to investigating  compared  study  behaviour  in different  mechanisms  whether  The  coast of  of the  movement between t h e t r a n s p l a n t and  the r o l e s of v i s i o n  a method o f age  involved.  tidepool  o f chetaosensory  detect touch  clarify  particular  y e a r - c l a s s and one  to  i n homing  fish  included: interactions  pools, detection fins  mechanism  Variability  in  in  on t h e s o u t h w e s t  a r e a s , as w e l l as age, homing  attempt  homes;  c h a r a c t e r s between  of  i n t h e homing b e h a v i o u r  to  fish  i s t h e major s e n s o r y  conducted  Island,  the  s t u d y were t o f i n d t h e c a u s e s  o f age  groups  3 and  4  in  fish.  between a r e a s . in  homing  behaviour  t o wave a c t i o n  between  (turbulence)  and  iii  the  topographical  decreasing  turbulence  fidelity area  irregularity  i s  shown  and i n c r e a s i n g t o an i n c r e a s i n g  and, i n t i d e p o o l  homing b e h a v i o u r .  areas,  inlets,  little  homing  turbulence  decreasing  numbers o f f i s h  0.  can  be  related  maculosus i n t u r b u l e n t  No c o n s i s t e n t exposure  or  homing  that  there  parts in  of  these  of  t h e body, w i t h  With  terrain,  area.,  These  tide activity of (1971b,c).  which c o u l d  be  related  to  i n t h e m e r i s t i c and  in  different  areas.  length  i n t h e number o f c i r r i , and age.  There  areas but they  t o exposure.  Year-class  The f u n c t i o n  on a l l  are differences  do n o t seem t o  be  differences i n c i r r i  of the c i r r i  could  not  behaviour  are  determined. Age  apparent, of  high  0. m a c u l o s u s  increase  numbers a r e n o t a p p a r e n t . be  expressed.  r e g u l a r i t y of the  were f o u n d  pool.  on 0. m a c u l o s u s i n d i f f e r e n t a r e a s showed  i s a variable  directly  is  show  turbulent  t o t h e home  a r e a s shown b y G r e e n  r e l a t i o n s h i p s between  related  do home i n  t o t h e reduced  behaviour,  of c i r r i  which  i n the transplant  differences,  morphometric c h a r a c t e r s Investigation  remain  terrain,  percentages of f i s h  behaviour  and i n c r e a s i n g  Sfith  number o f p o o l s o r a w i d e r  percentages returning  decreasing  findings  of t h e t e r r a i n .  r e g u l a r i t y of the  increasing  However, t h e f i s h  r o u g h a r e a s show h i g h e s t In  (roughness)  the  classes  related  differences  although year-class age  than  differences one  year  homing  d i f f e r e n c e s are not.  in  age  in  homing b e h a v i o u r  groups  showed  improvement i n t h e p e r c e n t a g e s u c c e s s f u l l y  Examination  by s m a l l e r  that  there  homing w i t h  sizei s an  length  up  to  about  cm  5 cm.  (age  2)  Homing i s b e s t e x p r e s s e d and  after  percentage  successfully  returning  to  size  classes.  the  size  homing.  home  There  the t r a n s p l a n t  this  The p e r c e n t a g e  i s a decline  between 5 and 7  t h e r e i s a decrease i n t h e  p o o l appears  area with  i n fish  of  homing  t o be a b o u t  equal f o r a l l  i n t h e percentage  length  except  in  fish  the  remaining i n largest  size  c l a s s e s o f f i s h .. r  Juvenile into  lower  settle, begin  (about  from  the  2.3 t o 2.7 cm) which have moved high  show e x t e n s i v e movement  behaviour  and  tidepools  showing  period  fish  at  evidence about  of  3  cm.  tidepools  in  between t i d e p o o l s  home  range  It  o f e x t e n s i v e movement, t h e a r e a  they  and a p p e a r t o  fidelity  i s suggested  which  that  and  homing  during  i s i n some way  this  "learned"  "memorized". Investigation  homing b e h a v i o u r density  or  of  d i d n o t produce  "space",  found pool  to  suggest  that  Vision  homing, a c o m b i n a t i o n  are  interactions,  fins  movement  mechanisms i n v o l v e d i n  any e v i d e n c e t o show t h a t  or  on t h e body a r e i n v o l v e d  i s directed.  effective  sensory  behavioural  c l u e s d e t e c t e d by t h e p a i r e d elsewhere  the  taste  touch  to  vision  and s u b s e q u e n t l y  between t h e t r a n s p l a n t  and o l f a c t i o n  fish.  I t i s suggested  successful olfaction  that  adult  located  i n homing.,, Some e v i d e n c e was  appear  o f b l i n d n e s s and a n o s m i a  the  or taste  receptors  a n d home  t o be i n v o l v e d i n being  i n r e d u c i n g homing s u c c e s s t o low l e v e l s .  essential  pool  the Both  most  senses  homing o f j u v e n i l e f i s h b u t  become fish  unnecassary  a r e unable  in  older  t o home u n l e s s  V  one  of  these senses i s a v a i l a b l e .  of a p p r e c i a t i n g any  distance  olfactory pools  i n the t u r b u l e n t  used  " p o o l " t o home. nor  olfactory  be r e c o g n i z e d study  (Khoo,  sensory  how e i t h e r o l f a c t i o n  and v i s u a l ,  are  Because  t o home and t h a t However, n e i t h e r clues  emanating  by 0. m a c u l o s u s . 1971)  suggest  b e h a v i o u r and between a r e a s .  other  i t i s suggested  clues  the f i s h  from  moves f r o m  conspicuous  visual  over that  particular "pool" to landmarks  f r o m t h e home p o o l were shown t o T h e d a t a from t h i s  that  mechanisms may d i f f e r  o r v i s i o n c a n be used  intertidal,  and p e r h a p s  of the d i f f i c u l t i e s  the  relative  and a r e l a t e d importance  both i n t h e development  of  o f homing  vi  TABLE OF CONTENTS ABSTRACT ............... ..... .........................,.. ,. l i TABLE OF CONTENTS  vi  LIST OF FIGORES  xi  LIST OF TABLES  xiii  ACKNOWLEDGEMENTS I.  ................  ...........  ,.  xvi  GENERAL INTRODUCTION .................................. 1  II.  DESCRIPTIONS OF STUDY SITES AND CAPTURE BETHODS ......5 I.Major study s i t e .....................................5 F i r s t Beach  . ..... ..... 5  2.Other study s i t e s  10  Exposed study s i t e s ................................. 10 Moderately exposed study s i t e s  11  Sheltered study s i t e s ............................... 12 3. E n c l o s u r e  14  4. Capture methods ...................................... 18 III.  AGE DETERMINATION IN O. HACOLOSUS ................... 19  1. Methods .................... Age determination u s i n g o t o l i t h s  19 ..............19  Age d e t e r m i n a t i o n using v e r t e b r a e ................... 21 A n a l y s i s o f l e n g t h freguency data ................... 22 Age-length r e l a t i o n s h i p s  ............................24  F i r s t Beach .........................................2<4 Other areas 2. R e s u l t s Otoliths  ..........25  ..............................................26 ... . .  . . 26  Vertebrae ........................................... 28  vii  Graphical  analysis  Age-length  relationships  .........32 43  First  Beach  43  Other  areas  47  3.Discussion IV.  of length-frequency data  .....................................53  VARIABILITY 1. T a g g i n g  IN HOMING BEHA VIGOR  and r e c a p t u r e methods ........................57  Tagging  methods  Reacapture  ............57  methods  59  2. Homing i n d i f f e r e n t l o c a t i o n s First  57  Beach  ......60  .........................................60  Home r a n g e and a r e a f i d e l i t y  ........................60  Methods . . . . . . . . . . . . . . , . Results  61  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..... 62  Homing b e h a v i o u r  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66  Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Results  68  Inlets  71  Home r a n g e and a r e a f i d e l i t y  . . . . . . . . . . . . . . . . . . . . . . . . 71  Methods . . . . . . . . . . . . . . . Results  72  .............................................72  Homing b e h a v i o u r  along i n l e t s  74  Methods  . . . . . . . . . . . . . . . . . . . . 75  Results  . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . 76  Homing b e h a v i o u r  across i n l e t s  . . . . . . . . . . . . . . . . . . . . . . 78  Methods  ... .79  Results  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80  viii  3.Homing w i t h  respect  t o e x p o s u r e . . . . . . . . . . . . . . . . . . . . . . 80  Met hoas  83  Results  85  4.Measurement o f wave a c t i o n  and t o p o g r a p h i c a l  regularity  ...........90 Methods  92  Results  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95  5.Discussion V.  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99  MORPHOLOGICAL DIFFERENCES LOCATIONS LPort  FISH  IN  DIFFERENT  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106  Renfrew,  First  Beach, G r a p p l e r  Inlet  . . . . . . . . . . . . 106  Methods  106  Results  108  2. The r e l a t i o n s h i p  of c i r r i  t o exposure  110  Methods  110  Results  ......................113  3. E x p e r i m e n t a l  manipulation  of c i r r i  number . . . . . . . . . . . . 128  Methods  ........ 128  Results  .............................................132  4. H i s t o l o g i c a l  examination o f c i r r i  138  Methods  138  Results  138  5. D i s c u s s i o n 71.  BETWEEN  OTHER BEHAVIOUR 1. Age  ...........................................140 FACTORS  AFFECTING  VARIABILITY  IN  HOMING  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..... ........... 144 ............. 144  ix  Methods  144  Results  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145  2. Y e a r - c l a s s  148  Methods . . . . . . . . . . . . . . . » . . . . . . . . . . . . . . . . . . . . . . . . . . . , . 1 4 8 Results 3. L e n g t h  ,... 149  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154  Methods . . . . . . . . . . . . . . . . . . . . . ............. ........... 154 Results  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157  4. D i s c u s s i o n VII.  SENSORY  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172  MECHANISMS INVOLVED I N HOMING BEHAVIOUR  .....177  1 . S e n s o r y i m p a i r m e n t methods . . . . . . . . . . . . . . . . . . . . . . . . . . . 177 Vision  177  Olfaction Taste Fin  ............................  ........... 178  ..... .......  Removal  2.Interactions  179  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181 between r e s i d e n t  and i n t r o d u c e d  fish  ....182  Methods  ..... 183  Results  183  3. N a t u r e o f movement  between r e l e a s e and home p o o l  .....186  Methods . . . . . . . . . . . . . . . . . . . . . . . . .  186  Results  187  4. T o u c h a n d / o r c h e m o s e n s o r y  c l u e s . . . . . . . . . . . . . . . . . . . . . . 190  Methods  191  Results  192  5. R e l a t i v e i m p o r t a n c e  of d i f f e r e n t senses  . . . . . . . . . . . . . . 195  Methods . . . . . . . . . . . . . . . . . . . . . .  195  Results  196  X  6.Simultaneous impairment o f v i s i o n and smell  .......... 202  Methods  ........203  Results  ... ........ 204  7. V i s u a l and o l f a c t o r y t i d e p o o l c l u e s .................. 210 Methods  211  Results  ..216  8. D i s c u s s i o n VIII.  ...........................................220  GENERAL DISCUSSION  237  1 . V a r i a b i l i t y i n homing behaviour ...................... 237 2.Sensory mechanisms i n v o l v e d i n homing behaviour ......242 IX. X.  SUMMARY ...... ....  ......... .....245  LITERATURE CITED  XI. . APPENDICES  .....250 269  Appendix 1 .............................................269 D e s c r i p t i o n s of study s i t e s ......................... 269 Appendix 2 Total length  276 (TL) t o and from s t a n d a r d l e n g t h  (SL) ..276  Appendix 3 ......................... .... ................ 277 Algae and i n v e r t e b r a t e s from experimental tank ...... 277  xi  LIST  OF  FIGURES  Figure  Paqe  1.  General locations  of study  2.  Locations of  3.  Aerial  4.  E n c l o s u r e from  5.  O t o l i t h s o f 0. m a c u l o s u s s h o w i n g z o n e s ( a l l t a k e n on 30 A u g u s t )  study  phctoqraph  sites  6  sites of  First  outside  7 Beach  (above)  and  inside opaque  ....  16  hyaline 27  Length-age determined  7.  V e r t e b r a e o f age 3 0. m a c u l o s u s showinq h y a l i n e z o n e s ( t a k e n on 5 D e c e m b e r ) Lenqth-aqe determined  {below) and  6.  8.  reqressions for by o t o l i t h s ( m a l e s  8  F i r s t Beach and f e m a l e s  0. maculosus s e p a r a t e l y ) . 29 opaque  and 31  regressions for F i r s t B e a c h 0. maculosus by v e r t e b r a e ( m a l e s a n d f e m a l e s s e p a r a t e l y ) 33  9. 10.  11. 12.  13.  14. 15.  16. 17.  Lenqth-frequency method  analysis  of  aqe  groups  usinq  Cassie's 35  Lenqth-age r e q r e s s i o n s f o r First Beach C. maculosus d e t e r m i n e d by o t o l i t h s , v e r t e b r a e a n d l e n q t h - f r e q u e n c y a n a l y s i s ( m a l e s and f e m a l e s c o m b i n e d )  38  Length-frequency 0* 12.f2ili2Sil5  42  histograms  over  time  of  First  Beach  Age-length r e g r e s s i o n s of First Beach 0. maculosus determined by o t o l i t h s ( m a l e s and f e m a l e s s e p a r a t e l y and c o m b i n e d )  45  Age-length r e g r e s s i o n s of 0. j n a c u l o s u s f r o m eight • areas determined by otoliths (males and females combined)  50  Cement b l o c k tidepool Graphical characters  design  and  cement  block  anchored  in 94  a n a l y s i s o f e i g h t m e r i s t i c and o f 0. m a c u l o s u s a t f o u r a r e a s  mcrphcmetric 109  Total cirri-length regressions for 0. m a c u l o s u s f i f t e e n a r e a s ( m a l e s and f e m a l e s c o m b i n e d )  from  Total  nine  cirri-age  r e g r e s s i o n s f o r 0.  maculosus  from  116  xi i  areas 18.  19.  20.  21.  22.  (males  and  females  Total cirri-age 0. m a c u l o s u s from f e m a l e s combined)  by two  combined) year-class regressions selected areas (rcales  showing  Total cirri-length the c a l m and e x p o s e d  0.  cirri  and  exposed  from 0, maculosus and f r o m F i r s t Beach  regressions f r o m 0. rcaculosu s i d e s o f e x p e r i m e n t a l tank  from  the  lateral  line  and  136  head  of 139  Homing  performance  by  age  24.  Homing  performance  by  year-class  25.  Homing  performance  by  length  2f>.  Circular  27.  two s a m p l e s o f f i s h Sections of tips  plots  of  (mean  direction  ±  2 s.e.) (mean  classes over  first  ±  146 2  (mean  Landmarks  29.  Y-shaped  150  ± 2 s.e.) taken  d i s p l a c e d to c e n t r a l peel o f p e c t o r a l f i n (above) and  .  163  by 188  pelvic 194  installed choice  s.e.)  3 days  (below)  28.  134  s in  maculosus  23.  fin  calm  130  Total cirri-lenqth regressions i n t r o d u c e d i n t o e x p e r i m e n t a l tank  of  for and 126  Design of experimental tank s i d e s a n d wave m a k i n g t a n k  Sections  123  tank  in pools  at  First  Beach  212 214  xiii  LIST  OF  TABLES  Table 1. 2. 3. 4.  5.  Page First Beach 0. m a c u l o s u s (otoliths) functional regression statistics  length-age  First Beach 0. m a c u l o s u s (vertebrae) functional reqression statistics  lenqth-aqe  w e a n l e n g t h (cm) f o r e a c h t h r e e d i f f e r e n t methods  aqe  qroup  0. t n a c u l o s u s (First Beach) length-aqe statistics from three different methods determination C o m p a r i s o n o f a q e d e t e r m i n e d by o t o l i t h s o f t h e same 0. m a c u l o s u s i n d i v i d u a l s  7.  First Beach 0. m a c u l o s u s (otoliths) regression statistics and analysis of statistics (males vs females)  10. 11.  of taqqed  0.  maculosus  A l l areas 0. m a c u l o s u s regression s t a t i s t i c s (males A l l areas analysis females)  vertebrae 40 44 aqe-lenqth covariance 46  (males  48  vs 49  (otoliths) aqe-lenqth a n d f e m a l e s c o m b i n e d ) ...  12.  First  replacement  13.  Long-term pool fidelity f r o m 30 J u n e r e p l a c e m e n t performance  39  (otoliths) aqe-lenqth and f e m a l e s s e p a r a t e l y ) .  covariance statistics combined)  Homing Beach  reqression of aqe  individuals  A n a l y s i s of and f e m a l e s  14.  and  of c o v a r i a n c e s t a t i s t i c s  A l l areas 0. m a c u l o s _ u s regression s t a t i s t i c s (males  Beach  by 36  Growth  9.  34  determined  6.  8.  30  of  for a l l areas  (males 54  experiment  63  of individual experiment  u n t r e a t e d o.  52  0.  maculosus  i a c u l o = u s at  64 First 69  15.  Grappler Inlet  replacement  experiment  73  16.  Homing  behaviour along Grappler Inlet  77  17.  Homing b e h a v i o u r a c r o s s G r a p p l e r I n l e t  18.  Homing  behaviour  i n areas  of  different  81 exposure  86  xiv  19.  Area  20.  F i n a l study of r e l a t i v e of cement b l o c k s  21. 22.  23.  24.  25.  fidelity  in sites  of  different  exposure  -  exposure weight  89 statistics 96  Homing b e h a v i o u r a c c o r d i n g and t u r b u l e n c e  to topographical  regularity 100  0. m a c u l o s u s (15 a r e a s ) t o t a l c i r r i - l e n g t h r e h r e s s i o n s t a t i s t i c s (males and females and combined)  functional separately  0. m a c u l o s u s (9 areas) total cirri-age r e g r e s s i o n s t a t i s t i c s (males and females and combined)  functional separately  120  125  0. m a c u l o s u s (9 a r e a s ) t o t a l c i r r i - a g e by y e a r - c l a s s functional regression statistics (males and females combined)  127  Experimental manipulation cf cirri i n 0. functional regression statistics for i n i t i a l s a m p l e s and F i r s t B e a c h  135  maculcsus and final  26.  Homing  performance  by  age  147  27.  Homing  performance  by  year-class  151  28.  Summary o f e x p e r i m e n t a l enclosure  tidepool 159  Homing  30.  Analysis  31.  Homing  32.  Homing performance of j u v e n i l e 0. m a c u l o s u s k e p t i n the l a b o r a t o r y f o r v a r y i n g p e r i o d s p r i o r t o r e l e a s e ..  171  Homing p e r f o r m a n c e resident fish  after  184  34.  Homing  after  35.  Relative impairment  36.  37.  of  length  replacement  performance  performance  of  classes  juvenile  after  importance of  164  e x p e r i m e n t s by 0.  length  pools  removal of paired  four  168  maculosus  transplant  homing performance treatments  Homing p e r f o r m a n c e treatments Relative  by  into  29.  33.  performance  introductions  after  169  cleared  of  fins  four  193  different 197  simultaneous  impairment 199  blindness  and  anosmia  205  XV  38.  39.  Homing anosmia  performance  after  simultaneous blindness  and 206  R e l a t i v e homing performance a n o s m i a by l e n g t h  ao.  Homing  performance  41.  Choice  tank  tests  after o f arm  pool  following  blindness  and 208  "landmarks"  preference  exchanged  ..  217 219  xvi  ACKNOWLEDGEMENTS  I  wish  D.  McPhail,  for  their  Dr.  T.  advice  especially for  t o t h a n k t h e members o f my c o m m i t t e e . D r . J . G.  and  Northcote  criticisms  grateful  of this  work.  This  work  was  the  particularly  Phil  with in  t o Brodie  identifications the  Barbara  field  tank.  the  I  am  criticisms  Julie algae  of t h i s  Zittin  N.  J.  during  Celestino and to  am  the  Canadian  and I  f o r helping  I  Wilimovsky,  Dave  thank  h e l p and Peacock,  to build the  Bunting  and i n v e r t e b r a t e s  Scudder  and  for their  and B a r b a r a  and  f o r help  assistance  I appreciate t h e help o f  and  Myriam  Haylock  from  Bruce  f o r h i s thoughtful  Lea man  an  who  invertebrates  experimental  thesis.  Hards  prepared  the h i s t o l o g i c a l  drew most o f t h e f i g u r e s .  conducted  thesis,  Chundoma,  Stanley  Broderick  of algae  grateful  Daphne  was  Haylock,  on numerous o c c a s i o n s .  Bunting,  identified  Floy  Myriam  Rhynas, Kent R o s s a n d R i c k and  E.  S t a t i o n at Bamfield,  My t h a n k s g o e s t o Emmanuel  enclosure  G.  c o n d u c t e d o u t o f t h e Western  Marine B i o l o g i c a l  patience.  this  G.  and a s s i s t a n c e i n t h e f i e l d  Universities staff,  of  t o my s u p e r v i s o r , D r .  h i s guidance, advice  course  and D r .  while  The m a j o r i t y  I was on a CSIRO P o r t g r a d u a t e  s e c t i o n s and of this  work  Studentship.  1  I.  GENERAL  INTRODUCTION  V a r i o u s a s p e c t s o f homing in  about  a  dozen  different  b e h a v i o u r have  intertidal  G e r s b a c h e r and D e n i s o n , 1930; Beebe, Williams, 1970; of  1957;  Green,  Peppar,  1971a,b,c,  intertidal  fish  evidence o f l i m i t e d the  ability  For only  (Cuvier  and  clear  one  i n this regard  intertidal  The  majority  a p p e a r t o show some  a r e a when d i s p l a c e d , fish,  Bathyqobius  does t h e r e  fish  respect  to  mechanisms i n v o l v e d  California  maculosus  1973).,  of  at  of  that i s , soporator  a p p e a r t o be r e l a t i v e l y  range,  i n homing,  involved:  Kuril  least  10  showed homing  homing  ranges  Islands  vision  the  (1971b)  behaviour.  sculpin,  from  northern  Okhotsk  pool  showed  contrast  over  demonstrated  that  S u b s e q u e n t l y , Khoo  at  Port  a  relatively  a comprehensive examination o f v a r i o u s  b e h a v i o u r o f 0. m a c u l o s u s  Sea  s u g g e s t e d , on t h e  same  the f i s h  most  b e h a v i o u r and  and  (1930)  i n the  days, t h a t Green  received  i s the t i d e p o o l  which  t a g g e d O. m a c u l o s u s  0. m a c u l o s u s  in  has  G e r s b a c h e r and D e n i s o n  movement.  o f t h e homing  home  Sea,  restricted  undertook  which  Girard,  t o t h e Bering  basis of finding  found,  G i b s o n , 1967; S t e p h e n s e t a l . ,  to a small  intertidal  with  Oligocottus  1974)  1971;  1951, 1971) .  attention sensory  1951,  home r a n g e and i n some s p e c i e s , e v i d e n c e  Valenciennes),  The  period  examined  studied  (Hubbs, 1921;  1931; A r o n s o n ,  e v i d e n c e o f t h e p r i m a r y s e n s o r y mechanism  (Aronson,  (Hart,  fishes  1973; Khoo, 1971, 1974).  to return  homing.  1965;  been  (1971, aspects  Renfrew.-  He  t o t h e s u g g e s t i o n s o f G r e e n , and G e r s b a c h e r  2  and Denison, their  that  natural  range was  deep  action.  subtidal  extent  fish,  size  although a f t e r one  effect  (from 4.5  on  experiments  homing  he  weather,  rough  areas decreased homing s u c c e s s and  improvement i n homing performance  no  Adverse  a d i f f e r e n c e i n homing performance  experienced  and  maximum  He found that 0. maculosus could home from up t o  modified by wave  and  t h e r e was  restricted  and t h a t the e f f e c t o f i n c r e a s i n g d i s t a n c e on homing  a b i l i t y was  Sex  rarely  u s u a l l y a s m a l l group of p o o l s with a  away  terrain  individuals  movement t o one p o o l , and showed that t h e i r home  of about 30 m. 300m  0. maculosus  to 9.0  between  naive  displacement t h e r e was  of r e p e a t e d l y d i s p l a c e d  cm t o t a l length) appeared  performance.  and  From  sensory  fish. to have  impairment  concluded t h a t touch or t a s t e c l u e s d e t e c t e d by  the p a i r e d f i n s were not i n v o l v e d i n homing, but that v i s i o n o l f a c t i o n were, o l f a c t i o n being more important than v i s i o n . concluded  no  that  and He  0. maculosus uses o l f a c t i o n and/or some kind of  e x p l o r a t o r y search process t o home. T h i s study was investigating purposes  of  variability  the this in  designed t o continue the work of  homing study  homing  behaviour were:  to  of O. maculosus. find  success observed  the  causes  olfaction  is  The  two  of  the  by Khoo and t o examine  f u r t h e r the sensory mechanisms i n v o l v e d i n homing, t o whether  Khoo,  determine  the major mechanism i n v o l v e d and whether  other mechanisms are a l s o i n v o l v e d . The v a r i a b i l i t y i n the percentage o f f i s h homing observed by Khoo l e d  him  to  suggest  that  successfully there  were  3  homing  and non-homing members o f t h e p o p u l a t i o n .  elucidate  the  performance,  factors  several  that  cause  approaches  variability  were taken.  areas  variability  e x i s t e d between areas and i f i t could  particular  investigated  examined  factor.  In  to  determine  addition,  on  whether  behaviour  by age, year c l a s s and l e n g t h t o determine  differences  i n homing juvenile  development  of  performance,  fish.  It  was  felt  that  investigation  in  intertidal  homing,  the  make i t d i f f i c u l t t o  utilization  of  distinct  following  the  might be more i n adults  odour  any  role  in  homing  mixing  streams,  behaviour.  involved  the r o l e  processes  understand  mechanisms i n t h e homing behaviour of  in  developed.  of  of t h e  the e x i s t e n c e  and  as suggested by Khoo,  A d d i t i o n a l mechanisms were examined t o i n v e s t i g a t e played  whether  of the sensory mechanisms  since  was  was  i n homing behaviour was c h i e f l y designed t o c l a r i f y olfaction  to  emphasis  homing behaviour i n j u v e n i l e f i s h  p r o d u c t i v e than examination o f homing performance  The  any  I n the study of l e n g t h  particular  which t h i s behaviour i s r e l a t i v e l y well  homing  be r e l a t e d  homing  these f a c t o r s a f f e c t homing performance.  placed  in  Homing behaviour i n  different  a  was  To attempt t o  The  juvenile  whether  they  r o l e o f sensory fish  was  also  investigated. In t h i s study, home range i n t i d e p o o l a r e a s i s used i n the  way i n which i t was used by Khoo (1971) and Gerking  as t h e area covered by t h e f i s h during from  Hayne, 1949).  normal  travel  (1959), (modified  Thus, t h e home range i s regarded as an area  of  limited  extent,  0. m a c u l o s u s  individual  r a n g e was r e g a r d e d tidepools. way  covering  the  i s found.  as a small area  In this  study,  i t was used by Khoo  tidepools  in  which  an  In n o n - t i d e p o o l  areas,  home  comparable  to  a  group  of  t h e t e r m homing was a l s o used i n t h e  (1971),  a s t h e r e t u r n t o t h e home  range  when e x p e r i m e n t a l l y d i s p l a c e d . This  study  i s  composed  of  five  parts.  Since the  e f f e c t s o f a g e a n d y e a r - c l a s s on homing were t o be i n v e s t i g a t e d , it  was n e c e s s a r y  Thus  the  first  Investigations morphological  t o develop section of  a valid i s  concerned  behavioural  d i f f e r e n c e s between  o f age d e t e r m i n a t i o n . with  ageing  differences populations  in  homing  various  the  following  concerned  with  t h e i n v e s t i g a t i o n s o f age, y e a r ^ - c l a s s a n d  the  role  in  homing  sections.  in  studies.  comprise  differences  two  method  behaviour.  of various sensory  and areas  The f o u r t h s e c t i o n i s length  The f i n a l s e c t i o n c o n s i d e r s  mechanisms  i n homing.  5  II.  DESCRIPTIONS OF ST DDI SITES UND  CAPTURE METHODS  1.Major study s i t e  First  Beach  First Channel  in  site L). it  Beach i s s i t u a t e d on t h e  Barkley  Sound  T h i s s i t e was  east  side  of  Trevor  (48°49»N,125°10*W) ( F i g u r e s 1 and 2,  chosen as the major study  area  because  o f f e r e d a l a r g e s e l e c t i o n o f t i d e p o o l s at both the north and  south ends of t h e beach and because both  Trevor  it  permitted  Channel and the head of Bamfield I n l e t  The beach i s moderately exposed s i n c e p r e v a i l i n g (from  the  relatively  access  southeast large  and  waves  particularly breaking  the  outside  the  from  (Figure 3).  adverse weather north)  produce  bay.  Further  d e s c r i p t i o n of t h i s area i s p r o v i d e d i n appendix 1. The m a j o r i t y o f experimental work conducted Beach  was  of  First  c a r r i e d out i n two groups o f t i d e p o o l s a t e i t h e r end  of the southern rocky s h e l f . groups  at  pools  is  The  distance  about 60 metres.  between  these  There were s i x pools i n  each group, but f o r most experiments only about f o u r of the pools i n each group were used t o t r a p f i s h . depended  largely  on  Since  six  The c h o i c e of pools  the season and abundance  o f 0. maculosus.  Both groups of pools ranged between about the 1 and 2.5 level.  two  m  tide  the v e r t i c a l d i s t r i b u t i o n o f O. ma,culosus i n the  Figure  1  General  location  of  study  sites  125° 10'  Figure 2  Locations  of s t u d y  sites  Figure 3  A e r i a l view of F i r s t  Beach  9  intertidal there At  was some s i z e  Port  (total less  Renfrew,  length)  than  larger than  i s related  t o the size  selectivity Green  occurred  occurred  in  Beach,  than  10  cm,  so  0. m a c u l o s u s  at  high  levels.  largely  fish  Thus, a t F i r s t (total  length)  2  m tide  level  there  are Since  i n high  and  c a n be f o u n d  relatively settlement  shallow  down t h e i n t e r t i d a l  i n about  experiments.  l e s s than  tidepools less  3.5 cm  few  fish  Although  fish than  larger 10  few  cm  large  of 0. m a c u l o s u s  t i d e p o o l s , there  as t h e y  i s  a  increase i n size.  m a c u l o s u s between a b o u t 2.0 t o 3.0 i n t i d e p o o l s a t a b o u t t h e 1.5  mid-June.  The number o f f i s h  4.0 cm i n c r e a s e s i n p o o l s a t and below t h i s progresses.  in  1971a),  few p o o l s i n t h e u p p e r i n t e r t i d a l a r e  B e a c h , some 0*  cm  t h a t few f i s h  inhabit  deeper  of  used  tidepools at a l l levels,  At  movement  (Green,  below a b o u t t h e 1.5 m l e v e l .  deep.  appears t o occur  i n the fish  found  5.5 cm d i d n o t n o r m a l l y First  fish  below a b o u t t h e 2 m l e v e l  5.0 cm o c c u r r e d  fish  of the  level  less  as t h e  to than  summer  10  2. O t h e r s t u d Y  sites  Exposed s t u d y  sites  Benson southwest of the chosen  on  northeast oceanic study  t o east swells  (Figure and  1,  shelf  directly  Barkley  s i n c e e v e n on  While  shelf  facing  the  and  thus  the  site  s t o r m s come from t h e s o u t h e a s t ,  unprotected  i s s i t u a t e d at the  site  was  s e l e c t e d as  Sound.  a calm  day  Botanical  The  The  shelves. beach  San  beach Although  of  the  site  was  Cape B e a l e ,  Beach a t P o r t on  Juan I n l e t consists  Barkley  (48°47»N,125°13*«)  an  c h o s e n as  of  be s e e n  and  on  a  narrow,  facing  west  exposed  site  breaking  the  1,  shore  on  the  itself.  Renfrew, h e r e a f t e r r e f e r r e d t o the  west  coast  (48°32*N,124°27*tf) extensive  i t i s s i t u a t e d i n the  faces i n a westerly  an  Sound  (Figure  was  lighthouse  l a r g e waves c a n  Renfrew, i s l o c a t e d near  mouth o f  c h o s e n at Cape B e a l e  in front  offshore reefs surrounding  the  rocky  study  of  study  rocky  D).  The  majority  of T r e v o r C h a n n e l  The  Island  A).  side  B).  Port  Sound.  the  site.  site  as  site  severe  i s relatively  the east  across  i n Barkley  i s situated in  I s l a n d i s a narrow, s t e e p  Cape B e a l e on  (H8°53» N, 125° 22'»)  Broken Group  Benson  site  exposed  Island  and  north  of  ( F i g u r e 1,  sandstone Strait  Vancouver  and  o f J u a n de  westerly d i r e c t i o n  site shale Fuca, onto  11  the  Pacific  swells.  Ocean.  Thus i t r e c e i v e s  the full  effect of  F o r t h i s r e a s o n i t was s e l e c t e d a s a n e x p o s e d  situated kilometre  The  b e a c h d e s i g n a t e d as P a c h e n a P o i n t  on  the  west  north  coast  of  (48°**3»N, 125°07« H)  of  the  (Figure  Vancouver  Pachena 1,  C) .  study i s a b o u t one  Point  site  site.  i n this Island  lighthouse  The  beach  s o u t h w e s t d i r e c t l y o n t o t h e P a c i f i c Ocean and t h u s was as  an  exposed  steeply  conducted.  shelf,  i n t e r t i d a l i s composed o f a  in  which  most  which  Island M) .  consisting  lies  a  broad  Two  Point in  study  sites  of r e l a t i v e l y  were  shelves  boulders.  i s s i t u a t e d on  t h e Deer Group  flat  o f t h e e x p e r i m e n t a l work was  l a r g e r t i d e p o o l s and s c a t t e r e d  Diana  and  upper  faces  selected  On e i t h e r s i d e o f t h i s s h e l f a r e l o w e r r o c k y  Kirby  site  The  s l o p i n g g r a v e l beach, below  sandstone  with  site.  Pacific  the  southwest  corner  (48°51«N, 125°12»W) used  at  Kirby  narrow rocky s h e l v e s  (Figure  Point,  facing  of 1,  both  southwest,  thus r e c e i v i n g t h e impact o f oceanic s w e l l s . More d e t a i l e d d e s c r i p t i o n s o f t h e s e a r e a s a r e  provided  i n a p p e n d i x 1.  Moderately exposed study  sites  Haines I s l a n d i s s i t u a t e d towards t h e s o u t h e r n end the  Deer G r o u p o f i s l a n d s i n B a r k l e y  of Haines I s l a n d c o n s i s t s  of  a  Sound.  large  rocky  The s o u t h e a s t shelf  which  of side i s  12  exposed at low t i d e (48°49.9»N,125°11.7«W). prevailing  winds  are  Helby i n Barkley on  the  direction  was  (Figure 2, s i t e  chosen  as  a  H).  I s l a n d i s s i t u a t e d i n the Deer Group of i s l a n d s  Sound.  The  s i t e s e l e c t e d f o r study i s a rocky  Eagle Channel (48°51 'N, 125°11»5?)  Although  the  numerous d r i f t  on  moderately  the southwestern s i d e of the i s l a n d , f a c i n g Ohiat  Imperial  strongest  g e n e r a l l y from the southeast, an area  the s h e l f f a c i n g t h i s exposed study s i t e  Since  beach  (Figure  i s somewhat protected  l o g s and  absence o f low  Island  and  site  E) .  2,  by Ohiat  shelf  Island,  brush j u s t below the  the tree  l i n e suggest that the beach i s moderately exposed. More d e t a i l e d d e s c r i p t i o n s of t h e s e s i t e s are i n Appendix  Sheltered  1.  study  sites  Grappler Trevor  Channel  Inlet i s situated  in  Barkley  is  a  mudflat,  adjacent  One,  small  (48°49.9* N, 125°07.7'W) and to  the  on  Sound.  chosen as major study s i t e s . inlet,  provided  Two on  the  eastern  Government  of  s i t e s i n the i n l e t were  the  south  relatively  the second,  side  an  side  of  the  sheltered extremely  Wharf  in  bay  sheltered  Port  Desire  (48°49.8» N,125°07.6'S) (Figure 2, s i t e s I and J r e s p e c t i v e l y ) . A second s i t e sheltered  study  site  on  Haines  Island  was  (48°50.1»N,125°11.7»W).  selected On  the  as  a  eastern  13  s i d e of the low  tide.  extremely into  Between  the i s l a n d  Dodger C h a n n e l from  One selected  F).  The  and  on  f o r study was  the  calm  d u r i n g most s t o r m s .  the  Haines  was  south as a is  s h e l t e r e d study a  shallow up  with  study.  was  Inlet  water  the  1.  tide  relatively  rocky  outcrops  2,  (Figure  site  s i n c e i t remains  marina e x t e n d s  along  the s i t e  chosen  between Ranee I s l a n d and  Two  site  K).  specific  main c h a n n e l  a t low  tide  there  were  and  chosen  t o Ranee I s l a n d , tide.  adjacent to the  the  chosen  Sance i s l a n d  areas  adjacent  d e s c r i p t i o n s of these  was  At low  surrounding  a large tidepool  an a r e a i n t h e main c h a n n e l  Appendix  the  (48°49»N, 125°8 * W)  ( F i g u r e 2,  Inlet.  from  low  at  bed.  Bamfield  More d e t a i l e d in  of Zostera  of  which i s i s o l a t e d  i s an  I s l a n d s i d e o f Dodger C h a n n e l  intertidal  site  shelf  pool i s  o f Dodger C h a n n e l and  of the  the  was  The  a sheltered site  channel  One  drains  C h a n n e l by  Haines  A bed  east side of Bamfield  joining for  area of  G).  the rocky  at  sides.  as  shore  a t t h e e a s t e r n end An  and  of r o c k e x p o s e d  (-48°50. 1 »N, 125°11. 8»W)  selected  Island  site  Trevor  eastern  area  partially  ( F i g u r e 2,  site  site  itself  which  s w e l l s from  i t s southern  was  i s a large irregular  large tidepool,  protected on  island  The  other  mainland.  sites  are  provided  14  3. Enclosure  A spanning a  3.5  l a r g e e n c l o s u r e c o n t a i n i n g a r t i f i c i a l t i d e p o o l s and  the i n t e r t i d a l from l e v e l was  about the 1 m l e v e l  to  about  c o n s t r u c t e d i n the s m a l l bay i n G r a p p l e r  the Inlet  (48 49.9»N,125<>07.7*W) {Figure 2, s i t e I ) . o  The in  the  e n c l o s u r e was  study  experiments  of  homing behaviour  conducted  0. maculosus  designed t o serve  in  the  individuals  several  purposes  of 0. maculosus. , In homing  field,  a  large  percentage  of  which do not home are l o s t f o r study.  In a completely enclosed system of t i d e p o o l s , those  fish  do  differences  not home may  be c o l l e c t e d f o r examination  between homing and non-homing reason  of any  0. maculosus.  The  f o r the c o n s t r u c t i o n of the enclosure was  the pool f i d e l i t y and homing behaviour  other  i n t h e f i e l d as  entail  tagging  they  large  moved  of j u v e n i l e 0.  numbers  into of  lower  maculosus.  c o n t r o l l e d environment of the e n c l o s u r e o f f e r e d of  introducing  monitoring  large  their  numbers  movements  of  and  juvenile homing  juvenile  tidepools  small f i s h .  major  to i n v e s t i g a t e  F o l l o w i n g the development of homing behaviour of tagged fish  which  The the  relatively possibility  Q_. maculosus  behaviour  would  and  much  more  c o n s t r u c t e d on a r e l a t i v e l y  steep  readily. The rocky  outcrop  enclosure in  the  was  otherwise  gently  sloping  bay.  The  15  framework  of  the enclosure,  10 m l o n g , 3.5 m w i d e and 5 m h i g h f  a t t h e s e a w a r d end was made o f two 7.7  cm  galvanized pipe  top  of these  supports  the  h o r i z o n t a l pipes.  as  intermediate  embedded  two l o g s were l a s h e d  along  the  to increase s t a b i l i t y  mesh  chicken  screening. prevent  operation,  mesh mesh cm  of  juvenile  down This  the  proved  planks  bottom was  wire  embedded left  enclosure  window In  and window and  screening the  2. 5  walking  on t h e s i d e s were f a s t e n e d a t t h e base o f  window  of the on  the  summer, t h e  t o 2.5 cm  the  to  attached  bottom  In t h e second  cm  summer o f  were  the  sides  served  first  screening  across  of  of fibreglass  u n s a t i s f a c t o r y because  i n concrete  the four  x 10  sides.  The  dishpans  (40  uncovered.  Deep  (30 cm)  cm i n d i a m e t e r ) polypropylene  layer  fish.  sides  fish  an e x t e r i o r l a y e r  p r o d u c e d l a r g e numbers o f h o l e s . and c h i c k e n  served  of the e n c l o s u r e .  were added t o t h e  an i n t e r i o r  the c h i c k e n wire  continuously enclosure.  and  with  The s m a l l mesh o f t h e  escape  On  (Figure 4).  were c o v e r e d  wire  end.  i n concrete  sides  e l i m i n a t e escape of introduced  of t h e enclosure  at either  of  t o the outer edges o f  Wooden p o l e s embedded  supports  constructions  i n concrete  S e v e r a l d i a g o n a l and c r o s s s u p p o r t s  To  I shaped  served lines  as t i d e p o o l s . tied  with  h y d r a u l i c cement.  the  positions  s m a l l rocks with  of  and s h a l l o w  (15 cm)  plastic  T h e s e were  t o e y e b o l t s embedded  T h i s arrangement allowed  particular  attached  Fucus  tidepools. s p . and  attached  with  i n the substrate alteration  of  Gravel, pebbles  and  Nxtilus  edulis  were  Figure 4  Enclosure  from o u t s i d e  (above) and i n s i d e  (below)  17  placed  i n the A  shallow  s e a w a r d end return to level  tidepools.  of the  concrete  enclosure  a tidepool  to  at  low  to enable tide  of the  a b o u t one from  l o g s and boat  winter the  the  t i r e s was  wake, w h i c h  problem  was  screening. during  the  constructed fish  followed  which  at did  the r e c e d i n g  levels  lower  placed  on t o p  than  the not  water  that of  the  was the  top o f the e n c l o s u r e  o u t s i d e o f the  severe  effect  Several  to  effects of  severe  enclosure,  provide  waves  the  cross  to  enable  on  the and  a s m a l l boom o f  a partial screaning. strong  barrier  summer months.  the  to  A greater  winds caused  s t o r m s were s u f f i c i e n t t o  Thus, e x p e r i m e n t s i n the  of  tide.  constructed  had  the  storms.  feasible  metre f r o m  above a t h i g h  Around  was  enclosure.  A l a r g e wooden p l a n k  viewing  but  remain i n water at t i d e  s e a w a r d end  supports  trough  enclosure  by  destroy  were  only  18  4.Capture  methods  Two of  methods  0. f a c u l o s u s used  wire  minnow  of capture  i n homing  traps.  ft  were u s e d .  experiments  piece  of  The v a s t were  mussel  was a t t a c h e d  t o a hook c e n t r a l l y  trap.  were  i n pools  low  tide  When  ( d e p e n d i n g on  reciprocal  insufficient group  placed  of  necessary  weather  remaining  dipnets  cm  (total  smaller  fish  in  were  and  being  tide  and from  (fish  less  length)  to enter  few f i s h  l e s s than  about  because t h e s i z e of the wire  and e x i t  anywhere  t h e 2.5 m t i d e l e v e l , than  an one  to c a t c h the  in  the  3.5  mesh  Thus, i n pools  juveniles for specific  a b o u t 3.5 t o 4.0 cm)  were c o l l e c t e d  to  allows  trap.  s i n c e t h e m a j o r i t y o f homing e x p e r i m e n t s were c o n d u c t e d up t o a b o u t  around  height).  conducted  were s o m e t i m e s u s e d  the  fish.  Minnow t r a p s c o l l e c t 4.0  Mytilus  located  number o f 0. m a c u l o s u s had b e e n c a p t u r e d  pools, small  using  f o r about f o u r hours  conditions  transplants  caught  (usually  califqrnianus) Traps  majority  studies  with  small  dipnets. To  catch  fish  s t u d i e s a combination pool  population  little dipnets  for cirri  age  determination  o f minnow t r a p s and d i p n e t s were u s e d .  s t u d i e s , pools  water remained.. and by hand.  c o u n t s and  Fish  were b a i l e d  were  then  by b u c k e t  captured  until  using  In very  small  19  III.  To  AGE DETERMINATION  evaluate  b e h a v i o u r and o t h e r method  vertebrae  the  r i n g s on o t o l i t h s  method  of  ageing  and by u s i n g  groups devised  BACULOSUS  o f age and y e a r - c l a s s  f a c t o r s , i t was n e c e s s a r y  f o r determining  Initially, this  the effect  IN 0.  age  to develop a  o f 0. m a c u l o s u s  valid  individuals.  were i n t e r p r e t e d and v a l i d a t i o n  was  attempted  by  t h e g r a p h i c a l method  by C a s s i e  on homing  examining of  of  r i n g s on  separating  age  (1954).  !• Methods  Age  determination  using  Following preserved after  both  1  light  with  left  Otoliths  collection,  i n 10% f o r m a l i n .  soaking  length)  otoliths  in a dial  and  fresh  were examined  on a b l a c k  Within water,  caliper  right  0. m a c u l o s u s  collection  and (total  0.005 cm, s e x e d  removed a n d s t o r e d  immersed  surface, using  of  t h e f i s h were measured  t o the nearest  otoliths while  48 h o u r s  i n d i v i d u a l s were  i n water  and  i n glycerin.  under  reflected  30 x m a g n i f i c a t i o n . /  *A11 l e n g t h measurements o f 0. m a c u l o s u s i n t h i s s t u d y were made of total length, except i n the analysis o f m e r i s t i c and morphometric characters, where s t a n d a r d l e n g t h was u s e d . F o r c o n v e r s i o n s f r o m t o t a l l e n g t h t o s t a n d a r d l e n g t h and v i c e v e r s a , see A p p e n d i x 2.  20  Initially, determine was  least  two  Either  otolith  that  right  consequent  age o f t h e f i s h  and  second  without  First  1 June,  were e x a m i n e d t o  made  pattern of r i n g s o f one o t o l i t h  in  t h e number  was n o t e d .  until  a decision  was  each  difference or f o r m  pair.  between  zones  and  the  between t h e  by f u r t h e r  reached.  at  o f opague a n d  Disagreements  r e a d i n g s were r e s o l v e d  After i t  existed,  from  The numbers o f opague and h y a l i n e  examination  Otoliths  were  p r i o r r e f e r e n c e t o l e n g t h or s e x .  Six at  of o t o l i t h s  s i n c e t h e r e was no  otoliths  both o t o l i t h s  read  were  was u s e d ,  hyaline rings.  of  a consistent  readings  and  first  number  w h e t h e r t h e r e was e v i d e n c e o f g r o w t h r i n g s .  established  left  a large  collections  Beach.  totalling  The c o l l e c t i o n s  1976; 21 J u l y ,  1976; 29  365 0. m a c u l o s u s were made  were t a k e n  on 9 F e b r u a r y , 1976;  September,  1976;  5  December,  1976 and 12 F e b r u a r y , 1977. Functional constructed shown  by  males  and  length), in  females.  Regressions Since  and,  of  included  combined only  once.  the  of fish  regressions.  on  growth  were c a l c u l a t e d sex  regression  the  were  separately f o r  determination t h a n 3.50 cm  (22 f i s h ) The  t o form  age  relationship  were  a single  d a t a from  i s  (total  included  regressions  whether t h e y were s t a t i s t i c a l l y  i f n o t , t h e d a t a were c o m b i n e d the  length  juveniles less  age d a t a f o r t h i s g r o u p  to determine  of  external  f o r 0. m a c u l o s u s  b o t h t h e male and f e m a l e  compared  In  t o examine t h e n a t u r e otoliths.  impossible  regressions  were  different, regression.  juvenile fish  were  21  Age d e t e r m i n a t i o n  To  attempt  determination extracted  on  using  from  collections  using  fish  cut  otoliths,  of  age  t h e v e r t e b r a e and o t o l i t h s  were  made  the  Beach.  measuring  first  and  second  while  v e r t e b r a l c o l u m n b r o k e a n d i t was p o s s i b l e  solution After  vertebrae  of t r y p s i n  were r i n s e d  and water t o  s e v e r a l days, t h e v e r t e b r a e  solution, prior  The  rinsed  i n glass  Examination m a g n i f i c a t i o n on a single  vertebra  of  was  examined  f o r evidence  number  of  to  the  and fish  t h e number  were n o t e d .  to  sexing  pointing  d o r s a l f i n s and until the  extract  several  attached  in a  tissue.,  t o dry f o r s e v e r a l  away  using from  determine  consistency i n the patterns o f rings, read  and  were removed f r o m t h e t r y p s i n  o f growth r i n g s .  vertebrae  taken  twisting  the  vertebrae  surface  cut  were  days,  vials.  the  black  the  i n water and p l a c e d  digest  i n w a t e r and a l l o w e d  to being stored  of  An o b l i q u e f o r w a r d  f o r c e p s were u s e d t o h o l d t h e v e r t e b r a e  vertebrae.  two  1977 and i n v o l v e d 177 f i s h . ,  the o t o l i t h s . the  in  These c o l l e c t i o n s  were e x t r a c t e d a f t e r  between  results  individuals  1976 a n d 12 F e b r u a r y ,  and r e m o v i n g  was  validate  at F i r s t  Vertebrae the  to  0. m a c u l o s u s  made  5 December,  vertebrae  was  made  reflected  under  30 x  light.  A  t h e group and t h e c e n t r u m After  examining a  whether a l l the  there  was  vertebrae  large any were  o f opaque and h y a l i n e z o n e s and t h e age o f Two r e a d i n g s were made  of  a l l vertebrae  22  and  disagreements  single  vertebra  extracted. reference  to  reference  for  regressions,  otoliths (1940)  other  made  of  males  only  without  and v e r t e b r a e determine  prior  were  vertebrae  and  length  on  also  were made  females  age  were  t o examine t h e data  from  fish  were i n c l u d e d i n b o t h  male  and  once i n t h e combined  there  of  vertebrae  otoliths  of  t o the  readings.  was  regressions  comparison  to  readings  length)  but  which was c a l c u l a t e d i f  A  were  to  As i n t h e c a s e o f o t o l i t h s ,  (total  between t h e s e p a r a t e  farther reference  Although  regressions  growth r e l a t i o n s h i p .  female  fish,  separately  t h a n 3.50 cm  sex.  to otolith  Functional  less  vertebrae  and  these  by  and i f n e c e s s a r y ,  of  length  from  calculated  resolved  extracted  Readings  extracted without  were  no  regression,  significant  difference  f o r each s e x .  the  was  made  the  degree  age o f e a c h f i s h using  the  determined by  method  of  o f agreement between  Yasuda t h e two  methods.  Analysis  of l e n g t h frequency  During of  collections  various  purposes:  experiments, ageing last  of  o f t h e study  0. m a c u l o s u s  pool  population  were  a  Except  purposes,  studies,  i n the case of f i s h  a l l fish  considerable  made a t F i r s t  i n v e s t i g a t i o n o f morphological  studies. two  the course  data  homing  number  Beach f o r behaviour  c h a r a c t e r i s t i c s and collected  were measured w i t h  f o r the  a r u l e r t o the  23  nearest  0.1  cm  morphological  {total  and  length)  ageing  studies  c a l i p e r t o t h e n e a r e s t 0.005 cm this  analysis,  construct May,  1975  length  from  t o February,  sexed.  For the  by  curve  constructed  Cassie  collections and  and  original  graphical analysis. the  using  length-age  A  percent  The  the  significant  to  corresponding  used  a period  to from  difference t h a t age  l a r g e sample  cumulative the  method  frequency points  of  groups e x t r a c t e d .  as c l a s s i f i e d  obtained  1975  then by  using  calculated  age  from  compared  with  otoliths  between  the  determination using  the  and three  otoliths  technigue.  a further  population,  examined  data  regressions  might s u g g e s t  As  were  r e g r e s s i o n o b t a i n e d was  regressions  invalid  of cm.  graphical  paper,  v a r i o u s aqe  length  A  an  dial  the October,  the  probability the  vertebrae.  was  a  purposes  of the r e l a t i v e l y  groups  on  from  A l e n g t h - r a g e f u n c t i o n a l r e g r e s s i o n was the  with  for  were c o m b i n e d  spanning  histogram  (1954).  determined  these  (because  o f age  developed  using  used  1977.  selected  for analysis  inflection  measured  histograms  length-frequency  was  was  Fish  were  associated collections  length-frequency  collection size)  data  of c l o s e l y  The  and  sexed.  t h e l e n g t h s were r o u n d e d t o t h e n e a r e s t 0.1  The in the case  and  p o s s i b l e c o r r o b o r a t i o n o f age  the l e n g t h - f r e q u e n c y  determine  whether  the  t o d e r i v e d a g e s , c o u l d be  histograms  over  modes  size  of  f o l l o w e d over  groups time  in were  groups, the  months  24  represented  throughout  Finally,  data  from  caught  for  b e h a v i o u r a l experiments  initially  measured a t a l a t e r determine  by  collected  the  measured and s e x e d  were  outlined  above.  when  and s u b s e q u e n t l y investigated  These  to  rates  data  were  over t h e d u r a t i o n of t h e r e s e a r c h .  Beach  length  enable the  o f 0. m a c u l o s u s ,  were c a l c u l a t e d readings.  both  fish  when r e t r a p p e d ,  methods  intermittently  To  for  date  tagged  w h e t h e r t h e y p r o v i d e d any s u p p o r t f o r t h e g r o w t h  suggested  First  the population.  As  determination  from  data  obtained  (total  regressions since  possible.  The  they  significantly  were  regressions  any  given  from  otolith  were  used  were compared t o d e t e r m i n e  different,  and  i n t o one a g e - l e n g t h r e g r e s s i o n ,  juvenile  fish  once.  length)  data in  sex d e t e r m i n a t i o n was n o t  combined  only  for  t h e case o f t h e length-age r e g r e s s i o n s ,  0. m a c u l o s u s l e s s t h a n 3.5 cm male and f e m a l e  age  p r e d i c t i v e r e g r e s s i o n s o f age on l e n g t h  separately in  of  i f  not,  including  whether  they  the data  were from  25  Other  areas  Collections  of  0. m a c u l o s u s were made a t e i g h t  a r e a s on t h e west c o a s t o f V a n c o u v e r age-length  relationship  analysis.  Collections  Benson I s l a n d , tidepool), fish  fixed  collection.  Within  (total  length)  stored  in glycerin.  otoliths  were  with  The  and a f t e r  a dial  from  and  First  following water,  the  followed  in  for otoliths  removed and reading  the  e x t r a c t e d from  Beach. regressions  and f e m a l e s .  3.50 cm  I n cases  ( t o t a l length) i n both  had  were where been  male and f e m a l e  and o n l y o n c e i n t h e combined r e g r e s s i o n c a l c u l a t e d difference  regressions.  First  The  in  the  between t h e s e p a r a t e  regressions  Beach) were compared  were any d i f f e r e n c e s locations.  with  case,  t o t h e n e a r e s t 0.005 cm  d a t a were u s e d  t h e r e was no s i g n i f i c a n t  (including  (large  I n each  and r i g h t o t o l i t h s  f o r males  the age-length  female  rinsing  caliper  procedures  j u v e n i l e 0. m a c u l o s u s l e s s t h a n  if  the  in cirri  Island  immediately  area, p r e d i c t i v e age-length  separately  regressions,  f o r use  Haines  formalin  t h e same a s o u t l i n e d  For each  collected,  Island,  and t h e l e f t  0. m a c u l o s u s i n d i v i d u a l s  calculated  10%  HQ h o u r s  sexed  determine  (bay) a n d Ranee I s l a n d .  in  measured  location  to  were made a t P o r t Renfrew, P a c h e n a P o i n t ,  Grappler Inlet  were  each  Cape B e a l e , H e l b y  were  fish  in  Island  other  f o r a l l areas  t o determine  age-length  male  whether t h e r e  relationship  between  26  2. R e s u l t s  Otoliths  From first  inspection  otoliths  summer growth z o n e a p p e a r s  numerous  finer  c e n t r a l nucleus form  of  a  almost  complete  narrower h y a l i n e subsequent  broad  opaque  hemispherical  rings  surrounding  circle. band.  This  indirect  indication  juvenile  fish  laboratory the  in  caught i n l a t e  reared  h y a l i n e zones  hatching  m e t a m o r p h o s i s was  The otoliths  may  to have  to  zone  in  investigating found  seven  juveniles  been spawned  An  earlier  that weeks  12.2  mm  collected that  year  1939).  onset start  (1973)  Thus  first  readings i s the  hyaline  six  and  5).  d a y s , one j u v e n i l e was  w e l l advanced.  A u g u s t c a n be e x p e c t e d  ( a l s o see Atkinson,  51  occurred  a  zones are  (Figure  thin  summer., S t e i n  after  the  usually  of the  o. m a c u l o s u s l a r v a e i n C a l i f o r n i a  and t h a t  of  second  growth  of o t o l i t h  band and a  adoption of a b e n t h i c h a b i t a t  after and  opague  the  T h e s e opaque  o f the v a l i d i t y  o c c u r r e n c e o f a broad  not  narrower than t h a t  by t r a n s l u c e n t  band  w i n t e r zone i s { u s u a l l y )  Opaque g r o w t h z o n e s i n  summer a r e g e n e r a l l y  separated  opaque z o n e d o e s  The f i r s t  summer and f o r m c o m p l e t e c i r c l e s . clearly  that the  a  (Figure 5).  as  i t i s evident  o f growth i n t h e s e c o n d summer a s shown by  as e a r l y  as  February  or  as  late  as  Hay,  Age 1  Age 2  1.0 mm  Age 3  Figure 5  O t o l i t h s of O. maculosus showing opaque and h y a l i n e zones  ( a l l taken on 30 August)  28  depending appears first  on  location  that fish  appears  to  relatively fish  occur  growth  zones  sometime  w i t h wide g r o w t h  untransformed  data.  July.  {Figure values  regressions  Thus t h e l e n g t h - a g e  Again  otoliths, fish  growing  6) ,  log  to  with longer  f o r males  the  base  were made a s s u g g e s t e d  ( T a b l e 1)  confidence  results i n  f o r males and  females  10  by  the  limits  rejection  t h a t m a l e s grow more r a p i d l y  data  prior  summer. ,  r e g r e s s i o n s of length-age  I n s p e c t i o n o f t h e 95%  hypothesis  the  zones.  separately  (slope) of the null  after  It  for  summer's g r o w t h  s m a l l growth z o n e s a p p e a r t o c o n t i n u e  females  the  conditions.  c e s s a t i o n o f y e a r l y g r o w t h , as shown by  t r a n s f o r m a t i o n s o f t h e age  v  opaque  commence t h e s e c o n d  For the f u n c t i o n a l and  seasonal  s h o w i n g a w i d e growth z o n e f o r the f i r s t The  than  probably  showing s m a l l  summer*s g r o w t h  to f i s h  and  than  of of  females.  were c o m b i n e d  to  produce a s i n g l e r e g r e s s i o n .  Vertebrae  The in  vertebrae than  necessary angle  of  and  h y a l i n e zones are  otoliths.  t o permit  which  patterns on  opaque  allows  the  light  rings are  (1976)  moving o f  t o shine through  resolution similar  vertebrae there are f i n e  Chadwick  Frequent  of  the  to those  r i d g e s i n the  interpreted  more d i f f i c u l t  as  annual  to  see  the v e r t e b r a i s  the centrum  rings.  at  an  However  the  of o t o l i t h s e x c e p t  that  opaque marks)  zones  (which  (Figure 7).  The  29  First Beach O. maculosus female (otoliths) . Y= 4., 4 2 0 + 5 . 4 ^ 2 x log X  N= 2 2 8  8. 7. 6. E .0. c  5. 4. 3. 2.  0.2  A g e (years)  First B e a c h O. maculosus male (otoliths) Y= 4. 4 4 6 + 5. 1 3 8  x log X  .N= 1 6 0  8. 7. 6.  S  5.  /—\  o> 0)  4.  3  1.  0.2  Figure 6  1.  A g e (years) Length-age r e g r e s s i o n s o f F i r s t Beach 0. maculosus d e t e r m i n e d by o t o l i t h s (males and females s e p a r a t e l y )  Table  1  Beach O l i g oc o 11 u s m a c u 1 o sg s  (otoliths)  Length-age f u n c t i o n a l r e g r e s s i o n  statistics  First  log  x vs y: y = u + v l o g x  Statistic  Males  Females  4.420  5.138  5.499  Lower 95% c o n f i d e n c e l i m i t of v  4.511  5.021  Opper  5.765  5.977  159  228  95% c o n f i d e n c e l i m i t of v  31  Figure 7  Vertebrae o f age 3 0. maculosus showing opaque and h y a l i n e zones  ( a l l taken on 5 December)  32  p o s i t i o n o f these r i d g e s does not  always  correspond  with  edges of the opague growth zones which were used i n t h i s Figure for  8  shows the c a l c u l a t e d length-age  males and females s e p a r a t e l y  again,  inspection  the r e g r e s s i o n s hypothesis  as  determined  o f the 95% confidence  (Table 2)  results  in  study.  regressions  by  vertebrae,  l i m i t s o f v (slope) o f  rejection  of  the  that males grow more r a p i d l y than females.  length-age data  the  null  Thus the  f o r males and females were combined i n a  single  regression.  G r a p h i c a l a n a l y s i s of 1enqth-frequency d§ta  Graphical  a n a l y s i s o f a length-frequency  using t h e method of C a s s i e lenqths  (1954)  (Figure 9 ) , provides  the  mean  f o r each aqe group which can be compared with the mean  lengths f o r each age from o t o l i t h and vertebrae 3).  distribution  Although  the  standard  analysis  (Table  d e v i a t i o n s f o r the mean l e n q t h s a t  each age d e r i v e d from C a s s i e * s method are r a t h e r l a r g e , t h e r e i s fairly  good agreement i n mean l e n g t h s between the three The  length  s m a l l sample numbers  distribution  l e n g t h values percent  make  and  cumulative  fish  of  the  unevenly  absence  interpretation frequency  the i n f l e c t i o n p o i n t at length  the  four  at  of  the  upper  methods.  end  o f the  of any f i s h f o r s e v e r a l the  upper  end  of the  curve somewhat d i f f i c u l t .  98.5% t h e o r e t i c a l l y  the  upper  year age group, the p a u c i t y of data  (four  distributed)  f o r the  marks  While  remaining  four  lengths  33 First  Beach  Y=  O. maculosus female  3.535"^  7. 9 7 8  (vertebrae)  x log X  N=114  8, 7.  I O)  6.  51 4..  OJ  31 2 1.1 0.2  First  1. (years)  Age  Beach  Y=  O. maculosus  3. 3 9 4 +  8. 2 6 9  male (vertebrae) x  log X  N= 7 3  8. 7.  6.  S  5.  si  4  cn c OJ  3 2. 1 0.2 Figure  8  Age  1. (years)  L e n g t h - a g e r e g r e s s i o n s o f F i r s t B e a c h O. m a c u l o s u s d e t e r m i n e d by v e r t e b r a e (males and f e m a l e s s e p a r a t e l y )  Table First  Beach O l i g o c o t t u s  2 maculosus(vertebrae)  Length-age f u n c t i o n a l r e g r e s s i o n log  statistics  x vs y : y = u + v l o g x  Statistic  Hales  Females  u  3.394  3.535  v  8.269  7.978  Lower 95% c o n f i d e n c e l i m i t of v  7.047  7.214  Upper 95% c o n f i d e n c e l i m i t of v  9.492  8.743  73  114  N  100  OCTOBER, 1975 N=278  8.0  2 y i  6.0  r-  o  -z.4.0  LU  o  -o-  2.0 Jl  0.1  L  12  -1  5 10  1  1 1 1 1  CUMULATIVE Figure  9  Length frequency  50  I  I  I  90  FREQUENCY  I  I  I  I  I  I  99.99  (%>)  a n a l y s i s o f age g r o u p s u s i n g C a s s i e ' s  method LA  cn  Table  Hean l e n g t h s determined  Length Statistics  x  3  (cm) f o r e a c h age g r o u p  by t h r e e  Otoliths  different  methods  Vertebrae  Cassie's method  s.d. N  4.21 .73 127  4.02 .38 26  3.9 . 30 89  Age 2  x s.d. , N  5.01 .96 194  4.99 1.19 119  5.5 2.41 111  Age 3  x s.d. N  6.49 .60 43  6.14 .84 32  6.4 3.49 67  x s.d. N  7.30  Age 4  Age 1  Age 4+  x s.d. N  1  7.3 .21 7  7.8 .13 4  37  theoretically although  they  probably  In the  the  four  could  obtained  Inspection (Table  4)  Cassie^s  method, but  fit  this  of t h e  for  the  males  using  10),  since a  fish  definite  purposes and  of  females  a n a l y s i s a r e a l s o shown i n  relevant s t a t i s t i c s that  there  relationships  is  by  f o r the  good  described  t h a t growth d e s c r i b e d  may  for  of  the  age  fish  from  between t h e of  two  each  and  by  three  agreement  otoliths  vertebrae  and  may  not  but  (Siegel,  age  (p<.05).  Thus, there  is  otoliths  from  fish  derived  from  vertebrae.  method  of validating  age  Although  vertebrae  1956)  methods i n age group  a significant  older  by  d i f f e r e n c e between  and  a  shows  a  (Table  determination  individually  data  one  appear  determination  to  using  be  two  three  fish  derived  a  no  or  t o show more r a p i d growth  Vertebrae  5),  (p<.05).  reveals  d i f f e r e n c e f o r age for  is  significant  methods f o r age  tendency  there  otoliths  which b o t h were e x t r a c t e d  d i f f e r e n c e s between t h e  (p>.05),  vertebrae.  determined  Kolmogorov-smirnov t e s t  Examination  account f o r the  for otoliths  between t h e  fish  group.  (Figure  For  fish  pattern.  agreement  significant  method  them.  vertebrae  suggests  growth  regressions  difference  to  calculated  and  Several reasons  the  Cassie's  ascribed  otolith  group  of the length-age r e g r e s s i o n  from  regressions  age  y e a r age  be  the  86%  four  not  between  the  t o the  plus  g r o u p were n o t c o n s i d e r e d  10.  regressions  four  age  (combined) f r o m  into  belong  as  plus  comparison the  Figure  them  calculation  information  of the age  classifies  than  from that  successful  otoliths,  at  least  38  First B e a c h O . m a c u l o s u s otoliths  Y= 4. 5 0 9 + 5.076 8  First B e a c h O . maculosus C a s s i e c u r v e  x log X N=3 65 \ . _  Y= 4.340 + 4.892 x | 8.  7.  7.  6.  6.  5.  5 & 5.  4.  P cu  o g  X  N=274  4.  _J  3.  3,  2.  2.  0.2  0.2  A g e (years)  A g e (years)  First B e a c h Q . maculosus vertebrae  Y« 3. 5 7 0 +• 7.805 8  x log X  N=178  7. 6  a 5 fc  c  3. 2 1  0.2 A g e (years)  4.  F i g u r e 10 Length-age r e g r e s s i o n s o f F i r s t Beach 0 . maculosus d e t e r m i n e d by o t o l i t h s , v e r t e b r a e and l e n g t h frequency a n a l y s i s (males and females combined)  Table 4  Oligocottus  maculosus ( F i r s t  Beach)  Length-age r e g r e s s i o n s t a t i s t i c s three d i f f e r e n t log  from  methods o f age d e t e r m i n a t i o n  x v s y: y = u -f v l o g x  Statistic  Otoliths  Vertebrae  u  4.509  3.570  4.340  v  5.076  7. 805  4.892  Lower 95% c o n f i d e n c e limit of v  4.702  7.162  4.708  Upper 95% c o n f i d e n c e l i m i t of v  5.451  8.447  5.076  365  178  274  H  Cassie*s method  Table  Comparison o f by  otoliths  and  5  age  determined  vertebrae of  same O l i . g g c o t t u s m a c u l o s u s  Vertebral age (years)  0:  age age  -:  1  0  25  1  2  8  107  4  3  11  21  0  10.7  86.4  determined determined  age  i n d i v i dua I s  O t o l i t h age ( d i f f e r e n c e from v e r t e b r a e ) -1 0 +1  % of total  +:  the  by o t o l i t h s by  determined  otoliths by  2.8  greater than egual  otoliths  less  by  vertebrae  t o t h a t by  vertebrae  than  by  vertebrae  41  for age one and difference  two f i s h .  between  The  the  length) and few  possible  regressions  vertebrae a n a l y s i s c o n t a i n (total  other  few  is  reason  that  juveniles  for  the  the samples f o r  less  than  3.50  cm  f i s h i n the one year age group, compared  with the data f o r o t o l i t h and l e n g t h - f r e g u e n c y a n a l y s i s . „ Taking these f a c t o r s i n t o  consideration,  agreement  determination  in  age  there  appears  to  be  good  between o t o l i t h s and C a s s i e ' s  method of length-freguency a n a l y s i s , and good agreement  between  these methods and v e r t e b r a e f o r a t l e a s t age one  fish.  Further  validation  of  two  major  and two age  groups  p r o g r e s s i v e l y s m a l l e r numbers o f t h r e e and four year fish  existing  in  length-frequency variation the  histograms  for  over one  is  fish  were  experiments  (in  pools  produce more l a r g e r s i z e d  provided by a n a l y s i s of  time  caught. up  Collections  to about the 2.5  of  February  and  growth May  takes (probably  place  The  behavioural  m level)  young  starting  tended  to  collections  there  of  the  year  and J u l y , t h a t t h e sometime  A p r i l or e a r l y May)  u n t i l about J u l y or August, a f t e r which growth  The  F o l l o w i n g t h e modes over  pools below about 3 m between May  majority  for  f i s h than p o o l p o p u l a t i o n  the months shown makes i t e v i d e n t t h a t the in  11).  (Figure  length c l a s s i s a r e f l e c t i o n o f  pools around the 2 t o 3 m l e v e l ) .  appear  group  which, and the r e l a t i v e height of the pool i n  the  (in  population  i n the s i z e of any  purpose  which,  the  age  and  is  and  between continues  very  little  ( p a r t i c u l a r l y i n older fish) u n t i l the f o l l o w i n g spring.  data suggest  t h a t the young of  the  year  appear  in  lower  42 •  N=75  MAY  N = 135  15  J  N = 278  25, 26, 29, 1975  SEPT  4 5 1975  OCT.  30, 31,  NOV. 11, 1975 il  tl  n^ N=76  15  N = 217  Uy  MAY  12, 1976  JULY  20, 23, 28, 1976  A U G . 8,9,10,12, 1976  15  N=230  „  I |  N=129  m  P] rn PI i  N=92  j  A U G . 25,26, 1976  S E P T . 29, 30, -1976  D E C . 5, 1976  m 15  N=206  M  a  F E B . 12, 1977  h i 1.0  3.0  5.0  LENGTH  F i g u r e 11  L e n g t h frequency  7.0  9.0  (CM)  h i s t o g r a m s o f 0. maculosus  t i d e p o o l s a t a mean s i z e o f about of  about  second their  4.0  cm  year they third  growth  period  entire  t h e end  t o about  no f i s h and  7.5  i s not  may  be  thus  inconsistent  somewhat l o w e r  mid-summer  1976  and  September u n t i l  that  cm  In  size their  mean s i z e and  prior  f o r other f i s h  Age-length  relationships  fish  i n mid-summer  that the  by  very  to  the  of growth  summer over  shown by  w h i c h was 1977  the  little,  (Sicker,  the  tagged  tagqed  showed  i t in  growth  length-frequency i f  following sprinq.  that  in  size.  described  conclude  been found  any The  growth data  are  t a q q i n q r e t a r d s qrowth, as  has  1971).  Beach  For (Figure of  the  12 and both  significant  regressions calculated  T a b l e 7) age  (p<.01).  and On  and  female  f o r males  l o g t o t h e b a s e 10 length  data.  the b a s i s of the  c o v a r i a n c e , w h i c h showed no  male  6.0  d i r e c t estimate  recaptured  occurs a f t e r  of  growing season.,  ( T a b l e 6 ) . , One  I t i s evident  made  mean  with o t h e r d e r i v e d v a l u e s , although  histograms.  First  mean  grow t o a  t h e amount o f growth  with  to  cm  no  consistent  insufficient  o f one  and  have been t a g g e d  period i s available,  fish  cm  a p p e a r t o grow t o a b o u t  year  While  by  3.0  significant  and  females  t r a n s f o r m a t i o n s were  Both  regressions  results  are  from  analysis  differences  between  r e g r e s s i o n s i n v a r i a n c e s , s l o p e s or  intercepts  Table  6  Growth o f t a g g e d Q l i g o c o t t u s  Month f i r s t trapped  May  Month l a s t trapped  Initial length (cm)  Final length (cm)  Growth (cm)  October  4.0 5.2 5.7 5.7 4.7  5.7 6.1 6. 5 7.8 5. 4  1.7 0.9 0.8 2.1 0.7  5.1 5.5 6.3 6.6 3.9  5.2 5.7 6.6 6.6 6.2  0.1 0.2 0.3 0 2.3  5.4 6.1 6.2 6.6 6.7 6.8  5.6 6.2 6.3 6.8 6.8 6.8  0.2 0.1 0.1 0.2 0.1 0  4.6 4.8 6.9  4. 8 4.8 6.9  0.2 0 0  December August  October November July  September  October  maeulosus i n d i v i d u a l s  October  November  45  50 r  FIRST BEACH Q MACULOSUS FEMALE (OTOLITHS) LOG Y = -0.8787 . 1.423 LOG X N = 228  5"r  FIRST BEACH Q_ MACULOSUS MALE (OTOLITHS) LOG Y = -0.7427.1.202LOG X NU159  <  u  ^ 1.0 UJ  1.0  3  Q2  02  LENGTH (CM)  LENGTH (CM) FIRST 5 0  r  B E A C H  L O G Y =  Q  MACULOSUS  - 0 . 9 1 3 7 • 1.459  L O G X  (OTOLITHS) N =  365  CO  cr < UJ  1.0  .> UJ  3  0.2 LENGTH  F i g u r e 12  (CM)  A g e - l e n g t h r e g r e s s i o n s of F i r s t B e a c h O. m a c u l o s u s . d e t e r m i n e d b y o t o l i t h s ( m a l e s a n d f e m a l e s s e p a r a t e l y .and combined)  Table First  Beach  7  Oligocottus  maculosus  Age-length r e g r e s s i o n  log  Sex  (otoliths)  statistics  x v s l o g y: y = a + b (x-x)  a  b  x  N  s.e. of b  F  244.0  Females  .08  1.42  .67  228  .091  Hales  .05  1.20  .66  159  .12  Combinea  .08  1.46  .68  365  .085  analysis  of  249.7  covariance  Source  variance  Different slopes Different  (b)  intercepts  (a)  1.33  .024  2.07  .151  .464  .01  86.73 .01  Males vs f e m a l e s  Common  p<  .496  .01  til  (p>.01) length  (Table 7 ) , the  was  into  (Figure  12  a  and  single  Table  age-  7).  This  (p<.01).  areas  m a l e s and  regression  For  calculate combined  Ranee  separate  covariance  where t h e  (Table  places  except Grappler  calculated  variances females  are Inlet  shown.  and  (p>.01).  probably  relative  r e g r e s s i o n s are slopes  (Figure  to  agreement w i t h  was  due  in  there  males.  The  a  female  13 and  Table  (p<.01).  In  The  Analysis  regressions  places  except  case  three  which  t h a t f o r a l l areas a r e no  significant  in  10). the  variances,  regression  The  Pachena  agreement  between  difference  between  of  of G r a p p l e r  different for  were  combined  case of  preponderance  the case  to  single  Thus combined r e g r e s s i o n s  the In  data  area.  regressions  good.  to  thus  decimal  i s evident  evidently significantly  (p<.01).  two  were c o m b i n e d s i n c e t h e  intercepts  was  to  female  a l l significant  regressions  male and  Pachena P o i n t  f o r each a r e a are  It  significant  were i n s u f f i c i e n t  close to zero,  and  all  calculated for this  show v a l u e s  intercepts  regressions  slopes  was  was  were  female r e g r e s s i o n s ,  between male and  and  P o i n t the  9)  probability  differences  there  s t a t i s t i c s comparing  f o r each area  slopes  male and  calculated separately for  ( T a b l e 8)  Island,  regression only  decimal  statistics  f e m a l e s a t each a r e a  (p<.01).  and  Beach  also significant  The  of  were combined  regression for First  regression  Other  data  males  in  age  two  Inlet,  the  variances shows  r e g r e s s i o n s f o r both sexes f o r a l l places  good except  Table 8 All  areas Oligocottus Age-length  maculosus  regression  M a l e s and f e m a l e s  log  (otoliths)  statistics  separately  x v s l o g y: y =a -f b ( x - x )  Area  P<  s.e. of b  Cape Beale  F M  .21 .27  1. 18 1. 23  .63 .66  41 23  . 10 .15  140.32 57.153  .01 .0 1  Benson Island  F -.092 M -. 14  1.31 1.20  .55 .52  36 30  .12 .13  12 5.00 88.802  .01 .01  Port Renfrew  F M  .18 .18  .67 .66  49 18  .10 .17  68.239 20.4 94  ,01 .01  Pachena Point  F M  . .32  1.47 1. 42  .70 .76  46 26  .062 .14  556.91 95.790  ,01 ,01  Helby Island  F M  . 17 .21  1.43 1.56  .65 .68  50 37  .14 .19  109.83 64.981  .01 .01  Haines Island  F M  .23 .18  1. 33 1.36  .73 .71  42 30  .11 .21  146.43 40.338  .01 .01  Grappler Inlet  F M  .11 .22  .846 1.37  .67 .74  23 17  .16 .054  28.891 629.33  .01 .01  .796 .772  Table  All  areas a n a l y s i s of covariance Males vs  Beale  Benson Port  Island  Renfrew  Pachena Belby  Point  Island  Haines  Island  Grappler  va:  Inlet  statistics  females  P  F (b)  P  F (a)  P  1. 47  . 14  .092  .76  1.85  . 18  1. 17  .34  .41  .53  . 119  .73  1.08  .40  .15  .90  .602  . 44  2. 60  .003  . 15  .70  .263  .61  •1.57  .07  .32  .57  .059  .81  1.02  .49  .012  . 91  2. 22  . 14  5. 42  .001  .00 3  3.69  .06  F (va)  Area  Cape  9  variance,  10.07  b: s l o p e s .  a:  intercepts /  50  F i g u r e 13  Age-length r e g r e s s i o n s of 0. maculosus from e i g h t areas determined by o t o l i t h s combined)  (males and  females  51  H E L 8 Y I S L A N O Q M A C U L O S U S (OTOLITHS) L O G Y . - Q 7 6 7 7 . 1.4.47LOG X N * 87  L E N G T H (CM)'  ""l  'Of  RANCE I S L A N D Q M A C U I O S U S F E M A L E (OTOLITHS) L O G Y • - 0 . 4 6 9 1 . 0.8721 L O G X  MALE N . 39  L E N G T H (CM)  MACULQSLS  HAINES I S L A N D Q L O G Y • - 0 . 7 7 8 0 • 1.362 L O G X  (OTOLITHS) N = 72  G R A P P L E R I N L E T Q MACULOSUS (OTOLITHS) L O G Y . - 0 . 6 9 2 6 • 1.216 L O G X N*40 '  02  LENGTH (CM)  F i g u r e 13  (continued)  L E N G T H (CM)  Table  All  areas O l i g p c o t t u s Age-length  10  maculosus  regression  H a l e s and f e m a l e s log  area  Benson I s l a n d Port  Renfrew  Pachena  Point  statistics combined  x v s l o g y: y = a + b ( x - x)  a  Cape B e a l e  (otoliths)  b  x  N  s.e. of b  F  p<  .24  1.23  .66  57  .094  169.60  .01  -.057  1.33  .58  44  .11  146.12  .01  .68  62  .086 106.63  .01  .18  .872  .27  1.46  .72  72  .061  570.60  .01  Helby I s l a n d  .19  1.45  .66  87  .11  173.19  .01  Haines I s l a n d  .21  1.36  .72  72  .10  201.77  .01  Grappler  .16  1.22  .70  40  .088 189.96  .01  .69  39  ,14  Inlet  Ranee I s l a n d  .13  .872  40.570 .01  53  Port  Renfrew,  but  difference  may  year  male  old  be  the  at  least  and  s m a l l sample s i z e ,  regression  no  may  corresponding  good a g r e e m e n t betwen  other  places  significant  difference  relationships into  in  statistics for a l l sites the  different  significantly relevant  and  between  male  Inlet,  f o r the (Table  the  Ranee I s l a n d  are  four  given  the  female r e g r e s s i o n s at  a l l  obvious  and  reason  female  for a  age-length  r e g r e s s i o n s were combined Analysis  of  covariance  show s i g n i f i c a n t  differences  the  slopes  adjusted  that  which,  differences.  any  Inspection  suggests  one  view  the  11)  The  In  area.  lines;  (p<.01).  statistics  Renfrew and  of  regression  (p=,01)  male and  absence  Grappler  a single regression  between  females,  produce s i g n i f i c a n t  the  does n o t .  p a r t i a l l y a c c o u n t e d f o r by  of the  and  f o r females  group  of the  different  are  the  significantly means  lines  and  regressions  from t h e  differ their  for  other  Port  sites.  3.Discussion  The First age  Beach r e v e a l e d  group.  whose s t u d y at that  Port  there  of  age  three  determination  m a j o r age  i n 0.  g r o u p s and  T h i s i s i n c o n t r a s t t o the  work o f  o f age  vertebrae  Renfrew  collection July  study  determination  and  Bruels Point, California  were s i x age of  a t which  using  fish time the  he  groups at both made at P o r t young  of  the  a fourth  smaller  Chadwick  (1976)  o f 0.  l e d him  localities. Renfrew was year  majculosjjs a t  have  maculosus to  suggest  The  single  taken  in  appeared  late in  T a b l e 11 Analysis of covariance All  sites  males a n d f e m a l e s  Source  d. f .  D i f f e r e n c e between g r o u p means s l o p e and common s l o p e  combined  Mean Sgua re  1  .055  Group means from their line  7  ,50  Between i n d i v i d u a l lines  8  .034  820  .014  About i n d i v i d u a l lines  Probability  o f common v a r i a n c e  S l o p e o f group  means  Common s l o p e o f l i n e s  0 1.35 1.62  ,05  .01  55  tidepools.  Chadwick*s  length-frequency  distribution  ( u s i n g a moving a v e r a g e  o f t h r e e and s t a n d a r d  p r o m i n e n t mode a t a b o u t  2.5 cm and a n o t h e r  about  1.5  cm.  respectively.  These  correspond  I f age c l a s s  cm  be p o s s i b l e t o f i n d  (about  3.0  Reference no I  such  cm  total  I was r e c r u i t e d  analysis  Beach i s  histoqrams  for  July  between  p r o m i n e n t one a t  of l e s s than and  2.5 Hay.  o f F i g u r e 11 shows  a n d shows t h a t C h a d w i c k ' s 0 age g r o u p d e f i n e d  and  here.  of the length-frequency histoqrams f o r  supported  by  Green*s  (1967)  length-frequency  0. m a c u l o s u s a t B o t a n i c a l B e a c h , and  (1939) l e n q t h - f r e g u e n c y  classes,  i n t o account, i t  October  histograms  t o t h e one y e a r  one  to the population i n  class of f i s h  length)  s i z e c l a s s a t t h a t time  This First  a size  t o the length-frequency  y e a r - c l a s s e s belong  less  shows  t o h i s I and 0 age  t h e p r e v i o u s y e a r , and t a k i n g g r o w t h a f t e r should  length)  curve  histoqrams  for  Atkinson's  0. m a c u l o s u s  at  Puqet  Sound, W a s h i n g t o n . Other are  few., Chadwick  Clinocottus concluded  o f age d e t e r m i n a t i o n i n m a r i n e  (1976) c o n d u c t e d  globicep.s  (Girard),  age d e t e r m i n a t i o n and  as  t h a t t h e r e a r e s i x age c l a s s e s .  ageing  in  cottid  which  He  studies  examined  the  more  has been fish  b a s i s of o t o l i t h  up  euryhaline  25 cm  readings suggested  (1975)  examined  the  analis  (Girard)  in  age-length California.  Weiss  studies i n  O. , m a c u l o s u s  (1969) examined  L e p t p c o t t u s arm,atus G i r a r d , a  r e p o r t e d t o grow t o about  for  cottids  t o 46  cm  (total  l e n g t h ) and on t h e  t e n age  groups.  relationship On  the  (Hart,  of  basis  1973).  HcElderry Clinocottus of  otolith  56  examination few  fish  he c o n c l u d e d  were  conducted  more  ageing  t h e r e were s e v e n  than  two  studies,  Scor p a e n i c h t l j Y s mar mora t u s found  thirteen  frequency Sound is  data  suggest  recorded  using  that to  rotenone  and n i n e  to  population,  the approximate  those  0. m a c u l o s u s  O'Connell in  the  plus  essentially  two age g r o u p s  (1953) cabezon,  c o t t i d , and  males.  10  cm  (Starks),  Length-  {Hart,  a cottid  which  1973), c o m p a r a b l e t o  t h r e e age g r o u p s w i t h i n t h e  mean l e n g t h s o f w h i c h c o r r e s p o n d  age  groups.  Although  subtidal  cottid,  ( B r u c e Leaman, p e r s o n a l  appears  to  t h e r e a r e fewer  d a t a on w h i c h t o base c o n c l u s i o n s , A r t e d i u s l a t e r a l l y another  although  c o l l e c t i o n s i n k e l p beds i n B a r k l e y  0. m a c u l o s u s , shows two and p o s s i b l y  for  otoliths,  Artedius harringtoni grow  old.  (Ayres) , a l a r g e s u b t i d a l  year o l d females from  years  age c l a s s e s ,  (Girard),  t o show a t l e a s t  communication).  57  IV.  1.Tagging  Tagging  VARIABILITY  IN  HOMING BEHAVIOUR  and r e c a p t u r e methods  methods  Tags coloured  for  b e a d s and  were u s e d .  adult  maculosus  monofilament  line.  A p i e c e of monofilament  heavier  when  threaded  w i t h a bead  this  knot,  leaving  one  long.  Another  bead  line.  0.  A large  was  not  and  end  (usually  available)  3 cm  of  three  Nineteen c o l o u r s o f  beads  4 or 6 l b test,  about  attached with a double  about  was  consisted  20 cm  long  improved  long  number o f t h e s e were made p r i o r  end  was  cinch  l o n g and t h e o t h e r a b o u t  t h e n t h r e a d e d on t h e  but  15  of  to tagging  cm the  fish,  to e x p e d i t e the a c t u a l t a g g i n g process.  The a  fine  l o n g end  needle  and  o f the monofilament the  musculature  of the f i s h  right  sides  of the f i s h .  line  and  bead that long  was  the then  the end  line  on  A  loop f i v e  between t h e d o r s a l Another  bead  through fins  was  down t o t h e r i g h t  both s i d e s  of monofilament  through the  inserted  threaded  was  then  side  through  the  dorsal  from t h e l e f t  threaded  t h r e a d e d o n c e more t h r o u g h  manoeuvred  beads  needle  was  onto  the  t h e bead. of the  This  fish  s e r e a d j a c e n t t o t h e body. l o o p e d and  i t s free  end  to  so The  wound  or s i x times.  dissecting  needle  was  inserted  in  the loop  and  58  pushed ends  down t h e l i n e  t o the outer s i d e  of monofilament  tighten.  were t h e n  with  the technigue  tagged i n w e l l under All individual  one  fish  were  recognition  fish  g r e a t e r t h a n 2.5 cm,  overnight  almost  knot t o  a  fish  Once  could  be  unigue  Over  as i n d i c a t e d was  five  months.  fish  tags  2800 f i s h  were  that tagged  Tag l o s s ,  from  by r e c o v e r y o f f i s h very  low  f o r varyinq  after  with  (about  256}.  p e r i o d s from  The o n l y m o r t a l i t y  occurred  so  recorded  blockages  in  the  system. Several  juveniles.  methods  were  P i e c e s of v a s e l i n e  and l a r g e  coloured  (8 mm  surveyors  musculature the  with  musculature  u n t r e a t e d , tagged  seawater  tagged  were k e p t i n t h e l a b o r a t o r y to  free  were t h a n c u t o f f .  No a n a e s t h e t i c was u s e d .  on t h e d o r s a l  long,  causing the l a s t  gained,  was p o s s i b l e .  t h i s technique.  Taqqed f i s h  was  Both  minute.  using  from  pulled,  The f r e e ends o f m o n o f i l a m e n t  familiarity  lesions  of the bead.  with  taqs either  1  x 3 mm) tape  nylon  attempted  for  the tagging of  coated nylon thread and s m a l l (3 mm  were  attached  about  square)  through  5  pieces of  the  dorsal  t h r e a d . , However, w i t h i n s e v e r a l  came o f f  or  entangled  the  fish  in  cm  days,  assorted  debris. Finally beads lb  (one bead  test)  relatively  juvenile  each  side  fish  o f t h e body) and m o n o f i l a m e n t  i n t h e same manner satisfactory  were t a g g e d w i t h two c o l o u r e d  except  as a d u l t for fish  fish.  This  line  method  l e s s t h a n about  (2  proved 2.5  cm  59  (total was  length)  find  every  homing b e h a v i o u r  tagged  fish  f o u r hours around f o r capture  two methods were  r e l e a s e . ftt F i r s t  possible  thoroughly  f o r tagged  captured  release  and  pools  thoroughly,  combination Grappler  Beach a b o u t  experiment. made  trappings,  examined,  The  experiments and  is  conducted  there,  addition,  pools  were  pools  in  the  conducted  observation were  times  Simultaneously,  schedule  In  although  set,  24 h o u r s ) , a t l e a s t  for  for  and  inspected which  fish  latter  less  where o t h e r w i s e s t a t e d .  traps  several  tidepools  once  t r a n s p l a n t e d and p o o l s between t h e home and  of t r a p p i n g  (approximately  used  T r a p s were s e t o n l y i n p o o l s  0. m a c u l o s u s .  homing  Inlet,  freguently  between  were  except  In  los tide.  and t r a n s p l a n t i n g o f f i s h .  whenever  also  after  experiments  two weeks, b a i t e d minnow t r a p s were s e t i n  about  were  individual tag  methods  In  used  satisfactory  developed.  Reacapture  to  f o r w h i c h no r e a l l y  tagged  outlined  per  the  In  other areas, a  also two  every  other  used.,  tidal two  for the f i r s t  o b s e r v a t i o n s from  fish, in  week  was  over  once  in  In  cycles  weeks  and  month o f an  the surface  were  places, the recapture  descriptions  of  experiments  60  2. Homing, i n d i f f e r e n t  First  locations  Beach  Home r a n g e and a r e a  Since  fidelity  the  majority  b e h a v i o u r o f 0. m a c u l o s u s it  was  size  regarded  have been c o n d u c t e d a t  as e s s e n t i a l t o investigate  homing  studies.  u s e d a s i n Khoo  normal  at  First  into Port  homing Renfrew,  p o o l f i d e l i t y and Beach,  I n t h i s s t u d y , the term  prior home  (1971) a s t h e a r e a c o v e r e d by t h e f i s h  to  range during  travel, Gersbacher  suggested strict  that  fidelity  and  the to  neighbouring  majority  pools.,  He  Green's  metres from  determine whether  of f i s h  size  observations that their  Khoo's  home  (1967)  display  rather  (1971)  studies  a group o f  t h e amount o f movement  of  the  few f i s h  pool,  Green  move w i t h i n  suggested that on t h e  and  0. m a c u l o s u s  pools.  maximum e x t e n t o f t h e home r a n g e Three  (19 30)  of  the majority  between p o o l s may depend Despite  Denison  particular  however, showed t h a t  three  investigations  o f home r a n g e o f 0. m a c u l o s u s  commencing is  of  Khoo  pools  involved.  are found  more t h a n  proposed  that  the  was n o t more t h a n 30.5 m e t r e s .  experiments  were  conducted  0. m a c u l o s u s  exhibited  specifically  pool f i d e l i t y  at  to First  61  Beach.  These  indication other  experiments  were  also  designed  o f t h e s i z e o f t h e home r a n g e .  experiments  untreated,  on  tagged  the  pool  0. m a c u l o s u s  to determine t h e degree  to  A n a l y s i s of data  movements  after  homing  i n d i v i d u a l s was a l s o  of natural  give  movement between  an from of  undertaken  tidepools.  Methods  In t h e f i r s t were  tagged  captured. In  Fish  the second  and  returned  third two  were c o l l e c t e d  experiment,  groups which  a total  from  t h e home  f o ranalysis.  pool only, were  i n which  maculosus they  were  tidepools. were t a g g e d  t h e y were c a p t u r e d .  Fish  pools each.  were  In the  were c a p t u r e d  from  and r e t u r n e d t o t h e p o o l  pool  in  which  a  fish  was  i n any p o o l o v e r  four  pool.  o b s e r v a t i o n s o f tagged f i s h  months, 10 weeks and f i v e  0.  o f 76 0. m a c u l o s u s  o f 44 O. m a c u l o s u s  The  33  adjacent  o f t h r e e and f i v e  t h e y were c a p t u r e d . was t e r m e d  of  three  o f two and t h r e e p o o l s each  All  used  total  the tidepool  a total  f r o m two g r o u p s  experiment,  captured  weeks, r e s p e c t i v e l y ,  Percentages o f tagged  i n t h e home a n d n e a r b y  pools  were r e c o r d e d and  fish and  found in  i n t h e home  distant  pools  calculated. To  provide  an  i n d i v i d u a l 0. m a c u l o s u s , the  to  a  r e t u r n e d t o t h e p o o l i n which  collected  in  and  experiment  second  replacement  indication  location experiment  of  records  the of  pool f i d e l i t y of individuals  seen a t l e a s t  f i v e times  from over  62  a period  of at least  regarded  as  28 d a y s  having r e l a t i v e l y  p e r i o d o v e r which  the f i s h  which  was  the  p o o l were  were  fish  analyzed.  extensive  was s e e n ,  seen  These  location  the  and t h e number  number  fish  were  records. of  The  pools i n  of t i m e s seen i n each  calculated.  Results  The (Table fish  results  12) show  that  of  the  i n each  or  sliqhtly  pools.  replaced  found  In each case, a in  away  Several  from t h e home It  of  fish  show  This  about  were f o u n d i n d i s t a n t  the  were f o u n d i n t h e of  that  the  fish  (either the  i s , the  home  p o o l s from  30 t o 60 m  to a small  percentage  pool.  stray  i n addition  widely, equal proportions  to a particular  pool,  as  qroup comprises u s u a l l y  occasionally  about  t h e a r e a o f t h e home p o o l  thus appears that  which  fidelity  pools. but  fish  30 t o U0% o f t h e  majority  home p o o l o n l y o r t h e home and n e a r b y p o o l s ) , range.  experiments  i n t h e home p o o l .  h i q h e r p e r c e n t a q e s of f i s h  home and n e a r b y were  replacement  experiment, about  were s u b s e q u e n t l y f o u n d o n l y  same  three  up t o f i v e ,  coverinq  to  a  o f 0. small  maculosus qroup  a b o u t two o r t h r e e  of  pools,  an area with a d i a m e t e r o f  10 t o 15 m maximum. From t h e a n a l y s i s o f l o n q t e r m  second replacement experiment 0. m a c u l o s u s  show  strict  (Table  fidelity  p o o l f i d e l i t y from  13) i t i s e v i d e n t over  a lonq  that  period  the some  t o one  Tafcls  First  Date r e l e a s e d  12  Beach r e p l a c e m e n t  experiments  28 May 7 5 2 J u n e 75  Number r e l e a s e d  Total  30 J u n e 77  9 Aug 77  19  11  33  76  11  9  2  11  27  17  i n home  17  14  33  36  39  Number f o u n d i n home and n e a r b y p o o l s P e r c e n t f o u n d i n home and n e a r b y p o o l s  7  2  9  10  18  37  11  27  53  11  16  1  20  67  35  81  29  61  . 88  80  0  2  2  2  2  0  11  6  3  5  Nunber f o u n d pool cnly Percent found pool cnly  i n home  Number f o u n d i n home p o o l and home a n d n e a r b y p o o l s P e r c e n t f o u n d i n hone p o o l and heme and n e a r b y p o o l s  Number f o u n d i n d i s t a n t pools Percent found i n d i s t a n t Fools  Table  Long  tern  (long  One  pool  Cays o v e r which observed  61 61 75 75 74 61 38 61 52 5fi 34  Two  Number of times observed per p o o l  term:  75  in 75 71 75 48 58 58  71) 30 43  *  fidelity  always  of i n d i v i d u a l  from  30 J u n e  seen  at least  replacement  Three  poels  Days e v e r which observed  12 25 20 18 25 11 17 7 12 5 7.  pool  Number o f times observed per p o o l  17,1 15.3 18,7 9,8 26,1 17,1 17,1 7,2 6,1 6* 8,0  found  13  five  experiment over  reels  Days o v e r which cb s e r v e d  pocl  at least  Four  Number c f times observed per p o o l  19,1,1 5 1,2,1 17,4,2 7,2,1 7.5,2 4,3,3 2,2,1  75 75 75 42 60 41 37  in a different  times  Q l i g o c o t t u s maculosus  from  the pool  days)  pools  Days o v e r which observed  58 60 60 57  28  Five  Number of times observed per p o o l  7,4,4,1 15,7,2,2 13,5,4,1 5,2,1,1  of first  capture  pools  Days o v e r which observed  70 75  Number of times observed per pool  10,8,7,1,1 12,4,3,2,1  65  pool.  T h i s was  the 0.  fish  after  was  in seen  homing a l w a y s  their  e v i d e n t from homing 17  in  in  1976  after  another  the  of  the  majority  c a s e s , t h e r e i s one  records  the f i s h result  from  0.  of  times  one  p o o l and  pool in  each  and  maculosus  four  being  18,  6,  28 d a y s ,  pool  4  1.,  pool.  five  that  101  pool.,  that  in  i n two  days,  of pools  location  was  seen  fish  times  times  over a p a r i o d  p o o l and  o f t h o s e 56,  42  maculosus  56  (75%)  were  in  times  each  pool  involving and  of at  were least  found  were f o u n d  other pool.  one  o f 29  in  homed  of  15 t i m e s i n  11 t i m e s  a total  successfully  after  pools a t o t a l  one  a  i s shown.  pool  10 homing e x p e r i m e n t s  o n l y i n one  t h e m a j o r i t y o f O.  some  individuals  t h e group  of  were  that  greater f i d e l i t y  more than t w i c e as many t i m e s a s any  appears  which  months l a t e r i n  shows  analysis  days  over l o n g p e r i o d s of time  of  which  days,  to a particular  range  were s e e n  From  one  179  doubt  other, the other f i s h  seen at l e a s t  one  fish  no  d a y s , t h e number o f  fish  37 were f o u n d  than  fish  a period  and  tagged  subsequently  over  two  pool within  i n the other; a f i s h  p o o l s o v e r 83  untreated,  more  over  seen  of  41 t i m e s o v e r 83  is  home  i s s u p p o r t e d by  once i n the  five  the  moves, t o which  homing: f o r example, two 16  pool;  some  example,  number o f t i m e s t h a t  i n pools  Again, t h i s  seen  fidelity  were o b s e r v e d  which  for  homing  There  very s t r i c t  Examination  around  data of  were f o u n d e l e v e n and t w e l v e  display  of  was  home  o r i g i n a l capture pools.  maculosus  the l o c a t i o n  experiments:  times  i n t h e same p o o l ;  released  0.  used  maculosus  always  also  in  in  one  Thus  i t  show q r e a t e r f i d e l i t y  66  to  one  This  particular  p o o l than t o other pools i n  p o o l i s termed t h e p r e f e r r e d  the  home  range.  pool.  Homing b e h a v i o u r  Two  experiments  d e t e r m i n e w h e t h e r 0. behaviour,  that  displaced.  The  difference,  were  maculosus  i s , return  conducted  at F i r s t  any,  also  undertaken  involving to  homing  designed  to  compare  the  i n homing b e h a v i o u r between r e c i p r o c a l  non-reciprocal transplants of f i s h . homing e x p e r i m e n t s  demonstrate  to  t o t h e home r a n g e when e x p e r i m e n t a l l y  e x p e r i m e n t s were a l s o  i f  Beach  specifically  u n t r e a t e d , t a g g e d 0.  provide  homing b e h a v i o u r a t F i r s t  A n a l y s i s o f d a t a from  a  more  other  maculosus  comprehensive  and  was  picture  of  Beach.  Methods  In 0.  maculosus  to  a  any  the  taken from  p o o l about pool  first  over  60  experiment, a group  m away.  a  four  a  of f i v e  total  of  p o o l s were  42  transplanted  A l l o b s e r v a t i o n s o f tagged  month p e r i o d  tagged  were r e c o r d e d and  fish  in  used f o r  analysis. The effect in  second  and  subsequent  experiments i n v e s t i g a t e d  o f a p p r o x i m a t e l y m a i n t a i n i n g t h e number  the study t i d e p o o l s  experiment  0.  maculosus  while t r a n s p l a n t i n g were c o l l e c t e d  of  fish.  from two  0.  maculosus  I n the  groups  the  of  second pools  67  about pool  60  m apart.  i n one g r o u p  A total  and 34 from  Releases  were  most  fish  were t a k e n , s o t h a t  and  34  and  into  i n one p o o l  Data c o l l e c t e d  homing  20 f i s h  60  m  apart.  so that  egual  to  Reciprocal  removed  were removed month  period  the  number o f f i s h  other  group).  in  1977,  about  experiment  were l e s s t h a n 4.0 cm  untreated,  1977,  16  In 25%  the  of  1977  the  (4 J u l y , of  pools  to  each  pool  (and t r a n s p l a n t e d t o  first  the  (total  on 23 A u g u s t ,  July  two g r o u p s  removed  were used i n t h e a n a l y s i s  experiments s t a r t i n g  tagged  examine  experiment  transplanted  case f o r t h e other experiments.  month p e r i o d  involved  t r a n s p l a n t s were made a s f a r a s  t h e number o f f i s h  conducted  collected  which t h e  were  designed t o  were t a k e n f r o m  the  July  group.  experiments of  used i n s e v e n e x p e r i m e n t s  pools i n  4  other  over a four  performance  In a l l cases, f i s h  possible,  the  the  t h e p o o l i n each g r o u p f r o m  b a s i s o f homing, and one t r a n s p l a n t  1977).  the  in  a n a l y s i s o f d a t a from o t h e r  the  0. m a c u l o s u s  about  pools  i n the analysis.  examining  sensory  four  were t a g g e d , 20 f r o m one  and i n t h e o t h e r p o o l 2 2 f i s h  20 were i n t r o d u c e d .  The  was  made  introduced,  were used  o f 54 f i s h  fish  two  experiments  r e l e a s e d i n each  length),  which  Data c o l l e c t e d except i n  was  not  over a f o u r  the  cases  of  1976, 15 S e p t e m b e r , 1976,  and 9 A u g u s t  1977 f o r w h i c h  data  o v e r 3.5, 2.5, 2+, 2 and 1+ months, r e s p e c t i v e l y ,  were  used.,  Following regarded  the  as s u c c e s s f u l  method  of  Khoo  (1971),  i f a t a g g e d 0, m a c u l o s u s  homing  returned  was  to i t s  68  home range pool  ( e i t h e r t h e home p o o l or nearby  i n the home  range  pools).  The  i n which a tagged f i s h was found was  regarded as t h e pool to which the f i s h homed, even i f subsequently fish  was l o c a t e d i n another pool.  l o c a t e d i n pools  first  other  than  those  the  fish  A r e c o r d was kept o f of  the home  range.  C a l c u l a t i o n s were made o f the percentage o f 0. maculosus homing, the  percentage  of  homers  returning  t o t h e home p o o l and the  percentages o f r e l e a s e d f i s h remaining i n  the  transplant  area  f o r any period of time.  Results  Although  there  i s considerable  r e s u l t s from the ten homing experiments the  variability  (Table 14) and  i n the some of  percentage r e t u r n s are s m a l l , t h e r e i s d e f i n i t e evidence t o  i n d i c a t e that a m a j o r i t y of 0. maculosus i n d i v i d u a l s a r e a b l e t o r e t u r n t o a r e l a t i v e l y s m a l l area when d i s p l a c e d . In  nine out of the t e n experiments the percentaqe  successfully  hominq  fish  which  returned t o the home p o o l was  c o n s i d e r a b l y g r e a t e r than t h e percentage r e t u r n i n g to range.  Although  to  the home  many o f these f i s h were subsequently found i n  other pools i n the home range, returned  of  the  home  2» I§.£Slosus to r e t u r n t o  the fact  pool a  that  demonstrates  small  area  from  they  initially  the a b i l i t y  of  a considerable  distance. It  i s interesting  to  note t h a t i n a l l experiments,  Table  Honing  performance  of untreated at  Date r e l e a s e e  Number  released  Number homed P e r c e n t homed  Nuaber found i n t r a n s p l a n t area Percent found i n t r a n s p l a n t area  Days s p e n t i n t r a n s p l a n t area a. h e m e r s  non-homers  P e r c e n t o f homers returninq to home p o o l  26 Hay  11 &uq  4 Sept 1975  21 Sept  14  First  O l i g o c p t t u s magulgsus  Beach  16 Hay  23 Auq 1976  15 Sept  July  16 July 1977  9 Auq  42  54  69  33  38  31  20  60  45  32  15 36  30 56  52 75  24 73  20 53  15 4ft  14 70  31 52  19 42  20 63  6  8  17  3  8  8  4  27  26  14  14  15  25  9  21  26  20  45  58  44  1,1  1,1,2, 2,4,7  1,1, 1,1  1, 102  73  1 3 , 2 2 , 1 , 1 , 1, 4 25,41 1,1,4, 6,6,7, 17,18 1,21, 6,6,7, 3,78 42,82 13,22, 42  87  73  60  2,27  2,2, 73,73, 83, 110  70  2  1,2, 10,11, 46,80, 104  60  10,26  64  1,1,2,2, 2,4,10,10, 16,17,17, 18,21,21, 22,26,26, 44,54,57,58  65  2,2,2, 2,5, 42  3,3, 8,18  1,1,1,2, 1,3,3,3, 2,5,6,6, 8,8,11, 6,13,14, 11,18,30 14,14,27, 27,32,55,55 59,59  42  60  70  considerable transplant days)  pool  after  majority there this  numbers  being  of  i s  area  (from 9% t o 58%) o f f i s h f o rvariable released.  0. m a c u l o s u s  a  reasonable  behaviour.  periods This  number o f f i s h fish,  o f time  implies  individuals  Of t h o s e  remained  transplant  increasing  the density  be  homing  which  do n o t d e m o n s t r a t e percentage  area  i s influenced  A number o f f i s h at  l e a s t once i n p o o l s  range. home the  The m a j o r i t y ranges.  and  of f i s h  in  a  and t r a n s p l a n t e d  s e v e n d a y s and t h e h i g h e r  t y p e o f movement outside  pool  was o b s e r v e d  after  or  pool  release. between  and t h e home  the  day  of  days, t h e n e x t  home p o o l  pool i n  o f 3 t o 33 in a in  a  highest  many t i m e s f r o m displayed  the  following  captured  a f t e r release,  Not a l l f i s h pools.  located  on t h e p e r i p h e r y  originally  i t s home r a n g e a f t e r f i v e  after  i n the  displaced t o a high  lower  Another f i s h ,  appear  I t i s possible,  was back i n t h e home r a n g e f o r a p e r i o d  in  days  not  that  factor.  were found  pool  98  shows  remaining  between t h e t r a n s p l a n t  was f o u n d  days a f t e r r e l e a s e . tidepool  t h e non-  w h i c h s u c c e s s f u l l y homed were  F o r example, one f i s h  intertidal  release  by t h i s  fish  (up t o  area.  t h e "space" does  a f a c t o r i n f l u e n c i n g s u c c e s s f u l homing.  transplant  behaviour,  and t h e r e c i p r o c a l t r a n s p l a n t s  however, t h a t t h e number o f non-homing  the  show  a considerable  or reducing  1 t o 10 4  while  o f t h e p e r c e n t a g e s homed between  reciprocal  to  (from  that  17% o f non-homing f i s h ) r e m a i n i n t h e t r a n s p l a n t  Comparison  i n the  this  17  high low pool to  apparent  Some were o b s e r v e d i n p o o l s  at  t h e edges o f t h e home r a n g e s i n t h e m i d - i n t e r t i d a l .  71  A few f i s h  were o b s e r v e d  from  release  the  in a  pool  series  of  t o t h e home r a n g e .  were f o u n d  out of the transplant  were f o u n d  i n p o o l s up t o 30 m away f r o m  in  the opposite direction  mid-intertidal  from  and home  pools  Several fish  areas  which  after  release  the transplant  a r e a and  t h e home p o o l .  Inlets  Home range  and a r e a  Homing  fidelity  behaviour  been examined i n t i d e p o o l Green,  1967;  Khoo,  (Grappler I n l e t , particularly  of  areas  1971).  has t o date,  ( G e r s b a c h e r and  Denison,  only 1930;  I n s e v e r a l a r e a s o f B a r k l e y Sound  Bance I s l a n d  in  O. m a c u l o s u s  Dodger  and  Dodger  Channel,  Channel)  seasonal  large,  but  populations  of  O. m a c u l o s u s c a n be f o u n d .  If when  homing i s d e f i n e d a s t h e r e t u r n  experimentally  displaced,  whether i n l e t  0. m a c u l o s u s  area.  experiment  One  display was  0. m a c u l o s u s c a n be r e g u l a r l y inlet,  i t i s  comparable t o a group  a  to  to  a  range  particular  determine  found i n a r e s t r i c t e d of tidepools.  home  necessary t o e s t a b l i s h  fidelity  conducted  to  area  whether of  an  72  Methods  a total Grappler they  Inlet  were  captured  were  and  Port  Desire.  a line  Of  released  and 15 f i s h  these  with  fish,  i n the small  site  on t h e s o u t h  and r e p l a c e d  were c a p t u r e d  at e a c h  parallel  from two s i t e s  tagged  captured.  inlet  in  o f 58 f i s h  i n the area  43  side of i n which  0. m a c u l o s u s  bay a b o u t h a l f w a y  and r e l e a s e d  a t the  along  the  mudflat  in  a s e r i e s o f minnow t r a p s was  the shore l i n e  and  covering  linear  o f a b o u t 7 t o 10 ra. T r a p s were a l w a y s s e t a t low  however  the l e v e l  depended  on t h e h e i g h t o f t h e t i d e .  o f t h e i n t e r t i d a l a t which  were s e t a t a h i g h e r  perpendicular  to  Trapping  conducted  was  two s i t e s .  the  shore  as  the  low  tides,  the  b u t i n t h e same t r a n s e c t lowest  and o b s e r v a t i o n s  Data were c o l l e c t e d  tide,  t h e t r a p s were s e t  at h i g h e r  water l e v e l  spread  a  distance  traps  were  trapping  site.  were made o n l y  o v e r a f o u r month  a t the  period.  Results  The suggest the  that  relatively i n inlets  low  percentages  t o a s m a l l area  shown by 0. m a c u l o s u s i n d i v i d u a l s found  52 only  was l o c a t e d t h r e e  and 64 d a y s a f t e r once a f t e r  fish  resighted  0. m a c u l o s u s i n d i v i d u a l s do n o t d i s p l a y  same d e g r e e o f f i d e l i t y  One t a g g e d f i s h  of  comparable  i n tidepools  r e l e a s e , and w i t h i n t h e f i r s t  site  were s e e n  month.  that  (Table 15).  times a t the capture  r e l e a s e ) , but t h e m a j o r i t y  to  One  (50, again  tagged  Table Grappler  Inlet  15  replacement  Bay  Date r e l e a s e d  Percent  resighted  Mudflat  2 October  43  Number r e p l a c e d  Number r e s i g h t e d  experiment  there  there  12  28  Total  1975  15  58  6  18  40  31  74  0. M a c u l o s u s  was o b s e r v e d  varying tide  levels  about  0.6 m below  from  the  a number o f t i m e s a t t h e s m a l l b a y , a t 0.5 t o 3 m.  water  This fish  level,  was a l w a y s  apparently  found  following  the  tide.  It  is  possible  0. m a c u l o s u s i s l a r g e r  than  is  that  also  possible  that  much  better  the  defined  However t h e low p e r c e n t a g e s duration  of  the  m a j o r i t y range  display  a degree  Horning b e h a v i o u r  Although shown by i n l e t experiments displays area  displaced.  in  of than  under seen  t o suggest  of  inlet  observed. ,  It  trapping  and  in  tidepools  consideration. again  over  the  that  some  inlet  area  but  that  Even those f i s h  which do  w i t h a s m a l l area appear  to  do  of time.  inlets  the  degree  conducted  homing b e h a v i o u r  (as  or  area.  of fidelity  0. m a c u l o s u s a p p e a r s  were  range  to a particular  a wider  period  along  are  appear  of a s s o c i a t i o n  so o n l y f o r a l i m i t e d  in inlets  trapped  experiment  over  efficiency  areas  O' m a c u l o s u s show some f i d e l i t y the  home  the areas trapped or  observations i s considerably less where  the  tidepool  to  t o be s m a l l ,  determine  in inlets,  to a particular  that  0. m a c u l o s u s )  two  w h e t h e r 0.  area  transplant maculosus  i s , returns t o a small when  experimentally  75  Methods  In  the  were c o l l e c t e d bay and  36  from  from  transplanted  first  t h e two s i t e s the  Data  In  fish  were  The  numbers  o f 78 0. ma cu lapsus  fish were  were not  42 f r o m t h e reciprocally  quite  equal).  a n d o b s e r v a t i o n s were made o n l y a t  were c o l l e c t e d  the  were c o l l e c t e d ,  the  a total  i n Grappler I n l e t ,  mudflat.  (although  T r a p p i n g was c o n d u c t e d two s i t e s .  experiment  second  o v e r a f o u r month  experiment a t o t a l  period.  of 88 p .  maculosus  64 f r o m t h e bay a n d 24 f r o m t h e m u d f l a t .  released  at a s i t e  a t the  These  a p p r o x i m a t e l y h a l f w a y between t h e  capture  sites.  appear  t o o c c u r i n any numbers, a s t h r e e d a y s by t r a p p i n g  to t h e experiment release, all  trapping  three sites.  analysis successfully area.  release  produced was Data  of  only  point,  four  conducted  0. m a c u l o s u s  does  0. m a c u l o s u s .  not prior  Following  and o b s e r v a t i o n s were made a t  were c o l l e c t e d  data  these  involved  over  a four  month  calculating  period.  percentages  homing and p e r c e n t a g e s r e m a i n i n g i n t h e t r a n s p l a n t  76  Results  The low p e r c e n t a g e s suggest  that  homing  successfully  behaviour  0. m a c u l o s u s f o u n d i n i n l e t s . was  c o n s i d e r a b l y reduced  In  the  0*  were fiaculosus  experiment bay) of  were may  experiment  at  the  that  tagged  fish  suggests that majority  did  percentage  not  Compared  with  developed  in inlet inlet  tidepool  situation.  from  Grappler Inlet  First was  fish  Beach.  removed  site  remaining i n  Few  exposed  the mudflat.  no  untagged the  first  area  (the  The  second  absence  experiment  the transplant  An  of  area, the  a s shown b y t h e  0*  maculosus  homing b e h a v i o u r i s n o t a s  attempt improved  17 O c t o b e r ,  was  made  homing  to  of  0.  well  in a  maculosus  tidepools  I n t h r e e o f t h e t i d e p o o l s an e g u a l number o f  a s was r e p l a c e d (31 f i s h ) .  in  determine  performance  1975, 71 t a g g e d  were i n t r o d u c e d t o a group  movements o f t h e f i s h  home  small.  The d a t a from  i n the  to  experiment  site.  behaviour  that  showed On  were  t o the capture s i t e ,  homing  areas, i t appears  whether  distance  in  homing.,  tidepool  fish.  than  16)  developed  second  more  move away from  return  successfully  site.  the s l i g h t l y  at the transplant while they  the  transplant  i s a more d e s i r a b l e h a b i t a t  (Table  the percentages  In  trapped at t h i s  imply  well  t h e p e r c e n t a g e s homing  first  found  not  Even when t h e  t h e t r a n s p l a n t a r e a were s m a l l . fish  i s  homing  at fish  I t was i n t e n d e d t h a t t h e  between t i d e p o o l s be  monitored  for  some  Table  Homing b e h a v i o u r  Bay Date  released  Number  released  Number homed P e r c e n t homed Number s t a y i n g i n transplant area Percent staying i n transplant area  Days s t a y i n g i n transplant area a. homers b. non-homers  16  along Grappler  Mud f l a t  2 October  Total 1975  Inlet  Bay  Mudflat  4 February  Total 1976  42  36  78  64  24  88  5 12  3 8  8 10  2 3  3 13  5 6  5  6  14  8  1,15 20,22, 54  78  period  of  time  and  became a p p a r e n t , examine homing  first  two  of  were  introduced release, and  to  and about  egual  conduct  rockfish,  23  Of  water.  adspersus (1971) subtidal ability  Green  tidepool  0. maSiJiosus. were  those  In the  located. which  fish  Most showed some t e n d e n c y after  was l o c a t e d , the r e l e a s e  the f i r s t  54 d a y s area.  experiments  t h e y were  seen  at  flavidus  (1975)  t o move f r o m t h e  two weeks o n l y one  after release  i n a pool  T h u s i t was n o t p o s s i b l e t o examine homing  (1972) f o u n d  ability.  that  the  yellowtail  ( a y r e s ) was u n s u c c e s s f u l i n homing  found  and  that  home  sites  the cunner  P o r t Renfrew showed t h a t  areas appeared o f 0.  maculosus.  after cleared  included  Tautoqolabrus  (Walbaum) was u n a b l e t o home o v e r deep w a t e r .  work  The  inlets  and H a i g h t  Se b a s t e s  fish  natural  of p o o l s i n t o  t h e a r e a between t h e d i s p l a c e m e n t  deep  to  storms o c c u r r e d  p e r c e n t a g e s were i n t r o d u c e d i n t o  any t r a n s p l a n t  Carlson  both  tagged  pools.  pool t o other pools.  60 m away from  severe  in  and  release  adjacent  Homing b e h a v i o u r a c r o s s  if  decrease  f o u n d i n t h e group  t a g g e d 0. m a c u l o s u s about  were; t o be c o n d u c t e d  introduction,  after  uncleared pools.  release  the  0. m a c u l o s u s  weeks  majority  of pools  behaviour.  i n a noticeable  populations  t o a p o o l o r group  t r a n s p l a n t experiments  Following resulting  i f fidelity  rough  t o have a d v e r s e e f f e c t s  terrain on  the  Khoo's o r deep homing  79  To  d e t e r m i n e w h e t h e r deep w a t e r h a s a d v e r s e  the  homing a b i l i t y  on  t h e homing a b i l i t y  the  same t i m e a s t h e homing  were  conducted.  home i s n o t  i s  fish,  well a  two e x p e r i m e n t s were c o n d u c t e d  o f 0. m a c u l o s u s a c r o s s G r a p p l e r experiments  Although  as  apparently  of inlet  e f f e c t s on  the ability  developed  small  as  along  percentage  Grappler  of inlet  i n  are  Inlet  0. m a c u l o s u s t o  tidepool  which  Inlet, at  fish,  able  there  to  home  successfully.  Methods  A total from on  t h e mudflat  the north  bay. the  The depth  Trapping site four  Inlet directly the inlet  of the i n l e t  month a f t e r  months a f t e r  release  conducted  bay  opposite  12 m  at  and  to a the  26 site  small  80 m a n d  high  tide.  were made a t t h e r e l e a s e  r e l e a s e and a t t h e c a p t u r e  sites for  r e l e a s e . .,  the from  second  experiment  t h e b a y on t h e s o u t h  site  Trapping  the  i s approximately  about  was c o n d u c t e d a n d o b s e r v a t i o n s  f o r one  inlet.  across  (56 f r o m  I n l e t ) were t r a n s p l a n t e d  s i d e of Grappler  i n t h e middle  transplanted  for  i n Grappler  distance  In  the  o f 72 0. m a c u l o s u s  directly  and  f o r one  f o u r months a f t e r  opposite  observations month a f t e r release.  at  HU  0. m a c u l o s u s  side of Grappler on t h e n o r t h the  release  were  Inlet to  side o f the site  r e l e a s e and a t t h e c a p t u r e  were site  80  Results  Table O.  maculosus  interesting  17  makes  are  unable  i t to  abundantly home  to note t h a t although  o f 0.  maculosus at the  fish  was  found  site  after  transplant  site  none  from  density  d e p e n d e n t e x c l u s i o n c a n n o t be  resjaect t o  Khoo were r a r e l y shown  that  Beach, e g u a l pool is to  found  restricted  t o one  the  as t o a g r o u p o f  u s u a l l y one show  t h a t 0.  of  of t h e i r fish  pools,  the  It  Since to  is  numbers  transplanted no  fish  releasing  fidelity  maculosus at pool.  normal  t h a t i n the  In  inlets,  to a particular  0.  Port  This  were fish,  Renfrew  study  movements a t  show f i d e l i t y t o  and  preferred pool.  little  water.  eliminated.  particular  course  proportions  inlet  exposure  (1971)  in  of  prior  that  considerable  release.  removed  3. Homing w i t h  deep  there are  release site,  at t h i s  the  over  clear  a  latter  First  particular group  maculosus  area,  has  there appear  comparable  to  a  g r o u p of t i d e p o o l s . With r e s p e c t First  Beach and  P o r t Renfrew  p e r c e n t a g e s homing little  evidence  t o homing b e h a v i o u r ,  and of  0.  0,  maculosus at  show moderate t o h i g h , maculosus  homing.  The  in  Grappler  work o f G r e e n  but  both  variable  Inlet (1967) a t  show Port  Table  Homing b e h a v i o u r  Bay  Date r e l e a s e d  Number r e l e a s e d  17  across Grappler  Mudflat  16 O c t o b e r  Total  1975  Inlet  Bay  4 Feb 76  56  26  72  44  Number homed  0  0  0  0  Number s t a y i n g a t transplant site  0  0  0  0  82  Renfrew O.  showed  that  the  vast  majority  of  transplanted  l a c u l o s u s l e f t the t r a n s p l a n t pool w i t h i n a few  In  a  nomber of experiments Green found that only f o u r f i s h  of over 100 t r a n s p l a n t e d for  any  period  of  weeks.  The  f i s h remained in  time:  f o l l o w i n g r e l e a s e , two latter  one  was  transplant  pool  three  at  the  taken with poison  subsequently  the  end  of the f i r s t  thus  the  day  about Khoo  four (1971)  fish  left  two  weeks.  At  the  the First  transplant  periods o f time, e i t h e r p r i o r t o homing or  to t a k i n g up r e s i d e n c e It  pool  all  Beach a s i z e a b l e percentage of f i s h remained i n f o r considerable  one  homed.  that  out  transplant  remained t h r e e weeks and  t r a p p i n g on a two-weekly b a s i s found  area  tidal cycles.  i n the  appears  area. that  there  may  be d i f f e r e n c e s i n  homing behaviour of 0.  maculosus i n areas of d i f f e r e n t exposures  (substrates) .  from F i r s t  the  Results  work of Green and  pools  and  In s h e l t e r e d i n l e t in any the  0.  maculosus d i s p l a y s l e s s  area.  few  fish  home and  conducted.  and  fidelity  shown  r e l a t i v e l y few  to  area. than  remain i n  Thus a s e r i e s o f experiments designed t o  compare hominq behaviour simultaneously areas was  Inlet  fewer f i s h remain i n the t r a n s p l a n t  areas much l e s s area f i d e l i t y i s  t i d e p o o l area,  transplant  Grappler  Khoo at Port Renfrew suggest t h a t i n more  exposed t i d e p o o l areas, particular  Beach and  i n exposed and  sheltered  83  Methods  Four e x p e r i m e n t s i n v o l v i n g nine differing  exposures  behaviour  of  as  were  conducted  0. m a c u l o s u s .  exposed,  moderately  location  and  retrapping  and o b s e r v a t i o n s  all  to  Each s i t e  exposed  observation.  or In  of f i s h  Port  Renfrew  Grappler  were  behaviour  (exposed), F i r s t  Inlet  transplanted  the  was i n i t i a l l y sheltered  each  homing  classified  according  experiment  to  releases,  were made s i m u l t a n e o u s l y  was  compared  Beach  in  (sheltered).  a  as f o l l o w s : Grappler  reciprocally  transplanted  First  Beach  transplanted apart;  total  Inlet  about  60  Renfrew  up t o f o u r  groups  apart.  Subsequent  286  and  fish  were fish  t h e s m a l l b a y and t h e  49  fish pools  and  trapping  o f about  were each  - two g r o u p s o f 44 f i s h  five was  reciprocally about  were  120  60  m  reciprocally pools  each,  conducted  and  were made a t two week i n t e r v a l s a s f a r a s p o s s i b l e ,  months.  Homing b e h a v i o u r was Benson  of  exposed),  a distance  between two g r o u p s o f t h r e e  m  observations  two  of  from  - two g r o u p s o f 50  between  between two g r o u p s o f f o u r  Port  transplanted  -  i n 0. m a c u l o s u s  (moderately  m u d f l a t on t h e s o u t h s i d e o f t h e i n l e t ,  of  compare  locations of  l o c a t i o n s as f a r as p o s s i b l e .  Homing  m;  different  Island  103 f i s h  compared  (exposed) and Ranee I s l a n d  were t r a n s p l a n t e d  as f o l l o w s :  in  0. m a c u l o s u s  (sheltered). Ranee  Island  from  a -  total two  84  groups  o f 17 and  34  fish  each  were i n t e r c h a n g e d between t h e  study s i t e s  i n the i n l e t ,  a b o u t 50  groups  26  were r e c i p r o c a l l y  two  of  fish  groups o f t h r e e  retrapping  and  each  p o o l s each about  observations  occasions f o r almost  Homing south eastern  two  were t r a n s p l a n t e d from  fish  apart.  the western 70  m;  between two  groups of  three  Subseguent  retrapping  on  times u n t i l  t h e e x p e r i m e n t was  was  exposed)  the l a r g e  and  o f 111  fish  a g r o u p o f 52 f i s h tidepool  to  (sheltered) .  to eastern  - two  pools  each,  about  Point  reciprocally  pools  60  another  about of  transplanted  m apart;  nine  fish  between two  about First  60  fish  %pstera 30  m  apart.  between  - one  compared  Beach  m  Beach  transplanted  were  34  of  s i x weeks.  as f o l l o w s :  Kirby Point  the  transplanted 50  t i d e p o o l on H a i n e s I s l a n d  approximately  and  groups o f  were  (exposed). F i r s t  Channel,  the  were c o n d u c t e d a number o f  terminated after  were d i s p l a c e d  on  A total  end o f t h e  Haines Island  were t r a n s p l a n t e d  Dodger  number o f  compared  Homing b e h a v i o u r and a r e a f i d e l i t y  total  a  Dodger C h a n n e l -  and o b s e r v i n g  from K i r b y  two  Subsequent  e a c h were a p p r o x i m a t e l y r e c i p r o c a l l y  0. m a c u l o s u s  -  between  (moderately exposed)  as f o l l o w s :  i n the c h a n n e l , about 24  ffl  maculosus  s h o r e o f Dodger C h a n n e l  were t r a n s p l a n t e d  transplanted  attempted  s i d e o f Haines I s l a n d  88  and  were  b e h a v i o u r o f O.  Island  bed  80  Island  months.  Haines fish  m a p a r t ; Benson  two  in  (moderately  (sheltered).  A  Haines I s l a n d  -  from -  32 two  the  large  fish  were  groups  of  g r o u p o f 18 f i s h  and  approximately  g r o u p s o f p o o l s a t two  reciprocally  l o c a t i o n s on  the  85  point.  In  both  Percentages calculated  cases  the  homing a n d r e m a i n i n g f o r a l l nine  In  addition,  a total  i n which  fish  were r e p l a c e d i n t o  they  were c a p t u r e d ; F i r s t  fish  were  a t the  displaced transplant  o f 123 f i s h  which  areas  were  the large t i d e p o o l B e a c h - 44 f i s h  t h e y were f i r s t  Haines I s l a n d  i n t h e area i n  retrapping  was c o n d u c t e d  t w i c e weekly  at a l l sites u n t i l  after  five  weeks.  nearby  p o o l s and i n d i s t a n t  the  Percentages  - 53 which  were r e p l a c e d i n t o t h e  c a p t u r e d ; K i r b y P o i n t - 26  were r e p l a c e d i n t o t h e p o o l s i n w h i c h t h e y  Subsequent  50 m.  were r e p l a c e d i n t h e  t h e y were c a p t u r e d , as f o l l o w s :  in  about  sites.  area  tidepools  fish  were  captured.  and o b s e r v a t i o n s were made experiment  found  was  terminted  i n t h e home a n d home a n d  p o o l s were c a l c u l a t e d  f o r each  site.  Results  The 18)  show  r e s u l t s o f the four  that  in  spite  o f t h e low p e r c e n t a g e s  r e l e a s e i n some a r e a s , i n e a c h homing a r e g r e a t e r i n a r e a s (tidepool  areas)  no  than i n s h e l t e r e d areas  of v a r i a b i l i t y  relationship  determined  by  moderate . t o  greater  (inlets). areas  location  this and  variability  exposure  However t h e  show  observation.  a t the transplant  sites  and  after  successfully  a  greater  t h a n a n t i c i p a t e d , and t h e r e appears  between  0. m a c u l o s u s r e m a i n i n g  tidepool  (Table  located  case the percentages  of  p e r c e n t a g e s homing i n d i f f e r e n t degree  transplant experiments  exposure  t o be as  The p e r c e n t a g e s o f a r e v a r i a b l e , but  Table  Honing  P.R.  Date  5 Nov  released  F.E. G.I.  2 Nov  behaviour  i n areas  B.I.  4 June  released  Number homed P e r c e n t homed  Number s t a y i n g in transplant area Percent staying in transplant area  88  98  18 20  35 36  0  2  0  2  Days s t a y i n g i n transplant area a. homers b. non-homers  Percent of homers returning to home p o o l  P.P.:  15,26  42  74  100  of d i f f e r e n t  P.I.  2  1976  1975 Number  18  exposure  H.I.i  21  June  October  1976  52  54  34  K.P.  F.B.  H.I.z  Aug  9 Auq 1977  7 Aug  8  27  32  26  7  19 59  33  11  13  63  41  41  27 50  23,39, 90,94  D.C.  1x21, 28x9, 53x3  11,11 39  100  70  1,3,3,3, 3,3,10,22, 22,23,36 100  52  3,3,3,3 1,3,3,3, 11,11,18 18,18 68  R e n f r e w , F.B.: F i r s t B e a c h , G . 1 . : G r a p p l e r I n l e t , B . I . : B e n s o n I s l a n d , R.I.: Ranee I s l a n d , H.I. : H a i n e s I s l a n d (south e a s t side), D o d g e r C h a n n e l , K.P.: K i r b y P o i n t , H. I. : H a i n e s i s l a n d (large tidepool)  Eort  1  D.C:  2  co  87  do  not  show  transplant and  increasing  site  exposed  channels are the p r e d i c t e d In  the f i r s t  located  after  severe  storms  At b o t h  First  of  in less  numbers  0.  found  which Beach  maculosus  areas.  and  Renfrew  evident  (as measured  the  in inlets  the small percentage  be t h e r e s u l t  Port  Only  at  findings confirmed. of  fish  of a s e r i e s of extremely to  a reduction  following  transplanting. in  the storms.  r e l a t i o n s h i p between homing  state conditions  remaining  o c c u r r e d j u s t subsequent  was  an i n v e r s e  fish  tidepool  experiment  r e l e a s e may  of  the  number  Khoo  (1971)  performance  and  by p e r c e n t o b s e r v a t i o n s o f  sea  swells  g r e a t e r than or equal t o s i x f e e t ) . The produced  second  very  experiment  unexpected  results.  r e l e a s e t h e r e were numerous 0. Ranee  Island.  population  of  disappeared armatus.  release  0. m a c u l o s u s  in  to  be  At t h e t i m e  of  the  by  in  area  large  Beach  diet  and  reflection  of  Port o f 0.  Renfrew  maculosus  inlet  tagged  and  around  fish,  almost  the  completely  numbers o f L e p t o c o t t u s maculosus  when compared (Ruth,  of t r a p p i n g  with  1976).  a t Ranee  from fish  this from  The a p p a r e n t l y  Island  may  be  a  this.  Although Dodger C h a n n e l ) there  diversity  Ranee I s l a n d )  the  the  A n a l y s i s o f s t o m a c h c o n t e n t s of 0.  unstable population  when  replaced  I s l a n d and  maculosus  Following  a r e a showed a low First  (Benson  is  was a  the  third  conducted high  experiment f o r only  (Haines  Island  s i x weeks and a t  freguency o f storms, the  a  and time  percentage o f  88  O, maculpsus at Haines I s l a n d which had the  end  of  this  period  was  successfully  considerably  percentage homing i n Dodger Channel.  greater  end o f the e e l grass bed and  trapping  p r i o r t o the experiment  T h i s and the release  this low  site  percentage  homing  suggest  seen  intensive  produced  that  by  than the  No 0. maculosus were  or trapped a t the western of  homed  none.  while  the  s i t e was not f e a s i b l e as an a l t e r n a t i v e h a b i t a t f o r the  t r a n s p l a n t e d f i s h , homing behaviour i s not s t r o n g l y expressed i n 0. maculosus found i n i n l e t s . The Haines  fourth  Island)  experiment  first  that  the  majority  almost simultaneous  twice  time,  of  month f o l l o w i n q r e l e a s e .  possible  data  First  Beach,  although  from  First  this Beach  i n f o r m a t i o n i s acquired i n the S i n c e the weather was f a v o u r a b l e  weekly  visits  to  f o r the d u r a t i o n o f the experiment.  t h e other experiments, the data suggest (that  Point,  was t e r m i n a t e d a f t e r f i v e weeks.  i s a r e l a t i v e l y s h o r t p e r i o d of suggest  (Kirby  that  each  site  were  In aqreement with homing  i s , r e t u r n i n g to the area of capture) i s b e t t e r  behaviour developed  or a t l e a s t expressed, i n t i d e p o o l than i n l e t a r e a s , but f a i l to show that a g r a d i e n t of a b i l i t y  i n homing behaviour e x i s t s which  corresponds to a v i s u a l l y determined exposure g r a d i e n t . From t h e r e s u l t s Kirby P o i n t , F i r s t (Table  19)  it  of  the  replacement  experiment  Beach and t h e l a r g e t i d e p o o l on Haines I s l a n d  i s e v i d e n t that the percentage of f i s h found i n  the home p o o l only i s much g r e a t e r at Kirby P o i n t than a t Beach,  where  at  the  usual  First  s i t u a t i o n of equal percentaqes i n the  Table  Area  fidelity  i n sites  19  of different  K.P.  Area r e l e a s e d Date r e l e a s e d  exposures  F.B.  8 Aug 77 9 Aug 77  H.I. 7 Aug 77  Number r e l e a s e d  26  aa  53  Number pool Percent pool  f o u n d i n home only f o u n d i n home only  23  17  1  88  39  2  Number f o u n d i n home and n e a r b y p o o l s P e r c e n t f o u n d i n home and n e a r b y p o o l s  2  18  8  a 1  Number f o u n d i n home p o o l and home and n e a r b y p o o l s P e r c e n t f o u n d i n home p o o l and home and n e a r b y p o o l s  Number f o u n d i n d i s t a n t pools Percent found i n d i s t a n t pools  K.P.: K i r b y P o i n t , F. B.: F i r s t  25  35  96  80  0  2  0  0  5  0  Beach, H. I . : H a i n e s I s l a n d  90  home and n e a r b y fidelity  data suggest that  location  homing  and  (turbulence) distribution  of  has  and a c t i v i t y  in  my p e r c e p t i o n  0. m a c u l o s u s .  been  consideration of this  shore)  anticipated,  observation, bears l i t t l e  behaviour  terrain,  As  terms  of  of  the  relevant  exposure.  These  than factors  an  as  steepness  The  action  important  i n the  1971a,b,c),  (the roughness  ofthe  and i r r e g u l a r i t y  ofthe  problem  subjective  are considered  of  and t o p o g r a p h i c a l  measuring  the  o f wave a c t i o n  been  a t t a c k e d i n a number o f ways.  eight  biologically  based  on t h e abundance o f common s p e c i e s ,  sheltered  to  extremely  exposed.  of  section.  r e l a t i v e exposure i n  Ballantine  localities  categories,  essentially  r a n g i n g from  not  has  (1961) s u g g e s t e d  While b i o l o g i c a l  i n a d e s c r i p t i v e sense they a r e  may  regularity  or turbulence i n d i f f e r e n t  determined exposure  impression  i n t h e next  terms  easily  extremely scales are  generalized  wider a r e a s . Determinations  wind  based  wave  (Green,  factor  overall  4.Measurement o f wave a c t i o n  over  of exposure,  relationship tothe  0. m a c u l o s u s  and a r e l a t e d  Island.  Since  demonstrated  area  w i t h r e s p e c t t o t h e homing b e h a v i o u r o f 0. m a c u l o s u s ,  be more  useful  little  was shown t o t h e l a r g e t i d e p o o l on H a i n e s The  on  p o o l s was f o u n d .  of  measurements have been  exposure  or  u s e d by Moore  wave a c t i o n b a s e d on (1935)  and  Southward  91  (195 3 ) , u s i n g t h e the  exposed  percentage  o f winds o v e r  aperture of a given  winds e g u a l t o or g r e a t e r than least The and  one  locality  10 k n o t s  f o o t over the p r e d i c t e d  former  tidal  does not  take  into  into  and t h e p e r c e n t a g e  which  method d o e s n o t t a k e a c c o u n t  the l a t t e r  10 0 d a y s b l o w i n g  produced  height, o f wind  account  of  a wash a t  respectively. f o r c e or s w e l l s  the  slope  of  the  locality.  Green and  Green,  (1967) u s e d  1970),  a submersion-emersion  b o l t e d t o t h e s u b s t r a t e a t a known  h e i g h t and  a t t a c h e d t o a r e c o r d e r which r e c o r d e d  emergence  and  times with  predicted  pools two  submergence  into four  such  of t i d e p o o l s .  the  sensors are r e g u i r e d Demetropoulos  exposed  t o be  Jones  and  spring  dynamometer w h i c h r e c o r d s t h e No  Harger  wave i m p a c t  (1970) measured  of a steel  to sheltered.  At  least  simultaneously.  a drogue a t t a c h e d to a drag  o v e r one held  (1968) i n which P l a s t e r o f P a r i s b a l l s a r e  compared  i n t h e s u b s t r a t e and  t o weight  While  these  methods may  do n o t p r o v i d e a s i m p l e of widely spaced  loss figures  be  long  period  was  by  with a  described  loss of  the  velocities.  u s e f u l i n some s t u d i e s  period  of  C  to  the weight  exposure  the  attached  a t known w a t e r  means o f m e a s u r i n g  areas over a  exposure  by  i s provided.  in position  by Huus  balls  s h o r t term  tidal  A method  r o d s embedded  produced  of freguency  clip.  steel  f o r measuring  actual rate  maximum  plate  the  of  to  positioned  measurement  vertical  him  (1968) d e v e l o p e d  waves i n t h e l o c a l i t y . ,  (Druehl  times  Comparing  t i m e s o f submergence e n a b l e d  c a t e g o r i e s from  movement down a n a i l  sensor  in a  time.  they  variety In  this  92  study  a  the f i s h the  method  was r e g u i r e d w h i c h m e a s u r e d t h e e x p o s u r e where  a r e l o c a t e d over  method  used  was b a s e d on t h e w e i g h t  attached t o the substrate move  in  a  a r e l a t i v e l y extensive  period.  Thus  l o s s o f cement  blocks  ( i n p o o l s a n d on r o c k s )  able  to  h e m i s p h e r e a b o u t t h e a n c h o r a n d u n d e r g o a b r a s i o n by  t h e a c t i o n o f waves moving t h e b l o c k over Initially, approximately  a  two  pilot  months.  study  the substrate.  was  However,  conducted  Subseguently,  a l o n g e r term s t u d y ,  months was c o n d u c t e d .  lasting  the weakness o f t h e p i n s  used r e s u l t e d i n a h i g h l o s s o f cement b l o c k s areas.  but  in  more  exposed  lasting  almost  nine  Only t h e r e s u l t s o f t h e f i n a l  study  are  reported. The r e l a t i v e r o u g h n e s s o f t h e t e r r a i n was e v a l u a t e d by a  subjective estimate  The  steepness  irregularity  of  b a s e d on o b s e r v a t i o n s o f t h e s t u d y  the  of these  rocky  shelves  and  the  s h e l v e s were u s e d i n t h i s  areas.  degree  of  estimate.  Methods  Styrofoam high.  cups  The c u p s were t h e n  were  cut  greased  down t o make m o u l d s 4.5 cm  with  vaseline  and  eyebolts  6.35 cm l o n g w e r e i n s e r t e d i n t h e b o t t o m o f t h e c u p s o t h a t of  the  eye  was  inside  P o r t l a n d CSA Type 20  the  cement  cup.  in  the  Concrete following  cement : s a n d : w a t e r - 5 : 1 : 3.  This mixture  the  The s t y r o f o a m  cups  and  allowed  to  set.  was m i x e d  part using  proportions was p o u r e d  into  c u p s were  then  93  peeled  off leaving a concrete  (Figure  14a).  The  The  blocks  substrate at nine Cape  Beale,  blocks  Port  on  the  tidepool  each s i t e hydraulic the  used  a  pin  cement.  To  eyebolt in the calm  as  the  (Figure field,  and  14b).  constant  immersion  All 30 and  August and 28  Say,  washed and for  scrubbed the  with  attached  grant.  Vancouver  the  Island:  Point,  First  problem  the  were a n c h o r e d  eyebolt  pilot  to  Ranee I s l a n d and  blocks  an  and  of  in  t h e weakness o f  study,  halibut  I n a d d i t i o n t o t h e cement b l o c k s  anchored  e a c h were a n c h o r e d of  to  a tank used numbers  fish  i n an  provided  a  bricks  i n an  of  "exposed"  and  of  in  the  experimemt  to  could  be  cirri  measure  the b l o c k s  and  snaps  anchored  on  cement b l o c k  embedded  at  tank  as  opposed  the  effect  of  the r e l a t i v e exposure  of  t a n k r e c e i v i n g waves.  the 12  cement b l o c k s September,  1977.,  scrubbed  48 h o u r s .  to  eyebolt  the  greater  This  Ten  used i n t h e  p r o d u c e d on  s i d e of the  of  (bay),  overcome the  sides  whether  t o a calm tank.  the  attached  ten b l o c k s  experimentally  Inlet  p i n between t h e  "exposed"  determine  nearest  tidepools  west c o a s t  Haines I s l a n d .  shower c u r t a i n c l i p s  were  in  inserted  R e n f r e w , B e n s o n I s l a n d , Pachena  on  using  an  to the  anchored  Beach, Helby I s l a n d , G r a p p l e r large  with  were weighed  were  sites  block  1976  After  with  were p u t  steel  wool.  animals,  position  the  w o o l , and dry,  b l o c k s were  then  they  oven  as  21  were a g a i n  S p i r o r b i s s p . , The  in  May  uniformly  d r i e d at  T h i s procedure succeeded such  between  were removed between  removal  Keeping the blocks steel  and  into  55°C  uniformly removing  blocks  were  5.8  cm  -+—4.5 c m — * • Figure  14a  Cement  block  Figure  14b  Cement  block  design  anchored  i n pool  95  weighed t o the nearest gram. The ranks, the  Kruksal-Wallis  corrected  for  one-way  ties  there  were  any  of  ( S i e g e l , 1956), was  i n i t i a l weights o f the blocks  whether  analysis  at  each  significant  variance  by  used to compare  site  to  differences  determine between the  samples when they were i n s t a l l e d .  Results  The ranks,  K r u k s a l - W a l l i s one-way  corrected  for  ties,  samples (Table 20)  showed  samples  p>.05).  (H=.958,  on  no  analysis  the  initial  significant Weight  of  variance  weights f o r a l l  differences  loss  (mean  between  grams and  ranks) and the number of cement b l o c k s remaining are a l s o in  Table  20.  Examination  of  the  c l u s t e r s of ranks: Cape Beale and Pachena  Point  Grappler  I n l e t and  tank.  Since  and  only  First  Ranee  the wave tank; Haines  Port  shown  Renfrew,  I s l a n d , Helby I s l a n d , Island  and  a r e l a t i v e exposure r a t i n g was were  mean  ranks shows s e v e r a l  Benson I s l a n d ;  Beach;  s i n c e the i n d i v i d u a l sample s i z e s several  mean  by  small,  the  calm  reguired  especially  and for  groups, i t appeared t h a t c o n s i d e r i n g the samples i n the  groups d e l i n e a t e d above was  the most u s e f u l approach.  Dsing the m u l t i p l e  on  rank  Wolfe, 1973;  Dunn,  1964), the f o u r groups of samples were compared with each  other.  sums,  and  corrected for t i e s  comparisons  test  (Hollander and  Dunn suggests using a r e l a t i v e l y  based  high s i g n i f i c a n c e  level  (such  T a b l e 20  Final Weight  study o f r e l a t i v e statistics  Initial mean wt (grams)  Cape  Beale  Benson Port  Island  Renfrew  Pachena  Point  exposure  o f cement  Initial mean wt (ranks)  N  blocks  Mean wt Mean wt loss loss (grams) ( r a n k s )  N  175.9  56.1  10  32.0  80.3  2  174.6  47.2  10  53.8  75  41  4  176.2  56.7  10  21.3  59.6  7  175.6  54.6  10  22.1  57.0  10  First  Beach  176.1  56.4  10  29.0  63.4  7  Helby  Island  176.0  57.0  10  17.0  38.8  6  Ranee  Island  175.9  56.4  10  19.2  49.9  9  53.7  10  17.7  39.3  10  175.7  56.3  10  12.7  17.3  10  Wave Tank  176.0  57.0  10  17.1  38.3  10  Calm  176.4  59.4  10  12.2  14.0  10  Grappler Haines  I n l e t. 175.6  Island  Tank  97  as  0.20)  i n such m u l t i p l e comparison t e s t s , s i n c e t a k i n g a more  traditional level substantial  of  0.05  for  differences  example,  exceedingly  T h i s r e s u l t e d i n the f o l l o w i n g of  decreasing  blocks remaining i n  Cape Beale,  exposure  establishing  difficult".  comparisons were made u s i n g a value of of  order  "makes  Hence,  0.20. ranking  of  sites  in  (with the mean number of cement  parentheses):  Benson I s l a n d  (3)  Port Eenfrew, Pachena P o i n t , F i r s t  Beach  Ranee I s l a n d , Helby I s l a n d , Grappler  (8)  I n l e t , Wave Tank  (8.75)  Haines I s l a n d , Calm Tank ( 1 0 ) .  The  f o u r groups were a l l s i g n i f i c a n t l y  each  other  Pachena  except  Point, is  remaining at  these  indication  of  Cape Beale, Benson I s l a n d and  First  significance  different  Beach.,  primarily sites  relative  However,  due and  to  since  (p<.20)  Port Renfrew, the  this  in  exposure, Cape Beale and  itself  of  blocks is  an  Benson I s l a n d  are r a t e d as more exposed than Port Renfrew, Pachena First  lack  the s m a l l number of  since  from  Point  and  Beach. The  ranking  of  sites  according  to  the  relative  roughness of the t e r r a i n r e s u l t e d i n a somewhat d i f f e r e n t  order.  98  The  sites  a t Cape B e a l e ,  extremely  irregular,  sloping  s h e l v e s and  wider,  less  of Haines  Port  Renfrew and  Inlet  and  irregular Haines of  small  location and  Port  regularity site  which  rougher  the by  of  within  the  location  irregularity these  two  area  Pachena  observed  suggested  groups  factors  on  homing  tidepool  the  suggests  that  observation  and  because  In  there  were  the  of  the  study  the  a  following sites  and  initially  cement b l o c k r a t i n g s , by  a  which has  the  classified  an i n d i c a t i o n  behaviour.  <for  o v e r a l l exposure to  Beach).  divided  location  exposed  more s h e l t e r e d b u t  the  on  results  which by  may,  exposure,  estimates to provide  Grappler  with  most  and  by  further  by  Point)  have a s i m i l a r  homing b e h a v i o u r  shelves at  topographically  terrain  the  the  southeast  together  an a r e a  c l a s s e d among  appears  flat  large  simply  ( f o r example, F i r s t  groups  these  be  terrain^  homing b e h a v i o u r  into  the  the  C o n s i d e r a t i o n of the  somewhat m i s l e a d i n g .  Renfrew,  terrain  analysis the  of  of  are  with  areas,  more  o f roughness o f the  t h e e x p o s u r e o f an be  be  e v a l u a t i o n o f wave a c t i o n  o b s e r v a t i o n might  example,  flats  Dodger C h a n n e l .  estimates  may  Beach and  sheltered to  Point  compared  with the broad,  appear  t h e sand/mud  t h e cement b l o c k  evaluating  again  Kirby  narrow, s o m e t i m e s s t e e p l y  shelves at F i r s t  Island  I s l a n d and  and  channels,  Pachena P o i n t , , In  Ranee  subjective  surge  I s l a n d , and  than  Island  with r e l a t i v e l y  perilous  side  Benson  and  topographical of the e f f e c t  of  99  5»Discussion  From turbulence  Table  i t i s  evident  and i n c r e a s i n g r e g u l a r i t y  which f i d e l i t y from  21  one  of the terrain^  i s shown i n t i d e p o o l  pool,  t h a t with  areas,  apparently  rough  areas  turbulent, in  display  rough  p r e c i s i o n than return other  to  rough  steep  initial  the  o f homing t h a n  poor  few f i s h  (that  pool o f capture,  which  i n less do  weather  and a b r u p t l y The  and  few  fish  extend  kind  of  greater  percentages  shelves homing  home.  With  of t h e t e r r a i n , area.  Khoo's  and r o u g h t e r r a i n , sloping  home  t h e home p o o l , than t o  stay i n the transplant  confirm  In  turbulent  fish  i s , greater  and i n c r e a s i n g r e g u l a r i t y  results  ability.  apparently  areas  numbers o f f i s h  that  in  a r e a s r e t u r n t o t h e home r a n g e w i t h  turbulence  trenches  homing  evidence  However, t h o s e  i n other  These findings  size  f i d e l i t y i s shown  When d i s p l a c e d , f i s h  i n t h e home r a n g e ) i n i n l e t s  decreasing decreasing  less  areas.  the  pools  in  t o one p r e f e r r e d p o o l t o a g r o u p o f p o o l s .  l a r g e r areas.  turbulent  the area to  increases  more s h e l t e r e d t o p o g r a p h i c a l l y r e g u l a r i n l e t s , to  decreasing  (1971)  i n t h e form o f  adversely  behaviour  affect  exhibited  d e p e n d s on t h e d e g r e e o f wave a c t i o n and r o u g h n e s s o f  terrain. Turbulence  vertical  h a s been shown t o p l a y a major r o l e  distribution,  high  tide distribution  i n the  and a c t i v i t y o f  Table  Honing  behaviour  according  21  to t o p o g r a p h i c a l  regularity  and  turbulence  TIDEFCOIS  high  Turbulence  T o p c g r a phy  Area  highly  fidelity  Honing Percent  moderate  irregular  particular  pool  relatively  low  irregular  u s u a l l y one preferred pool  low  relatively regular  very  regular  group o f pools  large  area?  low  extremely regular  large  area?  behaviour homing  Percent remaining transplant area Percent homing  INLETS  of to  in  homers home p o o l  moderate  to high  high  moderate lew  to high  t c moderate  moderate  to high  moderate very  moderate  to high low  to  low  very  low  low  very  low  high  Ho o  1  101  0.  maculosus  0.  maculosus  in  (Green, tends  sheltered  Seasonal  1971a,b,c).  t o be  areas,  i t  occurs  and  out  pool or departs  (Green,  winter  intertidal (Gibson,  fish,  the  5.5  tide,  pool  cm);  isolated  (in  the  pool i s i n J u l y , year  a t which  (Green,  200C  water  to  (Green, being 0.  shown  in  11°C 1971c).  active  Q.  for  fish  another  (Euphrasen)  In  moderately  proportion  the  (1971a).  at  in  the  more  Reduced  areas,  fish  or  above  leave  months o f  15°  activity  to  tidal  temperature  from  w i t h i n one  hour  the  from  occurred  exposed  retreat C,  during  high t i d e  pool i s  exposed of  warmest p a r t s o f a p o o l  inactive  the  less  are also the hottest  temperatures  t h e change  almost  maculosus  September, t h e c a l m e s t these  areas  i s an  until  been shown t o r e d u c e t h e  inhabiting  pools  there  of O.  maculosus remain at  sheltered  they seek cover  Although  Green found to  in  areas  distribution  A u g u s t and  pools  in  maculosus  a significant  and  i n J u n e has  maculosus  similar  been  L=Eno£hrj;sJ b u b a l i s  1971b,c).  1971b).  months o f t h e y e a r ,  flooding  i s washed  i s greatest, a decline in  sheltered  areas  (Green,  only  deeper  induced  1971a; p e r s o n a l o b s e r v a t i o n ) .  also  0.  i n the  i n exposed  again time  only some  shift  the  the  when s t o r m  1967).  complete onshore thsn  has  Acanthocottus  At h i g h leave  This  intertidal.  the r e s i d e n t population  months, when t u r b u l e n c e  numbers i s e v i d e n t .  the  are evident  of p o o l s o c c u r s  transects  t i d e p o o l s , whereas  throughout  disruption  In  exposed  confined t o higher  changes i n d i s t r i b u t i o n  of the  In  of  fall first  in  the  two  otherwise  during  turbulent  102  p e r i o d s has a l s o been r e p o r t e d  in Clinocottis analis  (Williams,  1957} . There of  wave  action  0. m a c u l o s u s , to  a p p e a r s t o be a r e l a t i o n s h i p and  from  continual  tide. high  movement  and  fidelity  home, a l t h o u g h  those  out  of  the  turbulent,  home  less  in  display  transplanted,  inlets,  few f i s h pool  these  terrain;  areas area  in  in  areas,  are  appear  more  is  movement  appears  their  fidelity  topographical occurrence  of  pools  to  possible  and  follow the  When  at  depend areas  they  o f the t e r r a i n .  rarely  move  In l e s s  Where  on  the fish  the  between p o o l s i s  hazardous.  the  tide  When  roughness  fish i n the  of  the  leave the t r a n s p l a n t i s never g r e a t  and  a l l t i m e s , a much g r e a t e r amount o f place,  success.  irregularities fidelity  less  fish  i n the  o f movement a t h i g h  where t u r b u l e n c e  take  homing  precision  n a t u r a l movements.  t h e degree  regular  to  possibly  d i s p l a c e d , t h e numbers r e m a i n i n g  In sheltered i n l e t s ,  movement  and  fidelity.  r e t u r n , presumably because pool  of  leave the pool a t  which do home show g r e a t e r  rough  movement  i s c h a n g e d t o t h e new p o o l and few  and movement between  transplant  pool.  areas  i s more r e g u l a r , p r e s u m a b l y t u r b u l e n c e  reduced from  tide  a s the f i s h  strict  a p p e a r s t o depend on t h e r e g u l a r i t y terrain  high  tidepool  t u r b u l e n t areas,  hence  area t o which they  of  none i n an e x p o s e d t i d e p o o l a r e a s  i n sheltered  T h u s i n rough tide  amount  virtually  an o n s h o r e s h i f t  almost  the  between t h e amount  also  resulting The  in  absence  appears  to  little of  area  distinct  reduce  the  t o a s m a l l area comparable t o t h a t o f a  10 3  group of t i d e p o o l s . To  my k n o w l e d g e , t h e r e i s  comparing  home  range  {Blennius  johplis  bubalis)  in  fidelity  (Linnaeus)  two  suggested  as of  the  two  frequency  of  return  observed,  is  irregularly  producing  gullies,  to  areas;  the  divided  by  (Gibson,  between  tidepool fishes and  fairly  (Hubbs,  (Ayres)  bubalis,  B l e n n i u s £holis  (Jordan)  (Stephens small  (Williams,  and  (Sale,  1971), H o l a c a n t h u s  shore Haight,  rocky  Epinephelus  coral  which  a  being  vertical  areas, home  absent  rock  (Gibson,  edges rocky  comparing  home  reported f o r  au r o r a  (Jordan  analis.  Girella  Acanthocottus  L= E n o p h r y s l  1967), H y p s o b l e n n i u s  D a s c y l l u s aruanus  b e r m u d e n s i s Goode restricted  Tautoqolabrus  was  undivided  ranges  clinocottus  1957),  reef f i s h  higher  c o m p a r i s o n s c a n be  =  reef fish  but f a i r l y  and c o r a l  1972),  fih.olis)  qilberti  e t a l . , 1970), C l i n o c o t t u s q l p b i c e p s  1973)  somewhat l a r g e r  areas.  where  f- ili£E2E§trusJ  1921),  nigricans  1961)  1967).  limited  Am p h i g on op t e r u s  Gilbert)  area  t h e r e a r e few p u b l i s h e d d a t a  the  species  the topographical  home p o o l a f t e r  r a n g e i n t h e same s p e c i e s i n d i f f e r e n t drawn  to  i n contrast to the flatter,  at t h e other s i t e Although  (Blennia§ due  study  f =Enophrys 1  (Ballantine,  being  the  published  Acanthocottus  i n one s p e c i e s  dissimilarity  shore  and  possibly  one  i n t h e same i n t e r t i d a l  semi-exposed  D i f f e r e n c e s were n o t e d were  only  (Bardach,  home r a n g e s  Sebastes  s t r i a t u s •• ( B l o c h ) , E. g u t t § t u s  (Linnaeus) 1958) , t h e  shown by  flavidus  adspersus  (Green,  near  ( C a r l s o n and  (Green,  1975),  (Linnaeus)  (Bardach,  104  1958)  and  the  fidelity (Berra, Cottus  shown  by  bairdi  (Linnaeus)  (Girard)  Henidia  that  t h e r e are reefs, fairly can the  fish  area,  restricted  to  be  of  the  any  of  the  fidelity  species  globiceps  fish  L=Eng£hr;ysJ GiEglla to  al.,  in  (Green,  1970),  is liable  nigricans (Williams,  fishes  show  substrate example,  t o be  a p p e a r s t o be there  different  returns  mustela 1957), t o  analis similar  1973)  Blennius  bubalis, Ciliata  home i n C l i n o c o t t u s  marine  the  to  where pools, of  turbulence swept o u t  of  shown. do  not  appear behaviour  areas.  Different  h a v e been shown t o d i s p l a y v a r y i n g  (Beebe, 1931), m o d e r a t e t o low et  1975).  I n a d d i t i o n , where  fish  and  e t c . show e v i d e n c e  i n homing p e r f o r m a n c e , f r o m h i g h  (Stephens  creeks  s t u d i e s appears  for  patches  mega l o t i s  r e p o r t s o f d i f f e r e n c e s i n homing  intertidal  Clinocottus  1952) ,  Fundulns h e t e r o c l i t u s  with  t o homing b e h a v i o u r ,  published  of  boulder  (Bailey,  in t i d a l  these  associated  Macleay  L§£2ffiis  #  area  bongbong  Lotrich,  irregularities,  area  same i n t e r t i d a l  species success  the r e s u l t s  and  With r e g a r d  (Linnaeus),  fidelity.  considerable  and  1960;  closely  area  1972)  developed  (Gill)  1964)  Brattstrom,  boats,  restricted  be  (McCleave,  topographical  wrecked  Galaxias  punctulatus  menidia  of  poorly  fish  Gunning,  ( B u t t n e r and  Comparison suggest  bairdi  ( B e r r a and  marshes  comparatively  freshwater  1973), C o t t u s  (Rafinesque) salt  apparently  percentage  and  returns  Bathy g o b i u s  soporator  in Hypsoblennius p. h o i i s , (Linnaeus)  variability  (Gibson,  1957). from  gilberti  ftcanthocottus  almost complete  (Williams,  in  1967),  inability  Hear  relatively  shore high  105  percentage r e t u r n s i n  Tautogolabrus  moderate  Sebastes  returns  1972), low t o E. g u t t a t u s  in  moderate (Bardach,  considerable  flayidus  returns 1958)  adsgersus  in  and  distance  in  general,  freshwater  (Green,  (Carlson  EjDine^helus  inability  Holacanthus  1975),  and Haight, str^atus  to  and  r e t u r n from a  berjrudensis  (Bardach,  1958) . In  f i s h and f i s h  show much l e s s success i n homing than this  has  been  1973;  Gerking,  examined  fishes  in  creeks which  ( L o t r i c h , 1975; McCleave, 1964; B e r r a ,  1959; Gunning, 1959)..  While t u r b u l e n c e , topography evidently  marine  in tidal  appear  to  have  (and d i s t a n c e  displaced)  a general e f f e c t on homing  success,  other f a c t o r s such as changes i n area f i d e l i t y over t h e l i f e the  fish  and  significant.  existence  and  size  of  of  home range may a l s o be  106  V.  MORPHOLOGICAL  DIFFERENCES  BETWEEN FISH IN DIFFERENT  LOCATIONS  1.Port Renfrew, F i r s t  The  early  homing b e h a v i o u r prompted the  Beach, G r a p p l e r  suggestion  further  investigation  behaviour  in  examination o f m e r i s t i c populations  was  differences  between  present,  there  were  differences  o f 0. m a c u l o s u s i n a r e a s o f d i f f e r e n t  same t i m e a s t h e f i r s t  homing  that  Inlet  of behavioural  behavioural different  and  exposures  differences.  experiment locations  morphometric  in  At  investigating was  conducted,  characters  in  these  u n d e r t a k e n t o d e t e r m i n e w h e t h e r t h e r e were any  might  the  populations..  conceivably  be  Such  differences,  related  to  i f  behavioural  differences.  Methods  To total the  of  examine m e r i s t i c  304  Beach 1975).  in  collection  Inlet  Grappler  (2 November The  morphometric  characters,  0. m a c u l o s u s were c o l l e c t e d a s f o l l o w s :  bay i n G r a p p l e r  mudflat  and  fish  Inlet  (29 (1  October,  1975),  136 from from  November, 1 9 7 5 ) , 59 from  1975) and 56 from P o r t  Renfrew  were f i x e d i n 10% f o r m a l i n  and l a t e r t r a n s f e r r e d  53  (5  the First  November,  immediately  t o 37.5% i s o p r o p y l  a  after  alcohol,,  107  The conversion  fish  to  were  total  measured  length  (standard  distance with and  distance,  distance  all  d i a l c a l i p e r s to the nearest  standard  length.,  anterior  percentages spines  of  and r a y s  Cirri nature around  (single, eye,  operculum  and  Interorbital  and p o s t e r i o r n a r e s head  length.  and a n a l  head d e p t h ,  between  between p o s t e r i o r n a r e s .  e y e l e n g t h were e x p r e s s e d  head  and b e h i n d  anterior  0.005 cm.  for  as  nares  and  were  made length  percentages  of  a n d d i s t a n c e between  expressed  Counts  length,  Head d e p t h ,  examined distance  were  eye  measurements  and  examined  as  were made o f d o r s a l f i n  f i n rays.  were a l s o e x a m i n e d w i t h bifid,  -  s e e a p p e n d i x 2) and s e x e d , a n d s i x  c h a r a c t e r s were m e a s u r e d : head l e n g t h , interorbital  length  triple  respect  e t c . ) and l o c a t i o n  (orbit to occipit), the opercular  flap  t o number,  their  (lateral  line,  maxillary,  preoperculum,  above t h e  base  of  the  pectoral f i n ) . analysis according  of  a l l characters  except  t o t h e method o f Hubbs and Hubbs t h e number  of lateral  line  (1953). cirri  was made  The  number  of  cirri  and  p l o t t e d a g a i n s t l e n g t h t o d e t e r m i n e i f any r e l a t i o n s h i p  apparent.  and  cirri  were c a l c u l a t e d was  108  Results  From i n s p e c t i o n o f characters  examined  (Figure  obvious  or  the  populations  four  consistent  The against  asymptotic Inlet,  might and  be  the  and  could  receptors,  determined  of both the  Port  to  Hence  larger  distribution undertaken.  be  and  maculosus  water i n t h e the  study  the  To  nature  ( f o r example,  variability  cirri  in  was  If  of  the  in i t s  investigation  exposed  to  of  the there  performance  differences  sensory  receptors  an i n d i c a t i o n  be  the  number  and  exposure  was  whether c i r r i of  small  provided.  number i s juvenile  c o n t i n u o u s waves o r  examined f o r e a c h o f  the the  in  might  different  Exposure, using  in  number  perceptiveness  of  groups  either  raised  homing  the  Grappler  receptors,  i n homing b e h a v i o u r  areas  number  order  chemoreceptors),  scale  laboratory.  sites,  no  between  d i f f e r e n c e s i n the  determine experimentally  was  are  cirri  This  sensory  r e l a t e d to  a f u n c t i o n o f e x p o s u r e , each o f two 0.  in  receptors.  i f the  variability  of  there  r e l a t i o n s h i p , with  Renfrew.  contained  mechanism and  relation  that  line  some r e l a t i o n s h i p between e x p o s u r e ,  of sensory  a  lateral  increasing  i f cirri  behaviour  be  and  exponential  number o f s u c h s e n s o r y  homing  could  total  cirri  Beach  that  i t i s evident  eight  examined.  number o f  possibility  15),  s u g g e s t e d an  First  g r a p h i c a l a n a l y s i s of the  differences in these characters  analysis of  length  the  same method a s two  calm for  conditions.  DORSAL Botanical First  •  inlet  Botanical  (bay)  14 FIN  4i  I  i  I  9  l_  X  L  13  11  15  10  BETWEEN  22  1  26 DEPTH  r~is I  i  I  38  J  L  06 DISTANCE  i I  J  I  34  30  Beach  NARES  L  L  12  POSTERIOR  1  I  _J I  Inlet (bay)  18  LX=i:  (bay)  Beach  L~  14  DISTANCE  LENGTH  i_  G r a p p l e r Inlet (mudflat)  38  db  HEAD  F i g u r e 15  •  34  DISTANCE  L_  G r a p p l e r Inlet (mudflat)  Grappler  30  (bay)  Beach  Botanical  26  INTERORBITAL  RAYS  X  I  X  I  Beach  Inlet  X  22  18  16  HEAD  Grappler  L  X_l  G r a p p l e r Inlet (mudflat)  Botanical  I  Beach  inlet  i  X  Beach  Grappler  LENGTH  I  ANAL  First  EYE  Beach  G r a p p l e r Inlet (mudflat)  First  RAYS  Beach  Grappler  First  FIN  ^ A  I  1  10  BETWEEN  C  . 14  ANTERIOR  !8  NARES  X  I  i  16 20 .18 22 G r a p h i c a l a n a l y s i s of eight m e r i s t i c c h a r a c t e r s at four a r e a s  .26  .30  110  F i n a l l y , h i s t o l o g i c a l examination of the c i r r i  was undertaken t o  determine whether any sensory r e c e p t o r s were apparent.  2.The r e l a t i o n s h i p o f c i r r i  Since  the  t o exposure  p r e l i m i n a r y i n v e s t i g a t i o n of c i r r i  from three d i f f e r e n t areas suggested that f i s h i n areas  have  more  cirri  (although  little  more  on f i s h exposed  difference  was  subseguently found i n r e l a t i v e exposure between P o r t Renfrew and First  Beach)  undertaken.  examination  of  0. maculosus  this  were  apparent  collected  s i t e s on the west coast o f Vancouver  phenomenon  was  from s i x a d d i t i o n a l  Island  and  five  Alaskan  c o l l e c t i o n s were examined.  Methods  To  add  to  Renfrew, F i r s t Beach Grappler except  Inlet, the  O* l i c u l o s u s  the and  further  mudflat  initial both  t h e bay  collections  for cirri  for cirri  collections and  were  analysis.  made the  at  mudflat  Port in  taken at a l l s i t e s Collections  of  a n a l y s i s were a l s o made at Cape Beale,  Benson I s l a n d , Pachena P o i n t , Helby I s l a n d , Ranee I s l a n d and t h e l a r g e t i d e p o o l on Haines covering site  a  range  Island.  The  sites  were  chosen  o f exposures, t h e r e l a t i v e p o s i t i o n s of each  i n an exposure s c a l e subseguently being determined  weight  as  by the  l o s s of the cement b l o c k s i n s t a l l e d a t each s i t e , except  111  the  mudflat i n G r a p p l e r I n l e t .  Immediately 105S  formalin  The  fish  sexed  and t h e c i r r i  transferred  (total  determine  Fish  whether  collections  five  examined  fixed  isopropyl  in  alcohol.  0.005 to  cm, their  and  f  (BC63-1255;  from  i n a general  However,  since  Bay,  Harbor, Little  s i t e s h a s been  Kuiu Cohen  Port  made,  cirri  These  The  (BC63-151:  Island  (BC61-099:  Island  (BC63-87:  Walter,  Baranof  Island  a r e a s were l o c a t e d  exposures r e l a t i v e  manner t o t h e s i t e s  no o b j e c t i v e  to  (BC63-251:  Island  an e s t i m a t e o f t h e i r  o t h e r and  Island  Egg  56°23*N,134°38'30"W)•  charts to obtain  maculosus  with r e s p e c t  Shelter  Saginaw  Tee  o f 0.  phenomenon i s w i d e s p r e a d .  Fish  58°26 N,134°46«50"W)  on  t o each  on V a n c o u v e r  Island.  measurement o f the e x p o s u r e o f t h e the  exposure  rankings  must  be  cautiously.  The  fish  from  to the nearest according  Total for  were  recorded according  were examined  were  5 6 0 5 5 ' N , 13i*°16'W) ,  recorded  and  the apparent  55°28'30"N,133°11•15"W),  length)  t o 37.5%  Alaskan c o l l e c t i o n s  Huseum  58°26»N,134°52*W),  regarded  the f i s h  length) t o the nearest  were c o u n t e d  addition,  t h e O.B.C.  Alaskan  collection  location.  In  to  later  were m e a s u r e d  n a t u r e and  from  and  after  each  area  these c o l l e c t i o n s  0.005 cm,  t o n a t u r e and  cirri and  number was then  sexed  and  were measured t h e number  of  (total cirri  location.  examined  evaluated  with r e s p e c t  according  to  to l e n g t h exposure.  112 Functional regressions constructed the case  of  for  of  males  juveniles  total  male  and  number  vs  and f e m a l e s s e p a r a t e l y  ( l e s s than  c o u l d not be d e t e r m i n e d , t h e the  cirri  3.5 cm t o t a l  data  on c i r r i  female r e g r e s s i o n s  but  only  length  were  and combined. length)  In  where  sex  were i n c l u d e d i n  both  once  in the  combined  regression.  Using of  the  mudflat  areas  the age-length  on t h e west c o a s t  in Grappler I n l e t ,  were c a l c u l a t e d and t o t a l to  age  in  each of  were c a l c u l a t e d Juveniles  were  regressions  but  Using this  could  recorded. and  be  for  males  included only  t h e ages o f cirri  once i n  t h e age o f  Since  the  o n l y combined r e g r e s s i o n s  Again,  separately  fish  of of  sample  of  year-class  sizes  with  the area  respect  regressions  and  combined.  male and  the  cirri were s m a l l  were c a l c u l a t e d .  areas  of  each y e a r - c l a s s total  except  female  regression.  from each  the  nine  each  functional  t h e combined  calculated,  for  from  both the separate  fish  regressions  the f i s h  number was examined  and f e m a l e s in  calculated  of Vancouver I s l a n d ,  these areas.  For each a r e a ,  functional  calculated.  regressions  where  each f i s h were vs in  was  extracted age  were  some  cases,  113  Results  The variable  distribution  between  fish.  and Young  s m a l l e s t 0, maculjosus examined cirri  and  on  fish  nature  less  fish  (1.4 cm  than  of  cirri  possess total  are  few  c i r r i ; the  length)  2.0 cm few c i r r i  highly  had  no  are apparent.  There i s an i n c r e a s e i n the number o f c i r r i  and  branched  a s observed by B o l i n  cirri  (1944).  with  increasing  C i r r i i n c r e a s e i n number  although  multifid  cirri  do  and  on  on  not  l o c a t i o n s , even i n l a r g e r f i s h . eye  length,  a l l parts  number  of  the  of  body  appear to occur at a l l c i r r i  Cirri  appear f i r s t  around  the  the top o f the head p r i o r t o being found along the  l a t e r a l l i n e s and m a x i l l a r i e s ,  preoperculars,  Cirri  be  do  the  not  distribution.  appear  to  bilaterally  Uneven d i s t r i b u t i o n appears  to  operculars e t c . symmetrical be  more  in  common  than not. Along  the l a t e r a l l i n e c i r r i  begin to appear i n s m a l l  f i s h immediately i n f r o n t of t h e a n t e r i o r pores. the c i r r i  mid and upper  of  cirri them  fish  precede the a n t e r i o r h a l f t o two-thirds of the l a t e r a l  l i n e pores and a r e b i f i d  two  In l a r g e r  margin  and t r i f i d  i n larger f i s h .  Around  the  of the eye, s m a l l specimens may have one or  while l a r g e specimens may have up t o nine c i r r i , many  multifid. On the top of t h e  variable,  although  the  head  pattern  the  distribution  i s highly  found on t h e f i s h examined i n  114  this  s t u d y d o e s n o t a p p e a r t o be t h e same a s t h a t  Bolin  (1944, p.  "The  pattern i n large adults  of three  fronto—parietal three  two t r a n s v e r s e  One  the  to  three  multifid  often  and may  cirri  occur  be l i m i t e d  pattern  behind  each  rows of up t o f i v e multifid)  Occasionally  found  o f two p a r a l l e l  cirri  (sometimes  side  these,  in  extend region.  the  midline  on t o p o f t h e head  t o a s few a s f o u r on  of the l a s t  eye.  cirri  occipital  unbranched c i r r i Up  to  were f o u n d four  row  were  i n the occippital  absent  these,  were  in  six  found. and  One o r two  to  cirri  In the s m a l l e s t pairs  of  region.  occasionally  small  two  and  were  parallel  p o s t e r i o r end o f t h e m a x i l l a r i e s ; one t o t h r e e cirri  frequently  occurred)  anteriorly.  symmetrical  cirri  to  row o f c i r r i .  or  two  of  were f o u n d s l i g h t l y b e h i n d  fish  one  pairs  respectively  i n the midline  examined  large adults i n this  Posterior  (when t h i s  one o r two c i r r i  on  transverse  were s o m e t i m e s f o u n d  Maxillary  while  each  o f t h e head i n t h e o c c i p i t a l cirri  each  s e r i e s o f two o r  laterad to  s e r i e s o f up t o t h r e e  general  study consisted  the  slightly  along  side." The  parallel  extending  In j u v e n i l e s t h e c i r r i  a l l simple  each  cirri  i s u s u a l l y formed  r i d g e , and a p a r a l l e l  top  anteriorly. are  multifid  similar c i r r i  across  by  66) :  general  by a l i n e  described  was  found more  specimens,  a t the common.  along  the  115  preopercular three any  more  of  above the the  base o f t h e  mention  spine  and  several  Smaller  on  Hart the  fin.  In  occurred fish  in  16  order  These c i r r i  although  showed one,  and  22)  The  Island.  The  up  i f  to five  a l l cases, where  s e x e s and good  not  cirri  shown  by  i n Grappler  e x p o s u r e as  omitting  base of  the cirri  in larger  fish.  number vs  Inlet  is  cirri  the  is  block  tidepool  s i t e s are from  number w i t h  arranged  arbitrarily  classified  variability  l e n g t h , but  with  l a r g e sample  on  charts.  considerable  increases  reasonably  are  length  cement  large  determined  that there  of t o t a l  between  by  three  the  same e x p o s u r e as t h e  are  smallest  preopercular  or  were f o u n d  where t h e l e n g t h d i s t r i b u t i o n s  agreement  two  end  while  mentioned  f o r Vancouver I s l a n d s i t e s  t h e number o f c i r r i there  located upper  i n the  i f  any.  mudflat  i s evident  the r e l a t i o n s h i p  areas  the  above t h e  graphs f o r the Alaskan  order of decreasing  in  and  are  location,  exposure as  as about the  It  in  Table  rating.  classified  in  were  At  or  o r two,  (1973) m e n t i o n s them, w h i l e  last  two  to ten c i r r i ,  usual  m a x i l l a r y , eye,  this  of decreasing  exposure  Haines  cirri  spine.  F u n c t i o n a l r e g r e s s i o n s of t o t a l (Figure  o n l y one  showed up  f o u r was  although  p o s t e r i o r t o t h e o p e r c u l a r f l a p above t h e  pectoral commonly  to  apparent.  cirri  showed  preopercular  i n d i v i d u a l s one  (1944), a l t h o u g h  were f o u n d  fish  In l a r g e r f i s h ,  s p e c i m e n s none was  to  Small  opercular f l a p larger f i s h  smaller  Bolin  to s i x c i r r i  common.  were p r e s e n t .  just  in  was  m a r g i n , up  length. sizes for  are s i m i l a r ,  r e g r e s s i o n s f o r males and  there  that For both is  females.  116  200. C a p e  Beale  2QQ  150.  .b 1 5 0 . o  100.  _§100. E  o  Benson  Island  o  v_  CD  -Q E  C  c  J/— —> \  " r o  50.  o 0. L e n g t h (cm)  200..  First  Beach  L e n g t h (cm)  F i g u r e 16  0. 2.  200  ^  L e n g t h (cm)  o r t  R  e  r  ,  f  r  e  w  L e n g t h (cm)  T o t a l c i r r i - l e n g t h r e g r e s s i o n s f o r 0 . maculosus from f i f t e e n a r e a s (males and females combined)  10.  117  F i g u r e 16  (continued)  118  f i g u r e 16  (continued)  119  200  Saginaw Bay Alaska  Length  200. C o h e n  5o  (cm)  Island A l a s k a  150.  j§ 100., E c  2  1  50.  0.  Figure  Length  16  (cm)  (continued)  200. S h e l t e r Island  .  '  L  e  n  9  t  Alaska  h  (° ) m  Table  fiiAsocottus  (15 a r e a s ) t o t a l  maculosus  functional reqression  statistics log  Beale  Benson  Port  Island  Renfrew  Pachena  Point  number o f c i r r i  -  ( m a l e s , f e m a l e s and s e x e s  length combined)  x v s y: y = u • v l o g x  N  Cape  22  u  v  Upper  95* c . l . of v  Lower  95X c . l . of v  Hales Fe m a l e s Combined  25 42 58  -102.7 -145.6 -130.1  261. a 3 35. 3 318. 1  306.9 386. 9 360. 1  216. 6 283. 7 276. 0  Males Females Combined  16 34 50  -29.97 -107.9 -83.17  157.7 271.97 234. 2  218.6 332.8 279. 1  96.75 210.6 189.2  Males Fe m a l e s Combined  40 83 1 18  -63.13 -79.23 -78.69  210.7 230.2 229. 7  24 1. 7 254.1 249.9  179.7 206. 3 209.6  Males Females Co abined  26 46 72  -112.4 -46.80 -67.05  262.0 178. 1 205.0  325.0 216.4 273. 3  198.9 139.9 172. 6  Pirst  Beach  Males Females Combined  37 71 102  -105.7 -fi8.07 -96.94  262.7 234. 2 248. 2  300.7 264.4 273.0  224.7 204. 0 223. 5  Helby  Island  Males Females Combined  • 46 72 1 18  -'80.20 -94.74 -86.97  219.8 244.0 231.0  267.6 275. 9 257.0  172. 1 212. 1 205.0  Ranee  Island  Males Females Co m b i n e d  75 79 154  -90.76 - 142.5 -118.3  236 .6 305.5 274. 2  285. 2 356.2 308. 1  188. 0 254. 3 240. 4  Males Females Combined  98 179 275  -89.45 -117.2 -100.6  233 . 9 281.0 253. 2  261.5 311.5 273. 3  206. 4 250. 6 23 3.2  Grappler  Inlet  (bay)  to o  Tabl^  22  (continued)  Upper  Haines  Island  Grappler  Inlet  Fish  Island  Egg  (mud)  95X c . l . of v  Lower  9555 c . l . of v  Males Females Combined  1C7 79 186  •102. i* -74.60  253. 212. 226. 8  294. 3 243.8 250.6  213.6 181.9 203. 0  Hales Females Combined  26 28 54  -78.04 -174. 9 -143.5  213. 369. 317.  275. 2 447.7 372.6  152. 0 290. 3 263. 3  Males Females Combined  20 30 45  -74.83 - 8 0 . 10 -76.33  218.2 233. 1 224. 1  267. 1 276.3 258. 1  169. 2 189. 9 190. 1  159. 181. 168.  182.0 205.7 188.4  136. 0 156. 8 147.7  210. 266. 248. 0  255.5 333. 1 290.0  164. 199. 206.  -83.ec  Baranof  Islan-d  Mai es Females Combined  21 21 23  -35.31 - 4 1 . S7 -38.04  Saginaw  Bay  Males Females Combined  39 24 62  -7C.46 -103. 6 -94.89  Shelter  Island  Males Females Combined  22 45 54  -46.56 - 5 6 . 23 -52.45  174. 196. 187,  211. 224, 213,  137. 4 167. 9 161.9  Males Feaales Combined  13 26 39  -166. 4 -64.72 -98.31  315. 185. 229.  515.0 261. 9 302.5  115.8 108. 6 156. 2  Cohen  Island  122  for  example. P o r t Renfrew, F i r s t  cases  in  which  distributions classes, large in  the  are different,  and  where  one  number o f c i r r i  the  sample  divergent,  sample  for  sizes  are  Helby  in  where  the  length  larger  size  comparable  sex, the r e g r e s s i o n s are  Benson I s l a n d , Cape  In  i n a sample show a  a r e no f i s h o f  other  Island.  small,  two l a r g e f i s h  and t h e r e  f o r example,  and  particularly  or  the  Beach  Beale  and  size widely  Pachena  Point.  No and  there  from  consistent  i s no e v i d e n c e  more  d i f f e r e n c e s a r e a p p a r e n t between of a greater  exposed  areas.  regressions  of t o t a l  cirri  variability  between  consistent  with  but  17  and  considerable data show  show  variability.  i t  does  area.  for  of  total  not f o l l o w a  cirri  i t i s evident  f o r t h e same  are  Comparison  again  constructing  a  is  good  such  age  that there i s the  of the regressions  cirri  length  does not and  the  t o show any c o n s i s t e n t  number  illustration  r e g r e s s i o n s on few p o i n t s  variability  vs  r e l a t e d t o exposure.  regressions of t o t a l  simply  is  pattern  number  reasons that  t h e s e x e s combined f a i l  between a r e a s  The  fish  There  any c o n s i s t e n t d i f f e r e n c e s between s e x e s o r a r e a s  differences  The  23),  variability,  regressions  class  Table  on  exposure.  From t h e r e g r e s s i o n s (Figure  cirri  i s b o r n e o u t by t h e combined  v s l e n g t h f o r each  areas  relative  This  number o f  sexes  that  vs age by  year-  o f the h o p e l e s s n e s s  of  ( F i g u r e 18 and T a b l e 2 4 ) .  comparisons  are  rendered  Benson Island  200.  2QQ C a p e Beale  « 150.  1 100. C "TO  £ l—  50.  0.5  i Age (years)  4.  1  200  200. First Beach  \_  o  o  0) X>  100.  100.  3 C  "ro 50 .o  50  0-5  200 P  0.5  ^ A g e (years)  a c n e n a  4  Island  o  o _g 100.  o a) 100 E  E C  C  S' 50.  50  F i g u r e 17  ^-Age (years)  t 150.  150.  0.5  ^'by  200  Point  M—  S  Renfrew  E  0  ,o  t  o Q)  3 C  o  r  o  E "ro  o  4.  150.  150.  u—  \_  P  A g e (years)  .o  1.Age (years)  0.5  1- Age (years)  T o t a l c i r r i - a g e r e g r e s s i o n s f o r 0. maculosus from nine areas (males and females combined)  4  124 Rcince  Island  A g e (years) 2QQ Grappler Inlet bay  o  150.  M—  O  _g 100. E 3  C  J2 .o  50.  0.5  200.  1  - A g e (years)  Haines Island  .b 150. o u—  Q  | E 3  100.  £  50.  C  ,o  0.5  F i g u r e 17  (continued)  1.  Age (years)  .4.  Table  Oligocottus functional  naculosus  x vs  N  Cape  Beale  Benson  Port  Island  Renfrew  Pachena  Point  (9 a r e a s )  r e g r e s s i o n sta t i s t i e s loq  23  tctal  (males ,  y: y =  u  u •  number  of c i r r i  f e m a l e s and vlog  -  sexes  aqe  combined)  x  V  Upper  95? c . 1 . of V  Lower  9 5 * c, of V  Hales Fenales Combined  25 12 58  17.44 9. 18 3 8.847  214.2 270. 4 256.9  251.3 312. 1 290.9  177. 2 228.7 223. 0  Hales Fenales Combined  16 3D 50  69.02 62 . 15 63.44  117. 1 201.5 174. 2  162. 8 247.8 208. 1  7 1 . 40 155.2 140. 3  Hales Females Combined  UO 83 1 18  36. 66 30. C8 30. 45  238.6 259.0 258.6  273.6 286.3 281.5  203. 5 231.7 235.6  Hales Females Combined  26 • 46 72  29.74 48. 90 43.33  176.2 121.8 139.8  219.1 143. 8 16 2.1  133. 2 9 5 . 35 1 17.5  First  Beach  Hales Feu a l e s Combined  37 71 102  59.67 59. 75 59.60  170. 8 150. 4 159.9  197.0 170.4 176.5  144. 6 ,130. 4 143. 4  Helby  Island  Hales Females Combined  46 72 1 18  36.0 8 35.15 35. 82  152. 1 166.6 158.0  185.4 188.6 175.9  118.8 144. 6 140. 1  Ranee  Island  Hales Females Combined  75 79 154  38. 95 22.04 29.73  247. 5 352.0 311.0 -  299.3 409.7 349.6  195.7 294.4 273. 0  Hales Females Combined  98 179 275  43.9 7 42.89 41.70  190.9 229.3 206.8  213.8 254.5 223.4  167.9 204. 1 190. 2  Hales Females Combined  107 79 186  41.80 41). 00 4 6.61  182.9 151.8 162.0  212. 3 174.7 179.7  153. 5 128.9 144.-9  Grappler  Haines  Inlet  Island  (bay)  126 200.  •5  1 5 0  Port Renfrew 1973  Grappler Inlet bay 1973  200  1o 150.  '  o g 100. E c 2  _§ 100. E 3 C  50.  jB 50.  |2 '  ,o 0. 0-5  200.  1. A g e (years)  4.  0.5  Port Renfrew 1974  •5  1 5 0  4.  Grappler Inlet bay 1974  200  'E 150. o  1- Age (years)  -  o o3  X!  100.  .8 100. E  E  3 C  3 C  ro 50.  ro 50.  .o  0. 0.5  200,  1- A g e (years)  4.  0.5  Port Renfrew '1975  1  - Age (years)  • 4.  Grappler Inlet bay 1975  200  I  150.  £ 150. o  o  o _§ 100. E  _§ 100. E 3 C  3 C  ~2 50.  S 50.  .o  0. 0.5  F i g u r e 18  ,o  1  A g e (yeais)  4.  °-5  1  A g e (years)  . 4  T o t a l c i r r i - a g e by y e a r - c l a s s r e g r e s s i o n s f o r 0 ' raaculosus from two s e l e c t e d a r e a s (males and females combined)  Oliqocottus  naculosus functional  (9 a r e a s )  X  N  Beale  Benson  Port  Island  Renfrew  Pachena  Point  24  total  nuirber o f c i r r i  reqres sion loq  Cape  Table  st.atis t i e s  - a q e by  year-class  (sexes combined)  v s y: y = l) + v l o q X  u  v  Upper 95* c . l . of v  Lower  9 5 % c. of v  1974 1975  16 41  -10 1.3 23.60  533. 7 183. 2  788.2 207.7  279. 2 158.7  1974 1975  3 37  -18.21 5 5.91  359. 5 224. 0  2710 278.6  -1991 169. 4  1973 1974 1975  13 48 56  -150.4 20. 80 34.51  744. 4 301. 1 245. 6  1173 344. 4 277.0  315.3 257. 8 214.1  1973 . 1974 1975.  8 19 44  -363.4 -59.25 34.79  882. 3 395. 8 215. 3  1742 576. 5 271. 3  22. 27 215. 2 159. 2  1974 1975 1976  29 60 12  44.28 6 0.23 64.4 3  231. 8 154.7 204. 9  298.7 181.0 338. 0  164.8 128. 4 71.78  637. 6 166. 9  990. 9 190.7  284. 3 143. 2  311.3  349. 6  273. 0  First  Beach  Helby  Island  1974 1975  16 102  -129.7 35.26  Ranee  Island  1975  154  29.73  1973 1974 1975  9 223 43  -41.57 39.99 53. 45  417.0 238. 1 875. 4  787. 6 26 1. 7 1151  46.32 214. 5 599.4  1974 1975  15 170  -79 .95 4 1.67  500. 0 183. 9  708.8 207. 5  291. 3 160.4  Grappler  Haines  Inlet  Island  (bay)  128  impossible. The age in  by  year-class  with r e s p e c t  number o f c i r r i  there  is  respect a  analysis of t o t a l  a  great  to t o t a l  direct  with l e n g t h deal  cirri  of  number,  an  attempt  cirri by  maculosus  wave were  is  other  water  age.  but  that  length,  It also  the  age  and  increase  indicates  that  between p o p u l a t i o n s  with  this  variability  measured  h a l f of  to a  cirri  by  the  is  not  cement  block  placed  number  determine experimentally function  action),  t a n k , h a l f o f which was the  by  wave a c t i o n . ,  of  determined 0.  and  of v a r i a b i l i t y  3 . E x p e r i m e n t a l mani.p.uj.ation o f  In  number  t o exposure c o n f i r m s  f u n c t i o n o f exposure, as  evaluation  number  cirri  of  egual  into  two  subjected  whether  exposure  (essentially  numbers separated  the  of  juvenile  sides  of  a  large  t o a l m o s t c o n t i n u o u s waves  which r e m a i n e d c a l m ,  except  for  a  and  constant  inflow.  Methods  A divided of  1.9  tank.  large  plywood  tank,  i n h a l f l e n g t h w a y s and cm At  plywood, t o p r o v i d e one  end  of  the  to  183  a height  " c a l m " and  tank  cm  on  the  x  122  o f 3U  cm cm  x 31  cm  with a  "exposed" s i d e s of  was piece the  e x p o s e d s i d e a wave t a n k  129  was erected  ( F i g u r e 19) .  The  wave tank, based on one at t h e  Scripps  Institute  of Oceanography, was c o n s t r u c t e d of p l e x i g l a s s and was supported by  and p i v o t e d on two threaded rods p a s s i n g through the t o p end  of 5 cm x 10 cm wooden u p r i g h t s screwed t o the i n s i d e c o r n e r s o f the  exposed tank.  of  two  10  cm  By d r i l l i n g a guincunx of h o l e s through  x 10 cm x 5 cm b l o c k s o f p l e x i g l a s s a t t a c h e d t o  each s i d e o f the pivoting  each  wave  tank,  i t was  possible  to  alter  the  point o f the tank and thus manipulate t h e amplitude and  freguency  of  the  waves.  The wave tank was f i l l e d  from an overhead pipe u n t i l t h e maximum exceeded,  equilibrium  with water level  was  at which time the s l o p i n g edge of the tank would begin  to f a l l and the water would be emptied out as a wave.  The empty  tank  tank  would  then  right  itself  p o s i t i o n e d f o r the experiment every  20  seconds  established.  with  a  and  refill.  The  so t h a t a freguency volume  of  was  one  o f approximately 13 cm  wave was  3  A c o n s t a n t flow o f water was maintained i n t o  both  s i d e s of the tank as f a r as the seawater system permitted. The  end  of  p o s i t i o n e d was r a i s e d the  the  tank  at  which  10 cm t o f a c i l i t a t e  o u t l e t end of the tank, t h e l a s t  the flow of  being  waves t o  washed  out  An i d e n t i c a l p i e c e o f n i t e x was f i t t e d t o the calm  of t h e tank, t o maintain uniform c o n d i t i o n s . the  wave tank was  0.6 m o f which was covered  with n i t e x s c r e e n i n g t o permit f i s h from waves.  the  tank,  "shelter"  was  provided  in  styrofoam egg c a r t o n s and a sheet o f f r u i t  On  the box  each  side  by  side of  form of connected packing  stapled  F i g u r e 19  D e s i g n of e x p e r i m e n t a l tank s h o w i n g c a l m and e x p o s e d  s i d e s and w a v e m a k i n g t a n k  131  to  the  bottom.  In  addition,  ten  cement b l o c k s a t t a c h e d t o  e y e b o l t s anchored  i n h y d r a u l i c cement t o 5  throughout  each  side  o f each  side.  exposure  J u v e n i l e 0. and  divided  exposed  side  group  was  into  fixed  maculosus  were c o l l e c t e d  groups.  and  a  juveniles  o f 102 f i s h  preserved  measured  (total  examined  to  for  at  the  July  time  Beach  placed  i n the  and  cirri  i n each  First  the  third  counts. , and  19  Four August  of the three groups.  of  introduction  were  l e n g t h ) t o t h e n e a r e s t 0.005 cm and c i r r i  provide  distribution  preserved  side  spaced  the r e l a t i v e  from  One g r o u p was  were made between 6  total  were  tank, to determine  o f t h e t a n k , one i n t h e calm  giving  The  the  three  i n t r o d u c t i o n s of f i s h 1976  of  bricks  data  for  the  fish  remained  on  juvenile  initial  cirri  fish  t h e c a l m and  in  were  nature  and  exposed  tanks. The a period were  i n the tanks u n t i l  of approximately four  fed  pieces  of  mussel  months, d u r i n g  food. feed into the  Fresh  f o o d was  To e n a b l e t h e f i s h without  danger  t h e wave t a n k fish  which  1976,  time  they  (mostly M y t i l u s c a l i f o r n i a n u s but  o c c a s i o n a l l y Jf. e d u l i s ) , chopped amphipods.  16 December  frozen  always  used  i n t h e exposed  euphausiids  and  fresh  i n preference to frozen side  of  the  tank  o f b e i n g washed o u t o f t h e t a n k , t h e  was t u r n e d o f f f o r s e v e r a l  hours  each  to  water time  were f e d .  When  the f i s h  were removed from  each s i d e o f t h e t a n k  132  at  the  conclusion  preserved. the  of  the  experiment,  Subsequently,  0.005  nearest  cm  number and l o c a t i o n  s e e i f t h e r e were any d i f f e r e n c e s  calculated and  the  fish  conclusion  of  the  transformations untransformed  months  from  each  experiment.  of  length  were  the f i s h  the  were  of  and  fauna  several  weeks  identified. until  January,  length) to  examined  time  side  made  of  of  Loq  to  were  introduction  the  to as  vs length  tank  the  at the  base  10  by  the  suggested  removed  after  approximately  t a n k was m a i n t a i n e d f o r a l o n g e r p e r i o d .  cement b l o c k s were removed i n flora  and  data.  Although four  of t o t a l c i r r i  preserved at the  removed  fixed  between t h e two g r o u p s .  regressions  f o r the f i s h  were  t h e y were measured ( t o t a l  and c i r r i  Functional  they  which  1977  and  the  considerable  had d e v e l o p e d were removed o v e r a  i n late  The  May  August  tank  was  1977, a f t e r  and  September  period  1977  i n s i d e from i t s c o n s t r u c t i o n  which  r e c e i v i n g c o n s i d e r a b l y more  early  The  and  (July)  t i m e i t was moved o u t s i d e ,  thus  light.  Results  Subsequent number,  the  tank r e l a t i v e  to  exposure  of  this  experiment  t h e exposed  manipulating  side  of the e x p e r i m e n t a l  t o o t h e r a r e a s i n B a r k l e y Sound was shown  loss  of  weight  that  of  Grappler  cirri  by  the  o f t h e cement b l o c k s t o be a b o u t e q u i v a l e n t t o inlet.  Ranee  Island  and  Helby  Island,  133  significantly  more  significantly  exposed  side  of  indication chance  raising  great in in  deep  deal of l i g h t .  l e n g t h from  either  of tidepool  of  difference of  the  were c o n s t r u c t e d , distribution  of the d i s t r i b u t i o n s  used  in cirri  length  of c i r r i  with  25).  i s very  respect  t a n k and on  a n a l y s i s shows a This  fish  considerable  i s probably  little  the regressions o f t o t a l  a  result  the r e g r e s s i o n s  overlap  was  high  sides  respectively) .  mortality  any d e f i n i t e  cirri  in  length  vs length of f i s h  sides o f the experimental  25), i t i s evident  there  precludes  found  at t h e time of i n t r o d u c t i o n  d i s t r i b u t i o n s on which  since there  t h e c a l m and e x p o s e d  and Table  common  between t h e two s a m p l e s .  From from  not  to a  and s u b t i d a l .  ( F i g u r e 20 and T a b l e  different  effect  tank exposed are  the equal  and a r e c h a r a c t e r i s t i c a l l y  t h e sample p r e s e r v e d  Beach  was  f o u n d a l s o shows t h e  j u v e n i l e 0. m a c u l o s u s i n t o t h e e x p e r i m e n t a l First  presumably  found  in  biological  the tank  (20 m) i n t o a s h a l l o w  intertidal  some  since  side  areas.  and i n v e r t e b r a t e s  provide  Many o f t h e s p e c i e s  the lower i n t e r t i d a l  from  21  to  of species  water  t h e mid t o u p p e r  to  of algae  appears  in  The k i n d s  Comparison  of  tank  settlement  (Appendix 3 ) . of  the  of the l i s t s  o f t h e d e g r e e o f wave a c t i o n ,  of  side of the tank, but  l e s s exposed than t h e m a j o r i t y  Inspection each  than the calm  The  that  tank  i n both s i d e s o f  (Figure  the  tank  ( 7 7 % and &H% i n t h e c a l m a n d e x p o s e d fact  conclusions.  that  so  few  fish  survived  ^^Experimental tank initial cirri counts  1 3 4  ._ 100. o  CF)  75.  0)  -a 50. E ' f  25.  ro  0. 7.  Length (cm)  125.  First Beach  100. o o  75.  % E  50.  =  25.  CO  o  0. 1.  F i g u r e 20  H  Length (cm)  1  1  . 7.  T o t a l c i r r i - l e n g t h r e g r e s s i o n s from 0. maculosus i n t r o d u c e d i n t o e x p e r i m e n t a l tank and from F i r s t Beach  135  Table Experimental Functional  manipulation  of c i r r i  regression s t a t i s t i c s  for  initial log  25  and f i n a l  i n O l i q o c o t t u s maculosus  (total  samples  cirri  number -  and F i r s t  length)  Beach  x v s y: y = u • v l o g x  Statistic  First Beach  Initial cirri number  Calm side  -95.91  -43.27  -154.9  246.7  172.3  333.0  242.9  Lower 95% c o n f i d e n c e l i m i t of v  221.1  157.6  278.3  183.2  Upper 95% c o n f i d e n c e l i m i t of v  271.4  186.9  387.7  302.6  101  102  23  Have side  -99.24  16  F i g u r e 21  T o t a l c i r r i - l e n a t h r e g r e s s i o n s from 0 . maculosus i n the calm and exposed s i d e s o f e x p e r i m e n t a l tank  137  Comparison initial fish the  cirri  calm  and e x p o s e d  exposed  cirri  side  sides..  delayed c i r r i The p r e d i c t i o n  o f t h e t a n k would  placing  the juvenile  formation, that  produce  results  Inspection i n rejection  greater  note  that  o f t h e 95% c o n f i d e n c e l i m i t s of the n u l l  the  hypothesis.  relationship  l e n g t h shown by f i s h  i n t h e exposed  closer  to t h a t  resemblance  the relationship  o u t by of v  from  interesting  cirri  number and  First  from t h e calm  the  (slope)  s i d e of the tank b e a r s  shown by f i s h  shown by f i s h  both  numbers o f  It i s  between t o t a l  in  0. m a c u l o s u s i n  t h a n i n t h e calm s i d e o f t h e tank i s not borne  results.  does  with the r e g r e s s i o n o f  c o u n t s a p p e a r s t o show t h a t  i n t h e e x p e r i m e n t a l tank  the  to  of these regressions  much  Beach  side  of  than the  tank.  Thus tank can only  while  the  exposure  be r e g a r d e d as moderate,  formation  appears  experiment,  t h e numbers o f c i r r i  of  surviving  influence  have  suggest that  been  between t h e d e g r e e  produced.  side  and t h e p a t t e r n altered  w h i l e wave a c t i o n there  is  of the  of  somwhat  f o u n d on t h e s m a l l  on t h e p r o d u c t i o n o f c i r r i ,  relationship cirri  fish  to  o f the exposed  cirri  by  the  percentages may have  not  a  some  direct  o f wave a c t i o n and t h e number o f  138  4 . H i s t o l o g i c a l examination of c i r r i  To receptors  determine  whether  cirri  contain  s u c h a s t a s t e buds, s e c t i o n s o f c i r r i  any  sensory  were e x a m i n e d .  Methods  Small lateral to  lines  kept  length)  from F i r s t  eosin-haemotoxylin in  destroy  5  cirri  were t a k e n f r o m t h e  and h e a d s o f t e n male amd f e m a l e 0. m a c u l o s u s (3.9  6.3 cm t o t a l  with  pieces o f skin bearing  ppm  Beach, s e c t i o n e d  and e x a m i n e d .  Triton-X-100  for  c h e m o r e c e p t o r s , i f a n y were  and  stained  Five of the f i s h seven  had been  days i n an a t t e m p t t o  present.  Results  T h e r e i s no e v i d e n c e 22)  to  suggest  the  presence  They seem t o be composed Marshall,  personal  the  Blood  are  present  the  skin  large cirri  of  goblet c e l l s appears  to  surrounded  basement  membrane the  contains  and s m a l l contain  in  cartilage by  a  (W.S.  layer  of  of the e p i t h e l i u m .  cartilage.  While  e o s i n o p h i l i c granular  goblet c e l l s only  (Figure  o f chemoreceptors i n the c i r r i .  communication)  and  epithelium  any o f t h e s e c t i o n s  o f some s o r t o f e l a s t i c  chromatophores vessels  from  the e p i t h e l i u m  a few s m a l l ,  scattered  the cells,  of  the  goblet  Q5 mm  0.5 mm  0.5 mm  d  0.5 mm  F i g u r e 22 S e c t i o n s of c i r r i f r o m l a t e r a l l i n e (a, b) a n d h e a d (c, d) s h o w i n g s m a l l (g) a n d l a r g e ( G ) g o b l e t c e l l s , e o s i n o p h i l i c g r a n u l a r c e l l s (e), c h r o m a t o p h o r e s ( c ) , b l o o d v e s s e l s (b), l a t e r a l l i n e p o r e (p), b a s e m e n t m e m b r a n e of e p i t h e l i u m (B)  CD  i ao  cells,  and  cirri. cells in  these  Since  seem t o o c c u r o n l y the  eosinophilic  a r e mucous s e c r e t i n g  the  cirri  uncovered covered  by  cells  epithelium  towards  mucous.  r e c e p t o r s i n the  cirri,  their  and  ( M a r s h a l l , 1977)  suggests  Since  base  granular c e l l s  that  a mucous c o a t a l t h o u g h t h e  with  the  there function  of  the g o b l e t  their  the c i r r i  the  absence  may  rest  of  the  appear  to  be  remains  unknown.  remain  skin no  is  sensory  5.Discussion  The fish  i n areas of d i f f e r e n t  suggest fish  that  that  can  appears  to  cirri  0.  be  investigation  on  there  are  be r e l a t e d be  no  of morphological differences exposures any  to  produced  by  exposure..  evidence  number o f c i r r i ,  the  that  In  desired  to  between  particular,  there  t h e r e i s a g r e a t e r number o f  f a i l u r e of the  i n the  evidence  morphological differences  m a c u l o s u s i n more e x p o s e d a r e a s .  confirmed  no  between  This  attempt  direction,  by  appeared  to  t o manipulate  the  wave a c t i o n  i n the  laboratory. The morphometric  absence  of  characters  any  differences  between  surprising.  Even i f t h e r e l a t i v e  is  with r e s p e c t  important  totality presumably  of  attributes only  one  of  to  areas  placement  homing,  possessed these  by  in was  meristic not  altogether  of sensory  selection an o r g a n i s m  attributes.  acts and Any  and  receptors on  the  homing i s relative  141  d i f f e r e n c e s i n the placement o f sensory r e c e p t o r s and  may have l i t t l e e f f e c t on the p e r c e p t i o n While  latitudinal  have been demonstrated the  difference  1953;  The importance  determining numbers i n m e r i s t i c c h a r a c t e r s 1952),  but  are probably s m a l l . carbon  dioxide  meristic  determined  Decreased  pressure,  and  before  Taning,  1952),  of  temperature  oxygen  in  has been demonstrated  the temperature d i f f e r e n c e s i n v o l v e d  increased  and i n some cases, s a l i n i t y ,  have been  morphometric hatching,  and  pressure  here  and  shown t o i n f l u e n c e m e r i s t i c c h a r a c t e r s since  characters  i n t h e study areas on Vancouver I s l a n d  i s l e s s than h a l f a degree.  (Taning,  of c l u e s .  differences i n meristic  (Hubbs and Hubbs,  involved  would be small  (Taning, 1952) .  However,  characters  are  largely  the  are  pelagic,  larvae  d i f f e r e n c e s would not be expected over such a  small  change  in  latitude. Although between c i r r i the  there  and l e n g t h  were  d i f f e r e n c e s i n the r e l a t i o n s h i p  with d i f f e r e n t amounts o f wave  action,  r e l a t i o n s h i p does not take t h e form of a simple i n c r e a s e i n  number of c i r r i  with i n c r e a s i n g wave a c t i o n .  No r e c o r d was made  of any environmental parameters other than wave a c t i o n , although one  temperature  recording  in  between the s i d e s of the tank 9.2-10.2°C).  The  high  July  showed  little  (calm s i d e : 9.3-10.2°C, wave s i d e :  m o r t a l i t y observed i n both s i d e s o f t h e  tank may, however, obscure any p o s s i b l e r e l a t i o n s h i p be v a l i d  i n the f i e l d . ,  difference  which  may  142  There l e n g t h and  is  age.  differences  increase  The  and  characteristics  increase i n  age  t h i s respect, c i r r i  has  distributrion  since  cirri  d i f f e r from the  of  the  and  the  genetic  members  of  of  study  branching  where  has  differences  one  age  morphological constant  y e a r - c l a s s may  length  provided  no  number  of  in  have  parents  year-classes.  The  f u n c t i o n o f the c i r r i remains unknown.  of  structures  (Schulte  While  s i m i l a r t o t a s t e buds i n the  c i r r i - l i k e s t r u c t u r e s of other  tentacularis  and  ( B o l i n , 1944)., In  majority  This  with  obvious  are found throughout the  species.  against)  to s e v e r a l  presence  and  f i s h , for  Holl,  s e n s i t i v i t y of c a t f i s h b a r b e l s 1967)  number  investigated i n f i s h populations  evidence f o r (or  and  number of c i r r i  been noted before  p r o p o r t i o n s or ranges o f values  belonging  the  i n the r e l a t i o n s h i p between number o f c i r r i  length  cirri,  in  However, there do not appear to be any  by y e a r - c l a s s e s . with  an  1972)  and  tentacles  example,  Blennius  the chemoreceptive  (Hoagland, 1933;  suggested that such a f u n c t i o n was  the  likely  Bardach e t  al.,  f o r the c i r r i  0. maculosus. t h i s does not appear to be the case.  The  of  presence  of  s i m i l a r s t r u c t u r e s on other i n t e r t i d a l and s u b t i d a l c o t t i d s ,  for  example, Oligocottu.s s n y d e r i Greeley,  Clinocottus D§sycottus lateralis, and  embryum setiger  Bean,  for  and  Starks),  ftsemichthys  A. h a r r i n g t o n i , A.  agonids,  Asterotheca  (Jordan  example,  Agonus  (Greeley),  C. qlobiceps»  taylprj Gilbert,  fenestralis  p.entacanthus ( G i l b e r t ) ,  0. r i m e n s i s  Jordan  and  Artedius Gilbert,  acipenserinus  Tilesius,  A. i n f r a s p i n a t a  (Gilbert) ,  143  Bothraggnus interesting function  swani  (Steindachner)  to speculate  seems  to  be some k i n d  pressure) d e t e c t i o n . live  0. m a c u l o s u s  movement o f water  nebulosus currents Blennius touch. the  true  The c i r r i stand  (Lesueur)  o f water  The most movement  are flexible from  the  body,  barbels  responded t o minimal  gattpru^ine  suggested  Linnaeus  However, f u r t h e r  were  that  structures.  and  on  that the  appendages. r=Ictalurus 1  touch  or  water  t h e head t e n t a c l e s o f  particularly  investigation i s reguired  of these  plausible  such  ameirus  i t  (or possibly  these  that  (1914)  catfish  makes  structures  showed  and B a y l i s s  function  out  function.  might c a u s e d e f l e c t i o n s o f  (1933)  Hoagland  on t h e i r  1973)  (Hart,  sensitive to to  determine  144  VI.  OTHER FACTORS AFFECTING VARIABILITY  Since 0.  m a c u l o s u s and  differences  in  juvenile  0.  time they  first  homing  ability  i t  is  possible  that  there  homing  ability,  age,  length  and  year-class  emphasis  was  placed  on  of behaviour at  the  area  and  maculosus,  relatively  be  since  well  more  examination evidence  revealing  developed  change  Particular  b e g i n t o show  might  HOSING BEHAVIOUR  homing  d i f f e r e n c e s were e x a m i n e d .  may  IN  of  than  homing b e h a v i o u r  over  the l i f e  may  be  genetic  fidelity  examination of l a r g e r  of  of  the  fish.  1. Age  To behaviour  determine  is  a  untreated f i s h  whether  function  of  the age  variability the  homing  was  e x a m i n e d w i t h r e s p e c t t o age  on  homing  in  homing  performance and  of  sex.  Methods  Data maculpsns Beach  used  were  used  regression determined and  ten for  from by  the  of  analysis.  previously,  S i n c e i t was sexes,  only  untreated  experiments  known l e n g t h s and  sex.  between  homing this  calculated  initially  differences  in  behaviour  over  Using the  ages  Oligocottus 60  the of  m at  age-length fish  the data c l a s s i f i e d  apparent  that  t h e combined  First  were by  age  t h e r e were  no  d a t a were  used.  145  T h r e e a n a l y s e s were c o n d u c t e d group  i n each experiment:  the  percentage  remaining  percentage of s u c c e s s f u l facilitate standard  calculated are  percentage in  the  of  o f t h e means  transplant  merely  individually  to  and  the  t o t h e home p o o l .  To  t h e d a t a , t h e mean p e r c e n t a g e s and (Snedecor  reflect  tabulated  straightforward  age  homing,  area  and C o c h r a n ,  f o r e a c h a n a l y s i s i n e a c h age g r o u p .  used  f o r each  successfully  homers r e t u r n i n g  presentation  errors  the  on homing p e r f o r m a n c e  what  is  percentages,  1967)  were  The mean v a l u e s  apparent  but  from  in  the  a  more  manner.  Results  While experiments is  year  that  old  home  pool,  0. m a c u l o s u s is  variability under  between  ( F i g u r e 23 and T a b l e  26) , i t  are r e l a t i v e l y  and t o a l e s s e r  However, o f t h o s e f i s h  that  is  egual  which  percentages  in  results  homers  extent,  three  returning  precision.  With  w i t h aqe i n t h e p e r c e n t a g e  t h e y were  poor  do home, a l l age  remaining i n the t r a n s p l a n t area, i t  a decline  t o which  variability  year o l d f i s h  t o show e q u i v a l e n t  These  fish  one  fish.  appear  area  considerable  w i t h two y e a r o l d f i s h ,  groups  there  is  and w i t h i n age g r o u p s  evident  compared  there  to  respect appears  the to that  remaining i n the  transplanted.  suggest  that  homing p e r f o r m a n c e  consideration. , I t i s  at  least  i s related  possible  some  of  the  t o t h e age o f t h e  that  "learning*'  of  Mean Percentage Homing by Age  146  irjor 287  33 c  CD  50 104  0  Mean Percentage Remaining in the Transplant Area by Age c  CD u  100r  104  50 287  OJ D_  33  0  Mean Percentage of Homers Returning to Home Pool by Age  100r  18  16 190  c U  50  0  Age (Years) F i g u r e 23  H o m i n g p e r f o r m a n c e b y age ( m e a n + 2 s . e.)  Table  Hcming  Date  released  Percentage  26 May 1975  successfully  11 Aug 1975  26  performance  4 sept 1975  21 S e p t 1975  13 8  1 100  1 100  Age 2 Number r e l e a s e d P e r c e n t homed  21 08  44 59  59 78  30 77  Age 3 Number r e l e a s e d P e r c e n t hcmed  8 50  9 33  9 56  3 33  Age  i n transplant  Age 3 Percent  16 J u l y 1977  5 Aug 1977  9 Auq 1977  13 31  18 61  17 82  21 19  15 20  38 68  30 53  38 11  30 60  1 100  2 100  0  14  14  13  22  29  31  50  67  67  22  18  34  53  66  47  remaining 33  0  remaining  Percentage  o f homers  r e t u r n i n g t o home  pool  1  Percent  hcming  t o home  pool  100  100  100  Percent  hcming  t o home  pool  60  88  72  Aqe 3 Percent  homing  t o home  pool  100  67  60  Age  4 July 1977  area  15  remaining  2  Percent  Age  15 S e p t 1976  1  Percent Age  remaining  .23 Aug 1976  hcming  Age 1 Number r e l e a s e d P e r c e n t homed  Percentage  b y age  2  25  57  100  73  64  50  33  69  44  0  100  61  50  148  some  kind  tested  is  involved  s i m p l y i n t o age  account  for  a  in  homing,  groups  great  deal  e l u c i d a t e the t i m e a t which  but  d i v i s i o n s of the  are obviously too gross to of  this  either  the observed v a r i a b i l i t y learning  o f the  fish  or to  a r e a might  take  place.  2.Year-class  To behaviour function was  determine  is a result  whether of genetic  of y e a r - c l a s s ,  the  variability  d i f f e r e n c e s and  t h e homing p e r f o r m a n c e  examined w i t h r e s p e c t t o y e a r - c l a s s and  in  homing  specifically  of untreated  a  fish  sex.  Methods  used  in  were u s e d length  on  ten  displacement experiments  in this  homing p e r f o r m a n c e  analysis.  regressions,  conducted, homing  Data  behaviour  initially,  by s e x .  were  was then  untreated o v e r 60  0.  in  which  the  a s s i g n e d t o each analyzed  Beach  from  age-  experiment  was  fish.  by  maculosus  m at F i r s t  U s i n g t h e ages c a l c u l a t e d  and t h e y e a r  a year-class  of  The  data  year-class,  A g a i n , s i n c e t h e r e were no  sex  on and  differences,  t h e d a t a were c o m b i n e d . The identical  to  subseguent that  analysis  conducted  f o r the examination  on  the  data  o f homing p e r f o r m a n c e  was by  149  age,  except  that i n t h i s  case,  the  analyses  were  performed  by  year-class.  Results  While  again  y e a r - c l a s s e s and it  is  homing  evident  between e x p e r i m e n t s t h a t the  performance  differences year-class  there i s considerable v a r i a b i l i t y  in  are  homing  only a  apparent  performance.  percentages  homing and  greater  transplant  area  the  y e a r - c l a s s e s examined t h r e e age class  groups.  differences  (1974  of  year  t h r e e f a c t o r s examined.  homing  age  old fish,  27),  related  contain  performance  shows  remaining  year-classes. 1975)  Table  Thus f o r e x a m p l e , t h e  Thus, i n g e n e r a l , t h e r e in  the  percentages  other and  one  and  year-class differences i n  reflection  which c o n s i s t s o f o n l y  than  ( F i g u r e 24  within  a p p e a r t o be with  lower  in  O n l y two fish  1977  the  of  from no  the a l l  year-  respect to  the  Mean Percentage Homing by Year Class  lOOi  53 155  113  C <D  if  29  50  0'  Mean Percentage Remaining in the Transplant' Area by Year Class c  73  74  75  76  100r  113  50  Q_  29  155 53  I  0  vlean Percentage of Homers Returning to Home Pool by Year Class  100r  73  74  75  76  31  64  13  105  c  U  50  0  73  74  75  Year Class F i g u r e 24  H o m i n g p e r f o r m a n c e b y y e a r - c l a s s ( m e a n + 2 s. e.)  76  Table  a.  Homing  performance  , (percentages  Date  released  26 May  1975  11  Aug  1975  27  year-class  successfully  4 Sept  1975  hy  21  Sept  1975  homing)  23  Aug  1976  15  Sept  1976  Year-class 1977 Number r e l e a s e d P e r c e n t homed  'ear-class 1976 Fercent released P e r c e n t hcnted  Year-class 1975 Number r e l e a s e d P e r c e n t hcmed  13 8  1 100  1 100  Year-class 1974 Number r e l e a s e d P e r c e n t homed  21 48  44 59  59 78  30 77  Year-class 1973 Number r e l e a s e d P e r c e n t homed  8 50  9 33  9 56  3 33  4 July  16 J u l y  5 Aug  9 Auq  1977  1977  1977  1977  21 19  15 20  38 11  30 53  13 31  2 0  38 68  18 61  17 82  1 100  1 0  30 60  2 100  Table  b. Hominq (percentage  Date  released  26 May 1975  11 Aug 1975  27  (continued)  perfotuance remaining  4 Sept 1975  by y e a r - c l a s s  in transplant  21 S e p t 1975  area)  23 Auq • 15 S e p t 1976 1976  Y e a r - c l a s s 1977 Number r e l e a s e d Percent stayed  K e a r - c l a s s 1976 Percent released Percent stayed  Y e a r - c l a s s 1975 Number r e l e a s e d Percent stayed  13 15  1 0  1 0  Y e a r - c l a s s 1974 Number r e l e a s e d Percent stayed  21 14  44 14  59 29  30 7  Y e a r - c l a s s 1973 Number r e l e a s e d Percent stayed  8 13  9 22  9 0  3 33  4 July 1977  16 J u l y 1977  21 67  15 67  30 53  13 31  2 50  38 34  18 22  17 18  1 0  5 Auq 1977  9 Auq 1977  38 66  30 47  2 0  1 0  Cn  table  c. H o m i n g (percentage  Date  26 Hay 1975  released  Y e a r - c l a s s 1977 Number r e l e a s e d Percent h o m i n g t o home  pool  Y e a r - c l a s s 1976 Percent released P e r c e n t h c m i n g t o home  pool  11 Aug 1975  27  (continued)  performance  o f homers  4  Sept 1975  year-class  homing  t o home  2 1 Sept 1975  2 3 Aug 1976  pool)  15 S e p t 1976  13 25  Y e a r - c l a s s 1975 Number r e l e a s e d Percent h c m i n g t o home  pool  13 100  '1 100  1 100  Y e a r - c l a s s 1974 Number r e l e a s e d Percent h o m i n g t o home  pool  21 60  44 88  59 72  30 £7  9 67  9 60  3 100  Y e a r - c l a s s 1973 Number r e l e a s e d Percent h o m i n g t o home p o o l  by  8 100 .  18 73  17 64  14 J u l y 1977  16 J u l y 1977  21 50  15 33  38 69  30 44  5 Auq 1977  9 Auq 1977  38 100  30 61  2 50  154  3. L e n g t h  Since t h e r e appears performance size was  with age,  t o be  an  improvement  homing p e r f o r m a n c e was  c l a s s e s t h a n a r e encompassed by one done t o e l u c i d a t e more p r e c i s e l y  juvenile  fish  improvement best  commence appears  expressed.  enclosure  in  to  Grappler Beach  experiments at F i r s t conducted.  juvenile  0.  maculosus  was  capabilities  to juveniles  might  time.  imply that  the size  (and age)  classes  experiments and  homing  investigating  on  had  just  groups.  In  at  Inferior  had  homing i s  conducted  in  the  performance  by  size  addition,  several  homing i n j u v e n i l e  moved i n t o  been i n t h e  homing a b i l i t y  " l e a r n i n g " had  which which  from  lower  the  fish field  pools  r e p l a c e m e n t , t h e y showed s i m i l a r which  This  over  t h e s i z e s a t which  undertaken.  which  whether  of  y e a r age  size  homing  examined i n s m a l l e r  o f t h e s e i n v o l v e d removing  determining  period  of  Inlet  Beach  One  the  o c c u r and  Analysis  classes at First  were  homing,  in  field  homing  for  that  o f t h e removed  begun t o o c c u r a t t h i s  and  fish  time.  Methods  A number o f e x p e r i m e n t s tagged  and  conducted.  untagged 172  0.  tagged f i s h  involving  maculosus  into  were i n t r o d u c e d  where a t t e m p t s were made t o e x a m i n e  pool  the i n t r o d u c t i o n the  enclosure  i n seven fidelity  of were  experiments and  hcming  155  behaviour,  concentrating  on  juvenile fish.  About a n o t h e r  t a g g e d and u n t a g g e d 0. m a c u l o s u s o f a l l s i z e s varying  times  untagged  to try  to  establish  but r e c o g n i z a b l e  pool  by i n d i v i d u a l  100  were i n t r o d u c e d a t  residents,  which  markings, c o u l d  i f  later  be  t a g g e d and e x p e r i m e n t s c o n d u c t e d on them.  The collected  majority  at First  collected  in  of  the  of  into  bay  in  various  the  fish  determining  taking the size  them t o d e t e r m i n e this,  various  undertaken. survival  screening. the  In  highest  high  tides  chances  pool  of  the  were  Fish  used  tagged  in and  I n a l l b u t one a  available  homed.  homing  view  pools,  Conseguent  behaviour  conducted  to  with  tagged  fibreglass  i f fish  window  j u v e n i l e s were i n t r o d u c e d which would n o t be c o v e r e d  to  on  were t o be  determine  i n the p o o l s ;  T h i s was  returning  and l e s s  they  covered  experiments  monitoring  juveniles  d i d t h i s and then d i s p l a c i n g  residence  days of r e s i d e n c e  low t i d e s  was c o n d u c t e d  of  i n the enclosure  fish  Regular at  two  were  possible,  in  was  and t h e p o o l  f o r s e v e r a l days.  uninterrupted  daily  length  experiment  were p u t i n a p o o l  enclosure  i n t r o d u c t i o n s were made w i t h  residence  by  of  i n the enclosure.  investigations  affected  the  Inlet.  where  a t which t h e y  a t what  One  was  Grappler  pools  up  in  a number  sexed  the experiments reported,  to  used  Beach, a l t h o u g h  e x p e r i m e n t s were measured, introduced  fish  into by  undertaken t o enhance t h e that  pool  after  several  i ni t .  (usually daily  and s o m e t i m e s  f r e q u e n t l y a t high  to determine the l o c a t i o n  tides) of  of introduced  fish.  twice pools The  156  trough at the front determine without  which  of  fish  the had  enclosure followed  was  the  also  examined  receding  water  length  classes  was a l s o  Beach  from  t e n d i s p l a c e m e n t e x p e r i m e n t s o v e r 60 m a t F i r s t  used  i n this  4,0  cm  undertaken.  by  First  analysis.  (total  size  length). . This  addition  classes  The d a t a  on homing  E m p h a s i s was p l a c e d s i z e group  c l a s s e s - 2,5 t o 2.9 cm, 3.0 t o  3.4  cm  on  fish  was d i v i d e d and  3.5  were examined i n 1.0 cm s i z e  to calculating  p e r c e n t a g e s by l e n g t h  t o t h e home p o o l f o r each standard  errors  calculated  size  and p o o l f i d e l i t y  f o r each  length  pools,  less  than  into  three  3.9  by  In  returning  percentages  analyzing Beach  i n t h e home p o o l ,  i n t h e home r a n g e  cm.  successfully  in  and i n d i s t a n t  and  1967) were  The r e l a t i o n s h i p  experiments a t F i r s t  Percentages found o n l y  nearby  class.  was i n v e s t i g a t e d  the three replacement  classes.  t h e mean  were  intervals.  o f t h e means ( S n e d e c o r and C o c h r a n ,  also  and  experiment,  Beach  to  class  at  performance  homing, r e m a i n i n g i n t h e t r a n s p l a n t a r e a and o f homers  from  line  moving i n t o a p o o l .  A n a l y s i s o f homing p e r f o r m a n c e  Larger  to  between  the  data  by l e n g t h the  home  p o o l s were  calculated.  One analysis  was  performance juveniles  o f t h e homing e x p e r i m e n t s  of (less  specifically juvenile  fish,  weeks.  conducted A  t h a n 3.4 cm t o t a l  p o o l s a p p r o x i m a t e l y 60 m away.  included  Data  total  to of  in  the  examine 38  O.  above homing  maculosus  l e n g t h ) were t r a n s p l a n t e d t o were  collected  for  five  P e r c e n t a g e s homing and r e m a i n i n g i n t h e t r a n s p l a n t  area  157  were  calculated.  To examine whether juvenile  fish  have  just  c o l l e c t i o n s of juvenile Beach.  P o o l s from  vertical  collections cm  (total  seven  length).  collection (total  These  prior to release. 30 O. m a c u l o s u s  collections  fish  42  almost  between  collection  daily  £3liJoEfii§Sfis) and c h o p p e d  between  feeding  times.  about  60  the  The f i r s t  two  2.3 and 2.7  laboratory  2.7  for  The t h i r d and 3.3 cm  f o r t h r e e weeks  initially The  involved fish  over  i n the  with s m a l l p i e c e s o f mussel mussel  Following m.  three  spanned  between  were k e p t i n t h e l a b o r a t o r y  fed  when  pools,  prior to release.  0. m a c u l o s u s  The s e c o n d  place  made  were k e p t i n t h e  (i?Jtiia§  Percentages  lower  were  b u t a number o f t h e s e d i e d .  were  take  were made from s t u d y p o o l s a t F i r s t  weeks, r e s p e c t i v e l y ,  l e n g t h ) which  transplanted  into  24 and 14 0, m a c u l o s u s  involved  laboratory  moved  may  o f study pools at t h e s i t e .  involved  and f i v e  fish  which  distribution  "learning"  was l e f t  tagging,  Data were c o l l e c t e d  homing a n d r e m a i n i n g i n t h e  i n the tanks  the  fish  f o r four  transplant  were weeks.  areas  were  calculated.  Results  In enclosure, tagged fish  fish  each  of  the  seven  experiments  t h e movements o f t h e f i s h could  were s e e n  be f o u n d .  after  were  monitored  I t i s evident that  introduction  conducted  i n the  until  no  i n a l l c a s e s few  f o r any l e n g t h o f time  (Table  158  28).  I t i s apparent t h a t there  results  whether  the  fish  is  were  little  difference  introduced  into  in  the  high or  low  percentages  of  tidepools. In experiments 1 and juvenile  H r e l a t i v e l y high  f i s h moved from the pool of i n t r o d u c t i o n to the  trough  at the seaward end of the e n c l o s u r e p r i o r t o d i s a p p e a r i n g . i n two  experiments (6 and 7)  highest t i d e p o o l  was  However once the  there  pool was  l e f t the pool and  not  same  r e l e a s e and front.  evidence  and  pattern.  in  one  observed i n the Larger  introduced  s u c c e s s i v e days i n one (recognized  by  pool  fidelity.  t i d e s , the  fish  t h i s t a b l e from tagged  pool.  individual  adult &  enclosure  and  present  f i s h were r a r e l y found a f t e r  i f they were l o c a t e d i t was  Only  of  w e l l flooded by higher  included  untagged f i s h introduced the  any  i n t o the  e n c l o s u r e very r a p i d l y .  Results  much  where f i s h were introduced  Only  number  in  the  f i s h was of  trough  at  the  found on s e v e r a l  untagged  juveniles  markings) were observed f o r s e v e r a l  s u c c e s s i v e days i n v a r i o u s pools,  but  never  for  longer  than  t h r e e days. From concluded  these  r e s u l t s i t i s evident that nothing can  be  about the l e n g t h a t which j u v e n i l e f i s h adopt and home  to a t i d e p o o l . The First  Beach  considerable  r e s u l t s of the a n a l y s i s of homing (Figure  25 and T a b l e 29)  variability  within  and  experiments  at  show t h a t while there i s overlap  between  size  Table  Summary  of experimental i n t r o d u c t i o n s into  tidepool  Experiment 1 Date r e l e a s e d Length  28  enclosure  19 September  classes(cm)  <3.0  3.0-3.4  1975  3.5-3.9 >4.Q  Total  Number r e l e a s e d  6  15  13  Percent found release pool Percent found other pools P e r c e n t moved more s e a w a r d Percent found trough  in  0  7  8  6  in  0  13  8  9  to pools in  0  7  8  6  67  53  1. 3.  2 fish  7 October  classes(cm)  Number r e l e a s e d Percent found release pool Percent found other pools P e r c e n t moved more s e a w a r d Percent found trough  1.  Fish 3.  69  34  62  Fish introduced into a l ltidepools 2. No f i s h s e e n a f t e r 6 d a y s f o u n d i n bay o u t s i d e e n c l o s u r e f o r 13 d a y s  Experiment 2 Date r e l e a s e d Length  0  <3. 0 3.0-3.4  197 5  3.5-3.9 >4 .0 0  Total 47  10  23  14  in  0  9  0  4  in  0  9  0  4  to pools in  0  9  0  4  0  0  7  2  i n t r o d u c e d i n t o one r e l a t i v e l y h i g h p o o l 2. No f i s h s e e n a f t e r 8 d a y s 2 f i s h s e e n i n bay o n c e a f t e r 8 d a y s  Table  28  (continued)  Experiment 3 Date r e l e a s e d  2 October  Length c l a s s e s ( c m ) Number r e l e a s e d Percent found i n release pool Percent found i n other pools P e r c e n t moved t o more seaward p o o l s Percent found i n trough 1. 3.  <3.0 8  3.0-3.4 4  1975  3.5-3.9 >4.0 3  0  Total 15  A l l f i s h i n t r o d u c e d a s c o n t r o l s i n t o one c o v e r e d p o o l 2. A l l f i s h s u r v i v e d up t o 21 d a y s A f t e r 21 d a y s s t o r m d e s t r o y e d p o o l c o v e r a n d e n c l o s u r e Experiment 4 Date r e l e a s e d Length  8 July  classes(cm)  <3.0  3.0 -3.4  Number r e l e a s e d  10  4  Percent found release pool Percent found other pools P e r c e n t moved more s e a w a r d Percent found trough  in  40  25  in  30  0  to pools in  10 70  1.  1976 3.5-3.9 >4 .0 3  Tota  8  25  0  24  0  0  8  0  0  0  4  50  0  0  36  33  Fish introduced into highest pool 2. No f i s h s e e n a f t e r 7 d a y s  Table  28  Experiment 5 Date r e l e a s e d  19 J u l y  Length  c l a s s e s (cm)  Number  released  Percent found release pool Percent found other pools P e r c e n t moved more s e a w a r d Percent found trough 1.  Fish  <3.0 3.0-3. 4 0  Total  8  11  in  0  0  0  0  in  0  0  25  18  to pools in  0  0  25  18  0  0  25  18  24 J u l y  classes(cm)  Number r e l e a s e d  1.  3.5-3.9 >4.0 1  i n t r o d u c e d i n t o second 2. No f i s h s e e n a f t e r  Percent found release pool Percent found other pools P e r c e n t moved more s e a w a r d Percent found trough  1976  2  Experiment 6 Date r e l e a s e d Length  (continued)  <3.0  3.0-3.4  highest tidepool 14 d a y s  1976 3.5-3.9 >4.Q  Total  8  12  in  100  75  85  in  0  8  5  to pools in  0  8  5  0  0  0  0  0  Fish introduced into highest pool 2. No f i s h s e e n a f t e r 14 d a y s  20  162  T a b l e 28  Experiment 7 Date r e l e a s e d Length  10 August  c l a s s e s (cm)  Number r e l e a s e d Percent found release pool Percent found other pools P e r c e n t moved more s e a w a r d Percent found trough 1.  {continued)  <3.0  3.0-3.4  1976  3.5-3.9 >4.0  Total  6  9  5  in  100  100  100  100  in  0  0  0  0  to pools in  0  0  0  0  0  0  0  0  0  Fish introduced into highest tidepool 2. No f i s h seen a f t e r 3 d a y s  20  Mean Percentage Homing by Length Class  1 63  1 0 0  60 147  20  108 c  OJ u $_ OJ Q_  21  50 34 27  0  vlean Percentage Remaining in the Transplant A r e a by Length Class c  OJ  u l_ OJ Q_  /lean Percentage of Homers Returning to Home Pool by Length Class  J  l  Xl  -I  1  1  1  1 0 0  50  q 0  o  c  OJ OJ Q_  50 h  Length F i g u r e 25  H o m i n g p e r f o r m a n c e by s i z e c l a s s e s (mean +  (cm) 2 s. e.)  I  8  I  I  Table  a.  Homing  performance  (percentaqe  Date  released  length  class  26 May 1975  11 Aug 1975  29  by l e n g t h  successfully  4 Sept 1975  21 S e p t 1975  classes  homing)  23 Aug 1976  - 2.9  Number r e l e a s e d P e r c e n t homed  1 0  0  0  0  0  3.0  - 3.4  Number r e l e a s e d P e r c e n t homed  0  0  0  0  5 0  3.5 - 3.9  Number r e l e a s e d P e r c e n t homed  5 0  0  0  4.0  -  4.9  Number r e l e a s e d P e r c e n t homed  18 17  9 78  16 93  5.0  - 5.9  Number r e l e a s e d P e r c e n t homed  4 100  31 61  6.0  - 6.9  Number r e l e a s e d P e r c e n t homed  6 67  7.0  - 7.9  Number r e l e a s e d P e r c e n t homed  Number r e l e a s e d P e r c e n t homed  - 8.9  4 July 1976  16 J u l y 1977  5 Auq 1977  9 Auq 1977  (cm)  2.5  8.0  15 S e p t 1976  0  0  0  5 0  4 25  6 0  0  17 0  2  2  0  1  0  19  6 50  0  5 20  5 40  3 100  3 33  2 0  25 56  24 38  0  6 33  26 69  21 76  13 62  7 86  13 92  8 75  0  24 67  7 29  18 89  7 71  4 75  10 80  6 67  2 50  0  0  4 50  6 33  6 67  1 0  0  0  0  0  2 100  .4 50  1 100  3 33  1 0  0  0  0  0  0  1  0  0  0  0  Table  b.  Hosing  (percentage  Date  released  Length  class  26 d a y 1975  11 Aug 1975  29  (continued)  performance remaining  4 Sept 1975  by l e n g t h  classes  in transplant  21 S e p t 1975  23 Aug 1976  -  2.9  Number r e l e a s e d Percent stayed  3.0  -  3.4  Number r e l e a s e d Percent stayed  3.5  - 3.9  Number r e l e a s e d Percent stayed  5 0  4.0  - 4.9  Number r e l e a s e d Percent stayed  18 17  9 0  16 31  3 0  3 33  Number r e l e a s e d Percent stayed  4 25  31 19  26 35  21 48  13 31  - 6.9  Number r e l e a s e d Percent stayed  6 17  7 14  18 17  7.0 - 7.9  Nurater r e l e a s e d Percent stayed  4 0  6 0  6 0  1 0  8.0  Number r e l e a s e d Percent stayed  .4 25  1 100  3 0  1 100  6.0  - 5.9  - 8.9  15 S e p t 1976  <J J u l y 1976  16 J u l y 1977  5 Auq 1977  9 Auq 1977  (cm)  2.5  5.0  area)  1 0  70  5 80  4 75  17 71  5 20  6 67  2 50  21 62  6 33  5 80  5 80  2 50  25 48  63  7 14  13 23  25  10 20  6 67  2 50  '  24  8  6 50  24 46  Table  c.  Honing  (percentage  Date  released  Length  class  2.5 - 2.9  26 May 1975  11 Aug 1975  29  (continued)  pecformarce of hoiers  by l e n g t h  hcming  4 Sept 1975  21 S e p t 1975  classes  t o home  23 Aug 1976  pool)  15 S e p t 1976  4 July 1976  16 J u l y 1977  5 Auq 1977  9 Aug 1977  (cm) Humber r e l e a s e d P e r c e n t homing t o home p o o l  1  Number r e l e a s e d P e r c e n t homing t o home p o c l  0  Number r e l e a s e d P e r c e n t homing t o home p o o l  5  4.0 - 4.9  Number r e l e a s e d P e r c e n t homing t o home p o o l  18 33  9. 71  16 60  3 33  3 100  2  25 71  24 50  0  6 50  5.0 - 5.9  Number r e l e a s e d P e r c e n t homing t o home p e e l  4 75  31 89  26 61  21 63  13 88  7 83  13 58  8 50  0  24 63  6.0 - 6.9  Number r e l e a s e d P e r c e n t homing t o home p o c l  6 75  7 100  18 88  7 100  4 33  10 50  6 50  2 0  0  0  7.0 - 7.9  Number r e l e a s e d P e r c e n t homing t o home p o c l  4 100  6 50  Number r e l e a s e d PPeerr cc e n t hhoommiinng t o home p o c l  4 100  3.0 - 3.4  3.5 - 3.9  8.0 - 8.9  0  0  0  0  0  5  4  17  0  100  0  0  0  5  0  6  2  2  1  0  100  0  1 10C  0  6  0  1  6 0  0  5 100  5  0  0  1  0  0  0  0  0  0  0  50  3 100  1  0  0  0  2 50  0  H" °? CT\  167  classes,  there  i s a d e f i n i t e improvement i n homing  with s i z e from about homing  2.5 to 5.0 cm.  i s best e x h i b i t e d .  show some d e c l i n e i n sample  sizes  Between  Older f i s h  homing  and  Despite  ability  to  return  to  cm  the  small  i t g e n e r a l l y appears t o be t r u e  that o f those f i s h which home, a l l s i z e c l a s s e s show same  7.0  {7.0 t o 9.0 cm) appear to  performance.  f o r smaller f i s h ,  5.0  performance  a  about  the  small area from a c o n s i d e r a b l e  distance. The mean percentages remaining i n the t r a n s p l a n t by  length  show  a s m a l l e r percentages o f f i s h remaining i n t h e  t r a n s p l a n t areas with i n c r e a s i n g s i z e , except size  class  found  at  First  Beach.  relatively  greater  variability  beginning  to  home,  may  these  length  largest  data show  classes,  but  shown i n the 3.5 t o 3.9 cm  s i z e c l a s s , the s i z e at which any s i z e a b l e are  f o r the  Again,  c o n s i d e r a b l e v a r i a t i o n w i t h i n and between the  area  reflect  percentage  of  fish  the d i f f e r i n g staqes o f  development of homing a b i l i t y w i t h i n t h i s l e n g t h range. , Analysis replacement  of  the  experiments  data,  (Table  by 30)  length suggests  f i d e l i t y i s not well developed i n j u v e n i l e large  fish  classes,  from  that home range compared  with  fish. From  Table  31  successful juvenile f i s h majority  of  f i s h remain  i t i s e v i d e n t t h a t t h e percentaqe o f hominq i s small  and  that  while  the  i n t h e t r a n s p l a n t area f o r some p e r i o d  of time, many subsequently move elsewhere.  The  results  appear  Table A n a l y s i s of  Date r e l e a s e d Length c l a s s  Number r e l e a s e d P e r c e n t found i n P e r c e n t found i n P e r c e n t found in P e r c e n t found i n  heme p o o l o n l y home and near pools heme ranqe d i s t a n t pools  30 June 1977  9 August 1977  4  6.0 - 6.9 Number r e l e a s e d 3 P e r c e n t found i n heme p o o l o n l y P e r c e n t found i n home and near p o o l s P e r c e n t found i n heme ranqe P e r c e n t found i n d i s t a n t pools 7.0 - 7.9 Number r e l e a s e d 1 P e r c e n t found i n heme p c o l o n l y P e r c e n t found i n heme and near pools P e r c e n t found i n heme ranqe P e r c e n t found i n d i s t a n t p o o l s Number r e l e a s e d P e r c e n t found in P e r c e n t found in P e r c e n t found i n P e r c e n t found i n  heme p e e l only home and near pools home ranqe d i s t a n t pools  1 1 56  5.0 - 5.9 Number r e l e a s e d 1 P e r c e n t found i n heme p c o l only P e r c e n t found i n home and near pools P e r c e n t found i n heme ranqe P e r c e n t found i n d i s t a n t p o o l s  8.9  length  44  4.0 - a.9 Number r e l e a s e d 17 P e r c e n t found i n home pool o n l y P e r c e n t found i n home and near p o o l s P e r c e n t found i n heme ranqe P e r c e n t found i n d i s t a n t pools  -  by  (cm)  home fcol only home and near p o o l s heme ranqe d i s t a n t pools  8.0  experiments  28 May 6 2 June 1975  3.0 - 3 . 1 Number r e l e a s e d P e r c e n t found i n P e r c e n t found i n P e r c e n t found i n P e r c e n t found i n 3.5 -3.9  replacement  30  18  25  28 56  25  83  15  34  35  24  38 59  13  0 100  59  40 33  97  40 44  92  33  100  33  84  25  100  33  73  25  38 54  100  11  50  75  75 75  100  100  100 100  CTl  co  T a b l e 31  Homing p e r f o r m a n c e  Length c l a s s  (cm)  of juvenile Qliaocottus  2.5 - 2.9  Date r e l e a s e d Number r e l e a s e d Number homed P e r c e n t homed  Number s t a y i n g i n t r a n s p l a n t area Percent staying i n t r a n s p l a n t area Days s t a y i n g i n t r a n s p l a n t area a. homers b. non-homers  P e r c e n t o f homers homing t o home pool  3.0 - 3.4  5 August  maculosus  Total  1977  17  21  38  0 0  4 19  4 11  12  13  25  71  62  66  1#1,1»1, 1,1,1,2, 5,7,12,16  1,4,6,6 1,1,2,2, 6,7,7,12, 39 100  100  170  to  suggest  than  a b o u t 3.0  beginning the  t h a t homing cm,  to  at  b e h a v i o u r i s not which  length  a  show homing b e h a v i o u r .  a n a l y s i s of  pool  apparent  inability  associated  with  the  homing b e h a v i o u r  fidelity of  absence of  i s not  data  small  0.  possible for fish small  percentage  are  Taken i n c o n j u n c t i o n  with  by  length  maculosus  home  range  demonstrated  less  classes,  to  home  fidelity,  until  the  may  be  that  is,  a home r a n g e has  been  established.  Table 2.3  amd  kept 2.7  2.7  in and  three  cm  the 3.3  32  shows  when c o l l e c t e d  cm  when c o l l e c t e d  p e r c e n t a g e homing was that  i f  homing  moving a r o u n d anywhere on once  in  between p o o l s one  adults  was  not  beach.  Of  transplant i n the  pool.  The  from h i g h  kept  the  and fish  area  a l l the  pool  cm  or g r e a t e r  fidelity.  fish  Pish  being between  laboratory  release in  given  view  of  p r e s u m a b l y the  could  have  pools  released  for  the  small  the  fact  fish  were  moved  moved a r o u n d  rather  i s they  introduced.  after  a b l e t o home.  movement was  tidepools, that  the  majority  t r a n s p l a n t area amount o f  home  almost  s e e n a f t e r r e l e a s e more  the  r e s i d e n t s of the  were 2.9  high,  undertaken,  movement t h a n t h e Of  in  of being  surprisingly  were between  s e v e n weeks.  seen a f t e r  between t i d e p o o l s  the  the  fish  and  and  weeks showed some e v i d e n c e  number o f  m a c u l o s u s which  were u n a b l e t o  laboratory for five  The  the  t h a t 0.  than  than  widely  remaining  in  comparable t o t h a t  of  showed a w i d e r r a n g e  of  into  which  only seven  i n s i z e ) showed any  evidence  they  were  ( a l l of  which  of  strict  171  Table  32  Homing p e r f o r m a n c e o f j u v e n i l e Q l i g o c o t t u s the  laboratory  f o r varying  L e n g t h c l a s s (cm) when r e l e a s e d  Date  periods  o f time p r i o r  2.5-2.9 3.0-3.4  released  maculosus kept i n  3.5-3.9  to release  Total  19 a u g u s t 1977  Group A Number r e l e a s e d P e r c e n t homed Percent stayed i n t r a n s p l a n t area  17 0 47  7 0 71  24 0 54  9 0 67  5 0 60  14 0 64  12 17 42  29 14 62  Group B Number r e l e a s e d P e r c e n t homed Percent stayed i n t r a n s p l a n t area Group C Number r e l e a s e d P e r c e n t homed Percent stayed i n t r a n s p l a n t area  Time i n l a b o r a t o r y Group a Group B Group C  7 weeks 5 weeks 3 weeks  2 0 50  S i z e when  43 14 56  collected  2. 3 - 2. 7 cm 2 . 3 - 2. 7 cm 2.7 - 3.3 cm  172  Previous  experiments  shoved  that  b e g i n n i n g t o show e v i d e n c e o f home r a n g e and  3.0  cm,  and  to  0,  maculosus  are  between  2.5  fidelity  a g r e a t e r d e g r e e between 3.0  and  Homing b e h a v i o u r a l s o b e g i n s t o become  evident  in  classes  fish  of 27  3.0 O.  of  t o 3.4  fish cm  -  maculosus  resulted  fish  appears  that  range  four  from  in their  larger  (2.5 t o 2.9  the  field  learning  C) , ,  - one  out  of  at  being unable  (Group ^  fish  cm  34).  this  t o home  were  able  Since  time when  to  of the i n t e r t i d a l  begins to occur during t h i s  out  period  3.4  these  cm. size  (2.9  cm);  removal  (Groups replaced,  of  A and  B)  whereas  home when r e p l a c e d , i t and of  a d o p t i o n o f a home time.  4.Discussion  are  not  Differences  i n homing b e h a v i o u r  apparent.  Atkinson  three d i s t i n c t mature year.  (1939) s u g g e s t e d ,  that  I f t h i s i s the  0.  maculosus  case,  spawn o v e r s e v e r a l y e a r s , any homing b e h a v i o u r  and  may  on  the b a s i s  since  would be d i f f i c u l t t o  related  differences  Q.  macnlgsus  related  a  probably  differences in  distinguish.  in  homing  i n t o only three  g r o s s t o p r o v i d e much i n d i c a t i o n  from  of  spawn more t h a n o n c e a  genetically  apparent, although the d i v i s i o n too  year-classes  modes i n t h e d i a m e t e r f r e q u e n c i e s of ova  female,  Age  between  behaviour age  groups  o f t h e s t a g e a t which  are is fish  173  are  learning t o display  fidelity.  It  i s  successful  evident  percentage of s u c c e s s f u l decline  age g r o u p s  the  number o f f i s h  that  adopt 2.8  much  of  homing  a home r a n g e  increasingly  fish  Between t h i s s i z e  between 5.0 a n d in  classes  appear  remaining  to the  which  transplant  does  size  with egual  into  l o w e r p o o l s from  they  settle,  begin  when  the f i e l d not  a  a r e about  to affect  do n o t a p p e a r t o they  are  about  5 cm t h e y  fidelity  appears  and  to  show  homing  homing  fish.  (greater  occur  be  some  A l l size Percentages  with  length,  which  than  about  2  the high t i d e p o o l s i n  extensive  movement  between  home r a n g e and d e m o n s t r a t i n g  " L e a r n i n g " and "memorizing*  o f O. m a c u l o s u s  appear  fish  fairly  adopting  the f i s h  and  class.  moved  to  behaviour  a r e a show a d e c l i n e  having  prior  size  precision.  juvenile  show  fish  larger  that  homing b e h a v i o u r .  from  There  home,  smaller  successfully  appears  tidepools,  to  with  by  and a b o u t  range  cm.  homing  home  except f o r the l a r g e s t  cm),  7.0  percentages  in  It  home  do  a r e a with age.  to i t u n t i l  The h i g h e s t p e r c e n t a g e s  decline  pool  i s a decline i n  homing  Juvenile  homing  better developed  performance. in  detail.  or start  t o 3.0 cm l o n g .  behaviour  between  which  There  remaining i n the transplant  greater  and  an i n c r e a s e i n t h e  although o f the f i s h  classes reveals the r e l a t i o n s h i p in  i s  home w i t h e g u a l p r e c i s i o n .  Examination  age  there  behaviour  homers between age one and two and some  i n age t h r e e f i s h ,  all  honing  1  the  area  2.3 t o 2.7 cm l o n g . are  greater  homing p e r f o r m a n c e ,  than  relative  appear Removal 2.7  cm  to fish  174  o f t h e same s i z e  which h a v e a l w a y s been i n  suggested  that  juveniles  and  to a l e s s e r  about  3 cm,  Although 0.  maculosus  the  provides  the  e x t e n t , up  low  tide  evidence  t o about  of  more e x t e n s i v e i n some c a s e s .  5.5  cm  in  some  that  sheltered  their  advancing the  vertical t i d e and  case,  sheltered  distribution,  high  and  a r e a s were t h e o n l y g r o u p t o show an in  He  a r e a s , may  to  cm. of  juvenile  almost moving  acquisition,  at  continue to occur  tide  movements  found  that  moderately complete  tidal  until  shift  with  the  If  this  is  for  fish  in  tide. least  fish  sheltered  shorewards  seawards with t h e f a l l i n g  information  is  e x t e n s i v e movement between  may  l e s s than  4  distribution  (1971b) work s u g g e s t s  even  It  move e x t e n s i v e l y between p o o l s up  p o o l s . Green's be  field.  the f i s h  are  quite  large. , The increase the  i n the percentage  largest  sizes,  d e c l i n e i n the percentage  size  remaining  classes of f i s h  i m p a i r e d "memory", r e d u c e d  combination  of these f a c t o r s .  growing  tend  fish  earlier  Port  Renfrew 0.  m a c u l o s u s do  the  percentage  of s u c c e s s f u l  (Green, suggests  than  1967; that  of s e n i l i t y  slower  may  be  Khoo, they  1971)  and  not  due  swimming  Gerking  growing  fish  abilities,  in  some  that  a l s o become  senile  faster  o f the same b r o o d .  homers i n t h e l a r g e s t  i n the decline  or  (1957) f o u n d  a p p e a r t o show any  s i n c e the  area  and  t o the s m a l l sample  data  grow more s l o w l y , p e r h a p s  i s a factor  homing  i n the t r a n s p l a n t  t o mature e a r l i e r and  die  successfully  from the  i n homing  and Since  decline size  in  classes  this earlier  performance  study onset in  175  the  largest fish  at F i r s t  Beach.  A reduced r a t e of growth i n Port Renfrew f i s h to  First  Beach  fish  may  a l s o account f o r the absence o f any  d i f f e r e n c e i n homing performance Since  age  relative  i n the  or time i s presumably  smaller  size  classes.  more important with r e s p e c t t o  l e a r n i n g i n f o r m a t i o n than a c t u a l l e n g t h , i t i s probable t h a t the  time 0. maculosus  by  at Port Renfrew are 4 cm l o n g the m a j o r i t y  have had s u f f i c i e n t time t o a c q u i r e the i n f o r m a t i o n to home with ability  comparable  to  that  of l a r g e r F i r s t Beach f i s h .  examination of homing performance o f fish  (less than 4 cm)  age  at  which  may  Port  smaller  size  be necessary to determine  Renfrew  fish  classes  of  the s i z e and  adopt a home p o o l and  start  homing t o i t .  Another f a c t o r  the  of the two a r e a s , and i t s apparent e f f e c t on pool  topography  f i d e l i t y discussed e a r l i e r . freely  between  pools  which may  Thus  be of r e l e v a n c e here  Port Renfrew Q. maculosus  than  First  Beach  cottid,  Clinocottus  differences exist  globiceps.  (Green, 1973). homing  performance  (Green, 1975).  Gibson  (1967)  as  ( l e s s than 5 cm)  adults  but  ( l e s s than 5 cm) adults.  by  size  of  showed  of  classes  situation  may  irregular.  performance  A similar  the  fiholis  in  homing  move more  0. l a s u l o s u s and  a c g u i r e i n f o r m a t i o n e a r l i e r s i n c e the beach i s l e s s I n v e s t i g a t i o n o f the  is  was  another  showed  no  found  to  Tantoqolabrus adspersus that  juvenile  Blennius  d i s p l a y the same degree o f pool f i d e l i t y  that j u v e n i l e Acanthocottus r=Enophrys1 b u b a l i s appeared  Williams  to  show  less  pool  fidelity  than  (1957) found no s i z e d i f f e r e n c e s (between  36  176  and  100*  mm)  analysis  i n pool f i d e l i t y  of  his  disappearing Girella than  36  found  mm,  to  deeper that  smallmouth  bass  cm  higher  showed  Complementary that  bass  up  changes over  fish,  evidence t o 3 3 cm  of was  degrees  different  A j n b l o p l i t e s r.up_es_tris  by  fiUSctatus  intermediate size  percentage  also  as s m a l l as  pool f i d e l i t y l o n g term Both  true  for  the l i f e  Larimore  some  than by  move  which  suggests  showed  that  than  (1957)  mobile but fish.  fish.,  who  the  25.4  found  number o f Funk  also  a c c o r d i n g t o s i z e i n rock  (Rafinesgue)  and  channel  catfish  (Rafinesgue) , g r e a t e s t m o b i l i t y being c l a s s e s i n both  to  fish.  smaller  Funk  of  fish  (1952)  sharply i n larger of m o b i l i t y  provides  Lacepede l a r g e r  return provided  were  pool f i d e l i t y  these  of the  11 mm  occurs i n small  increase in size,  were i n c r e a s i n g l y  showed  the  Williams  H i c r o p t e r u s dolomieu  declined  l£i§J:.3£i§  fish  long),  water as t h e y  mobile i n d i v i d u a l s  bass,  term  not e x h i b i t e d .  rates  in  although  he d i d n o t t a g any C . , a n a i l s l e s s  28 mm  freshwater  analis,  T h i s was  although  short  (23 t o  pool f i d e l i t y  In  increase  showed t h a t  pools.  was  subtidal  an  Although  suggest  group  Clinocottus  t a g g i n g , with s i z e .  same  nigricans  size  shows  f i n clipping the  evidence  this  after  nigricans.,  in  Girella  data  of  species.  shown  177  VII.  SENSORY MECHANISMS  !• Sensory, i m p a i r m e n t  INVOLVED IN HOMING BEHAVIOUR  methods  Vision  Fish (1971). sharp  were b l i n d e d a c c o r d i n g  A slit  was made i n t h e c o r n e a  scalpel  forceps u n t i l Although  and  perception  anaesthetized  the  field  technigue, difference  The  and  operation  lens  was  then  with  the  optic  nerve  a  with  removed. after  ultimately  was p e r f o r m e d  (1 ml  per  4.5  i n 1976, f i s h litres).  seemed good as i n d i c a t e d by s u r v i v a l  and l a b o r a t o r y .  As  experience  of a n a e s t h e t i c only  i n recovery  anaesthetized  eyeball  1949).  in guinaldine  use  the  and d a r k i s p o s s i b l e s h o r t l y  the retina  this  the operation  of  s i d e s o f t h e e y e b a l l were p r e s s e d  of light  (Rasguin, When  from  the  t h e l e n s emerged.  lens removal, both degenerate  t o t h e method u s e d by Khoo  fish  experiments conducted  from  gained  increased handling  the operation  and t h o s e  was  was  observed  Recovery i n both i n the time;  was u s e d .  no  between  w h i c h were n o t a n a e s t h e t i z e d .  i n 1977 no a n a e s t h e t i c  were  In  178  Olfaction  Several Initially  heat  methods were a t t e m p t e d t o make c a u t e r i z a t i o n was u s e d  (Khoo,  fish  1971).  n e e d l e was i n s e r t e d i n t o t h e a n t e r i o r n a r e s and moved position  of  guinaldine  the olfactory rosette.  anosmic. A r e d hot into  the  F i s h were a n a e s t h e t i z e d i n  p r i o r to the operation.  Recovery satisfactory, laboratory.  from  as  the  indicated  Examination  0. m a c u l o s u s destruction  which  by of  had  in  appeared  olfactory  rosettes  this  However, s i n c e  examine  each  this  possible  that not a l l released  to  be  s u r v i v a l b o t h i n t h e f i e l d and  undergone  of the rosette. fish  operation  regard fish  treatment  six showed  i ti s impossible  prior were  of  to  fully  to  release, i t i s anosmic  after  the uncertainty  of the  treatment.  To total  overcome  destruction  of  the the  problem  of  olfactory  methods were a t t e m p t e d t o make t h e f i s h olfactory of  nerve  l a t e x proved  nares.  impossible  of  the  because  irritating  o f f when i t h a r d e n e d .  Adhesive  (Norcliff  either dissolved  several  other  anosmic.  Cutting  o f the  b u t was u n s u c c e s s f u l . of  Covering the nares with v i n y l  because peeled  was a t t e m p t e d  rosettes,  the  acetate  size  of  the  was u n s a t i s f a c t o r y  n a t u r e o f t h e f u m e s and b e c a u s e i t Orafix  Special  P r o d u c t s ) and L e p a g e * s  or f a i l e d  small  Injection  t o adhere.  Plastic  Plastic  Hodel  Denture Cement  179  I n one e x p e r i m e n t Eastman  910  Adhesive  the nares  which  adhered  t h r e e days i n t h e l a b o r a t o r y . Finally, (ESPE  Fabrik  the nares. seven off.  a dental  days  After  glue,  Pharmazeutische  T h i s adhered  were  blocked  with  satisfactorily  this  f o r about  time i t p e e l e d o f f .  Durelon  Carboxylate  P r a p a r a t e GMBH) was u s e d  extremely  w e l l i n the  t o two weeks, b u t a g a i n  No a n a s e t h e t i c was used  Kodak  after  when e i t h e r  t o block  laboratory  this of  Cement  from  period i t peeled these  treatments  was a p p l i e d .  Taste  contained  The  possibility  that  taste  buds, s i m i l a r  to those  Blennius tentacularis these  and  other  homing, r e s u l t e d in  an a t t e m p t  surface  of  susceptible effect  chemosensory  and  act that  damage.  as come  natalis  histological to ten days).  receptors  0.  1972)  (Lesueur)  disintegration  et  a l .  found  al,  that  (1965)  Detergents thus  contact  detergents and  and  might be i n v o l v e d i n  f a t solvents into  maculosus  i n the t e n t a c l e s of  t a s t e buds.  Bardach  o f low c o n c e n t r a t i o n s o f  Ictalurus  found  of  i n u s i n g t h e method o f B a r d a c h e t  fish  to  cirri  ( S c h u l t e and Ho.ll,  t o d e s t r o y any such  tension  membranes  Brunn  the  reduce  making a l l  with  detergents  (1965) e x a m i n e d t h e (1  to  5  that  o f t a s t e buds o c c u r r e d  Behavioural observations suggested  ppm)  observable  (after that  on  three  damage t o  180  the  olfactory  epithelium  and g i l l s may  have occurred although  h i s t o l o g i c a l examination d i d not support t h i s . taste  buds  was  clean  water.  not  observed  T h i s technique eliminate  was  Recovery  of  the  even a f t e r f o u r to s i x weeks i n  used  initially  in  the sense of t a s t e i n 0. maculosus.  attempts  Ten O.  to  maculosus  i n d i v i d u a l s were put i n low s a l i n i t y water {5 ppt) c o n t a i n i n g ppm  household  daily. was  detergent.  Nakamura salinity  to  the  freshwater  (1970) found  that  s u r v i v e d from  ppt t o 192 hours, when suggests of  The water and detergent were changed  However, a l l f i s h d i e d w i t h i n t h r e e days,  due  5  or  Whether  to the detergent was  O. ..maculosus -• exposed  this  unknown.  to  two  88 t o 102 hours and at a s a l i n i t y of the  t h a t the complete  experiment  was  terminated.  ppt 7.5 This  m o r t a l i t y recorded here was a r e s u l t  the combined e f f e c t of low s a l i n i t y water and detergent. k second attempt t o e l i m i n a t e chemosensory  made  cells  was  using octylphenoxypolyethoxyethanol ( T r i t o n - X - 1 0 0 ) , a  non-  i o n i c detergent which was  found t o induce i n v o l u n t a r y c o n v u l s i v e  e f f e c t s on the mollusc, Succinum undatum, s i m i l a r t o the induced  by  saponins  from  Marthasterias g l a c i a l i s response  is  suggested  r e a c t i n g with, or  (Mackie  or  starfish et  Astgrjas  a l . , 1968).  rubens  This  and  escape  by the authors to be caused by saponins  causing in  the  effect  damage  nervous  origin  muscle.  Damage i s caused by the  to,  chemosensory  cells  of  near the s u r f a c e e p i t h e l i u m of the f o o t  p r o p e r t i e s of the saponins,  detergent  or  surface  active  181  Ten 0. s a c u l o s u s were put i n seawater c o n t a i n i n g 5 ppm Triton-X-100.  Another t e n 0. maculosa s were run as c o n t r o l s i n  the same volume o f seawater X-100  were  conditions  changed for  (10 1) o n l y .  daily.  seven  days,  The water and  The  fish  were  at  which  time  experimental f i s h and a l l the c o n t r o l f i s h in  10%  formalin.  The  remaining  kept under these five  cells  although, which the  effect  examined,  no  separately  f i v e experimental f i s h of  7  evidence  days was  to  the  homing  were  recovery  immersion found  to  in show  experiments i n  Triton-X-100  was  t h a t the c i r r i o f  0. maculosus c o n t a i n any t a s t e buds, i t i s p o s s i b l e buds may  the  ( i f they were d e s t r o y e d ) . subseguent  of  of  were f i x e d  returned t o c l e a n seawater t o determine the period of chemosensory  Triton-  be l o c a t e d elsewhere on the body and may  that  taste  be a f f e c t e d by  detergent.  lis  Removal  Pectoral  and  2« maculosus • i n d i v i d u a l s Removal  pelvic by  fins  cutting  were with  removed sharp  from  scissors.  of one s e t o f p a i r e d f i n s appeared t o have no e f f e c t on  the s u r v i v a l of f i s h .  Moring  (1976) reported t h a t  0.  maculosus  whose p e c t o r a l and c a u d a l f i n s were p a r t i a l l y c l i p p e d showed  "no  unusual m o r t a l i t i e s o r l o s s o f movement over an extended p e r i o d " i n the laboratory,  although none was kept i n the l a b o r a t o r y f o r  182  any 0«  period  of  time  in  this  study,  field  I§SJ?i2sus w h i c h had p e l v i c f i n s removed  months a f t e r r e l e a s e ,  by which t i m e  fully  Fish  regenerated.  were o b s e r v e d also  appeared  partial  fin  reported and  up t o f i v e  t o be f u l l y  in  one  experiment)  manoeuvreability individuals One in  and  treated  0. m a c u l o s u s w i t h the f i e l d  behavioural  this  both s e t s  experiment  or  of  reported  Gibson  (1967) pholis  fins  ( a s was u n d e r t a k e n  an  obvious  movement.  loss  of  0. m a c u l o s u s  manner t e n d e d t o l i e on one s i d e . o f paired  f i n s removed  other  was  and i n t r o d u c e d  designed  interaction  was  seen  home  from pool.  the transplant  pool  fish  t o determine  between  0. m a c u l o s u s i n a t i d e p o o l  departing  to t h e i r  in  control  between r e s i d e n t  This  fish  fins  81 d a y s a f t e r r e l e a s e .  2.Interactions  introduced  removed  (1976)  and  be  bubalis.  resulted  in  to  a f t e r c l i p p i n g i n Blennius  Removal o f b o t h s e t s o f p a i r e d in  appeared  Boring  0. m a c u l o s u s  f i n regeneration  Acanthocottus [=Enophrysj  fins  by which t i m e t h e s e  regenerated.  of  were made up t o s e v e n  had had p e c t o r a l f i n s  months l a t e r ,  regeneration  full  which  the  observations  the  whether a  resident  and  r e s u l t s i n the introduced and  possibly  returning  183  Methods  A from  two  100  m  total  nearby  pools  away.  resident  fish  refilled  by  The  36 and  bucketing the  pools  21  and  respectively.  Oligocottus transplanted  transplant  collected  introducing 11  of  and  in  transplanted fish  t o two  pools  returned  water  m a c u l o s u s were  were  t o the  from  and  D a t a were c o l l e c t e d  lab.  From 10  The  26  about and  the  pools  pools the  and  f o r two  pools  b a i l e d out  other  fish.  were removed  other  collected  two  were  prior  to  transplant  were  replaced,  months.  Results  While  the  percentage  of  0.  homing i s r e l a t i v e l y  small i t i s evident  fish  the  remaining  larger In  a l l  of the  release subseguently  The be  partially  transplant  small a  pools  experiments.  fish  function w h i c h was  that  pool  the  successfully percentage  i s not  observed  i n the  of  significantly  homing e x p e r i m e n t s  (Table  33).  transplant  pools  homed.  percentage of  fish  of the  than  still  (1971)  successfully  homing  d i s t a n c e between t h e  greater  However i t was  m from w h i c h d i s t a n c e Khoo home.  transplant  t h a n i n many o f t h e o t h e r  addition,  after  in  maculosus  in  the  other  w e l l within the  found  that  0.  home  may and  homing  range o f  maculosus  300 can  Table  Homing  33  performance a f t e r t r a n s p l a n t cleared of resident  Date r e l e a s e d  fish  12 O c t o b e r 1976  Number r e l e a s e d  36  Number homed P e r c e n t homed  14 39  Number r e m a i n i n g i n t r a n s p l a n t area Percent remaining i n t r a n s p l a n t area Days r e m a i n i n g i n transplant area a. homers b. non-homers  P e r c e n t o f homers r e t u r n i n g t o home p o o l  1,1,1  86  pools  185  This  experiment  0. m a c u l o s u s i n o t h e r increase pool  homing  for  suggest  that fish  Since  that  of  there  departure  from  bailing from  that  while  some  the  transplant  because o f t h e removal i s  no  evidence  i n t e r a c t i o n s between r e s i d e n t and f o r the majority  out  a  tidepool  the pool,  by t h e i n t r o d u c e d the  pool.  does  i ts t i l l  diluted after bailing  of f i s h  not  leaving  remains a p o s s i b i l i t y  (from  the resident f i s h )  and r e f i l l i n g  fish  Although  remove a l l  the pool, i s  and  results  in  this  problem  might  the i n t r o d u c t i o n o f t r a n s p l a n t e d  several  t i d e c y c l e s , t o allow  possible  that  other  untreated  a n d homing.  overcome by d e l a y i n g  period.  remaining i n  i s some p h e r o m o n a l i n f l u e n c e  detectable  from  r e s i d e n t s , there  are responsible  water  which, a l t h o u g h still  pool  behavioural  t r a n s p l a n t pool  traces  data  some p e r i o d o f t i m e may o c c u r ,  introduced the  the  experiments suggest  i n the percentage of f i s h  o f some o f t h e t r a n s p l a n t to  and  fish  tidal  fish  f l u s h i n g of t h e p o o l ,  may h a v e moved i n t o t h e p o o l  over  their be for i t is this  186  3. N a t u r e o f movement between r e l e a s e a n d home, J2Qol  One  experiment  was  conducted  homing p . m a c u l o s u s showed any e v i d e n c e revealed the  by t h e i r  release  transplanted angular  the  home  to a c e n t r a l  pool  pools, were  determine  of directed  low t i d e d i s t r i b u t i o n .  and  to  whether  movement  The l o c a t i o n s ,  of  two  analyzed  groups with  as  between of  fish  respect  to  direction.  Methods  A one  total  g r o u p and 37 from  pool  about  home p o o l s ) .  all  directions  As f a r as t h e t i d e  were  When a f i s h  sample  sizes  were p l o t t e d  were  a  height permitted,  was l o c a t e d ,  were the  pools  in  searched angle  of  Data  from  days f o l l o w i n g r e l e a s e a n d , b e c a u s e  the  small,  on c i r c l e s ,  to  (27 and 33 m f r o m  p o o l r e l a t i v e t o t h e r e l e a s e p o o l was r e c o r d e d . three  (29 f r o m  transplanted  between t h e r e l e a s e and home p o o l s  each o f t h e f i r s t  used  t h e o t h e r group)  transplanted f i s h .  that  from two g r o u p s o f p o o l s  midway between t h e home p o o l a r e a s  the  for  o f 66 f i s h  from  the f i r s t  t h r e e days  combined,  s i m i l a r t o diagrams of vanishing  for birds i n orientation  the  case  of the  data,  more than  o n c e i n t h e same p o o l i n t h e a r e a u n d e r c o n s i d e r a t i o n .  t h e one f i s h  one f i s h  In  combined  For  a l l except  analysis.  points  h a d been s e e n o n l y o n c e , o r  seen i n pools i n d i f f e r e n t  directions  from  the  187  release its  pool,  the  initial  sighting, in the direction  home p o o l was u s e d , t o a v o i d  bias.  Two k i n d s o f a n a l y s i s were c o n d u c t e d . records  and  f o r t h e combined  compare t h e directions  home of  directions  opposite  records, (85° a n d  F o r each  sign tests 235°)  day's  were used t o  with  the  mean  e a c h o f t h e two s a m p l e s t o d e t e r m i n e w h e t h e r t h e  mean d i r e c t i o n  t a k e n by e a c h sample c o u l d be c o n s i d e r e d  as t h e  home d i r e c t i o n  ( B a t s c h e l e t , 1965).  t h e two  samples  for  each  day  and  f o r the  different,  test,  on a mean s g u a r e d e v i a t i o n was used  1962;  Batschelet,  Watson  combined  significantly based  the  To d e t e r m i n e w h e t h e r  data  nonparametric  were  two  sample  (Watson, 1961;  1965).  Results  For there  are  a l l b u t one o f t h e s a m p l e s sufficient  data,  d i r e c t i o n s t a k e n by f i s h pools day,  were  the  day  where  i t i s evident  that  the  mean  release  and  home  directions.  the  However, f o r o n l y  show a s i g n i f i c a n t  fish  directions  combined  is  considered  taken  for the f i r s t  only  once,  sample  between sizes.  t h r e e d a y s , i n which  shows  that  the  mean  a r e t h e home d i r e c t i o n s , and f u r t h e r t h a t t h e  samples a r e s i g n i f i c a n t l y may  data  the f i r s t  difference  ( F i g u r e 2 6 ) . T h i s may be due t o t h e s m a l l The  samples  each  l o c a t e d between  does t h e Watson t e s t  samples  each  home  from  different  be s i g n i f i c a n t l y  (Figure  different  26).  i n angular  Since  the  deviation or  Ts m e a n d i r e c t i o n (Q) h o m e d r e c t o n { 0 )? ;  0 = 235° «  Day 1  //  y  w  °° ^ % \\ ooooooo  homepoois (235°) n  ov o°°\  .  /  0  ;  S p o o l s (85°)  O  /•  o°f °V.  Day3  1  8  0  N=4 insufficient data  ^ ^  0  o.••••*.  J  0  0  O  Samples significantly different [mean d r'n] U = . 3 4 8 p < . 0 5 ( B u r r , 1964)  N = 1 2 , n =2 p=.038 <.05  ooo  \  0=0 O  n  90'4«  F270-  N=7, n =2 p = . 4 5 4l> . 0 5  0=0  x ^  Day 2  homepoois (235°)  homepoois (85°) •••••  . •  - 0 = 85°  N = 7 , n =2 l .05 p=.454> J,  S a m p l e s not s i g n i f i c a n t l y d i f f e r ent: °  U 1  ?  4  ' °" ^ \  N = 7  = -0994 > . 0 5  (Burr,  P  ' V  3  N  p=1.0).05 [270° homepoois (235°)  90°+  h o m e p o o i s (85°)  0  4.  -\  o°  fVV  :  ^  =  1964)  3  ' nsuffi c i ent d a t a  0  =  Q  tables a va' l a b l e f or U „  =.2019  7 ^  >  _180°  o^  "cT^X. Day 1* Day 2 • Day 3  / '  P  oo(oooo)  Figure  26  n  0  \  o°°o\ c  o° V  N=9, n =3 P=.508 >.05 !  0 =0  jo.« h o m e p o o i s (235°)  N = 2 2 , n =8 =.286>.05  .......... 1  y* ./  *  j  0= 0  S a m p l e s si g n f ' c a n t l y different: ;  2  ^  U__  22,9  _°  -  >  =.2787 p < . 0 5  •  Stephens,  l  n  ,  c  .  1965)  C i r c u l a r p l o t s of d i r e c t i o n o v e r f i r s t 3 d a y s t a k e n b y t w o s a m p l e s of f i s h d i s p l a c e d to c e n t r a l p o o l ( s u c c e s s f u l h o m e r s s h o w n u n d e r h o m e p o o l s ; t h o s e i n p a r e n t h e s e s found between r e l e a s e and h o m e pools)  189  mean d i r e c t i o n ( B a t s c h e l e t ,  1965)  (although  the  data  suggest  mean d i r e c t i o n ) , 150° was added t o t h e a n g u l a r d i r e c t i o n s o f t h e smaller  sample  b o t h samples) 1962).  This  difference  (so t h a t  and t h e t e s t s t a t i s t i c resulted  between  in U the  between t h e s a m p l e s l i e s  It between three  days  recalculated  by t h e i r  (Watson,  22,9 = .1168, s h o w i n g no s i g n i f i c a n t  2  samples  (p>.05).  Thus t h e d i f f e r e n c e  i n t h e mean d i r e c t i o n .  pool  following  and t h e home release,  t h a t f o r the f i s h  pools  the  mean  within  the  seen first  d i r e c t i o n taken, as  p r e s e n c e i n t i d e p o o l s , by t h e g r o u p s from  side  of the r e l e a s e  mean  directions  pool  was t h e home  taken  by  different.  Of t h e 31 f i s h  pools,  or  24  was  may be c o n c l u d e d , t h e r e f o r e ,  the r e l e a s e  revealed  t h e home d i r e c t i o n s were i d e n t i c a l f o r  77%  each  seen  homed..  direction sample  between  In t o t a l ,  and  were  the  each  that  the  significantly  release  and  80% o f t h e f i s h  home  released  homed.  While t h i s show the  t h a t homing O. m a c u l o s u s home p o o l ,  three  days  remained area  found  31  out  the r e l e a s e  i n the r e l e a s e  display  directed  truly  pool  and 12 f i s h had  landward pools,  been i n p o o l s  lower than  importantly,  while  the  evidence  those  homed.) the fish  uncovered  to  movement t o w a r d s In  the  o f t h e 66 O. M £ 3 l ° § J i § r e l e a s e d  and home p o o l s .  under c o n s i d e r a t i o n  i n  appears t o provide  two g u a l i f i c a t i o n s must be made.  only  f o u n d between  the  experiment  (Another  five  first were fish  w h i c h were n o t s e e n i n Since  no  fish  n o t s e e n may w e l l by  the  tide.  low t i d e d i s t r i b u t i o n o f f i s h  were have Bore  supports  190  the  idea  about  their  random than  of d i r e c t e d  homeward movement, n o t h i n g c a n  behaviour  search  is  involved  were o b s e r v e d  "wrong" d i r e c t i o n  at high t i d e .  might from  number  t h e home  of  (Walbaum) , P r i o n i t u s (Bardach 1967)  Khoo  fins,  o f 0.  1965)  pectoral of  the  sea  results  experiments paired  b a r b e l s and  for  example,  and  Ictalurus  evolans  sp.  body s u r f a c e s .  were  fins.  pelvic  bottom he  fish  i n pools i n the  Urophycis  ( L i n n a e u s ) , P.  several or  of  pool.  fish,  conducted  of  detection  of  found  Given  clues  obtained  conducted  homing fins  (Linnaeus)  the  paired  the  sea  floor,  a s a method  were  inconclusive,  study  involving  determine  i s used  in t h i s  buds)  that  experiments to  chuss  (Bardach, et a l . ,  m a c u l o s u s a r e i n c o n s t a n t c o n t a c t with  (1971)  removal  Since  kind  clues  carolinus  case,  some  h a v e been shown t o p o s s e s s s e n s o r y r e c e p t o r s ( t a s t e  on t h e i r fins  and  concluded  i n homing, g r e a t e r numbers o f  be e x p e c t e d t o be  4.Touch a n d / g r c h e m o s e n s o r y  A  However, i f  be  whether of  involving the  homing. several removal  191  Methods  Three 0.  maculosus  experiments without  c o n d u c t e d . ft t o t a l with  pelvic  and  subseguent  to  fins  as c o n t r o l  the second  third  experiment  both  experiments,  or  pelvic  were  fins  a total fins  a total  of were  transplanted,  65  (In t h i s  u n t r e a t e d 0. m a c u l o s u s a r e r e f e r r e d  e v e n though  pectoral  pelvic fins  pectoral  o f 134 0. m a c u l o s u s  experiment  both  either  t h e homing a b i l i t y  removed and 69 a s u n t r e a t e d c o n t r o l s .  fish,  with  investigating  t h e y were n o t sham o p e r a t e d . ) I n  o f 66 f i s h  removed  were  transplanted,  a n d 33 a s c o n t r o l s .  o f 113 f i s h  In the  were t r a n s p l a n t e d , 38  removed, 37 w i t h b o t h  pectoral  33  with  f i n s r e m o v e d and  38 a s c o n t r o l s . In  a l l three experiments  made a s f a r a s p o s s i b l e about  60  m apart.  reciprocal  between two g r o u p s  Data  were c o l l e c t e d  of  transplants four  pools  were each  f o r f o u r months i n e a c h  experiment. fts a f u r t h e r e x a m i n a t i o n fins  to  detect  examination 0*  chemosensory  (or  was made o f t h e p e c t o r a l  ISLSulosus.  Sections  examined t o d e t e r m i n e receptors.  of the a b i l i t y  stained  touch) and  with  of the  clues, pelvic  paired  histological fins  of s i x  eosin-haemotoxylin  w h e t h e r t h e r e i s any e v i d e n c e  of  were  sensory  192  Results  Compared paired  fins  (Table  The  t r a n s p l a n t area  on  the  are  homing  ability  t o detect  a l l relatively  percentages  t o t h e home p o o l  of  high,  a n y major d i f f e r e n c e s i n  between c o n t r o l and t r e a t e d  fish.  successfully  In a l l  homing  fish  exceeded the percentages r e t u r n i n g t o  home r a n g e .  None o f t h e examined In t h i s  showed  suggested  of  distributed  involved  of  pectoral  interesting  Scorpaenichthys  of  which  carry  to  marmo r a t u s ,  ramus  transplant  provide  no  lateralis  pelvic  note  that,  Freihofer  clues  in  i n t h e homing o f 0. m a c u l o s u s .  from (1963)  accessorius  (the  fibres) are  i ncottids.  and  evidence t o suggest  fins  (Figure 27).  taste or gustatory  experiments  chemosensory o r touch  and  receptors  t o t h e p e c t o r a l and p e l v i c f i n s  The  detecting  i t i s  the f i b r e s of the  branches  studies  sections  any e v i d e n c e o f s e n s o r y  connection,  examination  main  effect  release  i t i s not p o s s i b l e  returning the  little  following  cases  0. m a c u l o s u s , t h e r e m o v a l o f  p e r c e n t a g e s o f 0. m a c u l o s u s r e m a i n i n g i n t h e  percentages s t a y i n g three  untreated  a p p e a r s t o have  34).  although  with  the  histological  that sensory the  paired  receptors fins  are  193  Table Homing  performance a f t e r  Pect Date r e l e a s e d  Cont  21 September 1975  34 removal of paired  Pelv  Cont  fins  Pect Pelv  4 September 1975  Cont  16 May 1976  Number r e l e a s e d  33  33  65  69  38  37  38  Number homed P e r c e n t homed  23 70  24 73  41 63  52 75  21 55  18 49  20 53  4  19  17  10  12  29  25  26  14  21  Number s t a y i n g in transplant area Percent s t a y i n g in transplant area  Days s t a y i n g i n t r a n s p l a n t area a . homers 13  b . non-homers  Percent of homers returning to home p o o l  2,2,2  65  3,78  67  1,1,1, 1,1,1, 1,5,5, 1,1,4, 6,7 6,6,7, 17,18, 1,4, 6,6, 6,6,7, 22,43, 12,32, 13,17 42,43, 49,50 88  73  1,1,2  2,27  1,1, 1,2, 1,2, 2,56, 27, 56 85, 123  2,2 73,73, 83, 110  81  P e c t : b o t h p e c t o r a l f i n s removed P e l v : b o t h p e l v i c f i n s removed Cont: untreated  61  80  gure 27  S e c t i o n s o f t i p s o f p e c t o r a l f i n (above) p e l v i c f i n (below)  and  195  5.Relative  importance of d i f f e r e n t  To  determine the r e l a t i v e  i m p a i r m e n t s on homing which 0.  homing  maculosus,  each group  involving  experiment  was  significantly  impairment  was  was  compared was  to  sensory  there  a  different This  which  t r e a t m e n t s was  had  redundancy  compared  undergone with t h a t  was  treatment. in  i n homing, t h e homing p e r f o r m a n c e  maculosus  of  performance  one  is  in  groups  receptors.  homing  by any  sensory  conducted  between  subjected  of  affected  whether  involved  of d i f f e r e n t  experiment  destruction  determine  0.  effect  d e s i g n e d t o show whether  systems of  an  o f which  more a d v e r s e l y  To  group  ability  performance  treatment  sensory  senses  the of  a  a l l four sensory  of u n t r e a t e d  fish.  Methods  In were  the f i r s t  reciprocally  e a c h a b o u t 60 were  involved  anosmic, 100  one  experiment, a t o t a l  transplanted  m apart.  Five  having  been  set of paired  between two  egual sized treated  of  150  0.  maculosus  groups of f o u r  groups of  as f o l l o w s ;  f i n s removed, immersion  0.  pools  maculosus  blinded, in  made  Triton-X-  f o r s e v e n d a y s and u n t r e a t e d .  Ten laboratory  blind  fish  and  f i v e anosmic  to i n v e s t i g a t e the effect  fish  were h e l d  of treatment  on  i n the  survival.  196  Data  were c o l l e c t e d  In impairment  the  m  experiment  apart.  fish  release. treated  examining  simultaneous  A group  were  o f 20 f i s h  kept  impairments.  over  t h e same p e r i o d  two  and  half  experiments,  as t h e e x p e r i m e n t .  months.  In  o f pools  o f independence  about  period.  Another  to investigate Data  these  treatment,  f o r s e v e r a l days b e f o r e  and  ten  survival  were c o l l e c t e d f o r the  two  t e s t s o f s i g n i f i c a n c e were c o n d u c t e d  factor G test  0. m a c u l o s u s  Following  occurred i n t h i s  were k e p t i n t h e l a b o r a t o r y  a  46  sensory  were u n t r e a t e d c o n t r o l s a n d 26  i n the laboratory  One m o r t a l i t y fish  of  t r a n s p l a n t e d between two g r o u p s  were s u b j e c t e d t o a l l f o u r the  months.  o f t h e same f o u r s e n s e s , a t o t a l  were r e c i p r o c a l l y 60  f o r t h r e e and a h a l f  ( S o k a l and R o h l f ,  subsequent  using the  two  1969).  Results  Given  the  low  percentage  s u c c e s s f u l l y homed i t a p p e a r s treatment  effect  s u c c e s s f u l l y homing 0. m a c u l o s u s  (p>.05)  differences are not s i g n i f i c a n t  successfully for  any  blind  suggesting  homing f i s h and  that  anosmic  groups  with  fish  sensory  on  the  (Table  which  impairment  percentage o f  35).  Although  (p>.05), t h e p e r c e n t a g e s o f  t o t h e home than  t h e r e may be a s l i g h t  on t h e p r e c i s i o n  The  returning  untreated  no s i n g l e  significant  the  has  that  of  pool  are  f o r any o t h e r effect  o f these  lower  treatment treatments  which t h e home p o o l c a n be l o c a t e d .  percentages  of  0. m a c u l o s u s  remaining  i n the  T a b l e 35 R e l a t i v e homing performance a f t e r  four  d i f f e r e n t impairment treatments  B l i n d Anosmic F i n l e s s * 7 days i n C o n t r o l Triton-X -100 Date r e l e a s e d  23 August 1976  Number r e l e a s e d  30  29  30  30  31  Number homed Percent homed  21 70  12 41  15 50  22 73  15 48  30  20  26  Number remaining in transplant ar ea Percent remaining in transplant area  10 17  Days s t a y i n g i n t r a n s p l a n t area a. homers 1,1,2 b. non-homers  Percent o f homers homing t o home pool  34  1  1,11  24,31 1,2,2,2, 1,1,2, 11,49,73, 8,24, 74, 104 74, 101 48  42  1,1,1,1, 11,24  53  *: one s e t of p a i r e d f i n s removed  59  2 1,2,10, 11,46, 80, 104 60  198  transplant  area  are  high  particularly  anosmic, f i n l e s s and c o n t r o l groups. group  shows an e g u a l l y  transplant the  in  The f a c t that the  groups suggests l i t t l e  treatments themselves i n t h i s r e g a r d .  f i s h held i n the l a b o r a t o r y system  months.  Until  Mortality  cases o f the control  high percentage of f i s h remaining i n the  area as the t r e a t e d  seawater  the  caused then  was  low, u n t i l  total  mortality  no  i n the f i e l d  anosmic  was  and  effect  of  M o r t a l i t y of t r e a t e d a  blockage  a f t e r two  one  blind  in  the  and a h a l f fish  died.  unknown, although b l i n d , anosmic and  c o n t r o l f i s h were observed i n the f i e l d u n t i l the experiment  was  terminated. From t h i s experiment be  shown  no s i n g l e sensory impairment  t o have a s i g n i f i c a n t l y more adverse e f f e c t on homing  a b i l i t y compared  with t h a t o f u n t r e a t e d c o n t r o l s .  D e s p i t e the s m a l l numbers of c o n t r o l f i s h simultaneous  impairment  homing performance the  treatments  experiment,  of t r e a t e d  (Table  it  is  used i n  evident  the  f i s h i s substantially affected  by  36). , The d i f f e r e n c e i n the percentages Two  a combination of both may  Homing a b i l i t y  been  account  impaired because  destroyed so t h a t no system fish  which  successfully  mechanisms or complete may  the  that  s u c c e s s f u l l y homing i s s i g n i f i c a n t (p<. 05).  have  can  may  a l t e r n a t i v e sensory mechanisms were was a v a i l a b l e f o r homing. ,  homed  destruction  not have been e f f e c t e d .  for t h i s .  e x p l a n a t i o n s or  may  have  The  two  employed a d d i t i o n a l  of one of the sensory systems  T a b l e 36  Homing simultaneous  performance a f t e r sensory  impairment  Impaired Date r e l e a s e d Number r e l e a s e d  four treatments  Control  15 September 1976 26  20  Number homed P e r c e n t homed  2 8  14 70  Number r e m a i n i n g i n t r a n s p l a n t area Percent remaining i n t r a n s p l a n t area  4  4  15  20  Days s t a y i n g i n transplant area a. homers b. non-homers  P e r c e n t o f homers r e t u r n i n g t o home p o o l  10,10, 23,28 50  1,1 10,26  64  200  The  alternative  explanation  may  be  that  treated  0. m a c u l o s u s were u n a b l e t o s u r v i v e , r e g a r d l e s s o f b e i n g home,  after  percentage  such  area,  treatment.  (p<.05) o f t r e a t e d f i s h  time d u r i n g suggests  drastic  the experiment  were  predators  subject of  to  The s i g n i f i c a n t l y  of f i s h  heavy  predation  or  p . m a c u l o s u s a r e unknown, b u t p o s s i b l e  candidates  o f mink o b s e r v e d  However, t h e m a j o r i t y crabs,  and  gut  hexagrammids and i n s h o r e Artedius (Bruce  spp. a r e  Leaman, D a v i d  observation). that  is  Gut  only  as  parasitic  tidepools unlikely the  Although  fish  tidepool  in this  study  were  cottids  suggests  that  fish  predators  personal  but  never  Sebastes  spp.,  perhaps  personal  of diving birds  guillemots are  suggests  probably  t o e a t p . m a c u l o s u s and i f t h e y catch gulls  host  (Ian are  lingua)  (Ching,  suggests  Robertson,  the  final  that  only  predation of t h i s  O.  do, i t  for  fish fish,  impairment  a  maculosus  observations  kind i s very severe.  which had u n d e r g o n e f o u r s e n s o r y  the  personal  host  f o r which  1978) ,  only  of p . m a g u l o s u s  communications;  analysis  {Cryptocotyle  areas  i n tidepools,  i n the i n t e r t i d a l  may be v u l n e r a b l e and even f o r t h e s e that  and  of  content  incidental  trematode  in  birds  analysis  Zittin,  a c t as an i n t e r m e d i a t e  gulls  diving  content  major  diving birds likely  communication).  can  the  areas  p e l a g i c c o r m o r a n t s and p i g e o n  only  snakes,  no g a r t e r s n a k e s were o b s e r v e d  c r o w s were s e e n a t t h e s t u d y pool,  moved o u t o f t h e Specific  fish.  a  controls)  died.  garter  in  20% f o r  either  i n c l u d e mink, g u l l s , c r o w s ,  collecting  higher  w h i c h were n o t l o c a t e d a t any  {77% compared w i t h  that that the majority  able to  in i t  of high  seems  Certainly, treatments  201  appeared  more v u l n e r a b l e t o  tidepools.  Known  laboratory  of  treated  fish  untreated tank  the  tank.  system  their  treated  after  behaviour  fish  held  a n d one  two months.  The moved  former  about  exhibited  tended  until  Behaviour of from  that  of  t o group i n c o r n e r s o f t h e  i n comparison  no i n t e r e s t  in  i n the  escape)  i n t h e l a b was n o t i c e a b l y d i f f e r e n t  fish.  fish  of  by  low (one d e a t h  the seawater  and r a r e l y  Treated  mortality  was r e l a t i v e l y  failure  predation  with  i n mussel  T h i s c a n be c o n t r a s t e d w i t h  u n t r e a t e d 0. m a c u l o s u s a n d t h e a l m o s t  untreated  fish.  p i e c e s dropped  into  the r e l a t i v e a c t i v i t y  instant  response  of  t o mussel  pieces.  When t r e a t e d First with  Beach  their  initial  untreated f i s h ,  movements.  fish  release  point to shelter  stones  or  pool o c c a s i o n a l l y a the the  solid  open  fish area  fish  immediately  underneath  Treated  fish  or at merely  of  these  i s  the  of the pool into  four  on  definitely  the  which they  their  from of  the  rocks,  I f they i t .  By  swam  into  comparison  were r e l e a s e d o r t o  movements.  while the cumulative  impairment survival  impaired.  in  and f o r t h e most p a r t s a t i n  therefore that  sensory  compared  away side  at  s a t i n t h e open i n t h e  b e s i d e o r under  moved v e r y l i t t l e  considerable effect ability  swam  swimming back a n d f o r w a r d .  appears  tidepools  and u n s t e a d i n e s s  w h i c h t h e y swam d u r i n g t h e i r e r r a t i c  It  into  swimming was h i g h l y e r r a t i c  o b j e c t they remained  treated  released  showing r o l l i n g  Untreated  algae.  were  treatments of  may  O. m a c u l o s u s ,  The r e s u l t s  of other  effect have  a  homing sensory  202  impairment  homing e x p e r i m e n t s s u g g e s t t h a t  of  blindness  the  homing p e r f o r m a n c e  seven  days  and  anosmia  which  of fish  immersion  i tis  a  combination  t o g e t h e r p r e v e n t homing,  without  in Triton-X-100  paired  fins  or  since after  a p p e a r s t o be r e l a t i v e l y  unaffected.  §.•Simultaneous  impairment  To e x p l o r e smell are involved  further  the effect  on homing  performance.  a group  o f both  effect  with  a  blind  to  particular,  vision,  to  If  home.  and a n o s m i c  anosmic  o f such  view  fish  v i s i o n and  homing  fish.  t r e a t m e n t s on j u v e n i l e determining a r e used  singly  whether  these  that  of  with that o f experiment  was i n v e s t i g a t e d senses,  to a c g u i r e t h e i n f o r m a t i o n  impaired adult  or e x p e r i e n c e i s a v a i l a b l e  performance  In t h i s  fish  to  of t h e s e senses  was compared  and u n t r e a t e d  j u v e n i l e s cannot, i t i s p o s s i b l e  to  both  o f s i m u l t a n e o u s impairment  one e x p e r i m e n t , t h e r e l a t i v e  groups o f b l i n d , the  the suggestion that  i n homing, two e x p e r i m e n t s were c o n d u c t e d  determine  In  o f v i s i o n and s m e l l  p . maculosus  and  in  reguired  c a n home, b u t  insufficient  t o them t o u s e a l t e r n a t i v e  information mechanisms  home.  In fish  anosmic  the second was  simultaneously blind  used  experiment, a d i f f e r e n t to  compare  and a n o s m i c  fish  method  t h e homing with  that  o f making  performance o f of  untreated  203  controls.  Only a d u l t f i s h were used i n t h i s e x p e r i m e n t .  ;  Methods  A  total  of  180  0. maculosus  were  reciprocally  t r a n s p l a n t e d between two groups o f p o o l s about 6 0 m a p a r t . , group  of 45 f i s h were u n t r e a t e d c o n t r o l s , 46 f i s h were b l i n d e d ,  44 were made anosmic group  One  of  45  using Kodak  Eastman  910  Adhesive  f i s h were made b l i n d and anosmic.  and  In each group,  about 25% of the f i s h were l e s s than 4.0 cm, t o i n v e s t i g a t e e f f e c t o f the impairment treatments on j u v e n i l e s . 0. maculosus  a  the  A group o f 10  which were both b l i n d and anosmic were kept i n the  l a b o r a t o r y t o examine s u r v i v a l .  Data  were  collected  f o r two  months. In reciprocally approximately  t h e second  experiment,  a  between  two  transplanted 60  m  apart.  t o t a l o f 72 f i s h were groups  of  pools  One group o f 32 f i s h c o n s i s t e d o f  untreated c o n t r o l s and t h e o t h e r group  of  40  fish  were  made  b l i n d and anosmic, t h e l a t t e r by b l o c k i n g the nares with Durelon dental  glue.  A  were kept i n the weeks.  group o f 10 f i s h which were b l i n d and anosmic laboratory.  Data  were  collected  f o r five  204  Results  In  the  was e f f e c t e d it  first  by b l o c k i n g  was e v i d e n t t h a t  two  days  in  the  the  experiment  t h e n a r e s w i t h Kodak Eastman  field.  condition  persisted  ( T a b l e 38) D u r e I o n i t  considerably  longer period  blind  unusual  Thus  i t cannot  laboratory  the  and  ( T a b l e 37) i n which  remained  anosmic  while the f i s h be s a i d after  dental in  than fish  were  release. was  around  In t h i s  manner  i s  remained  on t h e  although  some o f t h e D u r e l o n  the period  that In and  of  fish  longer  results  fish  for a  into tidepools  exhibited  several  than  succeeded  three  be s t a t e d  applications of i t  days.  inside  w i t h any  suggest  Thus  again,  the nares,  certainty.  that  simultaneous compared  with  or "anosmic"  fish.  the percentage o f s u c c e s s f u l l y was s i g n i f i c a n t l y  in  I t was e v i d e n t , s i n c e t h e  may have r e m a i n e d  certainly  t h e g r a v e l and  fish  u n t r e a t e d c o n t r o l s and e i t h e r b l i n d  anosmic  nares  the  However  and " a n o s m i a " i m p a i r homing p e r f o r m a n c e  both experiments  In  adhesive.  none o f t h e e x t e r i o r  of anosmia cannot  The blinding  that  how  t o w a r d s t h e s u r f a c e and back  knocking t h e c o v e r i n g s o f f t h e nares. Durelon  certainly  used..  the  t h e Eastman released  one o r  In the second  towards t h e bottom o f t h e p o o l "nose d i v i n g " i n t o  pink,  adhesive,  with any c e r t a i n t y  glue  and  b e h a v i o u r , swimming r a p i d l y  rocks i n the pool.  anosmia  t h e a d h e s i v e came o f f t h e n a r e s a f t e r  a n o s m i c when r e l e a s e d , long  experiment  lower than  homing  blind  the percentages  fable  R e l a t i v e importance  Blind  Date r e l e a s e d  37  o f b l i n d n e s s and a n o s m i a  Anosmic  16 J u l y  B l i n d and anosmic  Control  1977  Number r e l e a s e d  46  44  45  45  Number homed P e r c e n t homed  23 50  18 41  9 20  19 42  Number f o u n d i n t r a n s p l a n t area Percent found i n t r a n s p l a n t area  28  27  20  26  61  61  44  58  1,2,2,5, 9,10  2,2,2,2, 5,42  1,1,2,5, 9,10,11, 22,26 ,26 , 27, 59,59, 59  1,1,1,2, 2,5,6,6, 6,13,14, 14, 14,27, 27,32,55, 55,59,59  Days s p e n t i n t r a n s p l a n t area a. homers b.  non-homers  1,1,2,2, 2,6,9, 18,35 1,1,1,1, 2.2.2.5, 5.6.6.6, 10,10, 14,18, 32,36,42  1,1,1,1, 1,2,5,6, 6,6,14 1,1,1,1, 2,2,2,5, 5,10,42, 45,45,59, 59,59,  P e r c e n t o f homers homed t o home p o o l  22  39  44  42  Percent o f f i s h seen a f t e r r e l e a s e  91  77  51  87  T a b l e 38 Homing p e r f o r m a n c e a f t e r simultaneous  b l i n d i n g and a n o s m i a  Impaired Date  released  Number  released  Control  9 August  1977  40  32  Number homed P e r c e n t homed  8 20  20 63  Number r e m a i n i n g i n t r a n s p l a n t area Percent remaining i n t r a n s p l a n t area  18  14  45  44  Days s t a y i n g i n transplant area a. homers b.  non-homers  3,3,3*3, 3,3,8, 18 9,14,17 1 ,2,2,2, 1,3,3,3, 3,3,3,14, 8,8,11, 14,14,20 11,18,30  P e r c e n t o f homers r e t u r n i n g t o home p o o l  38  60  Percent of f i s h seen a f t e r r e l e a s e  48  94  207  homing i n t h e o t h e r g r o u p s e m p l o y e d However  the r e s u l t s  i n each e x p e r i m e n t  must be t r e a t e d  uncertainty  concerning  percentages  of  fish  the  with c a u t i o n because o f t h e  anosmic  fish  and  because  the  seen a f t e r r e l e a s e a r e s i g n i f i c a n t l y  lower  (p<.05) f o r t h e s i m u l t a n e o u s l y b l i n d  and anosmic  48%)  t h a n f o r any o t h e r g r o u p s  to  this  t r e a t m e n t may have some i m p a c t  Consideration impaired  (p<.05).  of  {77%  on  94%),  groups  suggesting  the  t h e t i m e t o home i n t h e d i f f e r e n t l y  groups i n t h e f i r s t  61%,  transplant  blind  e x p e r i m e n t showed l i t t l e  a r e a do - c o n t r o l :  and a n o s m i c :  67%.  t a k e n t o home was i n g e n e r a l anosmic did  difference  20%  and  32%, b l i n d :  considerably  This  home u n t i l  for  fish  homed.  homing c o n t r o l f i s h  might s u g g e s t t h a t  the dental  39%, a n o s m i c :  blind  60% o f t h e c o n t r o l s which  anosmic  of the successfully  longer  t h e t r a n s p l a n t a r e a w h i l e 88% o f t h e b l i n d  were.  observed  In t h e second experiment, the time  than f o r c o n t r o l s ;  so b e f o r e any b l i n d  only in  fish  that  survival.  between t r e a t m e n t s , a l t h o u g h t h e p e r c e n t a g e s o f homers in  ( 5 1 % and  t h e anosmic  g l u e was removed  and fish  and homed  Furthermore were o b s e r v e d anosmic  fish  were u n a b l e t o  from t h e n a r e s ,  and  the  s e n s e o f s m e l l was a g a i n p r e s e n t .  The first fish  experiment  reguire  impairment  to  of  homing p e r f o r m a n c e  by l e n g t h  ( T a b l e 39) a p p e a r s t o show t h a t  a r e 3,5 cm l o n g  information not  analysis  use  vision.  some  of  them  have  by t h e t i m e t h e  acguired  sufficient  a l t e r n a t e s e n s e s t o home; a t l e a s t Until  that  size  a p p e a r s t o be s u f f i c i e n t  i s  reached  i n the  t h e y do  however,  t o p r e v e n t homing.  any  •Table  Relative  homing  performance  following  Length  Eate  class  (cn)  2. 5-2.9  3.0-3.4  released  Elind Number r e l e a s e d Number homed Percent hcmed  (percentage  blindness  s u c c e s s f u l l y homing)  and a n o s m i a  3. 5-3. 9  16  July  4.0-4.9  by  length  5.0-5.9  6.0-6.9  7.0-7.9  Toti  197 7  0  5 0 0  il 2 50  25 13 52  7 7 100  1 1 100  Anosmic Number r e l e a s e d Number homed Percent hcmed  5 0 0  2 0 0  3 0 0  21 12 57  12 6 50  1 0 0  0  44 18 41  E l i n d and Anosmic Nuirber released Number hcmed Percent hcmed  4 0 0  3 0 0  3 0 0  21 3 14  1 1 4 36  3 2 67  0  45 9 20  4 1 25  2 0 0  5 2 10  24 9 38  8 6 75  2 1 50  0  45 19 42  Control Number r e l e a s e d Number homed Percent homed  3  39  0  1  0 0  46 23 50  Table  Relative  homing  performance  following  Length  Date  class  (cm)  2.5-2.9  3.0-3.4  released  39  (continued)  (percentage  blindness  and a n o s m i a ,  3.5-3.9  16  remaining  July  4.0-4.9  i n transplant  area)  by l e n g t h  5.6-5.9  6.0-6.9  7.0-7.9  Total  197 7  Blind Number r e l e a s e d Number s t a y e d Percent stayed  3 3 100  5 3 60  4 3 75  25 15 60  7 3 43  1 0 0  1 1 100  46 27 59  Anosmic Nuirber r e l e a s e d Number s t a y e d Percent stayed  5 3 60  2 1 50  3 1 33  21 14 67  12 8 67  1 0 0  0  44 27 61  E l i n d and Anosmic Nuirber r e l e a s e d Number s t a y e d Percent stayed  4 1 25  3 0 0  3 2 67  21 11 52  11 7 64  3 1 33  0  45 20 44  Control Nunber r e l e a s e d Number s t a y e d Percent stayed  4 3 75  2 1 50  5 4 80  24 15 63  8 5 25  2 1 50  0  45 26 58  O 1£>  210  Laboratory anosmic far  fish  less  after  was  survival  good  active  untreated  a d h e s i v e and  nares, the f i s h  would  they would  desultorily  fish  feed  began  treated  to  fish  7. V i s u a l  in  after  recognition  was  investigated.  the  olfactory was  t i m e , the D u r e l o n  pools  by  landmarks  appear t o be  treated  home  to  determine  the pool  landmark.  To  determine water  whether  from  m.  two  of c a p t u r e  kinds  O,  the  a choice  of and  maculosus  or to the  0.maculosus  the tidepool  homing  i n tidepools  homing  investigate  tidepool  whether  "landmarks",  whether  involved  clues i n  were i n s t a l l e d  f r o m s e a w a t e r pumped i n t o  a d e p t h o f 20  though  clues  c l u e s e m a n a t i n g from t h e t i d e p o o l to  Even  weeks and t h e Eastman a d h e s i v e  To  determine  familiar  distinguish  captured from  to  would  conducted  could  this  o f v i s u a l and o l f a c t o r y  recognize  conspicuous a r t i f i c i a l  with  the  and o l f a c t i o n  role  individuals  were  s i x weeks.  both v i s i o n  exchanged  fish.  fish  and  t h e d e n t a l g l u e had come o f f  four  the  then  blind  days  after  homing,  0. m a c u l o s u s  of  U n t i l about seven  and o l f a c t o r y , t i d e p o o l  Since  groups  not respond to mussel p i e c e s .  die  after  both  up t o f o u r weeks, a l t h o u g h t h e  than  t h e Eastman  of  role  pool of  experiment individuals  i n which t h e y were  the Bamfield  Marine  Station  211  Methods  To similar  tidepools  difficulty the  investigate  at F i r s t  of f i n d i n g  pools  v i s u a l landmarks, Beach  were s e l e c t e d .  similar tidepools  selected  were  high  two p a i r s o f somewhat  low i n  relatively  with  cod  each p a i r  of  pools  landmarks  consisted  o r a n g e and i n v e r t e d After p.  line  pair  pans p a i n t e d  cm)  collected.  being  which  the f i s h  number  of small  I t was  would be more r e a d i l y i n f l u e n c e d 154  fish  from t h e f o u r  m away, 11 d a y s a f t e r Prior  to  high  exchanged. unchanged  tide,  to provide  almost  two  the  that  pool,  so t h a t The  flourescent 28).  f o r nine  days,  of the pools  i f  the  fish.  were  to fish  total  of  i n t o a pool about initially  from  other  A  4.0  pool  by l a n d m a r k s , j u v e n i l e  one p o o l  pool  d a t a on p o o l f i d e l i t y .  months.  was  p . maculogj|s- ( l e s s t h a n  landmarks  The l a n d m a r k s i n  landmark  The h e i g h t  p o o l s were r e l e a s e d  the  Two  (Figure  than a d u l t  the landmarks  pools.  landmarks.  installed  anticipated  homed was i n f l u e n c e d  intertidal,  painted  green  m a c u l o s u s were c o l l e c t e d and t a g g e d . i n a large  of  bricks  t h e l a n d m a r k s had been  resulted  for  one  fireplace  dish  one  and cement n a i l s i n e a c h  contained of  the  shallow  p a i r s o f c o n s p i c u o u s l a n d m a r k s were made and anchored  Because o f t h e  ;  pair  65  installed. pair were  were left  D a t a were c o l l e c t e d  P e r c e n t a g e s homing t o e a c h  pool  were  calculated.  To  investigate  olfactory clues,  t e n p. m a c u l o s u s  were  212  213  collected several water  from F i r s t months.  Beach a t v a r y i n g i n t e r v a l s  At  the  from t h e c a p t u r e  lab.  Testing  of  r e t u r n i n g to the  time a f i s h  p o o l was  each  was  collected  fish  began  over  the  and  returned  within  seawater  of  was  of  p l e x i g l a s s and  exterior  run  through  the  inlets  Burettes  control  ( s t a t i o n ) seawater i n t o  in  40 t o  50  ml  Khoo's  were  disturbance  arms.  of  per  was  placed  a holding  series  f o r two  of  20  or  seguence; h a l f trials. trials  A  40  test  break  to permit  run The  both  of  burette  throughout  the  out  burette  through test  The with  29).  ends  test  than  and  outer  12.7  of  a sliding  choice  trials  choice  a  rate flow  o f the  non-  allowed run  i n the  trials,  to  as  a  following half  allowed  test  between  tank. station  seawater  home t i d e p o o l w a t e r  water  fish.  Station  r a t e of  were  through  t i d e p o o l water was  for a l l  reduce  gate.  Tests  control  Khoo  (tidepool) or  cm  t e n m i n u t e s was  station  which  trials  test  to  the  the  control trials  of the  by  of the tank at  c o n s i s t e d of running In  of  h o l d i n g chamber and  all  at l e a s t  flushing  (Figure  higher  testing.  and  trials,  burettes.  t h r o u g h one  i n the  test  Control t r i a l s through  e a c h arm  chamber  hours before  black  so as t o d r i p  experiments.  F i s h were p l a c e d acclimate  the  hours  used  painted  at the  minute, s l i g h t l y  (1971)  c h o i c e arm  4 1 of to  several  of  lab.  constructed  effect  period  caught, about  A Y-shaped c h o i c e t a n k s i m i l a r t o t h a t (1971) was  a  In  run each  was  the  other.  was  changed  trial  the  F i g u r e 29  Y shaped choice tank  IS) —\  4s-  215  appropriate  water  was  run  r a i s e d t o allow the f i s h  through  the b u r e t t e s and the gate  to swim out and make a choice.  f i r s t s i x f i s h t e s t e d each t r i a l was continued made  a  choice e i t h e r r i g h t or l e f t .  consuming t h a t f o r terminated  after  recorded. tagged  t h e remaining a  period  On completion  and  released  of  until  the  60  fish  T h i s proved t o be so time  four  fish  each  trial  was  30 minutes and *no c h o i c e ' was  of the t e s t i n g seven of the  about  For t h e  fish  were  m away from t h e i r home pool t o  determine whether they were a b l e t o home. The control  proportions of l e f t  t r i a l s were c a l c u l a t e d and compared with the r e s p e c t i v e  proportions of c o r r e c t trials.  ft  comparison  left  and  right  choices  i n the  of  containing  choices  home  tidepool  water. .,  1  denoted  that  the  proportion  A ' p o s i t i v e * denoted t h a t the  p r o p o r t i o n of c o r r e c t t e s t c h o i c e s was greater than choices  in  that arm.  Even i f a f i s h  the  number  being t e s t e d  showed a b i a s towards a p a r t i c u l a r arm i n the c o n t r o l preference  of  i n t h a t arm was greater than t h e p r o p o r t i o n o f  c o r r e c t t e s t c h o i c e s i n t h a t arm.  control  In t h e  c o n t r o l and c o r r e c t t e s t choices f o r each arm o f  the choice tank, a ' n e g a t i v e  of  test  c o r r e c t c h o i c e i n a t e s t t r i a l was one i n which t h e  f i s h chose the arm  control  and r i g h t c h o i c e s made i n the  tests,  a  f o r home t i d e p o o l water would be shown as a g r e a t e r  p r o p o r t i o n o f t e s t c h o i c e s i n t h a t arm.  216  Results  Table (Pair 0*  1)  40  had  clearly  no  effect  returned the  small  t o the  majority  moving  to  "control' 0.  the pools  1  to  in  pair.  one  Hhile was  evident  capture  pools,  r e l e a s e and pools.  displayed  alternate (Pair  2)  first  of  support  i n the  that they  moved t o  pools  apparently  experiment  the  landmarks  this  may  be  were  landmarks c o u l d  a  of  to the  installed. also  conspicuous t o the  be  of  two  fish  data  from  the  transience  of  to the  were s e e n  home,  capture,  tidepools.  No  fish  other  pool  after  release,  i n the d i r e c t i o n  resident  e x p e r i m e n t may pools,  of  the  from  these  been  more  failure  period  for  i s not  as  meaning  which  that  so c o n s p i c u o u s  is of  apparent l a c k of e f f e c t  interpreted  human eye  have  total  brevity of the The  fish  the  where p o o l f i d e l i t y  g r e a t e r , however, the  due  the  infreguently  The  few  Of  amongst p o o l s i n between  more t h a n t h e  conducted i n lower  considerably  the  landmark  pools  about  high  only  fish  capture  if  these  the  did  pool  although  the evidence  the tagged  moved  f i s h which  pair.  of  pools.  w h i c h homed, o n l y  most o f  and  p a i r of  pools,  p a i r and  landmarks  behaviour  f i d e l i t y t o the  pool  p o o l i n the  exchange of  Of t h o s e  little  range  the  homing  fish  a l t e r n a t e pool.  I t i s possible that  successful  the  the  m a c u l o s u s i n d i v i d u a l s from  homed the  the  percentages of  moving between a wide  it  on  l§£2l£>sus i n d i v i d u a l s from  relatively  the  shows t h a t  to  of a a  Table Homing p e r f o r m a n c e  after  40  p o o l "landmarks"  Pair 1 I n i t i a l placement Brick  Date  released  Number r e l e a s e d Number homed t o "brick" P e r c e n t homed t o "brick" Number homed t o "pan" P e r c e n t homed t o "pan" Number s t a y i n g i n t r a n s p l a n t area Percent staying i n t r a n s p l a n t area Number f o u n d i n distant pools Percent found i n distant pools  Pair 2 Initial S final placement  Pan  20 J u l y  exchanged  Brick  Pan  1977  42  37  21  54  19  0  0  0  45  0  0  0  2  16  0  18  5  43  0  33  18  9  8  33  43  24  38  61  2  0  6  1  5  0  29  2  218  fish a  i n a tidepool or that  conspicuous v i s u a l In  the  the pool  choice  over s t a t i o n  tank  experiments,  T h i s i s n o t shown  plus  minus  0»  maculosus  pumped i n t o  signs  individuals the marine  These  no  distinguish  results  tidepool.  factor This  possible  would  in  with  presumably i fthe  eliminate  the  are  very  olfactory  back  to t h e i r  olfaction  (when  the  may  alter  their  fish  a  home as  I f olfactory  a  clues are  nature. ,  or  significance  substances  to the f i s h . ,  producing  olfactory  nature, t h e time d e l a y s i n v o l v e d the fish  might  of the substance being  these  not  from t i d e p o o l water t a k e n a t low  results  conspicuous landmarks n o r o l f a c t o r y tidepool  fish  tide  water  t h e w a t e r and t e s t i n g  although  water  water a t low t i d e a r e  high  clues  substance  any c h a n c e  of  to suggest that  home p o o l w a t e r f r o m  need n o t be o f g r e a t  clues are of a v o l a t i l e collecting  at  other  Recognition of olfactory  Further,  evidence  does n o t n e c e s s a r i l y e l i m i n a t e  mixing  distribution  appear t o s u g g e s t t h a t  guiding  emanate from t h e t i d e p o o l  tide  The  homing mechanism, f o r two r e a s o n s .  homing)  maculosus  station.  c l u e s e m a n a t i n g from home t i d e p o o l significant  0.  o f p l u s s i g n s would be  (Table 41),  provides  i f  p r e f e r e n c e f o r home t i d e p o o l  water, a predominance  expected. and  r e c o g n i z e d by  landmark.  i n d i v i d u a l s demonstrated a d i s t i n c t water  i s not s o l e l y  important  be s u f f i c i e n t t o  detected.  suggest clues  between  that  emanating  neither from  the  i n t h e homing of 0. m a c u l o s u s , a  219  T a b l e 41 Choice  Fish Control choices percent L  tank t e s t s  Test choices percent correct L  1  60  50  2  90  70  3  80  4«*  o f arm  preference  Sign C o n t r o l Test Sign Total choices choices number o f percent percent trials R correct R  40  50  *  40  -  10  30  •  40  60  -  20  0  70  100  +  30  60  +  20  5'*  80  40  -  20  100  •  20  6*  90  60  -  10  80  •  20  7*  10  20  •  0  20  •  20  8«  0  20  +  0  40  +  20  9'  30  20  -  60  0  -  20  10  0  -  10  0  -  20  10»  -:  percent percent  c o n t r o l choices > percent c o r r e c t c o n t r o l choices < percent c o r r e c t ': t a g g e d and r e l e a s e d *: homed  20  test test  choices choices  220  s m a l l amount o f e v i d e n c e  was o b t a i n e d  from t h e b e h a v i o u r  recognizable  a d u l t 0. m a c u l o s u s which had t a k e n  the  i n Grappler  Inlet,  of the pool  itself  enclosure  a t t r i b u t e (s) than  the position  fish  was  regularly home  moved found  pool  of the pool. to  pools  the  pool  i t returned  displacement  of  the  in  to  fish  i t i s  some  homes, r a t h e r when  the  t h a n t h e one i n w h i c h i t was i t had a l w a y s r e t u r n e d t o t h e  f o l l o w i n g low t i d e .  exchanged with another  that  On t h e t h r e e o c c a s i o n s  other  one  up r e s i d e n c e i n  t o which t h e f i s h  i n the enclosure,  by  displaced,  suggesting  of  the  the and  When t h e home p o o l was  enclosure,  and  the  r e l o c a t e d home p o o l . pools  resulted  in  fish  Further the  fish  behavioural responses  were  disappearing.  8.Discussion  A  number  of  examined t o d e t e r m i n e home r a n g e .  and  substance no  effect  resident transplant  clues  was f o u n d  detected  immersion  which  was d e s i g n e d  introduced  to  show  that  by t h e p a i r e d f i n s  f o r seven  on homing a b i l i t y . and  and  how 0. m a c u l o s u s f i n d s i t s way back t o i t s  No e v i d e n c e  chemosensory homing  senses  days  t o destroy Behavioural  fish  and  in  a  touch  are involved i n detergent-like  chemosensory  c e l l s had  interactions  "space"  and/or  between  or density i n the  p o o l do n o t a p p e a r t o be i m p l i c a t e d i n homing.  Evidence  that  homing  i s  a  directed  movement  was  221  suggested  by  Comparison  of  percentages travelled  lines,  low  homing the  described movement.  pattern  the  direct by  With  over  a  exploration  travelling  about  Khoo significantly fish  at Port  anosmic  direct  fish  being  fish.  Unilaterally  control  fish  it  faithful  does  although  fish.  percentages  more  be  straight  rectangular  travelled.  one  turn  blind  or  anosmic  success  successful  anosmic  These  this  fish  results  as o l f a c t i o n  were control  blind  f o rcontrol  or  fish,  than  blind  homed a s w e l l a s  suggested  t o Khoo  that  home  range,  generally support  Khoo's  i n homing.  of  olfaction,  for First  blindness  than c o n t r o l  fish  of  i n maintaining  study  the importance  reguires  home r a n g e s t h a n  t h e homing  or  ft  distance.  less  Bilateral  homing  to  than  to their  n o t a p p e a r t o be a s g r e a t  Renfrew  expanded  significantly  of  search  along  than  results  t o be  t h e concept o f  less  i s not as important  conclusions  scattering  o r an  a p p e a r s t o be i m p o r t a n t  The  various  significantly  (Khoo, 1 9 7 1 ) .  vision  using  distance  involving  blind  calculated  d i s t a n c e o f 60 m, t h e p r e d i c t e d  (1974) showed t h a t  was  fish.  t h a n 30% a n d t h e r e i s a two t o t h r e e  Renfrew and t h a t  anosmic f i s h  while  pattern  12 t i m e s t h e d i r e c t  less  with  (1952) s u p p o r t s radial  increase i n the predicted  spiral  homing  distance,  simple  search  of transplanted  r a t i o of predicted distance  Griffin  p e r c e n t a g e s homing a r e l e s s fold  distribution  percentages  and  a rectangular  search  tide  observed  with  patterns directed  the  fish,  Beach  fish  resulted  by  itself,  as for in  and had l i t t l e  Port  higher  e f f e c t on  222  the  percentage  conspicuous  remaining  visual  in  landmarks  the  transplant  do  not  area.  appear  to  However, be t h e p o o l  c h a r a c t e r i s t i c s r e c o g n i z e d by t h e f i s h . Anosmia r e s u l t e d experiment), control fish  but  fish  homing  homing  generally remaining  in  to  experiments  by Khoo  the  While  home  may  be  pool  in  aguarium study they  an  more  over  than  not  no  higher  of  of anosmic  than  those  The percentages  of fish  seem  of  homers  t o be a f f e c t e d by  remains  doubtful  i n the  h e a t c a u t e r y , t h e method u s e d by very  in  Although  different  results. suggests  The that  i n r e c o g n i z i n g t h e home p o o l guiding  the  fish  Khoo f o u n d t h a t  home  be  0. m a c u l o s u s  f o r t h r e e months showed a p r e f e r e n c e f o r  general source  showed  do  important  other pools)  aguarium  percentages  which a c c o m p a n i e s h i g h t i d e  means o f an o d o u r s t r e a m . kept  the  (41% i n each  a r e a and t h e p e r c e n t a g e s  t o produce  amount o f water m i x i n g  homing  The p e r c e n t a g e s  (0 t o 2 0 % ) .  adhesives,  Khoo, d i d n o t a p p e a r  to  were  t h e p e r i o d o f anosmia  using  (and p o s s i b l y  similar  (48% and 4 2 % ) .  i n the transplant  experiments  olfaction  were  these  observed  returning anosmia.  these  i n low p e c e n t a g e s  water,  preference  0. m a c u l o s u s t e s t e d  f o r w a t e r from  in  this  t i d e p o o l s i n which  were c a p t u r e d , compared w i t h water pumped i n t o t h e B a m f i e l d  Marine  Station.  olfactory  clues  guiding the fish  Although  this  emanating  from  home o r a t l e a s t  the reasons discussed e a r l i e r , olfactory  does  hypothesis.  not the  necessarily  tidepool  as  eliminate a means o f  r e c o g n i z i n g t h e home p o o l , f o r  i t does c a s t some  doubt  on  the  223  A combination the 20%  most  effective  of released  (removal  of  lack  of  fish  nares  the  with  until  was  homing s u c c e s s  nares the  anosmic  affected.  The d i f f i c u l t i e s  a d h e s i v e s was mentioned  a r e a s p r i o r t o homing  the treated were  suggest fish  freed  seen  to  control  small  that  was  affected  straying  is  or that  fish  at  least  obstructions.  found  i n the t o home  In  Although  laboratory  the e n t i r e duration of  blind  ofthe  and  and  w i t h 87%  either s u r v i v a l i n the f i e l d much  more  likely  in  fish  way.  vision  and o l f a c t i o n a p p e a r about  3.5 cm.  t o be e s s e n t i a l f o r  t h e homing o f f i s h  l e s s than  become u n n e c e s s a r y  b e f o r e o l f a c t i o n but i t does n o t a p p e a r  sufficient  this  t o t h e home p o o l  (51% and 48% compared  suggests  Both  that  percentages  a f t e r release  However,  ( 6 7 % and 88% compared  fish.  94% r e s p e c t i v e l y )  i n this  i n keeping the  may h a v e been u n a b l e  from  and  treated  but the  earlier.  p e r c e n t a g e o f homers r e t u r n i n g  the r e l a t i v e l y  fish  further,  may  was s a t i s f a c t o r y f o r a l m o s t  experiment,  even  s u r v i v a l i s good, s u r v i v a l i n t h e f i e l d  v e r y low r e l a t i v e  survival  treatment  i n d e t e r g e n t f o r seven  o f s i m u l t a n e o u s l y b l i n d and a n o s m i c  experiment  experiment  Additional  that  transplant  the  Beach.  and i m m e r s i o n  28% and 20% f o r c o n t r o l s )  latter  was shown t o be  i n d i v i d u a l l y suggests  with  the percentages in  fins  drastically  blocked  First  of t h e s e t r e a t m e n t s  although laboratory be r a t h e r  at  t o reduce  effect  and anosmia  method o f r e d u c i n g homing s u c c e s s t o a b o u t  paired  days) a p p e a r s  of blindness  Vision  (without o l f a c t i o n ) t o enable the f i s h  appears  t o home  to  t o be until  224  they  are l a r g e r .  shows  an  length.  cm  increase  (or  death)  and  that  which  results  appears  definitely  found  t o be  anosmia  anosmic  seems t o  between a r e a s .  both v i s i o n  the i n f o r m a t i o n t o some e x t e n t  less  be  the  and  in than  of  importance.  olfaction  I f a technique  of olfactory  olfactory  nerve)  results  indicate  that  f o r homing, t h e r e l a t i v e  irregular  The r e l a t i v e  possible that  topography..  importance  orientation  homing and o r i e n t a t i o n different  areas,  acquired)  Blennius  can  conclusive^  importance when  This  might  behaviour  e.g.  has  and  Q.  two  varying maculosus  i s better i n suggest  regular  kinds of v i s u a l  that  areas. clues  used  b e h a v i o u r h a v e been shown t o d i f f e r i n  ponds,  p o s s e s s i o n o f a t o p o g r a p h i c memory visually  clues  there are at l e a s t  i s more i m p o r t a n t i n t o p o g r a p h i c a l l y  homing and  although  conducted.  Thus i t appears  more  olfaction  homing,  u n a b l e t o s m e l l , b u t c a n s e e , homing p e r f o r m a n c e  fish:  with  n e c e s s a r y f o r homing in  precludes the perception  mechanisms a v a i l a b l e  areas  fish  result  percentage of f i s h  of g r e a t e r  (e. q. c u t t i n g  These  in  and  suggest that  both are i n v o l v e d  experiments can  is  and  p e r c e n t a g e s u c c e s s f u l l y homing  i n a greater  important i n a c q u i r i n g  olfaction  be  the  of blind  than the o t h e r treatments.  These are  in  Simultaneous blindness  straying 3.5  Homing p e r f o r m a n c e  in  different  streams,  reefs  (presumably a t  been s u g g e s t e d  ocellaris  fish  Linnaeus  occupying etc..  least  f o r a number o f  (Pieron,  1914),  The partly  littoral Fundulus  225  (Hast,  1915)  soporator  acguire  the  area  1951,  knowledge  at  orientation  The  1971).  The  local  documented, jumping  goby  topography  Bathygobius is  importance  by swimming  o f s m a l l landmarks  phoxinus  (Linnaeus)  believed  in  was  over  correctly  the  learned  demonstrated  by  (1956).  homing  of  (and  reef 1958)  inability  o f Sebagtes  to l o s s  and  flavidus  essentially  home and anosmic,  green  homed  (Linnaeus),  (Linnaeus); luscius  Haight  still Hasler  as  Rutilus  a n o s m i a and  anaesthesia resulted orientation between  and  final  rut ilus  (195 8)  treated  fish  and  controls.  the  might  be  more  suggested  visual clues to  (bream,  and  fluviatilus  Bljcca  a n o s m i a and in initial  controls  on  Abramisbjorkna  pike,  Linnaeus)  line anaesthesia,  determined  fish  Experiments  (Linnaeus);  difference  orientation  that  seems t o be  reservoir  simultaneous  in little  striatus  when H a s l e r made t h e  (Linnaeus)  t h a t b l i n d n e s s , anosmia, l a t e r a l and  vision Hisby  as  i n the Kiev  A. b a l l e r u s  roach,  E.  the  bottom.  and  readily  in  (1972) s u g g e s t e d  water,  L i n n a e u s ; and p e r c h , P e r c a  blindness  and  t o home o v e r deep w a t e r  (1959) r e p o r t e d t h a t  various s p e c i e s of f i s h brama  important  s u n f i s h , L e p o m i s m e g a l o t i s used  Gunning they  t o be guttata  c o n t a c t with the  important than o l f a c t i o n . the  appear  Epinephelus  C a r l s o n and  of v i s u a l  In  olfaction)  fishes  (Bardach,  that  best  a l l o w i n g i t t o o r i e n t i t s jumps  of Phoxinus  Vision  due  the  of the  high t i d e ,  tide.  Hasler  perhaps  (aronson,  to  a t low  and  Esox showed  simultaneous lateral  line  azimuth  of  by t r a c k i n g  floats,  (Abrosimova,  1975).  226  Simultaneous different was  b l i n d n e s s and l a t e r a l  orientation  established  novocaine  wore  taken  homing  cutthroat  of  Yellowstone  singly,  Lake,  anosmic o r c o n t r o l believed fish  was  trout,  fish  Perception demonstrated  B pec us c h r y s o p s (Linnaeus), Si  of visual  (Rafinesgue) ,  bluegills, parrot  !>• £2§I§§£ifi!JS  1965) and  cichlids  halfbeaks  polarized  light  they  than  pumpkinseeds,  Scarus (Winn  not  home  to  et  the  seem  from  any  (Khoo,  freguency  other information i s required  Rafinesgue,  (Hasler  guacamaja  Cuvier,  of  1964)  and  (Goodyear  and  cloudy  which  forthe fish  Waterman,  to polarized  direction  bass,  gibbosus  in  homing  has  Lejgomj.s  al. ,  feasible that the  white  sun) ; s a l m o n i d s  1972) ( o r i e n t a t i o n  t h e y c a n home on  the  in  The a u t h o r s  landmarks  ( B a i r d and G i r a r d )  o b s t a c l e s , and s i n c e 1971),  197 2 ) i  i n a number o f f i s h :  are important i n  can  Richardson  r e a c h e d t h e home a r e a a s o t h e r  fishes,  (orientation  (Waterman,  i t does  nor  l o n g e r t o home than  and L a B a r ,  (Waterman, 1958-1959, c i t e d  However,  since  took  clues other  Valenciennes  1969)  c^arki  fish  vision  be e s s e n t i a l f o r t h e  LeBomis m a c h r o c h i r u s  m o s q u i t o f i s h , Gambusia a f f i n u s Ferguson,  orientation  t o locate the trap.  as possible  1958),  Neither  to  Salmo  trout  in  were removed t h a n when t h e  shown  (McCleave  t h a t a s many b l i n d  but c o r r e c t  1976).  although b l i n d  but were u n a b l e  been  when e y e c a p s  (Abrosimova,  olfaction,  anaesthesia resulted  from c o n t r o l f i s h ,  faster off  line  celestial of  0.  (Groot, 1972) light).  bodies o r maculosus.  and c i r c u m n a v i g a t e days  i s high.  and  nights  I n any c a s e ,  t o r e c o q n i z e t h e home  227  pool in  or range.  of v i s i o n t h e r e f o r e ,  the r e c o g n i t i o n of landmarks.  and  Given the  t u r b u l e n c e o f the i n t e r t i d a l ,  how  this  sense c o u l d  In be  The r o l e  be u s e d  over  streams, the r o l e  the  c a s e f o r homing  any  That v i s i o n  by t h e f a i l u r e  of simultaneously b l i n d  importance  particular homing  was  scented  streams  rainbow  to  control fish,  and  however, anosmic  homing  well  Oncorhynchus  of  1968).  et  Cooper  and  fish  established. tshawytscha  return  vision,  to  appropriately  Richardson.  imprinted  (Salbaum) to  confirm Cooper  Hasler,  1976).  However  an  as s m o l t s o r bulb  s c e n t e d w a t e r , by homing i m p r i n t e d  1976;  i n the o l f a c t o r y  1969).  eliminated  Salmo g a i r d n e r i  to  the  although  electrophysiological studies of olfactory  al., and  In  (Walbaum),  almost completely  The  shown  of s a l m o n i d s t o  kisutch  appear  the and The  importance  S c h o l z , 1976; involvement  discrimination  of  and  olfaction  Scholz et a l . , of  o f home and  electrophysiologically  Bodznick  be  t o home.  Oncorhynchus  morpholine  to  to  was  c o h o salmon  w a t e r s h a s been d e m o n s t r a t e d al.,  a role  the  anosmia  seems  (1959) showed t h i s  c h e m i c a l ( m o r p h o l i n e o r phenet hy 1 a l c o h o l )  responses  memory  al.,  trout  fingerlings,  1976;  salmon,  b l i n d n e s s and  (Groves e t  (Cooper  in  to appreciate  i n homing  shown t o be more i m p o r t a n t t h a n  homing  organic  olfaction  Chinook  simultaneous  and  must p l a y  s t r e a m s seems t o be f a i r l y  of  olfaction  of  turbidity  Lejaomis m e g a l o t i s megaj-otis  (Rafinesgue).  The  t o be  distance.  Gunning  longear sunfish  appear  possible  i t is difficult  of o l f a c t i o n  more i m p o r t a n t t h a n v i s i o n .  would  long  term  other stream (Oshima  (1975) has s u g g e s t e d c a u t i o n  et be  228  exercised  i n the  i n t e r p r e t a t i o n of e l e c t r o p h y s i o l o g i c a l  results  r e l y i n g on the amplitude of o l f a c t o r y bulb d i s c r i m i n a t i o n s . The  ability  of  fish  other  than  d i s c r i m i n a t e between odours has been shown. Hybgrhynchus  notatus  Kafinesgue  between the waters of two between  rinses  of  were  anosmic  fish  Bluntnose  d i f f e r e n t streams (Hasler,  natural  plants  could  not.  minnows  (Walker and these  Angsiilsl.  1966)  Hasler,  of  natural  changing  and  salinity  (Creutzberg,  vulgaris  (Turton) = be  f l o o d t i d e water using  1959).  did The  not  produce  ability  to  the  same  response  0.  maculosus has  olfaction  been demonstrated and  postulate  the  turbulent  intertidal,  associated  of  existence  with  or  of the  particular  pools.  Since  involved i n the a c g u i s i t i o n o f i n f o r m a t i o n , may  be e s t a b l i s h e d which i s a s s o c i a t e d  clues may  from  other  senses, e.g.  in  odour  existence  with  the  of  the  homing  of  necessary  to  streams  in  discrete  the  odours  v i s i o n appears to  be  a t o p o g r a p h i c memory the  the l a t e r a l l i n e ,  be used t o a s s i s t i n r e c o g n i z i n g  by  compartments.  i t appears  discrete  shown  an  r a t e of flow i n the  odour r e l e a s i n g compartment r e l a t i v e to the other importance  Simply  l o c a t e the source of  (1967) t o depend on an i n c r e a s e d  The  able  additions  odour i n a c i r c u l a r tank d i v i d e d i n t o compartments was Kleerekoper  1949).  i n l a n d water, presumably through o l f a c t i o n .  the  and  discriminations  £132.1115. a n g u i i i a (Linnaeus) e l v e r s have been shown to t o d i s c r i m i n a t e between ebb  to  t r a i n e d to d i s c r i m i n a t e  P r e v i o u s l y t r a i n e d b l i n d f i s h c o u l d make but  salmonids  perception  of  and o l f a c t i o n  p a r t i c u l a r pools.  When  229  both  vision  using  and  other senses,  home r a n g e , can  olfaction  t o determine  thus being unable  home c o u p l e d  olfactory  are e l i m i n a t e d t h e f i s h  can  be  suggests the  (or s e n s e s )  when o n l y v i s i o n  Since t h i s that  additional  consideration system  of  seems  auxilliary  must  be  the  involved  pressure.  range  effect  displacement other  ear-swim to  is  effect  and  van  The  waves  (or  vice  the in  the mixed sense  possibility homing,  some  acoustico-lateralis mechanism ( i f  forms;  particularly  chiefly  has  displacement  evident at been  close  termed  the  d e t e c t e d by t h e a r r a y o f line. ,  as a " f a r - f i e l d "  effect  and  on  the  the i n n e r  ( T a v o l g a , 1971a) a l t h o u g h t h e  near-field  acoustical  t o be  Pressure,  sensitive  and  vestibular  apparatus  1962).  line  can  movements o f water p r o d u c e d  obstacle  how  primarily  Bergeijk, lateral  fish  of another  auxilliary  phenomenon  is  regarded  c a n s t i m u l a t e the  ( H a r r i s and  field  the  is  b l a d d e r complex i s b e l i e v e d  compression  The  i s p r e s e n t i n two  receptors i n the l a t e r a l  hand,  blind  i s eliminated.  involved  made.  to the  d i s t a n c e i n the  eliminated  are  that  appreciating  any  most l i k e l y  Displacement  t o t h e s o u r c e and  "near-field"  fact  involvement  not  be  unable,  i n homing.  A c o u s t i c energy and  of  or o l f a c t i o n  s t u d y has  them  The  over  mechanisms  to  t h e r e i s one)  t o home.  used  intertidal,  be  where i t i s , i n r e l a t i o n  with the d i f f i c u l t y  sense  may  versa)  d e t e c t c u r r e n t s and when  (Dijkgraaf,  a  fish  1962)  and  other  near-  approaches possibly  an  other  230  hydrographic  phenomena p r o d u c e d  the  of obstacles  vicinity  suggested  that  the  three-dimensional In  addition,  frequency  the l a t e r a l  line  within  defined  respectively  (Tavolga,  information  and  bearing  Localization  of f a r f i e l d  open t o d i s p u t e  received swimming  i t  seems  assess.  the l a t e r a l  into obstacles  maintaining  the general  It  is  characteristic  within  one  Low f r e q u e n c y  sixth  should  van of  s o u n d s o f 50  Because the l a t e r a l  quite  and  range o f about  the f i s h  be  readily  hiqhly to f a r  may be o f q r e a t e r  by  (Harris  a to  50 t o 3 m, line  able  i s an  to  (Tavolqa,  s o u n d s by f i s h e s a p p e a r s  0* m a c u l o s u s a r e r e s p o n d i n g effects  near-field  to  get  1971b). be  more  (Popper and F a y , 1973). ,  While  field  accurate  t o d e t e c t low  1971a).  range  a "fairly  (1957)  a p p e a r s t o be a b l e  near-field  receptors  Lowenstein  environment".  being  array of displacment  of the fish i n  o f t h e immediate  1962).  an e f f e c t i v e  1971b). could give  the as  (van B e r g e i j k ,  100 Hz p r o v i d e  line  representation  1962)  wavelength  (Tavolga,  lateral  vibrations  Bergeijk,  by t h e l o c o m o t i o n  line  water  field  sound  importance.  swept o u t  movement  movements  to  can  that  i s ,  i s difficult to have  be  the f i s h  specific  detected  by 0. m a c u l o s u s .  have s p e c i f i c  recognizable  weather  and  s e a s t a t e c h a n g e s seems more d o u b t f u l , a l t h o u g h i f  at l e a s t  general  direction  movement  (perpendicular  between  and  discriminated  water  Whether a r e a s  sea,  near  information  prevent  tidepools which  hominq  sources,  of t h e animal,  that  that  Whether  c a n do more t h a n  and b e i n q  possible  improbable  tidepools  characteristics,  to  tidal  flow)  given  and  231  g e n e r a l a r e a can be p e r c e i v e d , t h e u s e o f v i s i o n  or o l f a c t i o n  recognize  Although  specific  knowledge, t h i s that  tidepools  Wilimovsky,  pools  may  be  h a s n o t been i n v e s t i g a t e d , may h a v e " s o u n d s "  p e r s o n a l communication)  information  can  be  or s p e c i f i c  I f this  i s  d e t e c t e d and used  f u r t h e r means o f r e c o g n i t i o n the  i t has been  of specific  (Norman  harmonics  the  case  (Bruce and i f  by 0. m a c u l o s u s ,  pools i s  available  a to  fish.  The crude  possibility  echo-location  investigated, this  i s  sounds  (1971b) f o u n d  itself. (wenz,  no  find  m a c u l o s u s may i t s way  and  1962)  and  felis  It i s  that  (Linnaeus)  i n the i n t e r t i d a l  may  of  emitted  level  to conduct.  eliminating  cautery of  the  the  of  reduce  of  A major p r o b l e m perceptiveness  lateral  whether  line  pores  another  i s known  to  However, T a v o l g a the  sea  sound  shallow  o f water movement  of t h e r o l e  been  catfish  was u s i n g i n f o r m a t i o n from to  orient  water  noise  and d e g r e e o f  t h e e f f e c t i v e n e s s o f any  c l u e s p e r c e i v e d by t h e a c o u s t i c o - l a t e r a l i s  difficult  not  known  (Bitehill).  suggest  to  amount  Investigations  has  not  1956) w  ambient  the  home  Mowbray,  reverberations  and  use some k i n d o f  o f sound p r o d u c t i o n a l t h o u g h  evidence  However, t h e  turbulence  O.  pctodecimspinosus  (Barber  5 S i § i c h t h y s [_=AriusJ reflections  to  capable  gioxocephalus  produce  that  b u t i t seems u n l i k e l y .  species  cottid,  of  t o my  suggested  which a r e p o o l s p e c i f i c  Leaman, p e r s o n a l c o m m u n i c a t i o n ) . this  involved.  to  system.  the  lateral  line  are  seems t o be t o f i n d  a way  of the l a t e r a l l i n e .  Heat  and  cutting  nerves  are  232  difficult  on s u c h  f o u n d which  will  small  adhere  period  of t i m e .  The  wear  off  soon  too  satisfactory  fish, to the  effects  lateral  line  water the  was  be  movements might itself  First  pressure  allowing  p r e s s u r e was fluctuations  was  t h e r e any  the  i n poor  fish  being  the  perception  unable  of  living  to perceive  and  live  to  in  these  maintain  depth  that  i n a n a r e a where p r e s s u r e detection  may  be  of  some  these f a c t o r s are involved i n  cannot  be  excluded.  response to pressure, s i x f i s h  In  {from  1,2  brief to  5.0  were p l a c e d  in  which  could  be a d j u s t e d , by m a n i p u l a t i n g b y - p a s s  valves  column  and  the  pressure  to a c c l i m a t e f o r s e v e r a l from  then r a p i d l y of  i n a p l e x i g l a s s tank  a  Beach  increased  performed under  since  a  sensory  i f this resulted  i m p o r t a n t to a f i s h  fish  maculosus  the f i s h  slowly  the  i f  environment.  possibility  of  any  be d i f f i c u l t t o c o n c l u d e t h a t homing  intertidal  homing o f 0.  from  an  for  Even  eliminate  i n the  c o n n e c t e d t o a mercury  was  and  1976).  result  the  examination  the  in  c a n be c o n s i d e r a b l e  importance,  cm)  line,  been  s u c h as n o v o c a i n e ,  E l i m i n a t i n g the a b i l i t y  a t a l l i n such  changes  maculosus  zone.  Since  the  to  i t might  involved  0.  1975,  found  movements must be r e l a t i v e l y intertidal  of  of a n a e s t h e t i c s ,  perceptiveness of the l a t e r a l performance,  skin  (Abrosimova,  method c o u l d  homing  and t o d a t e , no a d h e s i v e has  4 m light  sea l e v e l  raised  {simulating and  and  to  1.8  lowered  atmospheres.  providing  l a r g e waves).  by t h e f i s h  After  minutes, t h e p r e s s u r e  dark c o n d i t i o n s .  overt reaction  tank.  to  pressure  Both t e s t s  I n none o f pressure  The  the  were tests  changes.  233  Tbe  majority  tank, and did  of f i s h  although a few  maculosus  display a t i d a l midpoint  of  suggested that the hydrostatic  salinity  the  of  laboratory  locomotor  altering  have  activity  possible synchronizer pressure,  since  Gibson  c y c l e s were capable in  movement  been shown to the  peak  and  c o i n c i d e with high t i d e (Green, 1971c).  synchronizers.  pressure  the  changes.  arhythmic  rhythm  weather c o n d i t i o n s  entraining pholis  He  turbulence  (1971) found t h a t  of  Blennius  the phase of an  of the a c t i v i t y  temperature,  are too dependent on sea and  reliable  rhythm  in  rhythm  which  p o s i t i o n s i n the  f i s h o c c a s i o n a l l y moved,  not immediately f o l l o w pressure 0.  was  remained in t h e i r i n i t i a l  a  and  to  be  experimental  tidal  individuals  activity as well  as  e x i s t i n g rhythm.  Thus while there i s some c i r c u m s t a n t i a l evidence, from Green's work, to suggest t h a t changes,  the  nature  of  d e t e c t i o n of pressure bladders,  p.  bladder.  have  Plf^u/r^nectes (Linnaeus), larvae  mechanism  changes i s  at  been  least shown  platessa  perceives is  possible  two to  in  al.,  swim 1966).  lacking  gas-filled  Morgan,  1966).  not c l e a r . fish  respond and  to  pressure  Cen tronotus  appear t o be  as  changes, gun n e l l us  sensitive  Pressure  responses  (1971c)  swim  possess a swim  f o r example. B l e n n i u s  are  While  with  bladders,  organs  Green  pressure  l a r v a l t e l e o s t s l a c k i n g swim  Linnaeus  although they do not  possessing  (Qasim et  the  maculosus  maculosus l i k e a l l c o t t i d s , does not  However  bladders  0.  well points  known out  in  as  phpjLis  invertebrates  (Knight-Jones  and  that  and  Morris  234  Kittleman  (1967)  piezoelectric pressure that  reported  otoliths  reception.  1  two  species  which t h e y  Further,  of  suggest  fish  which  have  may be a mechanism o f  Shamos and L a v i n e  (1967)  p i e z o e l e c t r i c i t y may be a u n i v e r s a l p r o p e r t y  suggest  of a l l l i v i n g  tissue. Although  the r o l e of pressure  homing c a n n o t  be e v a l u a t e d  from t h i s  reguired  home  no  since  to  pools  outward  the  homing  this  integrating  other clues  in  would be  i s t a k e n o f d i r e c t i o n and  height  with again  that  will  of  inertial  be  at  the  respect with  navigation  0. m a c u l o s u s  hypothesis,  movements ( d i r e c t l y  acceleration  order  possibility  in to  detection  same  tide.,  The implicated  study,  account  a t t h e same v e r t i c a l  depth a t high  According  since  and d e p t h  the  seems  respect  unlikely.  a n i m a l keeps t r a c k  recording acceleration, to  time  of a l l  integrating  to  obtain  v e l o c i t y , and  t o time t o  obtain  distance)  t o e i t h e r r e t r a c e p r e c i s e l y t h e outward journey  or t o c a l c u l a t e t h e c o u r s e  m i g h t be  back  home  (Barlow,  in  back home  1963;  Keeton,  1974) . The outward Animals  journey)  former can  collected  possibility easily  at First  be Beach  (precisely  ruled  out  for  retracing  the  0..maculosus.  (and a t t h e m a j o r i t y  of other  crystalline arrangement * 0 t o l i t h s with a such that when a mechanical s t r e s s ( p r e s s u r e ) i s a p p l i e d , an e l e c t r o m o t i v e f o r c e i s generated.  235  sites) by  boat,  kept for  were t r a n s p o r t e d t o a n d from t h e l a b o r a t o r y by taking  at least tagging  overnight  and o t h e r  The home)  either  geographical  the  coordinates  the  and  although  possibility  Keeton,  surgical  of  "know"  the  be  However  lesions  Moreover,  eliminated  respond  of  the  system  The has  (1969) f o u n d a u r a t us  that  possibility not  been  are nearly  the  conducted  with  negative,  (Schmidt-Koenig,  apparatus  showed  that  no  1965;  to  which  would  fish  maintains  symmetric)  (Keeton,  vertebrates  four  be  1974).  orders  of  tolerated in a  1968).  examined.  turns  with  ( t h e presumed  which  to  or  difference i n  from c o n t r o l s  may  make  However,  use  a  nearly  cues,  constant  (that i s , cumulative over  of  such  Kleerekoper  t h a t i n t h e absence o f d i r e c t i o n a l  and l e f t  calculate  i s largely  i s three  (Harden J o n e s ,  (Linnaeus)  between r i g h t turns  than  back  The m a j o r i t y o f  been  acceleration 2  precision  system  threshold  greater  the course  deep a n a e s t h e s i a  vestibular  (,5°/sec and 6 c m / s e c )  magnitude  around  b i r d s c a r r i e d t o t h e r e l e a s e s i t e s on  o r homing s u c c e s s s  the  were  r e l e a s e s i t e s and  home.  the a v a i l a b l e evidence  i n l i g h t p r o o f drums, under  bearings  and  have  principal detectors of acceleration) inital  or  home  possibility  cannot  1974).  turntables,  fish  (calculating  not n e c e s s a r i l y d i r e c t ,  investigations of this birds  The f i s h  i n t h e l a b o r a t o r y and were moved  alternative  that  "compute" a p a t h ,  and  purposes.  second  reguires  of two p o s s i b l e r o u t e s .  foot  et al.  Carassius  relationship  left  t h o u s a n d s of t u r n s  a  and r i g h t  of  varying  236  magnitude, direction turn (11  direction  is established  o c c u r s and hours).  the  The  established  by  other  the  perception soon be  to  suggest  fish  may  be  of  position  swims,  and  since  homing,  The  receptors For  the  the  (McCleave derived  tidal  et from  fe  y  Q.  is  reguired  the  ability  of  period may  be  direction  was  t i d e s and  currents  the  1975).  navigation  short  the  to recognize  the  Since  would  the  In home  without  movement  i n the  be  present  the  rather  electric  since  electric  majority  of t e l e o s t s .  involved,  favour  possibility  of  the that  i s used  e i t h e r case, pool.  than  u s e s an  unlikely,  distances in  be  seems u n l i k e l y .  maculosus  magnetic f i e l d  maculosus i s u n l i k e l y .  clues  the would  would  evidence  earth's  fish  currents  and  identified  1971),  the  flow  is  the  Other  below  hops r a t h e r  in orientation  any  rate  of s h o r t  0.  be  a  composed  that  al.,  angle  initial  at  directional clues  same r e a s o n and of  (Tesch,  be  possibility  have y e t t o  absence  altered  inertial  other  s e n s o r y mechanism  of  a d i f f e r e n t course.,  long  of  this  of  once a  f o r some  a n g u l a r a c c e l e r a t i o n and  maculosus appears t o  use  maintained  that  presence  o f 0.  extensive  be  the  maintain  a t t i m e s of  compensation  in  on  suggests that  clues.  threshold  swimming  reguired  d i r e c t i o n can  This  range o r i e n t a t i o n i f the  However of  seguence.  continuous  authors  important i n long  position  and  as  well  as  hypothesis information  in orientation  other  information  237  VIII.  1* V a r i a b i l i t y , i n homing  This 0.  With  shown  that  turbulence  and t h e p e r c e n t a g e  decreases  irregularity  and  with in  tidepool  remaining  t u r b u l e n t rough areas  not  seem  suggest  unreasonable that  behaviour the  at  observed  turbulence  topography  of  some  particular  topography,  size  exist,  t h a t homing s u c c e s s  the fish.  appears  the  which  y e a r s and t h e n  to  o r age r e l a t e d  declines.  transplant  of  Homing  area  i n the t r a n s p l a n t  v e r y few f i s h  The f i s h  the  which  do  percentages  from  this  variability  and  pools.  the In  related  variability  home  returning It  does  finding to in  homing  a function of  nature  of the  addition  to the  to  turbulence  and  i n homing a p p e a r s t o  i n c r e a s e s w i t h a g e up t o a b o u t 2  The p e r c e n t a g e  decreases  success  but i n i n l e t s ,  pools  be  terrain,  topographical  extrapolate  variability  so  the  w i t h i n any o n e a r e a i s p a r t l y  surrounding  topography.  with other t i d e p o o l areas.  to  least  the  and  show t h e h i g h e s t  t o t h e home p o o l , compared  of  v a r i a b i l i t y can  of  turbulence  areas,  behaviour  l a r g e area.,  of f i s h  decreasing  of  regularity  home o r r e m a i n i n t h e t r a n s p l a n t a r e a . in  homing  between a r e a s and t h a t t h i s  i s shown t o an i n c r e a s i n g l y  increases area  has  t o t u r b u l e n c e and t h e r o u g h n e s s  decreasing  fidelity  DISCUSSION  behaviour  study  maculosus d i f f e r s  be r e l a t e d  GENERAL  with  of  fish  age e x c e p t  remaining  in  f o r the o l d e s t  238  The v a r i a b i l i t y with  t h e s i g n i f i c a n c e o f such  been  put f o r w a r d  Williams from as  i n homing b e h a v i o u r  (1957)  behaviour.  f o r the function  suggested  that  dry  drainage.  land  Green  Two major r e a s o n s  o f homing i n i n t e r t i d a l  and Khoo  disappear through  (1971) s u g g e s t e d  that  mechanism  balanced  o f r e s o u r c e s i n the i n t e r t i d a l .  the a v a i l a b i l i t y involved  establishment  fidelity  adult  life,  t o t h e adopted  these  factors  inflexible  species,  since  annihilation may  homing  added  mechanism  turbulent  the  adult  i s  the  Space o r  main  resource  I f d i s p e r s a l and of the f i s h  maintained  t o a homing  mechanism  However,  recruits  in  result  to the  i n complete  t h e 'non-homers*  tidepools  would  Conversely,  would be d i s a d v a n t a g e o u s may  and i f  throughout  o f t h e environment.  conditions  of the population.  provide  i n the l i f e  home  ensure an b a l a n c e d e x p l o i t a t i o n an  t o be  o f p . jmaculosus.  o f home o c c u r e a r l y  strict  i t may  f o r p o p u l a t i o n d i s t r i b u t i o n and  of t i d e p o o l s appears  i n the d i s t r i b u t i o n  such  subsurface  s e r v e as a s t a b i l i z i n g utilization  have fish,  i n unfavourable s i t u a t i o n s  or i n pools t h a t (1971b)  associated  homing s e r v e s t o p r e v e n t t h e f i s h  b e i n g s t r a n d e d a t low t i d e on  may be  may  which  serve  a  d i s p e r s a l and r e p o p u l a t i o n f u n c t i o n .  Both selection would  suggested  against the kind  seem  t o be s e v e r e .  r e a s o n s have some v a l i d i t y . of  mistake  suggested  by  be a l l t h a t i s r e g u i r e d ,  Williams  Knowledge o f t h o s e c h a r a c t e r i s t i c s o f  a p o o l w h i c h make i t an a c c e p t a b l e p l a c e a t low t i d e , to  However,  a n d homing  to a p a r t i c u l a r  would  seem  pool f o r  239  this  reason  would seem t o s u b j e c t t h e  In i n l e t areas, behaviour range  a restricted  conceivably  Sith fidelity  and  could r e s u l t  regard  falling  of  risk  In rough a r e a s ,  may  result  and  displacement new  lost  from t h e t h e r e an  i s very  the home a r e a .  Only  provides  not  fish:  those  the guestion o f  directly  move  fish  concerning  from s h e l t e r e d  e x t e n s i v e l y between  is  of  being  reduced, behaviour.  shown  can  be  because the  resources.  their  be s e v e r a l c a t e g o r i e s  area  shown,  throughout  information  i n t u r b u l e n t , rough a r e a s  w h i c h a p p e a r t o show l i t t l e but  of  tide  'non-homing' f i s h i n t i d e p o o l  addressed,  appear to  when d i s p l a c e d , t h o s e  particular  l o c a l p o p u l a t i o n , and  distribution  a number o f s u g g e s t i o n s  f u n c t i o n . , There  with  danger o f  homing  behaviour  a l l o w s t h e even u t i l i z a t i o n of  was  to  pool  i n adoption  pools  movement and  the  maintenance o f  i n a balanced  where t h e  movement between  i n c r e a s i n g amount o f  although  a  movement i s n o t  pools appears to r e s u l t  related  intertidal,  the f i s h  associated  i n movement a r o u n d  homing  results  home  i t appears that  closely  r e d u c t i o n , such  p o p u l a t i o n by  to the  homing  i n s t r a n d i n g , unless the  Thus i n e a c h a r e a , t h e k i n d o f  areas  inflexible  where movement between p o o l s a t h i g h  to other  pool as  risk.  tide.  involved  i n population  the  it  and  to the second reason,  homing b e h a v i o u r  degree  area.  is  home r a n g e  encompassed a v e r t i c a l t r a n s e c t w h i c h a l l o w e d  move s e a w a r d s w i t h t h e  the  f i s h t o unnecessary  from t h i s  derivation of  which a d o p t  pools, those  or  and  non-hcming the  ( f r e g u e n t l y high)  fidelity  study  homing  f i s h which  new  pool  tidepools behaviour are  never  210  seen  again  after  relatively  displacement,  widely  and  juvenile  around t h e i n t e r t i d a l  fish  before  which  adopting  move a home  range. Since sort  t h e r e a p p e a r s t o be a  involved  0. m a c u l o s u s  in  (John  observation)  the  determination  Green,  personal  the r e l a t i v e l y  o f a home r a n g e  of  numbers o v e r  fish  population  w h i c h adopt  provide  a  the  transplant  dispersal  rough areas,  and t h o s e  this  However,  latter  when t h e y  encounter  accidently used  being  a  relatively pool  similar data  to  personal  as  fish  the " b a l a n c i n g "  wide areas.  the  new  Those  home  pool  mechanism i n t u r b u l e n t  function  from  in  rough that  c o n t i n u e t o move a r o u n d w i d e l y  even  they  t h i s study  less suggest  exposed  i n a homing e x p e r i m e n t ,  pools.  If  these  fish  are  p o o l a n d a r e c a u g h t and  may b e t h e f i s h  which  do  not  p o s s i b l y moving e l s e w h e r e on t h e b e a c h o r  m o r t a l i t y by moving o u t o f a t i d e p o o l d u r i n g The f a t e o f f i s h  i s unknown; m o r t a l i t y , d i s p e r s a l  predation  which  are  never  out of the study  seen  area or  appear t o be t h e a l t e r n a t i v e s .  In i n l e t s , are  may p e r m i t  of  s h e l t e r e d p o o l s which show e x t e n s i v e  more  turbulent conditions. again  communication;  d i s p l a c e d o r move t o a l o w e r  subject  numbers  from  of f i s h  show homing b e h a v i o u r ,  pool  some  repopulation  the limited  category  of  of  and  movement p r e s u m a b l y s e r v e areas.  factor  e x t e n s i v e movements o f j u v e n i l e  p r i o r t o the adoption pool  "space"  shown.  topographical  The  little  area  relative  regularity  of  f i d e l i t y and  absence such  of  areas  homing  turbulence allows  behaviour and  the  relatively  241  unrestricted vertical  movement.  seasonal largely food of  lateral  movement  I t i s suggested  instability a result  and  of  inlet  fidelity  fact  movement i s b o t h  the p o s s i b i l i t y  little  the  purpose.  p o s s i b l e and and  would s u b j e c t t h e f i s h  the  reason  contribute of  other  winter  deeper  presumably  armatus,  fish  least  disadvantageous  of  absence  areas.  subject  and  to  homing  risk. of  Another  t o be  In  little suggests  behaviour  This  many  factor  much  been o b s e r v e d  some  more  Herons, g u l l s ,  of  the  may  be  inlet  fish  which  may  (sometimes  and i n f l e x i b l e  f o r t h e avoidance  in  important  prey  in  k i n g f i s h e r s and  f e e d i n g i n such i s  Artedius fenestralis  i n a l l the sheltered  a s m a l l home range  be  o f homing b e h a v i o u r i n i n l e t s i s t h a t  predators,  have been c a u g h t  0. m a c u l o s u s )  instability  of t h e y e a r ,  dispersal  tidepool areas.  at  range  water).  b i r d s have f r e q u e n t l y  Possible  of  the  t o t h e absence  areas than  part  t o unnecessary  p r e d a t i o n , which i s b e l i e v e d  inlet  and  to  to  t i m e s o f y e a r , t h e m a i n t e n a n c e o f a home  range  (presumably  (believed  the  When f o o d and s h e l t e r r e s o u r c e s d i s a p p e a r  most t u r b u l e n t  for  and  o f b e i n g s t r a n d e d a t low t i d e , t h e  t h e a d o p t i o n o f a s m a l l home  serve at  populations  o f b e i n g swept away f o r a l a r g e  that  amount o f  that these f a c t o r s  and homing b e h a v i o u r i n s u c h  to  risk  considerable  resources) c o n t r i b u t e to the r e l a t i v e  addition that  a  o f t h e s e a s o n a l t u r b u l e n c e and  shelter  home r a n g e  and  0.  areas  maculosus.  and L e p t o c q t t u s  greater  areas s t u d i e d .  numbers  than  The a d o p t i o n  homing b e h a v i o u r  might  well  0. m a c u l o s u s  may  of predators.  T h u s t h e k i n d s o f movement shown by  242  be  closely  particular  related area,  distribution  g.Sensory  the  and i n t h i s  and r e s o u r c e  Movement  a p p e a r s t o be d i r e c t e d , search  the  in  successful  return The  Thus i t seems t h a t  of  possibly  of  distance.,  also.  range  Although  i s  a  the  to  the  some  kind  of that  rather  than  i t s acguisition. be  intertidal  vision)  essential for  that  suggests  cannot  or  study f a i l e d  nether  involves  t o show t h a t  can  from be  may  and p o s s i b l y  with  wider  that  be u s e d o v e r any  t h e use o f o l f a c t i o n  v i s i o n i s concerned  recognition  fish,  pool  suggestion  in  r e c o g n i t i o n o f t h e home p o o l ,  Presumably  this  t o t h e home  by t h e f i s h ,  appears  also  landmarks, or o l f a c t o r y c l u e s by  there  l a n d m a r k s o f some s o r t , a l t h o u g h w h e t h e r t h i s  home  are  t o t h e home r a n g e .  (and  pools  home  involving  particularly  neither  turbulence  to  to  O l f a c t i o n and v i s i o n a r e known t o be  olfaction  restricted  release  than  are recognized  adults  population  behaviour  0. m a c u l o s u s  the  and  i n homing b e h a v i o u r ,  However,  by  rather  p o s i t i o n of the pool.  involved  i n homing  from  process,  a t t r i b u t e s of t h e pool  by a  utilization.  mechanism(s) used  unclear,  expanded  demands f o r s u r v i v a l imposed  way d e t e r m i n e a b a l a n c e d  mechanisms i n v o l v e d  The still  to  other  the r e c o g n i t i o n i s confined  to  areas i s unclear.  e i t h e r v i s u a l home  t h e home p o o l  eliminated  be  were  pool  recognized  as p o s s i b l e , f o r the  243  reasons  discussed  landmarks the  of  particulars  intertidal  juvenile pools  may  areas  detection  of  of  the  and  water  may  may  of  their  established during  the  the  a l s o be  an  information  positions in of  learned.  senses,  the  pigeons  possibility  has  available  for  homing, be  Further,  a n i m a l s a t any conditions,  in  homing  s e n s e s may  First  the  opposed t o the of  the  intertidal, flow  be  of  and  the may  work on  that  relative  different  Khoo's  (1971)  i s more i m p o r t a n t  between work  and  due  are  different  conditions  (Keeton,  i n homing.  areas  homing  that  d i f f e r e n c e s between  between  very  be  home p o o l .  individual  r e g a r d l e s s o f changes o v e r t i m e  the  fish,  mechanisms  importantly^  different be  The  number  most  may  home  relative  currents,  r u l e d out. a  under  of  been f o u n d i m p l i c a t i n g o t h e r  mechanisms used  suggests  of  olfaction  time,  behaviour  topography,  results  used  be  that  and  there  one  has  cannot  demonstrated  mechanisms may 1974).  evidence  extensive  In a d d i t i o n ,  t o swim f r o m " p o o l " t o " p o o l " t o r e t u r n t o the  no  and  characteristics  indication  from t i d a l  odours  period  t r a n s p l a n t p o o l , as  provide  the  relative  movement  t r a n s p l a n t p o o l i n the  Although  in  "memory"  pools  Specific  when  using d i r e c t i o n a l able  A  between p o o l s  i s flooded  height  be  movement.  and  pool,  earlier.  to  The  this  t o homing a t P o r t  of  suggest  Renfrew  and  different  Certainly, study  in  variability  turbulence  importance areas.  or  than  the that at  Beach.  Emlen pointed  out  that  and  Keeton  c u e s w h i c h may  (Adler, be  1971;  Keeton,  e s s e n t i a l i n the  1974)  have  ontogenetic  2HH  development essential used,  of  orientation  to adults,  the  but  relative  importance  of  vision  and  T h i s study  at First  guite  olfaction. behaviour (ages) areas.  Beach.  This  is  being  differ for  both  v i s i o n and  0.  laculssus  apparently does  become  suggest  that  may be e s s e n t i a l t o " l e a r n i n g " t h e a r e a , b u t once  successfully  in  the  Associated with t h i s between  a t which  fish  The r e l a t i v e  different  may n o t be  may  juvenile  olfaction  o t h e r s e n s e s have a c g u i r e d s u f f i c i e n t used  cues  has shown t h a t  subseguently  at least  senses  behaviour  the  a r e e s s e n t i a l t o t h e homing o f  unnecessary, these  homing  and even i f t h e same s e t o f c u e s  j u v e n i l e s and a d u l t s . olfaction  and  areas  and  i n f o r m a t i o n , they  absence  of e i t h e r  are t h e d i f f e r e n c e s the  apparently  can  be  vision or in  different  hcming sizes  b e g i n t o show homing b e h a v i o u r  i n different  importance  may be  of different  i n t h e o n t o g e n e t i c development  areas of d i f f e r i n g turbulence.  senses  very  o f homing b e h a v i o u r i n  2H5  IX.  Variability  i n homing b e h a v i o u r  The evidence  SOMH&BY  i n different  m a j o r i t y o f 0. m a c u l o s u s  o f home r a n g e  showing f i d e l i t y  fidelity,  with  t o one p a r t i c u l a r  fidelity  t o one o f t h e p o o l s i n t h e g r o u p  evidence  of  percentages 0.  0. m a c u l o s u s behaviour,  homing  was  sometimes  observed.  considerable  showed  pools.  Beach  variability majority  of  i n t h e t r a n s p l a n t area,  showed i n the  of  of capture.  percentages  of  greater  than t h e o t h e r  The pool  group  homing  Variable  transplanted  some f o r e x t e n s i v e  o f time. In  evidence  Grappler  Inlet,  of r e s t r i c t e d  Homing a c r o s s  p . maculosus  home r a n g e f i d e l i t y  V a r i a b l e b u t low p e r c e n t a g e s area.  majority  although  t o the i n i t i a l  maculosus remained  periods  the  showed  proportions of fish  displaced at F i r s t  homing  maculosus r e t u r n e d  but 0.  of  group,  Beach  pool as t o a small  Of  majority  latter  First  egual  pools.  The  the  at  areas  of fish  displayed o r homing  remained  i n the  t h e deep w a t e r o f t h e i n l e t  little  behaviour. transplant  was f o u n d  t o be  impossible. Differences at  First  believed and  Beach, G r a p p l e r to  be  observation.  different  in  areas  homing b e h a v i o u r  Inlet  between 0. m a c u l o s u s  and P o r t Renfrew  were  initially  related  t o exposure,  as d e t e r m i n e d by l o c a t i o n  Further  examination  of  showed  that  homing  differences  behaviour  i n homing  at  behaviour  246  could be r e l a t e d t o wave a c t i o n of  the  terrain.  With  (turbulence) and  decreasing  turbulence  r e g u l a r i t y o f the t e r r a i n , f i d e l i t y i s shown number  of pools o r a wider area.  in  turbulent  behaviour  and i n c r e a s i n g  to  an  increasing  The  fish  which  do  rough areas show t h e h i g h e s t percentages o f  homing f i s h r e t u r n i n g t o homing  roughness  In t i d e p o o l areas, i n c r e a s i n g  percentages o f f i s h show homing behaviour. home  the  the  home  i s expressed.  pool.  In  inlets,  little  With decreasing t u r b u l e n c e and  i n c r e a s i n g r e g u l a r i t y of the t e r r a i n , decreasing numbers of f i s h remain i n the t r a n s p l a n t area. the reduced  high t i d e  areas shown by Green  Age,  These f i n d i n g s can be r e l a t e d t o  activity  of p. maculosus  (1971b,c).  length and y e a r - c l a s s d i f f e r e n c e s i n homing There  i s an i n c r e a s e i n the percentage  s u c c e s s f u l l y homing from age 1 to age 2 f i s h . d e c l i n e i n the percentage s u c c e s s f u l l y show pool.  equal  percentages  There  transplant  of  with  age.  c l a s s d i f f e r e n c e s i n homing Examination behaviour  by  percentage  of f i s h  cm. seems  homing  i s a d e c l i n e i n the area  of 0. maculosus  Age 3 f i s h show a  homing. fish  behaviour  A l l ages of f i s h  r e t u r n i n g t o the home  percentage  remaining  behaviour.  o f t h e age r e l a t e d d i f f e r e n c e s  smaller  size  i n the  There do not appear t o be any year  classes  shows  in  hcming  an i n c r e a s e i n the  s u c c e s s f u l l y homing with l e n g t h up t o about 5  Between 5 and 7 cm, homing to  i n turbulent  i s best  expressed  be some d e c l i n e i n the percentage  and  there  s u c c e s s f u l l y homing  247  with to  the l a r g e s t show  pool.  size  classes of fish.  egual percentages  There  transplant classes.  with  length,  Juvenile fish  moved down i n t o  lower  pools  is  and  o f homing f i s h  i s a decline i n the area  i t  long,  range  the  suggested  mechanisms There  the  paired  are  involved  "space"  suggest to  that  fish Both fish  i n homing.  i n homing  size just  between  they begin t o are  about  3  o f home  behaviour  that touch  There  i s  performance.  homing i s a d i r e c t e d  or taste  receptors  no  evidence  in  elsewhere  that  density,  i n the transplant  p o o l have  L i m i t e d e v i d e n c e was f o u n d t o movement from  the r e l e a s e  pool  pool. and o l f a c t i o n  are involved  in  homing,  a  o f b l i n d n e s s and a n o s m i a b e i n g t h e most e f f e c t i v e i n  are unable  t o home u n t i l  senses a r e e s s e n t i a l  Although  t h i s time  By t h e t i m e t h e y  homing s u c c e s s t o low l e v e l s .  but  have  and p o s s i b l y t a s t e r e c e p t o r s l o c a t e d  Both v i s i o n combination  which  the  behaviour.  involved  on homing  t h e home  reducing  at  or behavioural interactions  effect  largest  show e x t e n s i v e movement  that  i s no e v i d e n c e  fins  the  in  a r e b e g i n n i n g t o show some e v i d e n c e  f i d e l i t y and homing  Sensory  any  fish  for  appear  t o t h e home  remaining  {about 2.3 t o 2.7 cm)  tidepools,  classes  returning  percentage except  " l e a r n " a n d "memorize" t h e a r e a . cm  A l lsize  vision  I t i s suggested  one o f t h e s e s e n s e s  to t h e s u c c e s s f u l  homing  i s available. of  seems t o become u n n e c e s s a r y b e f o r e  i t appears  that  the  homing  fish  that the  juvenile olfaction.  recognize  some  248  attributes  of  pool, neither  the  home  pool, rather  conspicuous  visual  olfactory  clues  recognized  by 0. m a c u l o s u s .  importance of  emanating  of sensory  from  than the p o s i t i o n o f t h e  landmarks  in  home p o o l w a t e r  I t i s suggested  mechanisms may a l t e r  the  There morphometric studied, There  between  characters  were shown t o be  that  the  i n both  relative  t h e ontogeny  a  variable  d i f f e r e n c e s between  in different  differences  between  which c a n be r e l a t e d  is  fish  a r e no c o n s i s t e n t  p a r t s o f t h e body w i t h  0* m a c u l o s u s  t o exposure  increase  or  homing  classes  on a l l  i n t h e number o f c i r r i .  from  to  be  0. m a c u l o s u s i n d i f f e r e n t  by l i g h t  areas  behaviour.  i n t h e number o f c i r r i ,  age  unknown.  the  or  l e n g t h and a g e , b u t t h e r e a r e no a p p a r e n t  year  to  meristic  in  d i f f e r e n c e s i n t h e r e l a t i o n s h i p between c i r r i  related  areas  in  are  exposure.  T h e y do n o t p o s s e s s  There  and l e n g t h and  a r e a s b u t t h a s e do n o t a p p e a r  The  function  any s e n s o r y  of  the c i r r i i s  receptors  detectable  microscopy. ,  d e t e r m i n a t i o n i n 0. m a c u l o s u s Otoliths  determination vertebrae revealed the  nor  homing b e h a v i o u r a n d between a r e a s .  Morphological differences  Age  pools  shown  i n 0. m a c u l o s u s  and that  were  length  to by  be  a  valid  comparison  freguency  analysis.  method o f age  of  results  Ageing  a t any one t i m e t h e r e a r e two main age  p o p u l a t i o n with i n c r e a s i n g l y  from  studies  groups  in  s m a l l e r numbers o f a g e 3 and 4  249  fish.  Age-length r e g r e s s i o n s  areas  showed  related  for  d i f f e r e n c e s between  to a particular  factor.  0. l a o u l p s n s areas,  from  but these  different  could  n o t be  250  X.  A b r o s i m o v a , A. 1975.  M.  Study of f i s h o r i e n t a t i o n E n g l i s h summary]. Vestn.  Abrosimova, 1976.  Adler,  LITERATURE CITED  A.  in homing fin Russian, Zool, 1975(4):3-8.  M.  Role of sense organs i n o r i e n t a t i o n of carp-bream, z o p e , r o a c h and w h i t e bream i n the Kiev Reservior Tin Russian, English summary]. Vestn. Zool. 1976(3) : 40-44.  Helmut.  1971.  ftronson,  The d e v e l o p m e n t and e v o l u t i o n o f o r i e n t a t i o n : panel discussion. In Orientation: Sensory Basis. C o n f e r e n c e h e l d by New York Academy o f S c i e n c e s and American Museum of Natural History, 1971. Ann. N.Y. Acad. Sci. 188:393-407.  Lester  1951.  O r i e n t a t i o n and j u m p i n g b e h a v i o u r Bathygobius soporator. Am. Mus. 22.  Aronson, Lester 1971.  A t k i n s o n , C. 1939.  R. i n the Gobiid f i s h Novit. 1486:1-  R.  F u r t h e r s t u d i e s on o r i e n t a t i o n and j u m p i n g b e h a v i o u r in the Gobiid fish. 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Appendix  J  Descriptions  Major  of study  study s i t e :  situated  below  intertidal  a  end o f t h e b e a c h steep  sessile  there i s a  gravel is  beach.  rocky In  abundant.  shelf  the lower  Moving  up  O l v a s p p . , and Porph.yra p e r f o r a t a o c c u r , g i v i n g  t o t h e Fucjus z o n e i n  be  Beach  small  Hedophyllum  intertidal,  sites  First  At t h e s o u t h  Higher  APPENDICES  which  most  of  the  study  pools  the way  occur.  t h a n t h e F u c u s z o n e some p a t c h e s o f Ente^omo^pha s p . c a n  found. In t h e  Also  occurring  (especially  tidepools,  Prionitis  sp. i s  most  abundant.  a r e I r i d a e a s p p . , clumps o f P h y l l o s p a d i x s c o u l e r i  in  the  algae, p a r t i c u l a r l y  lower  pools),  Corallina  Cla^ophora  Vancouverensis,  sp.,  coralline  and i n t h e  higher  pools, G i g a r t i n a sp.. In  t h e F u c u s z o n e and l o w e r ,  californianus  occur.  t i d e p o o l s along with funebralis that  are  M. c a l i f o r n i a n u s some  are extremely present  e x t e n s i v e beds o f M y t i l u s  M. e d u l i s .  i s also  Littorines  numerous i n most s t u d y  include  Notoacmaea  present i n lower and  pools.  JB§£§21§»  Tequla Limpets  N. s c u t u m .  270  Collisella  digitalis  extremely  common,  ftnthopleura  and  and  C. p e l t a .  some  B. c a r i o s u s  xanthogrammica  abundant,  particularly  Pagurus  and  in  nirsutiusculus  Balanus  pools.  Some  Strongylocentrotus  and  Along species  being  primarily  and P h o l i d a e ,  families liparidae, Chadwick,  1976;  but  at F i r s t this  Jknoplarchus  Exposed  Pachena P o i n t , K i r b y  the  beds  goose  sessile  is  the  species families from  the  (Green, 1967;  The o n l y  was C l i n o c o t t u s  numerous.  Port  about  The most a b u n d a n t  were c a p t u r e d  t h e lower  barnacle,  abundant  as  Cape B e a l e ,  of the C a l i f o r n i a neck  from  26  fish other  globjeeps  0. s n y d e r i  and  occasionally. ,  Renfrew, Benson  Island,  Point  In a l l areas large  S.  about  the remaining  and Hexagrammidae  was n o t n e a r l y  sites:  and  oohraceus,  representatives  observation).  Gill  common  Island,  s t u d i e d was 0. m a c u l o s u s .  purpurescens  study  are  crabs,  occasionally  species  Beach r e g u l a r l y t r a p p e d  species  hermit  very  i n t i d e p o o l s , o f which  a few  Gobiesocidae  in a l l t i d e p o o l areas fish  of  with  personal  found, are  Pigaster  Vancouver  t o the f a m i l y C o t t i d a e ,  composed  be  i s  pools.  a r e commonly f o u n d  belong  Stichaeidae  i n lower  t h e west c o a s t o f  of fish  14 s p e c i e s  purpuratus  occur  The  P. g r a n o s i m a n u s  S§li3JE§£Sus nudus c a n a l s o be f o u n d .  drogbachiensis  can  A. e l e g a n t i s s i m a  lower  and  jlandula  intertidal  mussel,  Pollicepes  at these  i scharacterized  by  My.tilus c a l i f o r n i a n u s and pplymerns.  Hedophyllum  l e v e l s and P o s t e l s i a p a ^ m a e f a r m i s  271  occurs  on  Point,  exposed rocky  outcrops,  Moving  the  intertidal,  sp.  and  up  Endocladia  seaward s t u d y  pools,  californianus  shelves.  At h i g h e r  scattered  covering  at  these  fragile Beach  and and  corallines.  pools  Prionitis  themselves,  Phy.llospadix  are  Point  typically  i n a l l areas  purpuratus  Balanus glandula  included  tidepools.  cover  the  have  a  numerous  Beach.  scouleri, At  Codium  Botanical with  are  bare  comparatively  and  At B.  Clinocottus  the  fish  and  Pachena  l e v e l s i n and  tunicata around  at  embrxaj,  Gilbert.  the  present.  most a b u n d a n t f i s h  trapped C.  Point),  eleqantissima  Katherina  c a r i o s u s were  globiceps,  A s e l i c h t h y s rhodorus Jordan  at  pe I t a ,  higher  shores  i n c l u d e M. c a l i f o r n i a n n s ,  x a n t h o g r a m m i c a . A.  m a c u l o s u s was Other  exposed rocky  (particularly  Anthopleura  ochraceus.  0.  where  e x t e n s i v e l y covered  f o u n d on  at Kirby P o i n t ) , Col l i s e l l a  Pisaster  more  P o l l i c e p e s polymerus.  i n pools  droebachiensis,  the  Botanical  pools  Again, in  and  the  pools at Kirby  Point  L i t t o r i n e s are  Pachena P o i n t , The  Pachena  bare or  common i n a l l a r e a s .  Strongylpcentrotus  pools,  are  are  w h i c h were f o u n d  and  Kirby  £olymerus  sites  Pachena P o i n t  Invertebrates  (except  study  at  surrounds the  sp.  e x c e p t f o r clumps of  S.  the  Point.  (and  and  Pollicepes  E n t e r o m o r p h a spp.  l e v e l s a t both  I n the  at Cape B e a l e  and  levels, of  Fucus s p .  C l a d o p h o r a sp.)  except  Mytilus  except at Kirby  the 0.  species  study snyderi  sites and  272  Moderately  exposed  study  sites:  Helby  Island,  Haines  Island  (southeast side) Both  study  Fu cu s , Cladop_hora Haines and  Island,  mussel  and  at  and  spp.  few  the  Island,  areas  of the  lower  purpuratus.  At  B. c a r i o s u s and  N.  Fucus  O l y a spp. sites  Helby several  t o those Island,  Sheltered  Tggula  present.  with  At  algae  spp.,  at  at  include  Haines  both  Phyllosgadix  also occur.  and  littorines  at F i r s t  sites i s scouleri  calj. f o r n i a n u s .  A. e l e q a n t i s s i m a ,  digitalis,  Hemigrapsus  Mojealia s p p .  Island,  and  Invertebrates to  Myti,lus  funebralis  gravel  and  in a  Strongylocentrotus  h i g h e r l e v e l s i n both a r e a s , Balanus  qlandula.  a r e common.  or observed  Beach, except  at  these  that  on two  Gastergsteus aculeatus Linnaeus  sites  were  occasions at  were o b s e r v e d  in  tidepools.  study s i t e s :  I s l a n d , Dodger C h a n n e l , The  lined  scutum, C o l l i s e l l a  pools  high shallow  the  ,Halo§accign g l a n d i f o r m e  xanthogrammica,  F i s h s p e c i e s caught similar  are  predominant  Anthopleura  nudus, Pagurus spp.,  Amongst  occur.  Corallines,  Notoacmaea p e r s o n a ,  F u c u s zone.  of c o r a l l i n e s ,  i n pools at both  edulis,  l i e i n the  L e a t h e s i a d i f f o r m i s are  pools i n both  Haines  be f o u n d M.  also  shells  Prionitis  s p . , and patches  D i v a spp. The  sites  Grappler Inlet Haines  Island  s u b s t r a t e i n t h e bay  (bay and  mudflat),  Hance  (large tidepool)  i n Grappler Inlet  consists  of  273  boulders shell,  and  rocks  g r a v e l and mud.  Polxsiphonia sp,  of various s i z e s Clumps o f  collinsii,  a r e found  site  Rhodomela  larex  and  Pagurus  f o r Thais  Saxidomas g i g a n t e u s The  beached f i s h  nudus,  mudflat  boat  Pagurus  sp,  areas,  area  i s  somewhat  t h a t t h e Ranee  Balanus  glandula,  s p . , and  Pycnopodia  Notoacmaea  helianthodes  i s  but n o t a t Ranee I s l a n d and t h e r e v e r s e  i n Port marina,  and  shells  Desire supports covered  with  and s e v e r a l w a t e r l o g g e d of  The o t h e r  gchraceus,  sp,,  of  Tresus  capax.  and Cr ass o s t r g a g i g a s .  outcroppings  glandula.  Fucus  and E n t e r o m o r p h a  Ranee I s l a n d e x c e p t  lamellgsa  of e e l g r a s s . Zoster a  have  The  s p p . , Hemigrapsus  a r e common i n b o t h  true  edulis,  difformis  Gracilaria  common i n G r a p p l e r I n l e t is  o f crushed  i s m u d d i e r and t h e p r e d o m i n a n t a l g a e a r e F u c u s s p . ,  B, c a r i g s u s , persona  Leathesia  behind  a mixture  Utilus  on t h e r o c k s and s t o n e s .  s i m i l a r t o t h e channel Island  over  Mytilus  most o b v i o u s  sp.,  Pycnopodia  edulis,  Hopalia  a c o n s i d e r a b l e bed  S mi t h g r a  naiadum.  logs a t t h e study  A site  Fucus s p p . a n d B a l a n u s  i n v e r t e b r a t e s are  sp., Thais  Hemigrapsus  lamellgsa, Pisaster  h e l i a n t h o d e s , Cucumaj;ia m i n i a t a a n d T r e s u s  capax. The of sand  a n d mud w i t h  outcrops. and  There  are covered  o c c u r on r o c k y  on H a i n e s  small stones  I s l a n d have s i m i l a r  substrates  and embedded r o c k s  and  i s a s m a l l e e l g r a s s bed i n t h e l a r g e  i n summer months  Channel sp.  two s i t e s  this with  and  the  Smithora  patches  at  extensive  naiadum.  both  sites  bed  while  tidepool  in  Fucus s p .  rocky  Dodger  and Diva  Halosaccipn  274  gland i f or me the  large  i s common i n Dodger Channel and Enteromorpjia sp. i n tidepool.  B. c a r i o s u s  Mytilus  edulis,  are found i n both areas,  Pagurus spp., Hemigrapsus  sp. and  common  while  i n the tidepool  fiotoacmaea  Dermasterias  anjthc»plejjr.a  scutum.  imbricata are  elegant issima i s  numerous i n Dodger Channel. The  pool  .alanJfliS*  and s h e l l s o f T res us capax, Sa_xidom as gj,ganteus and  Protothaca staminea  extremely  Balanus  on  pool a t Ranee I s l a n d i s somewhat  Haines  similar  to the  I s l a n d with a muddy, sandy s u b s t r a t e and l a r g e  boulders s c a t t e r e d throughout. edulis,  Balanus  giandula,  Mopalia  sp. occur  Fucus  sp., Ulva  Notpacmaea  i n the  rocky  scutum,  areas.  sp., M y t i l u s N. persona and  £aau£us  sp. and  numerous  fish at  lemigrapsus sp. are numerous. 0 . maculosus  while  was  t h e most  c e r t a i n times of year i n a l l areas, r e l a t i v e to other f i s h Grappler  were more abundant i n s h e l t e r e d areas. Inlet,  Artedius  (Bean) and Nautichthys caught,  although  Gasterpsteus trapped  tidepool  fenestralis,  they  and  aselichthys  occasionally.  at  t h e mudflat  A. xhodorus  was  sometimes  and  were  nicholsi  frequently  were never as numerous as 0 . maculosus.  aculeatus  G. a c u l e a t u s  at the bay i n  Cory_phopterus  p c u l o f a s c i a t u s (Girard)  numerous,  Isopsetta  isQlepis  rhodorus  were  i n Grappler and  (Lockington)  At Ranee I s l a n d , L e p t o c o t t u s armatus  sometimes  in  greater  numbers  also Inlet,  a. f e n e s t r a l i s ,  o c c a s i o n a l l y caught. trapped  areas  than  G. a c u l e a t u s were f r e q u e n t l y caught t h e r e , and l e s s  were were  0 . maculosus. freguently.  275  Ph2i.i§ small  laeta  (Cope) ,  Porichthys  P.  notatus  H a i n e s I s l a n d and  at t h e  bed,  was  L.  armatus  ornata-  (Girard) ,  Girard. eastern  In  the  of the trapped  Clinocottus globicegs.  At t h e  w e s t e r n end  I § x a a i a S § o s decagramjus  ( P a l l a s ) was  a l g a e r e f e r r e d t o above, biology  such  Carefoot, 1977.  Kozloff, 1973.  Ricketts, 1968.  pictures  see  any  fenestralis  large  along  with  occasional bed,  the only  species  of  invertebrates  general reference  on  grass  of the e e l g r a s s  the  and  tidepool  Dodger C h a n n e l e e l  commonly  N o t e : F o r d e s c r i p t i o n s and  A.  caught.  and  on  intertidal  Douglas L t d . ,  Vancouver,  as:  Thomas. P a c i f i c Seashores. B. C. 208 pp.  Eugene  J.  J.  N.  Seashore l i f e o f P u g e t Sound, t h e S t r a i t o f G e o r g i a and t h e San J u a n A r c h i p e l a g o . J. J . , Douglas L t d . , V a n c o u v e r , B. C. 282 pp.  Edward  F. and  Jack  Calvin.  Revised  Between P a c i f i c T i d e s . Stanford S t a n f o r d , C a l i f o r n i a . , 614 pp.  by  J o e l Hedgepeth.  University  Press,  276  Appendix  2  Conversions  of t o t a l  length  (TL) t o and  from  T L to S L : S L = -.08132 + . 8 3 4 3 T L (N=58)  '  8.  T  0. S L to T L : T L  2.  4. 6. Total length (cm)  .1180 + 1 . 1 9 3 S L (N=58) 8..  0  - <> p  ' 2. 4. 6. Standard length (cm)  standard  length  (SL)  AEE£I!S\iZ 2 Alsae  an£ iHZSE*£b_ra.te.s f_E°JD e x ^ e x i n e n t a l  Exposed  iank Cain  side  Chloropybceae H l ^ a exp.ansa ( S e t c h e l l ) s e t c h e l l a n d G a r d n e r I H i e r c r i o r D h a l i n ^ a ( L . ) C. Aqardh £li'°2sis e l u i p s a (Hudson) C. Aqardh Scongosorpha g a x a t i l i s (Ruprecht) C o l l i n s  side  Chlorophyceae M l i a exgansa ( S e t c h e l l ) Entergmgr^ha sp. Link  Setchell  and G a r d n e r  Phaeophyceae N£E£Ocjystis l u g t k e a n a ( M e r t e n s ) P o s t e l s a n d R u p r e c h t H a o r o c v s t i s i n t e g r i f o l i a Bory I3£g3i3. ™ § S 2 i " s i i (Turner) Areschouq D££!i:2.iest i a l i g u l a t a v a r . I L g u l a t a ( L i q h t f c o t ) L a m o u r o u x L^rcinaria enhengra S e t c h e l l Laninaria saccharina ( L i n n a e u s ) Laraouroux Jis§2i3 a r b o r ^ a A r e s c h o u q Agarum s p . ( E o r y ) P o s t e l s a n d R u p r e c h t Rhodophyceae Ant i t h a s i n i o n J s i l i n i i Gardner P o i E l l I E a £ i n i a t a (C. A q a r d h ) C. Aqardh f H g d v x a n i a £ ° r t u s a ( P o s t e l s and Ruprecht) J . Aqardh C a l l o p h v l l i s f i i B i (Kylin) Norris l E l i i g a c o r d a t a (Turner) . Bory fauch°a l a c i n i a t a J . G. Aqardh *5tiiM2£ions22a g l i n d u l i f e r a ( K y l i n ) Hollaston P i a t X t h a - ^ n i on E t c t i n a t u s i K y l i n  Rhodophyceae C a l l o 2 h Y . l l i s firraa  Cirripedia BalaE-Us c a r i o s u s (?allas) .fialanus I Q ^ t r a t u s a l a s k e n s i s  Cirripedia Balanus carigsus ^alanus rgstratus  Pilsbry  Gastropoda S s i l i s e l l a p_Si£a R a t h k e i j o t g a c r e a Egrsgna Pathke iiStoacnea s c u t u n (Pathke) £ a l 2 i o s t g 2 a l i o a t u ^ Gould I i i i 2 E i D 3 . s s u l n l a t a r.ouid Iiit2rifi3. N n a s s e s ia£Una o a r i n a t a G o u l d .Lacuna J T n a s s e s £ 2 I S 3 l i i £ S n u p _ i l l u s (Gould) I l E l l I a r i a l i E l i l a t a (Carpenter) e c  ec  (Kylin)  (Pallas) alaskensis  Norris  Pilsbry  Gastropoda C Q l i i s e l l a a e l t a Rathke Ifotgacmga 2 2 i s g n a R a t h k e l i Q ^ o a c n e a Scutura ( R a t h k e ) £a 1 1 i g s ^ g n a l i g a t u n G o u l d I l i i 2 l i E a s c u t u l a t a Gould L i t t Q r i n a °qq m a s s e s Lacuna c a r i n a t a G o u l d L3.SUS2 ^ I Q i s s s o s " a r g a r i t P s Eiinillug (Gould) l i l ! l l 3 E i 3 l i r u l a t a (Carpenter) £2iatgstgma f o l i a t u m (Gra^lin) Velutina l a e v i g a t a Linnaeus Mflhissa coluiibiana Dall  ~J  Ascidiacea  Ascidiacea £°I°Il£ " i l l ^ e E i a n a  £°I!2ll2 " i l l u s r i a n a  Herdman  Bivalvia Chlamvs r u b i d a Hinds Hl'tilus g d u l i s Linnaeus H X t i l u s ca l i f o r n i a n u s C o n r a d Holothuroidea C u c u r a r i a o s a u d o c u r a t a Deichraann  Bryozoa  Bryozoa  Anph i p o d a Caprellidae  Arcph i p o d a Caprellidae  S  Bivalvia ^illaBYS rubida Hinds J5Xi-iIl!§ §3tilis L i n n a e u s i i l t i l l i s c a l l forr.ianus Conrad Anthozoa Antho2iSi!ra e l e g a n t i s s i m a Brandt Calcarea Scygha s p .  Caleerea Scycha sp.  Folychaeta Nereis procgra Ehlers 55il2rbis sp. Piatvn rsis bicanaliculata  Herdman  (Baird)  Polychaeta Eaoit^lla ^agitata (Fabricius) S p i r o r b i s sp. PI§£ynereis b i c a n a l i c u l a t a ( B a i r d ) N e r e i s n e o s j a z t h e s Hartman Harnothoo irabricata (Linnaeus)" Oliqocha eta Enchytraeidae Isoooda Manna s p .  

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