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Age and growth of the mosshead sculpin Clinocottus globiceps Girard with an assessment of its role in… Mgaya, Yunus Daud 1989

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AGE  AND GROWTH OF  GLOBICEPS  GIRARD  THE MOSSHEAD SCULPIN WITH  PRODUCTION  AN ASSESSMENT OF TIDEPOOL  CLINOCOTTUS  O F ITS  ROLE  FISHES  by YUNUS DAUD MGAYA B.Sc.  (Hons.), The University of Dar es Salaam,  1986  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE  in  T H E FACULTY OF GRADUATE STUDIES Department of Zoology We accept this thesis as conforming to the required standard  THE UNIVERSITY  OF BRITISH COLUMBIA  September  1989  © YUNUS DAUD MGAYA,  1989  IN  In  presenting this  degree at the  thesis  in  University of  partial  fulfilment  of  of  department  this or  thesis for by  his  or  requirements  British Columbia, I agree that the  freely available for reference and study. I further copying  the  representatives.  an advanced  Library shall make it  agree that permission for extensive  scholarly purposes may be her  for  It  is  granted  by the  understood  that  head of copying  my or  publication of this thesis for financial gain shall not be allowed without my written permission.  Department The University of British Columbia Vancouver, Canada  Date  DE-6  (2/88)  ABSTRACT  Age  otoliths  taken  during to  of Clinocottus  a n d growth  the period  verify  also  from  formula  ageing.  i n this  and  obtained  from  Girard  a  Data  from  study.  Growth  Gompertz  growth  were  tidepools  M a y 1988 to J u n e  otolithic  included  fish  globiceps  investigated  at H e l b y  Island,  1989. Length-frequency fish  collected between  rate  w a s estimated  curve.  with  the aid of  British  analysis  Columbia w a s used  1980 a n d 1987 were  using  Instantaneous  a  back-calculation  growth  rates  were  determined.  Results from  indicated  less  younger (about  than  that  one y e a r  age groups 9  C.  the  of  (about  m m per year).  globiceps age to  Growth  w a s obtained:  Lt  exp{1.58(l  26.7mm  Instantaneous of  *  growth  instantaneous  period The  that lowest  rates  growth  water  -  years  of  w a s composed age. G r o w t h  and declined  w a s described  by a  of  individuals  w a s faster  at older  Gompertz  for  age groups  model  a n d the  exp[-0.30t])}.  were highest  rates  for the 0 +  occurred during  temperatures  instantaneous  5  17 m m per year)  following equation =  population  reach  growth  a  rates  age class.  the spring  maximum occurred  a n d early  a n d food during  T h e highest  s u m m e r , the  is most  the  levels  fall  abundant.  and  winter  months.  The  age —length  relationship  relationship  for  C.  globiceps  of the species is described.  is  presented.  N o differences  ii  The  i n growth  length—weight between  sexes  as  revealed  by  length — weight  relationship  were  observed,  thus  the  expression described the length — weight relationship for C. globiceps  following  population at  Helby Island: W  =  1.5913  *  10~  L  5  3  -  1  5  5  2  Overall tidepool production with regards to C. globiceps  was assessed by direct  comparison  maculosus  abundant  with  production  tidepool cottid.  of  sympatric  Production  was  Oligocottus  estimated  by both  the  Girard,  an  instantaneous  growth rate and size-frequency methods. Annual production as computed by the instantaneous and  0.  growth rate method was 6.9 and 11.0 g/m /year 2  maculosus  contributed respectively.  33  respectively.  and  Estimates  was suggested  65% of  Young-  age  production  made by the  for  groups C.  size-frequency  for C.  (between  globiceps  and  1+ O.  globiceps and  2+)  maculosus  method were higher and it  that these estimates may be more accurate, since the method is  not affected by nonsynchronous cohort development.  Production was analyzed by zones on the intertidal area and the results reflected the distribution pattern of the two species, at the upper intertidal pools for 0.  i.e., higher production was observed  maculosus  and at the lower pools for C.  globiceps.  The  relationships  between  the  physical  characteristics  of  the  tidepools  and  production of the two species are given. None of the physical variables examined were significant predictors of production.  iii  TABLE  OF CONTENTS  ABSTRACT TABLE  ii  OF CONTENTS  iv  LIST  OF TABLES  LIST  OF FIGURES  LIST  OF APPENDICES  x  ACKNOWLEDGEMENTS  xi  I.  II.  vi viii  G E N E R A L INTRODUCTION 1.1 B A C K G R O U N D 1.2 D E S C R I P T I O N O F T H E S T U D Y 1.2.1 P h y s i c a l C h a r a c t e r i s t i c s 1.2.2 F l o r a a n d F a u n a  1 1 5 5 7  SITE  A G E AND GROWTH 2.1 I N T R O D U C T I O N 2.2 M A T E R I A L S A N D M E T H O D S 2.2.1 S a m p l i n g 2.2.2 L e n g t h M e a s u r e m e n t s 2.2.3 W e i g h t M e a s u r e m e n t s 2.2.4 S e x D e t e r m i n a t i o n 2.2.5 A g e D e t e r m i n a t i o n A g e D e t e r m i n a t i o n u s i n g O t o l i t h s A n a l y s i s o f L e n g t h - F r e q u e n c y D a t a A g e - L e n g t h R e l a t i o n s h i p 2.2.6 G r o w t h D e t e r m i n a t i o n M e t h o d o f B a c k - C a l c u l a t i o n o f F i s h I n s t a n t a n e o u s G r o w t h R a t e s T h e o r e t i c a l G r o w t h M o d e l 2.3  2.4  Length  9 9 11 11 12 12 13 13 15 16 17 17 18 19 20  RESULTS 2.3.1 S i z e C o m p o s i t i o n 2.3.2 O t o l i t h D e s c r i p t i o n 2.3.3 O t o l i t h A n a l y s i s 2.3.4 L e n g t h - F r e q u e n c y A n a l y s i s 2.3.5 T i m e o f A n n u l u s F o r m a t i o n 2.3.6 G r o w t h i n L e n g t h B a c k - C a l c u l a t e d L e n g t h s T h e o r e t i c a l G r o w t h C u r v e s L e n g t h - W e i g h t R e l a t i o n s h i p G r o w t h R a t e s  21 21 21 24 28 28 33 33 40 45 49  2.3.7 2.3.8  49 52  A g e - L e n g t h Relationship Total Length - Standard Length  2.3.9 S e x R a t i o DISCUSSION  Relationship  52 56 iv  III.  FISH PRODUCTION 3.1 I N T R O D U C T I O N 3.2 M A T E R I A L S A N D M E T H O D S 3.2.1 C o l l e c t i o n o f F i s h e s 3.2.2 A b u n d a n c e E s t i m a t e s a n d D e t e r m i n a t i o n o f A g e 3.2.3 P r o d u c t i o n I n s t a n t a n e o u s Growth Rate Method S i z e - F r e q u e n c y M e t h o d  64 64 66 66 66 67 68 70  3.3  73 73  RESULTS 3.3.1 D i s t r i b u t i o n  ,. within  the  Study  3.3.2 A g e o f Oligocottus maculosus 3.3.3 3.3.4 3.3.5 3.3.6 3.3.7 3.3.8 3.3.9  3.4 IV.  Area  Density Biomass Growth Production Size-Frequency Estimate of Production A n n u a l P r o d u c t i o n to B i o m a s s R a t i o s Estimation of P r o d u c t i o n from Initial B i o m a s s  DISCUSSION  GENERAL  LITERATURE  DISCUSSION  76 76 84 85 86 90 91 93 94  A N D CONCLUSIONS  CITED  102 106  APPENDICES  120  v  LIST OF TABLES Table  1. S u m m a r y Clinocottus  Table  Table  Table  statistics  globiceps  for standard  population  lengths  at H e l b y  a n d body weight for the  Island  23  2. S a m p l e size a n d m e a n length ( + S E ) males, sexes of C. globiceps of each age group  females,  a n d combined 27  3. P a r a m e t e r s of the component n o r m a l distrbutions identified i n the population length distributions b y the method o f M a c D o n a l d & Pitcher (1979) 4. S u m m a r y of standard length—otolith radius (y = a + b x ) a n d analysis of covariance between  regression statistics male a n d female C .  globiceps Table  30  35  5. Back-calculated lengths  at each age for combined data  of C.  globiceps 36  C. globiceps  Table  6. Back-calculated lengths  at each a'ge for male  Table  7. Back-calculated lengths  a t each age for female  Table  8. P a r a m e t e r  Table  9. A n a l y s i s  Table  10. S u m m a r y of m e a n growth i n length f r o m  estimates,  derived  from  37  C. globiceps  the Gompertz  growth  38  model, for  C. globiceps  Table  Table  43 of variance  statistics for G o m p e r t z  model  43  observed, back-calculated a n d theoretical the present study for male a n d female C.  globiceps 44  11. S u m m a r y of m e a n observed, back-calculated a n d theoretical growth i n length for combined sexes 12. L e n g t h — weight relationship  of m a l e ,  female  a n d combined sample  of C. globiceps Table  Table  Table  47  13. F u n c t i o n a l length — weight regression described as L o g W  a  +  44  = Log  (b/r) L o g L for C. globiceps  47  14. A n a l y s i s of covariance between sexes for length — weight regression and t-test for comparison of calculated regression coefficient with regression coefficient of the cube l a w  48  15. Instantaneous growth rates of C . globiceps in their age 0 + through age 5 + . Rates were calculated for the intervals between the 1987  a n d 1988 collections  50  vi  Table  16. S u m m a r y analysis  of age—length  regression statistics  of covariance between  male  a n d female  (y = a + b x ) and  C. globiceps  of C. globiceps by age groups  Table  17. S e x ratios  Table  18. A g e - l e n g t h data f r o m C h a d w i c k ' s (1976) study showing the discrepancy in the former  Table  19. A g e structure of Oligocottus maculosus from determined b y age —length regression  Table  20. M e a n annual density, production and production  maculosus Table  Table  Table  study study  a n d the  present 58  the present  study, as 77  initial biomass, m e a n biomass, annual to biomass ratios for C. globiceps and O.  statistics  for growth,  a n d zones for C.  22. A n a l y s i s of variance production b y years  53  79  21. A n a l y s i s of variance production b y years  Table  53  statistics  for growth,  and zones for O.  density,  biomass and  globiceps density,  80 biomass and  maculosus  80  23. S u m m a r y of regression analysis variables (defined in text). Correlations of each set of variables were tested for significance 24. A n n u a l production, m e a n biomass and P : B ratios as estimated instantaneous growth rate method a n d size-frequency methods  vii  83 by 92  LIST Figure  1. M a p showing the general  Figure  2. Photograph  Figure  3. Length-frequency  Figure  Figure  Figure  Figure  FIGURES  location  a n d female  of study  Clinocottus  site  6  globiceps  14  o f C. globiceps  distribution  22  4. (a) Illustration of the l a b y r i n t h , showing the position of the otolith c h a m b e r s , (b) otolith of C. globiceps showing opaque a n d hyaline zones 5. Length-frequencies age  Figure  of male  OF  of male  a n d female  C. globiceps  b y estimated  groups  26  6. Length-frequency distribution A p r i l 1987 a n d October 1988  of C. globiceps  7. Length-frequency distribution f r o m otolithic ageing  of age 0 +  8. Length-frequency of modes  sampled  between 29  through  5+  fish  derived 29  distributions  of C. globiceps  showing progression  .  Figure  9. Otolith  Figure  10. Relationship  marginal  31 increment  plotted  of saccular otolith  b y month  radius  to  for the three :  fish  standard  age groups ...32  length for  C. globiceps  34  Figure  11. G o m p e r t z  Figure  12. L e n g t h — weight relationship  Figure  13. Instantaneous  Figure  14. A g e —length  relationship  Figure  15. Relationship  of total length  Figure Figure  Figure  growth  function  growth  for C. globiceps  of C. globiceps  distribution  17. Length-frequency  distribution  42  of C. globiceps  curves for age 0 +  16. Length-frequency different levels different  25  to standard  46 through  determined length  of C. globiceps  3+  C. globiceps  b y otoliths  a n d vice  from  versa  .51 54 55  tidepools a t 74  of O. maculosus  from  tidepools at  levels  75  18. D e n s i t y of C. globiceps a n d O. maculosus zone for the period 1 9 8 6 - 8 7 to 1 9 8 7 - 8 8  viii  b y age group, i n each 81  Figure  19. Age-frequency and  Figure  Figure  Figure  distribution  curves for populations  from  Island  Helby  of C.  globiceps 82  20. Production of C . globiceps a n d O. maculosus each zone for the period 1 9 8 6 - 8 7 to 1 9 8 7 - 8 8  b y age group, i n 88  21. Relationship between production a n d initial biomass of 1987 a n d 1988 y e a r classes of C. globiceps a n d O. maculosus 22. Relationship year  Figure  O. maculosus  classes  between  production  of C . globiceps  23. Relationship  between  a n d density  of 1987 a n d 1988  a n d O. maculosus  P : B ratio  ix  and mean  89  97 biomass  (B) of C.  globiceps 100  LIST OF APPENDICES Appendix  1.  Length-frequency  distribution  table  for  combined d a t a  of C .  globiceps  120  Appendix  2.  Length-frequency  distribution  table  for  female  Appendix  3.  Length-frequency  distribution  table  for  male  C. globiceps sample  of  121  C. globiceps  122 A p p e n d i x 4. A g e — l e n g t h key for H e l b y Island n u m b e r of fish per age-class b y sex  C.  globiceps,  showing  the  A p p e n d i x 5. M e a n annual density, initial b i o m a s s , m e a n b i o m a s s , m e a n instantaneous growth rates and a n n u a l production of C. globiceps  O. maculosus growth  rate  b y age method  groups as computed b y  the  123  and  instantaneous 124  x  ACKNOWLEDGEMENTS  This  thesis  Special  would  thanks  encouraging  not  are  me  programs  study  highly  is  Drs.  Timothy  guidance  has  Heggenes  for  My  sincere  Marine  R.  work  fishes  His  would  support  like  to  this  assistance in  thanks  are  of  for  the  financial  support  and  would  to  during  Centre  Ms.  John  Canadian  well  as  financial  duration  all  Alistair  thank  Dr.  field  his  J.  my  every  his  are  people.  data  phase  supervisory  Antony  Thanks  many  Wilimovsky,  making  through  and  and trips.  and  Blachford Matseliso  E.  for and  of  this  committee,  R . E . Sinclair also  Mclnerney,  Universities staff  support  of m y  Wilimovsky  Computing  Nightingale  as  the  entire  N.J.  to  thesis.  from  due  whose  to  Dr.  Jan  of  Bamfield  field.  Western  support  the  Dr.  due  the  for  thank  improved  his  help  Norman  and  considerably  to  like  I  me.  the  Dr.  G . E . Scudder  grateful  Columbia  supervisor,  tidepool to  without  Geoffrey  acknowledge  Scholarship  my  on  available  about  Parsons,  logistic  gratefully  come  to  appreciated.  Station  extensive  due  to  computer  have  The  of  provided Morapeli  thesis.  xi  a  staff  Data  for  the  University  technical  typing  the  I  Commonwealth  Station  Centre,  for  LaCasse.  Canada.  Marine  of  essential  in  Society,  Steve  Canadian  studies  Bamfield  Biosciences  Biological  particularly  graduate the  Director  I  am for of  their British  especially  support.  also  Jon  Lastly,  manuscripts  of  I  this  I. G E N E R A L  1.1  BACKGROUND  Cottidae some the  (commonly  known  as  species  in  the  marine  Cottidae  are  demersal  50  Family  from  the  Wilby  The  subtidal  to  the  throughout  tidepools the  on  the  much  waters in  intertidal  of  Pacific  G u l f of A l a s k a  about  190  mm  ecological data  a  large  of  B.C.  habit  and  zone  of  the  diverse (Hart  family  1973).  occupy  Most  diverse  northern  represented  by  members  habitats  hemisphere  of  ranging  (Clemens  &  in have  its  range  (Green  coast  of  North  (Miller  &  Lea  Girard 1971b).  America 1972).  total  length  (Hart  been  published outside  1857  Its  1973). of  It  from  is  is  found  the  size  this  manuscript  abundant  rocky  California  is  reported  wide  reports  intertidal  inhabiting  central  maximum  Despite  an  coast to  be  distribution with  few  regards  to  species.  Several habits  investigations of  a  contents  of  studied  food  Girard three  is  Clinocottus globiceps  mosshead sculpin,  this  sculpins)  1961).  cottid  to  INTRODUCTION  and  few  marine  three habits  0.  intertidal  have  conducted  species of Cottidae.  species of  been  of  two  snyderi Greeley,  tidepool  sculpins in central  Mitchell  sculpins  cohabiting and  concerning  tidepool  Yoshiyama  California.  1  in  age,  (1953)  analyzed  California. Cottidae,  (1980)  growth,  and  the  Nakamura  Oligocottus  investigated  food  stomach (1971)  maculosus  food habits  of  2 Moring  (1979,  tidepool  1981) 0.  cottids,  investigated  factors  aspects  life  have  of  also  with  Green  (1971a,  respect  (1978)  on  spatial  (1979)  on  (1984)  only  in  Miller  this  species.  &  species were age  Moring  The  tidepool  Green  maculosus on  (1972)  and  fishes  1971a,  and this  C.  temperature  et  specifically data  on  limits  Green  al.  1985).  have  their  1971b; R i c h k u s  information  exhibit it  Enophrys  with  two  1976b)  species.  of  and  salinity  distribution, biology  Other  tidepool  distribution  bison;  similar  (1976a,  and  three  gradients.  and  DeMartini  cottids  of  include  hemilepidotus,  was  Clinocottus  sensitivity of  this  to  homing  DeMartini &  Patten  Grossman  movement,  but  home 1978;  this  are  been  close  &  and  distribution  Chadwick  has  range  Clemens  taxonomy  and  1973).  globiceps temperature  species;  essentially  (1971b,  maculosus  globiceps,  the  very  &  synderi.  behaviour,  0.  reproductive in  the  movement,  Hemilepidotus  of  summarized  homing  Freeman  to  southern  established b y  1979;  Resident  Based  Lea  to  sensitivity  on  two  behaviour  analyzed  O.  dealing  and  of  1961)  studied  of  Nakamura  distribution  reproduction  biology  dynamics  (1960,  Studies  presented  C . globiceps  of  1967;  who relation  and  of  on female  publications  1961)  gradients  aspects  snyderi.  1973)  sculpins.  reproductive  deVlaming  (1960,  tidepool  O.  reproduction  Morris  1971c,  and  vertical  including  their  structure  and  influence  to  1971b,  of  The  that  reported.  behaviour  age  maculosus  history  been  cottids,  analyzed  by  Morris  and  salinity  Wilby  (1961)  distribution patterns  (1976a)  questioned  of  the  estimated  the  (Craik  1978;  to  the  tidepools  G r o s s m a n 1982)  and  some, notably  strong homing behaviour considered  feasible  to  (Green launch  of  1971a, a  (Gibson O.  1973).  study  on  3 tidepool  fish  a  basic  parameter  it  links  population  The  production  objectives  the  life  this  dynamics maculosus.  Two  C.  Island.  globiceps  biomass  statistics,  performance studies  have  is  Although  been  &  shore  estimates, provide  of  1972)  conducted Although  is  there  considerable is  very  combined  w a s known  about  age a n d growth  to  of  evaluate  production  distributed  Oligocottus  with  that  most  data  rate  fish ranging  on  of  found  at  the  been  (Le C r e n  fish  intertidal production  northern  documentation  California  great  abundance  west  coast  of  mortality,  1969).  Most  a n d the relevant  structure for  in  Vancouver  density,  and  assessing  the  fish  production  literature  in  this  aspects  of  the  in the present  the  study.  in  of  extensively,  of  production  means  ecosystems  adopted  for  from  age, growth,  understanding scanty  chosen  reasonable  reviewed have  were  Island  in freshwater not  on the  widely  a n d is  combining  the  little  was  hypothesis  1972).  tidepools.  Helby  by  since  the  maculosus,  Lea  used in such studies  there  communities,  was a  similar  with  of a species in its environment  enormous.  methodology  test  First,  of ecosystems because  (Le C r e n  objective  is a common intertidal  rocky  Production  to  a n d O.  S e a (Miller  the  second  production  species  rates  investigation  comparison  in physically  maculosus  at  The  on  a n d the ecology  two-fold.  globiceps,  in  in the ecological sense, is considered  and survival  were  conducted.  these  Bering  tidepools  study  globiceps  of  species,  the  area  C.  dynamics  growth  of Clinocottus  be similar  Oligocottus to  this  Information  distributions would  density,  was  of  Production  in population  of  history  species  dynamics.  of  study.  marine  production  fish  estimates  4 completed  on  individual  environment.  For  production of  Clinus  South an  South the  example,  fish  Africa.  Pacific  globiceps  Apart  from  of  including  age  for c o m p a r i s o n .  This  thesis  made  of  is age  to  density, tidepools together  composed and  estimate  Relationship and are the  In  growth  were  between growth)  growth  two C.  of  annual  and  and  and  sections. In of  parameters  sections is presented.  was  The  the C.  has  been the a  included in  first  a  depth general  and  of  Peninsula,  biology basis the  for  of  for  C. the  production  deals  with  attempt  O.  attributes  coast of  published  section  and  and  (biomass)  Cape  formed  globiceps  (e.g.,  Finally,  estimates the  tidepool  budget  second section, an  population  a  southwestern  aspects of and  in  energy  on  nothing  study,  various  discussed.  an  of the  pools  maculosus  globiceps.  physical  on  examined  major  production  production  analyzed two  of  rocky  this  (assemblage)  quantitative  studies, virtually  production d y n a m i c s . O.  of  reported  gave  intertidal  America.  and  taxocene  tidepool resident  (1984)  the  these  North  study  aspects  in  fish  (1984)  a  Griffiths  taxocene  coast  investigation  &  entire  Bennett  superciliosus,  A f r i c a . Bennett  entire  species or  (e.g.  surface  discussion  is  maculosus. biomass, area)  of  bringing  5  1.2 DESCRIPTION  OF THE STUDY  SITE  1.2.1 Physical Characteristics  The  study  site  Island  near  Helby  Island  Columbia.  the  consists  situated  The  typical  site  to of  of  bench The  marks height  from  tide  study to  along  with  surface  The  An  Deer  for  group  the  bench  a  rocky  (48°51'  N,  mark  given identifying level notes  pools  was  site  for  obtained the  into  to  width  four  the  vertical  Vancouver  Island.  Sound,  British  stretching  from  Figure  The  habitats.  several  1). The  the area  substrate  pools have  patches  isolation from  surveyed.  determine  the  zero  level  tide  zones  height  data.  physical  and length)  vertical  W;  pressure. A l l  levelling  general  shelf  extensively  barometric  from  Helby  Barkley  site is its  used to  relative  in  at  human  long series of fish collections.  was  symbols. T h e  describing  fall  of this  tidepools  coast of  intertidal  cobblestones although  study  rocky  1 2 5 ° 10'  rocky  advantage  corrected  west  is  previously established were the  the  of  Islands  inaccessibility, and a  at  series  of  semi-protected and  a  on  study  beach  dimensions (perimeter,  study  was  Station  the  important  information  tide  in  boulders  zone  of  were  zero  to  comparative  intertidal  Marine  selected  open  of s a n d y bottom.  The  investigation  northwestern  mainly  habitation,  this  Bamfield  is  northeastern is  for  A  level was  tidepools  total  features,  data  determined used in  were  maximum  20  of tidepools.  of each tidepool  These  of  the  relative recorded  depth,  and  of each tidepool.  along  the  shore:  upper,  middle,  Figure  1.  Map  showing the  (adapted  from  general  Craik  location  1978).  of study  site  7 lower 0.6  and  m  base.  and  from  positions  from  elevations  are  and  The 0.6  1.9  m  and  from  about  to  to  of  2.3  The  to  Predominant  organisms  sp.  corallines,  is characterized  Invertebrates edulis;  sea  Notoacmaea  Fucus by  that  anemones N.  pools located cariosus  0  m.  to  area  dominates.  to  to  positions  "middle" 3.5  m  and  with  from  degrees  maximum  seawater  —0.3  to  refer  to  "upper"  respectively.  different  Annual  the  in the  tidepools family  algae  scouleri kelp  the  These of  high  tidal  emergence tides  about  temperatures  and  upper are  are  and  shore forms.  range  pools  occurs.  is  The  sp.  Prionitis lower  digitalis;  crabs  Pagurus  middle  levels,  A.  and  elegantissima;  spp.,  barnacles  M.  limpets  Tegula  and  spp.,  sp.  californianus  gastropod  and  intertidal  sp. and Hedophyllum  and  Moving  Cladophora  mussels Mytilus  Collisella  hermit  the  also  Laminaria  rocky Fucus,  in  xanthogrammica  nudus; the  typical  Amongst  pools include  scutum,  at  were  predominant  beds of the  and littorines  found  study  Anthopleura  Hemigrapsus  belong  2.3  semidiurnal  about  The  occur in  spp.  species  sp.  large  persona,  Commonly  while  represent  spp.; Phyllospadix  crab  and B.  the  present.  grapsid In  they  mixed  tides  in  Fucus  are  and  from  refer  Fauna  intertidal,  Leathesia  respectively, and  is  "lower"  12.0°C.  and  the  m  and  because  low  1.2.2 F l o r a  up  m  tide  extreme 6.7  "base"  1.9  interest  submergence.  3.96  terms  funebralis;  chitons  Balanus  Mopalia glandula  common.  about  Cottidae.  26  The  species remaining  of  fish  species  of are  which  about  primarily  14 from  8 the  families  Liparidae,  Stichaeidae Gobiesocidae,  communication).  maculosus numbers  The  whereas  in the  and  Pholidae,  and  most  with  Hexagrammidae  abundant  fish  Clinocottus globiceps  middle  and  few  lower  tidepools.  in was  representatives (Dr. the  N.J. upper  regularly  from  the  Wilimovsky, tidepools  caught  and  is  families personal  Oligocottus occurred in  high  II.  2.1  INTRODUCTION  Age  structure  As  a  a  strong  result,  and a  growth  bias  towards  coast of N o r t h  species,  1979),  O.  synderi  and  small  sample  great with  length  age  at  fish  has  scales  not  weight  generally  concretions 1971)  and  maturity,  and  other  been  susceptible to  alteration  &  et  (1976a).  The  of confirmation  populations, Tesch  been hard in  based on  the  utilized  to  resorption  once formed  a  fish  (Mugiya  (Campana  have  validation)  stated  been  that  of  "age  and  production". A g e  of  the  body.  for  Watabe  1983a);  and  9  of  the 1977);  they  on  are  (Wells  rings  the  they  work.  is  determination appear  otoliths ear,  of  in  in on  (calcium  Lowenstein  reasons: they  undergo  available  to  conjunction  which  inner  following  due  growth in  a  stock composition,  Historically,  labyrinth  reliably &  growth  ageing  on  Moring  analis  data,  growth  of  1978;  and  information  presence  fishes  is questionable  age  wide  conducted on  Craik  of the  with  the  of these  Clinocottus  study  knowledge  subject but  give  membraneous age  growth  1985),  populations.  Despite  1939;  latter  fish  on the  can  the  structures  al.  (or  (1978)  measurements,  and  (Atkinson  Freeman  lifespan, mortality,  deposited  have  Studies  1981;  of  species.  limited.  fish  Bagenal  harvestable  maculosus  study  up  A m e r i c a is  sizes and lack  importance.  built  age  Chadwick  of  been  the  on the  (Moring  investigations  has  in  information  Oligocottus  C . globiceps  1986)  all  fishes,  notably  features  commercial  the few  essential  of literature  of cottid  Pacific  are  multitude  distribution  In  A G E AND GROWTH  no  are  chemical  species  which  10 lack  or  tested  have by  small  Six  &  scales Horton  (Six  &  Horton  (1977),  otoliths  1977).  Of  were  the  the  25  different  superior  structures  structure  for  age  determination.  Chadwick  (1976a)  populations along  at  northern  limited  sample  California)  that fill  two  used  vertebrae  locations,  California. sizes were  attempts  to  common  intertidal  structure  and growth  one  to on  However,  (i.e.,  41  at  assess  fish,  the  of a  age  southwestern Chadwick's  Port  Renfrew,  collected during a  some of the  the  gaps in primary  one-week  knowledge objective  C. globiceps  structure  Vancouver  study B.C.  was  Island  plagued  and  10  period in J u l y concerning the being  population at  to  at  and by  another relatively  Bruels  1973.  Island,  Point,  This  life history  present  Helby  C . globiceps  of  data B.C.  study of  this  on  age  11  2.2  MATERIALS  2.2.1  globiceps  intertidal  tides  a  with  sparse  complete  be  crevices,  sampled  Fish  specimens  an  search  cover  or  undoubtedly fixed  transferred  to  study  in  37.5%  pools  collected  over  included  in  the 1986  collected in  few  from  low  of  (n=105)  sampled years study. and  all  from  yield  rocks many  and  permanent  They  additional  storage  February same  comprised (n=164).  or  all  site  and by  1980 In  addition,  the  pools fish  numerous fish  were and  laboratory.  N.J.  (n=85),  where  freshwater  October,  Dr.  of  and  in  low  terminated  Collected  washed the  bailing  was  removed,  in  during  specimens. In  detection.  later  throughout  tidepools  molluscs, barnacles  formalin  the  scattered  Sampling  could be  avoided  from  the  dipnets.  fish  between  1987  to  tidepools  using ichthyocides  with  with  the  buffered  by  failed  where  27  sampled  tide)  pool  pools  set of  were  isopropanol for  past  a  collected  the  pools  10%  currrent  1988.  of  rocky a  of  then  in  were  the  hour  were  collected.' In  immediately  (n = 225),  were  within  Specimens  after  could  zone.  (usually  possible.  The  METHODS  Sampling  Clinocottus the  AND  1988.  Wilimovsky  1981 306  Material were  (n = 64),  1983  specimens  were  12 2.2.2  Length  Length  and  production.  Measurements  weight Total  as  the  distance  to  the  position  distance Lagler  measurements  formed  the  and standard lengths were from of  the  anterior  maximum tip  most  from  the  head  to  the  1958).  All  measurements  to  the  end  standard  posterior  were  for  estimating  measured. Total  tip  extension;  basis  the was  the  hypural  of  to  the  nearest  was  tail,  length  end  made  of  length  growth  measured  when  measured plate  0.1  and  pressed as  the  (Hubbs  mm  using  & dial  calipers.  Shrinkage after when  of  preservation compared  otherwise,  2.2.3  Weight  the  in  to  considered to  alcohol than human  be  in  error  negligible  formalin  in  (Parker  (Shetter  measurement  has been used in  all  1963)  1936),  (Balon  since it  and  1974).  a  minor  Unless  analyses throughout  is  less bias  specified  this thesis.  Measurements  fish  were  nearest  weight  was  s t a n d a r d length  Individual to  fish  0.05  against  weighed g  after  length  was  using blot  a  Sauter  d r y i n g the  calculated  balance.  Measurements  specimen. A  using  logarithmic  regression  equation  were  made  regression of of  the  form  the  slope  b  were  b W = aL ,  where  (regression  W  is  coefficient)  calculated  by  using  estimation  (Pienaar  a &  the  weight  and  a  is  computer Thomson  (gm), a  is  constant.  program 1969).  L  for  the The  length  (cm),  parameters  nonlinear  least  b a  is and  squares  parameter  13 The  length — weight  for  combined  females  sexes.  were  between  relationship Slopes  compared by  sexes  before  was  (b  examined  values)  Sex  Sexes  pooling  the  data  18.0 of  easily  of  mm  a  of  attempted  distinguished  genital  long  lack  papilla  (standard  in  smaller  Age  Because  equations  a  for  common  for  and  males  and  significant differences  regression  significance level of  females,  equation.  All  a=0.05.  or  of  this  penis can  the  (Figure  for  Males  ratios  are  characterized  2).  However,  males  be  mistaken  for  sometimes  genital  specimens. Sex and tested  species.  papilla. were  Therefore,  calculated  significant differences b y  smaller  females sexing  for  all  by  the than  because was  not  collections  of  chi-square analysis.  Determination  mosshead  sculpins  statistical  approach  analysis,  MacDonald  counting  in  length)  development  Clinocottus globiceps  2.2.5  for  males,  Determination  were  presence  regression  for  analysis of covariance to test  statistical inferences were based on a  2.2.4  of  separately  annuli  in  based &  have on  Pitcher  otoliths  no  their  length-frequency 1979),  (Brothers  cross-validation technique of the  scales,  latter  and  1987).  by The  method.  age  was  distributions an  otolithic  former  determined  (distribution approach  method  was  by  a  mixture based  used  as  on a  14  Figure 2. Photograph of male and female Clinocottus  globiceps.  15  Age  Although  Determination  each  used is the calcareous which  sagitta  has  probably cause  the  slight  removal,  Otoliths  from  preferred  to  density  illumination obliterated  the  differential  opacity,  sacculus  is  laid  Less  were  ocular  found  peripheral while  that  circuli. the  while  placing the  immersed  a  in  ear  likely  attached  one  (see  ordinarily  Figure  layers,  dependent  definite  a  4).  A  process  on  bands  source of black  water  region,  light  at  background.  and  the  with  The light  could  light  the more  interpreted.  an Both  one  with  annuli.  The  the angle  a  most  though  although  before  whole-view  reflected  could still be  tissue  made  transmitted  central  readability,  the  against  the  concentric  produce  therefore  In  frequently  otolith  in  micrometer.  light,  the  of  thin,  observed with transmitted  achieved b y  side,  inner  factors,  which  cleaned are  an  was  improved  down  known  each  of the  sections. Readings were  with it  on  food  or  (annuli)  in  1983b).  transverse  and  three  variations  globiceps  equipped  (X30)  the  otolith  otoliths C.  otoliths,  in  continuous.  species ( C a m p a n a  After  Otoliths  six  laid down  concretion,  is  seasons, most  teleost  using  best of  saccular greatest  examination  was  dissecting. microscope critical  factor  not  used  be  periphery opaque  Varying  or  50  otoliths  as  it  exhibited when  illumination was  usually  degrees were  definition  was  than  the  resolution 40  examination.  was  above  examined used  for  annuli counting.  Opaque  bands  remainder  of  were this  counted  thesis  to  as refer  to  rhythmic  term  "annulus"  growth  is  increments  used or  in  bands  the on  16 otoliths;  it  events. All  is  assumed  Criteria  rings  were  it  was  or  four  n+  different  in  age  and  parts  possible to  the  established  were broad  crowded on other  that  these  otolith.  that  a  at  annuli  study  anterior  count the  A c c o r d i n g l y , all beyond  as  but a  same  30  otoliths.  narrow  number  most  annual  starting  groups were  the  with  of  tip,  expanded tip  least  growth  coincide  subjective  on the  W i t h the  otolith. some  these  making  rings, or  places on the  indicating  after  of  reasonably distinct  of the  trace  formation  and point,  at  three  designated  recent  ring  as had  occurred.  Ages  were  between  the  eliminate and (in  in  determined readings  bias,  were  otoliths  random  micrometer  from  order. units  and  complete  resolved b y  were The  three  examined  following  series  of  readings.  further  examinations  without  prior  measurements  converting  to  distance  from  both  reference  were  millimeters)  of  to  taken  with  an  Disagreements otoliths.  length  from  or  each  ocular  To sex  otolith  micrometer  (X30): 1.  Otolith  radius  rostrum, 2.  Size  of  Analysis  As  further  a  otoliths,  the  the  the  nucleus  to  the  outside  rim  of  the  and annulus  each growth  —  of  —  the  distance  from  the  nucleus  to  the  outside  edge  of  band.  Length-Frequency  verification  of  length-frequency  Data  age  groups  distribution  in  the  was  population analysed  as  using  determined the  by  computer  17 program Pitcher  designed 1979;  of  fit  are  individuals.  obtained  from  April  1987  October  'The a  to  for  covariance  was  between  the  of  to  polymodal  data  basic assumption of the  normal  distributions,  mixtures  collected  during  were the  (MacDonald  program  for  is that  example  interpreted  sampling  of  from  period  & the  different  age—length  spanning  from  1988.  relationship  allows  predictive  sexes  from  performed  for  regressions  data  to  age  obtained  determine  sexes, and if not,  they  determination of  from  whether  were  age  of  on  length  otolith  there  pooled into  any  length  were  calculated  readings.  were one  given  of  Analysis  significant  of  differences  age —length regression.  Growth Determination  Growth  in  this  Measurements used in  growth  skeletal  hard  study  of  The  size  throughout  is  growth  defined zones  determination. parts  must  back-calculation of age 1.  The  resulting  otoliths  Ordinary  separately  distributions  Relationship  age —length  2.2.6  The  Age—Length  species.  1980).  mixtures  data  normal  MacDonald  length-frequencies cohorts  to  and  of  the  its  entire  be  the  (spacing  Three  and  lifespan.  change  between  of  size  annual  assumptions inherent  satisfied  growth  fish  as  before  (Van  Oosten  size  of  hard  a  body  of  a  marks) in  fish  with  age.  on  otoliths  were  growth  studies  using  part  can  be  used  for  1929): part  must  be  closely  related  18 2.  The  annuli  and at 3.  used for  of  Standard  length  Regression  2.  were  Annuli the  otolith  (marginal to  "the  of  and  Having  tested  was  life history Ln  =  O R n (Lc  how  from  different  age  globiceps  by  otoliths  otolith  y = a+bx  formed  year  in the  radius  were  analysis  and  recording the  of  The  in  term  otolith  monthly  outer  edge  to  once  yearly  classes  (by  of  the  at  an  The  +  way:  proportionality.  separately  covariance  to  change  the  each age  in  last  increment'  for  males  determine  whether  distance  between  individual  body  length  complete in  this  addition)  (Thorogood  were  of Fish  the  marginal  previous y e a r ' s b a n d "  large  S)  following  ascertain  calculated  of  'marginal  size  calculated with the -  be  hyaline  instance  consequent  zone refers -  to  the  1987).  compared for  various age  classes.  Length  above prerequisites, back-calculation here  determined. was  C.  form  of Back-Calculation  determining  1983)  for  by  increment).  the  must  different.  Back-calculated lengths  Method  given  compared  margin  increase  of  the  verified  completion of the 3.  analysis  time.  regressed on  statistically  were  a  tested  was  lines  females  they  same  at  growth  agree.  assumptions were  and  the  length  population m u s t  1.  and  approximately  Estimates  These  age  fish of  was  the  at  fish  defined  some at  any  as the process  previous  age  previous  age  (Smith in  its  following equation:  S  ORc where  Ln  and  Lc  refer  to  the  lengths  (in  mm)  at  age  n  and  capture  19 respectively, and  S  is  radius  One  the  the  1975;  length  slopes  of  versus  weight  describing to  =  where  Tesch  (in  mm)  regression of  at  age  standard  n  and  length  capture,  on  otolith  1978).  for  lengths  individual  =  to  relations  slope  (Ricker  1973).  between  variability  is  Fish  the  are  (Ricker  slope  (b/r)  estimates  (G) (In  calculated  the  were  for  relationship  for  geometric  Functional  weight  (Ricker  y  (b/r)x  were  estimate  from  -  rates  length—weight  y-intercepts  in weight G  of the  radii  estimation  the  of  differences,  growth between  Rates  the  length  natural  derive  These  otolith  back-calculated  growth  for  coefficients  y-int.  of  Growth  correlation  to  mm)  increments  instantaneous  subject  to  lengths-at-age.  Regression  for  refer  (in  uses  Instantaneous  Monthly  ORc  Bagenal &  primary  Annual  successive  and  y-intercept  (Ricker  of  rates.  ORn  and  1973).  individual were  mean  divided  functional  regressions  length  because  year  are  both  classes. by  their  regressions  recommended variables  are  F u n c t i o n a l regression slopes were used  equation  1973). employed  to  calculate  instantaneous  rates  of  growth  as: L  -  2  r=correlation  regression  slope  of  regression;  L^mean  In  L,)  (Ricker  coefficient weight length  on at  of  1975), the  length; time  t;  length — weight  b/r = slope and  L  2  of  the  =mean  relationship; functional  length  at  b ^ ordinary  length — weight  time  t+1.  20 Theoretical Growth Model  Attempts  were  made  Bertalanffy  and  pattern  growth.  some the  of  assumes  Bertalanffy  model  data  on both  von  length-at-age  growth  an  rate  tends  inflexion  (Ricker  The  describes only  part  an  of  situated  of  the  inflexion  growth  von  The  a  size  differ the  grow  the  length gets  Gompertz  the  curve  towards  model is  to  also  markedly  well  below  half  the  the  sense  that  von  in  curve  beyond  G o m p e r t z growth  point, including the  the  fish  closer  but  at  of  models,  generalized description of  size.  models  portion  two  that the  h a v i n g decreasing curvature.  sides  a  asymptote,  two  the  to  assumes that  and  change  point  1979).  model  size,  of  to  data  model offers  Bertalanffy  the  size  with  the  A  slower  fish  size  i.e.,  the  asymptotic m a x i m u m  asymptotic  point,  The  the  that  •asymmetrical  fit  Gompertz.  theoretical  maximum  to  early  the  inflexion  curve describes  years  of increasing  increments.  Preliminary  results  indicated  that  and  to  the  Bertalanffy  failed  considered  fit  logical  model which was Lt  =  L  *  0  to  von  describe  exp{G(l  -  t  in  years,  L  and  g  a  second  growth  0  al.  G,  a n d g)  1987)  parameter  which  a  0  were  is  a  parameter.  (Conway  to  Following  growth  a  Lt  is  the  length  at  t=0,  The  parameters  the  Gompertz  this  attempt,  curve  using  the  length  of  the  a  growth  model, it  was  Gompertz  al.  G  of  is  Gompertz  fish  (mm)  1970).  algorithm  for  growth  nonlinear  at  parameter, equation  microcomputer p r o g r a m F I S H P A R M (Prager  Marquardt's et  well  of length:  hypothetical  derived b y  fitted  model.  exp[ —gt])} where  implements  estimation  data  theoretical  written in terms  age  (L ,  the  least  et  squares  21  2.3  RESULTS  2.3.1  A  Size Composition  total  of 9 4 9 specimens  specimens  were  too  impossible  for juveniles the  of C.  small less  entire  to  globiceps  be  than  sample.  sexed 18  were  analysed;  out of this  since  external  m m . There  females  in  The  females,  a n d combined sexes of C . globiceps  were  length-frequency  unimodal.  Size composition statistics are s u m m a r i z e d i n Table  Statistical  differences  to  larger  than  males  (t=3.77;  49.9 m m ) contributed  2.3.2  Otolith Description  otolith  molecular largest  is  composed  weight  protein,  of the three  rounded traversed  deep  primarily otolin  otoliths  at the anterior by a  detected df=917;  of  (Degens  were  is  459  males  smaller  face  than  fish as  Females (5.0 m m  exemplified  49.9 m m (Figure 3).  carbonate)  T h e sagitta, i n shape,  is concave, while point  males,  1.  a n d females.  is elliptical  T h e anterior  for  distribution is  T h e smaller  (calcium  a n d 460  1980 a n d 1988 are  to the population  et a l . 1969).  i n C . globiceps,  is convex.  males  p<0.001).  aragonite  e n d . T h e lateral  sulcus,  between  densities  5 6 . 1 % of C . globiceps  the fact  determination  1, 2 a n d 3; the frequency  the greatest  from  An  that  were  sex  collected between  in Figure  i n size  30  distributions  illustrated  were  3 a n d Appendices  total  tends  and a which  high  is the  a n d bluntly  the medial  face,  to be short i n  Figure  3. Length-frequency  distribution  of  C. globieeps>  23  Table 1. Summary statistics for length and body weight for Q. globiceps population at Helby Island. GROUP Female Male Combined  QUANTITY SL (mm) W T (gm) SL(mm) WT(gm) SL(mm) WT(gm)  RANGE 18.1-139.1 0.20-98.50 18.4-121.9 0.13-60.50 8.5-139.1 0.01-98.50  MEAN±S.E. 52.7±0.972 6.70±0.372 47.5±0.977 5.42±0.347 49.0±0.702 5.87±0.249  N 460 459 949  24 very  young  otoliths  was  fish  and  long  best  reflected  older  specimens often  2.3.3  Otolith A n a l y s i s  A  regular  pattern  variations many  band  (Panella  while  the  480  was pairs  were  chemical fishes  of  were  location.  than  older  fish.  A  total  of  A  total  of  than  reasonably  zones  range  among  otoliths  the  on the  and of  generally  otoliths  were  a  zone  the  examined  out  of  because  broad  diffused  accurately  425  C.  The  otoliths  groups age  were  proportion  globiceps  age  estimated  some  5+  represented,  to  of  unreadable  comprising were  established,  (Figure with  214  the  5).  largest  55 too  although 1+  number  211  second  A  3+  age  total  of  opaque  and/or  number  higher  were  the  individuals  and  differed  males of  of  opaque  The  slightly  96.9% to  slow  11.5%  readings  and  In  or  because  was  4).  and  pairs  their  otoliths  seasonal  one.  determine  females  The  from  bands.  were  of  a broad  first  which,  otoliths tip.  band.  of their  of  fast  as  the  shape  (Figure  of  hyaline  they  eliminated  seen  appears  clarity  The  resulting be  than  the  of this  periods  narrower  narrower  in  can  seasonal  occasionally  and  rings  in  point.  dorsal m a r g i n  growth  usually  anterior  otolith  represent  is  Variations  hyaline  the  summer  are  of  unreadable,  ring.  well  rings  zone  winter  one  five  these  first  Occasionally,  more  specimens.  variability  components  first  saccular  too  older  opaque  1974). T h e  wide  considered  marks  less  a  the  alternating  subsequent opaque  There  by  the  showed protrusions  of  the  temperate  growth  and  in  in  by in  aged.  fish  were  groups  were  in  the  1 +  25  Figure  A.  (a) of (b) and  I l l u s t r a t i o n o f the l a b y r i n t h , showing t h e p o s i t i o n the o t o l i t h chambers ( a f t e r B l a c k e r 1 9 7 4 ) , o t o l i t h ( s a g i t t a ) o f C. globiceps s h o w i n g o p a q u e hyaline zones.  26  20  |  MALES  M=6  Q  FEMALES  F  =  7  10 0 20  M=II  10  F=16  Jim.  0 >• 20  0  M  10  ^  M=37  1  Ik ~  k  o- 20 LU  1  M=44  20 10  n  0  F=55  1  M=78 F=74  20  M=35  10  F=22  0  10  20  30  40  STANDARD  5. age  -  +  iJ  0 30  Figure  2  Length-frequencies groups. ( F i s h  50  LENGTH  of  male  collected  60  •  1  70  80  90  100  (mm)  and  female  during  C. globiceps  1987  a n d 1988)  by .  estimated  Table  Age (yr)  2.  Sample s i z e  and mean l e n g t h  C.globiceps  o f each age g r o u p .  Age-length Sex  n  data  females  + SE  t  d.f.  P  n  +  1  +  2  +  3  +  4  5  +  +  M F  35 22  26.9+ 0.728  M  78  F  74  M F  Mean  of  combined  length  + SE  (mm)  (mm)  0  and combined s e x e s  Sexes  Sex c o m p a r i s o n  by sex  Mean l e n g t h  (+ S E ) f o r m a l e s ,  Length  range  (mm)  0.84  55  0.41  57  27.3 +  0.542  18.4  -  32.2  40.0+ 0.472 40.5+ 0.496  0.71  150  0.48  152  40.3 +  0.341  32.2 -  47.2  44 55  53.1+ 0.716 55.1+ 0.707  1.95  97  0.06  99  54.2 +  0.513  47.0 -  66.0  M  37  70.4+ 0.654  F  40  69.6+ 0.555  0.88  75  0.38  77  70.0 +  0.426  61.6  -  76.6  M F  11 16  80.7+ 0.699 80.2+ 0.677  0.53  25  0.60  27  80.4 +  0.486  77.0 -  85.9  M F  6 7  91.5+ 2.848 90.5+ 1.235  0.33  11  0.75  13  90.9 +  1.411  86.1  105*1  of  freedom;  n=sample  27.9+ 0.801  size;  t=t-statistic;  d.f.=degrees  P=probability  -  28 and  2+  age  overlapped first  groups.  in  opaque  length zone  Also  worth  (Figure  had  not  5,  noting  Table  yet  is  2,  been  the  and  fact  that  Appendix  delineated  were  age  4).  classes  Fish  considered  in to  greatly  which  be  the  age  0 +  fish.  2.3.4  Six &  Length-Frequency  modal  classes  Pitcher  1979)  corresponded  age  group  fish  to  0+ ,  groups  the  Following  the  modes  The  distance  time of  annulus  annulus  more the  of  over  subsequent  3)  and  4+ , over  smaller  year  numbers 8).  analysis shown  and peaked  5+  of  reveals  there  and  of  three  year  age  were  not  sample  recruitment  (around  7).  are  five  small  data,  (Figure  histograms  that  summer  classes  four  These  (MacDonald  age-at-length  that  because  in  method  the  indicated  (Figure  months  from  and time  mixture  sizes.  began  in  August).  Formation  annulus  formation  the  3+ ,  population  the  distribution  Table  followed  and M a y )  last  formation  recently  2+ ,  the  Annulus  from  and  the  MacDonald—Pitcher  (between A p r i l  of  by  progressively  in  by  Time  1+ ,  and  existing  2.3.5  6  histograms  separable  spring  identified  (Figure  Length-frequency major  were  Analysis  by  annulus  formation  to  the  (Thorogood this was of  edge 1987).  distance. formed. the  new  of the Capture  Hence, The  as  otolith  is  an  date  is  related  this  delineation  opaque  zone,  distance of  the  evidence  indicator to  of  the  decreases,  hyaline of  zone  the  the date the and  regular  Fig. 6  N =  462  60 50 . 40 . 30 20 . o  10 -  0  Figure  10  6.  7.  30  Length-frequency  April Figure  20  1987  and  40  otolithic  60  70  80  90  Standard  Length (mm)  distribution  of C.  October  Length-frequency  from  50  ageing.  globiceps  100  sampled  110  120 130  between  1988-  distribution  of age  0+  through  5+  fish  derived  Table 3. Parameters of MacDonald-Pitcher length-frequency analysis. p S.D. Mean length (mm) Age  0.0351 3.76 25.9  0.2984 4.68 38.8  0.3482 5.13 49.4  0.1964 3.88 67.7  0.0799 2.52 78.4  0.0420 5.09 88.9  0+  1+  2+  3+  4+  5+  p - r e l a t i v e abundance o f mode a s p r o p o r t i o n S.D.=standard d e v i a t i o n o f mode.  of total  sample  31  A p r i l 1987  October 1988 N=38  |—|_j—j 10  20  30  40  50  Standard  Figure  8. Length-frequency of  modes.  60  70  80  90  100  110  120  130 140  Length (mm)  distributions  of  C. globiceps  showing  progression  32  Jan  Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Time (Months)  Age groups  Figure  ——  Age-1 +  —H-  Age-2+  —*-  Age-3+  - s  Combined  9. Otolith  marginal  increment  plotted  by  month  for  the  three  age  groups.  33 formation mean  of  a  single  marginal  (Figure  9).  band  increment  All  age-1,  per in  age-2  year  the  was  fish  and  age-3  indicated  by  a  sampled  between  fish  completed  had  sharp April  decline  and  annulus  in  May  the 1988  formation  by  May.  Rapid  growth  increments summer  of  fish  age by  each  growth  with  early  several  individual  that  opaque  months,  entire  that  3  winter  3+ )  (Figure  9).  by  and of  B a s e d on  increased  area) in  marginal  throughout formed  Despite  annulus  deviation these  the  were  spring.  seasonal little  in  occurred zones  earl}'  (incremental  numbers  increases  rings  hyaline  showed  sequence  incremental  opaque  examination  to  stock  represented  Likewise,  late  an  (1+  as  and  months.  hyaline—opaque  and  2,  during  classes  the  zone,  1,  fall  predominantly  for  displayed  the  fish  and  annually, data  of  a  the once  lack  formation  from  of for  the  pattern  observations, I  conclude  represented  number  with  an  one  year's  increase  in  size.  2.3.6  Growth  in  Length Back-Calculated Lengths  A  strong  and Table and  positive  otolith 4).  radius  linear for  relationship the  entire  A n a l y s i s of covariance  female  fish  for  standard  was size  found range  indicated  length  and  no  between of  fish  body  available  significant  otolith  length  radius  of  the  fish  (Figure  10  and  differences  between  relationship  (Table  male 4).  20  r  2  3  Otolith Radius (mm*30)  Figure  10.  Relationship  C. globiceps.  of saccular otolith radius  to  fish  standard  length  for  35  Table 4. Summary of standard length-otolith radius regression statistics (y=a+bx) and analysis of covariance between male and female _C_. globiceps. GROUP Female Male Combined  N  a  b  191 175 366  -22.68 -27.23 -24.97  28.51 29.81 29.17  r  2  0.85 0.91 0.88  F 1096.12 1723.53 2653.89  P 0.001 0.001 0.001  Analysis of covariance (Males vs Females). Source of variation Equality of adj. means Error Equality of slopes Error  D.F.  SS  MS  F  P  1  87.44  87.44  2.89  0.09  363 1  11000.81 40.36  30.31 40.36  1.33  0.25  362  10960.45  30.28  Table  5.  Age  Back-calculated lengths  No.  at  capture  (yr)  fish  of  at  Mean at  each age  for  combined data  Back c a l c u l a t e d  length  capture  (mm)  +  57  27.3  1  +  152  40.3  32.0  2  +  99  54.2  31.4  3  +  77  70.0  4  +  27  80.4  5  +  13  90.9  lengths  (mm) a t  successive 4  3  annuli 5  47.4  32.2  50.2  63.3  32.4  51.7  66.3  76.7  33.8  50.5  65.9  78.3  86.7  32.4  50.0  65.2  77.5  86.7  425  G r a n d mean Growth  l  globiceps.  2  1  0  Total  of C.  Increment  17.6  15.2  12. 3  9.2  CO OS  Table  Age  6. Back-calculated lengths  at  capture  No. (yr)  fish  of  at  each age  Mean at  for  length  capture  male  C.  globiceps.  Back c a l c u l a t e d (mm)  2  1  lengths  (mm)  at successive 4  3  0  +  35  26.9  1  +  78  40.0  31.9  2  +  44  53.1  31.1  46.7  3  +  37  70.4  32.3  50.2  63.2  4  +  11  80.7  31.9  51.6  66.2  76.8  5  +  6  91.5  34.3  51.8  66.7  78.6  32.3  50.1  65.4  77.7  Total  87.4  211  G r a n d mean  Growth  5  increment  17.8  15.3  12.3  87.4  9.7  annul!  Table  Age  7.  Back-calculated lengths  at  capture  No. (yr)  fish  of  Mean at  at  each age for  female  Back c a l c u l a t e d l e n g t h s  length  capture  C. globiceps.  (mm)  1  2  (mm) a t  successive 4  3  annuli 5  0  +  22  27.9  1  +  74  40.5  32.0  2  +  55  55.1  31.6  48.1  3  +  40  69.6  32.1  50.1  63.4  4  +  16  80.2  51.7  66.4  76.7  5  +  7  90.5  33.3  51.6  66.8  78.4  86.0  32.4  50.4  65.5  77.6  86.0  I  Total  32.8  214  G r a n d mean Growth  1  increment  18.0  15.1  12.1  8.4  39 Therefore, SL  =  a  c o m m o n regression line  -24.96  This  +  equation  29.17  demonstrates  proportional.  However,  and  radius  otolith  females  an  175  lengths  1+ ,  3+  and  larger  than  larger  than  the  observed  lengths  reversed  Lee's  phenomenon used  for  the  for  than  those  Growth it  calculated  is the  to  be  females,  6  and is  situation lower  when  from  4+  length  by  the  end  increased in length b y mm  in  length  by  the  of  end  of  5+ . the  whereby the  the  value  fish  amongst  mm  and  standard  total of  OR)  length  10).  The  366  is  (SL) above  specimens  tend  whereas  males  The  first  age  older  3 + . Age  year,  from  1979).  third  1+  whereas  trend  fish  are is  slightly slightly  reflected  Rosa  whose 1969).  older  fish  Negative  in  1+  and  members  second y e a r .  Age  year  5).  of 3+  Mean  Lee's  otolith  is  Reversal  of  are  greater  size-selective  C. globiceps  individuals  (Table  be  Within  C. globiceps studied,  (Ricker  groups  to  briefly,  the  obtained  age  of  Stated  phenomenon in  during the their  same  sample  (Ricker  the  close agreement.  average,  back-calculated lengths  their  (SL  (Figure  in  the  lengths  In  cause of this  15.2  fish  are  demonstrated.  younger  faster  on  and  7).  decreased slightly, especially after  in  between  larger  classes  groups  most likely  appeared  age  age  calculation, the  phenomenon occurs  with  parameters  used for back-calculations.  different  5,  two  variability  back-calculated  a  Lee's  mortality  were  phenomenon to  these  for  (Tables  refers  in  basis of back-calculation. A  groups, the  males  females  the  from  age  growth  evident  males)  Back-calculated 2+ ,  was  equation  p<0.01).  2  that  formed  and  (r =0.88,  calculated and provided the  increase in the  (OR)  regression equation (191  OR  was  2+  population.  after  which  grew  17.6  mm  age  group  2 +  fish  grew  12.3  back-calculated  40 lengths  (Tables  approximately  6  the  and  7)  same  illustrated  that  both  sexes  grew  in  length  at  rate. Theoretical Growth Curves  A  plot  of  fish  relationship  length  which  part  of growth,  The  rate  curve  of  The young  were  plotted, was  sigmoid  from  in  a  reason  very  model)  age  age  better for  a  which  show  an  model is u s u a l l y fish  has  0+  to  to  and  von  applied to  concordance  (Figure  the  pattern  to  the  the  pattern  the  in  final  stanza  fishery because  and  (in of  empirical  indicating  exhibited  by  Gompertz  curve  known  life  that  this  size  age  was  linear  the  early  greatest. growth  von Bertalanffy  growth  more  complete,  When  these  by  i.e., data  Gompertz  model  does not  conform  to  the  point  early  the  of  the  to  occur  curves  Gompertz Helby  von  is taken and  yield shows  model Island.  life  short-lived  The  stocks), in  in  in  1987).  application  fitting are  a  Gompertz  described  of life which  species at  by  were  best  (Moreau  growth the  In  included.  commercial its  11).  the  were  growth  is  (Figure  data  feature  showed  3 + . The  than  inflexion  limitation  in  age  age  (a  an  early  from  groups  otoliths,  ages  data  the  age  from  increase  length  from  the  work  theoretical 11)  older  Bertalanffy  point  of  at  that  older  inflexion  comparison  the  being  resulting  fishery  determined  2 +,  sigmoidal  recruited  from  off  This  in  estimated  level  1975).  extensively  growth  to  fit  observed. T h e  (Yamaguchi  as  this  slight  length-growth  age,  decreased gradually  groups  fishes  the  tended  acceleration  resulted  curve.  against  to  species,  Bertalanffy start  has  been  when used  computations.  A  almost  complete  adequately  describes  Growth  presented in T a b l e  8.  parameters  41 The  null  males  and  each the  hypothesis females  parameter limits  growth  the  was  (Table  between  between  therefore  that  examined 8).  It  sexes,  males  computed  and  three  Gompertz using  can  be  suggesting  growth  the  95%  confidence  seen  that  there  that  there  is  and  females.  The  the  following  Gompertz  parameters  growth  is no  a  are  attached  to  overlap  of  difference  in  complete  for  equation  in  limits  significant  curve  equal  combined  described  data  the  was  growth  of  C. globiceps: Lt  =  26.7mm  *  Goodness  of  that  model  the  model  is  growth  mm  a  mm. and  From  were two  in  11  compared lengths  support to  age  it  lengths  Predicted  are  5+  can  analysis  not  lengths by  of  variance  a  highly  significant  11.  Mean  observed,  Tables  10  1+  and  females  averaged  be  predicted  seen from  apparent, from  using t-test  compared  the  an  back-calculated length  of age  sexes  in  exp[-0.30t])}.  by  Figure  Estimated those  -  accounted for  mean  theoretical  sexes.  tested  compared  Table  between  was  shown  were  obtained 87.4  fit  exp{1.58(l  the for  reasonably  accuracy of the  of  11.  had  86.5  a  which  variation.  back-calculated, estimated  mm  mean  and  age  5+  The  and  age  indicated fitted  theoretical  1+  male  males  fish  averaged  back-calculated length  observed  Gompertz therefore, Gompertz paired  model.  of  9)  of  32.3  back-calculated,  and  mm.  that  well  amount  By  32.0  (Table  model  lengths, agreed  closely.  comparisons were growth  model  comparisons (Sokal  (t=0.23,  df=5,  made  and &  Differences on  combined  observed Rohlf  p = 0.824)  lengths  1981).  adding  The  further  42  100  0  1  2  3  4  Age (years)  Curves Gompertz  Figure  11. G o m p e r t z  growth  function  for C .  — — Empirical 1  globiceps.  5  6  43 Table 8. Parameter estimates derived from the Gompertz growth model, for C. globiceps.  Group  Parameter  Male  L (mm)  Estimate  Asymp. S.E.  1 .96(S.E.)  26.5  0.6100  1 .1956  0  Female  Upper 1 imit  25.3  27.7  G  1.64  0.0648  0.1270  1 .51  1 .77  9  0.29  0.0258  0.0506  0.24  0.34  0.6917  1 .3557  L (mm) 0  Combined  Lower 1 imit  27.1  25.7  28.4  G  1 .53  0.0498  0.0976  1 .43  1 .63  9  0.31  0.0268  0.0524  0.26  0.37  0.4569  0.8955  L (mm) Q  26.7  25.8  27.6  G  1 .58  0.0395  0.0773  1 .50  1 .66  9  0.30  0.0185  0.0363  0.27  0.34  Table 9. Analysis of variance for Gompertz model.  Source of variation FEMALE Model Error MALE Model Error C O M B I N E D SEXES Model Error  D.F.  SS  MS  F  P  3 211  659363.13 3823.40  219787.71 18.12  12129.56  0.001  3 208  579219.21 4031.92  193073.07 19.38  9962.49  0.001  3 422  1238518.59 7919.07  412839.53 18.77  21994.65  0.001  44  Table 10. Summary of mean observed, back-calculated and theoretical growth in length from the present study: Males and Females. Observed length (mm) Estimated age  Male  Female  0+  26.9 40.0 53.1 70.4 80.7 91.5  27.9 40.5 55.1 69.6 80.2 90.5  1+ 2+ 3+ 4+ 5+  Back-calculated length (mm)  Predicted length from Gompertz model  Male -  Female -  32.0 49.8 65.2 77.7 87.4  32.3 50.1 65.0 77.4 86.5  Male 26.5 40.0 " 54.5 68.6 81.6 92.8  Female 27.1 40.9 55.2 68.8 80.8 90.9  Table 11. Summary of mean observed, back-calculated and theoretical growth in length from the present study: Combined sexes. Estimated age 0+ 1+ 2+ 3+ 4+ 5+  Observed length (mm) 27.3 40.3 54.2 70.0 80.4 90.9  Back-calculated length (mm) -  32.2 50.0 65.1 77.6 87.0  Predicted length from Gompertz model 26.7 40.4 54.9 68.7 81.2 91.7  45 Length—Weight Relationship  The  length—weight  sample are  of 460  plotted  length  growth  significance  correlations  are  data  (Table  test  whether  between  12). the  sexes  Software  males,  and  and  the  rates.  and  Figure  The  a  12.  slope  calculated  using  pooled sample of 944 Functional  estimates  statistics  Since  a  3,  found  between  The  by  was  that  was (i.e.,  length  for  these  used  a  random  The  results  weight  versus  fish.  regressions of  were  method  from  slope  by  allometric  of  are t-test  weight  for  males  in  the  regressions  analysis  (Table  difference intercept  not which  growth)  by  calculation  of  presented  in  are  of  t-tests. males,  and  intercepts of  intercepts  each other  and  and  and  and  significant  determined  lines  values)  slopes  slopes  tested  (b  the  The  no  was  regression  slopes  common  hypothesis growth)  correlations  different  there  sexes,  of  1983).  significantly  14).  12  C. globiceps was  of  13.  The  from  Table  459  calculated  instantaneous Table  females,  in  were  relationship  of  (a  females,  females values)  were  significant and  pooled  compared  differed  covariance  to  significantly  (BMDP  Statistical  males  and  females  are  not  slopes  and  intercepts  of  both  The  null  14).  between were  calculated  significantly  different  revealed  the  indicating  Highly  a  that  deviation  (Table from  b  value  from  3  12). (i.e.,  differs  the  cube  isometric  significantly law  (Table  46  COMBINED DflTR  1  <D  1  O  -2  1  2  Log Length (cm)  Figure  12.  Length-weight  relationship of C . globiceps.  47  Table 12. Length-weight relationship in C. globiceps males, females and combined data. Regression equation Group Mali  !  W=9.8698 * l f T L Log W=-5.0057+3.2674 Log L W=1.1363 * 10~ I Log W=-4.9445+3.2323 Log L W=1.5913 * 10~ L Log W=-4.7982-f3.1552 Log L 6  Female Combined  3 2 6 7 4  N 459  r 0.981  p 0.001  2  5  3 2 3 2 3  460  0.978  0.001  5  3 1 5 5 2  944  0.975  0.001  Table 13. Length-weight relationship described as Log W=log a'-f-(b/r)log L (functional regression). a' Group Male Female Combined  1.4058 * 10~ 1.3630 * i o 1.9901 * 10"  5 5  5  b/r  N  r  3.3004 3.2683 3.1968  459 460 944  0.981 0.978 0.975  48  Table 14. Analysis of covariance between sexes for length-weight regression and t-test for comparison of calculated regression coefficient with regression coefficient of the cube law. Source of variation Equality of adj. means Error EquaUty of slopes Error  D.F.  SS  MS  F  1  36.94  36.94  2.77  0.096  916 1  12231.80 34.10  13.35 34.10  2.56  0.110  915  12197.70  13.33  P  t-test Group Male Female Combined  b-3 0.2674 0.2323 0.1552  S.E. of b D.F. 0.0236 457 458 0.0199 0.0110 942  t 11.34 11.62 14.11  P 0.001 0.001 0.001  49 Growth Rates  Instantaneous otolithic  growth  ageing.  rates  Male  and female  sizes for the calculations  The  highest  monthly  August  (Table  August  is  there  slightly  was  confounded decreased age  0+  15  results  were  than  the  specimen for  August  in  the  significant group.  difference  significant; 14).  the  growth  rates.  rates  were  value  for  preceding  age  age  obtained age  4+ .  in  June  group  in  question.  Generally,  age  so as to  5+  and October.  initially  It  for  c a n be (p>0.05)  Because there  between  age—length  16).  pooled  The  age  individual  5+  It  and  increase  from sample  during  June  and  during  June  and  noting  that  is  worth  this  more  Growth  growth  groups  than  rates  rates  were  likely  markedly higher  for  of each  age  with age.  Relationship  analysed  (Table  Figure  13).  for  were  growth  and decreased progressively  Data  data  Figure  higher  the  Age-Length  age  samples  of instantaneous  and  one  2.3.7  group  calculated  instantaneous  only  between  were  sexes  regression regressions  seen  males  in  length  analysis  the m e a n  for  were  length  combined  combined  and females  between  sexes  of covariance  to be no apparent  data  calculated for  from  between  appears these  difference  sexes  are  of each  difference for  also  that  there  is no  sex within  each  in the age —length  regression (Figure displayed  analysis.  14)  was  (Table  The  highly 16  and  50  Table 15. Instantaneous growth rates of C . globiceps in their age 0+ through age 5-)-. Rates were calculated for the intervals between the 1987 and 1988 collections. Instantaneous rate Age Collection dates 0+ 2+ 1+ May 1987 0.793 0.115 June 1988 0.712 0.324 0.229 August 1988 0.045 0.039 October 1988  3+  4+  5+  0.144  0.057  0.019  0.166  0.112  0.131  0  -  0.032  51  80  i  60  ho D) C  (no  data)  CO TJ C CO  co 20  0  Jan  Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Time (Months)  Figure  13.  Instantaneous  growth curves  for  age 0 +  through 3 +  C.  globiceps.  52 2.3.8  Total  It  possible  is  vice  Length-Standard  versa  to  convert  (Figure  15).  Length  Relationship  secured  with  data The  relationship  total  length  between  the  to  standard  two  was  length  defined  and  by  the  and  460  equations: Total SL  length  =  =  2.3.9  The  standard  -1.5183  Standard TL  to  +  length  2.1267  Sex  sex  0.8327  to total +  ratio  the  entire  for  Chi-square  on 17). for  a  fairly  =0.001,  sample From  2  =0.995,  n = 941).  (r  =0.995,  2  n = 941).  Ratio  females. 2  (r  length SL  was  (X  TL  1.1950  females  1:1  length  the  balanced, test  425 table  combined age  having  revealed  p = 0.97).  of  sample  Analysis  individuals it  that  can  an  not  that  comprising  overall  ratio  of  459  ratio  did  not  differ  sex  ratio  by  age  groups  sex  214  ratios  significantly  females for  age  different  males  0.998:1  sex  consisting of  seen  groups were  of  (n = 919)  and  favour  significantly was 211  groups  from  in  unity  0+  from  carried males to  (1:1).  5+  of  out  (Table and  53  Table 16. Summary of age-length regression statistics (y=a+bx) and analysis of covariance between male and female C. globiceps. Group  N  a -1.83  Female 214 Male . 211 Combined 425  -1.71 -1.76  b 0.07 0.07 0.07  r  2  F  0.935 0.937 0.936  3022.23 3103.84 6176.79  P 0.001 0.001 0.001  Analysis of covariance (Males vs Females). Source of variation Equabty of adj. means Error Equality of slopes Error  D.F.  SS  MS  F  P  1  0.055  0.055  0.55  0.457  422 1  42.27 0.129  0.10 0.129  1.29  0.257  421  42.14  0.10  Table 17. Sex ratios of C. globiceps by age groups. Age group 0+ 1+ 2+ 3+ • 4+ 5+ Combined age groups  M 35 78 44 37 11 6 211  F 22 74 55 40 16 7 214  Sex ratio 1.60:1 1.05:1 0.80:1 0.93:1 0.69:1 0.86:1 0.99:1  X  2  2.965 0.105 1.222 0.117 0.926 0.077 0.021  P 0.085 0.746 0.269 0.732 0.336 0.782 0.884  54  4  Combined N=425  3  0 5  co  3  D)  2  CD  <  0 5 4 3  2 1 0 20  40  60  80  100  Standard Length (mm)  Figure  14.  Age-length  relationship  of  C. globiceps  determined  by  otoliths-  Total Length vs. Standard Length  Figure  15.  Relationship  of total length  to  standard  length  and  vice  versa.  56  2.4  DISCUSSION  Clear  annual  which  made  preferred 1974;  The  distinguishable  to age these  fish.  T h e sagittae  for age determination  1980) a n d have Horton  been  1977).  (Bagenal  suggested  In the present  8 8 . 5 % of the specimens  of C.  i n sagittae  with  globiceps  of teleost  (Figure fish  1974; W i l l i a m s  to be the most study  are the  &  zones  Bedford  accurate  it w a s possible to  the annual  clearly  4)  ageing estimate  evident  in  otoliths.  of C . globiceps,  otoliths  such  as hearing  record and  possible  (Six &  age of  were  structure  Panella  whole  a  large  details  provide  associated spawning, exact  amount  of its life  ring  with  low  a n d altered  the onset  beginning  those  in  &  teleosts,  Coombs  about  Likewise,  fish  serve  1980).  a n individual's  otoliths  obscure zone  seasonal  study  did not look  hyaline  zone formation,  during  temperatures,  photoperiod  of hyaline  current  occurs  water  remains  of a  (Popper  of information history.  i n other  from  a  many  functions  Furthermore, condition  collection  otoliths  over  time  of individuals  population d y n a m i c s .  formation  mechanism  Island  like  a n d equilibrium  insights into  Hyaline  the  it  hard  structure the  patterns  into  therefore  (e.g. W i l l i a m s (e.g. P a n e l l a  &  of slow food  Bedford  1974).  growth,  generally  availability,  seasonal  1974),  (Figure  temperature  to  9)  about  a n d gonadal activity  no support is offered  although the  F o r C . globiceps  i n October  i n water  food intake  period  reduced  formation  drop  a  at  Helby  coincides  with  10.1°C.  The  as they  relate to  for these hypotheses.  57 Chadwick the of  (1976a)  time  of  annulus  Vancouver  occurred  months  northern  2+ ,  classes age  present fish  with  correspond to  A  major  from  the  by  of  lengths  18).  small is  age  0+  in  the  between  small  by  that  present  this  sample  age  June  to  of  increment  other  temperate  Betsill  1987).  of  revealed  not  5 + ).  4+  classes  study,  groups  five  many  Renfrew,  age  this  as  age  as  six  Vancouver observed  but  with  the  using  age  Whereas  observed Port  in  Island,  three major  and  at  fishes  Helby  comparable  were  the  those  in  the  on  a  single  collection  fish  at  California  and  and  1+  in  his  study  by  lengths  based  41  0+  from  otolith  seasonality  (1976a)  and  of  The  displayed in  (10  formation  rates  rapid  coast  remaining  and  were  west  the  (4+  0+  the  been  over  rings,  analyses  sizes  of  population  Chadwick  individuals  possible  &  groups  those  Since his  growth  analysis,  attained  sample  highly  discrepancy extremely  3+  rate  have  annulus  Observed was  most  may  on  that  growth  May.  vertebral  May  location  9).  globiceps  age  study,  counts  and  (Table  to  Maceina  smaller  this  the  with  C.  closely paralleled  2+  it  two  in  on  in  a  to  speculation  that  April  1979;  at  the  in  consistent  April  (Figure  length-frequency  and  Standard  very  respectively),  is  observed  (1976a)  study  confirms  being  from  globiceps  indicated  determination  3+ )  1+ ,  C.  (April/May)  (Grossman  based  B.C.  age  age  and  were  Chadwick for  here  verified  classes  Island,  lowest  hemisphere  and  period  significant . decrease  the  of  study  spring  the  for  deposition  a  shown  study  (1 + ,  the  with  with  deposition  otoliths  This  increment  September,  that  formation  Island.  during  marginal  The  speculated  of  B.C.  actually  study.  study  sizes he  and used.  Chadwick's For  ageing  example,  as  work  emanates  shown in  Table  58  Table 18. Age-length data from Chadwick's (1976a) study and the present study, showing the discrepancy in the former study. Chadwick's study Age groups N 0+ 1+ 2+ 3+ 4+ 5+ 6+  8 21 3 6 1 1 1  Present study Standard length (mm)  N  Standard length (mm)  26.1 32.2 43.7 54.2 79.0 113.0 106.0  57 152 99 77 27 13 0  27.3 40.3 54.2 70.0 80.4 90.9 0  59 18,  age groups  for  each  lives  up  to  for  remains  The  older  agreement and  by  adjacent  the  6  Both  of  Other  studies  of  years  (e.g.  Scorpaenichthys another  up to  four  fish.  classes both  Rare  at  deviate  (McNew  highly  T h e otolith  &  there  by  of the  modal  age-class failure  length  represented  assumed  but there  in  marine  cottids  snyderi,  Freeman  marmoratus,  subtidal  cottid,  a  intertidal  Wells  when  not be  Clearly,  from  Summerfelt  (e.g. Oligocottus  years.  could  of  overlap  age groups  overlapping  the  collections,  distributions  normality)  are  1978; M a c D o n a l d  appear  to provide  is no reason  & the  to suspect  incorrect.  determination  abundant  the  m a y also be missed b y  interpretations  identification,  in  identified  times  of poorly  from  globiceps  methods  age classes  of  C.  readable.  discrepancies resulted  in definition  analysis  of age class  age  on these  modes  specimen  determined,  not  misclassification  1987).  were  age  single  do suggest that  were  of  on a  specimens  supports  age  as being substantially  1 to  survive  otoliths  Some  mixed  i n modal  means  from  maculosus,  2  ages  location  distributions  1979; M a c D o n a l d  method  whose  interpretation  a n d 7).  study  larger  ageing work  the  are based  some  their  the difficulty  the  recognised problems  either  in this  age groups w a s great.  if  reliable  those  since  identify  possibility  (particularly  more  than  otolith  to  analysis.  Pitcher  years,  between  (Figures  i n his study  the results  5+  analysis  analysis  modal  least  much  identification  between  While  a need for additional  analysis  and  at  otolith  good  modal  5 + , and 6 +  age group.  presumably used  4+ ,  cottid,  (1986)  suggest et  al.  determined  1985)  O'Connell  w a s estimated the  lifespan  by  to  1953). Craik  ranging 13  years  Oligocottus (1978)  age of Clinocottus  to  analis  60 using was  otoliths 6  at  years  Point  for  lifespan between were  no  females  deviations  in  of C . globiceps  was  when  peaks  1984)  occur. R a p i d  growth  Oligocottus  maculosus  cottids, al.  in  approached  and  1979,  1981)  that  during  winter  might  during  the  winter  d u r i n g the  months  strongly  in  (Moring  evident  in  all  during  activity this  globiceps  was  October,  thought  to  study  (Figure  productivity reported  was  has  caused  found  in  responsible for  et  as  al.  (Moring  wave  Gonadal  et  winter  suggested  action  development  reproductive reduced  time  intertidal  non-reproductive  winter  no  Freeman  increased  reduction.  there  the  other  reduced  been  by  13),  1981;  in  was  (Parsons  for  (Moring  since  There  summer  It  therefore,  be  this  lifespan  Differences  longer.  Growth  growth  males.  sex lived  snyderi  winter.  maximum  collections.  also been  1986).  explain  (C.  also  for  in  O.  that  years  primary  1979),  foraging  A p r i l through is  unity  has  (Wells  ceased  months  not  and  summer  partially  8  spring and  nutrients  reduced  suspected)  from  concluded  suggest that either  analis  probably  were  confined to  temperature,  Clinocottus  1985),  ratios  would  and  approximately  globiceps  sex  which  California  and C.  sexes in  discernible pattern  Growth  Fermin,  condition activity  growth  is  (Moring  1981).  The the the  radius time  of  of  otolith  their  agreement  seasons  age of  growth group  life  provided  formation. between  back-calculated previous  annuli  were  of  The  accuracy  observed a  fish  (Ricker greater  of  lengths  species  1969). in  reasonably  The older  accurate  back-calculations and  quantifies  the  for  C.  growth lengths  globiceps  of  was  back-calculated  back-calculated fish  estimates  length  at  confirmed  by  lengths.  attained for (Table  in  the 5).  The each earlier This  61 relationship and  is  appears  manifested  described  for  put  of the  rapidly  or the  the  use  stock  frequency towards uptake &  as  large  1974) hard  to  there  indirect  fairly give  The  justified  data  include:  (Lee  1920),  3)  of  and  as  result  a  part  false 5)  This  errors  in  1956;  way  directly  Other  mortality  back-calculation sampling  of  increase  in  which of  the  annuli  or  active  dissolution  O u c h i et  been  (1969).  non-random  passive  1979)  has  size-selective  contraction  (Yamada  1969,  Weiss  annuli  the of  by  1)  2)  older,  accurate.  due  to  indication  Lee's  fish  be  made.  al.  or  this  cannot be it  collecting  technique  other  as  1972;  satisfied, but  phenomenon  in  this  in  during  was  that  Mugiya  also  of  is  most  in  this  lack  of  also  period.  different  year  of the  likely  study skewed  appeared In  classes  data The  to  addition is  Unfortunately  observed phenomenon.  species  observed  non-representative  study  reasonable.  ideas.  back-calculated  study  upon examination  the  the  noticeable  this  between  above  employed  a  the  the  concordance of  suggestive  annuli  formation  explain  was  evidence  assumption  does not  The  suggest  bands  annual  testing  strongly  of back-calculated lengths  that  that  can  of  samples as there  opaque  their  close agreement  strong feeling  The  distributions of  no  lengths  representative  usage  fifth explanation  is  evaluations  back-calculated  frequency  reversed  grow  (Bilton  armatus,  1958),  presence  (Ricker  phenomenon.  (Jones  method  the  mortality  Lee's  phenomenon  individuals  4)  the  Rosa  Leptocottus  Lee's  individuals  nucleus  size-selective  of  cottid  for  1969),  current  are  further  reversal  inappropriate  the  and  sampling.  negative  growing  an  several  appeared  a  1977).  the  lengths  the  a  of calcium f r o m  lengths  be  of  the  However,  length  as  forward  (Ricker  Watebe  Given  be  another  explanations more  to  a  the  there is presence  attributable  to  a of  the  62 greater  survival  of  of the  slower  Mean  observed,  faster  growing  and relatively  Growth  in  length  appeared model  growth  models except  to  imply  Gompertz von  more  that  growth that  point,  g  point  One  with  of  For  many  rate  for  than  that  from  indicating  is  showing  fit  the  rate  and  two  (Figure  first  11).  used and this  most  appropriate.  the  early  and  'goodness of fit'  that  especially is,  up  This  to  the  reflected growth  rate  life  Other  intended of  the  data.  years  The when  corresponding  with by  of  use  empirical  age  11).  Gompertz  is not  early  be  &  years.  The  the  increases  is  representing  in  the  later  to  10  years  Thus  not  progressively  0.30  the  were  data,  decreases.  (Tables  in  was  inflexion,  aspects  others cube  combined  a  of  species it  the  showed 3  not  examined  to  age,  the"  whereas  parameters  before  to  and  after  G the  respectively.  important  but  mortality  appeared  growth  assessed on its  did  1.58  the  length,  lesser  best  progressively  values  size-selective  lengths-at-age  during  thereafter  model  growth  of inflexion  weight. of  it  (exponential)  von Bertalanffy  point  the  subsequently and  a  negative  C. globiceps  the  reflected  Gompertz  model  displayed  i.e.,  theoretical  asymptotic  the  was  Bertalanffy  rapid  accurately  the  model  very  become  growth  and  consistent for  was  to  individuals,  individuals.  back-calculated,  realistic  but  growing  a  it  has has  (Le  growth been  been  Cren  regression  deviation  positive  of  found  the  relationship  that  shown that  1951).  The  coefficient  from  allometric  is  the  growth  cube  weight weight  between  increases accrues at  length —weight  for law  both  sexes  (Table  (b = 3.1552)  that  length as a  the  and cube  greater  or  regression analysis differed  14). is,  significantly  Therefore the  fish  the  fish  becomes  63 heavier  for its length  length — weight different  not  Sex  sexes.  ratios  differences  poorly to  reasonable  result  &  (Emlen There  of either  not seem  easily  obtained  in  the  1961; Potter 1973).  is  males  no  more  zone  selectivity,  In  sex,  the case of C . were  found  considering the  be important  b u t were  evidence  or females  to  with  w a s the case statistically  for C.  from  a  one sex. M o v e m e n t s a n d small  sample  mate  1:1 of  sizes  ratio, sexually  over  a  increasing  globiceps.  a  age (Table  even  Although  the overall  segregated  account for a predominance of one sex i n some samples.  pairs  0.998:1  towards  a  17). It  is  a n d have  sex ratios  individual  restricted  (Warner  was  trend  or f o r m  have  few  species  globiceps  suggest  randomly  b u t they  a n d as a consequence  of C.  T h e sex ratio  statistics  fish  et a l . 1974),  the sexes, would  intertidal  Differences i n  relationship  atypical  between  by  1978).  of all fishery  or no size difference  dominated  gear  seasons might  which  not deviate  1978).  to be important  species,  a n d such  of  Tesch  to assume that  1973),  Tesch  length—weight  different  except  (Hardisty  understood females).  a  &  study.  undiscussed,  and lampreys  as  in  does  are one of the most  preponderance  did  occur  (Bagenal  of d a y (Bagenal  selectivity  in this  remain  (males  little  no  larger  of fish collection, however,  usually  are  Gear  investigated  1975),  may  a n d times  population  between method  relationship  seasons,  globiceps  as it grows  sex ratio  collections  groups  period  (Emlen  within  of time  were the could  III.  3.1  term  1946;  'fish  Allen  the  of  died  during  best  epitome  species. such  Ivlev  fish  a  Production  the  population  species in  the  (Ricker  1946)  method  (Hynes  &  Garman  &  tracking  energy  concept  investigators drew  different  introduced  have  be  many  a  to  &  of  Gill  including  growth  indices  for  1987)  (Ricker  is  defined,  produced  growth  by  production  environmental all  authors  tissue  considered  and for  many  body  time,  comparative  estimated  size-frequency Hamilton  Other  the  Weatherley  (1969)  account  1968;  by  1969;  approaches  by  in  a  fish  that  be  "the  to  of  population  assessing  the  a  fish  performance"  major  concept its  energetic  ability  route  in  fish  the  possibility  64  of  formerly  a as  ecological  or  an  organism's  to  fix  and  studying  Hynes  1980;  studies.  involve  organisms.  success  retain  production  the  Hynes  of  that  rate  the  production  communities  production  growth  called  1979;  studying  approaches to  this  instantaneous  Benke  to  populations  of  the  method,  in  function  of the  taken  attention  Cren  was  the  1983).  by  environments.  study  and  and  of  dynamics  transformations  might  underlies  Le  thesis  weight  interval  valid  Coleman  Waters  (1942)  environment  as  present  method  total  estimates  serve  this  1978b;  as  given  interval.  production  in  Lindeman  a  in  1978a,  time  could  of  used  (1945)  during  of  Since  as  Chapman  of  the  they  success  production'  1951;  sense  population  first  PRODUCTION  INTRODUCTION  The  in  FISH  in  energy.  This  studies, but Winberg  bioenergetics  an  few  (1956) of  fish  65 populations.  Mann  quantitative  relationship  productivity (1984)  (1965)  of pupfish  estimated  between  production  present  Because  of  maculosus, for  lake  study of  study the  it  attempts  strong  an  enclosed  watershed  are focused on (i) estimating  C.  globiceps  and  O.  maculosus  biomass  and  influence  of p h y s i c a l characteristics  area)  production  by  their  an  food;  energetic  intertidal  estimate  patterns  w a s considered that  and  and  approach  to  the  Naiman  viewpoint. Clinus  clinid  study  of the  (1976)  studied  Recently  Bennett  superciliosus  by  budget.  to  homing  bioenergetic  from  of  energy  a  fish  population  developing the population  The  described  the  of  the best  production  of  Clinocottus  globiceps  estimates  production  should follow  estimates.  Basic  and comparing annual by  age  populations (depth,  of' tidepools on production of these  group as  two  in a  shoreline  two species.  cottids. Oligocottus  and  the procedures  emphases  in  this  biomass a n d production  the  population,  whole,  index,  tidepool  (ii)  also  evaluating  perimeter,  total the  a n d surface  66  3.2 MATERIALS  3.2.1  The  Collection  technique  Production 0.  and  AND  METHODS  of Fishes  used  i n collecting  estimates  maculosus  were  which  fish  carried often  has been  out on two tidepool  occur i n s y m p a t r y .  on the basis of n u m b e r s , biomass, a n d the ease  A  total  of  globiceps  17  collections  made  (n = 218);  out of  (n=169).  Between  August  comprised  C.  in  16 collections  3.2.2  growth  July  O.  cottid  T h e latter with which  maculosus  a n d October  and out of this  were  sections. globiceps  species, C .  species w a s selected it could be aged.  1986 a n d J u l y found  1987 yielded in  1988,  a  total  of  total,  O.  maculosus  C.  6  collections  29  collections  were  caught  should be sampled regularly  i n fish  studies  (Chapman  reliable,  (Chadwick  a n d Determination  advocated  abundance  be less  accurate  (n=352)  a n d abundance,  estimating  total  1987  Estimates  it is being  production  this  between  i n the earlier  (n=1382).  Abundance  Though  to  globiceps  described  that  populations  1978a)  often  this  so as to account  for seasonal  is  The  not  e.g. m a r k — r e c a p t u r e moreover  they  of A g e  practical.  and removal  require  1976b; P o t et a l . 1984).  intensive  variability  in  methods  of  classical  methods sampling  have to  be  been  found  reasonably  67 Since  fish  species was  were  were  based  has  been  structure  the  same  relationship  using a  as  for  on  87  age and  ordinary individuals  of  structure for  length earlier  by at  age data  described  populations  censuses  the  abundance of the  the  only  determined  an  of  the  weight  those  was  by  sample  for  tidepool,  using  caught  in  C. globiceps  of  O. maculosus  C.  for  will  be  in  O.  globiceps.  Age  age—length  Island.  predictive  chosen species  measurements  the  Helby  resident  She  relationship  described  regression  of  the  the form  1976.  Production  mortality)  on density, of  information the  a  is  growth  rate  biomass  are  equal  plotted  method  during  mean  to  a  biomass  methods  rates  be  against  area  (Ricker  given by  will give  time the exact  used  1951)  are joined the  and  rate  of change  in  production  fundamental  can  (Allen  These points  interval  is  methods  A l l e n curve  intervals.  growth  species  two  individuals  Both  (1978)  thus  therefore  were  Information  the  previously,  Craik  each  Information  procedures  age —length  of  censuses.  O. maculosus  from  accurate,  The  by  y = a+bx,  be  here.  of  developed  first,  these  presented  maculosus  removed  considered to on  presented  3.2.3  totally  to  is a  under 1946), period.  a  individual curve  that  and  curve.  involves  for  is  growth identical  (e.g.  calculations.  production  weight  a  With species.  whereby  this The  numbers time  production during  a  given  time  The  instantaneous  rate data.  two  to  more  then  at  of  due  or  integration  Production  instantaneous results  numbers  graphical representation  mean by  estimate  in  second, to  the  determine  calculated during  by  that  the  mean  multiplying time  period.  68 A  more  by  recent  Hynes  and  (1961)  Benke  has  been  &  of  and  (1979)  found  Freeman ageing  method,  the  developed  for  to  be  size-frequency  estimating useful  Freeman  in  1985).  individuals),  by  Hynes  &  secondary  fish  This  therefore  method  it  is  was  Coleman  studies  does  useful  originally  (1968),  production  production  method  which  in  suggested  Hamilton  aquatic  (Garman  invertebrates,  &  not  require  cohort  where  ageing  of  (1969),  Waters  1983;  separation  (i.e.,  sampled  fish  poses  problems.  In  this  study  I  chose  attempted  to  results  as  a  Instantaneous  The  (1948)  fish  The  method  also  for  of  collection, question  that are  this  method  over  the  studies.  interval when  the  time  year  assumption  that  G/Z  first  time  interval  is  short,  usually  a  has  of  for  down  become  method  method.  and  I  also  compared  the  method.  the  to is  year,  (1946)  one  of the  most  ageing  to  sampled  production  is  calculated.  are  on  based  abundance  a  be  month  single  individuals.  rate  (G/Z)  is  Furthermore complete  the  is  often  not  considered  valid  when  (Chapman  1978a).  that  &  models  of  assumption may  a  Ricker  common  mortality  an  rate  of  and  to  numerical  constant  fortnight  Ricker  growth  which  and  by  requires  estimates  structure  from  laid  method  ratio  constant  The  were  years  the  size-frequency  rate  Method  production  age  the  growth  accuracy of each  This  of  instantaneous  by  Rate  assumes that  assumes,  the  production  Growth  and  the  use  of checking the  production  constant it  way  fundamentals  Foerster in  compute  to  fish  species  in  valid. the  69 Keeping  track  of  the  continuous  monitoring  monitoring  process  studies  (Thomson  Wilimovsky  the  long  to  a  run.  estimate  Basic are  months  of  (1982)  production in this  statistics  pertinent  listed  below:  •  Ni  —number  •  "Wi  -mean  to  weight  •  Zi,i—1  --instantaneous  -(InNi  -  Gi,i—1  -instantaneous  InWi •  Hi,i—1 equals  •  -  biomass in  redundant,  that  removal  and  al. of  fish.  were  the  several  1986;  N.J.  tidepool  fish  species richness  tidepools  of  by  but  However,  sampling  densities  age-class  are  I  the  completely  therefore  only  in  decided  practical  way  environment.  this  method  in g of an  B i —initial  on  one  Y o s h i y a m a et  collections probably  of fish in each group  •  •  complete  of  mortality.  regular  demonstrated  following  single complete  effects  and  1982;  that  (N)  assumption  growth  shown  significant  density  second  Grossman  have  have  and  the  affect  1976;  Grossman  three  series  Lehner  not  (W)  render  might  comm.)  does  repopulated that  &  weight  would  itself  pers.  populations  mean  as  outlined  in  Mahon  &  Balon  (1977)  i,  individual in  age group  i,  g of age group i w h i c h equals  Ni*Wi,  mortality  and  i— 1,  this  equals  and  i—1,  this  equals  i—1  and  i,  this  multiplying  the  rate  between  ages  i  InNi-l), growth  rate  between  ages  i  InWi-l, -instantaneous G —Z for  _  Bi,i — 1 - m e a n  this  rate  of increase  in  biomass between  interval,  biomass between  i — 1 and i,  this  pj  equals Bi(e  — 1).  H  Total  production  as  calculated  by  this  method  was  obtained  by  70 mean  biomass  summing  up  collection  (and  accurate  (B) for  were  the  each  not  method  species  by  age  class.  of  spawning)  (Halyk  &  Balon  initial  The  growth  Production  time  unknown.  production to  instantaneous  ratios  and  of  (G)  was  since this  1983)  biomass (P:Bi)  rate  on  a  yearly  calculated  has  been  since the  production  shown  times  to  from  of  mean  were calculated for  basis,  the  to  be  and  time the  most  spawning for  biomass  of  this  (P:B)  and  each species. Size-Frequency Method  This  method  requires  production is simply size  group  solution  to  to  the  the  required  by  accuracy  was  that  calculated b y next.  tedious  both  Since  and  Allen  determined  by  growth  This  assumes that  class  (Hamilton  1969).  groups,  chosen to  length as  a  by  the  were  rate  way  of  chosen  frequency  expensive the  has task  some  length  of individual  the of  potential ageing  instantaneous  losses of  production estimates  and  from  being  many  growth  groups,  one  a  real  individuals  rate  method,  with those  of  as its the  method.  animals  reflect due  technique.  for  into  summation  method  and  Garman  according  distributions  this  grouped  c o m p a r i n g the  reducing errors  size-frequency  be  annual  often  curve  instantaneous  method  individuals  to  the  spend an  equal  Waters  (1983)  & ageto  or  assumption  the  number  predetermined  C. globiceps  of  time  suggested  "use  size-dependent  this  Thus,  amount  and  changes  in and  the  each  of  lengths-at-age,  of  length based  size  nonequal  growth  production  size  O. maculosus.  in  in  rate"  estimates classes on  size  Production  w a s calculated  the  notation  was  required:  1.  of Newman  T h e annual was  mean  using &  number  (1983).  o f individuals  Waters  (1983)  formulation  T h e following  pertinent  the kth length  within  following information  group,  which  2,..., c) denotes  length  fvny  i(i=l,  where groups,  2,..., a) denotes  Nik represents  and Di is a weighting  2.  Martin  &  estimated b y  =-  and  Garman  the first  T h e annual  the density factor  s a m p l i n g date mean  s a m p l i n g dates,  weight  value  k(k=l,  the kth group  within  representing  the interval  between  on date  i  the ith date  (in days).  o f each length  group i n the population,  which w a s  expressed a s  where group  Using  Wik is the average  number  formulation,  the population  o f average  of size classes:  P-t  o f those  individuals  within  the kth size  o n date i.  the above  lost f r o m  weight  production  through  cohorts which  Hamilton  is calculated  mortality develop  between during  a s the s u m o f the biomass  size classes,  the period  multiplied  (same  b y the  a s the n u m b e r  1969) a n d w a s expressed a s  = [ ^ [ ^ . i ( / V - . i - / v . ) + g W. (N ^ 2  k  k  - N ) k+1  +  W. (N ^ C  C  -  tf )]](^) c  72 where  CPI=cohort  production  average  maximum  age  (Garman  &  1983).  Total  Waters  production  collections made year')  and  Production  were  from  from  by  production  the  levels  estimates  were  between  zones  Analysis  estimated July  for  is  that  to  vertical were  calculated  by  years)  each  was  zones  carried  S y s t e m (SAS) computer  out.  area  programs  in  the  time  this as  corrected  to  annual  on  an Cren  middle,  lower,  to  zero  tests  were  area  The  level.  1988).  with  'first year').  values  basis,  significant  performed  ( S A S Institute  span  as  by  most  tidepools  and base. T h e tide  for  between  'second  1972).  the  population  name  to  the  by  production  (referred  (Le  statistical  Analyses  represented  summing (I  described  relative and  1987  were  on  is  individuals  1988  zones: upper, fixed  fish  by  July  year  depend  in  species b y  October  second  to  which  through  Production  known  four  were  1986  the  (in  for  1987  365/404.  grouped into  follow  attained  August  estimates  multiplying aquatic  was  interval,  zones  Production differences Statistical  73  3.3  RESULTS  3.3.1 Distribution within the Study Area  Clinocottus occur  globiceps  a n d Oligocottus  sympatrically, pools.  C.  zero  level,  b u t occur  tide  were  rarely widely  (Figure  17).  Distribution factors. but  also  higher pools in  A  within great  with  vertical  the  fish.  Larger  concentrated usually less  than  (and/or  25  at tide  level  levels  as 2.70 m  above  (Figure  influenced  growth,  16). O.  by  rocks  C.  globiceps  abundance cover.  many  It  They  maculosus  level  large  allows  i n fair  tidepools.  tidepools  interacting a n d crevices to  inhabit  i n the more  rocky  m a y frequently  occur  of some tidepools.  although  between  lower  this  level  and often  m i d to  a n d lower  of macrophytic  than  the  i n middle  i.e., primarily  b y C. globiceps  globiceps,  m m were  cover,  areas  favour  pools as high  c a n be  eel grass)  amounts  to  bottom-dwelling  i n the m i d a n d higher  c a n be found  inhabited  on the lower  found  than  area  of habitat  i n open sandy  C.  from  numbers  higher  intertidal  moderate  range  collected  are both  tends  are concentrated  maculosus  numbers  The  the  algal  only  were  i n pools  amount  O.  pools.  globiceps  i n great  distributed,  dense  great  globiceps  observed  though  C.  although  intertidal  maculosus,  a n d O. maculosus  found  were  smaller  0.70 m  not common  in  throughout fish.  pools  the a r e a ,  Young  a n d 1.50 m inhabited  reflects  were  more  C. globiceps  were  (Figure by  the size of  16). Juveniles  adults.  Young  O.  74 N=12 60  UPPER  40 • 20 • 0  N=295  80 L  MIDDLE 60  N=241 LOWER  N=21 BASE  _l  0  10  20  30  40  50  60  Standard Figure  16. Length-frequencj' different levels.  70  80  90  100  110  L.  120 130  Length (mm)  distribution  of  C. globiceps  from  tidepools  at  75  N=412  200  UPPER  150 100 50 0 N=614  200 0  MIDDLE  150  1 ioo 3  o- 50 cu  L °  N=323  200  LOWER  150 100 . 50 0  N=0  50 0  BASE 10  20  30  40  Standard  Figure  17. Length-frequency different  levels.  50  60  70  80  90  Length (mm)  distribution  of  O. maculosus  from  tidepools  at  76  maculosus inhabiting of  do not occur pools  exposure,  affect  as high  i.e.,  as those  how long  the distribution  of the vertical  as low as do older  a  inhabited  pool  of intertidal  is  fish.  b y juveniles  actually  fishes  Larger  position of the pool in relation  (Figure  emerged  (Green  or  1971b).  to tide  O. maculosus 17).  factor  found  T h e degree  submerged,  This  were  may  is a  also  function  level.  3.3.2 A g e of Oligocottus maculosus  The  results  presented used  show  in T a b l e  otoliths  in  O. maculosus  that 19  are compared  ageing  this  species.  survives  with  up  a n earlier  A g e groups  to  four  study  as  years.  by Craik  computed  The  data  (1978) who  from  age —length  regression Log  age  otolithic were 3,  =  -0.7677  ageing  3.3.3  1.447  as shown  in age group  with only  +  0,  in  log total  Table  50% were  0.2% in age group  19.  length Only  i n group  1,  (Craik  1978)  15% of the 30% in group  compared  collected  well  with  O. maculosus  2, a n d 4% i n group  4.  Density  Mean  densities  two-year  of  period  and  middle  had  high  and  Appendix  both  (Table  levels  densities  than  species  20).  fluctuated  Densities  in the lower  in the upper  5). T h e density  of  considerably  in  of all ages  were  zone  globiceps  to middle  for C . zones  C. globiceps  than  ranged  all  usually  zones  less i n the  whereas,  the lower from  1.2  during  zone  the  upper  O. maculosus (Figure  18  (at the base level)  77  Table 19.  Age-structure of 0. maculosus from the present study as determined by age-length regression: Log(age)=-0.7677-fl.447Log(total length). Craik's (1978) data are presented for comparison. Present Study Age N % total 232 15.0 0+ 51.4 796 1+ 29.7 2+ 461 3.7 3+ 58 0.2 4+ 3 Total 1550  SL 18.9 29.7 43.2 55.2 61.3  Craik's (1978) Study %total TL N SL 23.7 127 34.3 36.6 34.8 194 53.2 52.7 41.0 67.0 11.8 53.3 43 74.3 1 0.3 60.1 365  TL 42.1 50.1 64.9 73.0  78 to  2.7  fish  (base  level)  (Table  20).  level) 9.8  to  per  m  to  7.5  (lower  fish  level)  as  and  per  the per  to  m  22).  the  lack  pools  1988.  were  The  globiceps, and  upper  1+  the  and  lowest age  The  absolute  of  survival  for  from  the  number  intervals at  was  possible to  19).  The  rate  during  than  derive  first  density  the  zones  between  the  highest  (Appendix  life  consequently  the  two  of  of age  survival curves years  of  any  quantitative  groups  in  the  curves for  the  life  older  was  year-class  shown  the  species  was  no  years,  and  22).  1987,  most  August  1987  and lower  substantially age  was  (Tables  among 21  (base  whole,  There  July  C.  for  more  groups were  O. maculosus in history  no  On  between  held  0.1  range  (Tables  (middle  year  the  either  0.6  age  lower  5).  C. globiceps and their  of  from  20).  and  from  second  constant  1986  densities  zones. Densities  groups  duration  for  interaction  O. maculosus)  for  the  C. globiceps.  fairly  July  and  ranged  (Table  of  were  in  second y e a r  level)  those  year,  level)  based on collections made  middle  survival  first  among  were  frequency  empirical the  and  entire  relative  in  (middle  2  sampled  throughout  Helby, the  m  and  higher  density  those of younger  tidepools  the  O. maculosus  zones with  than  successive  (lower  pool  of significant zone X y e a r  involving densities  than  area  per  differences  analyses  fish  of  level)  were  fewer  October  in  year  fish  Because  and  level)  in densities a m o n g zones and y e a r s  Relative  indicated b y  2  (middle  (lower  2  O. maculosus  of  m  first  15.7  significant difference 21  pool a r e a  fish  During  5.3  densities  of  2  in  (note  not pattern be  population 19  of  of as  the  mortality  and  of  both a a  the  at  for  illustrated.  indicate  steepness  year-class  estimated  population  Figure the  can  each  However, species,  whole high right  it  (Figure mortality limb  of  T a b l e 20. Mean annual d e n s i t y (N/m ), I n i t i a l biomass ( B i , g/m ), mean biomass (5, g/m ), mean i n s t a n t a n e o u s growth r a t e s (G), a n n u a l p r o d u c t i o n (P, g/m /year) and a n n u a l p r o d u c t i o n t o biomass r a t i o s ( P : B i , P:B) o f C. g l o b i c e p s and 0. maculosus. P r o d u c t i o n and biomass f o r 1987-88 i s based on c o r r e c t e d a n n u a l v a l u e s o b t a i n e d by m u l t i p l y i n g by 365/404. 2  2  2  2  J u l y 1986 - J u l y 1987  August 1987 - O c t o b e r 1988  Zones Cllnocottus  N Bl  B  G P P:Bi  P:I  Zones  olobiceps  Upper  Middle  Lower  Base  Upper  Middle  Lower  Base  -  2.7 18.6 21.1 0.51 10.8 0.58 0.51  1.9 13.0 11.3 0.78 8.8 0.68 0.78  1.2 13.5 8.3 0.57 4.7 0.35 0.57  0.7 2.4 1.8 0.94 1.7 0.71 0.94  1.0 4.1 2.6 0.77 2.0 0.49 0.77  7.5 29.5 26.0 0.66 17.1 0.58 0.66  0.6 6.7  5.3 5.7 4.9 0.76 3.7 0.65 0.76  0.1 0.03 0.03 1.00 0.03 1.00 1.00  15.6 10.5 9.3 1.20 11.2 1.07 1.20  15.7 14.3 13.1 1.04 13.6 0.95 1.04  9.8 9.4 7.6 1.09 8.3 0.88 1.09  -  -  -  O l i g o c o t t u s maculosus N Bi B G P P:B1  P:B  _  -  2.2 2.5 1.8 1.00 1.8 0.72 1.00  -  -  Table 21. A n a l y s i s of v a r i a n c e s t a t i s t i c s f o r growth, d e n s i t y , biomass and p r o d u c t i o n by years and zones f o r C . QJJQ  Source Year Zone Zone*year Error Year Zone Zone*year Error Year Zone Zone*year Error Year Zone Zone*year Error  Growth  Density  Production  Biomass  D.F.  SS  F  P  1 1 1 16  0.1070 0.1709 0.2326 1.8262  0 .94 1 .50 2 .04  0 .35 0 .24 0 .17  1 1 1 19  0.4671 0 .04 28.7909 2 .40 35.1182 2 .93 227.5485  0 .85 0 .14 0 .10  1 1 1 19  122.7273 1 .64 82.4755 1 .10 75.8039 1 .01 1419.168  0 .22 0 .31 0 .33  1 1 1 19  371.4819 1 .20 237.7747 0 .77 478.5106 1 .54 5891.303  0 .29 0 .39 0 .23  Table 22. A n a l y s i s of v a r i a n c e s t a t i s t i c s f o r growth, d e n s i t y . biomass and p r o d u c t i o n by year and zones f o r 0. maculosus. -  Source Year Zone Zone*year Error Year Zone Zone*year Error Year Zone Zone*year Error Year Zone Zone*year Error  D.F.  SS  F  P  1 1 1 12  1.2496 0.0329 0.0195 0.8917  16.82 0.44 0.26  0 .002* 0 .52 0 .62  1 1 1 11  210.2645 1.79 45.3809 0.39 23.6696 0.20 117.6899  0 .21 0 .55 0 .66  Production 1 1 1 10  99.2201 1.74 34.2435 0.60 10.1877 0.18 571.7371  0,.22 0,.46 0,.68  145.6370 1.90 33.1877 0.43 32.4460 0.42 765.0710  0..20 0..53 0. 53  Growth  Density  Biomass  1 1 1 10  81  o  Upper  Middle  Lower  100  Base  Oligocottus m a c u l o s u s  1987-1986  80 60  I  40 20  0  Upper  Middle  -,  Lower  Base  Zone Age groups  tm o  +  Figure  18. zone  Density for  the  of  \zzu =i_B 2  C. globiceps  period  1986-87  +  _ S 3 Csa 4+  and 0 . to  +  maculosus  1987-88.  o  by  5+  age  group,  in  each  82  60  0  1  2  3  4  5  Age (years)  Species Q. globiceps  Figure  19. and  Age-frequency  O. maculosus  distribution from  Helby  —I— Q. maculosus  curves  for  Island.  populations  of C .  globiceps  6  T a b l e 23. Summary o f r e g r e s s i o n a n a l y s i s v a r i a b l e s ( d e f i n e d i n t e x t ) . C o r r e l a t i o n s o f each s e t o f v a r i a b l e s were t e s t e d f o r s i g n i f i c a n c e , r - s q u a r e d v a l u e s a r e t h e p r o p o r t i o n o f t h e t o t a l v a r i a t i o n a c c o u n t e d f o r by t h e independent v a r i a b l e . Oliaocottus  maculosus  Clinocottus  alobiceDs  Variables  D.F.  r  r^  P  D.F.  Densities N w i t h depth N w i t h area  10 10  -0.68 -0.25  0.46 0.06  0.016 0.442  14 14  0.19 0.56  Growth G with B G l w i t h BI G2 with B2 G3 w i t h B3 G with N G l w i t h NI G2 w i t h N2 G3 w i t h N3 G l w i t h PI G2 w i t h P2 G3 w i t h P3  11 6 11 8 11 6 11 8 6 11 8  -0.13 -0.35 0.24 -0.03 -0.10 -0.43 0.22 -0.04 -0.01 0.42 0.57  0.02 0.13 0.06 0.001 0.01 0.18 0.05 0.002 0.0001 0.18 0.32  0.663 0.391 0.438 0.945 0.734 0.292 0.473 0.906 0.985 0.154 0.086  16 6 11 7 16 6 11 7 6 11 7  Production P with B P with B i P with G P w i t h depth P w i t h area P with perimeter P w i t h SI  11 11 11 10 10 10 8  0.98 0.96 -0.14 -0.45 -0.04 -0.10 0.21  0.96 0.93 0.02 0.20 0.002 0.02 0.04  r  r  2  P  0.04 0.31  0.480 0.024  -0.06 -0.10 0.02 -0.68 0.03 -0.32 -0.15 0.73 0.04 0.32 0.54  0.004 0.01 0.001 0.47 0.001 0.10 0.02 0.53 0.002 0.10 0.29  0.809 0.809 0.936 0.040 0.913 0.440 0.620 0.030 0.928 0.292 0.130  0.92 0.96 0.06 0.23 -0.25 -0.37 -0.24  0.85 0.93 0.004 0.05 0.06 0.14 0.06  0.0000 0.0000 0.815 0.383 0.357 0.159 0.469  i  0.0000 0.0000 0.645 0.139 0.902 0.690 0.565  15 15 16 14 14 14 9  84 the  curves).  were  not  The  very  adequately  youngest  fishes  represented  in  ascending left limb  a n d the dome  The  of  surface  globiceps; density  there  correlated  3.3.4  a  negatively these  were  weak  with  area  studied.  was  significantly  but  non-significant  surface  area  of  mostly  This  still  is  correlated  pools  density  23).  there  by  the  19.  correlation  (Table  (Table  planktonic  depicted  with  negative  O. maculosus;  for  C. globiceps  for  group),  of the s u r v i v a l curves in F i g u r e  density  two variables  large  different  species, middle was  was  pool  the  age  Depth  was  of  C.  between of pools  no  correlation  23).  Biomass  There at  the  O. maculosus and  of  between  area  (0+  differences  zones  (Table  substantially  (Table change  20).  the  biomass  Maximum 11.4 g / m  21 for  biomass 2  for  in  When  higher  Biomass the  years,  of  lower  differences  two  (Tables  5).  exhibits  20).  more  Relative  over  interaction highest  (Appendix  O. maculosus zones  in biomass  as  the  O. maculosus  is  all  age  zone  A g e groups and  in  middle  the  1+  &  the  was  lack  of  a n d upper  zones  species did not  &  respectively 2  both  C. globiceps  significant  g / m , for  for  the upper a n d  for  2 + , and 2 +  10.5  for combined zones.  at  zones for both  C. globiceps  second y e a r  zones  C. globiceps combined  among  the two species  by  groups  than  by  between  analyzed  biomass than  indicated  O. maculosus  age groups  biomass  in biomass  and 22).  during  by  C.  zone X y e a r  3+  had the  (Appendix  globiceps  5). and  85  3.3.5  Growth  Annual  instantaneous  (Appendix  5).  significant  differences  significant  There  growth were  to  rather  than  lack  There  was no relationship  the  amount  low  (r  lack  in  a n d 0.01  globiceps  negatively  (Table  species p<0.05) 2+ by  were  age  and  1+  and  a n d 22).  ANOVA vertical  b u t no  T h e lack  was  of  probably  location  of  pools  (Table  growth  indicated  and  23);  was  very  density  of  C. globiceps a n d O. maculosus age groups  O. maculosus were  for  age groups, growth  and m e a n  that  growth  23).  growth  T h e correlations  23);  the correlation  positive,  but  weak,  in growth  2+  (Table  changes  with  in  rate  by  Correlations  that a m o n g  (Table  differences  among  explained  statistically.  biomass  computed  groups.  by  zones,  species  2+  and  positive  and  w a s usually not  with its density.  mean  between  and 3 +  species was  of  21  within  population density - a n d growth  growth  23) indicating  correlated  Correlations  indicated  for all age groups  3+  insignificant  as  in growth  (Tables  differences  between  a n d insignificant  C.  zones  zones  increasing age for both  variability.  respectively) for  differences  among  consistent  of variability  of 0.001  2  of  decreased with  yearly  found  among  attributable  of  large  were  differences  rates  Significant biomass  for  these  of  for  all  for C .  age  that  there  age groups.  age  related were  1+ are  between  to  for  age groups  population  for  large  both  (r=—0.68,  insignificant  and 2 + no  groups  correlation  globiceps  groups  age group  all  negative  was negatively  suggesting  between  rates  biomass for  both  O. maculosus density-related  86 3.3.6  Production  Total  production for  year  and  6.9  production  in  5)  the  the  same  was  because of lower  overlap.  was  For  year  data  was  estimated  situation.  were  The  comparable effect  to  as  in  12.3  of  these  the  given  one  that the  bigger  size  similar  to  of the  high fish  in  from  some  of  estimate  almost  for  residents  lower (7.5  twice was  a  were for  per  were  the  the 11.7  both  first  and  11.0  C. globiceps  of  whereas  during  65%  the  species  these  same  (Appendix  not  level  computed.  C. globiceps  m )  here  a  because  sympatric that on  is  the  production  it  is  attributable  per  two  value  value  However,  fish  the  a  Speculations  could be  (7.8  when  2  for  2  not  pools. Second  production  estimate  case.  do  O. maculosus  devoid of  upper  be made  density  species  g/m /year,  sympatric  pools since fish  in  2+  the  1.9  2+ ,  C. globiceps average  analyses.  above  and  and  33%  and  where  level  production value the  tidepools  on production cannot  tidepool  1+  in  2  age groups.  O. maculosus  for  and  lower  2  1+ 1+  the  g/m /year,  estimate  other  noting  absent  g/m /year  Almost  age age  0+  those  some pools at  used  between  ages  for  8.1  C. globiceps,  for  occurred between  analyzed  of species interaction  estimates  occurred  less between  example,  zones was  O. maculosus.  for  densities of these  also  all  second y e a r  period  C. globiceps were  whereas  the  production  Production  Production  species in  second year  O. maculosus  period.  in  2  over  of  two  g/m /year  g/m /year 2  the  m ) 2  species  was  worth to  the  very  occurred  together.  Differences  in  production  among  zones  were  not  large  except  for  lower  zone  in  87 the  second y e a r  zone  than  20). to  in  The  greater  in the  production  annual and  (Tables constant  groups for  less  11.2  O. maculosus  g/m /year  tended  variance  and  over  (Table than  vary  zone  and  among  both  zones  22),  time  was  to  by  there  was  and  13.6 1.5  due  no  20).  O.  Annual  respectively,  g/m /year the  with  respectively.  2  in  was  O. maculosus in  2  C. globiceps  Table  were  (Table  for  2.0  and  O. maculosus  g/m /year  and  lower  probably  zone; there  estimates  zones  year  revealed  and  (Figure  as  years  relative  lower  zone  were  no  for  significant  the  above  significant  the  two  of  among  suggest,  differences  species  interaction  differences  values  (Tables  zone  with  zones  were  in 21 year'  fairly  20).  correlated  0.4  and  2%  of  respectively. for  was  second y e a r  among  that  age  biomass  being  indicating  all  mean  the  and  of  For  with  of  zones;  lack  23).  O. maculosus  in  a  poorly  Production  20  the  respectively.  2  and  85  were  zones  lower  in  (Figure  middle  in the  by  for  Again,  Production  zones  20).  zone  estimates  production  21  bottom  greater  C. globiceps  and  zones t h a n  C. globiceps  of  22).  the  (Table  Production was  for  upper  species differed  middle  production  analysis  pools of  and middle  two  zones  for  17.1  Though  middle  density  from  the  (Figure. 20).  values  corresponding  and  lower  caught  and  Production  and  production in  upper  for  upper  8.3  upper  obviously  maculosus  the  the  lower  their  C. globiceps  for  both  with  instantaneous  groups changes  combined, in  year  Production  was  species.  Fluctuations  growth  rates  variability  class  in  production  significantly in  mean  for  growth for  age  accounted  C. globiceps  positively  biomass  most  correlated  accounted  and 90% of fluctuations in production rates of C. globiceps and O.  for  maculosus  Upper  Lower  Middle  100  Base  Clinocottus globiceps  1987-1988  80  5  60 40 >—  CO CD 20  E 3  o  C  10  o o  __L Upper  Middle  Lower  1986-1987  Base  Oligocottus maculosus  8-  T3 O  Upper  I  Base  Lower  Middle  60 rOligocottus maculosus 50 40 30 20 10 0 -  I  1.  Upper  Middle  Lower  Base  Zone Age groups  Figure  20. each  Production zone  for  !o+-1+  1Z21+-2+  of  C . globiceps  the  period  KS4+-5+  a n d O.  1986-87  to  maculosus  1987-88.  by  age  group,  89  Figure  21.  Relationship  and  1988  year  between classes  of  production  and  C. globiceps  and and  initial  biomass  0. maculosus.  of  1987  90 respectively was  (Table  investigated  23).  by  The  relationship  between  regressing the former  production  on the latter  and  initial  for both  biomass  species.  A  plot  of the production and initial biomass values for C . globiceps a n d O. maculosus is presented variation  Depth,  in  Figure  21.  in production for both  a p h y s i c a l character  ranging  from  0.1  to  1.9  between  tidepool  However,  the correlation  enough area,  irregularity apparent  between in  production for  estimates  maculosus presented  the  this  in  surface  a  (a  positive  made  for the  Table  was  24.  and area  and  a negative measure  these  p>0.05)  of  of  less  the  accounted  for  important.  O. maculosus  6  C. globiceps.  for  (Table  degree  (Table and  values  correlation  O. maculosus  relationship  variables  was high  23).  of  Surface  regularity  or  N o correlation 23).  0.2%  For of  was  example,  changes  in  respectively.  method  spanning A u g u s t  are  substantial  instantaneous  growth  method  are generally  higher  for  93%  a m o n g pools with  a n d production for  a n d the  observed  for  of Production  period  There  accounted  but non-significant  (r = 0.24,  of production b y the size-frequency  method  trend  biomass  substantially  seemed to be somewhat  C. globiceps  size-frequency  same  was  depth  index  production  pool  varied  to suggest  Size-Frequency Estimate  The  by  pool)  initial  production  between  a n d shoreline  of a  variations  There  and  p<0.14)  in  species.  of pools,  m.  depth  ( r = —0.45, perimeter  3.3.7  Variability  1987  differences rate for  O. maculosus,  but  for  C. globiceps  through in  method.  the  values  1988  middle  O. are  obtained  T h e estimates  C. globiceps the  October  and  from  in all zones. T h e zone  had  lower  -  91 estimates. growth and  Annual  rate  from  method)  to  (instantaneous  instantaneous  growth  might  have  development  O.  for rate  been  growth  maculosus  method  are  6.9 g / m / y e a r  as  recruitment  for  a  for C.  method) to  24).  The  both  species  result  (usually  (instantaneous  2  rate- method)  (Table  lower  underestimated  a n d prolonged  from  (size-frequency  2  2  method)  ranged  15.8 g / m / y e a r  11.0 g / m / y e a r  (size-frequency  rates  production estimates  of  starting  19.5  globiceps g/m /year 2  estimates  by  because  growth  nonsynchronous around  the  April  cohort through  August).  3.3.8 A n n u a l Production to Biomass  Annual  production  globiceps  a n d higher  biomass larger  ratios  mean  followed  species a n d higher i n both  than  the  second  both  species  middle  ratios year.  (Table  a n d lower  exhibited  the  observation  that  ratios  trend  were  with  with  also  globiceps  16 a n d 17).  (Table  juvenile  the first  being lower  20). T h e r e year  having  considerably  w a s higher during  in  trend  C.  the  upper  is  annual  lower  values  zones for  followed O.  levels  by  maculosus  consistent  i n the higher  for the  were  between  the second y e a r .  This fish  for the larger  20). Production to initial  as P : B ratios,  varied  P : B i ratio.  more  lower  species (Table  T h e P : B ratio for C.  (P:B) were  maculosus  (P:B a n d P:Bi),  trend  there  (see F i g u r e s  0.  the same  These  zones  ratios  for the smaller  20).  same  biomass  for the smaller  (P:Bi)  variations  levels  to  Ratios  with  than  the lower  92  Table 24. Annual production, mean biomass and P:B ratios as estimated by instantaneous growth rate (IGRM) and size-frequency methods (SFM) for 1987-88. Mean biomass and production are expressed as g.m~ and g.m~ .year respectively. 2  Zone Upper Middle Lower Combined zones Upper Middle Lower Combined zones  2  C. globiceps IGRM B P P:B B 0.94 1.8 1.7 4.7 5.7 2.6 2.0 0.77 17.1 26.0 0.66 27.7 10.1 12.7 6.9 0.68  SFM P 6.2 7.7 33.6 15.8  P:B 1.33 1.35 1.21 1.24  0. maculosus 9.3 11.2 1.20 13.1 1.04 13.6 7.6 1.09 8.3 10.0 11.0 1.10 ~  31.7 8.0 18.7 19.5  1.89 3.81 1.68 1.95  16.8 2.1 11.1 10.0  -1  93  3.3.9  Estimation  of Production  Mathews  (1970)  curve  instantaneous  an  or  that  (Waters  suggested estimate a  easy  1969,  that  the  production  given to  site  by  determine  throughout mortality  the  year  rates.  from  a  single  does  riot  The  suitability  production  of  a  since or  changes in  while  variability  it  is  the  1970; to  fish very  of  on  abundance  ratio.  sample  hand,  mean  (P:B)  mean  to  1978b) be  estimate  Moreover,  its  to  biomass is  not  sampling  growth  be  have  used  biomass  regular)  of  and  determined estimation  individuals collected be aged.  initial  biomass. T h e  biomass accounted for  in m e a n  number  mean  easily  biomass in predicting production was  and  A  may the  (and  Allen  time c o n s u m i n g ,  study.  can  weight.  either  Chapman  However,  a  by  and  multiplying  continuous  other  and  1971;  biomass by  either  the  tedious  Allen  mean  P:B  production  course of this  invertebrates  requires  biomass,  that the  initial  in  Mathews  and  of  method  apparent  Biomass  ageing of individuals in  of initial  that  rate  predetermined  estimate  mean  estimation  production  fishes  Initial  require  on  1977;  ratio of  the  growth  observation which became  authors  at  noted  from Initial  biomass explained  results 93%  85%  tested  presented  of the  in  variation  of fluctuations  b y regressing  Table  23  show  in production,  in production.  94  3.4  DISCUSSION  Production which  rate  is regulated  in t u r n  accuracy  are regulated  of calculating  density  are  subject  to large  determined.  this  assumed  that  possible or not  to  fish  to differentiate  other  factors  entirely  closed  affect  accuracy (Green  density  fluctuations.  population  Beverton  1971b,  natural  and  and C .  Holt  mortality  C. globiceps Gibson  While  (1982)  therefore  in  that  biomass  therefore  that the  particular  growth  are  and  sometimes  source of inaccuracy  that  since  of  the  homing  it  w a s not  to  mortality  study  pools  tendencies  is  least  a minute  c a n be  tidepools  less important  at  it  the  factor  only  4  could these  in c a u s i n g  and 5  fraction  of  are  years  of the  for tidepool  18 a n d 19).  have  is  the  no direct  by other  has rarely an  out  However,  attributable  that  strong  lifespan  are eaten  predation  accurate.  or  predation I  collection, therefore  argue  the  respectively,  populations.  not constitute  in  in abundance  the movement  an age (Tables  0. maculosus  notes does  fish  size  are  into  but  potential  believe  follows  the principal  O n e could  estimates,  globiceps  (1957)  and  fluctuations  movements  the  It  and their  on how accurately  complete  estimates  1973) m a k e  approach such  &  involved  e.g. predation. systems,  depends  present  1978b).  abundance between  fish  a n d density.  a n d are often  sampling  of density  species  O. maculosus  errors  of  of population  (Chapman  numerical  numbers  production  Estimates  sampling  study,  the  b y growth  annual  in production calculations  During  by  important  been factor  most  general  evidence  tidepool observed  to  cause  suggest  that  In  context  fishes.  this  in intertidal  controlling  of  the  fishes  populations  95 of intertidal (Paling  1968),  (Gerking both  fishes. A p a r t senescence  1957;  species  Craig  the  and  may  to  However,  the  demonstrate,  growth  rate  and  was  remarkably  g/m  and  2+ )  made  contributed  for  the  10.2  O.  significant  to  (Mathews fish  g/m  for  are  (1976)  all  Mann  most  of  15.8  10.0  groups  g/m /year 2  Mean  contrast, This  1971;  productive  in  and  qualify  as  in be  size  other difficult  unless  the  by  the  by  growth  24).  2+  observation  for  the  rate)  in  1976b)  contribute  two  and  age  both  species g/m  and  groups  (1 +  65.4%  consistent which  have  up  95%  2  rate)  species,  groups  to  growth  14.05  growth  Young  the  size-frequency  instantaneous  contributed  is  by  2  C. globiceps age  may  by  g/m /year  biomass  production  Chadwick  and  suffered  described  (instantaneous  2  to  1+  would  11.0  2  maculosus (Table  contribution  populations  rate  population  g/m /year  g/m  fish  death  measured.  (instantaneous  2  production.  1971;  the  In  and  respectively.  globiceps and  species.  32.8%  production for  Moring  methods  19.5  causing  populations would  production:  and  19)  mortality  wild  estimates  wild  mortality  (Figure  the  factors  some  in  senescent  highest  6.9  O. maculosus; age  in  production  young  C.  a  life  could be  method  produced  (size-frequency)  2  obviously  studies  rate  the  in  increase  of  resemble  of mortality  similar: for  year  absolute  had  size-frequency  (size-frequency) 11.0  thus  extrinsic  demonstrated  presumed  second  factor  C. globiceps  method.  The  with  Oligocottus maculosus instantaneous  been  and other  onset of senescence in  even  p r e s u m e d predation  predation  has  1985).  after  age-dependent, fishes.  from  most  to  1+  total  and  with  several  shown of  2 +  that  the  total  ages.  reported  average  densities  of  1.64,  2.66  and  2.86  fish per  m  2  of  96 "effective in  sampling  Trinidad  0.1  to  study  fish  of  from  present  0. the  A  analis from  a  study  the first  tidepool  area  first  for C .  of their  (1986)  from  to  to  range  globiceps appear  the  (Yoshiyama densities  the average  from  8.5  study  to  densities of  represent  total  lower  et  al.  that  northward  1986). T h e  C. globiceps  density 27.0  fish  per m  fish  in  Clinocottus  of  per m  the a r e a  initial  than  i n the present  latitudes  of  considering  sites  observation  of 0.6 to 7.5  reasonable  densities  2  with  to higher  determine  estimated  with  for the second y e a r  range  at three  are substantially  consistent  the present  (460 m ) . T h e above  they  compared  as one moves  attempt  California  Estimates  year,  is  0. maculosus  for  densities  reported  2  increase limits  area)  these  difference  b y Wells  southern  surface.  of  2  2  of  that w a s  biomass  of 11.4  for O. maculosus and 10.5 g / m for C . globiceps during 1987 — 88. 2  Annual were  This  southern  is  tidepool  sampled  5).  maculosus  study  tidepools.  2  Although  in  2  as tidepool  of 9.8 to 15.7 fish per m  densities  g/m  per m  (Appendix  away  (same  B a y , California.  5.3  densities  area"  variations closely  a n d positively  tidepools  at H e l b y  for  most  the  globiceps directly  structure  and  Island  part  linked  (Figure  to  C. globiceps  within  and  of both  zones remained suggestions that  of C. globiceps a n d O.  changes  22). A n n u a l  did not depend  age structure  consistent with  a n d production  on  O. maculosus respectively).  affected  Population  i n biomass  in  variation  density  2  Whatever  relatively  varied  controlled  i n the  w a s little, a n d  a n d 0.01 density  for  C.  therefore  production.  from  constant  c a r r y i n g capacity  numbers  i n growth  (r =0.001  O. maculosus  species  population  maculosus  year  (Figure  to y e a r ,  b u t population  18). These  m a y be determined  results are b y available  97  6  8  14  Density (N/m ) 2  10  15  20  Density (N/m j 2  Figure  22. Relationship between production a n d density y e a r classes of C. globiceps and O. maculosus.  of  1987  and  1988  98 habitat  and  having  available  Bohlin age  possibly  (1978)  suitable  study  has  individuals  These  observations  patterns is  as  of  2.2  zones  differ  zones  the  similarity  There  than  at  that Depth  present.  For  plaice  and  live  was  of  lower  related  long  deeper  of  Heincke water.  the  by et  at  al.  the  be  among  size or  zones.  a  The  tendency  pools  and  log P  on  (Figure  The  of 16).  will  be  and  influence  (1971b)  for  as  the in  (1.7  the  a  two  way  of  regressions did  support  in  differences  partly  (cited  log B  zones  the  physically upper  null  similar  and  production  middle among  component of production.  area  to  in  C. globiceps  middle  results  similar  distribution  for  slopes of  These  Again,  could  the  covariance  O. maculosus  20).  pool  reflects  upper  of  1986).  numerical  (1913)  Furthermore,  may  production estimate  analysis  for  known  Green  species  (p>0.52).  pools  been  two  production  the  higher  regressions of  sites  between  area  the of  (Table  to  structure  pools  above suggestion.  estimate  production  correlation  has  for  (Portt  zone  than  the  The  fish  available  level  compared  high  example, in  the  between  rates  surface  density.  of  age  within  1984).  rather  feature  variation  primarily  significant  indicated  the  the  One  Bachman  habitat  or  retention  exhibited  zones  respectively).  interzone  size  and  globiceps populations  lower  by  species.  computed  zones were  The  examined  that  C.  to  1966;  of suitable  consistent with  significantly  hypothesis  that  migration  (Chapman  different  favour  are  2  were  habitats.  to  g/m /year  determining not  shown  with  amount  in  these  apparent:  and  habitat  resulting  larger  Production  sites  suggested that  dependent,  present  foraging  in  density explain the  Cushing  observed  of the  size  (C.  globiceps)  variability and  1981)  that  fish  age  stated  0-year  O.  in  fish  of  fishes  that  larger  maculosus  99 were  concentrated  pools  in  made  for  depth  of  the  species is  of  The  this  habitat. and  P:B  ratio,  Hopkins  mean  lower  is  1971)  biomass is  and  suggested in  and  resultant  the  P:B  the  varies for  the  similar the  ratio  regression was  on  to  P:B  study  present  mean  significant  and  decrease virtue  biomass  ratio  of  with  the  with  on  r = —0.45). leads  to  distribution  species for  some  respect to  age  classes  were  made  the  any  P:B  the  biomass takes  the  increasing  (P:B)  ratio  of  Generally  and  have  (Chapman  1965;  C.  form  zones  pointed to  this  study  might This  be  idea  globiceps  the with  are  out  initial  those  that  biomass  Hunt  1966;  (Appendix related was  C.  and  O. maculosus)  production  in  0.62 the  of  trophic  larger . size,  zone.  for  production  their  ratio  given  of  of  0.98.  to  rate with  researchers  similar  that at  turnover  By  Other  trend  younger  log  identical  C. globiceps  for fish.  a  to  mean  observations  present  correlated  which  with  between  were  O. maculosus  of  1976b)  consistent  the  1973).  middle  in  earlier  of pools ( p < 0 . 0 5 ,  species are  value  higher  of juvenile  biomass of individuals  regressing  for  a  Production  differences  indicates  (Emlen  had  higher  two  out  deeper  distribution.  production  (upper  which  no  these  inhabited  significantly  (Nakamura is  fish  pointed  not  depth  water  correlation  intertidal  longevity  populations  ratio,  shallow  Furthermore,  to  was  with  larger  as  C. globiceps.  correlated  negative  tendency  a  ratios  large  This  The  had  higher  (P:Bi)  production  observations  for  prefer  whereas  production  suggest that  O. maculosus  smaller  the  species.  greater  globiceps  with  to the  seem to  of  However, r = 0.24)  that govern their  ratio  level  that  tidepools  Similar  negatively  known  suggestion  factors  the  was  upper  zone.  (p>0.05,  hand  correlations  the  intertidal  pools  patterns  shallow  C. globiceps.  other  This the  the  in  to  tested  (Figure  23).  5). the by The  100  Figure  23.  Relationship  between  P:B  ratio  and  mean  biomass  (B)  of  C. globiceps.  101 P:B  =  1.37  The  log of m e a n  can  be  seen  biomass  that  indicating  In  this  0.30  from  Figure  is,  turnover  study,  a  variation  in m e a n  also lower  when  statistically  relative  ease  ratio  of  than  P:B  Production compare of things not of  production.  nonsynchronous production  Bi  the  size-frequency method  is  biomass (r  (P)  2  (Bi)  was  can  be  estimated  2.83,  s  2  the  contributed  to  instantaneous  Based  and on  the  were  it  a  implies  rate  method  recruitment,  could be considered more  accurate.  explained  of the  by  estimate  as  the  B  and  given  useful  than  the P:Bi  (Table  of P : B i  for  range  and  gave  is  subject  both  of  a  better  1986).  on  the  24).  A  size-frequency it  predictor  is  al.  not  higher  was  21).  (Portt et  thus  biomass  biomass (Figure  that  method  differences,  was  variance  did  mean  C. globiceps  more  less  It  survival.  error  and  discrepancy: the  protracted these  P  method  development,  growth  of  biomass data  rate  and  than  standard  range  This  growth this  =0.93)  with  increasing  production of  makes  size-frequency  instantaneous  with  on growth  predictor  which  =0.400).  production f r o m  2  P : B ratio.  correlated  regressed on initial  good a  =0.033)  in  The  as  2  in the  negatively  (r  =0.85).  variation  declines  of variability  is  s  development  estimates.  ratio  production  nonsynchronous cohort The  of the  corroborated b y  0.89, to  30%  p<0.05).  changes probably  initial  from  have  of  production  (0.46  with  by  n=17;  P:B  biomass (B)  to predict  might  affected  in  to  estimates  well  rate  is further  (0.22  P:B  that  proportion  which  of production. T h i s  variance  23  speaking, B i  with  globiceps  (r=-0.55;  dependent  higher  variation  Since  B  biomass accounted for  by  was  C.  In  some density  explained the  -  is  estimates effects  which  estimates  number  method  higher to  whole,  of  depress by  the  IV. G E N E R A L D I S C U S S I O N A N D C O N C L U S I O N S  The  objectives  of  this  Clinocottus globiceps comparison  thesis  and  to  of  otoliths  in  globiceps.  This  work  has  of  individual  190  mm  1973), ratio about  The  but from  maximum  unity  population  as  of  recruits coast  to of  lacking.  in It  in  were  July. would  total  17,  Helby  groups  in  modes  over  followed for  the  entire  year.  of  this  study  growth  information  west  In  The  has  at  of  lack  for  least  suggests that  any  males  of  species  in  a  years,  the  total  length  Island  (Hart  deviations and  the  Clinocottus  5  Vancouver  of  growth  demonstrated  B . C . waters,  coast  time  Is.  consisted of  dominant  newly 8).  Similar  1978),  in  females  conduct  from  102  in  maculosus at although data  sex have  winter  length-frequency  a  of  dominated proportion location  on  dominated  winter  sampling  and  times  the  recruits for  major  certain  juveniles  changes  distribution  to  three  during  recruited  Oligocottus  (Craik  instructive  of  are  1988  (Figure  found  progression  dynamics  and  Island.  Length-frequency be  the  age  present  length.  reported.  Table  August  Island  and  the  C. globiceps lives  from  age  distributions  adults  mm  was  Helby  in  Vancouver  population  age  Some  example,  length-frequency  168  shown  investigate  production  age  that  C. globiceps at  year-classes. For  found  recorded  equal lifespans at  smaller year.  no  been  to  maculosus. T h e  providing  measured  has  been  evaluate  Oligocottus  with  usefulness  largest  have  months so  distributions  that could  two the the of west the are the be  103 Age  determinations  and  unavoidably  age  (Beamish  still  a  zones  1979).  The  could  Exploited more  fish  prone  otolithic  otolith  radius,  particular in  its  possible  to  where  new  most  the  fish  present  thin  technique  work  to change  the  flat  of  with  has been  such  ageing &  as fish  i n otoliths  experience  of  clearly  reading  age classes,  (MacDonald  spring  after  and summer  which  it  is the  visible  whole  otoliths.  conducted  m a y be  a n d hence  it would be  as  to  cottids  was  Pitcher  on the  otolith,  was some  the body  easily  phenomenon.  C.  1969) probably  to  on  length  improve  corroborated 1979),  explore  rapid  by  a  a n d the two  the  lines  nucleus  the  demonstrated  the  production  time  for body  at  present dynamics  to  it  on a  was  demonstration  negative  of  size-selective  factors.  study of  when  a n y previous age  the  mortality  the  length  otolith  back-calculations  through  natural  during  the  of the  of the fish  From  mainly  is probably  regression  mortality  arising from  where  which  from  globiceps  environment  opportunity  the  calculated.  information  a n d was  slowed,  distance  tidepool  and  perfect  well.  to  history  on  techniques  otolithic  are not  of the zonation  populations  study  annuli  is known  extent  ageing  the  (Ricker  an  developing  and measuring  gather  growth  great  plotting  mortality  provided  a  By  Lee's  marine  to  maturity.  reversed  The  In  of otolith  the interpretation  unexploited  of life  point  life  use  reach  otolith  because of the disrupted  confined  C. globiceps  in  agreed  years  since  C. globiceps being  distribution  ageing methods  interpretations  relying  of  useful  to  length-frequency  two  process  to error  was  errors,  populations  ageing.  Growth  on the  Furthermore  otoliths  be  advantageous  first  contain  subjective  observer.  based  was  two  undertaken  cottid  species.  104 Production cohort  in  of  rates  the  the  to  study  in  study  the  estimate  an  to  over  the  This  study  high  was  whole  it  very  reasons.  First,  samples  (Figure  (Grossman  up  1982;  dispersal  to  production  O. maculosus habitats fewer,  for  more  the into  fish.  and age  turbulent  high  to  the  of  support my  al.  was  best  and  (Appendix  tidepool  C. globiceps  C.  to  of  and  is  probably  phase  of  these  between  age  Second,  was  ages  found  difficult  to  in  this  study  the the  tidepool following in  tidepool  and  cottids  older  the  in  1+  study,  the  to  which  turn  can  than  substantial. pools  be  however,  C. globiceps  pools were  sub tidal  sample.  1+  the  production.  attributable  0+  biomass  and  represented  species  and  not  high  O. maculosus  in  other  In  0+  the  in  O. maculosus  production  thus  1971).  A  figure.  for  adequately  The  production  production,  underestimate  1986)  and  its  of  production  shown  survival  species.  secondary  been  globiceps are  and deeper,  fish  has  5).  in  rate  each  successful species  high production  not  (Mathews  12.2%  and  a  very  seen  substantial  an  were  larval  production  of  a  growth  estimate  probably groups  et  contribution of  that  biomass of  mortality  success  tidepools. Production between  11.3  especially  as  estimate  may  was  and  Clinocottus globiceps.  of  well  discrepancy  respectively  their  that  planktonic  total  only  older  hence  individuals, are  of  of  was  stressed  This  growth,  ecological  success  as  tidepools  Yoshiyama  recruitment  95%  times  quantitative  younger  19).  the  ecological  preliminary the  of growth  density,  lifespan contributed  a  be  synthesis  Oligocottus maculosus  1.5  that  must  of  numbers)  of their  However,  influences  over  attempted  is  its  population  indicate  passive  and  a links  picture  shown that  site  estimates  habitat  overall  has  which  (due  is  population,. therefore  give  present  ecological sense  not age  Perhaps where  and  optimal 4+  were  large waters  105 Production  estimates  compared other  one  growth  and  physical  a  studies  and  a n d the  Weatherley  &  fish  production  in  (1987)  to  ecosystem of which  fish form  looked  annual  not  into  take  that  is  into  as  one  production  the  of a  environmental  variable  of  winter  growth  in  affecting would  to  research  a n integral  part.  effects  a  the  undoubtedly  relate  as  of  they  in this  and  the  with  in  the  entire to  (1972) a n d to  F o r example,  T h e present  aquatic  study  did  is worth suggesting  the  environment  reduction or  view  needs to be  season could become  little  production  be desirable  aspect which  to  tidepool  processes of the  of  habitat  predation)  Le Cren  in production. It  in  are  estimates  relate  area.  Another  (e.g.  production  Production  result  and  processes  competition  parameters  production.  they  study)  physical  on production.  particular  when  as a consequence of fish  of production  physical  species,  (e.g.  species  consideration seasonal variation  dynamics  present  production  from  23).  the complex  the  the  only  pointed out that it would  seasonal  incorporates  the  production  biotic  between  correctly  in  of  (Table  for future  relation  case  attempted  interactions  themselves,  isolation  and  tidepools  possible avenue  in  treated  in  study  could be a  the  herein  dynamics,  of  value  understanding  cited  present  characteristics  Gill  was  temperature)  The  taxocene  (as  better  population  environment.  not of m u c h  another  yield  Many  dimensions  fish  with  factors  ecosystem.  are  no  into  an  important  (or  cessation)  production  of the  fish.  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Length-frequency distribution for C . globiceps: Males. Length interval 0-9.9 10-19.9 20-29.9 30-39.9 40-49.9 50-59.9 60-69.9 70-79.9 80-89.9 90-99.9 100-109.9 110-119.9 120-129.9 130-139.9 Total  Frequency 0 16 95 93 75 44 47 53 23 10 1 1 1 0 459  Percent 0.0 3.48 20.70 20.26 16.34 9.59 10.24 11.55 5.01 2.18 0.21 0.21 0.21 0.0 100  Appendix 3. Length-frequency distribution for C. globiceps: Combined data. Length interval 0-9.9 10-19.9 20-29.9 30-39.9 40-49.9 50-59.9 60-69.9 70-79.9 80-89.9 90-99.9 100-109.9 110-119.9 120-129.9 130-139.9 Total  Frequency 5 49 159 172 147 100 116 114 64 18 2 1 1 1 949  Percent 0.53 5.16 16.75 18.12 15.49 10.54 12.22 12.01 6.74 1.90 0.21 0.11 0.11 0.11 100  123  Appendix  4. Age-length key f o r Helby I s l a n d C l i n o c o t t u s g l o b i c e p s . number of f i s h per age c l a s s by sex.  Length Interval (mm)  showing the  Length d i s t r i b u t i o n of age groups M  0+  F  15-19. 9  4  2  20-24. 9  6  1  25-29. 9  11  10  30-34. 9  14  9  M  1+  F  8  7  35-39. 9  29  24  40-44. 9  26  29  45-49. 9  15  14  M  2+  F  13  11  50-54. 9  19  19  55-59. 9  7  12  60-64. 9  4  13  65-69. 9  1  0  M  3+  F  3  2  12  21  70-74. 9  15  13  75-79. 9  7  4  M  4+  F  4  9  80-84. 9  6  7  85-89. 9  1  0  M  5+  F  3  3  90-94. 9  2  4  95-99. 9  0  0  100-104 .9  0  0  105-109i.9  1  0  6  7  TOTAL  35  22  78  74  44  55  37  40  11  16  Appendix 5. Mean annual d e n s i t y (N/m ), i n i t i a l biomass ( B i , g/m ), mean biomass (B, g/m2), mean instantaneous growth r a t e s (G), annual p r o d u c t i o n (P, g/m /year) and p r o d u c t i o n t o biomass r a t i o s (P:B) of C. g l o b i c e p s and 0. maculosus by age groups. Age groups were designated 0+ t o 5+. Biomass and p r o d u c t i o n values f o r 1987-88 are based on c o r r e c t e d annual values by m u l t i p l y i n g by 365/404. 2  2  2  Clinocottus globiceps.  July  1986. t o J u l y  1987  Zone: M i d d l e  N Bi  B  0+  1+  2+  3+  4+  5+  0. 7 0. 3  5 .4 9 .9  3.9 16.9  2.7 25.4  1.6 23.3  1.8 35.7  G P  P:B  1.0 1.76 1.8 1.8  10.3 0 .72 7.4 0.72  14 .1 0.89 12.5 0.89  50.9 0.45 22.9 0.45  29.1 0.32 9.3 0.32  Lower N Bi  B  2. 2 0. 6  P  P:B  2.5 6.8 2.72  2 .4 5 .1  7.8 5.7 0.73  3.4 12 .7  12 .8 10.0 0.78  1.7 16.6  19.0 10.0 0.53  1.3 21.7  14.6 11.2 0.77  0.7 21.4  Base N Bi  B  0 .5 0 .4  1. 0 0. 3  G P  P:B  _  2.2 2.64 5.8 2.64  -  -  -  0.5 6.2  6.3 0.05 0.3 0.05  0.8 12.3  16.3 0.5 8.1 0.5  A l l zones P Percent  2.9 7.7  6.3 16.9  7.5 20.1  11.1 29.7  9.5 25.6  3.1 48.1  Appendix 5  —Continue.  Clinocottus globiceps  August 1987 to October 1988  Zone: Upper  N Bi  5  0+  1+  2+  3+  1 .3 0 .5  1. 5 2. 8  0.2 0.8  0.3 4.2  G P  P:B  0.9 1.44 1.6 1.44  0.6 0.67 0.4 0.67  2.5 1.4 3.5 1.4  4+  3.2 0.16 0.5 0.16  5+  0.2 3.5  -  0.3 4.5  0.1 2.0  Middle 1 .6 0 .9  N Bi 1 G P P :B  1 1.06 1.1 1.1  1. 8 3. 0  5.1 0.98 5 0.98  1.7 6.4  3.3 0.82 2.7 0.82  0.8 8.1  3.2 0.31 1 0.31  0.5 0.6 0.3 0.6  Lower 8 .2 4 .7  N Bi 1 G P P :B All  7.7 1.14 8.8 1.14  21 .7 37 .9  31.7 0.88 27.9 0.88  7.4 34 .2  35.9 0. 65 23.3 0.65  5.4 55.3  41.6 0.41 17.2 0.41  1.6 25. 5  13.2 0.63 8.3 0.63  zones  P Percent  3. 83 11.3  11.1 32.8  9.83 29  6.23 18.4  2.87 8.5  0.6 18.4  Appendix 5  —Continue. J u l y 1986 t o July  O l i g o c o t t u s maculosus  1987  Zone: Middle 0+  1+  2+  3+  4+  1.6 0.3  4.6 2.0  4.2 7.5  0.4 1.3  0,2 1.2  N Bi  B  G P P:B  0.8 0.75 0.6 0.75  3.8 1.47 5.6 1.47  3.3 0.73 2.4 0.73  1 0.,5 0..5 0..5  Lower 10.2 7.5  N Bi  I  G P P:B  7 0.69 4.8 0.69  5.3 7.4  2.8 0.93 2.6 0.93  0.5 2.3  -  Base 0.1 0.01  N Bi B G P P :B All  0.03 1 0.03 1  -  0.1 0.1  -  zones  P Percent  0.21 3.9  3.47 64.9  1.67 31.2  -  Appendix 5  --Continue.  O l i g o c o t t u s maculosus  August 1987 to October 1988  Zone: Upper  N Bi  5  0+  1+  2+  11.0 1.8  37.0 14.1  13.8 24 .0  G P P:B  2.9 0.83 2.4 0.83  17.4 1.45 25.3 1.45  3+  7.4 0.8 5.9 0.8  4+  0.5 2.3  -  Middle N Bi B G P P:B  9.3 1.1  5.7 1.23 7 1.23  40.4 18.6  25.6 1.22 31.3 1.22  26.4 42.3  19.8 0.81 16 0.81  2.1 8.1  1 .3 0. 11 0. 14 0. 11  0.2 1.4  Lower N Bi B G P P:B All  2.7 0.5  3.8 1.13 4.3 1.13  27.9 15.2  11.4 1.51 17.2 1.51  6.7 14 .4  7.6 0.46 3.5 0.46  zones  P Percent  4.57 12.2  24.6 65.4  8.47 22.5  2.0 7.6  _  -  


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