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Social behaviour in a non-pupping colony of steller sea-lion (Eumetopias jubata) Harestad, Alton Sidney 1973

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SOCIAL OF  BEHAVIOUR  STELLER  I N A NON-PUPPING  SEA-LION  (EUMETOPIAS  COLONY J U B ATA)  by Alton  B.Sc,  Sidney  University  A THESIS THE  Harestad  of B r i t i s h  Columbia,  SUBMITTED IN PARTIAL REQUIREMENTS MASTER in  FOR OF  THE  1971  FULFILMENT DEGREE  OF  OF  SCIENCE  the Department of Zoology  We  accept  required  THE  this  thesis  as conforming  to the  standard  UNIVERSITY  OF  August,  BRITISH 1973  COLUMBIA  i  In  presenting  this  requirements  for  British  Columbia,  freely  available  that  permission  scholarly  shall  I agree for  that  degree the  reference  may  be  of  without  and  8,  study.  Canada  shall  granted  by  the  thesis Head  written  of  for financial  permission.  of i t  agree  I t i s understood thesis  the  make  I further this  Columbia  of  the U n i v e r s i t y  of  of of B r i t i s h  at  copying  this my  fulfilment  Library  his representatives.  allowed  University  Vancouver  partial  advanced  publication  n o t be  in  for extensive  o r by  or  Department The  the  purposes  Department copying  thesis  for my that gain,  i i  ABSTRACT JjJl^iociias segregate  during  consisting colonies studies  of  of  social  males mainly  social  only  polygyncus.  breeding  adult  the  i s  the  consisting  concerned the  jubata  with  season  and of  behaviour  and  into  sub-adult 1  of  sea-lions  pupping  females,  behaviour  pupping  The  and  non-pupping  males.  A l l previous  E_. j u b a t a  colonies.  This  organization  colonies  have  study  of  a  been  describes  non-pupping  colony. The  colony  i s  Columbia  ( L a t . 52016*  100-150  animals.  several  adult  concerning One of  method 34  and  method  as  well  were  taken of  an  as  though  present.  Data  using  two  ethogram  spacial  notes  on  methods. consisting  of  social  organization  territorial  population  of  males  observation  involved  British  It consists  sub-adult  the  Qualitative  behaviour  peck-dominance  i s related  more  socially  for  males  behaviour are  development  other  H.).  sexes were  Island,  and  structure  were  recorded. A  and  The  mainly  classes  the  Mclnnes  128<>43»  both  patterns  activity.  reproductive  are of  s i x age-sex  interactions.  also  They  involved  at  B. , L o n g .  sea-lions  behaviour  and  located  not  Contact  to  many  involved  than becomes  i t  behaviour.  of  the  than i s  more  aggressive.  hierarchy  Males  behaviours  females.  than  the  measured.  by  aggressive  males  of  mature  lack  towards  are  voluntary  females.  their  classes  Males  i s more  As  that  i s reflected are  between  This  f o r females.  complex  This  exists  their  Females of  Body  other  males  and  use  more  Contact As  behaviour  males  behaviour social  Body  mature which  complex  adept  adults at  Sub-adult males  Territories activity This  act  as  Non-pupping colonies  classes.  organized Is.  than  infes!. i 3  inhibited  by  s  that  are  pupping are  females.  Body  aggressive  and  interaction.  Non-body  displays.  males  better  not  Contact  The  results  able  to  intense in  cope  territorial.  i s related  to  i i i  more  with  the  However,  some  colonies.  refuges  to  the  f o r females by  the  presence who  sexually  mature  grouping  of  of  avoid  females. areas  of  sub-adult  males.  females  around  males. colonies except  are  f o r the  Socially, pupping  promoted the  threats  than  with  sub-adult  harassment  adult  physical  replaced  of  This  contributes  pupping  i s  males  are.  and  territorial  sex  i t  i s largely  society  behaviour  i s mainly  conditioning  socially  adult  Contact  I  by  spatially relative  non-pupping  colonies. the  disruptive  tranquil behaviour  organized proportion colonies  Social  to  of  age-  are  less  organization  behaviour of  similar  of  adults  sub-adult  males.  in and  iv  TABLE  OF  CONTENTS  Abstract  i  List  Of T a b l e s  List  Of F i g u r e s ,  i v  1  viii  Introduction  1  Materials  2  and methods  Results  16 Territorial  Behaviour  Reproductive  17  Behaviour  19  Hierarchy  20  Duration  24  Initiation  26  Variety  Of B e h a v i o u r  Beginning  and T e r m i n a t i o n  Investigative Non-Body  Eehaviour  Behaviour  and Body  Population Class  Patterns  Contact  Structure  Association  27 29 39  Behaviour  41 47 49  Activity  50  Transiency  52  Interaction  Rate  53  Interaction  Frequency  57  Discussion  59  Summary  76  Acknowledgements  79  Literature  80  Cited  V  LIST  OF  TABLES  Table 1.  2.  3.  4.  5.  6.  7.  8.  9.  Page, Determination of hierarchy using the retreat,  21  Determination df hierarchy using i n i t i a t i o n of interactions.  23  Variety of behaviour patterns used i n i n t r a - c l a s s b e h a v i o u r .  28  Use o f t h e Advance classes.  by a l l 30  Use o f t h e Advance other males.  by m a l e s t o  Use o f t h e Advance L a r g e Cows.  by m a l e s t o  Use o f t h e Advance Yearlings.  by  Use o f t h e Advance S m a l l Cows.  by  31  32 males to 32 males t o 33  The r e l a t i o n s h i p o f t h e Advance t o h i e r a r c h y .  33  Use o f t h e A t t a c k by m a l e s , f e m a l e s and Y e a r l i n g s .  35  Use o f t h e A t t a c k by m a l e s t o m a l e s . Y e a r l i n g s and f e m a l e s .  35  12.  Use o f t h e A t t a c k  36  13.  Use o f t h e R e t r e a t by m a l e s , f e m a l e s and Y e a r l i n g s i n i n t r a - c l a s s behaviour.  36  Relationship of the Retreat to hierarchy.  37  10.  11.  14.  by m a l e s .  vi  15-  16.  17.  18.  19.  20.  21.  22.  23.  24.  25.  26.  27.  28.  Use o f t h e C h a s e by m a l e s , f e m a l e s , and Y e a r l i n g s . Use o f t h e C h a s e L a r g e Cows.  by  Use o f t h e C h a s e S m a l l Cows.  by  males  37 to 39  males  to 39  Use o f I n v e s t i g a t i v e behaviour by m a l e s , f e m a l e s and Yearlings in intra-class behaviour.  40  U s e o f I n v e s t i g a t i v e by to males, females and Yearlings.  41  males  Use o f I n v e s t i g a t i v e behaviour by f e m a l e s a n d Y e a r l i n g s to m a l e s , f e m a l e s and Y e a r l i n g s .  41  Use o f Non-body and Body C o n t a c t b e h a v i o u r by a l l classes in intra-class behaviour.  43  Use o f Non-body and Body C o n t a c t b e h a v i o u r by m a l e s other classes.  to  Use o f Non-body and Body C o n t a c t b e h a v i o u r by females to o t h e r c l a s s e s .  43  -  45  Use o f Non-body and Body C o n t a c t b e h a v i o u r by Yearlings to other c l a s s e s .  45  Use and  46  o f m o u t h by m a l e s , females Y e a r l i n g s to a l l c l a s s e s .  U s e o f M o u t h by to a l l classes.  male  classes 46  Use o f M o u t h i n g by m a l e s females to a l l classes.  and  Use all  to  of Mouthing classes.  by  males  47  47  vii  29.  30.  31.  32.  33.  34.  Age-sex pupping  structure colony.  Interaction rate i n t e r a c t i o n s per  of  a  48 given as minute.  Time each c l a s s spends i n t e r a c t i n g with other lions. A c t i v e time interacting lions.  each with  non-  class other  sea56 spends sea57  F r e q u e n c y o f male, f e m a l e , Yearling interactions Interaction frequency Cows w i t h m a l e s .  54  of  and 59 Large 59  viii  LIST  OF  FIGURES  Figure  Page  1.  Peck-dominance classes  2.  Mean  hierarchy  interaction duration  of 24 (in  seconds).  25  Percent of i n t e r a c t i o n s i n i t i a t e d by t h e c l a s s on t h e o r d i n a t e when i n t e r a c t i n g w i t h the c l a s s e s on t h e a b s c i s s a .  27  4.  Association.  50  5.  Activity.  52  6.  Transiency.  53  3.  1  INTRODUCTION Euffletqpias Aleutian south and are  jubata  Islands,  colonies  usually Ii  they  harassment occurs  during  will  excluded  be  animals  1966)  which  males  and  females.  There  have  distribution Belkin  in  (1966).  account  of  and young.  male  the  and e a r l y  occupy  these  social  t  holder.  i n this to  to  pupping  E.. j u b a t a .  organization  (1970)  Gentry  o f a pupping  describes  (1970) and  colony.  groups  and  maintain These  (Belkin,  1966; adult  behaviour i s  described  give  a  of a  pupping  of The  a n d a few  I t s  more  and and by  detailed colony  the behaviour  emphasizes  female  without  on t h e b e h a v i o u r  Islands (1966)  Only  colonies.  colonies  studies  which  colonies.  males  the  summer.  These  pupping  of sub-adult  during  of the breeding  establish  within  colonies  areas  Most  way.  as  as f a r  territories  spring  Kurile  territoriality  organization  establish  several of  Sandegren  certain  the  gregarious  The  colonies  O r r and P o u l t e r  the  California. females  been  organization  o u t on l a n d .  non-pupping  consist  are  ftt  unable  form  America  islands.  referred males  of North  through  and exposed  are not tolerated  Pike,  social  organized  and a d u l t  territories  haul  the t e r r i t o r y  i n colonies  immature  can  Japan,  of the species  males  late  yearlings from  sea-lions  on r e m o t e  the adult  maintain and  they  i s polygynous.  season  females  the coast  Members when  located  jubata  breeding  of  a n d down  as C a l i f o r n i a . form  i s d i s t r i b u t e d from  of  in the  the s i g n i f i c a n c e  distribution  in  the  2  The  aggregations  have  only  study  i s to describe  a  been  non-pupping  with  a  more  The  of  shows  the  get  older.  of  Small  Cows  i s followed  to Old Bulls.  social  of  occuring  behaviour Female and  from  Trends  sexually  mature,  processes  a r e r e l a t e d t o and a f f e c t behaviour  and  become  and  of social  to  more  large  Bulls  as  the  occur  along  sea-lions  Male  and  patterns  organization.  then  ontogeny with  the  get larger,  experienced.  and t h e i r  as  from  Cows.  the behaviours  These  distinct  i s followed  t o Young  The  as  between  progressively  behaviour  of things  of the classes behaviour.  as w e l l  are discussed  behaviour.  social  the discussion  understanding  E. j u b a t a ,  changes  Yearlings  of  for  of  of  colony  i n interactions  then  ontogeny  of s o c i a l  this  organization  a better  similarities  A number  patterns  of  non-pupping  organization  behaviour.  The  a  yearlings  objective  and s o c i a l  facilitate  behaviour  of this  through  and  polygyny.  Much  behaviour  The  may  differences.  to  anecdotally.  of the s o c i a l  o f JB. j u b a t a  Yearlings  males  Comparison  colony  social  animals  non-breeding  the behaviour  colony.  generally  classes  mentioned  pupping  specifically  of  These  measured.  development provide  a  give basis  3  MATERIALS jubiiii! terminology The  will  Fisheries  British  The 150 the  males  summer  (Lat.  and f e m a l e s  were  blinds  A  started  During  made  from  this  time,  determined. devised  were  consisting  120  of the year  and  most o f  remain. neighbouring  these  heard,  using  (20 t o 60  the long  the  the  from t h e  blinds  distance  sources.  of general  Vocal  until  observations  May  27,  c l a s s e s a n d 34 b e h a v i o u r  May  The  reason.  1972 a n d l a s t e d  From  between  t o 250 m e t e r s  locate  for this  6 age-sex  a n d more  easily  Island,  Yearlings.  and a s p o t t i n g scope  readily  study  o n Hay 9,  and  of many  128°43'W.).  islands  were  was n e g l e c t e d  preliminary  periods  50  sounds  containing  on  portion of the colony.  from  20x60)  Department  males  were  and  to  on  was b a s e d  Mclnnes  Long.  a few s e a - l i o n s  erected  Observations  behaviour  near  minor  certain  blinds  i tdifficult  colony  i s i n an a r e a  a  This  sections.  i n 1971 a n d 1972 w e r e  Three  Although  "sea-lion".  Canada  immature  form  During  (8x30  and  located  of  and o n l y  The  as  52«16»N.,  leaves  power).  were  and  colony  binoculars  were  The c o l o n y  populations  to  non-pupping  surveys  mainly  sea-lions.  colony.  was  a  islands  Columbia  colony.  made  of  records.  METHODS  i n the remaining  aerial  consists  Mature  referred  be u s e d  from  exposed  colony  be  selection  information  small  will  AND  28, to J u l y  9,  structured observations  1972, were  1972.  patterns techniques taken.  4  Classes The classes. The  A  brief  i n  curves  the  compiled  the  colony  description  age o f t h e a n i m a l s  comparing  1.  sea-lions  sizes  by P i k e  Old Bulls  o f each  comprising  relative  are  divided  class  each  class  into  i s given  six  below.  i s estimated  of the classes  by  to the growth  (1966). -  much  larger  than  a l l  ether  very  stout  classes. 6 -  years  large  and  older.  forehead  and  neck. thick long the 2.  Intermediate  Bulls  -  lower  jaw.  mane. testes  are obvious.  larger  than  smaller  than  a  typical  the to  size  Young  Bulls  Old Bulls.  Intermediate  Bull  o f an O l d B u l l  2'times  but  the  size  and  of  i s 2/3 1  1/2  a  Young  and  stout  Bull. 4  t o 6 years o l d .  accentuated  forehead  neck. the  lower  massive  as that  noticeable testes  jaw i s n o t a s  thick  o f an O l d  or  Bull,  mane.  a r e not as obvious  a s c n an  5  Old 3.  Young  Bulls  Bull.  1/2  the  size  of  Intermediate  Bulls. 2  to 4 years o l d .  some  indication  o f an  accentuated  forehead. neck  i s not  testes mane  are not r e a d i l y  i s short  lower  4.  Large  Cows  i s  visible  the  size  although  some  slightly  larger.  4 t o 5 years slim  neck.  slim  lower  teats  Cows  1/2  and  of  Young  Large  Bulls  Cows  are  older.  jaw.  noticeable, a  the size  slim  neck.  slim  lower  mane.  especially  young. of Large  2 to 4 years o l d .  no  the animal  profile.  suckling Small  when  mane.  smooth  5.  thick.  moving.  about  no  seen.  or non-existant.  jaw i s n o t  penis is  stout.  jaw.  Cows.  i f  6  6.  Yearlings  -  smooth  profile.  1/4  1/3  or  to  Large  1 year no  the  size  of  Young  Bulls  Cows.  old.  obvious  second  sex  charac-  teristics. The Young  above  Bull  Sea-lions too  The  class  that  small to  Bulls.  classes  difficult  and  to  determined,  i s  Unclassified  Small.  a  i s  If  is  Occasionally In  to  are  Bull  classed  known,  large  applied  the  groups.  Intermediate  Bulls  Intermediate  i t  distinct  the  too  also  separate.  then  sea-lion  into  ordinarily  scheme  Bull  form  Intermediate  This  Young  not  grades  are  be  do  be  Bull  Young  the  classes a  are  of  as  Unclassified  neither  these  as  other  sex  sex  but  Young  classes. the  sea-lion  the  class.  Bulls  classed  to  The  most  cannot  be  Large  or  nor  size  of  instances i t i s recorded  as  Unclassified. Old  Bulls  Yearlings which  of  and  males  weights  the  There  sample  collected.  found  epididymides  were  no  and  are  individual of  and  males  males  sexually  not  mature  sperm  This  Bulls  are  Cows a r e  s i x years  1962).  (1971)  Cows  sexually  same  aged  Intermmediate  Large  Small  become  are  males  and  is  older  meant capable  than  spermatozoa years  least  i n smears older.  at  Testes  epididymides  s i x years  producing  and  age  known.  i n the  at  female  The  ( T h o r s t e i n s o n and  that of  1966).  not  i s present  younger  five  (Pike,  mature,  Lensik,  eld in the  sperm. taken He  older Perlov  from also  the  the found  7  that  i n four  While  the  coupled  year  with  Bulls The  the  are of  morphological  two  do  months  (Farentinos, and  female  sexual  probably  sexually  immature.  Y e a r l i n g s i s not  intuitively  seem  of  age,  of  On  used  these  as  Behaviour  Patterns  divided  The  of  34  into  Miscellaneous.  the  be from  behaviour  individual at  the  behaviour  similarly  same.  which  male  least  of  male  Yearling and  female  determined.  These  and  Body  Termination, Contact,  patterns  were  and  observed  Copulation.  Beginning  And  Advance  Termination -  A  sea-lion  seems  as  intent. medium 2.  of  and  obvious  behave  Beginning  Contact,  by  Until  the  that  Yearlings  sexes  p a t t e r n s were  groups;  Non-body A l l  to  base  behaviour five  Investigative,  a  Male  different.  criteria,  this,  suggests  behaviours  both  absent.  definative,  distinguishable  not  pups  were  dimorphism, mature.  1971).  A total  i s not  sexually  originates.  1.  spermatozoa  characteristics.  i s  spermatozoa  are  behaviour  except  the  Y e a r l i n g s i s assumed  behaviour  are  of  obvious  Bulls  sex  Yearlings  males,  presence  Intermediate Young  old  Attack  -  This  approaches  another.  i f i t i s approaching It  occurs  at  a  It with  slew  or  shuffle. i s  similar  to  the  Advance  8  except 3.  Chase  -  A  that  fast fietreat  -  very  quickly.  chases  or  follows  sea-lion  another.  4.  i t occurs  This  run  or  A sea-lion This  slow  to  to  at  either  medium  moves away  appears  other's  occurs  be  a  a  shuffle.  from  another.  result  of  the  behaviour.  Investigative 5.  Noseing  A  -  A sea-lion the  head,  another 6.  Noseing  B  -  Noseing  C  -  Licking  A  -  Licking  Non-Body 10.  11.  B  -  i t s nose  puts  or  near  back  of  or  near  another.  i t s nose where  cn  cn  or  near  another  has  past.  A sea-lion  A  or  region of  ground  head, 9.  puts  A sea-lion  recently 8.  body  on  sea-lion.  peri-anal  the  i t s nose  anterior  A sea-lion the  7.  puts  licks  anterior  sea-lion  another  body,  licks  peri-anal  region.  The  mouth  of  and  directed  or  cn  the  back.  another  in  the  Contact -  Mouth  Snobbing  A  -  A  sea-lion  45  degree  a  sea-lion  towards sits  with  angle.  is  opened  another. i t s head The  eyes  on  a  are  9  sometimes 12.  Snobbing  B  A  sea-lion  90  Turn  Away  Nodding  A  angle.  head  15.  Houthing  The  on a  eyes  are  back  from  nods  i t s This  then  are  together.  be g u i t e  slow  interacting  open  and  directed  There  i s shaking  and moving  movements  close  are  sometimes  These  sea-  very  The v i b r i s s a e  a  action.  and  head.  turns  head,usually  may  or relatively  m o u t h s o f two  lions  and  away.  sea-lion  violent  Contact  The  sea-lion  successively.  Body  i t s head  moves s l i g h t l y  other  its  with  closed.  A sea-lion the  14.  sits  degree  often 13.  closed,  erect  forward.  may  of be  the quite  rapid. 16.  Necking A  A  sea-lion  stroke 17.  18.  Necking B  Necking  C  A  Necking D  the neck  sea-lion  i t s neck.  A  sea-lion  A  with  neck  pushes  i t sneck  with  to  another  vigorously  sea-lion  another  i t s own  of another.  lightly  with  another 19.  uses  and  pushes shoulder.  vigorously i t s  pushes neck  and  10  frequently  tries  opponents  bite  the  fore-flippers.  20.  Biting  A  - A sea-lion  21.  Biting  B  -  A  to  lightly  sea-lion  another.  bites  another.  vigorously  This  bites  appears  quite  vicious. 22.,  Lunging  - a  sea-lion,  lunges snap and  with  i t s mouth  at another.  shut.  The  jaws  The v i b r i s s a e  extended  open,  are  may erect  forward.  Miscellaneous 23.  Down  -  A  sea-lion  Its  body  leisurely  does  n o t have  appearance. seem 24.  Ground  -  The  A sea-lion  lies  mostly  The  neck  -  the  ground.  Chinning  -  does not  sea-lion  the  and  body  touching  appears  The c h i n  may  seems  t o be  touch  slightly.  the The  alert. i s walking  a t a slow  or  rate.  sea-lion  chin,  with  The body  animal  A  stretched  i n the f o r e f U p p e r s .  o r be r a i s e d  medium 26.  down  ground  A  animal  i s extended  stretched.  Moving  a  down.  alert.  weight  25.  lies  throat,  succesively and neck  of  nips  the  another.  11  27.  -  Backing  During goes  an to  animal and 28.  Behind  -  the and  neck  During  interaction  tries,  on  an  Scratching  -  A  of  sea-lion  30.  Looking  -  A sea-lion  31.  Suckling  -  A  back.  one  behind  scratches  sea-lion another.  itself.  i s looking  sea-lion  ether  t o l a y i t s head  interaction goes  sea-lion  the  the other's  deliberately 29.  side  a  around.  sucks  the  teat  of  another. 32.  attempted  Mount  -  A  sea-lion  but  falls  tries  to  o f f almost  33.  Mount  -  A sea-lion  mounts  34.  Copulation  -  A sea-lion  copulates  Types  o f Data After  the preliminary  collect  the  i n f o r m a t i o n they  1.  Encounter  using  sea-lion  data  i s  approximately  immediately.  another. with  o b s e r v a t i o n s , methods  pertinent  another.  yielded  to the study.  The  were  devised  techniques  are described i n this  and  section.  Method  behaviours  what  another  Collected  to  The  mount  will  of the be  selected 30  various  referred from  minutes.  classes  are  to as the Encounter the  colony  and  The  following  determined Method.  watched informatiicn  A  for i s  12  recorded: 1.  Class  of  2.  Classes  3.  Duration  4.  Which  sea-lion initiated  5.  Types  of  of  actions The This  Encounter  i s outlined  the  sea-lion  the  of  sea-lions  observed.  i t interacted  with.  the i n t e r a c t i o n s .  behaviour and  being  their  Method  the  patterns sequence  gives  a  interaction. used  of  in  the  inter-  use.  v a r i e t y of  information.  below.  Hierarchy A which  class  hierarchy  suggest  can  be  determined  by  using  criteria  dominant-subordinant r e l a t i o n s h i p s .  Duration The This  mean  allows  classes  comparisons  tend  indicative exchanged be  used  duration  to  of  interacting  the  between  the  The  length  length  or  kind  1972).  The  duration  other  proportion  of  of  amount  c a l c u l a t e the with  i n t e r a c t i o n s can  interact.  (Caldwell, to  of  cf  of  be  determined.  the  an  time  interaction  information of  the  being  interactions  time  a  class  is  can  spends  classes.  Initiation The  classes  interactions determine  are  of  the  recorded.  class hierarchy.  sea-lions This  It  may  i s used also  which as  a  indicate  initiated criterion i f  to  classes  13  have  preferences with  Intra  And The  of  times  Class  types  of behaviour  they  were  the  inter-class one  and  patterns  are  used  The  data  ontogeny  of  and  are  class  behaviours  The  help  This  number  information another.  are considered  way  behaviour.  The  for a l l  Miscellaneous i s because  many  that  The  best  types  shows  and  the  relative  behaviour  patterns  between  the  classes.  These  describe  the  colony's  social  existing will  the  significance.  in a  o f use o f c e r t a i n  This  patterns.  a r e c f minor  and  towards  are not discussed.  social  relationships  recorded.  o f one  are presented  relationships  p a t t e r n s employed  behaviour  behaviour seldom  used  intra-class of  interact.  Behaviour  behaviour  group  frequency  they  Inter  describes  but  whom  indicate  the  organization.  Interaction  Rate  Sea-lions number  of  the using  time  particular  i s  they  a  a class  class  of the active spent  the Looking  class had  calculated  the Duration  o f time  that  remainder  rate  with  proportion time  a  interactions  interaction combined  of  i s  data  f o r each  are  used  displaying  a t no  used  i s as  class.  data  calculate  the  proportion of calculated. an  An  These  estimate  specific social  pattern.  and t h e  recorded.  to  The  interacting c a n be  observed  were  i s active.  time  behaviour  were  this The of  partner or  11  Interaction The which with to  Frequency  number  the observed which  the  classes  expected  association  of  information  2.  series  Encounter  class  classes  (page  selection  Distribution  These  transiency  of the classes.  repeatedly and  other  searched following  When  determine  were  f o r these  features  was  1.  Date  2.  Individuals  and  the  data  give  will  be r e f e r r e d  to as  technique  was  The  were could  file  were  activity,  selected be  was u s e d  recorded.  approximately  with t h e  from  and each  easily  and  i n which  scars  The c o l o n y  every  hour.  was The  time. looked  f o r and t h o s e  I further  3.  Its location.  4.  The  classes  of  greater  were  considered.  not  situation  found.  recorded;  i t s five  Neighbours  this  cn  recorded;  was f o u n d  did  based  concurrently  association,  that  A card  sea-lions  information  made  Sea-lions  animals  identifying  compared  partners.  Method.  the  recognized.  a sea-lion  were  Distribution  to  These  of social  frequency  i s  Frequency  observations  designed  This  interaction 49).  The  with  Method  of observations  Individual  class.  of  of sea-lions  a r e known.  i s calculated.  frequency  Method.  and c l a s s e s  interacted  interact  on c l a s s  Individual A  the  of interactions  than  3  nearest body  In only  arise.  neighbours.  lengths a few  away  instances  15  5. These  Which  data  of the sea-lions  result  were  i n information  active.  as o u t l i n e d  below.  Association The  association  totalling  the  o f one c l a s s  number  individuals  being  proportion  of  determined  whether  of  that  the  another  neighbours  observed  these  to  recorded  and  then  belong  to  classes  were  i s  found near  the  calculating each  the  class.  clumped  or  by  This  randomly  distributed.  Activity The by  proportion  dividing  the  of time  t h e number  a class  of active  total  number  of neighbours  considered  active  i f when  i s active  neighbours i n that  observed,  i s calculated  in a  class.  class  with  A sea-lion  i t had i t s head  i s  raised.  Transiency Transiency the  colony.  areas and  were e i t h e r cracks.  calculated  ,  The i s l a n d  observations  moved  i s a measure o f a s e a - l i o n s  by  was  natural  platforms  If a sea-lion this  was  dividing  by t h e n u m b e r  divided  moved  counted  of times  areas.  These  20  different  movement.  of times  i t was  within  or bordered  to a  a s one  t h e number  into  movements  a  sighted.  by r o c k area  faces  between  Transiency i s sea-lion  class  16  3.  Other Descriptive  reproductive the  colony  behaviour was  Presentation The figures  notes  also  and  were  abbreviations  -  Old  IB  -  Intermediate  YB  - Young  a r e used  in  The  Cows  LC  -  Cows  Large  are presented  classes  involve  class  interactions  section  on  status  different  format  i n age f r o m  female  a r e shown  easily.  i s used.  The  more  one  t o age  o f two  are ordered For  classes  Yearlings.  behaviour  according  tables  are net involved.  the c l a s s e s  most  i n contingency  are ordered  interactions  hierarchy,  Bulls  Bulls  - Small  class  relative  Bulls  SC  where  and  of  Yearlings  data  male  structure  f o r the classes  OB  of the data  increasing  age-sex  tables:  histograms. in  The  and  Data  YY -  Most  taken.  territorial  recorded.  of the  following  .concerning  are  grouped  apparent.  that  In the  show  remaining  makes  sex  Data  the  This  and  ways.  to  and  their data  by  a  sex,  differences  in  17  RESULTS  Territorial  Behaviour  Territorial by  Gentry  (1970)  classes. this are  behaviour recognized. of  Bull  the  #2  was  Old Bull  Bull  contact  not  #1  with  occasions, filter  Bulls  the  Old Bull  Bulls  occasionally  territories. the  #2  colony  occupied  a territory  island.  exhibit  boundaries  more t e r r i t o r i a l l y  Old Bull  area  a n d no  male  Old  inclined  an a r e a  there.  Bull  #1  When  Old Bull  usually  without  were  on  When O l d eventually  #2 r e t u r n e d . any  physical  loosely  are  usually  lies  down  maintained. excluded. and  Young However  the  and  on  sub-adult  many males  and Y e a r l i n g s .  The O l d B u l l i s  presence.  Eventually the  Old  to their  up,  nods  h i s head,  The  other  vocalizes  quickly  leave  and approaches the territory.  enters  the t e r r i t o r y  Females  and Y e a r l i n g s a r e not a g g r e s s i v e l y  territory  the  the females  oblivious  males  between  #2.  are  among  was  #2's t e r r i t o r y .  the Old Bull  i n  defend  and m a i n t a i n e d  left  described  frequenting  #2  the  Bulls  to those  i s defended  Old Bulls  Originally  Territories Intermediate  and Young  Bull  on  similar  i n the interactions  Old  #1.  of the island  acguired Old  Some  Old  Old Bull  displays  b u t no t e r r i t o r y  territorial.  top  occur  Intermediate  Two  than  boundary  by t h e O l d  and s u b s e q u e n t l y  Bull.  lies  Bull  the other Any down  gets males.  male also  chased  that  leaves.  from  the  18  Females island  where  females end  were Old  were  where  seen  Bull  seen  Old  #2  in a  Bull  #2,  established  occasionally  occupied  he  did  groups Old  not of  although  #1 he  island's and as  his  Bull. The He  nose  persisted.  behaviour  was  now  Intermediate then  that  territorial Large  The  Cow  lay  down  either  her  the  not  Intermediate  again.  so  and  Bull  peri-anal  region.  Cow  being  moved to  15  10 feet  the  did  not  feet  Bull  #1  areas,  joined  these  the  times  around  him  any  but  males  are.  Yearlings.  For  down  the  near  approached She  her  moved  away  to  closer  the  the  to Old  further.  female.  Old  feet. It  recognized the  this  Bull.  for several  Bull pursue  v  maintained  subjected  from  female  approach  but  best  lying  after  previous  above  from  harassed,  Intermediate and  the  was  Later  them.  f o r females  followed  did  the  males  the  described.  During  with  of  north-east  Old  elsewhere,  subadult  Cow  minutes 10  of  Bulls.  top  his  previously  occupying  An  Large  Bull  boundary  as  interacted  within  l a y down a n d  possible  on  the  Large  near  for several  She  areas  aggressively  a  island's  territory.  Yearlings  25,  the  the  territory.  new  create a refuge  June  a  on  abandoning  Intermediate  not  on  territory  chase  study  after  not  a  n o r t h - e a s t end.  put he  and  to  on  was  occasionally  are  tends  example  and  d i d not  Females territory,  #1  the  established  a  these  establish  Young  Bull  This  Bull  had  in  small area  territory,  When O l d  early  is the The  19  A Bull  similar  instance occured  attempted  territory and  went  did  not  of  Old  closer  No  pups  copulations  In mount  were  Bull  lay  her  Unclassified  Old  Bull.  an  The  Intermediate  Small  near  the  Small  moved  away  Intermediate  Bull  away.  The  although  did  his  1972.  However  On  June  not a  Small  was  After  a  A  (1970)  16,  an  Cow.  some  few Bull  similar  also  the  then but  on  as  June on  described  of  Small  16,  top  the Cow  mounted less  Bull  Intermediate  was  neck  to  Unclassified  The  movements  minutes  attempted  The  He and  displays  Intermediate  Small.  i t .  No  occured.  or  territory. pursue  common.  precopulatory  mounted  foreflippers  There  Small.  in  not  Sandegren  Bull's  Intermediate  Unclassified  was  Unclassified  mounted  (1970).  region.  behaviour  Old  with  Island  Yearlings.  an  and  Bull  pelvic  Mclnnes  colony.  mount  down  at  observed,  the  down  Sandegren  The  d e s c r i b e d by  into  Intermediate  An  Unclassified  instances sea-lions  on  to  moved  born  sexual  several  Small  the  suckled  those  attempted  #2.  20.  follow i t .  were  others  held  to  females  to  Bull  an  June  Behaviour  Involved  similar  mount  towards  attempt  Reproductive  several  to  on  an and by  male's sguirmed  a  nearby  involved  sequence  occured. On  June  20,  Unclassified squirmed  away.  an  Intermediate  Small. The  The  Bull  attempted  Unclassified  Intermediate  Bull  Small  approached  to  mount  an  immediately i t  again.  20  The  Unclassified  The  Intermediate  to  mount  Small Bull  another  moved  into  Old Bull  d i d not attempt  Unclassified  to follow  Small  Small  approached  and  the  two  Unclassified  Small  was  forgotten.  The  Bull,  his  extended,  then  Young  Bull  penis  lying  nearby  A  territory.  i t but  i n t h e same  Unclassified  with  resisted.  #2's  tried  area.  The  second  Intermediate  Bull  began  interacting.  The  males  b u t t h e Young  first  Intermediate  attempted  Bull  moved  t o mount  a  away.  Hierarchy The  Retreat  relative use  status  the Retreat  elephant lions  seal  should  recieve  behaviour  of the classes.  (Hirgunga  1).  During  uses  the Retreat;  they  More  in  than  o f two  the  the  often  dominant classes,  other,  to  the  (1968)  northern  Subordinant sea-  p a t t e r n more with  index  and P e t e r s o n  dominance  interact  the Interactions  1.  an  an^ustirostris).  the Retreat  when  i s  LeEoeuf  f o r determining  give  i t  pattern  then  than  animals i f  they (Table  one  class  i t i s considered  subordinant. 2.  Less  than  the  other,  then  i t  i s  considered  dominant. 3.  Equal  to  the  other,  then  i t i s considered a  peer. A second hierarchy. with  criterion Sea-lions  subordinants  more  can should than  be  used  tend they  to  determine  tc initiate do  with  class  interactions  dominants.  In  21  OB  IB  LC  2/23  IB LC YB SC  Table  YB  SC  YY  0/35  0/8  0/1  0/14  3/6  5/39  0/11  3/35  0/1  0/0  0/2  0/0  0/12 0/2  1.  Determination of hierarchy using the r e t r e a t . The r a t i o i s the number o f i n t e r a c t i o n s where t h e R e t r e a t was u s e d b y t h e v e r t i c a l l y l i s t e d c l a s s e s o v e r t h a t u s e d by the h o r i z o n t a l l y l i s t e d classes.  22  Table  2,  i f one 1.  class  Wore  initiated;  interactions  considered 2.  Fewer  Old Bulls  An  interactions  Bulls  and  Large  Intermediate  not a  are  number,  only  Cows  Bulls.  as  well  result  Cows  are  Small  Cows.  status.  i t  i s  Small  From  Cows,  Young  Part  Small  Bulls  Cows  hierarchy i s represented  and  Yearlings.  Large  available  Small  Bulls  to  and  peer.  Bulls, Old  classes.  the  to  and  to  a  Intermediate  Young  are dominant  the other  Bulls  class  i ti s  then  i t i s considered  subordinant  dominant  Young  another,  to a l lc l a s s e s .  Bulls,  samples.  slightly  The  then  A l lclasses  d e l i n e a t e d as  of small  than  to Old B u l l s .  are  h i e r a r c h y o f Young  then  subordinant.  dominant  are subordinant  Cows,  The  equal  another,  dominant.  considered 3.  than  Cows a r e  cf this  data. and  are  Large  peers of  i s  with  similar  i n Figure 1 .  Duration The female  male and  classes  Yearling  classes  involving  males  of  involving  other  male  interact they  do  periods with  with with with  peers.  Intermediate  interact  the  peers.  class  classes  or  animals  Intermediate  subordinant Young  (Figure  same  subordinant  Bulls  Bulls.  and  show  That  f o r longer  a  periods  2).  than  Interactions  are longer  Yearlings.  than  those  Old  Bulls  for shorter intervals Bulls  do  than  interact  for  shorter  dominant c l a s s e s  than  they  trend  similar  i s , the duration  to  Old  decreases  do and  as the  23  OB  IB  LC  YB  SC  YY  25/6  38/5  12/2  3/2  17/0  16/4  80/20  32/1  71/6  4/0  11/3  21/11  IB LC  1/0  YB SC  Table  40/5 6/0  2.  Determination of hierarchy using i n i t i a t i o n o f i n t e r a c t i o n s . The ratio i s t h e number o f i n t e r a c tions initiated by t h e v e r t i c a l l y l i s t e d c l a s s e s over those initiated by t h e h o r i z o n t a l y listed classes.  24  l  OB  IB  dominant  Fig.  1.  LC  YY SC  YY  subordinant  Peck-dominance of c l a s s e s  hierarchy  25  100 OB  H= 6 6 . 6 p< .05 df= 5 n~=  10  26  19  36  46  100 IB rti  0  0=  "D C  167  101  84  37  23  100  o «  27  H= 3 4 . 3 p< .05 df= 5  YB  C  H= 8 9 . 1 p< .05 df= 5 n=  o  12  100  192  47  100 3 Q  YY  H= 17.6 p< .05 df= 5  c o n=  21  82  46  2  28  7  100 SC  H= 13.0 p< .05 df= 5  0  3 7  n=  20  13  100 LC  H= 3 0 . 9 p< .05 df= 5  I n=  Fig.  2.  46  23  4  28  12  8  OB  IB  YB  YY  SC  LC  Mean i n t e r a c t i o n d u r a t i o n ( i n s e c o n d s ) . The i n t e r a c t i o n d u r a t i o n i s g i v e n f o r t h e c l a s s on t h e o r d i n a t e when i n t e r a c t i n g w i t h t h e c l a s s o n the abscissa.  26  difference  i n the  status  between  the  interacting  classes  increases. The show  Large  the trend  Bulls  Large  short  Bulls  long  periods  interact  Intermediate  Cow,  above.  for fairly  for  Cows  Small  mentioned  interact  interact  Cow,  for  than  and Y e a r l i n g Old Bulls  periods with  longer  they  do  with  and  with  Small  classes  Intermediate  Large  Cows  do n o t  Cows.  and  They  Yearlings.  periods  with  Old  other  classes.  and  Initiation The class  proportion  i s given  behaviour  with with and  i n Figure  status more  Large  Cows.  variety  use  each  Patterns  observed  interacting  a r e used  i s related  classes.  with  this to the  Sea-lions  subordinants  do n o t i n i t i a t e  initiate  using  with  any  each  than  interactions  behaviour  u p w a r d s o f 15%  of Old  of  their  Patterns variety  (Table 3).  females.  This  i s defined  by  Cows.  a greater  considered. other.  who  initiated  of the i n i t i a t o r  interactions  Large  and Y e a r l i n g s than  Hierarchy  the  Bulls  with  interactions  T h i s c o n t r a s t s with  Of B e h a v i o u r  Males females  3.  Young B u l l s  Intermediate  interactions  of  of their  dominants.  Variety  the  and so the c l a s s  relative initiate  of  removes used  Only  of behaviour Yearlings  intra-class  any b i a s caused more  than  patterns  use  a  than  greater  behaviours  are  by c l a s s e s  mimicing  1% o f t h e t o t a l  patterns  i n the construction of Table  3.  27  100 X =  23.0 05 p< . 5 df=  OB  2  5  4 3  X2 =  IB  P< df=  65. 4 05 4  YB  c CD  u  a.  YY  SC  LC  OB  Fig.  3.  IB  YB  YY  SC  LC  P e r c e n t o f i n t e r a c t i o n s i n i t i a t e d by t h e c l a s s t h e o r d i n a t e when i n t e r a c t i n g w i t h t h e c l a s s e s the a b s c i s s a . & The sample s i z e i s too small s o t h e d a t a a r e not included.  on on  28  OB  IB  YB  YY  SC  LC  B e g i n n i n g and Termination Investigative  3  4  2  2  0  0  0  0  1  1  1  1  Non-body  5  4  3  3  3  2  5  7  6  0  1  1  2  4  4  4  2  2  15  19  16  10  7  6  Body  Contact  Contact  Miscellaneous  Total used  Table  patterns  3.  Variety of behaviour used i n i n t r a - c l a s s behaviour. X2= 1 7 . 7 , p<.05, d f =  patterns  5  29  Males  and  Termination,  Beginning  females  Non-body  differ  Contact  and  Termination  use  of  mostly  and  Body  in  Beginning  Contact  and  behaviour.  Behaviour  Advance The are  compared  females  and  (Table  5).  Old  Bulls  Bulls. Bulls  (Table  Old  and  The  freguency  Cows  only  slightly  Use  than  use  the  of  the  Advance  involved  pattern with  more a  6.  Advance  and  males Old  with  the  (Table  use  they  when  dominant.  in  this  with  most  other  most  with Young  with  Young  Intermediate  often  used  in  in  the  use  Bulls  Advance  with  use  this  Bulls.  Large of  the  Young  Cows.  this  pattern  There  behaviour  interact  with  Bulls is  no  pattern  Yearlings  8).  i s related (Table  than  frequently  seldom  Intermediate  when  more  It i s occasionally  which  in  much  Bulls.  Advance  Cows  most  with  Bulls  Young  than  classes Small  i t does  often  a l l classes  selective  and  the  Bulls.  in Table  more  or  animals  behaviour  of  difference  male 7)  Old  Advance  use as  Advance  more  Bulls,  half  with  i s given  i f ever  the  the  Intermediate  behaviour  significant  use  pattern in  the  are  Bulls  with  with  use  Males  i t about  Large  seldom  Males  Bulls  seldom  intra-class  behaviour  Intermediate  use  interactions  the  4).  Yearlings.  and  but  (Table  Advance  Intermediate  Bulls  by  the  9).  to  the A  relative  sea-lion  interacting  with  a  status uses  of  this  subordinant  30  advance no  Advance  freguency o f use  Table  4.  OB  IB  YB  YY  SC  LC  50  149  105  7  3  4  116  343  292  227  35  135  .264  .030  .024  .029  .301  . 302  Use o f t h e A d v a n c e by a l l c l a s s e s . The number o f i n t e r a c t i o n s where t h e A d v a n c e d i d and d i d n o t o c c u r i s g i v e n f o r each class. X = 130.5, p<.05, d f = 5 2  Old  B u l l s to OB  IB  YB  5  12  1  11  25  20  .312  . 324  .050  Advance no  Advance  frequency of use  X2=  6.1,  p<.05,  Intermediate  YB  1  45  62  35  151  46  .027  .229  .574  X2=  5 3 . 6 , p<.05,  Young  Advance no  Advance  frequency of use X = 2  Table  5.  Bulls  IB  Advance  frequency of use  df= 2  OB Advance no  males  Bulls  t o males  df= 2  t o males  OB  IB  YB  0  14  84  14  97  134  .0  .126  .385  3 2 . 5 , p<.05,  df= 2  Use o f t h e Advance by m a l e s t o o t h e r m a l e s . T h e number o f i n t e r a c t i o n s where t h e A d v a n c e d i d and d i d n o t o c c u r .  32  OB  IB  YB  Advance  30  12  0  no  18  11  4  Advance  .625  frequency of use  Table  6.  2  6.0, p<.05, af= 2  OB  1  Advance Advance  frequency of use  Table  7.  .0  Use o f t h e Advance by m a l e s t o L a r g e Cows. T h e number o f i n t e r a c t i o n s when t h e A d v a n c e d i d a n d did notoccur i s given.  X =  no  .521  IB  YB  18  7  36  74  42  .027  .24.3  . 166  Use o f t h e Advance by males t o Y e a r l i n q s . The number o f i n t e r a c t i o n s when t h e A d v a n c e d i d a n d did not occur i s given.  X = 2  5.64, p>.05, d f = 2  (  33  OB  IB  Advance  1  11  no  6  26  .142  .297  Advance  frequency of use  Table  8. -Use o f t h e A d v a n c e b y m a l e s t o S m a l l Cows. T h e number o f i n t e r a c t i o n s when t h e A d v a n c e d i d a n d did not occur i s given. p>.05  dominant t o subordinant  subordinant t o dominant  Advance  157  25  no  359  477  .304  .049  Advance  frequency of use  Table  9.  The r e l a t i o n s h i p o f t h e Advance t o h i e r a r c h y . T h e number o f i n t e r a c t i o n s when t h e A d v a n c e d i d and d i d n o t o c c u r i s g i v e n . X2= 1 1 2 . 2 , p<.05, d f = 1  3U  Attack The (Table  Attack 10).  It  (Table  4).  The  animal  the  male  this  behaviour  never  to  behaviour i s less use  classes  use  12).  Old  Bulls  than  and  the  Attack  Attack this  Bulls,  Young  used  employed  depends  with  different  behaviour  and  to  than the  males Advance  class  11).  to  by the  of  Males  other  occasionally  in  only  on  (Table  frequently  only  use  Intermediate  frequently  the  p a t t e r n most  females  i s  frequently  i s interacting  male  than  of  pattern  direct  males,  but  Yearlings.  The  frequencies  p a t t e r n more  Intermediate  the  Bulls  use  (Table  frequently i t  more  Bulls.  Retreat The is  Retreat  shown  in  considered by  Table  in  more  The  than  (page  Subordinants Peers  use  to  pattern in intra-class Only  reduce  relationships. females  Retreat  hierarchy  13.  order  hierarchical  pattern  behaviour  is  20)  use  i t more  a  and  bias  that  Males  behaviour  would use  i s  be  introduced  this  behaviour  Yearlings.  criterion i s  this than  and  intra-class  behaviour  related  action dominants  more but  used to  to  determine  status often  less  than  class  (Table  than  14).  dominants.  subordinants.  Chase The males when  Chase (Table  they  behaviour  p a t t e r n i s used  15).  Male  classes  interact  with  Large  use Cows  most  this  frequently  pattern  (Table  16).  by  differently Old  and  35  males Attack no  Attack  frequency of use  Table  '  73  0  982  461  .074  .0  10. O s e o f t h e A t t a c k b y m a l e s , f e m a l e s and Y e a r l i n g s . The n u m b e r o f i n t e r a c t i o n s when t h e A t t a c k d i d and d i d n o t o c c u r i s g i v e n . p<.05  males Attack no  Attack  frequency of use  Table  f e m a l e s and Yearlings  YY  females  70  3  0  687  175  120  092  017  .0  11. Ose o f males. number Attack given.  t h e A t t a c k by m a l e s t o Y e a r l i n g s and f e m a l e s . The o f i n t e r a c t i o n s when t h e d i d and d i d not o c c u r i s X = 2 2 . 8 , p<.05, d f = 2 2  OB  IB  YB  Attack  18  42  13  no  93  Attack  frequency of use  Table  . 162  390  379  .097  .033  1 2 . O s e o f t h e A t t a c k by m a l e s . T h e number o f i n t e r a c t i o n s when t h e A t t a c k d i d and d i d n o t occur i s given. X*= 2 4 . 3 , p<.05, d f = 2  males Retreat no  Retreat  frequency of use  Table  females Yearlings  86  1  430  54  .166  .018  13. U s e o f t h e R e t r e a t by m a l e s , f e m a l e s and Yearlings i n intra-class b e h a v i o u r . T h e number o f i n t e r a c t i o n s when t h e R e t r e a t d i d and d i d n o t occur i s given. p<.05  V  37  dominant to subordinant Retreat no  Retreat  frequency of use  Table  peer  13  87  189  503  398  313  .025  .179  .376  14. R e l a t i o n s h i p o f t h e R e t r e a t t o h i e r a r c h y . T h e number o f i n t e r a c t i o n s when t h e R e t r e a t d i d a n d d i d n o t o c c u r i s g i v e n . X = 2 0 3 . 0 , p<.05, d f = 2 2  males Chase no  Chase  freguency of use  Table  subordinant to dominant  f e m a l e s and Yearlings  94  4  961  457  .089  .008  15. U s e o f t h e C h a s e by m a l e s , f e m a l e s , and Y e a r l i n g s . The n u m b e r o f i n t e r a c t i o n s when the Chase d i d and d i d n o t occur i s given. X*= 3 4 . 3 , p < . 0 5 , d f = 1  38  Intermediate  Bulls  use  Bulls  use  Chase  much  male  classes  with  the  Small  do  Cows  Investigative The  avoid  as  males than  a  similar  is  used  Non-Body The  and  as  a  base,  Non-body  variety  when  not.  they  Old The  interact  behaviour females 19).  and  by  in  males  behaviour Females  (Table 20).  Investigative this  more  than  behaviour  frequently those  i s directed and  with  with  more  Yearlings  Investigative  interactions  to  show  behaviour  females  and  males.  Contact of  Behaviour  Non-body  and  used  i s shown  i n Table  the  ontogeny  of  increases.  of  Yearlings  Yearlings.  males  with  i s used  This  frequently  females  i s considered  more  use  i s  mature.  to  Body  behaviour  behaviour  intra-class  Contact  Yearlings,  Intra-class  The  variety  patterns  do  Bulls.  Investigative  males.  i t i s to  than  of  Y e a r l i n g s use  with  most  Yearlings  difference  and  (Table  females trend  Bulls  Intermediate  Females  males  interactions  other  Young  17).  18).  Investigative in  than  this  freguency  (Table  than  decreases  while  Behaviour  bias.  behaviour  more  show  (Table  occurence  calculated to  not  i t ,  sea-lion  behaviour and  3.  Young  Body  contact  If  consider Yearlings  we  behaviour  patterns are Bulls  but  as  about males  behaviour  is  apparent.  the  same  mature  for the  OB  IB  YB  Chase  28  3  0  no  20  20  4  Chase  frequency of use  Table  .583  .130  16. U s e o f t h e C h a s e b y m a l e s t o L a r g e Cows. T h e number o f i n t e r a c t i o n s when t h e Chase d i d and d i d not occur i s given. X2= 7 3 . 3 , p<.05, d f = 2  OB  IB  Chase  1  2  no  6  35  .166  .057  Chase  frequency of use  Table  .0  17. U s e o f t h e C h a s e by m a l e s t o S m a l l Cows. T h e number o f i n t e r a c t i o n s when t h e Chase d i d and d i d not o c c u r i s g i v e n . p>.05  40  OB  IB  YB  Females  and  Yearlings Investigative  0  11  32  14  Non-investigative  79  834  1694  68  freguency of use  .0  .013  .018  ,206  Table  18. U s e o f I n v e s t i g a t i v e b e h a v i o u r b y males, f e m a l e s and Y e a r l i n g s i n i n t r a class behaviour. T h e number o f I n v e s t i g a t i v e and N o n - i n v e s t i g a t i v e b e h a v i o u r p a t t e r n s a r e compared. X2= 9 4 . 9 , p<.05, d f = 3  41  i  males  YY  females  81  80  137  Non-investigative  3729  301  286  frequency o f use  .021  .209  .323  Investigative  Table  19.  U s e o f I n v e s t i g a t i v e by m a l e s t o m a l e s , f e m a l e s and Y e a r l i n g s . The number o f I n v e s t i g a t i v e and Noni n v e s t i g a t i v e behaviour patterns are compared. X = 7 2 3 . 0 , p<.05, d f = 2 2  males  Investigative Non-investigative  frequency o f use  Table  20.  f e m a l e s and Yearlings  37  27  599  184  .058  .128  U s e o f I n v e s t i g a t i v e b e h a v i o u r by f e m a l e s and Y e a r l i n g s t o males, f e m a l e s and Y e a r l i n g s . The number o f I n v e s t i g a t i v e and Non-investigative b e h a v i o u r p a t t e r n s a r e compared. X2= 1 1 . 0 4 , p<.05, d f = 1  42  Yearlings use  only  not  i s  acquired  class  behaviour  is  use  more  behaviour.  They  also  than  do  males.  Contact  with  Young  Old  behaviour.  pattern.  Bulls  and  Females  Body  Contact  i s retained  (Table  Non-body use  21).  Contact  more and  behaviour  as the  than  Non-body  Body  Intermediate  Bulls  Bulls,  who  similar  to females  u s e more  Males  are  direct  Body  22).  Males  mature,  and  behaviour  use l e s s  This  males  intra-  Contact  Contact  behaviour.  As  i n  Females  Contact  behaviour  Contact  compared  Contact  Body  Contact  less  Body  than  behaviour.  behaviour  and  Young  Contact  with  behaviour  by  use o f Non-body  Yearlings  use  Contact  matures. The  body  u s e Body  the the Mouthing  behaviour male  do  t h e amount  Non-  contrasts  of  Non-body Non-body  increases. selective  Contact use  t o which  sea-lions  and  Non-body  Contact  behaviour  more  Non-body  Contact  than  i n interactions in interactions behaviour  i n regards  when  with with  females males.  interacting  and  (Table  Body  Contact  Yearlings.  Males with  they  u s e more  females  They  Non-body  than  they  do  Yearlings. Females  behaviour proportions different males.  u s e more (Table of  Non-body  23).  Non-body  classes.  This  Contact  There  i s  Contact contrasts  no  than  difference  behaviour with  Body  Contact in  directed  the behaviour  the to  of the  43  Non-body Body  Contact  Contact  proportion of Non-body C o n t a c t  Table  21.  OB  IB  YB  YY  44  346  671  26  30  35  475  992  0  5  .556  .421  .403  1.0  .857  Use o f Non-body and Body C o n t a c t b e h a v i o u r by a l l c l a s s e s i n i n t r a c l a s s b e h a v i o u r . T h e number o f Nonbody C o n t a c t and Body C o n t a c t b e h a v i o u r p a t t e r n s used a r e c o m p a r e d . X*= 7 0 . 5 , p < . 0 5 , d f = 4  males  YY  females  1707  185  201  Contact  1935  96  72  proportion of Non-body C o n t a c t  .468  .658  .736  Non-body Body  Table  Contact  22.  Females  Use o f Non-body and Body C o n t a c t b e h a v i o u r by m a l e s t o o t h e r c l a s s e s . T h e number o f N o n - b o d y C o n t a c t a n d Body C o n t a c t b e h a v i o u r p a t t e r n s u s e d are compared. X2= 1 0 3 . 2 , p < . 0 5 , d f = 2  44  Yearlings class the  behaviour  sexes In  each  Mouth  Body  the  to  behaviour. that  of  use 26).  this  and Body  The  and  i s very  common.  and  patterns.  The use o f t h e  in  more  pattern Mouth  between  behaviour. Contact This  i s  and t h e M o u t h i n g f o r  Yearlings  Males  intra-  Body  u s e o f t h e Mouth  u s e t h e Mouth  behaviour  Contact  Contact  behaviour  females  behaviour  behaviour  do n o t d i s c r i m i n a t e  pattern  Contact  and Y e a r l i n g s  classes  They  Non-body  one b e h a v i o u r  Contact  (Table  24).  f o r Non-body  compared  body  of  Contact  Females  (Table  Contact  i n t h e use o f Non-body  behaviours, the  o s e a l l Non-body  than  by m a l e s i s (Table  25).  the other  use l e s s .  i n different decreases  Non-  The  male  frequencies  as  the  males  mature. The used  Mouthing  by  females  behaviour the use  where  Body  i s the only (Table  of t h e Mouthing  Population The known,  forms  behaviour  contrasts  only  There  by t h e m a l e  classes  number  of sea-lions  however  estimates  with  the  varied  colony.  At t h e t i m e  the  This  behaviour.  structure  well  Contact  a small  pattern  with  male  proportion  of  i s no d i f f e r e n c e i n t h e (Table 28).  Structure  familiarity  as  27).  the Mouthing  Contact  Body  a s most  actual  with  i n the d i f f e r e n t c a n b e made.  colony  (Table  immigration  classes  This  29).  i s based The  population  Method  structure  was  data  were  probably  not  c n my  population  t o and emmigration  the Individual Distribution  of the Encounter  i s  from the  Method  data  collected,  close  to that  45  Non-body Body  Contact  Contact  proportion of Non-body C o n t a c t  Table  23.  Non-body Body  Contact  Contact  24.  YY  females  177  23  54  37  9  10  .823  .718  .843  Use o f Non-body a n d Body C o n t a c t b e h a v i o u r by f e m a l e s t o o t h e r c l a s s e s . T h e number o f Non-body C o n t a c t a n d Body C o n t a c t behaviour p a t t e r n s used a r e compared. X2= 2 . 5 , p>.05, d f = 2  proportion of Non-body C o n t a c t  Table  males  males  YY  females  154  26  19  84  0  9  .647  1.0  .678  Use o f Non-body and Body C o n t a c t b e h a v i o u r by Y e a r l i n g s t o o t h e r c l a s s e s . T h e number o f Non-body C o n t a c t a n d Body C o n t a c t behaviour p a t t e r n s used a r e compared. X*= 1 3 . 4 , p < . 0 5 , d f = 2  46  f e m a l e s and Yearlings  males Mouth  378  39  o t h e r Non-body Contact  683  17  .356  .696  proportion o f Mouth  Table  25.  Use o f mouth by m a l e s , f e m a l e s and Y e a r l i n g s t o a l l c l a s s e s . T h e number o f Mouth and o t h e r Non-body Contact behaviour patterns are compared. xz= 2 6 . 3 , p<.05, d f = 1  Mouth  o t h e r Non-body Contact  proportion o f Mouth  Table  26.  OB  IB  YB  5  89  284  39  257  387  .113  .257  .423  Use o f Mouth by male c l a s s e s t o a l l c l a s s e s . T h e number o f Mouth and o t h e r Non-body Contact behaviour patterns are compared. X2= 3 9 . 2 , p < . 0 5 , d f = 2  47  males  females  280  5  o t h e r Body Contact  1222  0  proportion of Mouthing  .186  1.0  Mouthing  Table  27.  Mouthing o t h e r Body Contact  proportion of Mouthing  Table  28.  O s e o f M o u t h i n g by m a l e s and f e m a l e s t o a l l c l a s s e s . The number o f M o u t h i n g and o t h e r Body C o n t a c t behaviour patterns are compared. p<.05  OB  IB  YB  6  104  170  26  341  822  . 171  .218  . 171  U s e o f M o u t h i n g by m a l e s t o a l l c l a s s e s . The number o f M o u t h i n g and o t h e r Body C o n t a c t behaviour patterns are compared. X = 4.8, p > . 0 5 , d f = 2 2  Class  Proportion of  OB IB  .05 \  .30  YB  .40  YY  . 15  SC  .05  LC  . 05  Table  colony  29. ftge-sex s t r u c t u r e of non-pupping colony.  a  49  estimated.  Class  Association For  that It  each  belong  would  class  to the different  be  expected  association, neighbours whole  and  then  would  colony  sub-adult  males  the  classes i f  i s calculated  there  were  proportions  be s i m i l a r  to those  29).  That  and Y e a r l i n g s ,  of  no  (Fig.  found  as  of  the  characteristic  only  neighbours a  4).  assortitive  classes  i s , most  with  sea-lions  few  would  adult  be  males  females.  Yearlings.  associate associate associate  Old Bulls  Intermediate  themselves  and with with with  a s s o c i a t e mainly and  Yearlings Old Bulls, other  more  sub-adult  males  with than  Large  with  Intermediate with  Cows and  Large  expected.  the  other  Bulls.  They  Intermediate  as  Small  expected. classes  Bulls. Cows  than  and with  tend  Cows.  to  They  b u t do n o t Small  and l e s s  a s s o c i a t e with  Cows with  Old Eulls  Yearlings associate as except  a s s o c i a t e more w i t h  Bulls  Cows  expected,  and S m a l l They  females  associate  Large  and  Young  Cows  with  Bulls  expected.  Yearlings  Yearlings a t the l e v e l s  expected  Young  as  Intermediate  associate  less  that  (Table  Territorial  and  the proportion of neighbouring  expected.  Yearlings  and  t h e m s e l v e s and  50  100 OB  0 11  n =  37  14  13  100 IB  YB  c  67  23  IS  36  47  100  u «  52  YY  0 n=  100 SC  100 LC  0  1 3  n=  OB  Fig.  H.  IB  YB  42  20  YY  SC  LC  Association. F o r e a c h c l a s s on t h e o r d i n a t e , i t s neighbours that belong to d i f f e r e n t classes i s g i v e n i n p e r c e n t . Sample s i z e s o f l e s s than five were n o t i n c l u d e d i n t h e c a l c u l a t i o n o f X . X = 3 7 1 . 7 , p < . 0 5 , d f = 15 . T h e e x p e c t e d values are c a l c u l a t e d u s i n g T a b l e 29 a n d a r e s i m i l a r t o the Young B u l l and IB f r e g u e n c y distributions. 2  2  51  Activity The  proportion  calculated than  of  ( F i g . 5}.  females.  active Hales  As s e a - l i o n s  That  i s ,older  males  Large  Cows a r e l e s s  and  they  active  than  for  each  Yearlings  mature  are less  active  time  Small  a r e mere  become  than  class  less  younger  i s  active active.  males,  and  Cows.  Transiency Yearlings specific colony  (Fig. 6).  Females  Yearlings they  i n one a r e a  site  are less  on t h e  transient  than  specific  as they  mature.  i n t h e non-pupping  colony.  Using  females  become  more  site  specific  Bate  interaction  interactions  per  interactions derived observed.  i s given  minute  the  This  Intermediate as o f t e n  half  times  using number  Bulls  as O l d B u l l s  as often  decline  rate  in  An  and Large  Yearlings  and  number  expected  and  other  Cows,  and one  Small  Cows.  In  males  to a high  minutes  rate  rate. sea-lions and  There  i t  of  rate  observed  with  of  number  interaction  the  interact  mature.  mean  of interactions to  as Yearlings  as females  the  an o v e r a l l  compared  Young  as  (Table 30).  total  i s  and  twice  low  rate  i s calculated  from  a  more  one a r e a  site  mature.  The  from  become  words n o t  move f r o m  and females  as a comparison,  Interaction  gradual  or i n other  frequently  Males  Males  remain  transient  They  to another.  Yearlings.  as  are very  one i s a  increases  i n Young and  50  18  49  28  18  3  19  OB  IB  YB  YY  SC  LC Cn CO  Fig.  6.  Transiency.  number Jfrlhe  The  proportion  of s i g h t i n g s . sample  size  of m o v e m e n t s to the  X =33.0, P<.05. df = 4. 2  is too s m a l l so the d a t a  are  not  used.  54  # of interactions  class  minutes observed  interaction rate  OB  149  1188  .125  IB  283  1051  .269  YB  165  654  .252  YY  118  665  .177  SC  60  353  .169  LC  129  1003  .128  Table  i  30.  Interaction rate given as i n t e r a c t i o n s per minute. X2= 101.5, p<.05, df= 5 '  55  Intermediate The  between  decreases ontogeny  as males  Yearlings  with  Young  i n this  active all  time  As m a l e s manner.  spent  the time  values  that  there  This  negative  i s  is  very  the  significance  If the the  for  and  of there  of their  active  time  active  the  spends  (Table 32).  active  mature,  6%  of  a  i n  time i s  proprotion  Large  Zeros  i n Table  32 when  i t i s unlikely  way  then  Young  minor.  large,  less  d e f i n e d i n such  calculated,  i s  a sea-lion  portion  increases.  are interacting,  values  Interaction  mature,  Cows  (Table 31).  calculated  their  i n the  Cows  of  interact  are active.  i s a minimum  error  a l l  As females  they  i s  a small  interacting  that  two a n i m a l s  The  sea-lions  that  I t then  trend  Small  interacting time  increases  stages.  Yearlings,  active  spend  Bull  i s nc d i s t i n c t  time  f o r only  Bulls  Activity if  of  interacting  and Young  There  equal  other  interact  interactions. used  about  i n Old Bulls.  spent  behaviour.  proportion  interacting  time  mature.  o f female  The  of  decreases  the Yearling  Cows s p e n d  time.  I t then  proportion  sharply  Large  Bulls.  they  were  i n most  are  Bulls  error  X  that,  also  and L a r g e  involved used  in  that  these  active. Cows  show  (Table 32). place of the  i s calculated.  2  cases,  Since  adjustments  X  2  affect  o f the data.  Frequency  sea-lions  frequency  interact  randomly  of interaction  same a s t h e f r e q u e n c y  that  with  with  surrounding  a particular  the  classes  class  are  animals will  be  associated  56  OB minutes interacting minutes not interacting proportion of time spent interacting  Table  31.  IB  YB  YY  SC  LC  48.9  208.3  252.2  27.6  9.6  60.4  1139.1  842.7  401.8  637.4  343.4  942.6  .041  .198  .386  .042  .027  .060  Time each c l a s s spends i n t e r a c t i n g with s e a - l i o n s . X = 6 5 1 . 6 , p<.05, d f = 5  other  2  i  57  OB minutes interacting minutes a c t i v e but not interacting  proportion of a c t i v e time spent interacting  Table  32.  IB  YB  YY  48.9  208.3  252.2  27.8  137.6  138.5  -16.*T  .261  .600  1.072  153.3  .154  SC 9.6  LC 60.4  32.4 -2.2  .227  1.034  A c t i v e time each c l a s s spends i n t e r a c t i n g w i t h o t h e r s e a - l i o n s . X = 4 5 7 . 2 , p<.05, d f = 5 T h e s e e r r o r s a r e p r o b a b l y c a u s e d by an over estimate o f a c t i v e time. '0* was u s e d i n the c a l c u l a t i o n o f X . 2  2  58  (Fig.  4).  classes class more  t h e male  (Table  33).  classes  and  frequencies  frequencies  Sea-lions less  with  of  the  (based  on  tend  to interact  the  female  and  classes.  The  interaction  detail the  (Table  male  association. less  interaction  to the expected  association)  Yearling  of  observed  a r e compared  with  more  The  than  frequency  34).  classes  The expected  Cows  while  Bulls  i s considered  interaction  i s c a l c u l a t e d only  O l d and Young  expected,  o f Large  Intermediate  frequencies  on t h e b a s i s  interact  with  Bulls  in  c f male  Large  interact  Cows  more.  DISCUSSION  Hierarchy A the  peck-dominance  classes  i n non-pupping  California  sea-lion  Bartholomew,  1967).  dominant  the  to  chickens, amount This  less  a  also  class  Sandegren,  (Zalophus The  that  californianus)  and  would  fprsteri)  and  (Boyd 1970).  and  and  older  younger  otherwise New  (Stirling,  reduce  may  The  classes In  1971;  and are  domestic  reduce (Guhl, fur  the 1953). seal  establishment  function  socially  and  (Peterson  Zealand  1970).  i n 12. j u b a t a  Campbell,  occur  between  jubata  ones.  relationships  for  thereby  1951) e x i s t s  of Eumetopias  larger  suggested  hierarchy  fighting  mortality  colonies  smaller  fighting  (Arctgcephalus of  (Allee,  dominant-subordinant  of i s  hierarchy  i n causing  induced  Bartholomew, '  pup 1953;  59  Observed  Expected  Males  597  485  YY  145  194  47  108  Females  Table  33. F r e g u e n c y o f male, f e m a l e , and Yearling i n t e r a c t i o n s expected i s c a l c u l a t e d on t h e b a s i s o f random i n t e r a c t i o n w i t h neighbours. X*= 7 2 . 7 , p < . 0 5 , d f = 2  Observed  Expected  OB  37  51  IB  22  4  YB  ,4  8  Table  34.  I n t e r a c t i o n freguency of Large Cons with males. Expected i s c a l c u l a t e d on t h e b a s i s o f random i n t e r a c t i o n s w i t h male neighbours. X«= 8 6 . 8 , p<.05, d f = 2  60  Hierarchy and  i s r e l a t e d to  i s discussed  where  many  of  the  behaviours  measured  appropriate.  Duration Hany are  behaviour  patterns  Investigative.  keeping Young  aware Bull  of  This their  stage,  establishing  Hale  i n t e r a c t i o n s are  exchange the  than  those  small  because The between  more  and  the  to  in  stone and  Caldwell,  assert  peers.  of  the  more  i t ' s  or  and  very  involved.  i s being  This Males  status sheep  shrimp  The  information  dominance  or  1960).  information  The  duration  of  of  information and  results interact  as  dalli  the  female from  the  longer  animals  to  difference  behaviour  stonei)  i s exchanged both  the  Similar  (Gonodactylus  closer  as  exchanged.  decreases. (Ovis  the  neighbours  more  Yearling  short.  By  (Mason,  type  only  well  their  Yearlings.  1972).  are  as  skills  involve  amount  Yearlings  environment. this  so  and  females  interactions increases  mantis  1969).  doing  and  the  usually  animals*  1971)  hierarchy,  on  information  social  motor  females  information  duration  reported 1968,  of  that  and  (Caldwell,  are  amount  of  females  r e l a t i o n s h i p s with  complex  depends  communicated  interactions  are  communicative  interaction  being  immediate  hierarchical  developing  by  suggests  males  and  used  in  accept  zacae) are  is  (Geist, (Dingle in  the  order  for  one  each  other  as  61  The  association  of duration  interactions  i s  males  females  towards  Bulls Cows  interact and long  stone  that  with  for  periods  sheep  communicated  further  information  demonstrated  by  and Y e a r l i n g s . short with  (Geist, with  to  Large  with  Cows.  1968, 1 9 7 1 ) .  Large  Cows  i s  Y e a r l i n g s and Small  and  This  i s  The type  sexually i sn o t .  sheep  (Geist,  of  Intermediate  Y e a r l i n g s and  Cows  of  the behaviour  Old  periods  content  Small  similar  to  of information  oriented  whereas  Initiation A  J i j u b a t a and stone  s  initiate  more i n t e r a c t i o n s  classes. social  Older  older  animals  behaviour.  younger  contest,  would  classes  are not prevented  younger  classes  Differences evidenced Bulls, The  Intermediate  for  sexual  interactions sexual  with  immaturity.  they  can assert maturity  and Large  may  initiate  reasons. Large  starting  Cows.  Young This  hierarchy.  The  interactions  with  i s a  hierarchical The o l d e r  interactions  their  Large  classes are Cows.  Cows a r e s e x u a l l y interactions Bulls reflects  with  dominance.  o f t h e male with  of  voluntary  o r be i n j u r e d .  of interactions  Bulls  classes  lose  from  sexual  by i n i t i a t i o n  o l d e r male  Cows  in  do n o t s t a r t  they  variety  toward  to  I f the interaction  the subordinant  because  increasing  i s related  classes  classes.  an  a r e more i n c l i n e d  Initiation  subordinant  dominant  with  1968) m a t u r e ,  do  mature.  with not  Old  Large  initiate  t h e Young  Bulls'  62  Variety  o f Behaviour  The males  variety  i s  arises  than  different  interactions  involve  (Farentinos, between  of behaviour  greater  from  Patterns  1971).  that  patterns i n the  hierarchical type  the participants  complicated  threat  behaviour  Yearlings.  The  variety  communicating  their  more c o m p l e x  The one of  and T e r m i n a t i o n Advance  sea-lion  an  effort  this  action  This  of  more  indicates  While  with  of  the simple  females  signals  being  males  similar  and  assists  in  approach  of  do  to  initiation  The sender  a particular  f r e q u e n t l y than that  than  messages.  i t i s more i n t e n s e .  interact  fighting  transferred  pattern i s a definite  to another.  to  male  Male  Behaviour  behaviour  an i n t e r a c t i o n  information  i s more  and  This  relationships.  c o n t e s t s a n d mock  of  investigative  Beginning  minor  of  o f Y e a r l i n g s and f e m a l e s .  intra-class  The  repertoire  animal.  Yearlings  a r e more  i s  willing  making  Males use  and  females.  t o be  socially  involved. The Cows  increases  sexual Small of  frequency  with  maturation. Cows,  there  the  Advance.  sexually  immature  o f t h e Advance t h e male's  When  males  i n interactions age and r e f l e c t s  interact  a r e no s i g n i f i c a n t The  males  ( p a g e 66)  may  with  with  a n d s o do  the male's  Yearlings  differences  recognize  Large  i n their  these  not approach  classes them.  and use as  63  Males  are  Advance. is  used  selective  This most  and  interactions  behaviour  behaviour Bekoff,  interact  an  with as  by  peers well  provide as  continually Although would  be The  to  approach  more  Attack  being  become  obviously  Young  Bull  be  be  If  the  developing  1971;  with  favour  by  extreme  males  animals  that  and  peers.  subordinants subordinant  exposed  be  to  animals  Little  new  However  the  new  behaviour  Animals  selected  gained,  and  Interactions  acquired.  would  very  the  that  against.  subordinant  injury. pattern  between  as  with  Farentinos,  dominant.  already  male  more a g g r e s s i v e  stages,  avoid  social  practised.  would  risk  behaviour  differences  to  male  learning  interactions.  be  dominants  and  major  the  can  that  would  Immediately to  most  males  interaction  injured  i n these  experience  defeated  of  subordinant  opportunity use  w e l l as  1968;  Selection  acquired  immediately  Since  smaller  the  Advance  experiences.  as  value  similar  is  peers.  of  The  classes functions in  peers.  i s encountered already  that  new  Schaller,  of  use  Interactions  skills  the  chance  behaviourally  behaviour  yield  immediately  chance  behaviour  ones.  male  i s minimal.  is little  with  their  hierarchy.  aggressive,  motor 1960;  on  in  class  often  not  the  then  with  egual  are  of  (Mason,  subordinants  do  and  1972)  most  the  larger  animals  males  i s based  quite  with  communicative  There  next  are  interactions  intense  selection  other  f r e q u e n t l y with  subordinate  subordinant  of  demonstrates and  female  between  indicated  by  one  of  behaviour. the  the  the Males  Yearling  first  use  of  and the  64  Attack.  The  increase  use  as  of  the  this males  sequence  i s evident  1972).  The  use  of  mature.  The  adult  they about  as  Attack. This  are  In  Ei  i s also  (Adenota sheep  the  true  kob)  (Geist,  potential  of  important  in a  of  feral  and  1964;  Bartholomew,  be  does  approached  which  (Shank,  not  change  colony  used  they  1972),  Schloeth,  1956)  females  where by  would This  direct  against  (Shank,  lost.  colony  1952;  to  i s not  goats  Otherwise  pinniped  immediately  hircus)  non-pupping  classes  Attack  would  almost  developmental  as are  Yearlings.  (Buechner  mates  (Cajjra  at a  aggressiveness  similar  goats  females  the  1971).  A  also  Attack i n females  selective jubata  ana  mature.  in feral  a g g r e s s i v e as  Males  pattern  females.  Uganda and  females  n e i g h b o u r i n g males et  and  particularly  unattended  Carrick  kob  mountain  withdraw  i s  the  are  (Paulian,  a l , 1962b;  Sandegren,  1970). The pupping  Attack colony,  (Matheson,  would  If  The  use  subordinants, males  and  males  f o r c e , them reduce  as  they  the  7%  Lopp, were  away  of  1962)  their  mature.  elephant  1972)  non-breeding  the  on  in  (Mirounga  adult  male  1970)  females  of  the  E^  and are  follow.  255?  i t This  especially  a g g r e s s i v e n e s s of  s p a c i n g out  gngustirostrig) jubata  a  males.  males,  the  In  suckling  Yearlings  might  other  changes  Yearlings.  towards  Aggression causes  seals  on  mothers  success  Attack  indicates  of  aggressive  and  the  used  (Sandegren,  reproductive  northern and  occasionally  between  Badde  Yearlings. would  i s only  (Orr and  of  male  (LeBoeuf, Poulter,  65  1967).  In  E._  maturity  jubata  may  , the  increase  accomodate  the  in  aggressiveness  development  of  with  territorial  behaviour. The could  lack  be  as  females around  intense  adaptive  were  as 1967).  would  be  tend  to  lost,  trampled  or  Retreat  to  in  do  a  hierarchy  pattern  dominant.  If  are  The that of by  a an  an  pattern  sea-lion  is trying  interaction.  of  and  This  Yearlings.  Initiation  that  male  the  of  effect  Yearlings  behaviour  is  of  This male  would  chances  often  when  of  can  pattern It  is  has  behaviour  and  use  the  interactions  results  Sea-lions they  defeated  prolong  sea-lion  Males  from  the  and be  or  Class  use  interact  is agonistic,  pattern  a  interactions.  Retreat.  be  by  intra-class  the  to  used  interaction.  of  to  behaviour  mates,  also  and  If  groups  fewer  increasing  Yearlings.  going  males.  close  and  their  Yearlings  pups.  proportion  most  have  females  because  an  using  of  peripheral  colony,  interaction  surely  Chase  females  because  aggressive  or  and  form  success,  the  against  not  would  Highly  pupping  defined  male  of  characteristic i s  could  result  greater  behaviour  animals  The  behaviour  females  aggression  they  terminate  females  a g o n i s t i c behaviour  neglected  Retreat  attempting  than  the  in  reproductive  selected  disrupt  The  his a  (McLaren, also  the  males.  reducing  females  as  aggression  aggressive  territorial  thereby the  of  with  subordinant  so  to  try  escape.  i n t e r p r e t e d to  by  already the  a  then  r e i n f o r c e the  used  this  mean  result  males  more  been  suggested  advance,  that  than  males  66  are  more  voluntarily  classes. ~  The  The  sexual  relationships  use  of  Chase.  more  the  use  socially  Although  significant classes  older  males Cows  use  i t  females age.  in  males  distinguishes  sea-lions explains  constant  that  the  permits  being  Bulls  small,  As  of with  prolonging  use  the  interacting  use  Cows.  a  by  pattern  with  there  Large  are  no  this  action  the  Advance,  the  interactions  with  their  do  males.  This  young  from  used  to  use  by  only  by a l l  old  by  maturing  contact  the  use  increasingly  to  not  are  It of  with  that  age,  and  sex  and  the  of This  greater  also  possible  and  Yearlings  animals  behaviour.  themselves.  with  readily  visible.  females  are  the  Investigative  is  interacting  as  obvious  not males  Yearlings  change  more  Investigative  defend  and  maturity  maturing  nature  aggressive  order  the  between of  does  animals.  females.  aggressive  Investigative  characteristic  features  use  of  Females  becomes  identify  physical  decreasing  amount  interactions.  i s the  be  in  decreasing  size  closer  interacting  when  the  dimorphism  less  subsequently  i s reflected  are  Male  when the  use  than  females  may  the  classes  conclusion.  reasons.  intra-class  mature.  behaviour  probably  ether  Behaviour  more  In  Small  for sexual  Maturing  in  the  this  Bulls  samples  than  complements  Intermediate  Young  towards  Investigative  behaviour  and  the  are  Chase  between  differences  males  Large  the  Old  f r e q u e n t l y than  Cows.  but  of  inclined  separated  and Males, when  This separation  67  does  not permit  Body  Contact Body  and Son-Body  Contact  aggression.  patterns  other and  but  classes and  i n  the  pinniped  between an  of  result  several  the Yearling  a n d Young  increasing variety males  the  mature.  type  of  Yearlings many  are  of these  males  hierarchical  a n d mock  life  of  establishment  (1971)  these  fighting  adult  Most  i s mock who  their  differentiates  1. t h e patterns  the use of patterns  i s related  to  Females  and  and t h e r e f o r e  lack  repertoire. i n  i s  socially  of territories  This  male  The  their  highly complex  interactions.  of the males'  fighting.  a n d 2.  62).  patterns  males  and maintenance  interactions. colony  use  in  sources:  behaviour  as males  1956;  between  patterns  (page  Peterson  by t h e v a r i o u s  behaviour  Contact  of behaviour  not as a g g r e s s i v e patterns  used  stages,  interactions  aggressive  The  o f Non-body  Variety  social  Bull  of  1952).  two m a j o r  Contact  Many o f  Laws,  Differences  from  Body  1971;  1962b;  patterns  previously.  classes of  a l ,  also  repertoires  1965; Bartholomew,  behaviour  are  and d i s p l a y s .  (Stirling,  et  expressed  patterns  behaviour  species  1967; C a r r i c k  discussed  development  as  a r e found  variety  female  behaviour threats  1964; P e t e r s o n ,  was  are physically  to  Bartholomew,  The  patterns  Contact  behaviour.  Behaviour  limited  polygynous  Paulian,  Contact  behaviour  Non-body  aggressive these  t h e use o f I n v e s t i g a t i v e  behaviour  term  i t  from  i s  used  true  complex. involves  The  intense  in a  non-pupping  by  Farentinos  fighting  because  68  wounds a c e play low  not  inflicted.  fighting intensity  sub-adult intense  can of  males  as  colonies  by  recognized  Mock  fighting  is  reviewed  by  experience  Males and  not  would  use  more  expressed  larger  and  potential  and older  Body  males  aggression  pupping  (1970).  male  These  interactions  O r r and  Poulter  are  (1967).  of communicative  and  1972).  in  life,  show  marked  early (Gilbert,  social  1968;  Mason,  interactions  adapted  gaining  1960;  and  play  early  social  t o t h e complex  social  Contact behaviour as  males  than  mature,  aggressive, a  but  rather,  different  do  for  inflicting  injuries  fight  very  younger  close  restrictions  their  separated  would  result  in  are  males so  The  because  because From  Males aggression  i s greater.  injury.  Body  and  prevent serious  remain  of  Older  ones  together  must  females  t h e amount  manner.  than  behavioural  a  or  Shank,  experienced  to  jubata  1968;  more  are able  13.  the  vicious  Contact behaviour decreases.  in  by  life.  However,  less  and  as at  Sandegren  Sea-lions  better  adult  being  males  be  t o Non-body  becoming  (1972).  not  of  or  Schaller,  abilities  importance of  Yearlings.  Contact  restricted  Bekoff  of t h e i r  (1970)  mock  fighting  males  sub-adult  1960;  social  The  adult  are  i n the development  (Mason,  of  and  Gentry  socially  1971).  system  by  adult  true  that  interactions  colonies  and  states  from  The  for  (1970)  assists  skills  impairment Fox,  at non-pupping  between  also  Animals,  interactions.  Gentry  (1972)  distinguished  those described  differences  motor  be  Bekoff  remote  are  their younger  physical  injuries. their  i s  The  intense positions  69  they  interact  behaviour. itself  the  If  or  Contact  each  and  male  part  unaccustomed not  to  been  males  mature,  they  display  the  a  1970).  in  reserves  conserves  energy  colony.  in  tenency  of  feed  males  1967;  are  Peterson  establishing this  Contact  Geist, the  so  the the  Contact  of  rely  behaviour i t s  at  a  use  1970). pupping  reproductive  Non-body  behaviour  long  but  Gentry,  stay  or  territory  and  males'  use  a  Contact  1966;  males*  for  time  behaviour  1971)  Contact  remain  Non-body  1970;  (Geist,  Non-body  males  during  increases  Body  during  relationships  sheep  of  after  and  case  when  Rand,  stone  adult  prolong  1970)  Body  1971).  1970).  (Stirling,  asserting then  the  social  1970;  amount  not  Body  could  i s often  Contact  from  peer,  season  -jubata and  colony  male  as  and  (Gentry,  the  do  than  place  This  Non-body  one  other  breeding  (Gentry,  (Gentry,  behaviour  E.  They  saved  Longer  success  used.  Stirling,  pupping  energy  permit the  E.. j u b a t a  strenuous  using  neighbours  increase  (Gentry,  The  of  both  in  less  be  1967;  periods  is  may  established  of  to  accepting  behaviour.  fat  fails  their  Bartholomew, is  communicate  this  behaviour  early  have  and  is  Contact  selectively  advantageous. Males less  with  Contact so  may  Males order  use  more Non-body  Yearlings behaviour  be may to  s  and  Non-body  attract  least  patterns are  interpreted use  Contact with not  differently Contact  them  and  behaviour males,  physically by  males  behaviour promote  more  female  with  females,  The  Non-body  aggressive and with  and  females. females  in  gregariousness.  70  This  would  Body them  tend  Contact to  to  i n c r e a s e the  behaviour  males*  would  disrupt  amounts  treat  a l l  Body  and  of  classes  This  indicates  behaviour  their  relationships  and  Yearlings  use  females,  but  Yearlings  are  interact  with.  none  It  and  only  i s the  behaviour The  use  most  Bull  stage The  males  of  Body  important  simple  reflects  constant  by  the  classes.  with  males  that  the  animals  they  argument  of  early  experience. to  females  Lunging  and  and  or  Yearlings  pattern  simple  and  suggests  the  behaviour  Mouth  is a  simple  develop  i t s use  behaviour.  in  their  non-aggressive  their  Mouth  of  Contact  This  indicates  before  display  the  (Sandegren,  relative  to  t h e r e f o r e not Contact  more  more  mature.  Body  Young  further.  Non-body use  other  i s acquired  decreases are  to  males  threat  Males of  as  behaviour  not  the  used  female  This  in gaining  relative  does  use  to  are  classes.  Contact  variety  behaviour  of  relative  of  other  p a t t e r n compared  Contact  and  and  the  Yearlings.  further  Mouthing  mature,  with  behaviours  adds  Body  these  increasing  Females  cause  behaviours  simplicity  Contact  infrequently  remains  that  the  is a  This  of  behaviour  also  and  when  Contact  the  other  being  i s  repertoire.  use  It  Mouthing  Body  with  mimicing  Necking.  the  some  involvement  The  Contact  females  similarly  Non-body  considered.  As  the  success.  disperse.  Females  social  reproductive  complex  than  other only  behaviour do.  Non-body  increasing  behaviour  males  1970).  As  but  are  patterns. with  the  71  Mouthing,  this  demonstrates  the s i m p l i s t i c  nature  cf  female  behaviour.  Association  and I n t e r a c t i o n  Sea-lions colony. around  a r e not randomly  Females, Old Bulls  selective. reach  begin  to  pattern  at the Small and o t h e r  Immature  males  form  i s evident 1967)  classes  are  distribution social  the  partners.  Large  no  colonies  i s  not  sea-lions  Selective  evident  i n many  Selective distribution females  more  of  than  i f  randomly  from  initiate  interactions  Intermediate them. in  surrounding  involve  males  Bulls  Harassment  the  following  and  of  bighorn  inclined  animals. males,  attempt  of females  sheep  than  by  species: E  A  more  (Geist,  females.  This  behaviour.  chose  with  clumped  i n further  Intermediate  they  The  the  results  a r e an i m p o r t a n t  classes.  of  interaction  o f male  interactions  1970; O r r and  available  socially  aspects  distribution  their  true  is  they  restricts  This  a r e more  territorial,  obvious.  than  Males  more When  proportions  , interactions  1971).  group  clumping.  (Gentry,  I n E_. j u b a t a i s also  to  a r e even  T h e same  restrictions. females.  Cows  a n d become  ncn-pupping  begin  distinct  the r e l a t i v e  :i t  in a  stage,  females.  a t pupping  but because  of  show  with  different  Cow  stage  groups  distributed  females.  males  the Old B u l l  Poulter,  Frequency  Bulls  their  they  sub-adult  in  interact  social  Although  harassed  (Gentry,  i s  with  seldom  interactions  males  the  partners  females  are  to avoid  jubata  factor  by with  reported  1970),  Alaska  72  fur  seal  (Callorhinus ursinus)  elephant  seal  northern  elephant  (iJSiilpjsIpra Bromley show  territories. females with  near  Old Bulls  Behaviour  others  females.  and  males  Females  This  they  be  (Gentry,  be  no  sub-adult (Gentry,  and  male  males  are  access  to females.  less  tend  males  than  to  conclude  excluded  the by  Since  areas  of  territories.  colonies are  i s related a  to  releasing  the  1967;  Kenycn,  Paulian,  1 9 7 1 ; Hewer are  not  advantage  i f  mechanism  eliciting in  1960; P e t e r s o n  and  1967;  Bartholomew,  and Backhouse, at  colonies.  they  of  behaviour  territorial  Intermediate  territorial  presence  Y e a r l i n g s a r e s e x u a l l y immature,  territories.  from  i n aggregations  leave  created  1970) o r n o n - p u p p i n g  reproductive  maintained  their  1970;  1954; Cameron,  Bulls  and  influence territorial  1952;  Laws,  males  Transiency  1967; Rand,  colonies  by t h e  a t non-pupping  may  since  pinnipeds  jubata  are harassed  they  Bartholomew,  E.  territories  i n the refuges  are not.  territoriality many  They  a r e harassed  males,  Some a d u l t  Only  harassment.  sub-adult  and c o l l e c t  Territorial  while  ,  This r e s t r i c t s  sub-adult  harassment  1973).  a c t as s a n c t u a r i e s .  E. j u b a t a  n  pronghorn  this  courtship of non-territorial  territories I  outside  and  and K i t c h e n ,  quantify  southern  e t a l , 1962a,b),  1972)  (Bromley  (1973)  1953),  (Carrick  (LeBoeuf,  americana)  females  overzealous  leonina)  seal  and K i t c h e n  that  that  (Mirounga  (Bartholomew,  1960). pupping  Since  Young  there  would  established Bulls  are  and  sexually  73  mature.  However,  colonies  i t  refrain  from  may  Intermediate 1966).  on  colony.  growth. time,  be  stage  Those  would a  socially  with  Non-territorial interact  thereby  adapt  Ji This  may  the  JJjbtSta  habituation become one  the  socially  place.  This  relationships  to  Females reflects in  system  the  their  non-pupping  feeding  have  and  Bulls, able  to  stay  consequently  free to  tc  feed  any  establish  a  would  be  of  experience  familiar are  free  neighbours  be  at  able  pupping site  with  mutual  the  Animals  around colony,  with  more  communicate  and  specific of  as  they  male the  territorial their  move  colonies.  discusses of  from  neighbours.  to  of  to  development  stability  allows  are  territory  experiences.  (1970)  familiar  be  phase  Bulls  more  Gentry in  to  a c c e l e r a t e d growth  gain  number  better  become  accommodate  Males  territorial  they  social  would  complex  behaviour.  are  unfamiliar  new  experience to  that  Intermediate  gaining  f o r them  colony.  small  with  non-pupping  the  sooner  since  a  a  their  and  at  in  Intermediate  Bulls  deprived  interactions  social  restricts  pupping  females  advantageous  i n an  maintain  larger  Intermediate  few  behaviour.  are  that  This  be  at  are  selectively  Non-territorial  territory  and  there  territorial Bull  (Pike, the  since  territorial  importance systems.  neighbours  boundaries  mature.  and  by  of Males  staying  in  hierarchical  established. more  adjustment colonies.  site to  the  specific limited  Harassment  than  males.  territorial by  sub-adult  This system males  74  forces are  females  available.  adjacent 1970;  Peterson  The sub-adult less  and remain  pupping  i n  whatever  colonies  females  where  there  and I n t e r a c t i o n Time  high  levels  of  activity  males  affect  class  than  other  and  remain  activity  contributes  to  the  (Sandegren,  and i n t e r a c t i o n  distribution. areas  in  sub-adult  i n the t e r r i t o r i e s  (Bartholomew,  a r e many  1967).  I n t e r a c t i o n Rates  of the exclusion of  territories  are not s t a t i o n a r y  and Bartholomew,  disrupted  because enter  In  territories,  Activity,  of  to occupy  rates of  Territories  a non-pupping males.  The  i n order  are  colony females  to avoid  areas  1953; C a r r i c k e t a l , 1962b).  clumped  distribution  This  of Old Bulls  and  females. The  high  interacting more than  interaction  of  social  contact  females  experience  sub-adult  and  with  rates  and  males  results  a broader  Yearlings.  t o the sub-adult  longer  range  The  males  times  i n their of  spent  receiving  colony  adaptiveness  has a l r e a d y  members of  been  social  discussed  (page 6 3 ) . Transiency observed move  in activity  around  neighbours. familiar  may  less  unfamiliar  a factor  and  so  the colony  rate.  become  (1970)  interact  animals  c o n t r i b u t i n g to the  and i n t e r a c t i o n  Gentry's  animals  move a r o u n d  be  less  habituated  data  suggest  frequently.  and a r e always whose  sex  The o l d e r  and  being  trends animals  to  their  that  socially  Younger  sea-lions  confronted  status  are  with  unknown.  75  Establishment interaction  of  rates  Yearlings not  display  do  spend  explains social spent  social and  long  time their  little periods  using large  the  as  social  males  the  harassing In  males time  with  (Stirling,  two  behaviour time  get more  time  is  their  initiated are  This  they  pattern.  This  the  displaying spend  older  and  the  time  an  interacting.  the  disturbed  direct  active  spent  time  complements  no  females  by  do  However  with  older,  active  males.  colonies for  in  of  The  non-pupping The  close  species  the  21.9$  1971).  respectively. the  Bulls.  of  females  active  Ii. j u b a t a  higher  male  forced  to  observation  of  females.  pupping are  Old  Maturing  i n t e r a c t i o n s are  interact  in  i n t e r a c t i n g and  partners.  Since  inactive  time  active  and  proportion  classes,  do  males  increasing these  as  of  i n t e r a c t i n g decreases specific  active  Looking  amount  involvement.  without  results  activity.  spend  for  relationships  are  New  and  Zealand  fur  seal,  interacting for  values  for  colonies  agreement  of  operating  3.1%  territorial are  15.7$  these  under  data  adult of  adult  the male  and  4.19?  suggest  that  similar  behavioural  systems. Stirling of  the  This  adult  prolongs  their  males the  at  a  is  the  pupping  attributes to  the  males'  reproductive  explanation have  (1971)  the  adaptiveness  presence  potential. stabilizing  colony.  relatively  at  of  the An  conserving  colony  and  equally  e f f e c t reduced  Interacting  low  males  activity energy. increases plausible  activity  disrupt  would  females  76  and  cause  al,  1962b)  would with  them and  favour  more  becoming passive  move away  thereby adult  a minimum  males  to  males  amount less  they  of  lose who  potential  could  and  Contact  1953;  Carrick  mates.  maintain  disruption.  active  Non-body  (Bartholomew,  This  Selection  their  territory  i s accomplished  interactive  and  et  by  using  by the  behaviour.  SUMMARY Non-pupping into  a  peck-dominance  classes of  c o l o n i e s p f E±  dominant  the o l d e r  females  and  differences  females  Non-body This  result  behaviour than  are and  the by  agonistic. use  an  mock  Body  Contact a  between  fights.  exclusive  use  result  The  This  and The  whereas  older status  those  males  of  they  interact of  be  territorial mature  of  Non-body  Contact  become  the  interactions violent  between more  Contact  agonistic the  Body males  aggressive  behaviour  behaviour  in  use  females.  where  adaptive  males  with  tranguil  Contact  allows  These  increase their  o f Non-body  Non-body  would  and  Interactions  mature  and  others.  sexually  i n more  only  amount  behaviour  distance.  Males they  males  ones.  hierarchical  predominates. As  larger  towards  when  involving  increasing  their  from  behaviour  females  organized  are not.  sexual,  almost  behaviour  younger  differently  from  the  distinct  are non-agonistic.  interactions  Contact  and  are  Interactions  Contact  and  males  behave  relationships. and  classes  younger  are s o c i a l l y  h i e r a r c h y , with  to the smaller  male  Classes  jubata  in  replaces  communication territorial  77  situation  at  Males  at  interact  and  so  is  voluntary  shown the  are  by  more  the  development  and  able  in  to  cope  Adult that  of  larger less  of of  areas  of  males,  of  as  that  group to  and  areas males  of in  and mix  the  and  the  behaviours The  and  males  colony's are  the  males  not  at  subsequent  of  This  so  better  males  vigorous and  than  v  females  use  behaviour,  are  less  time  Territories  where  adult  tranquil  than  females  avoid  are  more Since  sexually  which  mature  provide  sub-adult  refuges.  This  organization.  territorial.  Some  but  behaviour  pupping  the  spend  territories  expulsion  together  by  spacial  This territorial  the  less  aggregate.  harassment  maintain  adult  are  use  society.  and  in territories  males  adult  and  i s  social  skills.  less  males.  and  The  abilities  freguently  This  contributes to  Contact  concentrated  involvement  interactions  motor  adult  females  females.  p a s s i v e Non-body  sub-adult are  for  fights  in social  males.  sub-adult  females.  results  female  interact  Sub-adult  near  the  and  activity  establish  mock  c o m p l e x i t i e s of  females  contributes  by  than  patterns.  with  than  they  behaviour  separated.  social  initiate  adept  where  areas  to  so  animals  amounts  and  less  are  frequently  This  communicative  sub-adult  interacting  but  males  more  involved.  Chase males  where  and  tendency  their  male  active,  males  longer  males  males'  conditioning  colony  socially for  Advance  results  pupping  only  aggregation  outside territories  when  i s not  colonies.  sub-adult  adult  males of and  as  they  are  intense  Territoriality from  females. do  males  not  certain Sub-adult segregate  78  into  age  specific  Sub-adult free  to  move  transient,  males  do  about  the  are  established.  more  maintain  colony.  results  is  t e r r i t o r i e s and Since  hierarchical  continually  This  social.stability are  not  individual  relationships  who  groups.  being in  accomodated  stationary  and  and  adult  so  are  other  broken  social by  they  so  and  are very  social new  ones  disruption,  whereas  males  females  maintain  and  prolonged  social  relationships. Non-pupping pupping sex  colonies  classes.  organized Ji  jubata  adults males.  colonies  and  than  except  are for  Non-pupping pupping  colonies  is  inhibited  s p a t i a l l y organized the  colonies  colonies.  promoted by  relative  the  by  the  disruptive  similar  proportions  are Social  less  of  to age-  socially  organization  in  behaviour  of  tranguil behaviour  af  sub-adult  79  ACKNOWLEDGEMENTS I  am  very  supervision Krebs, Maser, Allan their  Tsitika  I.  and t h e f a m i l y  Research  support. Council,  States  to  Murray  Fisheries  Dean  Br.  Kennedy,  o f Maurice  The r e s e a r c h  H.  thesis.  Cowan,  The Canada  National  Dr.  of this  McTaggart  Harestad,  assistance.  logistic  United  and c r i t i c i s m  Dr.  Edie  grateful  I thank H.C.  Geographic  Dr.  Eoard  Society.  his  J.R. 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