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Some ecological aspects of social behaviour in the song sparrow, Melospiza melodia Knapton, Richard Walter 1973

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C.I SOME E C O L O G I C A L IN  T H E SONG  ASPECTS  OF S O C I A L  BEHAVIOUR  SPARROW, § § l o s p J L z a m e l o d i a  by  RICHARD B  Sc., Lakeheaa  A THESIS THE  WALTER  KNAPTON  University,  SUBMITTED IN P A R T I A L REQUIREMENTS MASTER  in  FOR OF  1971  FULFILMENT  T H E DEGREE  OF  OP  SCIENCE  the Department of Zoology  We  accept  this  thesis  required  THE  UNIVERSITY  OF  as conforming  to the  standard  BRITISH  September  1973  COLUMBIA  In p r e s e n t i n g an the  advanced degree at Library  I further for  this thesis  shall  the  of  this thesis  written  University  of B r i t i s h  permission for  s c h o l a r l y p u r p o s e s may his  f u l f i l m e n t of  make i t f r e e l y a v a i l a b l e  agree that  by  in partial  representatives.  be  for f i n a n c i a l gain  permission.  Department  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, C a n a d a  the  I t i s understood  Columbia  shall  requirements  Columbia,  for reference  extensive  g r a n t e d by  the  and  Head o f my  be  thesis  Department  copying or  that  study.  copying of t h i s  that  not  I agree  for  or  publication  allowed without  my  i  ABSTRACT  The  purpose  experimentally Song  of  some  Sparrows,  this  ecological  Helosjaiza  melodia  questions  Two  A.  Does  the  temporal  influence  the  number  of  territory  size,  and  breeding  B.  the  the  opportunity  arises,  hierarchies  established  ft  necessary  answered  is  that  non-territorial are  and  the  showed area one  there and  that of  the  probably  Two  of  of  a  in  for  both  of  were  c a r r i e d but  were  was  types  surplus  of  birds  were  and  given  area?  season?  to  but  by  the  are which  resident  fall and on  and  of  1972  breeding  the  breeding.  juveniles,  be  investigated.  the  Sparrows  the  dominance  breeding,  in  of  when  in  replacements  Song  hence  which  experimentally  p h y s i o l o g i c a l l y capable  replacement local  a  territories  birds  territories  Subsequent  in  British  questions  of  experiments  there  a  pre-breeding  is  1973.  on  individuals  also  of  Island,  territories,  surplus  taking  behaviour  territories,  p o t e n t i a l l y capable  from  social  settling  density  the  investigate  :  obtain  during  to  Reifel  asked  dominant  exists  of  This  spring  that  on  occupied  that  the  holders.  Removal  are  pre-requisite  prevented  territory  ,  pattern  juveniles  is  aspects  Columbia.  Are  main  study  study All  a l l  but were  origin.  of  removal  experiments,  simultaneous  and  i i  successive,  were  carried  out  i n both  the  Simultaneous  Removal  nine  to  Significantly  and  the  Further,  ten  days.  increases the  mean  significantly boundaries  The three the  was  both  were  completely  replacement  on  days.  number  of  after  the  than  size that  and  replacement  territories and  fall  after  were  the  before.  Further,  nor  taken,  was  territory removals.  took  upto  difference  i n the  the  about  removals  Finally,  significant  taken,  took  40%.  areas  no  On  about  S u c c e s s i v e Removal was  fall.  were  the  territories removals.  more  spring  rearranged following  the  There  total  spring  territory  smaller  to four  size,  in  areas,  the  mean  in  territory  territorial  pattern  retained.  Breeding same  before  several these  density and  unmated  unmated  territories  after males  than  the  experiments. territories  proved  mated  which  a l l a r e a s , however,  with  males  Factors results  on  to  remained  much  Therefore, there after  have  the  the were  removals,  significantly  and  smaller  ones.  could  have  of  the  simultaneous  experiments  are  discussed,  accounted and  and  the a  f o r the  different  successive  proposed  removal  explanation i s  given.  Dominance of  juvenile  certain  hierarchies  Song  Sparrows  localities  along  were that the  determined congregated  hedgerows.  in  the  over Each  loose  the group  groups  winter tended  at to  i i i  be  a discrete  (both  dominant  themselves reversals  of  unit,  although  and  were  some  interchange  subordinate)  stable  and  and t h e o c c a s i o n a l  essentially  removal  experiments  the  members  of the hierarchies  that,dominant  successful largest obtained  ones  presented  males  in  i n establishing  hierarchy,  Factors  the  position  of the bird  the  possible  outcomes  The  hierarchies  linear,  with  an o p p o r t u n i t y  to obtain the  few  hierarchies  which  males, could  i n the hierarchy  of the hierarchy  were  Further, 4  from  possibly  are  are  f o r some  territories.  territories.  o u t o f 12 j u v e n i l e  territories.  individuals  triangle.  The  was f o u n d  occurred.  of  I t the  i n the  the top  5  influence  discussed,  considered.  and  iv  TABLE OF  CONTENTS PAGE  ABSTRACT  i  TABLE OF CONTENTS  iv  L I S T OF TABLES  ..  L I S T OF FIGURES  >  v  ACKNOWLEDGEMENTS GENERAL  vi i  i  viii 1  INTRODUCTION  1. . AIMS OF THE STUDY 2. THE STUDY AREA 3. L I F E HISTORY U. METHODS PART A.  THE EFFECTS OF TEMPORAL PATTERNS ON TERRITORY S I Z E  1 3 5 11  ..  SETTLING  INTRODUCTION 1. F o o d s u p p l y and t e r r i t o r y s i z e ..... 2. Other environmental factors and t e r r i t o r y s i z e ..................... 3. Temporal p a t t e r n s o f s e t t l i n g on territories I I PROCEDURE I I I RESULTS 1. R e p l a c e m e n t o f removed p a i r s . . . . . . . . 2. Number o f o c c u p i e d t e r r i t o r i e s ...... 3. Size of t e r r i t o r i e s 1. C h a n g e s i n number of- t e r r i t o r i e s on t h e F a l l Removal A r e a o v e r t h e winter 5. Breeding d e n s i t y i n the s p r i n g of 1973 6. Differences in territory size between mated and unmated m a l e s .... 7. The r e p l a c e m e n t b i r d s  16 16 18  I  IV  DISCUSSION ... A. The r e p l a c e m e n t Song S p a r r o w s and the s u r p l u s B.  The t e m p o r a l p a t t e r n s o f s e t t l i n g on t e r r i t o r i e s . . . . . . . . . . . . . . . . . . . . .  2  2  2  4  2  ? 32 32 33 35  40  42 44 47 51 51  55  V  1.  C.  Fluctuating environmental resource 2. P r e s s u r e from i n t r u d e r s 3. Inexperience of the j u v e n i l e s and the temporal patterns of s e t t l i n g 1. Disruption of territorial boundaries 5. Genetic basis f o r the size of t e r r i t o r y .................. Proposed explanation f o r the different results of the simultaneous and t h e s u c c e s s i v e removal experiments ................  I INTRODUCTION , I I PROCEDURE I I I RESULTS A. The h i e r a r c h i e s B. Success i n gaining t e r r i t o r i e s i n r e l a t i o n t o h i e r a r c h y p o s i t i o n ..... DISCUSSION a. The functional significance of dominance h i e r a r c h i e s .............. B.  Factors  determining  1 2 3 M 5 6 7 8 9  67  68  72 75 80 8  0  88 94 94  position i n 95  Outcomes o f t h e h i e r a r c h i e s  102  LITERATURE CITED APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX  63  dominance h i e r a r c h y  the C.  59  72  PART B"... DOMINANCE HIERARCHIES  IV  55 57  106  .-. .,  .  118 120 120 123 124 126 127 128 129  vi  LIST OP TABLES TABLE 1. fall  PAGE  D i s p e r s a l d i s t a n c e s o f j u v e n i l e s from 1972 ,  site  2. R e s u l t s of removal experiments: numbers t e r r i t o r i e s , o f both mated and unmated males  of  birth: 8  of  occupied  3. R e s u l t s o f removal experiments: d i f f e r e n c e s i n numbers o c c u p i e d t e r r i t o r i e s o f mated and unmated males 4. R e s u l t s o f r e m o v a l experiments: mean t e r r i t o r y s i z e . Removal Area • 5. R e s u l t s o f removal* experiments: mean t e r r i t o r y S p r i n g Removal A r e a ...................................  34  of 36 Fall 38  size. 39  6. R e s u l t s o f r e m o v a l experiments: breeding d e n s i t y ....  43  7. D i f f e r e n c e s in males: s p r i n g 1973.  46  t e r r i t o r y s i z e between mated and unmated Simultaneous Removal areas ........  8 . V e s t i n g d a t a f o r t h e C o n t r o l s and the E x p e r i m e n t a l 1973 9. Number o f j u v e n i l e s banded, f a l l  areas:  1972 t o e a r l y s p r i n g  1973  10. L i n e a g e : t e r r i t o r y s i z e s from one g e n e r a t i o n t o the next 11. Numbers o f e n c o u n t e r s (displacements, avoidances f i g h t s ) i n f o u r dominance h i e r a r c h i e s .................  and  12. Analysis Hierarchy  B-C  of  the  avoidance  interactions  in  the  54 ^  84  13. Rank i n h i e r a r c h i e s and sex o f Song Sparrows i n t e r g r o u p movements  performing 87  14. A n a l y s i s o f f i g h t s i n the B-C H i e r a r c h y 15. Change i n s t a t u s of a j u v e n i l e female d i s a p p e a r a n c e o f an a d u l t t e r r i t o r i a l female  49  89 following  16. Feeding o r d e r o f h i e r a r c h y : appearance o f f i r s t f e e d i n g s t a t i o n s o f the B-C H i e r a r c h y  the  92  b i r d at 1 0 4  vii  L I S T OF  FIGURES  ,  FIGURE 1. The s t u d y a r e a : Mestham I s l a n d  Reifel  Island  PAGE and  an  adjacent  part  of 4  2. Numbers o f j u v e n i l e Song S p a r r o w s r e s i g h t e d on t h e S p r i n g R e m o v a l A r e a f r o m mid-December t o t h e end o f May . . . . . . 10 3.  Singlengths  o f male and  12  f e m a l e Song S p a r r o w s . . . . . . . .  «». Mean w e i g h t s o f male and months  female  Song  Sparrows  for  a l l 13  5. Fall Removal A r e a : t h e movement and e x p a n s i o n o f C o n t r o l male C6 a f t e r t h e r e m o v a l e x p e r i m e n t and d u r i n g the subsequent winter 41 6. Spring boundaries 7.  The  B-C  Removal b e f o r e and  Area: Simultaneous a f t e r removals  Removal  area 65  Hierarchy  81  8. Number o f i n t e r a c t i o n s v e r s u s distance f o r a l l j u v e n i l e s i n t h e B-C H i e r a r c h y 9. P o s i t i o n i n h i e r a r c h y and Hierarchy  within  hierarchy  number o f i n t e r a c t i o n s :  89 the  B-C 103  1 0 . B u l l d o z e d Area: the arrangement of territory boundaries b e f o r e t h e dyke r e b u i l d i n g o c c u r r e d ................... 11*1 11.  Cottonwood  Stretch Hierarchy  12.  Sedge P a t c h  13.  Bulldozed  14.  First  15.  No  Hierarchy  Path Hierarchy  B i g A l d e r and  Marsh N a r r o w s H i e r a r c h i e s  Name C l e a r i n g H i e r a r c h y  1*9 1£° 131 132 1£3  . viii  aCKNOWLEDGEMEHTS  I for  am m o s t  h i sadvice,  work. read  Dr.  I  in are  C.  J.  a t the start  should  who g a v e  nest-finding.  assistance  and Dr.  and a d v i c e  Dr.  J.  R.  a t a l l stages N.  R.  was r e c e i v e d  Liley from  Krebs, of  this  critically  Dr.  J.  F.  o f the study.  t o thank during  P.  the  Taitt removal  who a s s i s t e d i n t h e b a n d i n g  Their D.  help  due  to  with  t h e computer  Finally, patience  Krebs  assistance  a l l the people  supervisor.  and guidance  further like  valuable  also  t o my  criticism  the manuscript,  Bendell  and  grateful  I should  and t o l e r a n c e  was  truly  Lauriente,  like  S.  a n d W.  Zales,  experiments, operations and  appreciated. Borden  a n d B.  Thanks Webb f o r  programming.  t o thank  t h e study  my  would  wife, n o t have  without been  whose  possible.  1  GENERAL  1.  Aims  of  The  purpose  ecological HelosjDiza of  the  first  of with  density  and  1970;  are  evidence some  settling of  study,  settlement test  this  Tompa  and  Sparrows,  classical  examples  and  the  1964).  and  some  Song  i s divided  has  been  dominance  This  into the  shown  territories occupied  patterns, hypothesis.  perplexing in  given  territory  any  I  in a  of  mean area  environmental  Krebs  then,  A  variability  density,  (see  territories  the  the  supporting  on  in  territory  1943;  1972).  influence  evidence  investigate  two  territoriality,  several  conceivably  show  in several populations  been  population  There  the  behaviour  Klomp  has  to  behaviour  Sparrows  both,  is  two  study  sections,  second  dealing  hierarchies.  breeding  however,  Song  aspects dealing  study  social  1937,  Territorial  hence  of  (Nice  dominance  Moss  this  interaction:  considers  with  of  melodia.  hierarachy  the  Study  aspects  social  INTRODUCTION  of  them  1971)  given  Watson  problem  in  birds,  territory from  year  and  size, to  year.  that  could  the  produced successive,  the  There  pattern  i n determining In  and  but  temporal  area.  experimentally simultaneous  (see  unequivocal.  the  the  animals  annually,  i s not  i s important in a  determine  resources  size  that  to  the  first two  is of  number part  of  types  of  i n order  to  2  In  the  second  hierarchies  in  congregated  over  area.  The  the  hierarchies  were  hierarchies  yearlings  of  but the  on  from  Song  1964).  that  Sparrows The  provided  a  a  of  surplus  fall  of  test  and  that  doing  so  process  to  the  i t  by  the  spring  classical  independant  for  the  was  study  is  hierarchies  were  the  outcomes spring In  the  of  the  when  the  particular,  opportunity,  the  spring  hierarchies  the  there  I the  are  the  established  investigation  existed  worth  birds  on  of  of  the  of  breeding, of  descriptive  another  population  (Tompa  on, there  both  behaviour  Columbia  that  of  surplus  good  instigated  (and  1962, indeed  would  be  such the  study  area  from  that  the  dominance  1973.  mentioning  territoriality  features  in  British  assumption  of  capable  is  operating  was  a  territorial There  southwestern  for),  in  that  the  physiologically  non-territorial  Finally,  be  was  present  1972  hierarchy  in  Sparrows  study  in  dominance  dominance  the  the  given  individuals.  this  of  at  season.  birds,  territorial  evidence  the  Song  localities  and  that,  pre-requisite  problems  looked  disintegrate.  pre-breeding  prevented  certain  territories  in  I  juvenile  during  to  obtain  non-territorial  of  winter,  followed  necessary  study,  composition  the  individuals  these  at  hypothesis  that  the  A  the  the  groups  and  began  investigated  of  winter  structure over  during  loose  the  determined  dominant  part  a  bird's  are  not  social  considered behaviour  to  (Brown  3  1963;  Davis  1959).  major  parts,  with  not  be m u t u a l l y  2.  The Study  The  the  part  north  west  by  Westham to  of  part  Topography f o r most  into  parts  two need  1).  Island  The  of  the  a n d an  area  i s  in  and i s bounded  to  arm o f F r a s e r  River,  t o the  C.  Eeifel  George  by a g r i c u l t u r a l f a r m l a n d  i s flat,  t h e water  o f the year,  and  various  the  with early  in fall  and  i s tidal,  hedgerows.  vegetation  table  of  i s close  and t h e l a n d  i s  of  o f woody Appendix  species. i s  and  herbaceous  1 gives  a  list  The s t r u c t u r e  similar  along  and  a l l the  identical.  t h e dykes, Strait  species  and i m p o r t a n t  but i s not  varies  sloughs  two  Columbia,  to the south  grown t o form  Georgia  levels  the  Reifel  (Figure  British  and  the dykes,  Beyond  spring  and  m o r e common  hedgerows,  level  i s divided  dyked.  have  the  on  by t h e s o u t h  Strait  surface  composition  and  east  Island.  Along  of  that  out  Island  of Delta,  Refuge,  necessity  carried  o f Westham  and  the s o i l  plants  the understanding  was  Georgia  Waterfowl  thesis  Area  municipality  the  this  exclusive.  study  adjacent  Therefore,  t h e marshes  are  essentially  the season: summer winter. rising  bordering  river  the Fraser  freshwater.  The  l e v e l s are high  the  to f a i r l y  low  the  marshes  and  daily.  In  but decrease  The  in  and f a l l i n g  twice  water  during  run-off,  water  River  summer.  4  FRASER  RIVER N  South Jetty  FALL  REMOVAL AREA  SPRING REMOVAL AREA  Figure 1  The Spudy A r e a : R e i f e l I s l a n d and an a d j a c e n t p a r t o f Westham Island  5  low  tides  The partly  occur  land  as  in daylight,  enclosed  pasture  The  crops  grown  to  season,  the  for cattle,  on  and  by  the  i n winter  dykes and  on  year  to  study  as  land  year,  night.  the  partly  agricultural  from  at  area  was  used  agricultural  changed  owing  to  from  the  land. season  rotation  of  crops.  The  territorial  vegetation marshes  at  along low  fall  and  water  levels  fields,  3.  the  tide.  winter were  either  Life  Song  Sparrows  hedgerows, The  foraged low.  No  pasture  or  were  found  occasionally  non-territorial extensively Song  at  any  in  foraging  juveniles  i n the  Sparrow  crops,  mainly  was  marshes  seen  to  the  in  the  in  the  when  the  use  the  time.  History  Song  Sparrows  have  a  wide  breeding  range  in  North  o  America, 1963)  and  the  suggests  a  environmental Beer  et  (1937),  a l .  very  variety  Nice  (1943)  attachment  of  the  high  (1956),  of  these lower  more  The  Phillips  (1956a,b),  habitat  and  of  plasicity  conditions.  Johnston  wide  In  recognition  types  Tompa birds  mainland  of  30  the  studies and  and  the of  show  a of  (Dickerman  s p e c i e s to Song  Dickerman  Marshall  site  races  on  for breeding,  (1964) to  than  Sparrows  (1957),  (1948a b) #  and  varying  the  Nice  reveal  studies  highly  by  a of  developed  birth.  British  Columbia,  the  Song  6  Sparrows Hi  I i  are  ™2££^BS#  Lowlands, and east,  and study  The in  by  however,  by  Nice  flocking  Nice,  behaviour  Sparrows  on  during  winter  i n c e p t i o n and bird;  place  probably  in  Weather snowfall  biotic  August  data  are  occurred  The  patterns  Sound  areas  to  the  (Munro  north  and  caught  in  Jjorphna.  has  birds  i n many  therefore a  Puget  A l l birds  Sparrows  1943).  the  occur  nu  Song  concerned,  on  to  discussed  Heifel  respects  detailed  relevant  been  Island  to  those  description  this  study  (Nice  1941).  is  are,  here.  have on  in  Type  Heifel the winter  months,  described  and  given  lasted  September  of 40 (cf.  in  Appendix  in late  December,  Tompa  whole  the  on  of  their to  Reifel  the for  Island  territorial  inclement  weather  post-breeding  moult.  50  Nice 2;  The  (1964)  year,  moulting to  within  contrast  birds  periods  during time  in by  The  the  during  moult  remained  round,  Island.  length of  and  territories  Island  f o r almost  only  the  A  year  Mandarte  territorially ceasing  to  Forest  Certain aspects  behaviour  The  Coast  of  and  boundaries  the  in  subspecies,  behaviour  holders  territory  resident  subspecies  (1937,  Sparrows  territory  behaved  the  discussed  Song  is  Other  behaviour  identical  unnecessary.  Song  of  general  described  subspecies  morphologically distinct.  were  detail  showed  I s l a n d s , and  1947).  are  The  Oberholser,  Gulf  Cowan,  this  non-migratory.  varied  from  bird  days,  and  took  1937; the  otherwise  Tompa  only the  v  1964).  sizeable winter  was  7  relatively  mild.  Territory and one  were pair  boundaries  linearly would  have  From  observations,  males  into  and  each  arranged  there  Island  along  i n general  others  becoming  Sparrows  perform  end  of  the  were  banded  long  were  well-defined,  the hedgerows.  only  was  two  little  territories;  independent  Hence,  neighbouring  were  any  pairs.  t r e s p a s s i n g by  boundaries  this  of  their  e x p l o r a t o r y movements  post-juvenile shortly  distances  resighted  adjacent  recognized  after  were  moult.  leaving  (Torapa  In of  on  most  parent's  Song  until young  suggested  fact,  their  young  1964),  Observations the nest  covered.  within the boundaries  parents,  the that  that  young  no were  territory  at  time.  In become  the f a l l ,  after  territorially  dispersal  of  the  individuals  whose  for  a  on  discussion  Mandarte  Island,  coincided  with  females, of  the post-breeding active  young.  by  some  Reifel  respected.  On  some  on  rise  to neighbouring dispersal  evidence  to suggest  from  and  the study  more,  1 gives  birthplace  the emigration  the  emigrate  Table  longthe  once  was  and  the  the  this  adults  results  distances  known  in of  fall  ( s e e Howard  territorial  many  young,  50%  On  Reifel  Island,  d i s t a n c e s were most  area.  of the  quite  of  1960,  For example,  On  behaviour which  large,  juveniles  in  covered  short-distance dispersers).  islands.  that  moult,  were  although there i s did  o u t o f 52  not  banded  8  TABLE 1  Dispersal distances of juveniles from s i t e of b i r t h ;  Numbers of juveniles  Distance (m)  4  0-150  2  150 - 500  1  500 - 750  1  750 - 1000  2  1000 - 1500  1  1500 - 2000  f a l l 1972  9  juvenile Area  during  October within to  Song  and  Sparrows August  areas  juvenile  amongst  young  individuals  in  favoured  centres  of  hierarchical  In  as  yearlings spring,  to  Sparrow  Waterfowl  groups  ( s e e PART  Island  and  only  one  two  areas.  Song  April.  ones  from  and were  banded  seemed  Sparrows Figure  the  on  bird  t o be,  Tompa  their  on  2 shows  any  fall  banded  of concentration hedgerows. the  groups began hostility  1964:26). began  May,  the  hedgerows  at  These winter  as  study  then,  a sharp  area  no  in  t h e numbers  of  3)  area.  was  the  in  the the  early  The  April  Reifel  the study and  area  April  located  decline late  between  non-territorial  i n March  (see Appendix  up,  exclude From  adjoining  to break  Also,  to  1973,  surveyed,  the study  i n the  non-territorial  territories.  i n late  also  activity,  B).  birds  recorded  as  throughout  increasing  (see  of the study  Westham  of  of  visits  in  present.  the  persisted  territorial  was  along  Removal  resighted  to reveal  centres  the h i e r a r c h i c a l  result  Refuge  There  formed  localities  themselves  the end  Song  remained  birds  non-territorial  failed  birds  fall  Fall  were  occasional  area  that  the  of t e r r i t o r i a l  Sparrows  young  a  the  t h e peak  Further,  on  19%  t h e many S o n g  the spring,  possibly  present  September,  the study  the  certain  were  after  area.  outside  The  and  November,  the study  that  March  juvenile  the  numbers  and  Song  1973,  in  i n the  on  early  Sparrows  10  Each month i s d i v i d e d i n t o two two-weekly time i n t e r v a l s  Figure 2  Numbers o f j u v e n i l e Song Sparrows r e s i g h t e d on t h e S p r i n g A r e a from mid-December  to the end o f May  Removal  11  resighted of  Hay,  two-weekly  showing  spring. in  in  Tompa  Song  a  sharp  (1964)  Sparrows,  and  been  found  in  1963;  Dhondt  1971b;  1967;  Watson  left  on  the  4.  Methods  The  of  the  banded  area  of  each  a  282 few  the  Each tarsus,  and  each was  banded  Sparrows  weighed,  culmen.  and  Males  winglengths  f o r the  Although  to  72  males  throughout  the  on  1972  mm  to  one  (Figure  Island  Carrick  Song  study May  have  1961;  Smith Sparrows pairs.  area  1973.  necessary  was  Because  to  have  a  recognizable.  aluminium  and  During  mainly  in  for length  females  can  winglength banded  one  the  or  study  mistnets,  1937).  from  more was  of  sexed  birds;  females  there  be  (Nice  slightly  4),  the  traps.  measured  weighed  in  territoial  the  caught,  in winglength,  average year  on  (e.g.  only  the  end  in spring  individually  with  and  3  67  on  i n p o r t a b l e sparrow  was  the  i t was  were  and  from  Orians  bird  differences  the  a l ,  February  behavioural gives  birds  apparently,  the  Mandarte  species of 1963;  to  seen  i n numbers  population  from  on  i n various combinations.  Song  bird  same r e s u l t  experiments,  bird  caught  et  numbers  i n m i d - A p r i l were  Sparrow  c o l o u r e d bands,  period, with  declines  population,  Therefore, two  sharp  other  mid-December  i n the  the  Thus,  continually  from  found  Jenkins  1965).  study  nature  decline  several  Song  observed  intervals  too  males 63  to  than much  wing, both  on  Figure ranged 68  mm.  females overlap  NUMBERS  Figure 3  W i n g l e n g t h s o f male and female Song  Sparrows  13  M E A N WEIGHTS (gm)  4  Figure 4  Mean w e i g h t s o f male and female Song Sparrows f o r a l l months  14  between be  'light*  useful  of  area  Song  than  and culmen was u s e d  Juveniles fall  plumage feature  i s that  retained  colour  (Nice of  yellow  in  decreased proved  identified  their  the  dusty,  the  1937).  fleshy  colour.  i n size,  have  a t t h e base  parts  In  juveniles  ,  plumage  males  I also  assume a  which a r e  have  used  rounded was  the  of the mandibles.  the  and a r e fleshy  colouration  c h a r a c t e r i s t i c when  in  distinguishing  are extensive,  but the yellow  t o be an o b v i o u s  which  other  area.  retrices,  adult  the  subspecies  and then  useful  pointed  whereas  case  no s u b s p e c i e s  moult, A  A criterion parts  f o r other  finely-streaked  adults.  the winter,  in  on t h e s t u d y  post-juvenile  a l l juveniles  these  taken  earlier,  the fleshy  fledgelings,  were  As m e n t i o n e d  to  through  retrices  of sex.  ground  loose  similar  f o r this characteristic to  measurements  o n e was  after  females  as a wintering  Sparrows.  the resident  late  'heavy'  i n determination  Tarsus study  and  bright  parts  have  i s retained,  the  In  bird  was  and in  fe"  the in  hand.  Certain  1972) r e t a i n e d  The  fleshy  dark  grey.  response Song  of  Sparrow's  described  the yellow  parts  Territory  in  a  yearlings  (possibly colouration  a t t h e base  were  territorial  male  on a t a p e  Dhondt  1966).  into  of late  broods  February  1973.  o f the mandibles  boundaries!  song  members  determined  i n adults are  by  noting  to the playback  recorder This  (Sony  method  the  of a  stranger  110A)  (method  proved  very  useful.  15  Males the a  would  tape  territory boundary  any  from  various  clearly  and  sing.  of  of by  the  usually  this  song  or by  had  f l y to  a  song  of  perch  along  the  particularly  as  Thus,  the  relatively  needed.  cross  be  neighbours,  not  recorder  could  two  was  to  elicit  not  boundaries  time,  in  tape  would  only  size  close  I could  the  Males  territory  territory (1960)  flying  playing  hedgerow.  way,  by  display.  short space  hedgerow.  Suthers  birds  would  very  new  sing  the  but  In  measuring  the  either  along  defined i n a  linearity  proposed  spots  pair  to  territorial  boundaries,  one  method  quickly  r e c o r d e r , and  response  at  react  Song  owing  to  the  complicated Sparrows  as  16  PART  A  .THE  EFFECT  OT  TEMPORAL  SETTLING  TERRITORY  I.  breeding  of  Brown  1970a;  SIZE  Klomp  five  in  population  .  far fully  removed  means  by  follows.  and  to  then  The a  population  in  limited and  they  1  by  feel  need  show  no  a  study  in  resident territory  replacement, (a)  behind  and a  i f (b)  any, in  removal  and  to  date  holders  8  experiment  new  be  breeding  occurs,  Appendix  list  to  i f territorial individuals removed,  other  (1970a)  limits  which  animals,  some  behaviour that  on  populations  which  Moss  Moss  opinion  conditions.  area, are  to  of  g.  Chitty  and  a l l five  conditions rationale  way  Watson  that  lines  the  e.  1966;  breeding  the  which  proceed  satisfied  given  limit  been  years,  Watson  main  been  out.  show  subsequent  which  In  have  8)  experiments,  determined.  stable  order  two  determining has  Rowan  1969;  i t i s simply  (Appendix  animals,  1966;  Lucas are  in  recent  Lack  i t does  themselves  They  Removal  (b)  populations  space  satisfied  (a)  of in  1965;  There  behaviour;  conditions  so  Crook  F r e t w e l l and  v e r t e b r a t e s , or  factor,  i t s importance  populations  1972).  whose b r e e d i n g  and  discussion  1962;  1969;  territorial  are  in  much  Wynne-Edwards 1967;  behaviour,  density  subject  has  ON  INTRODUCTION  Territorial  of  PATTERNS  are  can  be  is  as  from  a  individuals  I  17  subsequently  replace  replacements  had been  in  that  prior  area  on  by t h e p r e s e n c e  for  presence that  breeding,  removal  of species;  1964),  i n fish Healey  1961;  Watson  Elliot  1967; H a r r i s  i n  holders  of  that  these  territories individuals,  a  territory deduced  limits  breeding  been  carried  i s that  density  territory between  1968),  there  one  year  and  1955;  Moore  (e. g.  particularly and Cope  Lockie  in  birds  1951;  Orians  and J e n k i n s  1968; B e n d e l l  1970; Young  1970; K r e b s  i f  of t e r r i t o r i a l minimum  which within  a  between territory  This  variability  size  i s however  associated species:  would  be  and  1971;  not so.  with  and  within  a  habitat  in territory  size  in  are differences  territory  (c) t h e r e  and  There a r e  variations  (a) t h e r e  i n t h e mean  easier  were c o n s t a n t ,  one p a r t i c u l a r p o p u l a t i o n  habitats; size  behaviour  territory  are  are differences  This  Jacobs  i n mammals  1951; H e n s l e y  1970; Holmes  individuals within  (b)  mean  size  (e.g.  o u t on a  1972).  importance  problems  have  Clarice 1970),  for a l l individuals.  years.  on  i t c a n be f u r t h e r  1965, 1967; Watson  demonstrate  three  holding  insects  and A l d r i c h  1971; Young  The  the  then  1967; Smyth  Stewart  equal  settling  of the t e r r i t o r i a l  experiments  (e.g.  (e.g.  to  from  the  of the territory  variety  Peek  If  i t c a n be d e d u c e d  area.  Such  1966;  then  prevented  to the removals.  necessary the  them,  and  size  year; within  are differences i n over  successive  has prompted  Lack  18  (1966)  to reject  the  regulates  breeding  territories  have  in  limiting  major,  density.  He  minimum  the breeding Wood,  that to  minimum  year  size,  since  year  territory  territorial pointed  population  Oxford,  c o n s i d e r a b l y from  argues year  that  a fixed  i n Barley  varied  hypothesis  this  for territorial  out  that  not  operated  of the Great the breeding  t o year. size  has  However,  need  n o t be  behaviour  behaviour  Tits,  i f  Parus  density Krebs  has  (1971)  constant  from  t o determine  breeding  a possible  mechanism  density,  My causing year  study  was i n t e n d e d  the differences  within  considering  an now  to investigate  i n mean  area  territory  (problem  some f a c t o r s  (c)  which  taken  from  year  stressed  that  the  possible factors  Further,  mechanisms my  differences  1.  Food  approach between  su££ly  Territory if  size  of  availability  only,  t o the study  individuals  and t e r r i t o r y  size  could  territory of  year.  not  food.  above).  I t  year i s  It  should  was  worth  however  t o be d i s c u s s e d  to  to  influence the size  ultimate  (a)  of be  refer  to  mechanisms.  not intended  (problem  to e x p l a i n  above).  size  fluctuate was  from  could  territory  proximate  to  size  annually  proximately The  evidence  with  food  adjusted here  i s  supply, to  the  far  from  unequivocal.  Differences  i n mean  territory  size  in  various  habitats  19  have  been  related  some  instances,  in  three  Tinbergen 1960),  species  and  and  ,  Mason  and  numbers there other  of  other,  a  one  that  over  Oyster-Catcher,  from  taken to  defend,  Orians  size  a  territory, confines, factor  by  the  a  1969),  and  the  Egglishaw  times  species  biomass  of  richer  Turnstone,  1968).  Oncorhynchus  Dunlin,  Laqppus l - _  three  (Smith  (Glas  the  Grouse,  biomass  was  In  higher  kisutch,  where  on  salmonids food  of  fish,  and  (1967),  in  their  suggests  which  the  of  years  Harris  a  a  the  in  two  than  the  territory  size  and  general  Blackbird,  that  territory  food  i s related  (1970)  compromise  proportion  mean  ostralegus,  young,  (1961)found  the  in  Haematopus  probably  feed  and  number  conclusive.  Red-winged  but  Red  and  coelebs, in  in  indirectly  (Kluyver  food)  greater  differences  for  needed  the  1973),  a  spp.  Moss  supply.  three  i s far  of  1966;  that  supply  birds  a l .  showed  food  population  territory  the  g.  (and,  differed.  Evidence  the  ,  e.  Fringilla  measuring  1970)  quantity  species,  Tamiasciurus,  (1965)  found  streams,  a  et  genus  greater  hardly  within  (by  food  Parus  j u v e n i l e Coho Salmon,  hand,  Scottish  Chaffinch,  (Settleship  the  Chapman  was  the  (Miller  of  various  titmice,  (Holmes  M § £ § l i l lBf®I£I®s, squirrels  in  of  directly  alpina,  Scoticus  differences in  quality)  1953),  Calidris  to  of  is  size the  area  A ^ e l a i us  not  regulated  that, of  the  area  the  they between  can the  Eljoe n i c e us,  obtained  i s probably  size  the  correlation  food  food  suggested  between the  to  the  within i t s proximate and  that  20  territorial  behaviour  There  i s some  area  can  1958),  evidence  determine  Ovenbird,  Se^urus  although  experience  was  Evidence can  be  increase result  this  that  food  increased  Salmon,  demonstrate differing  in  Salmo a  laboratory  availability  the  territories  were  non-territorial).  that  fish  receiving reverse whist  deprived an to  be  superabundance  determine in  of  true (1962) of  latioes.  ways  food of in  Brook  found  If food  for  the  caused  size  (the  then  species.  not  1971)  that  fry  (1962)  showed  than  those  (1970)  found  in  do  the  ineonstans,  absence  work  they  the  subordinate  complete  aggression  in  of  S t i c k l e b a c k , Culaea  aggression  might  sizes  aggressive Smith  the  could  the  Yamamoto  whereas  in fish,  different  and  that  (Symons in  the  their  territory he  changes  size  of  deprivation  feeding  salmon  a  or  causes  suggested  food  more  reduced and  supply sizes  experiment  that  i f age  territory  However,  were food,  food  the  with  young  the  size.  the  Keenleyside  food  territory  different  food  dominant  abundance  Bagnusson  Ory.zias  of  of  known  influence  salar.  (Stenger  not  {1968)  in  given by  i t  is  a  taken year  change  size  in  one  of  associated  food  important.  within  can  Symons  of  more  territory  territory  to  fish,  i n aggression  Atlantic  #  i f manipulation  In  an  of  study  supply  i s much  quantity  auroca£illus  individuals  in  the  way  size  influencing  territories.  some  that the  in  obtained  territorial  in  the  or  Medaka,  together so  a  to  apparently  21  Bustard  (1969)  gekkonid  lizard,  vary  differing  with  social (1970a) and  not  cracks  used  the  territory  birds  were  supply,  or  i f  young,  whilst  production food,  in a  of  new  from  young  however,  1966),  in  the  not that  Watson  Moss  and  environments,  a l l the  suitable  i t  is  to  year,  not  found  to  size  that  which  not  the  et  mean  that  i f the  In  food  number  of  food  Peromyscus supported  survival population food.  d e c l i n e of  the  settled.  food  additional the  was  influence of  excess the  to  the  uneguivocal.  found  prevent  the  known  in  a l l of  for  improved  with  not  territory  year  evidence  showed  Pitelka  between  of  S t e r c p r a r i u s £omarinus,  difference  (1970)  winter.  relationship  in  availability  spring  a  and  the  the  in  Jaeger,  i s also  fluctuations  (1971)  previous  but  increased  failed  did  a  concluded  Krebs  was  the  area  to  and  size  inverse  (1959)  Fordham  stump  the  because  related  adjusting their  Bendell  as  been  lemmings,  density  but,  year-to-year  Pomarine  mammals,  population  fiSSicuAatus,  an  there  breeders  small  population  supply  i n the  breeding  on  He  the  territory  suggested  of  limiting,  (Lack  food  limit,  a  of  lizards.  not  that  density  be  Tit,  have  size  potential  the  young  the  (1955)  by  Great  size  to  could  upper  food.  p h y s i c a l s t r u c t u r e of  out,  f o r the  In  of  females  inhabiting  the  were  the  supplies  of  point  experimentally  and  variegata,  number  an  behaviour,  related  the  set  territory  al.  Gehyjra  that  behaviour  In  food  showed  a  a  of  the  size  and  Excess  population  22  of  Microtus californicus  and  o f Mus  musculus  Finally, appears within  i n  attributable nutritional Watson  2.  Other  levels  are  which  of  the  some  (1968),  (1967),  Zimmerman  (1970)  felt  tadorna, area,  was  influenced  (1961),  lag.  of  heather  not  This  directly  has  been  through  and  Moss  high  1970b;  factors size  been  (1956),  other  from  suggested  and  Snow  e.g.  Lin  (1958),  (1963),  than  year  of territories  (1961),  food  to year. as  being  i n a  given  D i n g l e and  (1963),  Davies  and  a n d Snow  Watson  (1964),  Snyder  and T e s t e r and Watson  (1973).  Young  territory  that  size  invertebrates  Calhoun  (1971),  the shape,  Warblers,  both  determined  suggested  has  t h e numbers  Dixon  and by t h e l i m i t s  (1972)  Reed  that  taken,  and t e r r i t o r y  territory  Alexander  e.g.  Buechner  quality  (Watson  environmental  animals,  (1965),  year  chicks  habitat  area  vertebrates  the  a g g r e s s i v e n e s s working  influence  i n influencing  Caldwell  one  1966),  1970).  of t e r r i t o r y  factors  important in  a  the  several  of  1965; K r e b s  1971).  might  nature  Grouse,  the size  to decreased  and Belong  1 9 6 7 ; Newsome  Red  environmental  There  The  the  but with  and M i l l e r  supply  (Delong  to influence one y e a r ,  (Krebs  size by  in  topography  Tadorna  topography  o f t h e owner.  but not the size,  AcrocePhalus  Shelduck,  t h e immediate  of tolerance  the l o c a l  the  of the  Catchpole  o f reedbeds  greatly  of the territories  scirp_aceous, and  Sedge  of  Warblers,  23  schoenobagnus. and  agonistic  spacing  for  a  can  Tit  (Lack  LY.£2.1§]J£I»  (  study  to  nestsites  has  Sparrow  is  the  in  of  a  of  1971;  1961;  Zaret  passerine Yeaton  a  shelter for  pomacentrid  the  fish,  of  the  supply  shown  in  (refs.  in  habitat  or  the  of type  of  Great  Ficedula 1972).  Tompa a  1961)  shortage  territory of  nest  the  Sjostrand  size  of  and  a  population  Flycatcher,  and  the  factor  has  and  Band  comm.)  for  some  larger  territories  an  and  There effect  of  taken  habitat  hence  that  Brown  on  limiting Song  territory  the  density  and  a  Sheppard Song that  resource Sparrows  size,  that  of  more  in a  given  be  (e.g. Ayala  1971).  the  the  another  1966;  Sparrows,  will  the  vertebrates  Michielson  1971;  hypothesised  the  influence  i s some e v i d e n c e  1966;  particularly  of  could  invertebrates  Greenberg  1971;  and  which  animal,  both  competing the  of  the  occupy).  Inger  (pers.  Sparrows  competition.  birds,  Pied  Song  obviously,  species  and  the  limit  been  Enemar  nature  territorial  one  has  1956;  possible  populations  Connell  of  birds,  This  i n f l u e n c e on  will  interspecific  density  that  responsible  n e s t s i t e s can  in  on  limits,  further  density  and  date  any  the  of  Haartman  V o n  that  A  heads  density.  1966),  suggests  Song  suggested  factors  hole-nesting  increase  (within  (1972)  were  coral  shortage  example,  boxes  No  over  Sale  aruanus.  Further, of,  behaviour  patterns  Dascyllus  Finally,  8.  For I.  species area,  the  ('horizontal'  24  rather  than  'vertical*).  circumstantial conducted  to  Krebs in  date  which  each  method  minimum  at  and  territories  can  Temporal  In  this  that  temporal  influence on  birds  and  Van  the  males  introduced  theoretically density,  some be  and  methods  been  possible discusses in  which  related directly  methods  settling  deals  pattern  on  to  the  with  of  the  settling  on  territories  densities in degrees limit  a  there  Assem  of  different settling  set  to  the  v a r i a b l e one. of  in  on  supporting  years  could  those  years.  number  For  settling  i s some  (1967)  gasterosteus  considerably territorial  have  on  of  the  settled  hypothesis  territories evidence  in  can work  fish.  den  stickleback,  of  be  not  these  patterns  density,  is built  density.  upper  can  some  temporal  different  owners  however,  experiments  considers  of  the  an  no  influence  need  breeding  of  way,  territory  could He  one  patterns  result  and  presented  influence  Fluctuating the  has  size  that  hypothesis,  test i t .  length.  territory  hypothesis  be  to  territory  environment,  3.  evidence,  (1971:17)  ways  His  with males  were  the  found  that  aculeatus, pattern  accomodated  introduced  successively.  in  of a  the  three-spined  territory  sizes  settling.  The  tank  simultaneously He  in  was as  interpreted  nearly when the  varied  number  of  double  when  they  were  results  as  25  suggesting in  the  male  settling males  that  the relative sticklebacks  and b u i l d i n g  were  Turdus  (1965)  population number compared  the  was r e d u c e d  of  b y 55%  200  the pattern of s e t t l i n g  year,  Great  and,  compressibility February more Krebs mean  showed February  and  Sedge  conclusive  adults  a  might  he o b s e r v e d He  that  evidence  size.  with the  Tits  occurs  influences  birds  in the warm  to  might  settle  increase.  correlation  breeding  between  density.  patterns  of  f o r the variations in  i n a three  admits,  244  consistent  temporal  account  to  forward  year  was a p o s i t i v e  suggested  was 93 a s  put  density  and s u b s e q u e n t  The  consistently  and f i r s t  the breeding  there  Warblers.  has  settling and  stable  rose  territory  least  of  a  as suggestive  (1971)  at  i n  but  synchrony  then  (1972)  evidence  influenced  Blackbird,  snowstorm  by G r e a t  both  size  the  i s taken  i s  territories  territory  found i f  snowstorm.  on t e r r i t o r i e s  temperature  on  after  territories,  that  Catchpole  mean  i f  synchronously,  settling  that  of  causes  a severe  T i t , Krebs  Settling  February,  European  territories  after  and t h i s  evidence  hypothesis.  of  the  i n the previous year,  that  suggestive  for  held  following  the  levels  i n the process of  t e m p o r a r i l y s m a l l e r than  that  the  For  a l l o f them  number  territories  t o about  i n aggressive  successively.  relates  merula,  -  - were  introduced  Simms  differences  year  however,  study to  to substantiate the hypothesis.  of  having  Reed no  26  Possibly in  birds  i s from  1971). the  The  study  year  caused  by  that  and  these  the  and  number  of  of  study.  I  the  the  and> t h e  from  number  therefore  of  settling of  territories  breeding  density?  the  birds  Song  catastrophic  peak  active.  for  Sparrows,  and  territorial  a  the  settle,  now  birds  increased.  the  given  had The  fourth  year  investigate  does  hence  amongst After  territorial  high  the  birds.  mortality  experimentally  occupied,  of  to  of  mortality,  the  territories  in  in  territorial  allowed  1964;  years  of  were  territories  two  substantially  established  to  hypothesis (Tompa  first  little  remained  out  on  at  the  Island  territorially  occupied  also  a  March,  spring  not  the  in  for  caused  of  set  question:  number  in  the  were  territories  following pattern  in  resistance  Mandarte  following  snowstorm  that  on  supports  increased  non-territorial  gone,  the  but  apparently  yearlings  best  territories  study  occurred  behaviour,  storm,  of  the  which  Sparrows  constant,  freak  storm  those  Song  of  a  example  number  was  third  The  the  the  the  temporal  area  influence  territory  size,  27  II.  PROCEDURE  In  order  temporal number to  of t e r r i t o r i e s  occurred,  there  of  juveniles holding  the  taking  territory for  territorial  types  •Simultaneous'  empty  then  on  necessary The  i f  came  were  territories for  pairs.  o f Song  birds  holders  i t was  that,  surplus  Sparrows from  preventing the  study i s  area.  i t c a n be f u r t h e r capable  a  As  necessary  of  birds  deduced certain  then  of the resident  loose  i t c a n be  breeding,  behaviour  on t h e  the  Sparrow  were  removal  replacement  Song  juveniles  that  can influence the  a  of  removal  and  t h e removal  females, creating  show  hypothesis  deduced  breeding, limited  the  density.  Two  involved  area,  juveniles,  territory  i f the replacement  breeding  a  non-territorial  pre-requisite that,  i n a given  I f the replacement  from a  on t e r r i t o r i e s  would  existed  the resident  the  of the t e r r i t o r i a l  themselves  area.  groups  experimentally  taken  out removals  experiments  that  test  patterns of settling  carry  study  to  i n  as  an a r e a  of  'Successive'.  simultaneous  Song  Sparrows.  a  space  suitable  territorial  allow  Song  settling  This  A  type  of  birds, time  habitat  some  o f removal  carried  of  both  out:  Removal  males  and  as possible,  thus  which  Sparrows. of  were  Simultaneous  of the t e r r i t o r i a l  short of  experiments  was  completely  This  was i n t e n d e d t o  the  non-territorial  was i n t e n d e d  to simulate  28  a  catastrophic  mortality  but  i t  void  of t e r r i t o r i a l  for  example  same  time  are  could  here  A  when  to a  well  birds  migratory  non-occupied  treated  at  as  regular  situation time  allow  be a p p l i e d  i n an a r e a  of  birds area.  to other  birds,  instances cf a  of suitable  habitat,  return  i n the spring  These  types  involved  intervals,  the periodic  of t e r r i t o r i a l successive  territorial  of  as  at the  occupancy  equivalent.  S u c c e s s i v e Removal  pairs  of  equally  o f non-migratory  the removal  simulating  disappearance  individuals.  settling  of  the over  This  some  of  territorial  more  normal  a longer  period  was  intended  non-territorial  to Song  Sparrows.  Simultaneous the  fall  o f 1972 a n d i n t h e s p r i n g  coincided  with  Sparrows were Each  of  s e t was  such  most  were  removed  successful  a s t r a n g e r Song  they  concerned  Both  responded the  that  either  Sparrow,  to  removal  Two  These  behaviour different  experiments a Control  conducted  area  by m i s t n e t t i n g  and f e m a l e s the of  times  of  Song  hedgerows  was  were  areas.  o r by  tape. females  The  strung over  caught only  1).  sandwiched  shooting.  t o be t h e u s e o f a t a p e d  and a m i s t n e t  in  two  (see F i g u r e  and t h e S u c c e s s i v e Removal  means p r o v e d  males  o f 1973.  1964).  s e t o f removal  arranged  were  of t e r r i t o r i a l  Tompa  the Simultaneous  recorder. as  1943;  f o r each  Birds The  t h e two p e a k s  (Nice  used  between  and S u c c e s s i v e removals  song  t h e tape  i n the mistnets major  i n the f a l l  problem  (see below).  29  Appendix  4 gives  difficulty  The  Fall  possible  i n locating  Removal  This 1972.  some  females  reasons at this  accounting  time.  Area  hedgerow  contained  8 territories  were  used  24  territories  i n the f a l l  f o r the Successive  for  the Control,  a n d 10 f o r t h e t h e S i m u l t a n e o u s  10  territories  on  adult  males from  mates,  that  intrusion on  the Simultaneous  t h e summer,  had  during  the  with  and  established the early  t h e S u c c e s s i v e Removal  mateless,  a  2  fall.  juvenile  Removal.  6 The  contained 8  males,  without  f o r themselves  contained  established  of  Removal,  area  Similarly,  area  territory  Removal  territories  the 8 1  by  territories  juvenile  by i n t r u s i o n  male,  early i n  fall.  9 o u t o f t h e 10 m a l e s o n t h e were  f o r the  removed  removed  i n the period  on O c t o b e r  October  1  to  locating  him c o u l d  completed  the  October.  Also,  possible  8  explanations  12.  11,  This  Simultaneous  October last  4 t o 7; t h e l a s t  male  was  and i t i s p o s s i b l e be  due  to  post-breeding  the moult  on t h e S i m u l t a n e o u s  females, for this  only  4  were  Removal  not  that  male  recorded  was from  the d i f f i c u l t y  fact  that  until  t h e second  Removal  area  he  area,  removed.  had  d i s c r e p a n c y a r e g i v e n i n Appendix  not  week o f  out  Some  in  of  a  possible 4.  30  The from  8  males  October  3  alternately. Out  of  a  Appendix  The  6  after  area day  an  female  and  March  Two  7.  of  taken  to  2.  males and  Successive alternately, apparently  days  locate.  removed  i n the  (see  spring  a  from were  One  the  of  Removal,  Simultaneous  apparently missing  territorial  territory  watching a  from  paired  her mate's male  h i s mate;  i s this  male,  i n the f a l l  disappeared, d i d not appear  from The  as  removed females  over  reveal  disappeared  i n PART  was  to  f o r the Simultaneous  experiment.  hours  length  one  territories  taken  were  had  at  only  four  explanations).  5 females  male  female  difficult  removed  6 f o r the S u c c e s s i v e Removal.  adult  juvenile  6  were  had  failed  were  o f t h r e e and  proved  females,  c o n t a i n e d 19  the  February  females  area  Area  that  territory  and  on of  (six  female  7  territories  yearling  29  Removal  males and  the  at intervals  for possible  f o r the Control,  Removal  27,  Once a g a i n ,  hedgerow  7  7  on  to  again  Spring  1973.  t h e S u c c e s s i v e Removal  possible  4  This  on  a hide  1972  t o have  in this  female).  territory  of  a  a  male's  The  second  between  January  thought  to  have  taken  a  incident  i s c o n s i d e r e d more  B.  4 females  Removal  area,  between missing  were  at intervals  March on  removed  3  the days  and  i n  pairs  o f t h r e e and  23.  Two  from  the  four  days  females  of the experimental  were  removals.  31  One  female  and  was  mate in  The  disappeared  not  seen  before  early  shortly  afterwards.  the  before The  experiments,  the  second  and  removal female  remated  with  a  experiments, deserted  her  Control  male  March.  Controls  Controls undisturbed  i n both  during  the  Spring  and  removal  Fall  Removal  experiments.  Areas  remained  32  III.  RESULTS  1.  Replacement  The  Sparing R e m o v a l  On  the  were  Pairs  Area  Within 9  Thus,  On  Removed  Simultaneous  were removed. settled.  Of  the  17  days,  birds  had  Successive  removed.  Removal 10  replaced  pairs  7  males  Removal  Replacement  area,  males and  the  12  area, 6 settled  7  females  females  had  removed.  males  on  5  and  and  vacant  4  females  territories  I, during  the  t h r e e or  removals.  By  Harch  replaced  the  The  Removal  Fall On  were male  the  10  they  were  were  until  the  On  experiment, the  7,  14  caught  spring  removed.  during  Within  :\  pairs  of  Removal 10  days  settled.  in  after  10  the  Thus,  males  the  successive  10  i n the  fall  of  with  the of  birds  1972,  number  of  4  had  of  females the  Although i t was  replacement  the  and  removal  settled.  not  birds females  ninth  several known i f or  was  not.  not  done  1 female  were  1973.  the  replacement three or  area,  had  S u c c e s s i v e Removal As  between  males  assessment  the  5  intervals  Area  paired  Therefore,  27,  Simultaneous  October  females  day  removed.  removed. on  four  four  area, 8  Successive birds day  males  Removal  settled  intervals  on  and in  the  vacant  between  the  spring  territories successive  33  removals. Once but  By O c t o b e r  again, their  the  After  in  a  settling  both  and  both  the  were  the  was  of  not  1972, known.  deferred  territories  of  i n both  until  slight  and  covered Fall  expansion area  Krebs  showed  who  were  the  Removal occurred  removals,  (Watson 1967;  met  (see  there  was  1971).  the existence prevented  by t h e r e s i d e n t t e r r i t o r y  of  from  holders.  Territories  of the removal  Removal Areas. mated  and  Numbers  unmated, a r e  Removal a r e a s ,  occupied  spring  Successive  Some  experiments  area  both  Spring  the  breeders  Simultaneous  number  only  was  an E x p e r i m e n t a l  before  the r e s u l t s  Fall  males,  and  expansion  Occupied  Spring  Increases On  Of  2 gives  On  that  potential  Table  by  occurred,  a Control  on t h e s t u d y  occupied  in  of  Numbers  therefore  example).  the replacement  surplus  2.  where  preventing  Thus,  had  6 as an  suggesting  pressure  i n the f a l l  males  o f t h e hedgerows i n both  (see F i g u r e  below),  to the t e r r i t o r i a l  resettling  instances  caught  territories.  1973.  length  Areas  were  of t e r r i t o r i a l females  spring of  entire  9 m a l e s had e s t a b l i s h e d  s e v e r a l females  relation  Assessment  30,  territories  and f a l l  Removal changes  there  were  areas in  the  experiments f o r of  territories  given.  was  after  an  the  i n the region  and on  increase removals. of  40%.  the Controls,  there  numbers  of  occupied  34  TABLE 2  Results of removal experiments: numbers of occupied of both mated and unmated males  Spring Removal Area  Before  After  Percentage change i n number of t e r r i t o r i e s  Simultaneous:  7  Successive:  6  -16.7%  Control:  6  +16.7%  10  +42%  F a l l Removal Area Simultaneous:  10  Successive:  8  Control:  14  +40% +12.5%  -16.7%  territories  35  territories  The after  (see  differences  the removal  differences Removal the  below).  experiments  between  areas,  and  Controls,  in  compared.  The  Simultaneous than  i n t h e numbers  the  those  on  the  i n  Spring in  areas  and  Fall  occupied  the  and  3.  The  Successive  Removal  areas  and  Removal  Areas,  are  territories  are s i g n i f i c a n t l y  Successive  before  Table  and  the Simultaneous  differences Removal  a r e compared  Simultaneous  between both  of t e r r i t o r i e s  Removal  on  higher  areas  the  (P<0.05)  and  on  the  Controls.  These more  data  suggest  territories  the simultaneous  taken  than  i f  settling  the  results  settling  in  occurs  successively.  3  •  Size  An  Of  increase  necessarily In  the  before  Territories  implies  present and  Removal  in  after  numbers  that  study, the  Area.  This  boundaries  territories  adjacent  removal  measure  of  experiments  t h e mean the  territory  areas  were  experiments,  territorial  accurate  of t e r r i t o r i e s  to the  was  due  of  those the  size  to  not e x a c t l y  the  the  the  expansion  Control  differences  fixed  males  areas. before  mean t e r r i t o r y  same  Spring of  that Hence  and  area  decreases.  particularly  removal  i s t o compare  in a  the had  a  more  after  the  sizes.  36  TABLE 3  R e s u l t s o f removal experiments: d i f f e r e n c e s i n numbers of  occupied t e r r i t o r i e s  o f b o t h mated and unmated males  Simultaneous  Successive  Spring:  +3  -1  Fall:  +4  +1  t = 3.13  P < 0.05  (one-tail)  Simultaneous  Control  Spring:  +3  +1  Fall:  +4  -1  t = 3.13  P<  0.05  (one-tail)  37  Tables  4  Simuntaneous Controls,  and  the  between  t h e mean  the  Fall  Simultaneous  territory  instance  Removal  sizes  smaller  are  much  There and  areas,  after and  on  Area,  P<0.05  sizes  on t h e  after  differences  the  i n  mean  the  removals  the  C o n t r o l s , b u t i n no  on  both  significant.  support  results  territory  the  the removals  smaller  are  and  differences  and a f t e r  territory  further  mean  the  Area. areas  settling  areas,  T h e mean  i n the  experiments.  Removal  are the differences  simultaneous  removal  before  sizes  areas,  f o r the Spring  before  data  territory  Removal  Removal  before.  Removal  These  with  P<0.01  sizes  mean  Successive  territory  than  Successive  the  s p r i n g and f a l l  Removal  experiments  give  Simultaneous  significant;  for  5 the  f o r both  On  are  and  the  i n more  size,  than  hypothesis  territories i f  the  that  occupied,  settling  i s  successive.  So  f a r , the  territories, the  removal  the  doing that to  and t h e s i z e s experiments.  breeding  males  be  remained so, took  analyses  density unmated  over  considered.  dealt  with  of t e r r i t o r i e s , I t i s necessary,  before during  however, c e r t a i n  place  have  and a f t e r the  before however,  and  spring  of  on t h e F a l l  after  a s some  1973. of  Removal  of  t o compare  the removals,  c h a n g e s i n numbers  the winter  t h e numbers  Before  territories Area  need  38  TABLE 4  R e s u l t s o f the removal experiments: mean t e r r i t o r y F a l l Removal A r e a .  N  Simultaneous: B e f o r e After * t =  Mean T e r r i t o r y  10  2323  14  1659  2.01  P <  Size  2  (m )  2  S(m )  972 652  0.05  Successive: Before  8  3238  969  After  9  2878  548  t =  o.96  0.2 > p  y 0.1  C o n t r o l : Before  6  1745  406  After  5  2094  520  t =  1.25  * Statistically significant  0.2  > P >  0.1  size.  39  TABLE 5  R e s u l t s o f removal e x p e r i m e n t s : mean t e r r i t o r y s i z e . S p r i n g Removal A r e a  N  Mean T e r r i t o r y S i z e  Simultaneous: Before After  10  * t = 2.69  P < 0.01  2 (m )  2 S(m )  3638  370  2572  887  (one-tail)  Successive: Before  1890  766  After  2143  608  t = 0.60  Control:  0.45 > P > 0.3  (one-tail)  Before  3956  611  After  3480  1229  t = 0.86  * Statistically significant  0.25 > P ^ 0.2  (one-tail)  40 \  4.  Changes I n Numbers Of T e r r i t o r i e s  Over The  the  the F a l l  the  Removal  Area  Winter  In on  On The F a l l  p r e c e d i n g a n a l y s e s , numbers Removal A r e a were d e t e r m i n e d  removal  territories  experiments. on t h e F a l l  Changes  of replacement  males  d u r i n g the f a l l  after  in  Removal A r e a  the  numbers  of  however o c c u r r e d o v e r t h e  winter.  By  October  territorial  was  spent  the  the  14  replacements  Only  Simultaneous  Removal  were s t i l l  i t  was  apparent  a 'natural'  'originals'  from  from  the f a l l ,  non-existent,  partitioned,  by  incorporated  into  the  two  had  the t e r r i t o r y of  place.  of  these  and one was a newcomer  been males,  territories.  expanded and  of a Control  events  The p a r t i c u l a r  1973, d e f e n d e d  9  the  Of  had been o c c u p i e d i n t h e f a l l  neighbouring  sequence  incorporation. of  that  of  had t a k e n  a l o n g t h e hedgerow;  four  territories  surveys  removal  the  spring  In  (See PART B ) .  had t a k e n o v e r one o f t h e ' o r i g i n a l ' s '  gives  area.  a t t h e dominance h i e r a r c h i e s on t h e S p r i n g  population that  now  10  time  that  were  replaced  t o mid-March, I d i d n o t s u r v e y t h i s a r e a a s looking  were  had  From  10 males had t e r r i t o r i e s  males  males  present.  mid-March,  breeding  juvenile  on  Removal A r e a . In  14  males  mid-December, mid-December  17,  male  into,  two  had  male.  leading involved  to  and been  Figure the  (C6),  apparently successfully  and  a  5  latter in  the  territory  Figure 5  F a l l Removal A r e a : the Simultaneous Removal A r e a - C o n t r o l . The movement and e x p a n s i o n o f C o n t r o l male (C6) a f t e r the removal experiments and d u r i n g the subsequent w i n t e r  42  whose than  size any  the  14  in  the  the an  the  10  area  that  the  males  actually 1972.  In  that  fact,  and  on  covered  supported  removal,  square  territory  replacement  of  8755  12 8  metres,  Reifel  their  less  adult  territories area  combined  replacement males  than  the  territories  males  in  larger  Island.  occupied  much  far  in  the  fall  males  out  the  before  removal.  On  the  Successive  replacements  were  neighbouring males,  5-  males.  Breeding.  number Fall  Density  6  of  gives  1972  and  Area, the  Despite after both  the Spring  pairs after.  the  In  the  The  remained There  1973  that  the  on  Removal  essentially  were  winter. and  in  S p r i n g . Of  of  unmated  spring of  Fall  2  the  Both  of  9  of  the  incorporated,  by  spring  of  1973,  7  territories.  breeding  experiments and  into,  number  the  fact  the  Therefore,  territorial  Removal  area,  over  expanded  a l l paired, held  Table  Removal  disappeared  territories  of  Sparrows  1973  fall  after  unprecedented  Song  spring of  covered fall  an  other  Thus, in  was  ,1973  breeding  males,  of  the  territories  Simultaneous  Areas, the  between  On  the the  spring  compared.  number  the  and  for a l l areas.  densities are  pairs,  the same  therefore several  (5)  Removal  number -  14  increased  of  pairs  unmated  areas  on  breeding before,  15  territorial  43  TABLE 6  R e s u l t s o f the removal e x p e r i m e n t s : b r e e d i n g d e n s i t y  S p r i n g Removal A r e a  Breeding p a i r s Simultaneous:  Territorial unmated males  Number o f territories  Before  6  After  7  3  10  Successive: Before  6  0  6  After  5  0  5  Before  6  0  6  After  7  0  7  Before  8  0  8  After  8  2  10  Successive: Before  7  0  7  After  7  0  7  Before  5  After  5  Control:  1  7  F a l l Removal A r e a  Simultaneous:  Control:  1 0  6 5  44  males  present.  It same  i s of i n t e r e s t  result  mortality  on Mandarte of  of occupied  whilst  the breeding  previous  was  not  females of  years  until  As there  the  sex  unmated  males  unmated  males'  1  The  areas,  47  the f a l l  with  and  expansion  study  the  roughly  much t h e  catastrophic  i n March nearly  i n 1960 a n d  1962, t h e  401  to  a t t h e same  69, level  1 9 6 1 , 44 i n 1 9 6 2 .  the s e t t l i n g  of  males  the  terminated  the  either  or through  found  raised  It  juvenile number  t o 69.  i n t h e summer  breeding  through  the  density  o f 1973,  achieved  the disappearance  settling  of  females  a  of the  on  these  territories.  of  on t h e the  as explained  breeding  Controls, the  pairs  Controls territory  on  remained  removal can  the Successive essentially  experiments. be  accounted  boundaries  of  Removal  the The  for  certain  same slight  by  the  Control  earlier.  D i f f e r e n c e s In T e r r i t o r y  Males  by  o f unmated  that  ratio,  after  of  rose  o f 1962 t h a t  pairs  present  number  differences  males,  pairs  1971)  Sparrows  d e n s i t y remained  -  and on t h e  before  6.  territories  i s no e v i d e n c e  balanced  (1964;  Following  Song  on t h e t e r r i t o r i e s  territories  Tompa  Island.  territorial  number  as  that  Size  Between  Mated  And  Unmated  45  An the  interesting  Simultaneous  attract those  a  did  that  choice  vacant  on  shortage  with  of  or  smaller  from  banded  in  the  Song  that  Nice  in  March  else  the  but  does  in  did  not  territories  than  and not  Tompa  (1964)  discriminate  male.  For  there  as  females  that  s e t t l e s wherever  either 1973,  territories  males  (1943)  unmated  Island,  of  smaller  Sparrows  an  size  was  several  there the  an  is  Song  overall  males  discriminated  in  did  not  against  males  territories.  9  banded  or  Both  with  females  is  territory,  Reifel  mates,  Table  7).  female  male  areas  on  significantly  (Table  the of  had  territory  Sparrows  obtain  Removal  mate  that  state her  a  sidelight  gives  the  fall this  1972  number to  period,  of  non-territorial  early  32  February  were  1973.  known  to  speculative  and  females Of  be  the  alive  57 in  mid-February.  The  evidence  Although in  the  i t appears  spring  males  with  that,  on  females  1964).  of  that  1973  smaller  and  there that  in  spring  of  through  in  the  even  the  concluded  that  spring  of  in  other  the  i t  (25)  obtained  settling there  was  circumstantial.  females  available  discriminate i s  males  1962  passive  several did  territories, those  the  were they  Island,  Tompa  Yet,  i s both  Mandarte  apparently  females  here  against  i n t e r e s t i n g to  note  that  not  have  the  fall,  did  mates  in  of  new  females  (Tompa  an  o v e r a l l shortage  of  1962.  years  of  his  study,  there  were  46  TABLE 7  Differences i n t e r r i t o r y s i z e between mated and unmated males: spring 1973. Simultaneous Removal areas.  Simultaneous Removal Areas  2 Mean T e r r i t o r y Size (m )  N  2 S(m )  T e r r i t o r i a l males: mated  15  2828  1401  unmated  5  1736  867  * t = 1.75  * Statistically  significant  P < 0.05 (one-tail)  47  unmated 7)  males  that  with  unmated  territories.  males  had  He  also  smaller  shows  (Ibid:18,Table  territories  than  mated  males.  The  situation  substantiate  7.  The  or  replacement  accurately  study was  known  length  in  before  i t  The  not  and they  Sparrows  the  fall  the  the  3  g.v.).  had  has  i s about  the  experiment  20  more t h a n  1000  metres  from  less  than  a l l non-territorial, attempt  to  fall,  as  establish  the  parentage more  at as  nest,  and  In  Song  1943).  removed  a l l of most  local  juveniles  birthplace,  metres.  juvenile  the  on  birds  their  500  (or,  defined  (Uice  probably  the  the  of  parents.  territorial  during  disperse  is  the  days  that,  majority  bird  i t s  the  of  banded their  left  were  3)  Some  been  young  from  of  considered  1 shows  not  to  juveniles  hence  is A  i t  replaced  Appendix  yearlings).  lineage  after  aspects  were  j u v e n i l e s , and  interval  removal  areas  females)  independence  time  some  (see  spring,  of  time  one  removal  Discussion  probably do  but  males,  juveniles that  disperse  were  in (5  Table  that  both  gains this  the  origin.  on  question  the  investigation  speculations.  considering  A l l  dependent  during  Sparrows,  during  as (the  dependent  do  birds  area  preceding  i t i s worth  birds  replacement  the  further  Birds  birds.  replacement more  refute  Replacement  Finally,  requires  Further, males  Song  territories  for  48  themselves and  after  coincidentally  behaviour at  until  which  On  time  the removal  the  Spring  males  Successive  and It  hence i s  juveniles  of  early  winter,  1972,  were  Results  The slightly  that  and  breeding  between  the  are given  pairs  on  for  Experimental  i n Table  earlier,  the pairs  success  the  hierarchies, ( s e e PART B ) . dispersal  non-territorial  Introduction), Interestingly  the  The r e m a i n i n g  young  Several of the  of the  some  of the  spring  areas  enough,  fall  or  i n t h e summer  6 were  the  these  territorial  preceding  young  the  preceding  as dependent  area.  birds.  during  5 from  considering  with  1964),  replacement  the  the winter  one as a dependent area.  during  (see G e n e r a l  banded  fall,  territorial  15  i n dominance  that,  paired  of  Removal,  banded  over  a l llocal  5 had been  Finally,  than  the winter  on t h e s t u d y  compared  suggested  females  only  banded  1) a n d t h e b e h a v i o u r  males  males,  out •o f  the study  followed closely  during  12  Area,  involved  the  carried out.  the Simultaneous  were  in  1 9 3 7 , 1 9 4 3 ; Tompa was  and 2 had been  yearlings  (Table  replacement  (Nice  experiment  o f 1 9 7 2 , om  tentatively  distances  out  winter,  were  birds  moult  recrudescence  ) , 11 h a d b e e n  t h e summer  replacement  the  Removal  (10 f r o m  Removal  or early  during  during  of the resident  yearling  fall  the post-juvenile  of  a l l unhanded.  of  1973  was  and the C o n t r o l s .  8.  Controls  and had a h i g h e r  on t h e E x p e r i m e n t a l s .  laid rate  slightly  more  of successful  The d i f f e r e n c e s  eggs nests,  however  49  TABLE 8  N e s t i n g d a t a f o r the C o n t r o l s and the E x p e r i m e n t a l  N  Nest  success  Experimental:  10  40%  Control:  11  46%  N  Experimental:  a r e a s : 1973  Clutch size  3.50 t = 0.683  Control:  P y 0.5  3.66  N  Mean l a y i n g date  Experimental:  April  19  Control:  April  17  (start of f i r s t  brood)  50  are  not s t a t i s t i c a l l y  Tompa  (1964)  unbalanced, Song in  however their  first  Experimentals  unmated  In were  of  territorial  summary,  a l l  t h e new  then,  (except  non-territorial replacement  of local  remaining  replacement  suggestive  that  species 1957; the  Island  could  Yearlings  and , t o  lay  of birds (e.g.  Perrins  Controls  than  of  interference  date  1965). and  the  of laying i n  be a t t r i b u t e d  rather  highly  mortality  males.  a n d i n t h e mean  i t was f o u n d  that  juveniles,  that  origin.  they  some  between  birds,  before  obviously  much  was  to  interference  the from  males.  one)  juveniles  was  clutches,  Richdale  size  on R e i f e l  there  in  differences  Sparrows  the catastrophic  smaller  1958;  i n clutch  the sex r a t i o  by t h e u n m a t e d  later,  Snow  the  inexperience  when  Island,  t o have  clutch  1966;  Song  that,  activities  are reported  Therefore,  the  on Mandarte  breeding  Coulson  the  found  a s i n 1962 f o l l o w i n g  Sparrows the  significant.  birds  the were  the replacement  and  removal banded  were  experiments.  of l o c a l  stock.  a l l Those  young  f o r the origin  i s circumstantial,  t o o were  probably  as dependant  The evidence  birds  were  of the  but i s strongly  51  IV  DISCUSSION  &.  The  Replacement  It  was  necessary patterns  stated  on  so  The  on  on  show the  the  replacements the  study  the  replacement  removals  had  The  not  been  breeders  have  been  carried  Krebs  1971)  since  removal summer Hensley  of of  and  and  Cope  the  do  and  not  carried  by  a  first  1950  (1951).  a  that  have  prevented  the  s u r p l u s of  Song  breeding  took  capable  out  the  bred  breeding  subsequent  experiments  and  Song  shows  that  of  breeding.  the  subsequent  Song  Sparrows  possibility  elsewhere  that  i f  the  out.  experiments  i s not  new.  variety large  birds by  of  a  settlement  but  surplus of  rule  removal  exists on  a  of  the  existed  the  that  s u r p l u s of  behaviour  experiments  not  a  breeding  fact  after  removal  would  territorial  1949  and  e x i s t e n c e of  out  the  of  physiologically  demonstration  potential  Introduction  existence of  indeed  areas  Surplus  investigation  territorial  area, they birds  the  there  the the  the  area,  were  show  on  of  that  although  for  The  experiments  removal  replacements  However,  the  study  General  capable  removal  replacements  the  by  And  the  was  physiologically  birds.  place  in  territories  doing  Sparrows  Sparrows  pre-requisite  Sparrows, from  Song  scale  was  of  Removal animals  the  and  a surplus  experiments  (references  experiments  conducted  Stewart Many  of  that  i n the  Aldrich  removal  on  spring  in the and  (1951)  and  experiments  have  52  shown  conclusively  exists.  The  Island  that  present  further  a  surplus  findings  add  to  the  on  of  potential  Song  material  Sparrows  on  the  breeders on  Reifel  existence  of  such  surpluses.  The  age  determined Sparrow be  of  the  in a  few  on  2§£dracjapus  In  (1967)  and  small  predominated (1971)  Micrptus  replacements  (Bendell  Boss  (Orians  e.g.  and  1970b),  the  1961),  and  for  j?eromv.scus  onto  an  not  have  been  found  the  Blue  the  support  this  Song  the  Tit  clear;  (Krebs Healey  maniculatus,  from  but  data  Red  Blackbird,  Great  plot,  to  Grouse,  1967) ,  situation i s less  empty  been  the  Red-winged  that,  could  also  Like  Elliot  the  in dispersal  removal  replacements exists  1970),  the  experiments.  mammals,  experiments  replacement  (Orians  removal  has  young  Myers  on  and  dispersing  spp.  Few  which  individuals  i n some b i r d s ,  obscurus,  claimed  Krebs  the  phgeniceus,  1971).  the  yearlings  (Watson  Ag;elaius  of  R e i f e l Island,  primarily  Grouse  replacement  1961; and  a  descriptive  are  individuals. drawn  at  Sparrows  Watson similar  the  case  in  on  from  the  studies  Song  have  Some r e m o v a l s nonbreeding  periphery  and  of  Jenkins  (e.g.  Great  Delius  Island  the  1968;  s i t u a t i o n has  Mandarte  the  a  established  by  been  the  origin  have  shown  population  breeding  population  Young shown  to  1970; exist  Carrick  1963;  Tompa  1964).  This  Wood  that  surplus  1965;  T i t i n Bean  of  (Krebs  Harris in  some  and i s  1971) ,  in not  where  53  vacant  woodland  already  had  Sparrows local  territories  on  on  from  which  by  the  patterns  study  area  that  were  was  should to  that  be  m o r t a l i t y and/or  of  birds,  out  Area,  Table  birds  Simultaneous  spent on  the  one  Spring areas  on  the  Removal or  the  known  dominance  areas,  the  Song  is a  of  surplus  are  apparently  derived,  week o f  to  and  April  the  surplus  Table over  which  (Figure  2;  9  the  further  accounted on  Area)  on  Controls. the  problem  Finally,  area  the  of to  appear  on  both  necessarily  either  more  birds  tended  time  so was  Successive moved  to  individuals another  be  (particularly  the  tended  juveniles  some  Spring  There  areas  A l s o , some  majority  for  settle  not  Removal  of  mid-February.  to  may  number  in a l l areas.  Proportionately  than  territorial  explanation.  this  Sparrows  shows t h e  in  Simultaneous  The  Song  winter  available  but  of  banded  alive  investigation  those  emigration  be  h i e r a r c h y i n one  i n P A R T B.  for  juveniles  reasons.  although  localities.  considered  82  potentially  following  hedgerows,  certain  were  Removal  trapping  Removal the  63  of  banded  9 r e q u i r e s some  more  the  In  the  removed.  winter  for  hedgerows.  n u m e r i c a l l y exceed  Although  be  that  pre-reguisite  juveniles  to  birds  second  non-territorial  Removal  mainly  1964).  settlement  these  by  replacements  necessary  birds  occupied  Island, there  further  the  were  i n adjacent  disappears  Tompa  A of  Reifel  birds,  apparently also  territories  around  remain moving  elsewhere to  in from is  congregate  54  TABLE 9  Number of juveniles banded, f a l l 1972 to early spring 1973  Spring Removal Area Males  Females  Undetermined 1  Total  Simultaneous:  29  20  Successive:  7  1  Control:  11  10  1  22  Totals:  47  31  4  82  2  50 10  Fall Removal Area Males  Females  Undetermined  Total  Simultaneous:  22  18  1  41  Successive:  11  7  1  19  Control:  4  1  0  5  Totals:  37  26  2  65  55  at  certain  Spring  favoured  Removal  Area,  the  junction of  Out  of  than  82  34  Thus,  there on  establish  is  not  The  Temporal  to  show  that  the  area  Spring this  be  a  and  the  On  the  was  at  Control.  Area,  no  less  locality.  large  number  The  s e v e r a l of fate  locality  Removal  areas.  The  hedgerows.  favoured  Removal  at  removal  Patterns  removal  that  a  removal  size,  i s now  resulted  number  can  be  of  not  Simultaneous  the  of  potential  results  these  in  birds  rest  of  of  the  were  able  the  surplus  enguire  areas,  Removal  smaller  mean  Territories  subsequent  in a  more  into  and  with  given  show  a  smaller  area  that  size,  in  f a c t o r s which  mean  territory  in  significant  no  replacements  a  a  mean  after  a  successive  territories,  territory  or  a  given  area.  could  have  size  on  the  differences  areas.  F l u c t u a t i n g Environmental  The  the  also  smaller  Removal  On  territories,  mean  to  the  and  They  smaller  Successive  of  result  necessary in  Settling  accomodated  removal.  does  Of  experiments  greater  significantly  1.  appeared both  The  the  most  the  banded  territories.  simultaneous  on  on  along  known.  territory  It  and  experiments  to  show  single  Simultaneous  caught  replacements  B  the  j u v e n i l e s banded  were  removal  the  localities  Resource  territory  size  on  the  Simultaneous  56  Removal  area  could  environmental  be  resource,  related  to  some  fluctuating  on the argument t h a t t e r r i t o r y  are a d j u s t e d a n n u a l l y to some environmental  variable.  were the case, then the t e r r i t o r y s i z e s on  the  Successive  mean  Controls  and  This  d i d not  happen.  t e r r i t o r y s i z e s on the S u c c e s s i v e Removal areas and  on the C o n t r o l s s p r i n g and f a l l in  If this  Removal areas should have f l u c t u a t e d i n accordance  with the Simultaneous Removal a r e a s . The  sizes  territory  fluctuated removals, size  in  opposite  although  before  and  directions  i n the  i n no case was a d i f f e r e n c e after  a  removal experiment  significant. I t i s p o s s i b l e t h a t some r e s o u r c e changed along only p a r t of t h e hedgerow, i n  both  hedgerows  are  t h e r e are  substantial  Simultaneous  Removal  areas.  The  not homogeneous i n v e g e t a t i o n a l s t r u c t u r e , and differences  in  mean  territory  between, and w i t h i n , the S p r i n g and F a l l Removal Areas  size  (Tables  4 and 5 ) .  A s u b j e c t i v e assessment o f the v e g e t a t i o n along the  hedgerows  suggested  that  smaller  mean  territory  r e l a t e d to a higher i n c i d e n c e of brambles, although a n a l y s i s of reasonable in  the  vegetation  was  carried  out.  s i z e was no f u r t h e r ,  However,  assumption would be t h a t a resource would  parallel  along  the  whole  j u s t i f i a b l e t o r e j e c t the argument  hedgerow. that  a  Thus, resource  a  fluctuate i t seems changed  along only p a r t o f t h e hedgerow from 1972 to 1973. The  observed  r e s u l t s on mean t e r r i t o r y s i z e , t h e r e f o r e ,  57 /  cannot  e a s i l y be  environmental nature the  of  study  Sparrows in  resource,  the  along  )  on  such  habitat.  period  numbers  bewickii  explained  on  the  as  the  basis  of  food  supply  or  Further,  species  of  hedgerows  birds  did  those  species  (e.g.  that  might  compete  for  interspecific  a  fluctuating change  in  observations  co-existing  not  in  Therefore,  casual  a  reveal  any  Bewick's  with  the  was  on  that  Song  changes  Thryomanes  some e n v i r o n m e n t a l  competition  during  major  Wren,  the  resource.  also  apparently  unimportant.  2-  Pressure  In  his  territory might  be  the  suggestion  Krebs  pressure are  is  given  (Perrins  of 1965;  in  in  Wheatear,  found  survived  the  the  spring  next  to  and  Oenanthe  winter on  A  did  a  not  Mandarte  a  the  relationship density (Conder  number  of  influence Island.  the  time  when  behind  this  Tit  between  breeding  density  number  has  been  also  1956)  Song  and Tompa  of  found  the  Tree (1964)  Sparrows  breeding  in  winter  between  1965). young  factor  Great  relationship  (Weeden  the  rationale  For  determine  possible  subsequent  oenanthe,  the  The  7.  breeding  arborea,  that  be  and  1966).  Spizella  however  Appendix  that  might  s e t t l e r s at  established.  appears  spring  suggests  potential  juveniles  birds  Sparrow,  the  from  in  Lack  factors  (1971)  being  Wood, t h e r e  population  the  Intruders  discussion  size,  territories  Marley  From  density  that of  58  For  the  population it  was  9).  For  of  example,  82  Area,  birds  If areas  an  but  areas  conclude  that,  the  then  would the  any  of  areas  (see  one  might  been  these  were  the  that  on  Table  on  the  there  settle  were  on  both  this  may  given.  the  Simultaneous  of  a  suppose  exerted No  and  Spring  banded  explained earlier,  latter  areas,  on  to  reasons  Controls.  for these  banded 50  pressure  have  and  Removal  i t appears  as  winter  large,  available  f o r the  by  settlers,  in  on  large  number  that  a  both  the  similar  significant areas,  pressure  Removal of sort  Successive differences  t h e r e f o r e one  from  can  intruders  was  important.  It formed and  Thus,  the  was  were  these,  areas,  so,  caused  occurred  not  of  area.  Removal  pressure  Removal  juveniles  Island,  juveniles  Simultaneous  potentially  be  Reifel  increase in territories  was  potential  the  and,  necessarily  on  non-territorial  Removal  Simultaneous  of  banded on  Simultaneous  not  Sparrows  largest  Removal  more  Song  may  be  worth  dominance  retained  consequence individual  has  subordinates apparently territory  hierarchies  them of  through the  the  the  collect to  in a  that  the  (see  PART  winter.  dominance  precedence  in  holder  mentioning  a  spring.  s h r i n k the  B)  in  the  vacant  The and  try  late  I t i s concluded  hierarchy i s that  to  group  non-territorial  boundaries  force of  males an  fall,  that  a  dominant  territory  yearling to  the  birds  over do  not  existing  his territory,  to  59  accomodate however, the  one  that  birds  dominant (PAST  Song  B),  mean  i t or  cannot  just  of  from  to  see  a l l  that  the  Successive  the  i t  males  i s  not  most  territories  competed  for  competed.  pressure a  areas,  that  Although  resulting  from  reduction in  Simultaneous  alone  the  7).  stressed,  imply  obtained  caused  Removal  be  not  individuals  settlers  on  does  Appendix  i f  dominant  taken  (It should  hierarchies  known  potential  Inexperience Of The  3-  -  i n the  the  explain  group.  intruders'  groups  eliminated,  sufficient  the  hypothesis  size  the  be  of  not  the  number  not  in  i s  territory  (but  up  Sparrows  Although large  more  'pressure  gang  territories,  a  or  Removal  or  the  the  areas  Controls),  considered  nearly  results.  Juveniles  And  The  Temporal  Patterns  Of  Settling  If  changes  potential then of be  the  settlers,  cannot  factor.  (Southern  than and  Dhondt  l§M£2£llH2* Chickadee,  fully  the  (simultaneous  territories  1968;  environmental  inexperience of  settling a  i n an  adults Morley  explain  have  i n the 1950),  been Marsh  the  White-crowned  (Ralph  and  Pearson  atricapillus,  on  shown Tit,  Great  the  or  T i t  (Stefanski  and  to  the  take  Parus  pattern  1967).  could  smaller  palustris,  (Dhondt  the  from  results,  territories  Sparrow,  1971) , a n d  pressure  observed  birds  successive)  1971a),  Parus  the  replacement  and  Yearlings  resource,  and  Huble  Zonotrichia Black-capped' Nice  (1937)  60  and  Tompa  that  there  territory and  (1964) was  a  sizes  weeden  , however,  could  significant  of f i r s t  (1965)  found  year  n o t show f o r S o n g  difference birds,  similar  between  and t h o s e  results  Sparrows the  of older  i n the Tree  mean birds,  Sparrow i n  Alaska.  It was  i s  playing  possible a part  Simultaneous  on  inexperience sizes  were  although  one;  territorial  the  was  section  again  system  Successive  i n this  one  unimportant,  3).  The  respect,  explanation  could  a tendency  to take  than  adult,  but  this  vacancy  appears  vacancy  may  is  ready  to defend,  in  two  would with  initially  between  be m a s k e d  i n  however,  the  were  territory removals,  a l l yearlings  areas  be l a r g e r  up  of  the  i s probably  than  i n general,  smaller  territorial  territory when  enough This of  occurred  only a  pattern.  the replacement  birds.  disappearance  replacements,  a  that,  i s not apparent  the  two r e p l a c e m e n t  subsequent  were  boundaries.  be  but i t i s not large  i f the  birds  retention  of t e r r i t o r i a l  that  the  and i s c o n s i d e r e d a t l e n g t h i n  do have  single  birds  birds  on  a s mean  b e f o r e and a f t e r  juveniles an  taken  areas,  t h e S u c c e s s i v e Removal  on d i s r u p t i o n  However,  Removal  the replacement  Appendix on  territories  as a l lthe replacement  apparently  see  important  areas,  t h e same b o t h  once  (except  very  the  much  inexperience of the s e t t l i n g  i n the smaller  Removal  yearlings.  that  to  be  initial  territory  The  juvenile divided tendency holders,  periodically,  as  61  happened  i n the  (catastrophic  studies mortality  experiments  i n the  masked  a  empty  i f of  From after  a  between  large  results of  each  neighbours, present)  tendency with less  to  then  levels  of  the  a  take  territory.  If  and  a l l  the  then  a  set of  risen  three to  four  days  the  (  removal) , the  that  hedgerow  the the  Tompa removal not  be  completely as  The  in  the  the  or point  four  days  where  he  as  be  they  take  have  this  removal  could result.  a  tendency  of  can  the  The  might  be  are  on  smaller  initial  low  experiment, If  successive  his level claim  with  connected  juveniles  to  in a  juvenile  disappeared.  because  simultaneous  i s involved,  interval(s)  initial  might  replacement  however,  boundaries  had  replacements  in a  time  original  possibly  areas,  replacement  perhaps  smaller territories  juvenile  might  present,  therefore inclined  a g g r e s s i o n , as  have  was  a smaller territory  of  after  suitable  and  successive  tendency  smaller territory  level  then  or  initially,  ground,  Nice  S u c c e s s i v e Removal  aggression.  unfamiliar  of  established  take  to  i n the  The  Sparrows  indicating  aggressive  replacement  study.  successive  his  and  of  experiments.  three  a p p a r e n t l y had  (if  Song  on  male  Sparrows  apart),  stretch  removal  period  Song  present  territorial  simultaneous  on  just  removal,  aggression whole  one  of  might the  62  territory  he  Van  den  explained into did  has  a  acquired . 1  Assent  (1967)  his observations, tank  resulted  relative  in a  differences  sticklebacks  -  settling  building  males  and  were  Mandarte large  of  resulted  the  1971)  that  more  likely  considered  in a  larger  in  i s  animal  greater and  no  familiar will  detail  position  an  levels in  smaller  of  animal be  i n PART  i f  Sparrows  on  there 1965;  i s with  an  subordinate B  under  in  non-territorial  territories  Dixon  of  found  settling  Further,  1963;  male  process  Song  previously  the  than  the  than  probably, of  the  In  number  Brown  territories  that  mortality.  1949;  less  that  experience  There  subordinate,  Brian  the  showed  introductions  manner;  aggressive  temporarily  (1964)  of  similar  successively.  catastrophic  (e.g.  a  number  simultaneously  were  introduced  numbers  evidence  -  in  three-spined stickleback  simultaneous  larger  in  them  I s l a n d , Tompa  juveniles after  a l l of  the  that  successive introduction,  the  and  for  prior  taken i s some  Phillips area, (this  the is  occupancy  i n the h i e r a r c h y ) .  conclusive  evidence  from  the  study  to  1. An i n d e p e n d a n t m e a s u r e o f ' l e v e l o f a g g r e s s i o n ' s h o u l d be taken i n order to v e r i f y t h i s hypothesis. One p o s s i b l e method w o u l d be t o m o n i t o r a b i r d ' s r e s p o n s e t o a t a p e p l a y b a c k a method w h i c h m i g h t be p a r t i c u l a r l y u s e f u l i n t h e c a s e o f S o n g S p a r r o w s , w h i c h p r o v e d t o be v e r y responsive to a stranger Song Sparrow's song.  63  substantiate  the  hypothesis  initially  less  smaller  territory.  hypothesis  a g g r e s s i v e , and  i s ;  non-territorial in  the  the  retention,  boundaries, section, time  of  On to  to  the  days  and  inclined  most  be  were  (PART of  discussed  i t may  evidence  were  A  further  Before  considering  young, adults i s  territory  leaving  the  a the  factor  traditional  next.  worth  take  subordinate to  B).  the  to  are  supporting  replacements  (b)  disruption,  i s suggested  replacement with  neighbours  with  the  the  guidelines  to  territory  to  longer  to  f i x .  days  needed  for total  Of  in  This  and  may  time  the  this  problem  interval  length  very  of  thus  partly  present  quickly  Removal  that  of  three needed  territory there  were  might  have  establishing  areas,  there  boundaries explain  of  time  the  fact  boundaries  bird  follow,  a  re-establish The  Simultaneous  much  Disruption  being  neighbours. fixed  On  areas,  as  birds  settling  boundaries.  4.  more  the  S u c c e s s i v e Removal  boundaries  such  hence  birds  replacement.  the  aided  settling  hierarchies  be  however,  four  for  a l l  birds,  or  the  Suggestive  (a)  dominance  that  may  the  his  were  have  nine  no  taken  or  ten  that  the  replacement.  Territorial  Boundaries i  Both increase  Simms in  catastrophic  (1965)  number  of  mortality  and  Tompa  occupied was  (1964)  noted  territories  accompanied  following by  the  a  total  64  rearrangement territorial broke  down  of system  hedgerows  The  Simultaneous in  of  completely  the  given  territory  were  the  divided  6,  Song  The  up i n t o  area  with  on  traditional  S p a r r o w s on t h e s t u d y  on t h e S i m u l t a n e o u s  Removal  Figure  boundaries.  Removal  territories  the Spring  t h e o l d a n d new  area  areas,  entirely  Removal  and anew.  area  territory  i s  boundaries  shown.  It were  i s apparent  not  removals. occur  The only  remained new  territory  pattern  i s i s  much  birds  settled  that  pattern  to  such  account  boundaries  according  territories  from  year  to  a n d Snow  the the  Removal  Removal  traditional to  the  the the  birds  Song  the  some  of  of the  areas.  of  landmarks.  The  experiments.  considering  features;  i n  hedgerow.  f o r the retention  territorial  year  boundaries  contrast  stability  to certain  before the  Successive  in  birds  i s preserved.  and a f t e r  I t i s worth  the  the  the  the removals,  topographic  that  of  retained,  that  system  territory  the  i n the Successive  possible  (Davies  along  might  replacement  arrangement  before  areas.  structured  philgmelgs,  i n the  after  was  the  territorial  section  t h e same  Removal  due  the  boundaries  explanations  territorial  6 that  of the territory  pattern  Simultaneous  It  by  narrowest  territory  Although  the  the  Figure  similarities  1  linearity  The areas  influenced  along  general  from  territory  habitat f i x  i s  their  The s t a b i l i t y i n Thrush,  1965) a n d t h e E u r o p e a n  Turdus  Blackbird,  Figure 6  S p r i n g Removal A r e a : b o u n d a r i e s b e f o r e and a f t e r the removal  .  e x p e r i m e n t s on the Simultaneous Removal a r e a  66  T_«. m e r u l a ,  (Snow  1958)  habitat.  Song  Sparrow  presumably  the  birds  themselves  and  boundaries,  however,  interactions directly  to  occurs,  to  birds  replacement territory  area.  and  after  schedule, been  the  retain  one  fact  i n an  some s e d e n t a r y their  of  i s not  traditional  of the  related  how  how  into  i t with  boundaries  deduce  that  interval  of  passerines  on  throughout  would  of  the  for  the  males  the  Removal and  wait  vacated d i d not areas, four  boundaries  of four  the  existing  boundaries  a three  a maximum  area.  along  takes  into  territory  that,  males  the  the Successive  conducted  before  Sparrow  the adjacent  the t e r r i t o r y on  settle  linearity  long  When a  the vacated  Song  the  long  their  were  can  male  to  territory  individuals.  bird  familiar  that  territories  product  neighbouring  owing  known  t o expand  can  two  ,become  the removals then  the  to encroach  only  or  between  recognition  and  and  habitat.  the experiments  determined  In  to  boundaries,  attempting  that  not  a  themselves,  a replacement  areas,  i s  bird  From  change  likely  replacement  with  The  the t e r r i t o r i a l  begin  study,  It  by  new  Removal  hedgerow.  before  a  contend  Successive  the b i r d s  respected  the present  have  most  defined,  boundaries  neighbours.  i s  to features of the  are c l e a r l y  the  explanation i s that  neighbouring In  attributed  recognize  features of the  i s  vacancy  been  territories  their  between  A second pattern  have  day had  days.  like  t h e Song  the  year,  as  Sparrow, i n the  67  European  Robin,  Erithacus  Wren-tit,  Chamaea  fasciata,  activities  of  newly  territory  boundaries,  former  boundaries  behaviour  has  M££2£fii£Ki# the  newly  show  such  areas,  been (Young  This  could  further  to  It  in  the  same  explain  one  of  to  explore  Similar  on  On  and  Reifel  the  first the  of  the  exploratory  Skua, to  Catharacta argue  Island  on  that  would  Simultaneous male,  the  the  re-establishment  replacement  would  is  Mccormick's  Sparrows  one  194 3)  seems r e a s o n a b l e  patterns.  the  birds  neighbours.  Song  any  at  1938),  leading  1972).  boundaries,  same t h i n g  (Erickson  described  settled  however,  (Lack  settled  with  behavioural  territory the  the  rubecula,  also  Removal  exploring  find  other  time,  with  no  guidelines  to  follow.  length  of  time  to  fix  the  replacement  males  the  taken  doing  the  boundaries.  5.  Genetic  It Klomp  Basis  i s not  known  (1972) that  evidence  that  size  i s  (1962). probable  lends  Of  Territory  territories  the  problem  support  individuals  of  fixed  Klomp  territory that  genetic  far  genetically  indications  their  how  Size  i s genetically fixed,  However, that  The  discusses  territory  that  For  the the  make-up.  to from  states  size  has  a  same s p e c i e s of  the  be  an  gives  Schoener  of  (1968)  basis,  of  this  are  based  size  of  circumstantial  although  genetic  compressed.  optimum  existence  that,  variations  Most  of  and  the  can  an  and  i t is there  attribute on  differences  optimum Gibb highly are  no  among  differences  in  seem  of  to  be  68  a  modificatory  a  territory  one  generation  of  lineage  particular  tested  the  of  a  are the  C.  large to  a  i s not  the  after  territorial  The  on  immediately  the  pursued  a  have  but,  territory into  For  the  could  Different  a be  i s that  the  from appear  one to  brambles,  and  involved.  The  to  be  drawn,  Results  Of  The  Experiments  not  show  a  or  mean  in  significant territory  replacement and  Each  the  occurred traditional  replacement  defend  because  area,  obtained  here.  basis,  holders  a  to  on  conclusions  did  to  of  of  few  territories  a l s o be  because  perhaps  vacant  small  territories,  prepared  too  from  examples  male  passed  on  parents  impression  Removal  areas  only  the  are  i s  The  retained.  been  which  data  passed  the  particular  firm  one-to-one was  in  further  removals  be  incidence  Successive  of  may  gives  general  high  Removal  pattern  territory,  neighbouring  The  f o r any  number  the  initially  vacant  that  However,  Explanation And  10  territory  i s a  sufficient  successively  may  next.  Successive in  but  size  area,  and  small  there  matter  The  size,  known,  or  the  Proposed  change  study  vegetational factor  not  Simultaneous  Table  f o l l o w i n g year.  where  therefore  and  the  are  particular  next.  statistically,  taken  data  a  the  from  generation be  of  to  male  territory  taking  nature.  of  only the  a  juvenile of  the  reluctance  of  their  part  to  expand  boundaries  and  p o s s i b l y because  of  the  69  TABLE 10  Generation 1  L i n e a g e : t e r r i t o r y s i z e s from one g e n e r a t i o n t o the n e x t  Territory size  2 (m )  Generation 2  Territory size  W/0  2708  R/BL  1824  W/0  2708  LP/BL  1444  W/DP  1625  Y/LP  2166  DP/M  1264  M/W  G/R  5054  B/B  722 3249  2 (m )  70  proposed  exploratory  boundaries the  of  the  replacement  behaviour  territory bird,  of  of  the  probably  the  new  removed in a  bird, bird  matter  the  were  of  size  and  taken  by  three  to  four  days.  The  Simultaneous  significant  changes  Removal  i n number  territory  sizes.  The  completely  broken  down,  entirely with  anew,  the  the  replacements  argue  that  were  possible  It  levels  immediately the  were  factors that  of  claim  removed  a  size  adults.  attempting  to  settle,  territories  the  summary,  study  area,  physiologically  then,  of  with  a  the  capable  size  the  have  of  a  birds  surplus breeding.  days).  A l l  reasonable the  less to low  a l l  to  area taken. had  likely  to  that  of  level  of  simultaneously  in a  larger  territory  experiments  the  initially  were  resulted  formed  territory  temporary  s m a l l e r mean  removal  was  ten  was  f i x  with  of  mean  comparison  to  equivalent  birds,  in  were In  i t seems  hence  territory  could  there  the  and  replacement  taken,  and  to  showed  system  taken  (nine  replacement  Thus,  the  time  unfamiliarity  aggression,  and  boundaries  the  birds,  hand,  territorial  longer  affecting the  of  In  much  other  guidelines.  areas,  young  aggression  of  apparent  took  the  territories,  territory  i n e x p e r i e n c e and/or  i s postulated  lower  and  without  boundaries  of  on  traditional  S u c c e s s i v e Removal  territory  areas,  number  size.  showed  that,  on  of  birds  that  were  The  replacement  birds  71  were  almost  a l l j u v e n i l e s , and  were  probably  a l l of  local  origin.  The show  successive  significant  territory  smaller account  changes  sizes.  territories  explanation  The  resulted  mean for  pattern  i n numbers  results  i s given.  numbers  sizes. are  on t e r r i t o r i e s  of territories,  simultaneous  i n higher  territory these  of settling  pattern of  Possible  d i dnot  o r i n mean  of settling  territories, f a c t o r s which  discussed,  and  a  on and  might  proposed  72  PART  I  B  DOMINANCE  HIERARCHIES  INTRODUCTION  Schelder'up-Ebbe  (1922)  biologist  to  recognize the  in  of  animals.  groups  peck-order  He  i n domestic or  subordinate  situation  was  essentially  then, species  Wood-Gush  1965)  been  and  vertebrates, Witter  1969);  Cynomys  in  laboratory  mice,  primates,  e.g.  spp.,  Leopard  Krebs  1973).  (e.g.  Ito  1970)  be  Lemur  Devore  Indeed, to  both wild  of  a  linear  was  that  animals,  the one  and  Frog,  either  the  dyadic  Rang  of  1965);  have  the  the  Mantis  and  most  Cockroach,  1969);  the  the and  and  (Jolly  hierarchy the  hierarchies  e.g.  e.g.  in 1953;  pipiens,  (McBride (Levick  observed  (e.g. Guhl  Caldwell  1955),  spp.,  been  domestic  and  (Anthony  lemurs.  first  hierarchies  bird  have  vertebrates,  musculus,  and  each  o t h e r , and  1970),  truncatus,  Mus  (Hall  For  higher  Tursiops  existence  hierarchies  (Dingle  ludoyicianus,  Dolphin,  the  the  irreversable.  (Ewing  the  being  dominance  invertebrates,  bredini,  as  of  that  any  animals,  in  e.g.  such  to  wild.  cinerea,  Gongdactylus  Papio  of  documented  Nauphoeta  discovered  dominance  numerous  credited  importance  fowl,  dominant  Since  i s  Shrimp, in  lower  (Boice  and  Prairie  Dog,  Bottle-nosed Hebb  1948),  and  Beilharz  1973);  in  1966),  i n man,  and  baboons,  (Sommer  1967;  i s c o n s i d e r e d by important  concepts  some in  73  animal  social  There  i s  hierarchical been  found  herons  behaviour.  wealth  i n such  diverse  Worm  1971; B r u l l ,  (Bennet  1939),  cited  parrots  i n corvids  in  (Brian  parids  fringillids Wessell  under  half  possibly  Brown  sentiments imposing  when  social  Several winter area  flocks from  although subject  Gartlan  the  of birds in  the function of  Wynne-Edwards  much 1962),  1952; L o c k i e  1956),  1942),  and  in  1964; S a b i n e  1939).  which  the  that  however,  animal's also  care  should  conducted  outcome  in  a  (e.g.  the consequences  taken  Hence,  hierarchy  were o f  geographic there  hierarchy.  Lack  i n  units.  different  the  o f t h e dominance  be  (see  similar  i n the wild  winter. of  behaviour  expresses  are not s o c i a l  they  discussion  various  i n laboratories, a factor  that.breed  on t h e f i n a l  and  done  studies  which  1934b),  pigeons  were  (1968)  o n what  (Watts and  1967),  Odum  studies,  influenced  models  that  information  bird  have  tree-nesting  1 9 3 3 ; Tompa  1941; Shoemaker  he s t r e s s e s  of  1966;  1954; Tompkins  conditions,  1963:460).  a n d Hogan  1963; L o r e n z  on  relationships  lek species  and A l l e e  Dixon  material  colonial  1938),  (Masure  o f these  have  These as  by K r u i j t  1949;  and Leigh  unnatural  could  observational  groups  (Brown  (Tordoff  Almost  of  and Schmidt  passerines;  1959;  v  relationships i n birds.  (Noble,  Stokes  a  ^  i s no  Indeed,  has been t h e 1954,  are not well  1966;  understood.  74 (  A  few  studies  hierarchy  through  spring.  Odum  flock the  the  status  of eight  Mandarte alpha,  Parus  four  beta,  two  alpha  birds  territories,  of  birds  I  therefore  opportunity the  dominant  during  the on  Reifel  to  at  the  fall,  area  had been  present  territories.  of  Finally,  by l a t e  chance  f o r  question; obtain  in  neither  established of  and  associations  Song  experiment  their i n  the  along the  Sparrows enabling  Song  opportunity  localities  fall,  when t h e  territories  population  loose  several  o n how spring,  and  an e x c e l l e n t  of the  o f two  difficulty  hierarchies  favoured  t h e removal  the  hierarchies,  formed  proportion  In  on  territories.  The  the  at special  no d a t a  no  that  presented  in  Sparrows  two o f t h e b e t a  season?  colour-banded  a  had  in a  i n Carolina  was c o m p o s e d  the following  The j u v e n i l e s  A large  recognition. would  nature  o f Song  i n obtaining  males  determined the  although  i n dominance  Island  and c o n g r e g a t e d  hedgerows.  but only  pre-breeding  (1964)  are given.  are the juveniles  individuals  consequences.  birds,  i n the  territories  results  t h e group  apparently  investigated  Sparrows look  that  succeeded  arises,  Tompa  determined  establishing t h e gamma  similar  i n a group  a n d t w o gamma were  the top three  established  found  males  t h e dominance  i t s breakdown  that  carolinensis.  and found  relationships  the  (1965)  followed  until  reported  juvenile  Island,  have  Chickadees  and Dixon  Chickadees,  birds  winter  (1942)  of Black-capped  spring,  the  in  i n the  individual  i n  the  spring  yearlings  to  obtain  75  II  The  PROCEDURE  Spring  Removal  hierarchies  were  Feeding male's set  millet, each  stations  they  measuring  by  barley  the  (about  three  stations, principally  left  early  flat  which  the  adult  tables  were  of  of yellow  worst a  after  December,  was  put out a t a s t a t i o n  of other  amount  Sparrows,  to  of on  the  seed-eating  spp.,  eaten  by  close  to a feeding  one  feeding  passerines, Pipilo  Passerella iliaca. day  and  bait  any  Towhees,  a t t h e end o f t h e  at  (see monthly  Only  attracted  white  intervals  weather  small  plywood,  and  2).  Fox  presumably  i n each  pieces  mainly  Juncg  and  up  put out a t i r r e g u l a r  a number  at the station  morning,  along  Nineteen  (a m i x t u r e  readily  Juncos,  erythrophthalmus, food  were  were  of  the winter's  Appendix  set  1972.  out  was  handfuls)  as  hedgerow  were  the winter,  of  Sparrows  of  Bait  oats)  period  Song  made  throughout  summary.  day.  (tables)  40cm.  and  weather  the only  i n the f a l l were  60cm  station  after  was  determined.  territory  up;  Area  had  Any  gone  by  rodents.  i  A  hide  station effected  was by  recorder activities given  was under the  (Sony  erected  observation.  The  presence  the  110a)  and  at the station;  of a  birds  i n this  to t h e i n t e r a c t i o n s between  A  were  way,  when  d i d not appear  hide.  microphone  station  used  full  the b i r d s .  that t o be  cassette to record attention  tape the was  76  All  interactions  Interactions feeding  stations  encounters between  and  individual  were  two a  causing  retreating  displacements,  below.  be  Appendix  clearly  noted  at  the  5). A l l  took  place  and i n a l l e n c o u n t e r s  to retreat  was  recorded.  birds  invariably  a time,  as  divided  avoidances,  A i s feeding Bird  flight  B  at Bird  1943:156)  determined.  was  noted  as  a  The  dominant;  subordinate.  into  three  categories:  and f i g h t s .  beak  directly  either  at  i s made.  dominant  bird  On  A,  being  i t would  Distances  ft.  driven  or t o the ground  displaced.  be d r i v e n  between  Should  o f f by B i r d  the birds  threat  and Bird  determined  B  below,  (Nice  has  wings  lowered,  flies  would  another  usually f l y  depending  a subordinate  and  o f f before  providing A  ground  posture  o u t , head  off,  the  a rapid  invariably  present.  on  Bird  fluffed who  or  makes  in  Bird  i s not already  been  hops  feathers  Bird  to the table  had j u s t  at  body  table  and e i t h e r  A, o r q u i c k l y  pointing  contact  at the bird  appears,  •out-of-pocket*,  then  at  were  other  (see  Sparrows  the other  were  and  noted  could  individual  Sparrows  Displacements  Bird  it  also Song  loser  Song  Sparrows  birds  Encounters  (a)  Song  involving  only  winner  the  between  between  be  on  where  present,  A.  varied  from  one-half  to  one  77  metre,  (b)  and  appeared  to  increase  A  is  feeding  the  table  appears  and  hops  below.  Bird  area,  whilst  adopting  •out-of-pocket*. 1943:225), down, and Bird  A  hops  raised  away  one  from  once  mild A  approached.  assumes  out  the  an  up.  feeding  more o f t e n ,  towards  body  As  the  posture,  alarm  feathers,  and  again  or,  threatening  crest  stretched  Distances roughly  a  Bird  with neck  B  at  ground  on  feeding  with  wings  posture feathers  Bird  B  the  (Nice sleeked  approaches,  area.  between  birds  varied,  directly  facing  each  but  were  metre.  Fights  Two have  birds tails  'square  out,  and  quite  close  to  Two  generally  close  bill  each  remain  seconds.  and  body  frequently  often  off*,  fanned  •out-of-pocket',  and  spring  Avoidances  Bird  (c)  as  open.  other,  motionless possible  to  in  i t s bill,  turns  the  other  in  pursuit.  fight.  through They  the  shrubbery,  spring  into  20  are  i t s head, There and the  (2) air  Both  depressed, ground,  two  these  outcomes  closes  chase  The  about  bird  close  the  other.  feathers  birds  to  30  wings  generally  centimetres  postures  for  then  then the  flies  follows birds  together,  a  are apart,  several  distinguishable: and  puffed  (1)  off  one with  prolonged physically  pecking,  and  78  attempting The  two  to  birds  ground,  through  the  Each fight)  the  the  of  that  Sparrows in  gives by  to  twelve  and  the  surmount feet  flies  victor  the  and  above  the  with  the  off, a  other.  prolonged  chase  encounter equal  (displacement,  weight  were  at  the  combined. did  analysis,  This not  and  that  the  encounter  was  not  in  interactions  seemed  involve  time,  the  a  in  avoidance, and  more  the  juvenile  than  as  doubt.  some o t h e r  a l l three  justifiable  decision  in  and  to  two  who  This  on  was  is  fringillids  in (e.g.  1959).  mentioned  congregated  at  earlier,  special  favoured  These  localities  frequently  male's  territory,  and  feeding  stations  observations of  would  inevitably and  the  female. group  involving certain  involved  individuals  of  the  feeding  in  Interactions juveniles adult  stations  group  were  more  more  than  one  than  females  and  only  at  as the  the  one  one  adult  were  adult  different  Further,  different  combined,  hedgerow.  juveniles.  than  involving  attracted  of  analysis.  over  with  more  Sparrows  the  observations  same the  Song  along  encompassed  ranging  interact  different  localities  therefore  were combined  group  and  as  way  interactions  an  to  As  male  trying  much  pursuit  given  grounds  Sabine  as  one  encounters  contrast  each  underbrush.  type  victor  claws,  rise  close  was of  Song  can  before  accompanying  types  clench  These  the  territory territorial adult  males  interactions  juveniles. adult  same  male  Also, and  his  1  79  female was  during  the observation  and hence  no  hierarchy  resolved.  Matrices (see  period,  were  Results).  during  the  particularly  constructed  The outcomes  spring they  for  each  hierarchical  o f t h e h i e r a r c h i e s were  a s t h e h i e r a r c h i e s began  were  followed  experiments  presented  an o p p o r t u n i t y  members  of the h i e r a r c h i e s to obtain  followed  t o break-up, and  i n the removal  removal  group  areas  f o r some  territories.  where t h e of  the  80  III  RESULTS  (A)  The  Hierarchies  Seven and is  matrices  included  The ranged time  bias due  i n the text  i n the  number  from  amount  to the fact  over  an  feeding  area  down group  involving  one.  the  The  7 ) , and t h e  involved  to five  hedgerows,  B-C  Hierarchy  remaining  the group  i n the  six  hierarchies  individuals.  of birds  spent  incorporating  i s  given.  The  amount  The  heavy  on  t h e B-C  Hierarchy  i s partly  of  birds  concerned  wandered  three  territories,  As n o t e d  earlier,  t h e same  group  a l l hierarchies, Table  (displacements, an  1153  example,  and  observations  of birds  fights, t o be  males)  were  11  t h e B-C  and  shows  hence  three  at different  were  combined  birds  only  at  the in  numbers  was 9%  in  based were  encounters top  more  of  of  in  four  hierarchies.  on a grand  On than  the  involved  encounters  avoidances,  displacements.  i n more  involved  the  Hierarchy  the rest  of i n t e r a c t i o n s  and f i g h t s )  o f which  involved  Hierarchy,  the majority  avoidances,  interactions,  tended B-C  each  that  displacements.  ^%  (Figure  along  analysis.  In  As  f o r each  Sparrows  of time  stations.  stations the  o f Song  with  determined  Appendices.  twenty-seven  spent  in  were  were c o n s t r u c t e d  represented  are  of  hierarchies  total less  t h e whole, feaales.  hierarchy  interactions  than  birds  of  than males  In the (mostly lower  81  Figure 7  B-C  HIERARCHY  LOSER 9  tf  tf  tf  LP  G Y  w®  0 • G  R Y  4  5  7  22  13  7  3  6  2  7  7  6  5  8 1  <f  tf  B 8  G B8I R YR  W R ROB 8 B  \  G B Bl W R \ R Y R R DB  6  LP  2. Y  £ OG R  T w  BG 8  BS  tt  3  Y  W ®  Y  1  tf w  9  w© R®  B RW  LP  6  11  1  4  11  2  5  5  16  11  5 '6  8  2  8  1  2  2  1  5  1  3  7  2  5  5  3  1  11  2  22  2  3  24  1  1  5  17  3  1  15  6  5  5  9  3  10  11  9  9  16  6  6  \  10  2  16  6  5  2  6  1  2  1  \  1  7  \  2  BG  B BG  BY BY  6  11  3  4  4  1  4  13  3  4  5  1  1  34  3  14  9  17  \  5  \  1  2 4  BY BY  c  1  1  2  2  10  tf  \  \  W © R ®  5  1  1  B RW  LP  1  BO  6  Y®  3  9  tf  tf tf  5  1  1  9  80  tf  B y<B BY  \ 1  10  9  9 G  \ 1  Q© 1  G OB  DB  1  6  13  2  13  4  6  5  3  4  10  6  2  8  4  6  2  9  6  18  1  12  2  1  5  3  18  4  20  1  3  1  2  6  6  3  B 08  IS. R  8©  7  2  1  3  10 12 5  1  1  3  4  2  6  1  14  2  20  1  2 1  3  1  1  3  2  3  1  3  8  9  1 4  2  1  2  4  16  7  \  2  1  4  \  7  1  2  2  6  \| 1  7  1  AVOIDANCES FIGHTS  3 1  1 2  1 1  1  4  1  1  0  B©  B©  2  ,  9  9  G©  B®  1153 I N T E R A C T I O N S 80 M A N H O U R S  9  JL. LO DB R  1  B BY  8©  9  B®  3 7  9  \  \  \  82  TABLE 11  Numbers o f e n c o u n t e r s  ( d i s p l a c e m e n t s , avoidances and f i g h t s )  i n f o u r dominance h i e r a r c h i e s  Hierarchies  B-C  Sedge P a t c h  Bulldozed Path  Cottonwood S t r e t c h  Displacements  Avoidances  Fights  Total  1043  101  7  1153  142  41  3  186  160  11  3  174  153  17  0  170  83  down  (mostly  fights and  in  the  very  the  B-C  seventh  slightly (Table  females)  more  9  below).  H i e r a r c h y , s i x were  male-female.  involved  the  male-male  However,  i n avoidance  Of  seven  encounters,  males  were  interactions  than  only females  12).  All  seven  hierarchies  few  reversals  involved  corresponds  with  the  not  the  (1922),  and  (1934a).  Nice she  Sparrows  pass  clear-cut  peck-order  Reifel  was  this  should  of  by  period  early after  I t seems  been  to  As  December, likely  that  young  Song Song to  three  old.  months Song  linear  amongst  one  more  might  a  Sparrows  'peck-right*  observations  then  allee  peck-dominance  established  winter.  and  juvenile  juvenile  stable  B-C  arrangement  on  that  are  the  the  Masure  through  they  are  Schelderup-Ebbe  observations  time  There in  of  of  of  concluded  then, have  type  * peck-right*  from  the  time  Island,  many r e v e r s a l s .  This  supercedence by  linear.  a l l interactions  •peck-dominance*  raising,  started  of  classical  relationships  hierarchies  strikingly  reversals.  from  juveniles  4%  (1943),  Sparrows  Extrapolating  are  ; Only  1  Hierarchy  on  (see F i g u r e  the on  groups the  not  expect  reversals  would  1. In encounters between a n y two b i r d s (a a n d B ) , a w o u l d p r e d o m i n a n t l y b e t h e w i n n e r o v e r B. O c c a s i o n a l l y , B would win s u c h an e n c o u n t e r , a n d t h i s w o u l d be c l a s s e d as a reversal. In the matrices, numbers of reversals appear below the diagonal lines.  84  TABLE 12  Analysis of the avoidance interactions i n the B-C Hierarchy  Won  Lost  Total  Males:  83  26  109  Females:  18  75  93  Total:  101  101  -%  2  = 64.74  P < 0.001  85  have  been recorded  b i r d was being Reversals often.  e a r l i e r i n the winter,  decided. d i d not i n v o l v e i n d i v i d u a l s t h a t were seen l e s s  In the B-C Hierarchy,  involved  in  members of reversals  as the rank of each  those i n d i v i d u a l s (9) that  were  more than one r e v e r s a l were a l l prominent, d a i l y  the  hierarchy,  when  under  observation.  Also,  d i d not tend to occur l a t e r than e a r l i e r during the  observation  period.  40 man hours before  Out of 43 r e v e r s a l s on the B-C February 12 showed 24  Hierarchy,  reversals,  40  man  hours a f t e r February 12 showed 19 r e v e r s a l s . •Triangles', dominates  A,  frequently  the  where  and  dominates B, B dominates C, and C  relationship  observed  e. g. i n domestic fowl Juncos  A  phenomena (Masure and  remaining of  stable,  dominance  Allee  1934b);  stelleri, itself.  (Odum 1942);  Jay,  In Song Sparrows, t r i a n g l e s occurred and remained s t a b l e during  Brown  Cyanositta  the h i e r a r c h y  only,  period.  middle  of  the a d u l t male and h i s female were  the dominant b i r d s a t the f e e d i n g The  the  B-C  with both sexes i n v o l v e d .  a l l hierarchies,  territory.  only i n the  the o b s e r v a t i o n  was found t h a t t r i a n g l e s i n v o l v e d b i r d s i n  In  Oregon  were t r a n s i e n t , the l i n e a r s i t u a t i o n soon a s s e r t i n g  Hierarchy, It  in  (Kikkawa 1961).  (1963) found t h a t t r i a n g l e s i n the S t e l l e r ' s  a  hierarchies,  (Sabine 1956); i n Black-capped Chickadees  i n White-eyes, Zosterops l a t e r a l i s ,  are  only  time  station that  located  pairs  from  in  their  adjoining  86  territories  f e d a t t h e same s t a t i o n  placed  on,  pairs,  as  or very  The  the  the  Cottonwood  respected  ranging  might  of  each  table  was  s e p a r a t i n g t h e two Stretch  territory  hierarchy  i n numbers from  determine  Hierarchy.  boundaries,  and  was  composed  two t o t w e n t y - t w o .  t h e dominant  or subordinate  of  Factors status i n  hierarchy are included i n the discussion.  Although intact, did  the  some  (Sabine  Some  maintained  of  juveniles  of individuals  of  Chickadees  et  i n v o l v e d b i r d s o f low rank  Tompa  (1964)  were  involved  low  mentioned  This  i n  position that  only  was n o t t h e c a s e  Song  same s t a t u s i n t h e new old  one.  These  Song  groups  of  low  which  joined. Sparrows  on  Mandarte  on  Reifel  rank  of middle  wandering  group as t h a t  they  Sparrows  movements, b u t a l s o i n d i v i d u a l s 13).  Wood  birds  intergroup  (Table  and  o n l y , and t h e s e  between  individuals  Juncos  group  Not  ranks  a l .  the  1966),  Island.  upper  did  1965),  subordinate  for  i n  I n Oregon  i n t h e new g r o u p  interchanges  only  involved  (Dixon  (fiurton  relatively  between h i e r a r c h i e s  a l l juveniles  palumbus,  their  remained  i n t e r g r o u p movements.  Carolina  Columba  interchanges  Island.  14%  performed  1959),  Pigeons,  groups  interchange  occur.  hierarchies  the  the  d i d not occur.  remainder  juveniles, which  i n  adults  trespassing  when  c l o s e t o , the boundary  happened  Otherwise,  was  perform and  even  b i r d s assumed t h e  they  Most examples a r e o f low ranked  had  held  birds,  in  and o f  87  TABLE 13  Rank i n h i e r a r c h y and sex o f Song Sparrows p e r f o r m i n g intergroup  movements  Sex Hierarchy  position  Top t h i r d  Male  Female  Total  2  0  2  Middle t h i r d  2  1  3  Bottom t h i r d  0  5  5  Total  4  6  10  Each h i e r a r c h y was d i v i d e d i n t o t h r e e s e c t i o n s i n o r d e r  to e l i m i n a t e  the d i f f e r e n c e s i n numbers o f j u v e n i l e s i n each h i e r a r c h y . The a d u l t males and females were n o t used i n the a n a l y s i s ; hence o n l y the j u v e n i l e s were n u m e r i c a l l y m i d d l e , and bottom.  d i v i d e d i n t o the t h r e e c a t e g o r i e s , t o p ,  88  females the  in  particular,  Bulldozed  early  February  mid-February Stretch male, male  Path high was  in  a  An shows  of  that  fights  the  analysis  hierarchy than  of  and  the  gives  the  results  There  i s no of  to  was  are  run  more of  the  in  the  interact  than  hierarchy  Success  in  birds  January, B-C  was  the  found  in  and  in  Hierarchy,  male  in  the  1000 only  a  was  Cottonwood  metres. dominant  territory  in  This  juvenile  (see  (B)  The  (Table  14)  ).  the  B-C  Hierarchy  are  closer  together  more  likely  to  be  apart.  B-C  Hierarchy birds  those  farther  the  involved  in  Therefore,  a  (displacements, to  closer  in  investigate  together  apart.  in  Figure  the 8  analysis.  correlation  are  about  that  interactions.  together  of  {Red/Yellow)  a l l interactions  the  than  the  farther  on  in  in  in  that  that  late  obtain  fights  possibility  interact  (B)  the  fights)  v  hierarchy  "numbers  enough, not  in  male  dominant  significantly  analysis  further  the  Hierarchies  those  avoidances,  scale  as  individuals are  complete  the  hierarchies  Consequences  dominant  distance  interestingly the  the  Hierarchy  found  Hierarchy,  in  but  between Thus,  distance birds  hierarchy  which  are  not  necessarily  which  are  farther  Gaining  in  are  more  and  closer  likely  to  apart.  T e r r i t o r i e s in r e l a t i o n to  Hierarchy  Position  Of  the  seven  hierarchies  (Figure  7  and  Appendix  8)  NUMBER OF INTERACTIONS 10  CO  o  o —I—  _1_  J>0 Oo o o • >• • MOOo DO D X 4^J>t> >&•• • ipt> O D D C J O o • J>o oo o• > " • •a a o• oo-Jpaa (tOOO  •  a*  $00  "D  a  ro co  b 00 x  +  o  CD CO  o o o o•o o o o  -< 11 1  p p  >  oa  J>a oo < > J Oa o o d> • o o JI o o o <po oo  o o  o o > X o •  DO  N)JP O  mm> > 2 s r r > > m rn mm .< .< < 5: "n 2 • m> 5 >>m ? "m > m r~ m r r  Figure 8  Number o f i n t e r a c t i o n s v e r s u s  I  a l l j u v e n i l e s i n t h e B-C H i e r a r c h y  distance within hierarchy fo  89  TABLE 14  Analysis of f i g h t s i n the B-C Hierarchy  Number of juveniles i n the B-C Hierarchy = 22 Number of f i g h t s observed = 7  Random sample s i z e ('pairs' from Random D i g i t s Tables) = 70  Distance i n hierarchy*  1 2  3  Numbers of 'pairs' of b i r d s : Observed  1 0  1 2  Expected  D = 0.76  P  .3  .5  .2  4  .3  5  6  2  1  .7  .4  7-22 0 4.6  0.01 Kolmogorov-Smirnov one-sample test  * Distance i n the hierarchy refers to how f a r apart i n numbers one member was from the other member of a 'pair'. Thus, i f the distance i n the hierarchy i s 1, then the two birds involved i n a f i g h t were next to each other i n the hierarchy. I f the distance was 5, then there were 4 other birds i n the hierarchy between the 'pair'.  90  determined, Sedge  four  Path)  Narrows)  were was  (Big  Alder,  i n the Simultaneous i n  the  Stretch)  was  the Simultaneous  and area (1)  incorporated  Simultaneous  Removal  explained  i n PART  the Simultaneous  holders the  had been  area,  The  area  i n the  one  (Marsh and  seventh  one (B-C)  and t h e C o n t r o l Simultaneous  met,  Removal  Area  A, t e n m a l e s  Removal  removed.  area  established  after  Of t h e s e  territories  the resident  t e n , seven  territory  had f i g u r e d  in  hierarchies.  The  dominant  territories: second  males  obtained Hierarchy  The Hierarchy  males  i n  obtained  a  dominant  male  stretch  hedgerow.  was c o n s p i c u o u s member  this  male  was  when  the removal  a n d B-C  the f i f t h  and  B-C.  The  Hierarchies also  male  i n  the  B-C  territory.  the only  prominent  Big Alder,  Path  and f i n a l l y  Removal  of  Patch,  the Bulldozed  territories,  was  i n the following hierarchies obtained  No-Name, S e d g e  Simultaneous  male  Removal  two t e r r i t o r i e s  area,  Removal  was o n t h e C o n t r o l .  No-Name, a n d  and one on t h e C o n t r o l .  As in  Path,  Removal  Successive  (Cottonwood where  Bulldozed  top bird  area  i n  not t o obtain  by i t s wandering  not present  in  the  As e x p l a i n e d  of three  took  (Red/Yellow)  hierarchies. along  place.  the Bulldozed  hierarchies  a territory  earlier,  this  behaviour,  on  the  along  this  particular  having  been  I t i s conceivable  the Simultaneous  Path  Removal  a  that area  91  Three part on  males  that  established  i n the hierarchies. the  Control,  and  One  was  simultaneous  removal  occasionally  throughout  Removal  area,  b u t was  during  the time  unbanded  male  hedgerows quite the  study  males  position  of  Island this  join  made.  of the seen  Simultaneous  feeding The  station  last  was  i n the region  the study  individual  December  was  the  any  territory  that  paired  the removals,  only  one  in  the  February Table  interactions in  seen  on  January  the  adult  15  This  Sedge  3.  spent  shows  area,  and  the winter  29,  female's male  and  and  date,  an  where i t  i s  outside  after  February  during  replacement  (2)  S u c c e s s i v e Removal  i n early  the removals,  member  had  a  low  Hierarchies  moved  difference  that  territorial  prominent  she r a p i d l y  i t i s thought  position  a  B u l l d o z e d Path  of the Bulldozed Path  the  one  lost  t h e new  (Black/Blue)  a significant  and  area  this  with  had ^ e e n  female  Patch  After  won  the  the adult  Only  along  x  t h e week  second  at  females  female  of  recorded  a  until  The  winter  established  hierarchies.  scale.  in  banded  take  area.  after  before  was  o b s e r v a t i o n s were  that  the seven  the  the  not  flestham  possible  Of  in  from  d i d not  not r e s i g h t e d  experiment.  that  that  of thes  territories  up  the  i n the  number  The  adult  3.  H i e r a r c h y was Black/Blue  assumed  February.  On  she  with  paired  last  the  death  one  of  males. Areas  hierarchy  was  determined  along  the Successive  92  TABLE 15  Change i n s t a t u s o f a j u v e n i l e female f o l l o w i n g d i s a p p e a r a n c e o f an a d u l t  territorial  B e f o r e Feb.3  Interactions:  X  The a d u l t  2  the  female  A f t e r Feb.3  Total  Win  2  23  25  Lo*se  11  5  16  Totals  13  28  41  = 17.09  female was  P < 0.005  l a s t seen on January 29. J u v e n i l e  ( B l a c k / B l u e ) presumably  female  assumes the a d u l t female's r o l e i n e a r l y  F e b r u a r y . The j u v e n i l e female wins s i g n i f i c a n t l y more i n t e r a c t i o n s , not  o n l y w i t h b i r d s below h e r , b u t p r i m a r i l y w i t h b i r d s above h e r ;  hence h e r change i n s t a t u s .  93  Removal were  area.  juveniles,  succeeded other  female  winter  across  with  which  Ewen  birds,  females.  a new  male  One after  It i s possible were  Slough  of which  unbanded,  i n the F a l l  of  these  the that  two  females  removal, the  could  Removal  only  the  replacement  have  Area  spent  the  (Figure  1).  Control  11  Of  Hierarchy,  In corpse this  juveniles  that  took  none  were  seen  after  May,  however,  early was  found  on  territory,  replacement  The  B-C  the only  amongst  five  have  and  male  had  4).  one  males  obtained  given  territories hierarchies  the are  one  5,  been  third  took  bird  more  chance  part  established  than  This  (the  taken  ever  female.  This  Hierarchy.  of the hierarchies, one  territorial  hierarchy,  i n the h i e r a r c h y , i s  highly  four  and male  out of  12  from  the  unlikely  to  0.01). strongly  opportunity, the  had  the adult  Analysing this  (P =  then,  a male  males d i e d  i n t h e B-C  the largest  Stretch  experiments.  of the adult  May  territories.  evidence,  i n the Cottonwood  the removal  On  supplying  that  part  apparently also  H i e r a r c h y was  o c c u r r e d by  The that,  May  the replacements.  juvenile top  of  of five  both  disappeared.  most  was  and  in pairing  birds,  (3)  I t consisted  the  supports juveniles  dominant  individuals  during the  pre-breeding  the hypothesis that in  obtain dominance  season.  94  IV  DISCUSSION  (a) The F u n c t i o n a l S i g n i f i c a n c e of Dominance H i e r a r c h i e s Gartlan social  (1968) suggests  dominance  available  can  subordinate although  food  function  when  l i m i t , regarded  the  (1962)  suggests  a  social  guillotine  times  of  as  whilst  food  that for  shortage, few  starve  or  numbers of the s p e c i e s are up a g a i n s t  the  bottom  of  concurring  to  the  the the  f u n c t i o n of  to r e l a t e p o p u l a t i o n s i z e  during  (1966),  i n d i v i d u a l s near  is  Wynne-Edwards  individuals Lack  emigrate  hierarchy  resources.  hierarchies  that the primary  the  that  peck-order  t h i s as a consequence,  not  a  function.  Lack  argues t h a t the h i e r a r c h y a r i s e s as a response to s c a r c e  food  resources,  and  Wynne-Edwards, pre-adaptive  not  however,  however,  differentiate  the  food  that  the  is  cause  shortage  of  hierarchy  frequently  and  effect  females!  food shortage, than  males.  The  is  guillotining  both  difficult of  I t i s even more  hypothesis  supporting  the  to  dominance  difficult  to  with the h i e r a r c h i e s i n  Song Sparrows, where a l l the bottom, hence subordinate are  them.  privation.  t h a t can be put forward it  between  anticipation  maintains  h i e r a r c h i e s i n most i n s t a n c e s . relate  an  f o r c o n d i t i o n s of  There i s evidence arguments;  as  of subordinates  birds,  i n times  then, would presumably e f f e c t females f a r  of  more  95  A  further  reducing  aggressive  conserving  and  aggressive Calhoun  states  that  the  and food  because  i t saves  lose  with  anyway,  elsewhere. between the are  and  animals. the  identify that  a  might  any  one  of  (b)  Factors  wasting  energies  are  on  can  Sparrows  i s  as  the  to  not  determining  fights  various  are  for  he  both they  those  lower  they  would  for  that  searching aggression  because  of  rare, reversals  like ways  functional  reduce  aggressive  determine  territory  for  reduced  behaviour,  Sparrows, to  been  when  stable.  in  be  has  Guhl  because  which  argue  1955;  g.  lines  conserved  to  reducing  (e.  value  fights  be  reasonable  s u r p l u s , or  them  survival  time  function  could  a  similar  and  hierarchy  obtain  birds  fighting,  little  hierarchical  hierarchy  along  has  particularly  Song  domestic  in  is  Anthony  very  Song  could  (e.g.  i s a  There  important  mammals  a l s o argued  in thus  this  hierarchy.  are  on  that  higher  seems  For  and  the  argued  f o r those  It  the  on  be  function  individuals,  individuals,  their  Dominance behaviour,  1971),  and  hierarchies, few,  i n work  them  individual  between  i t can  peck-order  weaker  obtain  again  hierarchies  f u n c t i o n , of  (1954)  that  that  hierarchies  Kinsey  Lack  stronger  a  encounters  1971;  1956).  but  not  evidence  is  encounters  energy,  consequence, good  argument  given  the  territorial for  different  significance  of  interactions,  to  dominant  individuals  the  opportunity.  the  Dominance  Data  unequivocal. position  in  Hierarchy  for  96  It  has  Sparrows  already  been  i s essentially  considering dominance  some  of  shown  linear the  that  the hierarchy  and s t a b l e .  factors  or subordinance of the  which  I t  i s  might  individuals  i n Song  worth  now  determine the  comprising  the  hierarchy. Sex  and p o s i t i o n  In  Song  juvenile 22  is  Sparrows,  females.  juveniles,  arrangement (e.  i n hierarchy  g.  from  c a n be  o r i n Steller»s  takes  the  rank  M£ii£2ia2[» are  graded  of i n  some  the  male  males.  Black-crowned  pairs  this  species  1942; B r i a n  where  the  with.  female  In c e r t a i n Nycticorax  1939) , m a l e s  sex,  of  1955)), i t  Night-Herons,  and Schmidt to  passerine (Tordoff  over  out  Although  1 9 6 6 ; Odum  1964),  she  dominant  f o r example,  other  (Dixon  (Brown  regard  were  curyi.rostra,  species  Jay  ( N o b l e , Worm without  i n  Loxia  i n Parus  1949),  as  t h e t o p 14 w e r e  found  Red C r o s s b i l l ,  instances,  males  I n t h e B-C H i e r a r c h y ,  12  not the case  juvenile  and females  suggesting  an  asexual  fowl that  cocks  situation.,  Guhl normally two  (1956) peck  hens,  hierarchies,  Sparrows involved  An  found  cannot  for  domestic  and t h e r e f o r e  one f o r each be d i v i d e d  a breeding flock  sex.  i n this  The  usually  hierarchies  manner,  do n o t has  i n  Song  sexes  were  by p a i r i n g  with  as both  i n the interactions.  individual  can climb  the s o c i a l  ladder  97  a  dominant  instance pairing  of  of  a  with  juvenile adult  bird  a  male  female  had  mate,  moving  scale  the  female.  disappeared  paired  this  with  after  the  the  up  the the  in  adult  spring  the  replacement  young  Size  and  position  hierarchy  (1963) of  the  relates  dominance Song  hierarchy period  (see  male  on  removal  the  an  after  Sparrows, after  January  interactions  period  with  in  In  in  success  after  and,  up  adult  Her  significantly  his  Brown  ( B l a c k / B l u e ) moved  3.  apparently  opposite sex.  dominant  female  February  the  a the  29  to  increased  Table  14).  She  disappearance  experiment,  she  of  paired  male.  f  Size  has  hierarchical Hebb  been  found  position  1948),  and  Forest  (1969)  found  that,  i n the  wings  than  subordinates.  weight  (Figure  and  relationship  between  hierarchy  was  found.  largest  or  heaviest  be  a  factor  in Bottle-nosed  i n New  4)  to  or i t  birds  1972).  vary  were  the  be  longer  considerably 3) ,  and  ones  but  position  argued  and  Fretwell  had  (Figure  winglength cannot  (McBride  dominants  Sparrows  winglength  weight Thus,  (Tyler  Junco,  Song  in  Dolphins  Ponies  Oregon  in determining  that  that  in no in the  obtained  territories. Age  and  position  Age domestic passerines  and fowl (R.  in  hierarchy  dominance  have  (Shoemaker I.  been  1939),  Stewart  pers  found but  not  coram).  to  correlate in A  in  short-lived more  subtle  98  relationship Grouse, is  a  Centrocercus centripetal  mating  centre  In of  between  Song  a t most  o f t h e ^Lek^)as  Age  juveniles  the  first  prior  movement  two a d u l t  the  birds  were  of  only  dominant  over  could  evidence,  hierarchy.  T h e a g e o f t h e two  B-C  Hierarchy  determined  (see  second  was  was  bird  the  calculated  of  he  6  was  those  the  banded  when  respect to  brood,  a second younger  or third  the ideas of  that,  position  males  i n  in the  i n t h e summer age  be  of age).  The  juvenile  top  brood,  the  of  could  but the  or third than  from  next.  calculations  dominant  rest  to suggest  their  a  comprised  second  with  meagre,  were  of a f i r s t  60 d a y s  from  i n determining  for  either  t o be a b o u t  Nevertheless,  as they  Appendix  product  i s  with  top juvenile  development  a product  towards  i f the individuals  albeit  the  of  there  the years.  group  t o be d i s c u s s e d  age i s n o t i m p o r t a n t  at a stage  over  be c o n f u s e d  juveniles,  a r e known,  where  males  be i m p o r t a n t  for  1972,  arise  i n t h e Sage  i n any one t e r r i t o r y , and t h e  and e x p e r i e n c e ,  some  1970),  territorial  f o r example  R e s u l t s , however,  i s  occurs  (Wiley  vacancies  could  themselves,  occupancy  dominance  any one h i e r a r c h i c a l  then  broods  There  and  urophasianus,  Sparrows,  juveniles.  broods.  age  bird  and  was  second  bird.  i n the  largest  hierarchy. Prior  occupancy  Experience familiarity  and e x p e r i e n c e  and/or  with  prior  territory)  and p o s i t i o n  occupancy and  the  i n the hierarchy  (prior  residency,  relationship  with  99  hierarchy These  rank  has  relationships  (1971); Krebs  in  birds,  e.  have g.  much a t t e n t i o n been  the  1965),  Great  Blue Jay  dominance  order  the  Brown  (Brian  Parus  (Brown  place;  with  i t s  the  distance  Territorial  adults  could  even  food  seems  juncos their  The  at  was  u n l i k e l y that  food,  not  recent  fish,  and  e.  even  feeding  territories recognition  juveniles  they  years.  g.  in  Phillips  man,  e.  (Colguhoun  p o s i t i o n of  the  distance  of  the  g.  its  enticed in  were  by  not  the  of  their  of  the  feeding  The  incentive  was  apparently  not  strong  of,  and  and  the  rest  therefore  the  of  enough  the  not  in  territory varied  be  In  Song  apply.  territories territory.  presence  of  behaviour  of  then  territorial  would  bird  not  neighbour's  aware  for,  and  its  does  noisy  respect  1942),  territory.  from  their  the  (Dixon  individual  probably be  Chickadee  stations.  comprising  non-territorial,  Carolina  from  supplied  particularly the  the  rank  phenomenon  when  1949),  with  this  the  (1964);  1963),  Sparrows,  the  in  caeruleus,  corresponds  feeding  inversely  Tit  Tit,  Steller's  It  found  in  (1973).  In  from  received  to to  leave  overcome  boundaries.  hierarchy involved  in  were this  idea.  Sabine first  on  groups  that  thought  (1959) the  by  suggested  wintering  arrived Kikkawa  that  grounds  later. (1961)  Prior to  those were  juncos  in,general  residency, be  a  that  dominant  however,  factor in  arrived  was  to not  determining  100  hierarchy  position  advantage area  was  conferred masked  The  situation  position  in  juveniles their  Song  can  own  formed  birds  with  the  large  number  regarding  same  during  the  of  the  case,  an  area,  to  Introduction),  would  have  supporting  an  or  It  seems,  a  factor in  however,  cannot  factors  The lower  top  rank.  the  first, one  during of  idea  that  hierarchy.  and  ruled the  birds  out  Further,  be  the  are  as  a in  birds  in  likely  along  fall  territorial  the  suggestion to  that  is  arrange  have  those  of  the  familiarity  dispersal  that  juveniles phase  (see  concentration  arrivals.  prior  position  possible  No  data  of  occupancy a  juvenile  prior  is Song  occupancy,  factor.  hierarchy  genetically  social  top  most  only'  available.  long-term the  that  would  centres  later  fact  previous  Short-term  position  may  the  over  this  argue  an  areas  appear for  with  hierarchical  are  this  individual  determining  be  groups  If  regard  possibly  therefore,  Sparrow  Other  no  advantage  in  adults.  and  were  of  any  present,  the  certain  juveniles  at  refuting  by  recrudescence  could  General  birds  adults  that  familiarity  occupancy  in  without  arrive  of  juvenile  time  because One  happened  all  The  the  positions  suggested  complicated  territorial  then  familiarity.  not  is  he  previous  prior  considered.  the  hierarchical  that  on  Sparrows  be  about  behaviour  with  White-eyes;  territories.  hedgerow,  the  by  in  rank  different could  be  from  birds  of  genetically  101  inherited. not  an  by  hypothesis juvenile  based  aggressive  shows t h a t  each an  at  Sparrow's  the  an  argued  that  individual,  evidence features  position  dominance  in  that the  dominance  but  supporting  a  or  might  is  'role*  refuting  determine  dominance  is a  bird  was  and  is a the  the  involved.  behaviour, the  bird  top  the  a a  hierarchies  not  number  show  because  incidence  of  end  of  the  then,  there  based  difficult  dominant  of  as  to  on  determine  bird.  Figure  interactions in  Krebs  the his  i n Song  are  9  p o s i t i v e c o r r e l a t i o n between  However,  hierarchies  higher  is  significant  i n d i v i d u a l need  that  hierarchies  i t i s frequently  aggressive  there  aggressive  there  no  genetic  in hierarchy  alpha  fact  on  have  encounters,  most  position  I  of  has  Island.  Although  the  however  feature  him.  Song  Reifel  if  (1967)  intrinsic  acquired  on  Sade  (1973)  which  points  out,  highest  freguency  position i s  stable.  Sparrows  are  interactions  stable,  hierarchy,  requires  The  and  concerning  of  yet birds  further  investigation.  In  summary,  evidence  in  this  determine  the  rank  hierarchy. prior  Males  occupancy  short-term  prior  study that  an  dominated were  not  occupancy  appears  supporting  to  be  any  little f a c t o r (s)  i n d i v i d u a l Song females. thought and  Sparrow  Size, to  possibly  conclusive  be  age,  that held  and  or  in  the  long-term  important,  genetic  might  but  aggression  102  features  were  not ruled out.  Finally,  i t i s worth  hierarchies, individuals obtaining  (c)  and  territories  Dixon  subordinates  position were  at  idea  that  the  of  the  t h e dominant  successful  ones  in  arose.  as often  from  as  bird  was  Guhl  first,  followed  appeared  the  B-C  just  1956;  of the  not related  Table  that  i s a  1955;  The appearance  station  there  16  by  first  to i t s  gives  the  on t h e t e n d a y s  that  Hierarchy. as o f t e n  In  fact,  as dominates,  and  males.  found  Serinus  that,  serinus,  success the  feeds  hierarchy.  first  (1939)  breeding  order.  feeding  of animals,  Anthony  individual  observing  arrived  canaries,  evidence  the  groups  (e. g.  dominance  i n  Shoemaker  higher  the  of the f i r s t  subordinates  of  order  i n descending  i n the  spent  females  outcomes  the opportunity  The dominant  Sparrow  position  when  hierarchical  feeding  1965).  Song  the  of the Hierarchies  certain  pronounced  particular  at  i n t h e h i e r a r c h i e s were  Outcomes  In  in  looking  than  present  in a  laboratory  the dominant  males  subordinates.  study  population had a  There  to support  much  i s  or refute  no this  suggestion.  In not  only  certain  lek species,  i s i t important  (e. g.  f o r males  Sage  Grouse,  to obtain  a  Wiley  1970),  territory  to  103  POSITION  Figure 9  IN  HIERARCHY  P o s i t i o n i n h i e r a r c h y and number o f i n t e r a c t i o n s : t h e B-C  Hierarchy  The l i m i t a t i o n s o f t h i s a n a l y s i s a r e acknowledged. Each b i r d d i d n o t spend an e q u i v a l e n t amount o f time a t t h e f e e d i n g s t a t i o n . Some b i r d s f e d w i t h o u t an i n t e r a c t i o n o c c u r i n g . The m a n i f e s t a t i o n s d i s c u s s e d i n the t e x t .  o f aggressive behaviour are  104  mate,  but  i t  i s  territories  to  species  birds  of  pre-requisite many in  of  the  in  for  successful which  species,  in  v  obtaining  winter  Odum  flock  a  the  of  results  correlation,  in  the  that  the  dominant  ones  pre-breeding  summary,  then,  in  the  at  Hierarchies  presence  stable,  with  In  the  few  long-term  in  determining  Dixon did  a  are  several  territory  is  breeding.  dominance  in  the  been  that  a In  hierarchy  hierarchy  shown  the  top  in  (1965)  found  (1964)  study  in  obtain  hierarchies  a  few  males  in  established  similar results Song  further  that  and  only  three  Chickadees  Tompa  dominance  a  favoured  determined.  has  individuals  in  There  hence  of  dominance  reported  as  groups  of  juvenile  favoured  formed,  Sparrows  strengthen  this  territories  established  .  are during  Song  localities  which  by  Sparrows along  congregated  the  mid-winter  hedgerow.  were  linear  reversals.  hierarchies,  and  been  present  certain  were  and  certain  season.  In  fall  mate,  has  spring.  obtain  a c q u i s i t i o n of  Black-capped  Chickadees,  Carolina  the  to  matings.  territory  (1942)  in  the  a  c o r r e l a t i o n between  in  the  the  season  territories  The  important  obtaining  pre-breeding  populations. a  ensure  these  The success  also  prior the  males  occupancy rank  dominated  were  that  an  not  females.  thought  individual  to  Size, be  held  age,  important in  the  104  TABLE 16  F e e d i n g o r d e r o f h i e r a r c h y : appearance o f f i r s t a t f e e d i n g s t a t i o n s o f the B-C  bird  Hierarchy  Sex Position of b i r d i n hierarchy  Totals:  Number o f days  Male  Female  F i r s t seven  3  3  0  Middle eight  4  2  2  L a s t seven  3  0  3  22 b i r d s  10  105  hierarchy. aggression  The  Short-term  prior  occupancy  features  were  removal  experiments  showed  that  those  spring  on  the  not ruled  yearlings Spring  individuals  in  the  pre-breeding  season.  that  possibly  genetic  or  out.  and  subsequent  obtained  Removal  hierarchies  and  Area  replacements  territories were  established  the  in  the  dominant  during  the  106  LITERATURE CITED  Alexander, R. 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Auk  d i s p e r s i o n of Ecology 53:  feeding  territories  Competition between E c o l o g y 52: 320-29.  1939. Social 56: 381-406.  hierarchy in  two  among  chipmunk  flocks  of  the  Simms, E. 1965. E f f e c t s o f t h e c o l d w e a t h e r o f 1 9 6 2 - 6 3 on the B l a c k b i r d p o p u l a t i o n of D o l l i s H i l l , London. Brit. B i r d s 58: 33-43. Smith, C. C. organization E c o l monogr.  1968. The i n the genus of 38: 31-63.  adaptive nature of social tree squirrels Tamiasciurus.  S m i t h , R. F. J. 1970. Effects of food availability on aggression and n e s t b u i l d i n g i n Brook S t i c k l e b a c k , Culaea inconstans. J. Fish. Res. Bd. Can. 27: 2 3 5 0 - 5 5 . S m i t h , S. M. 1967. Seasonal changes i n the s u r v i v a l Black-capped Chickadee. C o n d o r 69: 344-59. Smyth,  M.  1968.  The  effects  of  removal  of  individuals  of  the  from  a  115  population of Bank Voles, Anim. Ecol. 37: 167-83. Snow,  D.  W.  S n y d e r , W. scaled 19.  1958.  A study  Clethrionymus  of blackbirds.  glareolus.  London.  D. 1967. Experimental habitat improvement quail. Tech. Publ. Colo. Game F i s h P k s .  Sommer, R. 1967. 145-52.  Small  Stefanski, R. territory in 259-67.  A. the  Stenger, J. Ovenbirds  group  ecology.  J.  Psychcl.  1967. Utilization of Black-capped Chickadee.  Bull.  for Dep.  67:  the breeding Condor 69:  1958. Food habits and available food of i n relation to territory size. Auk 7 5 : 3 3 5 - 4 6 .  S t e w a r t , R. E. and J. W. Aldrich. 1951. repopulation of breeding birds i n a spruce community. Auk 6 8 : 4 7 1 - 8 2 . Suthers, R. A. territories 232-37.  of  1960. the  Removal and - f i r forest  Measurement of some Song Sparrow. Wilson  lakeshore Bull. 72:  S y m o n s , P. E. K. 1968. Increase in aggression and in strength o f t h e s o c i a l h i e r a r c h y among j u v e n i l e A t l a n t i c salmon d e p r i v e d o f food. J. Fish. Res. Bd, Can. 25: 2387-401. S y m o n s , P. E. K. 1971. density to available J. Fish. Res. Bd.  B e h a v i o r a l adjustment of population f o o d by juvenile Atlantic salmon. Can. 28: 569-87.  ° Tester, J. R. a n d A. Watson. t e r r i t o r i a l i t y o f Woodcock, S c o l o p a x roding behaviour. I b i s 115: 135-7. Tompa, F. S. 1962. Territorial controlling factor of a local Sparrows. Auk 7 9 : 6 8 7 - 9 7 .  1973. Spacing rusticola based  behaviour: population  the of  and on  main Song  Tompa, F. S. 1964. F a c t o r s d e t e r m i n i n g t h e numbers o f S p a r r o w s , M e l o s p i z a m e l o d i a , o n M a n d a r t e I s l a n d , B. Canada. Acta Zool. Fenn. 1 0 9 : 1-73.  Song C ,  Tompa, F. S. 1971. Catastrophic mortality population consequences. Auk 8 8 : 7 5 3 - 5 9 .  i t s  Tompkins,  G.  1933.  Individuality  and  and  territoriality  as  116  displayed  by  three  passerine  species.  C o n d o r 35:  98-106,  T o r d o f f , H. B, 1954. S o c i a l o r g a n i z a t i o n and b e h a v i o u r in a flock of captive, n o n - b r e e d i n g Red C r o s s b i l l s . Condor 56: 346-58. T y l e r , S. J. the New 187-94.  1972. Forest  The b e h a v i o u r and Ponies. Anim.  W a t s o n , A. 1964. A g g r e s s i o n and Grouse. N a t u r e 202: 506-7.  social organization Behav. Monogr.  population  of 5:  r e g u l a t i o n i n Red  Watson, A. 1965. A p o p u l a t i o n study of Ptarmigan, Laggpus mutus , i n S c o t l a n d . J. Anim. Ecol. 34: 135-72. W a t s o n , A. 1967. Red G r o u s e .  T e r r i t o r y and p o p u l a t i o n N a t u r e 215: 1274-75.  regulation i n  Watson, A. a n d D. Jenkins. 1968. Experiments c o n t r o l by t e r r i t o r i a l b e h a v i o u r i n t h e Red Anim. Ecol. 3: 5 9 5 - 6 1 4 . Watson, A. a n d G. R. Miller. a g g r e s s i o n i n a f l u c t u a t i n g Red Anim. Ecol. 40: 367-83.  the  on p o p u l a t i o n Grouse. J.  1971. Territory size Grouse population.  and J.  W a t s o n , A. a n d R. Moss. 1970a. Dominance, s p a c i n g behaviour and aggression i n relation to population limitations i n vertebrates. IN: Animal p o p u l a t i o n s i n r e l a t i o n t o t h e i r food resources. A. Watson (ed) Blackwell, Oxford: 167-218. Watson, A. a n d R. Moss. t e r r i t o r i a l behaviour, Grouse. IN: Behaviour b y a n i m a l s a n d men. A. York - London. W a t t s , C. R. a n d A. W. of T u r k e y s . Scient. Weeden, J. Sparrow.  S. 1965. Condor 67:  1970b. Spacing as a f f e c t e d by habitat, and nutrition in Red and environment: t h e use o f space H. E s s e r (ed) P l e n u m P r e s s , New  Stokes. 1971, The social Amer. 224 ( 6 ) : 112-8. Territorial 193-209.  behaviour  of the  Wessell, J. P. a n d H. Leigh. 1941. Studies of the organization of the White-throated Sparrow. Wilson 53: 222-30. Wood-Gush, D. G. M. 1965. domestic b i r d communities. 14: 2 1 9 - 3 1 .  The Symp.  order  Tree  flock Bull.  s o c i a l organization of Zool. Soc. Lond.  1 17  W i l e y , R. H. Jr. mating in the Onpubl. Ph. D. York. W y n n e - E d w a r d s , V. C. s o c i a l behaviour. Y o u n g , C. W. 1970. Tadorna tadorna.  1970. Territoriality and non-random Sage Gronse, C e n t r o c e r c u s urophasianus. T h e s i s , The R o c k e f e l l e r U n i v e r s i t y , New  1962. Animal d i s p e r s i o n i n r e l a t i o n O l i v e r and Boyd. Edinburgh. Territoriality in I b i s 112: 3 3 0 - 5 .  the  Common  to  Shelduck  Young, E. C. 1972. Territory establishment Mccormick's Skua. I b i s 114: 2 3 4 - 4 4 .  and  Zaret, T. M. a n d A. , S. Rand. 1971. tropical stream fishes: support for exclusion theory. E c o l o g y 52: 336-42.  Competition in the competitive  Zimmerman, J. L. dependant e f f e c t  1971. The territory and i n S p i z a americana. Auk 8 8 :  stability  in  i t sdensity 590-612.  118  APPENDIX plants  1.  List  along  The  occasional  J*i  Finally,  Populus  Pseudotsuga  herbaceous  plants  important  ones,  Vicia  spp.,  speciosa,  of  :  SE®ricanus,  Care x  of  small  racemgsa,  consisted  mainly  R.. l a c i n i a t u s ,  albus,  trees  the  and  were  A In us  Lonicera  principally rubra,  and  .  and  detailed  was n o t however;  vegetational  attempted. are listed  lanceolatum, margaritacea, species  adjacent  spp.,  o f mature  P.. t r i c h o c a r p a ,  and s e v e r a l  Salix  species  spectabilis,  stands  with  Sambucus  Shrubs  the  Cirsium  marsh  hookeriana,  Rubus  frequently  shrubs,  T h e main  spp.  important  thick,  and  spp., Symphoricarpos  Anapholis  aguilinum,  The  Salix  a  Rosa  menziesii  complete  trees.  and most  marsh.  of  trees  and Prunus  tremuloides,  A  small  stolonifera,  macrophyllus,  common  and a d j o i n i n g  o f mature  douglasii,  Cornus  o f most  comprised  of  AInus r u b r a ,  were  spp.  was  tangle stand  Crataegus of  t h e hedgerows  hedgerow  impenetrable  trees  of species  The  Urtica  lyalii,  Eguisetum  lyngbyei,  Typha  I^ysichiton  the and  lanatum, Petasitgs  aryense,  i n cleared  of  common  : Heraclgum  of grass  gale,  most  here  t o t h e hedgerow  Hyrica  survey  Pteridiura areas.  was c o m p r i s e d 1atifolia,  k a rots h a t c e n s e  mainly Scirpus  .  119  APPENDIX 2  Weather data from the M e t e o r o l o g i c a l O f f i c e , Vancouver I n t e r n a t i o n a l A i r p o r t  Mean monthly temperatures  Jan.  Feb.  Mar.  Apr.  May  June  July  Aug.-  Sep.  Oct.  Nov.  Dec.  —  39.0  44.6  44.6  54.7  58.3  62.3  64.0  54.3  46.1  43.0  35.2  1973: 36.0  40.5  43.2  47.4  53.7  Normal (37 y r ) :36.3  40.0  42.5  48.1  54.3  59.5  63.4  62.7  57.6  50.1  42.9  38.9  Apr.  May  June  July  Aug.  Sep.-  Oct.  Nov.  Dec.  1972:  Monthly temperature :ranges  1972 Max.  Jan.  Feb.  Mar.  —  56.0  62.0  ft 60.9  75.0  73.4  81.4  80.7  77.9  62.1  56.6  51.8  45.9  47.9  48.0  35.0  30.2  25.4  12.0  17.4  30.0  30.8  41.1  1973 Max. 52.9  58.1  53.2  60.6  76.6  Min. 13.1  24.3  28.1  33.9  36.8  Jan.  Feb.  Mar.  Apr.  May_  June  July  Aug.  Sep.  Oct.  Nov.  Dec.  —  5.25  6.98  45.1  1.17  1.84  3.20  0.92  3.14  2.09  4.61  11.34  1973: 5.43  2.69  2.69  0.51  1.91  Normal: A.95  4.29  3.51  2.39  1.87  1.78  1.17  1.46  2.41  4.81  5.47  5.96  —  0.3  0.4  1.9  1973: 4.7  TR  0.1  TR  Normal: 8.8  3.1  2.1  TR  5.28  7.05  4.70  1.27  1973: 5.92  2.69  2.70  0.51  1.91  Normal: 5.80  4.57  3.69  2.40  1.87  Min.  ~  Precipitation  R a i n f a l l : 1972:  Snowfall: 1972:  Totals:  1972:  —  TR  6.4  0.9  5.7  1.84  3.20  0.92  3.14  2.09  4.61  11.82  1.78  1.17  1.46  2.41  4.81  5.56  6.51  120  APPENDIX  3  Destruction  of  a  feet  of  stretch  of  dyke  ( Bulldozed ,  Area').  In  June,  Removal of  the  This  stretch 10  of  gives  bulldozing  3 4  was  males  birds  rebuilt.  the  Area  and  no  in  less  than must  bird  along  when  the  resident  the  fall  1973  as  about  2000  The Control  a  spring.  the  with  a  on  total  gravel  number  of  that  the  Fall  detruction and  Song  rock.  Sparrows;  existed  before  a  of have  the  this  were the  hedgerow  stretch.  A l l  relocated.  summer  close  adjacent was  to  to  removed  as the  the  on  a  One  cryptic  Bulldozed Bulldozed  October  3/1972  early  March  experiments.  territorial south  third  bird,  the  female  male  about this  4  spent  male  the  after  along  these  (Green/Green)  strongly metres  3  territory  removal  male  known  June.  to  apparently  i t  boundaries  were  late  over  A second  entailed  supported  1 female  took  during  he  hedgerow  started.  non-territorial as  This  territory  (Black/Orange)  Area,  dyked  replacing  hedgerow  disappeared  However, male  800  v e g e t a t i o n , and  Figure the  Area  1972,  300  bird on  metres  male's  was  located  in  in a stretch  Westham  (LP/White) from female  her  of  shrubbery  Island.  ,  paired  destroyed  disappeared  with  a  territory, in  early  F i g u r e 10  Bulldozed before  A r e a : the arrangement o f t e r r i t o r y  the dyke r e b u i l d i n g o c c u r r e d  boundaries  122  These non-breeding,  data  suggest  that  adults  can  survive  as  birds  through  the  summer,  in  the surplus  of birds  non-territorial  spite  of the fact  seemed  t o be  that  composed  only  of  juveniles.  \  on  the study  area  7  123~  APPENDIX locate  4  Possible  i n the  Some locate 1.  fall  fall  October,  found  are  that  September,  Island  were  which 4  partial 2. and  females  why  are  not  given  removals  1972,  and  were  to  to  of  to  seemed  Removal  by  Moulting  Nice  (1937)  i n August  finishing  moulting  However,  moult  place.  not  September  Simultaneous  out  moulted  bird  occurred.  difficult  find.  i n Ohio  occasional  were  took  difficult  Sparrows  most  the  difficult  here.  Casual observations of  from  were  females  a l l females  the  the  made i n  until  birds  on  to  the  be  and  Beifel month  one  male  removed  on  area  was  s t i l l  in  moult.  Females  are  generally  apparently  spring  that  Song  moulting  October  1972  extremely  with  mid-October.  reasons  when  the  why  1972.  of  I t i s possible  individuals  in  of  possible  i n the  early  reasons  (Nice  3.  Females  and  early  Tompa  desert  their  Thus, territorial  Nice  100-700  i t males  mates  to  d i d not  most  showed females  adjacent  metres  seems  fall  than  males,  than  in  the  1964).  did  3056 w e n t  to locate  responsive i n the  (1943)  instances  territories. territories  less  1943;  fall.  possible  are  more d i f f i c u l t  often that  i n the in only  return  to  late 31%  out  their  territories,  and  summer of  55  former 33^  to  distant.  quite  likely  that  have  females  i n the  several  of  fall  1972.  of  the  124  APPENDIX and  5  other  Interspecific seed-eating  Song of  Sparrows  birds that  common  frequented most  the  stations,  at  of  the  observations,  inetrspecific  dominance  most  which  and  tsong  stations.  There  Sparrow,  the  Sparrows  of the other  stations.  J u n c o , Fox  mexicanus,  occurred  From  feeding  between  the feeding  Occasionally, other  Carjgodacus  lincolnii,  at  i n t e r a c t e d with  Oregon  abundant..  Finch,  passerines  fed at the  species,  interactions  junco  were  and  three  Towhee,  was  by  seed-eaters  species  that  far  such  L i n c o l n ' s Sparrow,  as  the House  Helgspiza  stations.  a  very  could  general  be c o m p i l e d  list thus  in  order  of  :  Towhee Fox  Sparrow  Song  Sparrow  Lincoln's  however,  certain  Song  Fox  Sparrows.  On  Fox  Sparrow,  but  precedence  over  indifferent and  would  to  feed  Juncos  House  Finch  Oregon  Junco  Sparrows  would  occasion, this Song  was  a  Song  Sparrows.  were  leave  threaten  Sparrow  r a r e , and  the t h r e a t e n i n g and  Sparrow  would  Towhees behaviour  driven  to  o f t h e Song  by  and  off  Sparrows  tended  o f such  off  Towhees  drive  u s u a l l y Fox  independently  readily  both  a  took  remain Sparrow,  behaviour.  a l l  the  other  125  /  seed-eaters, relatively  but  close  retaliation.  often to  sheer  potential  numbers  allowed  antagonists  them  without  to  feed  inducing  126.  APPENDIX the  6  B-C  age  Hierarchy  of  the  a r e known.  banded  and  on J u l y  i t had p r o b a b l y  (see  Nice  35-40  i n hierarchy:  t h e t o p two b i r d s i n  Hierarchy  The  was  Age a n d p o s i t i o n  top  The f i r s t 25, w i t h just  19^3).  two  dominant  no s i g n s  gained  Its.age  juvenile  males  juvenile  t h e B-C  (Blue/Blue)  of post-juvenile  independence  on J u l y  in  25  was  from  moult,  i t s  parents  put t e n t a t i v e l y a t  days.  The banded (Stage  second on  May  4 of Nice  male  on  25,  the dominance  at  the  1943)..  scale  fluttering  Therefore  i t was  (Yellow/LP)  stage  of  between  was  development 10  and  16  days o l d .  Thus, were  i n  determined,  Yellow/LP  January,  Blue/Blue  would  be a b o u t  product  o f most  product  of a first  of  the largest  when  likely  a  brood.  hierarchy.  the positions  would  be about  260 d a y s second  old. brood,  Yet, Blue/Blue  of these  200  days  Blue/Blue and was  two  birds  old, would  Yellow/ the top  and be a  LP  a  juvenile  127  APPENDIX  If to (N  7.  Pressure  each  potential  a resident's =  nos. o f  proportional T i s also T  assumption to  increase  makes x i n t r u s i a n s  potential  settlers).  The  the resident  spends  the time  ,when  T.  settler  Explanation.  the total  to  being  intruders:  territory,  proportional  constant,  from  Nx  that  to territory goes  number o f i n t r u s i o n s  area,  up, t e r r i t o r y  the bird  does  p e r d a y on  value  of  n o t have  Nx i s  i n defence (T).  so i n order size  Nx  =  goes  much  to  keep  down - t h e spare  time  128  8  APPENDIX  The  Watson conditions a  five  and which  c o n d i t i o n s of  Moss need  (1970a) t o be  Watson  put  met  to  and  forward show  Moss.  the  that  following  behaviour  five limits  breeding population.  A.  A  either  substantial because  breeding  even  part  animals though  of  the  die; or  they  p o p u l a t i o n does  because  survive,  they  and  are  may  not  breed,  inhibited  breed  in  from later  years. B. if  Such the  are C  non-breeders  more  dominant  The  breeding such  the  resource  D.  The  as, If  change both  as  territorial  itself  other  (B) ,  the  are  space,  capable  ( i . e.  of  breeding  breeding)  animals  or  not  completely  nest  sites.  using If  up  they  some are,  limiting.  depressed in  causes  following  limiting  i s  or  changes  (A) ,  animals food,  mortality  factor(s)  E.  or  physiologically  removed.  resource,  rate  are  an of  (C) , a n d changes  r e c r u i t m e n t due  opposite  mortality (D)  are  i n food,  sense  or  to,  to and  depressed  fulfilled, then  food  the  and and  at  limiting the  same  recruitment. the  numbers  behaviour  are  breeding population.  129  COTTONWOOD  STRETCH  LOSER  \  6 \LP  G LP LP LP  M LpUnb R Y Y ©  LP LP jjBUnt)  \  R Y  14  \  o"  6"  Y © «  G G  9  3  7  10  5  2  5  \  2  2  1  4  1  2  3  1  LP RW ©  Black  <D RW RW G  1  1  G G  w  2 d* d LP Black RW w © ©  2  11  11  7  - 11  2  2  3  1  9  ?  ?  2  G ®  LP L-D  B 0 0 B  BO B  5  3  5  1  1  3  4  5  2  1  4  1  1  1  1  2  2 1  1 2  5  2  1  \  2  1  1 1  ®  LP L-D  1  BO OB  1  BO B  170 I N T E R A C T I O N S  13  MAN HOURS  F i g u r e 12  SEDGE  \  GB GB G BY R Y  cf cf GB G GB BY  \  2  \  cf  R Y  15  ? Black  B 1 1  0 Y  LO SEF ? ? cf R R Y G B B  1  3  cf  4  2  1.  ? '? ? cf ? cf ? B LD Y W BO W B © BY BY DG Y RW DG 4 8 9 6 9 4 7  6  1  4  2  2  7  5  2  2  1  2  8  1  3  1  2  2  2  2  3  3  3  11  2  \  2  1  (W)  1  1  Black  B O Y R G R B DC Y LU B Y DG it W Y BO RW W DG  PATH  1  1  1 1  1  1  2 1  \  2 2  3  \  4 4  B©  2  B BY LD BY  1  \ 186 INTERACTIONS  15 MAN  HOURS  121  F i g u r e 13  B U L L D O Z E D  LOS  . \  cf M  s _o_  DP  R  M DP  0  R  \  2  Y  w  10  5  2  10  1  5  1  20  9  2  6  34  «  11  1  DG  2  DG  cf  Black B Y  5  Y  \  2  2  RW B ® BY 1  6 1  2  1  Black  2  cf Black Black L P ® DG B  LP®  LU  E R  cf R Y  R Y  oc  PATHS  10  3  4  2  DG  W Y  6  5  1  3  2  1  2  3  3  \  B ® RW BY 174  INTERACTIONS  22  MAN  1  \  HOURS  1g2  F i g u r e 14  F I R S T BIG  ALDER  LOSER  \  1©  LD LD  R©  B W  0_ Y  Y DG  Bl BO  1  9  5  2  1  1  6  2  6  1  9  5  3  4  1  LD LD  cr  LU  z  R © B W  0 Y Y DG Bl BO 61 I N T E R A C T I O N S ;  MARSH  1  2  1  \  1  \  6 MAN HOURS  NARROWS  LOSER \  4  Y  i f © M B  ? UNB  ^  UNB BO OB  ? ? BO OB  Bl©  2  4  1 1  Bl© 8 INTERACTIONS;  5 MAN HOURS  1  ^  F i g u r e 15  NO-NAME  CLEARING  L O S E R  42 INTERACTIONS  11 M A N H O U R S  

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