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Some aspects of snowshoe hare behavioural ecology Graf, Ronald Paul 1981

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SOME  ASPECTS  O F SNOWSHOE  BEHAVIOURAL  HARE  ECOLOGY  by RONALD  B . S c , A  The  THESIS THE  University  B r i t i s h  Columbia,  1978  FACULTY OF GRADUATE STUDIES (Department of Zoology)  accept  THE  Of  GRAF  SUBMITTED IN P A R T I A L F U L F I L L M E N T OF REQUIREMENTS FOR T H E DEGREE OF MASTER OF SCIENCE in  THE  We  PAUL  this thesis required  UNIVERSITY  as c o n f o r m i n g standard  OF B R I T I S H  December  (c)  Ronald  Paul  COLUMBIA  1981  Graf,  to  1981  the  In  presenting  requirements of  B r i t i s h  i t  freely  agree for  this for  an  available  that  I  understood  that  f i n a n c i a l  by  his  or  reference  and  study.  I  extensive be  her or  s h a l l  DE-6  (2/79)  copying  granted  by  the  of  p u b l i c a t i o n  not  be  allowed  Columbia  /9p/  of  make  further this  head  representatives.  of  >£L«^ ^  University s h a l l  The U n i v e r s i t y of B r i t i s h •2075 W e s b r o o k Place Vancouver, Canada V 6 T 1W5  ^te  the  the  Library  permission.  Department  at  of  the  may  copying  gain  degree  fulfilment  that  for  purposes  or  p a r t i a l  agree  for  permission  scholarly  in  advanced  Columbia,  department  for  thesis  It  this  without  thesis  of  my  is thesis my  written  i i  ABSTRACT The  objectives  of  interactions  of  describe  social  to  the  determine  dynamics. initial  built  in a  performed  results  of  to  penned food,  and  water,  males  being  station A  to i n the  male  to  be  males  were  dominant several  were  hares,  might  and  experiments  hare  population  enclosures and  I observed the  to  by  interactions,  in  agonistic  americanus,  affect  field  stereotyped  Interactions and  i n the  and  to  gather  to  perform  wild  to  hares  confirm  the  and  occurred  in  occurred  over  right-of-way.  pens  and  i n the  winter  and  females  summer.  interactions  per  Hares  hour  The  hares  wild,  with  being  were  observed  access  most  aggressive at  a  feeding  wild. experiment  The  after  i n an  hour).  the  end  for  most  prevented  from  itself female,  between  the  each to  complete,  the  further  dominant  Subordinate  copulating  involved with  males  female's  courtship  showed  copulations.  oestrous  of  aggresssive less  enclosure  courtship  an  occurring  but  Lepus  situation.  i n the  spring  aggression  Similar,  d e s c r i b e the  metre  spots,  dominant  males.  extremely  30  hares.  behaviourally  At  the  experiments  responsible  males  of  agonistic  resting  the  23  mating  of  x  hierarchies  most  during  up  30  controlled  wild  dominance  with  two  to  hare,  interactions  interactions  formed  dominant  were  observations.  Agonistic both  structure  some  pen  study  snowshoe  descriptions  experiments and  the  i f these I  the  (94  fast  high  chases  the by  levels  of  interactions  per  oestrous  she  became  by  males.  approaches sequences  by  were  seen  in  the  wild  which  reasonable  description  densities  the  have  effect  some Wild  resident  disperse were  juveniles.  mechanism juveniles  on  juveniles  juveniles  status.  extreme  I in  were  and  adults  to  a by  animals  At  for  new  similar  wild,  but  population.  into  mimic  the  preventing a  events.  introduced  experiment  for  captive  competition  into  observed the  the  mating  male  harrassed  This  into  of  that  high  provided  a  population  oestrous  females  may  mortality.  juveniles  juveniles  juveniles  suggests  enclosures  a  situation  area.  A l l  residents, interactions  was  unaware  suggests the  containing  of  there  recruitment  in  30  introduced  both  adults  between their is  a of  which  adults  and and  r e s i d e n t i a l behavioural non-resident  i v  TABLE  OF  CONTENTS  ABSTRACT  i i  LIST  OF TABLES  v i i i  LIST  OF FIGURES  ix  ACKNOWLEDGEMENTS CHAPTER  1:  '.  GENERAL  x i  INTRODUCTION  .  1  A.  INTRODUCTION  1  B.  STUDY  4  Chapter  2:  AREA SOCIAL  BEHAVIOUR  AND S O C I A L  ORGANIZATION  5  A.  INTRODUCTION  5  B.  METHODS  6  C.  AND M A T E R I A L S  1 .  ENCLOSURES  6  2.  OBSERVATIONS  OF WILD  HARES  a.  Addition  of  A r t i f i c i a l  b.  Addition  of  Natural  c.  An  Unmanipulated  10 Food:  10  Food:  Situation:  11 ,  11  RESULTS 1.  2.  12  AGGRESSIVE  INTERACTIONS  a.  Description:  b.  Interaction  Intensity  c.  Interaction  Rate  SOCIAL a.  .12  ORGANIZATION  SUMMER: i . i i .  Penned Wild  12  Comparisons:  Comparisons:  . . .  18 20 20 25  Hares: Hares:  25 28  b.  WINTER: i.  Penned  i i . 3. D.  .  CHANGES  30  Hares:  32  I N DOMINANCE WITH  SEASON  37 39  1. S O C I A L BEHAVIOUR  39  2. DOMINANCE H I E R A R C H I E S  40  T H E CHANGE  I N DOMINANCE WITH  SEASON  42  SUMMARY  Chapter A.  Wild  Hares:  DISCUSSION  3. E.  30  50  3: DOMINANCE AND  T H E MATING  SYSTEM  52  INTRODUCTION  52  B. METHODS  53  1 . PENNED H A R E S :  53  2. WILD H A R E S :  55  C. R E S U L T S 1. A T Y P I C A L  MATING  2. C O U R T S H I P  I N WILD H A R E S :  3.  D.  *  DOMINANCE AND Male  Hierarchy  b.  Reproductive  55 59  SUCCESS:  63  i n t h e Pen  63  Success  64  DISCUSSION  75 MATING  2. DOMINANCE 3.  FEMALE  AND  SYSTEM: REPRODUCTIVE  77 SUCCESS:  78  CHOICE:  80  SUMMARY  CHAPTER A.  PEN:  REPRODUCTIVE  a.  1 . T Y P E OF  E.  IN THE  55  4: F A C T O R S  INTRODUCTION  85 AFFECTING  S U R V I V A L OF  JUVENILE  HARES  .. 87 87  vi  B.  C.  METHODS  88  1.  EXPERIMENTAL  INTRODUCTIONS  2.  OBSERVATIONS  OF WILD  89  JUVENILE  HARES:  RESULTS  89 90  1.  PARENTAL  CARE  2.  LITTER  3.  INTRODUCTION  SIZES  IN  SNOWSHOE  AND GROWTH OF  HARES:  RATES:  JUVENILES:  90 92 93  i  4. D.  E.  OBSERVATIONS  OF WILD  JUVENILES:  DISCUSSION  97  1 .  PARENTAL  3.  INTRODUCTION  4.  OBSERVATION  CARE: OF WILD OF WILD  97 JUVENILES JUVENILES  SUMMARY  CHAPTER APPENDIX  5:  100 103 104  GENERAL DISCUSSION  AND CONCLUSIONS  I  Postures  96  106 ..111  and  Intraspecific  Locomotion Communication  111 113  Chinning  113  Thumping  114  Vocalization  114  Appendix  II  116  Appendix  I l i a  117  Appendix  I l l b  118  Appendix  IVa  119  Appendix  IVb  120  Appendix  Va  121  Appendix  Vb  122  vii  Appendix  Vc  123  Appendix  VI  124  Appendix  VII  125  REFERENCES  CITED  126  ]  vi i i  LIST  TABLES  Table  1.  Table  2a.  F u l l  Table  2b.  Summer  Table  3.  Summer  hierarchy-  Table  4.  Summer  hierarchies-  Table  5a.  Winter  hierarchy-  Pen  2  31  Table  5b.  Winter  hierarchy-  Pen  2  31  Table  6a.  Winter  hierarchy-  1980-  Pen  1  33  Table  6b.  Winter  hierarchy-  1980-  Pen  1  33  Table  6c.  Winter  hierarchy-  1980-  Pen  1  34  Table  7a.  Winter  hierarchy  at  Single  Feeder  35  Table  7b.  Winter  hierarchy  at  single  feeder  36  Table  8.  Winter  hierarchy  Table  9.  Winter  hierarchy-  Table  10.  other  Agonistic  OF  interaction  data  for  Pen  25  1-summer  hierarchy-  Comparing  rates  1979-  Pen  1979-  Pen  1980-  at  hierarchy  27  2  multiple  aggressive  1  28  Pens  natural  26  1  & 2  29  feeders  37  food  39  posture-snowshoe  hares  to  Leporids  41  Table  11.  Schedule  of  Table  12.  A l l  hierarchy  Table  13.  Hierarchy  Table  14.  Distribution  of  copulations  Table  15.  Interactions  of  juveniles  Table  16.  Comparison  male  female  after  of  introductions in  Pen  dominant  behaviours-  54  1  '.  removed  64 64  between  the  males  . .  65 96  Pen  vs  Wild  108  LIST  design  the  FIGURES  Figure  1.  The  Figure  2.  A  Figure  3a.  A hare  exhibiting  an  Figure  3b.  A  exhibiting  another  hare  of  OF  marked  hare  enclosures  with  a  7  plastic  disc  aggressive type  tag  9  leap  14  of  an  aggressive  leap Figure Figure the Figure wild Figure  14 4.  6.  male  a  defensive  showing  the  leap  16  i n q u i s i t i v e  posture  and  posture of  17 types  of  interactions-  pen  vs  . . .  19  Frequencies  of  types  of  interactions-  male  vs  hares  21  Frequencies hares  9.  in  of  the  hares  10. vs 11.  Figure  12a.  Figure  12b.  in  the  Differences female  of  interactions-  male  vs  of  22 types  of  interactions-male  vs  winter in  23  individual  interaction  rates-  hares  Seasonal  13a.  types  summer  Frequencies  Figure  Figure  diagram  Frequencies  7.  8.  exhibiting  down  hares  female Figure  A  ears  female Figure  hare  5.  female Figure  A  24  change  in  dominance  Mating  success  of  males-effect  Mating  success  of  males-dominant  Attempted  copulations  between  by  of  the  sexes  injuries  . . .  removed  males-effect  13b.  67 68  of  injuries Figure  38  72 Attempted  copulations  by  males-dominant  removed Figure  14.  73 Relationship  in  young  of  age  to  weight  94  xi  ACKNOWLEDGMENTS I  would  America  for  Kluane  Lake  families people  the  Zorn  cutting  my  Stan  for  draft  of  gratitude  project,  support  thank  varied  the  greatly Boutin,  to  mark the  for I  his  on  my  thank  Most  supervisor,  especially employment a and  to  and  since in  it  the  U.B.C. a  from  U.B.C.  of  North.  G i l b e r t ,  would the  like  suggestion  Moses  offer  Jean  for  special  beginning  to  to of his  thanks end  of  and  J.N.M.  and  reviewing  an  earlier  I  would  A . R . E .  like  extended Thank two  to  extend  S i n c l a i r ,  support  Fellowship, Teaching  assistance.  C . J . Krebs  moral became  Scott  Many  enclosures,  their  Rich  the  Upton  Drs.  a l l  Dr.  the  I  also  and  the  support.  of  with  and I  committee,  assistance,  from  animals,  like  thesis.  up  North  at  Williams of  T a i t t .  assistance  my  types  M i l l e r ,  photography.  would  this to  the  the  Steve  coming  of  f a c i l i t i e s  appreciated  for  Mary  Institute  construction  thank  serving  Fellowships,  their  Mary  with  guidance,  use  and  project.  taking  Stan  Boutin  Smith  and  with I  Arctic  especially  many  and  to  the  Dundjerski,  hair  assistance  and  me  include  especially  to  their  etc.  thank  opportunity  helped  Carey,  to  Base,  for  trapping, They  like  as  you  my  for  his  throughout  this  a  result  a l l .  of  my  I  received  MacMillan  Fraser  assistantship.  1  CHAPTER  1:  GENERAL  A.  The North  snowshoe  America,  line,  and 40th  much  of  year  cycle  its  range, of  have  They  range  two  from and  predation.  Two  and  (1964)  to  explain  Both  the  Christian  hypotheses,  year  and  Davies  behaviour  C h r i s t i a n  conspecifics  of  stress-related rates,  and  and  found  to  diseases, therefore  in  Chitty  some  in  proposed  increased resulting  of  adrenal  to  death and  lower  hypothesis  based  population  C h r i s t i a n e a r l i e r  microtine  species.  their  individual  and  of  These  shock  densities  crowding  and  stress,  and  high  disease  increased numbers.  would  non-cycling  increased  hormones.  high  proposed  increased  in  with,  factors,  levels  result  phenomena.  respectively,  that  through  hypotheses  forest  cycling  in  miles  and  suggested  both  ten-  of  coupled  been  behaviour-genetic  Davies  lead  food,  had  consistent  sunspots  an  tree-  Throughout  natural  to  in  approximately  various  i n t r i n s i c  (1960),  regulation  levels  then  and  on  of  the  hundreds  this  proposed  only  to  1974). a  the  weather  lack  cycles  aggression,  hormones  and  based  and  lead  higher  explain  Chitty  3-4  reviews  (1978)  hypotheses  species.  therefore  to  to  (Banfield,  across  found  north  hare'exhibits  (1963)  factors-  and  extends  synchronous  Windberg  population  therefore  snowshoe  is  ecosystem,  U.S.A.  disease  explain  of  forest  the  proposed  extrinsic  Davies  in  Keith  been  Keith  the  the  americanus,  d i s t r i b u t i o n  abundance,  1963).  which  on  within  INTRODUCTION  Lepus  its  p a r a l l e l  (Keith,  fires.  and  south  the  hare,  INTRODUCTION  levels or  other  mortality  Chitty  (i960)  2  agreed  on  the  increased  increased,  but  d i f f e r e n t i a l  selection  one  morph  at  a  high  rate  densities.  populations  at  other  hazards  in  types  of  the  mechanism  which  snowshoe  hare.  examining  spacing  no  such  cycle.  It  ecology  was of  behavioural My  study  behaviour in  the  ecology  is of  female  in  conjunction of  l i t t l e snowshoe  descriptive,  with  the  and  and  The  an  to  withstand  the  changing  polymorphism  Keith  behaviour  et as  in  hares,  assess  but  phase  a the  investigation  the  by  found of  the  behavioural if  i n t r i n s i c  the  ten-year  cycle.  major  research  project  snowshoe  faculty  information hares.  a  mutual  regulation  to  at  decline  of  increase  to  well  and  1978;  study  contribute  peak  explore  an  order  survive  cycles.  snowshoe  to  reproduces  stable  such  during  which  suggested  not  of  population-  able  observed  started  in  students  less  Windberg,  therefore,  the  in  result  the  not  population  spring  in  is  this  did  to  in  densities  a  levels  Chitty  and  (1980)  done  Columbia.  anecdotal,  lead  could  B r i t i s h  high  the  1979)  mechanisms  graduate  behaviour  Keith,  does  favoured  but  in  (Keith  intent,  several  There  result  hares  investigating  is  maintain  snowshoe  was  but  as  is  populations  densities,  Boutin  my  morphs  environment.  could  behaviour  genetic  withstand  and  and  cycle  morph  pressure  aggression  the  densities,  studies  Cary  that two  other  high  population,  ,1978;  low  the  of  increasing  can  selection  Keith's a l .  at  and  on  in  The  interference  in  proposed  favoured  high  levels  hare,  from  The  available available  unquantified,  on  data with  involving  University  the are  of  social generally  almost  no  3  mention  of  conspecific  Criddle,  1938;  Severaid,  understandable Snowshoe moving  hares  are  between  snowshoe c r y p t i c ,  1971),  possible  constructed hares,  at  large  and  lack  hares  are  in  are  it  the  for  is  not  same  possible  basic  vegetative but  maintain Rongstad  observe  disputes  after  species.  observe  description  fast-  1961;  day  t e r r i t o r i a l to  to  is  observe. and  dense  Bider,  location  to  secretive, of  1932;  information  t e r r i t o r i a l ,  1980;  enclosures  a  of  (Grange,  d i f f i c u l t  areas  not  some  outdoor  obtained  This  (Boutin,  so  interactions  nocturnal,  found  they  ranges,  individuals be  1942).  usually  home  Tester,  might  wild  A d d i t i o n a l l y ,  overlapping and  as  creatures,  cover.  agonistic  day,  I  therefore  introduced  of  as  their  wild  social  behaviour. The  overall  behaviour  of  design  penned  conditions,  and  s u f f i c i e n t l y  to  the  hares.  penned  -social as  found A  in  of  is  and  comm.)  population  of  3  a  provides  or  and  spring  snowshoe  wild,  the  2  of  the  found  in  description  in  numbers  of  this  between  the  hares  reached  description  from of  snowshoe  the  mating  for  spring  area  some  (Krebs  decline  males  similar  near  various  the hare  summer.  suggested  a  the  free-ranging  organization  decline  hares  under  a  and  observe  conclusions  characteristic  found  to  presents  spring  competition  was  intensively,  deny  social  (1978)  has  study  observe  Chapter  winter,  Krebs  aggressive  (pers.  a  to  confirm  demographic  microtines 1974),  the  my  hares  then  behaviour,  of  of  system  may  cycling  and be  Myers, a  mates. decline  my  study.  of  L .  result Boutin in  a  Chapter  americanus,  4  and  discusses  chapter  the  also  relationship success  in  between  Changes  1978), provide the  in  in  interactions  of  in  study,  5 as  both  of  Arctic  by  study  snowshoe  white  as  male  a  an  This  of  the  reproductive  and  vegetation  STUDY  North  Picea  Salix  many may  herb be  the  America  Canada.  but  and  snowshoe  of  the  of  the  most  overall Windberg,  Chapter  4  hares,  I and  behavioural juveniles.  overall  design  of  results.  Kluane Mile  Lake  Base  1054  of  vegetation  is  has  a  varied  A more  Douglas  detailed (1974).  the  of  the  Alaska  dominated  understory  buffaloberry,  species. in  in  dispersing  at  The  spp.,  found  in  In  the  AREA  near  glauca,  be  examined  and  summary  (Keith  19.74).  which  evaluation  to  changes  hares  care  hares  brief  Yukon,  appear  Myers,  parental  conducted  willows,  canadensis, the  spruce,  and  resident  of  south-west  several  of  was  Institute  Highway,  mates.  analysis  and  c y c l i c  experiment  provides well  survival  B.  This  for  experimental  dominance  (Krebs  an  between  Chapter  competition  explaining  description  results  this  male  microt ines  a  an  juvenile  numbers  and  male  hare.  factor  population  of  provides  the  important  extent  of  Shepherdia description  5  •  CHAPTER  2:  SOCIAL  BEHAVIOUR  A.  Christian proposed  and  individual cycling  animals and  lead  of  these  hare in  hares,  interactions  on  the  Snowshoe  to  enclosures  to  my  over  observe  more  periods.  I  hares  but  in  than  a  population sequence,  a  added  wild  state.  I of i .  unmanipulated  pens  a  thus  able  was  a  fallen  completely  was  I  10m  able  to  then  large known  extended  unable  only  to  white  this to  two  and  for  to  brief  concentrate  natural,  food  often  d i f f i c u l t  later  switched  Through  it  s t i l l  to  a r t i f i c i a l I  was  food  and  such  and  observed I  a r t i f i c i a l  hares.  to  and  time.  but  the  quantitative  and  source,  situation,  of  Subsequently,  snowshoe e.  period  of  of  constructed  hares,  the  species.  make  animals,  area,  observed  this  gather  of  both  agonistic  effects  features  population,  few  local  food  numbers  wild  the  therefore  of  describe  of  between  cycle  fast-moving,  and  I  extended  the  concentrated  F i n a l l y ,  a  ten-year  examine  have  investigate  to  c r y p t i c ,  data.  small  therefore  from  animals  to  the  behavioural  regulation To  organization  are  an  to  to  populations,  house  observations  species.  (1960)  interactions  population  and  social  Chitty  aggressive  necessary  behavioural  individuals  the  and  is  These  wild  meaningful  to  hares  animals.  observe  which  hypotheses  it  interactions  secretive  (1964)  non-cycling  a p p l i c a b i l i t y snowshoe  in  ORGANIZATION  INTRODUCTION  Davies  hypotheses  AND S O C I A L  observe to  many  natural,  spruce  tree.  unmanipulated observational  natural  provide  a  the  a  food  to  an  description  of  6  the  social  hare,  behaviours,  and  to  of  wild  B.  near  constructed the  America  behaviour  in  of  the  the  snowshoe  enclosures  was  hares.  METHODS  1.5  in  m.  the  from  ground  wire  was  of  11  AND M A T E R I A L S  buried  to  and  was  through manner  to  the as  the  50  Pen  was  cm) cm,  The  link  2.5  inside  the  pens  grids above  using  The  flagging  ground  discourage  t e r r e s t r i a l  with  was  as  hares  the  poultry One  each  of  predators.  blind  with  The  ground  were An  around  level  link  ground.  The  into  in  the in  the out link out same place  plexiglass  floor  of  the  level.  Vegetation  to  f a c i l i t a t e  subdivided  into  e l e c t r i f i e d  wire  a l l  to  slipping  used  pen.  was  chain  was  trimmed  markers.  placed  from  1  digging  larger  wire  large  pens  from  constructed  plywood  m above  the  North  chain  digging  was  selectively  tape  level  captured  2  in  from  Pen  1.5  interiors  cm  the  ha)  of  Pen  ground  with  leverets,  cm  for  approximately was  overlapped 15  1).  from  hares,  fencing.  was  fencing  0.1  Institute  Figure  buried  the  approximately  Arctic  predators  mesh.  blinds  observation.  was  young  constructed  the  link  overlap  except  30m,  (See  and  prevent  prevent  1,  1979  prevent  to  x  of  chain  (1  larger  chain  windows,  gauge  to  (30m  Base of  wire  compounds.  fencing  Lake  level  enclosures,  pens  spring  Poultry  the  two  Kluane  constructed  of  if  organization  ENCLOSURES  I  of  social  determine  characteristic  J_.  and  compounds  3 x 3 50  cm to  7 Figure  1:  A diagram showing the design of the c o n s t r u c t e d in the Yukon.  ^BLIN  HOLDING  PENS  enclosures  8  In  1980  discourage great  I  chow  willow  boughs  accepted and  the  65cm),  and  disc Band was  winter  the  above  4  Tag  allowed  and  at  Observations binoculars,  gentilis),  The  netting  pens  with  to and  was  only  water,  ad  supplement  commercial  l i b , and  the  p r e f e r r e d the  added  chow. willow  I  with  a  (See the  other  Hares leaves,  the  Marked to  Another  tag  numbered During  the  supplemented  with,  or  white  outer  pile  method  the  hares their  metal  back-up.  cutting  ^generally  during  20-45x  sheets,  was  x  multicoloured  numbered  2). as  (23cm  new  worked  year,  were  and  guard  well  making  released  the  into  surroundings  as  the  for  3-  observations.  taken  Nightscope.  a  ear  throughout  habituate  were  night  by  Figure  method  dark  them  ear  This  to  a  marked  designs.  began  with  traps  an  Co.) to  live  of  were  prepared  to  discernible.  Observations  onto  netting  i n Tomahawk  cutting  remained  before  Starlite  and  but  hares  tagging  the  easily  sunset,  wild  individual  underfur  days  quickly,  attached  by,  and  the  to  individually  and  tag  pens,  days  attached  metal  designs  fish  bark.  x  (National  the  1 with  (Accipiter  inside  protein),  2-3  chow  live-trapped  in  Pen  virginianus).  hares  every  I  hairs  goshawks,  (Bubo  (16-20%  the  replaced  covered  successful.  rabbit  plastic  by  owls,  provided  twigs,  partially  predation  horned  partially  23cm  I  periods with  spotting  taken of the  close  to  substantial naked  scope,  eye,  and/or  Observational  r e c o r d i n g was  except  periods  during  of  sunrise  moonlight. with with  done  high  and  8x50 a  x5  directly activity,  ure  2:  A penned hare showing the type o f p l a s t i used f o r i n d i v i d u a l identification.  10  and/or  2.  night  watches,  OBSERVATIONS  a.  Addition  In hares  OF WILD  of  a  condition, present  weight,  about  75  kg  near the  an  hares  of  the  chow,  1m  high,  The  feeder  The  as  with was  3m  feeder  habituated  of  the  a  used.  placed City  was  a  I  feeder,  number  feeder,  of  the  60cm) edge  vehicle.  After  and  of  then  the did  3  a  on  km e a s t by  observed Hares  weeks  dominant  further  of  willow  away.  two  a  clearing  I  40m  three  held  cylinder  a  station.  parked  wounds  which  surrounded  vehicle  removed  of  resting  approximately  feeding  wild  reproductive  upright  p a r t i a l l y  from the  The an  at  marked  sex, and  diameter  the  observations,  area  was  from  to  length,  trapped.  Silver  clearing  individually  recorded  hind-foot  rabbit  known  was  of  hares  four  days  observations. In  were  February,  placed  description Boutin this  right  I  of  the  preliminary from  station.  approximately  became  observed  were  area  near  I  animals  enclosures.  thickets  soon  base.  taperecorder  Food:  the  (approximately c i r c u l a r  portable  HARES  1979,  feeding  when  a  A r t i f i c i a l  December,  at  when  was  on  the  of using  procedures  trough  feeders  ha  the  grid  may  be  found  food  addition  to  test  to  observe  feeders. were  ten  9.3  opportunity  a r t i f i c i a l  1980  as  Telemetry  Tagging,  previously  Boutin  other hare  weighing,  described.  x  grid.  in  further  (2m  I  8cm A  x  15cm)  detailed  (1980).  hypotheses,  While I  interactions and  observed  used at  observational hares  from  11  a  vehicle  approximately  alternated apart  b.  observations  between  Addition  In spruce  of  18  1980  tree  the  in  at 1979.  and  I  repeated.  observed  for  c.  three  weeks  c a l l e d  the  Highway years. and these the again  the  which This  Silver wild wild  hares,  of  been was No  and  population the  feeders  29,  a  the  feeders.  located  about  I  120  m  1980.  freshly  location  the  in  this  this  situation  Trapping hares  at  area  10m  feeder  are  to  and this  felled  upright  trees  parked  just  had  City.  of  white  had  often  provide  further  a  been blown  natural,  marking  location  was  intermittently  vehicle.  Si tuat ion:  location  observed  the  summer  "Ditch",  March  placed  same  a  An U n m a n i p u l a t e d  During  I  source.  from  two  one  Food:  mimicked food  I  and  Spruce  concentrated  from  between  Natural  February,  situated down,  March  40m  1980  I  off  a  ear-tagged  closed about  portion to  I  used  this  p e r i o d i c a l l y  wild  hares  from  at  the for  between  to  the  during  monitor the  parked  a  location  old  the  manipulations  area  a  of  t r a f f i c  half-way  experimental  hares  Alaska  previous  5  enclosures were  done  behaviour  summer vehicle.  of  1980.  on of I  1 2  C.  Hares their  introduced  new  disturbed 2-10  surroundings, when  minutes  reaction I  into  of  I  entered  after  the  here  to  .only  the  after  the  b l i n d .  entered my  are  described  Appendix  I  some  soon  became  several  accustomed  days  were  Observations  the  blind,  not  were  depending  to  unduly started  upon  the  entrance.  the  behaviours,  locomotion  pens  and  had  hares  present  interactive  I  RESULTS  interactive such  in  as  grooming, I.  Appendix  brief  behaviour  I  data.  postures,  also  observations  Non-  on  and  include  in  i n t r a s p e c i f i c  communication.  J_.  AGGRESSIVE  a.  INTERACTIONS  Description:  Aggressive observed were  in  later  through hare  of  this  an  within  if  3m  agonistic.  was  In  was  one  a l l  change  recorded  in  these  an  other  judgment,  an not hare  to  interaction. before the  I  out  change Two  of  in  hare, another  movement,  this  interaction. of its  hares  c l a s s i f i e d  interactions  one  resulted of  f i r s t  interactions  If  aggressive moving  were  of  hares.  d i r e c t i o n  as  this  hares  types  posture,  or  through  caused as  of  two  free-ranging  hare,  hare,  each my  or  between  location,  classed  of  but  wild,  its  oncoming  also  i n  events  addition,  of  pens,  movement,  changing  sequence In  the  found  its  interactions  their  the  pathway  d i r e c t i o n , had  to  actions  snowshoe  be as  hares  13  followed  a  gradient  interactions  hare,  avoidance  required  interactions. less*  Up-  or  the  stress  trace  s l i g h t l y  approached  hare  was  intense  the  loser  i .  e.  loss  to  than  did  did  gradient  more  as  intense  more  intense the  least  an  probably  of  from  by  loser  interactions than  intense  input the  descriptions  this  less  energy  action,  less  as  "ears a l l  up"  its  shoulders  (Figure  ears change  form  of  snorting down"  ears  six  intense  to  by  the  as  involved  sufficient of  slight though  most  its  ears  hare.  If  approaching.hare the  approaching  hare  did  approached  the  with  another  d i r e c t i o n ,  certain  aggressive a  the  with  against  not  and hare  was  this hare  move,  moved  d i f f i c u l t  the  and  and  around winner.  subjective  hare.  cause  hare  top  A  of a  the  its  mere  loser  slightly  to  animal head  and  dipping  of  move  more  was  accompanied  by  An  even  intense  rocking  the  aggressive  occasions,  to  type  the  the  movement.  interaction  included  backwards,  was  flat  direction  this  the  hare,  judge.  5 ) . On  appeared its  then  one  approached its  interaction  pressing  the  the  interaction  This  from  agonistic,  If  to  when  approached  away  changed  hare,  interactions  Down-  forwards,  as  winner. hare  occurred  moved  c l a s s i f i e d  stationary  Ears  and  to  less  energy  drastic  interaction  approaching  the  or  less  subsequent  types  This  c l a s s i f i e d  of  require  addition,  directed  action  The-  less  intensity,  intense:  Ears  the  to  caused  The  interaction  up  and  In  interactions.  most  aggressive  appeared  aggressive  caused  of  motion was  more of  about  the to  away, intense  grunting  body,  or  "ears forwards  "lunge",  as  Figure  3b:  An a g g r e s s i v e " l e a p " e x h i b i t e d h a r e a t t e m p t i n g t o l a n d on t o p beneath i t .  by of  a snowshoe the hare  15  described  next.  This  during  mating  season  the  discourage Lungehare  the  The  propelled strike  the  i t .  During  a  against  its  Leap-  A  leap  over  the  aggressive the  or  hare.  leaps,  and  generally  an  aggressive loser  interactions  the  which were  not  Chase-  This  after  another  hare.  losing  hare  more  with  on  3b).  an  the  to  aggressive  apparent or  attempt  actually its  to  bite'  ears  by  down  or  In  a l l  always  were  when The  avoid  a  hare  leaps  have  been  its  where  an  had  approached  most  frequently  the  close  responses  passed  with  females leap  was  by  an  "lunge"  up  leap,  it  defensive  legs  the  as  pregnant  its  as  winner, to  its  body  considered of  hares  the  in  losing  recorded.  the 3m,  loser  over  involved  than  other  leap  of  follow,  If  leaps  to  paw  observed,  situations  eventual  a  aggressive  hind  the  males.  a l l  an  leaps  passing  aggresssive  of  defensive  and  In  courting  the  ground of  one  season  hare  the  types  top  the  mating  interaction  for  attempted  which  held  off  two  Aggressive  During  some  analyses  in  leaps.  out  observed,  attempt  Only  were  land  and  3a  the  hare.  4).  to  discourage  tucked  (Figure  There  winner.  to  in  hare  jumping  lashed  attempted  during  attempted  hare  defensive  hare  was  observed  males.  forepaws  aggressive  a  other  leap  of  hare,  its  only  females  interaction  with  was  shoulders.  involved  eventual  observed  the  down"  pregnant  another  hare  and  (Figures  aggressive  the  head  it  forepaws  the  lunge,  an  at  other  ears  approaches  was  itself  aggressive  over,  when  courtship  lunge  to  "rocking  an  aggressive  winning the  hare  did  interaction  ,was  hare not  running chase  c l a s s i f i e d  the as  A snowshoe hare e x h i b i t i n g a d e f e n s i v e " l e a p " by t u c k i n g i t s paws up t i g h t to i t s body as i t passes over a l u n g i n g hare.  17 F i g u r e 5:  A male hare ( r i g h t ) i n the " i n q u i s i t i v e " p o s t u r e a t t e m p t i n g to s n i f f a female ( l e f t ) to d e t e r m i n e her r e p r o d u c t i v e c o n d i t i o n . Female i s d i s p l a y i n g the a g g r e s s i v e " e a r s down" posture.  18  a  "lunge",  males,  the  high, down  rather  as  aggressive  while tight  than  the  male  against  a  chase.  male  hare  doing  being  i t s rump.  If  the  the  chasing  chased  Chases  chase  kept  ranged  involved had  its  from  only  its tail tail  3m  up  tucked  t o more  than  200m. Hit-  This  another with  interaction  hare.  one  The  involved  h i t was  hindpaw,  or  during  result  of  a  lunge,  leap.  A  few  a  a  fights  raking  bite  with  a  were  one  hare  with  the  the  chase, observed,  actually forepaws,  teeth.  Hits  or  during  and  during  striking a  raking  o c c u r r e d as an  a  aggressive  these  bouts  the r  forepaws,  teeth,  devastating to  grasp  teeth,  and  technique  and  action  hindpaws  occurred  were  when  one  its  opponent's  f u r near  then  claw  its  has  been  cuniculus,  (See  with shown  of  the  the  XVI,  used.  The  combatants  back  hindpaws.  i n the  Plate  a l l  of  A  European  the  p.110,  was  neck  similar rabbit,  most able  in i t s  fighting  Oryctolagus  Mykytowycz  and  Hesterman,1975).  b.  Interaction  The  above  gradient  of  seasons, the  observed  i n the  (174.5  interactions  least these  compares  pens  Comparisons:  description  from  frequencies sexes,  Intensity  of to  (249.0 There  observed  the  a  aggression  intense  (pens  different  hours)  was  hare  to  or  (Chisquare=103.48,  varied forest).  types  those  difference  traces  interactions.  interactions  localities  f r e q u e n c i e s of  hours).  most  different  and  wild  snowshoe  of  d.f.=5,  The  between Figure  6  interactions  observed  between  a  the  in  the  types  of  p<.0l),  and  19  Figure  6:  A comparison of found i n penned  the d i f f e r e n t types vs w i l d h a r e s .  of  interactions  6050-  FREQUENCY  40  OF  30  V  INTERACTIONS 2 0 /  10 0 EARS UP  0  EARS LUNGE DOWN  INCREASING  PENNED I  I  HARES  WILD HARES  (N=  LEAP  CHASE  INTENSITY  661  (N= 6 5 0  c  interactions)  interactions)  HIT  20  from  Figure  intense in  the  6  can  be  seen  interactions, wild  rather  Figure intense d.f.=5, the  it  7  This  in  the  during  the  chases,  males were  (Figure  and  were"  of  h i t s ,  more  occurred  involved  females  in  8).  winter  frequencies  pens.  that  than  higher  leaps,  difference  months  i n t e n s i t i e s  g.  i l l u s t r a t e s  p<.0l).  summer  e.  than  interactions  that  months  and  were  more  (Chisquare=136.11,  intensity Male  in  occurred female  quite  during  interaction  similar  (Figure  9).  c.  Interaction  Table rate  1  shows  observed  in  observed  in  In  males  t o t a l ,  Rate  Comparisons:  no pen  difference  populations  manipulated  and  interacted  (Chisquare=82.35,  d.f.=1,  individually,  involved  females the  in  study,  were 14  e.g.  of  the  Pen  1  1979,etc.  2.  SOCIAL  ORGANIZATION  between  at  and  the the  non-manipulated a  higher  p<.0l). in'  rate  Figure  more  hourly  interaction wild than  10  interactions  different  situations  I  in  summer  1979,  2  Pen  females  that  males,  than  observed in  rate  populations. did  shows  15  of  interaction  were during  winter  of  21  Figure  7:  A comparison interactions  of the d i f f e r e n t types of found i n male vs f e m a l e h a r e s .  60 50  FREQUENCY  40  OF  30  INTERACT IONS 20  V  10 V  10  0  2L3L  EARS . EARS LUNGE UP DOWN  INCREASING  IZZ2J CZ3  MALE  HARES  FEMALE  LEAP  CHASE  INTENSITY  (N= 7 5 2 i n t e r a c t i o n s )  HARES  (N= 5 5 9 i n t e r a c t i o n s )  HIT  22  Figure  8:  A comparison of the d i f f e r e n t types of i n t e r a c t i o n s f o u n d i n t h e summer w i t h m a l e a n d .female hares. Notice the high frequency o f . " c h a s e s " e x h i b i t e d by m a l e s .  60 50  FREQUENCY 4 0 OF  30  n  V  /  INTERACTIONS 2 0 - /  0  10 0  EARS UP  EARS LUNGE DOWN  INCREASING  EZ22  MALE HARES  I  FEMALES  1  (N=  HARES  rJZL LEAP  CHASE  NTENSITY  175 (N=  interactions) 348  interactions)  HIT  23  Figure 9:  A comparison of the d i f f e r e n t types of i n t e r a c t i o n s found in the winter with male and female h a r e s . Females are a l s o e x h i b i t i n g a high frequency of " c h a s e s " i n t h i s season.  60 50  FREQUENCY OF  40 30  INTERACTIONS 20 io  "A  o  EARS UP  i si  EARS  LUNGE  I  LEAP  DOWN  INCREASING  I  0ZL  CHASE  I NTE NSITY  E223 MALE HARES  (N = 577 i n t e r a c t i o n s )  FEMALE HARES  (N= 192 i n t e r a c t i o n s )  HIT  5  4  3 -I  o  MALE-FEMALE INTERACTIONS PER HOUR  o  o  o  o  °  o  -H -2 -3  **  -4 -5  JL  J L  SUMMER Figure  10:  WINTER  D i f f e r e n c e s i n i n d i v i d u a l i n t e r a c t i o n r a t e s d u r i n g t h e 15 d i f f e r e n t s i t u a t i o n s observed throughout the s t u d y . On t h e Y - a x i s a r e r e c o r d e d t h e i n t e r a c t i o n r a t e o f m a l e s , m i n u s t h a t o f f e m a l e s d u r i n g t h e same observation period. In o n l y one c a s e d i d t h e f e m a l e s i n t e r a c t a t a h i g h e r r a t e t h a n t h e m a l e s , a s i n d i c a t e d by t h e " * * " d a t a p o i n t .  25  Table  1.  Agonistic interaction rates. were g a t h e r e d over the e n t i r e  Category  No.  of  Hr .  interactions  These interactions p e r i o d of the study.  of  Observation  #  i n t . / h r  Pens  642  174.5  3.7  Wild  888  249.0  3.6  Males  1 781  423.5  4.2  Females  1279  423.5  3.0  a.  SUMMER:  i .  Penned Hares:  Snowshoe therefore type  of  I  hares  f i r s t  social  are  observed  animals.  I  adult  into  1,  per  ha,  wild  which  occurred willow the  is  (Keith at  and  data  and  only  linear  ranking.  (1975)  animals.  18  over  the  Table  2b  i l l u s t r a t e s  to  before adult  studying  wild,  density  pathways. hours  hares  and  of  hares  of  in  the  at  piles 2a  f a l l s  hierarchy  of  presents  observations. defeated  observations.  interactions  hierarchy  hares  interactions  Table of  two  50  consistently  weeks 242  what  females  of  and  find  sources,  relations the  pens  forest,  Aggressive  organization  aggressive-submissive  the  density  83  dominance  in  actual  water  six  of  a  an  during  social for  three  regular  the  (7.4%) This  occur,  1978).  certain  study  large  peak  and  along  that  d e f i n i t i o n  sustained  food  recorded  hares  fact,  the  Windberg,  show  other  would  creating  to  in  introduced  times  a r t i f i c i a l  browse,  certain  Pen 3-4  interactions  These  hares  organization  free-ranging males  d i f f i c u l t  did  not  within which  In f i t  a  Wilson's  is  a  among  a  i t s e l f  with  set  group hare  of of 1,  26  Table  2a:  F u l l d a t a on i n t e r a c t ! ' o n s o b s e r v e d d u r i n g summer 1 9 7 9 ( J u n e 1 9 - A u g u s t  i n Pen 1 2).  c  Wl N.N ER  Hare  L  O  S E R  1  2  3  4  5  Totals  1  X  3  0  0  0  3  2  40  X  5  8  0  53  3  18  18  X  2  0  38  4  29  21  4  X  0  54  5  13  62  4  15  X  94  100 104  13  25  0  2 42  F  F  M  Totals  F  Sex Wt.(g) %Won 242  1720 1390 1640 1400 1020 97.1 66.2  Interactions  % W0N=  M  25.5 31.6 0.0  i n 83 hours*  (#won/#participated  in)  2.9/hour X 100  —  —  —  27  a  female,  being  interactions hereafter, with  the  f u l l  Data  as  A l l  of  evenly  In  in  from  Tests.  percentages not  dominant.  as  Table  this  Hare  a l l  i l l u s t r a t i o n  hierarchies  Chisquare  was  the  data,  Appendices.  of  most  whatsoever. only  subsequent  the  in  Table to  hierarchy,  male,  won,  d i s t r i b u t e d  between  an  a l l  in  the  data  analyzed  indicating  no  presented,  found  uneven  the  won  presented  are  be  been  showed  interactions  2b  and  have  hierarchies  a  hierarchies  2a,  presented  5,  from using  d i s t r i b u t i o n  that  hares  dominance  in  any  given  group.  Table 2b. The dominance d u r i n g the summer o f 1979 observation2.9/hour).  Hare # %  1  #  interactions  Weight(g)  In  Pen  observe  densities, as  in  Pens  density  I  female  F  M  1 720  1 390  1.  30  3).  Thus, hares  in  As  in  both  were  pens  25.5  31.6  0.0  F  .1 0 2 0  adult  female  and  may  occur  also  Pen  M  1 400  ha  compared formed  1,  a  dominance during  of  5 94  per  dominant  Pen'1 83 h o u r s  79  one  was  >  4  1640  hares  in in  51  which  A hierarchy  3  F  introduced  i.e.  >  66.2  differences  (Table  dominant. and  2  2 1 57  97. 1  sex  to  >  1 03  won  relations observed ( 242 interactions  in  two  because to  50  Pen  female  summer  of  hares 2  at  hare  hierarchies the  adult  of  males lower  per  ha,  the  lower  was  most  were  formed,  1979.  28  T a b l e 3. The dominance r e l a t i o n s observed in Pen 2 d u r i n g the summer o f 1979 ( 82 interactions in 41.2 observation2.0/hour).  Hare #  #  1  interactions  98. 1  Sex  Complete  In  May  females  and  two  males),  to  if  Females  as  i i .  WiId  these  4.7  they  As  hares  and  2(two  the  in  in  only  cover  top  of  only  a  summer  1  ranked  be  Table  4,  to  males  in  short  the  period  two  formed,  and  in  the  hierarchies both  interactions, of  adult  and  position  in  the  (two  females  again  throughout  of  Pens  adult  11.5%  continued  the  both  would  dominant  88.5%  II.  in  i l l u s t r a t e d  being  males  results  Appendix  attained  females  1040  hierarchies  again.  in  M  pens. whereas  interactions. in  the  May,  summer  female in  the  hares  at  1979.  Hares:  the  location of  if  females  had  in  observed  hierarchies  During  (14  42.0  males),  see  dominated  dominated pens,  I  hares  with  dominated  Although  total  to  once  formed  found  adult  female  hierarchies  the  43  1090  be  1980,  adult see  may  of  3  M  1700 data  >  69  F  Weight(g)  males  2  52  % won  were  >  hours  187  females  summer  of  described  one  I  observed  e a r l i e r  interactions, and  1980  but  male).  as  only Of  the  75  the  tagged  "Ditch".  involved 29  wild I  observed  tagged  interactions  a  hares  involving  29  Table 4. The dominance r e l a t i o n s observed i n Pens 1 and : d u r i n g the summmer of 1980. PEN 1 57 i n t e r a c t i o n s i n 6 .75 hours- 8.4/hour Hare #  1  Sex Weight(g)  2  >  3  >  4  24  23  42  23  87.5  43.5  57. 1  0.0  # interactions % won  >  F  F  M  M  1560  1 530  1 430  1 100  PEN 2 40 i n t e r a c t i o n s i n 6 .5 hours- 6.2/hour Hare #  1  # interactions % won  18  We i ght(g)  2  >  20  100.0  Sex  Complete  >  65.0  3  >  4  33  9  27.3  0.0  F  F  M  M  1410  1 250  1 1 50  1 320  data may be found i n Appendices  I l i a and  Illb.  tagged animals of opposite sex, 28 were won by the females, and one  i n t e r a c t i o n was won by the male. However, d u r i n g the summer  breeding season i t i s o f t e n p o s s i b l e  to  differentiate  between  male and female hares on the b a s i s of appearance, as a l s o found by  Lincoln  (1974)  heavy and paunchy whereas to August  males  from  pregnancy  and/or  snowshoe hares are  lactational  tissue,  are s l i m , as they do not gain weight from A p r i l  ( K e i t h and Windberg,  differences between  i n L. europaeus. Female  1978). On  the  i n appearance, I recorded a f u r t h e r  basis  of  these  34 i n t e r a c t i o n s  known females and unmarked males, of which 32 were won  30  by  the  The  known  tagged  females, total  male and  probable  females  b.  won  64  1979 was  two  were  involved a l l  in  he  lost  67  interactions  (95.5%),  observed  and the  i^.  early same  Penned  four  four  and  won  by  the  unmarked  interactions  interactions. between  the  males  males won  with  males. unmarked  Therefore, and  three  of  the  females,  the  (4.5%).  total  data  show  effect.  see  female,  during  if the  winter  months  their  social  summer  months.  the  i l l u s t r a t e s  in  late  organization  time  was  quite  1  was  winter as  I  the  the  new  the  dominant  organization  observation  in  or  social one  summer  it  the  and  This  by  dominant, hierarchy  a  male  months  one  of  the  dominant one  of  and  a  was  was had  adult  the  most  found  holding  pens.  the  females  I be  males.  Hare  male,  in  interactions  the won  Table  16.7  He to  the  was  in  summer  hare,  whereas  at  the  top  hare,  of and  observations over  5b  immediately  hierarchy. from  hours),  male  taken  Pen  the  continued  remaining a  from  in  Although  hierarchy  dominant  would  5,  (4.25  dominant  only  the  position  short  different  found  female.  dominance  removed  formed.  position of  I  therefore  hierarchy  percentage  determine  during  males  male  if  hares  adult  hierarchies. placed  to  occurred  three  organization, during  1980  as  5a  involving  the  snowshoe  Hares:  Table  to  was  and  WINTER:  I  2  females  by  the  i l l u s t r a t e s took  over  increased  his  92.5  the  after  31  removal  of  the  o r i g i n a l  dominant  animal.  Table 5a. The dominance r e l a t i o n s observed in Pen 2 d u r i n g the winter of 1980, before the removal of the dominant male ( 46 i n t e r a c t i o n s in 4.25 hours of observation10.8/hour).  Hare # %  #  1 37  interactions won  >  Complete  1450 data  may  30  15  found  in  3 10 0.0  F  1560  be  >  26.7  M'  M  Weight(g)  2  16.7  100.0  Sex  5  M  1 600 Appendix  1340  IVa.  Table 5b. The dominance r e l a t i o n s observed i n Pen 2 d u r i n g the winter of 1980, after the removal of the dominant male, #1 ( 53 i n t e r a c t i o n s in 4.5 hours of observation11.8/hour).  Hare #  #  5  interactions  % won  >  2 45  21  92.5  35.6  0.0  M  Complete  In  Pen  from  the  were  added  F  1560 data  1  may  two  previous to  the  3  40  Sex Weight(g)  >  be  1600  found  hares,  in  male  summer. pen.  M  The  Two  Appendix  2  1340  v  IVb.  and  female  3,  wild  hares,  female  hares  again  formed  were  a  s t i l l 1 and  alive male  hierarchy,  4, as  32  shown  in  dominant  Table animal.  determine dominant 6b,  had  most in  The  female  apparently  overcome,  removed been 2.  in  from  held  in  Table  dominant  interactions  i i .  Wild  I  Pen  one  of  the  in  It  move  is  hare  to  shown  Pen  2,  hare  should  be  noted  most  the  most  in  3,  to  Table  a  female,  that  hare  3  the  other  hares  had  weeks  before  the  observations  were  of  residency  any if  sex  introduced pen,  male  on  who  into  Pen  male  2  100%  the  effects  Pen  his  by  dominance  had  since  winning  was  dominance  residency  introduced pens  on  bias  these  dominant  holding  he  in  the  dominant  would  hierarchy  was  whilst  was  the  hare  male  Male  removal  been  1  had  from  became  of  could  e a r l i e r  1.  as  the  the  Pen most  observed  involved.  Hares:  wild  behaviours,  hares,  also  A  the  a  months,  see  the  which  social  Single  To  (#1),  in  6  o r i g i n a l  shows  removed  results  length  2  observed  ADDITION  2.  2,  female  two  Pen  hare  or  overrode  the  6c  I  Pen  animal.  for  of  2,  the  only  effects  i l l u s t r a t e d be  1  in  resulting  dominant  introduced  taken.  male  to  Pen  as  Pen  The  contrary  the  and  in a  p o s i t i o n .  resided  been  As  whether  but  became  6a,  populations  and  existed  organization  in  wild  hares.  OF A R T I F I C I A L  FOOD  :  Feeder:  station  hares  observations.  hares I  to  had  test  whether  found  in  the  penned  :  feeding  concentrate  of  I  in  a  observed  containing small  area,  i n d i v i d u a l l y  rabbit and  chow  was  thereby  tagged  wild  set  up  to  f a c i l i t a t e hares  (seven  33  Table 6a. The dominance r e l a t i o n s o b s e r v e d i n Pen 1 d u r i n g the w i n t e r of 1980, before the removal of the dominant animal (53 interactions i n 10.25 hours of observation5.2/hour).  Hare #  #  interactions  Sex Weight(g) data  may  Table 6b. The during the wint dominant animal 14.25 hours of  #  61 F  1 480  1 640  found  #  3  interactions  Weight(g)  1430  1 350  '  >  1  be  4.0  females)  Table  7a,  F  at  relationships  the  females,  in  the  1320  Appendix Vb.  S i l v e r  t.op  between  hierarchical similar  City  ranked  resulting  were  M  1420  found  four  thereby  4  52.2  four  the  >  95.8  and  dominating  F  25  may  c h a r a c t e r i s t i c s ,  21.4  23  data  consistent,  29.0  Appendix Va.  1640  dominance  28  M  F  males  31  24  Sex  Complete  1  dominance r e l a t i o n s o b s e r v e d i n Pen 1 e r of 1980, after the removal of the , m a l e #2 (36 interactions during observation2.5/hour).  % won  in  in  >  4  .9  M  be  >  21  96.2  Complete  3  26  % won  Hare  >  x 2  location.  hares  were  individuals in  a  social to  those  As males.  were  hierarchy.  These  structure,  and  found  in  penned  shown The quite two males hares  34  T a b l e 6c. The d o m i n a n c e r e l a t i o n s o b s e r v e d i n Pen 1 d u r i n g t h e w i n t e r o f 1980, a f t e r t h e a d d i t i o n o f t h e m a l e p r e v i o u s l y d o m i n a n t i n Pen 2 (32 i n t e r a c t i o n d u r i n g 8.5 h o u r s o f o b s e r v a t i o n - 8 . 5 / h o u r ) .  Hare  Pen2M  #  #  interactions  %  won  28 100.0  Complete  in  the  different and of  of  from  hares  movement,  categories Although  body  of  1 640  observed  dominant  unknown  I  described  population if  the  male I  by  vacated  hares,  removed  ranked unmarked  the  a  as  removing ranks had  total  hares hare  not  the  identify  etc. I  of  in had  three  the a  were the  of  data  and  be  from  be  over  i n the  pens.  males-  the  p o p u l a t i o n , and  61%  of  rate  the  the (22.3  on  marked  involved feeder.  experiments",  hares,  hares.  taken  in  interactions  removal  penned  additional  gathered  hares  speed  "Unidentified".  interaction to  was  hares,  their  two  involved  high  which  unmarked  because  dominant  temporary  situation of  "Unmarked"  the  occurred  1 320  t h e r e f o r e had  -"dominant  would  M  wild  h i e r a r c h y . Most  the  0.0  1420  presence  enough  for  19  F  supplanting subordinate  replicated  previously  the  hares  the  4  37.5  of  as  allowed  >  Appendix Vc.  (N=673),  formulate hares  was  known  interactions  to  in  feature  position,  interactions/hour) hares  found  I could  hares  these  11.1  pens,  which  8  1 370  One  the  9  F  d a t a , m a y be  winter.  1  M  Sex Weight(g)  3  >  This  by  a  first  distinguishing  this  was  female  Over  one  on  as  done  wild to  hares,  two  day  and  unmarked feature,  see  or  by  period,  third  top  hare.  This  and  was  35  Table 7a. The dominance r e l a t i o n s observed at a single feeder containing a r t i f i c i a l food, before the removal of the dominant animals (673 interactions observed •i n 30.25 h o u r - 2 2 . 3 / h o u r ) . Marked hares consisted of seven males and four females. P e r i o d o f o b s e r v a t i o n was f r o m N o v e m b e r 29 to December 7, 1979.  SEX  *-  #  males  17;101*  85-100  2  males  136;49*  50-60  1  male  1  female  17;95  40-49  3 2  females males  (RANGE)  &  & 42;34;1;10;14  0-30  indicates t h i s a n i m a l was r e m o v e d a t the end of this o b s e r v a t i o n p e r i o d as an e x p e r i m e n t a l manipulation. See t e x t for explanation.  found  it I  be  then  days,  males,  most  to  postions,  they  % WON  2  dominating  four  INTERACTIONS  dominant,  a  the  remaining  and as  found  shown  in  involved.  the  that  winning  The  manipulated four  Table more  One  although  interactions.  hares.  I  shot  this  animal,  and  male.  observed  each  were  of  7b.  than  of he  other  males The 97%  these  were two  of  the  been had  ranked  top  males  had male  population  for in  ranked  the  had  been  further top  hares  interactions  six were  in  which  previously  been  involved  just  a  in  only  tagged  as  17 the  0 o r i g i n a l  dominant  interactions, the  interactions  removal won  the  56%.  period  the  males  most in most  were  dominant which  she  dominant  of  removed. the  was  In  pre-removal  females  won  only  involved,  and  in  female,  a  different  44%  the  of  post-  animal,  36  Multiple  Feeders:  Table 7b. Winter dominance r e l a t i o n s observed at single containing a r t i f i c i a l food, after the removal of three dominant male hares (452 interactions in 13.0 hours34.8/hour). Marked hares consisted of six males and seven females. P e r i o d of observation was from December 7-11, 1979.  SEX 2  males  2  males  2  females  1  male  1 5  male & females  I apart  #  a l l  the  of  in  involving  was  construct ADDITION  a  in  my  had  males  number  8  of  the  the  only female  interactions No  FOOD  hares :  1,  I  feeders  male  120m  recorded  involving  area  unidentified  one  trough  was  one  360  male  dominant  '  and over  resulted.  obstructed,  and  of  separate  Location  Table  recorded  hierarchy.  OF NATURAL  At  0-30  (4.6/hour),  view  somewhat  I  two  hierarchy  involving  five  a  grid.  shown  2  at  hours  and  Location  low  hares  78  As  observations  This  wild  Telemetry  interactions  30-45  27;19;25;18;5;4  females,  feeder  45-60  50  the  At  90-100  45;43;18;18  females.  the  WON  &  interactions eight  %  74;48  observed on  INTERACTIONS  feeder  and  immediately this  resulted  animals. 46  After  interactions (one  (46)  regularly  surrounding in  many  39.75  hours  (1.2/hour),  interaction did  not  v i s i t e d  only).  allow  both  me  to  feeders.  37  T a b l e 8. The dominance r e l a t i o n s found.at the site of one of t e n feeders containing a r t i f i c i a l food located on Telemetry grid (360 interactions in 78.0 hours of observations4.6/hour).^ P e r i o d o f o b s e r v a t i o n was from F e b r u a r y 29 t o M a r c h 2 9 , 1980.  Hare #  21  #  spruce  95.7  68.0  45.5  1 500  feeder tree  1980,  tree  the  had to  observed were  been the  in  34  males  females.  The  1 420  1 490  1 460  IN  Figure found  in  season, dominant almost  the the at a l l  the  but  males end  11  did  WITH  females  hare.  the  The  four  not  254  enter  into  the  single  10m  seven  spruce were hares  males:four  43.5%  this  Two  white  dominant  only  in  3m  interactions  was  won  1979.  a  most  ratio  hares  where  added  of  of  her  hierarchy  previously  had  tagged  interactions.  SEASON  the  annual  During  most  the  cycle  winter,  dominant.  f i r s t  became  later  female  felled  City  total  December  were of  I  sex  i l l u s t r a t e s  snowshoe  the  A  freshly S i l v e r  e a r l i e r .  dominant  DOMINANCE  11  at  The  observed,  CHANGES  a  9).  hares  28. 1  1 490  placed  in  13.2  1360  /hour).  marked  40.0  F  (7.5  been  57  F  ground  of  53  15  F  hours  Six  1  F  location.  most  5  24  F  same  previously were  I  situated  (Table  interactions.  3.  1  February, on  >  77  M  In  7  25  Sex Weight(g)  >  2  93  interactions  % won  >  breeding  pregnant.  The  of  the  switch  I  non-breeding  females  session  The  dominance  in  became  April  back  to  when male  38  Figure  11:  The s e a s o n a l c h a n g e s i n d o m i n a n c e between t h e sexes o f snowshoe hares o v e r t h e p e r i o d of t h e study. P= PENNED HARES W= WILD HARES  zza  M a l e won i n t e r a c t i o n Female  n  1OO  won i n t e r a c t i o n  P W  \y  #0F INTERACTIONS 1 79  over  Ji  0  SUMMER '79  over  P W 1  v/ v/  female male  P. W i  r  P w  v/  J  r7-  141 64  0 73  , 52 67  WINTER (NOV.V9)  WINTER (FEB.'80)  SUMMER '80  39  T a b l e 9. The dominance r e l a t i o n s observed at the site a concentrated source of n a t u r a l food, a f a l l e n white spruce (254 interactions observed in 34.0 hours7.5/hour). Marked hares c o n s i s t e d of seven males and four females. f r o m F e b r u a r y 21 t o M a r c h 7 , SEX  #  1  male  3  males  1 1  male & female  2  females  1  male  1 1  male & female  INTERACTIONS  in  31-50  31,-29,-4  21-30  after  August, late  s i m i l a r l y  has  agonistic "aggressive during of  0-20  the  or  males  possibly  went  out  after  the  of  breeding  females  ceased  September.  provided  interactions  (1938)  51-80  DISCUSSION  BEHAVIOUR  have  occurred  81-100  25;46  D.  and  WON  26;11  in  I  %  was  &  condition  SOCIAL  observation  39;27;43  occurred  j_.  of  74  dominance  lactating  Period 1980.  of  the  descriptions  of  the  in  both  previously  leap"  and  breeding  interactions  are  the  snowshoe penned  provided  interactions.  of  He  wild  and  what  states  My d a t a  c e r t a i n l y  .  These  hares.  description  describes  fighting, season.  hare.  and  any  aggressive  more  I  of  interactions Only  these that  prevalent  Criddle  snowshoe  have  indicate  postures,  hare  labelled  the  only  occur  these  types  during  the  40  breeding  season,  The  gradient  members  related  passive  of  the  interactions reported  to  been  not  in  rabbits  1958;  2.  found the  most  a  and  those  One 10,  was  it  in  in  most the being  studies  interactions  of  has  never  comparative  apparent  this  and  of  not  rabbits,  within  the  f i e l d  aggressive 1978;  in  reported  subjective  facet but  have  occurred  been  for  and  found  postures  "boxing"  hares  compares  in  the  same  interactions study),  whereas  sexes  (Mykytowycz,  sustained  aggressive-  1964).  In  a l l  the  spruce  a  a  I  in  tree.  a In  of of  in  natural two  animals  different  observed  situations  wild  set  group  nine  addition,  hierarchy  is  among  in  feeder  fallen  most  The  .  10  interaction  the  (Lindlof,  hierarchy  experimental  a  have  increasing  HIERARCHIES  hierarchy  in  of  hares  Holler,  with  especially  1978).  interactions  Table  account  Table  sexes  hare  up"  may  an  aggressive  was  hare  that  relations  pens.  and  in  along  americanus  "ears  species  snowshoe  dominance  submissive  other  covered  Marsden  A  The  seasons.  "hit".  snowshoe  Lindlof,  between  DOMINANCE  the  studies,  was  and  in  to  Most  other  in  publications, within  Leporids.  which  some  observed,  ethology-  L .  judge,  1958;  in  a l l follow  up"  in  species.  in  found  "ears  interactions,  (Lechleitner, as  the  found  closely  during  interactions  found  of  interactions  occur  from  interactions  other  do  aggressive  intensity the  but  the  (Wilson,  groups  of  hierarchies wild.  I  situation  situations  in  hares at  also at the  the  3  1975).  I  kept  in  different  observed  a  location  of  wild,  one  at  a  41  Table  10.  AGGRESSIVE  A comparison of the a g g r e s s i v e snowshoe hare w i t h t h o s e found POSTURE  RELATED  SPECIES  postures in other  found i n the Leporids.  AUTHOR & A U T H O R ' S  PHRASE  Ears  up  S. S.  floridanus aquaticus  Ears  down  L.  europaeus  S.  s.  floridanus a q u a t i cus  L. S. S.  europaeus floridanus aquaticus  Leap  L. S. S.  c a l i f o r m ' cus f 1 o r i danus & a q u a t i cus  Lechleitner (1958) Marsden & H o l l e r (1964) - jump o r jump s e q u e n c e  Chase  L.  europaeus  L.  cal i f o r m ' cus  S. S. 0.  floridanus aquaticus cuniculus  Lindlof (1978) - c h a s e , but unusual Lechleitner (1958) -chase Marsden & H o l l e r (1964) - aggressive chase Mykytowycz (1958) - chase  Lunge  Hit  or  Fight  S. S. 0.  a method on t h e i r  &  &  &  europaeus L.  boxing-  &  californicus floridanus aquaticus cuniculus  &  Marsden & H o l l e r - avoidance Lindlof(1978) - threat posture Marsden & H o l l e r - threat posture Lindlof (1978) Marsden & H o l l e r - female charge  (1964)  (1964)  (1964)  Lindlof (1978) Schneider (1978) - b o x i n g ** Lechleitner (1958) - biting Marsden & H o l l e r (1964) ' - f i g h t i n g o r b o x i n g ** Mykytowycz (1958) - b i t i n g & r a k i n g with claws  of f i g h t i n g in which h i n d paws a n d s t r i k e  opponents both out with t h e i r  stand forepaws.  42  feeder  (Multiple  (Summer  1980-Ditch),  because  of  a  lack  Dominance phylogenetic (Breed,  et  (Taylor, 1971).  Feeders), I  of  hierarchies boundaries 1980),  1966),  to  Gauthreaux  hierarchies  which  show  animals  group  together  species  hierarchies musculus, (Ewing,  crowded  (Anderson,  1972-from at  t e r r i t o r i a l  raccoon,  the  overlapping  3.  home  T H E CHANGE  An switch to  IN  male  feature  female  that  of  dominance  dominance  during  not  mean  over  a l l  individuals  dominant  (Lindlof,  a l l of  animal  or  summary  schools,  as  Some  the  which  in  usually  the bands.  as  Mus  cockroach have  the  Sharp,  most  dominance  such  animals  and  or  form  in  as  In  of  shown  normally 1956),  or  maintains  1978).  SEASON  dominance  during  the  winter  individuals the  (Geist,  canadensis,  0.  (Sharp  the'  the  carolinenis  as  stations  lotor,  cinerea  kingdom.  a r t i f i c i a l l y ,  1980).  many  animal  t e r r i t o r i a l ,  feeding  WITH  across  canadensis  naturally,  a l . ,  hierarchy  animals.  Sciurus  herds,  europaeus,  DOMINANCE  does  most  L .  -ranges  unusual from  et  Procyon  hare,  such  or  a  extensive  the  usually  situation  Nauphoeta  Ovis  conspecific  experimental  European  in  1961),  known  species  an  together  Breed,  hierarchies  in  sheep,  found  are  identify  many  provides  in  to  s q u i r r e l s ,  hierarchies,  which  when  in  natural  cockroaches,  grey  (1980)  a  between  exist from  to  in  unable  bighorn  species  Some  was  one  interactions  a l . ,  dominance  and  animals,  summer  one sex,  was  sex but  was  breeding  non-breeding  of  opposite  hierarchy  the  predictable  season  season.  were  the  This  dominant  gender  of  according  the to  43  the  time  of  year.  contrary  to  the  a)  The  female and  female  which bias b)  in The  below  sex  long the  term  3,  It  3,  would  important in  Pen  animal  in  way the  over  the  than  they  reverse the  behaviour?".  I  the  #3,  long  who  term  about  however,  as  when  new  2,  was  introduced,  the  pen  this  percentage  did  and  of  of this  higher in  the  the  of  the  male  4  of  dominated of  the  male  previously  male  dominance  intersexual study  summer,  became  the  switch  is  11  (See of  while  also  interactions  shows  Figure  won  11).  interactions the  females  this  switch  by The  in  the  showed occurred  wild.  immediately suggest  seasonal  percentage  Figure the  of  male, new  in  .  examine  period  the  sex  dominant.  female  experience  any  d e f i n i t e l y  f a m i l i a r i t y  this  female  dominant,  male,  became  a  male  The  third  previous  to  a  overridden  experience  a  leaving  became  dominant  previous  brought  weeks.  ranked  was  because:  and  the  pattern.  question  the  appear  much  pens  the  had  was  female,  of  and  female  winter  The  dominant.  resident  combination  a  both  the  two  months  residency  1b)  removed  months,  only six  Pen  occurred  been  six  of  of  won  in  length  for  for  removal  in  before  males  the  pen  #4,  compare  each  the  this  had'  of  pen,  male  Another to  residencies  the  dominant dominant  pen  new  to  most  male  the  (Winter,  believe  became  hierarchy. was  in  in  that  female  losing  I  who  the  by  and  determining  believe  pen  lived short  suggested  hierarchy  dominant  l i v e d  hierarchy  'I  had  with  had  one  pattern,  previous  which  which  Only  the  arises, major  "Which  change  sex  occurs  changes in  the  its males,  44  and  that  it  is  the  manner  females  during  dominate  males.  approach  females  female's  reproductive  this  by while  Throughout in  keeping  the  female  inquisitive  posture  down  or  approach that  lunge,  by  males  manner-  by  in  in  a  by  in  species  Tamiasc iurus Crocuta rebuffed,  the  the  mating  season,  and  well,  the  to  allow for  douglasi i ,  crocuta,  the  (Chapter  male  3).  breeding,  female  in  if  a  ears  i n q u i s i t i v e  a  these  manner,  of  in  the  similar to  or  which reacts  into  the  aggression many  Douglas  more  than  During  further  snowshoe  several  any  an  female  and  in  This  allows  resource  the  occurs the  reported  reacts  lowering  1968), Male  in  the  continues  Such  1972).  season.  then  to  towards  different  subordinate  season,  (Smith,  a  a  access  example  (Kruuk,  approached  in  This  is  do  towards  g.  males,  female  over  mating  manner,  other  A  away  male.  winter  males  1964).  thus e.  the  Sylvilagus  Holler, and  rebuff  females,  been in  manner,  non-mating  interacting  females  often  the  moving  During  as  to  The  forward  and  non-aggressive,  constantly  s l i g h t l y  has  (Marsden and  aggressive  order  by  sequence  oestrous  an  posture  to  determine  cues.  directed  the  females  males  leaning  1958),  aggressively.  non-aggressive  courtship  and  the  to  olfactory  similar  appears  during  males  dispute.  A  season  closely,  up,  approaches  allows  attempt  using  aquaticus  approach  aggressively  is  ears  during  usually  non-courting  breeding  female  male  which  apparent  5).  in  males,  observed  winter,  S.  retaliate a  the  (Lechleitner,  and.  to  a  the  season  status  the  floridanus  female  an  (Figure  c a l i f o r n i c u s  which  mating  approaching  her,  L.  the  in  spotted hares, times  other  s q u i r r e l , hyena, even  if  before  45  becoming which  discouraged.  shows  male.  a  female  Severaid  snowshoe  hare,  Such  winning  (1942) as  a  also  one  sequence  of  or  interactions  refers  follows,  more to  "Although  a  dominance  she  allowed  dominated  before  pregnancy,  she  the  of  pen  pregnancy."  ruler If  summer be  the  males when  lost  being  considered  behaviour.  females  when  spots.  Although  observed  in  females and  hare  generally my  should  food, lose  know  the  change  occurs  exactly the  switch  that  such  males in  that  in  is  males  food,  over  water,  the go Yukon  for  when  this  were  dominant the  f a l l  in  the  wild spots  as  that  it  for  males  was  during  the  switch to  courting  the  seen  summer  in  the  the  wild,  water  were  interactions water  also  interact  a  in  summer.  November. females  pens.  in  male  dominance  snowshoe  hares  condition.  This  Although  occurred,  with  summer,  the  observed  late  during  the  change  male  to  resting  and  was  of  could  interactions  occurred  mating  treat  to  during  of  switch by  who  as  ground  food  behaviour  in  be  favourite  not  and  resting  out  to  female:female  responsible  they the  were  These  males and  and  a  herself  only  lose  over  the  during  also  food  correct,  was  dominance,  hares  data  in  non-courting  interactions,  spring,  when  dominance  penned  competition  water,  which  the  interactions  some  suggest  females  yields  switch  doubt  to  from  populations.  hypothesis  in  generally  such  is  behaviour  switch  show  I  over  If  for  interactions  there  summer.  of  however,  wild  then  switch  competing  female:female  the  a  Observations  indicate  l i t t l e  interactions  i n q u i s i t i v e ,  as  months  indicate  during  left  events  my  After the  I  do  not  observations this same  as  f a l l they  46  treat  any The  male  hare.  other  p o s s i b i l i t y  aggressive  after  levels.  must  It  are  probably  in  L .  be  also  results  in  the  determine  sex  however,  1974  for  ),  and  A  both  sexes  changing  was  the  basically  become  increased male  this  hormone  may  their  in  responsible  levels  as  occurs  increase  their  is  the  behaviours  which  Further  occurs  more  hormone  p o s s i b i l i t y  switch.  switch  hares  testosterone  third  dominance  when  of that  well.  observed  female  result  as  exactly  which  the  especially  (Lincoln,  of  a  remembered  level  combination  that  as  higher,  europaeus  aggressive  us  mating  is  research  the  for  f a l l  may  the  to t e l l  dominance  reversal. The  switches  beginning have  and  sometime  suggested  responsible blend  of  explain  for  why  the  such  an  is  individual  members  of  seasons.  both This  overlapping competing with  a  This  is  between  after  the  end  change  in  a  and  hare  sexes  occur  breeding  behaviour  hares  evolved.  the I  is  probably  a  certain  have  feature,  at  season.  characteristics  has  has  constant  throughout because  resting  female(s)  he  previously many  agonistic the  which  such  This  may  as  blend  a of  spots  in  species  with  and  non-mating  hares  have  intersexual  the  with  mated,  interactions  mating  snowshoe  Therefore,  or  to  sexes  follows:  ranges.  contrast  the  behavioural  food  in  the  male  ecological  switch,  as  of  Snowshoe  unusual  occurs  home  for  dominance  switches.  dominance  characteristics An  that  behavioural  seasonal  1)  in  winter  male  complete  and/or  which  a  lower  his  may  be  strangers, own  their  progeny. a c t i v i t y ,  47  or  actually  Tamias  hibernate  striatus,  competition  is  different lowered, the  Snowshoe  and  Windberg,  for  hares 1978;  (Severaid,  oestrous  determine This  requires  their  range  of  that  one  breeding  3)  This  Male  snowshoe  this  study).  mates  as  described  minutes  until  once  the  (Severaid, about  use  of  1971),  t e r r i t o r i e s ,  as  or in  to  find  check  were  the  a l l  only  males  begin  post-partum  females,  d r a s t i c a l l y  the one  within females'  l i t t e r , soon  the  and  go  treat  shorten  and  condition.  females  would  to  (Keith  successfully  determine  the  and  immediate  reproductive  and  there  year  compete  f i r s t in  per  out  females period  of  dominance.  1942;  5  are  session,  would  the-  (Geist,  l i t t e r s  have  are  summer, If  mating  and  constantly  condition,  indiscriminately. female  the  Intersexual  through sheep  four  must  females  condition.  only  to  males  males  chipmunks,  1972).  individual  up  they  the  bighorn  study), If  Smith,  as  1968).  produce  1942).  such  species  the  (Smith,  throughout  reproductive  other  separate  this  of  winter,  and  in  females,  which  therefore  in  as  squirrel  2)  mating  avoided  maintaining  Douglas  the  (Smith  ranges, by  during  Few  This  allows  above.  the  Females  for  nursing  leverets  are  weaned  This  feeding to  display  day  1942).  and  available  a  hares  the  other  feature  within  advances  species  of  show  no  parental  males only  at  v i s i t  the  home  courting of  their  and  the  25  females  range,  for  and  other  l i t t e r s  Tester,  approximately  means  (Severaid,  search  (Rongstadt  their  a l l  to  care  for  1971),  days  of  are  moving  are  age  therefore  males. above  characteristics  48  (or  have  the  been  various  fairly home  studied  features),  close  ranges,  sequential  breeding  and  the  to  grey  1966).  dominance  determine  squirrel  Grey  the  at  feeders, on  not  separate  the  non-breeding periods.  I predict  they  such a s w i t c h ,  if  but d i d  my r e a s o n i n g  (pers.  comm.)  is  to  one  ground  I  so  immediate  dominance  o c c u r r e d i n the  ranges via  throughout  hare  (Schneider,  post-partum mating, care that  dominance  switch.  European hares Sweden,  and  the  at  and m a l e s  found  formed.  Lindlof's  wrong,  but  an  and  European hare However, a  feeding no  sex  all  for  interactions  three  of to  intersexual  in  to the  several  with  their  periods,  Maaskamp, should  Lindlof station bias  in  of  his  data  the  evolution  litter not  exhibit  during  that  ecological  1980).  (1978)  the  the  o v e r l a p p i n g home  a p p a r e n t l y do  f i n d i n g s - suggest examination  the  similar  Both sexes  p r o d u c e more t h a n one  (Broekhuizen  therefore,  in  hare.  They have  1978),  into  s h o u l d show  m a t i n g and n o n - m a t i n g  fulfills  switch.  non-  data  switch  territories  which I suggested c o u l d l e a d  dominance  parental  snowshoe h a r e .  separate  snowshoe  European  characteristics a  the  i n d i v i d u a l s had i n t e r s e x u a l neighbours  The  of  in  2  correct.  reported a  s q u i r r e l maintained  years,  as  observed  come  interactions  Columbian ground s q u i r r e l , Spermophilus columbianus, the  of  overlapping and  reported  year,  Taylor  status  appears  s q u i r r e l s have  hierarchies  periods.  a full  Murie  but  (Taylor,  litter  throughout  sufficiently  year  show  any  I predict a  seasonal  observed the  dominance my  per  marked  winter  hierarchy  prediction  show t h i s  in  may not  was be  49  true. for  The  either  1)  the  England;  in  The  and  would  mating  appear  had  for  season  various  levels  the  of  the  mating  courting  "unmarked"  occurred  males  The  several  of  he  may  and  switch  to in  the  dominant  concluded  there  that  before  probably described and  could  interactions occurred and  males, males  in  the and  to the  therefore i n i t i a l  females  resulting  in  at  hares  behaviours.  highest  have  males  mated,  thereby  females, females  that  dominance  non-courting  these  by  lose  were  October  those  jackrabbit,  involved  won  to  the A p r i l  to  courting  to  courting  condition,  of  the  females  season  and  be  is  to  January  courtship  appearing  the  may  posture  hare  which  February  interactions  mated.  hares  If  sex,  (1980)-  towards  interactions. one  have  March  Maaskamp  major a l l  during 1974-  losing  males  of  both  His  may  (Lincoln,  Therefore  aggressively  session  exhibiting  and  been  when  mating  hierarchy  taken  snowshoe  in  hare  reacted  his  "inquisitive"  the  Although  snowshoe  2)  were  be  similar  resulted  females.  to  already  e a r l i e r  in  reasons:  Netherlands).  a  a l l  bias  Broekhuizen  involves  have  2  sex  observations  of  which  of  of  His  middle in  lack  percentage  unmarked was  a  hares  sex  bias  prediction  of  a  europaeus  be  non-mating  seasons,  of were  to  the  hierarchy. I  therefore  dominance should than  be  suggest, switch>  gathered  during  a  I  shown  have  individually  occurring during  transition in  involved  this in  my  in  L .  mating  and  period, chapter many  as  in  that  agonistic  L i n d l o f ' s snowshoe  seasonal  judged,  data rather  case. hares  interactions  may  leading  be to  50  the  establishment  exhibited  of  o r i g i n a l l y  Chitty  being some some and  in  dominant limited  behavioural  such  food  involved  access  fitness,  dominance  aspect  access of.the  examine  in  the  I  sources.  In  penned  in  snowshoe  hare's  following  both more  penned  and  Hares  stations view  further  in of  the  the  hypotheses  is  and  but  females  breeding  season.  I  is  access  the  winter  to  natural  In  this  ecology  of  interactions  most  more  in  of  were  agonistic  Male  terms  of  c r i t i c a l  in  reproductive which  I  shall  a  natural by  Males most  the  in  most  pens, in  feeding  the I  in  wild. suggest  regulation  dominant during  in  than  at  species  switch  in  involved  population  dominant  that  were  situation  this  were  behaviours  interactions,  hierarchies  behavioural  hypothesized  hares  intense  exhibited  warranted.  winter,  It  that  SUMMARY  stereotyped  in  aggression  investigation  some  seem  behavioural  dominance  wild,  in  spots.  would  populations.  and  formed  energetic  predict  access  hares,  and  chapter.  showed  wild  interactions,  females.  In  hares  (1964) and  I  that  over  females.  hypotheses,  preferential  resting  males  reproductive  further  stress  explained disputes  E.  Snowshoe  to  in  rank  warrant  interactions, lead  favourite  americanus  Davies  probable  have  L .  regulation  and  occurred  to  to  the  therefore  resource(s).  a r t i f i c i a l  Christian  aggressive  should  interactions  gaining  by  to  population  Considering  involved  hierarchies.  aggression  proposed  (1960).  costs  dominance  . sufficient  investigation as  of  in  the  the  summer  dominance  51  occurred  because  of  changes  towards  females.  Year-round  l i t t e r  breeding  seasons,  ecological led of  to  the  other  switch data  characteristics evolution  species w i l l  have  been  were  be  of  in  gathered.  in  approach  overlapping and of  the  the  lack  the  snowshoe  and  I  European  behaviour  home  of  dominance  examined,  found  the  ranges,  parental hare  switch.  males multi-  care  which  are  may  the have  Characteristics  predicted hare  of  once  a  dominance  the  necessary  52  CHAPTER  3:  DOMINANCE  A.  The  decline  feature Boutin L.  of  ame'ricanus  a  two  that  situation  spacing shall mates  mates which  to  exists  during  in  Wilson states the  this  may  lead  that  in  During  necessities  an  breeding  the is  to  to  reproductive  success.  shown  sources  have  aggressively of  food.  hares  interacting  over  forms,  and  shall  describe  assume  this the  examine  the  effects  access  to  oestrous  I  for  male  (1978)  male  has  voles, and  hares,  I  competition  in  to  dominance females.  of  for  food, Chapter  the hare  dominant.  being  dominant,  of  access  He shelter,  2  that and  and  snowshoe  concentrated of  penned  "bolt-holes"  wild.  this  mating  was  in  to  further  preferred In  but  appear  observations  ranking This  system  dominant,  p r i o r i t y  experimental,  access in  are  females  mean  some  occurs  hares  social  reproduction".  have  snowshoe of  of  aggression,  extreme  possess  "necessities"  natural  such  for  of  1974).  population  snowshoe  advantages  elaborates  compete  in  Myers,  Krebs  resource  season,  and  I  a  hierarchical  l i f e  hares  of  and  in  levels  c h a r a c t e r i s t i c  species.  winter  dominate  (Krebs  mating  whether  a  enclosures.  high  describing  discussing  "to  to  exhibit  summer  my  is  decline  l i m i t i n g  cycling  year.  (1975),  species  of  the  By  spring  spring  are  hares  the  the  a  determine  Snowshoe throughout  found  km n o r t h  behaviour. try  during  microtine  comm.)  SYSTEM  INTRODUCTION  numbers  cycling  (pers.  suggested  in  AND T H E MATING  chapter  system, males  and  and in  accomplished  I  then  gaining by  f i r s t  53  determining  the  receptive  females.  predicted  that  p o s i t i v e l y obtained  penned  most  correlated  per  the  hares  hares  PENNED  In  with  were  I  as  d i f f i c u l t  therefore  introducing  of  dominance,  have  success  the  was  number  possible,  features  to  observe  obtained  as  I  highest  assumed  of  then  t r i e d  in  wild  "hares.  existed  for  much  and  to  be  copulations  to  any  length  information verify  that  of  from these  METHODS  HARES:  April  1980  I  trapped  four  placed  them  in  Shortly  season  of  hares-and  Pen  1.  1980, placed  interactions  I  of  with  trapped  them the  in  in  reproductive  multi-coloured the  s i m i l a r l y  2.  to  hares  before  and  Pen  males  male  I  i n i t i a l tagged  observed  determine  discs,  breeding  seven  the  their  and  female  aggressive  individual  ranks  hierarchy.  Females according females removed  would  then  would  total  them  been  and  d e f i n i t i o n  male  the  ear-tagged  the  pen,  Reproductive  condition,  in  a  Wilson's  B.  J_.  in  male.  wild.  behavioural  males  dominant  success.  Snowshoe in  of From  the  reproductive  time  ranks  were  to  were the  the in  to  schedule  oestrus  dominant  introduced, move  i n d i v i d u a l l y  the  to  by  determine  dominant  to  the  experiment  started.  the  pen  after  females  the  position,  next and  three  highest to  I  pen  time  i f  11.  male  in  in  Table  the  shown  the  male  introduced  assumed  ranked  determine  if  a l l I had male  such  a  54  move  would  through #1,  the  and  4m)  a  his  experiment,  placed  with  U n t i l  increase  him  female  that I  13,  at  same  than  one  the  in  hare  time,  a c t i v i t i e s .  I  to  introduced  receptive  trapped one  male  time  success  to  of  see  in the  the  if  was  1  had  not  the  last  two  would  most  holding  capable been  reduce  Midway male,  pens  of  (1Om  in  mating  numbers  the  presence  competition  3  and  of  more  between  the  males.  Table  11.  Schedule pen.  Introduced A p r i l A p r i l A p r i l  of  Female  18 19 20  #  female  introductions  Female  1 4* 7**  Removed  A p r i l April April  #  into  the  Females  21 21 21  a l l  male  Present 1 2 3  Removed dominant m a l e , and females 1, 4, 7. Observed re-establishment of male h i e r a r c h y . A p r i l Test  of  April April * **  23 male 24 25  11 #1  3  with 5 & 13  D o m i n a n t m a l e #7 T h i r d - r a n k e d male  A p r i l female  #5,  24  prior  A p r i l A p r i l injured. #13 injured.  26 26  1 to  her  x  copulation.  involved  females,  whether  females.  subordinate  smaller  he  determine  female  obtaining  introduction. 1 3  55  2. WILD HARES: Observations  on  the  mating  from my own  studies,  and  from  working  other aspects of snowshoe hare ecology i n the same  on  of w i l d hares were compiled those  of  other  researchers  a r e a . These o b s e r v a t i o n s were obtained so that I c o u l d the mating of w i l d to penned  compare  hares, to see i f the e n c l o s u r e  was  having any e f f e c t s on snowshoe hare mating behaviour.  C. RESULTS  J_. A TYPICAL MATING IN THE The  following  PEN:  i s a general d e s c r i p t i o n of the sequence of  events I witnessed a f t e r a female had been  introduced i n t o  male pen. Although there were v a r i a t i o n s i n d e t a i l , pattern  of  events  appeared  similar  with  the  the o v e r a l l each  female  introduction. When the female was she would usually  investigate during  this  first various  introduced i n the e a r l y parts  of  the  interior,  the ground over which she had passed, would slowly and  the female orientate  attempt  and  i n v e s t i g a t i o n be approached by one of the  males. The female would move away, and the male, a f t e r  her,  morning,  sniffing  lope  after  to approach her. When the male got c l o s e to  he  would  his  ears  lean  forward,  stretch  out  his  neck,  up and towards the female, and a p p a r e n t l y  t r y to s n i f f her. If approaching the female from the f r o n t , the male would assume t h i s " i n q u i s i t i v e " posture,  and  attempt  to  56  sniff  noses  behind, would The  with  he  would  generally female  male,  and  c i r c l e being  then  f a i r l y  her  would  thrust  one would  his  again, would  the  his  male  male  move  be  and  ground.  ears up,  down,  ten  while  male,  emit  a  attempt  again,  but  at  twice  the  at  time  the  other  males  of  the  chase,  and  increase  other  join  even  The  in  during  the  them  with  the  become  each  h i t t i n g  away  from  away  The  speed  addition  her  the  female, The  male  onto  the  teeth.  Upon  legs,  and  hind  The  female  of  path  him.  in  be  the  during the  the  female.  The  chase  pen  became by  chase  other  chase.  b i t i n g  The  attracted the  female  previously  the  aggressive  and  the  speed  of  courtship  others  approach  his  from  apparently  extremely  subsequent  across  male  would  immediately.  would  cut  dominant  They  further  males  constantly  chasing  female.  and  mount  once  aware  quickly  to  begin  this  often  male.  would  was  the  stop,  squeal.  she  It  for  constantly  holding  his  the  male.  hindquarters.  but  observed.  as  onto  under  move  her,  back  out  immediately  would  with  the  the  while  sometimes  rear  from  her  female  would  would  male  times),  and  would  she  from  The  wait  She  logs  mount  raised  by  s l i g h t l y ,  under  female  area.  closer.  The  the  followed  speed got  probably  would  genital  Eventually  forepaws,  hares,  quickly The  to  and  he  over  usually  the  the  as  (approximately  with  of  on  approaching her  her  male.  with  ears  ejaculation,  the  low  behindhand  slowly,  hop  the  If  sniff  increase  and by  5).  to  away  continue  followed  female  attempt  would  with  (Figure  move  bushes,  crouch from  her  would  males.  towards They  the  the  each would  chase,  with  subordinates,  and  dominant  male  would  57  usually  be  caught  and  knocked  off  also  lowest  her to  the  the  behind  to  trees,  female,  he  look the at  or  the  around  a  high  approach occur  obtained  It  her,  chasing  had other  chosen males  A  male  sometimes  be  dislodgment).  male.  The  chases  about  the  pen  and  three in  in  an  which hit  This the  males,  a l l  right  The  their  and  could  pen  would  while  apparent  the  attempt  this one  the  the  chase  While  the  and  behind  him,  of  them  other  males  also  in  apparent  up  the  an  to  hide  under  logs,  ground.  ran  by  as  her,  they  rather  he  had  would  lost  the the  and  attempt  the  seemed  than  stop,  If  to  a l l find  that  males  would  also  interact  males  would  find  the  female,  begin  again.  Some  and  copulations  these  were  would  generally  male.  He  male  away  would  would  from then  keeping  dominant  male  would  often  locomotion  which  I  labelled  have  on  "realized"  dominant  rest.  holes,  female  front  progressed,  subordinates  away.  along  point  of  herself  the  in  bolt  attempt  male  ears  dominant  to  of  often  into  flattening  immediately  at  would  diving  sight  chases  the  female  by  by  lost  Soon  Eventually, by  males  stop.  was  the by  (a  or  leading  with  rate.  as  would  two  the  follow  male  would  female.  female.  ranked  moved  just  male  the  Once  male  top  chase,  would  female.  the  female  dominant  the  pursuing  follow  the  to  chases. the  males  closest  another  male  leading  other  stay  the  involve  During  the  by  ranked  avoid  from  to  mounted  happen  generally  to  able  monopolize  the act  the  display "thump  location as  other  a  the at  which  consort, males  female  and  away,  she keep the  an  unusual  method  of  and  hop".  he  at  As  ran  58  subordinates, to  0.8m)  t h e dominant male  and  long  hops,  would make e x a g g e r a t e d h i g h (up  which  "thump". He w o u l d do up t o f i v e sequence,  and  then  chase  began  of these  after:  oestrous  observed context force time and  f e m a l e , once a g a i n  the  female  then q u i c k l y continue  from  return until  p e r i o d s between Finally,  allow  late  female  by b e c o m i n g  extremely  leaps,  and  by  3b).  i n the normal  with  longer  end  hares  periodically  little  foreplay,  the  evening,  by  would  with  the  and l o n g e r . mating  s e s s i o n by would  t o w a r d s t h e male, and  responding frequently  with  lunges,  observed at t h i s  by t h e f e m a l e i n w h i c h she l e a p t  The dominant male  on him w i t h h e r f o r e p a w s would  s u b o r d i n a t e m a l e s would a l s o  be met w i t h  from t h e f e m a l e . T h i s would c o n c l u d e  (See  soon move away t o a n o t h e r  o f t h e p e n . Any s u b s e q u e n t a p p r o a c h e s t o w a r d s  female.  i n any  t h e dominant m a l e . She  c h a s e s . An i n t e r a c t i o n leap"  of  t o c o p u l a t e . At t h i s  or  aggressive  the male, and t r i e d t o l a n d  Figure part  with  h i s approaches  t i m e was t h e " a g g r e s s i v e over  copulation  would  do  rebuff  spot  afternoon  further  in  to the l o c a t i o n  i n t h e penned  c o p u l a t i o n s becoming the  hops"  t o h e r s a n c t u a r y . The c o n s o r t s h i p  t o -copulate  physically  return  her r e s t i n g  refusing this  and  t h a n m a t i n g . The dominant male would  she would g e n e r a l l y  often  "thump  audible  "thumping and h o p p i n g " . I n e v e r  t h i s method of l o c o m o t i o n other  a very  a subordinate  f a s h i o n . The dominant male would t h e n the  with  intense  the  female  aggression  the mating s e s s i o n  f o r the  59  2.  COURTSHIP  I  to  enclosure  penned was  WILD  observed  behaviours any  IN  parts  those  saw  I  those  obtained  wild  and be  observe  to  certain  in  could  to  courtship  mating  effects,  hares  unable  HARES:  in  the  the  so  from  complete in  to  if  hares.  as  determine of  Although  sequence  hares,  these  were  to  these  behaviours  behavioural  and  These  compare  hares,  wild  free-ranging  sequence,  pens.  could  decide  extrapolated  a  I  penned  thereby  copulation of  hares  I  were  did  I  from observe  identical  to  follows:  r  Males  "inquisitive"  observed twice (See the  this  again Figure  summer  Males and  posture  at  consorting by  determined  by  the  in  feeding. that  to  An  the  female from  a  different  location  in  posture  with  in  a  femalesMarch,  dominant  observed  which  male  many  I  f i r s t  and  then  in  A p r i l  male  times  were  of  (both  accompanying  oestrus  collected other  and  throughout  the  another a  new  though  (Severaid,  and  female  1-2  found  was  away  from  tagged)  days  are  In to  one be  observed  I  (both  was  however,  hares  1942).  myself  male  were  females  by  situation  female,  females  the  even  the  In  once  situations  dissected,  case,  1980.  and  these  p a r t u r i t i o n ,  observed  chased  female  feature  was in  appearance), his  post-partum  the  This  researcher  physical  reached  In  was  females-  consorting  unusual  was  term.  male  males  female show  dominant  another  the  area  approaching  a  This  in  when  1980.  observed,  the  a  5). of  once  posture  was before  reported case 1-2  the days  over  a  60  period  of  escorted.  2-3 After  be  described  her  l i t t e r .  days  days,  and  having later,  The  before  been she  male  was  observe  such  a  the  females  were  already  the  male  attracted  to  as  chase.  I  u n t i l  he  and and  therefore  a  paunchy  courtship  and  found  escorting  mating  in  in  trapped  consortship  to  several  follow  area,  (same  was  and  would  in  the  oestrus  to  penned  when  being  chase, have  the  actually  while  dropped  female  occur.  I  experiment  f i r s t  to  1-2 could  because  introduced  to  pen.  situation able  heavy  involved  post-partum  not  Males  appeared  a  he  times  at  him  as  least  example  of  a  male  the  pens  during  ran  35m  across  he  115m  the  the  pen  Courtship  chases-  In  hares,  a l l  in  other  through  and  to  I  observed  one  occasion,  slowly  apparently bush  was  through  an  listened  and  involving  a  female male,  I  hear  hares  from  attracted  experiment, investigate  30m to  a  when a  chase  chasing  away.  An  chase  occurred  the  a  marked marked  the  to  clearings,  another  could  this I  above),  undergrowth  mating  on  moved  chase  hares.  being  and  through  paragraph  hare  the  as  stopped,  other  through  chasing-  bush,  courtship  the  four  the  for  a  in  hares  male  repeatedly  joined  crashing  in  tagged  followed  female  other  extreme  dominant  in  male  involving  two  red  researcher  saw  six  squi r r e l s .  courtship  a  chase  A p r i l  line  like  the  forest.  I  observed  1980  boxcars The in  the  another in most  wild  a  train, detailed  was  the  one  chasing example  each of  mentioned  a in  61  the  previous  paragraph.  I  and  then  saw  four  (two  come  out  hares) was  similar  had  lost  the  sight  of  hares  the  marked  away.  males  This  bush,  and  the  other  hare,  by  a  male  in  to  penned  that  by who,  to  road,  one  marked  I  road  as  glimpses  bush  a  look  a  after  female,  had  the  a  short  male,  a l l  one  of  hare.  slowly  moved  went  the  to  courtship  be  back  ground  stopped.  (presumed  they  untagged  hare  sniff  This  when  marked  unmarked hare to  around.  pens  an  bush,  unmarked  thereafter,  untagged  stopped  the  two  the  after  bush  in  Following  Shortly  the  in  in  female.  hare  early  and  and  males  the  as  another  the  a  my  leading  to  who  They  the  Such a  into where  loping  female),  sequence  hares of  and  chases.  he  from  moved  involved  I  left  my  in  was  displayed  in  the  the  an  hare  forest,  location, by  and  followed  unmarked  hare  zig-zag  I  followed  and  found  completely.  He  came  and  loped  courtship  (Severaid, in.the  of  unmarked  a  scent.  sniffed  crashing As  edge  presence  observers  an  followed  their  my  as  the  away  ground,  ignored  above,  to  were  following  location  presence  moved  farther  female.  snowshoe  two  incident  female),  thumping of  same  though  of  Penned  heard  the  the  male  presumably  nose  male  3m  ignored  be  its  the  road.  of  again  tagged  the  tagged  with  within  in  a  the  the  forest.  close,  found  of  chases  hares.  (presumed up  the  got  after  Later  loped  the  males  unmarked hare  out  occurred  marked  oestrous  an  to  short  of  loped  male  the  similar  by  several  edge  the  returned  marked  the  behaviour  chase  four  the  to  to  aggressive  As  hares  saw  1942;  along  was  the  chases this  undergrowth, light  pattern  and  almost  up  old often  study). caught gone  62  (12:00  midnight),  l i s t e n i n g , caught  and  the  concluded the loose. an  had  six  For  unmarked  observed not  within  next  across males  contact  with  times  in  followed moved  "Thump males the  one an  and was  hop"  hares within  about  On and  "thumping 1m o f  Eurinating-  me  I  did  not  trapped, minute not  as  a  they  I  above  the  next  morning.  I  250g  in  she,had red,  had see  nor  lost  swollen,  been the  the  period.  and  very  accompanied  male  day  appears  when  by  escorting  next  It  lactating  to  locate chase. and  be  follow  males,  i t .  This  oestrous  I  observed  another  exhibiting  her  when  I  her  l i t t e r  trapped  twice  a  may  appears  females this  back  and  be  one  have  lost  to  they  behaviour  researcher  similar  sniff  behaviour  several  in  a  vehicle,  for  100m  as  it  roadway.  locomotion-  described  wild.  moving  old  chases.  was  thought  wild,  hare  the  vagina  was  to  during  of  her  she  hare  area  as  days,  unmarked  days.  t r a i l use  the  l i t t e r  five  she  ten  the  down  as  a  another  her  was  hares, a  method  she  over  Snowshoe forth  but  see  mentioned  previous  before  survive, the  and  male,  her  into  female  dropped  two  did  I  moving  days,  the  immediately  did  then  marked  she  last  but  e a r l i e r .  one  It  occasion  sniffing and  This  in  while  I  Eurinating  was  was  was in  a  hopping".  type  1979,  4-5  their  marking  of  by  several  dominant times  unmarked hares  c l e a r i n g ,  different  observing  the  locomotion  observed  large Two  of  with  hares  one  in were  or  two  approached  to  behaviour.  a  female  by  a  male  63  using  his  leapt S.  over  Eurinating  occurred  female  (Marsden  a  floridanus;  observed  3.  urine.  once  DOMINANCE  Lechleitner, in  the  penned  object  dominant  males  number  of  copulations,  f i r s t  determining  a.  of  The  four  on  I  did  hierarchy  #8,  becoming  Pen as  the  quickly most  based  into  on  pen.  twice  the  in  study  wild  was  each  males. male,  with  formed  the  in  Table  (8.0/hour),  of  a  linear the  or  1964-  in  This  was  hares.  to  find  success, I  did  and  then  if  i .  e.  this  by  observing  introduced  females.  linear  hierarchy  only  two  was  of  the  ranking. male,  the  13).  interactions after  r i g i d  This  and  with  (Table  most  12.  dominant  reformed  observations the  by,  c a l i f o r n i c u s ) .  success  shown  dominant few  and  Holler,  reproductive  of  1  follow  removal  was  subsequent  introduced  in  not  the  the  during  males  L .  ran  Pen  interactions  interactions  o r i g i n a l l y  rank  the  observed,  110  After  in  male  and  subordinate  reproductive  Hierarchy  hierarchy based  the  of  higher  than  the  SUCCESS:  portion  obtained  individual  Male  this  in  hares,  AND R E P R O D U C T I V E  The  their  1958-  as  a  #7,  second This  (18), new  from  Pen  ranked  male,  conclusion but  female  was had  1,  was  upheld been  64  Table 12. before the 110 Hare #  The male dominance r e l a t i o n s i n t r o d u c t i o n of the females.  interactions  #  in  13.7  - 7  Interactions  hours-  >  8  observed  in  Pen  8.0/hour >  '13  >  1  70  75  31  44  % Won  98.6  41.3  29.0  2.3  Weight(g)  1500  1230  1340  1330  Complete  Table  data  13.  The  previously 18  in  male  in  male 4.75  8  Complete  data  may  Reproductive  I  observed  introduced Table  injured  on  occurred  female  14  by  40  (mean=  the  with male  removal  of  the  3.8/hour >  13  >  1  10  9  100.0  10.0  0.0  1200  1270  1250  found  in  Appendix  V I I .  Success  females,  just  copulation  be  the  17  Weight(g)  in  hours-  Interactions  after  #7.  #  % Won  b.  Appendix V I .  hierarchy  dominant  interactions  Hare #  found  1  copulations  between  during  hours  5.7,  second as  the  female 8.  73.3  range day  2-9).  of  4.(Table  Female  4,  The  the  dominant  males  and  observations  the  seven  as  shown  dominant  male,  experiment.  This  injury  the  second  male  14), in  of  the  and  completed was  attempting  knocked to  escape  #7,  was  off  the  from  the  65 Table  14:  C o p u l a t i o n s observed during the mating experiment i n w h i c h seven o e s t r o u s f e m a l e s were i n t r o d u c e d t o P e n 1 w h i c h c o n t a i n e d a known h i e r a r c h y of males. ,  INTRODUCED Rank  FEMALES  1 3 &13 Total  of  Male  1  4  7  1  7  3.  *1  0  2  8  1  4  3  7  6  6  1  28  3  13  0  1  **2  0  0  0  0  3  4  1  0  0  0  0  1  1  5  4  6  5  7  7 Ii 2  40  Males  Total Copulations  11  5  4  REMOVED  9  injured  after  he o b t a i n e d  one  copulation  injured  after  he o b t a i n e d  two  copulations  I t e s t e d m a l e # l w i t h f e m a l e # 5 i n a h o l d i n g pen p r i o r to her i n t r o d u c t i o n i n t o the l a r g e e n c l o s u r e c o n t a i n i n g the o t h e r m a l e s .  66  melee,  raked  minutes  7's  Although did  male  to  a  short  male  13  any  A  was  s t i l l  of  injuries  similar with  not  on  to  while  Figure  indicated  completed  male  was  male  was  the  remaining  injured,  and  four  the  The  did  ranked  males  in  would  allow  the  removal  copulations  male  each  was  #1,  male  not  the  third  of  was  33.3%.  the  though  of  male  he  effects completed  as  shown  in  successful, second  After  ranked  4  injury,  even  most  the  injured  this  uninjured,  ranked  female  3  13  he  direct  apparent  while  removed,,  the  to  second  ranked  any  80%  further  even 25%  dominant  evenly  the  in  f i r s t  After  was  success  gain  male  blood.  injury,  was  the  10  the  male  ranked  dominant was  was  most  responsible  (33.3%).  accounted  an to  male  second  was  manage  pen,  of  7  his  male, the  the  This  examination  and  other  the  new  not  an  present  the  his  copulations, of  Within  with  obtained  when  copulations,  for  red  over  day.  view  dominant  and  increased  12b).  In  copulations  dominant  (Figure  further 1.  however,  (66.7%),  further  lowest  the  of  next  claws.  after  he  occurred the  was  rank  when  success,  male  66.7%  injured  After #8,  a  responsible  successful for  male  mating  copulations  and  any  was  exhibited  day  7  hind  copulations.  had  male  her  dominant  result  manage  dominant  12a,  he  previous  fight  did  his  with  surface  further  control  the  copulations.  ventral  retained  the  introduced  in  7  manage  contrast  underbelly  complete  male  not  7's  for  new  (20  of  male,  male).  d i s t r i b u t e d  copulations) had  been  copulations,  and  the  20%.  With  the  the  25  male,  #13,  remaining  d i s t r i b u t i o n of  dominant  of  copulations The  between  copulations (33%  after  d i s t r i b u t i o n the  males  of when  67  Figure  12a  Mating success of males with i n t r o d u c e d o e s t r o u s f e m a l e s , before the removal of dominant male#7, i l l u s t r a t i n g the e f f e c t o f i n j u r i e s on m a t i n g s u c c e s s .  100-,  %OF  66. 7  OBSERVED  66.7  V  5CH  33.3  33.3  COPULATIONS  V  0.0  MALE* RANK  1 ZZ3  I  7 1  BEFORE AFTER  INJURY INJURY  o.oK/1  0.00.0  8  13  1  2  3  4  A  TO TO  DOMINANT DOMINANT  MALE#7 MALE#7  (N=6 (N = 9  copulations) copulations)  68  Figure  12b:  Mating success females, after , m a l e . M a l e #1.3 period but s t i l  N= 2 5  of males with introduced the removal of the dominant was i n j u r e d t h r o u g h o u t this l remained dominant to male  COPULATIONS  10O-, 80.0  %OF OBSERVED  5(H  COPULATIONS  20.0  0.0  MALE #  8  13  RANK  1  2  #1  69  four  males  were  nor  when  three males  p<.01).  It i s clear  highest  mating  limit  a  male's  success  without  male  accounted  of  immediately  times  would  taking  mating  copulation female in male  a  was  up  returned.  have  be  not  male.  performed  copulation copulation  of was  i n some  him  chase  once  close  cases  by  "sneaky  each  the  5 such  A third  which  female's  and at other a  copulation  Secondly,  the  copulations".  A  sneaky  males male  male.  chasing  became  the  involved  The t h i r d - r a n k e d her  before  the  d i d not  necessarily  chase  immediately  a l l performed  accounted  male,  performed  male  obtain  male  mating  was  to  copulations,  way  It  strategy  courtship  dominant  generally  male,  again.  three  sneaky  the  the dominant  and c o p u l a t e d w i t h The  in  the  mating  copulations by.  complete  followed a  t h e two o r  male  I observed  which  Firstly,  to complete and  watch  as  dominant  third-ranked  present  often  the female,  beforehand.  able  the second-ranked  involved  his  the  by t h e d o m i n a n t  ways.  of h e r , and the dominant after  has  injuries  lower  the second-ranked  found  to  different  why  males  o c c u r r e d when  long chase  3  male  known  sight  d . f . = 2,  male  that  drastically  sometimes  the  courting  lost  then  in  s i t and  success  25.14,  dominant  evident  .05>p>.0l),  h i s rank.  dislodge  merely  subordinate  may  understand  would  most  It i s also  for  male  to  (Chisquare=  c o p u l a t i o n s not performed  the dominant  difficult  the  d.f.= 1,  30.49,  present  then,  altering  were  before  were  mobility  20-30%  with  (Chisquare=  success.  The  second-ranked even  present  f o rcopulations  involved session.  by t h e f i r s t  by t h e  the The  male  that  first initial found  70  the  female  seven  after  she  females  subordinate  had  been  introduced  introduced,  males,  and  four  three  into  the  f i r s t  copulated  f i r s t  pen.  Of  the  copulated  with  the  with  dominant  male. I  have  hierarchy  obtained  subordinate females,  male, male  immediately males.  to  male,  #7, In  mating,  I  capable  felt of  before  three  had  of  times  the  not  male  1 was  show  that  female  5.  time,  feature  completed of  female  Within  in  Within  pen,  a  of  30  a  a  before  these  I  For saw  the the  prevented  was  of  her  A p r i l  and  minutes,  male  the 1  the  was  most  time  had 1  experiment  by  and  involved, obtained  day. which the  in  became  increased  immediately  f i r s t  1  1  male  mating 1,  24,  male  male  became  chase, of  from  physiologically  re-introduced  male,  by  courtship  placed  end  male  chases  into  morning  hare  of  dominant  enter being  courtship the  most  was  dominant  re-started  subordinate  males,  and  the  he  I  snowshoe  copulation.  female  after  six  minutes  re-introduction by  5.  and  t y p i c a l  copulations  the  female  pen.  with  larger  aggression  observation  did  I  Another  level  to  1  5,  smaller  four  a  Even  new  a  f i r s t  the  courtship  that  necessary  of  the  male  state  most  avoided pen.  the  the  two  the  in  about  the  introducing  f i r s t  with  to  males  by  the  "sneaky"  evident  in  approached area  1  male  introduction  on  three  the  the  ranked  what  Therefore,  copulated  introducing  of  top  copulating.  in  into  three but  the  once  removed,  it  together  for  out  areas  order  the  During  only  chased  was  chases.  back  #1? 1  vacant  how  copulations,  Thereafter,  moving  1980,  described  63  after  he  hours  of  subordinate  male  71  defeat five  the  more  always  most  times  as  7  in  On males the  a  defined  male  but  successful, end  of he  the did  at  the  he  other  male  back,  terminated  by  examination  of  a  remarkable  the  most  after  of  p<.0 1 ) .  A  but  attempted male  defeated  a 13,  dominant  male onto  the  male  dominant  male  32  on  the  by  10.56,  Chisquare=  the  as  an  appeared ground out  d.f.=  and from  On  the was  male.  An  shows  completed  responsible and  as  1964).  copulations for  to  attempted  another  also 13a  at  copulation  e a r l i e r  Figures  Chisquare=  Holler,  copulation.  attempted  was  legs  female  attempted  male  attempt  moving  his  dislodged  involved  recorded  actually  dominant  observed  recorded  the  flat  found  (See  was of  that  an  hind  and  laying  performing to  his  the  the  in was  Marsden  complete  I  sequence  because  by  being  copulations  of  or  occasions,  males  dominant  removal  or  experiment  action  by  could  male  i.e.  attempted  removal  the  this  he  s i m i l a r i t y  copulations,  occurred  E a r l i e r ,  thrusting  Twenty-five  did  seven  the  once  This  back  unsuccessful  before  remaining  completed  mating  (Severaid,1942;  hindquarters,  the  the  reared  not  under  had  This  experiment,  paragraph.  usually  She  her  next time,  uncooperative. raising  had  unsuccessful.  rear  were  1  mating  copulations".  copulation.  copulations  the  female,  if  thrusting  attempted  of  during  being  not  end  male  interaction.  situation.  a  only  an  the  "attempted  mounting  copulate,  in  occasions  perform  in  after  similar  32  male  the  t h i r d - r a n k e d hare  male  If  before  immediately  copulation, the  dominant  for  13b--before 1,  p<.0l;  and  14.95,  d.f.=  3,  72  Figure  13a:  A t t e m p t e d c o p u l a t i o n s by m a l e s w i t h introduced o e s t r o u s females before the removal of dominant male #7, i l l u s t r a t i n g the e f f e c t of i n j u r i e s on m a t i n g a t t e m p t s .  100-.  100.0  %OF  66.7  50A  ATTEMPTED COPULATIONS  27.3  A  1  9.1 oo  A  oo[7]  o.o.o.o  MALE*  8  13  1  RANK  2  3  4  1  BEFORE  AFTER  I N J U R Y TO DOMINANT MALE #7 (N= 5 a t t e m p t e d c o p u l a t i o n s ) INJURY (N=  TO DOMINANT MALE #7 11 a t t e m p t e d c o p u l a t i o n s )  73 Figure  13b:  A t t e m p t e d c o p u l a t i o n s by m a l e s w i t h introduced o e s t r o u s f e m a l e s , a f t e r the removal of dominant male #7. M a l e #13 was i n j u r e d t h r o u g h o u t this p e r i o d , but s t i l l remained dominant to male #1.  N= .16 A T T E M P T E D  COPULATIONS  lOOH 80.0  %OF OBSERVED 50  ATTEMPTED COPULATIONS  9.0  0.0  MALE*  8  13  RANK  1  2  74  The male it  pen  was  late  at  in  the the  the  though  (See  Table  two  females,  same  time.  that  after  day,  the  mated  chase  even  was  the  evident  towards up  last  when  the  somewhat  females  at  matings.  In  female,  and  female a  experiment  aggressive dominant males, most  levels male  of  obtained  the one  copulation  of  with  day  with  the  I  males, may the to  both  be  by  play  a  other, would  both  Table  between  female  quickly, This in  took  occurred the  pen female  introducing role  in  monopolized this  should  suggest  did  that  increasing not  females  occur.  from  the  and  generally  females.  The  subordinate  one  female,  and-  a  2 the one  reduce  14),  with  female.  a  oestrous  a r t i f i c i a l l y  protect  by  larger  male  the  possessiveness  he  that  This  (See  as  one  males.  copulation other  and  no  to  experiment,  rebuffed  than  dominant the  the  remained  felt  ignored  in  introduced.  more  may  introduced  showed  s t i l l  males the  running  sneaky  was  of  and  if  attempted  copulated  female  other  enclosure  next  presence  the  being  morning,  natural,  between  juncture  the  female(s)  basically  were  male  more  competition  this  13,  dominant  The  addition,  and  eventually  11).  once,  By  new  e a r l i e r  3  this the The other spent male  regular  75  D.  Snowshoe Introduction and  to  this  aggression  dramatic The  hares,  and  courtship  aggressive proceed  lunges,  during  chase  males  chases,  occurred  during  with from  the  and  hits  upon  while  occurs,  males  at  even  act  as  does  zig-zag  pattern  ground,  the  by any  or  penned  snowshoe  competition of The hares  until  becoming other  for  portions were  Characteristics  male  male she  is  who  the  similar which  a  the  to  male of  towards  manifested  i t s e l f  those found  I  observed in  both  interact  session.  She  dominant her.  of  this  its  and  the  in  often  female.  Thus,  malermale the  highest  study.  observed in  before  female,  approaches level  sequence  are  the  mating  during  they  holding  the  high  observed  my  searching  defend the  in  period  This  scent  subsequently  mating  were  the  males  short  with  aggressive  which  to  a  female.  terminates  interactions of  for  able  exhibited  mates,  agonistic  the  this  movement  overtly  hares  rate  for of  If  following  dominant  consort  male,  rate  higher  searching  her  this  an  begin  Eventually  injuries  female  aggressively  the  Several  the  other.  keep  through  lose  of  to  this  penned  each  to  subordinates.  sight  Chases  attempting does  a  highly  female.  to  with  nose  in  interactions  interacting  a  He  is  americanus.  such often  entails  female.  L .  oestrous  the  competition  males  male  in  mating  in  several the  of  and  events  dominant  away  Males  a l l  of  discussed  levels  Courtship  for  the  mammals  high  involves  competing  experiment.  they  exhibit  sequence  often  speed  small  mating.  fast-moving  high  subordinate  other  chapter  states,  at  like  DISCUSSION  in  penned  circumstances  wild hares. were:  76  the  males'  being the  inquisitive  attracted  to  aggression  and not  in  penned  hares,  the  females  could  not  have  females  several  copulations),  s i m i l a r i t y  of  pens,  in  jackrabbits, presented 1958). when  L.  following  from  each  jackrabbits react of  in  either  do  an  in  wild, are  related a  also  other,  and  male  appear  to  hares  manner  as  (e.g. of  the  wild  and  features  found  hares.  to  snowshoe  hare  b l a c k - t a i l e d  those  I  met  have  (Lechleitner,  "inquisitive" with  posture  aggressive  to  those  described  to  6  or  7  jackrabbits  is  high.  B l a c k - t a i l e d  t e r r i t o r i a l .  to  I  oestrus.  the  free-ranging  aggression be  hares,  view  in  similar  up  e a r l i e r ,  hares,  In.  but  females  in  penned  wild  wild,  penned  hares.  often  involve  in  defending  the  and  locomotion,  already  similar  are  hop"  courtship  similar  Chases  not  other  in  and  females  antagonistic  approaches  Pregnant by  for  females  conspecifics  sex.  Several characteristics hare.  the  exhibit  females,  in  males  themselves  explained  mating  occurred  closely  and  with  wild  between  that  jackrabbits  americanus.  with  c a l i f o r n i c u s ,  approaching  postures  observed  and  females,  observed  presumably  not  mating  the  As  situation  I  and  for  Male  things  also  L .  chases  "thump  oestrus.  hares,  Courtship  the  consorting  were  suspect  penned  chases,  this  but  approaching  chases,  males  behaviours  I  when  characteristic  in  introduced  actual  only  was  observed  were  the  one  were  There  in  these  The  before  a l l  courtship  during  eurinating.  posture  Smith  other  species  to  I  those  (1968)  has  have  shown  have  •  described  that  in  the  similar for red  courtship  the  s q u i r r e l  snowshoe females  77  maintain  s t r i c t  t e r r i t o r i e s  their  individual  search  the  their  local  males  to  areas  lasts  aggressively  courtship the  chase,  female.  year,  t e r r i t o r i e s  t e r r i t o r i a l  generally  a l l  for  aggression  while  for  one  compete with  The  during  receptive  only  the  for  the  the  are  each  The in  abandon  season,  From  relax  is  responsible  S.  a b e r t i ,  which  one  female  usually  and  females  oestrus,  year.  oestrous male  males  breeding  females.  day  male  the  they  dominant  dominant  while  to  ten  during  staying for  a  closest  most  of  the  copulations. Tassel-eared  squirrels,  t e r r i t o r i e s ,  but  (Farentinos,  1972),  squirrels  have  s q u i r r e l .  Neither  to  a  show  courtship  TYPE  similar  mating  Smith  but  OF MATING  1977).  I  order  to  ways have  followed  for-  if  their  mating  in  (1972)  of  ranges  hare.  found  species  rankings  the  red  was  prior  the  These  able to  the  males  from  systems  in  sessions.  categorized 1972;  Emlen  which  dominance  wild  penned  hares hares.  Crook,  and  mating  sub-category,  americanus,  described  have  (Selander,  male  the  during  that  Farentinos  in  home  snowshoe  to  maintain  SYSTEM:  identify  The  nor  determined  occurred  the  system  (1968),  not  overlapping  in  hierarchy  authors  assemblages L.  found  which  Various  hares.  as  chases,  different  intersexual  dominance  interactions  J_.  a  have  do  mating 1977;  Oring's  system  was  polygyny, appears follow This  Emlen  followed  most  type  Oring,  c a t e g o r i z a t i o n , i n  explosive  the  and  by  snowshoe breeding  appropriate mating involves  to  system both  as  sexes  78  converging period. unable then  for  a  Emlen to  and  control  male:male  result  in  congregate  in  shall  the  2.  chase  f a l l  I have  into  be  paternity.  differences  in  of  sired  hierarchies breeding,  in  ova,  proportion  males  subsequent to  select  shortly.  the  the  each  both  wild  female,  Males  and  compete  have  already  consortship  by  a  (epigamic  opinion,  the  and  dominant selection),  red  b l a c k - t a i l e d  s q u i r r e l ,  jackrabbit,  category.  male  success.  I  had  no  a l .  (1974)  of  In  males  most are  animals  a  with  method  to  that  number  male,  species  and  (Geist,  for  1971;  were  sperm  between the  exhibiting  responsible  sperm  there  males'  of  actual  mixed  although  existed  would'  reproductive  the  in  different  each  hierarchy  determine  found  relationship by  in  assumed  positively  contributed  dominant  w i l l  selection),  dominant  a b i l i t y  male.  females  relationships.  mate  are  defense,  (intrasexual  the  chickens  positive  sperm by  in  domestic the  a  et  males  SUCCESS:  correlated I  for  relationships  my  system  most  as  In  if  mating  resource  oestrous  her  and  reproductive  Martin  inseminations  f e r t i l i z e  the  mating  obtained,  an  that  competition  dominance  AND REPRODUCTIVE  to  copulations  young  this  highest  success  during  s q u i r r e l ,  predicted  the  and  discussed  DOMINANCE  through  range The  female  tassel-eared  also  home  between  the  be  females  of  established  allow  suggested  the  attentions.  courtship  to  synchronized  (1977)  dominance  been  as  access  highly  d i f f e r e n t i a l  her  male  Oring  encounters  for  the  s h o r t - l i v e d ,  number  to the of  dominance  most  of  LeBoeuf  the and  79  Peterson, and  1969),  Holler,  and penned  1964).  dominant  males  receptive  females  rank,  have  I  responsible dominant  the  was  was  have  from  and  that I  have  as  a  penned I  have  o f snowshoe  hares,  a n d most  responsible  more an  dominance  Peterson,  four  opportunity  exist  most  densities, (Boutin,  f o r males  male  overlap 1979).  t o encounter  that  chases, and even  with  a  in wild  dominance There a r e  which  in wild,  suggest First,  as  observed  (Geist,  This  from  hares.  other  home  courtship  population.  exist  in  though  observed  between  however,  i n the wild most  explained  evidence  i n the pens.  matings  high  several  of c o u r t s h i p  observed  Second,  females  does  current  equally  also  in a wild  was  copulating.  i n penned  ecology,  features  hierarchies  1969).  near-peak  than  for  were  no d i r e c t  hierarchy  were  during  hare  The  males,  of  o f known  male  courtship  section  which  as found  r e l a t i o n s h i p may a dominance  also  the r e l a t i o n s h i p  success  a r e s i m i l a r t o those  and  hares  an  ranked  i n the previous  wild,  exhibit  I have  by h i g h e r  males  (N=40).  for  by  introducing  dominant  a c t u a l l y avoided  to substantiate  shown  penned  copulations.  However,  features such  responsible  breeding  from  reproductive  three  current  females  By  four  of the copulations  male  described  hares.  population  the  1980; M a r s d e n  to  hare.  t o be p h y s i o l o g i c a l l y c a p a b l e  characteristics wild  access  a pen c o n t a i n i n g  also  ranked  found  I  50-80%  of  et a l .  i n t h e snowshoe  that  of attempted  prevented  he  holds  shown  (Breed  priority  (7) i n t o  for  lowest  was  also  male  percentage  This  animals  well i nthe  Dominant  males  species  which  1971; L e B o e u f  ranges,  at  t h e home  ranges of  indicates  several  least  there  oestrous  is  females  80  during in  the  mating  snowshoe  and  Keith,  weeks  on  1979).  This  involved home  influence ecology  3.  three  or  span  most Thus,  with and  mate, rank  to  that  The the  area  allow  a  home  summer  of  1-2  (Boutin,  and  around  his  p r e f e r e n t i a l  access  to  in  .and  the  male  extended  dominance  c o r r e l a t i o n the  period  be  a  in  the  Meslow  to  ranges, for  a  dates  1979;  dominant  These  success  Keith,  occurring  provide  high  conception  covers  during  chases may  and  success.  and  or  features  of  mating  ranking  snowshoe  between  to  hare  dominance  wild.  1977; suggest  this,  to  participants.  Geist  bighorn  Cox  sheep.  species  in  females.  In  she  which snowshoe  given  is  to  observe  determine (1971) and this  choice  at  to  mate  to  do  provide  the  ranks  (1977)  strategy  was  chases  of  similar  a l l ,  because  her  are  choice:  the  female  interactions, various  situation  followed allow  (Cox several  the  l i s t e d  also  female  make-a  intrasexual  a  question  the  with  There  male  reported  the  does  hares  seems  LeBoeuf  hares,  a  e.  1980)?  snowshoe  i t s e l f  interesting i .  allowed  Farentinos, female  an  success,  not  who  chase  opportunity  asked  mating  is  courtship  from  have  determines  LeBoeuf,  1)  may  opportunity  authors  dominance  a  features  (Cary  occasions  weeks  though  CHOICE:  male  and  the  lead  even  local  overlapping  reproductive  regarding  only  1-2  a  dominance  provide  Several  select  four  reproductive  FEMALE  in  courtship  the  should  and  of  Third,  synchronous  mating  and  may  are  1968),  range.  period  rank  hares  in  females,  season.  in  several by  the more  receptive  females  to  81  ensure last  the  point  dominant  dominant became  obvious  male  became  copulations males  in  even  though  obtaining  resting  spots.  further  copulations  2) may  Any  which  because  the  again  one  in  good the  injured. he  after of  his  which  female  male  is is  males  together  further  over  a l l  food,  water,  f a i l e d  increases  other  to  By  allows  being  male  further  dominant.  stationary,  area.  also  with  most  small  to  no  the  and  obtain  injury.  courtship  the  access  This  when  obtained  dominant  male  condition.  experiment  He  remained  ranking  physical  pen  preferential  female  the  is  during  third  provide  determine  brings  The  feature  then  in  male  aggression  opportunities  Attempted the  males  uncooperative,  which  leads  to  many  to  mounts, to  gather  a  female  malermale  interactions. 3)  I  have  breeding too not  season  quickly follow.  thereby Upon  from One  after  assess  the  hypothesis  Hares is  are  or  a  fast  this  snowshoe  into  the  known  the  moves  away  male  generally  does  male  follows  quickly,  and  is  allow  male,  chase,  If the  to  copulation,  and  any  other  the  chase.  ranking  of  question  late  during  chases.  a  i n i t i a l  female,  induced  not  to  ensure  dominance  early  male  inquisitive  to  enter  raises  ova--  it  the  and  an  into of  that  attracted  way  enters  attracted,  but  are  completion  moves  the  suggested  of  The the  which  hares a  female  him  the  males  to  copulate.  male in  the  female various  quickly area  then males.  copulation(s)  are  could This  f e r t i l i z e  ones. ovulators how  (Martinet  long  after  and  Raynaud,  coitus  that  1973), ovulation  82  occurs.  In  European  copulation. into  a  Spermatozoa  female,  but  reproductive ova.  This  a l . ,  1967).  rabbits  and  known  the  uterus  The  capacitation  time  et  a l .  after  (1971),  snowshoe  hare  rabbits.  So,  copulation  always order  from  and  rats  copulations  did  produced  two  different  not  affect  more  offspring  copulation. (mixed  inseminations  that  others:  in  Beatty  et  a l . ,  actual  timing  determining  than  it for  males  1969;  rabbits-  copulation  et  may  than  other  copulations  appear  to  4-6  in  hours  a l . ,  1967).  but  Chang  does  with  longer  than  for  some  the  i n i t i a l  f e r t i l i z a t i o n s . copulations  found  but  that  the  some  have  than  produced a l . ,  less  of  the  found  in  more  male) offspring  in  1960).  cattleThus,  important  factors,  of  males  one  1974;  of  order  irrespective  Beatty,  paternity  et  et  and  be  f e r t i l i z e  (Dukelow  appears  more  always  to  female  rabbit  researchers  from  a  known  be  others,  Martin  of  is  consecutive  chickens.in  not  may  paternity,  sperm some  it  males,  Other  time  is  standpoint,  allowed  time  after  introduced  in  a b i l i t y  European  responsible  (1975)  f i r s t  (Dukelow  hare  suggested  been  Hays  oviduct  snowshoe  time  the  capacitation  the  hours  time  capacitation  inseminating  a  heterospermic  than  in  for  have  by  of  hours  the  spermatozoa,  could  Sharma female  ten  the  10-13  when  incubation  attaining  as  rabbits,  occurs  f e r t i l i z e  an  before  is  In  cannot  require  tract  period  ovulation  the in  e.g.  sperm  female  choice  competition. 4) but might  Sneaky  this judge  is  not  which  necessarily male  is  so.  I  dominant,  contradict have  suggested  but  I  assume  how she  a  female cannot  83  judge  the  v i a b i l i t y  subordinate  male,  female  may  sperm.  In  Beatty  (1960)  has  as  than  to  shown  inseminate male,  increase  conception  f e r t i l i z e  ova  offspring  when  found  under its  the  from  benefit  homospermic  only  sperm  males  female,  . in  by  rabbits to  the  mating  with  both  the  but  she  may  from  both  males  are  sperm  would  not  whose  mixed  a  viable  the  increases  insemination, were  to  a  males,  with  rate  sperm,  some  with  contributing  not  if  two  insemination  Therefore,  even  spermatozoa  sperm  males  viable  that  copulating  conception  of  rate,  also  possible  female  obtaining  also  of  increases. a  of  Beatty  above  that  By  obtaining  number  probability  viable.  sperm.  probability  the  the  one  her  his  thereby  the  addition  heterospermic more  and  increase  increases  of  actually  with  those  also  produced  of  another  male. It to  can  determine  dominance  thus actual  imperative  more  than  one  stone  sheep,  Panthera found  that  population beldingi. number 5)  and  because  of  during  a  d a l l i  of  of  27  have  Belding's  timing  of  Subordinate  and  were  they  did  hares  found also  in  and  and  actual  that  are  the  in  even with  in  the  the  l i o n ,  (1981)  have  in  wild  a  Spermophilus  dominance,  offspring  completed  a  elephant  1977 ) ,  sired  relate  from  mate  Sherman  s q u i r r e l s ,  not to  to  LeBoeuf,  multiply  ground  necessary  produced  1971),  Hanken  are  experiments  session  (Geist,  copulations male  been  (Cox  1972).  l i t t e r s  Unfortunately,  These  mating  stonei  experiments  offspring  system.  females  (Schaller,  78%  f i e l d  angustirostris,  Ovis  leo  that  mating  male  Mirounga  seen  paternity  related  more  seal,  be  nor  the  produced. a  sneaky  84  copulation  showed  post-copulation female  has  male's  rank.  successful eared a l .  also  s q u i r r e l s . found on  female  cockroach.  after  a  should  rank  to  or  sperm  mating  mortality The apply  to  a  change  5  of In  and  mating a l l  response  of  attract  any  s t i l l  present,  phases by  males  increase dynamics Adult  levels in  the  mortality  during  the  in  penned  hares.  had  the  had  the  change  proposed  during for  might  that  present  in  at  high  competition of  densities between  competition  mortality,  and  use  dominance of  genetic  low  should  densities, the  positive  females  area.  males  of  periods.  americanus  and  This  many  are  certainly  males  affect  to  system  males  would  between  a  Future  examination  allow  thereby  male  occurred  the  when  et  introduced  mating  the  of  the  females,  chases,  are  Breed  of  At  courtship  of  tassel-  female.  L .  cycle.  receptive  levels  the  closer  the  dominant  mine,  they  through  factors  the  high  to  with  and  Thus,  rankings  confirmation  ten-year  for  similar  cases,  i n i t i a l  subordinate  dominance  5  of  similar  after  in  male.  test  by  copulated  have  levels  males the  I  of  to  several  these  mortality  males  of  the  f i e l d  successful  High  shown  techniques,  males be  has  relationship,  system  although  intensify.  of  include  rates  another  occasions,  male  marking  yet  success,  dominant  experiment in  29 4  the  copulations  an  success  searching  would  In  and  with  (1980)  after  subordinate  research  1.  provided  Farentinos  cockroaches'  aggression,  interactions  aggression  (1980)  the  unusual  could  population  ways: may  increase  courtship Such  through  interactions  injuries  may  actual and  make  injuries  chases these  as  to  occurred  animals  more  85  susceptible  to  predation,  even  if  the  injury  i t s e l f  was  not  levels  of  chases,  by  f a t a l . 2.  Subordinate  aggression,  3.  moving  their  Juvenile  l i t t e r  of  marked l i t t e r Pen  of  2  a  female  six  during  dominance  oestrous the  during female.  introductions  of  the  indicated  the  even  though  suggest (male  mating, they  that  SUMMARY  of  both  lower  were  and  62,  1980.  and  wild  snowshoe  levels  were  males  show  a  both  selection  the  because  a  animals,  I  of  show  male  malermale  males,  and  an  that  of  Experimental  known  few of  this  in  hierarchy  responsible  capable  through  to  resembles  gained  the  trampled  hare  several  to  or  and  in  high  to  l i t t e r  jackrabbit.  males  epigamic  or  suggest  apparently  females  hares  the  described  p h y s i o l o g i c a l l y  snowshoe  I  was  involving  ranked  area.  page  the  system  injuries  on  involved  ranked  increase  female,  of  chase  b l a c k - t a i l e d  and  competition),  loss  penned  of  receptive  may  oestrous  old)  E.  mating  higher  This  immediate  courtship  chase  The  of  days  Courtship  a  the  the  mating  c l o s e l y - r e l a t e d  males,  (<7  courtship  through  described  observations  polygyny.  aggression  as  high  predation.  the. a p p a r e n t  overnight  that  to  the  during  increase  in  leverets an  Through concluded  of  avoid  areas. to  born  l i t t e r  to  males  new  may  just  because  wild  most  into  mortality  to  p o s s i b i l i t y  by  attempt  s u s c e p t i b i l i t y  leverets  injuries  may  exhibited  temporarily decrease  males  for  most  copulations,  copulating.  intrasexual (female  of  I  selection  choice).  I  86  consider at  a l l  hare  densities, effect  that  on  this  polygynous  densities  of  the  competition  male  mortality.  the  mating  system  ten-year for  would  cycle, females  be and  could  in  effect  at  high  have  some  87  CHAPTER  4:  FACTORS  AFFECTING  A.  Snowshoe changes  hares,  in  population 1978).  juvenile numbers  Keith  survival  is  best  hare  population  the  lower  and  is  a  early  and  social  the  may  HARES  the  which  and  this  relate  rate  factor,  as  it  increase  of  the  the  the  Davies, I  the  of  possible  Chitty,  w i l l  in f a l l  concepts  as  1964;  that  decline  in  behaviour  juvenile  juvenile  hypothesized  disregards  of  Windberg,  that  available  chapter to  of  major  trend  and  precipitates  being  the  found  They  spacing  In  Keith  demographic in  food  exhibit  follow  also  cycle.  (Christian  1974).  1974;  hypothesis  and  factors  behaviour  the  less  which  (1978)  change  This  stress,  Myers,  Myers,  survival of  species,  rates  important  throughout  winter.  regulating Krebs  with  result  behavioural  and  Windberg  juvenile  c y c l i c  survival  most  OF J U V E N I L E  INTRODUCTION  other  (Krebs  and the  correlates  numbers  like  SURVIVAL  1960;  examine  survival  in  on  early  how  snowshoe  hares. There history growth  is  of  l i t t l e L .  rates,  presents  americanus.  and  that  from  two  lactating  night,  some  l i t t l e  juvenile:juvenile data  information  and  returning  to  available  Severaid  interactions. radio-tagged  females  suggest nurse.  (1942)  observations  information  v i s i t the  Boutin  on on  Rongstad female their  females (1979)  the  provides  parental  l i f e leveret  care,  but  juveniletadult and  Tester  or  (1971),  on  snowshoe  hares,  have  shown  l i t t e r s  only  once  each  wean  their  suggested  l i t t e r s that  by  not  juveniles  88  weighing  less  than  500g  move  to  a r e a s of  low  adult  female  density. The  first  gathered  part  this  f r o m p e n n e d h a r e s on  g r o w t h r a t e s . The weaned  of  parental  second p a r t  juveniles  when  chapter  already  interactions  resident  the  cycle, life  ranges,  and  j u v e n i l e s . This question  was  into and  agonistic  stage.  what  behaviour  I  j u v e n i l e s w o u l d be  interactions  with  is  occurred  which also  involved resident  the  in  the  that period  in determining predicted  of  during  important  e f f e c t i v e during  Therefore,  pens  juvenile hares. I  i n w i l d h a r e s a s had  be  large  whether s i m i l a r t y p e s  s t a g e w h i c h i s most i m p o r t a n t  (non-resident)  that  of  densityintroduced  i n , and  lose,  adults,  than  more would  juveniles.  B. Immediately  upon  METHODS  finding  a  (leverets)  i n the  Thereafter,  I caught the  by  them i n t o t r a p s , o r by  chasing  They were a g a i n weight  to  w i l d t o see  then i t should  juvenile  resident  happens  examine  adult  i n t r o d u c t i o n s . If spacing  hare the  occurred  litter  they begin expanding t h e i r  observed j u v e n i l e s i n the  pen  and  introducing wild juveniles  contained  care,  information sizes,  will  e n c o u n t e r i n g u n f a m i l i a r a d u l t s and s t u d i e d by  provides  of  identification.  of  p e n s , I w e i g h e d , s e x e d , and l e v e r e t s w e e k l y by  w e i g h e d and  I50g,  litter  I  added  ear-tagged  regular  plastic  i f they ear-disc  had  hares them.  trapping,  n e t t i n g them w i t h a d i p  s e x e d , and a  young  net.  reached  a  for i n d i v i d u a l  89  J_.  EXPERIMENTAL  I  ~"~~~————————————  live-trapped  plastic  disc  smaller into  INTRODUCTIONS  wild  ear-tags.  holding  pens  introduced  10 w i l d  more  t h e summer  15  females  I090g, These  and  with  weights  ranged Snowshoe study).  28  hares  introduction,  The  sexes, the  and  pen.  throughout  2.  at  was  population  The  weights  of  wild  the following  by t h e i r  body  and r e l e a s e d  of from  of leverets the  3-5  resident  days.  1942;  this  and  non-  after  the  morning.  into  The  the.wild.  adults  which  796g).  of  had been  both  born i n  leverets  changed  t h e summer  o f 1980  of the experiment.  juvenile  (location between  resident  more  OF WILD J U V E N I L E  observed  the  520g t o  juveniles  (Severaid,  immediately  consisted  from  a g e o f 45  hours  recaptured  numbers  the course  the "Ditch"  litters  2-3  then  varying  differentiate  25-28 d a y s  included  females=  a mean  I  1 9 7 9 , a n d 20  introduced  with  i n the  hares.  juveniles  782g;  the  of  of  with  the evening  resident  i n weight  (males=  reactions  during  t h e summer  of  a n d f o r 2-3 h o u r s  OBSERVATIONS  I  o f 789g  them  individually  contained  and ranged  from  marked  introduced  t o 60 d a y s ,  for  held  during  t h e ages  the  and  1980. T h e 3 0 ~ w i l d  a r e weaned  hare  resident  were  which  males,  days  hares  introduced  of  weight  I observed  resident  pens  indicate  from  They  juveniles  15  a mean  juveniles,,  before being  one o f t h e l a r g e r  during  ^  HARES:  hares  described leverets  size  until  during  i n Chapter from  late  the  2 ) . I was a b l e first  i n t h e summer.  and  to  second  90  C.  J_.  PARENTAL  CARE  Trivers by  increases  the  reproductive in  includes and  the  other  in  after three  of  the  sat  other.  Within  group,  and  down  ran  to  her  hindquarters,  ground, 2-5  and  minutes,  observed nights,  and  and  the  became  active.  was of  same  in  the  a b i l i t y  this  feeding  is  sex  parental  c e l l s .  cover. the  huddled  The  following  leverets  300g.  In  Approximately l i t t e r s  to  investment as  then,  when  hence  were  the  two  present)  late hours  moved  to  pen.  The  leverets  at  any  one  together,  and  at  groomed  each  10-15 her  her  (and  well  weighed  (three  the  as  under  and  that  p a r t u r i t i o n ,  obtained  minutes  The  young  as  terminated  leverets  times  female  female  forelegs  these  a  hindquarters.  suckling.  with  licked  after  o l d ,  offspring  that  the.primary  weeks  on  began  nursing and  next  adds  care  were  "any  parent's  Parental  shortly  close  the  c e l l s  of  as  surviving  sex  leverets  huddled  sat  of  Trivers  cost  locations  the  of  depressions  12  investment  individual  cost  primary  the  hares the  an  chance  the  lactation  different  her,  than  parental  in  young.  three  sunset  location  the  l i t t e r s  approximately a l l  at  ground  observations  evening  of  the  found  together  parent  offspring".  cost  guarding  I  defines  success),  the  HARES:  offspring's  other  investment  SNOWSHOE  (1972)  investment  invest  IN  RESULTS  The  when  fed.  the  leverets  remained  extended  they  approached  but  female  three  l i t t e r s  on  were  usually  found  quickly  s i t t i n g  resting  Nursing  three at  on  lasted  moved  a  away.  on the from I  consecutive the  same  91  location, three  and  groups  with  were  suckled  from  complete  l i t t e r  this  awaiting  the  other  l i t t e r  Five  appears  minutes  may  to  have  been at  usually  However.,  unlike  rested  in  were  from  One the  screamed  ^the  her  leverets  identify  juvenile  occasion  one  10  minutes.  On  quite  close  young  began  charged  in,  doe  to  feeding, and  approached,  leverets  these  one  exceptions  the  that  one  stopped  own  parent  on  within  had  These  pens, by  hind  legs),  suggested  up  or adult  that  of  to  who  up  to  three  handled  began  and  the  indicated  the  before  an by  had  allowing  cover,  six  them,  on  within  females,  2m.  reacted  moving  towards  the  hares  to  the  adults.  the  leverets of  these  to  species  Leverets  such  in  screams  as  protect  on  their  leveret. of  their  the  hares  upright  c a l l  is  often  occasions  Other  distressed  distress  attempted  hours  the  many  several  (standing  the  in  predators.  to  with  l i t t e r s . found  and  hares  do  Members  posture  to  did  s o l i t a r y ,  young.  from  to  daylight  as  remained  snowshoe  l i t t l e  alert  hares  adult  and  female  During  investment  approached  males  they  different  offspring  quickly  assuming  by  adults  by  times.  under  parental  I  both  well,  reaction  the  as  other  remained  the  when  parent  any  of  of  investment  lactation,  groups  type  guarding  female  not  of  the  again.  parental  leverets  groups  female  though  and  females  five  later  once  major  leverets  often  two  one  containing  even  then,  feed.  The  the  as  assumed  females,  from  and  I  l i t t e r s ,  non-parent  l i t t e r ,  suckled  to  three  female.  different  the  lactating'female them  same  suckled  l i t t e r  suckling.  the  two  occasion,  the  the  a  Such  leveret young.  On  92  one  occasion  I  pecking  a  l i v e  hopped  around  prevent  the  predator. in  its  observed  one the  death The  beak,  Twenty  and  nest,  merely  a  ground  I  was  the one  kindling,  lactation,  but  attempted  to  2.  SIZES  LITTER  leverets  and  per  Keith  smaller.  Figure  by  the  canadensis,  female  excitedly  physically  attempted  attacking  the  up  did  the  to  smaller  struggling  young  i f  both  a  a  l i t t e r  young  leveret  of  from  or  penned fur-lined  some  under  cover  a  damp  fur  in  w i n d f a l l .  recently  born,  for  as  up  to  Parturition,  and  night.  male  hares  their  or  provide  parental  indications  had  up  the  season  N=7)  any  care  the  to  females  predators.  than  to  four  l i t t e r s  was  smaller  the  later  per  breeding  (mean=  l i t t e r s  (mean=  3.7 6.8  (Mann-Whitney  U-Test:  p<0.00l).  Cary  f i r s t  in  wild  hares  also  three  to  nine.  increase  in  weight  ranged  i l l u s t r a t e s  or  1942).  day  the  RATES:  hares  found  under  was  have  limited  weak  by  structured  tree,  penned  were  N=8)  the  hares  their  sizes  of  a  together  leverets  female  l i t t e r ;  produced  prepare  any  l i t t e r  were  base  A N D GROWTH  (1978)  never  (Severaid,  during  l i t t e r ;  14  the  there  f i r s t  L i t t e r  the  birth  female  per  not  furred  protect  Penned The  did  f u l l y  The  Perisorcus  while  picked  determine  occasion care.  but  leverets  at  after  no  leveret,  offspring  clumped  occurred  parental  leverets  of  to  jay,  away.  females  p r e c o c i a l ,  season.  flew  able  hour  its  grey  eventually  depression  often  On  of  l i t t e r s  The  old  bird,  bird  females. but  day  a  l i t t e r s  from  the  over  were  time  of  93  those  leverets  determined while  L.  showed  known  a The  b i r t h  to  14.4g  day If  rabbit  the  the  chow  at  INTRODUCTION  Wild pen  determine from born  i f  the  agonistic  hares  had  resident  days  the  hares  that  wild  surroundings.  over  my  in  pens  at  weight  day  from  penned  the  hares  to  days  60  than  applies  on  birth  f i r s t  higher  25-28  feeding  and  the  his  for  At  per  was  (1942)  weaned  wild  weigh  425-  vegetation  and  age.  adult  were  juveniles  hours).  moved  Agonistic  when  about  were  resident  would  Resident  summer.  (44  and  juveniles  summers,  juveniles  This  their  period  Age  17.ig  (non-residents)  resident  occurred  of  from  JUVENILES:  interactions of  of  (N=13).  I  leverets  study).  (r=0.985)  in  f i r s t  this  birth.  the  gestation  59g  age.  found  leverets  juveniles.  pen  introduction throughout  OF  of  of  c a l c u l a t i n g  1942;  of  date  finding  The  Severaid  leverets  by  average  days  rate  dispersing  resident in  60  growth  juvenile  containing  an  exact  by  relationship  by  seven  or  average  recorded  observed  either  (Severaid,  gained  Yukon,  the  mating.  an  linear  knew  damp,  days  approximately  in I  36  leverets  Maine.  480g.  is  I date  of  weighed  per  hares  s t i l l date  strong  days.  3.  were  young  which  parturition  americanus  the  in  the  they  mother's  for  be  data  no Upon the  interactions  hours),  both  to  investigating  between  the  to  hares hares' the  other  introduction, pen  of  during  at  manipulations  a  d i f f e r e n t l y  resident  and  into  hares  consisted  on  gathered (140  juvenile  treated  juveniles  The  introduced  the  times penned  most  non-  their  resident  new  hares  94  Figure  14:  The r e l a t i o n s h i p , o f a g e vs w e i g h t l e v e r e t s - N = 7 1 ; r= 0 . 9 8 5  in  penned  LEVERETS AGE VS WEIGHT  AGE(DAYS)  95  and  the  introduced  investigation, introduced  and  encounters, water,  or  when  hare,  approached  juvenile  occurred  residents  or  approached  conversely  sniffed  a  resting  spots,  30  the  when  resident  interactions  As  sniffed  the hare  resident:resident  over  meeting  this  introduced  in  occurred  when  during  and  the  hare.  also and  often  access  along  to  food,  w e l l - t r a v e l l e d  pathways. A l l aggressive  of  interactions  non-resident hour  than  P<.001).  were  by  then  dividing  Table  15  also  interactions juveniles  by  the  shows  hares,  residents,  and  56  hares  lost  adult with  residents, juvenile  they  10.83,  Of  residents.  interactions  more  resident  number  involved 1,  of  the  lost  50  345 of  in  shows per  U-Test: was  interactions hours, in  lost  more  than  did  and  the  pen.  of  the  resident  p<.00l). with  both  interactions  interactions were  of  present  juveniles  401  15  juveniles  observation  interacted  (14%)  and  d.f.=  (86%)  312  in  juveniles  were  the  345  Table  for  of  introduced  juveniles  residents.  of  involved  (Mann-Whitney  total  number  number  which  introduced  Introduced  the  the  that  (Chisquare=  Introduced juvenile  by  rate  were  hares.  juveniles  dividing  in  involved  interaction  juvenile  juveniles  resident  were  resident  hourly  calculated a  with  juveniles  The  involving  introduced  with  56  and  involving  the  were  with  resident  interactions the  adult  adult  juveniles.  (90.4%)  interactions  with (89.3%)  96  Table won  15.  by  Hare  Number  introduced  Type  #  of and  interactions resident  Interactions  #  p a r t i c i p a t e d  i n ,  and  #int./hare  hr.  number  juveniles.  Hare  H r . *  % Won**  Introduced  401  140  2.86  10.8  Residents  296  681  0.43  57.4  *  #hare  hours=  (#hours of o b s e r v a t i o n ) X (# juveniles of that c l a s s i f i c a t i o n p r e s e n t , i . e . introduced or  **  4.  % Won=  (# interactions won/total participated in) X 100.  OBSERVATIONS  I  wild  Approximately ear-tagged  juveniles one-third  wild  the  hares  discs.  August  20,  1980.  juveniles  in  of  juveniles  were  to  30  36  dominant  interactions in  the  "hits",  and  juveniles.  >  pens,  to  i.e.  approaches  interactions  hour  for  Adult  second  were  juveniles,  adults under  at  a l l  over  by  been  of  62.5  present)from dominant  and  to  had  agonistic  f i r s t in  similar  up"  and  "Ditch".  period  juveniles  were  i n i t i a t e d  a  were  (83.3%),  "ears  70  juveniles  hares  l i t t e r  the  observation  recorded  Interactions from  wild  hares  interactions  (77.3%).  observed  I  juvenile per  25  between  and  interactions  June  (1.1  of  plastic  involving  #  JUVENILES:  interactions  with  interactions hours  OF WILD  observed  between  resident).  17  l i t t e r of  to  22 those  "chases"  both  to  adults  and and  97  D.  Keith survival  and is  with  the  year  cycle.  the  l i f e  annual  describes penned  j_.  Windberg  the  changes  portion  of  is  hare  known  the its  most  of  l i f e  juvenile  closely  associated  numbers the  i n i t i a l  that  over  f i r s t  part  of  few  this  history,  the  as  ten-  months  of  discussion  observed  in  CARE:  observed  evening,  1-2  during  suckling  hours  this  after  daily during  additional  nursing  (pers.  found  comm.)  during  a  Through  (1971)  found  the  l i t t e r s  Maaskamp once  per  1-5  minutes. hours  day,  and  2-5  two  1-2  sunset,  of  in  female  that  hours the  24 L .  the  breeding  the  telemetry  the  as  hare  mentioned  hours  snowshoe  and  Tester  hares  v i s i t e d  Broekhuizen also for  a  time  view  and  nursed  of  above,  only  period  continued  actual In  an  24  penned  snowshoe  few  S i n c l a i r  per  Rongstad  once  say  follow.  of  late  have  cannot  sunset,  season. data  I  hours.  European  though  But,  europaeus  after  the  only  period  telemetry,  every  even  not  during  nursed  I  observation  lactating  concluded  minutes.  did  radio  once  l i t t e r  suckling  hours  only  darkness,  one  of  Lactation  during  observations  use  usually  after  for  Each  period  48  only  (1980)  sunset.  only  that  leverets  complete  continuous  hares.  by  period  observations  changed  shown  feature  snowshoe  l i t t l e and  have  animals.  I  1-2  in  hares,  this  PARENTAL  their  (1978)  demographic  However, of  DISCUSSION  of  to  of  occur sunset  S i n c l a i r ' s and  the  98  once  daily  likely,  lactation  that  L .  Five  by  two  americanus  minutes  or  huddled  together  of  groups  (3)  assumed  each  females  than  nursing. a l l  "that  40m If  they  accepted  approached nursing  bout  Rongstad  the  b i r t h  was and  site.  they  same  authors  site  of  the  Male  evidence  of  precocial  leverets  were  fallen observed females  the  even  pens  by  the  and  Maaskamp  to  my the  nursing  the  non-related wild  when  born  gathering  for  arrives,  to  nursing (1980)  in  they  lactating  the  female The  area. was  have  was  In also  numerous  hares  snowshoe  location  are  nurse.  site  European  findings  though  for  leaving  the  have  l i t t e r s  them  female  study,  I  by  the  the  location  reported  doe  allowed  began,  which  hares. the  The birth  l i t t e r .  no  good  In  hours.  and  when  the  free-ranging  suggested  hares  under  when  24  appears  therefore  nursing  together  then  Broekhuizen  snowshoe  found  of  mixed  (1971)  similar  and  l i t t e r ,  some  mix  terminated  also  stable,  different  and  are  make-up  young  nursed."  wild,  The  per  it  nursing  (1980)'  s t i l l  of.  once  Maaskamp  may  Tester  species,  actual  groups.  allow  and  l i t t e r ,  observations indicate  are  only  separate  the  and  the  the  a  apart,  nurse  quite  did  Broekhuizen  e.uropaeus  less  was  related  before  in  was  apparently  leverets. L.  group  also  more  leverets the  closely  cover  trees.  at  the  Neither  their  no  parental  nest-building  nest-building dropped  provide  found bases  by  huddled of  hares.  together  in  trees,  or  (1942),  in  L .  l i t t e r s  almost  whatsoever.  female  Severaid penned  care  under nor  in  of  depressions the  boughs  Grange  americanus, anywhere  L i t t e r s  I  but  (1932)  found  their  of  the  pens,  99  but  usually  care  by  under  nest-building  c l o s e l y - r e l a t e d holes,  and  a l t r i c i a l 1972), nest  provide  building and  is  young  by  hares  from  attempts  never  which  a  and the  the  and  is  by  to  made  female  spp.,  Rongstad  determine  l i t t e r  to  to  if  protect  parturition  she  a  protect  the  shallow for  their  et  a l . ,  difference  differences to  in  between  production  naked,  female but  have  ravens,  protect  however,  her be  possible  Thus,  protect have  from  interpreted  identify  offspring.  young  no  the  their  and  of  a l t r i c i a l  of  to  to  by  adult  rather  than  whether  or  is  indicate  not  s t i l l how  complete  f i r s t be  an  actions hares as  to  some rabbit attempt could, locate  guarding  female  to week  of  European in  a  data  enough  the  coronoides, The  k i l l e d  not  near  for  female  incident  and  were  enough  predators.  effectively  observations  a  the  telemetry  always  close  their  but  one  off  The  appeared  movements  question  recorded  (1971) was  protect  myself),  young.  Corvus  to  fought  reported  these as  it  been  predators,  young  her  wild  chased  two  attempts  (1942)  Tester  repeatedly  I  This  in  or  (Sorenson  and  jays  and  (1959)  hares  parental  found  l i n i n g  linked  successfully  them, may  weak  Severaid  Mykytowycz  and  1961).  hares,  of  burrows,  ecological  (grey  succeeded.  penned  in  some  assistance.  to  that  rabbit  probably  young  lack  vegetative swamp  main  from  nesting  (Lockley,  predators  Mustela  presented  after  of  This  rabbits.  leverets  her  fur in  cover.  different dig  rabbit  one  of  which  found  precocial  Female  enough  form quite  a  rabbits,  furred,  weasel,  as  European  fully  in  is  rabbits  young,  and  hares  some  of  snowshoe  open  to  debate.  weaning  in  snowshoe  100  hares  took  place,  returning have to  to  data  one  female,  to  its  unlike  summer,  periods that  From  the  OF WILD  may  not  return  to  the  themselves  at  about  this  age,  increasing stated  is  factors  for  passively, s i t e .  days  by  early  2-3  (Christian I  the  of  however,  kept  nursed  been  This  last  l i t t e r  for  of  extended  (1980)  not  the  amount  of  state  returning  address  the  left  and  that  winter  remaining  fend  for  Windberg  this  browse  480g.  1960;  portion  in  (1978)  factor  in  survival  available  possible  Chitty,  does  vegetation  hypothesis as  just  about  important  and  1964;  to  of  on  snowshoe lactation.  female  weight  most  that  than  feeding  behaviour  Davies,  are  a  This  and  seen  the  Keith  numbers,  spacing  be  other  and  weeks. is  can  i.e.  age,  winter.  and  and  Maaskamp  Juveniles  have  population  stress  1974).  care  survival  hare  and  behavioural  Myers,  amounts  return  night,  last  by  (1971)  not  days. its  nursing  parental  juveniles  determined  f a l l  terminate  and  it  weaned  28  71  often  discussion,  nursing  juvenile  determining  the  be  27th  nursing  are  did  not  JUVENILES  l i t t l e  Juveniles  was  she  merely  Tester  female,  least  Broekhuizen  above  provided  one,  at  by  and  the  Another  1942).  site.  INTRODUCTION  after  for  l i t t e r  indicates  l i t t e r s  b i r t h / n u r s i n g  a  Rongstad  these  hares  3.  rate  site  European  the  site  previous  and  wean  which  l i t t e r .  nursing  the  to  By  hare  her  could  location.  nursing  (Severaid,  female  hares  female  weaned  returning  females  nursing  apparent  therefore  the  the  on  the  but  in  disregards regulating Krebs of  and this  101  discussion  towards  a p p l i c a b i l i t y  to  Boutin weighing with  Boonstra,  Dhondt t i t ,  f a l l .  in  and  great  t i t  study  the  mechanism  i t s e l f .  laboratory  habitat  found  the  juveniles. spacing the  resident residents) juvenile juveniles  I  and  attempted  moving hares. into hares would  into I  did a  Peromyscus  to a this  large  the  mimic new by pen  involved  both  of  I  predicted in,  and  the  vole  of  adult  and  at  s  new and  introduced  mechanism a  1  the  into  of  mechanism of  a  already  in  weaned  contained  juveniles  contained  lose,  The  deermice,  the  the  the  S a d l e i r  the  such  in  during  while  wild  already  of  survival  effects  situation which  a l . ,  males  adult  of  et  by  mechanism,  only  1978b;  apparently  sexes  towards  introducing which  a l .  juveniles  effects  area  et  r e s t r i c t  maniculatus.  resident  the  to  behaviour  apparent  the  overlap  recruitment  recruitment,  examined  (residents). be  the  avoid  are  juvenile  introduced  not  hares  (Redfield  of  low  effect  I  snowshoe to  sexes  oregoni  aggressive  study  behaviour,  f i e l d .  juvenile  this  their  (Redfield  juvenile  contained  were  and  shown  aggressive  juvenile  which  been  adults  examined  Sadleir  adults In  M.  cause  the  areas  both  affect  season  both  new  have  vole,  suggested  on  at  into  of.  deermouse,  behaviour looked  adults  studies  juvenile  townsendii,  suggested  mating  period  M.  major,  (1965)  this  move  mechanisms  hares.  that  females  another  Parus  the  spacing  vole,  (1970)  Sadleir  during  500g  while  in  snowshoe  Adult  a  1978),  responsible  great  than  in  behavioural  suggested  females.  recruitment  1978a).  juvenile  (1979)  less  adult  these  adult  (nonand  non-resident more  agonistic  J  1 02  interactions, There that  of  were  the  juvenile  than  hares,  p a r t i a l l y  in  grid  (the  non-removal  animals  on  rates  (lO)were  same  the  a  new  their of  that  f i r s t  into  those  a  non-dispersers, facing  method  test  Interactions occurred of-way  along  involved Both  over  resident  introduced involved introduced  be  as  and  to  adults  and  more  juveniles  It  a  the  a  total  run  been  The  those hourly  non-removal This  of  was  resident  hares  analysis  were  seems  by  trapped  c l a s s i f i e d  juveniles  situation than  had  p>.05).  manner.  on  I  reacting  l i k e l y  that  dispersers more  like  hare.  I  than that  found  no  assumption. introduced  food,  A l l  lost,  vs  c h a r a c t e r i s t i c  their  between  juveniles, much  more  second  access  U Test;  similar  which  (10)  area.  being  dispersers.  hares  t y p i c a l  an  trapped  (1971)  non-removal  disperser  pathways.  in,  a  as  were  dividing  experiment  Krebs  1)  dispersing  into  by  been  those  grid  removal  in  a  the  and  of  juveniles  had an  and  (Mann-Whitney  would  normally  of  experiment:  t y p i c a l  recruited  which  area)  and  this  assumption  Myers  of  in  resident  part  removal  environment  to  was  the  were  already  the  removal  response  the  had  in  interaction  to  that  areas.  trapped  indicates  2)  grid  researchers  i m p l i c i t  juveniles  which  hares  juveniles.  assumptions  and  tested  introduced  other  two  introduced  juveniles  removal  resident  30  water, of  the  agonistic resident although  than  were  interacted  and  resident  resting  spots,  introduced  interactions juveniles  resident  the  resident  at  a  and  higher  r i g h t -  juveniles with  interacted  the  hares  residents. with  adults juveniles. rate  were  than  the were The did  103  resident  juveniles,  resident  and  juveniles,  and  also  thus  lost  more  f u l f i l l e d  my  interactions  than  did  predictions  for  the  experiment.  4.  OBSERVATION  I  was  OF WILD  unable  JUVENILES  to  replicate  observed  wild,  but  I  found  the  juvenileradult  agonistic  those  I  had  in  interactions  involving I  of  did  know  not  the  than  pens.  I  hares  status  in  the  in  of  in  many  and  similar  70  to  agonistic  62.5  hours  encountered  non-dispersers  r e s i d e n t i a l  were  recorded  dispersers  did  experiment  wild" ^population,  snowshoe  state  pen  interactions  the  juvenile  cannot  aggression  above  unmanipulated  observed  observation. levels  an  the  of  higher  the  wild,  of  the  as  I  hares  involved. Adult rabbits  aggression,  has  been  towards  recorded  in  non-resident  small,  indoor  juvenile  pens  European  (Mykytowycz  and  o  Dudzinski, 1960;  1972),  Lockley,  Mykytowycz and  and  Gambale  were  wild  in  large,  1961), Gambale  (1965)  and  outdoor in  (1965).  also  show  wild  that  dispersing,  The'  from  pen  hares  suggest  already  populated  rates  agonistic  of  already aggression  become could  that with  juveniles  resident  interaction,  than  established  in  lead  to  further  kittens  and  hares are the  the  moving are  (juveniles) kittens.  observations into  faced  juveniles area.  dispersion  by  1948;  and Mykytowycz  non-resident  experiment,  weaned  (Mykytowycz, (Southern  (1960)  resident  towards my  populations  Mykytowycz  aggressive data  enclosures  This the  an  with which  of  area higher have  increased non-resident  1 04  juvenile,  or  either  which  of  F i e l d effective  to  increased  stress  may  its  lower  experiments the  in.snowshoe  form  adding  of  populations,  as  recruitment  to  Redfield  et  chances  should  discussed  recruitment  aggression hares.  Boonstra  this  affect  the  and  of  adult  female.  only  once  offspring. hours the  female. just  born  the  the  The  introduced The  care 24  does pens  I  l i m i t i n g  how  juvenile  could  take  the  and  control  monitoring  natural  populations,  and  not  and  to  as  in  17.1g  f i r s t  care  the  to  the  per  day  is  some  is  upon  four  for  of  the  breeding  which  had  been  season  a r r i v a l  is  that  The  f i r s t  of  site  in  60  in the  occurs  guarding  site.  the  to  which  passive  nursing  weeks  limited  the  may  Through  b i r t h / n u r s i n g  immediately  to  which  hares.  lactation,  possibly  weaning  return  the  Parental  suckle that  l i t t e r  studied  at  mechanisms  snowshoe  only  gather  gained  f i r s t  her 1-2 of the  leverets days  smaller  of than  l i t t e r s .  juveniles  introduced  by  control  in  hare.  hours,  suggested  into  determine  manipulated  behavioural  extends  leverets  following Weaned  snowshoe  sunset, I  examine  survival  Her  after  in  a  The  on  to  SUMMARY  experiments,  every  female  l i f e .  I  juvenile  l i f e  be  or  1964),  s u r v i v a l .  experiments  to  and  Davies,  (1978a).  section  observations  may  These  manipulated  and  performed  (1978),  E.  In  of  be  juveniles  in  a l .  (Christian  a  pen  containing  juveniles  were  caught  resident involved  in  adults in,  the and and  wild  were  juveniles. lost,  more  105  agonistic  interactions  introduced  juveniles  residents,  both  adult:juvenile unmanipulated of  the  could  that  lead  juvenile  were  adults and  wild  status  spacing  recruitment.  did  resident  involved  juveniles. in  juveniles.  from  of  hares  of  many  behaviour,  observed  and  of  the  adults might  I  30 with  similar  interactions  although  resident  A l l  interactions I  juvenile:juvenile  aggression to  and  population  residential  conclude  than  in  was  unaware  p a r t i c i p a n t s . and  an  I  juveniles  therefore  affect  106  CHAPTER The  major  aspects  of  relation  to  hypotheses  e.  Myers,  Similar  of  cuniculus1965; Myers  and I  research  and  have  Mykytowycz and  Sadleir  populations, degree  to  which  enclosures suggested  and  a l l the  1958,  1959,  1964— the  concluded  that  enclosures  were  not  reproductive  c a p a b i l i t i e s  of  maintained  3-5  peak  appears cause  at  l i k e l y females  times  that to  in  the the  terminate  between  out the  Myers'and  of  in  large  with  other  behavioural  1959; 1961).  to  the  in  my  feature  affecting  As of  this  wild  upon  example, and  hares.  the wild  densities In  Poole,  depended  For  0.  Gambale,  studies  a r t i f i c i a l l y  densities  (  Anderson, pen  fact,  as  Therefore,  Mykytowycz and  l i t t e r s ,  breeding.  (Krebs  counterparts  data.  high  been  wild,  experimentally  relevance  the  wild,  the  secretive.  wild  the  produced  in  was  population  recruitment  musculus-  the  in  in  (1960)  had  carried  relevance  extrapolated  which  been  1972;  hare  intent  populations.  1960;  Mus  My  wild  their  some  Chitty  study  and  them  Dudzinski,  females  to  differences  discussed  one  juvenile  have  and  and  important  observed  few  snowshoe  situations  d i f f i c u l t  programs  and  Schneider, (1969)  then  examine  regulation.  be  and  to  the  (1964)  nocturnal,  animals,  Myk'ytowycz  in  manipulated  found  penned  Davies  to  were  was of  population  cryptic,  hares,  study  mechanisms  Hares  enclosures.  species,  df  AND CONCLUSIONS  ecology  and  dispersal  mobile,  observed  ecology  g.  1974).  were  outdoor  this  hypothesized  regulation,  they  of  Christian  study'behavioural  and  DISCUSSION  behavioural  the  previously  I  GENERAL  objective  the  behavioural to  5:  alone Cary  the  hares  were  hares,  it  would and.  not  Keith  1 07  (1979) s t i l l  found 78%  the  during  extrapolate, from  the  by  be  the  were  the  enclosures  was  of  observations  their of  researchers  table  i l l u s t r a t e s ,  the  social  wild  populations, be  -  I as  these  access -  pens  that  of  the  or  on  most  in  2)  There  done  in  in  extrapolation  species.  my  based  on  As  behaviours, are  of  the  with  observations  hares  the  populations.  hares  social  the  to  mortality  hare  penned  because  were  wild  I  avian  pens  populations,  related  that  had  penned  wild  l i k e l y  there  in  avian  population,  the  juvenile found  hares,  found  I  rates  to  prey.  than  trimming  closely  organization  following  snowshoe  such  the  to  suggest  but  available  features  is  in  of  that  snowshoe  I  hares  wild  I  relevant  most  rates  from this and to may  appropriate.  The  which  than  on  the  suggest  relevance  wild  It  lost  was  if  mortality  juveniles  1)  l i t t e r  However,  juvenile  factors: on  f i r s t  cycle.  a  selective  various  the  to  in  factors  the  summer)  density  higher  other  not  the  of  a l l  two  for  high  the  spots  two  presents  opinion  high  of  These  16  of  least  hiding  because  the  concentrating  vegetation.  Table  at  rate  phase  (almost  end  unnaturally  forest,  decline  hares  began  fewer  pregnancy  example,  ignoring  predators of  the  for  penned  predators would  lowest  these  hares  arranged "ears  is  along  up"  food,  summary  performed  and  interactions to  a  the  down"  occurred  to  both  in  both  and  my  study:  interactions  non-violent  "chases"  among  of.  agonistic  from  right-of-way, found  results  stereotyped  gradient  "ears  water,  interactions,  a  of  and  and  penned  "hits".  between  resting and  postures  sexes  over  spots.  wild  hares,  were  108 Table  16:  Features  B e h a v i o u r a l f e a t u r e s o b s e r v e d i n t h e e n c l o s u r e s , and t h e i r a p p l i c a b i l i t y to w i l d p o p u l a t i o n s , as found in t h i s s t u d y , or o t h e r studies.  of  Penned  Hares  Applicability  to  Wild  Pop'n.  1. Stereotyped aggressive postures, i.e. "ears up" t h r o u g h t o " l e a p s " and " h i t s " .  1. Yes, a l l postures found pens were l a t e r o b s e r v e d in wild hares.  2.  2. Yes, at a r t i f i c i a l feeders and a t c o n c e n t r a t e d natural food sources.  Dominance  3. Seasonal swi t c h .  hierarchies.  dominance  3.  Yes,  as  4. C o u r t s h i p chases were f a s t , a n d i n v o l v e d many aggressive interactions.  4.  Yes,  observed  5. Dominant male d i d most of b r e e d i n g , and aggressively prevented subordinates from breeding.  5.  Not  known.  •6. Females mated w i t h t h a n one m a l e .  6.  Not  known.  more  7. Leverets gathered at n u r s i n g s i t e , and a w a i t e d lactating female. 8.  Lack  of  much p a r e n t a l  7. Yes, (1971). care.  8. but and  in  2  Rongstad  in  above.  in  and  the  wild.  Tester  N o t known f o r c e r t a i n , s u g g e s t e d by R o n g s t a d Tester (1971).  9. S i z e of f i r s t l i t t e r s were s m a l l e r than subsequent litters o f t h e same s e a s o n .  9. Yes, (1978).  Cary  1 0 . J u v e n i l e s and a d u l t s were involved in agonistic interactions.  10.  Yes,  observed  11. High j u v e n i l e mortality, m o s t l y by a v i a n predators.  11.  Mortality  12. N o n - r e s i d e n t j u v e n i l e s were h a r r a s s e d more t h a n resident j u v e n i l e s by a d u l t h a r e s .  12.  Not  known.  and  not  Kei th  in  the  wild.  as  high.  109  quite  stable  and  resulted  in  the  exhibition  of  a  dominance  hierarchy. -  male  hares  hares  were  a  seasonal  -  the  mating  Mating  oestrous  in  many  of  -  end  the  aggressive  of  Once  during hares  weaned,  wild adults  and  these  dominant  penned was  thereby  hares  male  chases chases  by  female  indicating  and  somewhat  dominance  polygyny.  several  the  males  were  continued further  males  males  were  after  involved  responsible  juveniles  for  most  adults  juveniles,  completely  were  were  obtained occurring  although  their  the  extremely  males.  This  spring  and  care  on  into  harrassed  juveniles than  the  parental  introduced  wild  by  became  by  female summer,  periods.  l i t t l e are  copulating  she  approaches throughout  were  from  oestrus  oestrous  exhibited  interactions  prevented  female's  further  These  resident  were  each  juveniles  observations of  summer,  in  dramatic  During  juveniles  juveniles. from  observed  and  the  to  aggressiveness  adult  as  males  males.  -  the  while  copulations.  dominant  except  winter  interactions.  hares  subordinate  at  the  dominance.  fast  agonistic  during  in  observations,  female.  penned  the  in  f i e l d  involved  dominant  dominant  system,  by  an  in  more  more  switch  v e r i f i e d  -  were  their  pens by  were  faced  indicating between  r e s i d e n t i a l  l a c t a t i o n .  own.  containing  both  resident  besides  resident with  more  juveniles. there  wild status  are  adults  and  aggression Some  similar  adults was  resident  and  unknown.  f i e l d types wild  110  I  concluded  mechanisms competition juveniles, f i e l d these  in  that  place, for  which  e.g.  high  breeding could  experiments mechanisms  snowshoe  are  are in  lead  hares  levels  females, to  have  of  necessary actually  to  behavioural  aggression,  harrassment  population  some  of  non-resident  regulation.  determine  c o n t r o l l i n g  extreme  how  population  However, effective size.  111  APPENDIX Some  non-interactive  Postures  and  (1932)  hare  observations  agree  a  they  branch  off  as  time  they  twigs and  The  high  as  possible  This  onto  a l l  behaviour  feeding the  with  it  would f e l l  fours was  hares  to  and  eat  observed  were  chew  the  in  in  this  and  the  ground, the  or  s t a n d i n g o n  forepaws,  then  My  additions  while  their  of  postures.  several  while  They  u n t i l  pens  and  in  a the  the  b l a c k - t a i l e d  an  sound.  s i t t i n g  Lechleitner  similar  (1958)  I  but  on  The  hares this  pose  hook  branch at  leaves  which and/or  both  the  hind  feet,  in  and  looked  on  pens  has  this  times  described and  for  I the  their  in  the  for  the  the  hind  their  forepaws  legs  (Severaid,  described  neither  observed  1958),  in  orientated  many  been  what  a r c t i c  1975).  flutter  their  also  pose  has  (Lechleitner,  also  its  posture,  observed  (Bonnyman,  up  c a l i f o r n i c u s ,  sound. I  wild.,A  hares  stood  "alert"  jackrabbit  Snowshoe  also  strange  a r c t i c u s ,  f l u t t e r i n g .  hare  as  of  towards  L.  hare:  description  grooming  with  out  phalanges.  snowshoe  ears  while  snowshoe  basic  and  hares  that  strike  settle  describe  L .  the  wild.  direction  hare,  also  their  the  in  good  but the  observed  with  obtained.  in  would  would  would  his,  describes I  a  feeding,  with  He  hind'feet.  pose,  provides  resting,  explanations. their  observed  Locomotion  Grange snowshoe  behaviours  I  author  was  a c t i v i t y  a  similar able  in  to  snowshoe  in  the  1942).  a c t i v i t y explain hares  a i r  in this  in  the  11 2  pens  and  in  beginning on  its  the  and/or  hind  then  commence  The  bout  of  forepaws,  and  appeared  to  from  the  wild.  It  termination  legs, to  flutter  groom  then used  forepaws  as  down  i n i t i a l l y  to  l i c k i n g  hairs  grooming.  This  interpretation  the  deep  snow  in  an  from  pens to  a  forest  On  several  and  movement while the  normal gait may  allow  predator not  so  do  in  This  stopped  to  spruce  and  (1932).  material  the  or  dust  perhaps drying  later  to after  observations  hares  flutter  up  fluttering  later  found  moving  their  needles  predator,  constantly  directions.  On  from  forepaws  picked  up  hares  had  one  locomotion  locomotion  (Bonnyman,  through a  was  and  quick  non-moving moving  explanation  would  moved  forepaws appears  the  stops,  to  their  eyes  this  be  s t i l t e d metres  between  s t i l t e d  something  optical  the  movement a  moving it  could  Physiologically, the  in  kangaroo-like  perceive  background,  require  about  continually to  This  move  several  unusual  1975).  i t s e l f .  would  occasion  its  of  the  orientating  with  type  after they  mid-stride  hare  i t „  a  pens  actually  the  against  by  the  hare  arcticus  behavioural  was  The.  better  the  L .  if  f l u t t e r i n g  explained  it  in  manner,  extreme,  quadrapedal of  also  forepaws,  by  and  the  sit  Grange  any  for  dislodge  frightened  various  stopping a i r .  paws  often to  dislodge  would  in  rest.  them,  when  occasions  jerky  in was  ground  to  its  with  l i t t e r .  been  ears  the  1980  attempt  the  s t i l t e d ,  on  A p r i l  dry  apparent  apparently in  in  ended  settling  hare  lick  described  often  the  The  forepaws,  was  before  associated  grooming.  its  separate  from  usually  of  i t s e l f  grooming  be  was  tools  this of  the  113  hare  include  such  detectors Grange  of  the  "local-edge-detectors". in  (1932)  types  produced in  fallen  trees  wild.  the  same  forms  summer,  in  the  location  and  hares  day  (1938)  provide  by  snowshoe  hares  winter.  Individual  rabbit.  used as  the  merely  under  the  Similar in  after  (1967)  related  Criddle  results  the  Intraspecific  closely  and  of  similar  cover  the  the  Levick  the  good  rested snow  showed  in  This  study  forms  covered were a  found  descriptions  hare.  situations  pens  has  under  bushes  or  observed  in  preference  for  day.  Communication  Chinning  Mykytowycz rabbit  as  (1975)  being  an  odiferous  system.  Mykytowycz  European  hare,  was  probably  occasions which  had  observed few  of  in  has  but  (1962) states  less  it  described  i n t r a s p e c i f i c also is  important  snowshoe  hares  to  introduced  adult  males  and  of  hares  the  in  t e r r i t o r i a l  marking  hare.  a  I  and  the  the also  wild  in  chin  gland  and  by  in  an  numerous animal  pens.  Chinning  in  juveniles.  were  the  therefore  observed  especially  into  females,  chinning  European  developed,  chinning,  been  the  describes  less  just  occasions  chinning  observed.  was A  1 1 4  Thumping  one  Thumping  in  of  hind  the  Severaid  different  fear,  the  wild  B)  Adults  often  a  new  thump  during  was  in  could  be  have  been  more  unsure  it  was  d e t a i l as  a  (c)  A  loped  by  in  the  an  Chapter  would  be  ground  d i s t i n c t i v e cannnot  with  sound.  provide  but  a  suggests  thumping  hare  released  in  it  several  from  a  trap  as  I  was  the  male  away. the  pen  pen.  observed  flee,  the  exagerrated  displacement to  but  around  in  also  a  observed  a)  hares  whether  Situation  I  moving  of  This  interpreted  i t s e l f .  as  of  behaviour,  follows:  accompanied  courtship.  described  the  while  l i t t e r  produces  anger.  thumped  s t r i k i n g  thumping  for  as  thumped  examining  quick  which  and/or  situations  in  the  describes  explanation  communicates  A  is  paws  (1942)  complete  C)  hares  in  2.  hop  the  by  wild,  Situations  behaviour.  freeze,  or  (a)  The  attempt  involved  in  and  is  and  (b)  hare to  may defend  intraspecific  communication.  V o c a l i z a t ion Grunt  or  snort  and  usually  one  hare  occasion, was an  black-  sound  occurred  when  no  This capped  of sound  was  during  another.  Grange  expression :  This  towards  struck.  Chirp  :  movement (1932)  produced an  actual  The  sound  was  and  made,  Severaid  but  by  an  aggresssive  aggressive was an  (1942)  hare  movement  by  heard,  on  also  aggressive explain  posture  grunts  as  displeasure. is  very  chickadee,  similar (Parus  to  the  sound  a t r i c a p i l l u s ) .  produced I  by  heard  the the  1 15  sound  only  dominant had  from  male  male  in  chased  a  the  about  the  heard  from  more  pen.  well  as  males. emitted  rabbit  and  the  Scream  :  Snowshoe  the by  they I  wild,  by  penned  Squeal  :  in  assumed  to  emitted  be  by  the  was  cannot  be a  S i n c l a i r this  state  the  male  reported  produce  a  female  hares  being  sound similar  cottontail  by  this in  and  a  male.  of  (1932);  being in  when  Severaid  handled the  it  in  pens,  and  completion  of  goshawk. upon (Marsden Poole,  study).  any  scream  handled  squeal  (Myers  squeal  has  the  buck  Alaska.  (Grange  flor idanus,  1942;  in  piercing  danger  attacked  S.  and  (1962)  hares  by  dominant  dominant  (1964)  a  leverets  cuniculus  the  hare  from  rabbit.  Lagomorphs  (Severaid,  loud,  courtship  scream  being  not Trapp  only  the  the  Holler  mortai  young  after  female  produce  including  produces  generally  hares  Several  0.  americanus sex  very  and  usually  while  is  7m.  penned  swamp  this  or  snowshoe  during  juveniles  copulation, 1964),  found  are  heard  male,  the  and  occurred  about  in  female  hares,  "chirp"  Marsden  is  (1942).  than  has  sound  appears  It  The  vocalization  (pers.comm.) as  pen.  subordinate  moving  similar  snowshoe  the  It  is  above  and  1961), not  Holler, and  known  species,  L . which  but  is  116  APPENDIX  II  F u l l d a t a on i n t e r a c t i o n s o b s e r v e d d u r i n g summer 1 9 7 9 ( J u l y 1 1 - A u g u s t on T a b l e 3 , p . 2 8 ' . '  i n Pen 2 4)-:Summary  Wl N N ER  L O S E R  Hare  1  2  3  To t a 1 s  1 2 3  X  i  o  1  32  X  Z  1 3>  zr  X  4-1  X •  Totals  5"!  Sex  F  Wt.(g) %Won  -  '  :  X  •  2.  H  M  — —  \o \v c  4.1-  -  117 APPENDIX  Ilia:  F u l l d a t a on i n t e r a c t i o n s o b s e r v e d i n P e n 1 d u r i n g May 1 9 8 0 - s u m m a r y on T a b l e 4 , p . 2 9 .  WINNER Hare  L O S E R  1 2  1  2 6  X  X  3 1 • •  -  10  Sex  F  F  To t a I s  Z 13 . 1* 23  X X  • • •  21  %Won  I?  -  Totals  Wt.(g)  3 H z .0. 4- o X o  .•5"<T —  M  —  IS30  o.o  —  118  APPENDIX  111b:  F u l l d a t a on i n t e r a c t i o n s o b s e r v e d i n P e n 2 d u r i n g May 1 9 8 0 - s u m m a r y on T a b l e 4 , p . 2 9 .  WINNER Hare  i_ L O S E R  1 X  2  3  0  O  O 3. 11 • IS X H 0 o X  Totals Sex  o o o  F  ^  •  X  rr  4-0 — —  Wt.(g) %Won  o  X 0  12 F  To t a 1 s  100.0 6S".0  o.o  —  .  119 APPENDIX  IVa:  F u l l d a t a on i n t e r a c t i o n s o b s e r v e d i n ' P e n 2 d u r i n g w i n t e r (November 1 3 - 1 6 , 1 9 7 9 ) , before t h e r e m o v a l o f t h e d o m i n a n t m a l e - s u m m a r y on T a b l e 5 1 , p. 3 1 .  WINNER Hare  1 L O S E R  1  5"  2  3  To t a 1 s  X  0 X  0  0  j X  0  G 25"  z 3  H  3  3  11  O X X  Totals  O  3^ F  Sex Wt.(g)  m-so  %Won  too. o  -  M.  l boo  — —  o.o  —  120 APPENDIX  IVb:  F u l l d a t a on i n t e r a c t i o n s o b s e r v e d i n ' P e n 2 d u r i n g w i n t e r (November 1 6 - 1 9 , 1 9 7 9 ) , after r e m o v a l o f d o m i n a n t m a l e - summary on T a b l e 5 b , p. 3 1 . ' .  WI N N ER Hare  5* L O S E R  5\ 2 X 3 X  Z  3  3  To t a 1 s  O  2< 21  O X X X  Totals  33-  Sex  H  V  Wt.(g)  IbOD  %Won  3S".4  0  S3  M  •—  15*4-6  —  O.O  —  121 APPENDIX  Va: ;  F u l l d a t a on i n t e r a c t i o n s o b s e r v e d i n Pe.n 1 d u r i n g w i n t e r (November 9 - 2 3 , 1 9 7 9 ) , before r e m o v a l o f d o m i n a n t m a l e - s u m m a r y on T a b l e 6 a , p .  33  W l N N ER  L O S E  Hare #  2  3  4  1  To t a 1 s  2  X  1  0  0  1  3  5  X  I  2  4  13  X  4  S  X  22.  X Totals Sex  M  13  i  Q  P  M  P  1.430-  Wt.(g) %Won  —  4,1.9  2 5 . 0 21.4  —  122 APPENDIX  Vb:  F u l l , d a t a on i n t e r a c t i o n s o b s e r v e d i n Pe.n 1 d u r i n g w i n t e r ( N o v e m b e r 2 3 t o D e c e m b e r 1, 1 9 7 9 ) a f t e r r e m o v a l o f d o m i n a n t m a l e - s u m m a r y on T a b l e 6b,p.33.  Wl N N ER  L O S E R  Hare  3  3  X  1 4 1 o X 1 11 X  To t a 1 s  1 VI 24 X  -  X Totals Sex Wt.(g) %Won  F  12  1  3£>  P  H  —  1-4  \\2-0 4-0  —  123 APPENDIX  Vc:  F u l l d a t a on i n t e r a c t i o n s o b s e r v e d i n P e n 1 d u r i n g w i n t e r (December 2 - 9 , 1 9 7 9 ) , a f t e r the a d d i t i o n o f a new m a l e - s u m m a r y on T a b l e 6 c , p . 3 4 ,  W l N N ER Hare #  Per As)  L O S E R  pen  2.rA  X  X  3  4  5" O IS" 1  0 X  0  3  X  Totals  o  o X  I 3 o  Totals Sex  3 1 4 0 0  M  M  32 —  Wt.(g)  [Vtv  lt><4o  \32-0  —•  %Won  100.0  II.1  O-O  —  124 APPENDIX  VI:  F u l l d a t a on i n t e r a c t i o n s o b s e r v e d i n t h e a l l m a l e pen b e f o r e t h e i n t r o d u c t i o n o f o e s t r o u s f e m a l e s and b e f o r e removal of d o m i n a n t male ( A p r i l 7 - 1 6 , 1 9 8 0 ) - s u m m a r y on T a b l e 1 2 , p . 6 4 .  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