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Territorial aggression among males of three syrphid species Fitzpatrick, Sheila M. 1981

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TERRITORIAL AMONG MALES OF  AGGRESSION  THREE  SYRPHID  SPECIES  by SHEILA  B.Sc.(Agr.),  University  M.  FITZPATRICK  of B r i t i s h  Columbia,  Vancouver,  A THESIS THE  SUBMITTED IN PARTIAL FULFILLMENT R E Q U I R E M E N T S FOR T H E D E G R E E OF MASTER OF S C I E N C E in T H E F A C U L T Y OF GRADUATE S T U D I E S (Department of P l a n t S c i e n c e )  We  accept  THE  ©  this thesis required  as conforming standard  U N I V E R S I T Y OF B R I T I S H November 1981  Sheila  M.  to the  COLUMBIA  Fitzpatrick,  OF  1981  1979  In p r e s e n t i n g  this thesis i n partial  f u l f i l m e n t o f the  requirements f o r an advanced degree a t the U n i v e r s i t y of B r i t i s h Columbia, I agree t h a t the L i b r a r y s h a l l make it  f r e e l y a v a i l a b l e f o r reference  and study.  I further  agree t h a t p e r m i s s i o n f o r e x t e n s i v e copying o f t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by the head o f my department o r by h i s o r h e r r e p r e s e n t a t i v e s . understood t h a t  copying o r p u b l i c a t i o n o f t h i s t h e s i s  f o r f i n a n c i a l gain  s h a l l n o t be allowed without my  permission.  Department o f The U n i v e r s i t y o f B r i t i s h 2075 Wesbrook P l a c e Vancouver, Canada V6T 1W5 Date  ^  £  Iti s  IO/&I  Columbia  written  ABSTRACT Among males and  are  territorial always  Eristalis  maintained all  fall  species  with  below  Possible  are  Males a  of  attempted  did  not  to  Differences  final  two  implications studies.  aggressive  (Fab.)  gradient  is  the  aggressive  intensity  increases  with  the  density  of  conspecifies,  if  temperatures  and  the  stage,  and  level  difference  mate to  with  the  in  ceases  or  i f sunlight  in  the  to  them  male's  of  disappears.  inherent  level  of  terms  these  and  of  behavioural  patterns energy  species,  energy the  results  females  cues.  intruders  i f , like  thresholds of  conspecifie  A  aggressively  females,  the  intruders  approach.  different each  recognized  a l l other  behaviour  sections of  visual  approached  Within  in  species  of  temperature  discussed  This  equestris  that  for  traced  strategies.  The  males.  Merodon  (Rond.)  fact  a l l three  respond  tentatively  than  tuberculatus  discussed.  male  but  is  the  combination  territorial  different  (L.)  certain critical  reasons  Eumerus  aggressive  i n s e c t age a  aggression  through  tenax  despite  three  varies  less  syrphids,  among  budgets  males  for  and  and  thesis deal with  This  and  with  are  mating  females  activity.  requirements  and  species  show finding  reserves.  the  agricultural  possibilities  for  future  i ii  TABLE  OF C O N T E N T S  ABSTRACT  i i  LIST  OF T A B L E S  v i  LIST  OF F I G U R E S  ...vii  ACKNOWLEDGEMENTS  .. v i i i  Introduction  1  Territoriality Insect  3  territoriality  Materials The  defined  6  and Methods  Narcissus  Bulb  22  F l y , Merodon  equestris  30  Importance Biology  30  and L i f e  History  30  Results  39  Characteristics Territorial Variations 1.  of a T e r r i t o r y  Behaviours  Sunlight  3. C r o w d i n g Insect  Recognition The  Lesser  Bulb  Results  Behaviour  and temperature weather e f f e c t s  by c o n s p e c i f i c s  age and stage of Intruders  F l y , Eumerus  Importance Biology  43  in Territorial  2. A d d i t i o n a l  4.  39  and L i f e  tuberculatus  Patterns  47 50 51 52 54 62 64 64  History  65 73  i v  Characteristics Territorial Variations  drone  a  73 74  in T e r r i t o r i a l of  Behaviour  Patterns  81  Intruders  fly, Eristalis  Importance,  Territory  Behaviours  Recognition The  of  Biology  83  tenax  and  Life  84 History  84  Results  89  Characteristics Territorial  of  a  91  Territorial  1 . Sunlight  and  2.  Crowding  by  3.  T e r r i t o r y type  4.  Insect  age of  Behaviour  97  conspecifics  98  and  100 stage  Intruders  in Aggression  Recognition  of  '  101 102  Implications  for  future  Literature  104  Females  Differences  for  97  104  Variation  Areas  Patterns  temperature  Discussion  Other  89  Behaviours  Variations.in  Recognition  Territory  111  Between  113  Agriculture  116  study  117  cited Solar  Species  120  Appendix  I.  elevation  Appendix  I I . Behavioural  data  131 132  A1  133  A2  1 34  A3  135  V  Bl.  . ...  ......136  B2.  137  B3.  . . . . 1 38  C1  1 39  C2. C3  ..140 . .. .  .... .141  vi  LIST  Table  I. Behaviours  Table  I I . Time  Table  I I I . R e s p o n s e s o f M.  Table  IV. Responses of E.  Table  V.  Time  of male  allocation  allocation  OF  TABLES  syrphids  by  an  M.  25  e q u e s t r i s male  equestris males.to t u b e r c u l a t u s males  by  an  E.  tenax  male  intruders to  50 ..  intruders  53 77 94  vii  LIST  Figure  1. M e r o d o n  Figure  2.  Figure  3. M.  Mating  equestris M.  4.  Map  Figure  5.  Behaviours  Figure  6.  Patrol  Figure  7.  M.  bumblebee  equestris  ...... 32 34  i n mating  37  site  41  o f t e r r i t o r i a l M.  routes  ..  pair  Stages  of the study  FIGURES  and  equestris  equestris:  .Figure  OF  o f M.  equestris  equestris  territory  size  males  ...  males  i n response  44 48  to  crowding  55  Figure  8.  Newly-emerged  Figure  9.  Eumerus  Figure  10. E .  Figure  11. M a t i n g  Figure  12.  Figure  13. B e h a v i o u r s  E.  M.  equestris  tuberculatus  tuberculatus E.  and  of  58  halictid  66  male  tuberculatus  tuberculatus  male  68 pair  71  territory  75  t e r r i t o r i a l E.  tuberculatus  males 78  Figure  14. E r i s t a l i s  Figure  15. A  Figure  16.  E.  tenax  hovering  Figure  17. E .  tenax  male  mating  tenax E.  and A p i s  tenax  on  me11ifera  pair  ..85 87 92  horizontal  territory  95  vi i i  ACKNOWLEDGEMENTS In  order  insects the me  realm with  flash He  and  to  begin  of  a  do  study  units,  site,  moral  many  of  changed  hats  from  and  h i s wife  extremely  grateful  department Elliott, their  had of  I  thank  Deborah  about  Mark  a l l o w i n g me  enjoyed  uncannily thank  and  and  guide  also  camera  much  good and  field son  and humour.  readily  assistant.  Stephen  encouragement;  to  provided  of  here,  able-bodied  and  I  assistantships am  Runeckles  Henderson Ridgway  quote  of  the  wit  grateful  to  for these  for  and  her  and  sharing  Lee  Gass  I  were  a  am  in  the  Drs.  Copeman,  positions  Peter  Cahoon,  wizardry  of  Morrison at who  was  just  Animal and  the  helped  me  her for  data,  unpublished  companionship  Institute  supportive  Peter  and  appear  Kathy,  My  who  combination  analyzing behavioural  to  the  at  particular,  doing.  them a l l .  i n e x a u s t i b l e computer  population  " l o o k i n g " at  and  support.  discussions  have  were  which  daughter  Science.  Todd  me,  whose  to  stop  support,  to  sustenance  Plant  U.B.C. w i t h  for  photographs  of  to  Wellington,  several teaching  Myers,  moral  they  financial  Margret,  had  elaborate  photographer  source  also  what  Dr.W.G.  an  and  the  continual  I  was  I  study,  "seeing"  seeing  took  He  this  of  field  the  Laura and  Ecology  Lertzman,  right  moment. what  Dave  Zittin,  graduate  Resource  see  Richards  appreciated.  Ken  to  at  helpful  work, much  plot  who  student and,  were  Finally, was  I  really  in  always I there.  1  INTRODUCTION Territorial  aggression  has  evolutionary  context,  reproductive  success  evolutionary  approach  facilitates  comparisions  and  now-familiar  Parker,  1978).  aggression continually  of  other,  with  in order  investigated  the  external  internal  of  and  territorial  aggressive I  to  of  behaviour  and  A  In  subsequent  three  (Fab.),  species  Eumerus  of  tuberculatus  and  were  third  second.  species These  hypothesis,  inherent  level  response  to  population  of  second  study  insect of  The  (e.g.,  territorial  animal  and  males  is  must  field  respond  study, to  I  changing  changes  a  and  and  arose  territorial  led  in  patterns in  male  the  have  a  The as  the  Eristalis  males  than  the males  of  my  species-specific which  such  as  such  as  varies  weather  in and  insect  combination  among  of  formation  of  age. my  entomologists  controversy he  and  aggressive  factors, from  Merodon  that  to  factors,  controversy  territoriality.  showed  aggressiveness  internal  a  flies:  (Rond.),  more  males  in external  hypothesis  observations  that  territorial  density,  always  observations  namely,  changes  field  intent  of  fluctuations  syrphid  first  My  this  observing  observations  first  aspects  territorial  (L.). Preliminary  the  the  o p t i m a l i t y arguments  tenax  of  an  theoretical  environment  by  in  animals.  interesting  territorial  factors  with  territorial  survive.  responses  deal  immediate  dynamic  studied  intensity.  observed  equestris  more  of  interest. a  been  hypotheses  fitness  equal  faced  appropriately  or  many  But  are  where  often  approaches  concerns  own who  the  i n t r u d e r s on  his  2  territory.  Some  approaches  a r e made w i t h  recognize instead and  intruders  that  that  workers  aggressive  territorial  from  a l l other sexual  Additional boundaries discussed  mating  males  toward  syrphid  in relation  disagree,  cannot believing  to distinguish  intruders my  there  females,  are truly  second  hypothesis  is  females  i s a difference  approaches.  of t h i s  study  fall  two h y p o t h e s e s .  to their  males  c a n a n d do d i s t i n g u i s h  i n t r u d e r s , and that  results  a l l such  because  Others  Therefore  and a g g r e s s i v e  that  are able  males  of the preceding  implications.  intent  a l l other  ( s e e pp.11.1-113).  that  between  postulated  at a distance.  territorial  approaches  have  outside the These  results are  p h y s i o l o g i c a l and e v o l u t i o n a r y  3  TERRITORIALITY Attempts as  the  known  (1971)  advanced  and  perhaps  by  can  as  animal  overt  more  either  to  "no  look  "an  area of  would  habitats",  occupied  animals  expected  and  by  added  aggression  or  markings.  Brown  and  Orians  (1970)  the  area  must  change  over  time;  location overt it  used the  be  or  1981)  e.g.,  the  In  situations.  i s the  behaviours  Leyhausen  are  (1968)  (1978a)  a  or  recognized  used  by  the  as  scent  to  defended  and  of  define  i t s exact  of  resident.  either  However,  chipmunks  i s not  and  but  exclusively  factor  space  by  defense,  overlap spatially  common of  such  result  territory the  spaced  maintained  criteria  be  1968)  which  are  were  although  a  through  occupation  three  as  exclusively  groups  random  i t must  i s defense most  repulsion  3)  a  augmented  of  and  Therefore,  definitions  definition  Defensive  cases,  (1939) d e f i n e d  (1975)  territories  (Leyhausen,  known,  means  fixed,  territories  such  resident.  preceding  concise  cats  maintain  temporally. by  exclusively  territoriality  less  used  2)  Kaufman  behaviour  advertisements  be  advertisement  almost  animals,  (Getty, not  may  defense  must  some  1)  by  of  numerous  or  from  that  fact,  Noble  animals  overt  territory:  of  as  more  Davies  by  a  In  area". Wilson  individual  be  kinds  f o r one".  advertisement".  "whenever  almost  definition  a l l related to  are  animals.  simplified  defended  group  or  than  suitable  territorial  cover  "any  or  defense  territories out  that,  definition  an  of  i t i s naive  territory  this  define territoriality  types  stated  yet  a  to  DEFINED  i n each  of  Noble's  comprehensive. not  unique  hypothesized  to  territorial  that  such  behaviours  4  are  an e x t e n s i o n  and  related  distance were  concept.  proximity  getting distance an  area  becomes  Huxley  elastic  disc  community  they  space,  (1934) which  outside  streams,  territory paths The  which  spines  made  and got c o l d . eventually  found  warm  a  without  i s individual attaches  itself  ridges  to  inside  to v i s i t "  distinction  between  with  a home  animal  range  exclusive  territory),  home  boundaries  by c h a n g i n g  In p r a c t i c e , usually  a  they  fixed  an a n i m a l ' s  range.  as "the area  home  o f an a n i m a l ' s  may  and Orians  territory  a n d i t s home occupy  part  (1970)  an a n i m a l  normally  or unusual  i s defended  o f two a n i m a l s  Sometimes a  b u t "a n e t w o r k  territory  i n which  range  natural  1968).  of migrations, emigrations  ranges  territorial follow  A territory Brown  a s an  shape as  boundary  (Leyhausen,  species.  I f the entire home  so.  t o t h e dynamic  and v e g e t a t i o n b o r d e r s .  and p l a c e s  varies  i s not r i g i d l y  responds  circular;  i s n o t an a r e a  wanderings".  the  they  fixed,  and r e t r e a t .  a l l o f an a n i m a l ' s  lives,  apart  an a n i m a l  i t s circumference  lines  like  although  continually  are rarely  defined  their  be c o m f o r t a b l y  halo  when  of porcupines  metaphorically described a territory  boundaries  or  the individual  a group  moved  could s t i l l  a territory  encroach  range  night,  i n and p u t ,  The moveable  neighbours  of  cold  distance,  (op. c i t . ) .  Territorial Julian  to explain  so they  repeatedly  pricked."  individual  f o r warmth. However,  uncomfortable,  a t which  to maintain  fable  "One v e r y  together  shuffling  distance  used  the following  huddled  After  of those  (i.e.,  erratic i sa  do n o t o v e r l a p . But i f a n d i t s home  range a r e  5  not  contiguous,  two  home  ranges  may  overlap.  6  INSECT In  his  review  territoriality occurs less  has  widely  of  territoriality,  "a  patchy  through  g e n e r a l , among  crustaceans insects,  and  territoriality previously The  the  the  the  however,  v e r t e b r a t e s and  arthropods,  evidence  i n the  defense  brood  nest  of  (Wynne-Edwards, areas  with  the  and  1962), by  more  Insecta  of  work to  far  especially  the  i s done  with  indicate  i s more  It  but  that,  widespread  types areas  mate-seeking of  by  larvae  males  territoriality, social  insects  ants  (Wilson,  t e r m i t e s and  mantid  territoriality (1899, of  territorial  in  in Jacobs,  (MacKinnon,  than  i s of e.g.,  like  bees  1971),  1969)  insects  was  first  1955),  who  noticed  brightly-coloured  s u b j e c t was  1952).  i s common,  that  will  or  not  of be  here.  Williamson  on  As  behaviour Other  behaviour  stated  believed.  interest.  Male  (1975)  including  accumulates  phylum  particular  dealt  Wilson  phylogenetic distribution.  insects".  territorial  foraging  TERRITORIALITY  done  behaviour  Since  then,  territoriality,  by  of  more  dragonflies.  ornithologists  birds  (Noble,  entomologists Odonata,  and  dragonflies  become  territorial  Nice,  become  by  aggressive  interested  have  i n the  the  However,  1939,  first  described  early in  the  1941, aware  subsequently  work  Moore, of  in  other  orders. Male their  emergence  resident  male  objects,  chasing  (teneral  patrols  molt)  his  intruders  site  territory, and,  to  an  after  dispersing  aquatic  approaching  infrequently,  area.  from  A  conspicuous  darting  after  7  prey.  When  copulates, (Jacobs,  met  either  i n the a i r within  Later  with  a  he g u a r d s  trying  the water  each  other  resident larger Similar  males  Alcock, Because  a  All  researchers  are  However, fact  agree  1964),  frequent  that  males  former a r e attack turn  left  height  grasp  and  to  face  and  drop  to the  but those  alone  among  and b i t e a t  in size  boundary,  much  (op. c i t . ) .  territorial 1966a,  Bick  aggressive  intruder  behaviour  and  Bick,  also  from  unimorphic  researchers attacks  on  i s modulated  (Kormondy,  that  conspecifics  male-male  occur  depends  i s primarily  by t h e o b s e r v a t i o n  in sexually  "mistakes"  t o an  recognition  and that  h a s l e d some  that  to lose  of the occupant"  distinguish aliens  which  conclude  reaction  ability  i s supported  male  The  repeatedly,  1962, P a j u n e n ,  identity, "territorial  (Pajunen, more  observed  Conspecific  usually  comparable  are usually  resident's  discriminatory  size  Both  i t (Pajunen,  1979).  the  apparently  from  of a head-on  other  territory  (Johnson,  latter's  which  each  Aliens  has been  the  fact  to a  the resident  behaviour  away  she o v i p o s i t s .  conspecifics  1964).  possible.  a r e chased  Zygopteran 1975,  to force  her and  his territory  a n d t h e two c i r c l e  (Pajunen,  when  than  or near  consisting  Intruding  resident,  into  he g r a s p s  different.treatment.  display  contact.  apparently  female,  her while  receive  threat  attacking  a  o r s e t t l e s on t h e g r o u n d  and.aliens  physical the  encounters  1955),  1966b). males  a male  by  1961).  visual,  a  males on t h e b a s i s  attempted  of  copulations  species  (Moore,  1952).  i n dimorphic  species  , a  (e.g.,  a r e always  Kormondy, sexually  1961) t o motivated.  8  There just  appear colour  instead type may  of  1979),  the and  flight  visible  pattern to  population  population became  to  very  1955,  density  aggression. or  rainy  continue  to  f l y during  bad  but  work  studies  odonate 1972).  that  (op.  on  tenacity  site  1960,  a  point,  flying  tolerant  periods  which  are  (Pajunen,  frequency)  social such  as  territorial cool, not  cloudy  aggressive,  1966b). after  1952). lepidopteran  "female  c i t . ) and  tenacity  and  increases  territoriality,  l e p i d o p t e r a n males,  defend  1964).  males  factors  during  and.  increasing  attachment  females,  on  site  site  influence  of  b),  Pajunen,  with  a  Pajunen,  1966a,  Calyopterix  also  intruder,  1961,  of  mutually  stop  mating  done  of  Environmental  (Moore,  been  indicate  Baker  males  (and  counterparts, Both  to  formed  species  increases with  their  cloudy  weather has  lost  although  periods  Less  and,  cues  1964).  Pajunen,  Ito,  male-  (Silberglied,  Kormondy,  1952,  1952,  1966a).  Territorial  intensity  of  1960,  precipitation  Aggressive  and  a  important  (Pajunen,  populations  and  (Pajunen,  intervals,  Other  than  encounter,  between  ultra-violet  sex  intensity  males  and  male.  (Moore,  "normal"  expected,  form  with  (Moore,  tendencies  temperature  he  Ito,  residence)  dense,  aggregations  a  i n t r u d e r s depends  d e n s i t y . When  aggressive  air  of  observed  i n the  body  varying  factors  length  in recognition  attack  only  resident's aggressive  (i.e.,  (i960)  and  (Jacobs,  environmental A  Ito  involved  normally-marked  reaction  resident  1964),  a  addition  resident's  more- f a c t o r s  aggressive  markings  In  the  be  pattern;  female be  to  search  Wellington  like  areas" (1974a)  their (Baker, found  that  9  nymphalid  males  definite bounded  establish  territories  topographical characteristics, by v e r t i c a l  edges.  territory  may v a r y  remain  constant.  Wellington  defending  t h e same a r e a  hypothesized  that  from  areas  e.g., s u n l i t  The g e o g r a p h i c a l  male's  related  in sunlit  (1974a)  observed  onset  t o the disappearance  1974) o r male  day, and  of t e r r i t o r i a l  of polarized  of a  a nymphalid  f o r t h e same p e r i o d e a c h  the d a i l y  areas  location  day t o day ( S c o t t ,  with  behaviour  light  was  needed f o r  navigat ion. Resident most  l e p i d o p t e r a n males  intruders,  Davies,  1978b),  (Wellington,  including drifting  1974a).  f o r 2-3 h b e f o r e  between  conspecific except  that  tries  to achieve  above  the other Territorial  related three and  leaves,  bees  the s p i r a l l i n g  (Baker,  Alpha  among  were  presence  alphas  and betas  which  territories.  not r e a d i l y  odonate  as each  slightly  behind  male  and  between when  by  beta  betas  t o both  males  a t low d e n s i t i e s  alpha,  males,  tolerated  not permit  i s often  1973) r e c o g n i z e d  submissive  The d i s t i n c t i o n  apparent  males  males:  the top-ranking  was o f t e n  would  (1972,  cockroach  a n d gammas  A gamma's  their  between  height  orthopteran  h i e r a r c h y . Ewing  to alphas,  gain  Encounters  1972).  behaviour  males  people  place.  to those  pair  1974,  a r e approached and  a position  c a t e g o r i e s of t e r r i t o r i a l  gamma.  and even  are similar  toward  1972, H e n d r i c k s ,  c o p u l a t i o n takes  males  betas.  was  (Pyle,  and m a i n t a i n  to a social  subordinate  on  birds  territorially  C o n s p e c i f i c females  courted  males,  behave  alphas and  territorial  of t h e i r the three there  own  rank  ranks  was a  10  surfeit  of  increased  territories, or  were  common,  lost  often Male  male on  and  died  young of  i s dominant  high  of  encounters (op.  increases  Whitford, other  defend  1979)  Males that to  lack  mate  emit  a  or  for  male  insects  females  at  Males  males  are  encounter  sites  the  But  same. genera,  bee  Alcock  et  D e t a i l e d work and  some  wasp  and  families, has  families:  been  battles is the  homosexual  crickets 1975,  Schowalter  females  and  to  also  although  . are  now  and  among  where  cit.).  allow  listed  Not  other 20  on  Megachilidae  sites go  of  a l l  only  site  may  female-  chosen males  species males  to  of  are  species (Raw,  best  1980).  at  kind  their  i n which done  or  females  latter  patrol  the  Ginsberg,  flowers,  the  (op.  Each  towards  grasshoppers  sites,  a l . (1978) 6  always  encounters  of  attract  male  s t a t i o n e d at  non-aggressively  representing  territorial. several  to  (Eickwort  -  and  lose  gradually  Joern,  bees,  nest  territorial  to  Male  female-attracting resource, rest.  apt  frequency  apparent  particularly  males  hierarchy.  conspecifics,  f e m a l e - a t t r a c t i n g pheromones  seeking  12  stridulate  not  but  and  intruders. Territorial  search  social  1961).  (Otte  against  repeatedly  aggressive  the  (Alexander,  territorial  a  lessens  whereas  h i e r a r c h i e s are  of  of  but  densities  1967).  territory  males  when  B a t t l e s between  fought  I n t e n s i t y of  both  Hymenopterans, studied  which  part  own  grasshoppers  territories  dominance  his  obvious  present.  (Ewing,  also  hierarchy,  c i t . ) and  repel  on  top-ranking the  were  stress are  became  males  territories.  among  bottom  females  crickets  others'  and  when  but  from  1975a,  do  bees  in  .11  Jaycox,  1967, Pechuman,  Severinghaus 1963),  and  e t a l . , 1981),  Anthophoridae  (Barrows,  1976),  Colville,  Pteromalidae  Sphecidae  1 9 8 0 , Gwynne  communally  when  territorial,  shape  (see Eickwort  males  males  will  noted  approach  objects  d i s c r i m i n a t i o n appears  are not  by  that,  using when  and pounce  Female  from  (Fox),  a c t i v i t y but,  of appropriate  1980).  in recognition  S c e l i o n i d a e and  territories  have  Colville  gloriosa  territorial  workers  1975,  males  Protoxea  their  Linsley,  1978), H a l i c t i d a e  1980),  Territorial  and Ginsberg,  important  i s some  . Fujisaki  cherry  evidence  (1980)  bug, A n t h o c o r i s  another"  where  females  stridulation  and  and 0 ' N e i l l ,  and  1978,  on a  size  visual  a distance,  t o be m o s t l y  wide  and  cues are  whereas  olfactory  ci t. ) . There  part  1963, A l c o c k ,  of i n s e c t s and other  close-range  one  (Lin,  a morning's  f o r females,  apparently  (op.  Wainwright,  F o r example,  Several  (Cazier  1975b,  a l l maintain  defense.  searching variety  after  1977, W a i n w r i g h t ,  Andrenidae  1961).  aggressive.  rest  physical  (Raw,  (Wilson,  continually  1967, E i c k w o r t ,  t o occupy tend  for territoriality  reported sordidus middle  spacing  individuals, females.  receptive  females  may  stems  (1973)  which  t o induce  plants  noted  includes  nudging  factor in  a precopulatory (Gerridae)  " v i e with  (Corixidae) i s  be a d e t e r m i n i n g  G e r r i s males  hemipteran  winter  of host  Cenocorixa  regime  and i s a l s o  and perhaps  of the  (Coreidae),  Jannson  i n t h e genus  Stridulation  receptive  Thunberg  to congregate.  by m a l e s  males  or lower  of an a g o n i s t i c b e h a v i o u r chasing.  that  among  signal  are able  oviposition  to  to attract  using  high  12  frequency  surface  specific spacing  s i g n a l s may behaviour  There the  she  also  a part  instance  male  guard  mates  these and  of t e r r i t o r i a l i t y  of the  quiescent  with  sex-  1978).  1980). Males  (Mulsant),  1979);  in territorial  comm.,  one r e p o r t e d  and each  (Wilcox,  in  coccinellid, female  h i s guarded  pupae  prior:,  female  after  emerges.  Male  territoriality  dipterans: Tephritidae  females,  they  Scatophaga  neither  "floating"  territory,  territorial  (Parker,  Gastrophilus  hover  after  falling  by a  to  as long  hover  between wind, During  (Catts,  3-4  coursing  1979). above  begin  weather,  the  o f some  other  for receptive  not  classified  the p a i r  defend  the ground  may  spiral  males  may  males  as high  as 3 m  i f female,  flies  i s not a before  contact  is  Males  continue  height  is  15-20 km/h.  They  i n a h o r i z o n t a l a r c o f <1 hover  most  the resident  at hovering of  (horses)  and pursue  on t h e g r o u n d .  i n winds  and f o r t h  sites,  i t . I f the intruder  and s e p a r a t i n g ;  even  host-sites  At h i l l t o p  as the temperature  back  f a m i l i e s of  l o c a t i o n nor a  t h e same way:  min c o p u l a t i o n  19 a n d 3 4 ° C ,  warm  20 cm  males  and c o n t a c t s  t o the ground  followed  a particular  and a r e t h e r e f o r e  A l lp u r s u i t s  female,  males  s t e r c o r a r i a ) search  defend  at approximately  conspecific  Although  intestinalis  the intruder  four  1970a,b).  at hilltops  intruders.  among  the Drosophilidae,  and the Syrphidae.  (e.g.,  sites  i s recognized  the Gastrophilidae,  families  or  play  (Richards,  galbula  emergence  communication  (Rollo, pers.  i s only  Coleoptera  Leptothea to  wave  face the m.  f o r 30 m i n a t a t i m e , b u t  13  when  temperatures  longer  periods  Most acting sex  are cool  than  and r e c e p t i v i t y  female male  any  continues  at oviposition sites, of d r o s o p h i l i d s  of  Hawaiian  females may  oviposit  their  there  surrounding  vegetation,  soon  feeding frond  spend spend  sites.  much  Each  pinna,  a  trunk.  The  male  type  of a  space.  less  gently size  female,  away  as a  the  without  1968).  both  no  But  males  males and  though  females  aggressive and f l y  become  feeding  highly  trunks, a  fern  frond  to intolerant  species  (e.g.,  D.  time  and  away  from  surfaces,  accumulate  at  a single  leaf,  or a p o r t i o n  by m a l e s  for light,  and o v i p o s i t i o n s i t e s .  females,  leaf  area:  stem  chosen  are also  crucigera,  on  and o f t e n  limited  than  of t h e i r  sitting  feeding  D.  by  definitely  the s i t e  they  proportion  preferences  Territories  site,  show  leave  time  of t e r r i t o r y  species-specific  are  of  proximity.  defends  section  turns  to feeding;  males  a r e found  s t i p e s and t r e e  sites.  species  however,  a great They  for  ascertains the  t h e same  (Spieth,  feeding  as males  individuals in close  therefore  and  At a  as w e l l ,  As  Males  behaviour  activities  tendencies.  of  at the s i t e  i f n o t , he  Drosophila  (op. c i t . ) .  restrict  a male  i s approximately  h i s courtship;  any a g g r e s s i v e  territorial  s i t  Instead  If the i n d i v i d u a l i s a v i r g i n  displaying several  are not t e r r i t o r i a l .  i n d i v i d u a l that  Drosophila.  a r e o l d , they  hover.  d r o s o p h i l i d males  aggressively  tapping  they  or males  these  grimshawi  males a n d D.  fern  tree  on  temperature  invariably established Territorial  a  of a  depends  humidity,  fern  of  near  some  engyochracea)  14  advertise over so. a  the  their  bubble  assume  of  assumption  a  fluid  dragging  of  this  physical  a  ritualized  from  non-receptive  with  by  substrate, depositing  Others  and  presence  their  females:  contact,  thin  of  i f not,  will  the  at  their  and  Often  repel  as  they  the  mere  owner  with  do  retracting  intruding  territory  intruders  abdomen  liquid  exuding  papillae.  stance  slashing  t i p of  film  posture,  anal  aggressive  the  males,  responds  h i s wings  and  legs. Because of  courtship  f e e d i n g and  females  to  behaviour True  oviposition,  their has  evolved  females  suggested  that  "The  of  tasks  physical  pattern  of  Males  to  the of  some  territories.  Males  R.  of  walnut  indifferens,  defend  in  female  the  territories  of  tephritids  others maintain at husk  least  cherry  (op.  cit.).  first  i f the  1934;  A  He  species. with  the  most  her  kinds  primitive  pertained."  species:  type.  the  proceed  defend  fruit  males  and  without  floating,  f l y (Boyce,  western the  two  reason.  mature  to  to  pertain  lek  strikingly  sexually  ovipositing,  would  s p e c i e s of  whereas  the  promiscuous  male-lunging-onto-the-female  territories,  completa,  that  for this  advantage  the  sites  postulated that  engaging  i s an  f e e d i n g and  harassment  of  from  receptive  f e e d i n g grounds"  pattern allows of  attract  Drosophila  attract  communal  separated  (1968)  number  p r o x i m i t y and  lek behaviour  drosophilid  important  a  which  must  Spieth  in Hawaiian  displays  receptive  spatially  males  "involves  in close  hypertrophied  are  territories.  lek behaviour  assembling  sites  fixed-space "personal  space"  Rhagoletis in Biggs,  1972)  f l y (AliNiazee,  territorial  male  and 1974)  walnut  15  husk  f l y "...walks  outward, a  moving  strutting  spryly  them  peacock"  male  f l y approaches,  both  flies  venter.  by  approach  Unless  (Boyce, either  one  flies  away. T e r r i t o r i a l  with  jerks  1934; the  other,  flies  until  towards  sudden  each  both  about,  fall  males  a  after  in Biggs,  rear  and  up,  o f f the in a of  nut,  the  upward  If  charges  similar  cherry  pattern  or  venter  to  continues  defeated position, the  of  another him  grapple combat  and  fashion  1972).  male  behaviour  follows  much  extended  resident  i s dislodged, crouches  other  wings  fruit  or  fly  (AliNiazee,  1974). The females  behaviour  is different  (op.  cit.).  from  a  male;  position female.  a  both not  R.  was  was  those  only  territorial  behaviour  conditions:  one  male  himself  into  jump  onto  i s observed  when  a  (op.  however  that  resident  non-aggressive  cit.).  were  often  seen  fights  between  between  males.  However,  i n the  i t normally  a the  aggressive behaviour  They  her  behave  although  observed  g r e a t e r than  males,  distinguishes  not  indicating  females.  other  intruding  mating  for females  females,  v i g o r o u s as  aggression density  and  does  quick  c i t . ) observed  indifferens  males as  (op.  a  male,  to  manoeuvers  behaviour  intruding mistaken  but  f l y towards  behaviour  resident  facilitate  AliNiazee young  female's  will  an be  a  females  this  fruit  h i s response  territorial  Sometimes  may  than  cherry  toward  that  approaches  a male  Apparently,  aggressively  males  of  laboratory, i s i n the  also  subsided  under  would  tolerate  another  in  fighting  females  were  female where  field.  insect Male  laboratory on  the  same c h e r r y  16  without  initiating  occurred  only  indicating became  so  space  when  that  1972).  Males'  charging,  got  very  close  densities,  to  Aggressive each  territorial  that  males  defended  the  apple  maggot,  and  individuals.  territory,  pawing  Female  of  a  (apparently manner, male.  Prokopy  only  displays  other, boundaries  small,  personal-  reacting objects  as  Female  unable  to  tactile  reaction  and  Bush  wings  at  primarily  (1973)  observed  either  flew  away,  reacted  or  reacted  i n an  the  wings  of  male), and  moving  behaviour and  Bush  to  insect  frass,  aggression-releasing  have  probably except  toward  the  dung,  sexes  inanimate  mechanism  the  (op.  objects  similar  to  that  inanimate of  an  males  are  stage  of  non-receptive cit.).  may  the  encounters  calyx  distinguish  behaviour  towards  females  because  until  the  aggressive  to  the  on  passively  to  even  or  evolved  cannot by  or  engaged  male  charges  restricted  females,  not  a  c i t . ) observed  bird  between  females  not  (op.  other  a g g r e s s i o n may  females  was  with  that  females  in short  behave  associated  encountered  she  also  other  female  c o n t a c t , and  of  their  a  (Biggs,  Females  If a  distinguish  receptive  waving  legs.  only  wing-waving,  activity.  aggressively  apple.  prothoracic  of  among  Prokopy  such  consist  one  common  ignoring  males;  spatially-fixed  most  fruit,  type  apparently defends  was  u s i n g her  This  a  a g g r e s s i o n seems  behaviour  oviposition  surface  with  c h a r g i n g and  non-receptivity. aggressive  not  aggressive displays  aggressively,  from  high  pomonella,  personal-space  with  males  at  reduced  aggressive encounter.  territories. R.  in  an  be  The  an  observed  in  17  territorial which  males  respond There  and  Bush  orders  aggressively  i s some  recognition  of other  may  to drifting  evidence  also  be  that,  chemical.on  that  sexes  which,  upon  tactile  contact  d e p o s i t e d by  females,  When  this  substance  aggressiveness  when  d e p o s i t e d by m a l e s ,  during  copulation  aggregation Prokopy  patrolling  away  i n Maier  piplus from  territorial  males  alone  hover  (Blickle,  [e.g.,  stimulate the  by S m i t h  and  Pantle,  1975) ] .  i t i s more are only Collett  and Waldbauer, chasing  behaviour.  hovering  a  widespread few  and  and Waldbauer  pursue spot  (Collett  and Waldbauer,  papers  Land  (1979).  1979)  observed  conspecific that  Station-keeping -  i n the Syrphinae  i n Maier  recently  f l o w e r s , and h y p o t h e s i z e d  Eristalis  1959)] and  1924,  there  (L.) males  or i n groups,  to the o r i g i n a l  Milesiinae  To d a t e ,  (1976) and M a i e r  constituted  and  also  has only  in syrphids, including  (1944,  Syritta  insects  (Gruhl,  may  males;  receptivity  observed  of e n t o m o l o g i s t s ,  believed.  Wellington  Parmenter  return  (leks)  territoriality  the attention  territoriality  other  The c h e m i c a l  of males  syrphid  previously  (1975),  i n c r e a s e female  in  (1980).  attracted  on  attempts.  of groups  Although  than  i t may  Prokopy  d e p o s i t an  as a m a l e - a r r e s t i n g substance. stimulate sexual  sex  cues.  acts  may  butterflies)  pomonella,  by c h e m i c a l  both  the f r u i t  nymphalid  leaves. .  i n R.  influenced  (1972)'discovered  unidentified  (e.g.,  passersby  - has been and Land,  and  this  i n which then  observed 1975),  and  i n the  Volucella  [e.g.,  Melanostoma  1979),  Syrphus  (Heinrich  18  Wellington Eristalis 2  m  tenax  and  territories,  rests  within  feeding Males  or  of  the  the  but  bulb-fly  and  Mallota homifera  study),  morning, In  the  to  mate  apparently periods. feeding In plants,  and  the  insects  (1979)  studied  principally decora  May,  June at  wait  to  the  same  not  fly, is  also  towards  learn  to  foraging avoid  Mallota Macquart in  and  Illinois  flowers  which  grow  to  (the In  habitats  in  larger  unreceptive  proportion  of  forests.  the  forest  are  only  of  site  the  near  o v i p o s i t . Females or  (Walker),  Spilomyia  July.  larval  among  bautias  early  receptive a  territoriality  and  near  arriving  spend  and  territory.  are  bulb  flights  1-  neutral  the  daily  defend  for  their  brief  time  males. morning,  searching  decora  beyond  to  aggressive  those  males  of  cit.).  mate  males  also  so  u n i v o l t i n e and,  and  females  m  narcissus  that  (op.  continually  than  the  ( F . ) , Somi11a  feed  They  autumn  is often  syrphids,  afternoon, with  several  in males  short  flights  generation  f l y during  they  between  the  of  Loew. A l l a r e  tenax  return  equestris,  bostica  E.  generation  Waldbauer  of  and  stations  megachilids  and  species  their  longer  territories  Maier many  territory,  territorial,  honeybees  equestris. time  those  M.  territoriality  dividing their  mid-summer  territorial. highly  recognized  Merodon  grooming  territory,  S.  (1976)  each  distance;  S.  generally  more  for  patrol  hamifera  males  of  a l l four  females. flowers males  sedentary.  M. of  bautias, 10-20  patrol  Males  species  of  M.  plants  smaller a l l four  patrol  flowering  posticata, within  areas  and  species  a  and  50  m  are patrol  the  19  same  flight  often a  repeatedly. Flight  overlap although  given  moment.  proximity, usually contact. the  pattern  lasts  no  One  male,  on  males  the other  longer  than  usually  foliage  a f t e r n o o n , M.  the  forest  near  often  the pursuer,  postieata  their  respective  circular  nearly  around  wet, d e t r i t u s - f i l l e d  pockets  nearly  always  openings.  have  similarly-sized elongate leaves an  trunks  conspecific  treehole flights bursts  not t r i g g e r e d of f l i g h t  f l yabout M.  the  posticata  filtering;  Maier  by p a s s i n g  activity,  t o an  males  M.  males  seek  1-2  m  physical  i n diameter, r o t pockets defend  holes  perches which  to oviposit. which  spontaneous insects).  as at  males  detritus-filled  insects  In  females i n  males  entrance  of  or  as w e l l  These  S. d e c o r a  hover  which  intruder.  postieata  in trees.  arriving  make  chase,  feeding  species prefer  and o t h e r  and a l s o  close  habitats,  the treehole  of females  males  entrance,  drier  of both  near  way  territories,  near  Males  u n o b s t r u c t e d view  chase  and  circular  openings  openings.  or tree  mating  Males  larval  defend  separated at  not involve  decora  Individual  individuals  returns to the site  t h e same  a n d S.  outside the forest.  into  i n . a high-speed  5 s and does  react  of  spatially  come  and c o n t i n u e s p a t r o l l i n g .  the  flowers  are usually  conspecific  one p u r s u e s  encounter  resting  When  males  courses  pass  on afford  Males near  flights  Between  with  the  (i.e.,  rests  at the treehole  and  opening  territory. males  are apparently capable  and Waldbauer  pursued  insects  s m a l l e r than  passing  insects  that  were  (1979)  observed  themselves,  t h e same  size  of s t i m u l u s that  they  but pursued or l a r g e r .  S.  rarely  80% o f decora  20  males  are less  often  include  head  contact.  other  moving  apparently  mate  physical Both  (1979)  with  Males  female  syrphids,  mating  that  1970c)  females  then  holding  high-quality  to  mate  with  oviposit.  females  Males  afternoon,  that  that  patrol  females  relegated  i n r o t c a v i t i e s i n the morning  may  Waldbauer  noted  oviposition and a  Maier  that  sites  by d o i n g  distinct  oviposit  selective  although  some m a l e s  lifetime,  maximize  their  be t h e  i n order  to  i n the  t o lower  fitness, may  no m a l e s a r e  at flowers  tree  last  i n the  c a v i t y . Maier  defend  only  some d e f i n i t e l y potential  and  flowers do  strategies  also exist  pointed  f i t n e s s and  out that  in territorial  similar  male  or  both gain  advantage.  and Waldbauer  mate-seeking  mated  last  receptive,  should  when  If  and  i n the morning  i n an u n d e f e n d e d  in their  s o , may  Females  attempt  have t h e  in their  especially  oviposit  afternoon  females,  the treehole  at flowers  sites.  and  rot cavities  by M a i e r  territories  Some  oviposition  Maier  would  continually  however.  defending  mated  and ,  (op. c i t . ) .  obtained  flowers,  are not a u t o m a t i c a l l y  near  dipteran  are almost  approach  male  males  and i f , as proposed  syrphid  leaves  of e i t h e r sex.  other  by s p e r m  head-to-  the sexes.  territories  cyclorrhapous  species  males a r e  a territorial  Waldbauer, males  posticata  conspecifics  fertilized  frequently,  between  f i t n e s s than  of both  t o wind-blown  M.  high-quality  like  of eggs  react  objects.  or l i v e  a higher  (Parker,  also  reported  holding  have  majority  will  by m a l e s  and, l e s s  to discriminate  dead  should  Pursuits  contact  inanimate  unable  Waldbauer to  disciminating.  dual  bees, e.g.,  21  in  the  anthophorid  the  megachilid  and  syrphid  evolution  Centris pallida  Osmia  ecology  of  dual  rufa(L.) may  be  mate-seeking  ...  resources:  flowers.for pollen  development resource increased  Both  bees  (or nests  areas, mating  (Raw,  largely  behaviour.  males  and  success"  (Alcock,  1975a).  syrphids and  Since  search (Maier  or and  and  females wait  and in  in bee  f o r the  particularly  exploit  nectar,  1976)  "Parallels  responsible  strategies,  in bees). that  Fox  two  well-defined  sites  for  utilize  there  Waldbauer,  these  probably 1979).  larval  have  22  MATERIALS The sites:  three  species  a Vancouver  British  Columbia.  the  garden,  the  field The  weather season  Males  permitting,  included  until  three  1976).  plot  a t two  study  at the U n i v e r s i t y of were  individuals  among  present were  Observations  1980 a n d  1980. The  weather,  a n d one  males  7,  and  of t h e one  tuberculatus,  of E r i s t a l i s males from  in  found  at  field  I  observed  (and only)  Second  are not t e r r i t o r i a l of  continued,  therefore  three  but of only  tenax.  the f a l l  1981  i n June.  e q u e s t r i s , a n d o f two o r  o f Eumerus  E. tenax  o n May 25,  i n May  o f Merodon  generation  August  days  generation  generation  began  inclement  behaviour  (summer)  observed  species  tenax  by  territorial  generations  field  of a l l three  period  cut short  only  were  .  observation  was  and a  Eristalis  plot.  METHODS  of s y r p h i d s  garden  but only  AND  the  first  (autumn) (Wellington,  1980 s u p p o r t  Wellington's  conclusion. Twenty-seven E.  tenax  males  Individual  coloured  Metron  marking,  since  they  the area were  marking male  were  with  observed  a  capture (also  marked  flies  30 E . t u b e r c u l a t u s  during  from  Insects  invariably  see Singer  worked  the course  usually  of the  on  one of f i v e  were  upset  >50 study.  them  and caused 1981).  basking  t h e two b u l b  too flighty  different  not caught f o r  and Wedlake,  s a t grooming, well  and  p o s s i b l e , marked f o r  of p a i n t  pens.  as they  were  when  spot  marking  procedure  drone  equestris,  i n s e c t s were,  identification  leave  M.  them t o  Instead,  or feeding.  This  f l ys p e c i e s , but  t o be m a r k e d a n d ,  therefore,  were  individually  colour  and  wear  patterns.  solved  by  marking  established  territories  1969),  period  obligatory  place  of  movements within  one,  and  their  recorder  135  taken mm  flash  with  to  by , d i f f e r e n c e s of  marking  before  cannot  they  kinds  of  their  adult  stage  which  size, be  have  most  during  in  insects  they  with  leave  a  their  return.] two,  flies Data and  observers  moved  were  from  maps  of  place  recorded  observations  to  were  supplement  ESII  camera  six extension  body  tubes,  to  with  individual  used  recorded  a  place tape  subsequently  home  ranges  these  and  observations  in conjunction and  three  with  electronic  units.  be  did  not  unusual  legs,  of  the  I  recorded  detailed or  male  the  short  wings,  rapid  feeding,  recorder  and  of  transitions grooming,  of  occurred.  groomed  then  duration  mutually  movements  movement  which  pair,  determined  D e t a i l e d movements  with  extremely  (e.g.,  a  last  "grooming".  and  record posture  then  reported  observations  recorded.  behaviour  s)  as  Pentax  four  like  sometimes  ranges.  a  males  begin  never  Photographs  lens,  should  an  and  detailed  Preliminary  but  flies  stopwatch,  onto  territories.  since,  behaviour  a  problem  dispersal  but  home  and  transcribed  were  drone  emergence  Usually  [The  newly-emerged  (Johnson,. of  identified  exclusive wings For  behaviours,  or  legs  example,  head,  then  was  noted  simply  not  stopwatch of  the  between  basking)  behaviours  his  could  many  which  be  the  first  pair  of  as  consistently  because  of  behaviours  t h e m . Many  occurred  unless  the (e.g.,  behaviours  whether  the  1-2  a  24  resident p.39).  was  For  "on-duty"  example,  to  intruders  an  on-duty  which  male  a  or male  passed  would  a l l interlopers.  on-  or  off-duty  status  flying  insects.  When  status  of  the  model,  or  flicked  the  Thus  the  was  of  pebble,  but  a  model,  after  or  two  passes,  by  a  respond  vision,  whereas  a  the  usually not  of  of  with  On-duty  insect  a fly  vision,  and  flies  learned that  an  other  behavioural  p r e s e n t i n g him field  to  resident's  presence  tested  his  see  not  of  response.  was  also  feeding bouts  the  I  across  absence  to  on  p.25;  would of  his  present,  or  responded  I,  distinction  in question  always  range  depended  small pebbles  presence  his  interrupt  often  resident  one  within  none  (Table  feeding off-duty  always  pursue  noted  "off-duty"  and  the retreated  from i t . Soil, and  leaf  Northrup  cm  above  determined latitude  by  Cahoon  of  used and  Resource  behaviours  a  by  of  of  angle  of  the  plus  i n the  model  1956;  behaviours.  early the  at part  light  of  was at  tended to  to  occur  determine  in  the  pattern-matching by  Peter  Institute  of  1981.  user  The  sequences  behaviour,  49°N  I).  requested the  Leeds  approximately  elevation  This  l e n g t h of  specified  at  Appendix  program  a  #8692-2. " A i r  solar  behaviours  Zittin  with  polarized  of  " s t r e a k " ) was  David  during  behaviour ending  the  taken  temperature  pattern-matching  (designated  written  the  presence  kinds  sequences  Ecology  specifies  a  to  Cleminshaw,  different  I used  frequencies  The  calculating and  were  potentiometer  refers  soil.  (Alter  seqences,  program  always the  Because  a i r temperatures  temperature  temperature" 25  and  and  Animal  of lists  the  25  Table  I. M.  B e h a v i o u r s of males of the t h r e e e q u e s t r i s , E. t u b e r c u l a t u s , and E.  syrphid tenax.  species:  26  BEHAVIOUR  TYPE  On-duty (responsive to i n t r u d e r s )  DESCRIPTION  NAME  watch  male a l e r t and crouched, on s o i l or v e g e t a t i o n , w h i l e watching f o r intruders  nbask (=watch/bask)  male on-duty w h i l e b a s k i n g angles body towards sun, spreads wings, and lowers body to s u b s t r a t e  nf eed  male on-duty w h i l e f e e d i n g  (=watch/feed) ngroom (=watch/groom)  male on-duty w h i l e grooming wings, head, and/or any p a i r of l e g s  hover *  male hovers; may be done w h i l e watching, or i n s p e c t i n g an i n t r u d e r or a f l o w e r  inspect  male f l i e s toward an o b j e c t to examine i t ; i f a f l o w e r , he hovers i n f r o n t of i t , i f an a i r b o r n e i n t r u d e r , he f o l l o w s or chases i t  patrol  male f l i e s around t e r r i t o r y , f o l l o w i n g one of s e v e r a l r e g u l a r s p a t i a l patterns  chase  male pursues i n t r u d e r , f l y i n g behind and s l i g h t l y above i t  chase-contact  male chases and s t r i k e s i n t r u d e r ; i f i n t r u d e r i s female, "contact" = "clasp"; i f not, "contact" = " s t r i k e "  mate  male c o p u l a t e s w i t h female  (continued on next page)  27  BEHAVIOUR TYPE  Off-duty (not r e s p o n s i v e to intruders)  DESCRIPTION NAME  sit  male s i t s , territory  fbask  male o f f - d u t y w h i l e b a s k i n g  o f f - d u t y , anywhere on  (=sit/bask) ffeed (=sit/feed)  male o f f - d u t y w h i l e  fgroom (=sit/groom)  male o f f - d u t y w h i l e grooming  goto  male f l i e s from p o i n t to p o i n t on t e r r i t o r y , but remains o f f - d u t y w h i l e doing so  goaway  male f l i e s away from  rest  male s i t s at an o f f - d u t y away from the t e r r i t o r y  feeding  u s u a l l y observed only among _E. tenax males  ft* never observed among E. t u b e r c u l a t u s  males  territory site  28  number  of  times  distributions Appendices these this  of  preceded  the  the  one  of  evident of  requesting sequences  Such  in  whereas  the  sequences  I  of  the  indicates  the  can  of  patterns  29  of  consistent three  behaviours,  shown  in a l l present  "w").  precedes  also  be  by  that  It  "watch"  elucidated  by  the 18  behaviour out  of  Such  territorial  29  times,  the  additional  defense  behaviours. Sequences  longer  information  about  frequencies  of  sequence  length  less.  For  of  than  behaviour  sequences  example,  four the  of or  three  behaviours  patterns length five  frequency  than  three.  only  a  requesting  ("nb") p r e c e d e s times.  For  of a l l  (designated  "chase-watch" bask"  specified  information.  ("c")  see  in  chose  information  "watch"  But,  out  of  I  a  frequencies  length  11  few  additional  "chase"  "on-duty  in  consistently  sequences  can  sequence  behaviour  of  two.  for  behaviours  contained  show  one  shown  Output  information  three,  Frequency  behaviours.  However,  behaviour  This  are  sequences  plus  IIA2,  "chase-watch"  information  three  which  With  two,  data  reasons.  behaviour  length  precedes  of  showed  the  the  "watch"  sequence  length  occurs.  A l l sequences, p r e s e n t e d  distributions  sequences of  these  11C1.  emerged.  times.  sequence  specified.  length  that  29  a  I had  ending  from  and  behaviours  in Appendix  sequences  unique  following  frequency  sequences  total  have  appendices.  example,  is  IIB1,  patterns  obtained  nine  for  two  behaviour  in  IIA1,  length  each  constructed  appendices  length  I  that  yielded that  provided  Thus,  occurred  distribution  less  a  given  four of  by  times  or  sequences  of  29  length  four  informative three, of  two  three  ending because or,  most  behaviours  extraction  of  in  the  each  behaviour of  often, was  the one  117  Hence  since  concerning  was  sequences  times.  selected,  information  "watch"  less  occurred  the  sequence  i t optimized  patterns  of  four, length  the  behaviours.  30  THE  NARCISSUS  BULB  FLY,  MERODON  EQUESTRIS  Importance  Merodon Europe. 1842,  It  equestris  was  first  apparently  reported  and  in  the  (Doucette  et  a l . , 1942).  bulbs,  narcissus  of  Cyrtanthus, Leucojum, oviposit  Pacific  Scilla, on  through  the  within  for  rest  bulb  tissue  larvae  also  Broadbent,  and,  soil  to  the  a  provide  result,  entry  Hodson,  1932,  France  biology (Lyon,  of  1973),  Columbia  (Doucette  regions.  Adults  continue  emerging  emerge live 17  an  days  before  M.  et  begin  average (Hodson,  Britain  to  of  11  the  but  1932).  end  are  days, Both  been  (Hodson,  appear  It  They  et  Iris,  Adult  females larvae  remain  feed  on  the  . W o u n d s made  by  (See  a l . , 1942.)  well-studied  1932) is  and  April  of  June  or  in a l l  and  three  or  early  May,  early  July.  Males  s h o r t e r - l i v e d : males females  in  British  similar  late  males  Amaryllis,  pathogens.  in  whereas  of  History  has  a l . , 1942).  of  in  1908  hatching,  bulb  Doucette  Life  equestris  until  females  and  plant  in  pest  i t , and  period.  for or  After  the  Britain  Habranthus,  enter  destroy  sites  Biology  The  larval  also  1970).  level.  bulb,  in  Columbia)  but  (Woodville,  southern  serious  Galanthus,  soil  of. t h e i r  as  1927,  Vallota  of  1738,  is a  particular,  f o l i a g e at  tunnel  the  in  in  (British  equestris  Galtonia,  and  plant  M.  native  in France  Northwest  bulbs  Eurycles,  is a  live  females  an  are  in  and  captivity  average robust,  of  31  pubescent small  flies  bumble  several  bees  coloured  which  and o f t e n  sound  like  grey,  tawny  eyes;  as  like  may  on  (Fig.1).  or black, while  of s e v e r a l Hodson  resemble There  are  of the face  the underside  other  found  Both  Both  hind  bands  that a l l -  sexes  the eyes  flies  of each  rusty-  circumferential  female.  (Fig.2).  from  with  others are  (1932)  Diptera, only  be d i s t i n g u i s h e d  process  tawny  invariably  i n most  them  segments,  or rust.  were  i n the middle  flies  entirely  abdominal  superficially  i n c o l o u r p a t t e r n . Some  by c o m b i n a t i o n s  individuals  reddish  variations  posterior  black,  meet  long,  a r e almost  distinguished  black  12 mm  individual  individuals  of  about  have  large  o f t h e male  sexes  of  bulb  by a b r o a d , femur  tooth-  (Woodville,  1970) . Males  are t e r r i t o r i a l  territories makes with  f o r mating  contact  with  his first  mating stage  process t h e male  were  his  The  for  approximately this  with  of l e g s  s o , moves  passive  o f f by p u s h i n g  5 min. Both  of stage  insects  the male's  three  the  to  stage,  him w i t h  the mating  above  attempt  i n the first  to locate  back  at this  male  stages  During  the c o u p l i n g of the g e n i t a l i a ,  stage,although  The o n s e t  done  pair  a  from  definite study.  establish  when  her thorax  my  his posterior  i s usually  begins  Three  during  i f she i s s u c c e s s f u l ,  begins  3a).  observed  t h e male  two  during  of legs.  1976) a n d  Mating  clasping  and, having  female  t o shake  swaying;  female,  extends  genitalia  attempt  purposes.  two p a i r s  female's own.  a  (Wellington,  the  insert b u t may  her l e g s and ends.  and  Stage  continues  are quiet  and  wings  vibrate (Fig.  i s marked  by a  may  motionless  loud, high-pitched  32  Figure  1. Merodon e q u e s t r i s and bumblebee. (a) A marked M. e q u e s t r i s m a l e , s h o w i n g t h e b r o a d h i n d f e m o r a . (b) An u n i d e n t i f i e d b u m b l e b e e f o r c o m p a r i s i o n .  34  Figure  2. M a t i n g M. e q u e s t r i s p a i r . Note that the male's e y e s meet i n t h e m i d d l e o f t h e f a c e , w h e r e a s t h e female's are widely separated.  35  36  whine  emitted  by t h e m a l e .  The whine  has been  Doucette  e t a l . (1942)  and i s s i m i l a r  syrphids  (pers.  Hodson  may  be p r o d u c e d  suggested the  that  beginning  male,  which  female, three  may  minutes  mated  usually  three,  the female  the noise  that  (1965)  noise  min, ending  and feed  o f f the  o f f the  ( F i g . 3b). Stage  when  o r groom  (but u s u a l l y  after  t o shake  sideways  other  t h e sound  but Oldroyd  tries  falls  the high-pitched  and e i t h e r  of  suggested  spiracles,  and u s u a l l y  bask  to that  in pitch  by t h e h a l t e r e s . S h o r t l y  f o r 0.5-3.0  female  (Hodson,  (Doucette  the insects  for several  the male)  may  f o r an a d d i t i o n a l 0.5-1.0 m i n  e t a l . , 1942).  They  are deposited  level,  the s o i l through  there  o r more  larvae October  (Hodson,  the bulb's  may  t h e same  (Doucette  1932).  basal  one l a r v a oviposit bulb.)  long  and chalk  singly  bulb  (Hodson,  plate,  at or  the subsequent per bulb;  two  site  Third-instar larvae  e t a l . , 1942) a n d o v e r w i n t e r  hatch  down A  larva  upward, and  on r a r e  t h e same  bases.  eggs  move  1932).  tunnels  just  desiccated  well-placed larvae  her  white  or leaf  u s u a l l y become  Viable,  on  throughout  foliage  first-instar  t o complete  i s only  plant  surface  to the nearest  females  occupy  on h o s t  whereupon  i n the bulb  (Usually  often  on t h e s o i l  10-15 d a y s ,  remains  o f 40 e g g s  a r e 1.5 mm  do n o t h a t c h  enters  an a v e r a g e Eggs  deposited  through  lays  1932).  ground  within  two  continue  t o emit  lifespan  and  emitting  by  obs.).  A  below  back  afterward,  continue (pers.  i t i s produced  moves  Both  (1932)  by t h e t h o r a c i c  of stage  still  separate.  Eggs  obs.).  reported  instars. occasions, so t h a t mature  i n the  their by  bulbs.  37  Figure  3. M. e q u e s t r i s : S t a g e s i n . m a t i n g . (a) C o p u l a t i o n , s h o w i n g m a l e wirig v i b r a t i o n . The f i n a l s t a g e , showing t h e f e m a l e s h a k i n g male. Note that the male's wings a r e s t i l  S t a g e 2: (b) S t a g e o f f the l.  3:  39  Early  i n the spring,  plate  a n d move  protrude become  the pupal  during after harden  Doucette  larvae  the bulb  the s o i l  until  Pupation  takes  (Doucette  days,  undergoes  e t a l . (1942) year  occurs  the season;  development  days, Adults  emerge  to f l y  and exoskeleton  one c o m p l e t e  reported  that  overwinter  few, i n d i v i d u a l s  e t a l . (1942)  development  hardens t o  and a r e ready  t h e wings  may  postulated that  l a r v a e may  i n the drying  year  such  from  be u n a b l e  tissue  approximately  to  require three  i n larvae hatched  such  generation per  a s (immature)  l a r v a e a n d r e q u i r e an a d d i t i o n a l  Doucette  35-60  just  1932).  i n a given  A  heads  skin  e t a l . , 1942).  sunny which  from  v i a the basal  their  s u r f a c e . The l a r v a l  o n warm,  e q u e s t r i s thus  development.  in  case.  morning  (Hodson,  instar  the s o i l  1-2 h p e r i o d d u r i n g  M.  the  through  on t e m p e r a t u r e  late a  year.  upward  through  depending  the larvae leave  second-  complete years  retarded  eggs  20% o f  .  larval  deposited  late  .to c o m p l e t e  characteristic  of mature  bulbs.  Results  Characteristics  I  observed  territorial  of a  Territory  that  a l l b u t o n e o f 27 m a l e s  boundaries  or  more  of the three  62  for a discussion  restricted  their  which  flower  approximated beds  territories  to a  the boundaries  i n the study  of the exception.)  established  area.  Fourteen  single  flower  of one  (See pp.54-  males bed, whereas  10  40  males 3  included  males  beds to  maintained  33.5 m  (three  2  frequently  prolonged made  with  used  only  home  within  (However,  with  males,  defended,  territories, duty  In  coincided varied: day,  they  on J u l y  males  space  rested  territorial  17. P e a k  territories  usually  defended  his territory  days,  or reappeared  defended on s u c h  from  territories territories  their  territorial  one  territory  was o b s e r v e d  a male  males  they  range.  weather.  Several  corners).  from  at least  o f sunny  died.  remained  of t e r r i t o r i a l  o f f by a n o t h e r  them.  sites  respites  periods  h e was d r i v e n  between  s i t e s ) away  home  within  territorially  the adjacent  male  numbers  territory.  i n two ( o r  particular  a larger  site  territories  2 and 3  maintained  within  t h e most  contiguous  of t e r r i t o r i e s  during  bed  on t h e s t u d y  flying  between  visited  ( b e d 1)  2  The f l o w e r  behaved  of beds  three  never  with  until  7.0 m  were  (or three)  not while  a l l males  site  with  and  of a l l  the favoured  t o as "off-duty"  1980, t h e f i r s t  the last  two  o f t h e number  referred  one o f f - d u t y  from  territory  Males  parts  repeatedly  ( F i g . 4 ) . Thus  parts  2  such  beds,  defending  where  ranged  a n d i t became  Usually  regardless  (subsequently  included  territories,  d a y . I n 1981, p r u n i n g  i n t h e 2.7-m  also  in their  a mean o f 1 7 m .  range.  the flower  territorial  sizes  maintained  range.  males  that  of a male's  o f h i s home  of t h e beds  beds  i n . 1980 was b e d 2. I t r e c e i v e d  each  boundaries  larger  and  beds),  sunlight  those  three)  and  territory  bed 3 t h e s u n n i e s t , The  All  territories  ( F i g . 4 ) . Thus  most  a  two o f t h e t h r e e  Length  male,  of  males  for at least moved  four  7,  residence one  on, or  f o r as long after  on May  ass i x  to five  days  41  Figure  4. Map o f t h e s t u d y s i t e , s h o w i n g p o s i t i o n s o f t h e three flower beds. E x a m p l e s o f two M. equestris t e r r i t o r i e s a r e i n d i c a t e d by d a s h e d l i n e s , a n d two o f f d u t y s i t e s a r e i n d i c a t e d by a s t e r i s k s .  42  0  2m  43  absence  due  All feed  on  plants of  inclement  territories  open  as  soil,  positioned  a  1 m.  toward  intruders  on  an  Once  I  became  (p.25;  see  also  behaviours intruders  by  Fig.5)  open  perceived.  on  away  the  and  bulbous  sites),  areas  Territorial  males  vegetation  from  space,  small  analyzed  the  not  vegetative  watched  Materials  and  behaviours behavioural  "watch"  "watch-patrol-watch" sequences, and  the  most  was  see  The  were  and  as  for  the  the  I  The  most  in  were  "chase-watch-patrol"(Appendix basic  off-duty  if a  program  the bulb  respond. (see of  frequent or  patterns  "patrol". were  Within  "patrol"  "patrol-watchIIA2,  behavioural  fly  distribution  "watch"  "watch-chase-watch".  the  I  f l y would  frequent  patterns  in Table  used  frequency  ending  most  and  in  intruders,  "streak"  IIA1.  those the  no  in doubt,  resulting  frequent  indicated that  shown  involved  r e a c t i o n to a l l  whether  using  sequences,  them  were  i n Appendix  sequences  behaviours  resident's  there  to  data  Methods).  i s shown  the  d i s t i n g u i s h e d on-  status  pebble the  with  the  When  model  a  . I  observing  behavioural  analysis  or  I classified  resident's  patrol"  soil  will  1932)],  (oviposition  vegetation.  faced  familiar  defense,  Within  (Hodson,  flies  Behaviours  territorial  I  of  [bulb  (Fig.5a).  Territorial  or  plants  of  the  they  area  plants  narcissus  backdrop  There  food  garden  and  themselves  than  backdrop  irises and  weather.  contained  most . f l o w e r i n g such  higher  to  IIA3).  pattern  The in  44  Figure  5. B e h a v i o u r s o f t e r r i t o r i a l M. e q u e s t r i s males, (a) " w a t c h " : A male o n - d u t y , f a c i n g t h e open s p a c e , watching f o r intruders; (b) "nbask": A male o n - d u t y , body a n g l e d t o w a r d t h e sun; ( c ) "ngroom": A male ond u t y , g r o o m i n g ; (d) " r e s t " : A male o f f - d u t y , resting u n d e r a l e a f away f r o m h i s t e r r i t o r y .  45  46  47  territorial "patrol", cool,  defense  where  i s an a l t e r n a t i o n  "watch"  between  i s preempted  and " p a t r o l " preempted  by  by  "chase"  "watch" and  "nbask"  i f the a i r  when  intruder  an  is  is  present.  Variations  The watch,  in Territorial  basic  territorial  watch-chase-watch)  established  territorial  territorial  males  in  areas  In  such  where places,  time  was  chasing  (Table  routine  repeatedly the  same  Individual  frequency striking to  males.  Such  males  only males  less  II). Spatial  site(s)  were  rare:  established  or other  was  left  patterns: route  patterns  well-  o f t h e 27  territories  intruders  routine  defense  were  rare.  interrupted  only  the t e r r i t o r y .  Most  were  an uncrowded and remained  of t e r r i t o r i a l local  behaviour  weather,  as  less in consistent  male faithful  to  varied  with  and crowding  and the i d e n t i t i e s  individual difference in life  i n uncrowded,  (Fig.6).  patterns  of i n t r u s i o n s  (watch-patrol-  i n p a t r o l l i n g , and s t i l l  t h e same  and s u n l i g h t ,  differences  three  or temporarily  as behavioural  watch  temperature  evident  watching,  patrolled  patterns  most  observed,  mated  Patterns  were  the "watch-patrol"  the resident spent  defense  conspecific  when  and  Behaviour  stage  i n response  (q.v. P.54).  (i.e.,  of intruders was  apparently  ) . One due  48  Figure  6. P a t r o l r o u t e s of M . . e q u e s t r i s males. (a) R o u t e s of an u n c r o w d e d m a l e ; (b) o f a c r o w d e d m a l e . Hatched areas indicate plants. Note that s p a t i a l patrol p a t t e r n s b e t w e e n u n c r o w d e d a n d c r o w d e d m a l e s do n o t v a r y significantly.  49  50  Table  II.  Time  allocation  b y M.  equestris  male.  Behaviour  Time  watch  allocated  (%)  73.6  nbask nfeed ngroom patrol chase chase  17.2 5.3 contact  inspect  J_.  Sunliqht The  and  temperature  presence  territorial when  a cloud  fall  from  1.2  of f u l l  activity  than  24 t o 2 2 ° C ,  it  returned.  (During  still-available  air  t h e sun caused  territorial  t h e s u n was o b s c u r e d ,  inactive  heat,  When males  curtailed  several sites,  or ceased  inspecting,  on t h e s o i l . min, males or simply  males  critical for  level  . F o r example,  a i r temperatures stopped  inactive  flying  to  as  soon  (off-duty)  males  basked  until i nthe  or fed.)  In c o n t r a s t ,  remained  active  as  a t an  22°C.  a i rtemperatures  (patrolling, sites  of  more  periods,  groomed,  a s t h e s u n was n o t c o v e r e d , temperature  males  and remained  such  radiant  seemed  the temperature  p a s s i n g over  as  long  sunlight  i n sunny  areas  territorial chasing)  basked  their  defense  below  remained  territories  on t h e s o i l  to basking  below  for their  in their  20°C,  activities  a n d f e d o r went  I f the temperature left  fell  20°C f o r resting  territories.  When  51  air  temperatures  hovered  patrolling  and  vegetation  within  increased,  the  chases  basking  males  were  22°C;  and  the  surface 33°C. in  of and  using  on  the  above  were the  m  i f they  were  basking  of  >22°C.)  temperature  Territorial or  full  23°C.  At  such  sites  on  begonia  A on  reached  males  a  the  leaves,  basked  on-duty  between  which  Basking  as  "thermometers": were  <20°C;  day  too  reached  and  outlook  leaf  i f males  20-  the  soil,  hot;.e.g.,  27°C,  sunlit  could  were  above  become  on  i f basking  temperatures  could  the  occurred  (Air temperatures  their  insects  on  still  50-55°C,  where  or  and  0.15-0.60 m  calm  soil  flights  soil,  male  the  temperatures  shorter.  at  on  patrol  males  the  interspersed  of  soil.  shifted  shade,  times,  air  temperatures  0.15  were  basking  frequency  basking  soil,  males  As  insects  interludes  a i r temperature  dappled  ground.  although  by  20°C,  on-duty  within  temperatures when  m  0.3-0.75 m  estimated  were  0.3  duration  basking  vegetation be  chasing with  increased;  flights,  around  soil  leaves  posts  averaged  to  temperatures  leaves were  only  avoided previously-frequented  were  as  warm a s  33°C  in  the  shade.  2.  Additional  Males periods  of  d i d not rainy  approximately weather fed  to  weather  repletion  f l y or  weather.  0930  improved  effects  feed  Therefore,  (depending  after and  during  one  basked  or  on  the  evening  i n the  morning  temperature),  more  before  days  of  becoming  or  bad  or  during until  when  the  weather,  territorial.  males  52  _3.  Crowding  by  conspecifies  The  most  interesting variations  occurred  when  a  challenged  with  conspecifics. flight  also  confronted conspecific  repletion. to  an  and  alien.  Such  increased of  the  new  resident.  was  not  only  or  the  intruding  constraints  male  was  shadows),  (Table  III).  of  his  male's  his  and  kinds  struck  more  of  males  conspecific  females,  to and  normally  conspecific  was  males  i t  to  became situation  also of  of  unable  to  such  heightened patterns:  intruders  his  three  to  from  periods  thereby  intruders  of  feeding  crowded  provided  therefore contact  of  feeding  rest  behaviour  the  intruders  to  result  in  more  aggressiveness,  Territorial  changes  but  was  both  forfeited  a  Such  chased  reaction  in  The  other  resident  which  territory,  male  resident  such  effectively  resident  and  from  the  during  males,  to  interrupted  resident  aggression. by  flies  territory  interruptions.  manifested  (e.g.,  A  to  a  conducive  intruders,  a  the  more  defending  prevented  resident  increased  on-duty,  of  patterns  therefore  or  wasps,  territory  effect,  without  was  stayed  In  also  conspecific  confined  groom  aggression  the  one were  the  preventing  his  and  intrusions  outside  left  activity,  one  feed  sites  i f he  so  Repeated  thus  by  bees,  number  intrusions  off-duty  undisturbed;  active,  increased  bouts,  of  behaviour  crowded  conditions  kinds  more  was  frequency  weather  many  were  grooming  going  When  by  male  increased  activity,  insects  and  resident  in  an  than  he  kinds  (which  of  usual  pursued  accurate  "crowded  the  index  state  ".  intruders:  usually  hit  53  Table  III.  State  R e s p o n s e s o f s i x M. e q u e s t r i s m a l e s t o intruders. E a c h row o f d a t a i l l u s t r a t e s r e a c t i o n o f one m a l e .  of male  Reaction  to  chase  c r o w d e d by conspecifics and a l i e n s  13 13  c r o w d e d by aliens only  26 1 2  uncrowded  back), several may  be  times  screen  apparently  ignore whereas  to  likely  from  males  0  26 14  1 3  9 6  2  the  the t e r r i t o r y .  intruders visually or merely  u n l i k e bulb  flies.  males  will  are  aggression closeup  Crowding  also  are usually  territory). males,  Thus  wasps,  chase  repeatedly less  chasing  uncrowded  males and  h i t everything difference  apparent  t o an  the  until  the  aggressive  them  butterflies  and  and have notice  behaviour  after  but  between observer  mating.  but  people, shadows the  two  trying  photographs! affected  or  whereas  intruders that  i n s p e c t e d . The is readily  although  Uncrowded,  before  h i t one  Intruding.bees  striking,  strike  inspect  crowded  which  (which  usually  flies,  obtain  24 20  to chase without  small  of  11 7  uncrowded  shadows,  people,  levels  .  bees  or  Total  chase-contact  leave  crowded  ignore  silhouettes  and  by  of  they  aggressive  is driven  males  kinds  until  a r e more  crowded, bee  various  struck  latter  intruders  8 3  and  the  Crowded  54  males  resumed  seven  such  males  watched  and  seventh  began  chased  a  sat  for for  (nbask), mating.  an  off-duty  posts  one  (nbask),  immediately  watched  and  two  behaviour  3-5  min  had  after  also  Therefore, prefer  after  basked  provided to  remain  to  The  Of  of  (nfeed),  but  The 5 s  remained  three  observed,  watched  the  Of  three  within  males  the  third  mating.  patrol.  uncrowded  f o r most they  after  fed  (fgroom)  mating.  mating.  and  began  c o n s p e c i f i c . However, s e v e r a l min  but  apparently  sentinel  observed,  basked  male  off-duty two  their  and  5 min  basked  preceding  are  not  harassed,  males  off-duty  for  s e v e r a l min  after  mating. Spatial patterns routes  as  and  crowding did  territories resident  patterns  not  grew  began  react  to  example  i s shown  to  include  both  at  p r e v i o u s l y - n e u t r a l bed  4.  Insect  All already  but  one  the  behavioural others  3 after  or  he  same  patrol  anything,  or  sites  near  male's was  the  as  the  those  sites.  territory  repeatedly  expanded harassed  stage  male  observed  else  were  observation patterns  .Although  a  at  behavioural  3.  established territories  observations, during  and  2 and  If  off-duty  harassment where  than  retained  areas.  previous  in Fig.7,  beds  changes  males  smaller  include  An  age  fewer  increased:  defend  to  to  showed  during before  attempting  period.  radically  observations  The  1980-81 the to  on  beginning  drive  exception  different one  either  from  out  had of  the  residents  exhibited those  individual  do  of not  the usually  55  Figure  7. E x p a n s i o n o f a n M. e q u e s t r i s t e r r i t o r y i n response t o c r o w d i n g by c o n s p e c i f i c s . S o l i d l i n e shows initial Bed 2 t e r r i t o r y ; d a s h e d l i n e shows t e r r i t o r y expansion t o i n c l u d e Bed 3 i n r e s p o n s e t o h a r a s s m e n t a t o f f - d u t y sites there. Hatched areas i n d i c a t e backdrop of high vegetation i n each bed.  56  57  provide  a. s u f f i c i e n t  of  particular  this  which an  The  for  the  period  in a  bulb  from  outside  e x c e p t i o n a l male  came  obviously  newly-emerged:  his  body  dense  m  fresh  and  above.the  and  vivid  lawn,  much  seldom  f l y as  high  except  during  chases,  higher  than  within  of  behaviour  s t r o n g l y suggest  dispersal  flight  the  area  as  male's  that, while  terminated  dispersal  1 m  that,  zenith  while this  intact  the  and on  was  area  males,  The  He  shiny,  pattern  3-5  which activity,  height  and  ensuing  obligatory  f l y became  trapped  waned  site,  initiated  during  and  study  his  his  in  area.  territorial  polarization  and  manner  study  ground.  the  behaviour  life.  the  into  the  to  the  his colour  path,  restricted  behaviour  fly's  their  of  approach  for  territorial  keep  the  in behaviour  were  flew  emergence,  midday  1974a) and  and  He  and  changes  wings  (Fig.8).  2m,  after  his  unbroken,  direction  study  this  as  noteworthy  of  was  still  was  hypotheses,  sequence  transition  pile  testing  individual  i t clarified  important  base  he  in  (Wellington, gradually  territorial  behaviour. The a  gap  male  entered  between  two  disappearance 17  (Appendix  when  the  days  by  (e.g.,  I ) . [That  the a  fact  the  open  at  that  lawn  that and  His  from  the  settling  southeast  settling  was  related  disappeared  insects hornet, moment  marked stopped  i n s t e a d began  to  at  1213  to  the  overhead  familiar  through  coincided with  plane-polarized light  his  bald-faced  area  trees.  overhead  previously),  across  corner  study  plane-polarized light  supported site,  of  the  with  and  observed  flying use  the  on  the July  time  was study for  10  diagonally  landmarks  along  58  Figure  8. T h e n e w l y - e m e r g e d M. e q u e s t r i s m a l e d e s c r i b e d p.55. Note the dense unbroken p i l e , shiny intact wings, and v i v i d c o l o u r p a t t e r n .  on  60  the  edges  of  the  lawn,  (e.g.,  gaps  in the  point-to-point  travels.] I  initially  male  as  f l y because  he  more  like  a  a  bulb a  bumblebee  prominent  flew  repeatedly  area,  slow  For  upward  begin  hovering  hovering  1962;  site.  or  first  behave  long,  The  at  time  This  by  four  min,  toward by  his  mostly  chasing flights.  above  saccades, of  through  the  study  which  number  of  he  he  had  the  long,  of  dispersing  spruce of  budworm  migrating  as  Each  7 m  5-7  of  towards edges by  his  ground  and  any  end  of  to  the  the  potential  exits  from  is characteristic  feed,  other  site  spirals  interruptions in  that  rapid  study  orientation  or  groom he  insects  insects  bumblebees  of  their  1974a).  period,  a l l airborne The  back  of  (Wellington,  alight,  landmarking  at  interspersed with to  et  insects  spiral,  m,  about  (Greenbank  began  above  were  disperal  the  repeatedly  "towering"  combination  not  minutes  the  well-marked  did  few  insects  polarized light male  the  of  flights  their  their  characteristic  downward  of  first  corners  a  for  recognize  Between  gap  1964)].  high  heights  during  during  were  the  the  executed  spirals  purposeful  this 10  so  slow,  upward  territorially  encountered that  a  aid,  Although  he  kinds  as  high.  the  [e.g.,  other  bounded  halted  navigational  those  are  m  corner  min,  1962),  corners.  space  of  for  to  f l y , hovering  1.5  Wellington,  flights",  apparently  insects  10  extended  with  boundaries  open  each  that  many  site,  terminated  an  next  (Hagen,  and  drone  daisies  failed  behaved  southeast  spirals  (Southwood,  and  a  directed flight  1980)  "escape  the  the  coccinellids al.,.  of  toward  favouring  arrived.  to  patch  or  shrubbery),  during began  to  he  flying  (Wellington,  during  61  1974a), each  a l l considerably  one v i g o u r o u s l y ,  contact,  until  The  he p u r s u e d  bees  failed  a  bulb  hovering,  territorial  h e was  not begin brief  -  area.  they  until  began  then  chased  his arrival,  small  insect he  heights,  airborne  while  h e was  a l l intruders  crawled  ignored  with  longer,  a l l bees  when n o t  spirals.  feeding  15, a n d 40  the onset  toward  of  He  bouts s patrol  frequent corners.  cirrus  clouds  passed  behaviour,  t h e male  only  hovering  or f l y i n g ,  and  During feeding  flying  bees,  right  for virtually  then  more  2  1953).]  insects.  when  to within  at the s i t e :  - 25,  37, w h i l e  h e was  e . g . , bumble  which  bouts  took  by  orientation  and f l i g h t s  of s e t t l i n g  airborne  ignored  feeding  a t min  confined  23, a n d e v e n  (Wellington,  to intruders  of v i s i o n ;  f l y was  min  spirals  the onset  range  were  and c o i n c i d e d  at lesser  responded  he  which  this  bees  followed,  the zenith  following  males,  thus  encounters  o r making  spirals  Despite  comparable  chasing  [However,  hovering,  until  manoeuvers  his arrival  and fewer  flower,  visible  a s 7-10 m, b u t  heights,  22 m i n a f t e r  to feed  chases  only  than  In f a c t ,  landings,  through  at those  and  chased  of the study  as high  territorial  (1-2 s ) . Longer  respectively flights,  f l y . He  audible  t h e edge  ascended  first  heights  of the f i r s t  were  often  fly's  m of the s u r f a c e .  did  reached  him o f f , even  at greater  established  all  the bulb  boundary.  This place  than  making, r e p e a t e d  the v i c t i m  t o shake  reached  larger  feeding  38 m i n  or grooming  or crawling  wasps,  over  the f i r s t  a n d one  on a  within h i s unidentified  him. Furthermore, on t h e f l o w e r s  while  beneath  62  him, At  although  min  39,  he  was  however,  close he  behaviour  by  intruder.  Thereafter,  established seeking height very  later  to  his in  Males  of  A  perhaps  butterflies, Intruders  and  not  shadows, bees  males  involved  a  female  a  time, and  chased and  overhead  male  and  like and  flies always  a  movements. territorial flying an  occasionally  restricting  i n t r u d e r s . He remained  on  polarization  the became  what  had  returned  c l a s p i n g and attempted  make  wasps  the  were  one-way  identifies  and,  they  did  usually  entered  not  look  contacted a  females  of  other  but  their  never to  males,  i.e., try  indicate that by  chasing  and  intruder)  prelude  "mistake",  mistakes  or h i t .  i f possible, hit;  ( u s u a l l y the  with  ignored.  chased  sequences  always  mate  reverse  were  long  one  was to  identify  which  inspected,  chased  females  to  i n t r u d e r s as those  were  until  cues  c o n s p e c i f i c s . Thus  involved  contact  Such  male  ignored  birds  were  males  away.  territorial  and  screened  sound)  off. Conspecific  saw  chase  a  (Fig.5d),  behavioural  usually  audible  never  the  visually  frequent,  Several  at  photograph,  and  confrontations  contact  leaves to  to  increasingly like  for minutes  when  male-male  driven  conventional  flower  behaved  and  visual  did  like  Conspecific  daisy  their  Intruders  male  territory,  (and  perceive  afternoon.  used  intruders.  he  approach  the  a  responded  territory  Recognition  his  from  under  he  to  e x h i b i t e d more  feeding  spots  which  easy  become  male;  rest at  rising  enough  was  hit:  mating. but  I  to  drive  as  well  a  behaviour  63  as  by t h e i r  All  male-female  female and  contacts - observed  airborne)  and c o n t a c t ,  mating  occurred.  I f s h e was  she remained  conspecific other  when  couple  males  male  on t h a t  fought  intruders .fled.  territorial  and  ( a n d ..perhaps b y t h e s o u n d  was  occurred,  -  appearance  back  involved a chase after  basking spot  when  by  repeatedly  i t . This  f l y away.  caused  or feeding  a  contact  bees and  the  feeding attempted  the i n t r u d e r t o shake  landed  By c o n t r a s t ,  while  observed,  t h e male  ( i f the  when  mating.  hitting  make).  the female  contacted,  the i n t r u d e r d i d not respond, with  which  during  In a l l "mistakes"  began  they  intruder to  o f f t h e male  64  THE  LESSER  BULB  FLY,  EUMERUS  TUBERCULATUS  Importance-  "Lesser similar  species:  narcissus, hyacinth, For  bulb  and  E.  i n both  species  have the  identified E. but  and  the  similar  only  as  E.  larvae  of  [e.g.,  bulbs  are  often  each  found  stage  in  iris,  narcissus. one  is  very  individuals  1927;  here  very  in  considered  life  (Hodson,  two  found  occasionally in  are  of  usually  in narcissus,  habits  i s not a  already  so  of  instars  Presence  of  larvae  symptom  of  disease  or  pest  of  Woodville,  were  by  only the  in bulbs pest  a  of  entering can  attacked  1927)].  serious  primary  capable  subsequent  other  two  usually  to  both  1970).  positively  tuberculatus.  tuberculatus  larvae  the  and  insects discussed  i t i s nonetheless  emerged  potato,  because,  same  name g i v e n  tuberculatus,  morphology  and  the  common  striqatus(Fall.),  purposes,  since  However,  Eumerus  parsnip  practical  species,  f l y " i s the  pest  as  M.  narcissus. healthy  although  unsound  eelworm  is therefore  problems  Newly-  bulbs,  penetrate stem  equestris,  bulbs  (Hodson, usually  (Woodville,  a  1970).  65  Biology  Adults halictid are  are  bees  black  approximately  (Fig.9).  with  a  Like  have  on  (Woodville.,  than  abdomen  females,  Life  6 mm  and  three  have  History  long.  halictids,  metallic-green  pubescence,and the  and  to  eyes  that  E.  white Males  meet  closely  resemble  tuberculatus  -bronze  p a i r s of 1970).  They  tint,  adults  lack  cresent-shaped are  usually  i n the  marks  smaller  middle  of  the  face  (Fig.10). Woodville the  field  late  generations  the  former  The  flies  the  live  third  days  f o r an  one,  complete  and  spring;  e q u e s t r i s males,  the  two  enter  notable  species.  by  clasping a  do  not  E.  tuberculatus  M.  equestris  their  or  per  year,  forceful  females  females mate  never  (pers.  overwinter  to  mate  of  each  attempt  "a  to  pupate  of  a  during  1927).  obs.)  and,  a l l conspecific However,  there  interactions in  species  In  15.  less  Larvae  (pers.  with  E.  for  (Hodson,  (Fig.1l).  female,  obs.).  and  male-female  contact.  repeatedly,  third.  in bulbs  emerge  captivity,  generations  territories  sitting  In  latter  partial  in  successive  Males  females.  territorial  males  of  appear  the  never are  first  days,  attempt  Although  flying  males  13  d i f f e r e n c e s among  make a u d i b l e ,  although  than  overwinter  males  flies  e a r l y October.  of  two  pupae  M.  three  individuals  occasionally a  tuberculatus  are  that  earlier  generation  which  adult  average  E.  females  and  that  active until  three  following  like  be  usually  frequently  states  in A p r i l ,  may  approximately  partial  (1970)  initiate  tuberculatus  mating males  contrast, shake  off  Hodson  (1927)  female  once  a  male  reported  mated  as  do  that,  avoids  66  Figure  9. Eumerus t u b e r c u l a t u s and E. t u b e r c u l a t u s male. ( b ) An for comparision.  halictid. ( a ) An unidentified halictid  bee  68  Figure  10. E. t u b e r c u l a t u s male. in the m i d d l e of the f a c e .  Note  that  the eyes  meet  69  70  further  advances",  prevented. that,  E.  The t h i r d  although  length  b u t d i d n o t s a y how difference  the mating  of time  i n both,  tuberculatus pairs,  the  oviposit  (Hodson,  oval-shaped clusters within the  region  through  the s o i l  neck  reach  down  i t . Feeding  Mature  e t a l . , 1942).  are  i n groups  found  singly  which  t o burrow  down  foliage  foliage,  and then  reduce  the bulb  in size  until travel  tissue  t o , a n d may  equestris.  to a  i s long,  tunnel  M.  confused  respiratory  and stubby,  slender  and  whereas  brick-red  E. t u b e r c u l a t u s l a r v a e  of the bulb  whereas  be  T h e two a r e  i n the posterior  a t t h e neck  to  1970).  In a d d i t i o n ,  in a distinct  enter  of bulb  i t i s s h o r t , dark  mass o f t i s s u e ,  i t through  surface  l a r v a e o f M.  (Doucette  homogeneous  and enter  the s o i l  (Woodville,  equestris,  white,  across  E. t u b e r c u l a t u s , the process  found  bulb  travel  rapidly  to  foliage..Larvae hatch  They  mass  begins  are deposited in  1927).  larvae  obs.).  long,  (Hodson,  clump  emitted  are shiny,  no a t t e m p t  by d i f f e r e n c e s  i n M.  female  equestris larvae,  o r move  ever  which  soil-level  larvae are similar  distinguished  in  M.  a  make  second-instar  process:  8 mm  t h e same  equestris (pers.  Larvae  the nearest  rotting  nor female  fertilization,  The eggs,  was  are discernible in  plate).  of the bulb,  they  approximately  stages  t o M.  to the nearest  (unlike  the basal  with,  1927).  move  through  pulpy,  after  o f 5-40 a t o r n e a r  neck  the  common  and approximately  3 days,  takes  separate  contact  t h e two s p e c i e s i s  and n e i t h e r male  high-pitched noise W i t h i n .2-3 d a y s  between  process  no  further  amid  a  pulpy,  equestris larvae are  o r burrow  which  opens  at or  71  Figure  11.  Mating  E.  tuberculatus  pair.  72  73  near  the bulb's E.  brown  p u p a r i a , , 6 mm  second  plate  tuberculatus  generation  and  basal  (op. c i t . ) .  larvae long,  pupate  i n tough,  a t or near  the soil  larvae  pupate  in early July,  generation  pupate  in early  pupate  during  pear-shaped,  fall  the following July  surface.  whereas  those  grey-  Firstof the  or overwinter  i n bulbs  (op. c i t . ) .  Results  Characteristics  Territorial first 25, by  coincided  probably  which  seven  A  days,  inactivity  beds  defended  with  defended  and returned  males  tuberculatus  were  boundaries  were  in  Both  and each  territories,  territory  M.  4  m  2  f o r up t o  periods  of  equestris ,  found  i n sunny  unlike  within  only  first  averaging  i tafter  However,  defended  twice:  suggested  generation.  2  usually  (August  as  weather,  t o 8.4 m ,  Like  territories  peaked  rectangular  t o defend  (Fig.4).  males  2  from t h e  the last  April,  sunny  t h e same  due t o bad weather.  up t o t h r e e  territorial  o f warm,  0.9 m  area  and e a r l y August.  of a separate  from  resident  in late  July  approximately  in size  1 and 2  1980) u n t i l  density  late  periods  i n the study  began  Population  indicative  tuberculatus  flower  present  (May 7,  between  defended  ranged  (Fig.12).  E.  were  probably  (1970).  May a n d a g a i n  Males  E.  males  Emergence  Woodville  peaks was  Territory  day of o b s e r v a t i o n s  1980).  late  of a  patches of  the former,  t h e home  range,  one t e r r i t o r y .  not contiguous  with  which  Although  those  of the  74  home  range  male's  (which  time  was  Although territorial  might  spent  the  obviously  a  sunlight.  At  "took  "floating  E.  required  M.  equestris  in  the  season,  territory (Despite  was  observed  male  retreated  included  a  always  Males described notably  same  of  was  the of  to  a  little,  resident  moved.  a  moving  were  areas  apparent;  the  new  Such  were  but  shifts  area.  of  resident  occasionally  were  much m o r e  t e r r i t o r i a l features  on  a  smaller  scale.  the. p r e f e r r e d E.  common  were short  and  territorial  when  by  the  tuberculatus food  Territorial the  early  territory.  separated  several  above  fact,  equestris  rare;  ended E.  plant.  0.20-0.25 m  were  usually  vegetation,  M.  as  In  tuberculatus  territory.)  Narcissus  restricted  the  was  1972)  preferred  conflict  a  of  varied  boundary  to  males,  species  from  most  2  males.  the  each  near  Territorial  him"  though  backdrop  space  as  territorial conflicts  of  only  with  center  also  this,  individuals  open  the  m ),  following  reason  (Biggs,  the  males,  shifted  no  equestris  11.2  territory  resident  times,  tuberculatus  Males  a.male's  the  territory  M.  as  corresponding  territories" by  large  territory.  often  and of  other  his  maintained among  result  as  his  of  boundaries  movements  just  on  size  Sometimes  be  time.  The  smaller territories  plants, activities  and were  ground.  Behaviours  exhibited for  rare  M.  almost  equestris  among  E.  a l l on(Table  tuberculatus  and  I,  off-duty  Fig.13).  males  was  The  behaviours behaviour  "chase-contact";  75  Figure  12. An E . t u b e r c u l a t u s t e r r i t o r y , c h a s e r o u t e s d u r i n g a 5 min p e r i o d . indicate plants.  showing p a t r o l Hatched areas  and  76  77  it  was  only  "contact" male  nor  observed  was any  away". M a l e s face  the  actually alien  The  to  switched  rest,  basic  E.  males  latter inspect  an  the  and  than  h i t most  intruder,  then  Responses  of  pattern and  was was  intruders,  the  again  that  the  another  behaviour  by  turning  seldom  syrphids  equestris  settle  was  therefore  so  struck  off-duty  "Inspect"  i n M.  ever  rare  to  defense  11B1 - 3 ) .  mating,  male  other  "watch-chase",  males  and  to  on-duty  backdrop,  (Appendix  chased  from  unlike  and  tuberculatus  prelude  " c l a s p " . No  territorial  "watch-patrol" all  the  intruder. Equally  vegetative  territory  as  left  a  here.  combination  clearly  evident  more, f r e q u e n t males: former  without  to  the  discussed  was  "go  in in  whereas might  chasing  the  rise  IV.  Male  E.  tuberculatus  no.  to  intruders.  Intruders Chased  1 2 3 4 5 6  males  8 20 1 6 6 9  Inspected  1 8 1  No  response  1 1 0  0 2 0  1 0 0  to  (Table  IV).  Table  of  Total  10 31 2 7 8 9  78  Figure  13. B e h a v i o u r s of t e r r i t o r i a l E. t u b e r c u l a t u s males, (a) "watch" ( o n - d u t y ) ; (b) " n b a s k " : o n - d u t y basking., w i t h body f l a t t e n e d t o s u b s t r a t e ; ( c ) " f f e e d " : o f f - d u t y feeding.  79  80  81  Variations  in T e r r i t o r i a l  Males  showed  correspondingly  little  little  Behaviour  Patterns  behavioural  variation  tuberculatus  variability,  and  in aggressiveness.  larger  bulb  f l y , E.  males  unless  full  s u n l i g h t p r e v a i l e d and  the  were  not  Like  the  territorial  a i r temperature  was  >17°C. E. evening  tuberculatus or  morning,  during  or  territorial  E.  rainy  f o l l o w i n g one  or  more  did  a  E.  after  basking  large  determinant  not  state  tuberculatus.  that  account or  in  There  tuberculatus  are  so  males  of  off-duty  breaks.  males  were  with  never  M.  two  as  three  never  exposed  to  short  periods.  which  could  on  reasons their  their  larger  17  males,  accomodate  m. 2  Thus  of  for  this.  E.  for  and  study  during deprive  but  only  2-3  equestris  no  more  than  territory  males then  site  that  three  a  could  were  only  for  smaller  which  territories M.  not  is  tuberculatus  tuberculatus  the  One  counterparts.  competed  e q u e s t r i s males,  became,  territories  maintained  M.  the  the  intrusions could i s that  in  behavioural  in  intensity  males  than  they  equestris  tuberculatus  tuberculatus  well-spaced  obvious  the  repletion.  male,  averaging  10  to  other  territories  2  rain,  variability  commonly  than  M.  during  Therefore,  one  (4  E.  m)  of  second  as  of  feeding  c o n s p e c i f i c s , E.  territories  potential  The  crowded  more  E.  and  repeated  e q u e s t r i s males  or  days  remained  that  feed  weather.  aggressive two  males  periods,  f l y or  f o r much  the  off-duty  Whereas  not  of  only  behavioural  did  periods  Crowding, variation,  males  defended contained accomodate  7-10 7-  territories  well-spaced  males.  An  82  E.  tuberculatus  usually  able  conflicts boundary two  M.  between  two  skirmishes,  they  "watch-chase" level  such  were  never  from  a  but, with  as the c o n f l i c t foraging  I thread  model  might  from  the process.  experiments  were  that  He  territory.  tuberculatus density  which  pattern  relatively  another  to  low  The e x c e p t i o n a l  male  over  became i n c r e a s i n g l y and r e p e a t e d l y  moderately  chased  a  a l l other aggressive  E. tuberculatus  to e l i c i t  changes  learned,  i t was shifting  with  in their  after  These  a behaviour  attempts  inspecting the  not a c o n s p e c i f i c and their  interfered  f o r 1981.  males  aggressiveness.  males  Bad w e a t h e r  of  study.  increase  i t , often  planned  E.  b e e . However,  only  of the  because  or twice,  t h e same  their  with  progressed  i n an a t t e m p t  which  once  territory.  territorial  unsuccessful  retreated in  the course  model  patterns were  individuals  presented  conflict  leafcutter  occasional  "watch-patrol"  d i d not increase.  aggressive  throughout  crowded,  was  Territorial  battles characteristic  one e x c e p t i o n ,  o f an e n t i r e  territorial  only  t o occupy  resident's  i n a n uncommon  larger  involved  very  territory  territory.  fluctuations in intruder  possession  much  males  attempting  of aggressiveness  engaged  an.occupied  not the constant  d i d experience changes  encountered  to a neighbouring  e q u e s t r i s males  prompted  was  that  t o move  Although males  male  territorial with  boundaries  a d d i t i o n a l crowding  83  Recogn i ti o n  Males ranging  most  were  from  (against  Intruders  of  an  able  0.3  m  various  tiny  flies,  (e.g.,  bees)  were  distance;  although  subjected  to  mating  a  distinguish a  males  definite  prolonged  head-to-head  characteristic  of  conspecific  a  in  never  attempted  react  to  to  M.  the  face  and  inspected  bulb  flies,  feathers.  Large  from  a  or  they  were  chased, could  distinguish  attempted  to  rare,  from  to  to  mate  to  An  not tiny  with  an  spirals.  Repeated  contact,  an  their  chase  a  the  confrontations, not  show  inspecting of lack  male,  close-up of  female  occurred  any  two  inspection.  engaged  in  a  never  they  may  pheromone  occurred.  have  and  -  response.  away. A t t e m p t e d the  male  discernable  behavioural  when  to  inspecting  and  did  cues  intruding  resident  to  resident's  behavioural  females.  response  equestris  but  on  the  combination  addition  were  males  females  response  sound  matings  male  Males  1m  recognized  also  relied  behavioural  turned  by  no  apparently  he  identified  since  to  conspecific.  conspecific  behavioural  Males  distances  large  shadows  inspected  attempts.  resident:  Since  sky).  apparently  were  conspecifics,  However,  showed  they  moving  at  backdrop)  conspecifics,  intruders  but  vegetative as  and  anything  a  intruders  such  bees,  from  moving  backdrop  of  flies  kinds  see  (against  unbroken  intruders:  to  intruding  been  or A  male  male-male male  did  not  84  THE  DRONE F L Y , E R I S T A L I S  Importance,  Eristalis known  as a Batesian  (Brower  genetic  Like  studies  insect,  other  at least  two  Males  but those  (Wellington,  1976).  and  mate  many  generation hibernation  eggs  1972). O v a r i e s has  been  oocytes  only  14 d a y s  i n 60  1979 a n d  the s u b j e c t of. attention  spatial  memory  lay their  10 d a y s  has been  clusters  100-200  batch  terminated eggs  (Heal,  i n summer,  are  from  not o n l y .1  emergence  before  begin  t o mature  and o v a r i e s  Females of eggs  winter  and S t r a d l i n g , after  a  female  contain  mature  o f t h e autumn 8-15  (op. c i t . ) and 1979).  l a y as  Second-  seeking  (Kendall  male,  Summer and  e t a l . , 1959).  and c r e v i c e s  (op. c i t . ) .  one  generation  after  (Dolley  oocytes  i t s phenology.  generation.are  i n t h e autumn  first  about  cosmopolitan  1979 a n d F i g . 1 5 ) .  out of h i b e r n a t i o n , later  this  t o use sperm  (Heal,  days  in caves  hibernation of  seem  once  and b a s a l  brought  generation  o f t h e autumn  f e m a l e s mate sites  i s well-  mellifera  per year,  o f t h e summer  Females  on  information  generations  females o v i p o s i t  a s 3000  but  received  in i t svisual  v a r i e t y of s t u d i e s  territorial,  generation  has been  and has a l s o  i s s c a r c e l y any  in fall.  probably  Apis  1976; H e a l ,  E. tenax  1979),  pest,  1975).  t h e wide  there  undergoes  the  equestris, (Heal,  History  of t h e honeybee  1965; H o l l o w a y ,  and Land,  Despite  and L i f e  agricultural  physiologists interested  (Collett  It  M.  i s n o t an mimic  and Brower,  Fig.14).  from  tenax  Biology  TENAX  days  after  lay several  85  Figure  14. Eristalis E. tenax female. comparision.  tenax (b)  and A p i s m e l l i f e r a . ( a ) An A honeybee, Apis m e l l i f e r a , f o r  86  .87  Figure  15.  A  mating  E.  tenax  pair.  89  In  various  human, o r nature,  rabbit eggs  decaying The  faeces  eggs  (Dolley  have  et  been  reared  a l . , 1959,  Heal,  usually  laid  on  organic  matter.  Eggs  hatch  often  respiratory  saprophagous after  studies,  are  larvae,  long  lab  the  referred process  to  at  medium. A d u l t  hatch,  flies  as  emerge  2-3  days  1957).  pupate  after  at  8-14  (Dolley  et  a l . , 1959,  Heal,  females  are  similar  in  appearance  and  females  are  usually  darker  separated 77  days  eyes in  the  (Heal,  1979).  laboratory  males  Both  and  oviposition. of  three  weeks  of  the  depending  may.  and  mm),  more  live  Golden,  upon  Males  (15  have  the  are  surface  length  sexes  (Dolley  to  In  other  because  1979).  and  or  end,  days,  moist  1979).  after  Two  the  temperature  than  manure  posterior  (Imms,  they  of  "rat-tailed"  their  in a l l stages  larvae  piles  on  although widely  for  as  long  as  1947).  Results  Characteristics  Males The m)  former  were  males  alternately intruders,  a  defended  broad-leaved  Resident  of  either  vertical  face.  clusters  of  low  vertical  established shrubs sat  on  watching then  Territory  along  (e.g., and  the  turning  to  Horizontal flowering  horizontal  vertical  faces  Rhododendron,  hovered open  or  near  space  inspect  few from  leaves  and  were  (e.g.,  of  tall  (1.5-3  Cotoneaster).  away  territories plants  a  territories.  particular the  shrub  flowers  established  Lobelia,  leaves, for  along where  Alyssum,  the  90  Aqeratum)  were  Territorial within  0.6  activity m above  surveillance leaf  interspersed  the s o i l .  o f t h e open  a  as  he d i d s o . T h e t y p e  All  Males that to  regularly area  heaps nor  every  one o f s e v e r a l  route,  sites,  returned  after  chase  sites  o r manure  piles)  females  they were  resources  (i.e., were  necessarily apparently  therein,  which  males  post  rotating  major (q.v. P.100).  feeding,  an a r e a  o f 0.36  sites  instead  m . 2  within  of  returning  edge  or p a t r o l  species.  part  within  attracted  sentry  had a  to  360°  and  a territory  stagnant  even  a  grooming,  within  along  never  a  hovering  or inspection,  posts  confined  to intruders  basking,  usually  was  from  defended  response  soil.  maintained  by w a t c h i n g  of the b u l b - f l y  Oviposition  were  resident  t o one o f t h e s e v e r a l  sentry  as d i d males  territories  of t e r r i t o r y  included  and mating  o f open  intermittently  on t h e r e s i d e n t ' s territories  A  space  o r on t h e s o i l ,  watching,  patches  on h o r i z o n t a l  on  influence  with  water  around  compost  of a male's 1 km  of i t .  to territories  regularly  territory, Passing  by t h e  inspected  food  and  defended. All small  territorial  a s 0.36 m  vertical  vertical  often  were  ranges  However,  extended  territories  Home  instead  in horizontal  territories.  territories  males  2  components  were  rarely  frequented  off-duty off-duty  often  boundaries  than  m  1.9  3.6  m, 2  in  and  m . 3  territories.  E.  territories,  as f a r as  3  horizontal  an a r e a  on t h e i r  sites  i n areas  a n d 0.22  of  encompassed  larger when  present  territories,  to include  sometimes much  were  15 m  tenax  and  beyond  as  91 territorial which  were  boundaries. scattered  ranges  were  because  of  their  the  Whether territorial  scanning  or  types  of  (on  male  to  For  crouched  both  from  a  a  m.  sites  Home  2  had  garden of  to  featureless  navigate  off-duty  to  h i s body  during  sites.  survey on  example,  a  more leaf.  i n two  of  males  himself  and  180°  also  only  they  or  at  him  most on  a  for time  duty,  commonly  he  at i t s  intruders was  spent  i f  during  vertical  than  a  location,  T h i s manoeuvre  observed,  employed  p r o p o r t i o n of  (on  rarely  vertical,  this  (Table V),  territory  males.  approximately  around  small  was  defense.  horizontal  territory).  Hovering  bulb-fly  in addition,  hovering  his  male  behaviour  (or basking  through  horizontal  a  hovered  marginal). A  allocated  the  territories  sentry post  were  of  f l y positioned  In  360°  or  stretches  territorial  was  above  temperatures  crouched  p a t t e r n s of  space  turned  500  such  or r e c t a n g u l a r  territorial  that  open  male  several  some  field  and  a  and,  his territory  either  was  of  males  hovered,  f l y males  drone  of  the  (Fig.16).  period  which  from  frequently  spatial  He  by  long  territorial  different  in bulb  different  range  imposed  across  visited  trapezoidal  included  between  males  observed  watching  home  b e h a v i o u r a l regime  tenax  often  Behaviours  considerably  center.  also  lawns)  travels  Territorial  E.  a  constraints  They  (i.e.,  The  over  male  a l l approximately  boundaries. ground  A  each which  duty the  territory) enabled  could  to  the  when  followed chasing.  hovering  followed a  chase  92  Figure  16. An E . t e n a x m a l e h o v e r i n g n e a r t h e c e n t e r o f h i s t e r r i t o r y , watching f o r intruders. T h e m a l e shown h e r e r o t a t e d t o s u r v e y b o t h t h e open s p a c e and t h e f o o d p l a n t shown ( A l y s s u m s p . ) .  93  94  Table  (or  V.  Time  allocation  territorial  Time  patrol hover chase chase-contact inspect watch nbask ngroom nf eed  0.02 3.60 30.30  a  may  patrolling allocated  serve  inspecting or  more  of  male's  to  inspecting  part %)  intruders. concerned All  with  frequented contact" "watch"  by  took  plants  the  males  were  almost  (Table V).  Thus  the  %  and  by  territories;  (Table V). ; Table  males  were  much  of  with  (see  male's  territorial  When  of  142  unlikely  searching more  in and  time  defense  on  discussion).  territories  a  flowers  i t is  "chase"  An  6 out  probably  females  therefore  V).  Rhododendron),  only  concerned  Males  (Fig.17).  (e.g.,  inspections,  solely  cases,  around.the  corollas  established  as  such  their  time  (7.3  feeding  bees;  In  flower. Since  inspecting  foraging up  male.  (%)  in his territory  tubular  discovering  observed  a  preceded  flights  Rather,  of  slowly over  into  were  inspection  tenax  procedure.  %  males  (4.22  times.  <1  food  way  35  reorientation  hovered the  of  patrolled  flowers with  chases  allocated  rarely  inspecting flew  out  a  males  time  male  29  as  occupied more  E.  7.30 6.00 31 .08 16.50 5.20  chase-contact)  Territorial  that  a  Behaviour  hovering  one  by  as  areas "chase"nbask"  pattern  and  among  95  Figure  17. E . t e n a x m a l e on h o r i z o n t a l t e r r i t o r y . l e g s d a n g l e l i k e a h o n e y b e e ' s , e x c e p t when t h e g r o o m s t h e m , a s shown h e r e .  The back insect  97  E.  tenax  males  was  a combination  "watch-chase-watch". temperatures integral  were  part  inspection  after  and  temperature  beyond that  appeared  On  males  when cool,  empty,  cloudy  because  warmth  >17°C  cool  Residents periods, sunnier  commonly,  becoming  on  a  activity  Patterns  the morning active  therefore, were  fell  most  sheltering  beneath  occurred  warm  of s u n l i g h t , the  flowers, again  below  territories  on a n y s u f f i c i e n t l y  few m o m e n t s  much  ate rapidly  when  unless  while  i t waned.  No  temperatures  were  periods.  warm  due t o h i g h  basked  days,  and  a i r temperature  disappeared  1980, p e r i o d s  periods  days,  basking  were  then  f o r prolonged  More  space,  an  during  a c t i v e before  to the nearest  defense  During rare.  there  lasted,  territorial  especially  the residents  Whenever  directly  rarely  the afternoon  object.  the  was.also  17°C, and d i d not remain  or unobtrusively  went  Behaviour  were  foliage  flies  when  chasing.  reached  the time  level.  "nbask"  and  temperature  Territorial air  by  17°C. Hovering  s u r v e i l l a n c e o f open  in Territorial  Sunlight  preempted  of t e r r i t o r i a l defense,  reorientation  j_.  " W a t c h " was  approximately  flights,  Variations  of "watch-inspect-watch"  o f warmth sunny  and f u l l  periods  fog or passing  low v e g e t a t i o n  active only  r e s i d e n t males  when first  were  sunlight  were  interspersed  with  altostratus clouds.  o r on t h e s o i l  during  t h e sun r e t u r n e d . fed to repletion  On  cloudy warmer,  before  98  beginning followed male  defense  immediately  basked  "watch"  activities. by  on a l e a f  position  changes full  i n radiant  watched  2.  from  Crowding  a  those  reach  87  bees) and  70  of s u n l i g h t  the the  between  leaf  behaviour  and  subsequent  were  on a  struck.  equestris  home were  sites,  >17°C,  in  frequently,  and  chases.  due t o c r o w d i n g males.  when range  Crowded  their  routes  as they  result,  similar  males to  or t e r r i t o r y .  harassed  and, as a  were  were  off-duty  Thus  attempted  were  to  often  off-duty. by c o n s p e c i f i c s c h a s e d  passed  through  through  a  The r e s t ,  honey-  their  during  resident's  a  a l l intruding  territories, 17 m i n  interval,  territory,  7 c o n s p e c i f i c males,  and bumblebees,  but  were m e r e l y  only 7  when  3 (all  syrphids,  chased  to a  boundary.  uncrowded  outside  patterns  one. F o r example,  passed  territorial  hover  that  foraging  An  males  not crowded  struck  were  resident's  and i n s p e c t e d  another's  going  and f l i e s  insects  a  a i r temperatures  constraints  off-duty  from  Males  rarely  f o r M.  to spatial  their  which  before assuming  hovered  post  in behaviour  territorial  prevented  bees  sentry  began,  When  after  was  center.  or absence  resident  overlapped  crowded  heat.  described  subjected sites  defense  session  by c o n s p e c i f i c s  Changes to  a  feeding  few m o m e n t s  at the t e r r i t o r y  on t h e p r e s e n c e  sunlight,  first  2-3 m i n o f g r o o m i n g ,  for a  Once, t e r r i t o r i a l depended  The  male  flowers  that  had r e c e n t l y  containing  foraging  been  off-duty  bumblebees  would  and wait  99  until  t h e , b e e s emerged  territory.  crowded  n e i g h b o u r s from  going  their  at  bumblebees  could  male  actually  pursued (L.)),  and t r i e d  not feign  attack  still  in less  to leave.  cm.  and a t t a c k e d b u t somehow  c o n f i n e m e n t on t h e t e r r i t o r y ,  conspecifics  polarized  light  potentially The attempts feeding  many  the  the zenith  of c o n s p e c i f i c  participants  males  Contacts  were  the  would, but males  would  were n o t male  even  maculata on h i m a n d  arising  from  from crowding  e.g., the l o s s of 1976)]  may  so t h a t ,  usually  after  affected  have  between  were  within  such males  and s t r i k i n g establish  were  1 h of such  each a  frantic  seen  a  per min), and often  could  scattered.  individual  12-18 m a l e s  territories  p e r male  no m a l e  also  Often  seen p u r s u i n g  such circumstances, food  pursuit,  that  [whether  (Wellington,  to establish  ( u p t o 18 c o n t a c t s  g e t enough  i t from  she t u r n e d  factors,  territories.  1 x 1 . 5 m.  males  strike  consequences.  and attempting area  a bee  (Vespula  aggression  or from other  to establish  as four  Under or  lethal  density  horizontal frequent  from  before  crowded  t o e s c a p e when  prolonged  repeatedly  One c r o w d e d  hornet  increased  prevented  the bees  male  predators  to sting.  male  would  These  attempted  by  Thus  until  a s an uncrowded  a bald-faced  dart  But even  circumstances.  managed  had been  In t h e e n s u i n g  and p o t e n t i a l  crowded  out of the  would  such a male  t h e bee r e p e a t e d l y .  butterflies  that  off-duty  >15  attacks  males  them  i n flowers,  itself,  i t tumbling  struck  harassed  were  extricate  sending  would  also  that  collecting  fully  above,  harrying  In c o n t r a s t ,  by  stopped  before  as other.  territory activity,  100  3.  Territory  Males fields  type  defending  of view  territories.  than  When  a vertical their  perched  see  and r e a c t  t o movements  and  overhead,  whereas  limited male on a  by t h e w a l l  vertical  As see  close  movements  intruders passed  These different  anywhere  below,  response  a vertical  2  Rhododendron  sarcophagids.  different interacted adjoining those and  territory  and thus  males  of eight  of view  was  as h i s  territory  severely view  counterpart's a male  on  of view.  responded t o those  which  of view  different  led to l e v e l s of  on t h e two k i n d s o f period  2 m high  wasp,  by  when  18 i n t r u d e r s :  t h e male  on a  5 honey syrphids  occupied  and  two  f o r 7 min, and  h i s neighbour  the intruders encounters with  t h e male  a n d 1 m wide  2 aphidophagous  period,  encounters  of a  of him.  field  o f 2 min w i t h  and with  to  i t , he c o u l d n o t  overlooking  side  was v i s i t e d  that  able  beneath  the overhead  only  i n one 15-min  2 m i n he h a d two b o u n d a r y  a total  in front,  to inspect  f o r 6 min, inspected  fora total  were  restricted field  and t h e r e f o r e  1 vespid  During  perches  more  in available  was  8 bumblebees,  Only  and t o e i t h e r  F o r example,  bees,  field  was a s b r o a d  flower  i n t e n s i t y , among  present,  side,  i n the flowers,  rates,  horizontal  p a t r o l l i n g or inspecting,  else,  territories.  flowering  on e i t h e r  had an even  differences  aggressive  the l a t t e r  When  to each  discovered  above,  or hovering,  territory  territory  he moved  on  the formers'  site.  h a d more r e s t r i c t e d  counterparts  of vegetation.  on a v e r t i c a l  a horizontal  territory  on t h e  he d i s c o v e r e d . with  intruders:  In  h i s neighbour, four  with  101  bumblebees, fly.  He  notice took  three  never 3  growing  or  which he  only  4.  16  age  M.  transition any  flights  when  their Late  intrusions,  which  and  a  sarcophagid  nor  did  he  s i n c e a l l of or  the  these  chasing  the  encounters  overlooking  entered  the  16  he  visitors  butterflies  for only  2 min  was  the  neighbouring  1 bee of  end.  the  and and  the  plants male.  He  challenged 2  muscids  territory  This  of  either  between  latter,  only  other  were  low-  male  while  overlooked  neighbour.  stage  e q u e s t r i s , E.  tenax  disperser to  drones  that  polarized  i n the  the  with  cabbage  min,  far corners  his  2  openings  with  c r o s s i n g by  and  13  i n t r u d e r s or  horizontal  there  perched  the  a  planted  syrphids,  remaining of  on  was  Alyssum,  the  transition  established  size,  interval  intruders at  from  observe  similar  visitors,  border  Insect  with  inspecting flowers  another  with  chasing  one  one  neighbour,,  r e s i d e n t male  or  boundary  Like  in  The  respectively  was  and  10  his  2 aphidophagous  i n one  the  .other  Lobelia  period. During  seven of  was  comparable  interacting  had 13  of  bees,  hovering  bees,  intruders.  4 muscids).  15-min  he  Ageratum,  honey  and  the  contrast, in a  territory  (8  honey  c r o s s i n g s by  while  individual In  noticed  border  place  with  had  males  undergo  a  r e s i d e n t . Although been  light  forced  waned,  to  I did  end  gradual I  did  their  observe  not dispersal  several  males  period.  morning  territorial  of  June  male,  24,  25-30  1981, drone  20  m  flies  from  an  (mostly  males)  102  were  observed  were  so  other;  feeding  intent  on  feeding  occasionally  another,  but  feeding.  These  completed  no  a  were  that  male  fight  their  establishing  voraciously  a  males  circled  ensued  and  presumably  dispersal  at  and  in  males  territories  or  thereafter,  several  of  daisies.  They  ignored  each  females  the  both  flights  clump  air  to  feint  subsequently and  and  females  were  searching  for  at  returned  which  feeding  to  had  before  oviposition  sites,  respectively. Shortly vertical other,  territories  became  very  nearby  and,  aggressive.  passed,  including  some  usually  chased  established,  These  by  observations  dispersal  period  they  fed  have  aggressive, Aggressive  do  not  even  when  They  that  other  i n t e n s i t y presumably  to  period, chase  males not  chased  aphids  become  no  these  although  and  males  staked  every  the  territorial  flies)  done  end  or  so,  on  of  they  males. their  become  conspecific  males  intrude.  diminishes  after  this  since  no  well-established  male  intruders  as  small  as  (see  E.  tenax  males  that  not  aggressive  aphids  each  intruder  territorial  at  out  bordering  tiny  uncrowded,  s u b s t a n t i a l l y . Having  establishment observed  (e.g.,  suggest  of  was  until very  ever  discussion).  Recognition  There females female until  of  i s no  from  Intruders  doubt  a l l other  in mid-air, she  landed  to  but  that  intruders.  A  more  often  followed  feed,  then  bask  or  male  can  distinguish  occasionally 10-15  hovered  cm above  clasped  behind her  her  for  a  103  several  seconds  entered  a  flower  Rhododendron), then  before  clasped  with  the  hesitated for  the  same did  back  males  before  become  male-male  between  over  was  often  impeded  to  a by  i n the  she  aliens  by  the  while  he  from  as  bees,  a  male,  No  female  emerged  unless  flower.  fled  i n the  lengthy  fights  s  while  an  alien  any  most  the  lasted  two  range. only  boundary,  and  the  male  attempting  various that  a  as  to  received struck,  male wait the they  characteristic two  to  kinds  of  often  chased  each  other  Confrontations long  as  i t  took  i t s escape  mate  with  Rather,  intruders, hovered  feeding  female,  and  chased  waited  f o r her  to  emerge.  of  conspecific  resident.  situations.  male  and  had  although  from  crowded  when  between  r e s i d e n t ' s home  he  Aliens  usually  30  wasps  territorial  they  blows  fact  such  treatment.  another  for  distinguish  containing a  competitors  until  territory  observed  even  illustrated flower  reach  rarely  able  outside  Confrontations  the  r e s i d e n t and to  were  hovered  since  lasted  alien  another,  a  as  emerge  but  the  I  the  (such  striking  interactions.  forth a  If  corolla  and  involved  sometimes and  c l a s p her.  tubular  received different  treatment,  males  male  i n t r u d e r to  not  a  to  her.  Conspecific butterflies  settling  males as  outside  away  a  potential  104  DISCUSSION My all  observations  three  which.is  species  show  a  the hypotheses  species-specific  i n f l u e n c e d and modulated  territorial intent.  support  males  do n o t a p p r o a c h  The o b s e r v e d  behaviour  also  capabilities  have  A in  variety  a l l three  among  males  a r e i n h e r e n t l y more  varies  species  with  below A  increasing E.  as  critical  subjected  males,  foraging  when  leafcutter  conspecific  or a l i e n .  to  by  crowding  and  Males  with  affected  the s u s c e p t i b i l i t y  intruders  were more  easily  gradient  equestris in  of c o n s p e c i f i c s , i f temperatures  on h i s t e r r i t o r y  disappears. responds  by  of p u r s u i t s .  pursued  but never  large struck  two  intruders an  species  intruding responded  of p u r s u i t s and  structure  of E. tenax  males a r e  intensity  or i f sunlight  of the other  hidden  intensity  a n d M.  Aggressive  and ceases  crowded,  intruders. Habitat  aggressive  E. tenax  i n c r e a s i n g the frequency  contacts  species.  aggressive  intensity  bees,  territorial  thermoregulatory  the density  level  t o crowding  the frequency  tuberculatus  such  age and s t a g e ,  a certain  male  whereas  with  of  E. t u b e r c u l a t u s  pugnacious.  increases  insect  heighten  maintained.  aggressive,  2)  Aggression  but the inherent  i s always  the least  three  fall  species,  the three  always  all  of circumstances  aggression  f a c t o r s and  patterns  s t r a t e g i e s of each  in  of  a l l i n t r u d e r s with, m a t i n g  i m p l i c a t i o n s f o r the  Variation  1) m a l e s o f  level  by e x t e r n a l  d i f f e r e n c e s among  and mating  that  (territory  males  to  by v e g e t a t i o n  type)  crowding:  on a  vertical  105  territory resident  than  on a h o r i z o n t a l  on a v e r t i c a l  tubular  of intruders  inspect  i t . Thus  noticed  fewer  crowding  intruders  pressure  influence  of habitat  territorial  dragonflies.  intruders  approaching  habitats  therefore  were  He a l s o  be  i n dense  site  dragonfly observed level  that  level,  males  "considerable among  with  here.  both The  high  t o behave  territory to  about the by  Leucorrhinia could  but that  from  many  correct  males  density  that  tenacity  see  residents  intruders  i n high-  of  and  density  sex r e c o g n i t i o n  habitat was m o r e  aggressive  When  may  increased density  This  and s i t e  effects  affected  important  only  to a  surpassed  response  of any o f t h e three i n aggressive  type  than  i n t e n s i t y . He  non-aggressively  tolerance".  Increases  crowding  to  horizontal  interactions that  flower  susceptible  habitats  meters,  intensity  density.  social  territorial  studied  site  aggressive  began  a  recognition  that  shielded  in increasing  of population  the v i s u a l  observations  intruder  postulated  and that  density  less  were  habitats.  (op.cit.)  tenacity,  on  for several  postulated  into  on a  similar  many a g g r e s s i v e  situations.  Pajunen  f l y male  He o b s e r v e d  visually  corollas  from  flew  in sparsely-vegetated  spared  impaired  a drone  structure  males  he  an E . t e n a x  on a v e r t i c a l  ( 1 9 6 6 b ) made  rubicunda  isolated  time male  flower  a n d was t h e r e f o r e  than  . Pajunen  where  became  each  an E . t e n a x  territory  dense  territory  (e.g.,Rhododendron)  stimulation  one. In a d d i t i o n ,  and  certain  that  high  exhibited  was n o t o b s e r v e d  syrphid  i n t e n s i t y were  species associated  tenacity.  of dense  populations  were  rarely  106  confined was  to  a  usually  and,  as  where  a  he  harassed  result, was  E.  aggressive  off-duty turning (1972) sites  to  more  face  observed between The  a  i t s afternoon  contain  nettles,  and  of  flew  to  were  non-aggressive  roost  with  other  prevented their  from  Although spatial shrink  in  entire M.  those  sites  parts  response  to  to  more  sometimes harassment  varied  their  not  Inachis  say  roosted  whether  by  Aqlais  flew  ca.  100  under  did  not  to  often  of  roosting  i o and  and  more  away  Baker  of  patch  sites,  became  roost.  sharing the  a  males  aggressive  in  little.  Territory  by  behaviour  crowding.  t e r r i t o r i e s , males  aggressively, extended at  them.  changes  confinement  of  simply  choice  latter  sites  deprived  off  behind  But  as  prolonged.  provoked  patterns  territories  equestris  was  nor  time  nettle  roosting  reaching  response  relinquishing  nearest  did  stay  intensely  if its territory  Baker  crowding  defense  at  or,  males.  territorial  crowded  The  well  territory,  off-duty  in  as  manner.  its territory  site.  his  became  sought  take  sites  male  frequent  this  butterflies,  the  Males  in  difference  territory  on  to  both  vegetation  left  territorial  off-duty  respite,  could  a  remain  neither  they  roosting  i t s own  to  rarely  were  always  to  harassed  which  wall  area,  equestris  nymphalid  former  on  likely  comparable  two  nettles  travels  M.  because the  crowded  without  and  males,  a  his  territories,  to  In  confined  periods  urticae. m  when  their  was  tenax  tuberculatus  from  on  subjected,  intrusions.  E.  territory.  their  and  E.  previously-neutral  size  Instead  did  and  areas  sites.  not  of  defended  tenax  defended  patterns,  in These  their  107  observations Aglais is  1972),  than  but  are  m  the  share  to  of  between  continuously  the  same a s  uncrowded  situations.  In  shorter  those  at  one  spacing  returned  It with  a  which  i s not male's  insects  (i.e.,  were  territorial each to  feeding  feed bouts  instance,  In  the  pursued  early part  established  and  males  to  observed frequent  at  least  distance between  Pajunen  between  3-  was males  in  (op. c i t . )  sites  but,  q u i c k l y d i s p l a c e d and  territorial  E.  tenax), was  gradual  dispersal  of  aggressive after this  other  he  such  the  r e s i d e n t s . The  i s the  in the  and  he  occur.  Although  transitions  ceased, to  territorial manoeuvers an  and  were  disperser  the  In  settling  bulb  fly  the each than  resident.  drone-fly  normally  steps  intense  established  to  male  intersperse  more  to  (one  behaviour.  i n s e c t s not  narcissus  from  in  few  characteristic  began  became  way  dispersal)  had  flight  varies  (i.e.,  transition  sequence, tiny  aggression  interest  stages  undergoing  bursts  of  more  non-aggressive  territorial)  occurred  aphids  that  v o r a c i o u s l y . Then with  This  maintained  was  He  maintained  sites.  of  (Baker,  (1964).  subjected  distances  s t a t e . Of  from  First,  initial  which  and  resident  case.  number  normal.  observed four  the  populations,  male  surprising age  equestris,  those  to  transitions  aggressive  began  least  when  situations,  dense  observed  cases,  Pajunen  distance  occasionally  in  of  occupied  the  when  that  suitable territories  those  in high-density  virtually  M.  Baker's-observations  territories  number  similar  to  L e u c o r r h i n i a caudal i s males,  interactions 5  in contrast  u r t i c a e males  greater  that  are  males  pursued  persistently  by  108  chased  bumble  observed  to heights  in resident  The of  bees  "overlap"  territoriality  M.  equestris  between was  h e was  still  light  forced  him t o stop  study  area  period  he  number  of rapid  site.  patch  of d a i s i e s  also  bees)  with  aerial the  began  area  aerial  longer  min a f t e r  of the urge  he a r r i v e d spirals  (Wellington,  Such  behaviour  aggressive  still  only  (mainly  down  briefly  to crawling to disperse  i n response  on by c i r r u s  when  bumble  to  while  were  to  leave  feed  basking,  to  feeding still  apparent  initiated  t o changes clouds  an  and then f o r  and  he  above,  to delineate h i s  to react,  i n the area,  again  the prominent  attempting  to settle  began  and f i n a l l y  each  s e t t l e d . As n o t e d  intruders  a  of the  After  above  As he b e g a n  at first  gradually  p o l a r i z a t i o n brought  zenith  to  He  began  a n d made  the f l y established  h e was  dispersing.  and t o bask,  Traces  orientation  while  that  corners  became e v i d e n t .  a l l airborne  polarized  During  himself,  t o hover  bulb f l y ,  10 m i n i n t h e  of the four  he h a d f i r s t  he a l s o  intruders  intruders.  of  even  boundaries,  sessions.  each  also  waning  His first  to orient  i n t e n s i t y . Thus  and c o n t i n u e  airborne  37  on w h i c h  when  and the onset  in this  to continue.  the f l y returned  unusual  voraciously  toward  urges  to chase  territory  travelling.  spiralled  flights  or s p i r a l ,  the end of d i s p e r s a l  attempting  But t e r r i t o r i a l  flight  he  spent  action not  males.  actively dispersing  frequently  an a g g r e s s i v e  e s p e c i a l l y pronounced  since  were  o f 10 m,  i n the plane  passing  through the  1953). suggests  resident  that  i s directed  t h e t r a n s i t i o n from by a  "biological  disperser  clock"  109  mechanism while  winding  changes For an  which  gradually  insect  i n other  there  emerges  from  lepidopterous  cocoons  (Dethier, requires  expanding make  aphids were  making  would.  These  redirected onset have  been  activity  partly  much  level  before  may  (Lorenz,  than  be a  fly's  supporting  would  the bulb  i n response intense  ignore  f l y which  or  to the  aggression  first may  also  by t h e d i s a p p e a r a n c e  Wellington's  the aggression  hypothesis  of  that  of t e r r i t o r i a l  male  1976). among  temperatures  males  of a l l three  temperature.  ( c a . 20°C  f o r M.  and E. t u b e r c u l a t u s ) , males  species  Below  equestris, ca.  ceased  territorial  activity  and basked  on t h e v e g e t a t i o n  temperature  was  marginally  below  only  towards the  established residents .  on s u n l i g h t a n d a m b i e n t  f o r E. tenax  newly-  learning procedure  1966),  The b u l b  syrphids  aggressively  a s was  adult  (e.g.,  The  established residents  of aggression  threshold  in their  stage  of aggression.  as  weather;  newly-territorial  adjustment,  increases  spring  about  t o i t s new  behaved  higher  histories.  as a newly-emerged  due t o h i s c o n f i n e m e n t  (Wellington,  depended  certain  which  which  adjustments  confinement  The  17°C  bees  polarization,  insects  also  flies  of aggression.  zenith such  flies  level  Such  physiological transition  t o move  just  i t s wings),  a behavioural  bumble  begin  And  in their  drone  and t i n y  pursued  1980).  and d r y i n g  male  pupae  stage.  stage  of i n s e c t ' s l i f e  long  some a d j u s t m e n t s  adjustments  arrived  aspects  diapause:  the "disperser"  aggressor"  i s a comparable  begins,  insect  down  up t h e " t e r r i t o r i a l  a r e common  example,  winds  ( i f the  the threshold)  o r on t h e  no  soil  ( i f the  increase  temperature  proportionally  species-specific seemed  to the  males  basked  thresholds, when  leaf  have  lower  three  as  the  too  f o r male  females  basked  between  basking  differences  periods.  1981,  to  1966b,  laboratory  males  (Thinopus  pictus;  food. time  This  or  Thus  difference  the  to  but  spend  related  1968,  i n the  time  remain  one  males  be  times,  be  active  1966b,  Buschman  activity.  1982),  not  thresholds  or  male  than  In males  although  in  the  species  same  "trading  cool  of  of  feeding  do  sex  high cost  the  were  Different  staphylinid  during  in temperature  these  mechanism  time  during  temperatures  1981).  therefore  active  of a l l  (Pajunen,  consume  may  Females  observed  the  shade Females  in feeding  more  1982)  and  have  to  were  33°C.  flying  when  Richards,  of  sought  continued to  Tengo,  much  and  During  workers  and be  that  f o r mate-seeking needed  feeding  thresholds  females  indicates  males.  activity.  Other  Richards,  reserves  than  to differences  and  temperature  temperature  reached  d i s a p p e a r e d or  may  Spieth,  the  occurred; i f not,  f l y which  sunlight  differences  females  (Pajunen,  threshold  upper  bulb  actively  Schone  aggression,  thermoregulation,  a  frequently,  thresholds  species,  by  territorial  sex  also  thresholds  sunlight  temperature  many  are  i n temperature  Patterson,  the  above  not  "on-off switch": i f temperatures  i n the  still  more  increased  aggressive behaviour  temperature  when  lower)..Aggression did  Instead,  demonstrated  were  much  temperatures  fed. There  periods  and  an  temperatures  species  low  as  threshold, or  as  threshold.  function  above  was  amount  off" have  feeding the  cloudy  between  of  energy  periods. sexes  may  111.  be  due t o t h e f a c t  energy  reserves  The  difference remains  density.  M.  unexplained,  larval  males,  whereas  exposed  to conspecifics  a l . , 1942).  E.  tenax,  however,  influences  adult  speculative  during  aggression  dragonflies  insects Maier  (1961)  wrote  to intruders workers  cannot  that  (Moore,  that  other  Bick  and B i c k ,  al.,  1981).  1979).  male  intruders  that  adults,  other  (Doucette  density  larval  of  density  beyond the  the approaches  Other  before  (1964)  contact  however,  (Pajunen,  females 1964,  1975, S e v e r i n g h a u s e t  postulated  that  He  that  with  1973,  1 9 7 2 , Raw,  failure  and that  to  accurate  territorial  to recognize  the evolution  the development  and  f o r other  and C o l l e t t ,  i s made  aggression.  combined  true  c a n a n d do d i s t i n g u i s h  a prerequisite  condition,  in nature",  investigators,  o f s e x was assumed  of a l l male  sexual  t h e same h o l d s  recognition  was  larval  Females  1974, L a n d  insects  1965, B a k e r ,  Johnston  of  that  1952, S c o t t ,  many  from  primitive  as  periods  be c a r r i e d  to  three  males a r e  the l a r v a l  "are apparently  believe  and Waldbauer,  believe  aggressive  larval  the  i n part  E. tuberculatus  about  the  stage.  Kormondy  other  be d u e  among  t h e company o f  so t h e h y p o t h e s i s  Recognition  some  from  lack  i n females.  aggressiveness  their  i s known  and t h e r e f o r e  present  a l l highly  less-aggressive  Nothing  less  b u t may  i n s t a r s away  larvae,  et  feed  i n inherent  equestris  their  males  ( i . e . , f a t body)  syrphids  spend  that  of  s e x was t h e territoriality  of sex r e c o g n i t i o n  so  that  112  sexual  reactions  were  released  observed  that  resident  butterfly discovered  offered  this  resident Most be  males  based  on  and  the  above,  (Eickwort  and  sexual  there  purely  type  pursuit  probably  in  until  aliens) never in  in  mistaken  combination  identify  her.  with  or  a  It  may  (Jacobs,  combination  of  quarters  association  have  males  same way  moment  identify  males  females  and  mate  driven  v i s u a l cues, never  turn  remains  of  the  (and,  make to  from  males  off. A  a  than  less  but  of  here  a  else.  These  often,  females  were  female's.behaviour,  i t possible  face  there  studied  anything  each  virtually  observer  males  them,  when  However,  for  with  1955,  distinguishing  a l l intruders  conspecific to  in  contact.  syrphid  aggressive  approaches  and  of  of  occurs  (Jacobs,  difficulty  the  mistook  Females  intent.  visual.  recognition  pre-mating  the  attempts  for  sexual  the  clasped  brief  mating  behavioural  close  Although  correctly  sometimes  at  mind  not  males  acts  the  did  never  1960);  and  that  colour  1964);  in  male.  they  than  a  male,  theory  wing  and  confusion  aggressive  distance,  the  (1972)  when  was  is primarily  territorial  in  the  always  cease  more  1963)  close  observers  begins  more  that  very  aggressive  indistinguishable is  is a  But  with  Baker  1980).  agree  motivations  between of  who  not  for  (Ito,  that  only.  intruder  (Pajunen,  pattern  Ginsberg,  1964).  (Lin,  pheromone  workers  that  Pajunen,  a  evidence  frequency  colour  plus and  Even concede  and  an  recognition  pattern  wingbeat  that  intruders  that  flight  differences  as  approach  agree  females  interactions did  observation  workers  1955)  male-male  for  male  or  for  a  male  engage  in  to  113  head-to-head obvious  spirals;  manner  to  in  et  instances  male-male  attack from  (and  the  close-up 1980) an  mate  ensures  like  a  like  i t . Such  prevent  would  be  time  their  a  energy  Other  species,  there  patterns  of  are  species and  bulb  regular  center  of  their  and  from  like  driving  a  watched  patrolled  routes. of  In  for their  female,  territories.  survey They  if  but  attempt  females  negate  to  which  away,  where  and  the  a l l the  Spec i e s  suggest  among and  different  i n t r u d e r s from  periods rarely  to  three  spatial  strategies.  territories  the  Males a  along  one  both post,  of  allocated  hovering  patrolled  of  sentry  c o n t r a s t , d r o n e - f l y males  their  a  defense.  Between  mating  emit  . However,  still  differences in behavioural  and  response  Ginsberg,  species  would  territorial  which  no  " d e f a u l t mechanism"  mistake  defense  that a l l  specifications,  differences in aggression  flies  large proportion  a  1973,  aggressive  females  r e s i d e n t may  Di f ferences  abilities  intermittently  several a  of  an  female  in aggressive by  Bush,  identification  be  males  to  (Eickwort  three  any  coincidence when  responds  may  away  territorial  thermoregulatory  of  the  valuable  alloted  a d d i t i o n to  bees  out  in  resident elicited  correct  attempts  females  and  i s possible that  female,  territorial  driving  In  two  respond  i t i s no  the  conspecific,  especially  and  do  satisfies  smell  with  of  by  some  would  act  contact)  as  not  clasping occurred  pheromone  looks  doesn't  often  do  ( c f . Prokopy  Thus  It  intruder  i.e.,  a l . , 1981).  i n t r u d i n g male.  which  they  h i s approach  Severinghaus of  fact  long  near  the  1 14  distances plant  without  or flower  individual Hovering energy  stopping.  to flower  leaves  is a  and  drone  flies  than  bulb  flies  suggests  incapable  of hovering  flies  "afford"  M.  need  and  lack  female  oviposition  mate  with  potential flies  females  they  inspection territory's  open  However, not  entirely  amount  long  on  f o r incoming  likely  than  are  physiologically  rate,  whereas  i s unlikely, of hovering males  and while  they  patrol  The  defend  opportunity  But  drone  out  food  females.  s i t on  t o be d i s c o v e r e d  by q u i c k  t o volume  t o watch  stake  feeding  be  which  territories. They  f o r long  1975).  of the  continue  because  may  area  males,  assured  sites.  hovering  great  surface  Since  leaves  during  flights  or  hovering  through  the  spaces.  the d i f f e r e n c e i n the type explain  of energy  time  The  drone  Bulb-fly  their  them.  o r b)  capable  as they  of  flies,  at a  are almost  are inconspicuous  flights  flies  large  inspected  i n which  ( H e i n r i c h and P a n t l e ,  likely.  sites  a r e more  bulb  to  ( H e i n r i c h , 1979).  E. t u b e r c u l a t u s  oviposition  and watch  females  flowers,  as  in front  suggestion  their  pile  sites,  do n o t d e f e n d  these  by  rates  plant  and  activity,  as drone  first  i s more  oviposition  resources  territories,  energy  although  o f body  rationale  to  as o f t e n  limited  second  a)  are certainly  (see p.60),  physiologically ratio  that  n o t . The  from  p r o p o r t i o n a l l y more  t o expend  e q u e s t r i s males  periods  spend  flew  by h o v e r i n g  a t near-maximum  that  flies  their  flowers  fact  bulb  on  they  "metabolically expensive"  i s expended  can  Instead,  why  a drone  ( i n terms  of s i t e  f l y should  of energy  defended  invest a  expended  does  greater  hovering)  than  a  •115  b u l b - f l y . male. concept  of  sperm  dragonflies Ginsberg,  (Alcoc.k,  and  and  a  from  female  (see  (Lin, 1981)  difference  in  drone-fly physiology behaviour those mate  are  and  may may  muscids  from  the  Raw,  also of  mate  only  worthwhile  for drone-fly females  maximize  fact  flies.  i t would  once  males  and  their  to  chances  of  eggs.  once.  between in  a  The bulb-  territorial resemble  the  which  before  they  useful  tactic  for  Drone-fly  their mating  and  reproductive  closely  Thus  Other  and  them  defend  expend  her  uses  of  1979).  to  female  most  sites.  (Heal,  a  B u l b - f l y females,  with  be  1970c,  (Buschman  that  more  obtain  mate  oviposition  apparently  to  bulb  to  to  order  the  and  and  mate  patterns  in  Waldbauer,  of  flies  this.difference  last  fly  and  most  only  that,  (Parker,  review),  and  and  could  Consequently,  newly-emerged  1974)  obviously  defend  for  the  (Eickwort.  orthopterans  defense  male  bees  fertilize  explain  of  evidence  (Maier  drone-fly. males  oviposit. males  to  monandrous  than  repeatedly,  syrphids  reflect  patterns  of  require  1963,  some  l i e in  anthomyiids  1970c,  mating  may  i s some  in  several  territorial  males  question  There and  1976),  Parker,  last  Patterson,  that  d r o s o p h i l i d s and  coloepteran,  her  bees  to  1979),  Parker,  lepidopterans sperm  answer  precedence.  1980),  Bloorman 1979),  The  sperm  resources  with  bulb-  females  i t would  energy  they  be  visited  hovering  virgin  by in  females.  1 16  IMPLICATIONS Territorial M.  equestris  foraging often  and  paths  to  the  bees  attempted E.  foraging their  tenax bees,  pollen  harassed  in  mnemonic 1971,  a  Wellington males  syrphid  species.  was  studies  involving  syrphids  E. from  blocked  defending  1 x  territorial outbreaks syrphids  of and  In  by and  and  struck  males  (1976)  bees,  fact,  of the  intended  males  U.B.C. for  in  syrphid  routes in  as  to  the  7  to  preventing  oviposition  subsequent  emergence  of  the  (von  Frisch,  aggressive  in  this  behaviour  However,  four x  8  m  territorial plot  by  i t s corners.  aphid  of  of  a l l aphidophagous  few  contribute  bees  used  females.  of  with  less-aggressive  site  as  each  emerged  oviposition  virtually  stunned  containing  view  Similarly, other  knock  syrphid  honeybees  field  use  Sometimes  territories could  1979).  to  et a l . ,  that  in  other  I  apecies  the  they  avoid  surprising  excluding  drove  as  to  particularly  Cmiralova,  as  flowers  reported  learn  i s not  force  Severinghaus outside  agents.  syrphid  manoeuver  repeatedly  quickly  which  plot.  see  the  control  with.enough  hovered them  disrupting  aggressive  the  (also  Wellington  a l l entrance 2 m  bees  repeating  often  of  particularly  biological  more  aphidophagous  tenax the  two  :  species,  capable  foraging  capable  originally  aggressive  males  are  are  the  fashion  fact  capacity  a l l three  f l y away  then  syrphid  study  and  loads.  territories,  of  males  this  AGRICULTURE  p o l l i n a t o r s and  ground, to  of  tenax,  males  airborne  them  1981).  E.  of  observed  striking  males  FOR  Such  population species  aphidophagous  of larvae.  117  AREAS The  work  questions the  presented  than  how  intensity  male  as  tightly  is linked  a  the to  and  contribute  substantially  of  the  parameter  three  to  Therefore,  I  i n the  behaviour adult  has  males  arranged  to  larval  not  been  a  the  behave or  be  is a  with  a  allowed  1973).  each  become  be  could  of  be  the  the  Territorial  l a b o r a t o r y , but  room  i f they  that  natural  urge,  of  difficult  monitoring  to  energy  aggression  adults.  rearing  to  to  parameters  i n the  about  aggressiveness  territoriality  dispersal not  seems  l a r v a e of and  gained  aggressive  King,  adult  resulting  should  do  see  be  more  conditions,  but  three  investigated  their  males  (also  rearing  bed,  in  territorial  other  chamber  raises  attempting  Inheritance  densities  flower  adults  larval  d e n s i t y on  probably  walk-in  exhaust that  of  by  variation  studied here,  different  mimic  Newly-emerged  indicates  the  l a b o r a t o r y by  would  in a  they  that  behaviour  released  until  species  of  species at  aggressive  to  and  i n f o r m a t i o n can  precedence.  the  effect  beginning  observed  suggest  The  STUDY  decision-maker  study  first.  three  More  q u a n t i f y and  explored  studied  is a  inheritance,  budgeting,  males  sperm  FUTURE  here  i t answers.  territorial  discover  FOR  had  light  f l y in a  territorial  until  been  source.  wind  since this  were  tunnel  study they  have  dispersed. Energy-budgeting males fat  and  body  females  and  could  d e p o s i t i o n of  be  thermoregulatory s t u d i e d by  field  animals  differences  comparing with  those  food  between  intake  and  of l a b o r a t o r y -  118  reared  animals  amounts [E. body  of  tenax  food  wind  are  they  overwintering  i f males  have  and  and  the  and  laboratory  been  found  (Kendall  females  to  activity,  flight  in order  one  could  to discern  consume  or  of  the  and  in a  with  energy  expenditure  during  sex-related  thermoregulatory  insects  changes  a i r temperature  also  be  useful  aggression, that  males  activity can  and  genus  a  f l y at and  Pantle,  and  would  energy  requirements  of  male  and  others  because  (op.  explain  E.  are  those  males  tuberculatus  surface  shivering  area and  10°C  shivering. described  able  males,  to volume  therefore  shown  thoracic  shivering  than  b a s k i n g and  their  i n b a s k i n g and  large  of  c i t . ) have  differences  a  ) have  non-aggressive  such  E.  (see p  a i r temperature  experiments  hovering.  also  It  of  tenax  would  inactive  differences.  combination  why  of  such  temperatures  regulate  as  temperature  shown  for  Syrphus  or  males  less  1975).  Pantle  Syrphus  the  study  ambient  and  of  h i g h e r ambient  females,  Heinrich the  this  require  do  absence  to quantify  since  than  (Heinrich  or  or  temperature  i n body  presence  By  laboratory  M o n i t o r i n g changes  elucidate  fat  i n body  quantify  i n ambient  as  with  1972).]  differences. with  energy  in caves  Stradling,  f l y freely  different  field.  storing  hibernating  m o n i t o r i n g changes  flight  females  apparently capable  females  males  tunnel  see  in either  males  since  allowing  to  temperature  might  between to  spend  which  spend  flies  of  through  Thermoregulation  above  ratio,  that  lack may  more  also  species  and  so  time  much  body  pile  r e l y , on  time  elucidate  close  and  have  basking  more  to  the  soil  a  119  where air  temperatures  20 cm  above.  are usually  equestris  males  as  suggested  by H e i n r i c h  may  Energy-budgeting  sperm not  i n E. tenax,  variations species.  be c o r r e l a t e d and P a n t l e  experiments,  (as per Bloorman•and  precedence  occurs would  i n behaviour  i n M.  pattern-recognition  in Materials  database,  would male  further  warmer, t h a n t h e among  warm-up  combined  Parker,  equestris  with  irradiation  and E. t u b e r c u l a t u s but explanations for intensity  i n conjunction with  (such as the streak  and Methods)  shivering,  1976) t o s e e w h e t h e r  p a t t e r n s and a g g r e s s i v e  applied  understanding  syrphids.  by  (op. c i t . ) .  provide partial  techniques  described  heard  with  Results of l a b experiments,  territorial  degrees  The h i g h - p i t c h e d whine  M.  experiments  several  to a  program  larger  of d e c i s i o n s  made  by  among  1 20  LITERATURE  CITED  A l c o c k , J . 1 9 7 5 . 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O l i v e r and Boyd, E d i n b u r g h . 653  to pp.  APPENDIX  DATE  LOCAL APPARENT  April May  I  21  21  (SUN) TIME  1100  1130  1200  1230  1300  50  —  53  —  50  57  60  61  60  57  June  21  61  64  64.3  64  61  July  21  57  60  61  60  57  50  —  53  •—  50  41  •—  39  August  21  September  21  39  A p p r o x i m a t e maximum s o l a r  elevation,  Vancouver,  Canada  N lat.).  disappears  from the  greater  than or  excessively varies  (49  z e n i t h when t h e  equal  turbid,  slightly).  16' to  60  (unless  i n degrees, Polarized solar the  i n which case t h i s  angle  sky  at  light is  is  relationship  APPENDIX I I BEHAVIOURAL DATA  The frequency d i s t r i b u t i o n s presented here a r e c o n s t r u c t e d from the output of the " s t r e a k " program ( d e s c r i b e d i n M a t e r i a l s and Methods). A l l b e h a v i o u r a l sequences graphed c o n s i s t of t h r e e b e h a v i o u r s . In s i x of the n i n e h i s t o g r a m s , the sum of the number of o b s e r v a t i o n s i n each column does not t o t a l "n", because t h e r e were too many sequences which o c c u r r e d o n l y once or twice to graph them a l l .  The  f o l l o w i n g symbols a r e used: w P g m h i c cc nb nf ng fb ff fg  watch hover goaway mate hover inspect chase chase-contact nbask nf eed ngroom f bask f f eed fgroom  133  A.  MERODON EQUESTRIS  140-  n = 60l  120-  100-f-  CO  80  W o w c/  w  CO  60  o  PS w  40-  20-  0 p  w  c  nb  cc  ng  g  nf  m  i fb  ff  fg  SEQUENCES ENDING IN  Al.  D i s t r i b u t i o n o f t h e t o t a l number o f b e h a v i o u r a l  sequences.  134  A.  32. 30  MERODQN EQUESTRIS  n = 117  r-nb-cc-wn w-cc-w  nb-c-w 254  204w-c-w  W-P-W  15 4-  10  0 MOST FREQUENT SEQUENCES ENDING IN 'w'  A2.  D i s t r i b u t i o n o f t h e most f r e q u e n t sequences ending i n 'watch'.  135  A.  MERODON EQUESTRIS  50  n= 1 3 2  45 403500  w u Z w o* w  3025  oo  O  3  20 15 + 10+  c-nb-p cc-w-p c-w-p  _n b-cc-p_ w-cc-p w-c-p  MOST FREQUENT SEQUENCES ENDING IN 'p'  A3.  D i s t r i b u t i o n o f t h e most f r e q u e n t sequences ending i n ' p a t r o l ' .  136  B.  EUMERUS TUBERCULATUS  10  n =328  100" 9080 -• CO  w o  70  25 W W CO  60t  fa o cr;  w  •3 3  50" 40 30 20 10 0  w  i  s  nb  ng  fb  g  fg  SEQUENCES ENDING IN  Bl.  D i s t r i b u t i o n of t h e t o t a l number o f b e h a v i o u r a l Note the absence o f ' c h a s e - c o n t a c t ' .  sequences.  137  B.  EUMERUS TUBERCULATUS  n =70  35-  304  25  20  15  0 MOST FREQUENT SEQUENCES ENDING IN 'p'  B2.  D i s t r i b u t i o n o f t h e most f r e q u e n t  sequences ending i n ' p a t r o l '  138  B.  EUMERUS TUBERCULATUS  5 0 45  n = 110 nb-p—w  co w CJ  w ow co Pn o  nb—c—w  erf W  w— p —w  w— e —w  I5 10 -  5 -0 MOST FREQUENT SEQUENCES ENDING IN 'w'  B3 •  D i s t r i b u t i o n of the most frequent sequences ending i n 'watch'.  139  C.  ERISTALIS TENAX  125  n =42  120  no 100 90 co w o is w cy w  CO  80f 70  Pu O  g  60  50' 40 30  20 10  0 nb  w  i  ng  h  cc  SEQUENCES ENDING IN  Cl.  D i s t r i b u t i o n o f t o t a l number o f b e h a v i o u r a l sequences. Note t h a t ' p a t r o l ' i s r a r e , 'hover' i s p r e s e n t , and ' i n s p e c t ' o c c u r s more f r e q u e n t l y than i n t h e o t h e r two species.  140  C.  ERISTALIS TENAX  25  n = 148 w-c-nb  201  CO  w  CJ S3 W  c w  n b-c-nb  CO  o  w-cc-w nb-c-w  w  *  104-  w-c-w  hb-i-n b  w—i —w  MOST FREQUENT SEQUENCES ENDING IN 'w' AND 'nb'  C2.  D i s t r i b u t i o n o f t h e most f r e q u e n t sequences ending i n 'watch' and  'nbask'.  141  C.  ERISTALIS TENAX  n » 122  25T  201  Icc-n b-c  CO  w c_> ts Pd &  o* w co  cc-w-c 15  Pu O erf W  I01  c-nb-c  c—w-c  0 MOST FREQUENT SEQUENCES ENDING.IN 'c'  C3.  D i s t r i b u t i o n of t h e most f r e q u e n t sequences  ending i n 'chase'.  


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