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Bird communities in relation to the structure of urban habitats Lancaster, Richard K. 1976

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BIRD  COMMUNITIES STRUCTURE  IN R E L A T I O N  OF URBAN  TO  THE  HABITATS  by  RICHARD B.Sc,  A  University  THESIS THE  K. L A N C A S T E R  of British  Columbia,  SUBMITTED IN P A R T I A L REQUIREMENTS MASTER  in  FULFILMENT  FOR THE DEGREE OF  1974  OF  OF  SCIENCE  the Department of Zoology  We  accept  required  THE  this  thesis  as c o n f o r m i n g  to the  standard  UNIVERSITY  OF B R I T I S H  May, Q)  COLUMBIA  1976  R i c h a r d K. L a n c a s t e r  1976  In  presenting  this  thesis  an a d v a n c e d  degree at  the  shall  I  Library  f u r t h e r agree  for  scholarly  by h i s of  the U n i v e r s i t y  make  that permission  purposes  written  thesis  for  may be g r a n t e d It  2^n-rr^^-v  Columbia  2075 W e s b r o o k P l a c e V a n c o u v e r , Canada V6T 1W5  SsLsodu~Ms  Zl  /976  the  requirements  B r i t i s h Columbia, for  I agree  r e f e r e n c e and copying of  shall  that copying  not  or  for  that  study.  this  thesis  by t h e Head o f my D e p a r t m e n t  is understood  f i n a n c i a l gain  The U n i v e r s i t y o f B r i t i s h  Date  of  for extensive  permission.  Department of  f u l f i l m e n t of  it freely available  representatives.  this  in p a r t j a l  or  publication  be a l l o w e d w i t h o u t  my  i  Abstract  This study i s an i n v e s t i g a t i o n of the hypothesis b i r d species'  abundances and  other  that  c h a r a c t e r i s t i c s of b i r d  communities i n urban areas are a f u n c t i o n o f h a b i t a t s t r u c ture.  To t h i s end,  hectares  e i g h t study p l o t s of approximately  each were s e l e c t e d i n Vancouver, B.C.,  i n terms of area covered by b u i l d i n g s , and  other man-made features o f the h a b i t a t .  estimated.  the p l o t s d e p i c t e d was  The  by  range o f h a b i t a t s represented  by  a gradient  of urbanization;  one  f o r one  diversity (H ), 1  the number of f o r a g i n g  these  breeding seasons.  The  i n f l u e n c e d by  number and  spring migration  and  d i v e r s i t y of b i r d  and veg-  summer  species  the d i v e r s i t y of man-made s t r u c t u r e s .  In most p l o t s , b i r d species than i n winter.  the number of s p e c i e s ,  g u i l d s decreased with decreasing  e t a t i o n cover, e s p e c i a l l y during  were not  a wood-  year.  B i r d species  i s not  extreme  the other was  B i r d s were counted at r e g u l a r i n t e r v a l s on  plots  In  for birds provided  a downtown commercial d i s t r i c t and  land.  described  d i f f e r e n t classes of f o l i a g e ,  a d d i t i o n , the amount of food intended people was  and  eight  d i v e r s i t y was  lower i n summer  Seasonal changes i n b i r d species d i v e r s i t y  r e l a t e d to u r b a n i z a t i o n .  ber o f species were greater Evenness of s p e c i e s '  However, changes i n num-  i n h a b i t a t s with more  abundances d i d not  vegetation.  change g r e a t l y with  season  i n most p l o t s , b u t  declined  noticeably  with  urbaniza-  tion,  i A  the  few  species  urban  Pigeon),  of birds  environment; Starling,  degree  of numerical  species  generally  Omnivorous habitats, ative were  House  were  the  i n a l l seasons. of  abundant  the  Rock  and  Dove  American  the  first  urbanization  most  abundant  T h e r e was  a  (Domestic  Robin.  three  with  insectivores  throughout  of  these  i n a l l seasons.  type  i n a l l urban  decrease  in winter  The  (many  i n the 0  f  rel-  these  migratory) .  to  residents food  the of  among p l o t s  amount o f  for birds,  winter.  response  to  response  hypothesized essential migrants.  their  total  Bird densities  w i n t e r , when  itat  and  the to  in total  vegetation  suburban p l o t s  n o u n c e d e f f e c t on  in  Sparrow,  d o m i n a n c e by  species  abundance  related  the  these were  increased  Differences  of  were most  density,  appeared  availability  of  that  factors  amount o f resident (e.g.  Migrants  are  to  a  are  habitat.  cues  have had  human  a  prothe  were h i g h e s t  t o be  food  by  believed  man,  available.  respond  greater  in  distributed in  provided  species  generally  c h a r a c t e r i s t i c s that s e l e c t i o n of  natural  bird  food)  food  not  amounts  especially in  i n most h a b i t a t s  some s p e c i e s  the  to  is  However,  substantial  i s thought  bird  density  cover.  provided  this  bird  It  is  directly  to  extent to  for these  be  others  than using  factors,  habin  i i i  TABLE  OF  CONTENTS  SECTION  PAGE  INTRODUCTION  1  METHODS  4  Plot  Selection  Methods Bird  4  of Analysis:  Habitat  Characteristics.  Counts  Methods  17  of Analysis:  Bird  Counts  Seasons Density  and  Questionnaire  Diversity  20  Analyses  21  Sent  to Plot  Residents  DISCUSSION  Characteristics Bird  Distribution Migratory  23 24  o f Study  Sites  Censuses  Resident  20 20  Statistical  R E S U L T S AND  13  24 36  and Abundance  53  species  54  Species  Domestic  60  Pigeon  61  Starling  61  House  64  Sparrow  Glaucous-winged House  Finch  American  Robin  Gull  64 65 65  iv  TABLE  OF  CONTENTS  (continued)  SECTION  PAGE Northwestern  Crow  Black-capped  Chickadee  Winter  ,..,  66 67  residents  67  Summary  68  Characteristics Diversity Total  of the B i r d  Communities  69  o f Species  Bird  Density  69  and T r o p h i c  Groups  72  SUMMARY  86  LITERATURE  CITED  ,'. 89  APPENDICES  92  1.  Sample  2.  a)  Questionnaire  - Letter  93  b)  Questionnaire  - Form  94  3.  Results Each  4.  of Bird  of Bird  Bird  Census  Mapsheet  Censuses:  92  Mean  Density  of  Species  Significant  95  C o r r e l a t i o n s between  Habitat  Features 5.  103  Statistical  Analyses  - Methods  and Results  a)  Results  of Multiple  Range  Tests  108  b)  Results  of Multiple  Regressions  113  V  LIST OF TABLES TABLE  PAGE  1.  A b b r e v i a t i o n s of P l o t Names and E n v i r o n mental C h a r a c t e r i s t i c s  15  2.  D e s c r i p t i o n of P l o t s : Features  25  3.  C o r r e l a t i o n s among P l o t s with Respect to H a b i t a t Features  26  4.  Groups of C o r r e l a t e d H a b i t a t C h a r a c t e r istics  27  5.  Summary D e s c r i p t i o n o f H a b i t a t  28  6.  Measures  30  7.  Q u e s t i o n n a i r e Results  8.  E s t i m a t i o n of Food Biomass Requirements  9. 10.  Cover by H a b i t a t  of H a b i t a t D i v e r s i t y  32  o f B i r d Communities i n Winter  35  L i s t o f B i r d Species  39  Seasons:  Dates of A r r i v a l and Departure  of Some Species  42  11.  Summary of B i r d Census Results  43  12. 13.  N u m e r i c a l l y Dominant B i r d Species Resident Status of B i r d Species i n Vancouver Comparison o f B i r d Community C h a r a c t e r i s t i c s with those o f Weber (1972)  45  14. 15.  T r o p h i c Groups o f B i r d s : P r o p o r t i o n of T o t a l Number o f I n d i v i d u a l s i n each . Group  55 76  77.  vi  LIST  OF F I G U R E S  FIGURE  TITLE  1.  Location  2.  A i r Photos  of Plots  3.  Mean T o t a l Vegetation  Bird Density Cover  4.  5.  6.  7.  8.  9.  10.  o f Study  Plots  8 versus  Total 37  a n d Number D o w n t o w n ..  47  Fluctuation of Total Density o f Species between censuses:  a n d Number False Creek  47  Fluctuation of Total Density o f S p e c i e s between censuses:  a n d Number MacKenzie  Fluctuation of Total Density o f S p e c i e s between censuses: Elizabeth  a n d Number Queen  Number o f M i g r a n t Vegetation Cover  Species  Total 56  versus  Estimates of Black-capped Chickadee s i t y versus T o t a l V e g e t a t i o n Cover versus  FHD2  12.  B S D v e r s u s Man-made D i v e r s i t y (Summer)  62 Den62  (Summer)  70  Component o f H a b i t a t  Number o f F o r a g i n g G u i l d s p e r P l o t s mer) versus T o t a l V e g e t a t i o n Cover Number versus  . 49  49  versus  Estimates o f S t a r l i n g Density T o t a l Vegetation Cover  BSD  14.  6  Fluctuation of Total Density o f S p e c i e s between Censuses:  11.  13.  PAGE  o f Species per Guild (Summer) T o t a l V e g e t a t i o n Cover  73 (Sum81  81  vii Acknowledgement  This sistance a  few.  study of  The  would not  a number o f p e r s o n s , criticisms,  of birds provided merit  my  Evans Advice  J. Dr.  Krebs  and  Lee  Gass  committee. for  their  and  Dr.  census tance  by  and  my  on  James  thesis  and  as-  mention  but  a i d i n the  s u p e r v i s o r , Dr.  census  W i l l i a m E.  Rees,  Dr.  Thanks counsel  Finally, helped the  analyses  Conrad Wehrhahn.  f o r the  Murray  i n c a r r y i n g out  bird  suggestions  was  provided  I thank they  Dr.  made a s  a l s o t o Thomas S u l l i v a n throughout  S p e i r s who I am the  I thank  this  provided  grateful  study,  the  that  were  efforts,  and  Dr.  Charles  J u d i t h Myers  and  my  Rudy B o o n s t r a  :  t o Wayne Weber advice  to E l a i n e Tasaka  on  the  f o r her  assis-  forms.  Canada W i l d l i f e  arose.  and  by  members o f  and  technical  questionnaire  f i n a n c e my  expenses  censuses  Hart.  statistical  in preparing  some o f  suggestions,  also helped  of birds.  arship  o f whom I s h a l l  the  deepest g r a t i t u d e .  T h o s e who Larry  have been p o s s i b l e w i t h o u t  my  S e r v i c e whose supervisor for  scholmeeting  INTRODUCTION  This density, birds, a  study  examines  and t r o p h i c  c h a r a c t e r i s t i c s o f communities  and the s t r u c t u r e  range  o f urban  bird  community  have  received  of  bird  age  height  of  foliage  habitat  intensive  (1961)  forest  examination  communities  i n ground,  in  areas  by  bird  that  of species,  abundances,  or the information  cities  i n many  than  other  that  d i v e r s i t y decreases  ance  and biomass  "natural" others loosely that  areas.  by b e i n g  1974).  be  City  that  largely  index  seed-eaters  of  1974).  while than  or cavity  species'  i s lower i n  and have  appear  or omhivores  bird  Emlen,  H',  of species  are tree  amounts  i n studies  the evenness  also  foli-  d i v e r s i t y , as  urbanization  avifaunas  1  F o r example,  1972 ;  in cities  (H )  by t h e  examined  species  theory  greater  composed  territorial,  are either  Emlen,  may  diversity  strata).  environments; with  and  on t h e r e l a t i v e  (.Weber,  bird  complexity  diversity, total  been  America  shown  t h e number  and tree  within  between  MacArthur  species  based  have  structural  be p r e d i c t e d  species  types  o f North have  index  environment  since  bird  could  shrub,  between  and h a b i t a t  studies  that  of  The r e l a t i o n s h i p s  and h a b i t a t  reported  density,  indexed  types.  d i v e r s i t y Can  relationships  urban  o f the p h y s i c a l  complexity  MacArthur  These  r e l a t i o n s h i p s between d i v e r s i t y ,  suggested bird  abund-  i n rural  to d i f f e r that  or from  are  only  nesters,  and  (Weber,  1972;  This  study  underlying ific 1)  such  research To a  3)  and of  of  undertake  of  species,  To  in  supply,  total  ities  within  readily itats,  year-long  study the  One  they  range  are  reason  information of  this  in  cities  in  some  that  for  count  often  factors  area.  Spec-  using  qualitatively  structural  to  of  estimate and  the  seasonal  of  fluctua-  food  provided  for  birds  by  the  between  d i s t r i b u t i o n of  habitat,  artificial  the  communities,  bird  food and  distributions.  and of  the  hypothesis  trophic urban  that  structure  habitat  of  types  species bird  are  i n non-urban studying be  The  situations health  commun-  the  to hab-  areas.  birds  applied  in to  abundance  economic  diver-  related  structural characteristics of  causes  number  plots.  correlations  could  "wildlife".  urban  the  follows.  types,  study  examines  quantifiable as  of  areas.  species'  density, a  an  abundances,  amount  features  thesis  some  q u a n t i t a t i v e l y and  c h a r a c t e r i s t i c s of  individual  sity,  as  habitat  within  the  structural  examine  habitat.  the  determine  This  a  were  species'  estimate  To  urban  these,  in  to  describe  the  To  people 4)  objectives  gradient  tion  designed  relationships within  identify  features 2)  is  and  hazards.  cities  is  ecological  of  to  obtain  management  certain bird  species  aesthetic  damage,  Birds  act  may  as  and vec-  3,  tors  of  such  Moreover  diseases  large  urban  v u l g a r i s ) , Domestic and  House  crops,  Sparrows  stored  (Princ.  of  of which  Murton and  et  may  the  but  at  Doves  domesticus)  the  buildings  avian  pest  control  the  Successes  of  time  in  habitat  regard.  have  that  been  nearby  they  roost  study  cities  and  favour  however,  habitat be  a  an  desirable  would  focuses  may  1964;  therefore,  pests,  This  attempted,  limited,  would  structure  been  (Elliott,  Clearly  discourage  f o r man. birds  have  livia) ,  1970).  poisoning  costs.  technique  same  between  or  damage  i n which  of  trapping  (Sturnus  C£olumb<i  can  cont.,  considerable  this  Rock  ^  e  encephalitis.  Starlings  pest  environment  in  s  or  of  anim.  manipulation  technique  s  and  management  relationships since  a  involved  involve  urban  CP  a l . , 1972).  ecological species  populations  and  Many m e t h o d s most  c o c c i d i o s i s and  Pigeons  food,  plant  as  on  improve the  structure, useful  METHODS  Plot Plots structure ization refers  representing habitats  used  b y man-made  and  pavement  character  that  with  related  with on  with  were  from  the City survey.  have  imity  o f major  was  using  Choice  composition  in  c o u l d be  i n physical  I  of bird  gradients  the former  cover.  considered travelling  environmental I preferred were  Department,  of plots  o f tree  t o be spent  the ocean.  "urban"  assoccor-  characteris-  1972 a i r p h o t o s  Planning  amount  of plots  building  sampled.  selected  the apparent  urban-  vegetation.  the environmental  changes  of  by v e g e t a t i o n  much  t o be more more  physical  of urbanization"  covered  and n u m e r i c a l  reflect  levels  Habitats with  relatively  quantifiable  would  and  o f area  u r b a n i z a t i o n , and that  a field  location  amounts  the species  o f the areas  obtained  different  The " l e v e l  are considered  those  would  with  Plots  study.  structures.  cover  than  communities  tics  i n this  to the r e l a t i v e  and  iated  with  and r e p r e s e n t a t i v e o f d i f f e r e n t  were  expected  Selection  within,  and that  effects  and the e f f e c t s  i n conjunction  was p r i m a r i l y  i n relation to these,  surrounded  that  were  by s i m i l a r  of internal  based  In addition, the t o time  which  and the prox-  discontinuities, areas  o f Vancouver,  such  as  parks  homogeneous habitat.  variation  Edge  of habitat  5.  were  thereby  level  minimized.  terrain.  streets,  or  Areas  near  the  Plots  without edge  of  were back  the  a l l located lanes,  on  with  a i r photos  fairly  curving  were  not  sel-  ected. Eight  plots  major  habitat  cial"  (Downtown,  "densely  types  (.Figures  "light  UB)  and  habitats.  (West  were  (2 0  ies  provides  compromise  which less  have  proportionately  daily  breeding  fluctuation  seasons)  homogeneous. by  Speirs  Plots  et  al.  some  authorities  used  (eg.  urban  plots  1973). more in  A.  of  eight  manpower.  acres)  reduced the  WE),  about  in  area.  edge  large  plots  might  bird  studies.  less not 20h  CWS,  than  personal 0. 5h  possible plots  in  have this  regularly  3  and  have  times  be  been  study  to  because  used  However  larger  been  than  count  those  while  censused  of  non-  more  communication), also  stud-  probably  (especially in  (.1970) i n u r b a n  or  and urban  size  2  Collingwood,  For  same  areas  MK;  effect  which  FC),  "single-dwelling  the  advise  Creek,  mixed-forest"  between  plots  "commer-  rectangular  avifauna  smaller  of  Erskine,  I t was  than  and  in  were  (False  "second-growth  A l l plots  a  represent  MacKenzie,  8 hectares  size  These  End,  QE;  approximately this  2). t o  industrial"  apartment"  SH),  1,  Vancouver.  (.Queen E l i z a b e t h ,  Shaughnessy,  (U.B.C,  chosen  within  DT),  populated  residential" CW;  were  (.Thomas,  birds  in  limitations  6.  Figure  1:  Location  of  study  plots  within  Vancouver,  B.C.  8.  Figure 2:  A i r photos of- study p l o t s . lcm  = 50 m.  Scale approximately  These photos are s m a l l e r s c a l e  copies  of 19 72 a i r photo enlargements, o b t a i n e d from the C i t y P l a n n i n g Department.  9. Figure  2:  A i r Photos  False  Creek  of  Plots  Queen  Elizabeth  Figure  2:  A i r Photos  Col1ingwood  of  Plots  U.B.C.  13.  Methods It  was n e c e s s a r y  physical vided was  of Analysis:  properties  by major  measured  ments,  laneous  paved  shrubs,  and weedy  further  broadleaved height 7.5m  vegetation  patches  1).  grass,  evergreen,  and were  assigned  0 t o 1.5m  and over over  strata  choices  made Weber,  observations  4.5m  7.5m.  1.5  Trees  i n height.  and h a b i t a t  i n related 1972;  (5'),  studies  Emlen,  of layering  coniferous,  t o one o f  (MacArthur  1974),  b u t was  of vegetation  4.5 t o  as woody  o f these  partially  or  four  (15'),  defined  The c h o i c e  features  herb-  categories  t o 4.5m  were  trees,  natural  The woody  deciduous,  (^apart-  and m i s c e l -  lawns, g a r d e n s ,  (e.g.t a l l  (see Table  structures  buildings  sidewalks  as b e i n g  etation  1961;  surfaces,  pro-  buildings  slant-roofed  lanes,  the  cover  a n d man-made  described  classes:  (25'),  roads,  and g r a v e l  vegetation)  Therefore  of flat-roofed  industrial),  residential),  were  plot.  that  Characteristics  and c h a r a c t e r i z e  of vegetation  including  (mainly  aceous  to inventory  o f each  types  commercial,  Habitat  veg-  reflects  the  and MacArthur, also  based  on  i n urban  areas  of  Vancouver. An  a i r photo  plots;  these  erable  detail  for each  each  were was  enlargement  was  o f the scale discernable.  o f the urban  o f the features  plots,  obtained  1cm = 24m These  mentioned  so t h a t  were  the percent  f o r each  used  o f area  above, w i t h  o f the  consid-  to  determine,  covered  by  the exclusion  of  shrubs, gardens,  and weedy patches.  The l a t t e r were o f t e n  i n d i s t i n g u i s h a b l e from and grouped with "lawn". A t r a n s p a r e n t g r i d o f t r a n s e c t l i n e s spaced  1cm apart  was p l a c e d over the p l o t area on each photograph,  and the  length o f these l i n e s i n t e r s e c t e d by each h a b i t a t f e a t u r e was measured.  The percent o f p l o t area covered by each  ture was then c a l c u l a t e d .  fea-  This method was s u b j e c t to e r r o r s  p a r t i c u l a r l y when e s t i m a t i n g the amount o f area covered by f e a t u r e s which were r e g u l a r l y d i s t r i b u t e d such as roads, l a n e s , and sidewalks.  Therefore the amount o f these  tures was estimated d i r e c t l y  from the a i r photos.  fea-  This more  accurate measurement o f the amount of area covered by road, l a n e , and sidewalk i n each p l o t was then s u b s t i t u t e d f o r the estimates d e r i v e d from the t r a n s e c t  sampling.  I v i s i t e d each o f the p l o t s i n order to estimate of t r e e s , p r o p o r t i o n o f t r e e s and shrubs f o l i a g e c l a s s , and area covered by shrubs  densit  i n each h e i g h t and and garden.  This  ' o n - s i t e ' survey covered at l e a s t 1/4 o f each o f the urban plots  (Table 2).  Since I had chosen  the p l o t s to be as i n -  t e r n a l l y homogeneous as p o s s i b l e , the areas sampled were s e l e c t e d s u b j e c t i v e l y .  'on-site  During these surveys, I paced  along s t r e e t s and lanes to measure one dimension  of features  such as hedges, and estimated the other dimensions  by eye.  Heights o f t r e e s and shrubs were i n i t i a l l y measured u s i n g a Suunto Clinometer, but I gained confidence i n e s t i m a t i n g  15.  Table  1:  Abbreviations of Plot Characteristics  Plot  Names  Names  and E n v i r o n m e n t a l  Habitat  Type  DT  -  Downtown  Commercial,  Retail  FC  -  False  Industrial,  Wholesale  WE  - West E n d  Residential  -  QE  -  Residential  MK  -  MacKenzie  Residential  CW  - Collingwood  Residential  SH  -  Residential  UB  -  Queen  Creek  Elizabeth  Shaunghessy U.B.C.  Mixed  Woodland  Apartment  16. Table 1 (continued) Environmental  Characteristics  D  Deciduous  C  Coniferous shrubs  BDEVG  Broadleaved evergreen  trees  E  Broadleaved  shrubs  <7.5.  Less (15 ) 1  shrubs  than  or trees or trees  evergreen  7.5m ( 2 5 ' ) b u t g r e a t e r t h a n 4.5m  i n height  >7.5  Greater than  7.5m  <1.5  Less  >1.5  G r e a t e r t h a n 1.5m b u t l e s s  SLANT FLAT  S l a n t - r o o f e d b u i l d i n g s (houses, churches) F l a t - r o o f e d b u i l d i n g s (apartments, o f f i c e s , warehouses)  ROAD  Paved road s u r f a c e  LANE  Paved o r g r a v e l back lane s u r f a c e  PVT+S  M i s c e l l a n e o u s pavement p l u s s i d e w a l k  LAWN  Ground c o v e r e d b y s h o r t (<30cm) g r a s s  HERB  Gardens  WEED  Untended herbaceous  SHRUB  Woody p l a n t  less  t h a n 4.5m  TREE  Woody p l a n t  over  4.5m  FHD  F o l i a g e h e i g h t s l i v e r s i t y - i n f o r m a t i o n t h e o r y measure  HFD  Habitat feature diversity-  TD  Tree  FOOD  Amount o f f o o d p r o v i d e d i n t e n t i o n a l l y b y r e s i d e n t s (kg/month/40h)  TOTVEG  Total  t h a n 1.5m ( 5 ' )  (herbaceous  t h a n 4.5m i n h e i g h t  p l a n t s p l u s bare  primarily earth)  vegetation  "  "  "  d e n s i t y (#/40h)  c o v e r by v e g e t a t i o n  17.  heights  by eye and t h e r e f o r e used  The cover the in  'on-site'  values  area  imated  Lands ing  from  was  the woodland  forest,  tape,  strip  visits  a plot  with  lines  vegetation quadrats,  roads cover  each  throughout  was  v e g e t a t i o n i n each  of  trees p e r quadrat.  located  laid  similar Emlen also  technique  made  used  techniques.  three  approaches  at  a "best  est-  were  and measur-  30m.  A  buffer  the p l o t  A l l estimates 'on-site'.  located  of  Fifteen  approximately  I estimated  the cover t h e number  Counts  strip  birds  count"  count"  i n this  method  i n an urban  transect  study  favoured  situation.  a n d "home  adjusting the results  t o account  Endowment  by eye, and counted  to count  After  estimate"  o f "lawn"  compass  areas.  In these,  stratum  f o r censusing  ping"  and weeds,  separated  subjectively  plot.  used  "converted  that  out using  12m s q u a r e  to the "direct  (1974)  shrubs,  i n t h e U.B.C.  60m w i d e  Bird The  transects,i . e .  c u t and f l a g g e d every  t h e census  to adjust the  method.  plot,  were  of  used  s u b t r a c t e d from  and r e s i d e n t i a l  12m x  were  by gardens,  of vegetation at least nearby  method.  the a i r photo  t o be c o v e r e d  by t h e t r a n s e c t  In  results  d e r i v e d from  found  the site  survey  this  by Emlen  range  map-  o f these  f o r known b i a s e s , h e  o f t h e d e n s i t y o f each  was  arrived  species.  How-  18.  ever  the r e s u l t s  mated  h i s best  appears  estimates.  t o be t h e b e s t  bird  communities  many  plots  here  have  Weber  method  survey  roads  r o u t e was  were  ersed  (Appendix  close  together to ensure  plots  a plot  was  was  census  2 1/2  census  The  species  of was  after  lines  similar  therefore urban  counts  t o t h e one  and Graber  plots  and  used  (JL963) , a n d  reduced  within  this  were  i n the plot  noted  species.  were  of birds,  trav-  coverage o f through  and the time 1/2  was  hour  occasional  or foraging  walked  pauses. fledged  r e c o r d e d on  census  particularly  for-  as i n d i c a t o r s  o f t h e u s e made birds  when i t  of impaired observability The " d i r e c t  to  done t o  The o b s e r v e r  I d i d n o t census  because  of birds.  This  with  of a l l perched  activities  days,  period. pace  lanes  sufficiently  the f i r s t  varied.  at a leisurely  o r windy  activity  was  were  were  o f passage  census  within  apart  visual  The d i r e c t i o n  i n which  o f the woodland  60m  complete  to sunrise,  Certain  plot,  and roads  on a l t e r n a t e  observed  by b i r d  raining  that  sunrise,  and n e s t i n g ,  f o r each  spaced  The l a n e s  and l o c a t i o n  birds  mapsheets. aging  approxi-  f o r censusing  or i n the case lines  obtained.  the counts  the  adult  1).  reversed  hours  defined  transect  relative  randomize  or  count  oft-repeated  by G r a b e r  traversed,  i n which  of  Methods  used  plots,  the  when  closely  strip  available  particularly  been  count  The d i r e c t  are involved. also  strip  (19 72) . A  and  o f the d i r e c t  strip  count"  and  method  is  reasonably  study not  quick,  i n areas  highly Four  where  applicable many  species  persons  helped  with  of  total  80%  may  in different  biased  throughout  the  observers. servers  The  successive  in  one  woody  by  an  plot  Time  season  but  about  opinion that  this  study  gave  populations very  in  the  er  degree  owing tion.  i n urban  rare.  woodland  to  the the  plot  plots,  and  may of  was  the  have  way  been  individuals of  low  generally  the  between  separate  the  survey  more  i n the  results  two  from  plots  time  urban  used  to  of  the  f o r those  was  varied  bird  species  species  birds species which  to  Counts a  great-  present,  i n dense  vegeta-  of v e g e t a t i o n i n urban  excellent.  with  plots.  count  underestimated  more  re-  confirming this.  and  ob-  G e n e r a l l y , the  detecting birds levels  a l l plots  in a plot  estimate  of  did  compared.  to  methods  I  observers  overlap  when  birds  except  i s no  difficulty of  the  Since  done.  minutes  but  censused  were  urban)  reasonable  There  number  Because  visibility  a  20%,  count  45  i s my  of  often  to  It  were  for  flocking,  d e n s i t y by  area  less  for counts.  I  observer  was  quired  in  suited  surveys,  provide  total  f o r one  ( i . e . the  averaged  of  to  same  this  bird  censuses.  average  i n the  and  the  i n order  possible  morning, the  loosely  directions,  estimates  counts  was  the  study  differed  It  are  and  territorial.  approximately be  year-round,  areas,  20.  Methods Several ties  of Analysis:  analyses  and h a b i t a t s .  were  These  Bird  employed  Censuses  t o compare  are b r i e f l y  bird  described  communi-  below.  SEASONS The based  census  results  on b e h a v i o u r a l  trends  and to reduce  While  I refer  and  Winter,  seasons. tion The  Spring  while dates  species set  number  be  followed total  were  i n the census Summer,  times  migra-  between  o f a number  census  Fall,  calendar  of bird  the times  results  results.  these.  of transient  and used  to  DIVERSITY are generally  p e r 40 h e c t a r e s here.  bird  reported  (100 a c r e s ) ;  T h e mean  community  species  interest.  where  peak  urbaniza-  density  within  i n the  this  o f each  each  plot  literature  convention species  (.for e a c h  will  and o f season)  calculated. Total  of  "seasons"  seasonal  precisely with  were  from  to detect  as S p r i n g ,  and departure  determined  densities  as  the  and W i n t e r  four  be a t t r i b u t e d t o  variance  were  into  lengths.  D E N S I T Y AND Bird  might  "seasons"  and F a l l  of arrival  season  i n order  do n o t c o i n c i d e  Summer  were  that  possible  t o these  they  categorized  criteria  i n c l u d i n g those  tion,  were  diversity  The i n f o r m a t i o n  p i i s the proportion  i s another theory  o f .the  i  th  community  measure  H'  species)  attribute  (j=" 2 P j ^ ° § e P i > has been  most  frequently of  used  comparison.  certainty an  The  synonymous  of  the  J'  species  and  as  ;  bird  this  ( J  1  of  amount to  of  un-  species  community  diversity  (BSD)  of  (MacArthur,  and  H'  are  paper. to  or  bird  number  , range of  0  each  a l l species  H'  to:  H ,  S,  1  to  species  diversity.  of species  CS)•  1), which  species, become  refer  respectively,  STATISTICAL  a  x  =  log  e  S  is a  and  The measure  approaches  equal.  (Pielou,  Habitat  diversity  or  foliage  and  These  that  indices  SAV  FHD  to  diversity  J ' , and  1961),  1974).  and  m  community  MacArthur,  A  species  a  purposes  It is  1969).  max  calculated;  H*,  s e l e c t e d from  richness  in a plot).  (Tomoff,  as  for  as:  ^  referred  here  identification  proportions  = _H1  Four  used  the  components  proportions  be  the  evenness  relative  calculated  been  two  will  i n t e r p r e t e d as  bird  species  i s the  the as  terms  so  be  randomly  are  i s the  other  may  throughout  There One  H'  surrounding  individual  1965) .  1  for this  the have  HFD  were  or  (the average  height  diversity  habitat  strata  recognized  number  i n d i c e s have  feature  c a l c u l a t e d i n the  foliage been  in a l l will  and by  be  of also  (MacArthur diversity  same  habitat  manner features  the  observer.  r e l a t i o n s h i p s between*habitat  structure  ANALYSES  number  of  and  c h a r a c t e r i s t i c s o f b i r d communities were examined  statis-  tically.  tech-  A n a l y s i s of v a r i a n c e  and m u l t i p l e range  niques were used to examine v a r i a t i o n between p l o t s between seasons with respect species  to the t o t a l b i r d d e n s i t y ,  d i v e r s i t y (H'), number o f s p e c i e s , and  of s p e c i e s '  abundances.  and  the  M u l t i p l e r e g r e s s i o n was  the  evenness  used to  derive " p r e d i c t i v e " equations f o r the abundance of each species  as w e l l as d i v e r s i t y , d e n s i t y , and  teristics features plexity  evenness  charac-  of b i r d communities, u s i n g area covered by and  habitat  d i v e r s i t y measures of h a b i t a t s t r u c t u r a l com-  (see Table 1 ) .  The variances  assumptions i m p l i c i t are homogeneous and  i n these techniques,  that  that the abundance data (dep-?  endent v a r i a b l e s ) have a normal d i s t r i b u t i o n , were v i o l a t e d i n many instances robust  by my  data.  these t e s t s are i n the  methods used i n c a r r y i n g out obtained  from them, w i l l be  than i n the  Since  i t i s not known  face o f these v i o l a t i o n s , the the t e s t s , and  described  text of t h i s t h e s i s .  the r e s u l t s  i n the Appendix  r e s u l t s are presented i n Appendix 5b.  with but  been used i n the  a few  sented i n the  references Appendix.  rather  M u l t i p l e range t e s t re-  s u l t s are p r e s e n t e d i n Appendix 5a and m u l t i p l e  approach has  how  regression  A nonstatistical  i n t e r p r e t a t i o n o f the r e s u l t s ,  to the s t a t i s t i c a l r e s u l t s pre-  23.  Questionnaire  Sent to P l o t  A mail-back q u e s t i o n n a i r e  Residents  (Appendix 2) was d e l i v e r e d  to r e s i d e n t s o f the urban p l o t s to gather  i n f o r m a t i o n on  the amount and types o f food p r o v i d e d by people  for birds.  This q u e s t i o n n a i r e was a l s o used to determine the s p e c i e s o f b i r d s which r e s i d e n t s n o t i c e d u s i n g t h i s food most f r e quently,  the frequency  o f b i r d s n e s t i n g i n b u i l d i n g s , and  the general a t t i t u d e s o f r e s i d e n t s towards urban a v i f a u n a . A q u e s t i o n n a i r e was d e l i v e r e d to every household i n the suburban p l o t s (Queen E l i z a b e t h , MacKenzie, Shaughnessy).  Collingwood,  One hundred more were d i s t r i b u t e d i n the  West End p l o t , p r i m a r i l y to r e s i d e n t s o f o l d houses and small apartments.  The high p o p u l a t i o n d e n s i t y and l i m i t e d  access to new apartment b u i l d i n g s made i t impossible to send a q u e s t i o n n a i r e to every household i n the West End.  R E S U L T S AND  DISCUSSION  Characteristics Each  plot  has been  ered  by v a r i o u s  from  Table  vegetation The  of  cover  theoretical  tree,  shrub,  maximum  cover  to other  similar  plots  habitat  features,  3.  , was Plots  (QE,  MK,  fell  Therefore  groups  Thus of  the gradient  highly  urban  some  In order other  These  features  with  groups  to  were  gradient similar just  with  how  ' r ' (Woolf,  are presented  at a =  total  t o a l l 19-  coefficient,  by v i r t u e  seen  175%,  a rough  respect  cov-  between  to determine  i n Table  of being  .01) w e r e :  highly  Downtown-  Elizabeth-MacKenzie-Collingwood  were  highly  features  cover  o f these  of urbanization  o f independently  been  (Table  into  4) a n d e a c h  plot  (Table 5).  described  habitat  and FC, h a d v e r y  (Appendix  placed  groups  c a n be  varying  (CW, S H ) .  correlated  have  correlated features  DT  respect  plots  results  significant  plots,  I t c a n be  5% t o o v e r  evident,  into  by t h e t o t a l  3 groups  than  Although  the habitat  of highly  described  with  and Collingwood-Shaughnessy  Many h a b i t a t 4).  2).  layers.  (DT, F C ) , Queen  CW),  less  the c o r r e l a t i o n  (r>.90,  Creek  (Table  o f the area  i s 3 0 0 % , due t o o v e r l a p  t o each  calculated.  correlated False  was  from  Sites  i n terms  a gradient  features.  were  which  was  varying  respect  1968)  features  there  and ground  vegetation  described  habitat  2 that  o f Study  i n terms  features.  little  tree  and  The  Table  2:  Description  of  Plots:  Cover  by Habitat  % Habitat  Cover Plot  Features*  DT  SLANT FLAT ROAD LANE PVT+S LAWN D<7. S D>7. 5 C<7. 5 C>7.5 DBEVG D<1.5 D>1.5 C<1. 5 C>1. 5 E<1. 5 E>1. 5 HERB WEED  Total  2 35 15 4 40 0 0 0 0 0 0 0 0 0 0 . 0 0. 0. 0  Cover  •7 1 0 8 2 9 8 1 0 0 0 0 0 1 0 1 0 3 3  ' FC 2 43 19 4 27 1 0 0 0 0 0 0 0 0 0 0. 0. 1. 0  2 4 7 1 1 9 2 5 0 0 0 0 0 1 0 0 1 1 4  '  WE 2 33 13 8 13 26 0 2 1 0 0 0 0 0 0 0. 0. 0. 0  QE  9 3 4 0 8 8 9 4 7 3 1 1 2 3 2 1 1 8 6  20 0 11 4 6 53 3 0 1 0 0 0 0 0 0 0. 0. 0. 0  0 0 7 6 8 3 5 6 2 2 0 0 3 4 6 2 1 6 0  MK 22 0 14 4 6 48 3 0 1 0 0 0 0 0 0 0. 0. 2. 0  1 0 6 2 7 3 7 0 0 4 0 0 4 1 3 3 1 2 5  100. 4  101. 3  106. 1  106. 3  106. 0  2. 6  4. 3  34. 6  62. 0  57. 3  Area sampled by ' O n - S i t e Method 1001  100%  .26%  27%  25%  Total Vegetation Cover  *For  meaning  Features  of  abbreviations  see  "  Table  1,  pg.  15.  22 0 8 5 7 43 8 8 2 1 0 0 0 0 0 0 0 3 0  115  CW  SH  9 0 8 1 7 8 2 3 6 1 0 0 5 3 3 3 4 3 4  14 4 0 5 10 4 3 0 8 2 51 1 10 9 8 5 3 3 2 6 • 1 6 0 6 0 3 0 3 0 3 0 3 1 0 2 6 0 8  3  121  UB 0. 0 0 0 0. 0 0. 0 0. 0 0 .0 12. 2 52. 1 4. 3 18. 4 0. 0 0. 0 29. 9 2. 6 1. 3 12. 2 6. 4 0. 0 31 . 9  7  187. 1  69. 5  84. 2  187. 1  .32%  33%  0.3%  26.  Table  3:  Correlations Features  among P l o t s  Correlation Plots DT  DT  FC  WE  .95  QE  with  Respect  to Habitat  Coefficient (r) MK  CW  SH  UB  .54  - . 31  - . 32  - .44  - . 40  - .46  .68  - .27  - .28  - .40  - . 36  -.42  .40  . 37  . 26  . 33  - .43  -  .99  .93  ,89  - . 27  MK  . 93  . 87  -.29.  CW  -  . 9.7  - .20  FC WE  -  QE  SH  -  UB  Significance  Level:  a=  .01;  r=  0.575  .09  27.  Table  Based  ally  Groups  of Correlated  on c o r r e l a t i o n s  indicated where  4:  significant a =  b y *, where  necessary  .05.  to indicate  Habitat  at a =  Groups  Characteristics  .01, e x c e p t  have  relationships  been  which  subdivided were  margin-  significant.  Group  1  a)  ROAD,  LANE  B)  PVT+S*,  FLAT*  Group  2  :  SLANT,  LAWN, HERB  Group  3  a)  D<7.5,  C<7.5  b)  D>7.5,  C > 7 . 5 , D<1.5, D>1.5, D<1.5,  E<1.5,  E>  see  where  Table  1, p g .  15  FOOD  FHD1, H F D 2 , FHD3, T O T V E G ,  D>1.5,  1.5, WEED, TDD, TDC  f o r meaning  of  HFD  abbreviations.  28.  Table  5:  Summary  Description  Total Groups  of Habitat  Percent of  Cover  Habitat  by  Features  *  Habitat 1(a)  1(b)  Commercial, Industrial. (DT, FC)  221  71%  Apartment (WE)  21%  Typical Residential ( Q E , MK)  2  3(a)  3(b)  5%  1%  1%  47%  31%  3%  4%  18%  7%  74%  5%  2%  Older Residential (CW)  14%  8%  70%  11%  12%  'Woody' Residential (SH)  13%  9%  68%  14%  15%  0%  0%  0%  17%  171%  Woodland (UB)  Based  on c o r r e l a t i o n s between  habitat  plots  are included  4) •  *  (see Table  Legend  1)  typically  urban  features  2)  typically  suburban  features  3)  typically  woodland  features  features  when  a l l 8  shrub  cover;  over  ered  by pavement  also  had a high  many h i g h - r i s e and  shrubs  tures for  former  by  cover  urban  was  trees,  even were  over  most  the area.  in  MK  and  features,  Area  covered  plots  20%  cover with  extensive  with trees fea-  accounted covered  a n d SH.  deciduous CW  30%  was  The  an  and  con-  o l d e r sub-^  features, while  cover  a n d SH,  percent  by  were  CW  by w o o d l a n d  i n QE  The h i g h e s t  10%.  about  plot  h a b i t a t s i n Vancouver  (shrubs,  under  cov-  suburban  gardens)  a n d MK  o f suburban  'woody'  by  these.  c o v e r i n g about  cover  by houses  50%  was  found  CW. habitat  in  terms  of  structures  structure  of the d i v e r s i t y  diversity  o f these  lawns,  features  was  T h e WE  and t y p i c a l l y  QE  being  plots  buildings.  f e a t u r e s , as were  typical  more  of  The  cover.  weeds)  with  7%  buildings,  the l a t t e r more  70%)  i n these  buildings.  about  by woodland  area  SH was  (about  apartment  i n WE  were  iferous  Lawns  cover  o f the t o t a l  primarily  o f the area  and f l a t - r o o f e d  (slant-roofed  30%  with  90%  (foliage  indices,  and v a r i o u s  Foliage  height  diversity  vegetation  plots  and the h i g h e s t plots  cover,  were  The  FHD  and h a b i t a t  i s shown  followed  the lowest  similar  described  o f v e g e t a t i o n and  i n woodland,  quite  also  relationship  features  (FHD) with  was  diversity,  6).  habitat  total  residential  of layers  height  Table  indices  o f the p l o t s  types feature  between  these  i n Appendix  4.  the g r a d i e n t o f FHD  in highly  apartment  with  respect  and  urban typical  t o FHD,  as  30.  Table  6:  Measures  of  Habitat  Diversity  Index Plot  FHD1  FHD2  FHD3  HFD  DT  0.12  0.14  0.91  1.77  FC  0 .18  0.21  1. 00  1.53  WE  0. 55  0.65  1.47  1. 88  QE  0.59  0.70  1. 34  1.67  MK  0 . 59  0. 69  1. 38  1.67  CW  0. 77  1.02  1. 66  1.97  SH  0. 79  1. 04  1.63  2 .12  UB  1.03  1.49  1. 49  2 . 04  Habitat p^ Description  Diversity  = -Ep.log  i s the proportion  of  p.  ;  of the i*"* habitat 1  feature.  Indices  FHD1:  3 layers.  trees,  shrubs,  FHD2:  6 layers. t a l l trees, short trees, s h r u b s , s h o r t s h r u b s , LAWN, g a r d e n s  FHD3:  9 layers. flat-roofed buildings, slantr o o f e d b u i l d i n g s , pavement, p l u s those i n FHD2 .  HFD:  19 l a y e r s . T a b l e 2.  A l l habitat  LAWN  § gardens  features  tall  as p e r  31.  were  old  ersity tial  and  was  woody  lowest  plots;  vegetation  food  cover)  provided  provided  this  order  obtain  food  provided  that  people  feed  birds  turns put  was  out  (DT,  the  were  of  study in  ment)  than The  been  a  was  commercial  biomass  roughly  i n woody  these  on  the  div-  residen-  (or  percent  the  the  of  Therefore plot  in  and  notes.  and  industrial  food  r e q u i r e d by  using  figures  food  amount was  small  area  assumed  (and  of  probably  very  within  from  is  the  of  areas that  i n each Emlen  the  outside  food  perhaps the  por-1  received  perhaps  source  birds  not  re-  this  plots  intentionally  did  questionnaire  u n i n t e n t i o n a l l y , and significant  of  assumption  a  is  which  assumed  industrial  It  of  In  amount  i t was  (WE)  since  households  field  of  7.  questionnaire  rate  not  amount  i n Table  This  commercial  one  questionnaire  extent.  degree,  a more  estimated  the  the  results,  apartment  the  provided of  7).  greater of  of  was  estimate  return  (Table  provided  that  feature  urbanization  summarized  since  f o r the  areas,  birds  to  i n the  based  food  are  from  case  l/12th)  Figures  FC)  amount  to  the  results  significant  high  for birds  forms.  The  people  any  very  highest  appearance,  conservative  in this  (about  habitat  failed  underestimated tion  a  who  Habitat  d i s t i n g u i s h i n g h a b i t a t s , but  man.  by  and  parallel  information  to  reasonable  industrial  of  to  by  plots.  gradient.  means  related  to  in  i t d i d not  Another directly  residential  for apart-  plots. plot  (1974)  has f°  r  Table  *  A l l  7:  Questionnaire  Results  Plot  • A # Forms Distributed  DT  0  -  FC  0  -  WE  100  20  11  QE  72  39  29  MK  146  50  CW  128  SH UB  amounts  of  B % Returned  C % of A that Feed B i r d s  •  D *Amount Fed i n  o f Food Winter  E. I of D which is not Fed at o t h e r Seasons  -  (estimated)  1  (estimated)  0  -  (estimated)  .1  (estimated)  0  40  0  81  12  36  345  8  55  33  87  8  40  55  38  .75  23  0  -  -  food  presented  as  kg/40h/month.  •  (estimated)  0  (estimated)  0  Table  7  (continued)  Number o f r e t u r n s i n as v i s i t o r s a t t h e i r  Which the feeders.  following  Species Plot  CP  S  WE  1  1  QE  1  MK  HS  HF  AR  6  0  15  13  4  40  CW  3  SI I Total  species  (see  Table  were  c i t e d by  residents  9)  GWG  NWC  BCC  3  1  1  0  3  0  0  2  7  7  1  5  2  0  0  39  6  22  12  14  7  4  3  3  19  20  1  17  1  5  13'  5  5  0  1  4  9  4  4  3  1  5  4  2  0  10  79  87  13  53  24  28  30  18  10  3  .  OJ  RSF  BB  the b i r d biomass/density r a t i o and the r a t e of food consumption  (Table 8).  These c a l c u l a t i o n s i n d i c a t e  that  s i n c e i n one case much more food i s p r o v i d e d than b i r d s are consuming, that b i r d biomass and consumption  have  l i k e l y been underestimated u s i n g these f i g u r e s .  Never-  t h e l e s s the food p r o v i d e d by man  is substantial in rela-  t i o n to the t o t a l amounts r e q u i r e d by b i r d s i n the suburban  plots. Those s p e c i e s which were most commonly fed by people  were sparrows  (mostly House Sparrows presumably;  domesticus), the S t a r l i n g Robin  (Sturnus v u l g a r i s ) ,  (Turdus m i g r a t o r i u s ) , Black-capped  a t r i c a p i l l u s ) , Dark-eyed western  Junco  American  Chickadee  (Table 7).  CP arus  (Junco h y e m a l i s ) , North-  Crow (Corvus c a u r i n u s ) , and Glaucous-winged  (Larus glaucescens)  Passer  In a l l r e s i d e n t i a l  Gull habitats  these b i r d s comprised over 50% o f the b i r d d e n s i t y yearround  (Table 12).  V i r t u a l l y a l l b i r d s p e c i e s encountered  i n the urban p l o t s were c i t e d by r e s i d e n t s as having used feeders, but I f e e l that except f o r some of those  listed  above, t h i s c o u l d not have c o n t r i b u t e d s i g n i f i c a n t l y to these s p e c i e s ' d i e t s except during times of food shortage. Other s p e c i e s were i n f r e q u e n t l y seen by r e s i d e n t s at t h e i r feeders.  35.  Table  8:  E s t i m a t i o n o f Food Biomass R e q u i r e m e n t s B i r d Communities i n Winter  Plot  *  Food x  .25  kg  Food  of  I o f Food R e q u i r e ments p r o v i d e d a t Feeders  Requirements  WE  191  16  QE  67  91  MK  168  158  CW  103  63  SH  162  35  Required  = kg of birds/40h  (.correction f a c t o r  of birds  x 0.25  f o r food  = X2To\irds  X  b  i  r  d  kg/day  x  30  days/mo,  waste)  d  e  n  s  i  t  y (from  Emlen,  1974)  36.  Bird A one  total  year,  imates these mean  the  total  525  (Table  relatively  and  influenced pusilla),  40h  number  although at  migratory  migration.  these Song  dates  Sparrow  Sparrow  lowest  year,  to  species  Wilson's  (4)  Dark-eyed  residential during of  which  these  of,  of  the  most  of  strongly  Qfi.%1sonia  Violet-green  Swallow  leucophrys).  were  Golden-crowned  Junco,  (Z o n o t r i c h ! a  the  because  Warbler  (Selasphorus  in  d e l i m i t dates  melodia),  t h a i a s s i n a ) , Barn  observed  industrial  3).  percent  Hummingbird  were  in  (Hirundo  r u f u s ) , and Large  either in Spring  or  White-  pulses Fall  . ,  of  (Appen-  4) . The  The  the  (Melospiza  satrapa),  Rufous  Those  were  of  i n woody  used  Est-  Estimates  Presence/absence  was  of  from  40h  was  the  course  resulting  encountered  of  behaviour.  transient species  Fall  species  times  4. per  species  seventy-five  certain  (Tachycineta  dix  bird  the  observation.  (Figure  were  swallow  these  130  species  9),  of  species  Sixty-two  (Regulus  crowned  from  apartment  of  during  i n Appendix  varied in  each  (10.5)  Kinglet  rustica),  hours  highest  abundant  Spring  250  made  and  rare  example,  most  per  habitat  11).  some  presented  density  average  industrial  to  were  densities of  are  bird  to  The  study  censuses  mean  censuses  (Table  282  amounting  of  habitat  for  of  Censuses  dates  lengths used  to  of  the  various  seasons  d e l i m i t seasons  were  are  given  compared  i n Table  with  10  publishe  37.  Figure 3:  Mean t o t a l b i r d d e n s i t y i n each p l o t versus t o t a l percent v e g e t a t i o n cover.  Only b i r d  d e n s i t i e s f o r Summer and Winter seasons have been graphed.  Table  9.:  List  of Bird  Species  No. C o d e Common Name Common  Scientific  Pigeon  Name  Columba  livia  Sturnus  vulgaris  1  CP  2  S  3  CM  Crested  4  HS  House  5  GWG  6  HF  House  7  AR  American  8  SSP  9  OJ  10  BCC  Black-capped  11  RST  12  NWC  13  BB  Brewer's  14  VT  Varied  Thrush  Ixoreus  15  RSF  Common  Flicker  Colaptes  16  VGS  Violet-green  17  GCK  Golden-crowned  18  WW  Winter  19  S'J  Steller's Jay  Cyanocitta  20  EG  Evening  Hesperiphona  21  RG  Ruffed  22  BW  Bewick's  23  HWK  unidentified  hawks  A c c i p i t e r spp.  24  RCX  Ruby-crowned  Kinglet  Regulus  Starling Mynah  Acridotheres  Sparrow  Passer  Glaucous-winged  Song  Gull  domesticus glaucescens  Carpodicus  Finch  Turdus  Robin  Sparrow  Dark-eyed  Larus  Junco  mexicanus  migratorius  Melospiza  (Oregon)  cristatellus  melodia  Junco  hyemalis  Chickadee  Parus  atricapillus  Rufous-sided  Towhee  Pipilo  erythropthalmus  Northwestern  Crow  Corvus  caurinus  Blackbird  Swallow Kinglet  Euphagus  cyanocephalus naevius auratus  Tachycineta Regulus  satrapa  Troglodytes  Wren  Grosbeak Grouse  Banasa  Wren  thalassina  troglodytes stelleri vespertina  umbellus  Thrysomanes  bewickii  calendula  40. Table  '9  (continued)  Code  Common  Name  Scientific  25  RH  Rufous  Hummingbird  Selasphorus  26  WCS  White-crowned  Sparrow  Zonotrichia  27  AW  Yellow-rumped  Warbler  Dendroica  28  BS  Barn  29  CBC  Njo.  Name rufus leucophrys  coronata  Hirundo.rustica  Swallow  Chesnut-backed  Chickadee  rufescens  Dendroica  townsendi  30  TW  Townsend's  31  AG  American  32  BHC  Brown-headed  33  GCS  Golden-crowned  34  WWA  Wilson's  35  MGW  MacGillivray's  36  YW  37  OCW  Orange- crowned  38  OSF  Olive-sided  39  WT  Western  40  WV  Warbling  41  PSK  Pine  42  SVS  Savannah  43  FS  Fox  44  FC  unidentified  flycatchers  Epidonax spp.  Grosbeak  Pheucticus  Yellow  Warbler  Parus  Goldfinch Cowbird Sparrow  Warbler Warbler  WarblerWarbler  Flycatcher  Tanager Vireo  Sparrow  Molothrus  ater  Zonotrichia Wilsonia  45  BHG  Black-headed  46  BTP  Band-tailed  47  ST  48  REV  49  V  Pigeon Thrush  Vireo  unidentified  vireos  atricapilla  pusilla  Oporornis  tolmiei  Dendroica  petechia  Vermivora  celata  Nuttallornis Piranga Vireo  gilvus pinus  Passerculus  Columba Catharus Vireo  borealis  ludoviciana  Passerella  Sparrow  Red-eyed  tristis  Spinus  Siskin  Swainson's  Spinus  sandwichensis iliaca  melanocephalus  fasciate ustulatus  olivaceus  Vireo spp.  41.  Table  9  (continued) Scientific  Common Name  No.  Code  50  DW  Downy W o o d p e c k e r  51  PF  Purple  52  WF  Western  53  GHO  Great  .  Finch  Name  Dendrocopos Carpodacus  Flycatcher  H o r n e d Owl  pubescens purpure.us  Epi-donax . d i f f i c i l i s ' . Bubo  virginianus  54  PM . P u r p l e  Martin  Progne  55  CN  Common  Ni'ghthawk  Cliordeiles'minor  56  PW.  Pileated  Woodpecker  subis  Dryocopus  pileatus  57  RBN  Red-breasted  Nuthatch  Sitta  59  W  unidentified  warblers  Dendroica spp.  60  NT  61  CWX  62  CS  Hermit Cedar  Thrush Waxwing  Common S n i p e  Reference:  canadensis  Catharus  guttatus  Bombycilla Capella  American O r n i t h o l o g i s t s '  cedrorum  gallinago  U n i o n , 1 9 5 7 . a n d 197,3.  T a b l e 10:  Seasons:  Dates o f A r r i v a l and D e p a r t u r e o f Some S p e c i e s  Spring  :  April  Summer  :  June 1 - August 2 5  Fall  :  August 26 - October 21  Winter  :  O c t o b e r 22 - A p r i l 10  Dark-eyed Junco  :  d i s a p p e a r e d from p l o t s between March 19 and A p r i l 8  Song Sparrow  :  d i s a p p e a r e d from p l o t s between A p r i l 10 and A p r i l 22  :  appeared i n p l o t s from May'/' t o May! 18  B a r n Swallow  :  appeared i n p l o t s from May 7 t o May 20  White - crowned Sparrow  :  appeared i n p l o t s from A p r i l  Warbler species  :  appeared i n p l o t s from A p r i l 20  Spring  11 - May 31  •  Migration  Violet-green  Swallow  20 t o May 10  d i s a p p e a r e d from p l o t s by May 30 Fall  Migration  Dark-eyed Junco  :  appeared i n r e s i d e n t i a l p l o t s between  Sept. 14 and O c t .  Song Sparrow  :  appeared i n r e s i d e n t i a l p l o t s between  Sept. 3 and O c t .  :  d i s a p p e a r e d by August 5  :  d i s a p p e a r e d by Sept. 13  :  d i s a p p e a r e d by Oct. 6.  :  appeared by August 29  Violet-green  Swallow  Barn Swallow White-crowned  Sparrow  Warbler species  d i s a p p e a r e d by O c t o b e r 21  43. Table 11:  Summary o f B i r d Census Results  SPRING  Plot  Mean Bird Cumulative Mean Total Bird Species Number of Number o f Species Species Evenness Density Diversity CH'D  DT  1.05  6  4.57  0. 59  390. 0  FC  1.25  7  4. 33  0. 64  140. 3  WE  1. 72  12  7.67  0. 69  583.2  QE  1.32  12  6.50  0. 73  178.3  MK  1.62  16  7.63  0.58  391. 0  CW  1.93  24  8. 56  0.61  366. 5  SH  2 .10  23  12.17  0.67  380. 3  UB  2.57  23  12. 00  0.82  260.4  DT  1.11  7  5.25  0.57  393. 8  FC  0. 89  8  3. 38  0.43  151.6  WE  1. 72  10  6. 50  0 . 75  QE  1. 51  14  5. 90  0.57  16 8. 4  MK  1. 30  13  6.25  0. 51  411. 5  CW  1. 74  13  6. 31  0.68  253. 7  SH  1.99  17  9. 09  0 . 70  396. 8  UB  2. 62  22  0.73  211. 5  SUMMER  10.11  - 468.8  44. Table  11  (continued)  FALL  Plot  Bird Cumulative Mean Mean S p e c i e s Number o f Number o f Total Bird D i v e r s i t y Species? Species Evenness Density c m  DT  0. 84  5  4.17  0. 52  435.0  FC  1.16  8  3.80  0.56  111. 3  WE  1.63  12  7. 40  0. 66  481. 4  QE  2. 09  18  7.80  0. 72  173.6  MK  1. 72  20  6.57  0. 57  329. 2  CW  2.20  21  8.83  0. 72  339. 2  SH  2.27  25  1 1 . 14  0 . 70  472. 3  UB  1. 88  17  8. 00  0. 66  434. 8  DT  1.00  5  4.03  0.62  400. 0  FC  1. 34  8  4. 36  0. 64  114. 8  WE  1. 74  12  7.62  0 . 70  567.9  QE  1. 74  12  6.67  0. 70  190.9  MK  1. 73  15  8.17  0.64  470 . 9  CW  1.92  17  7. 70  0.68  366,5  SH  2.15  19  1 0 . 00  0. 73  342. 2  UB  2. 03  18  8.67  0. 70  477.1  WINTER  45. Table  12:  Numerically  Dominant-Bird  Species  P r o p o r t i o n s o f t o t a l b i r d d e n s i t y f o r the t h r e e abundant s p e c i e s i n each p l o t  most  Season Plot DT  QE  Spring  CW  SH  .63 -.20 . 13 .96  CP S HS  . 64 . 16 .13 .93  CP S HS  .75 . 11 .09 .95  CP S HS  .66 . 15 .13 .94  CP S HS  .52 . 31 . 09 .92  CP S HS  . . . .  75 11 09 95  CP S HS  . 53 .33 . 08 . 94  CP S HS  . 50 .28 . 08 . 86  S CP HS  . . . .  CP HS S  . 31 . 24 .22 . 77  S CP HS  . 31 .27 .25 . 83  S CP HS  . 34 .25 . 19 . 78  S HS  .32 .23 .20 TT5  S HS  .49 .25 ^_12 .86  S HS  .35 .35 ^_12 .82  S .43 HS. .41 AR 07 .91  S HS AR  34 26 14 74  s . 37 AR . 18 WWA . 14 .69 S  . 36  AR . 20 WWA .12 .68  UB  See Table  Fall  CP S HS  AR  MK  Summer  — — Winter  WW .17 BCC .15 AR .13 .45  AR  JL  .38 .17 WWA ^09 .64  S .42 HS .32 AR ^_07 .81  S AR HS  . 35 S .21 HS .19 AR .75 •  . 34 .15 . 10 . 59  AR  .25 .25 . 13 .63  AR  .30 . 20 . 09 . 59  S HF  BCC .33 WW .15 RST .11 .59  S HS  BCC .40 WWA .20 GCK ^K> .70  S HS OJ  Mean Total  .44 .22 _^08 .74  .75  S .44 HS .26 AR 07_ .77  .82  :L  S  OJ  HS  S OJ  AR  .41 . 13 .13 .67 . 26 .21 . 14 .61  GCK .27 BCC .26 PSK ^L5 .68  9 f o r meaning o f s p e c i e s ' a b b r e v i a t i o n s .  .61  migration  dates  (Campbell  e t a l . , 1974) a n d g e n e r a l l y  I  that  believe  justified  with  Despite  edly  4,  5,  6,  these  that  each  for  different  plots  the  estimates  o f mean  ilar  plot,  habitat than  was  plot  the estimate  was  (Figures' 6 and 7). differences but  that  plots  may  counts.  in total  small  density  (Table  These  three  significant,  density  times  between  owing  d i f f e r e n c e s between  such from  a mean  value  were  similar  In other  cases,  n o t i c a b l y between  11, F i g u r e s  comparisons  density  11).  see  i s undoubt-  factors  values  greater  bird  i n the est-  It i s clear  of birds  times  d i f f e r e n c e s between  n o t be Small  differed  of total  approximately  (Table  there  ( f o r examples,  around  T h e mean  be  i n the b i r d  seasons,  variation  centered  three  habitat  season  involved.  cases  density  approximately  each area  may  ignored,  error, but other  seasons.  i n some  seasons  into  of this  are also  F o r example,  in industrial  Similarly, zie  within  habitats.  Some  into  data  i n each  area  i n accordance.  n o t be  within  fluctuation  for  found  is variation  o f census  to sampling  behaviour  Figures  there should  density  and 7 ) .  attributable  flocking  that  fluctuation bird  i n the Vancouver  o f the data  that  the d i v i s i o n  of total  Figures  as  season  considerable  imates  species  the d i v i s i o n  on t h e grounds  communities  was  f o r various  that  i n commercial on t h e a v e r a g e 4 and 5 ) . i n t h e MacKen-  i n Queen E l i z a b e t h  indicate that plots  large  are probably  estimated  means  o r seasons  real,  f o r some  to the v a r i a b i l i t y plots  sim-  between  will  hot  47.  Figure  4:  F l u c t u a t i o n between counts o f t o t a l density  a n d number o f s p e c i e s  i n t h e Downtown  Figure  5:  (commercial)  p e r sample plot.  F l u c t u a t i o n between counts o f t o t a l density  and number o f s p e c i e s  i n the False  Creek  bird  bird  p e r sample  (industrial)  plot-  48.  FIGURE 4  49.  F i g u r e 6:  Fluctuation  between counts o f t o t a l  bird  d e n s i t y and number o f s p e c i e s p e r s a m p l e i n the MacKenzie ( " t y p i c a l  residential")  plot.  Figure  7:  Fluctuation  between counts o f t o t a l  bird  d e n s i t y and number o f s p e c i e s p e r s a m p l e i n the  Queen E l i z a b e t h  plot.  ("typical  residential")  50. FIGURE 6  "j  "  1975  F~"  M '  A  M  J  . J  A  S  0  N  D  J  D.ATE  FIGURE  7  be  discussed.  overall, and  Analysis  there  seasons  were  which  and  ones  which  numbers ies  of  within  estimates The ted  given  of  as  It  the  was  (commercial, vegetation species  Number though areas the bers  the  of  rather  highest  than number  i n Summer,  Number  of  mercial,  of  different I believe  i n Table total  diversity  highest and  the  11.  spec that  Estima-  number  and  of  evenness  also  with  species and  are  (Table  with  and  11).  vegetation  woodland);  was  with  cover  the  thus  most bird  urbanization,  i n wooded  i n some  seasons.  found  residential In  in Spring,  most  similar  and  between  seasons  in  apartment  plots  than  plot  while  a l l were  fluctuated less and  lower  urbanization.  found  Winter  Winter  i n those  decreased  was  i n Spring,  least  highest  i n woodland  industrial,  similar  compared.  and  residential,  number  Fall,  species  and  decreased  of  tests  estimates  of  consistent,  Fall,  plots with  species  highest  was  Summer,  (woody  diversity  plots  season.  industrial) cover  Since  abundances  number  range  that  significantly  5a).  species  diversity  i n the  were  summarized  mean  indicated  Multiple  meaningfully  are  and  through  lowest  be  bird  plot  species same  can  as  seasons  reasonably  results  well  density.  relative  density,  f o r each Bird  much  these  results  d i f f e r e n c e s between  (Appendix  and  p l o t s were  total  species,  and  weren't  census  mean  total  plots  species  variance  significant  i n mean  indicated  of  num  lower. com-  i n areas  with  52.  greater vegetation The  evenness  slightly  vegetation density  sonal  d e n s i t y was  cover  o f b i r d s was  change  and  changes were n o t  A  bird  example,  pro-  and  wood-  amount  of  in Fall  approximately  T h e r e was  and  However s e a s o n a l industrial, (Table  11,  same  a general  sea-  suburban h a b i t a t throughout Starling,  are  and  three nest  the  first  (Weber, 1972). each p l o t  are  The  listed  d e n s i t i e s as i n each  three  and  res-  typical  Appendix  the  12  year:  the  American the  in  Robin.  ground,  or w i t h i n b u i l d i n g s  along  a p r o p o r t i o n o f the  5a).  dominant  s p e c i e s w h i c h were most  i n Table  highhave  f e e d p r i m a r i l y on  e x t e n s i v e l y on  and  changes may  House S p a r r o w , and  omnivores,  densities in  Fall,  s m a l l number o f s p e c i e s were n u m e r i c a l l y and  the  the  lowest  densities i n Spring  These s p e c i e s  with  abundant  estimates  mean t o t a l  bird  of  den-  plot.  In summary, t h e study  to t h e  density, with  i n commercial,  Domestic Pigeon,  sity  For  to be  (Queen E l i z a b e t h ) p l o t s  urban  their  related  woodland p l o t s .  d e n s i t i e s i n Winter.  idential  not  estimated  in total  been n e g l i g i b l e  in  decreased  older residential,  (urbanization).  Summer, i n t e r m e d i a t e  the  Seasonal  in industrial,  bird  commercial  est  s p e c i e s ' abundances  plots. Total  in  of b i r d  with urbanization.  nounced except land  cover.  p l o t s was  abundance o f e a c h b i r d  surveyed  f o r one  year.  species  on  Survey r e s u l t s  eight were  53.  divided and  into  plots  with  diversity, Trends  evenness,  may  the  be  a l l but  differences  total  bird  numbers  of  in bird  related of  the  to  and  plots  degree  dominated  and  respect  between  numbers i.e.  seasons,  to  woodland  Distribution Hilden birds of  (1965)  states  is primarily  d i f f e r e n t environmental  "settling onmental  response"; factors  generally  is  believed  by  such  cover,  species  birds  few  Abundance  habitat innate  selection  in  behaviour.  A  in habitat "sign  of  other  Hilden,  Release of  of  a  ultimately  preferred  may  be  settling  Thus  habitat  established  the  range  of  than  breeding summation  release  response  to  the  a  envir-  selection. are  of  It  often  is  general  landscape,  and  the  specific  habitat  determine  survival  (e.g.  food,  1965).  internal motivation  but  rather  in  stimuli"  presence  shelter;  of  plots.  s t r u c t u r a l c h a r a c t e r i s t i c s of  which  examined.  vegetation  or  characteristics  were  species  of  textural  animals,  bird  species  psychological  important  seasons  d i v e r s i t y and  stimuli results  thus  that  A  and  that  governed  species  amount  urbanization. the  density,  species  the  between  in  to is  breed. in  on  the  ability  of  preferred  short  suboptimal  habitats  depends  response  specificity and  If  this  supply, habitat  i n which of  also  a  habitat  depends  on  the  motivation then  a  may  1949).  be  s e l e c t i o n , the  alternative habitats,  and  high  territory  (Svardson,  species  is  level  found avail-  internal  hormonal c o n d i t i o n s . the  The hypothesis examined here  importance o f s p e c i f i c i t y  stresses  i n h a b i t a t s e l e c t i o n , and  p r e d i c t s that d i f f e r e n c e s i n the s t r u c t u r e o f urban h a b i t a t s w i l l a f f e c t the d i s t r i b u t i o n and abundance o f b i r d s . The  b i r d s observed can be d i v i d e d i n t o three  based on t h e i r " r e s i d e n c e "  status and migratory  Throughout the year a l a r g e p r o p o r t i o n  groups  patterns.  o f the i n d i v i d u a l s  i n each p l o t were r e s i d e n t b i r d s that breed l o c a l l y . Spring,  During  Summer, and F a l l , migrants were a l s o present.  migrant species were t r a n s i e n t s ; out the Summer to breed.  Most  they d i d not remain through-  During F a l l , Winter, and S p r i n g ,  a group which I s h a l l r e f e r to as Winter r e s i d e n t s was present .  MIGRATORY SPECIES Migratory  species  are l i s t e d  i n Table 13.  Most o f . ,  these were found only i n p l o t s with high v e g e t a t i o n (Figure  8). These species  (with the exception  were probably s e l e c t i n g general the environment.  cover  o f swallows)  foliage characteristics of  Total vegetation  cover and f o l i a g e height  d i v e r s i t y were h i g h l y c o r r e l a t e d with the abundances o f many o f these s p e c i e s  (Appendix 5b).  However, the number  of migrants i n the woodland p l o t was lower than expected, e s p e c i a l l y i n F a l l , though t h i s may have been a r e s u l t o f the d i f f i c u l t y o f adequately sampling t h i s  habitat-  55. Table 13:  Resident S t a t u s o f B i r d  S p e c i e s p r e s e n t year-round  Transient Species (except  (resident)  f o r rare winter occurrences)  Common Pigeon Starling  S p e c i e s i n Vancouver  Barn Swallow  (Spring,  Fall)  Violet-green  Swallow  House Sparrow  Savannah Sparrow  C r e s t e d Mynah  Golden-crowned  Glaucous-winged American House  Robin  Gull  Brown-headed  (Spring,  Fall)  Rufous  Finch  Band-tailed Swainson's  Dark-eyed  Hermit  Black-capped  Chickadee  Pigeon Thrush  Thrush  Red-eyed V i r e o  Northwestern Crow V a r i e d Thrush  Cowbird  Hummingbird  Song Sparrow Junco  Sparrow  Warbling  (Spring,  Fall)  Vireo  Orange-crowned  Warbler  S t e l l e r ' s Jay  Wilson's  Evening  Townsend's W a r b l e r  Grosbeak  Brewer's  Blackbird  Warbler  Yellow-rumped  Common F l i c k e r  (Spring,  Golden-crowned  Kinglet  Fall)  Warbler  MacGillivray's Western  Flycatcher  W i n t e r Wren  Olive-sided  Ruffed  Cedar Waxwing  Grouse  Bewick's  Wren  Western  Ruby-crowned K i n g l e t White-crowned American Pine  Goldfinch  (Spring,  Siskin  Grosbeak  F a l l ) Common Nighthawk  Fall)  Purple  Finch  Common Snipe  Fox Sparrow ( S p r i n g , Chesnut-backed  Flycatcher  Tanager  Black-headed  Sparrow ( S p r i n g ,  Warbler  Fall)  Chickadee  Downy Woodpecker  (Seasons i n p a r e n t h e s e s  Great Horned Owl  t h a t some p o p u l a t i o n s a r e m i g r a t o r y )  indicate  P i l e a t e d Woodpecker :  Busht i t Red-breasted Nuthatch R u f o u s - s i d e d Towhee  (Fall)  from Campbell  et. a l . ,  1974.  56.  Figure  8:  Number o f m i g r a n t s p e c i e s p e r p l o t t o t a l percent  vegetation cover,  seasons o f peak  migration.  versus  during  FIGURE  20 16  40  80  Total Vegetation  120  Cover (%)  160  8  58.  Wilson's in  Spring  was  Warbler  and F a l l ,  common  i n some  was b y  f a r t h e most  but d i d not stay residential  to breed.  plots  during  ecially  i n Shaughnessy  and C o l l i n g w o o d .  related  with  o f deciduous  aging that  mainly  i n vegetation  t h e amount  stimulus", in  the cover  o f deciduous  so t h a t  Wilson's  selection of habitat  non-foliage-related does  n o t appear  distribution green  significantly since  these Hilden  may  respond  rather  habitat. certain were  states  species  that  social  factors  Smith  (1975)  discusses  and r e s i d e n t  contact  with  join these  as a  to this  foliage with  environment  a f f e c t i n g the Barn  construction  highly  of other  social  and V i o l e t which are of buildings,  species  individuals  configurations,  and  when  examined  whether  i n habitat  by r e s p o n d i n g  species  between  some m i g r a n t s  .  selection.  i n the tropics.  flocks  of birds  selecting  the r e l a t i o n s h i p between  birds  foraging  for-  "sign  Interaction  common m i g r a n t s  to assess  using  migrants  factor  c o r r e l a t i o n s were  i n order  seen  i n nesting.  to environmental  Therefore  passerines that  u s e them  to the presence  than  serves  Indeed,  b y man's  cor-  It i s possible  migration.  Warbler.  esp-  highly  a n d was  responds  a significant  influenced  (1965)  foliage  Warbler  be t h e o n l y  species  type.  species  migration,  I t was  trees,  migrant  This  c h a r a c t e r i s t i c s o f the urban  t o be  may  tree  during  o f Wilson's  Swallows  of this  abundant  of residents  to a noise  He  migrant  observed  and keep i n  resembling  59.  "spshing." attract local  This  birds  sound  into  Golden-crowned  and  Bushtit).  related  flocks  speci.es w i t h send's  present  were  are the Black-capped (possibly  a n d Town-  were  quite highly as w e l l social  i n habitat selection sample.  Perhaps  general  to this  to "spshing",  i s locally  directly  advantage  may  (and/or  interpretation  areas then  (1975)  cue on f l o c k s  of  predators).  of the migrants' respecies  know  where  i f the migrants'.diets are similar  of the residents, following  the migrants  the r e s i d e n t s .  resident species  i n response  Smith  in  of the resident's  the resident nucleus  b e n e f i t from whether  o f food.  i n the tropics  abundant;  to those  i n temperate  features of the landscape,  i n search  may  among some m i g r a n t s  migrants  f o o d resources  According  with.the  and n o n s o c i a l f a c t o r s  of local  uted  correlated  these  as w i t h v a r i o u s h a b i t a t  knowledge  then- a r i s e s  species  Wilson's  to take  therefore  Mixed  cor-  Indeed,  that migrants  i n . some way  highly  various warblers.  i n order  food  were  Junco  of  residents  sponse  Chickadee  also Dark-eyed  and k i n g l e t  of  composed  resident birds  reasoned  the nuclei  observed  recognize  follow  form  during m i g r a t i o n seasons.  so t h a t both  important  first  Kinglet  and k i n g l e t ,  features;  the  flocks  that  to  sometimes  Warblers  chickadee  Species  The c h i c k a d e e  (r>.90)  foraging  be  view.  mixed-foraging  and  i s o f t e n used by.birdwatchers  to local  i n general  availability  The  can question  arc distribo f food or  60.  other  essential  sponding cues  to features  which  shall  more  hypothesize  i n response  migratory  birds,  the  following  discussion  ed  along  in  the city  assumed  with  permanent  they  essential  Sparrow,  i n urban  are  re-  systems),  also  species  I can't  test  this  resident  portion  than  directly.  In  birds  are  consider-  The former  were  present  of the year,  selected  are distrib-  e s s e n t i a l resources  residents.  habitat  therefore  i t  factors  which  using  i s  survival.  species  were  areas:  Domestic  Glaucous-winged  Northwestern residents  are  resident  Winter  f o rtheir  following  residents  late  they  SPECIES  The  Thrush,  that  though  f o ra large  that  RESIDENT  the  o r whether  a r e (in u n d i s t u r b e d  to preferred  are  were  factors,  f o r these. I  uted  (ultimate)  Crow,  were  Gull,  t h e Song  Sparrow,  these  o f each t o some  essential  distribution.  o f these  common  Pigeon,  House  and Black-capped  and Golden-crowned  range  t h e most  Kinglet. species  S t a r l i n g , House  Finch,  American  Chickadee.  Dark-eyed  Robin,  Common  Junco,  I shall within  permanent  Varied  briefly  Vancouver  describe and r e -  c h a r a c t e r i s t i c s o f the environment  t o them,  a n d may b e d i r e c t l y  Winter  affecting  which their  61.  Domestic Pigeon Pigeons were most abundant i n "commercial"  habitat,  but were a l s o q u i t e abundant i n "apartment" and " i n d u s t r i a l " habitats.  They were found i n much lower  r e s i d e n t i a l areas.  density i n a l l  Two f a c t o r s are thought  to be c o n t r o l -  l i n g the abundance of the Domestic Pigeon:  food p r o v i d e d  by man, and s u i t a b i l i t y o f b u i l d i n g s f o r r o o s t i n g and nesting.  F l a t - r o o f e d b u i l d i n g s w i t h numerous ledges and eaves  were q u i t e common i n the commercial, trial  apartment, and indus-  areas, though the q u a l i t y o f these b u i l d i n g s f o r  r o o s t i n g was perhaps r e l a t i v e l y low i n the i n d u s t r i a l  area.  Murton et a l . (1972) found that s p i l l e d g r a i n and s i m i l a r f o o d s t u f f s were the most important the abundance o f pigeons  factors  determining  i n dockside areas.  Since the com-  m e r c i a l p l o t was the c l o s e s t one to g r a i n - s h i p p i n g f a c i l i t y i e s , i t was to be expected pigeons  that the h i g h e s t d e n s i t y o f  should be found there.  Starling S t a r l i n g s were most abundant i n the r e s i d e n t i a l with l e s s than 80% v e g e t a t i o n cover, peaking h a b i t a t s with 40 to 60% v e g e t a t i o n cover presence  o f grassy areas  i n abundance i n  (Figure 9 ) . The  (lawns) may be a c r i t i c a l  determining S t a r l i n g abundance i n c i t i e s ,  areas  factor  since this i s t h e i r  p r e f e r r e d f o r a g i n g m i c r o h a b i t a t (Weber, 1972).  Lawn was  62.  Figure 9:  Estimates o f S t a r l i n g d e n s i t y per p l o t t o t a l percent v e g e t a t i o n cover. sities  versus  The mean den-  i n Summer and Winter have been graphed.  Figure 10: Estimates o f Black-capped  Chickadee  d e n s i t y per  p l o t versus t o t a l percent v e g e t a t i o n cover. mean d e n s i t i e s graphed.  The  i n Summer and Winter have been  63. FIGURE  9  winter  40  "  80  Total  '  Vegetation  120 Cover  180  (%)  FIGURE  Total  Vegetation  Cover  (%)  10  plentiful  i n most h a b i t a t s where S t a r l i n g s w e r e a b u n d a n t .  H o w e v e r , my may  be  dant  results  i n d i c a t e t h a t f o o d s u p p l i e d by  of a d d i t i o n a l  i n the p l o t  residents  importance.  i n which  f o r b i r d s was  S t a r l i n g s were most abunr  t h e amount o f f o o d p r o v i d e d  g r e a t e s t CMacKenzie),  p l o t where p e r c e n t o f a r e a c o v e r e d by (Queen E l i z a b e t h ) . S p r i n g and  Fall,  roost  distribution  not  l a w n was  Regression results  the a v a i l a b i l i t y  influence Starling  people  by  i n the  greatest  indicated that i n  of coniferous trees  (Appendix  5b).  may  Starlings  communally i n t r e e s a t t h e s e times o f y e a r  (Jumber,  1956).  House S p a r r o w D i s t r i b u t i o n o f t h e House S p a r r o w was that o f the S t a r l i n g . residential were f o u n d in their  and  apartment  abundance when t o t a l  but  occurred i n typical  T h e r e was  a sharp  •  d r o p .'  v e g e t a t i o n cover exceeded  60%.  Gull  T h i s s p e c i e s was West End.  to  h a b i t a t s , t h o u g h House s p a r r o w s  i n a l l urban h a b i t a t s .  Glaucous-winged  plots.  Highest densities  very s i m i l a r  most a b u n d a n t i n Downtown and  Garbage i s p r o b a b l y t h e g u l l ' s  Food s c r a p s were a l s o a v a i l a b l e  only food i n these  in residential  g u l l s w e r e n o t n e a r l y as a b u n d a n t t h e r e .  d i s t r i b u t i o n of the Glaucous-winged  the  areas,  Therefore  Gull, primarily  a  the  coastal  species,  may  be  more a f f e c t e d by  former areas to  the  available.  presence  and  The  look-out  posts  o c e a n t h a n by  may  of t a l l  the  proximity  the  of  amount o f  buildings  the  garbage  f o r p e r c h e s .....  a l s o have a f f e c t e d t h e i r  distribu-  tion.  House  Finch This  species  residential structural  plots  the  was  d i s t r i b u t i o n of  both p l o t s .  creased  the  Summer, and  presence that  the  i n apartment  this  feature  tree  dropped  in typical  density.  it  was  habitat  in  In W i n t e r ,  density  of  i n woody r e s i d e n t i a l  habitat  and i n r  residential  seed a v a i l a b l e i n the  more d e n s e l y p o p u l a t e d  idential  habitat  American  Robin r o b i n was and  (Table  the  (MacKenzie).  to  commercial  plots  influence  however, s i n c e the  only  evergreen  actually  a t t r i b u t e of  woody  these  of broad-leaved  have been i n r e s p o n s e  The  The  distinguished  House F i n c h  insignificant  Fall.  and  Regression r e s u l t s indicated a relationship,  coniferous  House F i n c h  abundant  which c l e a r l y  It i s doubtful  a comparatively  with  most  i n Spring,  features  from a l l o t h e r s trees.  was  much g r e a t e r  This  quantity  the  change of  typical  may  bird res-  7).  commonly f o u n d  industrial.  in a l l habitats  Abundance was  greatest  except i n woody  residential 80%. and  In  habitat,  Winter,  woody  there  residential  robins  which  prefer  residential  tion  arrive  total  were  vegetation  relatively  few  cover  was  robins  in  about older  habitats.  This  indicates  that  i n Vancouver  each  Spring  to  breed,  more  than  habitats  which  have  70%  migrant may-  vegeta-  cover. This  choice  different  types  indicated  that  cover  of  sult  for not  habitat  perches  or  This  , I  and  in  cover.  that  the  amount  shrubs  and  robin  obtained  foraging conifers  or  similar and  nesting  may  be  between  density  a  habitat,  of  regressions  relationship  (1973)  either  on  Multiple  suburban  re-  conifers purposes  preferred  as  Crow was  found,  most  Spring  My  and  crows  were  least in  observations  and  Lawns  at  abundant  c h a r a c t e r i s t i c s which  purposes.  based  roosts.  plots.  during  be  positive  Thomas  suggest  I t was  residential  trees  for  species  habitats.'  a  censused  evening  Northwestern  was  Since  preferred  may  vegetation  there  robins  1972)  crows  of  5b).  (Weber,  mental  of  coniferous  (Appendix  are  where  may  garbage  may  typical,  older,  indicated  have  Summer w e r e  occasionally,  been  tall  have  of  that  of  and  woody  environ-  importance  trees,  been  in a l l  for  to  nesting  importance  in  Winter. Thus,  most  abundant  i n woody  residential  hab-  67.  itat  i n S p r i n g and Summer, and i n t y p i c a l r e s i d e n t i a l hab-  itat  in Fall  Black-capped  and Winter  (Appendix  4).  Chickadee  Although found i n a l l h a b i t a t s w i t h 351 v e g e t a t i o n cover or more, the Black-capped Chickadee was bird.  Its abundance was  p r i m a r i l y a woodland  roughly p r o p o r t i o n a l to the  v e g e t a t i o n cover (Figure 7).  Thomas (1973) and I b o t h i f o u n d  that the amount of c o n i f e r o u s t r e e s may the abundance o f chickadees  total  be used to p r e d i c t  ( r e g r e s s i o n r e s u l t s not p r e s e n t e d ) .  Coniferous t r e e s are f r e q u e n t l y used as r o o s t s by f l o c k s of chickadees i n F a l l , Winter, and p o s s i b l y e a r l y S p r i n g (Odum, 1942). Odum (1942) a l s o found that b i r d seed p r o v i d e d i n r e s i d e n t i a l areas a t t r a c t e d chickadees i n the Winter. tionnaire results ers  My  ques-  i n d i c a t e that chickadees eat at b i r d feedr  i n Winter to some extent, but the number of chickadees  i n urban p l o t s was  not h i g h e r i n t h i s season than at other  times of the year.  During F a l l , however, the abundance of  chickadees w i t h i n urban areas i s best p r e d i c t e d by the amount of  food ( b i r d seed) p r o v i d e d by man  Winter  (Appendix  5b).  residents  The Song Sparrow and Dark-eyed  Junco were found i n a  wide v a r i e t y of urban h a b i t a t s during Winter and m i g r a t i o n  68.  seasons. tial  They  habitats.  bined  with  gardens to  were  changes  in  results by  be  (,1972), of  the  responding  Thrush  and  Golden-crowned  habitat,  food  habitats  during  availability  from  urban  to  in a  though  or  the the  food  woody  (Appendix  Weber  features  and  are  urban  changes  may  in old  species  i n woodland  i n most  these  observations  Varied  abundant  abundant  statistical  shrubbery  these  The  found  field  and  which  My  most  4,  5b)  com-  indicate  that  environment  positive  manner.  K i n g l e t were Varied  most  Thrush  Winter.  was  Seasonal  preferences  woodland  residen-  may  cause  habitat.  Summary Migratory of  foliage  suitable  or  and  in direct  with  less  tic such  highest are  types  Pigeons as  species  the  70% to  of and  of  the  to  resident  than which  amount  are are  in  areas  other  birds  are  of  and  food  In  to  cover  of  resources of  this,  dominant  in  areas  be by  Resident  70%  I  distributed  distributed man.  d e n s i t i e s of  Black-capped over  hand,  support  to  provided  Gulls.  locating  essential  appear  influence  with  in proportion  of  numerically  cover  in  the  migrants.  Glaucous-winged Robin  On of  characteristics  species  availability  b u i l d i n g s may  abundances  structural  species  vegetation  American  distributed  use  migration.  resident  foods,  than  proportion  icular  during  response  particular resident  probably  presence  that  some  in  the  habitat  hypothesized more  birds  PartDomes-  species,  Chickadee,  vegetation vegetation  '  with  cover, or  part-  69.  i c u l a r types of  foliage.  Characteristics  of the  Community c h a r a c t e r i s t i c s of ways.  I will  description  the  of s p e c i e s d i v e r s i t y ,  trophic  DIVERSITY OF  be  d e s c r i b e d i n a number  and  discussion  description  a positive (H ) 1  Since f o l i a g e  and  linear relationship  and  evenness a l s o  gradient of u r b a n i z a t i o n .  between b i r d  (1961), Karr and  Roth (1971), W i l l s o n  relationship  MacArthur  (.1974), and  species  in natural  W i l l s o n both found a  between b i r d s p e c i e s d i v e r s i t y and  others  areas,  curvilinear t o t a l vege-  cover.  The birds  Roth, and  the  communities  between b i r d  f o l i a g e height d i v e r s i t y  though Karr and  support  MacArthur and  have observed a l i n e a r r e l a t i o n s h i p d i v e r s i t y and  diversity,  decrease l i n e a r l y w i t h  of urban b i r d  to h a b i t a t s t r u c t u r e .  (.Figure  t o t a l vegetation  These r e s u l t s  hypothesis that c h a r a c t e r i s t i c s  tation  density  f o l i a g e height d i v e r s i t y  height d i v e r s i t y  number of s p e c i e s , and  related  the  communities,  cover decrease with u r b a n i z a t i o n , b i r d s p e c i e s  are  to  SPECIES  species d i v e r s i t y  the  of  i n r e l a t i o n to t o t a l b i r d  s t r u c t u r e of the  There was  11).  may  Communities  r e s t r i c t t h i s aspect of the  abundance of i n d i v i d u a l s and  Bird  are  first  result  suggests that urban communities  fundamentally s i m i l a r  of  to those i n n a t u r a l areas  so  70.  Figure  11:  Mean  bird  versus (see  foliage  Table  regression fidence  species  6,  height page  line  level.  diversity  (Summer)  diversity.  30) h a s b e e n  is significant  The used  i n each index  plot FHD2  here.  The  a t t h e 0.05  con-  71.  F I G U R E II  B S D VS» F H • E (SLM/ER) Y =  0.B55B + 0«S5B3  *X  F H• E  N =  B  that a  the structural  prime  determinant  hand, w h i l e structural are or  of bird  man-made  of vegetation  species  features  complexity  to bird  species  add c o n s i d e r a b l y  o f the urban  found  that  bird  species  with  the diversity  lationship" diversity  environment  even  was  bird  than  Summer  (Table habitat  that  while  Winter  to the o v e r a l l (Table  6)  they  species  Similarly,  Emlen  considered,  varied  inversely  This  negative " r e -  that  foliage  t h e amount  height  of  cover  Winter  types  this  varied  considerably  similar  habitat  structure  d e n s i t i e s were  on t h e whole  In contrast, season  Weber's  densities  densities increased he c o n s i d e r e d  comparison bird  trophic  only  may n o t b e density  i n the relative  to specific  between  urbanization  in total  t o changes  GROUPS  between  breeding  14), but since  belonging  correlated with  no t r e n d  densities.  Variation related  with  therefore  density.  ization  the fact  TROPHIC  density  those  bird  suggested  reflect  On t h e o t h e r  structures.  between  There  simply  t o be  of bird  were  number  o f the environment.  D E N S I T Y AND  Total  12),  features  and s p e c i e s  i s negatively  BIRD  (Figure  w h e n man-made  diversity  may  man-made  TOTAL  diversity  seems  diversity.  i n v e r s e l y , o r n o t r e l a t e d t o t h e number  (1974)  of  complexity  and  u  r  e  3).  total  greater  (1972)  data with  urban  urbanization  a narrow  range o f  meaningful.  between  abundance  groups.  ( F i g  decreased with  plots,  Weber  plots  can be  of individuals Q972)  found  73.  Figure 12:  Mean b i r d s p e c i e s d i v e r s i t y (Summer) versus the man-made component  of habitat  structural diversity.  The l a t t e r has been c a l c u l a t e d by s u b t r a c t i n g  FHD2  (6 l a y e r s o f v e g e t a t i o n ) from FHD3 (9 l a y e r s , i n c l u d i n g b u i l d i n g s and pavement), s i n c e the a d d i t i o n of s t r a t a has an a d d i t i v e e f f e c t on d i v e r s i t y (personal communication P. B e l l e f l e u r ) . f i t t e d by eye.  The curve was  FIGURE  12  2.8  2.4  2.0  \  o  \ \  \  \  o  0  o  \  \  BSD  °  1.2  \  \ °  \  \  \ \  .8  * n o t e the addition of strata has an additive effect on diversity. :  \  .4  .2  ,.4  .6  F H D 3 - F H D 2 * (see  .8  1.0  Toble 6 )  -„.-  T  that  most  (Table  of  the  15).  As  birds  vegetation cover  creased  in relative  system,  I also  abundance.  found  ically  dominated  urban"  habitats  by  i n Winter  decreases  This  expected  larly  scarce  Weber's urban  study  and  habitat  amount  of  Winter  (Table  and 7)  may  the in  biomass a  vided  by  the  are  birds  a wide  forage  of the  particuT..  time  of  the  related  amount  are  year.  the  The  in  sub-  increased residents  of natural  foods  (90%  seed-eating  of bird, seed  in  increase in  dominance of  both  p r o p o r t i o n of  relative  the  In  omnivores  suburban  individuals)  Starling,  on  of  the  season,  data  foods  suggest  to  including  nest that  i n most  Sparrow,  suited  ground, and  birds  House  particularly  variety  mainly  My  the  are  breeding  1972).  (1974)  insects  while  of  "highly  habitat.  shortages  f o r some  in  proportion of i n -  increased.  with  i s numer-  i n woodland  proportion of  n u m e r i c a l l y dominant  Vancouver  eat  to  that  this  community  The  in-  of birds  and  bread  pro-  urban  areas  of  people.  The  These  area  at  omnivores  classification  particularly  except  p r o v i d e d by  combined  number  bird  15).  i n Winter,  bread  Emlen  and  suburban  the  Weber's  birds,  herbivores)  account  graminivores.  urban  (Table  areas  were  increased, insectivores  indicates  decreased  seed  Winter,  and  mine,  (plus  the  i n Winter,  i n urban  graminivores  in  that  habitats  Using  omnivorous  sectivores i s as  in residential  are  not  mainly  and  urban  Domestic  areas  since  those  p r o v i d e d by  highly  territorial  in buildings  variation  in total  Pigeon. they man, during  (Weber,  bird  density  76. T a b l e 14:  Comparison o f B i r d Community C h a r a c t e r i s t i c s with  those o f Weber  (1972).  SUMMER H a b i t a t Type  WINTER  T h i s Study  Weber  T h i s Study  Weber  Apartment  1. 72  1. 72  1. 74  1. 72  Typical  1. 30  1. 81  1. 73  1. 45  fX W W U <U > rH - H •H O PQ  Woody R e s i d e n t i a l  1. 99  2. 21  2. 15  2. 37  Woodland  2. 62  --  2. 03  --  --  2. 42  --  1. 87  Speci ichne  10 (D •r-l >-N O +J  Oi  Evenn  in in 0)  Residential  Rural Apartment  10  13  Typical  13  Woody R e s i d e n t i a l  17  Woodland  22  Residential  16  15  15  12  30  19  30  18  --  Rural  12  --  25  32  Apartment  75  67  70  62  Typical  51  67  64  58  70  65  73  70  Woody  Residential  Residential  Woodland  -  73  -  Rural  -  70  -  75  54  Apartment  469  314  f-c  568  370  Typical  412  411  471  501  m +->  Woody  397  439  342  319  Tota: Den:  •H  Residential  Woodland Rural  Residential  212  477  -  655  -  1021  T a b l e 15:  T r o p h i c Groups o f B i r d s :  Proportion  of Total  Number o f I n d i v i d u a l s i n each Group.  Habitat Trophic  Group  Commercial Industrial  Apartment  97V '  81%  Omnivore Graminivore +Herbivore  2%  Insectivore  1%  Omnivore  1001  .  Type  Suburban 82%  [:  1  Woodland  Rural  19%  -  -  8%  6%  17%  11%  12%  63%  91%  78%  24%  -  Graminivore +Herbivore  01  Insectivore  0%  Omnivore  •-  9% 0% •  16% : 6%  11% 65%  -  -  59%  92%  73%  1%  12%  -  7%  15%  -  59%'  -  2  Graminivore +Herbivore  -  Insectivore Omnivore  20% ' 21%  -  91%  +Herbivore  -  8%  31%  -  24%-  Insectivore  -  1*  10%  -  . 1%  •  75%  Graminivore  Based on f o u r s t u d y  areas.  B a s e d on two s t u d y a r e a s .  78.  i n areas n u m e r i c a l l y dominated by these s p e c i e s , was r e l a t e d to v a r i a t i o n i n the amount o f food made a v a i l a b l e by man r a t h e r than the s t r u c t u r e o f the environment as o r i g i n a l l y hypothesized.  For example, the t o t a l v e g e t a t i o n cover i n  " t y p i c a l r e s i d e n t i a l " p l o t s , Queen E l i z a b e t h zie  (MK), was q u i t e s i m i l a r  birds  (.QE) and MacKen-  (Table 2 ) , y e t the d e n s i t y o f  i n MK was roughly three times t h a t i n QE.  Correspond-  ing to t h i s was a 3 - f o l d d i f f e r e n c e i n the estimated amount o f food p r o v i d e d by r e s i d e n t s .  The h i g h e r t o t a l  density i n  MacKenzie was p r i m a r i l y a r e s u l t o f i n c r e a s e d abundance o f the S t a r l i n g and House Sparrow, s p e c i e s which are known to consume food p r o v i d e d by people. In areas with more v e g e t a t i o n cover, where these s p e c i e s were l e s s abundant, abundance o f n a t u r a l foods may have been r e l a t e d to t o t a l b i r d d e n s i t y .  Some i n t e r e s t i n g  relation-,  ships emerge i f we assume that the t o t a l v e g e t a t i o n cover i s an approximation o f the r e l a t i v e a v a i l a b i l i t y o f n a t u r a l  foods  The Winter b i r d d e n s i t y i n " o l d e r r e s i d e n t i a l " h a b i t a t ( C o l lingwood:  CW) was approximately twice that i n QE, but i n  both areas the amount o f food p r o v i d e d by people was about the same.  However, the t o t a l v e g e t a t i o n cover was 10% great-  er i n Collingwood than i n Queen E l i z a b e t h .  Perhaps  this  small i n c r e a s e i n v e g e t a t i o n cover, most of t h i s being i n the form o f deciduous b i r d density.  trees, resulted i n this  A critical  increase i n  amount o f p l a n t cover may be r e -  79.  quired before species which rely primarily on vegetation as a foraging substrate will 'recognize', in large numbers, the suburban habitat as suitable.  Between 60 and 75% vegetation  cover (.5 and 20% tree cover) , the abundances of many species changed sharply (e-g- Figures 8, 9, and 10).  The species  which accounted for most of the difference in total density in  Queen  Elizabeth and Collingwood plots in Winter were the  American Robin, Song Sparrow, and Dark-eyed Junco;  all of  which forage mostly in vegetation of various types for naturally occurring foods, though they also accept food from people.  Interestingly, Willson 0-974) derived a related  result:  "The addition of trees in a vegetational series  has a disproportionate effect on the addition of species, primarily by the addition rather than the expansion of guilds." Another useful comparison may be drawn between "older residential" (CW) and "woody residential" (SH) plots. ter  Win-  bird density was about the same in these (Table 11).  Total vegetation cover was 14% higher in SH than in CW, while food provided by man was approximately 13% less.  The  proportions of various species' abundances changed substantially, with those of the American Robin, Dark-eyed Junco, Black-capped Chickadee, and Varied Thrush increasing noticeably with vegetation cover (Appendix 3).  Thus, total  bird density may have changed little between woody and older  80.  residential (naturally quantity  because  available  of  species were  plots  food  most  These  abundant  uctivity"  of  foods,  is  related  trast,  Willson  between areas,  and  that  productive As and in  species  foraging  species  to  relative  by  number the  noted,  existing  data of  the of  food  was  guilds of  species  number  13)  as  of  to  s t r u c t u r a l complexity  can and  in  This not  as  natural  comparison  be  generally  i t is  in of  an  non-  as  an a  in  diversity increase  species.  14) , a n d  inhabit  urban  etc.).  increase  ground  con-  relationship  foliage height  of  "prod-  By  proportions  MacArthur's  which  of  natural  birds  may  well  (.Figure  omnivorous  conform  no  correlated with  corresponding  (Figure  total  and  habitat.  high  in  Junco  cities.  was  biomass  increasing  cover  a  that  in  (.pavement, b u i l d i n g s ,  was  guilds  evidence  there  by  d i v e r s i t y and  abundance  These the  there  Dark-eyed  environments  characterized  vegetation  prisingly, of  are  of  in natural  previously  bird  and  decrease  areas.  density  that  availability  "cover"  total  bird  productivity  factor  which  total  cover  assemblage  i n c l u d i n g man-made  abundance  the  a  modified  i n both  additional  (.1974) f o u n d  the  limiting  areas  to  A  by  S t a r l i n g and  species  are  in vegetation  countered man.  the  habitat,  estimated  suggests a  the  by  though  results  increase  f o o d ) was  provided  appeared,  the  the  Not the  surnumber  addition  of  decline  in  feeding  birds.  (.1965) p r o p o s a l an  area  productivity  of  the  is  that  limited  the  habitat.  81.  Figure  13:  Number  of foraging guilds  versus  total  curve  was  nized  were  percent  fitted  vegetation  by e y e .  t h e same  per plot  (Summer)  cover.  The g u i l d s  as t h o s e  used  by  The  recogEmlen  (1974).  Figure  14:  Number  of species  versus  total  graminivore  percent  vegetation  (Summer)  cover.  ( s e e d - e a t i n g ) , and ground-  trunk-gleaning graphed.  per foraging guild  insectivore  guilds  have  Only and been  FIGURE  10  40  80 Total  Vegetation  120 Cover  160 (%)  14  83.  He  states  and  that  therefore  dict  number  number  of species  Many  in  ler  vegetation,  vertically,  related  drops of  by  resource  are also  added  and that  suggests  t h e number  that  t h e number  a v a i l a b l e may  greater  species  of bird  than  be  5%  of In  resources  and n a t u r a l  areas  1972).  As  other  prod-  i n which  is functionally of  structural  The number  and  the important once  factors  vegetation  cover).  As  t h e amount  partitioning  i s found  preferences  tal-  can c o e x i s t .  o f ways  resource  species  microhabitat (Weber,  Hab-  behaviour  which  component areas.  increases,  bird  species  to p a r t i t i o n  of vegetation  vegetation  urban  of bird  example,  and  (less  foraging  For  by b u i l d i n g s , pavement,  low l e v e l s  Different  o f ways.  grassland.  of coexisting species  and t h e number  number  the  the  as i t i s i n n a t u r a l types  the  r e l a t i o n s h i p between  to very  occur,  able  pre-  partitioning.  i n simple  the negative  to  because  views.  i n feeding  to the vegetational  o f food  limiting  similar  i n c r e a s i n g t h e number  complexity  only  amounts  increases  c a n be p a r t i t i o n e d i n u r b a n  complexity,  i s needed  the coexistence  overlap  species  noted  This  resources  that  associated,  factor increases,  forwarded  niche  features,  uctivity.  this  and d i f f e r e n c e s  thereby  have  man-made  complexity  can coexist  is facilitated  minimize  structural  have  concluded  preferences  I  As  which  authors  grassland  species  structural  are usually  c a n be p a r t i t i o n e d i n a g r e a t e r  (J968)  itat  only  two m e a s u r e s  of species.  environment  Cody  these  to  increase.  are found  vegetation  can  among  height  and  84.  number  of  probably Cody  foliage enable  munities.  A  variety  izations (i.e.  tion  (1974)  examined  number  of her  by  others.  cover  was or  to  c o e x i s t , as  preferences  MacArthur  shared  in  way.  shared  not  Areas 60  comparisons, numerical  In  to  the  and  and such  conclusion,  in  similarity  of  of  few  similar  80% as of  of  counts  plots  the  than  "width  vegeta-  areas grassy  can  be  vegetation  species.  between  percent  which  percent  areas  species  However,  species  s p e c i e s , which  study  or  cover  habitat  my  with  compared  W i l l s o n made of  in  general-  study  of bird  com-  fur-  niche" was  not  investigate.  the  of  vegetation,  natural  foods  ered  as  components  that  by  providing  i t appears  avifaunas  urban  Both  areas  with  bird  like  diversity.  a  on  bird  similarity  proportion  dominance  characteristics factors  t h a t when  to  only  of  contradicted previous  found  height  I have  between  designed  f i n d i n g s based  related  a higher  areas.  this  characteristics  compared,  foliage  shrubby  ther  She  "wooded") were  trees  of  species  of n a t u r a l habitats  composition  and  more  increase, microhabitat  suggest. Willson  a  types  that  are  primarily  environment;  and  the  and  those  the  provided  greatly increase  for birds, the  total  and  by  of man  Emlen  human  the must  (19 74)  residents  abundance  diversity  r e g u l a t e d by  structural  "productivity"  of productivity. food  density  and  two  complexity habitat. be  consid-  concluded of  suburban  biomass  of  85.  birds,  over  that  in. n o n - u r b a n  this  further,  sity  (and biomass)  ing  and found  l a r g e l y on  effect  number  of bird  species  within  species  plexity do  studies  This urban  was  need  t h e number  be  only  depend-  have  The  habitat. shown  that  of  f o r the evenness the s t r u c t u r a l  considered;  of species,  the  complexity  f o r t h e number  as  den-  by p e o p l e .  to s t r u c t u r a l  as w e l l  However  areas,  o f the  areas  bird  man-made  and  comfeatures  or overall bird  spec-  diversity. I  able  suggested  i n the Introduction  t o e c o l o g i c a l l y manage  case,  then  i t i s clear  factors  that  should  of  made  available  food  amount of  structure  investigated  total  among  hypothesized  areas,  of vegetation  a city,  provided  is related  diversity.  not influence  ies  o f food  i n non-urban  species  I have  considerably  t h e amount  the habitat.  bird  varies  within  i s not correlated with  Numerous  of  that  habitat.  and s t r u c t u r e  species  from  birds my  results  by p e o p l e  that  be  desir-  I f this  i s the  the  environmental  a r e t h e amount  (to control  of vegetation o f many  i t might  in cities.  be m a n i p u l a t e d  and abundance  that  types  p e s t s ) , and t h e  (.to i n c r e a s e  "desirable"  and  t h e number  species).  86  SUMMARY  This study  examined t h e r e l a t i o n s h i p s between urban  b i r d communities and c h a r a c t e r i s t i c s i n a wide range o f urban h a b i t a t s . b i r d s were observed. Domestic Pigeon,  the b r e e d i n g  environment, species of  residents (e.g.  House S p a r r o w , S t a r l i n g ) w e r e f o u n d i n a  treed residential, Winter  Sixty-two  Some o f t h e y e a r - l o n g  wide v a r i e t y o f h a b i t a t s , r a n g i n g  of these.  of their  f r o m downtown  to well-  a n d were n u m e r i c a l l y d o m i n a n t i n many r e s i d e n t s , s p e c i e s w h i c h were a b s e n t  season but present  during  the r e s t o f the year (e.g.  Song S p a r r o w , D a r k - e y e d J u n c o ) , were a l s o common a t t i m e s , especially  i n t h e more wooded r e s i d e n t i a l  a l a r g e number o f t h e s p e c i e s o b s e r v e d as s w a l l o w s ,  w a r b l e r s , and v i r e o s .  resident species  However,  were m i g r a n t s ,  as w e l l .  s p e c i e s w e r e common o n l y d u r i n g m i -  g r a t i o n p e r i o d s , and n o t i n t h e b r e e d i n g e s p e c i a l l y the Wilson's  such  Some p o p u l a t i o n s o f  ( e . g . A m e r i c a n R o b i n ) were m i g r a n t  Many o f t h e s e m i g r a t o r y  areas  areas.  Warbler,  season.  A few,  w e r e common i n r e s i d e n t i a l  d u r i n g m i g r a t i o n s e a s o n s , b u t many w e r e f o u n d o n l y i n  woodland.  The a v e r a g e number o f s p e c i e s p e r m o r n i n g  i n an e i g h t h e c t a r e  p l o t v a r i e d f r o m 4,5 i n t h e c i t y  t o 10.5 i n a w e l l - t r e e d r e s i d e n t i a l  area.  Mean t o t a l  count centre bird  d e n s i t y v a r i e d f r o m 130 p e r 40h i n a n i n d u s t r i a l p l o t t o 525 per  40h i n an a p a r t m e n t  plot.  87.  The  major s p e c i f i c f i n d i n g s o f t h i s study  can be sum-  marized as f o l l o w s . (.1)  B i r d s p e c i e s d i v e r s i t y , evenness o f s p e c i e s ' abundances, and numbers o f species i n c r e a s e as t o t a l v e g e t a t i o n and  f o l i a g e height  diversity  cover  i n c r e a s e i n an urban e n v i r -  onment . (2)  There were no o v e r a l l  seasonal  trends  i n changes between  seasons o f b i r d species d i v e r s i t y , evenness, and numbers of s p e c i e s , though d i v e r s i t y was lowest mer  i n a majority of plots.  d u r i n g the sum-  There was l i t t l e  change i n number o f species i n areas with t o t a l vegetation (3)  The s t r u c t u r a l  less  seasonal than 40%  cover.  complexity  c o n t r i b u t e d to urban environ-  ments by man-made f e a t u r e s such as b u i l d i n g s i s negatively  (or not)  related  to b i r d species d i v e r s i t y .  i s i n c o n t r a s t with the trend f o r s t r u c t u r a l c o n t r i b u t e d by v e g e t a t i o n . complexity  complexity  This suggests that  o f v e g e t a t i o n i s a s s o c i a t e d with  This  structural  food compon-  ents o f niches whereas man-made f e a t u r e s g e n e r a l l y are not. C4)  Food p r o v i d e d by man probably t o t a l b i r d density  has a great e f f e c t on the  (and biomass) i n urban areas,  fore food i s a l i m i t i n g f a c t o r .  Resident  there-  s p e c i e s (e.g.  House Sparrow, S t a r l i n g ) which are n u m e r i c a l l y  dominant  i n most urban b i r d communities appear to be d i s t r i b u t e d  88.  in  relation  This  t o t h e amount  effect  density,  increases, with  however,  urbanization  (i.e.  decrease  total  bird  density,  and i s l a r g e l y  tural  changes  warblers,  that  in  r e l a t i o n . to  Thus  of diversity,  vegetation  but  not with  However  confirm  of urban.bird  structure.  Number  with  s t r u c t u r e , and w i t h  total  density  inance  by a p a r t i c u l a r  fected  more-by  of the  o f omnivores  of inscctivores  my  original  species  are a function and o t h e r  increasing  total  hypothesis,  in-  complexity  vegetation  cover,  c o n t r i b u t e d b y man-made s t r u c t u r e s . of birds, group  t h e amount  11abi.tat s t r u c t u r e .  such  are distributed  communities  of bird  d i dincrease  diversity  of struc-  species,  abundance that  of  i n Winter.  findings partially  of  independent  foods  urbanization, while  with  cover),  structure, or composition  the relative  especially  bird  determinant  Those  on n a t u r a l  amount,  characteristics  habitat  rely  Total  nor decreases  i s a major  i n the environment.  as  b y man.  i n vegetation  b y man  These  "the  increases  provided  decreases,  dices  with  urbanization.  food  increases w i t h  of  neither  provided  since  vegetation.  that  o f food  and degree  of species  o f food  of numerical  (omnivores),  provided  b y man,  i s  than  domafby  89.  LITERATURE CITED  American O r n i t h o l o g i s t s ' American Birds more P r e s s ,  Union.  1957.  (Fifth Edition).  Check-list 691 pp.  of North  Lord  Balti-  Baltimore.  American O r n i t h o l o g i s t s '  Union.  1973.  Thirty-second  plement t o the American O r n i t h o l o g i s t s '  Union  list  411-419.  of North American B i r d s .  C a m p b e l l , R.W., 1974.  M.G.  S h e p a r d , B.A.  Vancouver b i r d s  ural History Cody, M.L.  H.N.  In:  96 pp.  1964.  Sci.  E m l e n , J.T. ona:  a n d W . C . Weber. Vancouver  Nat-  On t h e methods o f r e s o u r c e d i v i s i o n i n  Starlings  Am?  N a t l . > 10.2':'  i n the P a c i f i c  "2nd V e r t e b r a t e p e s t c o n t r o l  Acad.  check-  Vancouver.  g r a s s l a n d b i r d communities. Elliott,  90:  MacDonald,  i n 1972.  Society,  1968.  Auk  sup-  pp.  19 74.  107-147.  Northwest.  conference."  Natur,  29-39. An u r b a n b i r d community i n T u c s o n ,  derivation, structure,  regulation.  Condor  Ariz76:  184-197. G r a b e r , R.R.  and J.W.  Graber.  of b i r d populations 1958. Hilden,  0.  Ann.  Bull. 1965. Zool.  1963.  A comparitive study  in Illinois,  111. N a t . H i s t .  Surv.  1906  - 1909  28:  Habitat selection i n birds:  Fen.  2:  53-75.  and 1956  383-528. a review.  -  9 0."  Hoerl,  A.E.  a n d R.W.  biased  estimation  metrics Jumber,  12:  J.F.  a n d R.R.  avian  diversity  105:  423-435.  MacArthur, Rev. Murton,  40:  R.K.,  ogical  Odum, E . P .  R.J.P.  Auk  Pielou,  E.C.  2 86 p p . Principles Acad. Smith, of  N.G.  42:  World  structure  areas.  1961.  On  Am.  bird  and  Nat.  species  594-598.  Patterns  Thearle,  of species  diversity.  Ecol.  9:  Biol.  pigeon  Columba  and methods  1972.  Ecol-  livia  var. I.  of  control.  835-874.  Annual  59:  cycle  o f Black-capped Chickadee  -  499-531.  1969. John  of plant  An  1975.  introduction  Wiley  and Sons,  and  animal pest  S c i . , Wash.  Acad.  and J . Thompson.  o f the f e r a l  D.C.  5:  Sci.  to mathematical ecology.  New  York.  control.  biological  and n o n a t t r a c t i o n 72:  1970.  Nat,  58-82.  Spshing noise:  i t s attraction  Natn.  New  in  411-426.  Vegetation  breeding biology,  1942.  3.  Techno-  510-533.  studies  Appl.  1971.  MacArthur.  1965.  Population, J.  73:  i n several  Ecology  R.H.  Auk  Roth.  a n d J.W.  diversity.  regression:  f o r nonorthogonal problems.  Pennsylvania.  R.H.  Ridge  Roosting behaviour of the S t a r l i n g  J.F.  MacArthur,  1970.  55-68.  1956.  central Karr,  Kennard.  1411-1414.  significance  by b i r d s .  Proc.  91.  Speirs,  M.J.,  G.  of  birds  i n urban  Field Svardson,  Biol. G.  birds. Thomas,  Ph.D. Tomoff,  Ecology W e b e r , W.C.  tat Woolf,  selection  of habitat  - a pilot  Ont.  in  species  requirements  study.  of Massachusetts,  245  pp.  Massachusetts.  diversity  1974.  in cities:  i n desert  a study  and n e s t - s i t e s  T h e s i s , Univ.  Nostrand, Zar,  Birds  ecology,  1968.  1969.  scrub.  396-403.  structure.  C.  areas  Avian  1972.  M. F.  Populations  County,  and h a b i t a t  determination  i n suburban  55:  foraging  Willson,  The  1974.  Ontario  1970,  157-174.  T h e s i s , Univ.  C.S.  M.Sc.  1:  1973.  Tozer.  1-12. Competition  Oikos  birds  a n d R.G.  habitats,  24:  1949.  J.W.  for  Markle,  of British  Avian Ecology  Principles  55:  o f urban  Columbia,  community  of populations, birds. British  269  pp.  Columbia.  o r g a n i z a t i o n and h a b i -  1017-1029.  of biometry.  359 ,pp.  D.  Van  Toronto.  J.H.  1974.  Hall,  New  Biostatistical  Jersey.  analysis.  620  pp.  Prentice-  Appendix 1:  —  Sample o f B i r d Census Mapsheet  U^r  Eaca —i  I  .  !  i i  1 i  A/icnt  A  t  !  i  J  Appendix 2 (a) : Q u e s t i o n n a i r e - L e t t e r  93.  Dear Resident: Birds i n c i t i e s are to some people a source of pleasure, but to others a bothersome pest. Studies have shown that urban b i r d s , notably pigeons, are often c a r r i e r s of diseases deadly to man. As w e l l , these birds sometimes f o u l stored goods, buildings, statues and destroy crops. Thus they not only damage the v i s u a l appearance of c i t i e s but may cause economic l o s s . On the other hand, birds, e s p e c i a l l y songbirds, bring a touch of nature to our c i t i e s which many people cherish. These sympathies and the apparent c o n f l i c t between those who l i k e birds and those who don't, led me to commence research at the University of B r i t i s h Columbia on the ecology of c i t y b i r d s . I hope to obtain information on needs and preferences of birds, which could be of use i n environmental planning. During the past year your neighbourhood has been censused r e g u l a r l y for b i r d s . A study of the types of buildings and amount of d i f f e r e n t types of vegetation has also been made. Perhaps you have noticed either myself or an assistant performing these. Bird censuses w i l l continue through the winter so i f you have not yet seen one of us you may i n the near future. You could help out greatly at t h i s stage by responding to the enclosed questionnaire. I am seeking information on the feeding and housing of birds i n your yard, as w e l l as on your attitude towards b i r d s . The questions are short and e a s i l y answered. A return envelope i s provided for your reply. Your co-operation would be very much appreciated. Thank you.  Richard Lancaster  Appendix 2 ( b ) :  Questionnaire  - Form 94.  QUESTIONNAIRE Please */ where appropriate.  Yes  Do you provide food f o r birds? If so, i s t h i s : store bought bird seed? bread table scraps other About how much of t h i s food do you put out i n an average month? lbs. What kinds of birds have you noticed eating i t ?  2a. What kinds of birds do you most enjoy? b. what kinds of birds do you consider as pests? 3.  Do you have any bird houses on your lot? If yes: How many pairs of birds could these accommodate? How many pairs of birds did these house t h i s summer? What kind(s) of birds were these?  4a. Did any birds nest: i n or on your house or garage? - how many? - what kind? i n your trees, shrubs or garden? - how many? - what kind? b. Did you deter  encourage  or ignore  t h i s nesting?  5a. Would you approve i n p r i n c i p l e of programs to increase the abundance of song birds: i n the c i t y centre? i n r e s i d e n t i a l areas? b. Would you approve i n p r i n c i p l e of programs to decrease the abundance of birds which are sometimes pests: i n the c i t y centre? i n r e s i d e n t i a l areas?  Thank you very much.  No  95. Appendix 3.  Results of B i r d Censuses:  Mean Density of each Bird Species (#/40 Hectares) SPRING Plot Species CP S  DT  WE  QE  MK  CW  17 .8  . 0. 6  SH  UB  244 3 . 79  72 9  150 0  42 8  197 2  57 3  137 4  137 .0  8. 1 136 3  51. 4  2 4 11 9  40 1  136 3  44 0  27 8  CM HS  FC  8. 6 .  1. 6  83 4 1 7  HF  6. 4  4.0  59 8  17 2  13 4  8 3  22 7  AR  1.4  "4. 8  45 5  35 8  45 1  65 7  74 7  1 7  0 7  1 1  GWG  SSP  0 6  -  OJ BCC RST  0. 7  NWC  1.4  BB  1 1  1 1  11 0  0. 6  1 1  13 2  1 2  25 6  0 6  RSF  2 2  3 6  38 8 30 3  VT VGS  32 7  2 2  0. 6 10. 1  9. 3  GCK  3 3  3 3  0. 6  4 5  5.1  WW  4 8 44 8  SJ EG  1. 5  RG  4. 8  BW  3.6  HWK RCK  2.2  3. 7  RH WCS  3.6  1.1  AW BS  4. 8 12. 1  31. 2  3.6  6. 1  2 1.1  1. 5  30. 6  9. 5  CBC  2.4 2.4  nv  0. 6  AG  0. 8  BHC •  0. 8  5.7  GCS  0. 8  1. 2  1. 1  2.9  7.3  1. 1 continued  7.3  4. 5  96. Mean Density of each Bird Species C*/40 Hectares) SPRING (continued) Plot Species  DT  FC  WE 0.8  WWA  QE 2.9  MK  CW  SH  UB  0.6  49.6  46. 2  31. 5  0.6  3.3  1.5  MGW YW  1  1.5  3.6  2.9 2.2  OCW OSF WT  1.7  WV  1.7  3 6 4 8  PSK SVS FS  1.7  FC  0.6  BHG  9 .7  BTP  1 .2 3 .6  ST 0.7  REV V DW PF WF GHO PM PW RBN BT  :  W HT CWX CS .TOTAL  390.0  140.3  583.2  178.3  391.0  366.5  380.3  260.4  97.  Mean Density of each Bird Species (#/40  Hectares)  SUMMER Plot Species CP S  DT  FC  WE  QE  MK  CW  SH  250. 6  113 . 6  143 . 1  0.9  7.4  1.9  22.0  63. 1  17 .2  101 . 1  82.9  178.0  89. 5  97.1  *167.6  47.8  45.6  CM  1.2  HS  51. 9  GWG HF  15. 0 9. 4  AR SSP  13 . 7  UB  4.3 113 . 3  42.1  0,. 5 3..0 - 0,.6  37,. 4 19.•  2  3. 0  2.2  7.3  51. 2  2.2  20. 6  28.9  54.4  98. 7  18.8  4. 7  5.9  .17.4  42. 8  68. 9  1.3  2.2  0.8  OJ BCC  2., 5  RST NWC  22.6 2. 5  1. 8  BB  8.8  7. 0  VT RSF  0.4 .  VGS  8. 6  0.9  1.9  3.5  GCK  7.6  WW  31.2  SJ EG RG  2. 7  BW  6.5  HWK RCK RH  0.4  wcs  1.7  0.4  3. 9  9.3  2.7  AW BS  0. 6  39. 5  27. 0  .17.6  5.4  2.3  1.2  7.5  0.4  1.2  2.7  CBC  TO AG ..  0.9  BHC GCS  0.4  continued  98. Mean Density of each Bird Species (#/40 Hectares) SUMMER (continued) Plot Species  D T  FC  WE  WWA  QE  MK  CW  0.9  MGW  SH  UB  0.8 0.8  YW OCW OSF WT WV PSK  1.1  3.5  SVS FS FC  0.4  BHG  2.7  BTP  0.4  ST  0.4  REV  5.4  V DW PF  0.5  0.4  WF  1.2  0.4  0.5 7.5  GHO • PM  1.1 19.4  0.5  1.3  PW  1.1  RBN  0.5  BT W HT CWX CS TOTAL  393.8  151.6  468.8  168.4  411.5  253.7  396.8  211.5  99.  Mean Density of each Bird Species  (It/40  Hectares)  FALL Plot Species  DT  CP  325 0  S  46 7  . FC 59 0 37 1  WE  QE  CW  SH  UB  4. 2 137 . 3 115. 3  12 .6 93 .6  1.2  30.1  103 .6  41 5  128 . 5 149 7 66. 2  CM  MK  5.2  HS  40. 0 22. 5  GWG HF AR  -  SSP OJ  8.6  121 4 3 0  1.9  44. 0  9.5  4 5  8. 1  9.5 6.9  22 9 7. 0  35.1  8.1  33. 3 142. 6  6.7  6. 9  7.1  11.2  1. 9  29.2  31. 4  4.1  2.6  16. 5  8.4  25. 1  1.0  2.6  7.0 2. 5  5.0  0.9  0. 6  0.8  1. 0  0. 8  BCC  •  RST NWC BB VT  51. 8  VGS GCK  8. 8 9.4  WW 2.5  EG  0.6  BW  3.6  HWK RCK  1.9  0.9  RH  CBC  '  BHC  5.0  2.6  1.0  1  . w 3.1  2.5  5.0  1.9  1-7  6.4  7.5  2.5 1.9  0.9  GCS continued  1.2  0.8 2.5  1-0 5-  1.3  6.0  TW AG  1.2  2.5  RG  BS  44.8 42.4  SJ  AW  173. 2 26.6  6.1  RSF  WCS  15.7  2.4  100 .  Mean.Density of each Bird Species (#/40 Hectares) FALL (continued) Plot Species  DT  FC  WE  QE  WWA  15.5  MGW  0.9  MK 0.6  CW  SH  UB  31.8  37.1  88.4  YW  1.3  OCW OSF  4.4  WT  2.5  WV PSK  7.0  SVS  1.9  FS  0. 6  1.3 0.8  FC  0.6  7.3  1.9  1.2  BHG BTP  20.1  ST  3.6  REV V  0. 8  DW  1.9  PF . WF GHO PM PW RBN  3.1  . BT W  1.0  HT  0.9  1.3  3.8.  3.3  cwx  14.5  CS TOTAL  2.5*  1.2  0.6 435.0  111.3  481.4  173.6  329.2  339.2  472.3  434.  101 Mean Density of each Bird Species (#/40 Hectares) WINTER Plot Species  DT  FC  WE  QE  MK  CP  264 1  57. 4  142 4  S CM  60 4  31. 9  191 4  83 .4 7 .2  8. 8  2 .6  HS  52 1  8. 5  110 1  41 S  GWG  22. 9  6. 1 0. 7 . 1.0  22 6 32 3 34. 2  1 1 11 1 11 5  0. 3  11. 3  12 2  17. 1  HF AR SSP OJ BCC RST NWC  0. 4  CW 6 5  206 . 1 150 9* 0 .4 3 0 121 . 2 48 1 4 .1  1 0 5. 0 43 1  SH  UB  12 .9 88 .9 13 .9  22. 6  14. 3  14 8 21 9  0 3 12 5 47 0 21 0  49. 1  71 7  1. 2  3. 6  10 4  4. 0 0. 5  39 2 1. 0  3. 5  3. 9  8. 2  7. 0  6. 8  3. 0 1. 5  15. 2  21. 8  .2. 4  0. 8  5. 4  128. 4  BB  23 7 31 1  13 . 7 8 1 4 8 124 3 14 1  16. 7  VT  1. 6  0. 7  RSF  1. 9  VGS  0. 4  GCK WW  50. 5  SJ  0. 4  0. 3  EG  1. 7  RG  18. 2 2.4  BW  8.1  HWK  0.8  RCK  0.3  RH WCS AW BS CBC TW AG BHC GCS ^Adjusted from a c t u a l was substituted.  count  cf  3195. The continued  mean  value  for  the  season  102.  Mean  Density  of  each  Bird  Species  (#/40  Hectares)  WINTER (continued)  • Species  DT  FC  WE  .  Plot  QE  MK  CW  SH  UB  WWA MGW YW OCW OSF WT WV PSK  12.5  69.0  SVS FS  1.4  FC BUG BTP ST  . . 0 . 8  REV  •:  V DW  1.0  0.4  PF WF GHO PM PW  0.4  RBN  0.3  BT W  7.4  HT  •..  0.4  CWX  7.5  CS TOTAL  5.1 '  10.5  • 400.0  114.8  567.9  190.9  470.9  366.5  347.3  477.1  Appendix  4:  t>  1  Correlations  c-  1  JO  .0 >  between nj <  +  Habitat  >; z  Features  0 A  aV  • tn  • tn  aA  n. V  tn  tn  W  0  EV  Habitat Features  Significant  M  A  O V  cn  tn  tn  a  M  ++  *  ++  + • ft  +  ++ +  ++  ++ + ++  ++  ++ ++  ++  lit  Appendix  4  (continued) r>  measures  a  a  tn  a  ++  + ++  ++ +  ++  ++ +  ++  ++ ++ ++  @  +'+  @  O  <  ++  + ++  •n o o a  TD  A l l  td  a  o  TD  = Positive = Negative  cn  a  TD  + -  cn  w  IJ T?  Explanation  cn  V M  HD  •SLANT •FLAT ROAD LANE PVT+S LAWN D<7. 5 D>7.5 C<7. 5 C>7. 5 BDEVG " D<1. 5 D>1.5 C<1. 5 C>1. 5 E<1. 5 E>1. 5 HERB WEED FIID1 FHD 2 FIID3 HFD TDD TDC TDE FOOD TOTVEG  t->  EE  Habitat Features  tn  M A  V  +  + ++  + + +  ++  ++  ++  ++  ++ + ++ ++ @  + ++ ++  @  ++ ++ +  *  ++ ++ ++ + ++  ++ ++ ++ +  ft  ++  + ++  + +  +  ++ + ++  *  @  @  -  @  ++ ++ +  ft  ++ ++  ft  + + +  ft  ++ ++  +. ++ ++ ++ + +  @  ++  + ++ ++  ++  ++  +  +  *  8  @  *  -  ++ ++ + + +  *  ft  * *  of  Symbols  (see  also  Table  1)  correlation correlation were  square  •+, - = s i g n i f i c a n t ++, - - = s i g n i f i c a n t @ = correlation root  transformed  prior  to  correlation  a t = .05 a t = .01 coefficient  except  for  greater  FHD, H F D ,  than  TDD,  .9800  TDC,  TDE  105.  Appendix 5 S t a t i s t i c a l Analyses A n a l y s i s of A two  Variance f a c t o r a n a l y s i s of v a r i a n c e  (,Zar, 19 74) was  used  to examine the variance w i t h i n four c h a r a c t e r i s t i c s o f p l o t avifaunas The  (rising the UBC  computing centre program BMD:10V).  c h a r a c t e r i s t i c s analyzed  species  were t o t a l b i r d d e n s i t y ,  d i v e r s i t y , number o f s p e c i e s , and  abundances.  The  nificant  evenness of s p e c i e s '  sources o f v a r i a t i o n were p l o t , season,  the i n t e r a c t i o n of these two. significant  (a = .05), with  one  exception.  There was  the presence of homogeneous subsets  Multiple  sig-  detect  of p l o t s , seasons,  respect to season, f o r the above-mentioned The  no  season.  Duncan's m u l t i p l e range t e s t s were performed, to  teristics.  and  A l l sources of v a r i a t i o n were  change i n evenness with  p l o t s with  bird  r e s u l t s f o l l o w i n Appendix  and charac-  5a.  Regression  A number of m u l t i p l e r e g r e s s i o n s were performed u s i n g the TRIP program, a l s o a v a i l a b l e at the UBC., computing These were forward stepwise r e g r e s s i o n s  and  centre.  derived predic-  t i o n equations f o r the f o l l o w i n g dependent v a r i a b l e s :  den-  s i t y of each s p e c i e s , t o t a l b i r d d e n s i t y , b i r d s p e c i e s  div-  e r s i t y , evenness, number o f s p e c i e s , and species.  average number o f  Four sets of equations were d e r i v e d f o r those  dep-  106 . endent v a r i a b l e s which, were present i n every The  independent  season.  v a r i a b l e s i n these r e g r e s s i o n s c o n s i s t e d  of the percent cover o f v a r i o u s h a b i t a t f e a t u r e s , t r e e dens i t y , t o t a l v e g e t a t i o n cover, amount o f food s u p p l i e d i n t e n t i o n a l l y by man, and a number o f h a b i t a t d i v e r s i t y  indices.  P r i o r to r e g r e s s i o n , a number o f data manipulations were made.  Transformations were used to c o r r e c t d e v i a t i o n s from  linearity.  Both dependent and independent  variables  (except  f o r d i v e r s i t y i n d i c e s and t r e e d e n s i t i e s ) were transformed by t a k i n g the square r o o t .  Test runs were made f i r s t  no t r a n s f o r m a t i o n s , then w i t h only the independent  with  variables  transformed, and compared to runs with both dependent and independent  v a r i a b l e s transformed.  independent  v a r i a b l e s alone improved  Transformations o f the the r e g r e s s i o n equations  2  (higher r ) .  The a d d i t i o n a l t r a n s f o r m a t i o n o f the dependent 2  v a r i a b l e s again i n c r e a s e d the r  i n many cases, but not s i g -  nificantly. A number o f s p e c i e s o c c u r r e d only or most abundantly i n the woodland p l o t .  To d e r i v e r e g r e s s i o n equations f o r these  s p e c i e s I r e g r e s s e d the data from a l l e i g h t p l o t s . the primary purpose  However,  o f t h i s a n a l y s i s was to d e r i v e p r e d i c t i o n  equations f o r those s p e c i e s which predominated  i n urban  areas.  Therefore r e g r e s s i o n equations were a l s o c a l c u l a t e d w i t h the woodland p l o t omitted. f o l l o w i n Appendix 5 b .  These sets o f r e g r e s s i o n equations  10,7.  A  possible  prediction method is  vectors  that  study  Seventy-eight  percent  Winter,  eighty-four  two  percent  o f those  =  which  .05).  most  element  small  deciduous  residents.  percent  abundance (fewer  The i n d i c e s  first  (Hoerl  related  studies.  regression  that  A  and Kennard,  of the species  i n Summer  of occurrence  cases.  was  not a l l orthogonal.  f o r ridge  o f those were  not predicted  h a d a l o w mean  frequency in  Those  regressions  1971) No  com-  at the time  c a r r i e d out.  and  (a  were  f o r future  available  was  i n these  regression  suggested  p r o g r a m was this  of error  (indices)  known as r i d g e  therefore  puter  source  shrubs,  weeds,  in Fall,  regressed  significantly (less than  most  of the regression  encountered  than three  and food  ninety-  significantly were  species  1 p e r 40h) o r a low sightings  frequently equations  and  i n Spring  chosen  were:  provided  per season), as t h e  road, by  trees,  human  108.  Appendix 5a Results o f M u l t i p l e Range Tests  The r e s u l t s o f these t e s t s , as summarized cate s t a t i s t i c a l l y  here, i n d i -  s i g n i f i c a n t s i m i l a r i t i e s among P l o t s and  Seasons with r e s p e c t to the mean values o f the f o l l o w i n g community  indices.  B i r d Species D i v e r s i t y (H ) 1  Evenness (J') Species Richness (S) Total Bird  Density  The f o l l o w i n g a b b r e v i a t i o n s are used. DT - Downtown FC - F a l s e Creek WE - West End QE - Queen E l i z a b e t h MK - MacKenzie CW - Collingwood SH - Shaughnessy UB - U.B.C. p - Spring s - Summer f - Fall w - Winter all  - a l l Seasons  bird  i.  Appendix Duncan's  Multiple  Significance  level  Bird  D i v e r s i t y (II )  Species Groups  Range T e s t  f o r a l ltests:  of Similar FC  2)  MK,  QE  3)  QE,  CW,  4)  UB,  SU  Results  .05  WE  of Similar  Seasons  Summer, F a l l ,  2)  Spring - Season  Winter  Interaction  1)  I'Cfs,  2)  D T ( a l l ) , FC(.p, f )  3)  DT(p,  . 4)'  a =  -  Plots  1)  Plot  (continued)  1  1) . DT,  Groups  5a  Oi).  f) , DT(s,  f,  Groups w)  f , w) , F C ( p , £,  w)  DT(p) ,. FC.Cp, f , w) , MK(.s)  5)  FC(p,  w) , Q E ( s ) , M K ( s ,  6) .  QE(.s), MK(s,  7)  Wli(f),  8)  Wli(all),  9)  WIi(p, s , w ) , . Q E ( p ,  f) , UB(f) '  f ) , CW(w) , U B ( f )  Ql-:(s, w ) ,  MK(p,  f , w)., CVv(w) , UB ( f )  Ql.!(p, ,.f, w) , MK(|»,  10)  C W ( f ) , SI I ( a l l ) . ,  11)  SI I ( a l i ) , U B ( p ,  f ) , CW(p,  UB(s) s)  f, w),  CW(.r.l.l),  f ) ,SH(all),  l)B(. f ,  UB(vv)  w)  110.  Evenness (J*) Groups o f S i m i l a r  Plots  1)  MK, FC  2)  FC, CW  3)  CW, UB, QE, SH, WE  4)  DT (lowest)  (highest)  Groups of S i m i l a r Seasons 1)  Spring,  Summer, F a l l , Winter  P l o t - Season I n t e r a c t i o n  Groups  1)  D T ( a l l ) , FC ( s ) , MK(s), UB (f)  2)  DT(p, w), F C ( s ) , MK(s), CW(w) , UB (f)  3)  DT(p, w), FC(p, f , w), WE(f), QE(s), CW(p,  4)  f , w), SH(p, f ) , UB(f)  DT(p, w), FC(p, f , w), WE(f), QE (s, f , w) ,MKCp,  5)  f , w), CWCp, f , w), SH(p, f ) , UBCf, w)  DT(p), FC(p, f , w), WE(p, f , w), QE(_all), MKCp, f , w) , CW(_all),  6)  MK(all),  S H ( a l l ) , UBCp, f , w)  FC(p, f , w), WE C a l l ) , QE Cp, £, w) , MKCp, f) , CW(p,  s, f ) , S H ( a l l ) , UB(p, s, w)  Species Richness (S) Groups o f S i m i l a r 1)  FC, DT  2)  QE, MK, WE  3)  MK, WE, CW  Plots  111.  4)  UB  5)  SH  Groups o f S i m i l a r Seasons 1) 2)  Summer, Winter, F a l l Spring  P l o t - Season I n t e r a c t i o n  Groups  1)  DT(p, f , w), F C ( a l l )  2)  D T ( a l l ) , FC(.P, W )  3)  DT (p, s, f) , FC(p, f , v\,  4)  DT(p, s ) , QE(p, s ) , MK(s), CW(s)  5)  DT(s), WECs), QE(p, s, w) , MK(s, f) , CWCs)  6)  WE (s , £, w) , Q E ( a l l ) , MK (p, s, f) , ..CW(s , w) , UB(f)  7)  WE C a l l ) ,  8)  WE(p, f , w), Q E ( f ) , MK(jp, f , w) , CW(p, £, w)., SH(s), UB(f,  9)  QECp,  QECs)  f , w) , MK(jp, f , w), CW(w) , UB (f).  w)  MK(w), CW(p, f ) , SH(s, w), UB(s, f , w)  10)  SH(£, w), UB(s)  11)  SH(f),  12)  SH(p, f) , UB(.p)  UB(p, s)  T o t a l B i r d Density Groups o f S i m i l a r  Plots  1)  FC, QE  2)  UB, CW, SH, MK, DT  3)  WE  112.  Groups  of Similar  Seasons  1)  Summer,  Spring,  2)  Spring,  Fall,  Plot  - Season  1)  2)  Interaction  s , W),  CW(p,  s , f ) , SHCp,  WE(s,  f ) , QECp,  QE(all),  MK(.p, s , f ) ,  W)  f , w), N K C a l l ) ,  UBCall),  :  CW(p,  s,  f) ,  DTCall) MKCall),  CW(.p,  f) ,  SH(all),  f , w)  DT(all), UB(f,  Groups  FC(all),  DT(all) , WECall), UB(p,  4)  Winter  DTCp,  SH(all), 3)  Fall  w)  WECall),  MK(p,  s , w) , CW(w) , S H C p ,  s, f) ,  113.  Appendix Multiple Regression Results  from  Spring,  Summer,  There  were  Species  CP  s  CM  a l l  28  HF  Fall,  seven  for  each  R e s u l t s : ...  plots;  U.B.C.  species  are  plot  omitted  presented  independent  probability  variables  in  this  Equation (variables least significant)  ranked  .00 .01  1 4 .. 99 1 3 .. 0 8  -  1 1 .. 7 0 D * 1 . 5 1 5 ,. 9 5 D > 1 . 5  +  24.24  . 81 .95  .01 .00  1 5 ,. 6 7 7.. 0 9  -  1 2 ,, 9 3 D > 1 . 5 1 9 ., 92 D > 1 . 5 +  33.87  . 75 .66 . 72 .57  . 01 .03 .02 .05  7,. 55 2.. 1 1 6, . 9 6 5. . 2 2  .92 . 72 .93  .01 .02 . .01 N.S.  1. , 4 1 2. . 4 4 1. . 3 4  + 0 .. 2 9 + 1 7 ,, 1 3 + 0 ,. 2 5 + 1 1 .. 2 3 _  -  6 . . 11 E < 1 . 5 + 1.06 3. , 1 6 WEED 3. . 3 0 W E E D + 2 . 3 9  . 80  - 9 ., 58 +  7. . 1 1  LANE  .96 .99 .63 . 78  .00 .01 .03 .01  - 5 ., 33 + - 9 . 42 + 5. , 0 6 6., 85 -  1. . 1 9 1. 4 9 3 . , 34 3. 61  PVTS-S PVT5S HERB HERB  + +  +  30  TDC  2.69  - 6 . 08 + 0 ., 0 8 + 0 ., 7 8 +  4. 46 28 23  .98 . 72 . 79  .00 .02 .01  the  order:  4. 93  most  E<1.S C<1.5  (see  Table  significant  -  1).  to  6.17  E<1.5  RDC E<1.S TDC D<1. 5  N.S. N.S. N.S. .01  .01  in  analysis  . 84 1.00  1.00  here  Winter.  potential  R  HS  GWG  seasons  on  Regression  5b  -  BDEVG TDC TDC  +  3.90 12.87  HERB C<1.5  D<1.5 E<1.S  -  1.82  WEED 1.2 0 HERB . 1 3 LAWN 3.84 LANE  E>1.5  -  114.  Species  R  2  probability  v  A  R  _ oo ioo 96 [99  .01 00 .00 . 00  4.49 -.54 -.78 - 2 . 53  + +. + +  .69  .02  0.03  +  -  SSP  Equation (variables least significant  ranked  most  7 . 8 2 FHD2 + 1 . 0 8 TDC 5 . 5 4 0 1 . . 5 + . 3 4 LAWN 7.57 C>7.5 . 2 2 T D C .+ 3 . 6 9 H E R B .07  significant . .  to  2.28 HFD  _ _ _ _ _ _  TDC  , -  • 1.00 .96  .00 .00  6.77 -2.02  +  1.06 3.09  PVTSS - 1.83 D>1.5 .' H E R B + .. 1 2 T D C  .97 1.00  .00 .01  12.40 -5.26  +  2.92 .14  ROAD + . 7 4 TDC + 1 .44  .91 1.00 .99 . 74  .00 .00 .00 .01  -.80 -.81 -.39 -1.02  + + + +  3.86 2.86 .20 1.76  E>1.5 C>7.5 + .80 FOOD + 1 . 3 9 D<7.5  .93  .01  0.50  -  1.22  WEED  1.00  .01  -.64  +  .72  .01 .00 .02 .00  - .65 + 6.60 -.14 + -.40 +  3.90 2.42 .17 4.81  0.31 + 1.34 + 0.81 + -1.21 +  .39 .27 .16 .38  FOOD FOOD FOOD FOOD  B D E V G •-  +  1.24  D<1.5  OJ  BCC  RST  NWC  . 83.85 . 70 . .93  .88  +  D<7.5  +  D>7.5 D<7...5 +  2 . 0 5 HERB  D<7.5 C>7.5 +  1.14  BDEVG  C<1.5  .09  FLAT  - .1.6.3 E < 1 . 5  -  5.89 C<1.5 + .13 FLAT . 3 2 LAWN - . 7 1 H F D . 1 9 LAWN - . 4 3 H F D . 2 7 T O T V E G - . 1 8 LAWN  1.00 1.00 1.00 1.00  .00 .08 .01 .01  VT  1.00  .00  0.49  +  .82  1.00  .01  -6.59  -  1.42  LAWN +  1.00 1.00 .85 . 71  .01 .00 .00 .02  .08 .35 3.20 .59  F O O D - . 0 9 LAWN - . 2 0 H F D BDEVG - .18 LANE + .10 HFD E>1.5 D<7.5  0.38 + 0.21 + -.51 + -.45 +  +. . .  E>1.5 LANE FOOD D<1.5  B B '.  RSF  + .-. .  .42  1.61  LANE  +  '.23 H F D  TOTVEG + 2 . 5 3  LANE  115.  Species  R  VGS  1.00 1.00  .01 .00  1.00  .01  .19  +  .97  .00  - .57  +  3.18 E>1. 5  1.00 .94  .00 .01  1 .02 - .41  +  1. 70 BDEVG • .8 7 LANE + .49 HFD .60 TOTVEG - .64 LAWN  .25  +  GCK  2  probability  . 73  RCK  WCS  AW  BS  +  .98 LAWN - 4.01 D>1.5 - 1.47 WEED.10 TDC + 18 D>7.5 + .01 TDD .06  FOOD - .04 TOTVEG -* .03 LAWN +  .02 N .S.  _  . -45 D>7.5  1.00  .00  0.40  +  .67 BDEVG - .34 LANE + .19 HFD  1. 00 1.00  .00 .00  0.53 0.43  + +  .87 BDEVG .45 LANE + .25 HFD . 72 BDEVG - .37 LANE + .21 HFD  .00  - .58  +  0.18  +  0.31 0.92  +  -  .06 FOOD - .07 LAWN - .16 HFD 3.17 WEED + .91 FHD3 + .18 D>7.5  .02  1 .14  -  1.90 WEED + .05 FOOD  . S5  .00  - .37  +  1.69 E>1. 5  .97  .00  - .65  +  6.80 D<1.5 • 3.61 C>1.5  .98 • .81 1.00  .00 .01 .01  13 .43 0.62 -1 .11  + .  6. 75 FHD3 + 3.23 LANE . 26 TDC 7. 50 D<1.5 - .61 D<7.5 + .90 WEED -  . 88 1.00  RH  1 . 73 + - .21 +  ---  SJ  EG  Equation (variables ranked most s i g n i f i c a n t to least s i g n i f i c a n t  N  .s.  .00  :  2.61 E>1.5 .30 BDEVG • .16  LANE + .09 HFD  .  1.00 1.00 . 88  .01 .00 N  .s.  N  .s.  + +  •  116.  Species  R  TW  probability  Equation (variables least significant  ranked  most  N.S. 1.00  .00  AG  .91 .74 .93  .01 .01 .01  -2.19 + 4.77 0.55 -  8 . 9 5 C<1.5 - . 1 0 TDC 1 . 1 8 RAOD 1 . 3 5 WEED + . 9 8 C < 1 . 5  BHC  .87 .93  .00 .00  -5.24 + -.33 +  3.20 HFD 1.41 E>1.5  GCS 1.00  WWA  1.00  MGW  '.  N.S. -01  0.31  +  +  - -1.89 +  1.001.00 .93  .01 .00 .01  0.40 + 0.30 + 0.55 -  . 75  . 01  1.00  OCW  0.46  .01 N.S. .00  .86  YW  significant  1.00  .00  -1.85 +  .  - 1 . 32 +  .76 BDEVG  .06  FOOD  2 . 0 3 D>7.5 2.43  -  -  .39 LANE  . 0 7 LAWN  -  +  -  .22 HFD  .16 HFD -  .09 TDC +  1.07  D<7.5  D<7.5  .67 BDEVG - .34 LANE +..19 HFD .49 BDEVG - . 2 5 LANE + .14 HFD 1 . 3 5 WEED + . 9 8 C < 1 . 5  1. 50  HERB  0.37 +  .62 BDEVG  -  .32  LANE  +  .18 HFD  0.57  .95 BDEVG  -  .49  LANE  +  .27 HFD  +  +  to  117.  Species  R  probability  Equation (variables . least significant ji  ,  OSF  ranked  most  significant  :  1.00  .00  0.49  +  .82  BDEVG  -  .42  LANE  +  .23  HFD  1.00  .00  0.70  +  1.16  BDEVG  -  .60  LANE  +  .33  HFD  -2.92 0.16  + +  2.49 .32  LANE + FOOD -  -.27  +  .08  FOOD  WT  N.S. N.S.  WV  N.S.  PSK .99 .99  . 01 .01 N.S.  1.08 BDEVG . 6 9 LAWN +  - 1.39 HFD 3.02 C>1.5  SVS .66  FS  .03  N.S. 1.00 1.00  .00 .00  0.41 0.39  + +  .08 .65  F O O D - . 1 0 LAWN + . 2 1 H F D BDEVG - .33 LANE + .19 HFD  1.00 1.00  N.S. .00 .00  0.21 0.26  + +  .35 .44  BDEVG BDEVG  .68 .60  --.02 .04  -2.04 -.61  + +  1.23 1.10  HFD D>7.5  FC  BTP  -  .18 .22  LANE LANE  + +  .10 .13  HFD HFD  to  118.  Species  R  V  probability  1.00  .00  Equation (Variables least significant -  ranked  .24  most  0.28  +  .47  BDEVG  LANE  .01 N.S.  0 . 70  +  . 14  FOOD  .73  --.02  . ' 3.21  -  .79  ROAD  1.00  .01  - 2 . 85  +  .45  PVT5S  +  3.93  1.00. 1.00  .00 .00  0.59 0.18  + +  .98 .30  BDEVG BDEVG  -  .50 .16  .00  1.39  +  1.25  HERB  -  .92  LANE  N.S. .01  -.98  +  .84  LANE  +  .36  BDEVG  +  significant . . .14  to  HFD  N.S.  DW 1-00  PF  -  .17  L A M  -  .37  HFD  PM E<1.5  -  .25  C>7.5  RBN  BT  LANE LANE  + +  .28 .09  HFD HFD  -  .47  HFD  ---  N.S. W -  --•97  . •99 CWX  '  N.S.  -  119.  Species  R  probability  Equation (variables least significant  ranked  most  significant  to  cs 1,. 0 0  #/40 ,88  BSD  ,95 93 98 92  .01  0 .41  N.S. N.S. .02 N.S.  +  -1672 .69  .00 .00 .00 .00  1,. 0 8 0, . 89 0, . 6 1 1.. 0 8  .08  FOOD  + 1210.83  + + + . +  . 56. .02 .21 .58  -  .10  LAWN  HFD -  C<7.5 TDD TOTVEG C<7.5  -  -  .21  HFD  -  1 3 3 . 0 1 D>7. 5  . 0 3 FOOD  STOT  SAV  J  Note:  99 1. 00 93  .0.1 .01 .00  1. 00 74 96 • 85  .01 .01 .00 .00  4 .. 1 1 4 ., 3 4 1., 1 1 3 ., 4 3  1. 00 68 1. 00 • 71  .01 .02 .01 .02  0. 0. 0. 0.  1 1 ., 5 9 + 9 .80 1 0 ..20 + 21 . 5 1 3.. 7 8 + 1 4 . 0 8 + + + +  51 + 46 + 48 62 +  2 2 5 5  C>7.5 E>1.5 FHD2  +  6.95 2.10  WEED L A M  .93 . 74 .24 . 55  BDEVG + C>7. 5 E>1.5 + FHD2  5.82  D<1.5  8.60  C<1. 5  . 50 .01 .02 .05  C < 1 . 5 - . 0 0 3 TDD + . 0 7 . B D E V G TDD FOOD + . 0 4 LAWN + . 0 8 E > 1 . 5 + C<7. 5  -  1.36 D>7.5 1 0 . 4 1 FHD3 +  -  2.87  C<1.5  The f o l l o w i n g Summer: FHD2 variables.  w a r n i n g messages were generated by t h e computer. was a l i n e a r c o m b i n a t i o n o f o t h e r independent  Fall: C<7.5 variables.  was  a  linear  combination  of  other  independent  +..  

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