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Do  not  assume  content  reflects  current 
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1 

1 


/]rt.i  mm 

Effects  of  Harvesting  Ponderosa  Pine 
on  Nongame  Bird  Populations 


Robert  C.  Szaro  and  Russell  P.  Balda 


Research  Paper  RM-212 
Rocky  Mountain  Forest  and 
Range  Experiment  Station 
Forest  Service 

U.S.  Department  of  Agriculture 


Research  Paper  RM-212 


December  1979 


Effects  of  Harvesting  Ponderosa  Pine 
on  Nongame  Bird  Populations 


^  Robert  C.  Szaro,  Research  Wildlife  Biologist 
Rocky  Mountain  Forest  and  Range  Experiment  Station^ 

and 

Russell  P.  Balda,  Professor  of  Biology 
Northern  Arizona  University 


Abstract 

Bird  species  diversity  and  species  richness  in  the  ponderosa  pine 
forest  were  not  significantly  affected  by  forest  cutting  and  logging 
except  on  the  clearcut  plot.  Bird  population  densities  were  signifi- 
cantly increased  on  the  silviculturally  cut  and  irregular  strip  shelter- 
wood  plots  and  were  significantly  decreased  on  the  severely  thinned 
and  clearcut  plots.  Guidelines  are  recommended  that  will  allow  sub- 
stantial logging  of  the  ponderosa  pine  forest  while  still  maintaining 
bird  density,  diversity,  and  species  richness. 


'Central  headquarters  maintained  at  Fort  Collins  in  cooperation  with  Colorado  State  Univer- 
sity. Szaro  is  at  the  Station's  Research  Work  Unit  at  Tempe  in  cooperation  with  Arizona  State 
University.  Study  was  conducted  under  graduate  program  at  Northern  Arizona  University. 


Effects  of  Harvesting  Ponderosa  Pine 
on  Nongame  Bird  Populations  , 

Robert  C.  Szaro  and  Russell  P.  Balda 


Management  Implications 


The  forest  manager  can  remove  between  one- 
sixth  and  two-thirds  of  the  available  foliage  of  the 
ponderosa  pine  forest  either  in  strips  (and  prob- 
ably in  blocks)  or  by  thinning  without  detrimen- 
tally affecting  the  breeding  bird  community  in 
terms  of  species  richness,  density,  and  diversity. 
Increased  densities  on  silviculturally  cut  and  ir- 
regular strip  shelterwood  plots  are  probably  at 
least  partially  due  to  openings  (MacArthur  et  al. 
1962;  Marshall  1957,  1963).  However,  forest 
treatments  must  consider  that  the  quality  of  the 
bird  community  on  cut  and  control  areas  are  not 
equal.  Species  found  on  the  control  plot,  such  as 
the  hermit  thrush,  red-faced  warbler,  western  fly- 
catcher, and  pygmy  nuthatch,  are  replaced  on  the 
cut  areas  by  species  such  as  the  western  wood 
pewee,  yellow-rumped  warbler,  and  rock  wren. 

When  forests  are  managed  for  tree  and/or 
water  yield,  some  specific  guidelines  can  be 
followed  to  minimize  the  impact  of  habitat  modifi- 
cation on  bird  populations.  To  simply  state  that 
foliage  volume  can  be  reduced  by  one-sixth  to 
two-thirds  is  of  little  real  significance  to  forest 
managers.  Foliage  volume  is  important  to  the 
birds  because  it  is  related  to  the  resource  base 
but  is  difficult  and  time  consuming  to  measure. 
Therefore,  to  maintain  and/or  increase  (by  up  to 
35%)  ponderosa  pine  forest  bird  populations,  the 
following  guidelines  are  recommended: 


1.  The  total  basal  area  of  a  given  stand  can  be 
reduced  by  15%  to  50%.  However,  large- 
scale  removal  should  be  in  strips  or  blocks. 
In  a  uniform  thinning  operation,  consider 
removing  only  30%  of  the  total  basal  area. 

2.  Remove  no  more  than  45%  of  those  trees  with 
a  d.b.h.  of  9  inches  or  greater.  Leave  a 
minimum  of  32  trees  per  acre. 

3.  Remove  no  more  than  75%  of  those  trees  with 
a  d.b.h.  between  6  and  9  inches.  Leave  a 
minimum  of  17  trees  per  acre. 

4.  Remove  80%  of  the  trees  with  a  d.b.h.  be- 
tween 3  and  6  inches  leaving  approximately 
25  trees  per  acre. 

5.  Gambel  oak  should  not  be  removed  at  all.  If 
absolutely  necessary,  remove  no  more  than 
25%  of  the  oaks. 

6.  Several  overmature  trees  per  acre  should  be 
left  to  allow  for  adequate  snag  recruitment. 

7.  Snags  should  be  left  as  nesting  and  roosting 
sites  for  cavity  nesters.  Balda  (1975)  sug- 
gests 2.6  snags  per  acre. 

These  guidelines  are  based  on  a  comparison  be- 
tween the  control  plot  and  the  silviculturally  cut 
and  the  irregular  strip  shelterwood  plots. 


Introduction 

Avian  ecologists  have  long  been  interested  in 
relating  breeding  bird  populations  to  the  vegeta- 
tion of  an  area  (Beecher  1942,  Johnston  and  Odum 
1956,  Bond  1957).  As  the  structure  of  a  habitat 
becomes  more  complex,  the  number  of  different 
bird  species  increases  (Karr  1968,  MacArthur 
and  MacArthur  1961,  MacArthur  et  al.  1966, 
Recher  1969).  The  population  density  of  black- 
burnian  warblers^  and  myrtle  warblers  appears 
to  be  closely  correlated  with  foliage  volume 
(MacArthur  1958).  Moreover,  foliage  volume  may 

'Common  and  scientific  names  of  all  birds  and  trees 
referred  to  in  this  paper  are  listed  in  the  appendix. 


be  an  important  factor  limiting  the  densities  of 
parula  warblers  and  nuthatches  (Balda  1969, 
Morse  1967).  Data  by  Balda  (1969)  strongly  sug- 
gest that  removing  tall  ponderosa  pines  (40  to  70 
feet)  may  have  a  negative  effect  on  the  density  of 
Grace's  warblers;  whereas  the  removal  of  the 
understory  may  reduce  the  populations  of  the 
gray-headed  junco  and  the  chipping  sparrow. 

Bird  population  densities  in  a  particular  habitat 
are  believed  to  be  regulated  by  many  factors.  Any 
alteration  of  that  habitat  may  affect  the  suitability 
of  the  habitat  for  a  given  species'  niche  require- 
ments. This  study  examined  effects  of  timber 
management  practices  on  bird  populations  and 
ways  these  practices  can  be  used  to  manage  non- 
game  bird  populations. 


1 


study  Areas 

Five  study  plots  were  chosen  in  relatively  homo- 
geneous stands  of  ponderosa  pine  v^ith  a  buffer 
around  the  periphery  of  at  least  330  feet.  Study 
plots  contained  about  the  same  proportions  of  dif- 
ferent size  classes  of  trees  and  density  of  Gambel 
oak.  All  study  areas  were  set  up  as  35-acre  plots 
except  for  the  study  area  on  the  clearcut  water- 
shed, which  encompassed  100  acres. 

The  five  study  areas  are  in  the  Coconino  Na- 
tional Forest,  Coconino  County,  Arizona.  All  the 
areas  are  located  within  a  13-mile  radius  on  the 
Beaver  Creek  Watershed.  The  areas  included  a 
clearcut,  a  severely  thinned,  an  irregular  strip 
shelterwood,  a  silviculturally  cut  (individual  tree 
selection],  and  a  control  plot.  All  study  sites  were 
cut  before  the  study  began  except  for  the  silvicul- 
turally cut  area,  which  was  cut  during  the  spring 
of  1974. 

The  ponderosa  pine  vegetation  type,  which  was 
found  on  all  study  areas  before  treatment,  is 
found  primarily  in  areas  of  brolliar,  siesta,  and 
sponsellar  soils  (Williams  and  Anderson  1967). 


Control  Plot 

The  control  is  located  on  watershed  13  approx- 
imately 41  miles  southeast  of  Flagstaff  at  an  ele- 
vation of  7,200  feet.  The  study  area  is  on  a  south- 
west-facing slope  of  about  17°,  in  the  west-central 
portion  of  the  368-acre  watershed. 

Watershed  13  was  left  untreated  as  the  control 
area.  Ponderosa  pine  was  the  dominant  tree  spe- 
cies with  an  importance  value  ^of  253  (table  1]. 
There  were  approximately  262  trees  per  acre 
with  a  canopy  volume  of  276,800  cubic  feet  per 
acre  and  a  total  basal  area  of  116.3  square  feet 
per  acre.  Of  the  trees  of  the  plot,  78%  had  a  d.b.h. 
of  9  inches  or  smaller  (table  2).  In  fact,  the  control 
plot  had  3.7  times  as  many  trees  with  a  d.b.h.  be- 
tween 3  and  6  inches  than  any  other  study  plot 
(table  2). 


Silviculturally  Cut  Plot 

The  silviculturally  cut  plot  is  located  on  water- 
shed 8,  approximately  39  miles  southeast  of  Flag- 
staff at  an  elevation  of  7,400  feet.  The  study  area 
is  on  a  west-facing  slope  of  about  13°,  in  the 
southwest  corner  of  the  1,800-acre  watershed. 

The  prescription  called  for  stands  made  up  of 
trees  smaller  than  10  inches  d.b.h.  to  be  thinned 
to  a  growing  stock  level  of  60  square  feet  per  acre 


of  basal  area.^  Stands  consisting  of  trees  12  inches 
d.b.h.  and  larger  were  thinned  to  an  actual  70 
square  feet  per  acre  of  basal  area.  Trees  were  cut 
to  upgrade  the  stand  rather  than  to  obtain  uniform 
spacing.  In  most  cases,  Gambel  oak  were  left  intact. 

The  treatment  was  completed  in  early  spring 
1974;  ponderosa  pine  was  the  major  dominant 
tree  species  with  an  importance  value  of  263.4 
(table  1].  There  were  approximately  96  trees  per 
acre  with  a  canopy  volume  of  243,500  cubic  feet 
per  acre.  This  amounted  to  a  reduction  of  28.9% 
in  the  available  foliage.  The  total  basal  area  for 
all  tree  species  was  101.5  square  feet  per  acre. 

Irregular  Strip  Shelterwood  Plot 

The  irregular  strip  shelterwood  cut  plot  is 
located  on  watershed  14,  approximately  42  miles 
southeast  of  Flagstaff,  at  an  elevation  of  7,050 
feet.  The  study  area  is  on  a  south-facing  slope  of 
about  9°,  in  the  southeast  corner  of  the  546-acre 
watershed. 

The  objective  of  the  treatment  was  to  increase 
water  yield  while  at  the  same  time  providing  good 
timber  production  and  pleasing  esthetics  (Brown 
et  al.  1974].  Clearcut  strips  were  designed  pri- 
marily to  increase  streamflow.  The  alternative 
"leave"  strips  were  thinned  to  improve  production. 

The  pattern  was  one  of  alternate  cut  and  leave 
strips.  The  cut  and  leave  strips  averaged  60  and 
120  feet  in  width,  respectively.  Irregular-shaped 
spacers  of  uncut  trees,  50  to  70  feet  long,  at  inter- 
vals of  about  400  feet,  were  left  in  the  cut  strips  to 
break  up  the  visual  continuity.  Most  of  the  Gambel 
oak  were  left  in  the  cut  strips;  where  there  was 
enough  oak  to  break  up  the  continuity  of  the  strips 
it  was  not  necessary  to  use  spacers.  Width  of  the 
clearcut  area  within  any  strip  varied  as  much  as 
50%  (i.e.,  120  ±  60  feet]  to  provide  an  estheti- 
cally  pleasing,  irregular  pattern  of  elongated 
openings. 

The  treatment  was  completed  in  spring  1970. 
Ponderosa  pine  was  the  dominant  tree  species 
with  an  importance  value  of  228.2  (table  1].  There 
were  approximately  74  trees  per  acre  with  a 
canopy  volume  of  92,700  cubic  feet  per  acre  and  a 
total  basal  area  of  54  square  feet  per  acre. 


Severely  Thinned  Plot 

The  severely  thinned  plot  is  located  on  water- 
shed 17,  approximately  27  miles  south  of  Flagstaff 
at  an  elevation  of  6,860  feet.  The  study  area  is  on 

^Personal  communication  with  Fred  Larson,  Researcli 
Forester,  USDA  Forest  Service,  Flagstaff,  Ariz. 


2 


Table  1.— Composition  of  trees  on  all  forested  study  areas 

Species  Relative 

Relative 

Relative 

Importance 

Absolute 

Total  foliage 

density 

dominance  frequency 

value 

density 

volume 

[perceni) 

(percent) 

(percent) 

(inoex) 

{Trees  per  acre) 

{11  /acrej 

Control 

1      luci  uod  ^Jiiic;        s/U.  1 

OO.  f 

/  /.U 

o'XA.  Qnn 
^o^,yuu 

<■>  nr\  \r\r\  1        o                                  Q  A 

OdllliJcl  UdK  0.*f 

8.3 

19.3 

OD.U 

OB  7nn 
^o,  /  UU 

Alligator  juniper  1.5 

6.0 

3.7 

11.2 

4 

13,200 

^ i t\/i 1 1 1  f  1 1     1 1\/  ^iit 
oi  1  vlou  1  lu  1  cii  1  y  oui 

rUllUcrUba  piilc         y  1 .0 

7Q  A 

OO 

ono  i^nn 

VJIctlllUcI  UdtS  O.U 

7.5 

20.6 

OO.D 

Q 
O 

4  1  ,uuu 

Irregular  strip  shelterwood 

1    UllUCIUOCt    pilIC              f  s7.  1 

o2.0 

67.1 

oy 

CO  nnn 

15.7 

31.5 

ft7  R 

1  /I 
1 4 

'3n  inn 

Alligator  juniper  0.5 

2.3 

1.4 

4.2 

1 

400 

Severely  thinned 

Ponderosa  86.8 

91.9 

74.3 

253.0 

24 

48,500 

Gannbel  oak  13.2 

8.1 

25.7 

47.0 

4 

8,600 

Table  2.— Tree  size  distribution  on  all  forested  study  areas  (trees  per  acre) 

Over 


Area 

3-6  inches 
d.b.h. 

Percent 
control 

6-9  inches 
d.b.h. 

Percent 
control 

9  inches 
d.b.h. 

Percent 
control 

Absolute 
density 

Percent 
control 

Control 

135.5 

68.1 

58.4 

262 

Silvlculturally 
cut 

Irregular  strip 
shelterwood 
Severely 

36.7 

27.1 

15.5 

22.8 

43.8 

75.0 

96 

36.6 

24.9 
6.2 

18.4 
4.6 

17.2 
9.0 

25.3 
13.2 

31.9 
12.8 

54.6 
21.9 

74 
28 

28.2 
10.7 

thinned 


a  southwest-facing  slope  of  about  8°,  in  the  south- 
west corner  of  the  121-acre  watershed. 

Treatment  was  intended  to  provide  a  reason- 
able opportunity  for  increased  water  yield  while 
leaving  a  lightly  stocked  timber  stand  that  could 
be  subjected  to  even-aged  management  (Brown  et 
al.  1974).  Slash  was  piled  in  strategically  arranged 
windrows.  Windrows  were  piled  as  high  and  nar- 
row as  possible  to  maximize  snow  trapping  and 
retention.  Windrows  were  arranged  with  30-foot 
breaks  at  intervals  of  200  feet  or  less  to  reduce 
possible  fire  spread. 

Treatment  was  completed  in  spring  1969.  Pon- 
derosa pine  was  the  dominant  tree  species  with 
an  importance  value  of  253  (table  1).  There  were 
approximately  28  trees  per  acre  with  a  canopy 
volume  of  57,100  cubic  feet  per  acre  and  a  total 
basal  area  of  22.2  square  feet  per  acre. 

Clearcut  Study  Plot 

The  clearcut  plot  is  located  on  watershed  12, 
approximately  43  miles  southeast  of  Flagstaff  at 


an  elevation  of  7,040  feet.  The  study  area  is  on  a 
southwest-facing  slope  of  about  10°,  in  the  south- 
east corner  of  the  200-acre  watershed. 

The  treatment  was  designed  to  test  the  effects 
of  clearcutting  all  the  woody  vegetation  on  the 
watershed  and  windrowing  the  resultant  slash 
(Brown  et  al.  1974).  All  wood  products  that  could 
be  sold  were  removed  from  the  watershed.  The 
remaining  slash  and  debris  were  machine  wind- 
rowed  in  such  a  way  as  to  trap  and  retain  snow, 
reduce  evapotranspiration  losses,  and  increase 
the  drainage  efficiency  of  the  watershed.  In  areas 
of  heavy  slash,  the  windrows  were  at  least  5  feet 
high  and  were  spaced  about  100  feet  apart.  In 
areas  of  lighter  slash,  the  windrows  were  spaced 
further  apart  to  achieve  the  minimum  height. 
Windrows  were  placed  in  either  an  east-west  or 
northeast-southwest  direction. 

The  treatment  was  completed  in  spring  1967. 
Since  that  time,  there  has  been  considerable 
shrubby  growth  by  Gambel  oak  next  to  the  slash 
windrows. 


3 


Methods  and  Materials 

Tree  measurements  were  made  on  all  plots  ex- 
cept the  clearcut  site.  The  plotless  point-quarter 
method  (Cottam  and  Curtis  1956)  was  used  to  sam- 
ple trees  with  a  d.b.h.  of  3  inches  or  larger.  A  grid 
composed  of  104  points  (416  trees)  was  sampled 
on  each  plot.  These  data  were  then  analyzed 
using  the  standard  formulas  of  Cottam  and  Curtis 
(1956)  to  obtain  the  following:  absolute  density, 
relative  dominance,  relative  frequency,  relative 
density,  and  importance  value.  The  following  ad- 
ditional data  were  also  recorded  for  the  trees 
sampled  at  each  point:  total  tree  height,  height 
from  the  ground  to  the  lowest  live  limb,  and  outer 
crown  diameter  at  the  lowest  live  limb.  Tree 
crowns  were  classified  as  conical,  cylindrical,  or 
hemispherical.  Tree  crown  data  were  then  ana- 
lyzed and  expressed  in  terms  of  foliage  (or  crown) 
volume. 

Breeding  bird  counts  were  made  during  the 
1974  and  1975  breeding  seasons  using  the  spot- 
map  method  described  by  Kendeigh  (1944).  Ten 
censuses  were  taken  each  year  on  each  study  site. 
Population  densities  were  averaged  for  the  2-year 
period  to  eliminate  effects  of  climatic  fluctua- 
tions. 

Species  diversity  (H')  (Shannon  and  Weaver 
1948)  was  calculated  on  the  mean  densities  for  all 
plots  by  the  following  formula: 

H'  =  -IF.  In  (Pi ) 

where  F-  is  the  proportion  of  a  given  bird  species 
present.  Evenness  (E)  was  calculated  by  the  fol- 
lowing: 

E  =  H'/ln  S 

where  S  is  the  number  of  species  present  (rich- 
ness). 

Bird  Community  Composition 

The  effects  of  habitat  alteration  on  species 
composition  and  densities  have  been  examined  in 
areas  where  the  habitat  was  altered  by  logging 
(Hagar  1960;  Kilgore  1971;  Lack  1933,  1939;  Lack 
and  Lack  1951),  burning  (Blackford  1955,  Bock 
and  Lynch  1970,  Marshall  1957)  and  other  means 
(Karr  1968,  Yeager  1955).  The  effects  of  the  vari- 
ous treatments  on  the  breeding  bird  communities 
of  the  clearcut,  severely  thinned,  irregular  strip 
shelterwood,  and  silviculturally  cut  plots  were 
pronounced  (table  3).  The  openings  made  by  cut- 
ting led  to  an  increase  of  those  species  which 
appear  to  require  a  more  open  habitat  (rock  wren, 
robin,  western  wood  pewee,  and  yellow-rumped 
warbler)  and  a  decrease  or  elimination  of  those 


species  which  appear  to  require  dense  foliage 
(western  flycatcher,  red-faced  warbler,  hermit 
thrush,  black-headed  grosbeak,  and  pygmy  nut- 
hatch). Cutting  the  irregular  strip  shelterwood 
and  silviculturally  cut  sites  increased  population 
density  and  slightly  changed  species  composition 
when  compared  to  the  control  site. 

These  results  tend  to  contradict  the  idea  that 
the  greatest  bird  species  diversity  and  population 
densities  are  in  the  climax  forest  (Johnston  and 
Odum  1956,  Karr  1968,  Kendeigh  1948,  Shugart 
and  James  1973).  Studies  have  shown  that  popula- 
tion densities  were  highest  in  intermediate  stands 
(Bond  1957,  Kendeigh  1946).  Karr  (1968)  noted  a 
decline  in  species  richness  and  density  in  the  last 
forest  stage  in  Illinois.  The  impact  of  fire  on  vege- 
tation and,  in  turn,  on  breeding  bird  populations 
was  studied  in  chaparral  (Lawrence  1966),  in 
pine-oak  woodland  (Marshall  1963),  and  in  pon- 
derosa  pine  (Lowe  et  al.  1978).  The  more  open 
habitat  produced  by  burning  in  both  vegetative 
types  led  to  an  increase  in  numbers  of  species  and 
density  with  some  changes  in  species  composition. 
A  significant  increase  in  bird  species  richness 
and  abundance  followed  logging  in  the  Douglas-fir 
region  of  northwestern  California  (Hagar  1960) 
and  in  a  giant  sequoia  forest  of  northern  Califor- 
nia (Kilgore  1971).  Similarly,  the  cutting  and/or 
logging  of  the  habitat  in  the  ponderosa  pine  forest 
increased  bird  population  densities  and  altered 
species  composition. 

Bird  pairs  on  all  the  treated  plots  except  the 
clearcut  site  were  more  highly  packed  (the  aver- 
age amount  of  foliage  volume  per  average  pair  of 
birds  was  smaller)  than  bird  pairs  on  the  control 
plot.  Pair  packing  on  the  severely  thinned  plot 
was  67,800  cubic  feet  per  pair,  whereas  on  the 
irregular  strip  shelterwood  plot  there  was  64,200 
cubic  feet  of  foliage  per  pair.  In  contrast,  on  the 
control  site,  there  was  251,400  cubic  feet  of  foli- 
age per  pair,  and  on  the  silviculturally  cut  site, 
there  was  140,300  cubic  feet  of  foliage  per  pair. 
On  the  severely  thinned  plot,  bird  pair  packing 
was  higher  than  on  the  control  site  because  of  the 
great  reduction  in  foliage  which  was  not  accom- 
panied by  a  proportional  decrease  in  population 
density.  In  fact,  on  the  irregular  strip  shelterwood 
plot,  not  only  was  the  amount  of  available  foliage 
reduced  by  two-thirds,  but  the  densities  increased 
as  well,  resulting  in  much  higher  pair  packing. 
Birds  on  both  these  heavily  treated  watersheds 
might  come  into  greater  potential  competition 
with  each  other. 

Bird  pairs  on  both  the  severely  thinned  and  the 
irregular  strip  shelterwood  plots  were  equally 
packed,  suggesting  that  approximately  65,000 
cubic  feet  of  foliage  is  the  minimum  required  by  a 
given  pair.  If  the  bird  community  on  the  control 


4 


Table  3.— Breeding  bird  composition  of  the  study  areas  (2  year  average  pairs  per  40  ha) 


Species 

Control 

Silviculturally 
cut 

Irregular 
strip 

Severely 
thinned 

Clearcu 

IVlUUIIlCllil   L^IMOIaCIUCC  \\^LJ } 

^  3 

4.5 

0.8 

ryyiiiy  iiuuiciiom  \\ju j 

1fi 

6.0 

1.9 

nuuoc  wicii  ) 

3.0 

.^olitarv  virpo  ^FN^ 

3.0 

6.0 

9.0 

6.0 

YpI low-ri  1  m npd  warhlpr  ^FN^ 

1.5 

12.0 

3.0 

1.5 

Grace's  warbler  (FN) 

9.0 

19.1 

14.3 

6.8 

.... 

Red-faced  warbler  (GN) 

3.0 



\A/pc  +  prn  tananpr  /FM\ 
VVcoLcill  Ldilctyt;!  ^ilN^ 

1  S 

1  .o 

3  0 

Hpnatip  t^in^^npr  /PM\ 
nc|JaUL«  Ldiidyd  y n  1 N ^ 

1.5 

64  5 

44.3 

17.0 

Rnhin 

3  0 

5.3 

3.8 

0.5 

Ri  if ni  iq-qiHpH  tnvAvhpP  /PM^ 

6.9 

6  0 

9.0 

4.5 

Mourning  dove  (FN) 

3.0 

1.5 

5.3 

.... 

Rock  wren  (GN) 

7.2 

4.5 

5.0 

Wormit  thriiQh  ^f^M^ 

1  Q 

0  4 

r^ra\/  hpaHoH  iiinpo  /r^M\ 
va  1  dy "1  icdUfcJu  juiiuu  ^oin^ 

18  8 

1 1 .3 

6.4 

1.8 

1 1  i Iri  npn^ itv 

22.2 

29.7 

32.8 

24.5 

14.2 

W  ck m  mprorc  a  n H  t pa  rp TQ 

f^nmmnn  flipkpr  /f^D^ 

3.0 

3.0 

3.4 

3.0 

0.8 

ndiry  wutjupcurxci  \\^u} 

\Jm\J 

4.5 

2.3 

Acorn  woodpecker  (CD) 

3.0 

.... 

White-breasted  nuthatch  (CD) 

6.8 

11.3 

10.5 

7.5 

Qfpllpr'q  ipv  /FN\ 

oidici  o  Jdy  ^niN/ 

7.5 

4.5 

5.3 

5.3 

Rl^^pk-hpadpH  nrn<^hpflk  ^FN^ 

3.3 

3.0 

1.5 

Cni\\\r\  npn ^ i tv 

23.6 

24.8 

25.2 

21.1 

0.8 

\A/pctprn  flvp^itrhpr  ^C^D^ 

4.9 

4.2 



\A/ potprn  \A/ond  np\iUPP  /FM^ 

2.3 

9.0 

3.0 

— . 

Ody  o  pilUtJUt;  ^1  IN^ 

1.5 

V  lUf t; l-y  1       1 1  bWdllUW  \\^U) 

8  3 

3.0 

Western  bluebird  (CD) 

4.5 

7.9 

13.5 

5.8 

.... 

Mountain  bluebird  (FN) 

0.5 

Rroad-tailpH  hii m m innhirH  /FM^ 
DiUdU  idiiC'Vj  iiuiiiiniiiyuiiu  ^niN^ 

4.1 

12.0 

9.8 

OUI 1 1 1 1  lU  1 1  1 1 1  y  1 1  U  Id W r\ 

1  S 

3.0 

3.0 

(^iiilH  r^pnQitx/ 

ouiiu  ut^iioiiy 

?fi  7 

28.3 

42.0 

21.6 

0.5 

Nesting  guilds 

1.3 

L/dvliy  dilU  UcpicbolUn  vV-'l-'/ 

i^^n  1 

OU.  1 

48  4 

24.3 

roiiage  nesiers  ^riN) 

O  (  .o 

fi7  1 

74  4 

46.0 

12.4 

orouna  nesiers  ^oiNj 

90  7 

91  5 

13.9 

1.8 

Total  density 

110.1 

147.3 

144.3 

84.2 

15.5 

Diversity  (H') 

2.83 

2.80 

2.94 

2.81 

1.35 

Evenness 

0.93 

0.91 

0.94 

0.96 

0.69 

Richness 

21 

22 

23 

19 

7 

plot  was  as  highly  packed  as  that  on  the  severely 
thinned  plot,  it  should  support  approximately  425 
pairs  per  100  acres.  Since  foliage  volume  does  not 
appear  to  be  the  limiting  factor,  then  factors  such 
as  territoriality,  food  supply,  and  lack  of  openings 
or  other  habitat  configurations  may  limit  bird 
populations. 

The  foraging  and  nesting  guilds  v^ere  variously 
affected  by  forest  cutting  and  logging  (table  3]. 
The  pickers  and  gleaners  and  the  ground  feeders 
increased  in  population  density  on  the  silvicul- 


turally cut  and  irregular  strip  shelterwood  plots. 
The  aerial  feeders  increased  by  57%  on  the  ir- 
regular strip  shelterv^ood  plot  in  response  to  the 
open  strip  areas,  whereas  the  hammerers  and 
tearers  remained  relatively  stable  on  all  the 
forested  watersheds.  All  the  foraging  guilds, 
except  for  the  ground  feeders,  were  virtually 
eliminated  from  the  clearcut  plot.  The  cavity  and 
depression  nesters  and  ground  nesters  greatly 
decreased  in  population  density  on  the  severely 
thinned  plot,  whereas  the  foliage  nesters  greatly 


5 


increased  in  population  density  on  the  silvicul- 
turally  cut  and  irregular  strip  shelterwood  plots. 
Interestingly,  the  majority  of  the  ground  feeders 
on  the  clearcut  plot  were  also  foliage  nesters. 
These  birds  used  the  Gambel  oak  saplings  that 
were  growing  throughout  the  area  for  nesting 
substrate.  The  cavity  nesters  which  also  use 
cavities  for  roosting  comprised  between  60%  and 
94%  of  the  wintering  bird  community  (Szaro 
1976). 

Eleven  species  (solitary  vireo,  pygmy  nuthatch, 
Grace's  warbler,  white-breasted  nuthatch,  com- 
mon flicker,  hairy  woodpecker,  steller's  jay,  gray- 
headed  junco,  chipping  sparrow,  broad-tailed 
hummingbird,  and  western  bluebird]  were  pres- 
ent on  all  the  forested  plots  during  the  study.  The 
common  nighthawk  was  observed  on  all  areas  in 
1975  but  was  not  found  on  the  silviculturally  cut 
plot  in  1974.  Several  species  increased  their  den- 
sity in  their  typical  habitat  (the  foliage)  on  the 
treated  plots.  Of  the  species  found  on  all  the  for- 
ested plots,  eight  (all  but  the  common  flicker, 
hairy  woodpecker,  and  pygmy  nuthatch)  had  their 
highest  population  densities  on  treated  plots  indi- 
cating density  increases  in  response  to  openness. 
In  contrast,  population  densities  of  five  species 
(red-faced  warbler,  pygmy  nuthatch,  western  fly- 
catcher, violet-green  swallow,  and  black-headed 
grosbeak)  were  significantly  reduced  with  heavy 
alteration  of  the  habitat.  The  rock  wren,  robin, 
and  western  wood  pewee  bred  only  on  treated 
plots,  whereas  the  acorn  woodpecker  was  found 
exclusively  on  the  severely  thinned  plot.  These 
species  probably  required  the  increased  openness 
of  the  habitat.  The  common  flicker,  hairy  wood- 
pecker, and  steller's  jay  maintained  relatively 
stable  densities  on  both  the  control  and  treated 
plots. 

Bird  species  diversity  and  species  richness  in 
the  ponderosa  pine  forest  were  not  significantly 
affected  (based  on  an  importance  criterion  of  at 
least  a  15%  difference)  by  treatment  except  on 
the  clearcut  plot.  Bird  population  densities  were 
significantly  increased  by  forest  cutting  on  the 
silviculturally  cut  and  irregular  strip  shelterwood 
plots  and  were  significantly  decreased  on  the 
severely  thinned  and  clearcut  plots.  For  a  more 
detailed  description  of  treatment  and  climatic  ef- 
fects see  Szaro  (1976). 

Literature  Cited 

Balda,  Russell  P.  1969.  Foliage  use  by  birds  of  the 
oak-juniper  woodland  and  ponderosa  pine 
forest  in  southeastern  Arizona.  Condor 
71:399-412. 


Balda,  Russell  P.  1975.  The  relationship  of  second- 
ary cavity  nesters  snag  densities  in  western 
coniferous  forests.  USDA  For.  Serv.  Wild.  Habi- 
tat Tech.  Bull.  No.  1,  37  p. 

Beecher,  William  J.  1942.  Nesting  birds  and  the 
vegetative  substrate.  Chicago  Ornithol.  Soc, 
Chicago. 

Blackford,  John  L.  1955.  Woodpecker  concentra- 
tion in  burned  forest.  Condor  57:28-30. 

Bock,  Carl  E.,  and  James  F.  Lynch.  1970.  Breeding 
bird  populations  of  burned  and  unburned  coni- 
fer forest  in  the  Sierra  Nevada.  Condor 
72:182-189. 

Bond,  Richard  R.  1957.  Ecological  distribution  of 
breeding  birds  in  the  upland  forests  of  southern 
Wisconsin.  Ecol.  Mongr.  27:351-382. 

Brown,  Harry  E.,  Malchus  B.  Baker,  Jr.,  James  J. 
Rogers,  Warren  P.  Clary,  J.  L.  Kovner,  Frederic 
R.  Larson,  Charles  C.  Avery,  and  Ralph  E. 
Campbell.  1974.  Opportunities  for  increasing 
water  yields  and  other  multiple  use  values  on 
ponderosa  pine  forest  lands.  USDA  For.  Serv. 
Res.  Pap.  RM-129,  36  p.  Rocky  Mt.  For.  and 
Range  Exp.  Stn.,  Fort  Collins,  Colo. 

Cottam,  Grant,  and  J.  T.  Curtis.  1956.  The  use  of 
distance  measures  in  phytosociological  sam- 
pling. Ecology  37:451-469. 

Hagar,  Donald  C.  1960.  The  interrelation  of  log- 
ging, birds,  and  timber  regeneration  in  the 
Douglas-fir  region  of  northwestern  California. 
Ecology  41:116-125. 

Johnston,  David  W.,  and  Eugene  P.  Odum.  1956. 
Breeding  bird  populations  in  relation  to  plant 
succession  in  the  Piedmont  of  Georgia.  Ecology 
37:50-62. 

Karr,  James  R.  1968.  Habitat  and  avian  diversity 
on  strip-mined  land  in  east-central  Illinois.  Con- 
dor 70:348-357. 

Kendeigh,  S.  Charles  1944.  Measurement  of  bird 
populations.  Ecol.  Monogr.  14:67-106. 

Kendeigh,  S.  Charles  1946.  Breeding  birds  of  the 
beech-maple-hemlock  community.  Ecology 
27:226-244. 

Kendeigh,  S.  C.  1948.  Bird  populations  and  biotic 
communities  in  northern  lower  Michigan.  Ecol- 
ogy 29:101-114. 

Kilgore,  Bruce  M.  1971.  Response  of  breeding  bird 
populations  to  habitat  changes  in  a  giant  se- 
quoia forest.  Am.  Midi  Nat  85:135-152. 

Lack,  David.  1933.  Habitat  selection  in  bords  with 
special  reference  to  the  effects  of  afforestation 
on  the  Breckland  avifauna.  J.  Animal  Ecol. 
2:239-262. 

Lack,  David.  1939.  Further  changes  in  the  Breck- 
land avifauna  caused  by  afforestation.  J.  Ani- 
mal EcoL  8:277-285. 


6 


Lack,  David,  and  Elizabeth  Lack.  1951.  Further 
changes  in  bird  life  caused  by  afforestation.  J. 
Animal  EcoL  20:173-179. 

Lawrence,  George  E.  1966.  Ecology  of  vertebrate 
animals  in  relation  to  chaparral  fire  in  the 
Sierra  Nevada  foothills.  Ecology  47:278-291. 

Lov^e,  Philip  O.,  Peter  F.  FfolHott,  John  H.  Dieterich, 
and  David  R.  Patton.  1978.  Determining  poten- 
tial w^ildlife  benefits  from  wildfire  in  Arizona 
ponderosa  pine  forests.  USDA  For.  Serv.  Gen. 
Tech.  Rep.  RM-52,  12  p.  Rocky  Mt.  For.  and 
Range  Exp.  Stn.,  Fort  Collins,  Colo. 

MacArthur,  Robert  H.  1958.  Population  ecology 
of  some  warblers  of  northeastern  coniferous 
forests.  Ecology  39:599-619. 

MacArthur,  Robert  H.,  and  James  W.  MacArthur. 
1961.  On  bird  species  diversity.  Ecology 
42:594-598. 

MacArthur,  Robert  H.,  James  W.  MacArthur,  and 
J.  Freer.  1962.  On  bird  species  diversity.  II. 
Prediction  of  bird  census  from  habitat 
measurements.  Am.  Nat.  96:167-174. 

MacArthur,  R.  H.,  H.  Recher,  and  M.  Cody.  1966. 
On  the  relation  between  habitat  selection  and 
species  diversity.  Am.  Nat.  100:319-332. 

Marshall,  Joe  T.,  Jr.  1957.  Birds  of  pine-oak  wood- 
land in  southern  Arizona  and  adjacent  Mexico. 
Pac.  Coast  Avifauna  32:1-125. 


Marshall,  Joe  T.,  Jr.  1963.  Rainy  season  nesting  in 
Arizona.  Proc.  Int.  Ornithol.  Congr.  13:620-622. 

Morse,  Douglas  H.  1967.  Competitive  relation- 
ships between  parula  warblers  and  other 
species  during  the  breeding  season.  Auk 
84:490-502. 

Recher,  Harry  1969.  Bird  species  diversity  and 
habitat  diversity  in  Australia  and  North 
America.  Am.  Nat.  103:75-80. 

Shannon,  C.  E.,  and  W.  Weaver.  1948.  The  math- 
ematical theory  of  communication.  Univ.  111. 
Press,  Urbana.  117  p. 

Shugart,  Herman  Henry,  and  Douglas  James. 
1973.  Ecological  succession  of  breeding  bird 
populations  in  northwestern  Arkansas.  Auk 
90:62-77. 

Szaro,  Robert  C.  1976.  Population  densities,  habi- 
tat selection,  and  foliage  use  by  the  birds  of 
selected  ponderosa  pine  forest  areas  in  the 
Beaver  Creek  Watershed,  Arizona.  Ph.D.  diss. 
Northern  Ariz.  Univ.,  Flagstaff.  264  p. 

Williams,  John  A.,  and  Truman  C.  Anderson,  Jr. 
1967.  Soil  survey  of  Beaver  Creek  Area,  Ari- 
zona. USDA  For.  Serv.  and  Soil  Conserv.  Serv. 
Washington,  D.C.  [Available  from  Sup.  Doc, 
U.S.  Gov.  Print  Off.,  Washington,  D.C.  20402]. 
75  p. 

Yeager,  Lee  E.  1955.  Two  woodpecker  populations 
in  relation  to  environmental  change.  Condor 
57:148-153. 


7 


Appendix 


Bird  Common  and  Scientific  Names 


Common  name  Scientific  name 


Mourning  dove 

Zenaida  macroura 

Cnmrnnn  niffhthawk 

ChordeiJes  minor 

Rrnpid-triilprl  hiimminffhird 

f  1  t  wClLX    LCIXXO^^    XX  LXXXXXXXXXXg  LJXX 

.SpJnsnhnrns  nlnfvrprcAifi 

C^X  Cl  O  IL/X  L  C/X  LX  O     XyXdl'V  OwX  V^LXlJ 

r^nrnmnn  fliplcpr 

vjL/lllilil-'ll    XXX^-jJN.C'  A 

CnlriTitp'^  nurnfiis  rnfpr 

\_/X  cx  !_/  L  O         CI.  t-i.  X  CX  1/  CX  L7     Cv  CX  1  Cr  X 

Acorn  woodDGckGr 

MeJanerpes  /ormicivorous 

Hairy  woodpecker 

Picoides  villosus 

Say's  phoebe 

Sayornis  saya 

Western  flycatcher 

Empidonax  difficilis 

Western  wood  newee 

Contopus  sordiduJus 

Vi  nl  pt-prppn  ^wpi  11  nw 

VXVJXOl   gXOO  XX    iD  V  V  cx  Lxyj  V  V 

Tnrhvrinptn  thnlri'i'iinn 

X  Ci.  C/X  LV  C/XXIO  L  Ci     I/XL  CXX  CI  O  OXX  L  \JL 

Steller's  jay 

Cyanocitta  stelleri 

VToiintain  chickadee 

X  V X  \j  Lxxx  I. n  XXX       XXX  wxv cx  d. O  W 

Pnrijs  pombeii 

X     cx  X    CA.            £^  CXX  X  L  l_/ C/ XX 

Whitp-hreasted  nuthatch 

VV  XXXLO     UX  O  cx  LJ  L  O  VJ.    XX  U.  LXXU  I.  VjXX 

fiittn  rnrnlinpnsis 

l./Xl'LCX     C/CXX  C/XXX  L  Cf  X  L  L7X 

Pvffmv  nnthatch 

X     y  QXXXy     XX  LX  LXXf_X  LwXX 

.Siftn  nvQwciPci 

C^X  L  L  CX     Xy  y  £1  XXL  CX  Cy  CX 

FTousp  wren 

X  X\-f  XJLiJKj      VV  X  C/XX 

Trnp/ndvtp*?  npdnn 

X  X  C/ £1 X  C  CX  y  L  Cr  O     CX  Cf  CX  C/X  L 

Rock  wren 

SaJpinctes  obsoietus 

American  robin 

Turdus  migratorius 

Hermit  thrush 

Catharus  guttatus 

Western  bluebird 

Sialia  mexicana 

Mountain  bluebird 

Sialia  currucoides 

Solitary  vireo 

Vireo  soJitarius 

Yellow-rumped  warbler 

Dendroica  coronata  auduboni 

Grace's  warbler 

DendroicQ  graciae 

Red-faced  warbler 

Cardellina  rubri/rons 

Western  tanager 

Piranga  ludoviciana 

Hepatic  tanager 

Piranga  flava 

Black-headed  grosbeak 

Pheucticus  melanocephalus 

Gray-headed  junco 

Junco  caniceps 

Rufous-sided  towhee 

Pipilo  erythrophthalmus 

Chipping  sparrow 

Spizella  passerina 

Tree  Common  and  Scientific  Names 

Common  name 

Scientific  name 

Ponderosa  pine 

Pinus  ponderosa 

Alligator  juniper 

Juniperus  deppeana 

Gambel  oak 

Quercus  gambellii 

8 


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