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RESPONSE OF SONGBIRDS AND SMALL MAMMALS
T0 WHOLE TREE HARVESTING OF ASPEN

presented by

Laura Elizabeth Eaton

has been accepted towards fulfillment
of the requirements for

Master of Science degnwin Fisheries and Wildlife

2 Major professor é

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Dam August 8, 1986

 

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a.

RESPONSE OF SONGBIRDS
AND SMALL MAMMALS TO WHOLE

TREE HARVESTING OF ASPEN

BY

Laura Elizabeth Eaton

A THESIS

Submitted to
Michigan State University
in partial fulfillment for the requirements
for the degree of

MASTER OF SCIENCE

Department of Fisheries and Wildlife

1986

ABSTRACT

RESPONSE OF SONGBIRDS AND SMALL MAMMALS
TO WHOLE TREE HARVESTING OF ASPEN

by
Laura Elizabeth Eaton

Whole tree harvesting has become increasingly popular
as a harvesting technique in recent years. Questions,
therefore, have been raised as to the environmental impacts
of this technique. In this study, songbird and small mammal
populations were compared between 8 whole tree harvested
aspen clearcuts and 8 aspen clearcuts harvested by conven-
tional pulpwood techniques.

Songbirds were censused during the breeding season
using the variable-circular'plot method. Small mammals were
trapped in August using Sherman live-traps. Percent cover
of slash, percent cover of vegetation, and density of woody
stems were also measured on the sites.

Slash was found to provide an average of 380% more
cover on the conventional sites than on the whole tree
sites. INo differences were found in vegetation cover or
stem densities between the harvest types.

Sixteen species of birds and 7 species of small mammals
exhibited a preference for l of the harvest types.
Approximately half the species preferred the conventional
sites and half the whole tree sites. These results suggest
that in large forest tracts both methods of harvesting can

be used, with a mix of the 2 methods recommended.

ACKNOWLEDGMENTS

I would like to extend special appreciation to my major
professor, Dr..Jonathan Haufler, for his help and guidance
through my graduate studies. I would also like to thank my
committee members, Dr. Herald Prince and Dr. Carl Ramm, for
their input and guidance.

Acknowledgment must be extended to the staff of the
Pigeon River Country State Forest for their help and
interest throughout this project.

I would like to thank my interns, Jim Brown, Kevin
Burt, Scott DeFrain, and Ann Resta, for their hard work and
input during field collection.

I would also like to thank.my fellow graduate students
for the camaraderie and support that we shared for 2 1/2
years.

Special thanks must go to Thomas Kulowiec and Kevin
Burt for the friendship and support; to my parents for
always supporting what I wanted to do; and to my 2 aunts
that provided financial and emotional support when I needed

it most.

iii

LIST OF TABLES . . .
LIST OF FIGURES . .
INTRODUCTION . . . .

OBJECTIVES . . . . .

TABLE

STUDY SITE DESCRIPTION . .

METHODS . . . . . .

Experimental Design .

Songbird Census

Small Mammal Census .

Vegetation Sampling .

Data Analysis .
RESULTS . . . . . .

Slash Sampling .

Vegetation Sampling .

Songbird Census

Small Mammal Trapping

DISCUSSION . . . . .

OF CONTENTS

SUMMARY AND RECOMMENDATIONS . . . . .

APPENDIX A . . . . .
APPENDIX B . . . . .
APPENDIX C . . . . .

LITERATURE CITED . .

iv

10

10

10

ll

12

13

l7

17

17

17

39

46

65

67

72

76

80

LIST OF TABLES

Table Page
1. List of parameter combinations for correl- 16

ation analyses.

2. Percent cover of vegetation in 5 height strata 18
-on 8 conventional and 8 whole tree harvested
aspen clearcuts which were harvested in 1982
and 1984. Sites were measured in 1984 and
1985 (mean : S.E.).

3. Foliage height diversity values, calculated with 19
the Shannon-Weiner equation (Hair 1980), for 8
conventional and.8 whole tree aspen clearcuts
which were harvested in 1982 and 1984. Sites
were measured in 1984 and 1985 (mean : S.E.).

4. Mean stem density per ha (: S.E.) for 2 strata 20
(<1m in height, and >1m but <10cm dbh), measured
on 8 conventional and 8 whole tree aspen
clearcuts which were harvested in 1982 and 1984.
Sites were measured in 1984 and 1985.

5. Density of birds per ha, number of bird species, 22
and bird species diversity, which was calculated
with the Shannon-Weiner equation (Hair 1980), on
8 conventional and 8 whole tree aspen clearcuts.
Sites were harvested in 1982 and 1984. Birds
were censused in 1984 and 1985 (mean i S.E.).

6. Mean densities (: S.E.) of birds in 1984 which 23
differed significantly between 8 conventional and
8 whole tree aspen clearcuts which were in their
first and third growing seasons.

7. Mean densities (i S.E.) of birds in 1985 which 25
differed significantly between 8 conventional and
8 whole tree aspen clearcuts which were in their
second and fourth growing seasons.

8. Correlations (rs) between bird densities and % 29
cover of slash. Birds listed were found to have
significantly different densities between conven-
tional and whole tree aspen clearcuts. Sites were
harvested in 1982 and 1984. Birds were censused
in 1984 and 1985.

Table

9.

10.

11.

12.

13.

14.

15.

Correlations (rs) of bird densities with 5

strata of vegetative cover and 2 strata of stem
density, (<1m in height, and >1m in height and <
10 cm dbh), which were measured on conventional
whole tree aspen clearcuts. Sites were harvested
in 1982 and 1984, and measured in 1984 and 1985.
Birds listed had significantly different
densities on the 2 harvest types.

Correlations (rs) of bird densities with 4
strata of vegetative cover and 2 strata of stem
density, (<1m in height, and >1m and <10cm dbh),
which were measured on conventional and whole
tree harvested aspen clearcuts. Sites were
harvested in 1982 and 1984, and measured in 1984
and 1985.Bird species listed exhibited no
differences in numbers between the harvest types.

Results of partial correlations (r3) between
clearcut area and number of bird species, bird
species diversity (BSD), and density of bird
species. Percent cover of slash was the factor
held constant. Birds were censused in 1984 and
1985.

Total number of animals, mean number of species,
and mean diversity indices, which were calculated
with the Shannon-Weiner equation (Hair 1980), for
small mammals live—trapped on conventional and
whole tree harvested clearcuts. Sites were harv-
ested in 1982 and 1984. Mammals were trapped in
1984 and 1985.

Mean number of small mammals (i S.E.) live-
trapped on conventional and whole tree harvested
aspen clearcuts. Sites were harvested in 1982
and 1984. Mammals were trapped in 1984 and 1985.

Correlations (r ) of small mammal numbers with:
% cover of slas , 3 strata of vegetative cover,
and density of stems <1m in height, and density
of stems >1m in height and <10cm dbh. Sites
were conventional and whole tree harvested aspen
clearcuts which were harvested in 1982 and 1984.
Mammals were trapped and vegetation was measured
in 1984 and 1985.

Bird species of upland openings which appear
to be sensitive to opening size. Small
openings were <5 ha. Large openings were not
defined (Taylor and Taylor 1980).

vi

37

38

4O

41

44

57

LIST OF FIGURES

Figure

l. The location of the Pigeon River Country
State Forest in Cheboygan and 0tsego Counties,
Michigan.

vii

INTRODUCTION

Whole tree harvesting became a production reality in
the early 1970's as a means of attaining greater efficiency
in the use of forest resources. Its initiation related
directly to an increasing demand for fiber for pulp and chip
products, and for wood fuel for industry as a supplement to
conventional fossil fuels (Arola and Miyata 1981, Keays
1974).

Whole tree harvesting is a technique whereby all above-
ground portions of the trees are utilized. This is usually
accomplished by chipping the wood on site, immediately upon
harvest. This technique both enhances time efficiency and
decreases transportation costs. Herrick (1982) estimated
that substituting chipping technology for conventional
hardwood pulpwood harvesting increased annual production by
2 1/2 times and reduced cost by $6.45 per cord-equivalent
delivered as chips. Whole tree harvesting also has the
added benefit of reducing the site preparation necessary for
regeneration by considerably reducing the amount of slash
left on the site (Blyth and Wilhelm 1980). For these
reasons, whole tree harvesting has become increasingly
popular as a harvesting method in recent years.

However, since little residual material is left on the

site, some questions have been raised as to the potential

environmental impacts of this technique. INumerous studies
have been conducted concerning nutrient loss, increased
exposure of soil and litter, increased erosion, and in-
creased leaching. Kimmins (1977) and Van Hook et al. (1982)
provided literature reviews on these topics.

Only 2 studies, hence far, have dealt with the effect
of whole tree harvesting on wildlife habitat. A study by
Hahn and Michael (1980) investigated the response of small
mammals to whole tree harvested hardwood clearcuts. They
found a significant decrease in.small mammal abundance in
the clearcuts which lasted for 6 years after harvest. They
mentioned that these results differed considerably from the
results of previous studies which found that small mammal
populations typically declined initially, but then quickly
recovered, and even increased in abundance on clearcut
sites. Hahn and Michael (1980) were unable to demonstrate
in their study that the lack of logging residue on the
clearcuts was responsible for the decline of the small
mammal population.

A study by Beyer (1983) looked at the response of song-
birds and small mammals to whole tree harvesting of aspen.
He found a reduction in the diversity, numbers, and species
of songbirds in the clearcuts as compared to uncut plots.
The small mammal population exhibited fluctuations which
could not be attributed to the harvesting treatment. Again,
this study did not specifically investigate the importance

of logging residue to the wildlife populations.

Slash has been documented as being utilized as a source
of food or cover for a number of songbird and small mammal
species on recent conventional clearcuts. Hagar (1960)
discussed the importance of slash to the ground foraging
dark-eyed junco (Junco hyemalis) as resting and escape cover
on clearcuts, especially before the invasion of brush.
Franzreb (1978) attributed the existence of a high density
of gray-headed juncos (Junco phaeonatus) in cut sites to the
presence of slash. Franzreb and Ohmart (1978) described the
importance of slash to gray-headed juncos as foraging sur-
faces, observation posts, and protection for nest sites.

Hagar (1960) found that winter wrens (Troglodytes
troglodytes) had a preference for cut sites and were always
found in close association with logs. Similar results have
been found for the house wren (Troglodytes aedon) (Franzreb
and Ohmart 1978, Scott and Gottfried 1983), and for the
Carolina wren (Thryothorus hidovicianus) (Connor and
Adkisson 1975). Connor and Crawford (1974) found that
recently clearcut areas attracted downy woodpeckers

(Dendrocopus pubescens) and hairy woodpeckers (Dendrocopus

 

villosus) because the slash and debris from logging provided
an abundant source of insect prey.

Many studies have indicated a relationship between
protective cover and small mammal populations (Eadie 1953,
Morris 1955, Lovejoy 1971, MHCloskey and Lajoie 1975). Red-
backed voles (Clethrionomys gapperi) have specifically'been
correlated with the presence of slash (Tevis 1956, Gunderson

1959, Ahlgren 1966, Martell and Radvanyi 1977). Dimock

(1974) mentioned that residue may serve as a potential
source of food for insectivorous small mammals.

Removal of slash by burning on clearcuts has also been
documented to have dramatic effects on small mammal popula-
tions. Small mammal populations.are'typically'reduced.ini-
tially; However, Peromyscus populations have been found to
recover within months after a fire (Ahlgren 1966, Sims and
Buckner 1972, Krefling and Ahlgren 1974, Fala 1975). Species
requiring more cover, such as the red-backed vole or various
species of shrews, require extensive periods of time to
regain their numbers after burns (Ahlgren 1966, Dimock
1974).

Songbirds and small mammals were chosen for this study
because of their usefulness as indicators of habitat quality
(Graber and Graber 1976, Plunkett 1979, West et a1. 1981).
These populations are also integral elements in ecosystems.
Birds are important to the balance of the food chain, and
aid in the perpetuation of communities through seed disper-
sal (Marks 1974). Birds have also been documented as being
important in controlling pest insect plagues (Bruns 1959,
Plunkett 1979, Crawford et a1. 1983). Small mammals serve
as a prey base for larger predators (Hamilton and Cook 1940,
Chew 1978, Potter 1978). They have also been credited with
increasing soil aeration and fertility, and thus plant
productivity (Hamilton and Cook 1940, Chew 1978). Small
mammals may also aid in seed dispersal and in the dispersal

of mycorrhizal fungi (West 1968, Chew 1978, Maser et a1.

1978, Ream and Gruell 1980). In addition to their ecol-
ogical importance, nongame wildlife populations have been
receiving increasing attention and concern from the general
public in recent years (Zagata 1978). Aspen (Populus sppJ
sites were chosen for this study because aspen is typically’
harvested by clearcutting, and because aspen is important to
the pulpwood industry in Michigan (Blyth and Smith 1980).
It is important for wildlife populations to be given
proper consideration in forestry management plans. This is
not possible, however, without sufficient information con-
cerning wildlife habitat requirements. The purpose of this
study was to determine if there are significant differences
in small mammal and songbird populations between whole tree
and conventionally'harvested.sites. ‘Ultimately, the study
should indicate the importance of slash to these populations

on recently clearcut sites.

OBJECTIVES

The objectives of this research project were to:
Compare the relative population densities, species
composition and species diversity of songbird and small
mammal communities between whole tree harvested and

conventionally harvested clearcut sites.

Determine the importance of cover provided by slash to

these populations.

Determine the importance of other site characteristics,
such as site size, and vegetative structure and compo-

sition, to the species observed, in order to explain the

occurrence of the populations found on the sites.

STUDY SITE DESCRIPTION

Sixteen aspen clearcuts, situated in the Pigeon River
Country State Forest, Michigan were selected for the study.
Half of the sites were harvested by whole tree harvesting
and 1/2 by conventional pulpwood techniques. Half of the
sites within each harvest type were cut in 1982 and half
were cut in 1984.

The Pigeon River Country State Forest consists of
33,603 ha. It is located throughout the northeast corner of
0tsego County and the southeast corner of Cheboygan County,
which are located in the central northern Lower Peninsula
(Fig. l). The topography of the Pigeon River area consists
of morainic uplands, steep morainic slopes, sandy outwash
plains, and river bottoms. Three rivers, the Black, the
Sturgeon, and the Pigeon, originate from coniferous swamps
along the southern edge of the area and flow'north. The
soils range from dry sand of low fertility on outwash plains
to sandy loams of medium high fertility on till plains and
moraines (Moran 1973).

The typical climate of this area is characterized by
long cold winters, short cool summers, mild autumns, late
cold springs, prevailing'westerly'winds, fairly high.humid-

ity and low evaporation. The average rainfall for the area

Cheboygan County

 

0tsego County

 

i

* Pigeon River Country State Forest

Fig. l. The location of the Pigeon River Country State
Forest in Cheboygan and 0tsego Counties, Michigan.

is 10.24 cm. The average frost-free season is 116 days in
the south-central part of Cheboygan County (Foster et a1.
1939).

The clearcuts investigated ranged in size from 5 to 16
ha. The total area of the conventional sites consisted of
77 ha. The total area of the whole tree sites consisted of
93 ha. The soils of the sites ranged from well drained to
poorly drained, and the extreme moisture conditions were
evenly divided between the 2 harvest types.

.All of the sites had both bigtooth aspen.(Popu1us
grandidentata) and trembling aspen (Populus tremuloides)
present, with 1 species usually dominant over the other. It
variety of hardwood species were mixed with the aspen on the
sites. These included oaks (guercus spp.), maples (Age;
spp.), birch (Betula spp.), cherries (Prunus spp.), and
beech (Fagus grandifolia). Conifers, such as balsam fir

(Populus balsamifera) and eastern white pine (Pinus strobus)

 

were occassionally scattered over the plots as well. Common

shrubs in the clearcuts included hawthorn (Crataegus sppJ,

 

beaked hazelnut (Corylus cornuta), serviceberry (Amelanchier

 

spp.), maple-leaf viburnum (Viburnum acerifolium), witch-
hazel (Hamamelis virginiana), and brambles (gubgg sppJ.
Herbaceous vegetation was typically dominated by bracken
fern (Pteridium aquilinum), grasses, and a variety of field
flowers. A number of the sites still had large trees stand-
ing which were residual from harvest. Snags, when present,

were also left standing by harvest.

METHODS

Experimental Design

This study was arranged as a completely randomized
design with 4 replications. Sixteen clearcuts served as
study sites. These were divided into 2 treatments: har-
vesting by whole tree techniques and harvesting by con-
ventional techniques. Each treatment was divided into 2 age
classes: sites out in 1982 and sites out in 1984. The data
for the study were col lected in the spring and summer of
1984 and 1985. The sites were, therefore, in their first
and third growing seasons during the first field season, and
in their second and fourth growing seasons during the second
field season. This arrangement allowed.for the examination
of treatment effects over 4 age classes, and thus over a

range of vegetative development.

Songbird Census

Songbirds were censused from mid-May to mid-June of
each year by the variable circular-plot method, as described
by Reynolds et a1. (1980). As many censusing stations as
possible. were established on each site. Stations were
established at least 150m apart and 75m from an edge. The
number of stations established on the sites ranged from 2 to

7.

10

11

The period of time spent censusing at each station was
6 minutes. This period was determined from several 30
minute pre-counts. In the pre-counts, the number of species
observed was plotted against time. The period at which the
addition of new species began to level off was used as the
counting time. .All "singing males" were counted as 2 birds.

Sites were censused 10 times in 1984, and 9 times in
1985. Censuses for each site were distributed throughout
the breeding season. Three observers were used for the
censuses. Observers were rotated on the sites to limit
observer bias. Censuses began at sunrise each morning.
However, censuses were not carried out on exceptionally
windy, rainy, foggy, or cold mornings. Approximately 2
sites were censused per observer per morning. Therefore,
the time of the morning a site was censused was switched for
each observation period. That is, if a site was censused
first 1 period, it was censused second the next. Censusing
was also initiated at a different station for each obser-

vation period.

Small Mammal Census

Small mammals were censused.by trapping with Sherman
live-traps during August of each field season. A trapping
grid consisting of 6 rows and 6 columns was established on
each site. Stations were established 15m apart with 2 traps
per station.

Grids were established in locations which appeared as

similar as possible in terms of topography, drainage, and

12

vegetation. Grids were placed as far from the site edges as
possible to limit the chances of catching edge inhabitants.

However, some of the sites were small or narrow. Therefore,
for consistency, all grids were established with 1 side 60m

from an edge, which was the largest distance possible on the
smallest site.

Sites were trapped for 5 consecutive nights. .A whole
tree and a conventional site of each age class (4 sites)
were trapped concurrently to control variation due to
weather. Traps were baited with a mixture of oats, lard,
and anise extract. .All animals captured were marked with

ear tags or by toe clipping.

Vegetation Sampling

Vertical cover of vegetation was measured in each
clearcut by the line intercept method (Gysel and Lyon 1980).
Five height strata were measured: 0-10cm, 10-30cm, 30cm-1m,
l-4m, and >4m. Percent cover of slash was measured in a
0-30cm stratum. Vegetation and slash cover <lcm were
ignored. Gaps in cover <5cm were also ignored. Line inter-
cepts were run from randomly located points. .All lines were
run north, unless the line ran out of the clearcut, then it
was run south, east, or west, respectively. Measurements
were made at least 20m from the edge of the site. The
length of the lines ranged between 20 and 100m, depending on
the variability of the cover on the sites.

Density of woody stems was measured in rectangular plots

which were 2m wide and ran the length of the line intercepts.

13

Stems were counted in 2 strata: <1m in height, and >1m in
height and <10cm dbh. Stems were recorded by species.
Frequency of herbaceous vegetation was recorded from 2m x
5m plots. The line intercept was used as 1 edge of the
plot. .All measurements were carried out until a required
sample size, calculated at the 95% level of confidence,
was met.

Trees >4m in height were counted and recorded by
diameter classes. Snags 32m in height and 15cm dbh were

also counted and were recorded by diameter and height.

Data Analysis

Statistically'adequate sample sizes formall vegetative
sampling and slash sampling were determined.with.the follow-
ing formula from Snedecor (1956):

tzs2

 

n-
32

t - tabulated at the 90%
confidence level

s - sample variance
E - allowable error (mean
multiplied by a maximum of 20%)
Foliage height diversity, bird species diversity, and
small mammal diversity were determined by the Shannon -
Weiner diversity index (Hair 1980):
.H' a 'ZPi 109 Pi
Pi - decimal fraction of the total

individuals or total cover of the
i category

14

Analysis of the variable circular-plot method was
carried out as described by Szaro and Balda (1983). Density
of each species per band around census points on 40.5 ha
scale was calculated by:

4.365(105)(N)
BD-

 

“0122-1122) (p) (r)
BD - band density

N -number of observations in the
band

OR - outer band radius
IR - inner band radius
p - number of sampling stations

f - frequency each station was
sampled

The number of birds per 40.5 ha (by species) for each
point for each count period was calculated by:

4.355(105)(N)

 

birds/40.5 ha -
77 (192)

Overall density of each species was then calculated
by determining the mean and the standard error of the
individual estimates. In order to account for the correl-
ation between sampling periods, standard errors were cal-

culated by: , I

X
V(Y) - (E S: + 2§<jrij Si Si)/(pf2):
SE " V67)

S = among-station variance on the i
sampling period

r a correlation between sampling periods
i and j

15

Differences between the clearcut types, within age
classes, were analyzed with the nonparametric randomization
test. Differences were tested for: % cover of slash, %
cover of vegetation, foliage height diversity, stem
densities, bird species' densities, bird species diversity,
songbird abundance, number of species of songbirds, small
mammal abundance, number of species of small mammals, and
small mammal diversity. These tests were carried out at the
95% confidence level.

Songbird and small mammal populations were tested for
association with habitat parameters using Spearman-rank
correlation analysis. Parameter combinations are listed in
table 1. iPartial correlations, holding % cover of slash
constant, was further used to test for songbird associations

with clearcut size.

16

Table 1. List of parameter combinations for correlation

analyses.

 

x-variables

Y-variables

 

Foliage height diversity

Clearcut size

Cover of slash

Cover of vegetation
(height strata: 0-10cm,
lO-30cm, 30cn-ln, 1-4m)

Cover of vegetation
(height strata: >4m)

Density of woody stems
(strata: <1m in height,
>1m in height and <10cm
dbh)

Number of woody species

Bird species

diversity

Bird abundance
Number bird species

Bird species

densities

Cover of slash
Bird abundance

Bird species
Bird species

densities
diversity

Bird species densities

Small mammal
Bird species
Small mammal

Bird species

Bird species
Small mammal

Small mammal

species numbers
densities
species numbers

densities

densities
species numbers

species numbers

 

RESULTS

Slash Sampling

The mean % cover of slash was found to be significantly
greater (p<0.05) on the conventional sites. The cover of
slash on the conventional sites ranged from 17-56% with a
mean of 37.39 i_4.51%. The cover of slash on the whole tree
sites ranged from 6-16% with a mean of 9.75 i 1.26%. Slash,
therefore, provided an average of 380% more cover on the

conventional sites than on the whole tree sites.

Vegetation Sampling

No significant differences were found for vegetative
cover (Table 2), foliage height diversity (Table 3), or stem
density (Table 4) between the harvest types (p>0.05).
However, some trends are evident. Consistently more cover
was sampled on the conventional sites in the 30cm-1m and
1-4m strata in all 4 year classes. Stem density in the >1m
stratum was also found to be consistently greater on the
conventional sites.

Absolute and relative frequencies of herbaceous plants

are provided in appendix A. No differences were noted.

Songbird Census
The overall abundance of birds was found to range from

2-12 birds/ha in 1984 and from 5-12 birds/ha in 1985. The

17

18

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moawm 05mm on» one: mvsouseao cHqudezI: use In use .mvsouweao cHqumozIN use I« m

 

 

 

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I. I. .I I. I. :
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I. I. I. I. I N
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ame>um= use»

«pauam vendor

 

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e A emem .fl CUOSV

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use on Hmcouaco>coo m so museum pgwqe: m cm souamvewe> no ue>oo aceouom

.~ canes

19

Table 3. Foliage height diversity values, calculated with
the Shannon-Weiner equation (Hair 1980), for 8 conventional
and 8 whole tree aspen clearcuts which were harvested in
1982 and 1984. Sites were measured in 1984 and 1985 (mean
i S.E.).

 

 

Year Classa Conventional Whole tree
1 1.28 i 0.05 1.23 i 0.03
2 1.21 i 0.02 1.19 i 0.03
3 1.31 i 0.03 1.28 i 0.03
4 1.22 i 0.06 1.29 i 0.02

 

a1- and 2-year-old clearcuts, and 3- and 4-year-old
clearcuts are the same sites measured in different years.

'2'-

20

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memo one one: menohmoao oaouemmhI: use In use .mpsosmoao oHoIsmoaIm use Ia

 

 

 

 

 

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owe H man: 83 ... mat: mom... .... an? $3 H ~83 m
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.33

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EH A one .pzwwms cw EHV.V museum N pom A.m.mHg m: use AHHmCmo Seam cow: .3 manna

21

species which were consistently high in abundance on the
sites regardless of harvest type or age, were the chestnut-
sided warbler (Dendroica pensylvanica), indigo

bunting (Passerina cyanea), and song sparrow'(Melospiza
melodia). Other species were also abundant, but were
primarily so on 1 harvest type or the other, or only within
certain year classes.

‘Values for bird density, number of species, and bird
species diversity (BSD) are listed in table 5. Bird density
was found to be significantly greater (p<0.05)lon the
conventional sites in the first year class. Bird density
continued to be slightly higher on the conventional sites
through the fourth year class, but the differences were not
significant. .Also in the first year class, BSD was found to
be significantly greater (p<0.05) on the whole tree sites.
In the second year class, the number of species was found to
be significantly greater (p<0.05) on the whole tree sites.
In the third year class, the number of species was found to
be significantly greater (p<0.10) on the conventional sites.

A complete list of bird species occurring on the sites
with their mean densities on each clearcut type are provided
in appendices B and C. Bird species which were determined
to be present on the harvest types in significantly different
numbers are listed in tables 6 and 7.

A maximum of 27 species were observed in clearcuts in
their first growing season. Thirteen of these species were
found to have significantly different numbers between the

harvest types. .Alder flycatcher (Empidonax traillii),

.wsMoz sass“; moose pmo>sms soosvos “no.0uvav moososoeewo psmowmflsmfim 4*
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.msao> asosoewfln cw oomsmsoo

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zpwmso>wv mofioosm when use .wowoosm oswn ho soDEss .ms son woman ho zuwmsoo .m oHnme

 

23

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souummm uaofim

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.mmsao woo» s sasuaa Amo.ovmv moosonouufiu Osmowuasmam 44
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13a magmas

 

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I AwmwsHEoum mmwommmmv

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“unsouoofi musuwosmwv

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% Ammmmmmxm mmmmMMNHv
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I I AuHHmwm eaamwmv
a.n+m.oa v.a+m.n «4H.HH~.e o.oHo.o uuwnosan suoumom
Amadeosuau mannaouowv

«ec.oHo.o v.nHm.m v.oHv.o o.HHm.H pneumuou sooauoss
ooua oHosz HusoHuso>soo ooua oaosz Hmsoauso>soo mofioomm uuflm

 

V WMMHU “MON

 

m mmoao Moo»

 

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m cmmsumn saucaosuecosm emumuusu noses mam. as mouse no x.m.msc mmeusmcme cum:

.5 means

26

n.w+n.m N.@HO.NH

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.mmoao moo» o sasuwz Aoa.ovmv moosonomuwu assuamwsmflm e

h.vHN.NH o.NNHn.om

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someone uouoousquuwsz

 

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0098 QHOHE HflfiOHUCO>GOU

m mmoao Moo»

mowoomm uufim

16.»:00. .e manna

27

American redstart (Setophaga ruticil la), common yellowthroat
(Geothlypis trichas), mourning warbler (Oporornis
philadelphia), and rufous-sided towhee (Pipilo
erythrophthalmus) were observed almost exclusively on the
conventional sites. White-throated sparrow (Zonotrichia
albicollis) was observed primarily on the conventional
sites, but was also observed in relatively low numbers on
the whole tree sites. Eastern bluebird (Sialia sialia),
killdeer (Charadrius vociferus), northern oriole (Icterus
galbula), scarlet tanager (Piranga M), tree swallow
(Iridoprone bicolor), and vesper sparrow (Pooecetes
ggmineus) were observed almost exclusively on the whole
tree sites. However, scarlet tanager, tree swallow, and
vesper sparrow were observed in relatively low numbers.
Eastern kingbird (Tyrannus tyrannus) was observed in signif-
icantly greater numbers on the whole tree sites, but was
also observed on 2 of the conventional sites.

A maximum of 36 bird species were observed in clearcuts
in their second growing season. Nine of these species were
found to be present in significantly different numbers
between the 2 harvest types. The bird species found in
greater numbers on the conventional sites were mourning dove
(Zenaidura macroura), Nashville warbler (Vermivora
ruficapil la), rufous-sided towhee, and white-throated
sparrow. Species found in greater numbers on the whole tree
sites were eastern bluebird, eastern kingbird, field sparrow

(Spizella pusilla), northern oriole, and tree swallow.

 

28

However, field sparrow, mourning dove, and northern oriole
were all observed in relatively'low'numbers.

In the third growing season, a maximum of 28 species
were observed on the sites. Of these, 9 species were found
to have significantly different numbers between the harvest
types. In this year class, alder flycatcher, American red-
start, common yellowthroat, golden-winged warbler (Vermivora
chrysoptera), great crested flycatcher (Myiarchus crinitus),
mourning dove, and tree swallow were found to prefer conven-
tional sites, while field sparrow and northern oriole were
found to prefer whole tree sites. IHowever, American red-
start, great crested flycatcher, mourning dove, and field
sparrow were all present in low numbers in this year class.

In the fourth growing season, a maximum of 34 species
were observed on the sites. In this year class, American
redstart was found to prefer conventional sites and field
sparrow was found to prefer whole tree sites. Again the
field sparrow was observed in relatively low numbers.

The general pattern of abundances remained the same in this
year class in that the species which showed a preference for
1 harvest type in previous year classes still had a higher
mean density on the preferred type in this year class.

Of the 16 species which were found to have significantly
different densities between the harvest types, 10 were found
to be significantly correlated with % cover of slash for at
least 1 of the censusing years. These 10 species are listed
in table 8 with the correlation coefficients for each year.

The species which exhibited a preference for conventional

29

Table 8. Correlations (rs) between bird densities and %
cover of slash. Birds listed were found to have signifi-
cantly different numbers between conventional and whole
tree aspen clearcuts. Sites were harvested in 1982 and
1984. Birds were censused in 1984 and 1985.

 

Correlation Coefficient (rs)

 

 

Bird Species 1984 Census 1985 Census
American redstart .783*** .411
(Setsnbssa rutisilla)

Common yellowthroat .674*** .535**
(Gestblxnie trishas)

Eastern bluebird -.490* -.612***
(Sialia sialia)

Eastern kingbird -.536** -.557**
(Tyrannus txrannus)

Field sparrow -.491* -.509**
(Spizella pusilla)

Mourning dove .455* .270
(Zenaidura masreura)

Mourning warbler .516** .505**
(geezernis eniladelnbia)

Nashville warbler -.043 .476*
(Kermixera rufisaeilla)

Northern oriole -.383 -.518**
(IQEQIEE 3212913)

White-throated sparrow .499** .433*

(Zenetrishia albisellis)

* Significant at p<0.10
** Significant at p<0.05
*** significant at p<0.01

 

30

sites were found to be positively correlated with slash,
while species exhibiting a preference for whole tree sites
were found to be negatively correlated with slash.

Six of the 10 species which were found to be signif-
icantly correlated with slash were consistently so for both
of the censusing years. Northern oriole and Nashville
warbler were found to be correlated with slash only in 1985,
while American redstart and mourning dove were found to be
correlated with slash only in 1984. The greatest density
of redstarts in 1984 was observed on the 1-year-old conven-
tional sites. ‘These birds were most often observed near the
ground, moving in and around the slash, and were therefore
assumed to be nesting in the slash. In 1985, the greatest
density of redstarts was observed on the 4-year-old
conventional sites. On these sites the birds appeared to be
using the upper strata of vegetation. If slash was impor-
tant to this species in 1985 it was not detected by observa-
tions or by correlation analysis.

Nashville warbler was not found to be correlated with
slash in 1984, but was found to be correlated with slash
(p<0.10) in 1985. The observations of this species tend to
support these findings. This species was present on the
sites in much lower numbers in 1984 than in 1985. Also, in
1984 this species was primarily detected near the edges of
the clearcuts, while in 1985 it was detected in substantial
numbers throughout the area of the clearcuts. iNo explan-

ation is evident for the differing use of slash between the

31

2 years by the northern oriole and mourning dove, suggesting
that the significant correlations with slash for the 1 year
were due to chance.

Four species which were not found to exhibit a pref-
erence for l of the harvest types were found to be signif-
icantly correlated'with slash. However, these species were
only found to be correlated with slash for 1 of the
censusing years. Common flicker (Colaptes auratus)
(p<0.10), song sparrow (p<0.10), and house wren (p<0.01)
were positively correlated with slash in 1984, and brown
thrasher (Toxostoma rufum) (p<0.10) was negatively
correlated with slash in 1985. The house wren was not
observed on the sites in 1985, therefore the validity of its
correlation with slash is difficult to evaluate. Since the
other species were abundant on the sites in both years,
their correlations with slash for the 1 year are assumed to
be due to chance.

The species which exhibited differences in numbers
between the harvest types, but which correlation analysis
did not indicate to be associated with slash, were alder
flycatcher, golden-winged warbler, great crested flycatcher,
killdeer, rufous-sided towhee, scarlet tanager, tree
swallow, and vesper sparrow. All of these species, except
the golden-winged warbler and the great crested flycatcher,
exhibited differences in the first growing season. Of these
species, only the rufous-sided towhee and alder flycatcher
exhibited differences beyond the first growing season. The
rufous-sided towhee had significantly different numbers in

32

both the first and second growing seasons, which suggests a
real preference for conventional sites. IMale towhees were
observed singing on slash piles on some conventional sites.
However, it appears that this relationship was not strong
enough to be demonstrated by the correlation analysis.
Alder flycatcher had significantly different numbers in the
first and third growing seasons. This inconsistency, and
the lack of association with slash, suggests that this
species was responding to variables other than slash. .A
difference for golden-winged warbler and great crested fly-
catcher were only found during the third growing season.
This may have been due either to chance occurrence or to a
response by these species to variables other than slash.

Some trends evident in the data suggest a relationship
between vegetative development on the sites and.bird
utilization. Table 5 illustrates an increase in bird
abundance and number of species over the 4 year classes.
Although BSD did not increase over the 4 year classes, it
was greater on all the sites the second year, In examining
the density of individual bird species over the 4 year
classes (appendices B and C), a number of other trends are
also evident. White-throated sparrow exhibited a dramatic
decrease in numbers on the conventional sites and a slight
increase on the whole tree sites. Indigo bunting, golden-
winged warbler, and rufous-sided towhee increased substan-
tial 1y on both site types. Nashville warbler showed a

slight increase in numbers on the whole tree sites.

33

Table 9 provides the results.of correlation analysis
between vegetative parameters and the bird species which
exhibited differences in numbers between the harvest types.
Some trends are evident in these results. Common yellow-
throat, golden-winged warbler, mourning warbler, and white-
throated sparrow were found to be positively correlated with
slash, and.were also positively'correlated.with vegetative
cover and stem densities. Eastern bluebird, eastern king-
bird, and northern oriole were found to be negatively cor-
related with slash, and were also either uncorrelated or
were significantly negatively'correlatedwwith‘vegetative
cover and stem densities.

A few other relationships between vegetation and bird
species are also worthy of mention. American redstart was
found to be strongly correlated with vegetation in the l-4m
stratum in 1985, which corroborates the observations of this
species for this year. .Alder flycatcher was positively'
correlated with these parameters for both years. In 1984,
rufous-sided towhee was positively'correlated with all
vegetative parameters, except vegetative coverI>4m.and stem
density <1m. In 1985,rufous-sided towhee was positively’
correlated with vegetative cover in the 0-10cm and lO-30cm
strata. The conflicting results between the 2 years makes
it difficult to draw conclusions for this species. Vesper
sparrow was negatively'correlated.with vegetative parameters
in 1985, but uncorrelated with vegetation in 1984. No

important trends or associations with vegetation were

34

 

eeo.u ¢~4.I «as. How. mow. ...eoe.u ..omm.a no .mmsammmmmoumwamo mdsmwm.

 

 

 

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omn.n can.I «on. Hma.a soc. onn.I ev~.I no .msmmdom mmmommm.
ecvv.I can. nae. eemom.I eevmm.I evev.I eeccm.I vm oHoHuo suosuuoz
own. ...Hoo. eon.u odd. ea~.I ...ooe. ...eoo. no .aumommos mmmuwoqow.
awn. hmc.I mmo.l oav. Hoe. emvv. mnv. cm sedans: msasuso:
...o~e. can. oHo.I ..oom. nan. .onv. enn. mo .mmommomsumm mmomwsmow.
can. awe. ea~.I can. nov. ..ecm. .eoe. so possum: uomcsaIcouHoo
.~63.I men.I nee. Hm~.I sec. «no. sue. me Ammmmomsm mmoqmusac
HeH.I HnH.I emu. e-.I onn.I mun.I nem.I so oneness: :uoumam
H-.I ..e~m.u can. eno.I «ms. -n.I can.I no .qumflo mflwmsw.
mmo.I coo. cv~.I «oH.I HAH.I mno. mco.I «a uuenoasn :soumam
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..vem. mm~.I om~.I ..m~o. .mov. ...~oo. ...mmo. co uaousuaosaoa :oaaoo
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vac. anH.I Hoe.u «no. nos. one. who. «a uuaumuou caosuosa
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.noe. apo.n «we. .«mom. nan. nae. .«ov. «a uosouuoasu moods
EMA luv EVA 3?..." 3H Ifloon floonIOH EOOHIO H50» mafiuoam Guam
>uwosoo soum soaueuomo> no uo>oo a

 

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use Assauuso>soo so uousueofl ouoa sodss .Assu sooav use usuwos su BAA use .usowos sa savv .xuaosou
Ioum no eueuum w use uo>oo o>aueuomo> uo eueuum v sud: mouuamsou uuwn no .muv mso«ueaouuoo .m oases

35

.«Ho.ovm. mueoa susuda mama» umo>ues soosuos oosououuau useoquusmum see
.Amc.ovm. muse» s«su«3 momxu umo>uss sooauon oosououuwu assoauusmum ea
.Aoa.ovn. mueoa s«su«3 mom>u umo>ues sooauon oosououuau useoaufismum c

 

 

 

..omm. ...Heo. ma~.I «on. mac. .cemm. sue. me Amqsammamdu munmwuwmmom.
omo. moo. ono.I man. «as. mam. ”an. «o saunas» uouoousquuess

...mmo.u eo~.I omo. .eoe.n em~.I ..H¢A.I eon.u no .msooqsomm monommmmc
mvo. mm~.I aeo.I nou.u moo. was. «as. «a sausage nodmo>
suA lav qu chH saIsoon soonIoH scoAIo use» mowoomm uuam
auamson aoum sowueaouo> no uo>ou »

 

.u.u:ou. .m manna

36

found for field sparrow, great crested flycatcher, mourning
dove, Nashville warbler, or tree swallow.

Correlation analysis between vegetative parameters and
the bird species which did not exhibit differences in num-
bers between the harvest types may be useful in explaining
the presence of these species on the sites as well. The
results of the correlations which showed a consistent pat-
tern for both censusing years are given in table 10.
Chestnut-sided warbler was strongly influenced.by all the
vegetative parameters except cover >4m. Song sparrow was
most strongly associated with vegetative cover'in.the
lowest 2 strata. Indigo bunting and rose-breasted grosbeak
were strongly negatively'correlated with stem density <1m
for both censusing years. In 1984, indigo bunting was also
positively correlated with all strata of vegetative cover
except cover in the >4m stratum. No important associations
were detected between FHD and bird species' densities.

Correlations between clearcut size and % cover of slash
did not indicate an association between these parameters.
Partial correlation analysis, with.% cover of slash held
constant, indicated that number of species of birds and BSD
were positively correlated.with clearcut size for both
censusing years. Partial correlations with individual bird
species indicated that northern oriole and eastern kingbird
were positively correlated with clearcut area, while common
yellowthroat and mourning warbler were negatively correlated

with clearcut area (table 11).

37

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souo no avenue m use uo>oo o>«ueuooo> no eueuum m sud: moauamsou uses no .muv oso«ueuouuou

.OH wanna

38

Table 11. Results of partial correlations (r,) between clearcut
area and number of bird species, bird species diversity, and
bird species' densities. Percent cover of slash was the factor
held constant. Birds were censused in 1984 and 1985.

 

Correlation Coefficient (r3)

 

 

Bird 1984 Census 1985 Census
No. Species .579*** .519***
Species Diversity .415* .398*
Common yellowthroat -.378* -.394*
We misses)

Eastern kingbird .704**** .559***
(IXIIDHHI EXIEBBHI)

Mourning warbler -.635**‘* -.503***
(murals animus)

Northern oriole .726**** .400*

(M 931931.”

 

* Significant at p<0.20
** Significant at p<0.10
*** Significant at p<0.05
**** Significant at p<0.10

39

Small Mammal Trapping

A total of 186 small mammals were captured in 1984 and
a total of 705 were captured in 1985. Nine different
species were represented on the sites. The disparity in
numbers of animals captured between the 2 years was thought
to be due to natural fluctuations of the small mammal pop-
ulations.

Table 12 outlines the total number of animals caught,
the mean number of species caught, and.the mean small mammal
species diversity on each harvest type and year class. No
differences were found in small mammal abundance or diver-
sity between the harvest types for either 1984 or 1985.
However, a pattern is evident in that in both years the
greatest number of animals were caught on the youngest whole
tree sites and the fewest were caught on the oldest
conventional sites. Although there were no significant
differences in the number of species caught on the sites
within trapping years, 2 species, eastern chipmunk (Tamias
striatus) and short-tailed shrew (Blarina brevicauda), were
not found on the sites in 1984, but were captured in sub-
stantial numbers in 1985.

Table 13 provides the mean number of animals caught of
each species on each of the harvest types for each of the
trapping years. In 1984, only 2 species were caught in
significantly different numbers between the harvest types.
Meadow vole (Microtus pennsylvanicus) was found to prefer
whole tree harvested sites during the first growing season,

and masked shrew (Sorex cinereus) was found to prefer

 

40

Table 12. Total number of animals, mean number of species, and
mean diversity indices, which were calculated with the Shannon-
Weiner equation (Hair 1980), for small mammals live-trapped on
conventional (C) and whole tree (W) harvested clearcuts. Sites
were harvested in 1982 and 1984. Mammals were trapped in 1984
and 1985.

 

 

Yeara Harvest Total No. Species Diversity
Class type animals caught 2 1 S.E. 2 i S.E.
59 3.8 :I_- o.3 1.11 r 0.13
1
W 90 3.8 i 0.5 0.83 i 0.11
c 174 7.0 3; 0.6 1.35 3; 0.13
2
W 220 5.5 :4; 0.3 0.99 i 0.11
13 1.5 1: 1.0 0.48 i 0.30
3
W 24 3.0 i 0.4 0.92 3; 0.11
121 7.0 1', 0.6 1.35 3; 0.13
4
W 220 6.8 i 0.3 1.39 '_I-_ 0.09

 

aSites in year classes 1 and 2, and 3 and 4 were the same sites
trapped in different years.

41

 

 

 

 

 

 

 

 

 

 

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.nmms cam ammo cw aweampp

.mvcosmoao cocoa covmo>smz meow

.mH manma

42

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3mmfi vmvmm>hmz mmvwm Nmma Umvmm>hmr mmVMm

 

Ac.#:oov .mH manna

43

conventional sites during both the first and third growing
seasons. In 1985, a number of differences were found.
During the second growing season eastern chipmunk and masked
shrew were found to prefer conventional sites, and meadow
jumping mouse (Zapus hudsonius) was found to prefer whole
tree sites. Peromyscus, meadow vole, and woodland jumping
mouse (Napeozapus insignis) were found to prefer whole tree
sites during the fourth growing season. Red-backed vole was
found to prefer conventional sites in both the second and
fourth growing seasons.

A few trends are also evident. .Although no other
significant differences in numbers were found for
Peromyscus, this species was consistently trapped in greater
numbers on the whole tree sites. Similarly; no significant
differences were found for short-tailed shrewu IHowever, this
species was caught in greater numbers on the conventional
sites in both year classes in 1985. Meadow jumping mouse
also exhibited a fairly consistent preference for whole tree
sites.

Table 14 provides the results of correlations for’small
mammal numbers with % cover of slash and vegetative
parameters. Only masked shrew was found to be correlated
with slash in 1984, and this was a strong positive
association. In 1985, Peromyscus, meadow jumping mouse,
and woodland jumping mouse were found to be negatively
correlated with slash, while red-backed vole was found to be

positively correlated with slash.

44

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.Aoa.ovmv munch canvas mama» um0>ucz soosuon oocmuouuao accoauwcmam «

 

 

 

 

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o>aueuowo> no vacuum n .nuoau no uo>oo a «sud: mucous: Hasle- Annsm no Anny mcoauoaouuoo .vH manna

45

Eastern chipmunk was not found to be correlated with
slash. However, on one whole tree site chipmunks were only
caught at the base of large oak trees which were residual on
the site. If the animals caught on this site are removed
from the analysis, the correlation indicates that eastern
chipmunk was positively associated with cover of slash at
the p<0.10 level of significance.

Correlations with vegetation also indicated some assoc-
iations. Peromyscus was strongly negatively correlated with
all strata of vegetative cover in 1984, and with.cover in
the 30cm-lm and l-4m strata in 1985. This species was also
negatively correlated with stem density >1m for both years,
and positively correlated with stem density <lm.in 1984.
Red-backed vole was also consistently negatively correlated
with vegetative cover in 1984, but no associations were
evident for this species in 1985. Short-tailed shrew
exhibited strong positive correlations with vegetative cover
in the 0-lOcm, 30cm-lm, and l-4m strata in 1985, as well as
with the number of woody species on the sites (p<0.05).
Woodland jumping mouse was found to be negatively associated
with woody stems >1m, and vegetative cover in the O-lOcm and
l-4m strata in 1984. No associations with vegetation were

evident for the other species captured.

DISCUSSION

Significantly more slash was found on the conventional
sites than on the whole tree sites. No significant
differences in stem density or vegetative cover were
found between the harvest types. This suggests that the
observed differences in songbird and small mammal species
compositions between the harvest types were due primarily to
differences in the cover of slash.

Habitat characteristics which influence songbird
numbers and diversity have been investigated by numerous
researchers. MacArthur and MacArthur (1961) described the
importance of foliage height diversity to bird species
diversity. MacArthur et a1. (1962) further concluded that
horizontal variability, or patchiness, was a principal
factor. Karr and Roth (1971) added the importance of total
vegetation volume. Roth (1976) concluded that overall
habitat complexity was the key. Slash, therefore, was
expected to contribute to overall songbird diversity by
adding another component to the habitat. It was also
hypothesized that slash would be most important on recently
harvested sites before extensive revegetation had occurred.

The results of this study indicated that 8 species of
birds had a preference for conventional sites and 8 species

had a preference for whole tree sites. Based on the

46

47

correlations between bird densities and cover of slash, the
differences found for 10 species could be attributed to the
presence or absence of slash. The greatest differences in
species composition were found in the first growing season.
Differences were minimal by the fourth growing season.
Differences in bird abundance and BSD were also found, but
only during the first growing season. This suggests that
the influence of slash on the bird populations declined as
expected as the sites revegetated.

A number of the species found to prefer conventional
clearcuts in this study have been documented to use slash by
other sources. Titterington et a1. (1979) felt that bird
species that nest and forage on the ground were associated
with the open ground-slash seral stage of clearcuts.
Titterington et al. used the white-throated sparrow as an
example, however, mourning warbler, Nashville warbler,
common yellowthroat, and rufous-sided towhee also nest and
forage on the ground. Bent (1963b) and Bent (1968a) also
reported accounts of the mourning warbler and rufous-sided
towhee using cut-over areas or "slashings". Rufous-sided
towhee was not found to be correlated with slash in this
study, however, it was found to prefer conventional sites
during the first and second growing seasons. A possible
explanation for the lack of response by towhees to the slash
is that only the 0-30cm stratum of slash was measured.
Towhees were observed using slash on sites where the slash
had been placed in fairly large discreet piles. Perhaps if

the slash had been measured in strata, or if its

48

distribution had been analyzed, a relationship between
towhees and slash would have been determined.

Nashville warbler was only found to be correlated with
slash in 1985. Observations of this species indicated that
it utilized the clearcuts to a much lesser degree in 1984
than in 1985. It is not clear whether this was primarily
due to differences in population numbers between the 2
years, or to a general shift in habitat utilization.

The responses of some of the other species to the
harvest types are not as simple to explain. American
redstart, alder flycatcher, golden-winged warbler, great
crested flycatcher, and mourning dove were found to prefer
conventional sites for at least 1 growing season. American
redstart and mourning dove were also found to be correlated
with slash for l of the censusing years. The habitat
requirements of the American redstart have been described
differently by different sources. Crawford et a1. (1981)
described the American redstart as a closed-canopy-
obligatory species. Webb (1977), however, described this
species as l which is positively affected by habitat
disturbance such as heavy logging. Bent (1963a) reported
accounts of the redstart nesting in mature hardwood, mixed
deciduous, and coniferous woodlands, as well as open shrubby
vegetation, alder and willow thickets, and edges of wood-
lands. This species is obviously versatile in its nesting
requirements. However, there appear to be no previous

accounts of this species utilizing slash as nesting cover.

49

The mourning dove has been described as preferring open
lands with scattered tree, shrubs, and open fields (Terres
1980). It typically feeds on the ground and has been known
to nest on the ground where trees are scarce (Soutiere and
Bolen 1973). It is possible that slash provided cover for
mourning doves using the clearcuts. However, the small
number of mourning doves observed on the sites suggests that
the clearcuts were not highly preferred by this species.

Alder flycatcher, golden-winged warbler, and.great
crested flycatcher were not found to be correlated with
slash. They also did not exhibit a consistent preference
for the conventional sites over the year classes. .Alder
flycatcher was found to prefer conventional sites in the
first and third growing seasons, and golden-winged warbler
and great crested flycatcher were found to prefer conven-
tional sites only in the third growing season. Therefore,
the apparent preferences of these species are probably due
to responses to other habitat variables on these sites, or
to chance occurrences. The correlations with vegetative
parameters suggest that alder flycatcher and golden-winged
warbler were attracted to areas with dense vegetative cover
and stem densities in the l-4m stratum. Although no sig-
nificant differences were found for these variables between
the harvest types, it is possible that these species found
more patches of dense vegetation in this stratum on the

conventional sites.

50

Northern oriole, scarlet tanager, eastern kingbird,
eastern bluebird, field.sparrow, killdeer'and'vesper'sparrow
were found to prefer whole tree sites for at least 1 growing
season. Eastern kingbird and northern oriole both nest and
feed above the ground (Bent 1942, Bent 1958), which provides
little explanation for their aversion to conventional sites.

Connor and Adkisson (1974) documented the eastern blue
bird's use of young clearcuts when cavities were available
for nesting. Connor and Adkisson (1975) found that eastern
bluebirds preferred l-year-old clearcuts which suggested a
preference for open areas created by clearcutting. This
study found the greatest abundance of bluebirds on l-year-
old whole tree sites. On 4-year-old sites eastern bluebirds
were equally abundant on whole tree and conventional sites
where cavities were available. This suggests that sites
with less cover are preferred by eastern bluebirds, but that
availability of suitable nesting cavities is the most impor-
tant feature attracting this species.

Scarlet tanager is typically associated with dense
deciduous woods (Bent 1958). It is possible that this
species was actually associated with the surrounding woods,
rather than with the clearcuts. This species was observed
in very low numbers, and was typically’observed near the
edges of the clearcuts or in large residual trees. Its
association with whole tree clearcuts may have been a factor
of chance.

Bent (1968b) described the typical habitat of the field

sparrow as old fields and pastures, and mentioned that it

51

will use open "slashings" after logging operations. This
species was, therefore, expected to have a preference for
conventional sites, but was instead found in greater numbers
on the whole tree sites.

The killdeer is a bird of meadows and grazed pastures,
which nests in scrapes of bare ground (Terres 1980). Since
this species appears to prefer open sites, it might be
expected that it would prefer the whole tree sites. ‘This
species was indeed found almost exclusively on the l-year-
old whole tree sites. However, no association with slash
was found for this species.

Vesper sparrows typically nest on the ground where
vegetation is sparse and low (Bent 1968b). Since this
species appears to prefer to nest in areas with little
cover, it was expected that this species would prefer the
whole tree sites. ‘The vesper sparrow exhibited a preference
for whole tree sites in the first growing season, but no
correlation with slash was found. Several field observers
had difficulty distinguishing the song of the vesper sparrow
from that of the song sparrow. Therefore, it is possible
that this species exhibited preferences beyond the first
growing season, or had associations with habitat variables
which were not detected by this study due to confusion with
the song sparrow. '

The tree swallow exhibited preferences for both harvest
types. Tree swallows were found to prefer the whole tree

sites in the first and second growing seasons, and the

52

conventional sites in the third growing season. This may
suggest a preference for the whole tree sites during the
early stages of vegetative.development. However, this
species was not found to be correlated with slash. .Also,
since sites in the first and second growing seasons were the
same sites censused in 1984 and 1985, it appears more likely
that the tree swallows were attracted to the sites rather
than to the harvest types. Isince this species is a cavity
nester, it is possible that availability’of suitable snags
was the primary feature attracting this species. The
significant differences in numbers of this species between
the harvest types, may again, have been a factor of chance.
'A number of species that were expected to respond to
the different harvest types based on previous studies were
not found to respond in this study. Winter wrens have been
found to be associated with slash (Hagar 1960, Titterington
et a1. 1979). In this study, winter wrens were heard singing
along the edge of the clearcuts where sites abutted wooded
swamps, however, none were observed to use the clearcuts.
Franzreb and Ohmart (1978) felt that house wrens were
attracted to harvested plots because slash provided obser-
vation posts and foraging surfaces. The house wren was
found to be significantly correlated with slash in this
study. However, the low amounts of slash on the whole tree
sites did not reduce the use of these sites by the house
wren. It is possible that the availability of snags
with suitable cavities was the primary factor influencing

53

this species. Why the house wren did not return to nest on
the sites in 1985 can not be explained by this study.

Connor and Crawford (1974) recommended that slash and
logging debris not be treated or removed after harvest as
they felt it provided important foraging substrate for
hairy woodpeckers, downy woodpeckers, and common flickers.
In this study, none of these species were found to show a
preference for the conventional sites. Since snags were
present on most of the sites, it is possible that the snags
sufficiently attracted woodpeckers to compensate for the lack
of slash on the whole tree sites.

In general, the species observed on the sites in this
study were birds which typically nest or forage in brushy
openings or edge areas. 'The fact that certain birds are
associated with particular habitat types or successional
stages suggests the importance of vegetation to bird
distributions. The results of the correlation analyses
between bird numbers and vegetative parameters in this study
indicated the vegetative components important to each
species. The trends found in this study of bird abundance,
number of species, and BSD increasing over the 4 age
classes indicated an increase in utilization of the clear-
cuts with increasing vegetative development.

A number of studies have documented that bird abundance
and diversity increase with site ecological age (Johnston
and Odum 1956, Hagar 1960, Connor and Adkisson 1975,

Titterington et al. 1979). This has been attributed to the

54

increasing spatial complexity'and thus niche availability
with the increasing vegetative development.

Titterington et al. (1979) found that each seral stage
following clearcutting was dominated by a characteristic
group of bird species. They found that bird species that
nest and forage on the ground, such as the white-throated
sparrow, dark-eyed junco, and winter wren, were associated
with the earliest stage -- the open ground-slash stage. The
next stage -- the bramble-herbaceous stage -- was dom-
inated by species which typically nest and forage in vege-
tation close to the ground. Examples of these species
included.the white-throated sparrow, common yellowthroat,
chestnut-sided warbler, and mourning warbler. .All of these
species, with the exception of the dark-eyed junco and
winter wren, were observed on the sites in this study and
were also found to be associated with vegetative cover in
lower height strata. Titterington et al. (1979) used the
American redstart as an example of a species associated with
the third stage -- the shrub-sapling stage. In this study,
American redstart was found to be strongly associated with
cover in the l-4m stratum. The rose-breasted grosbeak was
used as an example of a species associated with the fourth
seral stage -- the immature second growth stage. In this
study, the rose-breasted grosbeak was found on sites of all
4 age classes. It was, however, found to slightly increase
in numbers over the 4 age classes, which may suggest a

preference for older sites.

55

In summary, it appears that slash had the greatest
influence on the bird populations on the l-year-old sites
because it provided structure on otherwise barren sites. As
vegetation developed on the sites, the cover provided by the
slash became obsolete. By the fourth growing season, the
bird populations were primarily influenced by the vege-
tation. As the vegetative structure was relatively’similar
on all the sites of the same age, the bird species compo-
sitions were also similar.

The effect of clearcut size on bird species was also
examined. Correlations with this variable suggested that
BSD and number of bird species increased with clearcut size.
Correlations with individual species indicated that 2
species were positively influenced by the area of the clear-
cuts and 2 negatively.

The issue of forest fragmentation, and thus the impor-
tance of stand size to bird species, has been of increasing
concern to researchers. According to Anderson and Robbins
(1981), there is a minimum size for each vegetation type at
which all the typical species of that type are likely to
be present. Most of the studies investigating the impor-
tance of stand size to bird populations have looked at
second growth and mature forest stands (Forman et al. 1976,
Whitcomb et al. 1977, Anderson and Robbins 1981, Ambuel and
Temple 1983). Only 1 paper appears to be available which
addresses the importance of forest openings to bird species
(Taylor and Taylor 1980). Taylor and Taylor felt that

breeding birds associated with large openings were different

56

from those associated with small openings. They, therefore,
attempted to list the bird species they felt were sensitive
to opening size, and the preferences of these species for
large or small openings. Species listed as sensitive to
opening size by Taylor and Taylor which were also observed
on the sites in this study are listed in table 15.

Only the results for the common yellowthroat.and
mourning warbler from this study corresponded to the
relationships suggested by Taylor and Taylor. .However, this
study was not specifically designed to address the
importance of site size to bird numbers. .Although an
attempt was made to compensate for the differences in
harvest method by using partial correlation analysis, other
factors, such as site configuration, were not held constant.
The range of site sizes examined was also limited. It
should also be noted that Taylor and Taylor derived.their
information from their own observations, and those of other
studies, rather than from a well designed study of the
subject. Therefore, their information should also be looked
at conservatively. Although more work is needed to specify
species relationships with opening size, it is apparent that
relationships do exist. This suggests that site size should
be taken into consideration in forest management planning.

A number of other factors which could have had a minor
influence on the bird populations were not considered in
this study. The importance of horizontal diversity has been

emphasized in a number of studies (MacArthur et al. 1962,

57

Table 15. Bird species of upland openings which appear to
be sensitive to opening size. Small openings were < 5 ha.
Large openings were not defined (Taylor and Taylor 1980).

 

Opening Size

 

Species Preferred
Black-and-white warbler Small
(Miotilta yarn)

Black-capped chickadee Small
(Rams W)

Brown thrasher Small
(IQXQESQEQ IEIBE)

Chestnut-sided warbler Small
(Dendroica W)

Chipping sparrow Small
(Spizella mine)

Common yellowthroat Small
(M menu)

Downy woodpecker Small
(W W)

House wren Small
(Treslsdxfse 55.4911)

Killdeer Large
(Charadrius mifems)

Morning dove Large
(Zenaidura W)

Mourning warbler Small
(92912131.: philadelmhia)

Nashville warbler Small
(Mixers 131112221111)

Vesper sparrow Large

(W W)

 

58

Roth 1976, Rotenberry and Wiens 1980). In this study, only
‘vertical cover was measured. However, horizontal cover'was
seldom uniform over the sites. There is little doubt that
the horizontal distribution of cover influenced the occur-
rence of some species and thus overall species diversity on
the sites. Swift et al. (1984) and Petit et al. (1985)
examined the influence of soil moisture on breeding birds.
Soil moisture is felt to have an effect on vegetative
structure, species composition, and invertebrate fauna.
No attempts were made to test for such associations in this
study. Other factors which may have been influential
include: the surrounding vegetation types: nearness to
special features such as open water, streams, or swamps: the
number and quality of snags: the number, size, and species
of residual trees: and availability of food such as fruits.

The factors influencing small mammal abundance and
diversity have not been as well defined as for birds. How-
ever, the need for cover by many small mammal species has
been well documented. Slash.was, therefore, expected to
enhance small mammal numbers and diversity by providing
cover on clearcuts, particularly'before.the sites revege-
tated. Also, by contributing another component to the site
slash was expected to provide additional niches on the
conventional sites.

The results of the second year of small mammal trapping
suggest a similar response to the harvest types as that
found for the songbirds in that approximately the same

number of species showed a preference for each of the

59

harvest types. Nine species of small mammals were caught on
the sites. Of these, 7 exhibited a significant difference
in numbers for at least 1 of the growing seasons. Three of
the species showed a preference for conventional sites and 4
showed a preference for whole tree sites.

An inadequate number of animals were caught the first
year to draw conclusions concerning the response of small
mammals to the harvest types during that year. iHowever,
trends found the first year are useful for corroborating
results found the second year.

Correlations with % cover of slash were useful for
explaining the differences found between the harvest types.
Peromyscus and meadow jumping mouse were found to prefer
whole tree sites and were also found to be negatively
correlated with % cover of slash. However, the preferences
of meadow voles and woodland jumping mice for whole tree
sites could not be attributed to the absence of slash on
these sites. In 1985, the red-backed vole was found to
prefer conventional sites, and.was also found to be posi-
tively'correlated with slash. In 1984, similar results
were found for masked shrew.

The red-backed vole is the only species which has
consistently been associated with logs and slash in the
literature. Gunderson (1959) felt that the distribution of
red-backed voles was most closely correlated with.the.pre-
sence of stumps, rotting logs, and root systems in forests.
Tevis (1956), and Martell and Radvanyi (1977) found slash to

be important to red-backed voles on recently cut sites.

60

Slash has also been suspected to provide a source of insects
for insectivorous small mammals such as shrews (Dimock
1974). Hooven and Black (1976) listed species of beetles
which are attracted to fresh slashings. Burt (1957) des-
cribed the habitat of the masked shrew as woodlands with
logs and litter on the forest floor. He also mentioned that
short-tailed shrews frequently nest under logs. In this
study, only the masked shrew was found to be correlated with
slash. Both species of shrews were also found to be
abundant on both harvest types. It is possible that the
shrews were positively influenced by slash. However, it is
apparent that slash was not the primary factor influencing
their distribution.

Both red-backed voles and short-tailed shrews have been
found to have high water requirements and are often assoc-
iated with moist sites (Gunderson 1959, Getz 1968, Miller
and Getz 1977). These species were caught on both moist and
upland sites in this study. However, there is some possi-
bility that slash could help maintain soil moisture, partic-
ularly before sites have revegetated. However, the poten-
tial importance of slash as a source of food and cover
suggests that its contribution to site quality extends
beyond this possible feature.

Peromyscus appear to have an affinity for barren sites.
This small mammal is typically the first to reinvade sites
after a severe disturbance such as clearcutting or burning

(Jameson 1955, Tevis 1956, Gashwiler 1970, Sims and Buckener

61

1972, Krefting and Ahlgren 1974, Fala 1975, Hooven and Black
1976). Therefore, the preference for whole tree sites was
not surprising for this species.

No previous information suggested explanations for
the preferences of the meadow vole, the meadow jumping
mouse, or the woodland jumping mouse for whole tree sites.
The correlations between these species and cover of slash
suggest that the preferences of the meadow and woodland
jumping mice for the whole tree sites can be attributed to
the lack of slash on these sites, but the response of the
meadow vole must be attributed to factors other than slash.

The correlations with the vegetation parameters
indicate that the small mammals were greatly influenced by
vegetation. Consistent with its response to cover of slash,
Peromyscus was found to respond negatively to vegetative
cover. This response perhaps explains this species' pref-
erence for the younger sites. Previous studies corroborate
these results. Hahn and Michael (1980) found Peromyscus to
be negatively'correlated with % blackberry (33235 sppJ
which grew in thick tangles on their sites. Krefting and
Ahlgren (1974) also felt that vegetative development made
sites less attractive to Peromyscus. Some other studies,
however. have found Peromyscus to be positively associated
with vegetation. M'Closkey and Lajoie (1975) found a posi-
tive correlation between Peromyscus abundance and ground
cover in the 0-736cm stratum. Hahn and Michael (1980) found
deer mice to be positively correlated with.% herbaceous

vegetation. Duesar and Shugart (1978) found Peromyscus to

62

be associated with a high density of shrub-understory vege-
tation in the forest. They felt that this was consistent
with the arboreal behavior of this mouse. This preference
by Peromyscus for shrubs may explain the positive correl-
ation found in this study with stem density <1m. However,
since this association was not repeated the second year it
is difficult to have confidence in this finding. Getz
(1961a) also found that Peromyscus only inhabited areas with
trees and shrubs. They found no indication that Peromyscus
avoided herbaceous vegetation. They suggested that habitat
preference of Peromyscus was primarily associated with food
supply.

No correlations with vegetation were found for meadow
jumping mouse. A study by Quimby (1951) found the greatest
number of meadow jumping mice in moist meadows and marshy
areas with woody and herbaceous vegetation. Nests were
found in grasses or sedges. It was, therefore, expected
that meadow jumping mice would be positively correlated with
the lower strata of vegetation.

Woodland jumping mouse was negatively'correlated.with
vegetative cover in the O-lOcm and l-4m strata, and.with
stem density >1m. These findings are opposite to results
found by Miller and Getz (1977). They found woodland
jumping mice to be less abundant in areas with lesser tree,
shrub, and herbaceous cover.

Meadow vole was not found to be associated with any

vegetative parameters. However, meadow vole has been

63

documented to inhabit dense grassy areas (Burt and
Grossenheider 1976). Getz (1961b) felt that the presence of
graminoids for food was the major factor responsible for the
occurrence of meadow voles in grassy areas. He also found
that meadow voles avoided areas with woody vegetation and
areas containing only forbs.

Red-backed vole was found to be negatively correlated
with all strata of vegetative cover in 1984, as well as
stem density >1m. However, since these results were not
replicated in 1985 their validity is questionable. Previous
studies have found red-backed voles to be associated with
availability of food, water, and slash (Gunderson 1959, Getz
1968). No previous studies have found red-backed voles to
be associated with vegetation.

Masked shrew was not found to be associated with any
vegetative parameters in either year. However, according to
Burt (1957), masked shrews are frequently found in brushy
or grassy areas where they build runways or use runways
built by other mice. Hooven and Black (1976) found 4
species of shrews to require a mat of ground vegetation for
cover. Therefore, a positive association with vegetative
cover was expected for this species.

Short-tailed Shrew was only caught on the sites in 1985,
therefore, no replication of the correlations was possible.
For the 1 year, this species was found to be positively
correlated with vegetative cover in the O-lOcm, 30cm-1m, and
1-4m strata, with stem density >1m, and with the total

number of woody species on the sites. A study by Miller and

64

Getz (1977) corroborated these results. They found short-
tailed shrews to be more abundant on sites with greater
vegetative cover, particularly'of the herbaceous layer.
They also suggested that the diversity of invertebrates was
positively'correlated.with the diversity of the leaf litter,
especially that from trees and shrubs.

Eastern chipmunk was not found to be correlated with
any vegetative parameters. :However. previous studies have
found associations. Duesar and Shugart (1978) found eastern
chipmunk to occur at sites with primarily a deciduous
canopy, high tree density, and low shrub density. They also
found them more commonly in forested than in shrub vege-
tation types. Krefting and Ahlgren (1974) found the eastern
chipmunk to be attracted to areas with a wide variety of
seeds.

In summary, it appears that the small mammal population
was influenced by the presence of slash. Correlations with
slash and differenCes in species numbers during the fourth
growing season indicate that slash continued to influence
the small mammals after the sites revegetated. Responses to
cover of slash and vegetative parameters were individual for
each species, and suggest the differing habitat require-

ments of each.

SUMMARY AND RECOMMENDATIONS

Cover of slash was found to influence both the songbird
and small mammal populations. This influence was equally'
negative and positive in that approximately 1/2 the species
that responded had a preference for 1 or the other of the
harvest types. Songbirds found to prefer conventional sites
were primarily species which typically'nest on or near the
ground. Bird species preferring whole tree sites were
species which nest and feed above the ground, or species
which are typically associated with open areas. Small
mammal species found to prefer conventional sites appeared
to be species requiring cover, species adapted to specific-
ally'woody cover, and insectivorous species. Small mammal
species found to prefer the whole tree sites appeared to be
species whose habitat preferences were for areas with little
cover.

Differences in songbird species composition moderated
as vegetation developed on the sites. By the fourth growing
season differences were minimal. Small mammals responded to
vegetation, but responses to slash were still evident in the
fourth growing season. Because small mammals are decidedly
more dependent on the ground layer than are birds, it is
perhaps logical that slash would.have a more lasting

influence on the mammals. How long slash continues to

65

66

influence small mammal populations will require further
study to determine.

The results of this study suggest that in large forest
tracts both methods of harvesting should be used. This may
be a minor consideration for the songbirds as differences
between the site types were minimal by the fourth growing
season. However, responses by the small mammals were
still evident through the fourth growing season. This indi-
cates that slash has a more lasting effect on small mammals,
and also leaves open to question whether other species are
affected by the presence of slash as well.

The results of the correlations with clearcut size
suggest that maintaining a variety of clearcut sizes is
desirable for songbird management. JHowever, further
studies of the importance of forest opening size to
songbirds is needed to corroborate the findings of this
study. Back (1980) felt that the minimum cut size of
4-8 ha needed to make pulpwood operations feasible is
probably compatible with songbird management. Back (1980)
also recommended that for cuts greater than 20 ha, efforts
should be made to maximize the ratio of edge to area. The
response of the bird species in this study further emphasize
the importance of maintaining snags in clearcuts that are

suitable for use by cavity nesting species.

APPENDICES

Appendix A. Frequency for herbaceous vegetation in conven-
tional and whole tree clearcuts that were harvested in 1982
and 1984, and were measured in 1984.

67

68

 

 

 

 

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mmue macs; HMCOMpcm>coo mmua macs; Hmcowpcm>coo mmdomam nuflm
m mmmflo Law» a wmmfio ham?

 

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73

 

 

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IV I I I Amzpmpzm mwpmmfioov
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amps macs; HmcoHpCm>cou wmue macs: Hmcowpcm>coo mmwommm cLHm

 

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74

 

 

 

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I. .I I I. Amwamwmvmawnm manpopomov
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u.o+m.o o.o+o.o m.m+o.m o.o+o.o ummcaafix
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75

 

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I Amzmcmsmpm mmpwomocmv
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I. I I I Apofioomn mzoopmocmpHv
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mmua macs; Hmcowpcm>:oo maps maonz Hmcowpcw>coo mmwommm npfim
m mmmau ham» H mmmao ham»

 

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76

 

 

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: mmmao pamw m mmmHo ummw

 

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m :0 mmofi c“ um>pmmno mmfiommm ouwn Ham mo A.m.m+v m: awn mmwpwmcmc cams .o xmccmma<

 

77

 

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I. I. I. .I Ammnowh» mmeHcpomuv
m.w+m.m N.NIo.m :.m+o.: :.m+:.om amounpzoaamh coesoo
I. I. I. .I Aaasomwzm m3HMomwzov
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I. I. .I I “mapmpsm mmpmmHoov
o.o+o.~ m.o+o.m ~.H+o.m m.o+m.a pmx0uam season
I I. I. I Answammmmm mafimuwmmv
m.w+:.m o.:+:.HH m.~+:.m w.m+~.m zoupmnm mewamwco
I I .I I. Amoacm>aamcmm.mowouc:mav
5.:H+~.Hm :.:~+m.mm ~.m+o.mm :.-+~.mm “manpmz cmuHmIpscpmmsu
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: nmdao paw» m mmeo ham»

 

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78

 

 

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.I I. I. I AmszMCmomm.m:memm<v
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I I I I Amzmpzmpmm msomcompmov
o.o+o.o o.o+o.o m.o+~.~ o.o+o.o nonwm maggzm
I I. I. I Amscfim m32wmmv
o.o+o.o m.m+m.m 0.:«+©.ma m.o+m.o :wxmww mcmm
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I I. .I I Amflzmamnmawzm mwcpopommv
5.9.0.S m.n+m.om :.N+m.ma :.m+m.:m pwanhmz WCwCLsoz
I I _I I. Ampsoucme muzvfimCva
N.o+:.o :.~+:.H o.o+o.o m.o+m.~ m>oc w2wcuzoz
I. I. I. I. hamcmxo acmummmmmv
m.~+m.~: m.:H+H.om H.m+:.m: m.m+m.~m mewpcsn owwncH
I I. I I Amzmoaaw> momooopc2wov
m.o+m.o m.o+:.o m.o+m.o o.o+c.a pmxomgcooz khan:
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79

 

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I I I I. AmwmcmCMHohmo muwwmv
:.o+:.o m.o+m.o m.o+m.o m.o+a.o savages: umpmmmpnImpwcg
I I I I. ”mamCMEmmm.mmwmooomv
w.H+:.m m.o+n.o o.m+:.m N.H+w.H Boupmam uwmmm>
I I I I. “~0H00wn w:ooummcprv
o.~+o.~ m.H+o.m m.H+o.: o.o+o.o 3oHHmzm mmpe
I I. I I Amwpmwdz> wscpzpmv
N.m+o.m :.H+m.m m.o+m.~ o.o+o.o mcfifipmwm
I I I I. hawnone mufimmoamzv
H.m+m.mm «.ofi+m.mm H.~H+H.m: m.mfl+w.~m gopumam mcom
I I I I Amsefimnpzaopcyagm oawmfimv
m.a+:.md o.mI~.~H ~.H+:.H n.m+o.ofi amazop omonImzogsm
I I I I Amflunsaoo mzzooawnop<v
m.o+o.fi «.H+H.d ~.~+m.~ o.o+o.o npflnmcfieesz ompmonspnznsm
wwpe macs; Hmcowpcw>coo mmue macs: Hmcowpcm>coo mmwomam cufim
d mmwao hwmw N mmmHo ham»

 

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