OVERDUE FINES:

25¢ per W per item

RETURMNG LIBRARY MATERIALS:

Place in book return to remove
charge from circuhflon records

 

 

 

AN ANALYSIS OF INTER- AND INTRA-ITREE EFFECTS
ON THE SIZE OF SPRugE BUDWORM EGG NSASSES
ON BALSAM FIR AND WHITE SPRUCE
by

Bruce Allen Montgomery
A THESIS

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

MASTER OF SCIENCE

Department of Forestry
l 98 l

 

rChoristoneura fumiferana (Lepidoptera: Tortricidae)
2Abies balsamea (L.) Mill.
3Picea glauca (Muench) Voss

 

 

 

 

1; //

a //

ABSTRACT

AN ANALYSIS OF INTER- AND lNTRA-ITREE EFFECTS
ON THE SIZE OF SPRUEE BUDWORM ECG I~¢3ASSES
ON BALSAM FIR AND WHITE SPRUCE
by

Bruce Allen Montgomery

Spruce budworm, Choristoneura fumiferana (Clemens), egg masses on balsam

 

fir, Abies balsamea (L.) Mill., and white spruce, Picea glauca (Muench.) Voss, were

 

examined to determine if populations oviposit larger or smaller sized egg masses at
(a) particular site(s) on a host tree, within a stand, within a forest complex and
between tree species within a stand. Five stands exhibiting negligible defoliation
and low budworm numbers were studied in the Ottawa National Forest in
Michigan's Upper Peninsula. The mean number of eggs per egg mass does not differ
significantly from one directional quadrant to another within a host tree, from one
height division to another within a host tree, and, in general, from one host tree to
another of the same species within a spruce-fir stand. The mean number of eggs
per egg mass did not differ significantly between the five stands for each tree
species. Egg masses found on balsam fir averaged M to 5.6 more eggs per mass
than eggs found on white spruce within each stand. However, in only two of the

five stands were such differences statistically significant.

'2).

Dedicated to my parents,

Dr. Keith and Rosalind Montgomery

ii

ACKNOWLEDGMENTS

I am deeply indebted to my advisor, Dr. Gary Simmons, for his exceptional
guidance, patience, and encouragement. In most facets of my education, Gary
gave me the freedom to make my own decisions and, consequently, learn from
my own mistakes.

Dr. Gary Fowler's comments and assistance on statistics, field
measurements, and general research techniques and analytical methods were
invaluable.

Special thanks to Dr. John Witter and Jack McCarthy for their suggestions
and assistance.

Much appreciation to our hosts at Camp Filibert Roth, Mac Smith and Arch
Cowan.

My most sincere appreciation to the best research crew that I've been
associated with--my assistant John Simon, and Ray Drapek, Andrea Erickson,
Bryan Hill, Maryanne Kendricks, William Overbaugh, Tom Shreiner, Erik
Sorenson, and Susan Todd.

Work leading to this thesis was funded by a USDA Forest Service-sponsored
program entitled Canada/United States Spruce Budworm Program administered
through the USDA Science and Education Administration (grant 904-lS-I I).

iii

PREFACE

This is the second thesis that l have written on the subject of spruce
budworm egg mass size. The first, An Analysis of Intra- and Inter-Tree Effects
on the Size of Spruce Budworm Egg Masses on Balsam Fir (Montgomery I98I),
was based on data collected in the Ottawa National Forest of Michigan in the
summer of I979. In the first thesis, I analyzed the distribution of mean egg mass
size within and between balsam fir trees in stands that were experiencing light-
moderate to moderate-severe defoliation.

The second thesis is based on data collected in Michigan‘s Ottawa National
Forest in the summer of I980. Here, I investigate the intra- and inter—tree
effects of balsam fir and white spruce on mean egg mass size. Unlike the first
thesis, these stands were supporting endemic spruce budworm populations and

were suffering very little defoliation damage.

iv

TABLEOFCONTENTS

List of Tables ............................... . .................. viii
List of Figures .................................................. xvii
I. Introduction ............ . . . . . . . ........................... 1
Life History of the Spruce Budworm ........................ 2
Spruce Budworm Egg Mass Size ................. . ........... 3
Objectives of the Study .......................... 7
ll. Field Methods ............... . . ........................... 8
III. Regression Estimates for the Number of Eggs Per
Egg Mass Length--Balsam Fir and White Spruce ...... . ........ 16
Methods ................................................. 16
Regression Estimates ....................... . . . . . . ........ 20
IV. Laboratory Methods ....................................... 21
V0 ReSUlts eoeeeoeeeeeeeeeeeeooeeee eeeeee one eeeeee 34
Section I. An Analysis of Quadrant Effects on the
Size of Spruce Budworm Egg Masses on .
Balsam Fir . . ......... . . . . ..................... 35
Section 2. An Analysis of Stratum Effects on the
Size of Spruce Budworm Egg Masses on
&lsomFir ..... OOOOOOOOOOOOOOOOOOOCOOOII.O... 37
Section 3. An Analysis of the Effects of Thirteen
Cells (predesignated compartments
within balsam fir trees) on the Size of
SpruceBudwormEggMasses.................... 39
Section 4. Analyses of Inter-Tree Effects on
Spruce Budworm Egg Mass Size on
Balsam Fir Within Five Stands , , , , , ,,,,,,,,,,,,, 41

TABLE OF CONTENTS, continued

Section 5. An Analysis of Inter-Stand Effects on
Spruce Budworm Egg Mass Size on
Balsam Fir Within a Forest Complex . . .............. 47
Section 6. An Analysis of Quadrant Effects on the
Size of Spruce Budworm Egg Masses on
White $ruce eeeeeeeeeeeeeeeeeeeeeeoeoeeoeoooeoee 49
Section 7. An Analysis of Stratum Effects on the
Size of Spruce Budworm Egg Masses on
White Spruce ..... 51
Section 8. An Analysis of the Effects of Thirteen
Cells (pre-designated compartments
within white spruce trees) on the Size
of Spruce Budworm Egg Masses . . . . . . . ............. 53
Section 9. Analyses of Inter-Tree Effects on
Spruce Budworm Egg Mass Size on
White Spruce Within Five Stands . . . . . . . . ........... 55
Section I0. An Analysis of Inter-Stand Effects on
Spruce Budworm Egg Mass Size From
White Spruce Within 0 Forest Complex .............. 61
Section I I. Analysis of Inter-Species (trees) Effects
on Spruce Budworm Egg Mass Size
Within a Forest Complex .......................... 65
Section I2. Analysis of Inter-Species (trees) Effects
on Spruce Budworm Egg Mass Size
Within 0 Forest Complex ........... . . . ............ 74
VI. Discussion .................. . . ....... . . . .................... 77
VII. List of References ........ . . . . . . . ...... . . . . . ............... . . 86
Appendices... ....... OIOOOOOOOOOOOOOOOOO...OOOOOOOOOOOOOOOOOOOOOOO 90
Appendix A. Six maps denoting the location of the

five sampled stands within the
Ottawa National Forest and the
location of each stand within the
corresponding townships . . . . . . . . . . ....... . ....... 90

vi

I

TABLE OF CONTENTS, continued

Appendix B.

Appendix C.

Appendix D.

Appendix E.

Appendix F.

Appendix G.

Diameter, age, height and total
number of egg masses for each
sampledfreeoeeoeeoeooeeeoeeeeeee

Distance, angles, diameters and
heights of the ten "nearest neighbors"
(trees with diameters at breast
height greater than l2.0 cm) of each
sampledtree............... ..... .

Stemmaps...... ...... ........

Normality and homogeneity test
results for egg mass groups used in

....... .. 107

....... 110

............ 118

parametriCStatiSticseeoeeeeoeeeeeeeeeeeeeeoeoe142

Normality and homogeneity test
results for egg mass groups used in

parametric StatiSfics0.000.000.0000. ........ 0.. 156

Number ofeggsper massper ceII 185

vii

Table I.

Table 2.

Table 3.

Table 4.

Table 5.

Table 6.

Table 7.

Table 8.

Table 9.

Table I0.

LIST (I: TABLES

Regression estimates for the number of eggs
per egg mass length from balsam fir trees in
MiChigan. e eeeeeee eeeeeeeoeeeeeeeeeeeeeeee eeeeeeeeeeee 22

Regression estimates for the number of eggs
per egg mass length from white spruce trees in
MiChigan. eeeeeeeeeeeeeeeeeeeeeeeoeeee eeeeeeeeeeeeeeee 24

The mean number of eggs per egg mass
(transformed back into original form) from the
north, east, south, and west quadrants of
balsam fir trees ...... . ...... . . . . . . . . . .................. 36

Anova table for data in Table 3—Analysis of
differences in the mean number of eggs per
egg mass between four quadrants within
balsam fir trees............................ ............ 35

The mean number of eggs per egg mass
(transformed back into original form) from the

lower, middle, and upper strata of balsam fir
trees... eeeeeee eeeeeeeeeeeeeeee eeeeeee oeeeee eeeeeeeeee 38

Anova table for data in Table S—Analysis of
differences in the mean number of eggs per

egg mass between three strata within balsam
firtrees. eeeeeeeeoeeoeeeeeeeeeeeeeeeee eeeeeeeeeeeeeeee 38

The mean number of eggs per egg mass
(transformed back into original form) from
thirteen cells within balsam fir trees. . - - . - - - ----------- . . 40

Anova table for data in Table 7—Analysis of
differences in the mean number of eggs per

egg mass between thirteen cells within balsam
fir trees. eeeeeeeeeeeeeoeeeoeeoeeeoeeeeeeeeeoe ooooooooo 40

The mean number of eggs per egg mass
(transformed back into original form) from
four balsam fir trees within Stand I. .. .- . .. . . ...... 42

Anova table for data in Table 9-Analysis of
differences in the mean number of eggs per

egg mass between four balsam fir trees within
Stand I. eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee ooooo 42

viii

LIST OF TABLES, continued

Table I I.

Table l2.

Table I 3.

Table I4.

Table I 5.

Table I6.

Table I7.

Table I 8.

Table I 9.

Table 20.

The mean number of eggs per egg mass
(transformed back into original form) from
two balsam fir trees within Stand 2. . . . , . , , , . , ............ 43

Anova table for data in Table Il-Analysis of
the difference in the mean number of eggs per
egg mass between two balsam fir trees within
Stand2. .. ........ ........... 43

The mean number of eggs per egg mass
(transformed back into original form) from
two balsam fir trees within Stand 3. , . , , , , , , , ,,,,,,,,,,,,, 44

Anova table for data in Table I3—Analysis of
the difference in the mean number of eggs per
egg mass between two balsam fir trees within
Stand 3. .......... 44

The mean number of eggs per egg mass
(transformed back into original form) from
two balsam fir trees within Stand 4. . . . . . . . . . . ........... 45

Anova table for data in Table I5-Analysis of
the difference in the mean number of eggs per
egg mass between two balsam fir trees within
Stand ll, ...................o................ ...... o... 45

The mean number of eggs per egg mass
(transformed back into original form) from
two balsam fir trees within Stand 5. . . - - - - - - . . . . . ......... 46

Anova table for data in Table l7-Analysis of
the difference in the mean number of eggs per
egg mass between two balsam fir trees within
Stands, eeeeeeeeoeeeeeeeeo ..... eeeeeeoooee ........... 46

The mean number of eggs per egg mass
(transformed back into original form) from ten
balsam fir trees in five stands in the Ottawa
Nationa] Forest.o................o........... ooooooo ..o 48

Anova table for data in Table l9—Analysis of
differences in the mean number of eggs per
egg mass between balsam fir trees in five
spruce-fir sfandg,........................... ........... 43

ix

LIST OF TABLES, continued

Table 2|.

Table 22.

Table 23.

Table 24.

Table 25.

Table 26.

Table 27.

Table 28.

Table 29.

Table 30.

The mem number of eggs per egg mass
(transformed back into original form) from the
north, east, south, and west quadrants of white
spruce "995- .... ........ 50

Anova table for data in Table Zl—Analysis of
differences in the mean number of eggs per
egg mass between four quadrants within white
spruce trees. ......... 50

The mean number of eggs per egg mass
(transformed back into original form) from the
lower, middle, and upper strata of white
Sprucetrees ...... . ...... ............... 52

Anova table for data in Table 23—Analysis of
differences in the mean number of eggs per
egg mass between three strata within white
sprucetrees. . ..... . ..... ................... .......... 52

The mean number of eggs per egg mass
(transformed back into original form) from

Anova table for data in Table 25—Analysis of
differences in the mean number of eggs per
egg mass between thirteen cells within white
spruce trees- ...... 54
The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce trees within Stand l. . , , , , , , , , , , ,,,,,,,, 56

Anova table for data in Table 27-Ana|ysis of
the difference in the mean number of eggs per
egg mass between two white spruce trees
within Stond l- ........... 56

The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce trees within Stand 2. 57

Anova table for data in Table 29—Analysis of
the difference in the mean number of eggs per
egg mass between two white spruce trees
within Stand 2. 57

LIST OF TABLES, continued

Table 3|.

Table 32.

Table 33.

Table 34.

Table 35.

Table 36.

Table 37.

Table 38.

Table 39.

Table 40.

The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce trees within Stand 3. 58

Anova table for data in Table 3 I-Analysis of
the difference in the mean number of eggs per
egg mass between two white spruce trees
within Stand 3. ............................. ........ ... 58

The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce trees within Stand 4. . . . . . . . . ............ 59

Anova table for data in Table 33-Analysis of
the difference in the mean number of eggs per
egg mass between two white spruce trees
within Stand 4. ............................. ........... 59

The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce trees within Stand 5. . . . . . . . . . . . . . . . . . . . 60

Anova table for data in Table BS—Analysis of
the difference in the mean number of eggs per
egg mass between two white spruce trees
within Stand 5. 60

The mean number of eggs per egg mass
(transformed back into original form) from ten
white spruce trees in five stands in the Ottawa
NationalForest.. .......... .......... 63

Anova table for data in Table 37-Analysis of
differences in the mean number of eggs per
egg mass between white spruce trees in five
stands in the Ottawa National Forest. 63

Results of the Kruskal—Wallis Test, comparing
mean egg mass sizes on white spruce between
a” five Stands. eeoeeeeeeeeeeeeeeeeeeeeeeeeoeeeeeeeeeeee 64

The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce and two balsam fir trees in

STOfld lo"'0'0"0000000eeeeeooeeoeeeeeeeeeeoooeeeeeeeo 67

xi

LIST OF TABLES, continued

Table 4|.

Table 42.

Table 43.

Table 44.

Table 45.

Table 46.

Table 47.

Table 48.

Table 49.

Table 50.

Anova table for data in Table 40—Analysis of
the difference in the mean number of eggs per
egg mass between two white spruce and two
balsomfirtreesinsmnd I .......... 67
The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce and two balsam fir trees in
StondZ- ............... 68
Anova table for data in Table 42—Analysis of
the difference in the mean number of eggs per
egg mass between two white spruce and two
bolsom fir trees in Stand 2. 68

The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce and two balsam fir trees in
Stand 3 69
Anova table for data in Table 44-Analysis of
the difference in the mean number of eggs per
egg mass between two white spruce and two
bolsom fir trees in Stand 3. .......... 69
Results of the Mann-Whitney Test comparing
mean egg mass size on white spruce with mean
egg mass size on balsam fir in Stand 3. , , , , ............... 70

The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce and two balsam fir trees in
Stand‘t- ............ 71

Anova table for data in Table 47—Analysis of
the difference in the mean number of eggs per
egg mass between two white spruce and two
bolsom fir trees in Stand 4 ......... 71
The mean number of eggs per egg mass
(transformed back into original form) from
two white spruce and two balsam fir trees in

Stands. O....0.00...O...........IOOOOOCOOOOOOO0...... 72

Anova table for data in Table 49—Analysis of

the difference in the mean number of eggs per

egg mass between two white spruce and two

bolsom fir trees in Stand 5 72
x11

LIST OF TABLES, continued

Table 5 I. Results of the Mann-Whitney Test comparing
mean egg mass size on white spruce with mean
egg mass sizeon balsam fir in Stand 5. 73

Table 52. The mean number of eggs per egg mass
(transformed back into original form) from ten
white spruce and ten balsam fir trees in five

Stands.a.eeeooeecone-0000000000....oeooeooooooooeoooo 75

Table 53. Anova table for data in Table 52—Analysis of
the difference in the mean number of eggs per
mass between ten white spruce and ten balsam
fir trees in five stands................................. 75

Table 54. Results of the Mann-Whitney Test comparing
mean egg mass size on white spruce with mean
egg mass on balsam fir over all five stands. . - . . . - - . - - . . . . 76

Table B I. Sampled trees' ages, heights, diameters, and
total number Of egg masses. eeeeeeeeeeeeeeeeeeeoeeeeeee 108

Table C I. Locations, heights and diameters of trees that
were within ten meters of each sampled tree. . . ..... . . . . . 111

Table F I. Normality and homogeneity test results for
egg masses from balsam fir, used for
regreSSim estimates. eeeeeeeeeeeeeeeeeeeeeeeeeeoeeeeee159

Table F2. Normality and homogeneity test results for
egg masses from white spruce, used for
regreSSim estimates. eeeeeeeeeeeeeeeeeeeeoeeeeooeoeoee 160

Table F3. Normality and homogeneity test results for
egg counts (transformed into square root) from
north, east, south, and west quadrants within
balsam fir trees. ..................................... 161

Table F4. Normality and homogeneity test results for
egg counts (transformed into square roots)
from lower, middle, and upper strata within
balsam fir trees, .....................................152

Table F5. Normality and homogeneity test results for

egg counts (transformed into square roots)
from '3 cells Within ba'sam fir trees. a. .00.. ee 0 o e. e e .0 e e 163

xiii

LIST OF TABLES, continued

Table F6.

Table F7.

Table F8.

Table F9.

Table F IO.

Table Fl I.

Table FI2.

Table F l 3.

Table FI4.

Table F I 5.

Table F I 6.

Normality and homogeneity test results for
egg counts (transformed into square roots)

from four balsam fir trees within Stand I. . . . . .. . . .. . . . . . . 164

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two balsam fir trees within Stand 2. . . . . . . . . . . . . . . . .

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two balsam fir trees within Stand 3. . . . . . . . . . . . . . . . .

Normality and homogeneity test results for
egg counts (transformed into square roots I
from two balsam fir trees within Stand 4. . . . . . . . . . . . . . . . .

Normality and homogeneity test results for
egg counts (transformed into square roots)
from four trees within Stand 5.

Normality and homogeneity test results for
egg counts (transformed into square roots)
from balsam fir trees in the stands in the
Ottawa National Forest.

Normality and homogeneity test results for
egg counts (transformed into square roots)
from north, east, south and west quadrants
within white spruce trees.

Normality and homogeneity test results for
egg counts (transformed into square roots)
from lower, middle, and upper strata within
white spruce trees

Normality and homogeneity test results for
egg counts (transformed into square roots)
from thirteen cells within white spruce trees. . . . . . . . . . . . .

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two white spruce trees within Stand I. . . . . . . . . . . . . . .

Normality and homogeneity test results for

egg counts (transformed into square roots)
from two white spruce trees within Stand 2. . . . . . . . . . . . . . .

xiv

165

166

167

168

169

170

171

172

173

174

LIST OF TABLES, continued

Table F l 7.

Table F I 8.

Table F I 9.

Table F20.

Table F2 I.

Table F22.

Table F23.

Table F24.

Table F25.

Table F26.

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two white spruce trees within Stand 3. . . . . . . . . . . . . . .

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two white spruce trees within Stand 4. . . . . . . . . . . . . . .

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two white spruce trees within Stand 5. . . . . . . . . . . . . . .

Normality and homogeneity test results for
egg counts (transformed into square roots)
from white spruce trees in five stands in the
Ottawa National Forest. ...............................

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two balsam fir and two white spruce
trees within Stand I. ..................................

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two balsam fir and two white spruce
trees within Stand 2. ..................................

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two balsam fir and two white spruce
trees within Stand 3. eeeeeoeeeeeeeeeoeeeeeeeeeeooeeooee

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two balsam fir and two white spruce
trees within Stand ll, o.......o........o.o.o.o......-...

Normality and homogeneity test results for
egg counts (transformed into square roots)
from two balsam fir and two white spruce
trees within Stand 5. eeeeeeeeeeeoeeeeeeeeeeeeeeeeoeeoee

Normality and homogeneity test results for
egg counts (transformed into square roots)
from ten balsam fir and ten white spruce trees
in five stands within the Ottawa National

Forest.eeeeeeoeeoeeeeeeeeeoeeeeeoeeeoeeeeeeeeeeeeoeee

XV

175

176

177

178

179

180

181

182

183

184

LIST OF TABLES, continued

Table G I. Distribution of individual egg counts within
eaCh sampled tree.eeeeeeeoeeoeeeeeoeeeeoeee eeeeee 0000. 186

xvi

Figure I.

Figure 2.

Figure 3.

Figure 4.

Figure 5.
Figure 6.
Figure 7.

Figure 8.

Figure 9.

Figure I0.

Figure I I.

Figure I 2.

Figure AI.

LIST OF FIGURES

Severely defoliated pockets of spruce and fir
trees, photographed from the Tepee Lake fire
tower, eight miles south of Kenton, Michigan. ........... 9

Severely defoliated balsam fir tree,
photographed in the Ottawa National Forest in
I980. ....... ...... ..... O ..... 0.0.0.... OOOOOOOOOOOOOO 10

Photograph of a sampled tree—White spruce 2,
StandZQ .0.........OOOOOOOOOOOOOOOOOO 0000000000000000 14

Photograph of a sampled tree—Balsam fir I,
Stond3................OIOOOOCOOOOOOOOOO.... ..... 0.0.15

Photograph of an eclosed 2-row egg mass. . . . . . . . ....... . . 17
Photograph of an eclosed 3-row egg mass. . . . . . . . . . . . . . . . . 18

Photograph of an eclosed 2-row egg mass with
oportiOI third rOWOOOOO......OOOOOOOIOOIOOOOO 0000000000 19

Plot of the estimated number of eggs per egg
mass length, comparing 2-row egg masses
from white spruce and balsam fir. . . . . . . . . . . . . . . ......... 27

Plot of the estimated number of eggs per egg
mass length, comparing 2-row egg masses with
a partial third row from white spruce and
balsam hr 29

Plot of the estimated number of eggs per egg
mass length, comparing 3-row egg masses
from white spruce and balsam hr 31

Diagram of a sampled tree—Side view.
Division of live crown into three strata and a

top.0.0.0.0000.........OOOOOOOOOOOOOOIOOO000......... 33

Diagram of a sampled tree—Top view.
Division of live crown into four quadrants" - - . . . . . . . ...... 33

Map of the Ottawa National forest with
locations of the five study sites. .... 91

xvii

LIST OF FIGURES, continued

Figure A2.

Figure A3.

Figure A4.

Figure A5.

Figure A6.

Figure D I.
Figure D2.

Figure D3.

Figure D4.

Figure DS.

Figure D6.

Figure D7.

Figure D8.

Figure D9.

Figure D ID.

Map of the five study sites within the Iron
River, Watersmeet, and Kenton Districts of

themfowoNationo'Forest.eeeeeeeeeeeoeeeee ooooooo eooo 93
Map of the locations of Stand l and Stand 2 in
Section 6 of Township T45N, R36W, Iron
county’MiChigm. 00............OOOOOOOOOOOO0.... ..... 97
Map of the location of Stand 3 in Section 30 of
Township T46N, R37W, Iron County, Michigan. . . . . . . . . . . . . 100
Map of the location of Stand 4 in Section 32 of
Township T46N, R38W, Ontonagon County,
MiChigon' oeeeeeeeeeeeeeeeeeeooeeeoeeeoeeeeoe ooooooooo 103
Map of the location of Stand 5 in Section I2 of
Township T44N, R37W, Iron County, Michigan. . . . , . ...... . 105
Key to symbols used in Figures D2-D23. . . . . . . . . . . ........ 11g
Stem map of the ten nearest trees that
encompassed sample tree BF l in Stand l. . . . . . . . .......... 120
Stem map of the ten nearest trees that
encompassed sample tree BF2 in Stand l. . . . . . . . . . . . . . . . . . 121
Stem map of the ten nearest trees that
encompassed sample tree BF 5 in Stand l. , , , , , ,,,,,,,, , , , , 122
Stem map of the ten nearest trees that
encompassed sample tree BF6 in Stand l. , _ , , , , , , , , , , , , , , , 123

Stem map of the ten nearest trees that
encompassed sample free WSI in Stand |., , , , , _ , , , , , , , , , , , 124

Stem map of the ten nearest trees that
encompassed sample tree WSZ in Stand I.. . . , . . , , . , , . . , . . . 125

Stem map of the ten nearest trees that
encompassed sample tree BFI in Stand 2. , , , , , , , , , , , , , , , , , 125

Stem map of the ten nearest trees that
encompassed sample free BF2 in Stand 2.” 127

Stem map of the ten nearest trees that
encompassed sample tree WSI in Stand 2., , , , . , , , , , . , , , , , , 123

xviii

LIST OF FIGURES, continued

Figure DI I.

Figure D I 2.

Figure D I 3.

Figure D I 4.

Figure D l 5.

Figure D | 6.

Figure D l 7.

Figure D I 8.

Figure D I 9.

Figure D20.

Figure D2 I.

Figure D22.

Figure D23.

Figure El.
Figure E2.
Figure E3.

Stern map of the

ten nearest trees

encompassed sample tree W52 in Stand 2.

Stem map of the

encompassed sample tree BFI in Stand 3..

Stem map of the

encompassed sample tree BF 2 in Stand 3..

Stem map of the

ten nearest trees

ten nearest trees

ten nearest trees

encompassed sample tree WSI in Stand 3.

Stem map of the

ten nearest trees

encompassed sample tree W52 in Stand 3.

Stem map of the

ten nearest trees

encompassed sample tree BFI in Stand 4.

Stem map of the

encompassed sample tree BF2 in Stand 4..

Stem map of the

ten nearest trees

ten nearest trees

encompassed sample tree WSI in Stand 4.

Stem map of the

ten nearest trees

encompassed sample tree W52 in Stand 4.

Stem map of the

ten nearest trees

encompassed sample tree BF I in Stand 5.

Stem map of the

encompassed sample tree BF 2 in Stand 5..

Stem map of the

ten nearest trees

ten nearest trees

encompassed sample tree WSI in Stand 5.

that

that

that

that

that

that

that

that

that

that

that

that

Stem map of the ten nearest trees that
encompassed sample tree W52 in Stand 5. . . . . . . . . . . . . . . . .

Thefieldcampat Stand 5.

Sampled tree BF2 in Stand 3.

&mpledtree WSI in Stand 2.0..........OOOOOOOOCOCOOOOO

xix

129

130

131

132

133

134

135

136

137

138

139

140

141
143
144
144

LIST OF FIGURES, continued

Figure E4.

Figure E5.

Figure E6.

Figure E7.
Figure E8.

Figure E9.

Figure E I 0.

Figure Ell.

Figure E I 2.

Figure E I 3.

Figure E I4.

Figure E I 5.

Figure E I 6.

Measuring the height of a branch—a research
OSSESthf on the sampled tree. a o o o o o o o o o o o o o o ooooooooooo 145

Measuring the height of a branch and noting

its directional quadrant—a research assistant
mheground. ona.nooooooooooooooooaooooooooso...so...146

Recording the branch number, height and
quadrant--a research assistant on the ground.- - - - . - - - . . - - - 147

cutting the sampled bronCh.oooooooooooooooooooooooooooo 148

Dropping the branch to the spotter on the
ground. 0......IO......OOOOOOOOOOCOOOOOO...0000......148

Catching a sampled branch that was dropped
frOm the tree.oooooa.ooooooooaoooooooooococo-coooooooo 149

Tying the identification tag on to the branch.- - - - - - ° ° ° ° ° ° - 150

A pile of severed and tagged branches near the
screen tent-coooooooooococoa-o0.0000000000000000.ooooo 150

Research assistants measuring the foliated
Widfhofobronch. ooooooooooooooaooo0.00.00.00.00 ccccc 151

Research assistant examining foliage for egg

mosses. one...cocoo.coco...0.0000000000000000o.000000152

Research assistant removing a needle and an
egg mossfromafoliageclipping-........................ 153
Measuring the foliated area of a sampled

branch on a grid board.

......OOOOOOOOOOOOOOOOO..0.00...154

The top of the sampled tree.

...OOOOOOOOOOOOOOOOOO ..... O. 155

XX

I. INTRODUCTION

The spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera:
Tortricidae) is native to the northern boreal forests of North America.
Outbreaks occur in 30-40 year intervals, spaced by 20-90 years, and are
considered a "phase in the natural cycle of events associated with the
maturing of balsam fir" (Blais I954, I960). As the preferred hosts, balsam fir

(Abies balsamea (L.) Mill.) and white spruce (Picea glauca (Muench) Voss),

 

approach maturation age, the spruce budworm population rises from the
endemic level. It reaches epidemic proportions when there are consecutive
years of dry, sunny weather (Greenbank I963a, b; Baskerville I975) and
extensive unbroken, mature spruce-fir stands (Mott I963).

The present epidemic started in the I960$ and encompasses
approximately 60 million hectares of spruce-fir forests in eastern North
America with resulting timber losses as high as 283 million cubic meters
(Witter et al. I980). Pulp and paper industries are dependent on the spruce-
fir forest for provision of a continual supply of usable balsam fir and white
spruce fiber. Unfortunately, these industries are competing with a highly
successful organism for the same resource. In the past, man has detained
budworm damage through chemical spray programs, first aimed at killing the
pest and more recently directed at short-term perpetuation of the tree's life.
Blais (I958) postulated, "saving the fir by spraying possibly helps to
perpetuate budworm epidemics that would otherwise die out in a short period

with the change in forest composition." New studies have been initiated to

find environmentally safe and economically feasible, long-term control

methods that are effective alternatives to chemical spraying. Improving

spruce budworm population sampling methods, particularly when population
levels are low to moderate, is essential to this task. "Although at this time it
is hard to imagine an alternative to chemical treatment in widespread
outbreaks that cover millions of acres, the potential for such an alternative
does exist. If it can be demonstrated that outbreaks arise from discrete
centers, and if a monitoring system can be developed that will detect these
centers early enough, it is possible that measures other than chemical may be
applied, singly or in combination, to extinguish embryonic outbreaks" (Blais

I973).

Life History of the Spruce Budworm

The spruce budworm has one generation each year. Female moths lay
egg masses on spruce and fir needles in July and early August. Each mass can
contain from I to 60 eggs, and each female can lay around 200 total eggs.
The preferred oviposition host is white spruce (Jaynes and Speers I949;
Wilson I963, Kemp I98I), although oviposition occurs to a great extent on
balsam fir and to a lesser extent on red spruce (Picea rubens Sarg.) and black

spruce (Picea mariana Mill. (B.S.P.)) (Wilson I963). Eclosion occurs within 8

 

to l2 days, and the small first instars, dropping from branches on silken
threads, are often wind-borne and dispersed among trees in a stand. Feeding
does not occur during this stage. The larvae find overwintering sites on
branches, among bark scales, lichen mats, and staminate flower bracts, molt

once and hibernate. In May the second instars emerge to mine older needles

and flower buds. A second dispersal occurs in which many larvae may be
redistributed over the entire forest. As the larvae grow older and larger,
they become free feeders, preferring the succulent new foliage of balsam fir
but also feeding on white, black and red spruce and occasionally on hemlock,
pine, larch, and blue spruce. They form feeding shelters on branch tips, molt
through a sixth instar, and pupate in the shelters in late June and early July.
After about I2 days, the adult moths emerge and mate. Typical female
moths lay two large egg masses at their eclosion and mating site, and are
then wind—dispersed on an "exodus flight" for up to 80 km downwind per night

(Harvey I977). Egg depositon continues until the female expires.

Spruce Budworm EggMass Size

The egg mass is a logical stage to sample since it can be expressed in
terms of a basic unit (e.g., branch‘surface) and an absolute unit (e.g., area)
(Morris I955) and is retained on the needle surface for several months. Also,
the egg stage can be sampled in the early fall, providing ample lead time for
planning the following summer's management program and for appropriating
funds.

Current egg mass sampling methods are inconsistent. Sampling
schemes developed by Morris (I949, I954, I955) and others since then
disagree on the type, size and number of sampling units required. There are
variable recommendations as to where to sample within a tree and how many
trees to sample within a management unit. The vital relationship between
egg mass density and defoliation has not been clarified.

Since female spruce budworm moths lay different size egg masses, it is

imperative that this distribution be assessed to optimize sampling accuracy
and precision. Studies on egg mass size have neglected investigating the
number of eggs per egg mass in different parts of trees, on different trees on
host species or in different stands. It has been cited that the average number
of eggs per egg mass is about 20 (Morris I963; Neilson I963; Eidt and
Cameron I970; Harvey I977) or about l8 (Morris I954) or variations between
II and 25 (Blais I958; Harvey I977; Otvos I977). It has been suggested that
certain interrelated factors, such as severity of infestation, food quality and
quantity, and population vigor may be responsible for observed differences in
egg mass size.

Blais (I952) reported that in a year when some sample plots were
heavily infested with the spruce budworm and some plots were lightly to
moderately infested, the average number of eggs per cluster was 20.3. The
following year, when all plots were heavily infested, the average number of
eggs per cluster was l7.8. Blais (I953) found that "adults from fifth- and
sixth-instar larvae reared on old foliage produced fewer eggs than insects
reared on the current year's growth." He noted that as populations increase
to the point where the current year‘s growth is destroyed prior to completion
of the larval stage, fecundity decreases. Fecundity increases again as
populations decline to the point where there is incomplete destruction of the
current year's growth.

Miller (I957) reported an average of l5.7 eggs per egg mass in sample
plots with severe infestation and an average of l8.5 eggs per egg mass in
sample plots with light infestations. He postulated that "if a population is

subject to larval starvation, the resultant small adults tend to lay smaller

masses." Thus, "mean eggs per mass is dependent on the degree of
infestation."

In a severe spruce budworm infestation in the Shickshock Mountains of
New Brunswick, Blais (I958) reported an average of 326 egg masses per I00
square feet of foliage in 75% of 52 sampled plots. From this egg mass pool,
I00 clusters were examined and were found to have an average of I l.0 eggs
per cluster.

Harris (I963) observed the identical relationship in the two-year cycle
spruce budworm. He observed a drastically reduced number of eggs per mass
after two years of severe defoliation. In one study area, the average number
of eggs per mass an alpine fir (Abies lasiocarpa (l-took.) Nutt.), was estimated
to be 5|.9 in I958 but dropped to 20.2 in I959. He SUggested "that moths
which developed from larvae starved because of severe defoliation were
incapable of laying as large a number of eggs as those that developed from a
healthier I958 population, or that only the weaker individuals with lower egg-
Iaying capacities remained in the area."

McKnight (I969) observed certain trends in egg mass size of the

western spruce budworm, Q. occidentalis Freeman. He reported that "in the

 

Rocky Mountain region, I959-I965 . . . egg-mass size fluctuated widely: the
average number of eggs per mass declined from 25.3 in I959 to 23.6 in I96I
when egg mass densities were increasing; rose sharply to 3|.9 in I962 when
egg mass densities were low; and fluctuated between 24.0 and 29.7 in I963,
I964, and I965, when egg mass densities were generally low but variable."
Washburn and Brickell (I973) feel that McKnight's (I969, I97l) observation

"suggests that egg mass size could be a good indicator of the place of a given

infestation in the epidemic cycle."

Outram “97” studied spruce budworm specimens that "were one
generation removed from field-collected stock originally taken from an
epidemic population." Twenty fertilized females, reared in a laboratory,
produced egg masses with a low average of about l6.7 eggs.

Otvos (I977) reported an average of 25 eggs per egg mass for I70
masses collected from 50 cm long branch tips taken from the mid-crown of
dominant balsam fir trees. He explained that the relatively large number of
eggs per mass was possibly a result of a small sample size.

Harvey (I977) found that size and mean weight of eggs in successive
clusters drops gradually during the oviposition of most female spruce
budworm moths. He also found that "dietary limitations during (larval)
development substantially reduce fecundity and steepen the curve of
decreasing egg weight." He reported that the typical moth lays 01 average of
slightly more than two clusters at the site of her eclosion and mating. These
are the first clusters she lays and comprise the heaviest eggs.

Recently, Kemp “98” reported an average of l7.3 and l9.I eggs per
egg mass from laboratory-reared and mated female spruce budworm moths on
balsam fir in two separate experiments in Maine. In what I believe is the
first reported test of egg mass size on white spruce foliage, Kemp (I98l)
found an average of l4.6 and l4.3 eggs per egg mass produced by laboratory-
reared budworm moths in two separate experiments. However, his detailed
study of the capabilities of budworm and forest variables as predicting agents
of budworm population size yielded no information on mean egg mass from

balsam fir or white spruce in the field.

Montgomery 0980 conducted an analysis of mean egg mass size within

and between balsam fir trees in four spruce-fir stands in Michigan's Upper

Peninsula. He found that the mem number of eggs per egg mass did not
significantly differ from one height division to another within a host tree,
from one directional quadrant to another within a host tree, from one of
thirteen predesignated compartments to another within a host tree, or from
one balsam fir tree to another within a spruce-fir stand. Mean egg mass size
was significantly different between the four spruce—fir stands. Balsam fir
trees in two light-moderately defoliated stands had averages of I8.l and I9.3
eggs per mass as opposed to an average of I7.I eggs per egg mass on balsam

fir trees in each of two moderate-severely defoliated stands.

Objectives of the Study

The overall objective of this study was to analyze differences in egg
mass size in spruce-fir stands that had very low spruce budworm population
and defoliation levels. Specifically, I wmted to determine if egg mass size
on white spruce and balsam fir trees was related to particular sites within a
host tree, within a stand, or within a forest complex.

For egg masses collected from balsam fir trees, I analyzed:

I. intra-tree quadrant effects;
2. intra-tree stratum effects;
3. intra-tree compartment (cell) effects;
4. inter-tree effects within a stand; and
5. inter-stand effects within a forest complex.

For egg masses collected from white spruce trees, I also analyzed:

l. intro-tree quadrant effects;

2. intro-tree stratum effects;
3. infra-tree compartment (cell) effects;
4. inter-tree effect within a stand; and
5. inter-stand effects within a forest complex.

I was particularly interested in the differences in egg mass size
between the balsam fir and white spruce tree species. Researchers have
consistently avoided studying spruce budworm population dynamics on host
species other than balsam fir. At this time, it is not known what effects this
oversight has had on the accuracy or precision of sampling predictions. Since
white spruce is the preferred oviposition host (Jaynes and Speers I949, Wilson
I963, Kemp I98I), it is imperative that we investigate this tree species as an
alternate and/or complement to the one traditionally sampled tree species.
Kemp (I978, I98I) and Kemp and Simmons (I978, I979) seem to have taken
the initiative to quantify the effects of non-host and alternative host species
on spruce budworm populations.

For all egg masses that had been collected in this study, I analyzed:

l. inter-species (tree) effects within a stand and

2. inter-species (tree) effects within a forest complex.

ll. FIELD METHODS

The study was conducted in the summer of I980 in Iron and Gogebic
Counties of Michigan's Upper Peninsula. My objective was to select stands
that had at least four balsam fir and four white spruce trees, 30 to 60 feet in

height, with very low defoliation levels (no top kill and little or no new

 

Figure I. Severely defoliated pockets of spruce and fir trees, photographed
from the Tepee Lake fire tower, eight miles south of Kenton,
Michigan.

10

 

Figure 2. Severely defoliated balsam fir tree photographed in the Ottawa
National Forest in I980.

11

foliage consumed by larvae). However, the severity of the spruce budworm

epidemic in the lake states limited my selection of stands to those which

were isolated from other spruce-fir stands md were protected by hardwood
cover. The pockets of dead and dying spruce and fir trees (Figure I) and the
dying balsam fir tree (Figure 2) illustrate the typical severe damage that
exposed host trees were experiencing in the Ottawa National Forest in I980.

After considerable time and effort, I located five stands that satisfied
the study criteria. (Appendix A presents the locations of the four study sites
within the Ottawa National Forest and the location of each study site within
the respective township.) Each stand was primarily composed of mature
hardwoods, with spruce and fir concentrated in one or two pockets along a
ridge or on a bog's edge. Other spruce and fir trees were scattered around
the pocket(s) and throughout the adjoining hardwoods. There was no definite
border to the stands and the largest spruce-fir pocket served as the center of
each stand. Within each stand four balsam fir and four white spruce trees
were selected for study. Stand I included an additional six balsam fir trees
for a total of ten balsam fir and four white spruce trees. Stand I could then
be utilized as a replication of the I979 study (Montgomery l98l).

Selection of individual trees from each stand was based on the following
criteria, listed in order of priority:

I. Least attraction to ovipositing female moths

- Trees that were overtopped by hardwoods
2. Lowest defoliation
- Trees that had the healthiest tops and the least percent of new

foliage missing

12

3. Closest proximity to the spruce-fir pocket
- Trees that were within the pocket were considered first, followed

by trees on the pocket's border

4. Within the height restriction
- Trees that were thirty to sixty feet tall

5. Specific crown structure
- One balsam fir and one white spruce with full crowns (these two
trees would have every branch cut and examined and their feasible
branches would be clipped in designated lengths and examined), i.e.
"every branch--full sample" trees
- One balsam fir and one white spruce with small crowns (these two
trees would have every branch cut and examined but n_9_ sub-
sampling would be conducted on their feasible branches), i.e.,
"every branch" trees.
- Two balsam fir and two white spruce trees with crowns ranging
from small to full (these four trees would have only their feasible
branches cut and examined), i.e., "feasible branch" trees.

In summary, each stand had a pocket that contained more white spruce
and balsam fir trees than the surrounding forest. I would first look for a tree
within the pocket that was overtopped by hardwoods; that was experiencing
little or no defoliation; that had a healthy single-stem top; and that was
thirty to sixty feet tall. If the pocket's supply of trees that met these
criteria was in excess of the required number then I would base my choice on
crown structure. If it was less than this number, then I would select trees

that met the other criteria and were closest to the pocket. Photographs of

13

two typical sampled trees, White Spruce 2 in Stand 2 and Balsam Fir I in
Stand 3, are presented in Figures 3 and 4, respectively.

Data collected from each sampled tree Included height, diameter at
breast height, and age at stump height. (Refer to Appendix B for a tabular
presentation of the data from each sampled tree.) The ten trees that were
closest to each sampled tree and had diameters at breast height of I2
centimeters or greater were recorded by species, height, diameter at breast
height, direction from the sampled tree, and distance from the sampled tree.
(Refer to Appendix C for a tabular presentation of this data base and
Appendix D for stem maps.)

All foliage from each of the two "every branch--full sample" trees and
each of the two "every branch" trees from each stand was examined for egg
masses. This was accomplished by severing each branch at its base, clipping
the branch into small segments, and thoroughly examining the foliage pieces.
Data collected from the sampled branches consisted of height above ground,
directional quadrant, total length, foliated width, total number of egg masses
on current year's foliage (new), total number of egg masses (judged to be
from the current population) on old foliage (old), total number of egg masses
that could not be judged as definitely from the current or definitely from a
previous year's population on old foliage (questionable), and foliated area.
The apical I5 percent of the tree was classified as top and examined as a
unit. However, branches longer than 70 centimeters that grew within the top
were examined individually. (Details of the sampling procedure are

graphically presented in Appendix E.)

14

 

Figure 3. Photograph of a sampled tree—White Spruce 2, Stand 2.

15

 

Figure 4. Photograph of a sampled tree—Balsam Fir I, Stand 3.

16

Ill. REGRESSION ESTIMATES FOR THE NUMBER OF EGGS PER EGG
MASS LENGTHuBALSAM FIR AND WHITE SPRUCE

The tediousness and unreliability of directly counting the number of
eggs in spruce budworm egg masses led to the development of estimating
techniques by Miller (I957), Bean (_l96l), and Leonard et al. (I973). Their
regression estimates were based on the relationship of mass length to number
of eggs. Because of the spatial and temporal variability associated with
spruce budworm population dynamics, I developed regression estimates from

a subsample of the egg masses that had been collected for this study.

Methods

Egg masses were collected from balsam fir and white spruce trees in
Michigan, beginning July I980. They were placed in I: dram vials and stored.
The indistinctiveness of translucent egg chorions made counting of hatched
eggs difficult and unreliable. Staining eclosed masses neither reduced the
tediousness nor increased the counting accuracy. Therefore, only egg masses
that were in such good condition that individual eggs were clearly
distinguishable from each other were used in this portion of the study.

Spruce budworm moths in Michigan nearly always lay egg masses in
rows of two (Figure 5), three (Figure 6), or two with a partial third (Figure 6).
I have rarely observed four-row egg masses. Therefore six regression
estimates were made—one for each of the three egg mass types from each of
the two oviposition hosts. Six groups of approximately l00 egg masses were
randomly selected from the egg mass pool. The masses were measured with
an ocular micrometer to the nearest 0.5 millimeter, and the eggs were
counted. Egg counts and corresponding egg mass lengths were entered into a

file on Michigan State University's CDC 6500 computer.

l7

 

Figure 5. Photograph of an eclosed 2-row egg mass.

18

 

Figure 6. Photograph of an eclosed 3-row egg mass.

19

 

Figure 7. Photograph of an eclosed 2-row egg mass with a partial third row.

20

Each group of l00 egg masses was tested to assure that the assumptions

of normality and homogeneity were satisfied. The Statistical Package _f_o_r LIE

 

Social Sciences (SPSS) (Nie et al. I975) was utilized for this and all
subsequent analyses in the thesis. Appendix F presents a description of the
statistical packages and computational formulae used to assess normality and
homogeneity. Because the data in each group departed significantly from the
desired normal distribution, the form of the data was changed in order to
implement the desired parametric statistics. Morris (I955) reported that the
frequency distribution for egg masses, as well as for eggs, follows the
negative binomial pattern. Sokal and Rohlf (I969), on page 384 of their text,
suggest that "when the data are counted, as of insects on a leaf or blood cells
in a hemacytometer, we frequently find the square root transformation of
value." I had used the square root transformation on data in the I979 egg
mass study (Montgomery I98I) and again found it to be the one method that
would transform the egg counts so that they met the assumptions of
normality and homogeneity. Tables FI and F2 provide the test results,
including statistics of skewness, kurtosis, and Bartlett-Box F, for egg counts
used in the regression estimates of the number of eggs per egg mass from

balsam fir and white spruce foliage, respectively.

_Fggression Estimates

Having met the assumptions of normality and homogeneity, the six
groups of transformed egg counts were incorporated in SPSS's (Nie et al.
I975) "Subprogram Regression," pages 320-367. For each simple bivariate

regression, the dependent variable was the square root of the number of eggs

21

in each egg mass and the independent variable was the length of the
corresponding egg mass to the nearest 0.5 millimeter. The regression
estimates of the number of eggs per egg mass length for each of the three
types of egg masses on balsam fir and on white spruce are presented in Tables
I and 2, respectively. The estimates have been transformed back into the
more familiar and relevant original scale and rounded to the nearest whole
egg. Regression equations, 95 percent confidence intervals for the
coefficients, correlation coefficients (r), and coefficients of determination
(r2) are presented below the regression estimates in Tables I and 2.

Figures 8, 9 and ID present plots, comparing the white spruce and
balsam fir estimated number of eggs per egg mass length for the 2-row, 2-
row and a partial third row, and 3-row egg masses, respectively. The slight
differences in the regression estimates for white spruce and balsam fir are
probably related to the general shape of each species' needles. The flatter
balsam fir needles could conceivably allow female moths more room to
stretch fewer eggs over a longer horizontal distance. White spruce needles
are often more curved and narrow which may force the ovipositing moths to
"squeeze" more eggs together over a shorter distance. This trend becomes

more noticeable as the length of the egg mass increases.

lV. LABORATORY METHODS

Egg masses had been collected from all foliated portions of I2 balsam
fir and I0 white spruce trees. Egg masses from each branch on each tree
were separately stored in labeled vials. A total of 286 masses from balsam
fir and 907 masses from white spruce were available for analysis.

The 22 trees were initially treated as separate units. Each live crown

22

Table I. Regression estimates for the number of eggs per egg mass length
from balsam fir trees in Michigan.

 

 

No. Eggs

 

2-row with a par-
Length (mm) 2-row* tial third row** 3-row***

 

—\O\O(DCD\I\IO\O\U1MJ>J>WMNN—
0 MOMOMOMOU'IOUIOU‘IOU‘IOM
G
N
U1
N
\O

O
O
0‘
N

 

*y = l.708l + 0.49I3(x)
95% Confidence Interval for A = l.5l63 to I.9000
95% Confidence Interval for B = 0.4527 to 0.5299

r = 0.927; r2 = 0.860; n = l06

**y = 2.3676 + 0.4702(x)
95% Confidence Interval for A = 2. l5l4 to 2.5839
95% Confidence Interval for B = 0.4304 to 0.5I0l

r = 0.923; r2 = 0.85l; n = 98
***y = 2.2650 + 0.5623(x)

95% Confidence Interval for A = l.9634 to 2.5665
95% Confidence Interval for B = 0.5I06 to 0.6I40

r = 0.906; r2 = 0.820; n = l04

23

Table I. Continued.
where:

y = A + B(x) V,

y = (No. Eggs per Mass)

A = a coefficient (the slope)
B = coefficient for X

X = Mass length to the nearest 0.5 mm.

24

Table 2. Regression estimates for the number of eggs per egg mass length
from white spruce trees in Michigan.

 

 

No. Eggs

 

2-row with a par-
Length (mm) 2-row* tial third row** 3-row***

 

2.0 6 II

2.5 8 I2

3.0 9 l4 I7
3.5 II l6 l9
4.0 I3 I8 22
4.5 IS 20 24
5.0 I8 22 27
5.5 20 25 30
6.0 23 27 33
6.5 25 30 36
7.0 28 32 39
7.5 3I 35 43
8.0 34 38 46
8.5 38 4|

9.0 4| 44

9.5 45

 

*y = I.386I + 0.5602(x)
95% Confidence Interval for A = I. I429 to I.6294
95% Confidence Interval for B = 0.5l l9 to 0.6085

r = 0.897; r2 = 0.804; n = l30
**y = 2.3084 + 0.48l6(x)

95% Confidence Interval for A = 2.06I3 to 2.5554
95% Confidence Interval for B = 0.4354 to 0.5279

r = 0.898; r2 = 0.807; n = :04
***y = 2.4702 + 0.542400

95% Confidence Interval for A = 2.0885 to 2.85I9
95% Confidence Interval for B = 0.4749 to 0.6l00

r = 0.847; r2 = 0.717; n =102

25

Table 2. Continued.

y = A + B(x) V2

y =(No. Eggs per Mass)

A = a coefficient (the slope)
B = coefficient for X

X = Mass length to the nearest 0.5mm.

26

Figure 8. Plot of the estimated number of eggs per egg mass
length, comparing 2-row egg masses from white spruce
and balsam fir.

27

I I I I [ITI T] I I T T I Til I T] TOT I F l

 

_ d d d 4 _ ‘14 q _ ‘14 d

o.mv

o.oe

 

o.mm

_
9.0m

 

5.35—3J34_.54_~_454d5_5.
o.m~ o.o~ o.m~ 0.0“

mwaE ..ma mama .6 59:3... cmuoEzmm

 

o.m

4.0 6.0 8.0 10.0
Length in millimeters

2.0

0.0

28

Figure 9. Plot of the estimated number of eggs per egg mass
length, comparing 2-row egg masses with a partial third
row from white spruce and balsam fir.

29

 

 

A d d d
0.0m o.m¢

 

 

.44344535 _saaAaaaad._5._a.maadda
o.oe o.mm o.om o.m~ o.o~ o.w_
39.: Lou ammo so ..onEac cBoEzmm

 

I I T IITI I I I II I I I I I I I I I I T I I T I

2.0

 

o.o~

6.0 8.0 10.0 12.0

Length in millimeters

4.0

30

Figure 10. Plot of the estimated number of eggs per egg mass
length, comparing 3-row egg masses from white spruce and
balsam fir.

31

 

IFIIITTTIIIITIIITTIITTTII

 

-444;4_43.fi_3;5__adds4a#4454ds.q._4454
o.om o.oe o.om o.om o.o¢ o.om o.o~ o.o~
39: Lou mmmm Co cmnEzc UmuoEsmm

 

0.0

8.0 10.0 12 .0
Length in millimeters

6.0

4-0

2.0

32

was divided into three strata and a top (Figure II). Egg masses from

branches that grew in a particular stratum were grouped into Stratum l,

Stratum 2, or Stratum 3.

Each branch had been recorded as residing in one of four quadrants--
North, South, East or West (Figure l2). Egg masses from branches residing in
a particular quadrant were reclassified into a corresponding quadrant within
the predesignated stratum group. In this manner, each tree would have l2
quadrant/stratum divisions and a top, for a total of I3 cells.

Four branch numbers were randomly selected from each of the l2
quadrant/stratum divisions of each tree. The egg masses from the
corresponding branch were obtained and arranged in a row. Parasitized and
mutilated egg masses were disregarded. One egg mass was randomly chosen
from each branch. The egg mass was measured with an ocular micrometer
and the number of eggs in the mass was estimated with the appropriate
regression table (I or 2). Four egg masses were also randomly selected for
measurement from each top. Frequently, cells had less than four measurable
egg masses. In those cases I would measure as many masses (0, I, 2 or 3)
from each cell that I could. The data were tabulated and the number of eggs
from a maximum of 52 egg masses per tree was used in the analysis of intra-
and inter-tree effects. (Refer to Appendix G for a list of the cells and their
corresponding egg counts.)

The subsequent tests of significance (one-way anova) in Sections I-l2
were performed on the transformed data, but estimates of means are given in

the more familiar and relevant untransformed scale.

33

   
   

__ __ ._E°E__ __ __.___ Highest
Branch
Stratum 3
Stratum 2
Stratum 1
_____ L t
”I; \\\ 3:23:11

Figure 11. Diagram of a sampled tree--side View. Division of live
crown into three strata and a top.

\\ North

West East

 

/’ South

Figure 12. Diagram of a sampled tree-top view. Division of live
crown into four quadrants.

34

V. RESULTS

Nine analyses were conducted to test for trends in egg mass size

distribution within the sampled trees, between the sampled trees of each
stand, and between the five stands. These analyses were conducted
separately for egg masses from each tree species (balsam fir in Sections l-5
and white spruce in Sections II-l2). Six analyses were made to test for
trends in egg mass size distribution between the two tree species within each
stand and between the tree species over all five stands (Sections l I- l2).

Prior to each analysis, the assumptions of normality and homogeneity
were tested with the appropriate data group. These tests were made with
SPSS's (Nie et al. I975) "Subprogram Condescriptive" (normality) and
"Subprogram T-TEST" (homogeneity). These computer programs were
incorporated earlier in the study and are described in Appendix F. The
normality and homogeneity test results for each analysis are presented in
separate tables in Appendix F and will be subsequently referred to in the
corresponding sections.

"Subprogram Oneway," SFiS, (Nie et al. I975), pages 422-433, was used
to compute the mean numbers of eggs per mass, standard deviations, standard
errors, and confidence intervals for the means of the specific groups of egg
masses under investigation in Sections I-l2. These estimated values are
reported in their more relevant and familiar untransformed scale.

One-way analysis of variance was also conducted with "Subprogram
Oneway," fl, (Nie et al. I975). The statistical package tested for
significant differences between the groups of transformed egg counts in each

section. In four cases the square root transformation did not yield

35

distributions that satisfied the assumptions of normality and/or homogeneity.

With those groups of egg counts, a nonparametric rank test was incorporated

to test for significant differences.

Section I. An Analysis of Quadrant Effects on the Size of Spruce Budworm

Egg Masses on Balsam Fir

 

This analysis was designed to answer the question, "Do spruce budworm
populations tend to lay significantly larger or smaller size egg masses on (a)
particular side(s) of balsam fir trees?" The estimated numbers of eggs per
mass from the data pool were transformed into square roots and the
assumptions of normality and homogeneity were satisfied (Table F3) for each
group of egg counts from the four quadrants-North, East, South and West--
over all l2 sampled balsam fir trees.

"Subprogram Oneway," §P_S§, (Nie et al. I975), pages 422-433,
calculates means, standard deviations, standard errors, and confidence
intervals for the means. These results are presented in Table 3.

One-way analysis of variance was used to test if observed differences
of egg mass size between the four designated quadrants were due to chance
or whether they were indicative of actual differences between the four
population means. The significance test was performed on the transformed
data, again with "Subprogram Oneway," _S_P_S§, (Nie et al. I975). The results
strongly indicate that the mean number of eggs per egg mass does not
significantly differ from one side of a balsam fir tree to another (p > 0.69)

(Table 4).

36

Table 3. The mean number of eggs per egg mass (transformed back into
original form) from the north, east, south, and west quadrants of
balsam fir trees.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Quadrant Size Eggs/ Mass Deviation Error f orlnttlf ; \ICIL an
North 49 22.6I 8.54 l.22 20. I6 to 25.06
East 60 25. I7 l0.34 I.33 22.50 to 27.84
South 59 24.67 ll.36 I.48 2| .7l to 27.63
West 60 23.85 I0. l0 I.30 2l.24 to 26.46
Total 228 24. I4

 

Table 4. Anova table for data in Table 3—Analysis of differences in the
mean number of eggs per egg mass between four quadrants within
balsam fir trees.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares * Squares* Rat io* Probability *
Between

Ouadrants 3 I.6l I 0.537 0.483 0.695 (ns)
Within

Quadrant 224 249.205 I . I l2

Total 227 250.8I6

 

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 66.399, mean squares (within groups) = I04.042, F ratio = 0.638, and F
probability = 0.59l.

37

Section 2. An Analysis of Stratum Effects on the Size of Spruce Budworm

Egg Masses on Balsam Fir

 

This analysis was designed to answer the question, "Do spruce budworm
populations tend to lay significantly larger or smaller size egg masses at (a)
particular height(s) within balsam fir trees?" The assumptions of normality and
homogeneity were met with the data groups transformed into square roots
(Table F4). Means were computed (Table 5) and egg mass size between 3
height divisions within the I2 sampled balsam fir trees was tested to determine
if observed differences were attributable to chance or whether they indicated
actual differences between the three means. The result of the one-way analysis
of variance test (transformed egg counts by strata) indicates that the mean
number of eggs per egg mass does not significantly differ from one height
division to another (p > 0.40) (Table 6) within balsam fir trees. However, there
may be biological significance in the larger egg mass size found on the lower

stratum.

38

Table 5. The mean number of eggs per egg mass (transformed back into
original form) from the lower, middle, and upper strata of balsam fir
trees.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Stratum . . . Interval

Size Eggs/Mass Devnation Error for the Mean
Lower 2| 26.74 I I.65 2.54 2 I.43 to 32.04
Middle 9| 24.52 I0.62 l. l l 22.3I to 26.73
Upper I I6 23.38 9.52 0.88 2|.63 to 25.l3
Total 228 24. I4

 

Table 6. Anova table for data in Table 5-Analysis of differences in the
mean number of eggs per egg mass between three strata within
balsam fir trees.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Between Strata 2 2.023 |.0I I 0.9l5. 0.402 (ns)
Within Strata 225 248.794 I. I06

Total 227 250. 8 I 6

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = l I0.886, mean squares (within groups) = I 03.479, F ratio = l.072, and
F probability -.- 0.344.

39

Section 3. An Analysis of the Effects of l3 Cells (Predeiignated
Compartments Within Balsam Fir Trees) on the Size of Spruce

Budworm Egg Masses

 

This analysis incorporated egg counts from the tops of balsam fir trees
and tested for effects between all cells within I2 sampled balsam fir trees
(cells were described on page 28). It was designed to answer the question, "Do
spruce budworm populations tend to lay significantly larger or smaller sized egg
masses at one or more of the l3 cells within balsam fir trees’?"

The assumptions of normality and homogeneity were satisfied with the
egg counts from the cell groups transformed into square roots (Table F5). Mean
number of eggs per egg mass were calculated for each cell over the I2 sampled
balsam fir trees (Table 7). One-way analysis of variance was used to test for
significance in observed differences in egg mass size between the I3 cells. The
results clearly indicate that the mean number of eggs per egg mass does not
significantly differ from one location to another within the sampled balsam fir

trees (p > 0.84) (Table 8).

40

Table 7. The mean number of eggs per egg mass (transformed back into
original form) from thirteen cells within balsam fir trees.

 

 

95% Confidence

 

Cell Sample Mean No. Standard Standard In terval
Size Eggs/Mass Devnation Error f or the Mean

Lower North 4 22.25 4.72 2.36 I4.74 to 29.76
Lower East 7 27.86 l2.05 4.55 l6.72 to 39.00
Lower South 3 34. I7 I3.99 8.07 -0.57 to 68.9l
Lower West 7 25.00 I3.60 5. I4 I2.42 to 37.58
Middle North l8 22.56 8.7I 2.05 I8.23 to 26.88
Middle East 2| 26.43 9.74 2. l3 22.00 to 30.86
Middle South 26 25.08 I2.35 2.42 20.09 to 30.07
Middle West 26 23.77 I0.92 2. l4 I9.36 to 28. l8
Upper North 27 22.70 9.08 l.75 l9.I l to 26.29
Upper East 32 23.75 I0.45 I.85 I9.98 to 27.52
Upper South 30 23.37 l0.08 I.84 I9.60 to 27.l3
Upper West 27 23.63 8.60 l .66 20.23 to 27.03
Top 22 2| .36 l l .02 2.35 I6.48 to 26.25
Total 250 23.90

 

Table 8. Anova table for data in Table 7-Analysis of differences in the
mean number of eggs per egg mass between thirteen cells within
balsam fir trees.

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability *
Between Cells I2 8. l83 0.682 0.60l 0.840 (ns)
Within Cells 237 268.829 I. I34

Total 249 277.0l2

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 69.987, mean Squares (within groups) = |07.050, F ratio = 0.654, and F
probability = 0.794.

41

Section 4. Analysis of Inter-Tree Effects on Spruce Budwormjgg Mass Size on
Balsam Fir Within Five Stands

Do spruce budworm populations exhibit a tendency to lay significantly
larger or smaller sized egg masses on particular balsam fir trees within a stand?
Data from the five isolated stands were analyzed to answer this question. Since
intra-tree differences in egg mass size were not significant (Sections I, 2 and
3), the egg mass measurements were pooled together for inter-tree study.

The assumptions of normality and homogeneity were met for the
transformed data from each stand (Tables F6, F7, F8, F9 and FIG). The mean
number of eggs per egg mass from each balsam fir tree within stands I, 2, 3, 4
and 5 are reported in Tables 9, II, I3, l5 and I7, respectively. One-way
analysis of variance was conducted to test if observed differences of egg mass
size on the sampled trees within a stand were due to chance or whether they
were indicative of actual differences between the population means. An anova
table was computed for each stand (Tables I0, I2, I4, I6 and I8). The mean
number of eggs per egg mass did not significantly differ from one tree to
another within Stand | (p > 0.77) (Table I0), Stand 2 (p > 0.I3) (Table I2), Stand
3 (p > 0.52) (Table I6). However, egg mass size was significantly different
between the two sampled balsam fir trees in Stand 5 (p > 0.0I) (Table I8). Egg
masses from Balsam Fir I had approximately 7 eggs per egg mass less than egg

masses from Balsam Fir 2 in absolute terms.

42

Table 9. The mean number of eggs per egg mass (transformed back into
original form) from four balsam fir trees within Stand I.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Tree . . . Interval

Size Eggs/Mass Devuatlon Error for the Mean
BFI 6 24.83 l2.46 5.09 I l.75 to 37.9l
BF2 30 25.30 l0.44 I.9l 2|.40 to 29.20
BF5 l l 22.00 9.38 2.83 l.570 to 28.30
BF6 23 23.30 l4.38 3.00 I7.09 to 29.52
Total 70 24.09

 

Table l0. Anova table for data in Table 9—Analysis of differences in the
mean number of eggs per egg mass between four balsam fir trees
within Stand I.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares * Squares* Rat io* Probability *
Between Trees 3 l.569 0.523 0.365 0.778 (ns)
Within Trees 66 94.5I l I.432

Total 69 96.080

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 36.494, mean squares (within groups) = I4|.848, F ratio = 0.257, and F
probability = 0.856.

43

Table l I. The mean number of eggs per egg mass (transformed back into
original form) from two balsam fir trees within Stand 2.

 

 

Sample Mean No. Standard Standard 95% confidence

 

 

Tree Size Eggs/ Mass Deviation Error f 0:11;; \IIIIIean
BFI 24 20.29 8.95 I.83 l6.5l to 24.07
BF2 22 24.68 l0.34 2.2l 20. ID to 29.27
Total 46 22.39

 

 

Table l2. Anova table for data in Table I I—Analysis of the difference in the
mean number of eggs per egg mass between two balsam fir trees
within Stand 2.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares * Squares * Ratio* Probability *
Between Trees l 2.3l l 2.3I I 2.278 0.I38 (ns)
Within Trees 44 44.648 I.0|5

Total 45 46.960

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) 2 22 l.226, mean squares (within groups) = 92.948, F ratio = 2.380, and F
probability = 0. I 30.

44

Table I3. The mean number of eggs per egg mass (transformed back into
original form) from two balsam fir trees within Stand 3.

 

 

Samp|e Mean No. Standard Standard 95% confidence

 

Tree Size Eggs/ Mass Deviation Error forInttlsevlaLan
BFI 2| 26.24 ||.40 2.49 2|.05 to 3|.43
BF2 l7 23.76 I I.80 2.86 l7.70 to 29.83
Total 38 25. I3

 

Table I4. Anova table for data in Table l3-Analysis of the difference in the
mean number of eggs per egg mass between two balsam fir trees
within Stand 3.

- fir - -
1 _ — — — t

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Between Trees I 0.56I 0.56l 0.4l2 0.525 (ns)
Within Trees 36 49.086 I.364

Total 37 49.648

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 57.474, mean squares (within groups) = I34. I35, F ratio = 0.428, and F
probability = 0.5I7.

45

Table IS. The mean number of eggs per egg mass (transformed back into
original form) from two balsam fir trees within Stand 4.

 

 

Sample Mean No. Standard Standard 95% confidence

 

Tree . . . Interval

Size Eggs/Mass Devration Error f or the Mean
BF I 36 23.58 9.57 I.60 20.34 to 26.82
BF2 32 25. l9 ID. | I l.79 2|.54 to 28.83
Total 68 24.34

 

Table I6. Anova table for data in Table I5-Analysis of the difference in the
mean number of eggs per egg mass between two balsam fir trees
within Stand 4.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares * Squares * Rat io* Probability *
Between Trees I 0.4l8 0.4l8 0.4I7 0.52l (ns)
Within Trees 66 66. I49 I.002

Total 67 66.567

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 43.596, mean squares (within groups) = 96.600, F ratio = 0.45l, and F
probability = 0.504.

46

Table I7. The mean number of eggs per egg mass (transformed back into
original form) from two balsam fir trees within Stand 5.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Tree . . . Interval

Size Eggs/Mass Devuation Error for the Mean
BFI 35 I9.9I 8.55 I.44 l6.98 to 22.85
BF2 27 27.02 I0.95 2. ll 22.69 to 3|.35
Total 62 23.0l

 

Table l8. Anova table for data in Table I7—Analysis of the difference in the
mean number of eggs per egg mass between two balsam fir trees
within Stand 5.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Between Trees I 8. I59 8. I59 7.905 0.007 (5)
Within Trees 60 6I .933 I.003

Total 6| 70.092

 

*These statistics were computed with the data transformed intosquare root
form. The statistics from the untransformed data were mean squares (between
groups) = 769.262, mean squares (within groups) = 93.350, F ratio = 8.24I, and F
probability = 0.006.

47

Section 5. An Analysis of Inter-Stand Effects on§pruce Budworm EgLMass
Size on Balsam Fir Within a Forest Complex

This analysis was designed to answer the question, "Do spruce budworm
populations lay significantly larger or smaller sized egg masses on balsam fir in
different stands?" The measured egg masses that had been pooled for each
balsam fir tree (Section 4) were re-pooled over each stand. The transformed
egg counts met the assumptions of normality and homogeneity (Table F II in
Appendix F).

"Subprogram Oneway," fl (Nie et al. I975) was again implemented to
calculate means (Table I9) and to test if observed differences of egg mass size
were due to chance or whether they were indicative of actual differences in the
five population means. The results of this one-way analysis of variance
indicated that the mean number of eggs per egg mass from balsam fir trees did
not significantly differ between five stands with similar levels of defoliation (p

> 0.62) (Table 20).

48

Table I9. The mean number of eggs per egg mass (transformed back into
original form) from ten balsam fir trees in five stands in the
Ottawa National Forest.

 

 

Sample Mean No. Standard Standard 95% confidence

 

Stand . . . Interval

Size Eggs/Mass Devnatlon Error for the Mean
l 36 25.22 I0.60 l.77 2|.63 to 28.8I
2 46 22.39 9. 79 I .44 I9.48 to 25.30
3 38 25.I3 ll.49 I.86 2|.35 to 28.9l
4 68 24.34 9.79 I. I9 2|.97 to 26.7I
5 62 23.00 I0.22 I.30 20.4l to 25.60
Total 250 23.90

 

Table 20. Anova table for data in Table l9-Analysis of the difference in the
mean number of eggs per egg mass between balsam fir trees in five
spruce-fir stands.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares * Squares * Rat io* Probability *
Between Stands 4 2.95I 0.738 0.660 0.62I (ns)
Within Stands 245 274.06l l. I I9

Total 249 277.0I2

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 7I.9I5, mean squares (within groups) = l05.808, F ratio = 0.680, and F
probability = 0.607.

49

Section 6. An Analysis of Quadrant Effects on the Size of Spruce Budworm
ggLMasses on White Spruce I

With this section I begin analyzing various intra- and inter-tree effects on
the size of egg masses from white spruce. This analysis was designed to answer
the question, "Do spruce budworm populations tend to lay significantly larger or
smaller size egg masses on (a) particular side(s) of white spruce trees?" The
assumptions of normality and homogeneity were satisfied for the transformed
egg counts from the four quadrants--North, East, South, and West-over all l0
sampled white spruce trees (Table FIZ). Table 2| presents the mean number of
eggs per mass per quadrant. The one-way analysis of variance test revealed
that the average number of eggs per mass is not significantly different between

the four pre-designated sides of balsam fir trees (p > 0.82) (Table 22).

50

Table 2 I. The mean number of eggs per egg mass (transformed back into
original form) from the north, east, south, and west quadrants of
white spruce trees.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Quadrant . . . Interval

Size Eggs/Mass Devuation Error for the Mean
North 7| 20.27 7.78 0.92 l8.43 to 22.| I
East ' 97 20.45 8. I6 0.83 l8.8l to 22.l0
South l00 2| .22 8.80 0.88 I9.47 to 22.96
West 79 2| .54 9.50 I .07 I9.42 to 23.67
Total 347 20.88

 

Table 22. Anova table for data in Table 2|-Analysis of the difference in the
mean number of eggs per egg mass between four quadrants within
white spruce trees.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares * Squares * Ratio* Probability *
same” 3 0 769 0 256 0 297 0 827 (ns)
Quadrants ' ° ° '

Within

Qua dran ts 343 295.996 0.863

Total 346 296 . 765

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 30. l8l, mean squares (within groups) = 73.92l, F ratio = 0.408, and F
probability = 0.747.

51

Section 7. An Analysis of Stratum Effects on the Size of Spruce Budworm
Masses on White Saruce

Do spruce budworm populations tend to lay significantly larger or smaller
size egg masses at (a) particular height(s) within white spruce trees? Egg
counts from the lower, middle and upper strata of live crowns from the ten
sampled white spruce trees were aggregated to answer this question.

The assumptions of normality and homogeneity were met with the data
transformed into square roots (Table Fl3). The mean number of eggs per mass
per stratum are reported in Table 23. Statistically, the differences in egg mass
size are not significant at the a—level of 0.05 (p > 0.05) (Table 24). However, as
in Section 2--stratum effects on egg mass size on balsam fir-—there may be
biological significance to the smaller size of egg masses found in the upper

stratum.

52

Table 23. The mecm number of eggs per egg mass (transformed back into
original form) from the lower, middle, and upper strata of white
spruce trees.

 

Sample Mean No. Standard Standard 95% Confidence

 

Stratum . . . Interval

Size Eggs/Mass Devnatlon Error for the Mean
Lower 73 2| .74 9. l6 I.07 I9.60 to 23.88
Middle I36 2| .76 8.62 0.74 20.30 to 23.23
Upper l38 I9.56 8. I0 0.69 I8.20 to 20.93
Total 347 20.88

 

Table 24. Anova table for data in Table 23—Analysis of the difference in the
mean number of eggs per egg mass between three strata within
white spruce trees.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares * Ratio * Probability *
Between Strata 2 4.952 2.476 2.9l9 0.055 (ns)
Within Strata 344 29l .8I3 0.848

Total 346 296 . 765

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = l99.739, mean Squares (within groups) = 72.808, F ratio = 2.743, and F
probability = 0.066.

53

Section 8. An Analysis of the Effects of Thirteen Cells (Predesignated

Compartments Within White Spruce Trees) on the Size of Spruce

Budworm Egg Masses

 

Egg counts from the tops of the I0 sampled white spruce trees were
incorporated with the egg counts from the other portions of the I0 sampled
trees to test for effects between all I3 cells. This analysis was designed to
answer the question, "Do spruce budworm populations tend to lay larger or
smaller sized egg masses at one or more of the l3 cells within white spruce
trees?"

Table Fl4 shows that the assumptions of normality and homogeneity were
satisfied with the groups of egg counts transformed into square roots. The
mean number of eggs per egg mass per cell are reported in Table 25. The
results of the one-way analysis of variance are presented in Table 26. The
mean number of eggs per egg mass does not significantly differ from one

location to another within the sampled white spruce trees (p > 0.20).

54

Table 25. The mean number of eggs per egg mass (transformed back into
original form) from thirteen cells within white spruce trees.

Cell Sample Mean No. Standard Standard 95% Confidence

. , , Interval
Size Eggs/Mass Deviation Error for the Mean

 

Lower North I0 22.50 I0. I0 3. I9 l5.27 to 29.73
Lower East 24 20.92 8.56 l.75 l7.30 to 24.53
Lower South 29 23.59 9.76 I .8l I9.88 to 27.30
Lower West I0 I7.60 7.29 2.3l l2.38 to 22.82
Middle North 28 20.32 6.77 l.28 l7.70 to 22.95
Middle East 37 22.32 8.53 I.40 I9.48 to 25. I7
Middle South 36 I9.94 8.09 I.35 l7.2| to 22.68
Middle West 35 24.20 l0. I2 l.7l 20.72 to 27.68
Upper North 33 I9.55 7.93 I.38 l6.73 to 22.36
Upper East 36 l8.22 7. I3 I. I9 I5.8I to 20.64
Upper South 35 20.57 8.54 I.44 I7.63 to 23.50
Upper West 34 I9.97 8.88 l.52 I6.87 to 23.07
Top 25 2| .48 l0.06 2.0l l7.33 to 25.63
Total 372 20.92

 

Table 26. Anova table for data in Table 25—Analysis of the difference in the
mean number of eggs per egg mass between thirteen cells within
white spruce trees.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Between Cells l2 l3.605 I. I34 I .3l5 0.208 (ns)
Within Cells 359 309.528 0.862

Total 37l 323 . I33

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = I 00.I98, mean Squares (within groups) = 74.3l6, F ratio = I.3482.743,
and F probability = 0.I89.

55

Section 9. Analysis of Inter-Tree Effects on Spruce Budworm EgLMass Size on
White Spruce Within Five Stands

Data from the five isolated stands were analyzed to determine if spruce
budworm populations exhibit a tendency to lay significantly larger or smaller
size egg masses on particular white spruce trees within each stand. Since intra-
tree differences in egg mass size were not statistically significant (Sections 6, 7
and 8), the egg counts were pooled together for inter-tree study.

Each data group did not depart significantly from the desired normal
distribution and each group of variances was significantly homogeneous (Tables
F I5, Fl6, FI7, F I8 and Fl9). The mean number of eggs per egg mass per white
spruce tree within Stands I, 2, 3, 4 and 5 are reported in Tables 27, 29, 3|, 33
and 35, respectively. The egg counts from each of the two white spruce trees
within each stand were tested to see if the observed differences in average egg
mass size were significant.

Five one-way analyses of variance tests were conducted and the results
are reported in Tables 28, 30, 32, 34 and 36. The mean number of eggs per egg
mass did not significantly differ between the two white spruce trees within
Stand I (p > 0.87 (Table 28), Stand 3 (p > 0.4I) (Table 32), Stand 4 (p > 0.06)
(Table 34), or Stand 5 (p > 0.45) (Table 36). Statistically, egg mass size was
significantly different between the two sampled white spruce trees in Stand 2
(pF 0.02) (Table 30). Egg masses from White Spruce I had an average of
approximately 5 eggs per mass greater than egg masses from White Spruce 2 in

absolute values.

56

Table 27. The mean number of eggs per egg mass (transformed back into
original form) from two white spruce trees within Stand l.

 

 

SClmPle Mean No. Standard Standard 95% Confidence

 

Tree . . . Interval

Size Eggs/Mass Devnatlon Error for the Mean
WSI 37 23.30 9.45 I.45 20.l5 to 26.45
W52 33 23.09 |0.29 I .79 I9.44 to 26.74
Total 70 23.20

 

Table 28. Anova table for data in Table 27-Analysis of the difference in the
mean number of eggs per egg mass between two white spruce trees
within Stand I.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Between Trees I 0.027 0.027 0.025 0.874 (ns)
Within Trees 68 7| .896 I.057

Total 69 7| .923

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mem squares (between
groups) = 0.7430, mean squares (within groups) = 97.066, F ratio = 0.008, and F
probability 2 0.930.

57

Table 29. The mean number of eggs per egg mass (transformed back into
original form) from two white spruce trees within Stand 2.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Tree . . . Interval

Size Eggs/Mass Devuatlon Error f or the Mean
WSI 46 2I.57 8.23 l.2l l9.I2to 24.0l
WS2 27 I6.89 6.96 I.34 I4.l3 to I9.64
Total 73 I9.84

 

Table 30. Anova table for data in Table 29-Analysis of the difference in the
mean number of eggs per egg mass between two white spruce trees
within Stand 2.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Between Trees I 4.937 4.937 6.808 0.0I I (5)
Within Trees 7| 5| .483 0.725

Total 72 56.420

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 372.056, mean Squares (within groups) = 60.648, F ratio = 6.I35, and F
probability = 0.0l6.

58

Table 3 l. The mean number of eggs per egg mass (transformed back into
original form) from two white spruce trees within Stand 3.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Tree . . . Interval

Size Eggs/Mass Devnation Error for the Mean
WSI 28 20.29 6.38 l.2l l7.8l to 22.76
W52 40 I8.95 5.66 0.90 l7. l4 to 20.76
Total 68 I9.50

 

Table 32. Anova table for data in Table 3l—Analysis of the difference in the
mean number of eggs per egg mass between two white spruce trees
within Stand 3.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Rat io* Probability *
Between Trees I 0.320 0.320 0.666 0.4I7 (ns)
Within Trees 66 3| .768 0.48I

Total 67 32.089

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 29.386, mean squares (within groups) = 35.600, F ratio = 0.825, and F
probability = 0.367.

59

Table 33. The mean number of eggs per egg mass (transformed back into
original form) from two white spruce trees within Stand 4.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Tree . . . Interval

Size Eggs/Mass Devnatlon Error for the Mean
WSI 48 22.27 l0.63 l.54 l9.I8 to 25.36
W52 44 I8.32 8.68 l .3l I5.68 to 20.96
Total 92 20.38

 

Table 34. Anova table for data in Table 33—Analysis of the difference in the
mean number of eggs per egg mass between two white spruce trees
within Stand 4.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares * Ratio* Probability *
Between Trees I 3.856 3.856 3.432 0.067 (ns)
Within Trees 90 IOI .096 I. I23

Total 9| I04.95l

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 358.660, mean squares (within groups) = 95.078, F ratio = 3.772, and F
probability = 0.055.

60

Table 35. The mean number of eggs per egg mass (transformed back into
original form) from two white spruce trees within Stand 5.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Tree . . . Interval

Size Eggs/Mass Devnatnon Error for the Mean
WSI 45 22.48 8.55 l.27 I9.92 to 25.05
WS2 24 20.79 7. 72 I .58 I7.53 to 24.05
Total 69 2| .90

 

Table 36. Anova table for data in Table 35—Analysis of difference in the
mean number of eggs per egg mass between two white spruce trees
within Stand 5.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Between Trees I 0.426 0.426 0.57I 0.453 (ns)
Within Trees 67 49.960 0.746

Total 68 50.385

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 44.85l, mean squares (within groups) = 68.4I6, F ratio = 0.656, and F
probability = 0.42I.

61

Section I0. An Analysis of Inter-Stand Effects on Spruce Budworm EMS
Size From White Spruce Within a Forest Complex

The egg counts from the two sampled white spruce trees within each
stand were combined for an average number of eggs per mass per stand. This
analysis was designed to answer the question, "Do spruce budworm populations
lay significantly larger or smaller size egg masses on white spruce in five
stands?"

Table F20 shows that the assumption of normality was met but the
variances were n_ot_ homogeneous (p = 0.003). However, "Subprogram Oneway,"
fl (Nie et al. I975) was again implemented to calculate the mean egg mass
size per stand (Table 37) and to test if observed differences of egg mass size
were due to chance or whether they were indicative of actual differences in the
five population means. Results reported in Table 38 indicate that the average
egg mass size did not significantly differ from stand to stand on the white
spruce trees (p > 0.07).

Because heteroscedacity may have affected the outcome of the test, a
nonparametric rank test, the Kruskal-Wallis Test, was used to reaffirm or
nullify the original test results. Conover (I980) provides an excellent
explanation and examples of this test on pages 229-237 of his text. Table 39
relates the sum of the ranks and sample sizes of the egg counts from each
stand.

The Kruskal-Wallis Test is designed to determine if the null hypothesis,
the k populations do all have identical means, should be rejected. Since there
were many equal ranks (ties), tie decision rule was based on the chi-square

distribution with 4 degrees of freedom at the a-level, 0.05. since the test

62

statistic, T = 7.394, was less than the 0.95 qumtile, x2 = 9.488, the null
hypothesis was again accepted and the mean number of eggs per mass per stand

on white spruce are not significantly different (p > 0. l0).

63

Table 37. The mean number of eggs per egg mass (transformed back into
original form) from ten white spruce trees in five stands in the
Ottawa National Forest.

 

 

95% Confidence

Interval
for the Mean

Stand Sample Mean No. Standard Standard
Size Eggs/Mass Deviation Error

 

I 70 23.20 9.78 I. I7 20.87 to 25.53
2 73 I9.84 8.06 0.94 I7.95 to 2I.72
3 68 I9. 50 5.96 0. 72 I8.06 to 20.94
4 92 20.38 9.90 I.03 I8.33 to 22.43
5 69 2| .90 8.25 0.99 I9.9I to 23.88
Total 372 20.92

 

Table 38. Anova table for data in Table 37—Analysis of the difference in the
mean number of eggs per egg mass between white spruce trees in
five stands in the Ottawa National Forest.**

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Between Stands 4 7.366 I .84l 2. I40 0.075 (ns)
Within Stands 367 3l5.768 0.860

Total 37I 323. I33

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = I69.825, mean squares (within groups) = 74.I2I, F ratio = 2.29l, and F
probability = 0.059.

**NOTE:The assumption of homogeneity of variances was {191 satisfied with
this data set. Refer to the alternative test (Kruskal-Wallis) on page
64.

64

Table 39. Results of the Kruskal-Wallis test, comparing mean egg mass sizes
on white spruce between all 5 stands.

 

 

StandIi) ni Ri
I 70 I4,789
2 I 73 I2,645
3 68 I2,042
4 92 I6,00 I
5 69 I 3,883

 

S2 i=1 n1 4

T r. _L(§ 513.44" “12)

= the test statistic

where:
2
52:..1__z R(Xt.)2-Nm—fl—I-—
N-l all 3 4
ranks
R. = sum of ranks of egg masses for stand i

i = stand number

k = total number of stands

= 5
N = total number of ranks (egg masses)

= 372
ni = number of ranks (egg masses) per stand
IZR(X..)2 : 2 (individual ranks)2

U

all
ranks

= 17,183,967.5
S2 = 11,441.969
T = 7.394(NS)

65

Section I l. Analysis of Inter-@ecies (Trees) Effects on Spruce Budworm Egg
Mass Size Within Each Stand

Five analyses were conducted to answer the question, "Do spruce
budworm populations tend to lay larger or smaller size egg masses on white
spruce or balsam fir in a stand?" The egg counts from the two trees of the
same species within each stand were aggregated for comparison of mean egg
mass size.

The data were transformed into square roots to test the assumptions of
normality and homogeneity (Tables F 2 l-F 25). In two cases, Stand 3 (Table F23)
and Stand 5 (Table 25), the variances were significantly different from the
desired homogeneous form (p = 0.000) and (p = 0.080), respectively. Each
significance test was performed with the one-way analysis of variance method;
however, data in Stands 3 and 5 were also analyzed with nonparametric
statisticsnthe Mann-Whitney Test for two independent samples.

Tables 40, 42, 44, 47 and 49 present the mean number of eggs per mass
per species per stand. Tables 4|, 43, 45 and 46, 48, 50 and SI present the
results of the tests for significant differences between mean egg mass size per
species in Stands I, 2, 3, 4 and 5, respectively. Differences in the mean
numbers of eggs per mass between the white spruce and balsam fir species was
statistically significant in Stand 3 (p < 0.0I) and Stand 4 (p < 0.0I). Mean egg
mass size was not significantly different between the two tree species in Stand
| (p > 0.35), Stand 2 (p > 0.I4) or Stand 5 (p > 0.65). However, there may be
biological significance in the larger average number of eggs per mass from
balsam fir as opposed to white spruce. This trend was evident in each stand,
with egg masses on balsam fir containing an average of LI to 5.6 more eggs per

mass per stand than the egg masses on white spruce in absolute terms.

66

Heteroskedasticity may have affected the results of tests on data in
Stands 3 and 5. Therefore, nonparametric statistics were utilized to re-analyze
the observed differences in egg mass size. Conover (I980) describes the Mann-
Whitney Test on pages 2I5-227 of his text. The Kruskal-Wallis Test, used in
Section I0, is merely an extension of the Mann-Whitney Test. Again, the
numbers of eggs per sampled egg mass were ranked in ascending order, with the
rank I assigned as the value of the smallest number of eggs per mass in the data
base. The individual ranks were summed for each species from each stand and
these sums are reported in Table 46 (Stand 3) and Table 5| (Stand 5), along with
the test statistic and computational formulae. Results of the two-tailed tests
confirmed the original result that had been computed with parametric
statistics. Balsam fir had significantly different size egg masses than white
spruce in Stand 3 (p <0.0I) and balsam fir and white spruce egg masses were not

significantly different in Stand 5 (p > 0.65).

67

Table 40. The mean number of eggs per egg mass (transformed back into
original form) from white spruce and balsam fir trees in Stand I.

Sample Mean No. Standard Standard 95% Confidence

Trees Interval

 

Size Eggs/Mass Deviation Error for the Mean
BFI & BF2 36 25.22 l0.60 l.77 2|.63 to 28.8l
WSI 8: W52 70 23.20 9.78 I. I7 20.87 to 25.53
Total I06 23.89

 

Table 4|. Anova table for data in Table 39—Analysis of the difference in the
mean number of eggs per egg mass between white spruce trees and
balsam fir trees in Stand l.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares * Squares* Ratio* Probability *
Within

Tree Species I 0.944 0.944 0.87I 0.353 (ns)
Between

Tree Species I l2.7l7 I.084

Total l05 l I3.662

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 97.2 l9, mean squares (within groups) = I0|.32 I, F ratio = 0.960, and F
probability = 0.330.

6.8

Table 42. The mean number of eggs per egg mass (transformed back into
original form) from white spruce and balsam fir trees in Stand 2.

 

 

95% Confidence

Sample Mean No. Standard Standard
Interval

Trees

 

Size Eggs/Mass Deviation Error for the Mean
BFI & BF2 46 22.39 9.79 I.44 I9.48 to 25.30
WSI & W52 73 l8.84 8.06 0.94 l7.95 to 2|.72
Total I I9 20.82

 

Table 43. Anova table for data in Table 4I-Analysis of the difference in the
mean number of eggs per egg mass between white spruce trees and
balsam fir trees in Stand 2.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares * Squares * Rat io* Probability *
Between

Tree Species I I.860 I .860 2. I05 0.I50 (ns)
“”1”" I I7 I03.379 0.884

Tree Species

Total I I8 I05.239

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = I84.3|0, mean Squares (within groups) = 76.829, F ratio = 2.399, and F
probability = 0.I24.

69

Table 44. The mean number of eggs per egg mass (transformed back into
original form) from white spruce and balsam fir trees in Stand 3.

SClmple Mean No. Standard Standard 95% Confidence

 

Trees . . . Interval

SIze Eggs/Mass DeVIatIon Error for the Mean
BFI &BF2 38 25.I3 ll.49 I.86 2|.35 to 28.9l
WSI & W52 68 I9.50 5.96 0.72 l8.06 to 20.94
Total l06 2| .52

 

Table 45. Anova table for data in Table 43—Analysis of the difference in the
mean number of eggs per egg mass between white spruce trees and
balsam fir trees in Stand 3.**

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Beiwee" I 6 566 6 566 8 354 0 005 (8)
Tree Species ° ' ° °

“"1”" l04 8I .737 0. 786

Tree Species

Total I 05 88 . 302

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = I84.3|0, mean squares (within groups) = 76.829, F ratio = 2.399, and F
probability = 0.I24.

**NOTE: The assumption of homogeneity of variances was not satisfied with
this data set. Refer to the alternative test (Mann-Whitne_)'Ton page 70.

70

Table 46. Results of the Mann-Whitney test, comparing mean egg mass size
on white spruce with mean egg mass size on balsam fir in Stand 3.

 

 

 

 

n
Tree Species Sample Size 22 R

i=I

Balsam fir 38 2380

White spruce 68 329I
N+1

T - n

Tl = '77-

 

 

f um I; R? _ nmIN+1I2
N(N-1) i=1 ‘ 4(N—1)
= the test statistic when there are many tied ranks

where:

n
T = S R (Balsam Fir)
i=l
= sum of the ranks assigned to the balsam fir egg counts
2 2380
n
2 = the sum of the squares of all N of the ranks
'2' = 402,093.5
n = sample size from balsam fir
= sample size from white spruce
N = n + m
= I06

 

T = 2.292(5)

71

Table 47. The mean number of eggs per egg mass (transformed back into
original form) from white spruce and balsam fir trees in Stand 4.

 

 

Sample Mean No. Standard Standard 95% Confidence

 

Trees . . . Interval

SIze Eggs/ Mass DeVIatIon Error for the Mean
BFI & BF2 68 24.34 9.79 I. I9 2|.97 to 26.7l
WSI 8: W52 92 20.38 9.90 I.03 I8.33 to 22.43
Total I60 22.06

 

Table 48. Anova table for data in Table 45-Analysis of the difference in the
mean number of eggs per egg mass between white spruce trees and
balsam fir trees in Stand 4.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability *
Between

Tree Species I 7.807 7.807 7. I92 0.008 (5)
Within

Tree Species I58 I7I.5l8 I.086

Total I 59 I 79 . 325

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 6l2.470, mean squares (within groups) = 97.056, F ratio = 6.3I0, and F
probability = 0.0l3.

72

Table 49. The mean number of eggs per egg mass (transformed back into
original form) from white spruce and balsam fir trees in Stand 5.

_f
_—

SClmple Mean No. Standard Standard 95% Confidence

 

Trees . . . Interval

SIze Eggs/Mass DeVIatIon Error for the Mean
BFI & BF2 62 23.008 I0.2I9 l.298 20.4I3 to 25.603
WSI & W52 69 2| .896 8.250 0.993 I9.9l4 to 23.878
Total I 3 I 22 . 422

 

Table 50. Anova table for data in Table 49—Analysis of the difference in the
mean number of eggs per egg mass between white spruce trees and
balsam fir trees in Stand 5.**

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Ratio* Probability*
Between

Tree Species I 0. I92 0. I92 0.206 0.65l (ns)
Wilh‘" I29 I20.477 0.934

Tree Species

Total | 30 I 20 . 669

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = 40.4I I, mean squares (within groups) = 85.263, F ratio = 0.474, and F
probability = 0.492.

**NOTE: The assumption of homogeneity of variances was go_t satisfied with
this data set. Differences were tested for significance with the
Mann-Whitney test on page 73.

73

Table 5 I. Results of the Mann-Whitney test, comparing mean egg mass size on
white spruce with mean egg mass size on balsam fir in Stand 5.

 

 

 

 

 

 

 

 

 

n
Tree Species Sample Size 2 R
i=l
Balsam Fir 62 4202.5
White Spruce 69 4443.5
T _ n N 42- 1
TI 3
N
nm 2 R2 mn(N+l) 2
N(N—l) i=1 i 4(N-1)
= the test statistic when there are many tied ranks
where:
n
T = ER (Balsam Fir)
i=I
= the sum of the ranks assigned to the balsam fir egg counts
= 4,202.5
g R 2 —=’ the sum of the squares of all N of the ranks
._ i
1‘1 = 757,309.5
n = sample size from balsam fir
2 sample size from white spruce
N = n + m
= I3I
Tl = 0.5I0 (NS)

74

Section l2. Analysis of Inter-Species (Trees) Effects on Spruce Budworm Egg
Mass Size Within a Forest Complex

The final analysis was designed to answer the question, "Do spruce
budworm populations lay larger or smaller size egg masses on balsam fir or
white spruce trees?" All egg counts were aggregated into their respective host
species group to test for significant difference between the two averages.
Table 52 presents the average number of eggs per mass on the I0 sampled
balsam fir trees (23.90) and the average number of eggs per mass on the I0
sampled white spruce trees (20.92) from all five stands.

The assumption of normality was confirmed, but variances were
nonhomogeneous. Both parametric and nonparametric statistics were
performed. Results of the one-way analysis of variance test strongly indicate
that the mean number of eggs per mass is significantly different between egg
masses on white spruce and egg masses on balsam fir (Table 53) (p < 0.000l).

Results of the Mann-Whitney Test confirm the results of the parametric test

(Table 54) (p < 0.00l).

75

Table 52. The mean number of eggs per egg mass (transformed back into
original form) from ten white spruce and ten balsam fir trees in five

stands.

 

 

Sample Mean NO. Standard Standard 95% Confidence

 

Species Size Eggs/ Mass Deviation Error forlntt: er 1:; an

Balsam Fir 250 23.90 I0.26 0.65 22.62 to 25.I8
White Spruce 372 20.92 8.67 0.45 20.04 to 2|.8l
Total 622 22. l2

 

Table 53. Anova table for data in Table 52—Analysis of the difference in the
mean number of eggs per egg mass between ten white spruce and
ten balsam fir trees in five stands.

 

 

 

Source of Degrees of Sum of Mean F F
Variation Freedom Squares* Squares* Rat io* Probability *
Between Tree
Species I I 3. 059 l3 . 059 I 3 . 49 I 0.000(5)
y'lhln Tree 620 600. I46 0.968

peCIes
Total 62l 6 I 3 . 205

 

*These statistics were computed with the data transformed into square root
form. The statistics from the untransformed data were mean squares (between
groups) = l322.263, mean squares (within groups) = 87.246, F ratio = I5.I56, and
F probability = 0.000.

76

Table 54. Results of the Mann-Whitney test, comparing mean egg mass size on

white spruce with mean egg mass size on balsam fir over all five
stands.

 

 

 

n
Tree Species Sample Size . £|R
l=
Balsam Fir 250 85,380
White Spruce 372 |08,373

 

 

 

T _ n N :- 1
TI 2
nm 23 R2 _ nm N-I-l 2
N(N-1) i=1 i 4(N-l)
= the test statistic when there are many tied ranks
where:
n
T = ZR (Balsam Fir)
I=I
= the sum of the ranks assigned to the balsam fir egg counts
= 85,380
n
2; 2 '= the sum of the squares of all N of the ranks
i=1 i .
= 80,350,396
n = sample size from balsam fir
m = sample size from white spruce
N = n + m
= 622

VI. DISCUSSION

This thesis presents an analysis of the distribution of spruce budworm egg
mass sizes within trees, between trees, between tree species, and between
stands. A number of important trends were evident from the study results.
These findings should improve the accuracy and precision of egg mass surveys,
particularly when the sampled populations are at very low densities.

As I expressed in my first thesis (Montgomery I98I), spruce budworm
investigators must refrain from incontestably applying quantitative results from
one region to another. Regional and local diversity in stand structure,
composition, and management activities span the boreal forest habitat of the
spruce budworm. These factors and the year-to-year variability associated with
the intricate effects of parasites, predators, rainfall, temperature, humidity, and
available foliage nutrients and biomass play important and interrelated roles in
determining population vigor, dynamics, and relevant egg mass size.

Therefore, I feel that it is important to restate the conditions of the five
spruce fir stands that I studied and the conditions of the surrounding forest in
general. The project was conducted in the summer of I980 in the Ottawa
National Forest, located in the western half of Michigan's Upper Penninsula. In
the majority of spruce fir stands, the spruce budworms were in their fifth to
tenth year at outbreak densities, as shown by severe defoliation, an abundance of
top-kill, and tree mortality. Management in the Upper Peninsula has generally
refrained from chemical control in spruce-fir stands and have rarely made a
conscientious effort to implement silvicultural control or prevention measures.
This stems from the spotty demand for spruce or fir in the market; however,

higher demand will apparently come into effect in the near future.

77

78

The maximum size of spruce-fir stands in the Ottawa National Forest is
about 80 hectares, with most in the range of I6 hectares or less. They grow in
isolated pockets, are often intermixed with hardwoods, and generally consist of a
variety of soil types, influenced by enormous variability in past ecological
perturbations within each site (particularly glacial deposits and fires).

The five stands that I sampled for this study were all experiencing low
spruce budworm defoliation and population levels. Defoliation of current foliage
ranged from 0 to 30 percent, using Fettes (I950) method of estimating
defoliation, as a tree average. Population levels were as low as seven egg
masses on all foliated portions of a sampled tree. Sampled stands were high in
hardwood content, with spruce and fir comprising only I0 to 40 percent of the
overstory growth in each stand. Most sampled trees grew in the vicinity of a
high density spruce-fir pocket; however, nearly all were overtopped by hardwood
cover.

The objective of this study was to determine if egg mass size was related
to particular sites within a host tree, within a stand, or within a forest complex.
From my analyses, I wmted to determine if estimates of population size,
computed solely from egg mass densities, would be significantly distorted by
neglecting to account for differences in the numbers of eggs per masses.
Another goal of the project was to analyze differences in egg mass size between
the preferred ovipositional host, white spruce, and the preferred food source,
balsam fir.

Twelve balsam fir and ten white spruce trees had every branch severed,
measured, and thoroughly examined for spruce budworm egg masses. The

position of each egg mass within a sampled tree and a stand was recorded. The

79

number of eggs were estimated by measuring mass length and using regresion
equations derived from a subsample of this study's egg mass pool. "Subprogram
Oneway" in SPSS: Statistical Packages for the Social Sciences (Nie et al. I 975)

 

was used to calculate the mean number of eggs per egg mass for the specific
group of egg masses under investigation. The same subprogram was used to test
for trends in egg mass size distribution, incorporating one-way analysis of
variance to test for significant difference(s) between mean egg mass sizes within

trees, between trees, between tree species, and between stands.

Regression estimates

 

Separate regression estimates were made for 2-row, 2-row with a partial
third row, and 3-row egg masses from balsam fir foliage and from white spruce
foliage, giving the number of whole eggs per egg mass length. Differences in the
regression equations between the two hosts were small and possibly related to
structural differences between the two species needles. Ovipositing moths may
compress their eggs more tightly into masses, over shorter lengths on white
spruce, than moths ovipositing eggs on flatter, wider balsam fir foliage.

Large differences are evident between my regression equations (|98l) and
Miller's (I957), Bean's (I96I), and Leonard's et al. (I973). Although no specific
statistical tests were performed that compare the four groups of equations, a
simple visual comparison reveals significant biological differences between the
numbers of eggs per mass length. It would be unacceptable for me to
hypothesize on the specific reasons for these differences, without concrete data
at my disposal. Suffice it to say that estimated numbers of eggs per mass length

for each kind of mass will vary from location-to-Iocation md from year-to-year.

80

These differences may be due to a multitude of factors, including nutritional
composition of the larvae diet, weather conditions during oviposition, differences
in sampling methods, stand structures, moth dispersal, or a combination of these

factors and more.

Infra-tree distribution of egg mass size

 

Statistically, no significant differences were found in the mean number of
eggs per mass between any loci within sampled trees--balsam fir or white spruce.
Quadrant differences were minimal. However, there may be biological
significance to the relatively large differences in egg mass sizes between the
three stata within sampled balsam fir and white spruce trees. Mean number of
eggs per mass in the lower, middle, and upper strata decreased from 26.74 to
24.52 to 23.38 in balsam fir and varied from 2I.74 to 2|.76 to I9.56 in white
spruce, respectively. This anomaly may have resulted from the small sample
size of eggs in the lower stratum. However, this trend might also indicate that
female moths in stands with very low populations fly up toward the preferred
oviposition sites to lay smaller egg clusters, after they have laid their first two
largest egg clusters at their eclosion and mating sites. This vertical migration
may be stimulated by the nutritous flower buds on tops of larger trees that would
eventually provide their offspring with a better initial food source. Likewise,
the maths may have been drawn from the lower branches in an attempt to
disperse from the stand. Or gravid female moths may have immigrated to the
stand and oviposited their smaller masses at or near the "attractive" spire tops
of host trees.

This trend is different from the distribution of egg mass size between

strata that I reported in my first thesis (|98l). In that study a larger mean egg

81

mass size (though not significant) was found in the middle stratum as opposed to
the nearly equal sizes found in the upper and lower strata. The first thesis (I98I)
investigated spruce budworm populations with much higher densities and trees
with heavier defoliation levels. Oviposition in those stands was possibly
governed more by the quantity of foliage than by the quality.

In this study, the nonsignificant, yet highly visible differences in the mean
number of eggs per mass between strata does not appear important enough to
alter sampling methods on individual trees. In the first place, the differences
would "balance" out in surveys that sample feasible branches from the mid-
crown, since mass size there is intermediate between the sizes in upper and
lower strata. Secondly, the differences are not excessive (approximately three
eggs per mass and two eggs per mass mean differences between the largest and
smallest sizes in balsam fir and white spruce strata, respectively). Thirdly, the

difference could be related to the small sample size from the lower stratum.

Inter-tree distribution of egg mass size

Differences in mean egg mass size between trees of the same species
within the same stand were for the most part variable and nonconclusive. In
general, mean egg mass size was not significantly different. Statistically, tree
pairs in eight of ten analyses had mean egg mass sizes that did not significantly
differ at the 0.05 -level. Two groups of paired trees, the balsam fir (BF l and
BF2) in Stand 5 and the white spruce (WSI and WS2) in Stand 2 had statistical and
perhaps biological significance to the relatively large differences in egg mass
size. These differences might be related to the trees' phenological and/or

physiological characteristics.

82

BF l and BF 2 in Stand 5 had averages of I9.9l and 27.02 eggs per mass,

respectively. BFI was 33 years old and |0.l meters tall, while BF2 was 43 years
old and |2.0 meters tall. WSI and W52 in Stand 4 had averages of 2|.57 and

I6.89 eggs per egg mass, respectively. WSI was 5| years old and I3.4 meters
tall, while W52 was 32 years old and I l.7 meters tall.

Kimmins (I97I) found that higher concentrations of amino acids and a
higher content of amino-nitrogen were in the foliage of flowering balsam fir as
opposed to the foliage of non-flowering balsam fir. This effect was found to be
small in white spruce foliage. However, it is not known "whether the changes in
protein and amino acid metabolism result from flowering and fruiting, which in
turn are induced by climatic factors, or whether the changes or directly induced
by climatic stress and are themselves important determinants of flowering and
fruiting" (Kimmins I97 I).

This could serve as the basis for an explanation of the significantly larger
egg masses found on the much taller and older hast trees, BF 2 in Stand 5 and
WSI in Stand 2. If the relative abundance of available nitrogen was substantially
greater in the foliage of the larger trees and if other factors had constant
effects between the tree pairs, then larvae, feeding on the taller trees' foliage,
would perhaps be more vigorous and would perhaps produce ovipositing adults
that had proportionally higher fecundity rates and would be more apt to lay
larger egg masses. I emphasize that it is certainly beyond the scope of this
thesis to analyze the plant stress-plant response-herbivore response complex.
Mattson (I980) reviews these relationships for general herbivores and plants in
detail and other investigators are specifically studying spruce budworm

population dynamics in relation to nutritional release.

83

There are many other factors that could also be responsible for larger or
smaller egg masses being distributed on particular trees throughout a stand.
Factors, such as larvae and/or moth dispersal within or between stands,
microhabitat variation between trees, structural differences of trees and their
foliage, variability of microclimate during oviposition, each could individually or
in combination produce significant effects on egg mass sizes.

Whatever the cause(s), large differences in egg mass sizes can exist
between trees within the same stand. These differences can have significant
effects on population estimates derived from egg mass sampling surveys for a
stand. The more diverse stands should have a larger number of trees sampled
that are representative of the variety of host trees within these stands. This
would improve the accuracy of estimated values of the number of eggs per egg

mass per stand, particularly in the diverse forests of Michigan's Upper Peninsula.

Inter-stand differences in egg mass size

The mean number of eggs per egg mass was not significantly different
between the five stands for each tree species. This relationship was expected
since defoliation levels in all five stands were very similar. Slight differences in
egg mass sizes between stands were possibly related to such interconnected
factors as stand age, tree height, species composition, foliar nutrient content,
soil type, topography, past ecological perturbations, larval and/or moth dispersal,
and the local effects of population size and vigor, parasites, predators, diseases,
rainfall, humidity, and temperature. However, any of these factors, singularly or

in combination, could have had much more dramatic effects on egg mass size in

84

any given stand, region or year. This can be best exemplified by comparing this
study's results with the results in my first thesis “98”. Egg masses from
sampled balsam fir trees had mean numbers of eggs of l7.l2 in two moderate-
severely defoliated stands (l98l), l8.72 in two light-moderately defoliated stands
(I98l) and 23.90 in the five very lightly-defoliated stands from this project. The
differences are obvious and the potential effects on the accuracy and precision
of sampling programs should be observed. To recapitulate the suggestion that
was made in my first thesis (I98I)--as a rule, the mean number of eggs per egg
mass should be calculated on a per stand basis to assure accuracy and precision

in egg mass sampling surveys.

Inter-species distribution of egg mass size

Egg masses on sampled balsam fir in each stand contained an average of
H to 5.6 more eggs per mass than egg masses on sampled white spruce.
However, in only two stands were the differences in the averages statistically
significant. Over all 20 sampled trees in the five stands, the ID balsam fir trees
had an average of 23.90 eggs per mass and the I0 white spruce trees had an
average of 20.92 eggs per mass.

This trend is interesting since white spruce has been long noted as the
preferred oviposition host for the spruce budworm (Jaynes and Speers I949,
Wilson I963, Kemp |98I). It, therefore, might be expected that white spruce
would also be the site of the largest egg masses. This is not the case for two
possible reasons.

Structurally, white spruce foliage is generally not as conducive to receiving

long or wide egg masses. White spruce needles are often curled and close

85

together in a 3-dimensional whorl around each twig. Balsam fir needles are

generally flatter and more 2-dimensionally oriented, particularly in our sampled

stand that were heavily shaded. Female moths would have much less difficulty
extending their ovipositors over a longer, wider balsam fir needle surface, than
trying to squeeze their ovipositors between needles that are curled and narrow.

Kimmins (I97l) found that balsam fir foliage had significantly greater
levels of amino acids than white spruce foliage. If moths emerge, mate and
oviposit most of their egg complement on the same trees that they had fed upon
as larvae, then the protein content of the host species would be crucial to female
development and fecundity.

Sampling programs should be cognizant of the consistently smaller egg
mass size on white spruce foliage as opposed to balsam fir foliage. If white
spruce is a relatively large component in sampled stands, then this tree species
should also be incorporated in the survey and their average egg mass size should

be estimated on a per stand basis.

LIST OF REFERENCES

LIST OF REFERENCES

Baskerville, G.L. 1975. Spruce budworm: super silviculturist. For.
Chron. 51(4): 138-140.

Bean, J.L. 1961. A method for estimating the number of spruce bud—
worm eggs per egg mass. J. Econ. Entomol. 54(5): 1064.

Blais, J.R. 1952. The relationship of the spruce budworm (Chorist—
oneura fumiferana (Clem.)) to the flowering condition of balsam
fir (Abies balsamae (L.) Mill.). Can. J. Zool. 30: 1-29.

 

 

Blais, J.R. 1953. Effects of the destruction of the current year's
foliage of balsam fir on the fecundity and habits of flight of
the spruce budworm. Can. Entomol. 85(12): 446-448.

Blais, J.R. 1954. The recurrence of spruce budworm infestations in
the past century in the Lac Seul area of northwestern Ontario.
Ecology 35(1): 62-71.

Blais, J.R. 1958. The spruce budworm situation in the Lower St.
Lawrence and Gaspe at the end of 1957 with special reference
to spraying operations. Pulp Paper Mag. Can. 59(6): 218-224.

Blais, J.R. 1960. Spruce budworm outbreaks and the climax of the
boreal forest in eastern North America. Rep. Quebec Soc. Prat.

Blais, J.R. 1968. Regional variation in susceptibility of eastern
North American forests to budworm attacks based on history of
outbreaks. For. Chron. 44(3): 17-23.

Blais, J.R. 1973. Control of spruce budworm: Current and future
strategies. Bull. Entomol. Soc. Amer. 19: 208-213.

Conover, W.J. 1980. Practical Nonparametric Statistics. 2nd Ed.
John Wiley & Sons, New York.

Eidt, D.C., and M.D. Cameron. 1970. Pretreatment of spruce budworm
eggs for counting. Can. Dept. For. BieMon. Res. Notes. 26(5):
46-47 a

Fettes, J.J. 1950. Investigations of sampling techniques for popu-
lation studies of the spruce budworm on balsam fir in Ontario.
£2_Ann. Rep., For Insect Lab., Sault Ste. Marie, Ontario.

86

87

Greenbank, D.O. 1963a. Staminate flowers and the spruce budworm.
.lE.R'F° Morris (Ed.) pps. 219-223, The Dynamics of Epidemic
Spruce Budworm Papulations. Mem. Entomol. Soc. Can. 31: 1-332.

Greenbank, D.O. 1963b. Climate and the spruce budworm. .12_R.F.
Morris (Ed.) pps. 174-180, The Dynamics of Epidemic Spruce Bud-
worm Populations. Mem. Entomol. Soc. Can. 31: 1-332.

Harris, J.W.E. 1963. Sampling the egg stages of the two-year-cycle
spruce budworm near Babine Lake, British Columbia. For. Chron.
39(2): 199-204.

Harvey, G.T. 1977. Mean weight and rearing performance of successive
egg clusters of eastern spruce budworm (Lepidoptera: Tortricidae).
Can. Entomol. 109(4): 487-496.

Jaynes, H.A., and C.F. Speers. 1949. Biological and ecological studies
of spruce budworm. J. Econ. Entomol. 42(2): 221-225.

Kemp, W.P. 1981. The significance of non-host and alternative host
tree species on populations of larval spruce budworm with empha-
sis on improving sampling techniques.

Kemp, W.P. 1978. Stand parameters affecting mortality of spruce bud-
worm eggs and dispersing larvae. Master's Thesis. Michigan State
University Pest Management Series. 134 pp.

Kemp, W.P., and G.A. Simmons. 1978. The influence of stand parameters
on the parasitism of spruce budworm eggs by Trichogramma minutum.
Environ. Entomol. 7: 685—688.

Kemp, W.P., and G.A. Simmons. 1979. The influence of stand factors on
population levels and dispersal losses of instar-I and -II spruce
budworm. Environ. Entomol. 8: 993-996.

Kimmins, J.P. 1971. Variations in the foliar amino acid composition
of flowering and non-flowering balsam fir (Abies balsamae (L.)
M111.) and white spruce (Picea glauca (Moench) Voss) in relation
to outbreaks of the spruce budworm (Choristoneuna fumiferana
(Cleml). Can. J. 2001. 49(7): 1005-1011.

 

 

 

Leonard, D.E., G.A. Simmons and G.K. Van Derweker. 1973. Spruce bud-
worm: Technique to improve counting of eggs. J. Econ. Entomol.
66(4): 992.

Mattson, W.J. 1980. Herbivory in relation to plant nitrogen content.
Ann. Rev. Ecol. Syst. 11: 119-161.

McKnight, MgE. 1969. Estimating numbers of eggs in western spruce
budworm egg masses. USDA For. Serv. Res. Note Rocky Mt. For.
Range Exp. Sta. RM+146. 4 pp.

88

McKnight, M.E. 1971. Natural mortality of the western spruce bud-
worm, Choristoneura occidentalis, in Colorado. USDA For. Serv.
Res. Papa RM-Blo 12 pp.

 

Miller, C.A. 1957. A technique for estimating the fecundity of natural
populations of the spruce budworm. Can. J. 2001. 35(1): 1-13.

Montgomery, B.A. 1981. An analysis of inter- and intra-tree effects
on the size of spruce budworm egg masses on balsam fir. Master's
Thesis. Michigan State University. Michigan Cooperative Forest
Pest Management Program. Info. Report 81-2. 183 pp.

Morris, R.F. 1949. Budworm population techniques in the Green River
project. Bi-Mon. Prog. Rep. For. Insect Invest. Dept. Agric. Can.

Morris, R.F. 1954. A sequential sampling technique for spruce budworm
egg surveys. Can. J. Zool. 32(4): 302-313.

Morris, R.F. 1955. The development of sampling techniques for forest
insect defoliators, with particular reference to the spruce bud-
worm. Can. J. Zool. 33(4): 226-294.

Morris, R.F. 1963. The spruce budworm. .lg R.F. Morris (Ed.) pp. 12-19,
The Dynamics of Epidemic Spruce Budworm Populations. Mam. Entomol.
Soc. Can. 31: 1-332.

Mott, D.G. 1963. The forest and the spruce budworm. .12_R.F. Morris
(Ed.) pp. 189-202, The Dynamics of Epidemic Spruce Budworm Popu-
1ations. Mem. Entomol. Soc. Can. 31: 1-332.

Neilson, M.M. 1963. The analysis of egg survival in the unsprayed
area. t1g_R.F. Morris (Ed.) pp. 37-41, The Dynamics of Epidemic
Spruce Budworm Populations. Mem. Entomol. Soc. Can. 31: 1-332.

Nie, N.H., C.H. Hall, J. G. Jenkins, K. Steinbrenner and D.H. Bent.
1975. SPSS: Statistical Package for the Social Sciences. 2nd Ed.
‘McGraweHill Book Company, NY.

Otvos, I.S. 1977. Estimating the number of eggs in spruce budworm egg
masses in Newfoundland. Can. For. Serv. Bi-Mon. Res. Notes. 33(3):
17.

Outram, I. 1971. Aspects of mating in the spruce budworm. Chorist-
oneura fumiferana (Lepidoptera: Tortricidae). Can. Entomol.
103(8): 1121-1128.

Sokal, R.R. and F.J. Rohlf. 1969. Biometry: The principles and prac-
tice of statistics in biological research. W.H. Freeman and Co.,
San Francisco, CA.

89

Staedler, E. 1974. Host plant stimuli affecting oviposition behavior
of the eastern spruce budworm. Ent. Exp. & Appl. 17(2): 176—188.

Washburn, R.I., and J.E. Brickell. 1973. Western spruce budworm egg
mass dimensions--an influence on papulation estimates. USDA For.
Serv. Res. Paper Intermt. For. Range Exp. Sta. INT-138. 20 pp.

White, T.C.R. 1974. A hypothesis to explain outbreak of looper cate-
pillars, with special reference to populations of Selidosema suavis
in a plantation of Pinus radiata in New Zealand. Oecologia. 16(4):

 

Wilson, L.F. 1963. Host preference for oviposition by the spruce bud—
worm in the Lake States. J. Econ. Entomol. 56(3): 285-288.

Witter, J.A., C.E. Olson, Jr., T.P. Mag, J. McCarthy and C. Karpinski,
Jr. 1980. the spruce budworm with emphasis on impact and damage
assessment studies in Michigan's Upper Penninsula. Univ. Mich.
Sch. Nat. Res. News. 19: 5-7.

Witter, J.A. 1981. Progress report and renewal cooperative forestry
research report: Impact on the spruce budworm in Michigan's Upper
Penninsula. Univ. Mich. Sch. Nat. Res. (unpublished)

APPENDICES

APPENDIX A.

90

APPENDIX A.

Six maps denoting the location of the five sampled stands within
the Ottawa National Forest and the location of each stand within the

corresponding township.

 

       

_ .f
I, ..
. T .
. . i - 4 9'
. v ‘
. . .
y) ,, . .v 4 ..
“5 34f. -... .. I I I
. _.\4I ) I \\ n
H / a r ._
L t / .I‘\I( . _
. fl D I - 4. . ,
. a
‘./..f . u 2...... .5
u .
_ / . I
,. , 545:... :1:_:/ 45::
. .,7
\

   

Il'

..JVviilm l
J

Eu” .

    

 

 

 

91

 

 

 

.mauwm zvdum a>Hw ocu mo mcowumuoa :ua3 umuuom HmCOHumz mamuuolosu mo max .H<.uu:wfim

92

Figure A2. Map of the five study sites within the Iron River,
Watersmeet and Kenton Districts of the Ottawa National

Forest.

93

at...

n";

 

I .4 ‘ t _J . NATION I.
g . ‘ , .r

94

Stand 1 was located in the NE 1/4, SW 1/4, Section 6, T45N,
R36W, Iron River District, Ottawa National Forest, Iron County,
Michigan. The stand was located on Forest Route 139, 1.25 miles
north of the intersection of Forest Route 139 and County Route
657. The spruce-fir packet was approximately 450 meters due

east of the road through a white birch-aspendmaple type forest.

95

Stand 2 was located in the NE 1/4, SW 1/4, Section 6, T45N,
R36W, Iron River District, Ottawa National Forest, Iron County,
Michigan. The stand was located on Forest Route 139, 1.00 miles
north of the intersection of Forest Route 139 and County Route
657. The spruce-fir packet was approximately 100 meters due east

of the road on an abandoned logging road.

96

Figure A3. Map of the locations of Stand 1 and Stand 2 in Section

6 of Township T45N, R36W, Iron County, Michigan.

97

’42 u. /.

. h

>1... . :1 ...ol

.i-ru:

 

§.£.A:
gt.

.0 \nltla
1/ \s

l

l ’

 

 

 

 

Cl... OOOIOOOIOOL
...
one...
.000.
8 '4
"‘

d7

\

"’

d 4 I1 \\\ \t 1
0 ‘ ,
~\\\ \\s \I‘ \\\ .7 \ 04'
w. + a . - \Lzr. \ \s \\ . a!“
.. n i x 2:. 4 \ . \.
a . ¥ \ * \ “\ I
8 . o \ If... \\ \|\ \
.. .. .. 8 . I. r . A . 8..
’0' \ ’ \ \
u a . s a II \\ e
M . u 5 .. \ x i ”it: I s (IHW\
. \
H I II\ s I, a \\ \\ s». \ N \I
\“ 0.. -51-- l, \ \\ .1 c
cl
“ .0 fII’ \ \ 4 \ 0-
r r .0. I. ‘ ‘\ \\ .— u \ I
a. 0 ~ \ . a \
o \ \ u D I \
.. 2 8... x. ~ a I. .
II . \'
\\0lo& ‘ “‘ "'- VI "’ ‘.\~ \A“
a. . . .u \\ \ II. \\'- "' \
~ .I L‘ '0. "\. \ \\\\
xiv \ \ 0t \
‘-|"‘ I...' \ \
. ‘
.4 II: I
n... {as},

 

 

 

new

I

 

 

IJ.

 

 

 

 

 

“\tlll
"'- 0",

A,
.7-
4W9“?

.-..I\V..

L

 

 

 

Ill!!!)
I I

Ra's-...!!!" U

Sol-1903 O u!-
IIIII I. I!
all}! ‘ val!-
IlanD. a liar-III."
ill-a O tilt-la}
III-II ® til-ail.
In": E ill-2|...

a: .a . o...-

22
z<0.ru_2 Shzacu 20¢.

.3006. ..zmvd.

e .83“. .22.... .588 @

...-0 0:30.! I «to.
C30... ......
eel-1.2.01 .0 ...-[.3100 o 3

98

Stand 3 was located in the NW 1/4, NE 1/4, Section 30, T46N,
R37W, Iron River District, Ottawa National Forest, Iron County,
Michigan. The stand was located on Forest Route 364, 1.8 miles
west of the intersection of Forest Route 364 and Forest Route 164.
Stand was on the opposite side of the road from Tinsel Lake. It

was approximately 40 meters due west of road.

99

Figure A4. Map of the location of Stand 3 in Section 30 of

Township T46N, R37W, Iron County, Michigan.

100

 

 

—‘. Lib
:-

1. I.
II
I

3‘3”
_4-
Ti.

 

”.—
_-

't

 

 

- 21.; . ...! .3].

...—I‘ll!!! U

...-.1 in I al..

 

......I. a.
— gist-i
. 1.02: B vocatoofiivvoic "
d.
. . . C

a

 

 

'0‘! j 9.]. .... .....d 3.1 .S .9}

.
_ ‘8'. r a m {3..-1...}!!! §§

     

 

 

 

. . F ,
n fl. 1‘ k
+ q o A o I‘II'FOOIIOOHHNM‘O

3:le :IIITII/HILTII

0 n50—
. m 23:83 .5228 22:

..m .33.: :on ....
_ .x G 39.0“. .3232 030:0 ®
.6 m: .

1 ... ...:‘2'508 O 3

 

     

 

 

 

 

 

101

Stand 4 was in the NW 1/4, NW 1/4, Section 32, T46N, R38W,
Ottawa National Forest, Ontonagon County, Michigan. The stand
was located on Forest Route 171, 5.9 miles north of County Route
208 (Old U.S. 2) or 0.6 miles south of the intersection of Forest
Routes 171 and 172. It was approximately 40 meters west of the

road and on the opposite side of the road from Interior Lake.

102

Figure A5. Map of the location of Stand 4 in Section 32 of

Township T46N, R38W, Ontonagon County, Michigan.

103

.831. .

Ill

“--‘---.-

 

0 '0’

'I'l'lj

 

.- .
tilt lost--

MN

 

 

 

 

 

 

III 2; p I"; 308.

i _Idloi.tol.} flMU

30:1 Out-0U
3..-c l
6933.593I.n£l¢

0.301 :00.

II.-
v!i.|:enl
‘0... 9.!—

..Ii :05... v0! 70.}.

.3 n2- 9.01}:- ’,..I $9.... I

roll?! Q!

:C. 37.1 no. .9 .9...

no; r:

mggi-«‘--

03.17.! out! . ...-foil §nwu§

:1... _ ..u}

:9—
z<u.IU.2 55.2300 ZOO<ZO~ZO

.3005. ..2 0‘...

® «0050“ .Qp—O—uflz 030:0 @

....e .. .6.) C ,1.
.....a: . o
. ......a: v:

 

 

 

 

 

 

 

3...... ....C .

104

Stand 5 was located in the SE 1/4, NW 1/4, Section 12, Iron
River District, Ottawa National Forest, Iron County, Michigan.
It was located on Forest Route 149, approximately 0.3 miles ESE
of the intersection of Forest Routes 149 and 342 (approximately
0.1 mile southeast of railroad tracks and the south branch of
the Paint River). Stand is 100 meters southwest of Forest Route

149 on a two—track road.

105

Figure A6. Map of the location of Stand 5 in Section 12 of

Township T44N, R37W, Iron County, Michigan.

1'06

 

 

 

 

 

 

 

 

Ale*..I.II
. a

 

 

 

 

 

 

2'! in Inn-80’".
i..'tl!3!l.tlcl U

 

...: £0
‘0’ .0.

...-II";

CS‘IOQ

I;

If A

VOICE

 

 

 

 

 

 

 

.H o < 1'... will”!!! .I‘
.V
C I; POI-‘1} l
I ‘4'. 2|! @ {SEC-...} I. 6’
Ila... a a m it’d-III. ixfi“
:
X , I ..L
s ... a: 2 _ ....-
I J\\\ \\IJ.. uno—
V. .../r... ... 23.5.: .Czaoo 20¢.
ll'---.l--- \
.u .350... ..Z VG .._.
/ ,,.. g 320“. .3232 2520 ®
_ P

 

00.49 ,I:I.-' C I...
I~.b.o¢ one...
Isl-.I-lld o! .90....IIIG I 3

APPENDIX B .

107

APPENDIX B.

Diameter, age, height and total number of egg masses for each

sampled tree.

108

Table Bl. Sampled trees' ages, heights, diameters, and total number of
egg masses.*

 

 

STD SPP Tree S—D DBH Age HT # EMS
1 BF 1 FS 21.0 41 17.8
1 BF 2 EB 23.0 44 16.7
1 BF 5 EB 16.1 43 15.9
1 BF 6 FS 23.4 63 14.6
1 WS 1 EB 27.9 60 14.2
1 WS 2 FS 21.7 70 13.7
2 BF 1 FS 19.3 40 14.8
2 BF 2 EB 17.3 37 13.0
2 WS 1 FS 23.9 51 13.4
2 WS 2 EB 15.9 32 11.7
3 BF 1 EB 15.7 63 11.8
3 BF 2 FS 18.2 63 14.6
3 WS 1 EB 16.2 72 12.5
3 WS 2 FS 18.0 73 14.0
4 BF 1 FS 17.1 45 15.5
4 BF 2 EB 14.3 41 12.3
4 WS 1 EB 16.2 42 12.1
4 WS 2 FS 24.3 45 17.5
5 BF 1 FS 13.2 33 10.1
5 BF 2 EB 12.6 43 12.0
5 WS 1 FS 12.3 42 9.5
5 WS 2 EB 11.5 38 9.4

 

*Refer to the following page for a key to the column headings and their
Units.

DBH:

AGE:

# EMS:

109

Stand number

Tree species:

BF Balsam fir
WS White spruce

Tree number

Sampling design:

1 = Every branch full sample (feasible branches were
disected into varying lengths)

2 = Every branch sample (but feasible branches were
not disected)

Diameter of the tree at breast height (1.37 meters

above ground), measured with a metric "D—tape" to

the nearest 0.1 centimeter.

Age of the tree in years, taken at stump height (8"

above ground) by counting rings.

Total height of the tree measured with a metric tape

after tree was felled. Units were taken to the nearest

0.1 meter.

Total number of egg masses found on all foliated sur-

faces of the tree by the research crew.

APPENDIX C .

110

APPENDIX C.

Distances, angles, diameters, and heights of the ten "nearest
neighbors" (trees with diameters at breast height greater than 12.0

centimeters) of each sampled tree.

111

Table C1. Locations, heights and diameters of trees that were within
10 meters of each sampled tree.*

STD TREE SPP STSP DIST DIR HT DBH
1 1 BE PE 02.3 080 17.7 19.5
1 1 B“ PB 04.9 123 17.4 20.1
1 1 BF PB 04.6 092 20.7 24.7
1 1 BB PB 05.6 034 17.4 24.1
1 1 BE PB 04.2 354 18.6 23.5
1 1 BF PB 00.9 207 19.8 26.2
1 1 BE PB 01.2 317 18.3 25.7
1 1 BE PP 02.5 129 29.1 24.3
1 1 BE PB 04.8 153 19.2 25.4
1 1 BF P8 06.0 315 16.5 15.4
1 2 LE P5 01.8 166 17.4 24.1
1 2 BF PB 02.7 072 18.3 22.8
1 2 BE PB 04.8 096 16.2 15.5
1 2 BE PB 05.4 135 18.6 17.9
1 2 BE PB 06.4 131 19.2 17.0
1 2 -F PB 03.7 234 18.6 23.5
1 2 BF PB 06.7 027 16.5 18.0
1 2 BF PE 06.3 016 18.0 21.0
1 2 BE PB 04.4 333 13.7 13.4
1 2 BE PE 03.6 312 14.9 13.5
1 5 BF PB 03.5 294 16.2 20.4
1 5 BF P3 03.1 315 15.2 19.0
1 5 BE PM 03.3 338 13.4 16.1
1 5 BF P8 03.4 006 15.8 17.3
1 5 BF PB 05.7 311 19.5 28.0
1 5 BF RM 04.7 156 12.5 12.2
1 5 BF PE 03.5 215 19.5 26.7
1 5 BF PB 03.] 261 16.2 21.5
1 5 BF PB 04.4 088 16.2 14.8
1 5 BE PB 04.7 093 16.2 18.4
1 6 BE PB 01.4 278 18.9 19.4
1 6 BE PB 01.5 201 20.7 30.2
1 6 BF PB 03.0 250 14.9 13.2
1 6 BF A 05.7 262 24.4 16.0
1 6 BE P8 03.6 337 14.6 16.0
1 6 BF PB 03.7 356 20.7 26.4
1 6 BE PB 04.8 039 17.1 18.2
1 6 BF RM 06.3 173 11.0 12.4
1 6 BE PB 05.5 221 14.3 21.7
1 6 BF RM 06.2 164 11.0 13.2

*Refer to page after table for key to column headings and their units.

Table C1. Continued

112

MNMMNNMMMMMNNNMMNMMNHHHHHHHI—H—Jt—oI—w—H-JL—al—awu—au—awu

NNNNMMMMMMHHHHHHHHHHNNNMI‘JMNVMNHHl—IHHHHHt—JI—I

NS
.35
WS
NS
NS
NS
118
NS
NS
WS
NS
NS
NS
NS
".JS
".33
NS
NS
NS
NS
BF
BF
BF
BF
BF
BF
BF
BF
BF
BF
BE
BF
BF
BF
BF
BF
BF
BF
BF
BF

PB
PB
PB
PB
PB
PB
PB
PB

B
PB
PB

BF
PB
PB
A

PB

BF
PB
RM
PB
PB
PF»
PB
PB
P8
P8
BF
PB
PB
PB
PB
PB
PB
PB
PB
BF
PB

3.
M

05.

01.
02.
04.
02.
04
04.
:34

gs
44

04.

3
UI

Table C1.

Continued

113

wwwwwwwwwwwwwwwwwwwwmwwmknown)MNNwwwmwwwmm

whom)NroanronJNhahaHFdhdwhdhlwhdmamroxJwrohgmrou>~tdhaHLahawtahaw

NS
NS
NS
NS
NS
NS
NS
3J3
NS
NS
NS
NS
US
NS
NS
US
NS
NS
NS
NS
BF
BE
BE
BF
BF
BF
BF
BF
BF
BF
BF
BF
BF
BF
BF
BF
BF
BF
BF
BF

RM
RM
RM
RM
RM
RM
HS
BF
NS
BF

14.6
12.8
20.4
14.6
13.4
15.8
18.0
16.5
12.5
11.6
20.4
15.8
12.5
14.0
17.1
17.4
18.9
19.8
17.1
17.1
16.1
08.5
19.8
15.5
13.7
17.7
06.7
18.3
14.6
14.3
13.4
16.2
21.6
17.7
07.6
18.0
08.2
11.0
07.9
11.9

18.9
18.4
19.9
15.2
14.1
13.8
20.3
16.3
12.1
15.5
27.9
25.8
16.6
13.2
20.0
17.0
15.5
18.2
15.7
21.7
24.7
14.8
29.7
24.4
16.4
28.9
13.6
26.9
18.2
19.8
19.4
24.7
27.0
28.9
12.0
26.9
15.3
17.2
13.5
15.7

Table Cl.

Continued

114

bbb:bbb~54>bb¢~bb;bbbb.b.bboawwwwwwwwwwwwwwwwwmw

MNMMNNMMMMHHHHHHHHHHNNMMNMNNMMHt—H—w-aI—H—H—H—u—as—a

SPP STSP
WS BF
NS RM
HS BF
'NS NS
NS RM
NS R11
’NS Fm;
2955 VJES
NS RM
WS Ffii
WS F31
1 13 Y E
US BF
NS NS
W3 RM
1} S R11
NS RD:
NS RM
WS Rd
W S Y B
BF BS
BF A
BF RM
BF A
BE' A
BF RM
BF RM
BE’ RF]
BE' BS
BF BS
BF .A
85’ RP
85' BF
BF A
BF BF
BF‘ BF
BF BS
BF RH
BF RM
BF RM

04.
03.
07.
07.
H7.
05.
02.
07.
m7.
07.

be-Jxommmmwosq

.9
U‘.

03.6
06.9
02.5
08.3
06.3
03.6
03.2
01.3
03.6
02.8
04.5
05.5
05.6
04.4
04.9
01.7
03.9
03.8
01.2
00.4
04.4
04.7
05.8
06.2
06.3

15.2
17.7
12.8
08.5
20.7
12.2
11.9
14.6
12.2
11.6
18.3
20.7
12.8
07.6
15.5
15.8
15.5
14.9
18.0
13.7
10.4
17.7
13.7
22.6
23.2
12.8
14.3
14.3
11.9
11.3
14.6
16.8
12.2
19.8
12.2
14.3
12.2
15.2
14.6
13.4

Table Cl.

7115

Continued
STD TREE

4

4

4

4

4

4

4

4

4

4

4

4

4

4

U‘UTU‘UTU'IU‘UTWUTkfiUIUlUTWUWU’IU’U‘UIU‘Dbxbo’bnbuh.

Nbdk)NPQKJNtQBJMFHF‘HFHFJHFAFJHFHN)NDOK)N§ORJM1JKJHfidbdwbdhdwtahaw

SPP STSP
145 BE‘
HS A
W8 A
NS BF
WS RM
NS A
W8 A
HS A
US A
US A
W8 Ru
NS RM
NS Rm
WS RP
195 PB
W8 RM
1.73 BS
ws RP
NS BS
W9 A
BF B
BF 8
BE 3
BF RM
BF E
BF E
BF C
BF C
BF WS
BF E
BF BF
BF WS
BF WS
BF NS
BF RM
BF A
BF A
BF RN
BF BF
BE‘ \JS

12.5
16.2
15.8
12.5
13.4
16.8
14.6
18.6
18.3
15.5
17.4
16.8
16.8
10.7
18.3
12.2
12.5
22.8
16.8
18.9
15.8
14.3
12.3
16.5
12.5
14.0
14.0
12.8
12.5
12.5
14.9
13.1
13.4
11.0
12.8
16.5
15.4
13.4
10.4
11.0

14.4
13.4
17.3
13.7
13.2
19.m
17.2
32.6
20.2
22.0
12.2
16.1
20.2
17.8
17.3
13.2
18.4
22.7
16.1
29.2
18.2
13.9
21.7
47.6
13.8
14.2
15.9
14.9
19.0
13.5
17.5
16.9
27.6
13.4
14.9
15.4
19.8
13.2
14.9
16.0

Table C1. Continued

116

28.4
19.2
20.9
17.8
13.7
13.6
15.0
29.3
14.0
19.0
26.9
17.4
16.6
17.3
13.2
16.5
18.4
19.9
12.8

FINAL EXECUTION

117

STD: Stand number
TREE: Tree number
SPP: Sampled tree species:

BF = Balsam fir
WS = White spruce

STSP: Surrounding tree species:
BF Balsam fir
WS White Spruce

 

 

 

 

A Aspen (Populas tremuloides Michx.)
PB Paper birch
RM Red maple (Acer rubrum L.)

BS = Black spruce
RP = Red pine (Pinus resinosa Ait.)
YB = Yellow birch (Betula lutea Michx. f.)

C = Cherry (Prunus sp.)

E = Elm (Ulmus sp.)

DIST: Distance from sampled trees outside bark at breast height

to surrounding tree's outside bark at breast height to

the nearest 0.1 meter

ANGLE: Direction of surrounding tree, taken from the sampled
tree with a Sylva "Type 15 T" compass, declinated at

4° east of true magnetic north

DBH: Diameter at breast height of the surrounding tree to

the nearest 0.1 centimeter

HEIGHT: Height to the nearest 0.1 meter taken with a Suunto

clinometer in feet and later converted to metric units

APPENDIX D .

118

APPENDIX D .

Stem maps

Figure D1.

K

 

 

*The tree
sizes or
of those
tions of

to Appendices B and C.

   

119

Key to symbols used in Figures D2—D23.*

BF*: Balsam Fir (sampled trees)

BF IBalsan'. Fir

WS*: \Nhlte Spruce (san'upled trees)

W8 :Whlte Spruce

BS : Black Spruce

RM: Red Maple
A : Aspen

PB: Paper Birch
YB: Yellow Birch

JP : Jack Plne

.C : Cherry

E :Elm

symbols on all maps are not representative of actual crown

shapes.

They represent relative stem sizes and positions

stems around the sampled trees. For exact sizes and loca—
the sampled trees and the trees that surrounded them, refer

Numbers that are next to the symbols and the

species abbreviations are the trees' diameters in centimeters.

120

 

Sf; STAN D 1
23F * BF 1'
5 Scale : 1cm = 1 m
WROXNAYEMEAN
DECUMWONJ968

 

P B 24.1
P3 23.5 r
P B 15.4
P8 25.7
P8 25.4
Figure D2. Stem map of the ten nearest trees that encompassed sampled

tree BF 1 in Stand 1.

Figure D3 .

121

 

 

, 5 STAND 1
:3 :1: BF 2
E; Scale : 1 cm = 1m
W‘Sifie‘:
P8 13.4
§
BF*2a.o PB 2“
P8 15.5
1!
P8 23.5
PB 17 o

 

Stem map of the ten nearest trees that encompassed sampled

tree BF 2 in Stand 1.

122

 

5E STAND 1.
:2 :1: BF 5
"E Scale : 1cm = 1m

won-ant nun
oeumn-on. 1960

PB 28.0

 

Figure D4. Stem map of the ten nearest trees that encompassed sampled
tree BF 5 in Stand 1.

123

Q

’

 

STAND 1-
i=£3l= £5
Shaale : 1 crn : 1 n1

NW?”

 

1M seam

“ACNE—rut

WROIIUA‘E «(AN
occunmou I96!

A 16.0

 

I’EBSOJ

F’B»2L7

 

Figure D5. Stan map of the ten nearest trees that encompassed sampled
tree BF 6 in Stand 1.

124

‘0

1 g STAND _1
§§ *ws 1

Won-uncut“ Scale 2 1 cm = 1 m
oechAnou. 1960

P8 15.5

P8 24.1

 

PB 20.1 P8 19.8

Figure D6. Stem map of the ten nearest trees that encompassed sampled
tree WS 1 in Stand 1.

125

$5 STAND 1
9
§§ * ws 2
Scale: 1 cm = 1 m

mom." nun
munnlou. 1961

BF 13.5

)1
w/l//

A 20.6 )&@11
I, '
All"

.1 «..x
\ llu' A
A 13.8 fiblt/ 22 25 o
'Z/‘v’s '
’ ' 5.:
\ ' .

P3 15.7

 

Figure D7. Stem map of the ten nearest trees that encompassed sampled
tree WS 2 in Stand 1.

126

‘.

 

if; STAND 2
a; * BF 1
"l; Scale : 1 cm = 1 m

women: nun
DICLIMTION. 196!

P3 20.9
P8 23.5

 

P8119

R M 12.2

BF 17.8

Figure D8. Stan map of the ten nearest trees that encompassed sampled
tree BF 1 in Stand 2.

127

a”: STAND 2
g}; :1: BF 2 -
”g Scale: 1cm=1m

MOMMA V: nun
O

ucuuum, 19¢,

 

P B 20.2
PB 10.3
PB 28.8
PB 21.8
P8 15.1
PB 20.7
P B 22.1

Figure D9. Stem map of the ten nearest trees that encompassed sampled
tree BF 2 in Stand 2.

128

if STAND 2
E; *ws 1'
3 Scale: 1 em = 1 m

MOMMA“ MEAN
“CLINANON. I96!

PB 12.1
P B 13.3

PB 20.3

P 915.5

 

Figure D10. Stan map of the ten nearest trees that encompassed sampled .
tree WS 1 in Stand 2.

129

SE STAND 2
g? * ws 2
g Scale: 1cm = 1 m

WAY! KAI
octuuAv-ou. 1963

PB 21.7

 

P B 20.0

PB 15.5

PB 17.0

Figure D11. Stan map of the ten nearest trees that encompassed sampled
tree WS 2 in Stand 2.

130

'1

g; STAND 3
gé’ * BF ‘_
3 Scale: 1 cm = 1 m

MOMMA“ NEAN
DECUNA'ION. 1960

RM 2” B F(2)13.2

B F13.6

RM24.7
RM 26.9

3|:
BF 15.7
B F 10.4

R M 24.4

 

B F 10.3

P329]
Stem map of the ten nearest trees that encompassed sampled

Figure D12.
tree BF 1 in Stand 3.

R M 28.9

Figure D13.

 

BF(1) 15.7

131

.0

E 8

{f3 STAND

i; * BF
Scale :

Wynn; “A"
MCUNAYION. l9“

 

Stem map of the ten nearest trees that encompassed sampled
tree BF 2 in Stand 3.

132

5511\flt) :3
*ws 1
£3CNBIG>: ‘1crn = 1 n1

 

7M noun:

“Acmicmv -.

 

man-an; nun
“tonne... ,9“

 

BF19.6
\\ RM1e.2
RMaoo
V /’ RM327
4
-.. I __
I
k‘\ f

\
A’I .

74/“ ' '17!le
WS*16.2 WS12.3

R M 20.0

 

RM 13.4

Figure D14. Stem map of the ten nearest trees that encompassed sampled
tree WS 1 in Stand 3.

133

‘

 

:5 STAND 3
:3}. a: we 2
"I; Scale: 1cm 2 1 m

mom; '. BLAH
mama'v~ :91:

 

 

 

 

 

 

RM 14.1

\

RM 23.1

Stem map of the ten nearest trees that encompassed sampled

Figure D15.
tree WS 2 in Stand 3.

134

5 STAND 4
:8 *BF 1

Scale: 1cm = 1 m

m”! at»:
match. 1963

 

Figure D16. Stem map of the ten nearest trees that encompassed sampled
tree BF 1 in Stand 4.

135

gig £31WAJ‘I3 1‘

2 *BF

" Scale: 1cm = 1
WA?!“

Mlm. l9“

 

Figure D17. Stem.map of the ten nearest trees that encompassed sampled
tree BF 2 in Stand 4.

136

STAND 4
Eg * WS 1
Scale 1 c m = 1 m
Wm:o¥.1‘..‘:
A 32.0
,<
RM 13.2
A 19.0
A 20.2

 

A 22.0

Figure D18. Stem map of the ten nearest trees that encompassed sampled'
tree WS 1 in Stand 4.

137

“STAND 4
*ws 2

Scale: 1cm=1m

 

lg ”at“
NU‘W '

MOMMA? I MEAN
“CUNAT‘ON. |965

JP22.7

A 29.2

 

R M 13.2

 

Figure D19. Stem map of the ten nearest trees that encompassed sampled

tree WS 2 in Stand 4.

138

Figure D20.
tree BF 1 in Stand 5.

tampon"
\' ~.
In"

“Mini vic‘

 

woman; MEAN
DCCUNATION. 1955

 

E 13.5

Stem map of the ten nearest trees that encompassed sampled

-STAND 5
* BF 1
Scale: 1cm = 1 m

 

139

 

S. is STAN o 5
33’ * B F 2
mm Scale : 1cm = 1m
mews:
R M 14.9
ws "5‘9 BF 14.9
$’% ws 1s.

\_

 

% WS 13.4

WS 27.6

A 15.4
A 19.8
Stem map of the ten nearest trees that encompassed sampled

Figure D21.
tree BF 2 in Stand 5.

140

J; STAND 5
gr 3|: WS 1
Scale: 1 cm = 1 m

I
.

 

ADIOWA'I “AN
DICLINAYION. 196s

 

WS 19.0

Figure D22. Stem map of the ten nearest trees that encompassed sampled _
tree WS 1 in Stand 5.

141

'1

STAND 5
* WS 2
Scale : 1cm = 1m

 

1M mu

———~

“6»ch

AWROWAYE MIA-4
DECuNAYION, 1965

 

_ WS 26.8
RM 13.2

RM 16.5

 

Figure D23. Stan map of the ten nearest trees that encompassed sampled
tree WS 2 in Stand 5.

APPENDIX E .

142

APPENDIX E.

A graphical presentation of the field methods-the procedure used

in egg mass sampling research.

143

At each stand, camp was set up at a convenient location. Two
screen tents housed the research crew, while they examined foliage for

egg masses. The other tent protected the sampled branches.

 

Figure E1. The field camp at Stand 5.

144

Trees that were chosen for sampling were flagged with orange
ribbon and their trunks were painted with the appropriate identifi—

cation letters and numbers.

Figure E2.g;»
Sampled
tree BF
2 in

Stand 3.

Figure E3..%}‘
Sampled
tree WS
1 in

Stand 2.

 

Sampled trees' diameters, heights, ages, and locations within

the stand were recorded.

145

Climbing spikes and harnesses were used in the measurement and
collection of branches. Here, a research assistant has positioned
the metric tape at the end of his pruning saw into the "center of

gravity" of the branch.

V

,5
...
_ ~ 3' o .
xii; .. .91.».th

 

Figure E4. Measuring the height of a branch-a research assistant on
the sampled tree.

146

A research assistant below the tree would extend the metric tape
perpendicular to the ground and would note the height of the branch
to the nearest 0.1 meter and the quadrant that the branch grew in.
Orange flagging had been layed on the ground at NW—SE and NE-SW angles,

thus forming the four quadrant areas.

 

Figure E5. Measuring the height of a branch and noting its directional

quadrant-a research assistant on the ground.

147

The research assistant on the ground would record the tree number,

branch number, branch height and directional quadrant on a small tag.

 

Figure E6. Recording the branch number, height and quadrant-a research

assistant on the ground.

148

The research assistant on the sampled tree would sever the branch

at its base.

 

Figure E7. Cutting the sampled branch.
and would gently drop the severed branch to the research assistant on

the ground.

 

Figure E8. Dropping the branch to the "spotter" on the ground.

149

The spotter would carefully catch the branch.

3%.?“
""2 "E. _.
\’ 1

 

Figure E9. Catching a sampled branch that was dropped from the tree.

150

The research assistant on the ground would tie the tag onto the

branch.

 

Figure E10. Tying the identification tag onto the branch.

and the branches would be piled near the tents.

 

      

V ’ a
“‘I _ _/

Figure Ell. A pile of severed
tents.

'
.I . L

r the screen

\

and tagged branches—nea

151

Two research assistants would measure the total length, foliated
length, and foliated width of each branch. This information would be

recorded on the branch tag.

 

Figure E12. Research assistant measuring the foliated width of a branch.

152

Each research assistant would choose the top branch from the pile
of measured branches and would clip the branch into pieces that were
convenient to examine-approximately 10-20 cm in length. Each foliage

piece would be thoroughly examined for egg masses.

 

Figure E13. Research assistant examining foliage for egg masses.

153

If an egg mass was found, the examiner would remove the needle,
holding the egg mass, from the foliage piece (with forceps) and would
place the egg mass and needle in a vial. He or she would label the
vial with the appropriate stand, branch, and tree number. The number
of egg masses and the appropriate data from the branch were recorded

in the research assistant's field book.

 

Figure E14. Research assistant removing a needle and an egg mass from
a foliage clipping.

154

After the foliage clippings had been examined, the research assis-
tant would measure the branches foliated area on a grid. The grid was
marked off in 5 cm segments. Theoretically, the foliage would be placed
on the grid so that none of the grid board was visible and none of the

foliage pieces overlapped one another.

 

Figure E15. Measuring the foliated area of a sampled branch on a grid
board.

 

155

The highest branches were measured and cut, so that only the top

of the tree was remaining.

 

Figure E16. The top of the sampled tree.

The tree was then felled and data was collected on tree height,

age, and heart rot.

 

APPENDIX F .

156

APPENDIX F.

Normality and homogeneity test results

for eggxnass groups used in parametric statistics

"Subprogram Condescriptive" (§E§§, pp. 185-193) (Nie et al. 1975)
computed skewness and kurtosis, measures of nonnormality for continuous
interval-level data. Skewness measures deviation from symmetry while
kurtosis measures the flatness or peakedness of the frequency distri-

bution.

 

N _ N
zxi-3x 2x
i=1 1=

F

Skewness

N

Z ((x. - 1'1) Is)‘+
i=1 1
N

- 3

 

Kurtosis

SPSS‘s (Nie et al. 1975) computational formula:

N N
2 x14. 2INi+6XL(1Z113XXZ21 -4X3(‘Z1Ixi/N +x
_ ,i=1 -3

Kurtosis -

NX—i -Nxz /(N-1)
i=1

A normal distribution has a skewness of zero and a kurtosis of

zero. A two-tailed t-test, as described by Sokal and Rohlf (1969)

157

(pp. 166-172), was used to test for significant deviations from zero
(i.e., do the observed skewness and kurtosis values for each egg mass
sample group indicate a significant departure from normality?) where:

the sample statistic for skewness,

00
F‘
II

- the sample statistic for kurtosis,

00
N
I

Y1 = the expected value of skewness for a normally
distributed p0pulation (i.e., 0),

Y2 = the expected value of kurtosis for a normally
distributed population (i.e., 0),

Sgl'= the standard error of g1,
sg2 = the standard error of g2, and
n = sample size.

Estimations of values were computed with the following formulas:

_ 6n(n-1) _
Sg1 i/(n-2)(n+l)(n+3) ’ df -

‘lg: for large n (>100)

=lfi 24n(n--l)2
ng \I (n—3) (n-2) (n+3) (n+5) ’

‘léé for large n (>100)

 

 

 

 

df =

t 1 = ‘1'“—
S g1
t (gz-YZ)
32 S32

Estimated values of tsl’ and t82 were evaluated with critical
values of t with degrees of freedom, v.

Homogeneity of variances (homoscedasticity) was tested for each
egg mass group with the Bartlett test (Bartlett-Box F statistic),
"Subprogram Oneway",.§§§§ (pp. 422-433) (Nie et a1. 1975). The statis-

tical tests that were used in the thesis require equality of variances

158

in a group of samples. "Subprogram Oneway" incorporates Bartlett's
test and computes Bartlett's-Box F statistic and the corresponding
significance probability, p, for each egg mass group. Although the
§2§§_manual does not provide definitional or computational formulas
for the statistic, Sokal and Rohlf (1969) provide a good description
and an example of the Bartlett test on pages 369-376 of their text.
The value p permits easy assessment of the rejection level for homo-
geneity of variances (i.e., if p was found to be 0.110, homoscedasti-
city would be rejected for m > 0.110 but not for a < 0.110).
Normality and homoscedasticity were not confirmed for groups of
randomly sampled egg masses that are marked with an (3) (significant
departure from normality or homogeneity). These groups required statis-
tical tests other than the parametrical test, oneway analysis of var-

iance.

159

uoom uoou mumsvma

 

 

 

 

 

 

 

 

OH.OAaAo~.o omq.HI ome.o mHm.on om.CAaAoo.o ooo.o1 oq~.o «SH.OI soMIm
9 e e e e e o e I 9 e I vHHSH Hafiuumm
om OAQAON o «mm o mwe o mmm 0 mo OAQAOH o mum H new o woe o m nuH3_3OMI~
OH.OAaAON.o mmq.HI onq.o mwo.o1 ow.OAona.o mmH.o1 wmm.o oqo.o1 BOMIN
w
sufiaanmnonm Nmu «mm mm suSHfinmnoum Hm“ H m Hm
HHMHIN HHmHIN
mHmouHSM mmmcawxm
339682
Hum.o who.H www.o omo.o oqq.m «OH Bealm
. . . . . oHHAH HmHuumm
«NH 0 mum H oao o Ono o Hem q mm m nuHa BOMIN
omm.o Hem.H mHm.o mwo.o ooo.q oOH Boalm
m m NomluumHuHmm COHumH>wn Houum one: wwm oNHm oaay mom: wwm
moosMHum> mo muncmwwaom oumosmum oumvcmum Hon wwwm mo wHaEmm
Hmnaaz new:
.moumaHumo

:OHmmoummu pom pom: .uHm ammHmn Scum mommma www MOM muHammu ummu %uHmcmono: one muHHmahoz .Hm mHan

160

uoom uoou mumsumk

 

 

 

 

 

 

 

an.0AaAow.o NcN.o qu.o BNH.OI OH.0AaAom.o wmm.H meN.o omm.o1 BoMIm
om.0AaAo~.o cq¢.o om¢.o ¢H~.o ON.OAaAom.o wMH.H oq~.o mn~.o ouHsH HNHuumm
m :uHS somlm
om.OAaAoo.o Noo.o omq.o mmu.o: 0N.OAaAom.o «mm.H «HN.o com.on BOMIN
m w
suHHHnmnoum Nan Nam mm suHHHnmnonm H u H m H»
HHMHIN HHmHIN
mHmouu5M mmoaawxm
huHHmahoz

own.o Hmc.o om©.o «00.0 owq.m NOH zealm
«No.o nqe.m moo.o moo.o NHw.q «OH ouHSH HMHuumm
m :qu 3oMI~
on.o Hom.H How.o ono.o oHH.q OMH zoMIN
m m xomluuwHuumm ¢:0HumH>ma «Hosea some: wwm mNHm make mom: wwm

vumocmum osmocmum use wwwm mo oHaamm

moo:MHum> mo Nmeamonom

Hmnabz one:

 

Ion new mom: .ooauam ouHca Scum mowmma wwo pom muHsmoH

.moumaHumo cOHmmouw
ummu zuHmawonos was auHHNEHoz .Nm mHamH

161

 

omm.o n m .eoo.H u UHumHumum m somuuuoHuumm
mooamHum> mo NMHoauwoaom

 

OH.OAaAON.o me.H moo.o mma.0| om.OAa oNH.o aom.o amo.c1 umo3

 

 

 

om.oAaAcq.o o~m.o mHo.o com.o1 ow.OAaAom.o moH.o HHm.o ooo.o1 nusom
OH.oxaAo~.o Nam.H woe.o mom.ou oa.oAaAow.o mmm.o oom.o mHH.o- ummm
00.0AaAo~.o oe<.o woo.o oom.ol om.0Aa amo.o oem.o OHo.o :uuoz
huHHHpmnoum qu Nwm mm huHHHnmnoum Hmu me Hm ucmuomao
HHmBIN HHmHIN
meouu=M mmmasmxm
huHHmauoz

 

.momuu uHm ammHmn cHauHB musmuowav umoz one suaom .ummm .nuuos
Scum Awuoou oumsvm oucH omauommcmuuv assoc wwm How wuHamou ummu muHmomono: can huHHmshoz .mm anmH

162

 

msa.o n m .mm~.o n oHumHumum a xomuuumHunmm

 

mouamHum> mo huHmcmono:

 

 

 

 

no.OAaAoH.o oqw.H mmq.o nmw.01 ow.OAaAom.o emu.o n-.o 50H.OI yoga:
0H.0AaAom.o ome.H oom.o MH~.o1 om.0AaAco.o moo.o mm~.o mmH.o mHosz
00.0AaAon.o owe.o Nnm.o mne.o1 oa.OAu omo.o Hom.o mHo.o uwaoH
muHHHnmnoum Hmu Nwm mm zuHHHnmnoum Hmu me Hm abumuum
HHMHIN HHmHIN
mHmouu=M mmosawxm
zuHHmauoz

 

.momuu HHm BNmHmn chuHB mumuum swam: can .mHooHE .um30H Baum
Amoco“ mumsvm cucH omENOMmcmuuv mucaoo www New muHamou ummu %uHocmonon one muHHmauoz .Sm oHan

163

 

omm.o u m .omo.o u UHumHumnm s womuuumHuumm

 

moocmHHm> mo muHmaoonom

 

 

 

 

OH.OAaAo~.o «so.H mmm.o acm.H No.0AaAmo.o oem.~ Has.o meH.H ace
om.oAaAos.o mmm.ou Nam.o «Hm.on om.OAaAoo.o qu.ou msq.o ms~.on saw: “one:
om.OAaAom.o soo.Hu mmm.o oww.ou om.oAaAoe.o Hem.o- aNe.o eeN.ou nuaom nmaa:
o~.0AaAcm.o omH.H1 mow.o mmm.ou om.OAa o~o.ou «He.o NHo.ou ummm “mans
04.0AaAom.o ama.ou mam.o meo.ou om.OAaAoe.o mmm.o- mee.o He~.o- nunoz “mans
oH.oAaAo~.o mmm.H- amw.o oom.Hu om.OAa o~o.ou omq.o Nmo.on umms mHquz
om.OAaAoe.o cem.ou aww.o sme.on om.OAaAoe.o Hoo.o oms.o s-.o zuaom meeHz
oe.OAaAom.o Hma.ou Nea.o HHa.o- oa.OAa soo.o Hom.o Noo.o “was mHueHz
ow.OAaAom.o am~.o wmo.H ee~.o om.OAaAom.o maH.H omm.o omo.o eunoz mHseHz
om.oxa OHH.o awm.H maH.o o~.o.aAow.o smm.o «ma.o mo~.o ummz nmsos

o.H ooc.o mum.~ ooo.o om.oAaAos.o omo.H m-.H mmm.H auaom mason
om.oAaAoq.o oom.o1 awm.H ame.Hu om.oxaAoe.o som.ou saa.o mHa.ou “mam “use;
om.OAaAoo.o Hmo.o mHe.~ mmo.H ON.OAaAom.o mmm.H msa.o use.H nunoz noses

m w

suHHHnmnonm Nan N m mm anHHHnmnonm Hmu H m H» HHmo

HHmsum HHmsuu

mHmouHSM mmmc3mxm
auHHmauoz

 

.momuu uHm ammHmn cHnuHs mHHoo MH
Scum Amuoou wumscm cucH oweuommamuuv mucsou www How muHsmmu ummu muHmamonon was muHHm8hoz .mm anmH

164

 

mo~.o u a .Hamm.o n UHumHumum a xomnuumHnnmm

mwucmHHm> mo huHmcwwoaom

 

 

 

 

 

om.OAaAoo.o mwm.o1 mmm.o Hmm.o1 om.cAaAom.o wo~.H ch.o Hmm.o 0mm
oh.0AaAow.o com.OI mn~.H mum.o om.OAaAoq.o NHo.H Hoo.o moo.o mam
oq.oAaAom.o New.o1 mmm.o Hem.01 om.OAaAoo.o nom.o nmq.o Nem.o mum
om.0AaAoq.o qw0.HI HSN.H www.H| om.0AaAoq.c Hmo.H| new.o www.01 Ham
zuHHHnmnoum qu Nwm mm SuHHHnmnoum Hmu me Hw cope
HHmHIN HHmHIN
mHmouu=M mwmaamxm
SuHHmahoz

 

aoum Amuoou mumswm cuCH pmauowmcmuuv

.H vamum cHsqu momuu uHm ammHmn anew
uczoo mwm you muHSme ummu auHmcoonos mam huHHmahoz .om mHan

165

 

 

 

 

 

 

 

mam.o u m .mHm.o u oHumHumum a xomuuanunmm

woocmHum> we szmcmmoaom
ON.OAaAom.o Ono.H mmm.o oNo.HI ow.OAaAom.o oo~.o qu.o moo.on Nam
on.OAaAoq.o mam.o wHa.o mew.on OH.oAaAo~.o Hw¢.H mnq.o mao.o Ham
muHHHnmnoum Nmu Nwm Nw zuHHHnmnoum Hmu me Hm omuH

HHmHIN HHmHIN
mHmOuusx mmmczmxm
SuHHmEuoz

.N vcmum :HnuHa momma amemn 03u

Eoum Amuoou meadow cucH omeuowmamuuv mucsoo wwo Mom muHsmmu umou zuHmcmwoaon was huHHmauoz .nm mHan

166

 

mam.o n a .oHo.o u oHumHumum a xomuuanuumm

mouamHum> mo huHmcuwoaom

 

 

 

 

 

oo.OAaAo~.o qu.o moo.H on.o om.OAaAom.o oqN.H omm.o Nwo.o mum
om.OAaAoq.o qmo.o mnm.o wom.on oq.oAaAom.o cc~.o1 Hom.o Hmm.o1 Hum
auHHHnmnoum Nmu Nwm mm muHHHnmnoum Hmu me Hm mmuH
HHmHIN HHMBIN
mHm0uu=M mmmcaoxm
%uHHmEhoz

 

om UGmum GHSUH3 mwmhu HHM gmHmQ O3“

Scum Amuoou ohmsvm ousH oosuommsmuuv mucsoo wwo pom muHammu ummu kuHocmmoEo: one zuHHmshoz .wm mHan

167

 

amn.o u m .eoo.o u UHumHumum a xomuuumHuumm

 

mmocmHHm> mo Nmeamonom

 

 

 

 

om.OAaAoc.o Homnom mom.o qmq.01 oo.OAa 00H.ol ¢H¢.o qqo.ou «mm
om.0AaAoo.o coo.o won.o qo¢.c ow.OAaAo~.o Noq.o mam.o me.o Ham
w w
huHHHnmnoum Nmu N m Nw huHHHnmnoum Hmu H m Hm ompH
HHMHIN HHmHIN
mHmouusx mmosBmxm
huHHmEHoz

 

.q occum aHsuHB momma uHm EmmHmn o3u
Scum Amuoou muwsvm OucH pmauommcmuuv mucaoo wwm new muHsmmu ummu huHmcwwoBo: vow huHHmahoz .mm oHan

168

 

 

 

 

 

 

mmm.o n m .cwm.o n oHumHumum m xomluumHuumm
woamHum> mo thmcuwoeom
om.0AaAoo.o o~c.01 www.o Hem.01 om.OAa Nao.01 wqq.o qu.01 «mm
om.OAaAo¢.o www.01 mun.o ome.on oa.OAa ooo.o wom.o «No.9 Hmm
huHHHnmnoum Nmu «mm mm muHHHnmnoum Hmu me Hm some
HHwHIN HHmHIN
meOuusx mmmcsoxm
suHHmeuoz

 

.m ocmum cHnuHs momuu uHm ammHmn oau
Scum Amuoou mumovm oucH omeuommcmuuv muasoo wwo mom muHsme ummu huHocowoaoz vow huHHmauoz .OHh oHan

169

 

mew.o n m .mHm.o u UHumHumum m xomluumHuHmm

mmucmHHm> molwuHmcwwoeom

 

 

 

 

 

om.oAaAoq.o www.o1 mom.o Hmm.01 oh.OAaAow.o mam.o com.o Hmo.o m
ow.OAaAom.o noN.OI qnm.o mHH.OI ow.OAaAom.o mm~.o Hmm.o coo.o e
om.0AAAo<.o nam.01 omn.o mmo.o1 ow.OAaAoo.o cwH.o mwm.o meo.o m
OH.OAaAoN.o mco.Hu wme.o wsH.Hu om.oxaAoq.o mem.o omm.o Nom.o N
om.OAaAoe.o mqo.H| wo~.o How.01 ow.OAaAom.o qu.o1 mmm.o mmo.o1 H
w w
zuHHHnmnoum Nmu N m mm zuHHHnmnoum Hmu H m Hm vamum
HHmHIN HHmHIN
mHmouusx mmo:3mxm
muHHmauoz

 

.umouom HmCOHumz mamuuo on» cH mocmum m>Hm :H mwmuu uHm ammHmn
Scum Amuoou mumscm cuCH omEMOMmsmuuv mucsoo www pom muHammu ammo auHmcwwoso; use huHHmshoz .HHm mHAMH

170

 

HHm.o u m .owm.o u UHumHumum m NomluumHuHmm
mmocmHum> mo thmcwwoaom

 

 

 

 

 

om.oAdAoo.o mHo.on mmm.o Hmm.o1 om.OAaAoc.o OHo.o Hs~.o moH.o ummz
oe.OAaAom.o Hea.on omq.o mam.ou mo.oAaAOH.o msa.H mq~.o ems.o susom
om.0AaAce.o Hma.01 mmq.o www.cn OH.0AaAo~.o coo.H m¢~.o wmm.c ummm
oa.oAa ooo.ou mom.o smo.o1 oe.CAaAon.o omm.o mm~.o wcH.ou nuuoz
zuHHHnmnoum mmu Nwm Nw huHHHnmnoum Hmu me Hm ucmuvmao
HHmHIN HHmHIN
mHmOUHDM mmw§w¥m
suHHmsnoz

 

.momuu mosuam mans aquHs mucmuomso umo3 one .ummm .zuaom .cuuoz
Scum Amuoou museum oucH ooEMOWmcmuuv mucaoo wwm Mom wuHammH ummu muncoonon one huHHmauoz .NHm mHQMH

171

 

an.o u m .No~.o u UHumHumum a xomnuumHunmm

mou:MHum> mQINuHmcowoaom

 

 

 

 

 

o~.oAaAom.o mmH.HI NHe.o qu.o1 oq.oAaAom.o Hms.o mo~.o an.o swan:
om.OAaAoo.o mum.01 o~¢.o me~.ou OH.OAaAoN.o Nom.H oHN.o wmm.o oHosz
oo.OAaAo~.o HNe.o1 mmm.o qmm.0| o~.oAaAom.o NmH.H HmN.o me.o um30H
w w
zuHHHnmnoum Nmu N m mm huHHHnmnoum Hmu. H m Hm snumuum
HHNHIN HHmHIN
mHm0uu=x mmmasoxm
quHmauoz

 

.mowuu moauam ouch cHnuHB mumuum woman can .mHooHE..Hm30H
Scum Amoco» mumswm ouaH omEHOchmuuv mussoo wwo Mom wuHsmmH umwu NUHodwwOEos was huHHmEuoz .MHm oHan

172

 

000.0 n 0 .000.0 - 000000000 0 000-000H0000

 

moosmHum> mo muHocmmwaom

 

 

 

 

 

00.0 0 00.0 000.0- 000.0 000.0- 00.0 0 00.0 00H.H 000.0 000.0 000
00.0 0 00.0 000.0- 005.0 0H0.0- 05.0 0 00.0 0H0.0 000.0 5NH.0 0003 00000
00.0 0 00.0 0H0.H- 055.0 005.0- 00.0 0 00.0 000.H 000.0 0H0.0 00000 00000
00.0 0 05.0 000.0- 005.0 000.0- 00.0 0 00.0 00H.H 000.0 000.0 0000 00000
00.0 0 00.0 0H~.0- 005.0 05H.0- 00.0 0 00.0 000.0- 000.0 H00.0- 00002 00000
00.0 0 00.0 000.0- 055.0 000.0- 00.0 0 00.0 000.0 000.0 005.0 000: 0H000z
00.0 0 000.0 005.0 000.0 0H.0 0 00.0 5H0.H 000.0 500.0 00000 0H00Hz
00.0 0 05.0 000.0- 005.0 000.0- 00.0 0 00.0 005.0 000.0 005.0 0000 0H000z
00.0 0 05.0 000.0- 000.0 000.0- 00.0 0 00.0 005.0- H00.0 500.0- 00002 0H000z
00.0 0 050.0- 000.H 000.0- 00.0 0 00.0 005.0- 500.0 000.0- 0003 00300
00.0 0 00.0 00H.0- 000.0 00H.0- 00.0 0 00.0 050.0 000.0 000.0 00000 00300
00.0 0 00.0 000.0- 0H0.0 000.0- 00.0 0 00.0 000.H 550.0 000.0 0000 00000
00.0 0 00.0 05H.0- 000.H 000.0- 00.0 0 00.0 00H.H 500.0 005.0 00002 00300
0
000H0000000 N00 0 0 N0 000H0000000 H00 H00 H0 HH00
H000-5 HH0s-0
mHmOuHDM wmmgmxw
000H0anoz
.mmmuu moauam ouHss cHnqu mHHmo MH
Eouw Amuoou oumsvm ouaH o0a00mmsmuuv mucaou wwm MOM muHsmmu umou zuHmsmwoeo: can huHHmapoz .qu mHan

173

 

 

 

 

 

000.0 u 0 .0H~.0 u 000000000 0 000-000H0000
000:0Hum> mo muHmawwoaom
om.0AaAoq.o mwm.01 man.o own.01 00.0AaAom.o 0am.o aoq.o HoH.o was
o~.0AaAom.o mno.HI mmn.o an.OI ow.0AaAom.o HMH.o wwm.o Hmo.o Hmz
muHHHnmnoum ~00 Nwm mm huHHHnmnoum Hmu Hmm Hm muse
HHmHIN HHmHIN
mHmouuam mmossmxm

 

000Hmauoz

 

scum Amuoou 000500 oucH omauommcmuuv

.H ocmum cHsuHa momuu movuam muHss 030
mucaoo wwm How muHammu umou muHocmwoaoz can auHHmehoz .mHm oHQMH

174

 

 

 

 

 

000.0 u 0 .000.0 u 000000000 0 000-000H0000
mmocmHum> mo memcwwoaom
o~.0AnAom.o mmH.H1 an.o moo.HI om.0AaAoo.o wm©.o w00.o . mm~.o ~m3
om.OAa ome.ou www.o mmo.o1 0H.0AnAom.o mom.H omn.o wmq.o Hm:
w
auHHHnmnoum Nmu Nwm mm 0uHHHnmnoum Hmu H m Hm moufi
HHmHIN HHNHIN
mHmouuax mmwcsmxm

 

00HH05002

 

Scum Amuoou mumsvm oucH omeuowmcmuuv

.N oamum cHnuHB moon» moauam ouHSB o3u
mucsoo was How muHsmwu umou huHmcwwoaon one auHHmshoz .lo mHan

175

 

050.0 u m .0H0.0 u 000000000 0 0n00-000H0000

 

moocmem> mo huHocuwmaom

 

 

 

 

00.0A0000.0 000.H- 005.0 H05.0- 00.0A0 000.0- 050.0 0H0.0- ~03
0H.0A000~.0 000.H 000.0 00~.H 0H.0A000~.0 0H0.H- H00.0 050.0- H03
mHuHHnmnoum Nmu Nwm mm huHHHnmnoum Hmu me Hw coma
H000-0 H000-0
mHmouusm mmocsoxm
00HH0suoz

 

Scum Amoco» mumavm oucH omeuommcmuuv

.m vamum cHnuH3 woman oceans mqua oau
mucsoo wwo How muHsmou umou auHmcwwoso: cam zuHHmahoz .mHm oHamH

176

 

oc~.o

m .005.H u 000000000 0 mom-000H0000

 

mmocmHum> mo hUHmcowoaom

 

 

 

 

om.oAaAo0.o H0O.H Non.o Hmn.o OH.0AaAom.o moo.H nmm.o oom.o mmz
0H.000005.0 000.H 050.0 000.0- 00.000000.0 000.0 000.0 000.0 H03
zuHHHnmnoum N00 Nwm mm huHHHnmnoum Hmu me Hm mouw
HHmHIN HHMHIN
mHmouu=M mmmsawxm
kuHHmahoz

 

Scum AquOH mumscm oucH awesommamuuv

.0 ocmum cHsuHB mowuu museum muHss oBu

mucaoo wwm How muHamou umwu muHosmwoaon cam huHHmshoz

.me mHama

177

 

 

 

 

 

000.0 u 0 .000.0 n 000000000 0 0mom-0003.000
mmucmHum> mo hUHmcowoaom
OH.OAaAoN.o 000.H mHa.o mom.H no.OAaAOH.o 0mm.H mnq.o mH¢.o Nm3
ON.OAaAom.o NNH.HI mac.o om~.ou ow.oAaAow.o NHm.o 0mm.o mmH.o Hmz
w
huHHHnmpoum mmu N 0 mm huHHHnmnoum Hmu me Hw some
HHMHIN HHMHIN
meouusx mmocsmxm

 

zuHHmshoz

 

Scum Amuoou mumsam oucH owEhommcmuuv

.m ocmum cHnuHs momma museum mana 030
mucooo wwm now muHammu ummu huHonwoao: use huHHmauoz .mHm mHan

178

 

Amvmoo.o n m .mmo.q n ugumaumum m xomnuuofluumm

muonmaum> mo hufimammoaom

 

 

 

 

 

om.oAaAoe.o oeo.cu o~m.o onm.on OH.OAaAo~.o mem.a aw~.o «mm.o m
om.oAaAo¢.o «mm.ou oom.o mhc.ou OH.OAaAo~.o Hom.H Hm~.o qu.o q
ow.oAaAom.o mma.o «hm.o mmo.o om.OAaAoq.o omm.ou Hm~.o mq~.on m
oo.oAaAo~.o Nm¢.o: Hmm.o aq~.ou om.OAaAom.o NoN.H Hw~.o 0mm.o N
oH.oAaAo~.o mmq.H- oom.o q~m.ou cu.OAaAow.o mmm.o ~w~.o Noa.o H
w
zugflfinmnoum Nmu N m mm muHHfiamDoum Hm“ me Hm vamum
Hfimaum Hameum
maneuusx mmmnamxw
3:3qu

 

.uwwuom Hchfiumz mamuuo wnu ca mvcmum m>wm ca mwmuu woauam mugs:
Baum Amuoou mumsvm ouufi wwahommamuuv muasou www How muasmmu umwu hufimamwoab: vcm hufiamahoz .omm maan

179

 

moh.o u m .mqa.o u uaumfiumum m nomnuumfiuumm
mwofimfi Hm> mo NUHMwaOEO—w

 

 

 

 

 

0H.OAaAom.o mmq.al oom.o «mm.OI oh.OAaAow.o mmm.o nwm.o NOH.o muaunm muwnz
om.OAaAoq.o mco.H| won.o How.01 ow.OAaAO¢.o mqa.on mom.o wmo.OI Ham Emmamm
kuwaanmnoum Nmu Nwm mm hufiawnmnoum Hmu me aw meumam mmue
HHmHIN HHMHIN
mHmouHSM mmmc3mxm
huHHmEhoz

 

.H wamum cfinufis mmmuu moauam muwsa o3u cam Ham ammamn Cay
Scum Amuoou muwavm oucfi vaHOMmcmuuv muaaoo wwm How muasmmu umwu hufimcmwoaon vcm %uaamauoz .Hmm mHan

180

 

qm~.o u m .an.H u ofiumfiumum m xomnuumauumm
mmucmfium> mo muamamwoaom

 

 

 

 

 

oe.OAath.o ~mq.o- Ham.o me~.o- ON.OAaAom.o NON.H Hw~.o omm.o monuam mafia:
OH.0AaAo~.o aoc.HI wwo.o wea.au om.OAaAoq.o mow.o omm.o Nom.o Ham ammamm
kufiawnmnoum Nmu Nwm mm hufiHHnmnoum Hmu me Hm mmfiumam owua
HHmHIN HfimHIN
mamouuax mmmaamxm
zuwamauoz

 

.N mufimum GHSUflQw mmmhu UUSHQm QHH£3 O3“ vflm HHM ngNn— 03“

Scam Amuoou mumsvm oucw vaMOMmcmuuv muasoo wwm How muHSmmH ummu huamamwoaos can kuwamauoz

.NNm «Hana

181

 

Amvooo.o u m .omN.MH u uaumfiumum m xomuuumauumm

mmucmfium> mo huamamwoaom

 

 

 

 

 

om.OAaAom.o mma.o cum.o mwo.o om.0AaAoq.o www.0I Hmm.o m¢~.01 mosumm mufinz
om.oAnAoc.o 5mm.OI omn.o mno.ol om.OAaAoo.o owH.o mwm.o ooo.o Ham ammamm
huHHHnmnoum Nmu Nwm mm huaawnmnoum Hmu me Hm wmwumam mmuH
HHQHIN HHMHIN
mfimouusx mmmaswxm
huwamahoz

 

.m vamum away“: mwwuu mosuam muwn3 oau can Ham Banana 03u
Eouw Amuoou mumavm oucfi vmeu0mmcmuuv mucsou wwm How muddmmu ummu huwwcmmoEon cam kufiHmEHOZ .mmm wanna

182

 

cam.c n m .mmq.o u oaumaumum m xomnuumauumm
mmocmfium> mo Nwamqmmoaom

 

 

 

 

 

om.0AaAoq.o cmm.on oom.o whq.ol CH.OAon~.o Honda Hmm.o qu.o monumm mufinz
om.oAaAca.o nom.ou cum.o mHH.ou ow.OAaAoa.o -~.o Hm~.o ooo.o “an ammamm
huwflfinmnoum Nmu Nwm mm mufiafinmnoum Hmu me Hm mmfiuwmm wmue
HfidHiN HHMHIN
mfimouusm mmmaamxm
%ufiamahoz

 

.e wcmum cfinufia wwwuu wuauam mufiLB oBu can Ham Banana oBu
Eoum Amuoou mumavm oucH wmshowmcmuuv mucsoo wmm How muaammu ummu >uwmamono: cam huflamshoz .qmm mHAmH

183

 

cmo.o u m .Hno.m n oflumfiumum m xomnuumauumm
mwu:MHum> mo hwwmamwmaom

 

 

 

 

 

om.OAaAoo.o oom.on cum.o onm.0| oa.oAnAo~.o mom.H mm~.o qmm.o monumm mugs:
om.0AnAoc.o cmm.01 ¢mm.o Hmm.01 on.OAaAow.c mmm.o com.o H¢o.o “Hm ammawm
hufiafinmnoum Nmu Nwm mm hufiawnmnoum Hmu me aw mwfiomnm mmue
HHmHIN HfiMHIN
mamouuax mmmasmxm
%uHHmauoz

 

.m Vanum cwnuHB mwmuu woauam wuass oBu van Ham ammamp o3»
scum Amuoou mumsvm oucw nmfiMOchmuuv mucsoo wmm How muaammu uwmu hufimawwoao: vam huwamahoz .mmh magma

184

 

Aqumo.o n m .eam.q n afiumaumum m xomuuumauumm

 

mmusmfium> mo huwmawwoaom

 

 

 

 

o~.oAqum.o eo~.H- qm~.o HNm.on Amvmo.oAnAmo.o ¢mH.~ “NH.o m-.o wuauam mafia:
Amvmo.OAaAmo.o «mo.ml oam.o mqo.o: om.0AnADo.o mmo.o mmH.o moo.o Ham ammamm
mafiafinmnoum Nmu Nwm mm muwafinmnoum Hmu Hmm Hm mmaowam mmuH
Hfimylm HwMHIN
mwmouusx wwwcsmxm
huwamahoz

 

.ummuom HMcOHumz mBmuuo mSu sfinufia mv:Mum m>Hm cw mmmuu monumm muwcs :mu was Ham EmmHmn amu

aoum Amuoou wumavm oucH vmahowmcmuuv muaaoo wwm How muasmmu ummu huamcwwoao: cam hufiamsuoz

.omm wHQMH

APPENDIX G .

185

APPENDIX G.

Number of eggs per mass per cell
Four egg masses were randomly chosen from four branches that had
resided in one of the predesignated cells. As reported in the text,
the egg masses were measured and the number of eggs were estimated using
Leonard's et al. (1973) technique. Table Gl reports these counts. A
key for the headings and units is provided at the end of the table (0
eggs per mass always indicates that there was not an egg mass measured

for that cell).

186

Table 61.
tree.

Distribution of individual egg counts within each sampled

STD SPP TREE QUADRANT STRATUM #EGGS/D ASS
1 BE 1 LOWER NORTH -
1 BF 1 LOWER NORTH -
1 BF 1 LOE-‘JER NORTH -
1 BF ] LOZIER NORTH -
1 BF 1 LOWER EAST 35
1 BE" 1 LOWER EAST -
1 BE 1 LOWER EAST -
1 BF ] LOWER EAST -
1 BE ] LOWER SOUTH -
1 BF 1 LOWER SOUTH -
1 BF 1 LOWER SOUTH -
1 BF 1 LOWER SOUTH -
1 BE 1 LOWER W1“... T 09
1 BF ' 1 LOWER WEST -
] BE 1 LOWER WEST -
1 BF ] LOWER WEST -
1 BE 1 HI DDLE NORTH -
1 BF 1 MI DDLE NORTH -
1 BF 1 MI DDLE NORTH —
1 BF 1 MI DOLE NORTH -
1 BE 1 MI DDLE EAST -
1 Bk 1 MI DDLE EAST -
1 BF 1 MI DDLE EAST -
1 BF 1 MIDDLE EAST -
1 BF 1 MIDDLE SOUTH
1 BF ] MIDDLE SOUTH {49
1 BF 1 MIDDLE SOUTH 5
1 Bl“ 1 MIDDLE SOUTH -
1 BF 1 MIDDLE WEST -
1 BF 1 MIDDLE WEST -
1 BF 1 11 DDLE WEST -
1 BE 1 MI DDLE WE S‘I‘ -
1 BE 1 UPPER NORTH 29
1 BE 1 UPPER NORTH -
1 BF 1 UPPER NORTH -
1 BF 1 UPPER NORTH -
1 BF 1 UPPER EAST -
1 BF 1 UPPER EAST -
1 BF 1 UPPER EAST -
1 BF 1 UPPER EAST -

187

Table Gl. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
1 BF 1 UPPER SOUTH 32
1 BF 1 UPPER SOUTH
1 BF 1 UPPER SOUTH —
1 BF 1 UPPER SOUTH -
1 BF 1 UPPER WEST -
1 BF 1 UPPER WEST -
1 BF 1 UPPER WEST -
1 BF 1 UPPER WEST -
1 BF 1 TOP TOP —
1 BF 1 TOP TOP -
1 Bk 1 TOP TOP -
1 LE 1 ‘POP ‘fOP -
1 BE 2 LOWER NORTH -
1 PF 2 LOWER NORTH -
1 3b 2 LOWER [‘JORTh -
1 BF 2 LOWE R NORTrl -
1 BF 2 LOWER EAST 12
1 BF 2 LOWER EAST 12
1 BF 2 LOWER EAST -
1 BF 2 LOWER EAST -
1 EF 2 LOWER SOUTH 5%
1 BF 2 LOWER SOUTH 29
1 BF 2 LOWER SOUTH -
1 BF 2 LOWER SOUTH -
1 BF 2 LOWER WEST -
1 BF 2 LOWER NEST -
1 BE 2 LOWER WE"? -
1 BF 2 LOWER WEST -
1 BF 2 MIDDLE NORTH 26
1 BE 2 HIDDLE NORTH 25
1 BF 2 MIDDLE NORTH 12
1 BF 2 MIDDLE NORTH 26
1 BF 2 MIDDLE EAST 32
1 BE 2 MIDDLE EAST 35
1 BF 2 MIDDLE EAST 26
1 BF 2 MIDDLE EAST 26
1 BF 2 MIDDLE SOUTH 19
1 BF 2 MIDDLE SOUTH -
1 BF 2 MIDDLE SOUTH -
1 BF 2 MIDDLE SOUTfi -

188

Table Gl. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
1 BF 2 MIDDLE WEST 1 2
1 BF 2 MIDDLE WEST 32
1 BF 2 MIDDLE WEST 12
1 BF 2 MIDDLE WEST 29
1 BF 2 UPPER NORTH 15
1 BF 2 UPPER NORTR 38
1 BF 2 UPPER NORTH —
1 BF 2 UPPER NORTH -
1 SF 2 UPPER EAST 29
1 BF 2 UPPER EAST 17
1 BF 2 UPPER EAST -
1 BE 2 UPPER EAST -
1 B 2 UPPER SOUTH 46
1 BF 2 UPPER SOUTh 24
1 BF 2 UPPER SOUTH 38
1 BF 2 UPPER SOUIJ 32
1 BF 2 UPPER WEST 41
1 BF 2 UPPER WEST -
1 BF 2 UPPER WEST -
1 BF 2 UPPER WEST -
1 BF 2 TOP TOP 2%
1 BF 2 TOP TOP 22
1 BF 2 TOP TOP 20
1 BF 2 TOP TOP 17
1 BF 5 LOWER NORTH -
1 BF 5 LOWER NORTH -
1 BF 5 LOWER NORTH -
1 BF 5 LOWER NORTH —
1 BF 5 LOWER EAST -
1 BF 5 LOWER EAST -
1 BF 5 LOWER EAST -
1 BF 5 LOWER EAST -
1 BF 5 LOWER SOUTH 26
1 BF 5 LOWER SOUTR -
1 BF 5 LOWER SOUTH -
1 BF 5 LOWER SOUTH -
1 BF 5 LOWER WEST -
1 BE 5 LOWER WEST -
1 BF 5 LOWER WEST -
1 BF 5 LOWER WEST -

189

Table GI. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
I BF 5 MIDDLE NORTH -
I BE 5 MIDDLE NORTH -
I BF 5 MIDDLE NORTH -
I BF 5 MIDDLE NORTH -
I BE 5 MIDDLE EAST -
I BE 5 MIDDLE EAST -
I BF 5 MIDDLE EAST -
1 BF 5 MIDDLE EAST -
I BE 5 MIDDLE SOUTH I 3
I BE 5 MIDDLE SOUTH -
I BF 5 MIDDLE SOUTH -
I BE 5 MIDDLE SOUTH -
I BE 5 MIDDLE WEST I 7
1 BF 5 MIDDLE WEST 20
I BE 5 MIDDLE WEST 18
I BF 5 MIDDLE NEST 22
I BE 5 UPPER NORTH 35
I BF 5 UPPER NORTH -
I BF 5 UPPER HORTH -
I BE 5 UPPER NORTH -
I BE 5 UPPER EAST -
I BE 5 UPPER EAST -
I BF 5 UPPER EAST -
I BF 5 UPPER EAST -
1 BF 5 UPPER SJUTh 42
1 BF 5 UPPER SOUTH 22
I BE 5 UPPER SOUTH IO
1 BF 5 UPPER SOUTH -
1 BF 5 UPPER WEST I7
I BF 5 UPPER WEST -
I BF 5 UPPER WEST -
I BF 5 UPPER WEST -
I BF 5 TOP TOP -
1 BF 5 TOP TOP -
I BF 5 TOP TOP -
I BF 5 TOP TOP -
I BE‘ 6 LOWER NORTH I 9
I BF 6 LOY-JER NORTH -
I BF 6 LOWER NORTH -
I BF 6 L01 E R NORTH -

190

Table Gl. Continued.
STD SPP

HHHHl—JHHL—JHHL—Il—‘HHHHHHI—JHHHHHL-JI—It—Jb—Il—JL—IL—ll—‘i-JI—Jl—IL—ll—ll—it—IH

mmmoommmmomo‘Gama-mommamommmommmmmmooomooom

LOWER
LO'» TE R
LOWER
LOWER
LOWER
LOWER
LOWER
LOWE R
LOWER
LOWER
LOWER
LOWER
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
MIDDLE
UPPER
UPPER
UPPER
UPPER
UPPER
UPPER
UPPER
UPPER
UPPER
UPPER
UPPER
UPPER

EAST
EAST
EAST
EAST
SOUTH
SOUTH
SOUTH
SOUTH
WEST
WEST
WEST
WEST
NORTH
NORTH
NORTH
NORTH
EAST
EAST
EAST
EAST
SOUTH
SOUTH
SOUTH
SOUTd
WEST
WEST
WEST
WEST
NORTH
NORTH
NORTH
NORTH
EAST
EAST
EAST
EAST
SOUTH
SOUTH
SOUTH
SOUTH

32
32
26
42
Q9
23
22

29

191

Table 61. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
I BE 6 UPPER WEST I3
1 BE 6 UPPER NEST G9
1 BF 6 UPPER WEST G9
1 BF 6 UPPER WEST IO
1 BF 6 TOP TOP -
1 BF 6 TOP TOP -
1 BF 6 TOP TOP —
1 BF 6 TOP TOP -
2 BF 1 LOWER NORTH 19
2 BF 1 LOWER NORTH -
2 BF 1 LOWER NORTH -
2 BF 1 LOWER NORTH -
2 BF I LOWER EAST -
2 BF 1 LOWER EAST -
2 BE 1 LOWER EAST -
2 Bf 1 LOWER EAST -
2 BF 1 LOWER SOUTH -
2 BF 1 LOWER SOUTH -
2 EF 1 LOWER SOUTH -
2 BF 1 LOWER SOUTH -
2 BF 1 LOWER WEST -
2 BF 1 LOWER WEST -
2 BF 1 LOX JER WEST -
2 BF I LOi‘JER NEST -
2 BE 1 MIDDLE NORTH -
2 BF 1 MIDDLE NORTH -
2 BF 1 MIDDLE NORT} -
2 BF 1 MIDDLE NORTH -
2 BF 1 MIDDLE EAST -
2 BF I MIDDLE EAST -
2 BF 1 MIDDLE EAST -
2 BF 1 MIDDLE EAST -
2 BF 1 MIDDLE SOUTH 15
2 BF 1 MIDDLE SOUTH 13
2 BF 1 MIDDLE SOUTH -
2 BF 1 MIDDLE SOUTH -
2 BF 1 MIDDLE WEST 10
2 BF 1 MIDDLE WEST 32
2 BF 1 MIDDLE WEST 17
2 BF 1 MIDDLE WEST 12

192

Table 61. Continued.

2 BF 1 UPPER NORTH 20
2 BF 1 UPPER NORTH 17
2 BF 1 UPPER NORTH 22
2 BF 1 UPPER NORTH 19
2 BF 1 UPPER EAST 38
2 BF 1 UPPER EAST 15
2 BE 1 UPPER EAST -
2 BF 1 UPPER EAST -
2 BF 1 UPPER SOUTH 38
2 BF 1 UPPER SOUTH 13
2 BF 1 UPPER SOUTH -
2 BF 1 UPPER SOUTH 13
2 BF 1 UPPER NEST 35
2 BF 1 UPPER WEST 29
2 BF 1 UPPER WEST 26
2 BF 1 UPPER WEST 17
2 BF 1 TOP TOP 12
2 BE 1 TOP TOP 12
2 BF 1 TOP TOP 32
2 BF 1 TOP TOP 13
2 BF 2 LOWER NORTH -
2 BF 2 LOEER NORTH -
2 BF 2 LOWER NORTH -
2 BF 2 LOWER NORTH -
2 BF 2 LOWER FAST -
2 BF 2 LOWER EAST -
2 Br 2 LOWER EAST -
2 BF 2 LOWER EAST -
2 B 2 LOWER SOUTH -
2 BE 2 LOWER SOUTH -
2 BE 2 LOWER SOUTH -
2 BF 2 LOWER SOUTH -
2 BF 2 LOWER WEST -
2 BE 2 LOWER WEST -
2 BF 2 LOWER WEST -
2 BF 2 LOWER WEST -
2 BF 2 MIDDLE NORTH 29
2 BF 2 MIDDLE NORTH -
2 BF 2 MIDDLE NORTH -
2 BF 2 MIDDLE NORTH -

193

Table Gl. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
2 BF 2 MIDDLE EAST 1 9
2 BF 2 MIDDLE EAST 29
2 BF 2 MIDDLE EAST 1 9
2 BF 2 MIDDLE EAST -
2 BF 2 MIDDLE. SOUTH 17
2 BF 2 MIDDLE SOUTH 38
2 BF 2 MIDDLE SOUTH 35
2 BF 2 MIDDLE SOUTH 17
2 BF 2 MIDDLE WEST 2
2 BF 2 MIDDLE WEST 46
2 BF 2 1‘11 DDLE WEST 12
2 Br 2 MIDDLE NEST -
2 BE 2 UPPER NORTH 35
2 PF 2 UPPER NORTH 24
2 BF 2 UPPER NORTH -
2 BF 2 UPPER NORTH -
2 BF 2 UPPER EAST 26
2 BF 2 IJPPEHZ .EAST? 12
2 BF 2 UPPER EAST 13
2 PF 2 UPPER EAST 17
2 EF 2 UPPER SOUTH $9
2 BF 2 UPPER SOUTH -
2 BF 2 UPPER SOUTH -
2 BF 2 UPPER SOUTH -
2 BF 2 UPPER NEST 32
2 BF 2 UPPER WEST 32
2 BF 2 UPPER WEST 17
2 BF 2 UPPER ”EST 38
2 BF 2 TOP TOP -
2 BF 2 TOP TOP -
2 BF 2 TOP TOP -
2 BF 2 TOP TOP -
3 BF 1 LOWER NORTH -
3 BF 1 LOWER NORTH —
3 BF 1 LOWER NORTH -
3 BF 1 LOWER NORTH -
3 BF 1 LOWER EAST -
3 BF 1 LOWER EAST -
3 BF 1 LOWER EAST -
3 BF 1 LOWER EAST -

194

Table G1. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
3 BF I LOWER SOUTH -
3 BF I LOWER SOUTH -
3 BF 1 LOWER SOUTH -
3 BF 1 LOWER SOUTH -
3 BE 1 LOWER WEST -
3 BF 1 LOWER WEST -
3 BE 1 LOWER WEST -
3 BF 1 LOWER WEST -
3 BF I MIDDLE NORTH 32
3 BE I MIDDLE NORTH 46
3 BF 1 MIDDLE NORTH 19
3 BF 1 MIDDLE NORTH -
3 BE I MIDDLE EAST 46
3 BE 1 MIDDLE EAST -
3 BE I MIDDLE EAST —
3 SE 1 MIDDLE EAST -
3 BF 1 MlDDLE SOUTH -
3 LP 1 MIDDLE SOUTH -
3 EP 1 MIDDLE SOUTH —
3 PF 1 MIDDLE SOUTH -
3 BF I MIDDLE WEST 35
3 BE 1 MIDDLE WEST 10
3 BE 1 MIDDLE WEST 29
3 bF I MIDDLE WEST -
3 BF 1 UPPER NORTH 22
3 BF 1 UPPER NORTH 35
3 BF I UPPER NORTH -
3 BF I UPPER NORTH -
3 BF I UPPER EAST 29
3 BF I UPPER EAST 35
3 BF 1 UPPER EAST 38
3 BF 1 UPPER EAST @9
3 BE 1 UPPER SOUTH 15
3 BF 1 UPPER SOUTH I7
3 BE I UPPER SOUTH G9
3 BE 1 UPPER SOUTH 29
3 BF 1 UPPER WEST 29
3 BE I UPPER WEST 32
3 BE 1 UPPER WEST 23
3 BF 1 UPPER WEST 12

195

Table G1 . 9311333322: ______________________________________
STD SPP TREE QUADRANT STRATUM #EGGS/MASS
3 BF 1 TOP TOP -
3 BF I TOP TOP -
3 BF 1 TOP TOP -
3 BF 1 TOP TOP -
3 BF 2 LOWER NORTH 22
3 BF 2 LOWER NORTH -
3 BF 2 LOWER NORTH -
3 BF 2 LOWER NORTH -
3 BF 2 LOWER EAS '1‘ -
3 BF 2 LOWER EAST -
3 BF 2 LOWER EAST -
3 BF 2 LOWER EAST -
3 BF 2 LOWER SOUTH -
3 BF 2 LOWER SOUTh -
3 BF 2 LOWER SOUTH -
3 BF 2 LOWER SOUTH -
3 BF 2 LOWER WEST 59
3 BE 2 LOWER WEST -
3 BE 2 LOWER WEST -
3 BF 2 LOWER WEST -
3 BF 2 HIDDLE NORTH -
3 BF 2 MIDDLE NORTH -
3 BE 2 MIDDLE NORTH -
3 BF 2 MIDDLE NORTH -
3 BF 2 MIDDLE EAST 18
3 BF 2 MIDDLE EAST -
3 BF 2 MIDDLE EAST -
3 BF 2 MIDDLE EAST -
3 BF 2 MIDDLE SOUTH 22
3 BF 2 MIDDLE SOUTH 09
3 BF 2 MIDDLE SOUTH 50
3 BF 2 MIDDLE SOUTH 26
3 BF 2 MIDDLE WEST 32
3 BF 2 MIDDLE WEST Q9
3 BF 2 MIDDLE WEST I7
3 BF 2 MIDDLE WEST 20
3 BF 2 UPPER NORTH -
3 BF 2 UPPER NORTH -
3 BF 2 UPPER NORTH -
3 BF 2 UPPER NORTH -

196

Table Gl. Continued.

3 BF 2 UPPER EAST 15
3 BF 2 UPPER EAST 17
3 BF 2 UPPER EAST 32
3 BF 2 UPPER EAST 17
3 BF 2 UPPER SOUTH 26
3 BF 2 UPPER SOUTH -
3 BF 2 UPPER SOUTH -
3 BF 2 UPPER SOUTH -
3 BF 2 UPPER WEST 22
3 BF 2 UPPER WEST -
3 BF 2 UPPER WEST -
3 BF 2 UPPER WEST -
3 BF 2 TOP TOP -
3 BF 2 TOP TOP -
3 BF 2 TOP TOP -
3 BF 2 TOP TOP -
4 BF 1 LOWER NORTH -
4 BE ] LOWER NORTH -
4 BF 1 LOWER NORTH -
4 BF 1 LOWER NORTH -
4 BF 1 LOWER EAST 35
4 BF 1 LOWER EAST -
4 BF 1 LOWER EAST -
4 b5 1 LOWER EAST -
4 BF 1 LOWER SOUTH -
4 BF 1 LOWER SOUTH -
4 BE 1 LOWER SOUTH -
4 BF 1 LOWER SOUTH -
4 BF 1 LOWER WEST 32
4 BE 1 LOWER WEST 12
4 BF 1 LOWER WEST 24
4 BF 1 LOWER WEST -
4 BF 1 MIDDLE NORTH 15
4 BF 1 MIDDLE NORTH 19
4 BF 1 MIDDLE NORTH 22
4 BF ] MIDDLE NORTH 13
4 BF 1 MIDDLE EAST -
4 BF 1 MIDDLE EAST -
4 BF 1 MIDDLE EAST -
4 BF 1 MIDDLE EAST -

197

Table GI. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS

4 BF 1 MIDDLE SOUTH 26
4 BF 1 MIDDLE SOUTH 29
4 BF 1 MIDDLE SOUTH 19
4 BF 1 MIDDLE SOUTH 24
4 BF 1 MIDDLE WEST 20
4 BF 1 MIDDLE WEST 42
4 BF 1 MIDDLE WEST 27
4 BF 1 MIDDLE WEST 35
4 BF 1 UPPER NORTH 16
4 BF 1 UPPER NORTH 23
4 BF 1 UPPER NORTH 35
4 BF 1 UPPER NORTH 17
4 BF 1 UPPER EAST 24
4 BF 1 UPPER EAST 13
4 BF 1 UPPER EAST G7
4 BF 1 UPPER EAST 29
4 BF 1 UPPER SOUTH 32
4 BF 1 UPPER SOUTH 26
4 BF 1 UPPER SOUTH 26
4 BF 1 UPPER SOUTH 18
4 BF 1 UPPER WEST 29
4 BF 1 UPPER WEST 22
4 BF 1 UPPER WEST 19
4 BF 1 UPPER WEST 22
4 BF 1 TOP TOP 54
4 BF 1 TOP TOP 19
4 BF 1 TOP TOP 10
4 BF 1 TOP TOP 23
4 BF 2 LOWER NORTH -
4 BF 2 LOWER NORTH -
4 BF 2 LOWER NORTH -
4 BF 2 LOWER NORTH -
4 BF 2 LOWER EAST 24
4 BF 2 LOWER EAST 42
4 BF 2 LOWER EAST -
4 BF 2 LOWER EAST -
4 BF 2 LOWER SOUTH -
4 BF 2 LOWER SOUTH -
4 BF 2 LOWER SOUTH -
4 BF 2 LOWER SOUTH -

198

Table 61. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
4 BF 2 LOWER WEST -
4 BF 2 LOWER WEST -
4 BF 2 LOWER WEST -
4 BE 2 LOWER WEST —
4 BF 2 MIDDLE NORTH 32
4 BF 2 MIDDLE NORTH 15
4 DP 2 MIDDLE NORTH -
4 BE 2 MIDDLE NORTH -
4 BF 2 MIDDLE EAST 42
4 BF 2 MIDDLE EAST 19
4 BF 2 MIDDLE EAST 32
4 BF 2 MIDDLE EAST 29
4 BF 2 MIDDLE SOUTH 22
4 BF 2 MIDDLL SOUTH 29
4 BF 2 MIDDLE SOUTH 46
4 BF 2 MIDDLE SOUTH IO
4 SP 2 MIDDLE WEST -
4 BF 2 MIDDLE WEST -
4 BF 2 MIDDLE WEST -
4 BF 2 MIDDLE WEST -
4 BF 2 UPPER NORTH 29
4 BF 2 UPPER NORTH 29
4 BF 2 UPPER NORTH 19
4 BF 2 UPPER NORTH 36
4 LP 2 UPPER PAST 26
4 BF 2 UPPER EAST 29
4 BF 2 UPPER EAST 19
4 BF 2 UPPER EAST 26
4 BF 2 UPPER SOUTH 26
4 BF 2 UPPER SOUTH 26
4 bf 2 UPPER SOUTM 19
4 BF 2 UPPER SOUTH 16
4 BF 2 UPPER WEST 15
4 BF 2 UPPER WEST 26
4 BF 2 UPPER WEST 24
4 BF 2 UPPER WEST 13
4 BF 2 TOP TOP U9
4 BF 2 TOP TOP 46
4 BF 2 TOP TOP 13
4 BF 2 TOP TOP 22

199

Table G1. Continued.

STD SPP TREE QUADRANT STRATUM #BGGS/HASS
5 BF 1 LOWER NORTH -
5 BF 1 LOWER NORT H -
5 BF 1 LOWER NORTH -
5 BE" 1 LOWER NORTH -
5 BF 1 LOWER EAST -
5 BF 1 LOWER EAST -
5 BF 1 LOWER EAST -
5 BF 1 LOWER EAST -
5 BF 1 LOWER SOUTH -
5 BF 1 LO'IJL'R SOUTH -
5 BE 1 LOVER SOUTH -
5 BF 1 LOWER SOUTH —
5 BF 1 LOWER WEST 26
5 BE 1 LOWER WEST -
5 BF 1 LOWER WEST -
5 BF 1 LOWER WEST -
5 BF 1 MIDDLE NORTH 19
5 BF 1 MI DDLE NORTH 1 7
5 PF 1 MIDDLE NORTH 13
5 BF 1 MIDDLE NORTH 26
5 BF 1 MIDDLE EAST 22
5 BF 1 MIDDLE EAST 16
5 BF 1 MIDDLE EAST 22
5 BF 1 MIDDLE EAST 15
5 BF 1 MIDDLE SOUTH 32
5 BF 1 MIDDLE SOUTH 24
5 BF 1 MI DDLE SOUTH 24
5 BF 1 MIDDLE SOUTH 17
5 BF 1 MIDDLE WEST 35
5 BF 1 MIDDLE WEST 32
5 BF 1 MIDDLE WEST 1 9
5 BF 1 MI DDLE WEST 1 3
5 BF 1 UPPER NORTH 17
5 BF 1 UPPER NORTB 67
5 BF 1 UPPER NORTH 10
5 BF 1 UPPER NORTH 69
5 BF 1 UPPER EAST 26
5 BF 1 UPPER EAST 13
5 BF 1 UPPER EAST 26
5 BF 1 UPPER EAST 42

200

Table Gl. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
5 BF 1 UPPER SOUTH 29
5 BF 1 UPPER SOUTH 23
5 BF 1 UPPER SOUTH 13
5 BF 1 UPPER SOUTH E7
5 BF 1 UPPER WEST 10
5 BE 1 UPPER WEST 29
5 BF 1 UPPER WEST 1a
5 EE 1 UPPER WEST 22
5 EE 1 TOP TOP 16
5 BF 1 TOP TOP 22
5 EE 1 TOP TOP -
5 BF 1 TOP TOP -
5 BE 2 LONER NORTH 19
5 R: 2 LOWER WORTH 29
5 RE 2 LOWER WORTH -
5 BF 2 LOWER NORTH -
5 BE 2 LOWER EAST 35
5 BE 2 LOWER EAST _
5 BF 2 LOWER EAST -
5 BF 2 LOWER EAST -
5 DE 2 LOWER SOUTH 23
5 E 2 LOWER SOUTH -
5 BF 2 LOWER SOUTH -
5 LP 2 LOWER SOUTH -
5 BF 2 LOWER WEST 22
5 BF 2 LOWER WEST -
5 RE 2 LOWER WEST -
5 EP 2 LOWER JEST -
5 BE 2 MIDDLE NORTH -
5 BF 2 MIDDLE WORTH -
5 BF 2 MIDDLE NORTH -
5 BE 2 MIDDLE NORTH -
5 BE 2 MIDDLE EAST 29
5 BF 2 MIDDLE EAST 3S
5 BF 2 MIDDLE EAST 15
5 BF 2 MIDDLE EAST 38
5 BE 2 MIDDLE SOUTH 54
5 BF 2 MIDDLE SOUTH -
5 BF 2 MIDDLE SOUTH -
5 EE 2 MIDDLE SOUTH -

201

Table Gl. Continued.

5 BF 2 [‘11 DDLE WEST -
5 BF 2 MIDDLE WEST -
5 BF 2 MI DDLE WEST -
5 BF 2 [~11 DDLE WEST -
5 BF 2 UPPER NORTH 32
5 BF 2 UPPER NORTH 26
5 BF 2 UPPER NORTH 1 {71
5 BF 2 UPPER NORTH 22
5 BF 2 UPPER EAST 1 3
5 BF 2 UPPER EAST 46
5 BF 2 UPPER EAST 38
5 BF 2 UPPER EAST 35
5 BE 2 UPPER SOUTti 26
5 BF 2 UPPER SOUTH 1 2
5 BF 2 UPPER SOUTH 32
5 Br‘ 2 UPPER SOUTH 35
5 BF 2 UPPER WEST 1 2
5 BF 2 UPPER WEST -
5 BE 2 UPPER WEST -
5 BE 2 UPPER WEST -
5 BF 2 TOP TOP 29
5 BF 2 TOP TOP 24
5 BF 2 TOP TOP 20
5 BF 2 TOP TOP 1 5
1 NS 1 LOWER NORTH 43
1 NS 1 LOWER NORTH 33
1 NS 1 LOWER NORTri -
1 WS 1 LOWER NORTH -
1 W8 1 LOWER EAST 35
1 1'18 1 LONE R EAST 2 5
1 WS 1 LOWER EAST 1 8
1 ws 1 LOWER EAST -
1 WS 1 LOWER SOUTH 27
1 NS 1 LOWER SOUTH O9
1 NS 1 LOWER SOUTH 29.5
1 HS 1 LOWER SOUTH -
1 NS 1 LOWER WEST -
1 L}! S 1 LOWE R WEST -
1 WS 1 LOWER WEST -
1 WS 1 LOWER WEST -

202

Table 61. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
1 US 1 MIDDLE NORTH 25
1 US 1 MIDDLE NORTH 22
1 US 1 MIDDLE NORTH -
1 NS 1 MIDDLE NORTH -
I ws 1 MIDDLE EAST 36
1 HS 1 MIDDLE EAST 32
1 RS 1 MIDDLE EAST 43
1 US 1 MIDDLE EAST 28
1 NS 1 MIDDLE SOUTH 13
1 US 1 MIDDLE SOUTH U9
1 US 1 MIDDLE SOUTH 2m
1 HS 1 MIDDLE SOUTH 15
1 Us 1 MIDDLE WEST 22
1 HS 1 MIDDLE WEST 18
1 US 1 MIDDLE WEST 13
1 WS 1 MIDDLE WEST 13
1 JS 1 UPPER NORTH 39
1 HS 1 UPPER NORTH 32
1 NS 1 UPPER RORTH 27
1 NS 1 UPPER NORTH 25
1 WS 1 UPPER EAST 11
1 us 1 UPPER EAST 22
1 W5 1 UPPER EAST 2D
1 NS 1 UPPER EAST 11
1 US 1 UPPER SOUTH 31
1 US 1 UPPER SOUTH 36
1 US 1 UPPER SOUTH 23
1 HS 1 UPPER SOUTH I3
1 NS 1 UPPER WEST 22
1 WS 1 UPPER WEST 15
1 US 1 UPPER WEST 18
1 US 1 UPPER WEST -
1 US 1 TOP TOP -
1 WS 1 TOP TOP -
1 WS 1 TOP TOP —
1 ws 1 TOP TOP —
1 WS 2 LOWER NORTH 15
1 WS 2 LOWER NORTH -
1 US 2 LOWER NORTH -
1 US 2 LOWER NORTH -

203

Table G1. Continued.

SID SPP TREE QUADRANT STRATUM #EGCS/MASS
1 US 2 LOWER EAST 28
1 NS 2 LOWER EAST -
1 HS 2 LOWER EAST -
1 us 2 LOWER EAST -
1 US 2 LOWER SOUTH -
1 us 2 LOWER SOUTH -
1 wS 2 LOWER SOUTH -
1 ws 2 LOWER SOUTH -
1 ws 2 LOWER WEST -
1 ws 2 LowER WEST -
1 ws 2 LOWER WEST -
1 HS 2 LOWER WEST -
1 US 2 MIDDLE NORTH 25
1 HS 2 MIDDLE NORTH 25
1 NS 9 MIDDLE NORTH 33
1 US 2 MlDDLE NORTH 22
1 WS 2 MIDDLE EAST 23
1 NS 2 MIDDLE EAST 11
1 us 2 MIDDLE EAST 43
1 NS 2 MIDDLE EAST 3m
1 US 2 MIDDLE SOUTH 15
1 NS 2 MIDDLE SOUTH 13
1 US 2 MIDDLE SOUTH 23
1 US 2 MIDDLE SOUTH 33
1 NS 2 MIDDLE WEST 43
1 US 2 MIDDLE WEST 27
1 W5 2 MIDDLE REST 15
1 us 2 MIDDLE WEST 23
1 us 2 UPPER NORTH 25
1 ws 2 UPPER NORTH 22
1 NS 2 UPPER NORTH 11
1 US 2 UPPER NORTH -
1 US 2 UPPER EAST -
1 us 2 UPPER EAST —
1 US 2 UPPER EAST -
1 W8 2 UPPER EAST -
1 US 2 UPPER SOUTH 31
1 WS 2 UPPER SOUTH 11
1 NS 2 UPPER SOUTH 13
1 ws 2 UPPER SOUTH 18

204

Table 61. Continued.

STD SPP TREE QUADRAHT STRATUM #EGGS/MASS
1 WS 2 UPPER WEST 28
1 WS 2 UPPER WEST 11
1 W 2 UPPER WEST 18
1 WS 2 UPPER WEST 09
1 HS 2 TOP TOP 24
1 WS 2 TOP TOP $9
1 WS 2 TOP TOP 46
1 WS 2 TOP TOP 39
2 W 1 LOWER WORTH 20
2 WS 1 LOWER NORTH 15
2 WS 1 LOWER NORTH 32
2 HS 1 LOWER NORTH 18
2 WS 1 LOWER EAST -
2 WS 1 LOWER EAST -
2 WS 1 LOWER EAST -
2 WS 1 LOWER E ‘T -
2 NS 1 LOWER SOUTH 25
2 WS 1 LOWER SOUTH 2D
2 WS 1 LOWER SOUTH 43
2 WS 1 LOWER SOUTH 18
2 WS 1 LOWER WEST IE
2 WS 1 LOWER WEST 18
2 WS 1 LOWER WEST 31
2 NS 1 LOWER WEST -
2 WS 1 MIDDLE NORTH 27
2 W5 1 MIDDLE NORTH 13
2 WS 1 MIDDLE NORTH 18
2 WS 1 MIDDLE NORTH 2W
2 WS 1 MIDDLE EAST 13
2 WS 1 MIDDLE EAST 25
2 WS 1 MIDDLE EAST 23
2 WS 1 HIDDLE EAST 18
2 WS 1 MIDDLE SOUTH 27
2 WS 1 MIDDLE SOUTH 39
2 WS 1 MIDDLE SOUTH 2D
2 WS 1 MIDDLE SOUTH 23
2 WS 1 MIDDLE WEST 41
2 WS 1 MIDDLE WEST 18
2 WS 1 MIDDLE WEST 22
2 WS 1 MIDDLE WEST 27

205

Table G1. Continued.

2 WS 1 UPPER NORTH 22'
2 HS 1 UPPER NORTH 1 3
2 WS 1 UPPER NORTH 1 3
2 WS 1 UPPER NORTH 2‘8
2 WS 1 UPPER EAST 27
2 NS 1 UPPER LAST 23
2 WS 1 UPPER EAST 1 1
2 HS 1 UPPER EAST 1 3
2 WS 1 UPPER SOUTH 13
2 HS 1 UPPER SOUTH 1 7
2 WS 1 UPPER SOUTH 39
2 NS 1 UPPER SOUTH 1 8
2 NS 1 UPPER WEST 1 1
2 WS 1 UPPER WEST 25
2 NS 1 UPPER WEST 1 3
2 WS 1 UPPER WEST 22
2 HS 1 TOP TOP 31
2 NS 1 TOP TOP 1 8
2 NS 1 TOP TOP 20
2 HS 1 TOP TOP -
2 HS 2 LOWE R NORTH -
2 WS 2 LOWER. NORTH -
2 HS 2 LOWER NO R'I‘H -
2 WS 2 LOWER NORTH -
2 NS 2 LOWER EAS T 1 'r3
2 US 2 LOWER EAST 25
2 'WS 2 LONE R EAST 1 1
2 WS 2 LOWER EAST 11
2 NS 2 LOWER SOUTH U9
2 NS 2 LONE R SOU T11 -
2 NS 2 LOWER SOUTH -
2 WS 2 LOWER SOUTH -
2 NS 2 LO'VJER WEST -
2 "NS 2 LOWER WEST -
2 WS 2 LOWER WEST -
2 WS 2 LOWER NEST -
2 HS 2 MI DDLE NORTH 1 5
2 HS 2 MIDDLE NOR"h 20
2 WS 2 MIDDLE NORTH -
2 WS 2 MI DDLE NORTH -

206

Table GI. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
2 WS 2 MIDDLE EAST 15
2 WS 2 MIDDLE EAST 18
2 WS 2 MIDDLE EAST 08
2 US 2 MIDDLE EAST 13
2 US 2 MIDDLE SOUTH -
2 WS 2 MIDDLE SOUTH -
2 NS 2 MIDDLE SOUTH -
2 HS 2 I’ll DDLE SOUTH -
2 US 2 MIDDLE WEST 08
2 NS 2 MIDDLE WEST 15
2 WS 2 MIDDLE WEST -
2 \‘iS 2 MIDDLE NEST -
2 HS 2 UPPER NORTH 3 1
2 NS 2 UPPER NORTH 1 5
2 NS 2 UPPER NORTH 25
2 WS 2 UPPER flORTH 20
2 WS 2 UPPER EAST 18
2 NS 2 UPPER EAST G9
2 NS 2 UPPER EAST 11
2 WS 2 UPPER EAST U9
2 WS 2 UPPER SOUTH 28
2 US 2 UPPER SOUTd -
2 WS 2 UPPER SOUTH -
2 WS 2 UPPER SOUTH -
2 NS 2 UPPER WEST 25
2 WS 2 UPPER WEST 13
2 US 2 UPPER WEST 23
2 WS 2 UPPER WEST 13
2 HS 2 TOP TOP 30
2 NS 2 TOP TOP -
2 HS 2 TOP TOP -
2 WS 2 TOP TOP -
3 WS 1 LOWER NORTH -
3 WS 1 LOWER NO RTH -
3 WS 1 LOWER NORTH -
3 WS 1 LOWER NORTH -
3 WS 1 LOWER EAS T -
3 WS 1 LOWER EAST -
3 WS 1 LOWER EAST -
3 WS 1 LOWER EAST -

207

Table G1. Continued.

3 WS 1 LOWER SOUTH 22
3 WS 1 LOWER SOUTH 22
3 WS 1 LOWER SOUTH -
3 WS 1 LOWER SOUTH -
3 WS 1 LOWER WEST 23
3 WS 1 LOWER WEST $6
3 WS 1 LOWER WEST -
3 WS 1 LOWER WEST -
3 WS 1 MIDDLE NORTH 28
3 WS 1 MIDDLE NORTH 22
3 NS 1 MI DDLE NORTH 2 3
3 W8 1 MIDDLE NORTH -
3 NS 1 MIDDLE EAST 15
3 NS 1 MIDDLE EAST 18
3 WS 1 MIDDLE EAST -
3 WS 1 MIDDLE EAST -
3 W3 1 MIDDLE SOUTH 18
3 NS 1 MIDDLE SOUTH 18
3 WS 1 MIDDLE SOUTH 18
3 WS 1 MIDDLE SOUTH 15
3 US 1 MIDDLE WEST 09
3 WS 1 MIDDLE WEST 22
3 WS 1 MIDDLE WEST 23
3 WS 1 MIDDLE WEST -
3 WS 1 UPPER NORTH 30
3 WS 1 UPPER NORTH -
3 NS 1 UPPER NORTH -
3 WS 1 UPPER NORTH -
3 WS 1 UPPER EAST 18
3 WS 1 UPPER EAST 18
3 WS 1 UPPER EAST 23
3 WS 1 UPPER EAST 25
3 WS 1 UPPE SOUTH 26
3 WS 1 UPPER SOUTH 3%
3 NS 1 UPPER SOUTH -
3 WS 1 UPPER SOUTH -
3 WS 1 UPPER WEST 18
3 WS 1 UPPER WEST 15
3 WS 1 UPPER WEST 22
3 WS 1 UPPER WEST 36

Table GI.

Continued.

208

wwwwwwwwwwwwwwwwwwwwwwwwwwwwwwwwwwwwwwww

WS
WS
NS
W8
W8
WS
WS
W8
W8
W3
NS
NS
NS
NS
HS
NS
NS
ws
48
NS
NS
us
us
NS
NS
WS
NS
NS
NS
NS
WS
WS
WS
NS
NS
NS
W8
NS
NS
NS

NNNMNNNNNNNNNNNNNNMNNNNNMNNNNNNNNIONNHHHH

TOP
TOP
TOP
TOP
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
LOWER
MIDDLE
MIDDLE
MI DDLE
MI DDLE
MI DDLE
MIDDLE
MIDDLE
MI DDLE
MIDDLE
MI DDLE
MI DDLE
MI DDLE
MIDDLE
MIDDLE
MIDDLE
UPPER
UPPER
UPPER
UPPER

TOP
TOP
TOP
TOP
NORTH
NORTH
NORTH
NO RT H
EAST
EAST
EAST
EAST
SOUTH
SOUTH
SOUTH
SOUTH
WEST
WEST
WEST
NEST
”NO RT 'd
NORTH
N O R'I' H
NORTH
EAST
EAST
EAST
EAST
SOUTH
SOUTH
SOUTH
SOUTH
WEST
WEST
WEST
WEST
NORTH
NO RT H
NORTH
NORTH

23

20

209

Table G1. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
3 HS 2 UPPER EAST 13
3 WS 2 UPPER EAST 18
3 WS 2 UPPER EAST 22
3 WS 2 UPPER EAST 27
3 WS 2 UPPER SOUTH 13
3 US 2 UPPER SOUTH 25
3 WS 2 UPPER SOUTH II
3 WS 2 UPPER SOUTH 28
3 WS 2 UPPER WEST 20
3 NS 2 UPPER NEST 1 3
3 WS 2 UPPER WEST 2f“.
3 WS 2 UPPER WEST 18
3 WS 2 TOP TOP (49
3 WS 2 TO? TOP 1 5
3 ‘u'S 2 TOP TOP 1 3
3 VS 2 10? TOP 26
4 NS 1 LC? L3H “IO RIB. -
4 'JS 1 LOWER 1301?] H. -
4. NS 1 LOWER WORK :1 -
4 US l LOU/ER NORTH -
4 V13 1 LOWER EAST 2 3
4 HS 1 LOWER EAST l 1
4 WS 1 LOWER EAST 1 8
4 \JS 1 LOWER EAST 24
4 \JS 1 LOWER SOUTH 43
4 NS 1 LOWER SOUTH 46
4 WS 1 LOWER SOUTH l3
4 US 1 LOWER SOUTH 39
4 WS 1 LOWER WEST 20
4 HS 1 LOWER WEST 69
4 WS 1 LOWER NEST 20
4 NS 1 LOWER WEST 26
4 WS 1 MIDDLE NORTH 18
4 HS 1 MIDDLE NORTH 33
4 NS 1 MIDDLE NORTH 15
4 WS 1 MIDDLE NORTH 08
4 WS 1 MIDDLE EAST 36
4 WS 1 MIDDLE EAST 25
4 WS 1 MIDDLE EAST 27
4 WS 1 MIDDLE EAST 35

210

Table 61. Continued.

4 WS 1 MIDDLE SOUTH 31
4 NS 1 MIDDLE SOUTh 11
4 WS 1 MIDDLE SOUTH 15
4 WS 1 MIDDLE SOUTH 15
4 WS 1 MIDDLE WEST 25
4 WS 1 MIDDLE WEST 30
4 WS 1 MIDDLE WEST 39
4 WS 1 MIDDLE WEST 36
4 WS 1 UPPER NORTH 11
4 WS 1 UPPER NORTH 15
4 WS 1 UPPER NORTH 13
4 WS 1 UPPER NORTH 98
4 WS 1 UPPER EAST 13
4 W3 1 UPPER EAST 39
4 WS 1 UPPER EAST 13
4 WS 1 UPPER EAST 15
4 WS 1 UPPER SOUTH 24
4 WS I UPPER SOUTH 15
4 WS 1 UPPER SOUTH Q9
4 WS 1 UPPER SOUTH 15
4 NS 1 UPPER WEST 39
4 WS 1 UPPER WEST ”5
4 WS 1 UPPER WEST 3O
4 WS 1 UPPER WEST 33
4 NS 1 TOP TOP 27
4 WS 1 TOP TOP 18
4 WS 1 TOP TOP 39
4 WS 1 TOP TOP 18
4 NS 2 LOWER NORTH -
4 WS 2 LOWER NORPH -
4 WS 2 LOWER NORPH -
4 WS 2 LOWER NORTH -
4 WS 2 LOWER EAST 1 3
4 WS 2 LOWER EAST 1 5
4 WS 2 LOWER EAST 18
4 WS 2 LOWER EAST 33
4 WS 2 LOWER SOUTH 11
4 WS 2 LOWER SOUTH I3
4 WS 2 LOWER SOUTH 18
4 W8 2 LOWER SOUTH 23

211

Table Gl. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
4 WS 2 LOWER WEST -
4 WS 2 LOWER WEST -
4 WS 2 LOWER WEST -
4 WS 2 LOWER WEST -
4 WS 2 MIDDLE NORTH 13
4 WS 2 MIDDLE NORTH 29
4 WS 2 MIDDLE NORTH 15
4 WS 2 MIDDLE NORTH 38
4 WS 2 MIDDLE EAST 13
4 WS 2 MIDDLE EAST 20
4 WS 2 MIDDLE EAST 18
4 WS 2 MIDDLE EAST 23
4 WS 2 MIDDLE SOUTH 08
4 WS 2 MIDDLE SOUTH 43
4 WS 2 MIDDLE SOUTH 25
4 WS 2 MIDDLE SOUTH 1 5
4 WS 2 MIDDLE WEST 20
4 WS 2 MIDDLE WEST 39
4 WS 2 MIDDLE WEST 43
4 NS 2 MIDDLE WEST 27
4 WS 2 UPPER NORTH 04
4 WS 2 UPPER NORTH 25
4 WS 2 UPPER NORTH 18
4 NS 2 UPPER NORTH 2%
4 WS 2 UPPER EAST O9
4 WS 2 UPPER EAST 13
4 W3 2 UPPER EAST 25
4 WS 2 UPPER EAST 20
4 WS 2 UPPER SOUTH 11
4 WS 2 UPPER SOUTH 11
4 WS 2 UPPER SOUTH 22
4 WS 2 UPPER SOUTH 13
4 WS 2 UPPER WEST 99
4 WS 2 UPPER WEST 25
4 WS 2 UPPER WEST 15
4 WS 2 UPPER WEST 15
4 WS 2 TOP TOP 20
4 WS 2 TOP TOP 13
4 WS 2 TOP TOP 13
4 WS 2 TOP TOP 13

212

Table Gl. Continued.

STD SPP TREE QUADRANT STRATUM #EGGS/MASS
5 WS 1 LOWER NORTH 26
5 WS 1 LOWER NORTH -
5 WS 1 LOWER NORTH —
5 WS 1 LOWER NORTH -
5 W3 1 LOWER EAST 39
5 WS 1 LOWER EAST 11
5 WS 1 LOWER EAST 2,0)
5 WS 1 LOWER EAST 38
5 WS 1 LOWER SOUTH 22
5 WS 1 LOWER SOUTH 15
5 WS 1 LOWER SOUTH 38
5 W5 1 LOWER SOUTH 22
5 WS 1 LOWER WEST 1 1
5 NS 1 LOWER WEST -
5 WS 1 LOWER WEST -
5 WS 1 LOWER WEST -
5 NS 1 MIDDLE NORTH 1 3
5 W8 1 MIDDLE NORTH 36
5 HS 1 MIDDLE NORTH 13
5 W3 1 MIDDLE NORTH 25
5 HS 1 MIDDLE EAST 28
5 W3 1 MIDDLE EAST 18
5 WS 1 MIDDLE EAST 15
5 WS 1 MIDDLE EAST 1 5
5 WS 1 MIDDLE SOUTH 30
5 NS 1 MIDDLE SOUTH 13
5 NS 1 MIDDLE SOUTH 25
5 WS 1 MIDDLE SOUTH 25
5 WS 1 MIDDLE WEST 15
5 WS 1 MIDDLE WEST 15
5 WS 1 MIDDLE WEST 23
5 WS 1 MIDDLE WEST 1 5
5 HS 1 UPPER NORTH 23
5 WS 1 UPPER NORTH 23
5 NS 1 UPPER NORTH 28
5 WS 1 UPPER NORTH 23
5 WS 1 UPPER EAST 1 1
5 WS 1 UPPER EAST 18
5 WS 1 UPPER EAST 2G
5 NS 1 UPPER EAST 33

213

Table GI. Continued.

5 WS 1 UPPER SOUTH 25
5 WS 1 UPPER SOUTH 20
5 WS 1 UPPER SOUTH 40
5 WS 1 UPPER SOUTH 22
5 WS 1 UPPER WEST 38
5 WS 1 UPPER WEST 09
5 WS 1 UPPER WEST 33
5 WS 1 UPPER WEST -
5 WS 1 TOP TOP 13
5 WS 1 TOP TOP 25
5 WS 1 TOP TOP 26
5 WS 1 TOP TOP 33
5 W3 2 LONER NORTH -
5 NS 2 LOWER NORTH -
5 WS 2 LOWER NORTH -
5 WS 2 LOWER NORTH -
5 HS 2 LOWER EAST 20
5 WS 2 LOWER EAST -
5 WS 2 LOWER EAST -
5 WS 2 LOWER EAST -
5 WS 2 LOWER SOUTH 22
5 WS 2 LOWER SOUTH 31
5 WS 2 LOWER SOUTH 28
5 WS 2 LOWER SOUTH 18
5 WS 2 LOWER WEST -
5 WS 2 LOWER WEST -
5 WS 2 LOWER WEST -
5 WS 2 LOWER WEST -
5 WS 2 MIDDLE NORTH 2%
5 WS 2 MIDDLE NORTH -
5 WS 2 MIDDLE NORTH -
5 WS 2 MIDDLE NORTH -
5 WS 2 MIDDLE EAST 18
5 WS 2 MIDDLE EAST 25
5 WS 2 MIDDLE EAST 13
5 WS 2 MIDDLE EAST -
5 WS 2 MIDDLE SOUTH 13
5 WS 2 MIDDLE SOUTH 22
5 WS 2 MIDDLE SOUTH 15
5 NS 2 MIDDLE SOUTH 11

214

Table Gl. Continued.

STD SPP TREE QUADRANT STRATUM #FGGS/MASS
5 ws 2 MIDDLE WEST 46
5 us 2 MIDDLE WEST 20
5 ws 2 MIDDLE WEST -
5 ws 2 MIDDLE WEST —
5 ws 2 UPPER NORTH 23
5 ws 2 UPPER NORTH -
5 NS 2 UPPER NORTH -
5 ws 2 UPPER NORTH -
5 ws 2 UPPER EAST 13
5 ws 2 UPPER EAST 23
5 us 2 UPPER EAST 15
5 ws 2 UPPER EAST 27
5 ws 2 UPPER SOUTH 28
5 NS 2 UPPER SOUTH 15
5 NS 2 UPPER SOUTH 13
5 ws 2 UPPER SOUTH 20
5 ws 2 UPPER WEST -
5 HS 2 UPPER NEST -
5 us 2 UPPER WEST -
5 WS 2 UPPER WEST -
5 ws 2 TOP TOP -
5 ws 2 TOP TOP -
5 ws 2 TOP TOP -
5 RS 2 TOP TOP —

STAND:

SPP:

TREE:

STRATUM:

QUADRANT:

# EGGS PER
MASS:

215

Stand number

Sampled tree Specie

BF 8 Balsam fir
NS = White spruce

Tree number

Stratum (Fig. 10) that the branch grew in from which
the needle(s) and egg mass(es) were retained and the
eggs counted.

Quadrant (Fig. 11) that the branch grew in from which
the needle(s) and the egg mass(es) were retained and
the eggs counted.

The estimated number of eggs per randomly selected
egg mass. Four egg masses were selected from each

cell, within each tree.