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This is to certify that the

- thesis entitled
Host Preference and Potential Impact of Pine

Shoot Beetle [Tomicns.9iniperda (L.)(Coleoptera:

 

Scolytidae)] in Michigan Pine Stands

presented by

Nathan Wade Siegert

has been accepted towards fulfillment
of therequirements for

Master of Science." degree in ‘Forestrx

 

   

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Date 1 May 2000

 

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moo chiRC/DdoDmpGSoM

HOST PREFERENCE AND POTENTIAL IMPACT OF PINE SHOOT BEETLE
[T omicus piniperda (L.) (COLEOPTEM: SCOLYTIDAE)] IN MICHIGAN PINE
STANDS
By

Nathan Wade Siegert

A THESIS
Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of
MASTER OF SCIENCE
Department of Forestry

2000

ABSTRACT

HOST PREFERENCE AND POTENTIAL IMPACT OF PINE SHOOT BEETLE
[Tomicus pim'perda (L.) (COLEOPTERA: SCOLYTIDAE)] IN MICHIGAN PINE
STANDS
By

Nathan Wade Siegert

The common pine shoot beetle, T omicus pim‘perda (L.) (Coleoptera: Scolytidae),
is a native pest of pines in Eurasia that was discovered in North America in 1992. More
information is needed concerning T. pinz’perda host preference as it becomes established
in North American forests. Distribution, host preference and potential impacts were
examined using red (Pinus resinosa Ait.), jack (Pinus banksiana Lamb.) and Scotch pine
(Pinus sylvestris L.) in laboratory and field experiments between 1997 and 1999.

T. pim'perda populations were well-established in southwestern lower Michigan,
beginning to immigrate into mid-Michigan, and not found in northwestern lower
Michigan between 1997 and 1999. T. pim'perda colonized Scotch pine logs more
frequently than red and jack pine in southwestern stands. In laboratory experiments, T.
piniperda parent adults preferentially colonized Scotch pine logs over red and jack pine
logs, while progeny adults preferred jack and Scotch pine shoots over red pine shoots.

Pine stands were surveyed to quantify the frequency and extent of shoot damage
attributable to T. pim'perda, native shoot-feeding insects, and non-insect factors to
evaluate the current impact of T. piniperda in Michigan. Non-insect factors accounted
for the majority of shoot damage. T. piniperda damaged shoots most frequently in
Scotch pine stands. Implications of this research for developing improved methods to

detect and manage T. piniperda in forest ecosystems are discussed.

ACKNOWLEDGEMENTS

I thank my advisor and friend, Deborah G. McCullough, for her enthusiastic
devotion and tireless diligence as a mentor and researcher. I appreciate Deb’s willingness
to allow me the freedom to explore my interests in pine shoot beetle ecology and
behavior during this project. Her guidance and unceasing encouragement motivated me
to strive for excellence, originality and balance in my research and in my life outside
academia. I would also like to thank Don Dickmann, Ed Grafius and Bob Haack for their
counsel and valuable advisement during this project.

Numerous folks helped to make the completion of this research possible. I
gratefully acknowledge Kirsten Fondren, Erin Smith and Angela Toma for their
assistance sorting insect specimens and debarking thousands of pine logs during this
project. I also thank Matt Davenport for his assistance throughout the ’99 summer. Greg
Kowalewski, Jim Curtis, John Vigneron, and Karen Bushouse, all of Michigan State
University’s W. K. Kellogg Experimental Forest, have been of inestimable value during
this project. I also extend my sincere gratitude to Rufus Isaacs, Jim Keller, and Jim
Miller, all of Michigan State University’s Center for Integrated Plant Systems, for their
technical assistance with the wind-tunnels experiments.

I thank the Michigan Department of Natural resources personnel who provided
much assistance locating field sites and permitted me to conduct research in the state
forests in their areas; Maria Albright and John Lerg of Allegan State Game Area, Mark
Bishop of Barry State Game Area, Courtney Bourgondy of Gladwin Forest Area, Joe

Fields of Traverse City Forest Area, and Roger Mech and Frank Sapio of Forest Health

iii

Protection, Forest Management Division. I also thank Clara Ward for allowing me to
collect pine shoot beetle at Fenner Arboretum in Lansing, MI.

The support and comments of my fellow graduate student colleagues are greatly
appreciated. Most specifically, I thank Lyle Buss, Beth Dankert, Piera Giroux, Heather
Govenor, Dylan Parry, Ararn Stump and Tim Work. My interactions with them have
provided me with many opportunities and distractions, which have served to further
enrich my graduate experiences.

This research was funded by the McIntire—Stennis Cooperative Forestry Research
Grant program and the Michigan Department of Natural Resources. Additional funding
was provided through the Ray and Bernice Hutson Endowment form the Department of
Entomology for partial support of travel expenses to several regional and national

entomological meetings.

iv

TABLE OF CONTENTS

 

LIST OF TABLES __________________________________________________________________________________________________________ viii
LIST OF FIGURES ________________________________________________________________________________________________________ xii
LIST OF SYMBOLS, ABBREVIATIONS AND NOMENCLATURE ____________________ xiv
INTRODUCTION ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 1
T omicus piniperda Biology _______________________________________________________________________________ 2
Introduction to North America ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 3
Scope of the Present Study ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 5
Thesis Organization ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 6
CHAPTER 1
Colonization of Scotch, Red and Jack Pine Logs by T omicus piniperda (L.)
(Coleoptera: Scolytidae) in Michigan Pine Stands ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 8
Introduction ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 8
Methods and Materials ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 12
Field Sites ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 12
Experimental Design ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 12
Statistical Analyses ________________________________________________________________________________ 14
Results ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 15
Distribution of T omicus piniperda in Michigan __________________________________ 15
T omicus pim'perda Attacks in Pine Stands in Southwestern
Lower Michigan ,,,,,,,,,,, , ______________________________________ 16
T omicus pim’perda Attacks in Pine Logs in Southwestern
Lower Michigan _________________________________________________________________________ 16
T omicus pim'perda Attacks in Pine Logs in Mid-Michigan
Stands ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 17
Log Quality ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 18
Discussion ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 20
Tables ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 23
Figures ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 30
Literature Cited ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 31
CHAPTER 2
Host Preference of Tomicus piniperda (L.) (Coleoptera: Scolytidae) Parent
Adults and Shoot-Feeding Progeny Adults ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 37
Introduction ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 37
Methods and Materials _______________________________________________________________________________________ 39
Overwintered Parent Adults ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 39
Shoot-Feeding Progeny Adults ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 40
Statistical Analyses ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 42
Results and Discussion 43

Overwintered Parent Adults 43

oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo

Shoot-Feeding Progeny Adults _____________________________________________________________ 44
Tables ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 48
Figures ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 50
Literature Cited ___________________________________________________________________________________________________ 57

CHAPTER 3

Survey of Shoot Damage Caused by T omicus piniperda (L.) (Coleoptera:

Scolytidae) in Michigan Pine Plantations _____________________________________________________________________ 63
Introduction ________________________________________________________________________________________________________ 63
Methods and Materials _______________________________________________________________________________________ 66

Shoot Injury Diagnosis ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 66

Autumn 1997 and Spring 1998 Damage Surveys ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 67

Autumn 1998 and Autumn 1999 Damage Surveys ............................. 69

Statistical Analyses ________________________________________________________________________________ 69

Results _________________________________________________________________________________________________________________ 71

Damaging Agents __________________________________________________________________________________ 71

Non—Insect Damage _______________________________________________________________________________ 72

T omicus piniperda Damage .................................................................. 74

Native Shoot-boring Insect Damage _____________________________________________________ 75

Discussion ........................................................................................................... 76

Non-Insect Damage ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 76

T omicus piniperda Damage .................................................................. 76

Native Shoot-boring Insect Damage _____________________________________________________ 79

Conclusions ____________________________________________________________________________________________ 81

Tables .................................................................................................................. 82

Figures _________________________________________________________________________________________________________________ 99
Literature Cited ................................................................................................... 100
MANAGEMENT IMPLICATIONS _____________________________________________________________________________ 104
APPENDICES 108

APPENDIX A ~ Table A1. Frequency of shoots damaged by non-insect factors

in red, jack and Scotch pine stands that were surveyed in lower Michigan

between 1997 and 1999. ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 109
APPENDIX B - Table Bl. Frequency of shoots damaged by T omicus piniperda

in red, jack and Scotch pine stands that were surveyed in lower Michigan

between 1997 and 1999. ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 120
APPENDIX C - Table Cl. Frequency of shoots damaged by Canopthorus

resinosae in red, jack and Scotch pine stands that were surveyed in

lower Michigan between 1997 and 1999. ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 127

vi

APPENDIX D - Table D1. Frequency of shoots damaged by Eucosma gloriola
in red, jack and Scotch pine stands that were surveyed in lower Michigan

between 1997 and 1999. _____________________________________________________________________________________ 131
APPENDIX E - Record Deposition of Voucher Specimens _______________________________________ 135
LITERATURE CITED 139

ooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo

vii

LIST OF TABLES

CHAPTER 1.

Table 1.1. Location and characteristics of pine stands in lower Michigan used
as field sites in 1997 and 1998. BA refers to total basal area, DBH

refers to average diameter at breast height and SI refers to site index. _________

Table 1.2. Mean number (1: SEM) of T omicus piniperda attacks in pine stands
(11 = 18) stands in northwestern, mid-, and southwestern lower Michigan
in 1997 (n = 484 logs) and 1998 (n = 470 logs). Significant differences
among regions in each year are indicated by different letters after the

value (P < 0. 05). _______________________________________________________________________________________________

Table 1.3. Mean number (:5 SEM) of T omicus piniperda attacks in jack, red
and Scotch pine stands (11 = 6) stands in southwestern lower Michigan
in 1997 (n = 159 logs) and 1998 (n = 161 logs). Significant differences
among species in each year are indicated by different letters after the

value (P < 0. 05). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,

Table 1.4. Mean number (i SEM) of T omicus piniperda attacks in jack, red
and Scotch pine logs in all stands (11 = 6) stands in southwestern lower
Michigan in 1997 (n = 159 logs) and 1998 (n = 161 logs). Significant
differences among log species in each year are indicated by different
letters after the value (P < 0.05). ________

 

Table 1.5. Mean number (: SEM) of T omicus piniperda attacks in jack, red
and Scotch pine logs in jack, red and Scotch pine stands (11 = 6) stands
in southwestern lower Michigan in 1997 (n = 159 logs) and 1998 (n =

161 logs). ___________________________________________________________________________________________________________

Table 1.6. Summary of regression equations for predicting number of T omicus
piniperda galleries per log using all logs combined and for each log
species in 1997 (n = 159 logs). Predictor variables (excluding intercept)
are listed in order of contribution to 1'2.

ooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo

Table 1.7. Summary of regression equations for predicting number of T omicus
piniperda galleries per log using all logs combined and for each log
species in 1998 (n = 161 logs). Predictor variables (excluding intercept)
are listed in order of contribution to 1‘2.

viii

.23

__24

25

O.

26

27

28

29

CHAPTER 2.

Table 2.1. Number of T omicus pim'perda parent adults that selected Scotch, red
or jack pine logs in trials conducted in a wind tunnel in 1997 (n = 98

beetles) and 1999 (n = 414 beetICS). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,

Table 2.2. Number of T omicus piniperda progeny adults that selected current-
year Scotch, red or jack pine shoots in laboratory trials conducted in

1997 (n = 382 beetles) and 1999 (n = 1052 beetles). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,

CHAPTER 3.

Table 3.1. Location and characteristics of pine stands in northern lower

Michigan surveyed for shoot damage between 1997 and 1999. ,,,,,,,,,,,,,,,,,,,,

Table 3.2. Location and characteristics of pine stands in southwestern lower

Michigan surveyed for shoot damage between 1997 and 1999. ,,,,,,,,,,,,,,,,,,,,

Table 3.3. Mean number (:1; SEM) of pine shoots (per m2) damaged in red,
jack and Scotch pine stands in northern and southwestern lower
Michigan between 1997 and 1999. Significant differences within

each column are indicated by different letters after the value (P < 0.05)...”

Table 3.4. Overall mean number (1: SEM) of pine shoots (per m2) damaged
by various agents in red, jack and Scotch pine stands in northern and
southwestern lower Michigan between 1997 and 1999. Significant
differences within each column are indicated by different letters after

the value (P < 0. 05). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,

Table 3.5. Mean number (1 SEM) of pine shoots (per m2) damaged by non-
insect factors in northern and southwestern lower Michigan in all pine
stands between 1997 and 1999. Significant differences within each

column are indicated by different letters after the value (P < 0.05). ,,,,,,,,,,,,

Table 3.6. Mean number (i SEM) of pine shoots (per m2) damaged by non-
insect factors in red, jack and Scotch pine stands in lower Michigan
between 1997 and 1999. Significant differences within each column

are indicated by different letters after the value (P < 0.05). ,,,,,,,,,,,,,,,,,,,,,,,,,,

48

49

82

83

84

85

86

Table 3.7. Mean number (:1: SEM) of pine shoots (per m2) damaged by non-
insect factors in red, jack and Scotch pine stands in northern and
southwestern lower Michigan between 1997 and 1999. Significant
differences within each column are indicated by different letters after
the value (P < 0.05). ___________________________________________________________________________________________ 88

Table 3.8. Mean number (: SEM) of pine shoots (per m2) damaged by T omicus
piniperda in northern and southwestern lower Michigan in all pine stands
between 1997 and 1999. Significant differences within each column are
indicated by different letters after the value (P < 0. 05). __________________________________ 89

Table 3.9. Mean number (1 SEM) of pine shoots (per m2) damaged by T omicus
piniperda in red, jack and Scotch pine stands in lower Michigan between
1997 and 1999. Significant differences within each column are indicated
by different letters after the value (P < 0. 05). __________________________________________________ 90

Table 3.10. Mean number (: SEM) of pine shoots (per m2) damaged by T omicus
pim'perda in red, jack and Scotch pine stands in northern and southwestern
lower Michigan between 1997 and 1999. Significant differences within
each column are indicated by different letters after the value (P < 0. 05). ______ 91

Table 3.11. Mean number (3: SEM) of pine shoots (per m2) damaged by T omicus
piniperda in Scotch pine stands in southwestern lower Michigan between
1997 and 1999. Stands were surveyed in Cass (Cass — l, Cass — 2),
Kalamazoo (Kal — 1) and Allegan (Al — 1, A1 — 2) counties. ,,,,,,,,,,,,,,,,,,,,,,,,,,, 92

Table 3.12. Mean number (: SEM) of pine shoots (per m2) damaged by
Conopthorus resinosae in northern and southwestern lower Michigan
in all pine stands between 1997 and 1999. Significant differences
within each column are indicated by different letters after the value
(P < 0. 05). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 93

Table 3.13. Mean number (: SEM) of pine shoots (per m2) damaged by
Canopthorus resinosae in red, jack and Scotch pine stands in lower
Michigan between 1997 and 1999. Significant differences within
each column are indicated by different letters after the value (P < 0. 05). ...... 94

Table 3.14. Mean number (: SEM) of pine shoots (per m2) damaged by
Canopthorus resinosae in red, jack and Scotch pine stands in northern
and southwestern lower Michigan between 1997 and 1999. Significant
differences within each column are indicated by different letters after
the value (P < 0. 05). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 95

Table 3.15. Mean number (i SEM) of pine shoots (per m2) damaged by Eucosma
gloriola in northern and southwestern lower Michigan in all pine stands

between 1997 and 1999. Significant differences within each column are
indicated by different letters after the value (P < 0. 05). __________________________________ 96

Table 3.16. Mean number (: SEM) of pine shoots (per m2) damaged by Eucosma
gloriola in red, jack and Scotch pine stands in lower Michigan between
1997 and 1999. Significant differences within each column are indicated
by different letters after the value (P < 0.05). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 97

Table 3.17. Mean number (i SEM) of pine shoots (per m2) damaged by Eucosma
gloriola in red, jack and Scotch pine stands in northern and southwestern
lower Michigan between 1997 and 1999. Significant differences within
each column are indicated by different letters afier the value (P < 0. 05). ______ 98

xi

LIST OF FIGURES

CHAPTER 1.

Figure 1.1. Michigan counties that are shaded were quarantined for T omicus
piniperda as of April 2000. Stands used as field sites were located in
Benzie county (a) in northwestern lower Michigan, Clare (b) and Gladwin
(c) counties in mid-Michigan, and Allegan ((1), Barry (e) and Kalamazoo
(i) counties in southwestern lower Michigan. ___________________________________________________ 30

CHAPTER 2.

Figure 2.1. Proportion of T omicus piniperda parent adults that selected either
Scotch pine or red pine logs in a wind tunnel experiment in A) 1997
(n = 53 beetles) and B) 1999 (n = 191 beetles). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 50

Figure 2.2. Proportion of T omicus pim'perda parent adults that selected either
Scotch pine or jack pine logs in a wind tunnel experiment in A) 1997
(n = 45 beetles) and B) 1999 (n = 131 beetles). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 51

Figure 2.3. Proportion of T omicus piniperda parent adults that selected either
red pine or jack pine logs in a wind tunnel experiment in 1999 (n = 92
beetles). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 52

Figure 2.4. Proportion of T omicus piniperda progeny adults that selected either
Scotch pine or red pine current-year shoots in a laboratory experiment
in A) 1997 (n = 87 beetles) and B) 1999 (n = 257 beetles). ,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 53

Figure 2.5. Proportion of T omicus pim'perda progeny adults that selected either
jack pine or Scotch pine current-year shoots in a laboratory experiment
in A) 1997 (n = 158 beetles) and B) 1999 (n = 527 beetles). ,,,,,,,,,,,,,,,,,,,,,,,,,, 54

Figure 2.6. Proportion of T omicus pim’perda progeny adults that selected either
jack pine or red pine current-year shoots in a laboratory experiment in A)
1997 (n = 137 beetles) and B) 1999 (n = 268 beetles). Proportions were
significantly different between 1997 and 1999 (G = 22.43, P < 0.001). ........ 55

Figure 2.7. Frequency distribution of diameter of shoots (cm) infested by
T omicus piniperda progeny adults in 1997. ,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,, 56

xii

CHAPTER 3.

Figure 3.1. Michigan counties that are shaded were quarantined for T omicus
piniperda as of April 2000. Stands that were surveyed were located in
Benzie county (a), Manistee (b), Clare (0), and Gladwin (c) counties in

northern lower Michigan, and Allegan (e), Kalamazoo (f), and Cass (g)
counties in southwestern lower Michigan. ________________________________________________________ 99

xiii

LIST OF SYMBOLS, ABBREVIATIONS AND NOMENCLATURE

°C

BA

ca

DBH

1x

SEM

SI

degrees Celsius

basal area

circa

diameter at breast height

lux; one lumen per square meter
standard error of the mean

site index

xiv

INTRODUCTION

Bark beetles (Coleoptera: Scolytidae) constitute one of the most destructive insect
guilds associated with forests worldwide. Some species have the ability to mass attack
and kill healthy trees over extensive areas, while others effectively function as primary
decomposers by colonizing recently cut timber, stumps and slash, often introducing
fungi, which further act to breakdown and degrade the wood. The economic and
ecological importance of bark beetles has resulted in intensive research on scolytids
throughout the world and, coincidentally, establishment of the forest entomology
profession in America during the late 1800’s (Furniss 1997). Bark beetles, historically
and presently, comprise an important part of forest entomology. As the utilization of
forest products intensifies, so does the need for increased understanding of the ecology
and behavior of these species to better manage their populations and our forests.

Natural barriers, such as oceans and mountain ranges, have historically restricted
distribution of the world’s biota. Increased international trade, travel and ecosystem
disturbance, especially in the last century, have resulted in a dramatic increase in the
number of introductions of new species (Liebhold et a1. 1995, Niemela and Mattson
1996). Biological invasions can be successful because indigenous flora may have not
evolved defenses against them and the exotic pests usually arrive without their complex
of predators, parasites or pathogens. European biota have been vastly more successful at
invading North American forests than the reverse (N iemela and Mattson 1996). Many of
these foreign species become important pests, causing substantial disturbance to forest
ecosystems and often significant socioeconomic impacts (Liebhold et al. 1995). The

environmental threat posed by the increasing number of biological invasions necessitates

greater efforts to examine the spread and evaluate the potential ecological impacts as

exotic forest pests become established in North America.

T omicus piniperda Biology

The common pine shoot beetle, T omicus (synon. Blastophagus Eichh., synon.
Myelophilus Eichh.) pim'perda (L.) (Coleoptera: Scolytidae), is a significant forest pest in
Europe, Asia, and North Africa, principally infesting Pinus species (Bakke 1968, Salonen
1973, Lekander et a1. 1977, Langstrdm 1980a, Wood and Bright 1987, 1992, Ye 1991,
Bright and Skidmore 1997). T omicus Latr. species are ecologically unique from other
bark beetles in the subfamily Hylesininae Erich, in that adults feed in the pith of live pine
shoots to complete sexual maturation.

Bakke (1968), Salonen (1973) and LangstrOm (1980a) previously described the
univoltine life cycle of T. piniperda. Adult T. piniperda locate and colonize brood
material during a swarming flight period in early spring, when temperatures reach 10 to
12°C. These monogamous parent adults colonize recently cut pine logs, stumps and
slash, preferring thick-barked materials. Beetles bore into the inner bark and each female
excavates a longitudinal gallery, laying eggs along the sides of the gallery. Larvae feed
in galleries that extend perpendicularly from the parent gallery, develop in the phloem for
6 to 12 weeks, pupate and emerge from the brood material in early summer. These newly
emerged progeny adults fly to living pine trees for maturation feeding in healthy, live
pine shoots. Some of the parent adults also leave the galleries after oviposition and feed

in pine shoots, a behavior known as regeneration feeding. Occasionally though, some

parent adults will feed for a short time and then attempt to establish a second “sister
brood.” Progeny adults continue to shoot-feed throughout the summer until autumn,
when they move to the base of pine trees and bore into the bark where they overwinter.
T. piniperda is an important forest pest because it colonizes fresh cut pine timber,
introduces blue stain fimgus which discolors sawlo gs, and, perhaps most importantly,
shoot-feeds in healthy pine trees causing aesthetic damage and, under certain
circumstances, significant growth losses (Andersson 1973, Nilsson 1974, 1976,
Langstrdm and Hellqvist 1990, 1991, Eidmann 1992). Wood and Bright (1987, 1992)
and Bright and Skidmore (1997) provide extensive bibliographies of the European and

North American literature related to T. piniperda.

Introduction to North America

T. pim'perda was first detected in North America at a Scotch pine (Pinus sylvestris
L.) nursery in New Jersey in 1913 (Headlee 1914), after which no further mention occurs
in subsequent reports. T. piniperda has been one of the most commonly intercepted
scolytids on wood products at United States ports with 112 interceptions occurring during
1985 — 1995 (Haack et al. 1997). Established populations of T. piniperda were first
discovered in North America in 1992, near Cleveland, Ohio (Haack and Kucera 1993).
Czokajlo et al. (1997), however, suggest T. piniperda arrived in a New York pine stand
prior to 1982 and unquestionably before 1989. Additionally, an adult T. piniperda was
collected in Lansing, MI, in 1991 , but was not identified until 1993 (Entomology

Museum, Michigan State University). Genetic testing suggests that T. piniperda

populations in the United States were established separately in Ohio near Lake Erie and
in Illinois near Lake Michigan (Carter et a1. 1996). The population established in Illinois
occupies only a small area, while the population established in Ohio encompasses most of
the known infestation (Carter et a1. 1996). As of April 2000, T. piniperda has been found
in at least 295 counties in 11 north central and northeastern states (National Animal and
Plant Inspection Service [NAPIS] 2000), 24 counties in Ontario and 8 counties in Quebec
(C. Markham, USDA Animal and Plant Health Inspection Service [APHIS], pers. comm.)
Federal and state quarantines were implemented in 1992 that regulate the transportation
of pine logs, Christmas trees and nursery trees from infested counties to uninfested
counties (APHIS 1993).

Much of the North American research on T. piniperda to-date has been conducted
in Scotch pine Christmas tree plantations because the Christmas tree industry was
initially most affected by regulations arising from T. piniperda quarantines. Also,
Christmas tree plantations are especially susceptible to infestation because of the annual
availability of abundant brood material (stumps and cull trees). An integrated
management and compliance program for Christmas tree production was developed and
implemented as an alternative to annual federal inspections (McCullough and Sadof
1996, 1998). As T. piniperda populations continue to spread and become established in
pine forests, more information is needed to examine their ecological impacts in forest
ecosystems and to develop management guidelines to reduce the impact of quarantine

regulations on the forest products industry.

Scope of the Present Study

Researchers have shown that T. pim'perda can breed and shoot-feed in many
North American pine species (Chararas 1968, LangstrOm and Hellqvist 1985, Zumr 1992,
Sadof et al. 1994, Lawrence and Back 1995, Amezaga 1997), though few studies have
examined T. pim'perda ecology and behavior in North American forests. Michigan has
790,840 ha of pine on public and private lands (Leatherberry and Spencer 1996). Red
pine (Pinus resinosa Ait.) and jack pine (Pinus banksiana Lamb.) are the two dominant
native species with 358,880 ha and 338,440 ha, respectively. White pine (Pinus strobus
L.) is the third dominant native species with 93,520 ha (Leatherberry and Spencer 1996).
Red pine (V 085 1972, Rudolf 1990) and jack pine (V 055 1972, Rudolph and Laidly 1990)
are abundantly distributed across the Upper Peninsula, the northern half of the Lower
Peninsula and less commonly along the Lake Michigan shore in southwestern Michigan.
Scotch pine is widely planted in Michigan for hedgerows and windbreaks (V 085 1972,
Skilling 1990). Michigan is also a leading producer and exporter of Christmas trees in
North America with pines comprising ca 50% of the 38 million trees sold annually
(Michigan Agricultural Statistics Service [MASS] 2000). Few large Scotch pine stands
exist on public land, but extensive areas of abandoned Scotch pine Christmas tree
plantings occur in lower Michigan.

As of April 2000, 74 of the 83 counties in Michigan were quarantined for T.
piniperda (NAPIS 2000), causing substantial negative economic concerns for the forest
products and Christmas tree industries. Major forestry-related studies in Michigan to

date have focused on phenology of T. piniperda, native scolytids and natural enemies,

impacts of natural enemies, and interspecific competition between native scolytids and T.
piniperda (Lawrence and Haack 1995, Haack and Lawrence 1995a, 1995b, 1997a, Haack
et al. 1997, Kennedy 1998). Many questions remain about T. piniperda in North
American forests. More knowledge is still needed, including an understanding of T.
pim'perda host preference and host suitability, and ecological impacts of T. pim'perda in
North America. The present study addresses these questions about T. piniperda in

Michigan forests.

Thesis Organization

In chapter 1, the distribution and host preference of T. pim'perda adults was
examined in a large-scale field study using red, jack and Scotch pine logs placed in pine
stands throughout lower Michigan. Two host preference studies that were conducted
under controlled conditions in laboratory settings are presented in chapter 2. One study
tested whether T. piniperda parent adults preferentially selected Scotch pine logs when
paired with logs of either red or jack pine, and the other tested whether shoot-feeding
progeny adults preferentially selected current-year Scotch pine shoots when paired with
current-year shoots of either red or jack pine. The frequency and extent of damaged pine
shoots attributable to T. pim‘perda in red, jack and Scotch pine stands versus native shoot-
boring insects and other factors was surveyed in lower Michigan and results are presented
in chapter 3. A concise section on the possible implications of this research is presented

in conclusion.

Thesis chapters were prepared as manuscripts for submission to scientific journals
and are formatted to the specifications for the respective journals. I anticipate submitting
chapter 1 to Environmental Entomology, chapter 2 to The Canadian Entomologist, and

chapter 3 to Northern Journal of Applied Forestry.

CHAPTER 1

COLONIZATION OF SCOTCH, RED AND JACK PINE LOGS BY T OMIC US

PINIPERDA (L.) (COLEOPTERA: SCOLYTIDAE) IN MICHIGAN PINE STANDS

INTRODUCTION

The exotic pine shoot beetle, T omicus pim'perda (L.) (Coleoptera: Scolytidae), is a
native forest pest of pine in Europe and Asia (Bakke 1968, Ye 1991, Eidmann 1992).
Established populations of T. piniperda were discovered in North America in 1992 near
Cleveland, Ohio (Haack and Kucera 1993, Lawrence and Haack 1995, Haack et al. 1997).
As of April 2000, it has been found in at least 295 counties in 11 north central and
northeastern states (National Animal and Plant Inspection Service [NAPIS] 2000), 24
counties in Ontario and 8 counties in Quebec (C. Markham, USDA Animal and Plant
Health Inspection Service [APHIS], pers. comm.) Currently, 74 of the 83 counties in
Michigan are known to be infested (Figure 1.1). Federal and state quarantines were
imposed to regulate the transport of pine logs out of infested counties and the processing of
these logs at mills in uninfested counties (APHIS 1993). These regulations have had
negative economic impacts on mills located in uninfested counties that receive pine from
quarantined counties. Much of the research on T. piniperda in North America to-date has
been conducted in Scotch pine (Pinus sylvestrzls L.) Christmas tree plantations. More
information about the impact of T. piniperda in forest stands is needed to form regulatory

and management policy in Michigan and nationwide.

The recent introduction of the T. piniperda into the Lake States also provides a
unique opportunity to evaluate the ecological impacts of an exotic forest pest in native pine
ecosystems. Virtually all native bark beetles in the Lake States are considered "secondary"
bark beetles; they rarely attack healthy trees and will infest live trees only when the trees
are severely stressed by factors such as drought or defoliation. Native bark beetles, along
with the exotic T. piniperda, typically colonize the phloem of highly stressed or recently
cut pine trees, stumps, or slash and lay eggs. Larvae feed and develop in the phloem,
pupate, and progeny adults emerge from the brood material. Suitability of brood material
for colonization declines over time as phloem moisture and nutrient levels decline. After
12 months, most pine material is no longer attractive or suitable for bark beetle
development. Interspecific competition may arise between native phloem feeders and T.
piniperda if brood material is a limiting factor. However, it is also possible that T.
piniperda may prefer to colonize Scotch pine, its natural host in Europe, over North
American red pine (Pinus resinosa Ait.) or jack pine (Pinus banksiana Lamb). If so, then
T. piniperda and native bark beetles may be spatially partitioned, the likelihood that T.
pim'perda will damage native pine forest stands may be low, and management should be
adjusted accordingly.

Significant behavioral differences exist between the exotic T. piniperda and
native bark beetles which may influence the potential impact of T. piniperda in North
American forests. The most important native scolytid that affects pines in the Lake States
is Ips pini (Say), the pine engraver (Raffa 1991, Schenk and Benjamin 1969). 1. pint is
multivoltine (Thomas 1961, Rudinsky 1962, Schenk and Benjamin 1969), while T.

piniperda has only one generation per year (Bakke 1968, Lingstrdm 1980a, 1980b,

1983). T. piniperda adults colonize suitable pine brood material in early spring, usually 2
to 6 weeks before 1. pint become active (Haack and Lawrence 1995a, 1995b). T. piniperda
are monogomous and each female excavates a single gallery that parallels the grain of the
wood and is distinctively crooked at the ends. 1. pint and related Ips species are
polygamous and 2 to 4 galleries radiate from a central chamber, resulting in a Y or
H-shaped appearance. When T. pim'perda progeny adults emerge from brood material in
June, they fly to live pine trees and tunnel down the center of current-year shoots for
maturation feeding. The native I. pim’ adults complete maturation under the bark and do
not feed in shoots.

T. piniperda and I. pini also differ in the mechanisms they use to locate suitable
brood material. T. piniperda is primarily attracted to alpha-pinene, a monoterpene in pine
oleoresin (Byers et al. 1985, Volz 1988, Byers 1989, Schroeder 1988, Schroeder and
Lindelbw 1989, Zumr 1989, Brattli et a1. 1998, Czokajlo and Teale 1999). Although
recent studies suggest that females may produce a short-range pheromone (Czokajlo 1998),
there is no known aggregation pheromone for T. piniperda (Schdnherr 1972, Byers et al.
1985, 1989, Lanne et al. 1987, Ldyttyniemi et al. 1988, Volz 1988). In contrast, I. pini
beetles are primarily attracted to a combination of host volatiles and two aggregation
pheromones, ipsdienol and lanierone, that the males produce upon finding suitable brood
material (Lanier et al. 1980, Miller and Borden 1990, Raffa and Klepzig 1989, Seybold et
al. 1992, Safianyik et al. 1996).

Some monophagous or oligophagous scolytids have the ability to successfully
colonize (i.e. produce a brood in) hosts that are closely related to their preferred host,

though attack densities and brood production are generally higher in the preferred host

10

(Amman 1982, Holsten and Werner 1990, Cerezke 1995). In Europe, the preferred host
tree for T. piniperda is Scotch pine (Bakke 1968, Salonen 1973, Langstréim 1980a,
1980b, Schroeder 1988, Gibbs and Inman 1991, Brattli et al. 1998). Previous research
has shown that T. piniperda is capable of colonizing and developing in many North
American pine species (Chararas 1968, Ohmart 1980, Lingstrdm and Hellqvist 1985,
Zumr 1992, Lawrence and Haack 1995, Sadof et a1. 1994, Amezaga 1996, 1997), but it is
not clear if T. piniperda preferentially colonizes Scotch pine brood material over North
American pine species.

The goal of this study was to evaluate host preference of T piniperda in red, jack
and Scotch pine stands in three regions of lower Michigan. T. piniperda populations are
known to have been well-established in southwestern Michigan for several years, but
related studies indicated that T. piniperda populations were very low, even undetectable, in
central and northern lower Michigan (N. W. Siegert, A. Kennedy, and D. G. McCullough,
unpublished data). I hypothesized that T. piniperda would preferentially colonize Scotch
pine, its natural host in Europe, when Scotch pine logs and native red and jack pine logs
were available for colonization. My specific objectives were to: (1) assess the distribution
of T. piniperda in lower Michigan forests; (2) determine if T. piniperda adults
preferentially colonize Scotch pine when presented with red, jack and Scotch pine in
Michigan pine stands; and (3) examine whether log quality (i.e. bark thickness, phloem

thickness, and surface area) affected T. piniperda colonization of these three pine species.

11

METHODS AND MATERIALS

Field Sites

Eighteen pine stands, with no thinning or harvest activity in the past 5 years, were
selected in lower Michigan in January 1997. Six stands were located in southwestern
Michigan (Allegan, Kalamazoo, and Barry counties), 6 were located in mid-Michigan
(Clare and Gladwin counties), and 6 were located in northwestern lower Michigan (Benzie
county) (Figure 1.1). Within each of the 3 regions, 2 stands were red pine, 2 stands were
jack pine and 2 stands were Scotch pine. Stands in Kalamazoo county were located at W.
K. Kellogg Experimental Forest, Michigan State University. The remaining stands were
located on state land and selected using the Michigan Department of Natural Resources
(MI DNR) inventory database. Stand selection was initially based on the following
criteria: stands were to be at least 2.02 ha (5 ac), tree diameter at breast height (DBH) > 10
cm (4 in), and basal area (BA) of 23 — 34.5 mz/ha (100 — 150 ftz/ac) (well-stocked) or
(second choice) a BA greater than 34.5 mz/ha (> 150 fiz/ac) (over-stocked). However,
relatively few Scotch pine forest stands were available for this study. Those that were
ranged in size from 1.2 to 2.4 ha (Table 1.1). The minimum size criteria was therefore

dropped for these stands, so they could be included to ensure adequate replication.

Experimental Design

In February 1997 and 1998, 9 freshly cut red pine, jack pine and Scotch pine logs,

ca 20 cm in diameter and 46 cm in length, were placed in each stand (27 logs per stand;

12

486 logs each year). The logs were cut from pine trees located in the same region in lower
Michigan as the stand in which they were placed. Each stand was divided into a northern
block, a central block, and a southern block, and each block was split into three sections.
Three pine logs of the same species, set upright against one another in a tripod, were placed
into one of the three sections within each block. Order of species among each section was
determined by random draw.

The logs were placed in the stands in February before T. pim'perda became active.
Logs were colonized in late March and April by the T. piniperda populations that were
present at each site, along with native subcortical-feeding insects (data collected but not
presented here). Logs were retrieved from the pine stands in July and August after T.
piniperda progeny adults emerged from them, returned to the laboratory and carefully
debarked. Galleries were identified as T. piniperda, Ips spp. or other species and
counted. Data including the number of T. piniperda attacks ( i.e. number of galleries),
and log length, diameter, bark thickness, and phloem thickness were recorded for each of
the logs. The number of T. piniperda attacks were extrapolated to 1m2 of surface area
and included in the tables. Bark thickness was measured at 4 points on each end of a log
and phloem thickness was measured at two points on each end of a log.

Three Lindgren 12-unit multiple funnel traps (Lindgren 1983, Lindgren 1997)
were used to monitor T. piniperda populations in each of the three regions (N. W. Siegert
and D. G. McCullough unpublished data). Traps were within 0.5 km of at least one of
the study sites in each region. Funnel traps were baited with two alpha-pinene lures each

to attract T. piniperda (PheroTech, Inc., Delta, British Columbia) and Vapona insecticide

l3

strips were used in collection cups to quickly kill the insects. Beetles were collected from

traps weekly, sorted and T. piniperda identified.

Statistical Analyses

Differences in the number of T. piniperda attacks in red, jack and Scotch pine
logs (per log and per 1m2 surface area) were analyzed for each year using three-way
analysis of variance (ANOVA) to test for effects of region, stand species, log species and
interactions (PROC GLM program, SAS Institute, Inc. 1989). Due to the substantial
regional differences, only data from the southwestern lower Michigan stands were used
for the remaining comparisons. When results of the ANOVA were significant, a
Fischer’s protected least significant difference (LSD) test was used to determine which
regions or pine species differed (T = LSD critical value). Least squared means were used
in the AN OVA to adjust for missing logs in a few cases, with a corresponding reduction
in error degrees of fieedom (Searle et al. 1980, SAS Institute, Inc. 1989). True means are
reported in all tables. All analyses were conducted at P < 0.05 level of significance,
using SAS statistical software (SAS Institute, Inc. 1989).

Forward stepping multiple regression analysis was used to analyze effects of bark
and phloem thickness, and log surface area on the number of T. piniperda attacks per log
(PROC REG program, SAS Institute, Inc. 1989). Surface area of the log was calculated

by multiplying mean diameter by length by pi.

l4

RESULTS

Distribution of T amicus piniperda in Michigan

Significant regional differences in the frequency of T. piniperda attacks on red,
jack and Scotch pine logs were observed in lower Michigan in 1997 (F = 100.7, P <
0.05) and 1998 (F = 73.0, P < 0.05). Pine logs were colonized by T. piniperda with
significantly higher frequency in southwestern pine stands than logs in northwestern
lower Michigan and mid-Michigan pine stands in 1997 (T = 1.97, P < 0. 05) and 1998 (T
= 1.97, P < 0. 05) (Table 1.2). In 1997, T. piniperda colonized 100% of the logs in
southwestern Michigan and 1% of the logs in mid-Michigan. In 1998, T. piniperda
colonized 100% of the logs in southwestern Michigan and 5% of the logs in mid-
Michigan. No pine logs were colonized by T. piniperda in northwestern lower Michigan
in 1997 or 1998. There was no significant difference between the number of T. piniperda
attacks on logs in mid-Michigan and northwestern lower Michigan in either 1997 (T =
1.97, P > 0.10) or 1998 (T = 1.97, P > 0.10).

A similar pattern of T. piniperda distribution was obtained from fimnel trapping.
Between 1997 and 1999, hundreds of beetles were trapped in southwestern lower
Michigan. In mid-Michigan, no beetles were trapped in 1997, 5 beetles were trapped in
1998, and 34 beetles were trapped in 1999. No T. piniperda were trapped in

northwestern lower Michigan between 1997 and 1999.

15

Tomicus piniperda Attacks in Pine Stands in Southwestern Lower Michigan

The frequency of T. piniperda attacks on pine logs in red, jack and Scotch pine
stands in southwestern lower Michigan differed significantly in 1997 (F = 10.1, P < 0. 05)
and 1998 (F = 3.1, P < 0. 05), though trends differed between years. In 1997, the overall
fiequency of T. piniperda attacks on all species of pine logs were significantly lower in
jack pine stands than logs in red pine (T = 1.98, P < 0. 05) and Scotch pine stands (T =
1.98, P < 0. 05) (Table 1.3). Frequency of attacks did not significantly differ between
logs in red pine and Scotch pine stands (T = 1.98, P > 0.10) in 1997.

In 1998, pine logs in Scotch pine stands were attacked by T. piniperda
significantly more ofien than logs in red pine stands (T = 1.98, P < 0. 05). T. piniperda
attacks on logs in jack pine stands did not differ significantly from logs in Scotch pine (T

= 1.98, P > 0.05) or red pine stands (T = 1.98, P > 0.10) (Table 1.3).

Tomicus piniperda Attacks in Pine Logs in Southwestern Lower Michigan

The frequency of T. piniperda attacks on red, jack and Scotch pine logs in
southwestern lower Michigan differed significantly in 1997 (F = 9.2, P < 0. 05) and 1998
(F = 9.9, P < 0. 05). In 1997, the frequency of T. piniperda attacks on Scotch pine logs
was significantly higher than on jack pine logs (T = 1.98, P < 0.05) and red pine logs (T
= 1.98, P < 0.05), while jack pine and red pine did not differ (T = 1.98, P > 0.10) (Table

1.4).

16

In 1998, the overall number of galleries per log was lower, but frequency of T.
piniperda attacks on Scotch pine logs was again significantly higher than on jack pine
logs (T = 1.98, P < 0.05) and red pine logs (T = 1.98, P < 0. 05) (Table 1.4). Number of
attacks on jack pine and red pine logs did not significantly differ (T = 1.98, P < 0.10) in
1998.

The interactions between stand species and log species followed similar trends but
were not significantly different in either 1997 (F = 2.2, P > 0. 05) or 1998 (F = 0.3, P >

0.10) (Table 1.5).

T omicus piniperda Attacks in Pine Logs in Mid-Michigan Stands

Results from this study appear to document expanding or building populations of
T. piniperda in mid-Michigan pine stands in 1997 and 1998. In 1997, four red pine logs
were colonized by T. piniperda with 2.0 (i 0.71) attacks per infested log. Three of the
four logs were in a jack pine stand, with 2.3 (i 0.88) attacks per log. The fourth log,
which had only one T. piniperda gallery, was located in a Scotch pine stand.

In 1998, 28 logs were colonized by T. piniperda with 3.5 (i 0.60) attacks per
infested log. Sixteen of the 28 pine logs were in the two red pine stands and there were
3.3 (:1; 0.81) attacks per log. T. piniperda colonized 8 logs in Scotch pine stands with 2.5
(i 0.89) attacks per log. Only 4 logs were colonized by T. piniperda in jack pine stands
with 6.0 (i 1.78) attacks per log.

Scotch pine logs were consistently colonized by T. piniperda more frequently

than red or jack pine logs regardless of the stand species. Thirteen of the 28 logs that

17

were colonized by T. piniperda in 1998 were Scotch pine logs and had 4.7 (i 1.09)
attacks per log. T. piniperda also colonized 8 jack pine logs, with 2.5 (1- 0.73) attacks per
log, and 7 red pine logs, with 2.3 (_+_ 0.68) attacks per log. In jack pine stands, two Scotch
pine logs, with 9 attacks each, and 2 jack pine logs, with 3.0 (i 1.00) attacks per log,
were colonized by T. piniperda. In red pine stands in mid-Michigan, 7 Scotch pine logs,
with 4.7 (i 1.64) attacks per log, 5 red pine logs with 2.8 (i 0.86) attacks per log, and 4
jack pine logs, with 1.5 (i 0.29) attacks per log, were colonized by T. piniperda. In
Scotch pine stands, 4 Scotch pine logs, with 2.5 (i 1.19) attacks per log, 2 red pine logs
attacked once each, and 2 jack pine logs, with 4.0 (i 3.00) attacks per log, were colonized

by T. piniperda in mid-Michigan in 1998.

Log Quality

Variables related to log quality, including surface area, bark thickness and phloem
thickness, were regressed on the number of T. piniperda attacks in red, jack and Scotch
pine logs in 1997 and 1998. In 1997 (Table 1.6), all regressions were significant (P <
0.05) and the amount of variation explained ranged from 36 to 56%. Surface area and
bark thickness, respectively, were consistently important predictors of T. piniperda
attacks, contributing to all of the equations except for T. piniperda attacks on jack pine
logs. Predictor coefficients were positive, indicating that T. piniperda more frequently
attacked larger pine logs with thick bark or, in the case of j ack pine, thicker phloem.

In 1998 (Table 1.7), regressions were significant (P < 0. 05), except for T.

piniperda attacks on red pine logs (P > 0.10). The amount of variation explained was

18

relatively low, except for the number of T. piniperda attacks on Scotch pine logs,
probably because the overall number of attacks per log was lower than in 1997. Bark
thickness was an important predictor of T. piniperda attacks, entering three of the four
equations. Predictor coefficients were again positive, indicating that pine logs with thick

bark were colonized more often than logs with thin bark.

19

If “

DISCUSSION

Results from this study show that the distribution of T. piniperda in lower
Michigan forests is not uniform throughout the quarantined area. Established populations
of T. piniperda are present in the pine forests in southwestern lower Michigan, where
they are known to have been established since 1992 and probably at least 7 years earlier
(Carter et al. 1996, Czokajlo et al. 1997). Michigan’s T. piniperda population probably
originated in Ohio (Carter et al. 1996) which explains the relatively high density in
southwestern lower Michigan. My data appear to document the expansion of T.
piniperda populations into mid-Michigan pine stands. Although mid-Michigan counties
have been quarantined for T. piniperda since 1994 and 1996 (NAPIS 2000), only a few
beetles colonized logs in two stands and no beetles were collected in funnel traps in 1997.
In 1998, the number of T. piniperda galleries in my stands increased twelve-fold, and
colonized logs were collected from 6 stands. F rom 1997 through 1999, no evidence of T.
piniperda was found in logs or funnel trap catches in northwestern counties, which were
quarantined in 1996. However, most of the pine in Michigan is in these northern counties
(V oss 1972, Rudolf 1990, Rudolph and Laidly 1990, Skilling 1990, Leatherberry and
Spencer 1996), so the impact of T. piniperda may potentially increase if populations
continue to expand.

I hypothesized that T. piniperda would preferentially select Scotch pine logs when
red and jack pine logs were also available in pine stands because of the long association
between Scotch pine and T. piniperda in Europe (Bakke 1968, Salonen 1973, Langstrtim

1980a, 1980b, Eidmann 1992). Overall, Scotch pine logs were attacked more frequently

20

than red and jack pine logs in both years, regardless of the species of pine stand in which
they were placed. In southwestem lower Michigan where T. piniperda populations were
high in both 1997 and 1998, Scotch pine logs were consistently attacked more frequently
than red and jack pine logs. Even at low densities, such as in mid-Michigan in 1998,
Scotch pine logs were attacked by T. piniperda more often than red and jack pine logs.
My results also indicate that T. piniperda exhibit a preference for pine logs with
thick bark, a pattern consistent with other studies (Bakke 1968, Salonen 1973, Langstrbm
1980a, 1980b). I suspect that this preference is because the phloem resource is less likely
to dry out in thick-barked logs than in thin-barked logs during T. piniperda larval
development. Thick, furrowed bark may also improve survivorship of T. piniperda
adults during colonization of brood material by increasing the foraging time of predators
such as Thanasimus dubius (F .) (Coleoptera: Cleridae) and Cucujus clavipes F.
(Coleoptera: Cucujidae). Additionally, T. piniperda coevolved on Scotch pine in
Scandinavia with a closely related bark beetle, T omicus minor (Hartig), which colonizes
the upper portions of the trees (Bakke 1968, Salonen 1973, Lekander et al. 1977,
Lingstrdm 1980a, Markalas 1997). The coevolution and resource partitioning between
these two European bark beetles is likely to have strongly influenced T. piniperda’s
preference for thick-barked Scotch pine. Therefore, large, thick-barked Scotch pine logs
should be used for trap logs during detection surveys for T. piniperda. Thick-barked
Scotch pine trap logs placed in the spring are likely to attract T. piniperda and, if
destroyed by late April, may be an effective method of reducing populations and the
potential impact of T. piniperda (Haack and Lawrence 1997, McCullough and Sadof

1998).

21

Scolytid species that aggressively attack and kill healthy trees usually rely on a
multiple component aggregation pheromone to mass attack and overwhehn a tree’s
defenses, and are not strongly attracted by host volatiles, if at all (Byers 1989).
Secondary bark beetles, such as T. piniperda, which colonize recently killed, dying or
decaying trees, however, generally are strongly attracted to host volatiles and ethanol
(Klimetzek et al. 1986, Schroeder 1988, Schroeder and Lindeldw 1989) and less ofien use
an aggregation pheromone. Synergism between various monterpenes and ethanol have
been reported for secondary bark beetles (Tilles et al. 1986, Vité et al. 1986, Phillips et al.
1988, Volz 1988, Schroeder 1988, Chénier and Philogene 1989, Schroeder and LindelOw
1989, Phillips 1990, Czokajlo and Teale 1999).

I suspect that T. piniperda preferentially selects Scotch pine logs over red pine
and jack pine logs due to differences in pine oleoresin chemistry, but more research is
needed to address this. Future research should address how ratios of monoterpenes and
ethanol vary as logs dry out and how this affects attractiveness of logs to T. piniperda to
understand more about the sensory stimuli of T. piniperda. The exotic T. piniperda may
be detected more effectively and better managed with this information as it disperses and

becomes established in North American forests.

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Figure 1.1. Michigan counties that are shaded were quarantined for

T 0micus piniperda as of April 2000. Stands used as field sites were
located in Benzie county (a) in northwestern lower Michigan, Clare (b)
and Gladwin (0) counties in mid-Michigan, and Allegan ((1), Barry (6)
and Kalamazoo (0 counties in southwestern lower Michigan.

30

 

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1.... “fl

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I]:

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35

l "-'-|
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36

CHAPTER 2

HOST PREFERENCE OF T OMIC US PINIPERDA (L.) (COLEOPT ERA:

SCOLYTIDAE) PARENT ADULTS AND SHOOT-FEEDING PROGENY ADULTS

INTRODUCTION

The common pine shoot beetle, T omicus piniperda (L.) (Coleoptera: Scolytidae),
is a native scolytid of Europe, Asia, and North Africa (Bakke 1968, Ye 1991, Eidmann
1992). T. piniperda was first discovered in North America in 1992, near Cleveland, Ohio
(Haack and Kucera 1993, Lawrence and Haack 1995, Haack et a1. 1997). As of April
2000, T. piniperda had been found in at least 295 counties in 11 north central and
northeastern states (National Animal and Plant Inspection Service [NAPIS] 2000) and 32
counties in the Canadian provinces of Ontario and Quebec (C. Markham, USDA Animal
and Plant Health Inspection Service [APHIS], pers. comm). Federal and state
quarantines were imposed to regulate the transport of pine logs from infested counties
and the processing of these logs in uninfested counties (APHIS 1993).

Biology of T. piniperda differs from that of most North American conifer bark
beetles in that T. piniperda adults require maturation feeding in healthy pine shoots
(Hanson 1940, Bakke 1968, Lingstrom 1980a, 1980b, 1983). In spring, T. piniperda
parent adults use alpha-pinene and other host volatiles to locate recently cut pine logs,
broken t0ps, or slash to colonize for oviposition. Larvae feed and develop in the phloem

for 6 to 12 weeks, then pupate. Progeny adult beetles emerge from pine brood material in

37

June, fly to healthy pine trees and feed in the pith of shoots until autumn (Lawrence and
Haack 1995). In Scandinavia, T. piniperda shoot-feeding has caused significant growth
loss in trees near areas where brood material was plentiful (Andersson 1973, Nilsson
1974, 1976, Lingstrom and Hellqvist 1990, 1991, Eidmann 1992). T. piniperda is also
capable of exploiting severely stressed pines as brood material. In Europe and Asia, pine
stands stressed by severe defoliation (Amezaga 1997), years of drought (Ye 1991),
disease (Kaitera and J alkanen 1994), or poor management (Kaplan and Mokrzycki 1988,
Amezaga 1997) can be colonized and subsequently killed by this pest.

In Europe, Scotch pine (Pinus sylvestris L.) is the natural host of T. piniperda.
Scotch pine is widely planted for windbreaks, in Christmas tree plantations, and in forest
stands in Michigan, throughout much of the north central and northeastern regions of the
United States, and in areas of eastern Canada (Wright et al. 1976, Michigan Agricultural
Statistics Service 1999). In North America, red pine (Pinus resinosa Ait.), jack pine
(Pinus banksiana Lamb.) and eastern white pine (Pinus strobus L.) are the dominant
native pines within the area currently known to be infested by T. piniperda. T. piniperda
can breed and shoot-feed in these native pines, though white pine is only marginally
acceptable (Sadof et al. 1994, Lawrence and Haack 1995, Haack and Lawrence 1997).

I hypothesized that T. piniperda parent and progeny adults would preferentially
colonize and shoot-feed in Scotch pine over native red and jack pines. I conducted
laboratory trials to determine whether 1) T. piniperda parent adults preferentially selected
Scotch pine logs over red and jack pine logs, and 2) T. piniperda progeny adults

preferentially selected current-year Scotch pine shoots over red and jack pine shoots.

38

METHODS AND MATERIALS

Overwintered Parent Adults

T. piniperda were collected from infested Scotch pine stands in Allegan, Ingharn,
and Kalamazoo counties in southern lower Michigan using Lindgren multiple funnel
traps (Lindgren 1983, Lindgren 1997) baited with alpha-pinene lures (PheroTech Inc.,
Delta, British Columbia) during the spring dispersal flight of overwintered adults in late
March and early April of 1997, 1998 and 1999. However, beetles collected in 1998 were
not tested because of high mortality attributable to a pathogen that spread through my
laboratory colony before the experiments could be conducted. Also, an insufficient
number of beetles were collected in 1997 to test all pine combinations, so the preference
between red and jack pine logs was not examined the first year of this study.

Freshly cut red, jack and Scotch pine logs, 15 to 18 cm in diameter, were
collected from Kalamazoo county (W. K. Kellogg Experimental Forest, Michigan State
University) in early March. Shortly before conducting the experiment, logs were cut to
lengths of 46 cm and lightly trimmed with a drawknife to remove some layers of the
outer bark, but the inner bark was unaffected. This ensured that the logs appeared
visually similar and T. piniperda choices were not confounded by external bark features
such as color or texture (Bakke 1968, Salonen 1973, Lingstrdm 19803). Therefore,
beetle selection of a pine host was presumably based on the chemical composition of the
volatile plumes emitted from each pine. Bolts of two of the three pine species were

placed in a 61 x 61 x 61 cm aluminum framed collapsible cage with 16 x 18 mesh

39

| I l

aluminum screening (BioQuip Products, Inc., Gardena, CA), then placed in a wind tunnel
so the chemical cues from each bolt would not homogenize. New screen cages were used
for each trial to avoid possible contamination between different pine species. Before
conducting the experiments, the location and intersection of the chemical plumes emitted
from the bolts was determined using a cold smoke produced by a small amount of
titanium tetrachloride on cotton swabs. Bolts were placed 12 cm from the upwind side of
the screen cage and 15 cm apart. During the experiments, beetles were placed near the
downwind side of the screen cage where the volatile plumes began to merge, enabling the
beetles to experience the chemical cues from each bolt. Wind tunnel experiments were
conducted under the following conditions: air temperature of 20°C, wind-speed of 20
cm/sec, and light intensity of 40,000 to 45,000 1x. Each test was conducted with 20 to 30
female beetles. Only female T. piniperda were used in the experiments because they are
the pioneering gender, which initiate galleries. Each beetle was used only once. In 1997,
each trial was conducted twice. In 1999, the Scotch pine/red pine trial was conducted 8
times and the other trials were conducted 5 times each. Tests were run for 8 h, then the
bolts were removed and the number of beetles boring into the bark of each bolt were

counted. Beetles that did not select a pine host were recorded also.

Shoot-Feeding Progeny Adults

T. piniperda progeny adults were collected within 24 h of emergence from caged,

infested red pine brood logs in June 1997 and Scotch pine brood logs in June 1999 from

W. K. Kellogg Experimental Forest in Kalamazoo county. In 1997, groups of 10 progeny

40

 

beetles were placed in the center of plastic boxes (31.5 cm wide, 42.5 cm long, 34 cm
high) with mesh covered ventilation holes in each side, 24 to 36 h after their emergence
from logs. Two current-year pine shoots of different species were placed on two
opposing sides in each cage. In 1999, groups of 20 progeny beetles were placed in the
center of 61 x 61 x 61 cm alumintun framed collapsible cages with 16 x 18 mesh
alumintun screening, 24 to 36 h alter their emergence (BioQuip Products, Inc., Gardena,
CA). Similarly-sized piles of 12 to 15 current-year pine shoots were placed on two
Opposing sides of each cage. In both 1997 and 1999, pine shoots in each trial were of
similar length and diameter, and were clipped from healthy, young trees at the start of
each bioassay. The appearance and phenology of the pine shoots were intentionally
similar among species to avoid possibly biasing host selection by the progeny adults.
Cages were placed in well-ventilated areas void of any other pine terpenes. One cage
was placed in a wind trmnel, one in a fume hood, and 3 were placed outdoors in a mature
hardwood stand with no coniferous vegetation within 1 km. Each trial was replicated 37
times in 1997. In 1999, the Scotch pine/j ack pine trial was replicated 26 times and the
other trials were replicated 14 times each. Bioassays were run for 48 h, then shoots were
removed and carefully examined for evidence of beetle attacks. Number of tunnels
initiated by beetles was recorded. Beetles that were not feeding in the shoots, either dead
or alive, were recorded also. Each beetle was only used once in these choice
experiments.

In 1997, diameter of each pine shoot was measured with calipers and plotted
against fi'equency of T. piniperda attack. Based on these results, only shoots with

diameters of 0.4 to 0.5 cm were used in 1999.

41

[rm-‘1

Statistical Analyses

Between-year differences in proportion of attacks by parent and progeny beetles
were examined using G-tests of independence (Sokal and Rohlf 1995). Between-year
differences were not significant for parent beetles in the Scotch pine/red pine log trial (G
= 0.62, P > 0.10) or the Scotch pine/jack pine log trial (G = 0.05, P > 0.10), so data were
pooled for the remaining comparisons. The red pine/jack pine log trial testing the host
preference of parent beetles was conducted in 1999 only.

Between-year differences were not significant for progeny beetles (Scotch
pine/jack pine shoot trial G = 0.03, P > 0.10; Scotch pine/red pine shoot trial G = 1.41, P
> 0.10), except for the red pine/jack pine shoot trial (G = 22.43, P < 0.001). Data were
pooled for the remaining comparisons.

Pearson’s ,2} goodness of fit statistic (Sokal and Rohlf 1995) was used to
determine if T. piniperda parent adults preferentially discriminated between species of
pine logs and whether progeny adults discriminated between species of pine shoots. All

analyses were conducted at P < 0.05 level of significance.

42

RESULTS AND DISCUSSION

Overwintered Parent Adults

In Europe, Scotch pine is the preferred host for T. piniperda (Bakke 1968,
Salonen 1973, Langstrdm 1980a, Schroeder 1988, Gibbs and Inman 1991). Although T.
piniperda is capable of colonizing and developing in many North American pines
(Chararas 1968, Ohmart 1980, Langstrom and Hellqvist 1985, Zumr 1992, Sadof et al.
1994, Lawrence and Haack 1995, Amezaga 1996, 1997), my results indicate that it may
preferentially colonize its European host, Scotch pine, when given a choice.

In 1997, 98 out of 114 T. piniperda parent adults successfully began colonizing
one of the two pine logs they were exposed to, while 16 beetles died without selecting a
host (Table 2.1). No beetles were found alive without selecting a pine host in 1997. In
1999, 414 out of 523 beetles successfully began colonizing a log, 78 beetles died without
selecting a host, and 31 beetles were alive but had not made a choice between the two
given pine species within 8 h (Table 2.1).

Scotch pine logs were attacked with significantly greater frequency than jack pine
logs (j = 13.09, P < 0. 05) or red pine logs (f = 16.79, P < 0.05) in two-choice trials
conducted in a wind tunnel (Table 2.1). In 1997 and 1999, approximately two-thirds of
T. piniperda parent adults selected Scotch pine logs over red and jack pine logs (Figures
2.1 and 2.2). Scotch pine logs were selected 63% of the time when paired with red pine
logs and 64% of the time when paired with jack pine logs. There was not a significant

difference in the frequency of attacks in the 1999 trials between red and jack pine logs (j

43

 

= 1.09, P > 0.10) (Table 2.1, Figure 2.3). It is likely that beetles in my host preference
test responded primarily to volatiles because I ensured that log size, silhouette, and bark
characteristics were kept similar in wind tunnel experiments.

During colonization of brood material, T. piniperda parent adults use host tree
volatiles to locate suitable pine logs and stumps (Byers 1989). T. piniperda is primarily
' attracted to alpha-pinene, a monoterpene in pine oleoresin (Byers et al. 1985, Schroeder
1988, Schroeder and Lindelc‘iw 1989). There is no known aggregation pheromone for T.
piniperda (Schonherr 1972, Byers et al. 1985, 1989, Lanne et al. 1987, Loyttyniemi et al.
1988, Volz 1988), although the presence of a short distance female-produced sex
pheromone has been proposed (Czokaj lo 1998). This pheromone is not likely to have
influenced my results, however, because only T. piniperda females were used in my host
preference trials. Further evaluation of differences in pine oleoresin chemistry may

improve our ability to detect new T. piniperda populations or increase trapping efficacy.

Shoot-Feeding Progeny Adults

In 1997, 382 out of 911 progeny beetles successfully initiated maturation feeding
tunnels in pine shoots while 529 of the progeny beetles tested died before establishing
tunnels (Table 2.2). In 1999, 1052 out of 1454 progeny beetles successfully initiated
maturation feeding tunnels in pine shoots (Table 2.2), while 242 beetles died and 160
were alive but had not begun shoot-feeding afier 48 h. Beetles used in shoot-feeding tests

in 1997 emerged from red pine logs, while beetles used in 1999 developed in Scotch pine

logs. The relatively high survival of beetles in 1999 suggests that Scotch pine may have
been somewhat more suitable for T. piniperda development than red pine.

Jack pine shoots were attacked with significantly greater frequency when paired
with Scotch pine shoots (j = 15.49, P < 0.05), while Scotch pine shoots were
significantly preferred over red pine shoots (f = 52.20, P < 0.05) (Table 2.2). In 1997,
jack and red pine shoots were attacked at the same frequency when presented together (1}
= 0.01, P > 0.10), while in 1999, jack pine shoots were significantly preferred over red
pine shoots (j = 59.24, P < 0. 05) (Table 2.2).

Host preference of insects may be potentially modified by the host on which they
feed or develop as larvae (Smith and Cornell 1979, Rausher 1983, Singer 1983, Vet et al.
1984, Vet and Van Opzeeland 1986, Pennacchio et al. 1994). This larval conditioning
may be a modifying factor of host preference and was not directly tested in this
experiment. However, the T. piniperda progeny beetles that were reared from red pine
logs in 1997 and those reared fiom Scotch pine logs in 1999, had similar ratios for the
Scotch pine/red pine trials (Figure 2.4) and Scotch pine/jack pine shoot trials (Figure 2.5).
Results of the red pine/j ack pine shoot trial differed between years (Figure 2.6). This
may indicate that when larvae developed in red pine logs in 1997, they were more likely
to accept red pine shoots for maturation feeding. This result, however, could also be just
an artifact of the small sample size used in 1997. More research is needed to evaluate
effects of larval conditioning on subsequent host selection by T. piniperda adults.

Physical traits of shoots, along with host volatiles, may also influence T.
piniperda progeny adults. Approximately 70% of all the shoots attacked by T. piniperda

in my trials had a diameter of 0.4 to 0.5 cm and shoots that were 0.45 cm were most often

45

attacked (Figure 2.7). This pattern is consistent with other studies conducted in the
United States and Europe (Léngstrom 1980a, 1980b, Lawrence and Haack 1995,
McCullough and Smitley 1995).

In mixed-species pine stands at Indiana Dunes State Park, Sadof et al. (1994)
collected more T. piniperda-damaged red and Scotch pine shoots than expected based on
the respective basal area of all pines in the park. They suggested that red pine was more
susceptible to shoot-feeding by T. piniperda than other native pines. Shoot diameters,
however, were not examined. T. piniperda progeny adults may select shoots primarily
based on physical traits, such as diameter, in mixed-stands where the host volatiles are
homogenous in the air. Lawrence and Haack (1995) found that T. piniperda initiated
feeding tunnels more frequently when individually caged on red pine than when beetles
were caged on jack pine. Survival of beetles during the summer, however, was greater on
Scotch pine than on the native pines. Beetles were more likely to be pitched out and die
when caged on red pine than on the other pines (Lawrence and Haack 1995).

In many situations, the distribution of T. piniperda progeny may strongly reflect
the choice of the parent adults, in that the brood material from which the progeny emerge
will often be of the same species as nearby healthy trees that are available for shoot-
feeding (Butovitsch 1972, Léngstrdm 1979, Lutyk 1984, Legowski 1987, Sauvard et al.
1987, Ye and Li 1994). Shoot-feeding, in turn however, may determine where beetles
overwinter, thus affecting where the parent beetles are the following spring (Andersson
1973, Langsz and Hellqvist 1990, 1991). Additionally, the quality of brood material
is known to influence body weight and lipid content of bark beetle progeny (Amman

1971, Salonen 1973, Schmid 1972, Slansky and Haack 1986, Anderbrant and Schlyter

46

I"!

1989), which may affect viability or dispersal ability of progeny beetles (Anderbrant and
Schlyter 1989). As T. piniperda progeny adults continue to shoot-feed throughout the
summer though, their dispersal capabilities increase (Sauvard et al. 1987).

To-date, the only area in North America where T. piniperda is known to have
caused significant shoot damage to native pines is in southern Ontario. Each area with T.
piniperda damage is dominated by dead and dying stands of unmanaged Scotch pine (E.
Czerwinski and T. Scarr, Ontario Ministry of Natural Resources, pers. comm.) My
observations of these stands and discussions with Ontario pest managers suggest that T.
piniperda may have entered the stands secondarily, colonized the newly dead and dying
trees, and progeny beetles subsequently moved into adjacent or nearby stands of native
pines that were in better health to shoot-feed (N. W. Siegert and D. G. McCullough,
unpublished data). The consistent association of high T. piniperda populations with
Scotch pine brood material in Ontario is consistent with results of these laboratory
experiments.

Scotch pine may, therefore, be considered a high risk species, especially if dying
trees, broken branches or other forms of brood material are present. Managers should
perhaps emphasize activities to detect or monitor T. piniperda in these high-risk stands,
especially if jack pine is nearby, since significant shoot-feeding may be sustained in these
trees.

Additional research is needed to identify compounds used by T. piniperda to
differentiate among pine species and to understand more about the sensory stimuli of T.
piniperda. This information may be useful in determining how to better manage this

exotic scolytid in North American forests.

47

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Lindgren, B. S. 1983. A multiple funnel trap for scolytid beetles (Coleoptera). Canadian
Entomologist. 115: 299-302.

Lindgren, S. R. 1997. Optimal release rate of the host monoterpene alpha-pinene for
trapping the European pine shoot beetle, T omicus piniperda (Coleoptera:
Scolytidae). Proceedings of the Entomological Society of Ontario. 128(0): 109-
1 1 1.

59

Léyttym'emi, K., K. Heliovaara, and S. Repo. 1988. No evidence of a population
pheromone in T omicus piniperda (Coleoptera: Scolytidae): a field experiment.
Annales Entomologici Fennici. 54(3): 93-95.

Lutyk, P. 1984. Feeding of T omicus piniperda on shoots of Norway spruce and silver
fir. Sylwan. 128(3): 65-68. English summary.

McCullough, D. G. and D. R. Smitley. 1995. Evaluation of insecticides to reduce
maturation feeding by T omicus piniperda (Coleoptera: Scolytidae) in Scotch pine.
Journal of Economic Entomology. 88(3): 693-699.

Michigan Agricultural Statistics Service (MASS) Database. 2000. <http://www.mda.
state.mi.us/mass/mi_nurs97/>

National Animal and Plant Inspection Service (NAPIS) Database. 2000. <http://www.
cercis.purdue.edu/napis/pests/psb/mgif/psbcnty.gif/>

Nilsson, S. 1974. Increment losses caused by T omicus piniperda on Scots pine.
Rapporter och Uppsatser, Institutionen for Skogsteknik. 78(14): 1-64. English
summary.

Nilsson, S. 1976. Rationalization of forest operations gives rise to insect attack and
increment losses. Ambio. 5(1): 17-22.

Ohmart, C. P. 1980. Insect pests of Pinus radiata plantations: present and possible
future problems. Australian Forestry. 43(4): 226-232.

Pennacchio, F., M. C. Digilio, E. Tremblay, and A. Tranfaglia. 1994. Host recognition
and acceptance behaviour in two aphid parasitoid species: Aphidius ervi and
Aphidius microlophii (Hymenoptera: Braconidae). Bulletin of Entomological
Research. 84(1): 57-64.

Rauscher, M. D. 1983. Conditioning and genetic variation as causes of individual
variation in the oviposition behavior of the tortoise beetle, Deloyala guttata.
Animal Behavior. 31(3): 743-747.

Sadof, C. S., Waltz, R. D., and C. D. Kellam. 1994. Differential shoot feeding by adult
T omicus piniperda (Col.: Scolytidae) in mixed stands of native and introduced
pines in Indiana. Great Lakes Entomologist. 27(4): 223-228.

Salonen, K. 1973. On the life cycle, especially on the reproduction biology of
Blastophagus piniperda L. (Col., Scolytidae). Acta Forestalia Fennica. 70pp.

Sauvard, D., F. Lieutier, and J. Levieux. 1987. Spatial distribution and dispersal of

T omicus piniperda L. (Coleoptera, Scolytidae) in the forest of Orleans (France).
Annales des Sciences Forestieres. 44(4): 417-434. English summary.

60

Schmid, J. M. 1972. Emergence, attack densities and seasonal trends of mountain pine
beetle (Dendroctonus ponderosae) in Black Hills. USDA Forest Service
Research Note. 211: 7pp.

Schonherr, J. 1972. Pheromone in the pine bark beetle Myelophilus piniperda (Col.,
Scolytidae). Zeitschrift fur Angewandte Entomologie. 71(4): 410-413. English
summary. '

Schroeder, L. M. 1988. Attraction of the bark beetle T omicus piniperda and some other
bark- and wood-living beetles to the host volatiles a-pinene and ethanol.
Entomologia Experimentalis et Applicata. 46(3): 203-210.

Schroeder, L. M. and A. Lindelow. 1989. Attraction of scolytids and associated beetles
by different absolute amounts and proportions of alpha-pinene and ethanol.
Journal of Chemical Ecology. 15(3): 807-817.

Singer, M. C. 1983. Determinants of multiple host use by a phytophagous insect
population. Evolution. 37(2): 389-403.

Slansky, F., Jr., and R. A. Haack. 1986. Age-specific flight behavior of Ips calligraphus
(Coleoptera: Scolytidae) reared in slash pine bolts with thick versus thin inner
bark (phloem). Entomologia Experimentalis et Applicata. 40(2): 197-207.

Smith, M. A. and H. V. Cornell. 1979. Hopkins host-selection in Nasonia vitripennis
and its implications for sympatric speciation. Animal Behavior. 27(2): 365-370.

Sokal, R. R. and F. J. Rohlf. 1995. Biometry, 3rd ed. Freeman, New York.

Vet, L. E. M., C. Janse, C. Van Acterberg, and J. J. M. Van Alphen. 1984. Microhabitat
location and niche segregation in 2 sibling species of Drosophilid parasitoids:
Asobara tabida and Asobara rufescens (Braconidae: Alysiinae). Oecologia.
61(2): 182-188.

Vet, L. E. M. and K. Van Opzeeland. 1986. Olfactory microhabitat selection in
Leptopilina heterotoma (Hymenoptera: Eucoilidae), a parasitoid of
Drosophilidae. Netherlands Journal of Zoology. 35(3): 497-504.

Volz, H. A. 1988. Monoterpenes goveming host selection in the bark beetles Hylurgops
palliatus and T omicus piniperda. Entomologia Experimentalis et Applicata.
47(1): 31-35.

Wright, J. W., W. A. Lemmien, J. N. Bright, M. W. Day, and R. L. Sajdak. 1976. Scotch

pine varieties for Christmas tree and forest planting in Michigan. Michigan
Agricultural Experiment Station, Research Report 293. East Lansing. 15pp.

61

Ye, H. 1991. On the bionomy of T omicus piniperda (L.) (Col.: Scolytidae) in the
Kunming region of China. Journal of Applied Entomology. 112(4): 366-369.

Ye, H. and L. S. Li. 1994. The distribution of a T omicus piniperda (L.) population in the
crown of Yunnan pine during the shoot feeding period. Acta Entomologica
Sinica. 37(3): 311-316. English summary.

Zumr, V. 1992. Attractiveness of introduced conifers to xylophagous beetles and their
acceptance. Journal of Applied Entomology. 113(3): 233-238.

62

CHAPTER 3

SURVEY OF SHOOT DAMAGE CAUSED BY T OMIC US PINIPERDA (L.)

(COLEOPTERA: SCOLYTIDAE) IN MICHIGAN PINE PLANTATIONS

INTRODUCTION

The exotic pine shoot beetle, T omicus piniperda (L.) (Coleoptera: Scolytidae), is a
native bark beetle in Europe and Asia (Bakke 1968, Ye 1991, Eidmann 1992) that was
discovered in North America in 1992 near Cleveland, Ohio (Haack and Kucera 1993,
Lawrence and Haack 1995, Haack et al. 1997). As of April 2000, T. piniperda has been
found to occur in 74 counties in Michigan (Figure 4.1), at least 221 additional counties in
10 other north central and northeastern states (National Animal and Plant Inspection Service
[NAPIS] 2000), and 32 counties in the Canadian provinces of Ontario and Quebec (C.
Markham, USDA Animal and Plant Health Inspection Service [APHIS], pers. comm.)
Federal and state quarantines were imposed in 1992 to regulate the transport of pine logs
out of infested counties and the processing of these logs at mills in uninfested counties
(APHIS 1993). These regulations have negative economic implications for mills located in
uninfested counties that receive pine from quarantined counties. The majority of research
on T. piniperda in North America to-date has been conducted in Scotch pine (Pinus
sylvestris L.) Christmas tree plantations. The impact of T piniperda in forest stands needs

to be addressed to form regulatory and management policies in Michigan and nationwide.

63

Bakke (1968) and Léngstrém (1980a, 1980b, 1983) described the univoltine life
cycle of T. piniperda. Overwintering adult beetles become active in early spring, when
temperatures exceed 12° C. Adult beetles use pine volatiles to locate suitable brood
material including highly stressed or recently cut pine trees, stumps, or slash (tops, branches
and other debris left alter logging operations or storms). Beetles bore into the phloem and
lay eggs that hatch in 1 to 2 weeks. Larvae feed and develop in the phloem of the brood
material for 6 to 12 weeks, pupate, and progeny adults emerge in early summer. These
reddish-brown progeny adult beetles fly to live pine trees and tunnel down the center of
current-year shoots. This maturation feeding continues throughout the summer and
beetles darken to a shiny black color. Each beetle can feed in 2 to 6 pine shoots.
Researchers found that T. piniperda can breed and shoot-feed in all native pine species in
the Lake States including jack pine (Pinus banksiana Lamb), red pine (Pinus resinosa
Ait.), and white pine (Pinus strobus L.), although white pine is only marginally acceptable
(Lawrence and Haack 1995, Sadof et al. 1994). Tunneled shoots eventually die and fall
from the tree (Lingstrom and Hellqvist 1990, 1991, McCullough and Smitley 1995),
although the stout shoots on red pine may not always drop or fall from the tree during the
same year of attack (Kauffman et al. 1998). In autumn, beetles leave the shoots and
overwinter in the bark at the base of the trees. High populations of T. piniperda can
reduce tree growth and even cause tree mortality (Eidmann 1985, Lingstrom and
Hellqvist 1985, Ye 1991). Unlike T. piniperda, most native bark beetles complete
maturation before emerging fi'om the brood material, and do not shoot-feed.

A complex of other native insects in the Lake States, however, do cause damage to

shoots of pine trees. These include: cone beetles [(Conopthorus resinosae Hopkins and

64

Conopthorus banksiana McPherson) (Coleoptera: Scolytidae)], eastern pine shoot borer
[(Eucosma gloriola Heinrich) (Lepidoptera: Tortricidae)], red pine shoot moth [(Dioryctria
resinosella Grote) (Lepidoptera: Pyralidae)], and European pine shoot moth [(Rhyacionia
buoliana Schiff.) (Lepidoptera: Tortricidae)] (McPherson et al. 19703, 1970b, DeBoo et
al. 1971, Bright 1976, Wilson 1977, Hainze and Benjamin 1983, Katovich and Hall
1992). Little information is currently available on the frequency and extent of shoot-
feeding by these insects, and whether damage might be enough to affect growth rates of
trees.

The objectives of this study were to survey Scotch, red, and jack pine forest
plantations throughout lower Michigan to: (1) quantify the frequency and extent of shoot
damage attributable to T. piniperda versus native shoot-feeding insects, weather and other
damaging agents, (2) compare shoot damage in southwestern lower Michigan pine stands
to shoot damage in northern lower Michigan pine stands, and (3) to evaluate the potential

threat that T. piniperda poses to Michigan’s pine resource.

65

METHODS AND MATERIALS

Shoot Injury Diagnosis

Accurate diagnosis of shoot damage and identification of the causal agent were
based on specific distinguishing characteristics observed on the pine shoots. Shoots
damaged by T. piniperda are typically 10 to 15 cm in length, have a 2 mm diameter
hollowed pith that is free of fi‘ass and debris, and often have a resin tube surrounding the
entrance hole. Also, there is frequently more than one hole and tunnel per shoot. Cone
beetles generally shoot-feed at the distal tip of the shoot behind the bud, causing the shoot
tips to break off (Bright 1976, Wilson 1977). Their tunnels are 1.0 to 1.5 mm in diameter
and are also free of frass and debris. These shoot tips are usually still green when they
break off in the autumn. There is some question as to whether red pine cone beetle and the
closely related jack pine tip beetle are separate species (DeGroot et al. 1992). In this
survey, I did not attempt to distinguish between shoots injured by these two native
scolytids. The feeding tunnels of eastern pine shoot borer larvae are 10 to 30 cm long
with an abundant amount of frass packed in the gallery (DeBoo et al. 1971). The larvae
hollow out most of the pith in the shoot and emerge through a distinctive exit hole that is
3.2 to 6.4 mm in diameter (Wilson 1977). Shoots die and break cleanly off the tree, often
appearing to have been clipped off. Shoot damage caused by red pine shoot moth larvae
is similar to eastern pine shoot borer larvae except shoots tend to be shorter and have a
characteristic pitch mass at the point of attack (Hainze and Benjamin 1983, Katovich and

Hall 1992). European pine shoot moth prefers red pine but will readily attack Scotch and

66

Austrian (Pinus nigra Arnold) pines; it rarely injures jack and eastern white pine (Wilson
1977). After needle mining for a brief period, larvae injure shoots and produce
characteristic pitch-covered webs at the base of the buds where they feed (Wilson 1977).
Shoots may also be damaged from non-insect causes, such as squirrels, high
winds, and the weight of snow and ice (Wilson 1977). Non-insect damaged shoots
usually appear to be gnawed on, snapped off, or the base of the shoot appears to be torn
out of the branch. The absence of characteristic insect signs, such as tunnels, frass, and

emergence holes, aid in diagnosing shoots damaged by non-insect related factors.

Autumn 1997 and Spring 1998 Damage Surveys

In August and September 1997, ten Scotch, red, and jack pine forest stands (30
total), with no thinning or harvest activity in the past 5 years, were selected from Michigan
Department of Natural Resources (MI DNR) inventory database (Table 4.1 and Table 4.2).
Stand selection was initially based on the following criteria: at least 2.02 ha (5 ac), diameter
at breast height (DBH) > 10 cm (4 in), and basal area (BA) of 23 — 34.5 mZ/ha (100 — 150
ftZ/ac) (well-stocked) or (second choice) of a BA greater than 34.5 mz/ha (> 150 fiz/ac)
(over-stocked). A limited number of Scotch pine stands are managed by the MI DNR and
were available for this study. I included these stands in my survey but most ranged in size
fi'om 0.2 to 4.8 ha (Tables 4.1 and 4.2). Fifteen stands, 5 of each species, were selected in
southwestern lower Michigan (Table 4.1) in Allegan, Cass, and Kalamazoo counties where
T. piniperda populations are known to have existed since at least 1993 (Figure 4.1). Fifteen

additional stands were selected in northern lower Michigan (Table 4.2) in Benzie, Clare,

67

Gladwin, and Manistee counties (Figure 4.1). These counties were quarantined for T.
piniperda in 1994 (Clare Co.), 1996 (Gladwin Co. and Manistee Co.) and 1997 (Benzie
Co.), but related research has indicated that T. piniperda populations were either very low
or not sufficiently established in the forest ecosystems in these areas to be found (N. W.
Siegert, A. Kennedy, and D. G. McCullough, unpublished data).

Each stand was mapped and 100 m transects were randomly located along the
perimeter and within the interior of the stand. Stands were divided into 10 m by 10 m
quadrants which were randomly selected to determine the origin of each transect. The
heading of each transect was then selected along a cardinal direction which allowed the
transect to remain in the stand. Ntunber of transects in each stand was based on stand size.
Stands less than 2.02 ha (5 ac) were divided into 5 m by 5 m quadrants and had 3 transects
along the edge and 2 transects in the interior; stands 2.02 to 4.05 ha (5 to 10 ac) had 4
transects along the edge and 3 transects in the interior; two additional transects were
sampled for every 2.02 ha increase in stand size. The largest stand that was surveyed was
22.8 ha, which I sampled along 25 transects. At 20 m intervals along the transect, the
number of fallen shoots on 1 m2 of the forest floor was counted using a l m2 sampling
frame. Any damaged shoots found were inspected and dissected to determine the cause of
injury. In May 1998, forest stands were re-surveyed along the same transects that were
established in autumn 1997 to determine if more damaged shoots fell from the trees

following the winter snowfalls.

68

Autumn 1998 and Autumn 1999 Damage Surveys

In September and October 1999, new transects were established in randomly
selected localities in the stands to determine the frequency and extent of shoot damage
attributable to T. piniperda versus native shoot-feeding insects, weather and other factors
that occurred in 1998 and 1999. Shoot damage was quantified using the same methods as
in the previous surveys. I was able to distinguish between damage that occurred in the
summer of 1998 and shoots that were damaged during the summer of 1999. New damage
(i.e. damage that occurred in 1999) was differentiated from old damage (i.e. damage that
occurred prior to 1999) based on needle retention, firmness, and the appearance of woody
shoot tissue. Shoots damaged in 1998 had slightly decaying needles with poor retention
and their woody tissues were dark and more decayed than shoots damaged in 1999. Shoots
that were damaged before 1998 were much further decayed, retained fewer needles, and

were not recorded.

Statistical Analyses

Differences in the number of damaged shoots in Michigan pine stands, between
regions and among pine species, were analyzed for each damaging agent for each year using
two-way analysis of variance (ANOVA) (PROC GLM program, SAS Institute, Inc. 1989).
Data were not normally distributed in a few cases and transformations did not correct
normality. Therefore, the number of damaged shoots between regions and pine species

were also analyzed using the non-parametric Kruskal-Wallis tests (Kruskal and Wallis

69

1952) (Appendices A, B, C, and D). Results of the Kruskal-Wallis test were consistent with
ANOVA results in all cases.

When results of the AN OVA were significant, a Fischer’s protected least significant
difference (LSD) test was used to determine which regions or pine species differed. Least
squared means were used in the AN OVA to adjust for the partially unbalanced design of
the survey (Searle et al. 1980, SAS Institute, Inc. 1989). True means are reported in all
tables. All analyses were conducted at P < 0.05 level of significance, using SAS statistical
soflware (SAS Institute, Inc. 1989).

Number of shoots damaged by T. piniperda, native shoot-feeding insects, and non-
insect factors along the perimeter and within the interior of stands did not differ
significantly in 1997, 1998, and 1999, when data were analyzed using ANOVA (P > 0.50),
nor when analyzed by the Kruskal-Wallis test (P > 0.50). These data were pooled for

comparisons between regions and among pine species.

70

 

RESULTS

Damaging Agents

Mean total shoot damage in red, jack and Scotch pine stands in both northern and
southwestern lower Michigan between 1997 and 1999 ranged from 0.3 to 5.5 shoots per
m2 (Table 3.3). In autumn 1997, mean shoot damage ranged fiom 0.5 to 1.5 shoots per
m2. In spring 1998, the mean shoot damage ranged from 0.7 to 3.8 shoots per m2. When
new surveys were conducted in autumn 1999, it was determined that the mean shoot
damage for 1998 ranged from 0.6 to 5.5 shoots per In2 and the mean shoot damage for
1999 ranged from 0.2 to 4.8 shoots per m2.

In northern lower Michigan between 1997 and 1999, jack pine stands consistently
had more shoot damage than red or Scotch pine stands. In southwestern lower Michigan,
red pine stands consistently had less shoot damage than jack or Scotch pine stands (Table
3.3)

Biotic factors causing damage to pine shoots in the stands I surveyed included
insects and larger organisms such as porcupines, squirrels and birds. Insects affected the
shoots by turmeling in and feeding on the pith and other interior tissues; the larger
animals damaged the pine shoots by either chewing off or tearing the shoots from the
trees. Abiotic factors, such as wind, ice and snowfall, also damaged pine shoots.

The insects most commonly found to damage pine shoots in surveys between
1997 and 1999 included T. piniperda, red pine cone beetle, and eastern pine shoot borer

(Table 3.4). Non-insect factors, such as wind, ice, snowfall and squirrels, damaged more

71

 

shoots than insects in all pine stands except for Scotch pine in southwestern lower
Michigan, where T. piniperda caused the most damage (Table 3.4). Between 1997 and
1999, red pine stands in both northern and southwestern regions sustained the least
amount of damage due to non-insect factors and jack pine stands in the north had the
most shoots damaged by non-insect factors. Between 1997 and 1999, T. piniperda
damaged significantly more shoots in Scotch pine stands in the southwest than all other
stands in both regions (P < 0. 05). Between 1997 and 1999, red pine cone beetle damaged
significantly more shoots in northern red pine stands than in all other stands (P < 0. 05),
while eastern pine shoot borer damaged red pine shoots significantly more frequently in
northern than southwestern stands (P < 0. 05). Sphaeropsis tip blight (Sphaeropsis
sapinea (F r.) Dyko and B. Sutton), a fungal pathogen of conifers, and red pine shoot
moth damage were observed on trees in some pine stands but rarely caused damaged pine
shoots to fall on the ground (Table 3.4). European pine shoot moth was one of the most
destructive shoot-boring moths in the Lake States in the past (Wilson 1977), but was not

encountered during surveys between 1997 and 1999.

N on-Insect Damage

Most of the damage to pine shoots in Scotch, red, and jack pine stands in lower
Michigan was caused by weather, such as wind, snow or ice damage, and by squirrels or
other vertebrates (Appendix A). The overall number of shoots damaged by non-insect
factors between 1997 and 1999 in northern and southwestern regions of lower Michigan

differed significantly (P < 0. 05) (Table 3.5). Northern stands sustained significantly

72

more non-insect damage in spring 1998 and autumn 1998 (P < 0. 05) than did the
southwestern stands, but differences between regions were not significant in autumn 1997
and 1999 (P > 0.05).

Overall shoot damage attributable to non-insect factors differed significantly
among jack, red and Scotch pine stands (P < 0. 05) (Table 3.6). In autumn 1997 and
spring 1998, red pine had significantly fewer non-insect damaged shoots than jack pine
(P < 0. 05) or Scotch pine (P < 0. 05). Jack pine stands had significantly more damaged
shoots than Scotch pine stands (P < 0. 05) in autumn 1997. In spring 1998, the amount of
shoots damaged by non-insect factors increased in all stands. In autumn 1998 and 1999,
jack pine stands had significantly more shoots damaged by non-insect factors than red
pine stands (P < 0. 05) or Scotch pine stands (P < 0. 05). Red and Scotch pine stands did
not differ significantly (P > 0. 05) in 1998 or 1999.

Differences in shoot damage by non-insect factors differed among pine species in
both northern and southwestern regions (P < 0. 05) (Table 3.7). In autumn 1997, red pine
in both northern and southwestern stands had significantly fewer non-insect damaged
shoots than jack pine (P < 0. 05) or Scotch pine (P < 0. 05). Jack pine stands had
significantly more damaged shoots than Scotch pine stands (P < 0. 05) in southwestern
stands, but not in northern stands (P > 0. 05) in autumn 1997. In spring 1998, red pine
had significantly fewer non-insect damaged shoots than jack pine (P < 0. 05) or Scotch
pine (P < 0. 05) in northern stands, but not southwestern stands (P > 0. 05). Jack pine
stands had significantly more damaged shoots than Scotch pine stands (P < 0. 05) in
northern stands, but not in southwestern stands (P > 0. 05). In autumn 1998 and 1999,

jack pine stands in both northern and southwestern stands had significantly more shoots

73

damaged by non-insect factors than red pine stands (P < 0. 05) or Scotch pine stands (P <
0. 05). Red and Scotch pine stands did not differ significantly (P > 0. 05) in northern or
southwestern stands in autumn 1998. In autumn 1999, red and Scotch pine stands did not
differ significantly (P > 0. 05) in southwestern stands, but did differ significantly in

northern stands (P < 0. 05).

Tomicus piniperda Damage

In 1997, 1998 and 1999, T. piniperda damaged significantly more shoots in
southwestern than northern lower Michigan (P < 0. 05 ) (Table 3.8 and Appendix B). In
autumn 1997 and spring 1998, T. piniperda was not present in any stands that I surveyed
in northern lower Michigan. In surveys conducted in autumn 1999, a total of 3 shoots
damaged by T. piniperda were found in two Scotch pine stands in Clare and Gladwin
counties in northern lower Michigan.

Overall, the number of shoots damaged by T. piniperda was significantly different
between stand species (P < 0. 05). Scotch pine stands had a greater number of shoots
damaged by T. piniperda than jack pine stands (P < 0. 05) and red pine stands (P < 0.05)
in each survey period from 1997 to 1999 (Table 3.9). Jack pine stands consistently had
more pine shoots damaged by T. piniperda than red pine stands in absolute terms but
differences were not significant (P > 0.05) for any survey. Trends were similar when
region by pine species interactions were examined (Table 3.10). In particular, the number

of Scotch pine shoots damaged by T. piniperda in southwestern lower Michigan

74

increased dramatically in the two stands in Allegan county in 1998 and 1999 (Table

3.11).

Native Shoot-Boring Insect Damage

Red pine cone beetle caused significantly more shoot damage in northern than
southwestern pine stands in autumn 1997 (P < 0. 05), 1998 (P < 0. 05) and 1999 (P <
0. 05) (Table 3.12 and Appendix C). Differences between regions were not significant in
surveys conducted spring 1998 (P > 0. 05). Overall, red pine cone beetle damaged
significantly more red pine shoots than jack pine shoots (P < 0. 05) and Scotch pine
shoots (P < 0. 05) between 1997 and 1999 (Table 3.13). Relatively few jack pine and
Scotch pine shoots were attacked by this insect and differences between jack and Scotch
pine were not statistically significant (P > 0.10). Trends were similar when region by
pine species interactions were examined (Table 3.14).

Eastern pine shoot borer damaged significantly more shoots in northern stands
than southern stands in autumn 1997 (P < 0. 05) (Table 3.15 and Appendix D).
Differences were not significant in spring 1998 (P > 0. 05), autumn 1998 (P > 0. 05) and
autumn 1999 (P > 0. 05), however, overall differences were still significant between
regions (P < 0.05). Scotch and jack pine stands did not sustain any damage by eastern
pine shoot borer while red pine stands were consistently infested (Table 3.16) in the

northern and southwestern regions (Table 3.17).

75

DISCUSSION
Non-Insect Damage
The majority of shoot damage in the pine stands that I surveyed between 1997 and

1999 was attributable to non-insect factors. It is not surprising that most of the shoot
damage in Michigan pine stands was caused by factors other than insects because these I
factors ofien function on a large scale and rarely were beetle populations at high levels. 1
Animals, such as porcupines and squirrels, will typically break or chew off many shoots

over the course of a season, while a shoot-feeding insect may damage only one or, at

most, a few shoots. Abiotic damage from wind, ice storms and snow are common in

Michigan and often affect entire landscapes. Jack pine and Scotch pine, in particular,

have thinner shoots than red pine and may be especially vulnerable to wind, snow or ice

damage. Caches of pine cones, presumably stored by squirrels, were frequently observed,

especially in jack pine stands. This indicates that squirrels were actively gathering cones

in these stands and increases the likelihood that they may be responsible for some of the

damage caused by non-insect factors.

T omicus piniperda Damage

Initially, I hypothesized that there would be more damaged shoots along the edges
of the pine stands compared to the interior of the stands where shoots are more protected

from storm damage and probably less likely to be encountered by dispersing insects.

76

However, the number of damaged shoots did not differ significantly between the interior
and the edges of the pine stands. This may be explained by looking at the stands from a
landscape perspective. Many of the pine stands were located in contiguous woodlands.
Since they were surrounded by hardwoods, there may be less edge effect than if they had
been surrounded by non-wooded land where more storm damage or insect encounters
could have occurred.

The lack of an edge effect may reflect the dispersal behavior and spatial
distribution of T. piniperda. In France, Sauvard et al. (1987) occasionally found localized
concentrations of damaged shoots within the forest. These were always near available
brood material (i.e. dying pines or infested wood piles) where high densities of progeny
T. piniperda adults had emerged and shoot-fed. Sauvard et al. (1987) found that
continual shoot-feeding throughout the season gradually resulted in a more even
distribution of beetles, so that by winter, they were dispersed over the entire forested area.
Then, during the next spring flight, beetles could easily find available brood material.
This may partially explain the lack of edge effects for T. piniperda; by autumn, the
beetles may have dispersed through the forest area.

I expected that T. piniperda would not cause significant shoot damage in the pine
stands in northern lower Michigan. Although most of the northern counties of lower
Michigan were quarantined for T. piniperda in 1996 or 1997 (NAPIS 2000), results of
related research (Kennedy 1998) led me to predict that T. piniperda populations would be
absent or not sufficiently established in these forest ecosystems to be detected in my
surveys. In southwestern lower Michigan, where T. piniperda populations have been

established for at least 7 years and probably longer (Haack et al. 1997), Scotch pine

77

stands sustained more shoot damage by T. piniperda than jack or red pine stands. In
contrast, in a study in mixed-species stands in northwestern Indiana, researchers
concluded that red pine was more susceptible to shoot-feeding by T. piniperda than
Scotch pine and jack pine (Sadof et al. 1994). However, this finding was based on the
number of damaged shoots relative to the basal area of each pine species, which may not
be an accurate comparison of shoot damage between the pine species. Also, Scotch pine
and jack pine trees may have been less vigorous in these stands because of site conditions
and proximity to parking lots or campsites. If the trees were less vigorous, then the
shoots could have been too slender to be suitable for T. piniperda. Shoot damage
attributable to other factors was not included in the Sadof et al. survey. At an
experimental forest in southwestern lower Michigan, Lawrence and Haack (1995) found
that red pine shoots were tunneled in more frequently than jack pine shoots when beetles
were individually caged on shoots, but the percent of total attacks and T. piniperda
survival throughout the summer was greater on Scotch pine than on the native eastern
pines. My field data suggest that this exotic bark beetle preferentially selects, damages
more shoots or is more abundant in Scotch pine stands in a landscape where Scotch, red
and jack pine stands are intermingled.

In Europe and Asia, pine stands stressed by disease (Kaitera and J alkanen 1994),
drought (Ye 1991), defoliation (Amezaga 1997), or poor management (Kaplan and
Mokrzycki 1988, Amezaga 1997) can be colonized for brood material by T. piniperda.
Colonization of trees in severely stressed stands often results in increased populations
and, therefore, more shoot-feeding. In 1998 and 1999, T. piniperda shoot damage in two

small Scotch pine stands in southwestern lower Michigan (Allegan Co.) was 25 times

78

higher than in 1997. This dramatic increase occurred in just one year. These stands were
adjacent to severely-stressed young Scotch pine trees that were overstocked and dying
due to numerous pathogens. I suspect that T. piniperda secondarily colonized these dying
trees and utilized the available brood material. This would have led to emergence of
large numbers of T. piniperda progeny adults who then were responsible for the increased
shoot damage in the adjacent mature stands. Comparable levels of shoot damage have
been found to cause significant volume growth losses in Scotch pine in European stands,
which were also near areas with ample available brood material (Andersson 1973,
Nilsson 1974, 1976, Langstrom and Hellqvist 1990, 1991, Eidmann 1992). In the other
pine stands that I surveyed, however, slash that was suitable for T. piniperda brood
material was infrequently observed and did not appear to vary among pine species or

between regions.

Native Shoot-Boring Insect Damage

Cone beetles and eastern pine shoot borer were the most common native insects
that damaged shoots in the pine stands that I surveyed between 1997 and 1999. Cone
beetles bore into twigs and deposit eggs in young cones where the larvae develop and
subsequently kill the cones (Hopkins 1915). Most cone beetle species have been
described based upon the species of host they attack and cannot be recognized by their
own anatomical features (Bright 1976, McPherson et al. 1970a, 1970b). The preferred
hosts of red pine cone beetle (C. resinosae) are second-year cones of red pine

(McPherson et al. 1970a, 1970b), but occasionally broods develop in second-year cones

79

 

of jack pine (Lyons 1956, Wilson 1977). Some shoot-feeding may occur in current-year
shoots of mature red pine (Lyons 1956, McPherson et al. 1970a, 1970b, Wilson 1977),
mainly in the distal tips of the shoots (Hopkins 1915). Red pine cone beetle may affect
natural regeneration of pine when high populations destroy pine seeds (Bright 1976,
Wilson 1977), but generally this beetle does not cause significant loss of foliage. Jack
pine tip beetle (C. banksiana) will shoot-feed in red and jack pine also (Herdy and
Thomas 1961; McPherson et al. 1970a, 1970b). McPherson et al. (1970a, 1970b) found
that jack pine tip beetle did shoot-feed in Scotch pine in Michigan, however, I only found
one Scotch pine shoot that was damaged by Conopthorus spp in my surveys.

Eastern pine shoot borer is a lepidopteran that is larger than the shoot-boring
beetles. Larvae feed and tunnel in the pith of terminal or lateral branch shoots before the
shoots have fully elongated (DeBoo et al. 1971, Wilson 1977). Red pine shoot moth
feeds on newly expanding shoots and cones of red pine (Hainze and Benjamin 1983,
Katovich and Hall 1992). Populations of red pine shoot moth fluctuate widely
throughout the range of red pine, but damaging populations are usually restricted to red
pine planted on sandy, dry sites. Shoot-feeding is most prevalent in trees in open-grown
stands and along stand edges (Hainze and Benjamin 1983, Katovich and Hall 1992).
These larger lepidopteran shoot borers only damaged red pine shoots, perhaps because
red pine shoots are generally greater in diameter than Scotch or jack pine shoots. Large
diameter shoots may be needed to accommodate the larger size and nutritional needs of

the larvae (DeBoo et al. 1971, Hainze and Benjamin 1983).

80

Conclusions

Non-insect factors caused significantly more shoot damage in Michigan pine
stands than was attributable to T. piniperda or native shoot-boring insects, except in two
Scotch pine stands in southwestern lower Michigan (Allegan Co.). These stands are
unique because of the abundance of Scotch pine brood material adjacent to the stands. In

northern lower Michigan between 1997 and 1999, established populations of T. piniperda

 

were not present in any of the stands I surveyed. In surveys conducted in autumn 1999, a
total of 3 Scotch pine shoots damaged by T. piniperda were found in Clare and Gladwin
counties, suggesting that the range of T. piniperda may be expanding into central lower
Michigan. T. piniperda caused significantly more shoot damage in Scotch pine stands
than in red or jack pine stands in southwestern lower Michigan. Red pine shoots were
damaged more frequently by native shoot-borers than by T. piniperda.

I conclude that T. piniperda may have the potential to cause significant shoot
damage in Scotch pine in circumstances where suitable brood material is plentiful,
possibly resulting in grth losses. Scotch pine stands in areas where large amounts of
brood material are present, such as sawmills or severely stressed, unmanaged stands, are

at risk of damaging levels of T. piniperda shoot-feeding.

81

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Figure 3.1. Michigan counties that are shaded were quarantined for

T omicus piniperda as of April 2000. Stands that were surveyed were
located in Benzie (a), Manistee (b), Clare (c), and Gladwin (d) counties
in northern lower Michigan, and Allegan (e), Kalamazoo (t), and Cass
(g) counties in southwestern lower Michigan.

99

['T“_'l

LITERATURE CITED

Amezaga, I. 1997. Forest characteristics affecting the rate of shoot pruning by the pine
shoot beetle (T amicus piniperda L.) in Pinus radiata D. Don and P. sylvestris L.
plantations. Forestry. 70(2): 129-137.

Andersson, S. O. 1973. Increment losses after thinning [in Scots pine] caused by
Myelophilus piniperda. Sveriges Skogsvardforbunds Tidskrifi. 71(4): 359-379.
English summary.

Animal and Plant Health Inspection Service (APHIS). 1993. Pine shoot beetle. Federal
Register, 13 May 1993. 58(91): 28, 333-28, 335.

Bakke, A. 1968. Ecological studies on bark beetles (Col.: Scolytidae) associated with
Scots pine (Pinus sylvestris) in Norway with particular reference to the influence
of temperature. Meddelelser fra Det Norske Skogfosakvesen. 21: 443-602.

Bright, D. E., Jr. 1976. The insects and arachnids of Canada. Part 2. The bark beetles of
Canada and Alaska, Coleoptera: Scolytidae. Canada Department of Agriculture.
Publication # 1576. Ottawa, Ontario.

DeBoo, R. F., W. L. Sippell, and H. R. Wong. 1971. The eastern pine shoot borer,
Eucosma gloriola (Lepidotera: Tortricidae), in North America. Canadian
Entomologist. 103: 1473-1486.

DeGroot, P., G. T. Harvey, and P. M. Roden. 1992. Genetic divergence among eastern
North American cone beetles, Conaphthorus (Coleoptera: Scolytidae). Canadian
Entomologist. 124: 189-199.

Eidmann, H. H. 1985. Silviculture and insect problems. Zeitschrifi fur Angewandte
Entomologie. 99(1): 105-112.

Eidmann, H. H. 1992. Impact of bark beetles on forests and forestry in Sweden. Journal
of Applied Entomology. 114(2): 193-200.

Haack, R. A. and D. Kucera. 1993. New introduction — common pine shoot beetle,
T omicus piniperda (L.). Pest Alert. United States Department of Agriculture
Forest Service #NA-TP-05-93.

Haack, R. A., R. K. Lawrence, D. G. McCullough, and C. S. Sadof. 1997. T omicus
piniperda in North America: an integrated response to a new exotic scolytid. J. C.
Gregoire, A. M. Liebhold, F. M. Stephen, K. R. Day, and S. M. Salom, editors.
In: Proceedings of Integrating cultural tactics into the management of bark beetle

100

and reforestation pests. USDA Forest Service General Tech. Report. NE-236.
Pp. 62-72.

Hainze, J. H. and D. M. Benjamin. 1983. Bionomics and pattern of attack of the red pine
shoot moth, Dioryctria resinosella (Lepidoptera: Pyralidae), in Wisconsin.
Canadian Entomologist. 1 15: 1 169-1 175.

 

Herdy, H. and J. B. Thomas. 1961. The seasonal development of a species of
Conopthorus (Hopkins) (Coleoptera: Scolytidae) in the shoots of j ack pine, Pinus
banksiana (Lamb.), in Ontario. Canadian Entomologist. 93: 936-940.

Hopkins, A. D. 1915. A new genus of scolytid beetles. Washington Academy of
Sciences Journal. 5: 429-433.

 

Kaitera, J. and R. J alkanen. 1994. The history of shoot damage by T omicus spp. (Col.,
Scolytidae) in a Pinus sylvestris L. stand damaged by the shoot-disease fungus
Gremmeniella abietina (Lagerb.) Morelet. Journal of Applied Entomology.
117(3): 307-313.

Kaplan, M. and T. Mokrzycki. 1988. A contribution to the knowledge of the occurrence
of the pine shoot beetles T omicus minor and T. piniperda in Scots pine thickets in
Niedzwiady forest district. Sylwan. 132(6): 35-39. English summary.

Katovich, S. A. and D. J. Hall. 1992. How to identify and minimize red pine shoot moth
damage. USDA Forest Service # NA-FR-02-92.

Kauffman, W. C., R. D. Waltz, and R. B. Cummings. 1998. Shoot feeding and
overwintering behavior of T omicus piniperda (Coleoptera: Scolytidae):
implications for management and regulation. Journal of Economic Entomology.
91(1): 182-190.

Kennedy, A. A. 1998. Interactions of the pine shoot beetle [T omicus piniperda (L.)
(Coleoptera: Scolytidae)] with native pine bark beetles and their associated natural
enemies in Michigan. M.S. thesis. Department of Entomology, Michigan State
University. East Lansing, MI. 151pp.

Kruskal, W. H. and W. A. Wallis. 1952. Use of ranks in one-criterion variance analysis.
Journal of the American Statistics Association. 47: 583-621.

Langstrom, B. 1980a. Life cycles of the pine shoot beetles with particular reference to
their maturation feeding in the shoots of Scots pine. Ph.D. dissertation. Swedish
University of Agricultural Sciences, Faculty of Forestry, Division of Forest
Entomology. 123pp. Uppsala, Sweden.

101

Langstrom, B. 1980b. Distribution of pine shoot beetle (T omicus piniperda, T omicus
minor) attacks within the crown of Scots pine (Pinus sylvestris). Studia F orestalia
Suecica. 154: 1-25.

Langstrom, B. 1983. Life cycles and shoot-feeding of the pine shoot beetles. Studia
Forestalia Suecica. 163: 1-29.

Langstrom, B. and C. Hellqvist. 1985. Pinus contorta as a potential host for T omicus
piniperda L. and T. minor (Hart) (Col., Scolytidae) in Sweden. Zeitschrifi fur
Angewandte Entomologie. 99(2): 174-181.

Langstrom, B. and C. Hellqvist. 1990. Spatial distribution of crown damage and growth
losses caused by recurrent attacks of pine shoot beetles in pine stands surrounding
a pulp mill in southern Sweden. Journal of Applied Entomology. 110(3): 261-
269.

Léngstrom, B. and C. Hellqvist. 1991. Shoot damage and growth losses following three
years of T omicus-attacks in Scots pine stands close to a timber storage site. Silva
Fennica. 25(3): 133-145.

Lawrence, R. K., and R. A. Haack. 1995. Susceptibility of selected species of North
American pines to shoot feeding by an Old World scolytid: T omicus piniperda.
In: F. P. Hain et al. eds., Forest Insect Behavior, Population Dynamics and
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State Univ. Press, Wooster, Ohio.

Lyons, L. A. 1956. Insects affecting seed production in red pine. Part I. Conopthorus
resinosae Hoph. (Coleoptera: Scolytidae). Canadian Entomologist. 88: 599-608.

McCullough, D. G. and D. R. Smitley. 1995. Evaluation of insecticides to reduce
maturation feeding by T omicus piniperda (Coleoptera: Scolytidae) in Scotch pine.
Journal of Economic Entomology. 88(3): 693-699.

McPherson, J. E., L. F. Wilson, and F. W. Stehr. 1970a. A comparison between
Conopthorus shoot-infesting beetles and Conopthorus resinosae (Coleoptera:
Scolytidae). Part 1. Comparative life history studies in Michigan. Canadian
Entomologist. 102: 1008-1015.

McPherson, J. E., F. W. Stehr, and L. F. Wilson. 1970b. A comparison between
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Scolytidae). Part H. Reciprocal host and resin toxicity tests; with description of a
new species. Canadian Entomologist. 102: 1008-1015.

National Animal and Plant Inspection Service (NAPIS) Database. 2000. <http://www.
cercis.purdue.edu/napis/pests/psb/mgif/psbcnty.gif/>

102

Nilsson, S. 1974. Increment losses caused by Tomicus piniperda on Scots pine.
Rapporter och Uppsatser, Institutionen for Skogsteknik. 78(14): 1-64. English
summary.

Nilsson, S. 1976. Rationalization of forest operations gives rise to insect attack and
increment losses. Ambio. 5(1): 17-22.

SAS Institute, Inc. 1989. SAS/STAT users guide. Version 6, fourth edition. Cary, NC:
SAS Institute, Inc.

Sadof, C. S., Waltz, R. D., and C. D. Kellam. 1994. Differential shoot feeding by adult
T omicus piniperda (Col.: Scolytidae) in mixed stands of native and introduced
pines in Indiana. Great Lakes Entomologist. 27(4): 223-228.

"hf-l

1F

Sauvard, D., F. Lieutier, and J. Levieux. 1987. Spatial distribution and dispersal of
T omicus piniperda L. (Coleoptera, Scolytidae) in the forest of Orleans (France).
Annales des Sciences Forestieres. 44(4): 417-434. English summary.

Searle, S. R., F. M. Speed, and G. A. Milliken. 1980. Populations marginal means in the
linear model: an alternative to least squares means. The American Statistician.
34: 2 1 6-221 .

Wilson, L. F. 1977. A guide to insect injury of conifers in the Lake States. USDA
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Ye, H. 1991. On the bionomy of T omicus piniperda (L.) (Col.: Scolytidae) in the
Kunming region of China. Journal of Applied Entomology. 112(4): 366-369.

103

MANAGEMENT IMPLICATIONS

The range of T omicus piniperda (L.) (Coleoptera: Scolytidae) in North America
has substantially expanded in recent years. As of April 2000, at least 295 counties in 11
north central and northeastern states (National Animal and Plant Inspection Service
I [NAPIS] 2000), 24 counties in Ontario and 8 counties in Quebec (C. Markham, USDA
Animal and Plant Health Inspection Service [APHIS], pers. comm.) were regulated for T.
piniperda. To-date, the only area in North America where T. piniperda is known to have
caused significant shoot damage to native pines is in southern Ontario. Native pine
stands adjacent to or nearby areas with dead and dying stands of Scotch pine were
heavily shoot-fed on by T. piniperda (E. Czerwinski and T. Scarr, Ontario Ministry of
Natural Resources, pers. comm). My observations of these stands and discussions with
Ontario pest managers suggest that T. piniperda may utilize this brood material and
disperse into native pine stands to shoot-feed. Management guidelines need to be
developed for T. piniperda to prevent ecological impacts similar to those seen in Ontario
fi'om occurring in Michigan.

Results from this research indicate that T. piniperda parent adults exhibit a
preference for Scotch pine (Pinus sylvestris L.), its European host, over red pine (Pinus
resinosa Ait.) and jack pine (Pinus banksiana Lamb.) in laboratory and field experiments
in Michigan. T. piniperda populations were found to be well-established in
southwestern lower Michigan, where this beetle preferentially colonized Scotch pine logs
over red and jack pine logs regardless of the stand species. In laboratory experiments, T.

piniperda parent adults demonstrated a preference for Scotch pine logs when paired with

104

red or jack pine logs. Progeny adults tested in the laboratory preferred Scotch pine, along
with jack pine shoots, over red pine shoots for maturation feeding. In shoot damage
surveys, Scotch pine shoots were damaged by T. piniperda more fiequently than red and
jack pine shoots in southwestern lower Michigan stands, though non-insect factors such
as storms and squirrels accounted for the majority of shoot damage in all pine stands.

The apparent preference of T. piniperda for Scotch pine during selection of brood
material suggests that Scotch pine may be considered a high-risk species. Stands of
Scotch pine with available brood material, such as slash or declining trees, may be
especially at risk. Although T. piniperda is currently causing more shoot damage in
Scotch pine stands than in red and jack pine stands in Michigan, results from my
laboratory studies suggest that T. piniperda could potentially cause significant shoot
damage in jack pine stands. Fortunately, pine logs in jack pine stands were consistently
colonized at lower densities than logs in red and Scotch pine stands, indicating that jack
pine stands currently do not sustain damaging levels of T. piniperda. Red pine may be
considered a relatively low-risk species because it was least preferred by T. piniperda
during host selection of brood material and shoots and, in field surveys, trees infrequently
sustained shoot damage.

Emphasis should be placed on monitoring high-risk stands such as Scotch pine
stands, particularly stands containing or adjacent to areas with accumulated brood
material. Scotch pine trap logs with thick, corky bark should be the preferred type of log
used for detection or monitoring of T. piniperda populations. These logs are most likely
to be colonized by T. piniperda in early spring and may effectively trap out low

populations of T. piniperda in stands without other brood material available (McCullough

105

and Sadof 1998). These infested trap logs must be removed and destroyed by early to
late May to avoid emergence of T. piniperda progeny adults.

Despite T. piniperda’s preference for Scotch pine, all pine stands can be managed
to decrease the likelihood that T. piniperda or other scolytid p0pulations may build to
damaging levels. Emphasis should be placed on keeping pine stands clean of potential
brood material. Chipping or burning slash greater than 7 cm in diameter, running over it
with heavy equipment, or otherwise damaging the phloem to make it unsuitable for
scolytid colonization may be practical methods of maintaining clean stands. Tree vigor
should be encouraged to prevent predisposition to pathogens which could potentially
stress trees severely enough to attract and be secondarily colonized by T. piniperda.
Avoiding wounds during thinnings and maintaining appropriate stocking densities will
help to ensure healthy stands. Timing harvests or thinnings to occur after mid-summer,
late enough to avoid T. piniperda colonization during the current year and early enough
to cause most brood material to be unsuitable for colonization by the following spring,
may be an effective method of managing stands to reduce the potential impacts of T.
piniperda. In contrast, row thinning or partial harvests of pine stands during winter are
likely to create favorable T. piniperda brood material for the following spring.

Future research should involve analyzing pine volatiles to identify compounds
used by T. piniperda to differentiate among pine species and to understand more about
the sensory stimuli of T. piniperda. Host selection by T. piniperda of pine logs cut at
different times during the year should be examined and should involve analysis of pine
volatiles to determine how compounds and attractiveness of pine logs to T. piniperda

change over time. Host suitability and stage-specific survival of T. piniperda in different

106

 

pine hosts should also be examined at different population and attack densities. This
information may be useful in developing improved methods to detect and manage this
exotic scolytid in North American forests, and to understand the ecological implications

of this exotic pest’s establishment.

107

APPENDICES

108

 

APPENDIX A

109

 

Appendix A

Table A1. Frequency of shoots damaged by non-insect factors in red, jack and
Scotch pine stands that were surveyed in lower Michigan between 1997 and 1999.

 

No. Damaged Frequency

 

110

Year Region Stand spp. Edge? Shoots of
(per 1 m2) Occurrence
m 1997 North Red pine Yes 7 83
Autumn 1997 North Red pine Yes 1 23
Autumn 1997 North Red pine Yes 2 6
Autumn 1997 North Red pine Yes 3 1
Autumn 1997 North Red pine Yes 4 2
Autumn 1997 North Red pine No 0 93
Autumn 1997 North Red pine No 1 29
Autumn 1997 North Red pine No 2 11
Autumn 1997 North Red pine No 3 5
Autumn 1997 North Red pine No 4 2
Autumn 1997 North Jack pine Yes 0 83
Autumn 1997 North Jack pine Yes 1 23
Autumn 1997 North Jack pine Yes 2 17
Autumn 1997 North Jack pine Yes 3 12
Autumn 1997 North Jack pine Yes 4 3
Autumn 1997 North Jack pine Yes 5 3
Autumn 1997 North Jack pine Yes 6 3
Autumn 1997 North Jack pine Yes 8 1
Autumn 1997 North Jack pine No 0 66
Autumn 1997 North Jack pine No 1 46
Autumn 1997 North Jack pine No 2 16
Autumn 1997 North Jack pine No 3 20
Autumn 1997 North Jack pine No 4 14

Table A1 (cont'd.)

Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997

North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
South
South
South
South
South
South
South
South
South
South
South

Jack pine
Jack pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine

Jack pine

111

No
No
No
No
Yes
Yes
Yes
Yes
Yes
Yes
Yes
No
No
No
No
No
No
Yes
Yes
Yes
Yes
No
No
No
No
Yes
Yes
Yes

Gum-huts)

ObJNr—oOUI-bUJNv-t

OWN—-

p—o

68
15

34
27
19

Table A1 (cont'd.)

Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Autumn 1997
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998

South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
North
North
North
North
North
North
North
North
North

Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine

112

Yes
Yes
Yes
Yes
Yes
No
No
No
No
No
No
Yes
Yes
Yes
Yes
No
No
No
No
Yes
Yes
Yes
Yes
Yes
No
No
No
No

OWN OWNHO AWNHOOQUIAW

Ohmic—-

45
38
22

34
13

55
15

52
36
19

75
41
17

 

 

Table A1 (cont'd.)

Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998

North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North

Red pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine

Jack pine

113

No
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
No
No
No
No
No
No
No
No
No

Hos:

\OOONGUI-bUJN

F—‘Hh—‘D—II—i
\DM-br—tO

20

O

mflom-wat-r

36
19
26
14
19
15

 

Table A1 (cont'd.)

Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998

North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
South
South
South
South
South

Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Red pine

114

No
No
No
No
No
No
No
No
No
No
No
Yes
Yes
Yes
Yes
Yes
Yes
No
No
No
No
No
No
Yes
Yes
Yes
No
No

10
11
12
13
14
15
17
18
20
24

OLA-bulb)

Hows-wt»)

HON

NNr—‘NUI-P-

10

10

19
ll
15

40
19

54
27

 

Table A1 (cont'd.)

Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
1998
1998

South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
North
North

Red pine
Red pine
Red pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Red pine
Red pine

115

No
No
No
Yes
Yes
Yes
Yes
Yes
No
No
No
No
No
No
No
Yes
Yes
Yes
Yes
Yes
No
No
No
No
No
No
Yes
Yes

"O-bUJNt-‘OQUI-hwtq OUUUJN OhWN

ONMWN

43
34

43
33
19
14

20
15
17

45
20
l 1

48
41

 

lg ’ “q
h

I? .
I.

 

Table A1 (cont'd.)

 

 

1998 North Red pine Yes 2 9
1998 North Red pine Yes 3 1
1998 North Red pine No 0 53
1998 North Red pine No 1 51
1998 North Red pine No 2 13
1998 North Red pine No 3 7
1998 North Red pine No 4 1
1998 North Jack pine Yes 0 29
1998 North Jack pine Yes 1 22
1998 North Jack pine Yes 2 29
1998 North Jack pine Yes 3 9
1998 North Jack pine Yes 4 3
1998 North Jack pine Yes 5 1
1998 North Jack pine Yes 6 2
1998 North Jack pine No 0 35
1998 North Jack pine No 1 29
1998 North Jack pine No 2 40
1998 North Jack pine No 3 9
1998 North Jack pine No 4 6
1998 North Jack pine No 5 1
1998 North Scotch pine Yes 0 17
1998 North Scotch pine Yes 1 15
1998 North Scotch pine Yes 2 3
1998 North Scotch pine No 0 28
1998 North Scotch pine No 1 20
1998 North Scotch pine No 2 3
1998 South Red pine Yes 0 3 8
1998 South Red pine Yes 1 20

116

Table A1 (cont'd.)

 

1998 South Red pine Yes 2 7

1998 South Red pine No 0 56

1998 South Red pine No 1 26

1998 South Red pine No 2 7

1998 South Red pine No 3 l

1998 South Jack pine Yes 0 37

1998 South Jack pine Yes 1 3o """
1998 South Jack pine Yes 2 17 i
1998 South Jack pine Yes 3 4 '- t
1998 South Jack pine Yes 4 2

1998 South Jack pine No 0 43

1998 South Jack pine No 1 30

1998 South Jack pine No 2 22

1998 South Jack pine No 3 13

1998 South Jack pine No 4 4

1998 South Jack pine No 5 1

1998 South Scotch pine Yes 0 22

1998 South Scotch pine Yes 1 16

1998 South Scotch pine Yes 2 5

1998 South Scotch pine Yes 3 2

1998 South Scotch pine No 0 35

1998 South Scotch pine No 1 20

1998 South Scotch pine No 2 7

1998 South Scotch pine No 3 3

1999 North Red pine Yes 0 94

1999 North Red pine Yes 1 5

1999 North Red pine No 0 1 16

1999 North Red pine No 1 8

117

 

Table A1 (cont'd.)

1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999

North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
North
South
South
South
South
South
South
South
South

Red pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine

Jack pine

118

No
Yes
Yes
Yes
Yes
Yes
Yes
No
No
No
No
No
No
No
Yes
Yes
Yes
No
No
No
Yes
Yes
No
No
No
Yes
Yes
Yes

HON

OQMJiUJN—‘OOO-hU-JN

t—‘ONH

ON

40
32
12

56
4O
15

24
10

37
13

55
10
78
11

55
23

 

 

 

Table A1 (cont'd.)

1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999

South
South
South
South
South
South
South
South
South
South
South
South
South

Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine

Scotch pine

Yes
Yes
No
No
No
No
No
No
Yes
Yes
Yes
No
No

OUIJAUJN Ohm

ON

65
28

 

119

 

 

APPENDIX B

120

 

Appendix B

 

Table B1. Frequency of shoots damaged by T omicus pim'perda in red, jack and
Scotch pine stands that were surveyed in lower Michigan between 1997 and 1999.

 

No. Damaged Frequency

 

 

 

Year Region Stand spp. Edge? Shoots of
(per 1 m2) Occurrence

Autumn 1997 North Red pine Yes 0 1 15
Autumn 1997 North Red pine No 0 140 , -
Autumn 1997 North Jack pine Yes 0 145 L
Autumn 1997 North Jack pine No 0 170 g;
Autumn 1997 North Scotch pine Yes 0 50
Autumn 1997 North Scotch pine No 0 75
Autumn 1997 South Red pine Yes 0 65
Autumn 1997 South Red pine No 0 88
Autumn 1997 South Red pine No 1 2
Autumn 1997 South Jack pine Yes 0 86
Autumn 1997 South Jack pine Yes 1 9
Autumn 1997 South Jack pine No 0 1 13
Autumn 1997 South Jack pine No l 4
Autumn 1997 South Jack pine No 2 2
Autumn 1997 South Jack pine No 3 1
Autumn 1997 South Scotch pine Yes 0 43
Autumn 1997 South Scotch pine Yes 1 8
Autumn 1997 South Scotch pine Yes 2 2
Autumn 1997 South Scotch pine Yes 3 2
Autumn 1997 South Scotch pine No 0 64
Auturrm 1997 South Scotch pine No 1 1 1
Autumn 1997 South Scotch pine No 2 3
Autumn 1997 South Scotch pine No 3 1

121

Table B1 (cont'd.)

Autumn 1997

Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
1998
1998
1998

South
North
North
North
North
North
North
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
North
North
North

Scotch pine
Red pine
Red pine
Jack pine
Jack pine

Scotch pine

Scotch pine
Red pine
Red pine
Red pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine
Jack pine

Scotch pine

Scotch pine

Scotch pine

Scotch pine

Scotch pine

Scotch pine

Scotch pine

Scotch pine
Red pine
Red pine

Jack pine

122

No
Yes
No
Yes
No
Yes
No
Yes
Yes
No
Yes
Yes
Yes
No
No
No
No
Yes
Yes
Yes
Yes
No
No
No
No
Yes
No
Yes

HOWN—‘OUJN ow~oo~oooooooa

OOOWN

115
140
145
170

125
95

 

Table Bl (cont'd.)

1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998

North
North
North
North
North
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South

Jack pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine

Red pine

Red pine

Red pine

Jack pine

Jack pine

Jack pine

Jack pine

Jack pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine

Scotch pine

123

No
Yes
Yes
No
No
Yes
Yes
No
Yes
Yes
Yes
No
No
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes

t-‘OOHO

OD)

120
34

50

64

90

84

103

 

Table B1 (cont'd.)

1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999

South
South
South
South
South
South
South
South
South
South
South
South
South
South
North
North
North
North
North
North
North
South
South
South
South
South
South
South

Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Red pine
Red pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine

Jack pine

124

Yes
No
Yes
No
Yes
Yes
No
Yes
Yes
No
No
Yes
Yes
Yes

F‘O

\OWQONM-bUJN

OOHOOOOO

Ou—I

19
12

MWNW

{Audio

99
125
95
120
34

51
63

85

78
11

 

 

Table B1 (cont'd.)

1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999

South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South
South

Jack pine

Jack pine

Jack pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine
Scotch pine

Scotch pine

125

No
No
No
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
No
No
No
No
No
No
No
No
No
No
No

HON

\OOOQONUIhUJN

u—Au-dv—A
NHO

97
13

12
12

 

rr‘r—‘H
'1' a

 

Table B1 (cont'd.)

l 999 South Scotch pine No 1 3 1
1999 South Scotch pine No l 5 1
1999 South Scotch pine No 1 7 l

 

 

126

 

 

APPENDIX C

127

Appendix C

Table C1. Frequency of shoots damaged by Conopthorus resinosae in red, jack and
Scotch pine stands that were surveyed in lower Michigan between 1997 and 1999.

 

No. Damaged Frequency

 

 

 

 

Year Region Stand spp. Edge? Shoots of
(per 1 m2) Occurrence
Autumn 1997 North Red pine Yes 0 106
Autumn 1997 North Red pine Yes 1 7
Autumn 1997 North Red pine Yes 2 2
Autumn 1997 North Red pine No 0 117
Autumn 1997 North Red pine No 1 22
Autumn 1997 North Red pine No 2 1
Autumn 1997 North Jack pine Yes 0 144
Autumn 1997 North Jack pine Yes 1 1
Autumn 1997 North Jack pine No 0 169
Autumn 1997 North Jack pine No 1 1
Autumn 1997 North Scotch pine Yes 0 50
Autumn 1997 North Scotch pine No 0 75
Autumn 1997 South Red pine Yes 0 65
Autumn 1997 South Red pine No 0 90
Autumn 1997 South Jack pine Yes 0 95
Autumn 1997 South Jack pine No 0 119
Autumn 1997 South Jack pine No 1 1
Autumn 1997 South Scotch pine Yes 0 55
Autumn 1997 South Scotch pine No 0 80
Spring 1998 North Red pine Yes 0 105
Spring 1998 North Red pine Yes 1 9
Spring 1998 North Red pine Yes 3 1
Spring 1998 North Red pine No 0 134

128

Table C1 (cont'd.)

Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998
1998

North
North
North
North
North
South
South
South
South
South
South
South
South
North
North
North
North
North
North
North
North
North
North
South
South
South
South
South

Red pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Red pine
Red pine
Jack pine
Jack pine

Scotch pine

129

No
Yes
No
Yes
No
Yes
Yes
No
No
Yes
No
Yes
No
Yes
Yes
Yes
No
No
No
Yes
No
Yes
No
Yes
No
Yes
No
Yes

OOOOOH

HOt—a

HON—‘OOOOO

OOOOOOOOON

145
170
40
60

89

95
120
55
80
75
20

86
35

95
120
35
51
65
90
90
1 13
45

Table C1 (cont'd.)

1998
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999

South
North
North
North
North
North
North
North
North
North
North
North
North
North
North
South
South
South
South
South
South

Scotch pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine

Scotch pine

Scotch pine

Scotch pine
Red pine
Red pine
Jack pine
Jack pine

Scotch pine

Scotch pine

No
Yes
Yes
Yes
Yes
No
No
No
No
No
Yes
No
Yes
No
No
Yes
No
Yes
No
Yes
No

OWN

t-"C>OOO-l>u)t\.)u—n

OOOOOO

65
49
26
20

69
32 t":-
17

 

95
120
35
50

65
90
9O
1 13
45
65

 

130

 

APPENDIX D

131

Appendix D

Table D1. Frequency of shoots damaged by Eucosma gloriola in red, jack and
Scotch pine stands that were surveyed in lower Michigan between 1997 and 1999.

 

No. Damaged Frequency

 

Year Region Stand spp. Edge? Shoots of
(per 1 m2) Occurrence

Autumn 1997 North Red pine Yes 0 90
Autumn 1997 North Red pine Yes 1 17
Autumn 1997 North Red pine Yes 2 6 i
Autumn 1997 North Red pine Yes 3 2 E
Autumn 1997 North Red pine No 0 106 it
Autumn 1997 North Red pine No 1 26
Autumn 1997 North Red pine No 2 4
Autumn 1997 North Red pine No 3 4
Autumn 1997 North Jack pine Yes 0 145
Autumn 1997 North Jack pine No 0 170
Autumn 1997 North Scotch pine Yes 0 50
Autumn 1997 North Scotch pine No 0 75
Autumn 1997 South Red pine Yes 0 58
Autumn 1997 South Red pine Yes 1 5
Autumn 1997 South Red pine Yes 2 2
Autumn 1997 South Red pine No 0 80
Autumn 1997 South Red pine No 1 9
Autumn 1997 South Red pine No 2 1
Autumn 1997 South Jack pine Yes 0 95
Autumn 1997 South Jack pine No 0 120
Autumn 1997 South Scotch pine Yes 0 55
Autumn 1997 South Scotch pine No 0 80

Spring 1998 North Red pine Yes 0 91

132

Table D1 (cont'd.)

Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
Spring 1998
1998
1998
1998
1998
1998
1998
1998
1998
1998

North
North
North
North
North
North
North
North
North
South
South
South
South
South
South
South
South
South
South
North
North
North
North
North
North
North
North
North

Red pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Scotch pine

Scotch pine

133

Yes
Yes
No
No
No
Yes
No
Yes
No
Yes
Yes
Yes
No
No
No
Yes
No
Yes
No
Yes
Yes
No
No
No
No
Yes
Yes
No

F‘OOOOON—‘ONH

HON

OOOOON

OOOUJN

22

125
14

145
170

120
55
80
87
12

111
12

95
35
51

 

Table D1 (cont'd.)

1998
1998
1998
1998
1998
1998
1998
1998
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999
1999

South
South
South
South
South
South
South
South
North
North
North
North
North
North
North
South
South
South
South
South
South
South
South

Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Jack pine
Jack pine
Scotch pine
Scotch pine
Red pine
Red pine
Red pine
Red pine
Jack pine
Jack pine
Scotch pine

Scotch pine

Yes
Yes
No
No
Yes
No
Yes
No
Yes
No
No
Yes
No
Yes
No
Yes
Yes
No
No
Yes
No
Yes
No

HOOOOOO

OOOOO

COCO

63

85

90
113
45
65
99
121

95
120
35
51

89

90
1 13
45
65

 

134

 

APPENDIX E

135

Appendix E
Record of Deposition of Voucher Specimens‘

The specimens listed on the following sheet(s) have been deposited in the named museum(s) as samples of
those species or other taxa which were used in this research. Voucher recognition labels bearing the
Voucher No. have been attached or included in fluid-preserved specimens.

Voucher No.: 1999-10

Title of thesis or dissertation (or other research projects):

Host Preference and Potential Impact of Pine Shoot Beetle [Tomicus piniperda (L.) (Coleoptera:
Scolytidae)] in Michigan Pine Stands

Museum(s) where deposited and abbreviations for table on following sheets:

Entomology Museum, Michigan State University (MSU)

Other Museums: None

Investigator’s Name(s) (typed)
Wen

 

 

 

 

Date W

‘Reference: Yoshimoto, C. M. 1978. Voucher Specimens for Entomology in North America. Bull.
Entomol. Soc. Amer. 24: 141-42.

Deposit as follows:
Original: Include as Appendix 1 in ribbon copy of thesis or dissertation.

Copies: Include as Appendix 1 in copies of thesis or dissertation.
Museum(s) files.
Research project files.

This form is available from and the Voucher No. is assigned by the Curator, Michigan State University
Entomology Museum.

136

 

Appendix E

Voucher Specimen Data

Page 1 of 2 Pages

 

8.5 a
331“ X‘QM n «tuna-l WI
awe am

38%: 3% Swag: 2: a £88
3m «SE89. team: 26% 05 vozooom

 

ame Wm fiascofl 8am

 

 

 

O .— IGQQ m .02 H0£O§O>

tomomm 033 5:32
gas $.52 £8333
3.3330: a £85. 3:363 83

 

 

 

.00 005253

 

 

 

 

 

 

 

 

 

 

 

 

do Efis
.00 0.85
do Bowspfio
3 do «:82 ”EOEUE .m ESE? 35630
$2.325 “8898—00
Ewa
m fi .00 gang ”240503 SENSE fizzokfif .8633me
35:3 2 60 ooamEMWM
.00 Ed 5
“Ban 2 .oo 322 ”720503 T: nutmeg senses
anuboom ”gumoofioo
M d 0..
W m .m a m .m e m u s 3:8qu n95 no 0.8 880m
Mmm. % M M m. N .m “m.” vfivgsuevouoozoomuofimoommuomafigonj 5 . m
me 59:52

 

137

 

 

 

Appendix E

Voucher Specimen Data

Page 2 of 2 Pages

 

 

 

 

. 089 a
has has M 01.1th aflufi 82 .w bassoon 28
o .30 Em
£835 saw 5322 as a :88
8m 8083on can: 96% 05 3330M Ewomm 033 5852
0 fl -maa~ .oZ 8:025 Avonbv 325 Z {033333

Agaooou a 38% 38323 88

 

S

2

do egg—«M

.oo 82mg
.00 «:82

.oo $.83
.00 coaxed—«M
.8 3&5
do 0.85

.00 38380
.00 ofiaom

.00 gm

do 533%

.00 conga—«M
.00 0.35

do EQBSU
60 058m

”EOEUE

”720503

”EOEUE

Ammmv 382335“. nauseozoN

Q 53% asaszsfi

Ease
Ezmamxza 899%? «33.6on
3255 ”883200

 

Museum
where

deposited

 

Other

 

 

 

Pupae

 

Nymphs
Larvae

 

 

Eggs

338%

28 won: 8 38:8 3230on .5.“ 83 ~33

 

nous 850 B 86on

 

 

Adults (3‘

 

 

138

LITERATURE CITED

139

 

 

LITERATURE CITED

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Amezaga, I. 1997. Forest characteristics affecting the rate of shoot pruning by the pine
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Amman, G. D. 1971. Mountain pine beetle brood production in relation to thickness of
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Amman, G. D. 1982. Characteristics of mountain pine beetles reared in four pine hosts.
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Anderbrant, O. and F. Schlyter. 1989. Causes and effects of individual quality in bark
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Andersson, S. O. 1973. Increment losses after thinning [in Scots pine] caused by
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English summary.

Animal and Plant Health Inspection Service (APHIS). 1993. Pine shoot beetle. Federal
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Bakke, A. 1968. Ecological studies on bark beetles (Col.: Scolytidae) associated with
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Brattli. J. G., J. Anderson, and A. C. Nilssen. 1988. Primary attraction and host tree
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Bright, D. E., Jr. 1976. The insects and arachnids of Canada. Part 2. The bark beetles
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368 pp.

140

 

 

Butovitsch, V. 1972. Injury caused by feeding of Blastophagus piniperda on Picea
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summary.

Byers, J. A. 1989. Chemical ecology of bark beetles. Experientia. 45(3): 271-283.

Byers, J. A., B. S. Lanne, J. Lofqvist, F. Schlyter, and G. Bergstrom. 1985. Olfactory
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Byers, J. A., B. S. Lanne, and J. Lofqvist. 1989. Host tree unsuitability recognized by
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Carter, M. C. A., J. L. Robertson, R. A. Haack, R. K. Lawrence, and J. L. Hayes. 1996.
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Chararas, C. 1968. The power of adaptation by Blastophagus piniperda (Col.,
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English summary.

Chénier, J. V. R. and B. J. R. Philogéne. 1989. Field responses of certain forest
Coleoptera to conifer monoterpenes and ethanol. Journal of Chemical Ecology.
15: 1729-1746.

Czokajlo, D., R. A. Wink, J. C. Warren, and S. A. Teale. 1997. Growth reduction of
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Forest Research. 27(9): 1394-1397.

Czokajlo, D. and S. A. Teale. 1997. Synergistic effect of ethanol to alpha-pinene in
primary attraction of the larger pine shoot beetle, T omicus piniperda. Journal of
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Czokajlo, D. 1998. Semiochemicals for the larger pine shoot beetle (T omicus piniperda
L.) and its clerid predators. Ph.D. dissertation. State University of New York.
164pp. Syracuse, NY.

DeBoo, R. F ., W. L. Sippell, and H. R. Wong. 1971. The eastern pine shoot borer,
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Entomologist. 103: 1473-1486.

141

 

DeGroot, P., G. T. Harvey, and P. M. Roden. 1992. Genetic divergence among eastern
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Eidmann, H. H. 1985. Silviculture and insect problems. Zeitschrifl fur Angewandte
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Eidmann, H. H. 1992. Impact of bark beetles on forests and forestry in Sweden. Journal
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Fumiss, M. M. 1997. American forest entomology comes on stage. Bark beetle
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Gibbs, J. N. and A. Inman. 1991. The pine shoot beetle T omicus piniperda as a vector of
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Haack, R. A. and D. Kucera. 1993. New introduction — common pine shoot beetle,
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Haack, R. A. and R. K. Lawrence. 1995a. Spring flight of T omicus piniperda in relation
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Haack, R. A. and R. K. Lawrence. 1995b. Attack densities of T omicus piniperda and Ips
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1 18.

Haack, R. A., R. K. Lawrence, D. G. McCullough, and C. S. Sadof. 1997. T omicus
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142

 

Hainze, J. H. and D. M. Benjamin. 1983. Bionomics and pattern of attack of the red pine
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Hanson, H. S. 1940. Observations on the life cycle of the pine shoot beetles. Scottish
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Herdy, H. and J. B. Thomas. 1961. The seasonal development of a species of
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