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This is to certify that the

thesis entitled
Interact-.1013 of the Pine Short. Beetle [Tatrims pjmpemda
(L.) (Colecp‘bera: Scolytidaefl with Native Pine Bm‘k
Beetles and Their Peeociated Natural Enemies in Vddfigm
presented by
Pmy Pm Kemedy

has been accepted towards fulfillment
of the requirements for

M.S. __degree in B1tamlogy_

 

 

Date 16 Deoenber 1998

0-7639 MS U is an Affirmative Action/Equal Opportunity Institution

 

PLACE IN RETURN BOX to remove this checkout from your record.
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INTERACTIONS OF THE PINE SHOOT BEETLE [Tomicus piniperda (L.)
(COLEOPTERA: SCOLYTIDAE)] WITH NATIVE PINE BARK BEETLES
AND THEIR ASSOCIATED NATURAL ENEMIES IN MICHIGAN
By

Amy Ann Kennedy

A THESIS

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

MASTER OF SCIENCE
Department of Entomology

1998

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ABSTRACT
INTERACTIONS OF THE PINE SHOOT BEETLE [Tomicus piniperda (L.)
(COLEOPTERA: SCOLYTIDAE)] WITH NATIVE PINE BARK BEETLES
AND THEIR ASSOCIATED NATURAL ENEMIES IN MICHIGAN
By

Amy Ann Kennedy

The pine shoot beetle, Tomicus piniperda (L.) (Coleoptera: Scolytidae), native to
Eurasia, was discovered in North America in 1992. Interactions of T. piniperda with
native pine bark beetles and their natural enemies are not well-known in the Great Lakes
region. The phenology of T. piniperda and native insects in red pine (Pinus resinosa
Ait.) plantations were monitored in 1996 and 1997. Two species of phloem feeders and
four species of predators were active as early in the spring as T. piniperda. The impacts
of predators and parasitoids on the galleries and progeny of T. piniperda and of native Ips
bark beetles were also quantified. There was a negative relationship between natural
enemy density and the number of bark beetle progeny per parent female, but natural
enemies did not significantly affect mean scolytid gallery density, gallery length, or
progeny density. Interspecific competition between T. piniperda and native Ips bark

beetles was studied in laboratory and field experiments. Mean Ips gallery density, gallery
length, progeny density, and productivity were significantly reduced in the presence of T.

pimperda, but T. pim'perda was not affected by the presence of Ips.

This work is dedicated to the memory of Dr. George B. Craig, Jr.,
of the Department of Biological Sciences at the University of Notre Dame.
He passed down his fascination with insects and enthusiasm for science

to hundreds of the students he mentored, including myself.

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ACKNOWLEDGMENTS

I am grateful to my advisor, Dr. Deborah G. McCullough, for giving me the
opportunity to work on this exciting project. I thank Deb for introducing me to the
diversity of the field of forest entomology, and also for all of her assistance and expertise
with the field work. She has been an excellent mentor and role model throughout this
challenging project, and she has always encouraged me to do the best work possible.

My committee members have also contributed greatly to my graduate program.
Dr. Robert Haack worked with Deb and me from the very beginning to design the field
projects, and he has been invaluable in providing me with knowledge about how to work
with bark beetles in the lab and in the field. He generously supplied me with bark beetles
toward the lab experiment, and he also provided me with funnel traps when our shipment
was late. Dr. Edward Grafius has been very encouraging ever since I arrived at Michigan
State University (MSU), and I am grateful for his expertise on scientific writing. He
always makes time to talk with grad students, and I have especially appreciated our chats
about career planning. Dr. Donald Dickmann is an excellent teacher, and he has taught
me a great deal about silviculture, tree physiology, and red pine management. With the
knowledge that Dr. Dickmann has given me, I have seen how closely the biology of bark
beetles and their natural enemies is tied to forest management practices and forest health.

Funding for this study was primarily provided by the USDA Forest Service, North
Central Research Station and the USDA Forest Service, National Center of Forest Health.

I also would like to thank the MSU Department of Entomology for funding a portion of

 

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this research as well as funding my travel expenses to several regional and national
entomological meetings through the Ray and Bernice Hutson Endowment.

Many thanks are also due to the student employees who assisted me in field work:
Nathan Siegert, Kirsten Fondren, Angela Toma, Eric Black, and Philip Garvin. Without
their help in the logistics of maintaining nine field sites and de-barking hundreds of red
pine bolts, this research would not have been a success. The personnel at MSU’s WK.
Kellogg Experimental Forest (Greg Kowalewski, Jim Curtis, John Vigneron, and Karen
Bushouse) were especially helpful in providing red pine bolts, setting up the field sites,
and checking the funnel traps during spring 1996 and 1997.

I would also like to thank the organizations and their staff who helped me locate
the field sites, permitted me to work in their forests, and provided me with the stand
history data: Maria Albright and John Lerg from the Allegan State Game Area; Dr. Carl
Ramm and Greg Kowalewski of the WK. Kellogg Experimental Forest; Matt Sands of
the Huron-Manistee National Forest; Don Torchia of the AuSable State Forest; Roger
Mech of the Department of Natural Resources; and Clara Ward of the F enner Arboretum.

Deb Nelson of the USDA APHIS laboratory in Niles, Michigan and Drs. Robert
Haack and Robert Lawrence of the USDA Forest Service assisted with scolytid
identifications. Jim Zablotney of MSU assisted with identifications of cerambycid adults.
Their help was greatly appreciated.

Finally, I would like to thank my family for all of their support and love
throughout my education. My parents Jerry and Paula Christensen, sisters Becky, Erica,
and Mary, brother Joey, and husband Chris all assisted me with field work when things

were hectic. Their encouragement and love have been invaluable in my life.

 

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TABLE OF CONTENTS

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

CHAPTER 1
Phenology of Parent and Progeny Pine Bark Beetles and their Natural Enemies . . . .
Introduction ........................................................
Materials and Methods ...............................................
Study Sites ......................................................
Funnel Trapping .................................................
Progeny Rearing Study ............................................
Field Observations ................................................
Analysis ........................................................
Results ............................................................
Phloem Feeders ..................................................
Natural Enemies .................................................
Discussion .........................................................
Conclusions .....................................................
Literature Cited .....................................................

CHAPTER 2
Impacts of Natural Enemies on Native Ips spp. and Introduced Tomicus piniperda
(Coleoptera: Scolytidae) Bark Beetles in Red Pine Plantations in Michigan ........
Introduction ........................................................
Materials and Methods ...............................................
Field Sites ......................................................
Funnel Trapping .................................................
Exclusion Study ..................................................
Data Analysis ...................................................
Results ............................................................
Scolytid Progeny Density — 1996 ....................................
Scolytid Progeny Density — 1997 ....................................
Scolytid Productivity — 1996 ........................................
Scolytid Productivity - 1997 ........................................
Scolytid Egg Galleries and Exit Holes — 1996 ..........................
Scolytid Egg Galleries and Exit Holes — 1997 ..........................
Natural Enemies - I996 ...........................................
Natural Enemies — 1997 ...........................................
Discussion .........................................................
Conclusions .....................................................
Literature Cited .....................................................

vi

viii

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CHAPTER 3
Interspecific Competition Between Native Ips pim' and Introduced Tomicus

piniperda (Coleoptera: Scolytidae) Bark Beetles in Red Pine ................... l 14
Introduction ........................................................ 1 14
Materials and Methods ............................................... 117

Field Sites ...................................................... 117
Artificial Infestations .............................................. 118
Field Infestations ................................................. 120
Data Analysis .................................................... 122
Results ............................................................ 123
Artificial Infestations .............................................. 123
Field Infestations: Effect of Location on Ips spp ......................... 123
Field Infestations: Single Species vs. Both Species ....................... 124
Discussion ......................................................... 125
Conclusions ..................................................... 1 30
Literature Cited ..................................................... 131

APPENDIX 1 — Record Deposition of Voucher specimens ..................... 142

APPENDIX 1.1 — Voucher Specimen Data .................................. 143

LITERATURE CITED ................................................. 146

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Chapter 1

Table 1.

Table 2.

Table 3.

Table 4.

Table 5.

Chapter 2

Table 1 .

Table 2.

LIST OF TABLES

General characteristics and number of red pine bolts placed at the
1996 and 1997 red pine plantation field sites ....................

Total ntunber of insects collected from pheromone-baited funnel
traps at either the southern (1996: 4 traps each at KAL] and ALL];
1997: 2 traps each at KAL], KAL2, ALL] , ALL2), or northern
(1996: 4 traps each at WEXI and ROS]; 1997: 2 traps each at
WEX], WEX2, R081, R082) Michigan red pine plantations, or in
7 funnel traps in a red and Scotch pine plantation in Lansing .......

Julian dates (JD) and degree-days base 10°C (DD) (Baskerville-
Emin method) in 1996 when insects were collected from
pheromone-baited funnel traps in either southern (KALl and ALL])
or northern (WEXI and ROS 1) Michigan red pine plantations, or in
a red pine and Scotch pine plantation in Lansing, Michigan ........

Julian dates (JD) and degree-days base 10°C (DD) (Baskerville-
Emin method) in 1997 when insects were collected from
pheromone-baited fiInnel traps in either southern (KALl , KAL2,
ALL], ALL2) or northern (WEXl , WEX2, R081, R082) Michigan
red pine plantations, or in a red pine and Scotch pine plantation in
Lansing .................................................

Total number of insects reared from red pine bolts from the southern
(1996: KAL] and ALL]; 1997: KAL], KAL2, ALL], ALL2), or
northern (1996: WEX] and ROS]; 1997: WEX], WEX2, ROS],
R082) Michigan red pine plantations. A total of 204 bolts were
used in 1996; 304 were used in 1997 ..........................

General characteristics and number of red pine bolts placed at the
1996 and 1997 red pine plantation field sites ....................

Mean densities of scolytids and natural enemies per m2 of phloem
surface area from red pine bolts in 1996. There were a total of 60
bolts in batch 96-1; 88 bolts in batch 96-2; and 56 bolts in batch
96-3 ....................................................

viii

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Table 4.

Table 5.

Table 6.

Table 7.

Table 8.

Table 9.

Densities of scolytid progeny (number reared per m2 of red pine
phloem surface area) in 1996, for bolts with three or more natural
enemies and bolts with two or fewer natural enemies (NE). There
were a total of 60 bolts in batch 96-1; 88 bolts in batch 96-2, and 56
bolts in batch 96-3 ........................................

Mean densities of scolytids and natural enemies per m2 of phloem
surface area reared from red pine bolts in 1997. There were a total
of 112 bolts in batch 97-1; 128 bolts in batch 97-2; and 64 bolts in
batch 97-3 ...............................................

Densities of scolytid progeny (number reared per m2 of red pine
phloem surface area) in 1997, for bolts with three or more natural
enemies and bolts with two or fewer natural enemies (NE). There
were a total of 112 red pine bolts in batch 97-1; 128 bolts in batch
97-2; and 64 bolts in batch 97-3 ..............................

Scolytid productivity (number of scolytid progeny reared per parent
female) in 1996, for bolts with three or more natural enemies and
bolts with two or fewer natural enemies (N E). There were a total of
60 red pine bolts in batch 96-1; 88 bolts in batch 96-2; and 56 bolts
in batch 96-3 .............................................

Scolytid productivity (number of scolytid progeny reared per parent
female) in 1997, for bolts with three or more natural enemies and
bolts with two or fewer natural enemies (NE). There were a total of
112 red pine bolts in batch 97—1; 128 bolts in batch 97-2; and 64
bolts in batch 97-3 ........................................

Mean number of scolytid exit holes and egg galleries per m2 of
phloem surface area in 1996 and 1997. There were a total of 60 red
pine bolts in batch 96-1; 88 bolts in batch 96-2; 56 bolts in batch
96-3; 112 bolts in batch 97-1; 128 bolts in batch 97-2; and 64 bolts
in batch 97-3 .............................................

Number of scolytid exit holes and egg galleries per m2 of phloem
surface area in 1996, for bolts with three or more natural enemies
and bolts with two or fewer natural enemies (N E). There were a
total of 60 red pine bolts in batch 96-1; 88 bolts in batch 96-2; and
56 bolts in batch 96-3 ......................................

ix

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Table 1 1.

Table 12.

Table 13.

Table 14.

Chapter 3

Table 1.

Table 2.

Table 3.

Table 4.

Number of scolytid exit holes and egg galleries per m2 of phloem
surface area in 1997, for bolts with three or more natural enemies
and bolts with two or fewer natural enemies (N E). There were a
total of 112 red pine bolts in batch 97-1; 128 bolts in batch 97-2; and
64 bolts in batch 97-3 ......................................

Total number of scolytid natural enemies reared from red pine bolts
in 1996. There were a total of 60 bolts in batch 96-1; 88 bolts in
batch 96-2; and 56 bolts in batch 96-3 .........................

Percentage of red pine bolts infested with T omicus piniperda (Tp) or
Ips spp. (Ip) from batch 96-1 that also reared natural enemies. Of
the 60 bolts, 32 were not attacked by either scolytid ..............

Total number of scolytid natural enemies reared from red pine bolts
in 1997. There were a total of 112 bolts in batch 97-1; 128 bolts in
batch 97-2; and 64 bolts in batch 97-3 .........................

Percentage of red pine bolts infested with Tomicus piniperda (Tp) or
Ips spp. (Ip) from batch 97-1 that also reared natural enemies. Of
the 112 bolts, 34 were not attacked by either scolytid .............

General characteristics and number of red pine bolts placed at the
1996 and 1997 red pine plantation field sites ....................

Ips pini gallery densities, gallery lengths, progeny density, and
productivity in artificially-infested red pine bolts in 1996, for bolts
infested with I. pini only and bolts infested with equal densities of 1.
pint and Tomicus piniperda .................................

Tomicus piniperda gallery densities, gallery lengths, progeny
density, and productivity in artificially-infested red pine bolts in
1996, for bolts infested with T. piniperda only and bolts infested
with equal densities of T. piniperda and Ips pini .................

Ips spp. gallery densities, gallery lengths, progeny density, and
productivity in field-infested red pine bolts in 1996, for bolts from
southern sites (KF and ALL), where both T. piniperda and I. pim'
were present, and from northern sites (WEX and ROS), where T.
piniperda was absent. There were a total of 60 bolts in batch 96-1;
88 bolts in batch 96-2; and 56 bolts in batch 96-3 ................

100

102

103

104

105

134

135

136

137

Table 5.

Table 6.

Table 7.

Ips spp. gallery densities, gallery lengths, progeny density, and
productivity in field-infested red pine bolts in 1997, for bolts from
southern sites (KF and ALL), where both T. piniperda and I. pini

were present, and from northern sites (WEX and ROS), where T.
piniperda was absent. There were a total of 112 bolts in batch 97-1;

128 bolts in batch 97-2; and 64 bolts in batch 97-3 ............... 138

Ips spp. gallery densities, gallery lengths, progeny density, and
productivity in field-infested red pine bolts in 1996 and 1997, for

bolts infested with Ips spp. only and bolts infested with both Ips spp.

and T omicus piniperda ..................................... 139

Tomicus piniperda gallery densities, gallery lengths, progeny

density, and productivity in field-infested red pine bolts in 1996 and

1997, for bolts infested with T omicus piniperda only and bolts

infested with both Ips spp. and T. piniperda .................... 140

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Figure 1.

Figure 2.

Figure 3.

Figure 4.

Figure 5.

Figure 6.

Figure 7.

Figure 8.

Figure 9.

Figure 10.

Figure 11.

LIST OF FIGURES

Location of red pine forest plantation field sites in Michigan.
KAL], ALL], WEX], ROS 1 , and Lansing were monitored in 1996;
all stands were monitored in 1997 ............................

Seasonal distribution of scolytid adults collected in funnel traps
baited with alpha-pinene, ipsdienol, and lanierone at five Michigan
red pine plantations in 1996 and nine plantations in 1997 ..........

Seasonal distribution of scolytid progeny adults reared from 204 red
pine bolts in 1996 and 304 bolts in 1997 in Michigan .............

Seasonal distribution of scolytid adults collected in funnel traps
baited with alpha-pinene, ipsdienol, and lanierone at five Michigan
red pine plantations in 1996 and nine plantations in 1997 ..........

Seasonal distribution of scolytid progeny adults reared from 204 red
pine bolts in 1996 and 304 bolts in 1997 in Michigan .............

Seasonal distribution of scolytid adults collected in fiinnel traps
baited with alpha-pinene, ipsdienol, and lanierone at five Michigan
red pine plantations in 1996 and nine plantations in 1997 ..........

Seasonal distribution of scolytid progeny adults reared from 204 red
pine bolts in 1996 and 304 bolts in 1997 in Michigan .............

Seasonal distribution of adults collected in funnel traps baited with
alpha-pinene, ipsdienol, and lanierone at five Michigan red pine
plantations in 1996 and nine in 1997 ..........................

Seasonal distribution of progeny adults reared from 204 red pine
bolts in 1996 and 304 bolts in 1997 in Michigan .................

Seasonal distribution of predator adults collected in funnel traps
baited with alpha-pinene, ipsdienol, and lanierone at five Michigan

red pine plantations in 1996 and nine plantations in 1997 ..........

Seasonal distribution of predator progeny adults reared from 204 red
pine bolts in 1996 and 304 bolts in 1997 in Michigan .............

xii

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Figure 13.

Figure 14.

Figure 15.

Figure 16.

Chapter 2

Figure 1.

Figure 2.

Figure 3.

Figure 4.

Figure 5.

Seasonal distribution of predator adults collected in funnel traps
baited with alpha-pinene, ipsdienol, and lanierone at five Michigan
red pine plantations in 1996 and nine plantations in 1997 .......... 57

Seasonal distribution of predator progeny adults reared from 204 red
pine bolts in 1996 and 304 bolts in 1997 in Michigan ............. 58

Seasonal distribution of predator and parasitoid adults collected in

funnel traps baited with alpha-pinene, ipsdienol, and lanierone at

five Michigan red pine plantations in 1996 and nine plantations in

1997 ................................................... 59

Seasonal distribution of predator and parasitoid progeny adults
reared from 204 red pine bolts in 1996 and 304 bolts in 1997 in
Michigan ................................................ 60

Average degree-day (base 10°C) accumulations from 1 March in the
southern sites (KAL and ALL) and northern sites (WEX and ROS),
by Julian date ............................................ 61

Location of red pine forest plantation field sites in Michigan.
KAL], ALL], WEX], ROS], and Lansing were monitored in 1996;
all stands were monitored in 1997 ............................ 106

Relationships between natural enemy density and the productivity of

(A) Tomicus piniperda and (B) Ips pini. All scolytid and natural

enemy progeny adults were reared from batch 96-1. Note the

different scales ........................................... 107

Relationships between natural enemy density and the productivity of

(A) Ips pini and (B) I. grandicollis. All scolytid and natural enemy
progeny adults were reared from batch 96-2. Note the different

scales .................................................. 108

Relationships between natural enemy density and the productivity of

(A) Ips pini and (B) I. grandicollis. All scolytid and natural enemy
progeny adults were reared from batch 96-3. Note the different

scales .................................................. 109

Relationships between natural enemy density and the productivity of

(A) Tomicus piniperda, (B) Ips pini, and (C) I. grandicollis. All

scolytid and natural enemy progeny were reared from batch 97-1.

Note the different scales .................................... l 10

xiii

 

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Figure 6.

Figure 7.

Chapter 3

Figure 1.

Relationships between natural enemy density and the productivity of

(A) Ips pim' and (B) I. grandicollis. All scolytid and natural enemy
progeny adults were reared from batch 97-2. Note the different

scales .................................................. I 12

Relationships between natural enemy density and the productivity of

(A) Ips pim' and (B) I. grandicollis. All scolytid and natural enemy
progeny adults were reared from batch 97-3. Note the different

scales .................................................. 1 13

Location of red pine forest plantation field sites in Michigan.
KAL], ALL], WEX], ROS], and Lansing were monitored in 1996;
all stands were monitored in 1997 ............................ 141

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CHAPTER 1

Phenology of Parent and Progeny Pine Bark Beetles and their Natural Enemies

INTRODUCTION

The larger pine shoot beetle, Tomicus (=Blast0phagus =Myelophilus) piniperda
(L.) (Coleoptera: Scolytidae), is a well-known forest pest in Europe, Asia, and North
Africa, where it infests several species of pine (Pinus spp.) (Langstrom 1980; Ye 1991).
Established populations of T. piniperda were first discovered in North America in 1992
in a Scotch pine (Pinus sylvestris L.) Christmas tree plantation in Ohio (Haack and
Kucera 1993). Haack and Lawrence (1995a) speculated that this bark beetle probably
came to North America from Europe on cargo ships in wood packing material or dunnage
(logs used to support cargo). As of September 1998, T. piniperda had been found in 243
counties in nine northeastern US. states, and 21 counties in the Canadian province of
Ontario (NAPIS 1998).

With the introduction of an exotic insect comes the need to understand
interactions between the new exotic insect and native insects with similar habits.
Tomicus piniperda overwinters as adults, and early in spring, when temperatures reach
10-12°C, parent adults become active and fly to recently-cut pine stumps and logs for
breeding (Langstrom 1980, Haack and Lawrence 1995b). Tomicus piniperda typically
reproduce only in dead, recently cut, or severely weakened pines (Byers 1991, Schroeder
1992). Alpha-pinene and other monoterpene volatiles in pine oleoresin attract both sexes

of T. piniperda to wounded and freshly cut or fallen pine trees (Byers 1991, Schroeder

 

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I988, Schroeder and Lindelow 1989, Vite et a1. 1986), and a short range sex pheromone
for T. piniperda has recently been identified (S. Teale, State University of New York
Syracuse, personal communication). Afier locating a suitable host, females bore into the
pine bark and begin excavating individual egg galleries. After a male enters the gallery
and they mate, the female lays eggs along the sides of a long, straight gallery that
parallels the grain of the wood (Langstrom 1980). After establishing one brood, the
parents may emerge, excavate a new gallery, and produce a “sister” brood (Schroeder and
Risberg 1989). Larvae feed and develop in the phloem for 6 — 12 weeks, then pupate.
The progeny adults exit the bark and fly to the crowns of living pine trees. Each beetle
bores into the distal tip of one to six current-year or one-year-old shoots, and tunnels
toward the shoot tip, killing the shoot. This maturation feeding continues throughout the
summer (Langstrom 1980, McCullough and Smitley 1995).

Many other Great Lakes insects also utilize the phloem of weakened or recently-
killed pine trees. The pine engraver, Ips pini (Say) (Coleoptera: Scolytidae), is the most
economically important and well-studied pine bark beetle of the Great Lakes region
(Raffa 1991, Schenk and Benjamin 1969). This native bark beetle overwinters as an
adult in the litter of pine stands, and initiates spring flight in late April or May in
Michigan (Haack and Lawrence 1995b), when daily maximum temperatures are near
20°C (Schenk and Benjamin 1969). Like T. piniperda, 1. pim’ is a secondary pest that
breeds preferentially in recently-killed or damaged hosts, such as freshly cut pines,
logging slash, or pines recently damaged by wind or ice (Coulson and Witter 1984,
Schenk and Benjamin 1969). Ips pini is attracted to these host pines in large numbers via

host volatiles and two aggregation pheromones that I. pini males produce: ipsdienol and

 

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males create nuptial chambers in the phloem and mate with 2 -— 4 females (Schenk and
Benjamin 1969). Each female excavates an egg gallery that leads away from the nuptial
chamber, typically resulting in a “Y” or “H” shaped gallery that is distinct from T.
piniperda galleries. Ips pim' larvae and pupae develop under pine bark, but unlike T.
piniperda, I. pim' progeny adults do not require maturation feeding in shoots. Instead,
newly emerged adults locate new host material to colonize, unless it is late summer or
fall. In the Great Lakes region, 1. pini typically completes three generations per year
(Schenk and Benjamin 1969).

Other common pine bark beetles (Coleoptera: Scolytidae) in the Great Lakes
region include Ips grandicollis (Eichhoff), Orthotomicus caelatus (Eichhoff), and
Dendroctonus valens LeConte (Haack and Lawrence 1995b, Schenk and Benjamin 1969,
Raffa 1991). Larvae of weevils (Coleoptera: Curculionidae) and long-homed beetles
(Coleoptera: Cerambycidae) also compete with scolytids for pine phloem resources.

Red pine (Pinus resinosa Ait.), one of the most important and widely planted
species for pulp, sawtimber, and poles in the Great Lakes region (Rudolf 1990), is a
suitable host for T. piniperda and many native scolytids (Haack and Lawrence 1997b).
Management of red pine plantations typically involves multiple thinnings or partial
harvests during a rotation, each of which provides an abundance of suitable breeding
material for bark beetles and related groups (Haack and Lawrence 1995b). To date, most
of the research on T. piniperda in North America has taken place in Scotch pine
Christmas tree plantations (Haack and Lawrence 1997a, Kauffman et a1. 1998,

McCullough and Smitley 1995, McCullough et al. 1998, McCullough and Sadof 1998).

 

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Red pine plantations represent a more stable ecosystem with no insecticide use, and,
presumably, more natural enemies, and there is a need to learn more about the impacts
and the interactions of T. piniperda in a forest ecosystem.

Pine phloem in dead and dying trees is an ephemeral resource in both space and
time, and bark beetles must locate and colonize suitable brood material before the phloem
becomes too dry for larval development. Because T. piniperda becomes active in the
spring four to six weeks earlier than Ips spp. in the Great Lakes region (Haack and
Lawrence 1995b, Haack and Lawrence 1997b), it may have an opportunity to colonize
pine phloem for several weeks without interspecific competition from native bark beetles.
Determining the spring emergence of both native and exotic bark beetle species is the
first step in understanding the interactions between them and determining the degree to
which competition for breeding material occurs.

Interactions between a newly established exotic insect and native natural enemies
that may attack it are also important. Most predators can only prey upon one life stage or
upon a certain size of an insect because of physical limitations. For example, Anthocoris
spp. (Hemiptera: Anthocoridae) are egg predators of many pine bark beetles, but are too
small to prey upon large larvae or adults (Schmitt and Goyer 1983). Most parasitoids are
also stage- or size-specific (Jones and Stephen 1994, Kudon and Berisford 1980, Senger
and Roitberg 1992). Therefore, the timing of emergence of a natural enemy in relation to
its prey is vital to the natural enemy’s success.

Most bark beetle natural enemies in the Great Lakes region are phenologically
adapted to native bark beetles, becoming active in April or May (Raffa 1991). Haack and

Lawrence (1995b) predicted that T. piniperda would encounter no native predators or

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parasitoids until colonization and oviposition was mostly completed each spring. This
concern was large enough to warrant consideration of importing the exotic predator
Thanasimusformicarius (L.) (Coleoptera: Cleridae) to the United States soon after the
discovery of widespread T. piniperda establishment (Haack and Lawrence 1995b, APHIS
1996). Thanasimusformicarius is the earliest of the major bark beetle predators in
Europe to become active in the spring (Schroeder 1988). The native clerid predator

T hanasimus dubius (F .) is known to be well-adapted to the later-emerging native Ips spp.
bark beetles, and even uses Ips spp. aggregation pheromones as kairomones to locate
their hosts (Herms et a1. 1991; Raffa 1991; Teale et a1. 1991). It was unclear whether T.
dubius and other native natural enemies would emerge from overwintering early enough
in the spring to prey upon T. piniperda parent adults or progeny.

In this study, I examined the phenology of pine bark beetles and their arthropod
natural enemies in eight Michigan red pine plantations, using three methods: pheromone-
baited funnel traps, observations of activity on red pine bolts, and rearing from red pine
bolts. My objective was to compare the emergence of parent and progeny adults of the
exotic T. piniperda with that of native pine phloem-feeding insects and native natural

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MATERIALS AND METHODS

Study Sites

In 1996, I monitored phenology of overwintering and progeny bark beetles, and
their arthropod natural enemies at four red pine forest plantations in Michigan. Two
plantations were in southwestern Michigan and were known to be infested with both T.
piniperda and native Ips spp. These stands were located in Michigan State University’s
WK. Kellogg Experimental Forest in Kalamazoo County (KALl), and in the Allegan
State Game Area in Allegan County (ALLl) (Figure 1). Two plantations in northern
lower Michigan, where T. piniperda was not expected to be present, were also selected.
One stand was in the Huron-Manistee National Forest in Wexford County (WEX] ), and
the other was in the AuSable State Forest in Roscommon County (ROSl). Although T.
piniperda was later detected in these northern counties in regulatory surveys (N APIS
1998), I did not observe any T. piniperda life stages or evidence of infestations in the
northern plantations in either 1996 or 1997.

All four field sites shared the following characteristics: they were pole-sized red
pine plantations of similar size, age, and basal area (Table 1) with some hardwood
understory. Each field site was within 2 km of other red pine plantations, and each site
had been thinned during the 1994-95 winter. An abundance of slash (for example,
trunks, tops, large branches) was left on the ground from these thinnings and was
expected to attract scolytids in 1995. Temperature, precipitation, and degree day
accumulation (base 10°C) were monitored by Michigan State University weather stations

near all four sites.

In 1997, I monitored the phenology of bark beetles and their natural enemies in
the same four sites described above, which represented conditions two years after
thinning. I also selected four additional red pine plantations, each one within 6.2 km of
one of the original stands (KAL2, ALL2, WEX2, and R082). These sites had the same
general characteristics of the original stands (Table 1), but were row-thinned during the
1995-1996 winter.

In both 1996 and 1997, the phenology of overwintering bark beetles and their
natural enemies was also monitored at a mixed Scotch pine and red pine plantation in
Lansing, Ingham County, Michigan (Figure 1). This stand, located in the F enner
Arboretum, was chosen because of its proximity to the Michigan State University campus
and its intermediate location between the northern and southern sites. Observations of
galleries in 1995 indicated that this plantation was infested with both T. piniperda and Ips
spp. Brood material, consisting of dying and newly dead trees was available each year
because the plantation suffers from annosus root rot (Heterobasidion annosum (Fr.)
Bref.) and because of wind damage to trees and to tops.

Funnel Trapping

To monitor the spring flight activity of scolytids and their associates, two to four
12-unit Lindgren funnel traps (Lindgren 1983) (from PheroTech, Inc., Delta, British
Columbia) were placed in each red pine stand, and seven funnel traps were placed in the
Lansing Scotch/red pine stand. Traps were hung from metal poles to keep the trap
bottoms approximately 1 m above the ground. Each trap was erected and baited in
February of 1996 and 1997 with two or-pinene lures to attract T. piniperda (Byers 1991;

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1988), and with one lure each of ipsdienol and lanierone. Ipsdienol and lanierone are
aggregation pheromones of 1. pim' that are used as kairomones by predaceous clerid,
tenebrionid, and histerid beetles (Herms et a1. 1991; Raffa 1991; Teale et a1. 1991).

All lures were manufactured by Phero Tech, Inc., and the following information
came from the company. The or-pinene lures were made up of 15 ml polyethylene bottles
with a release rate of 150 mg/day at 22°-24°C, and a 10:90 ratio of (+) and (-)
enantiomers. The ipsdienol bubble-cap lures had a release rate of 100 ug/day at 25°C,
and a 50:50 enantiomeric ratio. The lanierone bubble-cap lures had a release rate of 10
pg/day at 25°C. I placed insecticide strips in each trap, and collected insects from traps
at weekly intervals from March through September in 1996 and 1997.

Progeny Rearing Study

A total of 204 red pine bolts in 1996, and 304 bolts in 1997, each approximately
61 cm long and 10 — 20 cm diameter, were cut from live red pine trees at the red pine
field sites in February, April, and June of both years (Table 1). These times roughly
corresponded to the initial activity of overwintering parent T. piniperda, initial activity of
overwintering parent Ips spp., and emergence of the first Ips progeny adults. February-
cut bolts were only placed in the southern sites because T. piniperda was not known to be
present in the northern sites. The bolts were placed on the ground in a shady area of each
plantation to reduce desiccation and were left for colonization by bark beetles and their
associates. Bolts were returned to Michigan State University at two week intervals in
1996 and at one week intervals in 1997. The cut ends of the retrieved bolts were dipped
in paraffin wax to reduce desiccation. Each bolt was placed in an individual emergence

container consisting of a cardboard tube (15 — 25 cm diameter, 0.32 — 0.64 cm wall, 61 —

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71 cm overall length [Michigan Can and Tube, Inc., Saginaw, MI]) with opaque plastic
endcaps and a clear plastic collection cup at one end. The emergence containers were
stored on the Michigan State University campus in East Lansing in a screened outdoor
insectary to expose the bolts to ambient temperature and humidity.

Bark beetle and natural enemy progeny adults emerging from the bolts were
collected from emergence containers at least twice a week from June through October.
When progeny emergence stopped in the winter, each log was de-barked and any insects
still under the bark or dead on the bottom of the emergence container were collected. All
insects reared from these bolts were assumed to be progeny adults, although it is possible
that a few parent adults were inadvertently collected. The number of parent insects was
assumed to be small and similar across all bolts and sites.

Field Observations

In addition to collecting insects from the funnel traps at regular intervals, I also
examined the red pine bolts that had been left in the stands for the progeny rearing study
in 1997. Each bolt was gently turned over to examine all sides. I recorded the presence
and quantity of the following variables: frass from boring scolytids, exit holes of bark
beetles, adult bark beetles or other phloem borers walking on the bolts, and adult and
larval predators on the bolts. The bark was not peeled away because the bolts used for
the observations were the same ones used to rear progeny, and 1 did not want to alter or
destroy the phloem environment in those bolts.

Analysis
Phenology of the parent bark beetles and their natural enemies was examined by

plotting the funnel trap collection (as proportion of total collected) of each species over

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time. Each year (1996 and 1997) was plotted separately. Phenology of the progeny bark
beetles and their natural enemies was determined by plotting collections from the
emergence tubes (as proportion of total reared) of each species over time, with each year
analyzed separately. Progeny insects that were collected during the de-barking are not

included in these graphs, since the date of emergence of these insects was not known.

10

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RESULTS

Phloem Feeders

Thirteen species of pine phloem feeders, including ten bark beetle species, one
weevil, and two long-homed beetles, were collected in the pheromone-baited funnel
traps. A total of 5296 adults were collected in 1996, and 2614 adults were collected in
1997 (Table 2). T omicus piniperda was the first pine bark beetle to be found in the
funnel traps at the southern sites (KF and ALL) and in Lansing in 1996 and in 1997
(Tables 3 and 4). Most native bark beetles, including Ips pini, Hylurgops rugipennis
pinifex (Fitch), and Ips grandicollis were first collected in funnel traps approximately 45
days after T. piniperda’s first flight. Twelve species of pine phloem feeders were reared
from the red pine bolts, including nine bark beetle species, Pissodes nemorensis Germar
(Coleoptera: Curculionidae), Monochamus scutellatus (Say), and M. carolinensis
(Olivier) (Coleoptera: Cerambycidae). In 1996, 19,045 progeny adults were reared, and
22,796 progeny were reared in 1997 (Table 5). Tomicus piniperda progeny were the first
pine bark beetles to emerge in both years. Ips perroli Swaine (Coleoptera: Scolytidae)
Progeny emerged at the same time as T. piniperda progeny in 1996, but were four weeks
later than T. piniperda in 1997. Monochamus scutellatus and M. carolinensis progeny
Were the last pine phloem feeders to emerge in both years; adults did not emerge until the
following spring.

Flight of T. piniperda parent adults was early and quite brief; over 90% of all
Parent adults were collected in the first two weeks of activity in 1996 and in 1997 (Figure

2). Parent T. piniperda adults were observed on red pine bolts in the southern sites

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during the same two-week period that they were first collected in funnel traps. Both
years, T. piniperda flew a few days earlier in Lansing than in KF or ALL (Tables 3 and
4). There were approximately 100 days between T. piniperda parent activity in funnel
traps and T. piniperda progeny emergence. Emergence of T. piniperda progeny adults
occurred during a brief period early in both summers; most of the progeny emerged by
Julian date 200 (Figure 3).

Funnel trap collections of I. pini overwintering parent adults varied dramatically
between 1996 and 1997. Parent 1. pini adults initiated spring flight approximately six
weeks after T. piniperda adults in both years. In 1996, the majority of the parent adults
were collected in late summer, after Julian date 220 (Figure 2). But in 1997, half were
collected within three weeks of first flight. In both years, there were three peaks of
collection (Figure 2), reflecting the three generations of I. pim’ in Michigan. Observations
of I. pim’ activity on red pine bolts coincided with the collection of I. pini in the fimnel
traps. There was little difference in dates of first I. pim’ flight between northern and
southern sites in either year (Tables 3 and 4).

The first I. pim’ progeny adults were collected from emergence containers about
60 days after parent adults were first collected in funnel traps; progeny emergence began
around Julian date 200 of 1996, and Julian date 180 in 1997 (Figure 3). [ps pim’ progeny
adults continued to emerge throughout the summer and into the fall of both years. In

1996, I. pim‘ progeny emergence peaked in early October, and in 1997, progeny adult
emergence remained steady through the summer, with no discernible peaks. Red pine is
not recorded as a host for I. pini by Wood (1982), but several studies report I. pim’

colonization in red pine (Raffa and Klepzig I989, Raffa 1991, Teale and Lanier 1991).

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In both 1996 and 1997, the parent adult flight for Ips grandicollis peaked soon
after flight began, and a second peak occurred seven weeks later (Figure 2) when F 1

progeny adults emerged. Observations of I. grandicollis activity on the red pine bolts
were consistent with the funnel trap collections and indicated that two generations of 1.
grandicollis occurred in both years. Flight of I. grandicollis parent adults occurred one
or two weeks earlier in the southern sites than the northern sites (Tables 3 and 4). There
were 50 - 60 days between flight of overwintering parents and flight of first generation
progeny adults. Ips grandicollis progeny emergence occurred at the same time as I. pini
in 1996, and a few weeks later than I. pini progeny in 1997 (approximately Julian date
200 of both years) (Figure 3). Progeny emergence began in late July of both years and
continued through November with no distinct peaks of emergence.

Hylurgops rugipennis pinifex was not an abundant scolytid in funnel traps at any
site (Table 2). However, I observed many parent adults actively colonizing the red pine
bolts early in the spring when T. piniperda was also colonizing bolts. The few Hylurgops
parent adults found in funnel traps were not collected until several weeks after they were
first observed colonizing the bolts (Figure 4). As the snow was melting in the early
Spring, HyIurgops adults were found along the cool, damp bottom of nearly every red
Pine bolt at all four (1996) or eight (1997) sites, constructing galleries and laying eggs.

In addition, Hylurgops rugipennis pinifex progeny adults were one of the first scolytids to
be reared from the red pine bolts in 1997 (Figure 5). There were about 50 days between
Parent flight and progeny emergence in 1996, but only 20 days difference in 1997. In
1996, there were two distinct peaks of progeny adult emergence: around Julian date 190

and again after Julian date 300 (Figure 5). In 1997, Hylurgops progeny emerged from

13

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Julian date 160 through the rest of the summer (Figure 5). H. rugipennis pinfex was
collected in funnel traps and reared from all nine field sites, which are new records in
Michigan according to Wood (1982). This scolytid appears to have at least two
generations per year, and since its generation time was only 20-50 days, a third
generation appears possible in late summer.

Orthotomicus caelatus parent adults had one peak of flight activity in 1996, just
after it was first collected (Figure 4). In 1997, however, this beetle had two peaks of
flight, approximately seven weeks apart (Figure 4). This bark beetle was observed on red
pine bolts only a few times, immediately after it was first collected in funnel traps, at the
same time as I. pini and I. grandicollis. There was no difference in date of first activity
between northern and southern sites (Tables 3 and 4). There were about 50 days between
first parent flight and first emergence of progeny 0. caelatus in both years. Orthotomicus
caelatus progeny adults were first collected around Julian date 190 and emergence
continued throughout the summer in both years (Figure 5). There were about seven
weeks’ difference between initial parent flight and initial progeny emergence, which
concurs with the seven week separation of the first and second peaks in the 1997 funnel
trap collection. There appear to be at least two generations of 0. caelatus per year.

Ips perroti parent adults were not collected in funnel traps in either year, but
Progeny were reared from the red pine bolts from the northern sites in both years (Table
5). In 1996, most of the progeny adults emerged quite early in the season, before Julian
date 200, similar to T. piniperda progeny (Figure 5). But in 1997, peak progeny
emergence did not occur until mid-summer, coinciding with the other Ips species (Figure

5). In both years, I. perroti progeny emerged in mid-summer and again in

14

 

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September/October, suggesting at least two generations per year. This small bark beetle
was not observed on the red pine bolts. Since overwintering parent adults were not
collected in funnel traps, no estimate of generation time can be made. Ips perroti was
reared from bolts from two of the northern sites (CADI and R082), and the CAD] site is
a new county record in Michigan (Wood 1982).
In both 1996 and 1997, overwintering parent adults of Gnathotrichus materiarius
(Fitch) (Coleoptera: Scolytidae) were first found in funnel traps around the same time as
the overwintering I. pini adults (Tables 3 and 4). Gnathotrichus materiarius parent adults
had one main peak of activity soon after first flight in 1996 (Figure 6). In 1997, it had
two peaks of activity: one immediately after first flight, and one six weeks later (Figure
6). I never observed this tiny bark beetle on the red pine bolts, so I do not know whether
flight activity coincided with mating and gallery formation in this species. About 50 days
separated the first parent flight and emergence of the first progeny adults. Gnathotrichus
materiarius followed the same pattern as did 0. caelatus: progeny emergence began just
before Julian date 200, peaked in mid-summer, and continued throughout the summer
(Figure 7). This scolytid appears capable of producing two or three generations per year.
Red pine is a new host record for G. materiarius (Wood 1982).
In 1996, Hylastes porculus Erichson (Coleoptera: Scolytidae) parent adults flew
much later in the northern sites than the southern sites (Table 3), but in 1997, there was
no difference between northern and southern sites (Table 4). In both years, most of the
H. porculus parent adults were collected soon after first flight, with a second smaller peak
immediately after the first (Figure 6). This beetle was not observed on red pine bolts in

either year. There were approximately 70 days between first parent flight and first

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progeny adult emergence. In both 1996 and 1997, H. porculus progeny first emerged
around Julian date 210. Another emergence peak occurred after Julian date 275 (Figure
7). As evidenced by the two peaks in progeny emergence and the 70 day generation time,
Hylastes porculus appears to have two generations per year. This bark beetle was reared
from all eight field sites and collected from funnel traps in Lansing, which are all new
county collection records (Wood 1982).
The tiny beetle Dryocetes autographus (Ratzeburg) (Coleoptera: Scolytidae) was
quite rare in the funnel traps; only two adults were collected in 1996 and three in 1997,
all from the Lansing funnel traps (Table 2), and at the same time as 1. pini (Figure 6).
Dryocetes autographus was not observed on the red pine bolts in the field, but it was
reared from bolts from the KF 1 and KF 2 sites (Table 5). Progeny adults of D.
autographus began emerging in late June of both years, and emergence continued
through September in 1996, and through August in 1997 (Figure 7). Forty days separated
first parent flight and first progeny emergence in 1996; there was only 30 days’
difference in 1997. Progeny of Dryocetes autographus had two distinct peaks of
emergence in both years of the study, indicating two generations per year. Red pine is a
new host record for this scolytid (Wood 1982).
Pissodes nemorensis, the eastern pine weevil, was not frequently found in funnel
traps (Table 2), but parent adults were commonly observed feeding and ovipositing on
the bottom of red pine bolts in KF l and ALL] early in the spring when T. piniperda and
H. rugipennis pinifex were active. Progeny P. nemorensis adults emerged approximately
100 days after parent adults were collected. Pissodes ncmorensis progeny emerged over

the course of the entire summer, but peaked around Julian date 225 in both 1996 and

16

 

 

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Michigan.

The predominant cerambycid was Monochamus scutellatus, with M. carolinensis
comprising only a small percentage of the funnel trap collections. The two cerambycids
could not be distinguished in the field, so the numbers represent combined totals.

C erambycids were the last of the pine phloem feeders to be collected in funnel traps
(Tables 3 and 4). Parent adult flight activity was irregular but lasted from mid- to late-
summer of both years (Figure 8). Monochamus beetles were very conspicuous from mid-
June through August on nearly all of the red pine bolts in both northern and southern
sites. Though not many adults were collected in funnel traps (Table 2), they were
frequently observed mating and ovipositing on the red pine bolts during this period.
Monochamus scutellatus and M. carolinensis progeny adults were not collected in the
emergence containers in either 1996 or 1997 until the bolts were de-barked the following
winter and spring (Table 5). Progeny larvae were also collected from the bolts during de-
barking; 523 larvae were collected in 1996, and 429 larvae were collected in 1997. These
cerambycids take a full year for development, as evidenced by the single flight period
and the large number of larvae that were collected during de-barking.

Overwintering parent adults of the red turpentine beetle, Dendroctonus valens,
were fairly common in the funnel traps in both years (Table 2). Dendroctonus valens was
collected in funnel traps from all nine sites, which are all new county records in Michigan
(Wood 1982). Parent adults had one main peak of flight activity in both years,
immediately after first flight (Figure 8), and were occasionally observed on red pine bolts

during the same period. Dendroctonus valens became active at approximately the same

17

 

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time in the southern and northern sites (Tables 3 and 4). Although D. valens parent
adults were commonly collected in funnel traps, D. valens progeny adults were not reared
from red pine bolts in either year, probably because this beetle is larger than the other
scolytids, and may prefer thicker bark or phloem, such as in pine stumps (Table 5).
Natural Enemies

Eight different species of bark beetle predators were collected in the pheromone-
baited funnel traps. In 1996, a total of 957 adult predators were collected, and in 1997,
930 predators were collected. Bark beetle parasitoids were not collected in the funnel
traps in either year. Staphylinid beetles (Coleoptera: Staphylinidae) and C ucujus clavipes
F. (Coleoptera: Cucujidae) were the first arthropod natural enemies of bark beetles to be
caught in funnel traps in both 1996 and 1997 (Tables 3 and 4), initiating spring activity
within approximately a week after peak flight of T. piniperda parent adults in both years.
Progeny of nine species of predators and six species of parasitic wasps were reared from
the red pine bolts. A total of 1113 progeny adults were reared in 1996, and 2258 were
reared in 1997.

The clerid predator T hanasimus dubius was among the most abundant and
Conspicuous of the scolytid natural enemies. In 1996, T. dubius adults were first
collected in funnel traps less than three weeks after peak T. piniperda parent activity, and
Continued to be collected throughout the summer and into the fall (Figure 10). In 1997,
T. dubius was not collected in funnel traps until about seven weeks after peak T.
Piniperda activity, corresponding to flights of native Ips spp. parent adults (Figure 10).

The 1997 funnel trap collections of T. dubius only continued until the end of July (Figure

10).

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In contrast to the funnel trap catches, however, I observed T. dubius adults
actively moving about on the red pine bolts early in the spring. In 1997, T. dubius was
observed actively preying upon T. piniperda and H. rugipennis pimfex adults in the
southern sites as early as April 18 (Julian date 108), only 2 '/2 weeks after T. piniperda
peak flight, again on April 25 (Julian date 1 15), May 7 (Julian date 127), and May 23
(Julian date 143), all before native Ips spp. flight activity began and before T. dubius was
collected in funnel traps. My observations of lab colonies of T. dubius adults (maintained
at 10°C) also show that T. dubius can be an active predator in cool weather, before it is
warm enough to fly to funnel traps. Thanasimus dubius adults held at 10°C actively
walked around the lab containers and would attack and consume any bark beetle adults
placed into the container, but I did not observe them flying at this temperature.

Larvae of T. dubius were observed walking on the surface of the red pine bolts at
the R082 and CAD2 sites on July 2, 1997 (Julian date 183). Thanasimus dubius progeny
adults were the first predator species collected in emergence containers in both years, but
these early individuals were probably parent adults. Thanasimus dubius progeny
emergence lasted from mid- to late-summer of 1996, peaking around Julian date 250
(Figure 11). In 1997, progeny emergence began much later in the summer, after Julian
date 225, and peaked nearly 50 days later (Figure 11). There were 90 days in 1996 and
60 days in 1997 between first collections of T. dubius parent adults in funnel traps and
first emergence of progeny adults from red pine bolts. Thanasimus dubius adults that
emerged from the rearing containers prior to August were probably not progeny, but
parent adults that were on the bolts when they were collected from the field sites.

Although T. dubius was active over several periods of the summer, it appears to only

19

 

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The histerid predator Platysoma (=Cy1istix) cylindrica (Paykull) (Coleoptera:
Histeridae) was commonly collected in funnel traps at all sites (Table 2). In 1996, P.
cylindrica was first collected only four weeks after peak T. piniperda flight (Table 3), and
over 70% of the beetles were collected within three weeks of initial flight (Figure 10). In
1997, P. cylindrica began flight activity ten weeks after peak T. piniperda activity (Table
4), and over 80% of the beetles were collected within four weeks (Figure 10). This tiny
predator was infrequently observed on red pine bolts in the summer months, and no
conclusions can be drawn regarding flight activity and predation activity. Emergence of
P. cylindrica progeny also differed greatly from 1996 to 1997. In 1996, progeny
emergence continued throughout the entire summer, with no distinct peaks (Figure 1 1).
In 1997, offspring emergence began in late summer, after Julian date 250, and peaked
within 25 days (Figure 11). There were only 25 days between initial parent and progeny
flights in 1996, and more than 100 days difference in 1997. As was the case with T.
dubius, the emergence of P. cylindrica adults from the emergence containers throughout
the summer of 1996 probably reflects numbers of parent adults that remained on the
field-infested bolts when they were brought to the MSU rearing facility. The peak
progeny emergence of this histerid occurred in September and October, and it probably
only has one generation per year.

C orticeus parallelus Melsheimer (Coleoptera: Tenebrionidae) is another tiny
predator that was frequently attracted to the funnel traps. In 1996, C. parallelus adults
were collected in funnel traps two weeks after T. piniperda peak flight (Table 3).

Although C. parallelus flight activity peaked early in the spring, adults were collected

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throughout the summer (Figure 10). In 1997, C. parallelus did not initiate spring flight
until ten weeks after T. piniperda peak flight (Table 4), and was then collected until mid-
July (Figure 10). C. parallelus was rarely observed on red pine bolts, and only during the
warm summer months. In 1996, emergence of C. parallelus progeny adults began around
Julian date 220, and peaked approximately 60 days later (Figure 11). Timing of
emergence was similar in 1997, but there was a second emergence peak near Julian date
275 (Figure 11). There were approximately 75 days between first parent flight and first
progeny emergence in both years. There were two peaks of progeny emergence in 1997:
one in September and one in late October, which suggests that two generations per year
are possible.
Two staphylinid species (Coleoptera: Staphylinidae) were active early in spring
1996; over 50% of these staphylinids were collected within two weeks of peak T.
piniperda flight (Figure 12). In 1997, parent staphylinids had two main activity periods:
early spring, near T. piniperda peak flight, and mid-summer, corresponding to flight of
first generation Ips spp. progeny adults (Figure 12). Since 1 did not observe many
staphylinids on the red pine bolts, nor did I observe any preying upon bark beetles, I
cannot confidently connect the dates of first flight activity with the dates of first
predatory activity. Staphylinids were collected earlier in southern sites than in northern
sites, and there was no early flight of staphylinids in the northern sites in 1996 (Table 3).
In 1996, staphylinid progeny adults emerged at three times: Julian dates 210 - 225, at
275, and again at 320 (Figure 13). In 1997, progeny adults emerged earlier; around
Julian date 175, and continued to sporadically emerge until Julian date 240 (Figure 13).

There were approximately 120 days separating first parent flight and first progeny

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emergence in 1996, and only 100 days difference in 1997. These predators appear to be
univoltine, but the results in 1997 show that two generations per year are possible.

Medetera sp. (Diptera: Dolichopodidae) was not collected in funnel traps during
either year, but was reared from the red pine bolts in both years. Timing of progeny
emergence was quite similar in both years, with activity beginning around Julian date 200
and ending about 50 — 60 days later (Figure 13). Since no parent adults were collected in
the funnel traps, I cannot estimate the length of time between generations. During de-
barking, I collected hundreds of Medetera sp. larvae, so it appears that Medetera sp.
produces two generations per year in Michigan: the first generation emerges in mid-
summer, and the second generation emerges the following spring.

A second histerid predator, Platysoma parallelum Say (Coleoptera: Histeridae),
began flying five weeks after peak T. piniperda parent flight in 1996 (Table 3), but was
most commonly collected during peak Ips spp. parent flight (Figure 12). In 1997, P.
parallelum was first collected ten weeks after peak T. piniperda flight, like many of the
other scolytid predators that year (Table 4). In 1997, it was collected through the end of
July (Figure 12). This predator was not observed on the red pine bolts. Progeny P.
parallelum adults were reared from all sites in both years (Table 5). In 1996, P.
parallelum progeny began emergence in September and continued through October
(Figure 13). In 1997, no progeny were collected until the bolts were de-barked in the
winter (Table 5). There were approximately 100 days separating first parent flight and
first progeny emergence in 1996. Since progeny of this histerid were only collected late
in the season of 1996, and not until de-barking in 1997, it appears that the funnel trap

peaks represent activity, not different generations. Plarysoma parallelum appears to be

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Cucujus clavipes was not a common predator in funnel traps, but it was among
the earliest predators to become active in the spring (Tables 3 and 4). Over 80% of the
1996 total and over 50% of the 1997 total were collected within two weeks of peak T.
piniperda parent flight in the corresponding years (Figure 14). In 1996 it was only
collected in the southern sites and in Lansing, and in 1997 it was collected in all sites, but
much later in the northern sites. Cucujus clavipes was not observed on the red pine bolts
or preying upon bark beetles, and it was not reared from any of the red pine bolts.

Another clerid predator, Enoclerus nigripes Say (Coleoptera: Cleridae), was much
less commonly encountered than T. dubius. Enoclerus nigripes was among the last of the
scolytid predators to be collected in funnel traps in the southern sites; the first individual
was collected in late July of both years (Tables 3 and 4). In the northern sites, E. nigripes
initiated flight at approximately the same time as Ips pini (Tables 3 and 4). In 1996, E.
nigripes was most often collected in July and August, whereas in 1997, it was mostly
collected in June and July (Figure 14). Enoclerus nigripes was often observed actively
searching for prey on the red pine bolts from all of the sites in the late summer months,
but it was only reared from bolts from the northern sites (Table 5). In 1996, 110 days
separated first parent flight and first progeny emergence; in 1997, there were 90 days
difference. Since E. nigripes progeny adults were only collected in September of both
1996 and 1997 (Figure 15) and during de-barking the following winters, this clerid is
probably univoltine in Michigan.

The anthocorid predator Anthocoris sp. was not collected in the funnel traps. but

nymphs and adults were reared from the red pine bolts in both years. In 1996, Anthocoris

23

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sp. progeny emerged from bolts three times between Julian dates 220 and 290 (Figure
15). In 1997, progeny were collected between Julian dates 185 and 275 (Figure 15). No
estimates of generation time could be made since no parent adults were collected in the
funnel traps.

Parasitic wasps (Hymenoptera: Braconidae, Ichneurnonidae, and Chalcidoidea)
were not collected in the funnel traps in either 1996 or 1997, but six species of parasitoids
were reared from the red pine bolts throughout the summers (Julian dates 175 — 300) of
both years (Figure 15). Since no parasitoids were collected in the funnel traps in either

year, the length of time between generations cannot be estimated.

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DISCUSSION

As expected, T omicus piniperda was the first bark beetle to initiate spring flight,
but, surprisingly, it was not the first phloem feeder to begin colonizing the red pine bolts.
Hylurgops rugipennis pinifex and Pissodes nemorensis were actively colonizing the bolts
at the same time as T. piniperda, even though they were rarely collected in the
pheromone-baited funnel traps early in the spring. The early spring activity of these
phloem borers has not been previously recorded, and T. piniperda was not expected to
have any competition for pine breeding material early in the season (Haack and Lawrence
1995b).

Four predators were also active early in the spring at the same time as T.
piniperda adults were colonizing brood logs. T hanasimus dubius adults were actively
walking on the bolts and preying upon T. piniperda and H. rugipennis pinifex, but were
rarely found in funnel traps early in the spring. The predators Cucujus clavipes and two
species of staphylinids were collected in the funnel traps soon after the start of T.
pim’perda’s flight, but neither were observed on the red pine bolts. Like P. nemorensis
and H. rugipenm’s pinifex, these predators have not previously been observed to be active
so early in the spring. Haack and Lawrence (1995b) speculated that T. piniperda would
not encounter predators or parasitoids until most of its colonization was completed.

Most of the phloem feeders were collected in the funnel traps around the same
time as Ips pini parent adults: Ips grandicollis, Hylurgops rugipennis pinifex,
()rthotomicus caelalus, Gnathotrichus materiarius, Hylastes porculus, Pisasodes

nemorensis, and Dendroctonus valens. With the exceptions of H. rugipennis pinifex and

25

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P. nemorensis, which were active before they were collected in funnel traps, these native
scolytids initiated both flight and colonization from mid- to late- spring of both years, as
was expected. Schenk and Benjamin (1969) reported similar activity periods for several
of these species during their study of 1. pini parent adults.

Most native predators, including T. dubius, P. cylindrica, C. parallelus, and P.
parallelum, were also first collected in funnel traps around the same time as 1. pini. This
activity was earlier than reported by Schenk and Benjamin (1969), who found that the
highest predator populations on I. pini-infested jack pine (Pinus banksiana Lamb.) bolts
occurred during the emergence of the first generation I. pini progeny.

Tomicus piniperda parent adults had an early and short spring flight in all
southern sites in both years. This spring flight behavior has been well documented in
Europe (Haack and Lawrence 1997b, Saarenmaa 1989, Salonen 1973) and in North
America (Haack and Lawrence 1995b), and reflects T. piniperda’s comparatively low
temperature threshold for flight (10 — 12 °C) (Langstrom 1980). Another explanation for
T. piniperda’s early flight is that since it overwinters in crevices in the outer bark of
standing pine trees, it is exposed to warm temperatures and solar radiation earlier than
scolytids that overwinter under the snow layer and duff. Tomicus piniperda was probably
collected first in Lansing because I checked the Lansing funnel traps more frequently
than the traps at the southern sites. The generation time of 100 days was longer than the
estimated 6-10 weeks described in Haack and Lawrence (1995a), and was likely due to
the shaded, cool conditions of the emergence containers and rearing shed in Lansing.
Emergence of progeny adults was characteristically brief, and earlier than most native

scolytids. 1 found no evidence of sister broods, since T. piniperda parent flight and

26

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progeny emergence both occurred in short, discrete peaks of time.

Overwintering I. pini adults initiated spring flight six weeks after T. piniperda
adults in both years of this study. Haack and Lawrence (1997 and 1995b) found that the
first flight of T. piniperda ranged from three to six weeks earlier than the first flight of l.
pini in Michigan from 1993 — 1996. The large funnel trap collections of 1. pini late in the
summer of 1996 paralleled the trends described by Teale and Lanier (1991) and Raffa
(1991) in New York and Wisconsin, respectively: the highest funnel trap collections were
obtained late in the flight season, and relatively few I. pini were collected in the spring
and early summer. This late season peak is hypothesized to occur when I. pini adults stop
colonizing new brood material, and begin a period of high-density feeding attacks, during
which they continue to produce and respond to pheromones (Teale and Lanier 1991).
Raffa (1991) also noted that the attraction of I. pini to ipsdienol seems to be activated by
cool temperatures in the fall. The absence of a late-season peak in funnel trap catches in
1997 may be explained by the cool weather in the spring that delayed the initiation and
development of the three generations. In 1997, degree-day (base 10°C) accumulations by
Julian date were approximately one week behind 1996, and a total of 100 fewer degree-
days were accumulated in 1997 compared with 1996 (Figure 16).

Ips pini progeny adults emerged about 60 days after initial parent flight; longer
than the expected 42 days (Schenk and Benjamin 1969). Although there were three flight
peaks for parent adults corresponding to the flight of the three generations of I. pini.
progeny emergence did not occur in such discrete patterns. The constant yet irregular
emergence of I. pini progeny throughout both summers may be due to the masking effect

of the cool temperatures in the emergence containers, since the containers were not

27

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exposed to sunlight.

Ips grandicollis parent adults were first collected in funnel traps at the same time
as I. pini, not earlier than I. pini as reported in Wisconsin (Schenk and Benjamin 1969).
Based on funnel trap collections, 1. grandicollis appeared to have two generations per
year, but the generation time of 50-60 days is similar to that of the trivoltine 1. pini, so
three generations per year could be possible.

Hylurgops rugipennis pinifex and P. nemorensis were exceptionally early
colonizers of the red pine bolts; they were both observed frequently on the bottom and
cut edges of nearly every bolt in March and April, as the snow was melting. The
preference of these insects to colonize bolt sections that were in direct contact with the
ground, suggests that this cool, moist environment is ideal for these species. Tomicus
piniperda primarily colonized the sides of the bolts, not the underside, so competitive
interactions between T. piniperda and these phloem feeders may be minimized by spatial
partitioning. These were the only pine phloem feeders that were active as early in the
spring as T. piniperda, and they could be important competitors of T. piniperda when
pine breeding material is scarce.

Dendroctonus valens parent adults were first collected in funnel traps in mid-May
of both years, a few weeks later than Haack and Lawrence’s (1995b) first catch in
Michigan in late April in 1993. In a Wisconsin study (Raffa 1991), D. valens had two
peaks of collection in funnel traps: May-June and August-September, but I did not
observe a second flight of D. valens in this study.

The clerid predator T hanasimus dubius was actively preying on T. piniperda and

H. rugipennis pinifex adults for a few weeks in the spring before it was first collected in

28

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the funnel traps. This was not anticipated, since T. piniperda was not expected to have
predation pressure until colonization was complete (Haack and Lawrence 1995b). 1
speculate that T. dubius funnel trap collections occurred after I observed predaceous
activity because T. dubius requires higher temperatures for flight than for walking.
Another possible explanation is the temperatures were too cool for the ipsdienol baits to
volatize sufficiently to attract T. dubius.

The first funnel trap catches of T. dubius occurred in late April 1996, and in early
May 1997, which agrees with the 1993 Michigan study by Haack and Lawrence (1995b).
Thanasimus dubius had several peaks of abundance: before I. pini parent adult
emergence, during 1. pini F1 progeny emergence, and in August, before I. pini F2
progeny emergence, which are all similar to T. dubius funnel trap catches by Raffa
(1991). These collection peaks probably do not represent different generations of T.
dubius, but rather differences in the activity of the adults. Since T. dubius adults prey
upon I. pini adults, it makes sense that T. dubius would be active at the same time as I.
pini adults are becoming active (i.e. overwintering adult emergence, F 1 progeny
emergence). Ips pini adults produce aggregation pheromones as they are colonizing their
hosts, and T. dubius uses these pheromones as kairomones to locate I. pini prey (Hansen
1983, Raffa and Klepzig 1989).

These findings on the biology of T. dubius have many implications for the
proposed introduction of the European clerid T hanasimus formicarius. The primary
reasons for the consideration of T. formicarius were that it is the earliest of the major
European bark beetle predators to become active in the spring (Schroeder 1988), and that

T. dubius was not expected to be active early enough to prey upon T. piniperda. 1 have

29

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clearly demonstrated that T. dubius is indeed active as early in the spring as is T.
piniperda, and that it will prey upon the exotic scolytid. The introduction of T.
formicarius or other biological control agents for the control of T. piniperda should be
postponed until the biology and impact of North American natural enemies is better
understood.

Platysoma cylindrica is another scolytid predator that has commonly been found
in funnel traps baited with 1. pini aggregation pheromones (Raffa and Klepzig 1989,
Raffa 1991), although a field study in Georgia indicated that P. cylindrica is not attracted
to ipsdienol, but is attracted to ipsenol, an aggregation pheromone of 1. grandicollis
(Turnbow and Franklin 1981). This histerid was first collected in funnel traps in late
May of both years, nearly a month later than Haack and Lawrence’s (1995b) 1993
Michigan study. Platysoma cylindrica was primarily collected in funnel traps during two
periods; corresponding to 1. pini overwintering parent adult flight, and I. pini F1 progeny
adult flight, respectively, which is similar to a Wisconsin study (Raffa 1991).

Corticeus parallelus adults followed the same trends as T. dubius and P.
cylindrica: the highest funnel trap catches occurred near peak I. pini parent adult catches
and near peak I. pini F 1 progeny adult catches. This tenebrionid is also suspected to use
I. pini pheromones as kairomones (Raffa and Klepzig 1989, Raffa 1991), so this
synchrony was expected. These tenebrionids were first collected in late May of both
years, about a month later than reported in another Michigan study (Haack and Lawrence
1995b)

Medelera sp. parent adults were not collected in the funnel traps, suggesting that

they are not attracted to OL-pinene or the 1. pini pheromone lures on the funnel traps.

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Fitzgerald and Nagel (1972) found that adults of M. aldrichii, a predator of Dendroctonus
pseudotsugae Hopkins (Coleoptera: Scolytidae), responded to or-pinene as an
ovipositional stimulant, but there are no references to whether any of the Medetera
species have been collected in a-pinene-baited funnel traps. Since I do not have funnel
trap or observational records, I could not determine when parent adults emerge and
oviposit. Studies of M aldrichii Wheeler in Washington and South Dakota (Hopping
1947, Schmid 1970) demonstrated initial parent activity in early June and early July,
respectively. I first collected Medetera sp. progeny adults from late July through
September. Numerous Medetera sp. larvae were collected from the de-barked bolts,
which agrees with several studies of M aldrichii and M. dendrobaena Kowarz that
report that these predators overwinter as larvae (Hopping 1947, Nicolai I995, Schmid
1970)

C ucujus clavipes, along with the staphylinids, were some of the earliest predators
to become active in the spring. The majority of C. clavipes adults were collected in
funnel traps within two weeks of peak T. piniperda parent flight, indicating that this
predator either has a low temperature threshold for flight or that it overwinters in
locations that warm up quickly in the spring. One such possible overwintering location is
under the bark of standing trees, which receives solar radiation and warms up much
earlier in the spring than does the snow-insulated forest litter layer. I did not observe the
overwintering sites of either C. clavipes or the staphylinids.

The funnel trapping method and the observations on the red pine bolts were
phenologically consistent for most of the phloem feeders: Tomicus piniperda, Ips pini,

Ips grandicollis, Dendroctonus valens, Monochamus scutellatus, and M carolinensis.

31

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Since these insects were attracted to the funnel trap baits, were present in large numbers,
and were large enough to be visible, both the funnel traps and the observations were
suitable for monitoring the initial spring activity of these species.

For two phloem feeders and one predator, however, the results of the funnel traps
and the observations were not consistent: H. rugipennis pinifex, P. nemorensis, and T.
dubius. These insects were observed on the red pine bolts several weeks before they
were first collected in the funnel traps, which suggests that walking activity and flight
activity are two separate events. A possible explanation is that insects require more heat
for flight than for walking, so they do not fly until it gets warmer. Another explanation
for the discrepancy between the two methods is that the funnel trap lures were not
volatilizing sufficiently enough in the cool spring weather to be attractive to these
species. For P. nemorensis, H. rugipennis pinifex, and T. dubius, direct observations of
activity on pine slash provide the best means of determining initial spring activity.

Observations were not useful in determining spring activity of other species,
though. Most of the smaller bark beetles and natural enemies were collected in the
pheromone-baited funnel traps, but were too small or too rare to be observed on the red
pine bolts: Orthotomicus caelatus, Ips perroti, Gnathotrichus materiarius, Hylastes
porculus, Dryocetes autographus, Plalysoma cylindrica, Corticeus parallelus, Plalysoma
parallelum, staphylinids, and Cucujus clavipes. In the cases of Medetera sp., Anthocoris
sp., and parasitic wasps, neither observations nor funnel traps were useful as indicators of
spring activity. These insects are not known to be attracted to a-pinene or pheromone-
baited funnel traps, and they were probably too small to be seen during the observations.

Of the three methods utilized to study the phenology of bark beetles and their

32

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natural enemies in this study, the pheromone-baited funnel traps were simple to use and

collected the greatest number of species of pine phloem feeders, as well as eight species

of their natural enemies. The funnel traps employed an effective pheromone/ kairomone
system that was utilized by many bark beetles and natural enemies of varied sizes.

There were several problems associated with funnel trapping, however. Some
bark beetle and natural enemy species were not attracted to a-pinene or the pheromones,
including parasitoids, anthocorid predators, and dolichopodid predators. Because 0L-
pinene, ipsdienol, and lanierone were deployed at the same time in all traps, I cannot
determine which of the baits was most effective for each insect. Many incidental insects
(neither phloem feeders nor natural enemies) were collected by this method — sometimes
up to 50% of each catch was made up of incidental species of beetles, moths, and flies.
Since I used insecticide in the collection cups, funnel trapping was a destructive sampling
method, and could have potentially impacted the population dynamics of the phloem
feeders or their natural enemies in my field sites (see Raffa I991). Funnel traps are
generally considered more of a qualitative than quantitative sampling method, since the
effectiveness of the lures varies depending on the temperature, humidity, wind, and other
factors (Lindgren 1983, Lindgren et al. 1983, Salom and McLean 1991). Finally, though
funnel trapping was an effective means of determining first flight activity for many of
these bark beetles and natural enemies, it did not determine first general activity (i.e.
walking, mating, preying) for two abundant phloem feeders, Hylurgops rugipennis
pini/12x and Pissodes nemorensis, and for an important predator, Thanasimus dubius.

Observing insect activity on red pine bolts was useful for large, conspicuous, and

numerous bark beetles and predators. Observations were an excellent means of

33

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determining the dates of the first walking and mating activity on the red pine bolts, the
first predator activity, and the emergence of the progeny. The observations of insect
activity were also useful when compared to the two other sampling methods — they
provided a real-life check for the funnel trap and rearing results. Observations are an
example of non-destructive sampling, and unlike the funnel trapping method, they
weren’t biased towards species utilizing host volatiles, pheromones, or kairomones.

However, this method underestimated activity of small, cryptic insect species and
did not include activity such as most mating, gallery formation, development, and
predation that occurred under the bark. The observations were also qualitative and
difficult to standardize; activity on the bolts varied greatly depending on the temperature
and weather.

Rearing pine phloem feeders and natural enemies from red pine bolts provided the
best means for studying progeny emergence. The rearing method yielded slightly fewer
numbers of phloem feeding species (twelve) than the funnel traps, but nearly twice as
many natural enemy species (fifteen) were collected. Unlike the other two methods,
rearing was not affected by insect size or utilization of lures. Very few incidental species
were reared from the red pine bolts; nearly everything collected was either a phloem
feeder or a natural enemy of phloem feeders. The rearing method was also the most
quantitative of the three methods, since progeny abundance could be standardized across
surface area measurements. Rearing provided an excellent insight on the number of
generations of phloem feeders and natural enemies.

The rearing method had several disadvantages, though. It was the most time-

consuming of the methods, since hundreds of emergence containers had to be checked

34

several times a week. The red pine bolts were cumbersome to transport, and they
required a large amount of room for storage. Because all of the bolts were subjected to
the same (ambient) temperatures and humidity in the emergence containers, phenological
differences between northern and southern sites were largely obscured. Also, the earliest
collections of each of each species were likely re-emerging parent adults, and these
parent adults confounded the actual date of first emergence of progeny. For the insects
collected during de-barking, I could not determine when or why the insects had died.
For some species, progeny were collected only during bolt dissection, so progeny
emergence phenology could not be determined.
Conclusions

Overall, in the Great Lakes region, most pine bark beetles and their associates
have not been well-studied (Raffa 1991). This paper has served to add phenological
information and number of generations to the limited knowledge accumulated on Great
Lakes pine bark beetles and their associates, but further research on the dynamics of this
insect complex is needed. Funnel traps were effective for determining first flight activity
of parent adults of bark beetles and coleopteran predators in the spring, but observations
were the best means of determining actual activity (i.e. colonization, predation) for H.
rugipennis pinifex, P. nemorensis, and T. dubius. Tomicus piniperda, despite its early
spring flight, will face a number of natural enemies and competitors for pine breeding
material in Michigan. I have demonstrated that two pine phloem feeders and four

predators are active as early in the spring as is T. piniperda.

35

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LITERATURE CITED

APHIS. 1996. Pine shoot beetle biological control project -- Project update. February
1996. Niles, Michigan, USDA-APHIS/PPQ Plant Protection Laboratories.

Byers, J. A. 1991. Simulation of the mate-finding behaviour of pine shoot beetles,
Tomicus piniperda. Anim. Behav. 41: 649-661.

Byers, J. A., B. S. Lanne, J. L6fqvist, F. Schlyter, G. Bergstrom. 1985. Olfactory
recognition of host-tree susceptibility by pine shoot beetles. Naturwissenschaften
72: 324-326.

Coulson, R. N. and J. A. Witter. 1984. Forest entomology: ecology and management.
New York, NY, John Wiley and Sons.

Fitzgerald, T. D. and W. P. Nagel. 1972. Oviposition and larval bark-surface orientation
of Medetera aldrichii (Diptera: Dolichopodidae): responses to a prey-liberated
plant terpene. Annals Entomol. Soc. Amer. 65(2): 328-330.

Haack, R and D. Kucera. 1993. New introduction - common pine shoot beetle, Tomicus
piniperda (L.). USDA Forest Service, Northeastern Area, Pest Alert NA-TP-OS-
93. 2 pp.

Haack, R. A. and R. K. Lawrence. 1995a. Attack densities of Tomicus piniperda and Ips
pini (Coleoptera: Scolytidae) on Scotch pine logs in Michigan in relation to
felling date. J. Entomol. Sci. 30(1): 18-28.

Haack, R. A. and R. K. Lawrence. 1995b. Spring flight of Tomicus piniperda in relation
to native Michigan pine bark beetles and their associated predators. Behavior,
population dynamics, and control of forest insects, Proceedings of the joint
TUFRO conference for Working Parties 8207-05 and 8207-06. Maui, Hawaii, 6-
11 February, 1994. Ohio State University Press, Wooster, Ohio. Pages 524-535.

Haack, R. A. and R. K. Lawrence. 1997a. Tomicus piniperda (Coleoptera: Scolytidae)
reproduction and behavior on Scotch pine Christmas trees taken indoors. Great
Lakes Entomol. 30(1&2): 19-31.

Haack, RA. and R.K. Lawrence. 1997b. Highlights of Forest Service research on
T omicus piniperda: 1992 — 1996. Pages 1 15-122 in 1997 Japanese beetle and the
pine shoot beetle regulatory review: Proceedings, Louisville, Kentucky, 24-26
February 1997. USDA APHIS, Riverdale, Maryland.

Hansen, K. 1983. Reception of bark beetle pheromone in the predaceous clerid beetle,
Thanasimusformicarius (Coleoptera: Cleridae). J. Comp. Physiol. 150: 371-378.

 

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--
lI -
l
‘U
‘-
.H‘.
.‘ I
< —

Herrns, D. A., R. A. Haack, and B.D. Ayres. 1991. Variation in semiochemical-mediated
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37

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38

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Kunming region of China. J. Appl. Entomol. 112: 366-369.

39

 

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Figure 1. Location of red pine forest plantation field sites in Michigan.
KALl, ALLl , WEXl, ROS] , and Lansing were monitored in 1996; all
stands were monitored in 1997.

46

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30 60 90 120 150 180 210 240 270 300

Julian Date

Figure 2. Seasonal distribution of scolytid adults collected in funnel traps baited with alpha-

pinene, ipsdienol, and lanierone at five Michigan red pine plantations in 1996 and nine
plantations in 1997.

47

.3
ax

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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Figure 3. Seasonal distribution of scolytid progeny adults reared from 204 red pine bolts in

1996 and 304 bolts in 1997 in Michigan

48

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ox

 

1996

.9
Ut

 

—O— Tomicus piniperda
- - I - -Ips pini

 

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a

 

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o
w

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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0.1
0.0 -
120 150 180 210 240 270 300 330
Julian Date
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1997 __. _ _, . .
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c
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0.1
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120 150 180 210 240 270 300 330

Julian Date

Figure 3. Seasonal distribution of scolytid progeny adults reared from 204 red pine bolts in

1996 and 304 bolts in 1997 in Michigan.

48

.—o
C

 

 

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30 60 90 120 150 180 210 240 270 300

Julian Date

Figure 4. Seasonal distribution of scolytid adults collected in funnel traps baited with alpha-
pinene, ipsdienol, and lanierone at five Michigan red pine plantations in 1996 and nine
plantations in 1997.

49

.0
ax

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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0 5 1996 - - I - - Orthotomicus caelatus
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Julian Date
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1997 +Hylurgops rugipennis pinifex
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E
‘5' 0.4
E'.‘
9
.o r'
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e l
a.
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a.
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120 150 180 210 240 270 300 330

Julian Date

Figure 5. Seasonal distribution of scolytid progeny adults reared from 204 red pine bolts in
1996 and 304 bolts in 1997 in Michigan.

50

 

 

] 996 + Gnathotrichus materiari us
- - I - -Hylastes porculus

.o f"
o o
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Julian Date

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o
l

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30 60 90 120 150 180 210 240 270 300

Julian Date

Figure 6. Seasonal distribution of scolytid adults collected in funnel traps baited with alpha-
Pinene, ipsdienol, and lanierone at five Michigan red pine plantations in 1996 and nine
Plantations in 1997.

51

.9
ON

 

 

 

 

—O— Gnathom'ch us materiari us

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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9 —O— Gnathotrichus
1997 :1 materiarius
0-5 f -. - - I - -Hy1astes porculus
:' : '
g 0 4 ' l '9‘. - O - Dryocetes H
: ° ll autographus
: .' 1 l I
o 3 z ! t
3': .- : ' I
e : - l I
3' . ' l I
: I ‘ ' A I
_' I i
I l
l

 

120

150

210

240

Julian Date

270

 

 

300

 

330

Figure 7. Seasonal distribution of scolytid progeny adults reared from 204 red pine bolts in

1996 and 304 bolts in 1997 in Michigan.

52

 

 

1996 +Pissodes nemorensis

 

Ll

- - I - ~Monochamus scutellatus
0. 8 . .
and M. carolmenszs

 

 

 

 

 

 

 

a 0.7 ' - O - Dendroctonus valens _
a l
._ 0.6 9
o l I
g 0.5 1
I

 

nun—III-—

opo
o

A

' — ‘-

 

 

 

HQ

 

 

02 t ll
.:. I\ . r‘h
0.0 -—I"IHMW—

3O 60 90 120 150 180 210 240 270 300

 

 

 

 

 

 

Julian Date
1.0
0 9 1997 —O-Pissodes nemorensis
' - - I - -Monochamus scutellatus and M. carolinensis
0.3 — O - Dendroctonus valens

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

30 6O 90 120 150 180 210 240 270 300

Julian Date

Figure 8. Seasonal distribution of adults collected in funnel traps baited with alpha-pinene,
ipsdienol, and lanierone at five Michigan red pine plantations in 1996 and nine in 1997.

53

.9
ca

 

 

-O—Pissodes nemorensis
1996

9
Vi

 

- - I - -M0nochamus scutellatus and M carolinensis

 

 

- O - Dendroctonus valens

 

 

.9
.h.

 

 

 

Proportion of Total
C:
U.)

 

 

 

 

 

 

0.2
0.1
0.0 W
120 150 180 210 240 270 300 330
Julian Date
0.6
1997 —O—Pissodes nemorensis

O 5 - - I - -M0n0chamus scutellatus and M. carolinensis

' - O - Dendroctonus valens

 

 

 

.9
a.

 

 

>

 

Proportion of Total
O
t»

.i, / \y/l
A ,

120 150 180 210 240 270 300 330

 

 

 

 

Julian Date

Figure 9. Seasonal distribution of progeny adults reared from 204 red pine bolts in 1996 and
304 bolts in 1997 in Michigan.

54

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

   

 

 

 

 

 

1.0
0 9 1996 + Thanasimus dubius __4
0 8 ~ - I - -P1alysoma cylindrica
' - O - Corticeus parallelus
B 0.7 Q
g ."
u- 0 6 ::
° -'.
2 0.5 § :
o i 1
gfl4 if
5 0.3 5' ‘2‘
L l.
0.1 i
0.0- - A-,
30 60 90 120 150 180 210 240 270 300
Julian Date
1.0
O 9 1997 + Thanasimus dubius a
' - - I - -Platysoma cylindrica
- C - Corticeus parallelus

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

180 210 240 270 300

30 60 90 120 150
Julian Date

Figure 10. Seasonal distribution of parent adults collected in funnel traps baited with alpha-
pinene, ipsdienol, and lanierone at five Michigan red pine plantations in 1996 and nine

Plantations in 1997.

55

 

F’
ox

 

 

1996 —O— Thanasimus dubius

- - I - -Plarysoma cylindrica .—

F’
u.

 

 

— O - Corticeus parallelus

 

.9
4;

 

 

 

Proportion of Total
o
w

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

0.2
0.1
0.0 - ‘
120 150 180 210 240 270 300 330
Julian Date
0.6
+ Thanasmzus dubius I
1997 . . z
0. 5 - - I - -Platysoma cylmdnca 3
- O - Corticeus parallelus .‘1
g 0.4 r A
c.- 1 .' 1
° .' 1
.5 0 3 : z
E : :
G a .
a. . .
2 0.2 ' ‘ :
a. \
0.1
00 ‘ ’ r
120 150 180 210 240 270 300 330

Julian Date

Figure 11. Seasonal distribution of predator progeny adults reared from 204 red pine bolts in
1996 and 304 bolts in 1997 in Michigan.

56

 

 

1 996 + Staph ylinidae
- - I - -Medetera sp.

 

 

- O - Platysoma parallelum

 

 

 

 

 

 

 

 

 

 

 

 

 

30 60 90 120 150 180 210 240 270 300

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Julian Date
1.0
0 9 1997 +Staphy1inidae __
0 8 - - I - -Medetera sp.
' — O - Platysoma parallelum
E 0.7
fl
.._ 0 6
G
‘7' O 5
o .
20.4 3.
a. l
e 1‘
Out 0 3 A i I
l
0 2 I 1
I i. 'm‘ a

 

 

 

30 60 90 120 150 180 210 240 270 300

Julian Date

Figure 12. Seasonal distribution of predator adults collected in funnel traps baited with alpha
pinene, ipsdienol, and lanierone at five Michigan red pine plantations in 1996 and nine
plantations in 1997.

57

 

 

.9
a

1996

 

 

.C
Ln

 

—O— Staphylinidae
- - I - -Medetera sp.

— O - Platysoma parallelum

 

 

 

.0
A

 

 

Proportion of Total
9 o
N w

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

120 150 180 210 240 270 300 330
Julian Date
0.6
1997 +Staphylinidae
0.5 - - I - -Medetera sp. *—
— O - Platysoma parallelum
3 O 4 —
a - -
a. .1
O a:
,5 0 3 3'.
2
g- .' '. ,I‘
a 0.2 : 1 iv 2.
0.1 5 K i. i K'-
.' 1; ' .l
0.0 4 ' ~ ' ‘ .
120 150 180 210 240 270 300 330

Julian Date

Figure 13. Seasonal distribution of predator progeny adults reared from 204 red pine bolts in
1996 and 304 bolts in 1997 in Michigan.

58

+ Cucujus clavipes

- - I - -Enoclerus nigripes
+Anthoc0ris sp.

- A - Parasitic wasps

 

3O 60 90 120 150 180 210 240 270 300

Julian Date

1.0
0.9

—O— Cucujus clavipes

- - I - -En0clerus nigripes
0.8 - O — Anthocoris sp.

7: 0.7 - i - Parasitic wasps

: 0.6

O

.5 0.5

O 0.4

E

a. 0.3
0.2
0.1

0.0

 

30 6O 90 120 150 180 210 240 270 300

Julian Date

Figure 14. Seasonal distribution of predator and parasitoid adults collected in funnel traps
baited with alpha-pinene, ipsdienol, and lanierone at five Michigan red pine plantations in
1996 and nine plantations in 1997.

59

.9
ox

 

 

 

 

 

 

 

 

 

 

 

 

 

—O— Cucujus clavipes
1996 - - I - -Enoclerus nigripes
0.5 - O - Anthocoris sp. I,
a - i - Parasitic wasps
*' o 4 "
[S 4 : '.
“a I ' ' '-
2: ? -' a : 1 i
Q 0 3 'l ' ' u 1
° ' I ' '- " rt ' '- ' l
a. ' '
o I | I; u [I I : l i
i. 0.2 , - f -, w . 1 +
a: l ' 1 ' .1 ' . I
l ‘ : ‘ " .0 I. . l ‘
l ,- '. : : .1 : - I
‘ Al .- : ‘4 : ' ' l

Julian Date

 

 

 

 

 

 

 

 

 

 

 

 

0.6
+Cucujus clavipes
1997 - - I - -En0clerus nigripes
()5 1—fi I
f. - O - Anthocoris sp.
'5 - i - Parasitic wasps
[:5 0.4 : . ,
"5 '. u
'3 o 3 Q : - n
'E H : '. H
2. ' ' : '- 9
i l O 1 '01‘
E. 0'2 l 1 f . [in
o I I ‘
, l l . . . .l
01 ’ {It 'Q‘:,A.'. H
a J =. t I # a
‘4 I ‘ I ‘ i
,x 1“” \ Ar i‘ {at} *i
o 0 W T

 

120 150 180 210 240 270 300 330

Julian Date

Figure 15. Seasonal distribution of predator and parasitoid progeny adults reared from 204
red pine bolts in 1996 and 304 bolts in 1997 in Michigan.

60

 

1400

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

1996 —0— Southern Sites /
1200 ' ' .- ' 'Northem Sites /
S’ 1000 "
S
i
m 800
9.
g 600
if
a 400
200
0
50 100 150 200 250 300
Julian Date
1400
1997 + Southern Sites
1200 - - O- - -Northem Sites
CU 1000
S.
i
m 800
9.
g 600
l-
3‘ 400
200
O I
50 100 150 200 250 300

Julian Date

Figure 16. Average degree-day (base 10°C) accumulations from 1 March in the
Southern sites (KAL and ALL) and northern sites (WEX and ROS), by Julian date.

61

CHAPTER 2

Impacts of Natural Enemies on Native Ips spp. and Introduced T omicus piniperda

(Coleoptera: Scolytidae) Bark Beetles in Red Pine Plantations in Michigan

INTRODUCTION
The subcortical life stages of bark beetles (Coleoptera: Scolytidae) are attacked by

many species of insect predators and parasitoids. Several studies in North America have
evaluated the importance of predation and parasitism in pine bark beetle populations in
the western (Dendroctonus ponderosae Hopkins) and southern US. (Dendroctonus
frontalis Zimmermann) (Dahlsten and Stephen 1974, Smith and Goyer 1980), but there
have been few quantitative studies on the influence of natural enemies on the pine bark
beetles of the Great Lakes region. Bright (1996) and Reid (1957) listed the predators and
parasitoids of T omicus piniperda (L.) and Ips pini (Say), respectively, but did not

examine their influence on the population dynamics of these bark beetles.

The recent establishment of an exotic pine bark beetle, T omicus piniperda, in the
Great Lakes region could potentially affect interactions between natural enemies and
endemic bark beetle species. The larger pine shoot beetle, T. piniperda, a well-known
Pest of pines throughout much of Europe, Asia, and North Africa (Langstrom 1980, Ye
I 991), was first discovered in 1992 in a Scotch pine (Pinus sylvestris L.) Christmas tree
Plantation in Ohio (Haack and Kucera 1993). As of September 1998, T. piniperda had

been found in 243 counties in nine states, and in 21 counties in the Canadian province of

Ontario (NAPIS 1998).

62

 

 

Although T. piniperda and several native bark beetles feed subcortically in pines,
there are important differences in their life cycles that could affect interactions with
natural enemies. For example, parent adults of the native scolytid Ips pini colonize
phloem of stressed, dying, or recently-cut pine trees, and there can be three generations of
I. pini per year in Michigan (Schenk and Benjamin 1969). Tomicus piniperda adults also
colonize recently dead or dying pines, but T. piniperda is univoltine in Michigan (Haack
and Lawrence 1995a). Teneral T. piniperda adults require maturation feeding, and spend
most of the summer feeding in shoots of live pines. Both 1. pini and T. piniperda
overwinter as adults, but T. piniperda has a lower temperature threshold for spring flight
(10-12°C) and can become active in the spring four to six weeks earlier than I. pini and
many other native scolytids and natural enemies (Haack and Lawrence 1995b). These
differences in the number of generations and the date of peak activity presumably
influence the quantity and species diversity of the natural enemies encountered by these

two scolytids.

Little is known about the impacts of predators and parasitoids on 1. pini and other
native scolytids in the Lake States region, and even less is known about how these natural
enemies will interact with T. piniperda in North America Previous studies of I. pini in
the Great Lakes region list its most abundant predators as Thanasimus dubius (F.)
(Coleoptera: Cleridae), Corticeus sp. (Coleoptera: Tenebrionidae), Histeridae
(COIeoptera), and Staphylinidae (Coleoptera), but do not present actual mortality rates of
1‘ Pihi due to these predators (Schenk and Benjamin 1969, Raffa and Klepzig 1989). A
Study in Ontario listed predators and parasitoids that emerged from a single Scotch pine

tree infested by T. piniperda, but no mortality rates were presented (Bright 1996).

63

Information on interactions of T. piniperda and native scolytids with their natural

enemies is needed to analyze scolytid population dynamics, predict population trends,

and develop management guidelines for endemic and epidemic bark beetle p0pulations

(Riley and Goyer 1986). Knowledge of natural enemy biology is also needed to develop

recommendations to conserve or enhance populations of natural enemies (Dahsten 1982).

Red pine, Pinus resinosa Ait., is one of the most extensively planted species for

pulp, sawtimber, and poles in the Great Lakes region (Rudolf 1990), comprising over

364,000 hectares of Michigan’s timberland (Leatherberry and Spencer 1996). Red pine is

a suitable host for T. piniperda as well as Ips spp. and many other pine bark beetles

(Lawrence and Haack 1995, Raffa 1991 ). During a typical rotation of red pine, multiple

thinnings or partial harvests are performed, each of which generates an abundance of

brood material for scolytid colonization. To date, most of the research conducted on T.
piniperda in North America has taken place in Scotch pine Christmas tree plantations
(Haack and Lawrence 1997a, Kauffman et a1. 1998, McCullough and Smitley 1995,
McCullough et a1. 1998, McCullough and Sadof 1998). I chose to conduct this study in
pole-sized red pine plantations because they represent a more stable ecosystem with no
insecticide use, and, presumably, a more diverse complex of natural enemies.

I examined interactions among T. piniperda, Ips spp., and natural enemies in red

Pine forest stands in Michigan in 1996 and 1997. My objectives were to (1) identify and
qUantify natural enemies that were commonly associated with T. piniperda and Ips spp.
in red pine plantations; (2) determine if these natural enemies affected bark beetle

reproduction or survival; and (3) determine whether the presence of T. piniperda altered

the impact of natural enemies on Ips 8131)-

64

MATERIALS AND METHODS

Field Sit“

In 1996, I conducted field studies in four red pine forest plantations in Michigan.
Two plantations were in southwestern Michigan and were known to be infested with both
T. piniperda and native Ips spp. These stands were located in Michigan State
University’s WK. Kellogg Experimental Forest in Kalamazoo County (KALI), and in
the Allegan State Game Area in Allegan County (ALLl) (Figure 1). Two red pine
plantations in northern lower Michigan, where T. piniperda populations were not present
(D. McCullough, unpublished data), were also selected. One stand was in the Huron-
Manistee National Forest in Wexford County (WEXI), and the other was in the AuSable
State Forest in Roscommon County (ROS 1). Although T. piniperda was later detected in
Wexford County in 1997 during regulatory surveys (NAPIS 1998), I did not observe any
T. piniperda life stages or evidence of infestation in the northern field sites in 1996 or in
1997.

In general, all four research sites shared the following characteristics: they were
Pole-sized red pine plantations of similar size, age, and basal area (Table l), with some
hardwood understory; other red pine plantations were located within 2 km of the sites;
and they were thinned during the 1994-1995 winter. An abundance of slash (for
example, trunks, tops, large branches), suitable for scolytid colonization, was left on the
ground in each stand from the thinnings, and was expected to attract scolytids in 1995.
Temperature, precipitation, and degree day accumulation (base 10°C) from March lwere

mOnitored by Michigan State University weather stations near all four sites.

65

In 1997, I used the same four sites described above, which represented conditions
two years after thinning (“old” sites). I also expanded the study to include four additional
red pine plantations (KAL2, ALL2, WEX2, and R082), each one within 6.2 km of one of
the original stands. These “new” sites had the same general characteristics of the original

stands (Table l), but were row-thinned during the 1995-1996 winter.

Funnel Trapping

To monitor the flight activity of scolytids and their associates, two to four 12-unit
Lindgren funnel traps (Lindgren 1983) (PheroTech, Inc., Delta, British Columbia) were
placed in each stand. The traps were hung from metal poles to keep the trap bottoms
approximately 1 m above the ground. Each trap was erected and baited in February of
1996 and 1997 with two a-pinene lures to attract T. piniperda (Byers 1991, Haack and
Lawrence 1997b, Schroeder 1988) and with one lure each of ipsdienol and lanierone.
Ipsdienol and lanierone are aggregation pheromones of I. pini that are used as kairomones
by predaceous clerid, tenebrionid, and histerid beetles (Herms et al. 1991, Raffa 1991,
Teale et a1. 1991). Vapona insecticide strips were used in each trap, and insects were

collected from traps at weekly intervals from March to September in 1996 and 1997.

Exclusion Study

In 1996, a total of 204 red pine bolts, and in 1997, a total of 304 bolts, each
approximately 61 cm long and 10 — 20 cm diameter, were cut from felled red pine trees at
each field site in February, April, and June of both years (Table 1). These times roughly
corresponded to the initial activity of overwintering parent T. piniperda, overwintering
parent Ips spp., and emergence of the F1 progeny Ips spp. adults. These times

represented three batches, or groups, and the bolts in each batch were divided evenly

66

among all field sites (Table l), with the exception of batch 1. Batch 1 bolts were only
placed in the southern sites since T. piniperda was not believed to be present in the
northern sites. Bolts were placed on the forest floor at least 50 m away from the funnel
traps, in a shady area of each plantation to reduce desiccation, just before the expected
flight of T. piniperda or Ips spp. (Table 1). Two weeks after the bark beetles’ peak flight
(as determined by funnel trap collections), half of the bolts in each batch were moved to
the floor of a nearby screened cage (1.8 x 3.7 x 1.8 m) constructed of 1 mm Lumite®
amber screen (Synthetic Industries, Inc., Gainsville, GA) to exclude natural enemies.
The other half of the bolts were left on the forest floor near the exclosure and remained
exposed to natural enemies.

Beginning two weeks after the bark beetles’ peak flight, two or three caged bolts
and two or three exposed bolts from each batch were returned to Michigan State
University. Bolts were retrieved at two week intervals in 1996 and at one week intervals
in 1997. The cut ends of the retrieved bolts were dipped in paraffin wax to reduce
desiccation. Each bolt was placed in an individual emergence container consisting of a
cardboard tube (15 - 25 cm diameter, 0.3 -- 0.6 cm wall, 61 — 71 cm overall length
Michigan Can & Tube, Inc., Saginaw, MI]) with opaque plastic endcaps and a clear
plastic collection cup at one end. The emergence containers were stored on the Michigan

State University campus in a screened insectary to expose the bolts to ambient

temperatures and humidity.

Adult bark beetles and natural enemies were collected from the red pine bolts in
the emergence containers at least twice a week until late October of both years. When

progeny adult emergence stopped in the winter, I counted the number of scolytid exit

67

holes per bolt, and then each bolt was debarked and any insects overwintering in the bark
or dead on the bottom of the emergence container were collected. All insects that were
reared from these bolts were assumed to be progeny adults. Although it is possible that a
few parent adults were inadvertently collected, the number of parents was assumed to be
small and similar across all bolts and sites.

For each debarked bolt, 1 determined the phloem surface area, calculated as a
cylinder using the mean of two length measurements and the mean of four diameter
measurements. I also measured the number and length of Ips spp. galleries (each egg
gallery radiating from the nuptial chamber was counted), and the number and length of T.
piniperda galleries for each bolt. T omicus piniperda is monogamous, and its galleries are
long and straight and parallel to the grain of the wood; Ips spp. are polygamous, and their
galleries consist of a nuptial chamber with 2 — 4 radiating galleries, usually in the shape
of a “Y” or “H.” The number of scolytid egg galleries per bolt was used as an indicator
of the density of female parent beetles (1 egg gallery = 1 female parent), and the number

of scolytid progeny per gallery was used as a measure of productivity.

Data Analysis

For each bolt, I calculated the number of scolytid and natural enemy progeny

reared per m2 phloem surface area (progeny density), the number of scolytid progeny
r cared per gallery (productivity), the number of scolytid exit holes per m2, the number of
T- Piniperda and Ips spp. galleries per m2 (attack density), the total length of T. piniperda

and Ips spp. galleries per bolt, and the average length of T. piniperda and Ips spp.

galleries per bolt.

68

If fewer than five beetles of a particular scolytid species were reared from a bolt,
it was considered a failed attack, and that species was excluded from that bolt’s data set.
Because the caging treatment did not effectively exclude all natural enemies, I instead
evaluated the effect of the presence/absence of natural enemies on the number of scolytid
offspring reared per m2 of bolt surface area. A bolt with three or more natural enemy
progeny was considered a successful attack (e.g. “with natural enemies”), and a bolt with
two or fewer natural enemy progeny was considered a failed attack (e. g. “without natural
enemies”). None of the natural enemy progeny data was excluded from the data set.

All variables were tested for normality with the Shapiro-Wilk W test, and were
log transformed to increase homogeneity of variances and tested again for normality.
None of the variables were normally distributed, even after log transformation, so the
nonparametric Mann-Whitney test was used (SAS Institute 1996). Each batch of bolts
was analyzed separately with the Mann-Whitney test to analyze the effects of exposure to
natural enemies (0 — 2 natural enemies per bolt or 3+ natural enemies per bolt) on each of
the variables. In addition, simple linear regression analysis was performed on the

productivity data to evaluate the association between the actual natural enemy density

and scolytid productivity.

69

RESULTS

In both 1996 and 1997, the mean phloem surface area per bolt did not differ

significantly between bolts with natural enemies and bolts without natural enemies

(Tables 9 and 10).

Scolytid Progeny Density — 1996

A total of 2294 scolytids were reared from 46 of the 60 bolts of batch 96-1,
including 569 Tomicus piniperda and 404 Ips spp. (I. pini and I. grandicollis). From
batch 96-2, 72 of 88 bolts were successfully colonized by a total of 3232 scolytids, with
Ips spp. comprising nearly 70% of the total. From batch 96-3, 1292 bark beetles were
reared from 38 of the 56 bolts, and over half of the bark beetles were either I. pini or 1.
grandicollis. The mean number of I. pini progeny per m2 phloem surface area was
highest in batch 96-3 (182 progeny/m2) and lowest in batch 96-2 (130 progeny/m2)
(Table 2). Ips grandicollis densities were highest in batch 96-2 (156 progeny/m2) (Table
2).

In batch 96-1, the mean number of T. piniperda progeny reared per m2 phloem
surface area was higher from bolts that also reared three or more natural enemies
(referred to as “with natural enemies” from this point on) compared with bolts with two
or fewer natural enemies (“without natural enemies” from this point on), but the
difference was not significant at the 0.05 level (Table 3). Ips pini and I. grandicollis
densities were higher from bolts without natural enemies, but the difference was not

Significant for I. pini, and there were not enough samples to test I. grandicollis (Table 3).

7O

 

In batch 96-2, no T. piniperda progeny were reared. The mean number of 1. pini
reared per m2 surface area was not significantly higher from bolts without natural
enemies compared with bolts with natural enemies (Table 3). Ips grandicollis density
was higher in bolts with natural enemies, but the difference was not significant (Table 3).

In batch 96-3, the mean number of I. pini reared per m2 surface area was higher in
bolts with natural enemies (Table 3). The results were the opposite for 1. grandicollis:
the density was higher in bolts without natural enemies (Table 3). In both cases,
however, there were not enough colonized logs to test for significance.

Scolytid Progeny Density - 1997

From batch 97-1, a total of 7518 bark beetles were reared fi'om 89 of the 112
bolts, including 3297 T. piniperda and 2656 Ips spp. (I. pini and I. grandicollis). A total
of 7549 scolytids were reared from 119 of the 128 bolts of batch 97-2, including 6062 Ips

spp. From batch 97-3, only 3140 bark beetles were reared from 52 of the 64 bolts, with I.

pini and 1. grandicollis comprising over 92% of this total. Mean T. piniperda density was
highest in batch 97-1 (224.6 progeny/m2) (Table 4). Mean 1. pini density was highest in
batch 97-3 (385.7 progeny/m2), and I. grandicollis mean density was highest in batch 97-
1 (1 38.7 progeny/m2) (Table 4).

In batch 97-1, the mean number of T. piniperda and 1. pini progeny reared per m2

Phloem surface area were both higher in bolts with natural enemies, but the differences
Were not significant (Table 5). Ips grandicollis followed the opposite trend: mean

2

Progeny per m was higher in bolts without natural enemies, though not significantly

(Table 5).

71

 

The results in batches 97-2 and 97-3 were similar. T. piniperda was reared from
only three bolts in batch 97-2, and all three also had natural enemies (Table 5). In both
batches, the mean number of I. pini progeny reared per m2 phloem surface area was
higher, but not significantly, from bolts with natural enemies (Table 5). And in both

batches, mean I. grandicollis progeny per m2 was higher in bolts without natural enemies,

though not significantly (Table 5).

Scolytid Productivity — 1996

Scolytid productivity (mean number of scolytid offspring per gallery) varied
widely between batches. Tomicus piniperda mean productivity in batch 96-1 was 8.7
progeny/gallery. Ips pini mean productivity was highest in batch 96-1 (28.5
progeny/gallery), and much lower in batch 96-2 (5.5 progeny/gallery) and batch 96-3 (6.7
progeny/gallery). The trend was similar for 1. grandicollis: mean productivity was
highest in batch 96-1 (21.0 progeny/gallery) and much lower in batch 96-2 (11.9
progeny/gallery) and batch 96-3 (9.4 progeny/gallery).

In batch 96-1, the mean number of T. piniperda progeny per gallery was higher in
bolts without natural enemies (Table 6), and there was a weak negative relationship
between natural enemy density and T. piniperda productivity (Figure 2a). For I. pini, the
mean progeny per gallery was more than eight times higher in bolts without natural
enemies, but the difference was not significant (Table 6). There was a negative but
“0118i gnificant relationship between natural enemy density and I. pini productivity
(FigUre 2b). Ips grandicollis productivity was also higher in bolts without natural

eneirrries, but there were not enough samples to analyze the data (Table 6).

72

No T. piniperda progeny were reared in batch 96-2. The mean number of 1. pini
progeny per gallery was again higher in bolts without natural enemies, but the difference
was marginally insignificant (p = 0.0576) (Table 6). There was a negative but
insignificant relationship between natural enemy density and I. pini productivity (Figure
3a). Mean 1. grandicollis progeny per gallery was higher in bolts without natural
enemies, but again, the difference was not significant (Table 6). The association between
natural enemy density and 1. grandicollis productivity was weakly positive, though not
significant (Figure 3b).

In batch 96-3, no T. piniperda were reared. Mean number of I. pini offspring per
gallery differed little between bolts with and without natural enemies, but there was a
significant positive relationship between natural enemy density and I. pini productivity (p
= 0.0124) (Figure 4a). This relationship may have been strongly influenced by one
outlying point, though (Figure 4a). The mean number of I. grandicollis progeny per
gallery was higher in bolts without natural enemies, but there was only one bolt with
three or more natural enemies present (Table 6). The relationship between natural enemy
density and l. grandicollis productivity was nonsignificant and negative (Figure 4b).
Scolytid Productivity - 1997
Scolytid productivity did not vary as widely between batches in 1997 as it did in
1996. Mean 7’. piniperda productivity in batch 97-1 was 10.5 progeny/gallery. For I.
pini, mean productivity was again highest in the first batch (6.4 progeny/gallery), and
lOWer in the later batches (97-2 mean: 3.4 progeny/gallery; 97-3 mean: 4.7

Progeny/gallery). Ips grandicollis mean productivity was highest in batch 97-1 at 9.9

73

 

progeny/gallery, and lower in batch 97-2 (4.8 progeny/gallery) and batch 97-3 (5.3
progeny/gallery).

In batch 97-1, the mean number of T. piniperda, I. pini, and l. grandicollis
progeny per gallery was higher in bolts without natural enemies for each species, but the
differences were not significant (Table 7). There was a significant negative relationship
between natural enemy density and T. piniperda productivity (p = 0.0338) (Figure 5a).
There was a weakly negative relationship between natural enemy density and I. pini
productivity, but it was not significant (Figure 5b). For I. grandicollis, the relationship

between natural enemy density and productivity was negative and significant (p =

0.0421) (Figure 5c).

Tomicus piniperda was only reared from three bolts in batch 97-2, and all three
bolts also had natural enemies (Table 7). Mean 1. pini progeny per gallery was slightly
higher in bolts with natural enemies (Table 7), and the relationship between natural
enemy density and I. pini productivity was weakly negative (Figure 6a), but neither was
significant. The mean number of I. grandicollis progeny per gallery was higher in bolts
without natural enemies (Table 7), and there was a nonsignificant, negative relationship
between natural enemy density and I. grandicollis productivity (Figure 6b).

Tomicus piniperda was not reared from batch 97-3. The mean number of progeny
per gallery for I. pini was higher in bolts without natural enemies, but not significantly so
(Table 7). The relationship between natural enemy density and 1. pini productivity was
negative, but nonsignificant (Figure 7a). For I. grandicollis, the productivity per gallery

Was significantly (p = 0.0372) higher in bolts without natural enemies (Table 7). There

74

 

was a negative but nonsignificant relationship between natural enemy density and I.

grandicollis density (Figure 7b).
Scolytid Egg Galleries and Exit Holes — I996

The mean number of exit holes/m2 phloem surface area was highest in batch 96-2,
and lowest in batch 96-3 (Table 8). The mean number of Ips spp. galleries per m2 and the
total length of Ips spp. galleries per bolt were both highest in batches 96-2 and 96-3
(Table 8).

In batch 96-1, the mean number of scolytid exit holes per m2 of phloem surface
area, the total length and average length of T. piniperda galleries, and the average length
of Ips spp. galleries were all higher in bolts with natural enemies, but the differences
were not significant (Table 9). The mean number of T. piniperda galleries per unit area
was also higher in bolts with natural enemies, and the difference was marginally
significant (p = 0.0539) (Table 9). The mean number of Ips spp. galleries per unit area
and the total length of Ips spp. galleries per bolts were both significantly higher in bolts

with natural enemies (p = 0.0266 and p = 0.0365, respectively) (Table 9).

In batch 96-2, the trends were similar. The number of exit holes per m2 of phloem
surface area, the number of Ips spp. galleries per unit area, the total length of Ips spp.
galleries per bolt, and the average length of Ips spp. galleries were all higher in bolts with

natural enemies, but the differences were not significant (Table 9).

In batch 96-3, the number of exit holes per m2 of phloem surface area and the
aVfiil‘age length of Ips spp. galleries were both higher in bolts without natural enemies.

thOugh not significantly (Table 9). The number of Ips spp. galleries per m2 and the total

75

length of Ips spp. galleries per bolt were both higher in bolts with natural enemies, but the

difference was not significant (Table 9).
Scolytid Egg Galleries and Exit Holes — 1997
The mean number of exit holes/m2 was much higher in batch 97-2 than in the

other two batches (Table 8). In batch 97-1, the mean number of T. piniperda galleries per
m2 and total length of T. piniperda galleries per bolt were similar to the results in batch
96-1 (Table 8). The average length of T. piniperda galleries per bolt, however, was much

lower in batch 97-1 than in batch 96-1 (Table 8). The mean number, total length, and

average length of Ips spp. galleries were all highest in batch 97-2, and lowest in batch 97-

1 (Table 8).

In batch 97-1, the mean number of scolytid exit holes per m2, the mean number of

Ips spp. galleries per m2, the total lengths of T. piniperda and Ips spp. galleries per bolt,

and the average lengths of T. piniperda and Ips spp. galleries were all significantly (p <

0.0001) higher in bolts with natural enemies (Table 10). The average number of T.
piniperda galleries per m2 was also significantly (p = 0.0003) higher in bolts with natural
enemies (Table 10).

In batch 97-2, the number of exit holes per m2 of phloem surface area was slightly

higher in bolts without natural enemies, but the difference was not significant (Table 10).
The mean number of Ips spp. galleries per m2, the mean total length of Ips spp. galleries
Per bolt, and the average length of Ips spp. galleries were all significantly higher in bolts
with natural enemies (p = 0.0052, p = 0.0034, and p = 0.0252, respectively) (Table 10).

In batch 97-3, all of the variables were higher in bolts with natural enemies, but none of

the differences were significant (Table 10).

76

Natural Enemies — 1996

The mean density of all natural enemy species combined was highest in batch 96-
2, and lowest in batch 96-3 (Table 2). For Corticeus parallelus Melsheimer (Coleoptera:
Tenebrionidae), Medetera sp. (Diptera: Dolichipodidae), parasitic wasps (Hymenoptera),
and staphylinid beetles (Coleoptera: Staphylinidae), the mean densities of these species

were highest in batch 96-2, and lowest in batch 96-3 (Table 2). Platysoma cylindrica
(Paykull) (Coleoptera: Histeridae) density was highest in batch 96-1 and lowest in batch
96-3 (Table 2). Thanasimus dubius density was highest in batch 96-2 and lowest in batch
96-1 (Table 2).

In batch 96-1, a total of 96 arthropod natural enemies (94% predators, 6%
parasitoids) were reared from 29 of the 60 bolts (Table l 1). The most common predators
reared from T. piniperda-infested bolts in batch 96-1 were Thanasimus dubius, Medetera
sp, and C. parallelus (Table 12). These three predatory species, as well as P. cylindrica
were the most common natural enemies in Ips spp. infested bolts (Table 12). Only one
bolt from batch 96-1 was infested with both T. piniperda and Ips spp., and no natural
enemies were reared from it. More predator species and slightly higher numbers of
Predators were reared from bolts infested with Ips spp. alone as compared with bolts with
T Piniperda alone (Table 12).

A total of 458 arthropod natural enemies (90% predators, 10% parasitoids) were
rtflared from 70 of the 88 bolts of batch 96-2 (Table 11). The most abundant natural
enemies from this batch were T. dubius, Medetera sp., and C. parallelus. Corticeus
Parallelus and staphylinids were more common in the southern sites, but T. dubius was

more abundant in bolts from the northern sites (Table l 1).

77

For batch 96-3, I collected only 101 arthropod natural enemies, 85 of which were
predators, from 40 of the 56 bolts (Table l 1). Medetera sp., T. dubius, parasitic wasps,
and C. parallelus were the most abundant natural enemies (Table 11). Once again, C.

parallelus was more abundant in the southern sites, and T. dubius and Medetera sp. were
more common in the northern sites (Table 11).
Natural Enemies - 1997

The mean density of all natural enemy species combined was highest in batch 97-
2, and lowest in batch 97-3 (Table 4). Anihocoris sp. (Hemiptera: Anthocoridae) mean
density was highest in batch 97-3, and C. parallelus and parasitic wasp mean densities
were highest in batch 97-2 (Table 4). Medetera sp. and T. dubius densities were highest
in batch 97-1 (Table 4). The mean densities of P. cylindrica and staphylinid beetles were
similar across all batches (Table 4).

In batch 97-1, I collected a total of 665 arthropod natural enemies (97% predators.
3% parasitoids) from 68 of the 112 bolts (Table 13). Overall, natural enemies were twice
as abundant from the “new” sites than the “old” sites (Table 13). The most common
predators reared from T. piniperda-infested bolts were T. dubius, P. cylindrica, and C.
parallelus (Table 14). From Ips spp.-infested bolts and from bolts infested with both T.
piniperda and Ips spp., I found four more predator species in addition to the three listed
above: two species of Staphylinidae (Coleoptera), Medetera sp., and Anthocoris sp.
(Hemiptera: Anthocoridae) (Table 14). Predators were more common in bolts infested

with Ips spp. alone or with both Ips spp. and T. piniperda than in bolts with T. piniperda
alone (Table 14).

78

ln batch 97-2, I collected nearly twice as many arthropod natural enemies, 1140,
from 116 of the 128 bolts (Table 13). Parasitoids comprised 10% of the natural enemies,
and predators made up the other 90%. Again, natural enemies were more abundant from
new sites than old sites, with the difference most pronounced in the northern sites (Table
13 ). Overall, the most common natural enemies were T. dubius, P. cylindrica, Medetera

sp., C. parallelus, and parasitic wasps (Table 13).

In batch 97-3, only 185 arthropod natural enemies (80% predators, 20%
parasitoids) were reared from 46 of the 64 bolts (Table 13). Unlike the other batches,
natural enemies in batch 97-3 were more abundant from the old sites than from the new
sites (Table 13). The most common natural enemies were A nthocoris Sp., P. cylindrica,

C. parallelus, and parasitic wasps (Table 13).

79

DISCUSSION

When I compared the density of T. piniperda, 1. pini, and 1. grandicollis progeny
reared from bolts with natural enemies to bolts without natural enemies, I found that there
were no statistically significant differences, but the trends for each of these species were
consistent. Tomicus piniperda progeny density was always higher in bolts with natural
enemies, which may have been an artifact of the relatively small number of bolts
colonized by this bark beetle. Another explanation for this trend is that natural enemies
were more attracted to or had higher survival in bolts that had higher densities of T.
piniperda. Though the natural enemies may have actually reduced T. piniperda progeny
density, the difference could not be detected since the original density of T. piniperda
parent adults (attack density) in those bolts was much higher than the bolts without
natural enemies.

Ips pini progeny density was more complicated: in the first two batches of 1996, it
was higher in bolts without natural enemies, suggesting that predators and parasitoids
were reducing I. pini density. But in batch 96-3 and in all three batches of 1997, I. pini
offspring density was higher in bolts with natural enemies. Several predators in the
Coleopteran families Cleridae, Histeridae, and Tenebrionidae are known to use I. pini
Pheromones as kairomones (Herms et a1. 1991, Raffa 1991, Teale et a1. 1991), so it seems
likely that these predators would be more attracted to bolts with higher densities of I. pini
Parent adults.

Ips grandicollis offspring density was higher in bolts without natural enemies in

all batches except batch 96-2. In this case, it appears that natural enemies may have

80

reduced bark beetle progeny density, although the low sample size and high variability
resulted in insignificant differences.

The mean number of exit holes per m2 of phloem surface area was used as a
second measurement of scolytid progeny density, where one exit hole was assumed to
correspond to emergence by one scolytid progeny adult. Occasionally, more than one
bark beetle will emerge from a single exit hole (Hanson 193 7, Salonen 1973), so the
number of exit holes may slightly underestimate the number of progeny. In this study,
the number of exit holes/m2 was higher in bolts with natural enemies in four of the six
batches, which was consistent with the results from actual progeny density for T.
piniperda and I. pini.

In both years, T. piniperda productivity (progeny per gallery) was higher in bolts
without natural enemies. In addition, the relationship between T. piniperda productivity
and natural enemy density was significantly negative in 1997; the higher the density of
natural enemies, the lower the productivity. Since T. piniperda galleries were longer in
bolts with natural enemies, and gallery length is correlated with numbers of eggs laid
(Anderbrant 1990, Foltz et al. 1976, Light et a1. 1983), the munber of eggs laid was not
affected by natural enemies. Therefore, native natural enemies affected T. piniperda
productivity by reducing survival of developing eggs, larvae, or pupae, but not by
affecting parents.

In four of the six batches, l. pini productivity was higher in bolts without natural
enemies, and in five batches, there was a negative relationship between productivity and
natural enemy density. These findings for 1. pini parallel the results for T. piniperda:

natural enemies reduced productivity even though they did not reduce overall progeny

81

density. And, like T. piniperda, 1. pini productivity was likely affected by natural
enemies during the egg, larval, and/or pupal stages, and not at the parent adult/oviposition
stage because mean egg gallery lengths were not affected by the presence of natural
enemies.

Ips grandicollis productivity was higher in bolts without natural enemies in all
batches of both years, and with the exception of batch 96-2, there was a negative
relationship between productivity and natural enemy density. Unlike T. piniperda and I.
pini, more I. grandicollis progeny emerged from bolts without natural enemies.
Therefore, natural enemies may have reduced productivity either by preying on parent
adults before or during oviposition, or by causing mortality of the egg, larval, or pupal
stages.

Scolytid productivity can be affected by more than just natural enemies; mortality
due to intraspecific competition can also play an important role (Langstrom 1980, Schenk
and Benjamin 1969). To determine whether the trend of reduced productivity in bolts
with natural enemies was due to natural enemies or competition, I looked at the attack
densities in bolts with and without natural enemies, and I compared these attack densities
with other field studies where scolytid competition was documented.

Tomicus piniperda attack density (galleries per m2 of phloem surface area) was
higher in bolts with natural enemies in both years. One explanation is that natural
enemies were either attracted to bolts with higher densities of parent adults or had higher
Survival rates in bolts with higher scolytid densities. Another explanation is that natural
enemies simply did not reduce the number of attacks per bolt. Tomicus piniperda is one

Of the earliest bark beetles to colonize pine brood material in the spring (Haack and

82

Lawrence 1995b, Langstrom 1980), so large numbers of natural enemies were likely not

active early enough in the spring to impact the number of parent adults colonizing logs.

Ips spp. attack densities were also higher in bolts with natural enemies in all six of
the batches, and these differences were statistically significant in three of the batches.
Again, natural enemies did not appear to reduce the number of colonizing parent adults;
in fact, they may have been attracted to the bolts by the high numbers of adult beetles.

Since the number of eggs can be correlated with the length of scolytid egg
galleries (Langstrom 1980, Salonen 1973), I looked at the effect of natural enemies on
total gallery length (and indirectly on the number of scolytid eggs laid). I also wanted to
determine if natural enemy presence affected the average gallery length excavated per
female, which would also affect egg production. Natural enemies did not reduce the total
or the mean gallery lengths for either T. piniperda or 1. pini, rather, the galleries were
longer in the bolts with natural enemies. Two inferences can be made: (1) natural
enemies did not affect the parent adults constructing the egg galleries, and/or (2) natural
enemies had higher survival in bolts where many scolytid prey were developing. These
bark beetle parent adults were protected from most natural enemies while they were
constructing galleries under the bark. T hanasimus dubius, the most abundant predator of
adult scolytids, was likely too large to enter these bark beetles’ galleries, and primarily
Preyed on adult scolytids on the outer bark surface.

Finally, to re-examine the question of whether competition or natural enemies
Caused the reduction in scolytid productivity, 1 compared the attack densities for T.
Piniperda and I. pini with those reported in the literature in this region (there were no

findings for I. grandicollis in the Great Lakes region). The mean of about 21 T.

83

piniperda galleries/m2 in this study was lower than Haack and Lawrence’s (1995a)
finding of 179 T. piniperda galleries/m2 in Scotch pine bolts in Michigan. The finding of
about 71 Ips spp. galleries/m2 was much lower than a Wisconsin study, where mean I.
pini attack densities of 186 galleries/m2 on jack pine (Pinus banksiana Lamb.) were
recorded (Schenk and Benjamin 1969). Ips pini attack densities in this study were
comparable to Hack and Lawrence’s (1995a) mean I. pini attack densities of 26
galleries/m2 [number of nuptial chambers, not individual egg galleries, were counted; two
to four females per male are typical (Schenk and Benjamin 1969)] in Scotch pine bolts.

It has been well documented that at high attack densities, the length of bark beetle
galleries decrease, the number of eggs laid per gallery decrease, and the number of
surviving progeny per gallery decrease as the phloem becomes more scarce (Langstrom
1980, Salonen 1973, Schenk and Benjamin 1969). At the relatively low attack densities
observed in this study, however, it is unlikely that competition played such an important
role. I hypothesize that the reduction in scolytid productivity in bolts with natural
enemies was primarily due to predation and parasitism, not competition.

Several native bark beetle predators and parasitoids successfully reproduced in T.
piniperda-infested red pine bolts. Three important scolytid predators had the highest
densities in batch 1, bolts colonized by T. piniperda: P. cylindrica (batch 96-1), Medetera
sp. (batch 97-1), and T. dubius (batch 97-1). But more species of predators and greater
numbers of predators were reared from Ips spp.-infested bolts and from bolts infested
with both T. piniperda and Ips spp. This suggests that natural enemies are (1)
phenologically better timed with native Ips spp. than the exotic T. piniperda, (2) more

attracted to bolts with Ips spp. than those with T. piniperda alone, or (3) can reproduce

84

more successfully in bolts with Ips spp. than those with T. piniperda. Based on
phenological data from a related study, I found that most native natural enemies are
phenologically well-timed with native pine bark beetles, but that four predaceous species
are active as early as T. piniperda.

Bright (1996) reared seven species of parasitoids and only four species of
predators from a single T. piniperda-infested Scotch pine tree in Ontario. I reared six
parasitoid species, but cannot be certain that they were from T. piniperda because other
scolytids and weevils were also present in the T. piniperda bolts. Further research is
needed to determine the relative effects of parasitoids on bark beetle pOpulations.
Conclusions

Tomicus piniperda, I. pini, and 1. grandicollis offspring density was not affected
by predators and parasitoids, but productivity of these species was reduced in bolts with
natural enemies compared to bolts without natural enemies. Attack density and gallery
length of these species were higher in bolts with natural enemies, suggesting that natural
enemies did not have an effect on colonizing parent adults. The field-imposed attack
densities in this study were relatively low, so interspecific and intraspecific competition
were likely not the important mortality factors; predation and parasitism appear to be the
primary cause of reduced scolytid productivity. Natural enemies were more common in
bolts with Ips spp. than with T. piniperda alone, but, in general, they were attracted to

bolts with high numbers of colonizing adults of either species.

85

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Teale, S. A. and G. N. Lanier. 1991. Seasonal variability in response of Ips pini
(Coleoptera: Scolytidae) to ipsdienol in New York. J. Chem Ecol. 17(6): 1145-
1158.

Ye, H. 1991. On the bionomy of Tomicus piniperda (L.) (Col, Scolytidae) in the
Kuruning region of China. J. Appl. Entomol. 112: 366-369.

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CHAPTER 3

Interspecific Competition Between Native Ips pini and Introduced

T omicus piniperda (Coleoptera: Scolytidae) Bark Beetles in Red Pine in Michigan

INTRODUCTION

Interspecific competition for food and space can be a major mortality factor when
more than one bark beetle species attacks the same host tree species. Dead, weakened,
and stressed trees are usually scarce and are distributed unevenly both temporally and
spatially (Schroeder 1992). Weakened pines may be attacked simultaneously by several
species of bark beetles, creating potential competition for food and breeding space.
lnterspecific competition has been demonstrated to increase mortality in numerous bark
beetle species. For example, interspecific competition between Ips pini (Say) and
Dendroctonus ponderosae Hopk. resulted in reduced success for both species when they
infested the same host material (Rankin and Borden 1991). Ips pini and I. paraconfusus
Lanier both had lower brood adult survival when the species were intermixed than when
alone at the same densities (Light et al. 1983). Exploitative competition is believed to be
the principal type of interspecific competition operating between bark beetles (Rankin
and Borden 1991). This occurs when members of one species utilize common resources
that are in short supply (i.e. larval breeding material) (Krebs 1994).

The recent establishment of an exotic pine bark beetle, Tomicus piniperda (L.), in
the Great Lakes region allows us a rare opportunity to evaluate the interactions between

native pine bark beetles and an exotic species that compete for the same resources. The

114

larger pine shoot beetle, T. piniperda, a well-known pest of pines throughout much of
Europe, Asia, and North Africa (Langstrom 1980, Ye 1991), was first discovered in 1992
in a Scotch pine (Pinus sylvestris L.) Christmas tree plantation in Ohio (Haack and
Kucera 1993). As of September 1998, T. piniperda had been found in 243 counties in
nine states, and in 21 counties in the Canadian province of Ontario (NAPIS 1998).
Tomicus piniperda has a low temperature threshold for spring flight (IO-12°), and is the
first pine bark beetle to fly in the spring in Michigan (Haack and Lawrence 1995a).
Adult beetles are attracted to stressed, dying, or recently-cut pine trees by host volatiles
formed in the dying tissues of conifers (Volz 1988, Schroeder 1992, Schroeder 1987) and
by short-range sex pheromones (S. Teale, State University of New York Syracuse,
personal communication). Tomicus piniperda is univoltine (Langstrom 1980, Haack and
Lawrence 1995b). Progeny adults feed in current-year or one-year pine shoots for the
remainder of the summer (Langstrom 1980, McCullough and Smitley 1995).

The pine engraver, Ips pini, is the most economically important pine bark beetle
in the Great Lakes region (Schenk and Benjamin 1969). Adults of this common scolytid
also colonize dead or dying pines, but]. pini can produce three generations per year
(Schenk and Benjamin 1969). Both 1. pini and T. piniperda overwinter as adults, but I.
pini typically becomes active in the spring four to six weeks later than T. piniperda
(Haack and Lawrence 1995a). Ips pini adults are attracted in large numbers to host
material by two aggregation pheromones that I. pini males produce: ipsdienol and
lanierone (Raffa and Klepzig 1989, Seybold et a1. 1992, Vite et a1. 1972). These

differences in the number of generations, the date of peak colonization activity, and the

115

use of aggregation pheromones could presumably influence the interactions between
these two scolytids.

Red pine (Pinus resinosa Ait.) is one of the most extensively planted species for
pulp, sawtimber, and poles in the Great Lakes region (Rudolf 1990), comprising over
364,000 hectares of Michigan’s timberland (Leatherberry and Spencer 1996). Red pine is
a suitable host for T. piniperda as well as 1. pini and many other pine bark beetles
(Lawrence and Haack 1995, Raffa 1991). During a typical rotation of red pine, several
thinnings or partial harvests may be performed, each of which generates an abundance of
suitable brood material for scolytid colonization.

I examined interactions between T. piniperda and 1. pini in laboratory and field
experiments in red pine forest stands in Michigan in 1996 and 1997. The objective was
to evaluate the effects of interspecific competition between T. piniperda and I. pini by
comparing attack rates, gallery lengths, progeny density, and productivity in (1) red pine
bolts artificially infested with one or both of these species, and in (2) red pine bolts

naturally infested in the field by one or both of these species.

116

MATERIALS AND METHODS

Field Sites

In 1996, I conducted field studies in four red pine forest plantations in Michigan.
Two plantations were in southwestern Michigan and were known to be infested with both
T. piniperda and native Ips spp. These stands were located in Michigan State
University’s WK. Kellogg Experimental Forest in Kalamazoo County (KALl), and in
the Allegan State Game Area in Allegan County (ALLl) (Figure 1). Two red pine
plantations in northern lower Michigan, where T. piniperda populations were not known
to be present, were also selected. One stand was in the Huron-Manistee National Forest
in Wexford County (WEXl), and the other was in the AuSable State Forest in
Roscommon County (ROSl). Although T. piniperda was later detected in Wexford
County in 1997 during regulatory surveys, (NAPIS 1998), I did not observe any T.
piniperda life stages or evidence of infestation in the northern field sites in 1996 or in
1997.

In general, all four research sites shared the following characteristics: they were
pole-sized red pine plantations of similar size, age, and basal area (Table 1) with some
hardwood understory; other red pine plantations were located within 2 km of the sites;
and they were thinned during the 1994-1995 winter. An abundance of slash (for
example, trunks, tops, large branches), suitable for scolytid colonization, was left on the
ground in each stand from the thinnings, and was expected to serve as breeding material

for scolytids in 1995. Temperature, precipitation, and degree day accumulation (base

117

10°C) were monitored after March 1 by Michigan State University weather stations near
all four sites.

In 1997, I used the same four sites described above, which represented conditions
two years afler a thinning (“old” sites). I also expanded the study to include four
additional red pine plantations (KAL2, ALL2, WEX2, and R082), each one within 6.2
km of one of the original stands. These “new” sites had the same general characteristics
of the original stands (Table l), but were row-thinned during the 1995-1996 winter.
Artificial Infestations

Red pine bolts, each approximately 61 cm long and 10 — 20 cm diameter, were cut
from live red pine trees in the W. K. Kellogg Experimental Forest in Kalamazoo County
in March 1996. A total of 120 bolts were used in the study: 40 bolts were infested with
T. piniperda, 40 were infested with I. pini, and 40 were infested with both species.
Tomicus piniperda and 1. pini adults used in this study were collected live from Lindgren
funnel traps in red pine stands at Kellogg Experimental Forest and in a mixed Scotch and
red pine stand at the Fenner Nature Center in Lansing, in Ingham County. Additional 1.
pini adults were collected from galleries under the bark of red pine in a recently-thinned
stand in Wexford County in late May. Tomicus piniperda were sexed on the basis of
stridulation (Bakke 1968), and I. pini were sexed on the basis of the third spine on the
elytral declivity (Wood, 1982).

Ips pini infestations took place six weeks after T. piniperda infestations, reflecting
the average time difference in initial flight and colonization between these species in
Michigan (Haack and Lawrence 1995a). Beetles were introduced into the bolts by

making a small “starter” hole in the bark of the bolts with the point of a nail sterilized in

118

95% EtOH. One beetle of the initiating sex (female T. piniperda or male 1. pini) was
placed in the hole, and one half of a size 00 clear gelatin capsule was placed over the
beetle and starter hole. The capsule was sealed to the bark with nontoxic clay. After 24
hours, the capsules were checked for the presence of boring dust as an indicator of a
successful attack. If boring dust was present, a single beetle of the other sex (male T.
piniperda or female 1. pini) was added to the gelcap. If the beetle did not successfully
bore into the phloem, it was removed, and another beetle was added to the same hole.
For the polygamous 1. pini, two females were added for each male; the second added 24
hours after the first. When only one species was introduced, five infestations were made
on small (10 — 13 cm diameter) bolts (about 23 females/m2), six infestations were made
on medium (13 — 17 cm diameter) bolts (about 21 females/m2), and seven infestations
were made on large (17 - 20 cm diameter) bolts (about 20 females/m2). When both
species were introduced, a combined total of ten, twelve, and fourteen infestations (46,
42, and 40 females/m2) were made, respectively, on small, medium, and large bolts. The
points of infestation were picked at random and were spaced over the entire bolt.

These bolts were taken to the field sites immediately after all of the infestations
were successfiilly completed. Bolts infested with I. pini were distributed equally to all
four sites, and bolts with T. piniperda or T. piniperda + 1. pini were distributed equally
between the two southern (KFl and ALLl) sites only. Bolts were placed in a shaded part
of each stand to reduce potential desiccation. Four to six bolts from each group (T.
piniperda alone, 1. pini alone, and both species) were returned to Michigan State
University at two week intervals to rear out the insects. Collected bolts were placed into

emergence containers and reared as described below.

119

l was not able to repeat this experiment in 1997. Unfortunately, most T.
piniperda beetles collected from the funnel traps in 1997 developed a fungal disease in
the lab and died before the experiment began, and few live I. pini adults were collected in
funnel traps.

Field Infestations

In 1996, a total of 204 red pine bolts, and in 1997, a total of 304 bolts, each
approximately 61 cm long and 10 — 20 cm diameter, were cut from live red pine trees at
the field sites in February, April, and June of both years (Table 1). These times roughly
corresponded to the initial activity of overwintering parent T. piniperda, overwintering
parent Ips spp., and emergence of the F1 progeny Ips spp. adults. These three times
represented three batches, or groups, and the bolts in each batch were divided evenly
among all field sites, with the exception of batch 1 (Table 1). Batch 1 bolts were only
placed in the southern sites since T. piniperda was not present in the northern sites. Each
batch of bolts was placed on the forest floor in a shady area of each plantation to reduce
desiccation, just before the expected flight of T. piniperda or Ips spp. (Table 1).

Beginning two weeks after the bark beetles’ peak flight, four to six bolts from
each batch were returned to Michigan State University. Bolts were retrieved at two week
intervals in 1996 and at one week intervals in 1997. The cut ends of the retrieved bolts
were dipped in paraffin wax to reduce desiccation. Each bolt was placed in an individual
emergence container consisting of a cardboard tube (15 — 25 cm diameter, 0.3 — 0.6 cm
wall, 61 — 71 cm overall length [Michigan Can & Tube, Inc., Saginaw, MI]) with opaque

plastic endcaps and a clear plastic collection cup at one end. The emergence containers

120

were stored on the Michigan State University campus in a screened insectary to expose
the bolts to ambient temperatures and humidity.

Bark beetle progeny adults emerging from the red pine bolts were collected from
the emergence containers at least twice a week until late October. When progeny
emergence stopped in the winter, I counted the number of scolytid exit holes per bolt, and
then each bolt was de-barked and any insects overwintering in the bark or dead on the
bottom of the emergence container were collected. All the insects that were reared from
these bolts were assumed to be progeny adults. Although it is possible that a few parent
adults were inadvertently collected, the number of parents was assumed to be small and
similar across all bolts and sites.

For each debarked bolt, 1 determined the phloem surface area, calculated as a
cylinder using the mean of two length measurements and the mean of four diameter
measurements. I also measured the number and length of Ips spp. galleries (each egg
gallery radiating from the nuptial chamber was counted), and the number and length of T.
piniperda galleries for each bolt. Tomicus piniperda is monogamous, and its galleries are
long and straight and parallel to the grain of the wood; Ips spp. are polygamous, and their
galleries consist of a nuptial chamber with 2 - 4 radiating galleries in the shape of a “Y”
or “H.” All Ips spp. galleries in the artificially infested bolts were assumed to be those of
I. pini. For the field-infested bolts, though, I was not able to distinguish which Ips
species excavated each gallery; Ips grandicollis (Eichhoff) and Ips perroti Swaine were
reared from the field bolts along with I. pini. Based on data from a related study, though,
I. pini was the most abundant Ips species reared from the field-infested bolts. The

number of scolytid egg galleries per bolt was used as an indicator of the density of female

121

parent beetles (1 egg gallery = 1 female parent), and the number of scolytid progeny per
female was used as a measure of productivity.
Data Analysis

For each artificially- and field-infested bolt, I calculated the number of T.
piniperda and Ips spp. galleries per m2 of phloem surface area (attack density), the
average length of T. piniperda and Ips spp. galleries per bolt, the number of scolytid
progeny reared per m2 (progeny density), and the number of scolytid progeny reared per
parent female (productivity). If fewer than five adults of a particular scolytid species
were reared from a bolt, it was considered a failed attack, and that species was excluded
from that bolt’s data set.

All variables were tested for normality with the Shapiro—Wilk W test, and were
log transformed to increase homogeneity of variances and tested again for normality.
None of the variables were normally distributed, even after log transformation, so the
nonparametric Mann-Whitney one-way test was used (SAS Institute 1996). The
artificially infested bolts were analyzed with the Mann-Whitney test to compare these
variables (attack density, gallery length, progeny density, and productivity) for T.
piniperda and 1. pini between bolts with one species and bolts with both species. Each of
the three batches of field-infested bolts per year was analyzed separately with the Mann-
Whitney test to analyze the effects of location (northern = without T. piniperda or
southern = with T. piniperda) on these variables for Ips spp. For batches 96-1 and 97-1,
which were placed only in the southern sites, 1 compared these variables between bolts

with one species and bolts with both species.

122

RESULTS

Artificial Infestations

Ips pini attack densities (number of galleries/m2 of phloem surface area), mean
gallery length, progeny density (number of I. pini progeny adults reared/m2 of phloem
surface area), and productivity (number of I. pini progeny adults reared/gallery) were all
significantly lower in the bolts infested with both 1. pini and T. piniperda than in the bolts
with I. pini alone (Table 2). An average of 43 I. pini females/m2 (= attacks) were
introduced into 80 of the laboratory bolts. In the bolts with I. pini alone, fewer than half
of these attacks resulted in successful gallery completion, and in the bolts infested with
both species, only 20% of the attacks were successful.

Tomicus piniperda attack densities, gallery length, and productivity were all
slightly lower in bolts with both T. piniperda and I. pini than in the bolts with T.
piniperda alone, but none of the differences were significant (Table 3). Significantly
fewer T. piniperda progeny/m2 emerged from bolts infested with both species than from
bolts with T. piniperda alone (Table 3). In the laboratory, about 22 T. piniperda

females/m2 ( = attacks) were introduced into each bolt. About two-thirds of these attacks

resulted in successful gallery completion.

Field Infestations: Effect of Location on Ips spp.

In the field-infested bolts of batches 96-2 and 96-3, Ips spp. attack densities were
lOWer in the southern sites than in northern sites, but differences were not significant
(Table 4). In 1997, Ips spp. attack densities were significantly lower in southern sites in

batch 97-3, and were lower in southern sites in batch 97-2, though not significantly

123

(Table 5). Ips spp. gallery lengths followed the opposite trend: galleries were
significantly longer in the southern sites than in the northern sites in batches 96-2, 97-2,
and 97-3 (Tables 4 and 5).

Ips spp. progeny density (number of progeny reared/m2) did not differ between
northern sites and southern sites in 1996 (Table 4), but in 1997, Ips spp. progeny density
was significantly lower in the bolts from the southern sites in batches 97-2 and 97-3
(Table 5). Mean Ips spp. productivity did not significantly differ between northern and
southern sites in either year (Tables 4 and 5).

Field Infestations: Single Species vs. Both Species

Only a single bolt was infested by both T. piniperda and Ips spp. in 1996. I could
not statistically analyze the 1996 data, although the results indicate that each of the
variables were fairly consistent from year to year when logs were only colonized by only
one of the species (Tables 6 and 7).

In 1997, eight bolts were colonized by both T. piniperda and Ips spp. in the field.
Ips spp. attack density, mean gallery length, and progeny density were lower in bolts
attacked by both T. piniperda and Ips spp. than in the bolts with Ips spp. alone, but the
difference was not significant (Table 6). Productivity for Ips spp. , however, was
significantly higher in bolts with both species (Table 6).

Tomicus piniperda attack density and progeny density were higher in bolts
attacked by both T. piniperda and Ips spp., but the differences were not significant (Table
7). Average T. piniperda gallery length was significantly higher in bolts with both
Species (Table 7). Tomicus piniperda productivity was lower in bolts attacked by both

Species, though not significantly (Table 7).

124

DISCUSSION

Several field and laboratory studies have demonstrated that interspecific
competition can occur between bark beetles, but the relative importance of interspecific
competition in regulating population dynamics has been debated. Berryman (1973)
found that Scolytis ventralis LeConte was a poor competitor against secondary bark
beetles that attacked the trees killed by S. ventralis, especially during a period of outbreak
decline. Light et al. (1983) demonstrated that the effects of interspecific competition
between I. paraconfusus and 1. pini were much greater than the effects of intraspecific
competition in bolts with similarly high densities of single species. Ips pini has been
found to reduce Dendroctonus ponderosae Hopkins progeny production in lodgepole
pine, Pinus contorta var. Iatifolia Engelm. (Rankin and Borden 1991). Interspecific
competition can also affect niche breadth and resource partitioning in bark beetles (Paine
et a1. 1981).

Although interspecific competition has been studied for several aggressive bark
beetle species, it has not been thoroughly investigated in the Great Lakes region, where
endemic pine bark beetles are secondary invaders of dying or cut trees. Schenk and
Benjamin (1969) reported that both interspecific and intraspecific competition were
major mortality factors for I. pini in Wisconsin, but that intraspecific competition was
more important, since 1. pini comprised 95% of the scolytid population. The recently-
introduced bark beetle T. piniperda represents an additional competitor for I. pini in the
Great Lakes region. Haack and Lawrence (1995b) found that at high densities, T.

piniperda colonized all sides of Scotch pine logs including upper and lower surfaces, but

125

at lower densities, T. piniperda preferred to colonize only the sides of logs. When T.
piniperda was already present in a log, I. pini was restricted to colonizing the tops of the
logs, which can be subject to extreme temperature fluctuations due to direct exposure to
the sun. Haack and Lawrence’s (1995b) results demonstrated that T. piniperda and I. pini
partitioned the phloem resource, but there was likely not much direct interaction between
the species.

In the lab portion of this study, the infestations of T. piniperda and 1. pini were
spaced over all sides of the bolts, so that the two species were not spatially segregated
from each other. Overall, the artificial introductions of bark beetles to the red pine bolts
were less successful than expected; 66% of the T. piniperda artificial introductions and
only 30% of the I. pini artificial introductions resulted in successful galleries. There are
several possible reasons for the poor success rates. I observed that the initiating sex of
both species (male I. pini, female T. piniperda) usually began boring into the outer bark
and phloem readily, but once the beetles were under the bark, some may have died. I also
witnessed a few cases of beetles boring into the phloem, and then promptly chewing a
second hole out of the bark, just a few centimeters away, suggesting that the region of
bark or phloem was unsuitable to the parent beetles.

The lab-imposed attack densities were selected to represent low field populations
and to reduce the chance of intraspecific competition. The artificial attack densities were
comparable with those seen in the field-infested batch 1 red pine bolts: about 74 Ips spp.
(primarily I. pini) attacks/m2 and 30 T. piniperda attacks/m2 (Tables 6 and 7). In another

field study of T. piniperda and I. pini attack densities in Michigan, Haack and Lawrence

126

(1995b) found similar I. pim’ attack densities, but much higher T. piniperda attack
densities (179 attacks/m2) in field-infested Scotch pine bolts.

Compared with other scolytid competition studies, the lab-imposed attack
densities in this study were much lower. In other interspecific competition studies with I.
pini, densities of 300 and 600 l. pini galleries/m2 (Rankin and Borden 1991) and densities
of 200, 400, and 800 I. pini galleries/m2 (Light et a1. 1983) were used.

Even at relatively low gallery densities in the lab-infested bolts, 1. pini galleries
(attack densities and gallery lengths) and progeny (density and productivity) were
significantly lower in bolts that were first infested with T. piniperda than in bolts infested
with I. pini alone. The field-infested bolts showed similar trends for Ips spp., indicating
that the lab study accurately represented the interactions taking place in the field. The
only inconsistent result in the field-infested bolts was that Ips spp. productivity was
higher in the bolts with both species. This finding may be an artifact of the small number
of bolts infested with both species and lack of replication in the field-infested bolts.
Further field and lab research on the interactions of these species is needed.

The most likely reason for I. pini’s lower attack densities, gallery length, progeny
densities, and productivity was that T. piniperda parent adults, with their six week head
start over I. pini (in the field and in the lab), colonized most of the available phloem in
the red pine bolts. This is an example of exploitation competition, where the first species
to arrive at the limited resource has better survival than the species that arrives later
(Krebs 1994). Interference competition may also be taking place, where one species has
the capabilitity to damage another; for example, through aggressive acts or cannibalism

(Krebs 1994). By the time I. pini larvae hatch, T. piniperda larvae would likely be much

127

larger, and would be capable of consuming any small 1. pini eggs and larvae they
encountered. A third possible explanation for I. pini’s poor success in the bolts with both
species is that T. piniperda may have introduced fungi into the bolts that inhibited either
I. pini parent adults or larvae (Paine et al. 1981, Yearian et a1. 1972). Further research is
needed to elucidate which, if any, of these interactions are causing the reduced success of
[pmi

T omicus piniperda, in both the lab-infested and the field-infested bolts, fared
much better than did I. pini in the bolts with both species. Tomicus piniperda progeny
density in the lab-infested bolts was the only variable that was significantly decreased in
the bolts with both species compared to the bolts with T. piniperda alone. In the field-
infested bolts, all of the variables except T. piniperda productivity were actually higher in
the bolts with both species. Having six weeks to colonize and begin development
without competitive pressure from I. pini appeared to be the key to T. piniperda’s success
in this study. In a related study, I found that two other pine phloem borers, Pissodes
nemorensis Germar (Coleoptera: Curculionidae) and Hylurgops rugipennis pim’fex (Fitch)
(Coleoptera: Scolytidae), were actively colonizing the field bolts as early in the spring as
T. piniperda. These beetles were primarily colonizing the cool, damp underside of the
bolts, though, not the sides of the bolts, where T. piniperda was more prevalent.
Interspecific competitive interactions are reduced when the phloem resource is
partitioned, and when colonization occurs simultaneously (Paine et al. 1981, Rankin and
Borden 1991), so it is not likely that severe competition was taking place between these

early colonizers.

128

I took advantage of the fact that T. piniperda had not yet spread to northern
Michigan, and compared Ips spp. galleries (attack density and gallery length) and
progeny (density and productivity) in the northern Michigan sites with the southern sites.
Although Ips spp. attack densities were lower in the southern sites, where T. piniperda
was also present, than in the northern sites, this trend cannot solely be attributed to the
presence of T. piniperda. Higher attack densities in the northern sites could be simply a
result of the larger Ips spp. populations in northern Michigan, where pines are more
abundant (Rudolf 1990). Although I can speculate that years of competition with T.
piniperda may have reduced the Ips spp. populations in southern Michigan, this
explanation was not likely, and it was not within the scope of this study to determine the
basis for Ips bark beetles’ regional population size.

Gallery length can be directly correlated with number of eggs laid by scolytids
(Anderbrant 1990, F oltz et al. 1976, Light et al. 1983), with shorter galleries I expected to
find fewer progeny per gallery (Anderbrant et a1. 1985, Rankin and Borden 1991). Ips
spp. gallery lengths were longer, on average, in the southern sites (where T. piniperda
was also present) than in the northern sites. Although this seems to contradict the
previous finding of lower attack densities in the southern sites, it can actually be
explained by intraspecific or interspecific competition. At high densities, bark beetle
galleries tend to be shorter, presumably because of the deterioration of the phloem, or
because females stop gallery construction when they sense the presence of other females
(Anderbrant 1990). At lower densities, such as in the southern sites, galleries tend to be
longer. At the highest densities in the northern sites (279 attacks/m2), densities were high

enough to expect intraspecific competition to occur, since Schenk and Benjamin (1969)

129

found intraspecific competition to be a major mortality factor at attack densities of only
186 I. pini attacks/m2.
Conclusions

Even at the relatively low attack densities used in the lab study, Ips pini attack
density, gallery length, progeny density, and productivity were significantly lower in
bolts first attacked by T. piniperda than in bolts with similar densities of I. pini alone.
Tomicus piniperda galleries and progeny were generally unaffected by the addition of I.
pini later in the season. Studies on field-infested red pine bolts confirmed these findings.
Interspecific competition, probably in the form of exploitation competition, is

hypothesized as the primary reason for I. pini’s reduced success.

130

LITERATURE CITED

Anderbrant, O. 1990. Gallery construction and oviposition of the bark beetle Ips
typographus (Coleoptera: Scolytidae) at different breeding densities. Ecol.
Entomol. 15: 1-8. '

Anderbrant, O., F. Schylter and G. Birgersson. 1985. Intraspecific competition affecting
parents and offspring in the bark beetle Ips typographus. Oikos 45: 89-98.

Bakke, A. 1968. Ecological studies on bark beetles (Coleoptera: Scolytidae) associated
with Scots pine (Pinus sylvestris L.) in Norway with particular reference to the
influence of temperature. Medd. Norske Skogforsoksvesen 21(83): 443-602.

Berryman, A. A. 1973. Population dynamics of the fir engraver, Scolytus ventralis
(Coleoptera: Scolytidae), 1. Analysis of population behavior and survival from
1964 to 1971. Can. Entomol. 105: 1465-1488.

Foltz, J. L., A. M. Mayyasi, F. P. Hain, R. N. Coulson and W. C. Martin. 1976. Egg -
gallery length relationship and within-tree analyses for the southern pine beetle,
Dendroctonusfi'ontalis (Coleoptera: Scolytidae). Can. Entomol. 108: 341-352.

Haack, R. and D. Kucera. 1993. New introduction - common pine shoot beetle, Tomicus
piniperda (L.). USDA Forest Service, Northeastern Area, Pest Alert NA-TP-OS-

93. 2 pp.

Haack, R. A. and R. K. Lawrence. 1995a. Attack densities of Tomicus piniperda and Ips
pini (Coleoptera: Scolytidae) on Scotch pine logs in Michigan in relation to
felling date. J. Entomol. Sci. 30( 1): 18-28.

Haack, R. A. and R. K. Lawrence. 1995b. Spring flight of Tomicus piniperda in relation
to native Michigan pine bark beetles and their associated predators. Pages 524-
535 in: F .P. Hain, S.M. Salom, F.W. Ravlin, T.L. Payne, and K.F. Raffa [eds],
Behavior, population dynamics, and control of forest insects, Proceedings of the
joint IUFRO conference for Working Parties 8207-05 and 8207-06. Maui,
Hawaii, 6-11 February 1994. Ohio State University, Wooster, Ohio.

Krebs, C. J. 1994. Ecology: the experimental analysis of distribution and abundance.
HarperCollins College Publishers, New York, New York. 801 pp.

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

131

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.
Pages 536-546 in F.P. Hain, S.M. Salom, F.W. Ravlin, T.L. Payne, and KR
Raffa [eds], Behavior, population dynamics, and control of forest insects,
Proceedings of the joint IUFRO conference for Working Parties 8207-05 and
8207-06. Maui, Hawaii, 6-11 February 1994. Ohio State University, Wooster,
Ohio.

Leatherberry, E. C. and J. S. Spencer Jr. 1996. Michigan forest statistics, 1993. Resource
Bulletin NC-170, USDA Forest Service, North Central Forest Experiment Station.
St. Paul, MN. 144 pp.

Light, D. M., M. C. Birch and T. D. Paine. 1983. Laboratory study of intraspecific and
interspecific competition within and between two sympatric bark beetle species,
Ips pini and I. paraconfixsus. Z. Ang. Entomol. 96: 233-241.

McCullough, D. G. and D. R. Smitley. 1995. Evaluation of insecticides to reduce
maturation feeding by Tomicus piniperda (Coleoptera: Scolytidae) in Scotch pine.
J. Econ. Entomol. 88(3): 693-699.

NAPIS (National Agriculture Pest Information System). Cooperative Agriculture Pest
Survey & NAPIS' page on PINE SHOOT BEETLE, Tomicus piniperda. [Online]
Available http://www.ceris.purdue.edu/napis/pests/psb/index.htrnl, September
1998.

Paine, T. D., M. C. Birch and P. Svihra. 1981. Niche breadth and resource partitioning by
four sympatric species of bark beetles (Coleoptera: Scolytidae). Oecologia 48: 1-
6.

Raffa, K. F. 1991. Temporal and spatial disparities among bark beetles, predators, and
associates responding to synthetic bark beetle pheromones: Ips pini (Coleoptera:
Scolytidae) in Wisconsin. Environ. Entomol. 20(6): 1665-1679.

Raffa, K. F. and K. D. Klepzig. 1989. Chiral escape of bark beetles from predators
responding to a bark beetle pheromone. Oecologia 80: 566-569.

Rankin, L. J. and J. H. Borden. 1991. Competitive interactions between the mountain
pine beetle and the pine engraver in lodgepole pine. Can. J. For. Res. 21: 1029-
1036.

Rudolf, P. O. 1990. Pinus resinosa Ait., Red Pine. Pages 442-455 in R. M. Burns and B.
H. Honkala [eds], Silvics of North America: 1. Conifers. Washington, DC, US.
Department of Agriculture, Forest Service. Agriculture Handbook 654.

SAS Institute. 1996. JMP start statistics. Albany, Duxbury Press. 52lpp.

132

Schenk, J. A. and D. M. Benjamin. 1969. Notes on the biology of Ips pini in central
Wisconsin Jack Pine forests. Ann. Entomol. Soc. Amer. 62: 480-485.

Schroeder, L. M. 1992. Olfactory recognition of nonhosts aspen and birch by conifer bark
beetles Tomicus piniperda and Hylurgops palliatus. J. Chem. Ecol. 18(9): 1583—
1593.

Schroeder, L. M. and H. H. Eidmann. 1987. Gallery initiation by T omicus piniperda
(Coleoptera: Scolytidae) on Scots pine trees baited with host volatiles. J. Chem.

Ecol. 13(7): 1591-1599.

Seybold, S. J., S. A. Teale, D. L. Wood, A. Zhang, F. X. Webster, K. O. Lindahl Jr. and I.
Kubo. 1992. The role of lanierone in the chemical ecology of Ips pini
(Coleoptera: Scolytidae) in California. J. Chem. Ecol. 18(12): 2305-2329.

Vité, J. P., H. A. Volz, M. R. Paiva and A. Bakke. 1986. Semiochemicals in host
selection and colonization of pine trees by the pine shoot beetle Tomicus
piniperda. Naturwissenschaften 73: 39-40.

Volz, H. A. 1988. Monoterpenes governing host selection in the bark beetles Hylurgops
palliatus and Tomicus piniperda. Entomol. Exp. Appl. 47: 31-35.

Wood, S. L. 1982. The bark and ambrosia beetles of North and Central America
(Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist
Memoirs 6: 1-1359.

Ye, H. 1991. On the bionomy of T omicus piniperda (L.) (Col., Scolytidae) in the
Kunming region of China. J. Appl. Entomol. 112: 366-369.

Yearian, W. C., R. J. Gouger and R. C. Wilkinson. 1972. Effects of the bluestain fiingus,
Ceratocystis ips, on development of Ips bark beetles in pine bolts. Ann. Entomol.
Soc. Amer. 65: 481-487.

133

 

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Figure 1. Location of red pine forest plantation field sites in Michigan.
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stands were monitored in 1997.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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APPENDIX 1

APPENDIX 1

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.

1998—8

Voucher No.:

 

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

InteractionsofthePineSfootBeetle [Tardous pinnpenda (L.) (Colemtera: Scolytidae)]
withNativePfleBarkBeetleeaflTleirAssmiatedthmlefieeinMdfigm

 

 

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

Entomology Museum, Michigan State University (MSU)

Other Museums:

Investigator's Name (8) (typed)
NmeKemedv

 

 

Date 16 Deoenber 1998

*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: Included 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 Miseum.

142

APPENDIX 1.1

APPENDIX 1.1
Vbucher Specimen Data

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145

LITERATURE CITED

LITERATURE CITED

Anderbrant, O. 1990. Gallery construction and oviposition of the bark beetle Ips
typographus (Coleoptera: Scolytidae) at different breeding densities. Ecol.
Entomol. 15: 1-8.

Anderbrant, 0., F. Schylter and G. Birgersson. 1985. Intraspecific competition affecting
parents and offspring in the bark beetle Ips typographus. Oikos 45: 89-98.

APHIS. 1996. Pine shoot beetle biological control project -- Project update. February
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