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This is to certify that the

thesis entitled

Host Preference and Host Suitability of Ten Hardwoods

for the Asian Longhorned Beetle, Anoplophora

glabripennis and of Four Conifers for the White-
Spotted Pine Sawyer Monochamus scutellatus

presented by

 

 

 

Laura Lenore Lazarus

has been accepted towards fulfillment
of the requirements for

M.S. degree in Entomology and

 

 

Ecology, Evolutionary Biology and Behavior

 

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Date / 20

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6/01 c-JCIRCJDateDuo.p65-p.15

HOST PREFERENCE AND HOST SUITABILITY OF TEN HARDWOODS FOR THE
ASIAN LONGHORNED BEETLE, ANOPLOPHORA GLABRIPENNIS
AND OF FOUR CONIFERS FOR THE WHITE-SPOTTED PINE SAWYER,
MONOCHAM US SC UT ELLATUS

By

Laura Lenore Lazarus

A THESIS

Submitted to
Michigan State University
In partial fulfillment of the requirements
For the degree of

MASTER OF SCIENCE

Departments of Entomology and
Ecology, Evolutionary Biology and Behavior

2003

ABSTRACT
HOST PREFERENCE AND HOST SUITABILITY OF TEN HARDWOODS FOR THE
ASIAN LONGHORNED BEETLE, ANOPLOPHORA GLABRIPENNIS
AND OF FOUR CONIFERS FOR THE WHITE-SPOTTED PINE SAWYER,
MONOCHAM US SC UTELLATUS
By
Laura Lenore Lazarus

Host selection behavior, and suitability of ten North American tree species for
Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae) oviposition, and
early instar development were examined during laboratory tests in 1999 and 2000.
Female A. glabripennis oviposited and larvae developed to the first instar on Northern red
oak, white oak, and hophombeam, while Ironwood, hackberry, honeylocust and
basswood were less suitable for oviposition and larval development. Tulip poplar,
sycamore, eastern cottonwood were not suitable hosts.

Relationships between host preference and host suitability were assessed for
Monochamus scutellatus (Say) (Coleoptera: Cerambycidae) in 2001 and 2002 in
Michigan. Female host preference and host suitability differed among jack pine, red
pine, white pine, and white spruce. Intra-specific competition for phloem was minimal,
but M. scutellatus deleteriously affected phloem-feeding scolytid bark beetles and
curculionids by reducing the area of phloem consumed and intercepting egg galleries.

Two-choice tests were conducted in 2002 to assess M. scutellatus female host
preference and larval performance on four conifers and larval conditioning. Adult M.
scutellatus preferred white pine and white spruce during the choice tests, while red pine

and white pine were most suitable for M. scutellatus larval development. Host preference

and larval performance was not entirely consistent for M. scutellatus.

ACKNOWLEDGEMENTS

I thank my advisor, Deborah G. McCullough, for her excellent example of what
hard work can achieve, her encouragement, and support throughout my Master’s
program. She helped me become a stronger person, to strive to reach my full potential,
and to take unforeseen events and surprises in stride. I would like to thank Bob Haack,
Rufus Isaacs and Jim Miller for their willing advice during this project.

I received an immeasurable amount of assistance and support throughout my
research program. Greg Kowalewski, Jim Curtis, John Vigneron, and Mickey Trimner
located, cut and shipped me the wood I tested during my Asian longhomed beetle study,
and they supplied me with logs for my White-spotted pine sawyer choice tests and field
study. They helped me select study sites, gave me assistance with biometry, and helped
to make my summer at W.K. Kellogg Experimental Forest fun. I gratefully acknowledge
Rachael Harris for her enthusiasm, for helping me haul logs, and for spending many hot
hours in the sun observing beetles at Kellogg Forest and the Tree research Center when
my knees wouldn’t allow me to do so. Abigail Sommers and Melanie DePoy did a super
job debarking and measuring hundreds of logs. I would like to give a special thanks to
John McCullough for taking the time to build me two custom designed olfactometers, and
a grid-writer to help with log measurements. Melody Keena provided me with Asian
longhomed beetles and some on the job advice. Jim Zablotney assisted with adult
Monochamus identification. I would also like to thank Randy Klevickas and Paul Bloese
for providing me with research space at the Tree Research Center, MSU, during 2001and

2002.

iii

I would like to thank all of my fellow graduate students and fi‘iends for making
my experience at MSU a pleasant one and for keeping things fim. In particular I’d like to
thank Emily Carlson, Kirsten Fondren, Meghan French, Amy Kearney, Andy Klein,
Rodrigo Mercader, and Nate Siegert for their companionship, advice and friendship.
Additionally, I am grateful and appreciative of Fred Moffitt for keeping me sane and
helping me remember to keep my life balanced. I wouldn’t have gotten anywhere
without the support of all my family and friends.

This research was funded by the USDA Forest Service’s North Central Research
Station (RJVA NC-23-99—24) and the McIntire-Stennis Cooperative Forestry Research
Grant Program. Additional funding was provided by the Ray and Bernice Hutson
Endowment from the Department of Entomology for travel support to national

entomological and ecological meetings.

iv

PREFACE

When the Asian longhomed beetle, Anoplophora glabripenm's, was first detected
in New York and Chicago and eradication programs were initiated researchers began to
realize the fill extent of the gaps in knowledge of cerambycid host-selection behavior,
the influence of the host on progeny survival, and about cerambycids in general. In this
thesis, I addressed relationships between host preference and host suitability of two
cerambycids, the nonindigenous A. glabripennis and the native Monochamus scutellatus.
In the case of A. glabripennis, researchers didn’t know how females select hosts, how far
they are capable of dispersing, or the potential for adaptation to novel hosts in the United
States. The white-spotted pine sawyer, M. scutellatus, is one of the most common
cerambycids in the US, nonetheless, there has been little research on M. scutellatus host-
selection behavior, host preferences and the suitability of different conifers for larval
development. Anoplophora glabripennis and M. scutellatus are both polyphagous and
are closely related species in the same tribe.

The host selection behavior and host range of A. glabripennis was studied in a
quarantine lab in Ansonia during 1999 and 2000. In chapter 1, specific behaviors of A.
glabripennis females associated with host selection and preferences among Northern red
oak, white oak, honeylocust, eastern cottonwood, sycamore, tulip poplar, basswood,
ironwood, hackberry and hophombeam for oviposition were evaluated, as was the
suitability of the ten hardwoods for early instar development. Researchers had a lot of

problems with the availability of beetles for laboratory studies in 2000 since colony

beetles kept dying or emerging malformed and unhealthy, and there were other problems
inherent when working with a non-indigenous species. We decided to leave the study of
A. glabripennis to others and to focus on the native wood-boring beetle, M. scutellatus.
There were no major constraints identified in obtaining adult M. scutellatus for laboratory
studies and this beetle could be studied in a natural environment.

In Chapter 2, the adult female host preferences of M. scutellatus, a common
cerambycid native to the lake states and much of North America were evaluated along
with the suitability of j ack pine, red pine, white pine and white spruce for larval
development in a field experiment. In chapter 3, a more controlled experiment was
conducted to assess the relationship between adult female host preference and larval
performance of M. scutellatus and to enable comparisons of results from both field and
controlled experiments. We hoped to gain valuable insight into the host-selection
behavior of cerambycids in the subfamily Lamiinae in general, or to identify similarities
in behaviors associated with host selection by studying this native beetle.

Each Chapter in this thesis is intended to be a prepared as a manuscript for
publication. We have published a part of the A. glabripennis work presented in Chapter 1

in The Journal of Environmental Horticulture.

vi

TABLE OF CONTENTS

LIST OF TABLES .................................................................................................... vii
LIST OF FIGURES .................................................................................................... ix
CHAPTER 1
Oviposition and Early Instar Survival of the Asian Longhorned Beetle
(Coleoptera: Cerambycidae) in Ten North American Hardwood Species ................. 1
Introduction .................................................................................................... 1
Methods and Materials ................................................................................... 6
Olfactometer Design ........................................................................... 6
Twenty—F our Hour Bioassays ............................................................ 7
F ifiy Minute Bioassay ........................................................................ 8
Host Suitabiliy ................................................................................... 10
Statistical Analyses ............................................................................ 12
Results ............................................................................................................ 14
Twenty-Four Hour Bioassays ............................................................ 14
Fifty Minute Bioassays ...................................................................... 16
Host suitability ................................................................................... 18
Discussion ...................................................................................................... 21
Tables ............................................................................................................. 26
Figures ............................................................................................................ 29
CHAPTER 2
Host Preference and Larval Performance of Monochamus scutellatus
(Say) and Competition for Phloem on Four Conifers .................................... 34
Introduction ........................................................................................ 34
Methods and Materials ....................................................................... 40
Host Preference and Adult Females ....................................... 41
Host Suitability ...................................................................... 43
Competition ........................................................................... 45
Statistical Analyses ................................................................ 45
Results ................................................................................................ 48
Host Preference ...................................................................... 48
Host Suitability ...................................................................... 50
Competition ........................................................................... 53
Discussion .......................................................................................... 57
Host Preference and Suitability ............................................. 57
Competition ........................................................................... 62
Tables ................................................................................................. 66

vii

Figures ................................................................................................ 70

CHAPTER 3

Effects of Host Preference on Larval Performance of Monochamus
scutellatus (Say) (Coleoptera: Cerambycidae) ............................................... 73
Introduction ........................................................................................ 73
Methods and Materials ....................................................................... 77
Statistical Analyses ................................................................ 79
Results ............................................................................................... 81
Host Preference ...................................................................... 81
Host Suitability ...................................................................... 82
Larval Conditioning ............................................................... 86
Diseussion .......................................................................................... 88
Tables ................................................................................................. 93
Figures ............................................................................................... 99
APPENDIX-Record of Voucher Specimen ............................................................ 101
LITERATURE CITED ............................................................................................ 103

viii

LIST OF TABLES

CHAPTER 1.

Table 1.1. Sequences of behaviors associated with oviposition for
three female A. glabripennis during 24 h in 16:8 (L:D), 23°C ...................... 26

Table 1.2. Mean (i SEM) percentage of time spent by A. glabripennis
females and the number of beetles out of the total tested for each
tree species that entered Olfactometer arms (first level choice) or
moved onto wood sections (second level choice), n=12 tol4
beetles per tree species ................................................................................... 27

Table 1.3. Mean number (i SEM) of A. glabripennis eggs, egg niches
and first instar larvae on 5 x 20 cm branch sections of ten North
American tree species; n=8 to 10 logs per species were used in
1999 and n=3 logs per species were used in 2000. Within columns
and year, means followed by the same letter are not significantly
different (P<0.05) ........................................................................................... 28

CHAPTER 2

Table 2.1. Characteristics of jack pine, red pine, Norway spruce and
white spruce compartments, and the number and size of logs of
jack pine, red pine, white pine, and white spruce set out in each
compartment May 2001 at WK. Kellogg Experimental Forest,
Kalamazoo, MI used to assess host preference of M. scutellatus.
Comparisons among conifers are made across columns ................................. 66

Table 2.2. Mean (1: SEM) number of M. scutellatus galleries per log for
colonized logs only, area of phloem consumed per log (cmz),
number of entrance holes into the sapwood, and the percentage of
larvae that died before entering the sapwood for Batch 1 September
to December 2001. These variables along with mean (:1: SEM)
number of exit holes and total survival are also shown for Batch 2
logs dissected in June to July 2002. Significant differences among
log species (rows) are indicated by different letters after the value ............... 67

Table 2.3. Number of adult M. scutellatus progeny and mean weight (g)
(1 SEM) of beetles that emerged summer 2002 from all jack pine,
red pine, white pine, and white spruce Batch-2 logs. Logs were
colonized summer 2001 at WK. Kellogg Experimental Forest.

ix

Significant differences among log species (within rows) are

indicated by different letters after the value ..............................................

Table 2.4. Mean number of M. scutellatus gallery interceptions,
M. scutellatus and scolytid bark beetle gallery interceptions and
M. scutellatus and curculionid gallery interceptions on Batch 1-
logs. Only logs with at least one interception were used in
analyses. Significant differences among log species (within rows)

are indicated by different letters after the value .........................................

CHAPTER 3.

Table 3.1. Mean (i SEM) bark thickness (inner+outer bark), diameter,
and length of logs used in 2-choice tests of M. scutellatus host

preference conducted summer 2002 ..........................................................

Table 3.2. Mean (i SEM) of variables associated with M. scutellatus
host preference and host suitability pooled for Jack pine (n=42),
red pine (n=49), white pine (n=57), and white spruce (n=52) logs
during 3-day trials in 2002. Significant differences within rows are

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

Table. 3.3. Mean (i SEM) of variables associated with host preference
and host suitability during 3-day choice tests in 2002. Each female
was presented logs of two conifer species. Significant differences
between the paired conifer species (in a row) are indicated by

different letters after the value. (P<0.05) ...................................................

Table 3.4. Mean (:tSEM) of variables associated with host selection and
host suitability during no choice trials to test larval conditioning
effects among conifers for adult female M. scutellatus host selection.
Jack pine (n=ltrial), red pine (n=9 trials), white pine (n=7 trials),
and white spruce (n=6 trials) in 2002. Significant differences

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

Table 3.5. Mean percentage of egg niches, eggs and number of galleries
(1- SEM) during larval conditioning trials in 2002. Female M.
scutellatus emerged from jack pine, red pine, white pine, and
white spruce logs. Each female was presented logs of 2 conifer
species. Significant differences within rows and between each

pair of conifer species are indicated by different letters alter the value.

..... 68

....69

...93

.. 94

....95

.96

(P<0.05) ....................................................................................................... . 97

LIST OF FIGURES

CHAPTER 1.

Figure 1.1. The olfactometer set-up used in 24 h and 50 min bioassays
in Ansonia, CT during the summer of 2000. The olfactometer
consisted of an acrylic central arena, four arms, and a glass lid .................... 29

Figure 1.2. Total percentage of time spent in specific behavioral
states by three A. glabripennis females over a 24 period .............................. 40

Figure 1.3. Percentage of time spent by three A. glabripennis females in
specific behaviors during 3 hour increments of 24 h bioassays ..................... 41

Figure 1.4. Duration and frequency (iSEM) of behavioral states for
three A. glabripenm's females in an olfactometer over a 24 hour
period .............................................................................................................. 42

Figure 1.5. Mean percentage of time (i SEM) that A. glabripenm’s
females spent in the central arena, in an olfactometer baited with
sugar maple or in the olfactometer arms that were baited with one
of the alternate tree species. Significant differences are designated
by different letters, n = 92 .............................................................................. 43

CHAPTER 2.

Figure 2.1. Frequency of behaviors associated with female M. scutellatus
host selection observed at Kellogg Forest from May 22 to
August 14, 2001 between 10 am. to 6 pm .................................................... 70

Figure 2.2. Mean area of phloem (:1: SEM) consumed by M. scutellatus,
scolytid bark beetles, and curculid weevils on Batch 1 logs that
were dissected in October to December, 2001 and Batch 2 logs
that were dissected in June and July, 2002. Only logs with M.
scutellatus galleries were used in analyses. Batch 1 included
55,76, 54, and 51 for jack pine, red pine, white pine and white
spruce logs. Batch 2 included 17, 24, 22, and 25 Jack pine, red
pine, white pine, and white spruce logs, respectively, in analyses.
Significant differences among log species are indicated by different
letters above columns for each type of phloem feeder ................................... 83

xi

Figure 2.3. Mean area of phloem consumed by M. scutellatus larvae,
scolytid bark beetles, and weevils on the top, sides, and bottom
of logs. Batch-1 logs were dissected September to December 2001
and Batch-2 logs were dissected summer 2002. Significant
differences among log species are indicated by different letters
above the column for each type of phloem feeder .......................................... 84

CHAPTER 3.

Figure 3.1. A graphic representation of the relationship between adult
female M. scutellatus host preference and larval performance on
white pine, red pine, jack pine, and white spruce logs from a
2-choice test experiment conducted summer 2002 ....................................... .99

xii

CHAPTER 1

Oviposition And Early Instar Survival Of The Asian Longhorned Beetle

(Coleoptera: Cerambycidae) In Ten North American Hardwood Species

INTRODUCTION

One of the most recent insects to become established in the United States is the
Asian longhomed beetle (Anoplophora glabripennis (Motschulsky); (Coleoptera:
Cerambycidae). This large, attractive beetle is an important forest pest in China and
Korea. In China, A. glabripennis is considered a major pest throughout a wide range
extending from 21°N to 43°N latitude and 100°E to 127°E longitude (Y an 1985), a band
that spans the climatic zones found in North America from southem Mexico to the Great
Lakes region (Haack et al. 1997). It reduces wood production in poplar (Populus spp.)
plantations, kills trees in windbreaks and is a pest on species of elm (Ulmus) maple
. (Acer) and willow (Salix). ‘

All life stages of A. glabripennis can be transported in logs, firewood and
untreated lumber. It was probably introduced to the US. in crates, pallets, or dunnage on
ships from China (Haack et al. 1997). Populations of A. glabripennis were first
discovered in 1996 in Brooklyn and Amityville, New York (Haack et al. 1996, 1997).
During the summer of 1998, three separate infestations were found in Chicago, Illinois
(Poland et al. 1998). Additional pockets of infested trees continue to be found in New

York and Chicago (Nowak et al. 2001). Quarantines and eradication programs have been

established in New York and Chicago to stop the spread of A. glabripennis (Nowak et al.
2001). The eradication program mandates that all trees with adult exit holes, egg niches,
or larval frass be cut down and chipped. The USDA Forest Service, state governments
and private groups have given millions of dollars to replace those trees cut down in
Chicago and New York (Haack and Mastro 1997). However, the full host range of this
beetle is still unknown and planting trees before host preference is understood could lead
to reinfestation if eradication is not successful.

The life history of A. glabripennis has been studied extensively in China (Gao et
al. 1993, He and Huang 1993, Li and Wu 1993, Zhao et al. 1997, Zhang et al. 2002) and
several studies have been conducted in the United States (Cavey et a1. 1998, Haack et al.
1997, Keena 2002, Smith et al. 2001, and Smith et al. 2002). Anoplophora glabripennis
is bi- or univoltine throughout its range in China, but only one generation a year occurs in
North America (Haack and Mastro 1997). Anoplophora glabripennis attacks healthy
trees, as well stressed trees, and recently cut logs or branches. Adults are diurnal and
volatiles (possibly sex pheromones) emitted from both males and females were shown to
have long range attractant properties (He and Huang 1993, Li and Wu 1993). Females
may also produce a short range contact pheromone (Zhang et a1. 2002). Most beetles
overwinter as larvae and new adults emerge the following summer through dime-sized
holes chewed through the bark (Haack and Mastro 1997). Newly emerged adults may
require up to 7 to 16 days of twig-feeding to reach sexual maturity. Adults live for 70 to
137 days and each female may oviposit from 50 to 193 eggs in the laboratory (Keena 2002,
Smith et al. 2001). Females chew through the bark to the cambial region, and lay a single

egg in a distinctive egg niche, often in branch junctures with the tree stem (Haack et al.

1997, Zhao et al. 1997). Eggs usually hatch in 1 to 2 weeks (Cavey et al. 1998). Larvae
feed in the cambial region for up to three weeks. Later instars score the sapwood and
tunnel into the sapwood where they overwinter. Adults emerge from June to September,
most frequently during the hottest part of the day (Kucera 1996, Nowak et al. 2001).
Adults can reinfest the larval host tree if it is still living, or, they may potentially disperse

1440 m or more (Smith et al. 2001).

Although adults can cause minor twig mortality during their maturation feeding,
the larvae cause significant damage to trees as they tunnel through branches and holes
(Gao et al. 1993, Haack and Mastro 1997). Individual branches or trees will die if larval
densities are high or if the infestation continues for several years. In North America, A.
glabripennis prefers maple (Acer spp.) trees and threaten the maple syrup and tourism
industries. It can also attack over 20 genera found in the eastern deciduous forests of the
US. (Haack et al. 1997, Nowak et al. 2001), larval feeding reduces the value of trees for
lumber (Haack and Mastro 1997). Homeowner property values in Chicago and New
York are also affected because all infested trees found on private property as well as on
public property are cut, chipped and burned (Haack and Mastro 1997). The estimated
maximum potential national urban forest impact of A. glabripennis is a loss of 34.9% of
total canopy cover, 30.3% tree mortality (1.2 billion trees) and a value loss of $669
billion (Nowak et al. 2001). Studies of several promising methods to control A.
glabripennis are underway using nematodes, predators and parasitoids, fungal pathogens
and systemic pesticides (Haack et al. 2001, Shimazu-Mitsuaki et al. 2002, Smith 2000,

Solter et al. 2001).

In Asia, the primary host trees of A. glabripennis include Acer spp., Salix spp.,
Populus, and Ulmus, although they will also feed on several other genera (Wu and J iang
1998). Boxelder (Acer negundo L.) is planted as a trap tree in parts of China to protect
more valuable hardwood trees (He and Huang 1993, Sun et al. 1990). In North America,
A. glabripenm's has primarily attacked species of maple, including Norway maple (Acer
platanoides L.), sugar maple (Acer saccharum Marsh), silver maple (Acer saccharinum L.),
sycamore maple (Acer pseudoplatanus (Bergahom)), and boxelder (Haack et al. 1997,
Poland et al. 1998). It has also infested species of birch (Betula spp.), poplar (Populus
spp.), willow (Salix spp.), horsechestnut (Aesculus hippocastanum L. ), and elm (Ulmus
spp.). At least three new host genera have so far been identified in the US: Aesculus,

F raxinus, and Hibiscus (Nowak et al 2001).

Host selection of most Cerambycidae of the Laminae subfamily may be
considered a three-step process where adults first first hosts using visual and olfactory
cues, then accept a host for oviposition using mostly chemoreception and host quality
characteristics, and finally offspring must be able to develop and survive (Bemays and
Chapman 1994, Hanks 1999). Little is known about the host-selection behavior of A.
glabripennis adults and how they respond to volatiles emanating from hardwood species.
Some nonhost plants may elicit feeding and be nutritionally adequate, but are not suitable
for development of egg to adult (Futuyma and Peterson 1985, Scriber 1984). Additional
North American tree species need to be tested to document whether they are attractive to
ovipositing females and suitable for larval development. Results will help to predict the
potential susceptibility of newly planted trees and of central and northeastern forests in

general. Evaluation of host suitability of North American tree species for A. glabripennis

could also help to develop guidelines for new plantings, monitoring protocols, and

management strategies.

In this study, I assessed the host-selection behavior of A. glabripennis females
during oviposition and evaluated the suitability of 10 North American tree species for A.
glabripennis early instar development. My primary objectives were to: 1) evaluate
preference of adult, virgin female A. glabripennis for six hardwood species native to the
north central region of the US; 2) evaluate specific behaviors of females during host
selection; and 3) evaluate the suitability of 10 North American hardwood species for

oviposition and larval development of the first instar.

METHODS AND MATERIALS

Olfactometer design. We developed a four-armed olfactometer to evaluate host
preference and host-selection behaviors of adult, virgin females (Figure 1.1). The
olfactometer was constructed out of acrylic and consisted of a central chamber 38 cm (15
in) tall with a diameter of 35.6 cm (14 in). Cylindrical chamber arms extended upward at
45° angles from the central chamber 42.7 cm (18 in) long and 7.6 cm (3 in) diameter.
The floor consisted of a circular piece of acrylic that fit tightly inside the arena, and
rested on brackets. A circular exhaust hole, 22.9 cm (0.9 in) diameter, was cut into the
center of the floor and covered with 0.33 cm (1/8 in) gauge wire mesh screen to allow
beetles to walk across the chamber floor. An acrylic lid, 38.1 cm (15 in) diameter was
placed on top of the central chamber. A Gast DOA-P104-AA vacuum pump (Grainger
Co., Lansing, MI) was attached to the exhaust hole on the chamber floor with 1.27 cm
(0.54 in) diam. Tygon tubing to pull air down through the chamber arms, into the central
chamber, and out the exhaust hole. Teflon tape was wrapped around all junctions to
ensure a tight seal. Incoming air was filtered through 0.64 cm (0.25 in) thick charcoal
filters attached to the distal opening of each arm. Smoke tests confirmed that the
chamber was completely sealed except for the designed flows. The olfactometer was
placed on a swiveling platform on the floor so that the position of the arms could be
rotated between each replicate, and the olfactometer was thoroughly cleaned with 75%
methanol before each bioassay.

All tests were conducted in an approved USDA Forest Service quarantine facility
in Ansonia, CT. We conducted preliminary studies to evaluate beetle behavior in the

olfactometer by placing individual female beetles into the olfactometer for 50 min. Two

arms of the olfactometer contained freshly cut sugar maple branch sections (5 x 20 cm)
and twigs and the other two arms were left empty. Of the five females tested, all entered
the arms containing sugar maple within 50 min. Therefore, we felt confident that beetles
were able to detect and select particular host volatiles in the olfactometer.

T wenty-Four Hour Bioassays. We monitored mating, host selection, and
oviposition behavior of a male-female pair of A. glabripennis adults in the olfactometer
for a 24 h. The length of the test was designed to allow adequate time for the completion
of behaviors associated with oviposition or host selection. Beetles used in the 24 hr
bioassays emerged from maple logs collected from infested trees in New York that were
reared at the Ansonia quarantine facility. Body weights were 0.62g, 1.05 g, and 0.49g for
female 1, 2 and 3, respectively and 1.00g, 0.68g, and 0.66g for males. Each 24 hour test
began at 0600 Eastern Standard Time. We used a 16:8 (L:D) photoperiod at 23C where
lights were turned off at 2200 hours.

Fresh sugar maple branch sections (5 x 20 cm / 2 x 8 in) and twigs were placed at
the top of two opposing arms of the olfactometer and the other two arms were lefi blank.
Seven to 10 fresh sugar maple twigs were placed with the wood sections to provide
material for attraction by the beetles.

We tested mating pairs of beetles so that all behaviors associated with oviposition
could be recorded in a more realistic sequence. Also, the presence of a male throughout
the entire test increased the liklihood that females would actively seek out oviposition
sites to deposit viable eggs. We placed a previously unrnated male and female A.
glabripennis pair in the olfactometer and allowed them to mate, move, excavate egg

niches, oviposit and feed throughout the duration of the test. We repeated the bioassays

three times, using a new pair of beetles each time. Female 1 was nine days post
emergence and was much smaller than the male or the other 2 females. Female 2 was 10
days post emergence and about the same size as the male. Female 3 was 10 days post
emergence and was larger than the male and the other females.

All behavior during the 24 hr (1440 min) trial was videotaped using a Sony
Digital camcorder, (model DCR-TRV103) (Park Ridge, NJ) mounted on a tripod. Video
from each bioassay was viewed later and the frequency, sequence, and duration of
behaviors were recorded. We determined the total time females were stationary and
engaged in walking, twig feeding, excavating egg niches, ovipositing, and copulating.
We also recorded the frequency and sequences of the behaviors throughout the 24 hr.

At the end of the test, branch sections from the bioassays were placed in a growth
chamber at 21° C (71°F) for 21 d and then dissected. We counted the number of egg
niches, eggs, and dead and alive larvae for each branch section.

Frfty minute bioassays. In this bioassay, we focused on host-selection behaviors
rather than behaviors associated with mating, so only females were included. We
monitored host selection of females in the olfactometer during 50 min bioassay using
similar olfactometer conditions and protocols for filming as in the 24 min bioassays June
2000. A total of 15 virgin female beetles was used in the bioassays. Females typically
initiate host selection and will excavate egg niches and oviposit even if they have not yet
mated. Beetles used had intact antennae and no evidence of physical damage. Each
female was tested with three to six host species because of the limited availability of

healthy beetles.

Potential hosts tested included eastern cottonwood (Populus deltoides L.),
honeylocust (Gleditsia triacanthos L.), northem red oak (Quercus rubra L.), sycamore
(Plantanus occidentalis L.), tulip poplar (Liriodendron tulipifera L.), and white oak
(Quercus alba L.). Sugar maple, known to be highly preferred by A. glabripennis, was
included in each bioassay as a positive control. Sugar maple, sycamore, tulip poplar and
white oak and twigs were collected from live trees in woodlots in Ansonia, CT. Northern
red oak, honeylocust, and eastern cottonwood branch sections and twigs were cut from
trees at W. K. Kellogg Forest, Augusta, MI and shipped in an insulated cooler by
overnight mail to the quarantine facility in Ansonia. Cut ends of all branch sections were
waxed with paraffin immediately after cutting to slow desiccation. Wood and branches
were stored in a growth chamber at 21° C (70° F) in the Ansonia facility until used for
bioassays (up to 1 wk). Paraffin was removed and wood sections and twigs were cut to
appropriate size with a bandsaw just before the olfactometer bioassays.

To evaluate female host preference, we suspended branch section of a test species
with bark attached (5 x 20 cm / 2 x 8 in) in two of the four arms of the olfactometer.
Seven to 10 fresh twigs of the test species were placed with the wood sections to provide
material to attract the beetles. Similarly-sized sections of sugar maple and twigs were
placed in the other two arms. Tests were conducted in the dark to prevent beetles from
simply responding to light. A glass Petri dish with a 1 g (0.002 lbs.) metal weight
attached to the bottom was used to introduce beetles into the central chamber. The
container was suspended with fishing wire through a pin-sized hole drilled through the

chamber lid. Each female beetle was kept in the holding container in the center of the

chamber for 5 min to acclimate to chamber conditions before the 50 min observation
period began.

To quantify behavior and attraction of female beetles to the alternative host
species, we observed beetles and noted if they stayed in the central arena or moved into
one of the arms, or onto the r branch sections. The central arena consisted of the lid,
wall, and floor of the olfactometer. To simplify analysis of female host selection, we
separated behaviors into first and second level choices. Beetles were said to have made a
first level choice if they walked into an olfactometer arm. Beetles that moved onto the
wood suspended in the arm were considered to have made a second level, or higher
choice, than those that only walked into the arms. Behavior associated with second-level
selection included: examining behaviors employing antennae or palps to sense the bark,
twig feeding, egg niche excavation, or oviposition.

Host suitability. Suitability of hackberry (Celtus occidentalis L.), hophombeam
(Ostrya virginiana (Mil) K. Koch), basswood (T iIia americana L.), ironwood (Carpinus
caroliniana Walt), and sugar maple for A. glabripennis oviposition and early larval
survival was assessed using mating male-female pairs of beetles in 1999 and 2000. One
pair of beetles was placed in a 3.8 liter (1 gal.) glass jar containing a fresh wood section
(2 x 8 in) and 7-10 twigs of one alternate host species. Branch sections were waxed on
the cut ends with paraffin to slow desiccation. The number of host species tested was
determined by availability of healthy beetles; a total of 10 pairs was available to test
basswood, hophombeam, ironwood, and sugar maple; eight pairs were available to test
hackberry. Age of females ranged from 9 to 48 days post emergence. The duration of

each test was 6 to 8 d for all species.

10

In 1999, mating pairs were randomly assigned to basswood, hophombeam,
ironwood, hackberry, or sugar maple. Beetles were allowed to mate, feed, excavate egg
niches, and oviposit on the branch section in each jar for 6-8 (1. Each pair of beetles was
exposed to only one test species and was used in only a single test.

At the end of each assay, wood sections were examined to determine the number
of egg niches on the wood, where the female beetle had clearly used her mandibles to
scrape the bark or create an egg niche. Wood sections were placed on end in a 21° C
(68° F) grth chamber, 16:8 (L:D) photoperiod and dissected 21 days later. Number of
eggs and first instars on each wood section were recorded.

In 2000, we followed similar protocols as in 1999 to assess suitability of eastern
cottonwood, honeylocust, northern red oak, sycamore, tulip poplar, and white oak for A.
glabripenm's oviposition and early larval survival. Sugar maple twigs were provided
during the assay instead of twigs of the test species to ensure that the beetles would
survive. A total of three male-female pairs was available to test northern red oak,
honeylocust, and eastern cottonwood (Test 1) and three pairs were used to test white oak,
sycamore, and tulip poplar (Test 2). Beetles used in test 1 were 25 to 48 days old, and
those used in test 2 were 54 to 55 days old. Beetles used in test 1 had all antenna]
segments intact, but one female was missing her left front tarsus and one female in test 2
had a malformed antenna. These beetles were used because of the limited number of
healthy beetles available for concurrent A. glabripennis research projects in 2000.

In Test 1, the three mating pairs were randomly assigned to northern red oak,
honeylocust, or eastern cottonwood, while one beetle pair remained on sugar maple for

the duration of the test. Beetles were allowed to mate, feed, excavate egg niches, and

11

oviposit on the wood in each jar for 4 days. Thereafter, each mating pair was placed into
a new jar containing sugar maple twigs and a sugar maple branch section for 2 days to
allow beetles to recover. Each mating pair was then assigned to a new species for 4 days,
followed by a 2 days recovery period on sugar maple. This process was repeated again,
so that each pair was exposed to all three different host species.

Test 2 included white oak, sycamore, and tulip tree along with sugar maple and
the methods were those of Test 1. A 61-day-old female of one of the mating pairs died
during the second resting period in Test 2. Another similarly aged mating pair of beetles
was used to complete the test. At the end of each 4 day period, the number of egg niches
were counted and branch sections were reared and dissected as before.

Statistical Analyses. Observations from the 24 hr bioassays were not analyzed
because of low sample size. However, we summarized the behavioral sequences,
frequency and duration of behaviors for each individual beetle. Data from fifty minute
bioassays and 1999 and 2000 host suitability trials were not normally distributed, based
on the Shapiro-Wilk test and residual plots (Shapiro and Wilk 1965). Data from 50 min
bioassays were pooled to compare beetles duration spent in arms baited with sugar maple
with duration spent in the arms baited with alternative host species using the
nonparametric Kruskal-Wallis test. Additionally, data were pooled to compare
differences in the percentage of time spent in arms of the olfactometer, and on wood in
the olfactometer, among sugar maple and alternate hosts using the Mann-Whitney Test
(Kruskal 1952). The Kruskal-Wallis test was also used to determine if the number of egg
niches, eggs, and larvae differed among alternative host species (Kruskal and Wallis

1952). All analyses were conducted at the P<0.05 level of significance, using SAS

12

statistical software (SAS Institute, Inc. 1989). The Kruskal-Wallis Multiple Comparison
procedure was used to determine which treatment pairs differed significantly (P<0.05)

(Conover 1971).

13

RESULTS

T wenty-four hour bioassays. The three adult females we monitored in the
olfactometer exhibited a range of behaviors during the 24 h period. Behaviors included
copulation, twig feeding, egg niche excavation, oviposition, and walking. Figure 1.2
shows the percentage of time spent three female A. glabripennis in each behavioral state
over 24 h. Females 1 and 3 spent more time twig feeding, about 20 % of the 24 h
bioassay, than Female 2 (8 % of the time), although female 2 spent more time excavating
egg niches (20 %) than females 1 (3 %) and 3 (12 %). Female 3 spent 20 % of the time
ovipositing while females 1 and 2 spent only 7 % and 8 % of the time ovipositing,
respectively. Female I spent the least time copulating, 3 %. All three females spent very
little time grooming during the 24 h, less than 5 % of the total time. Female I flew three
times during the test, while Females l and 2 never flew. Female I spent 5 h alone before
the male found her. She twig fed and excavated a single egg niche before the male joined
her and immediately attempted copulation and mate-guarding. We observed a similar
scenario with female 2. Female 3 was in close contact with the male for the entire test.

A more detailed analysis of behaviors observed over the 24 h bioassay was
achieved by dividing the 24 hours into 3 h increments and comparing percentage of time
allocated to each behavior by each female (Figure 1.3). During the 24 h, Female 1 was
not very active for the first three h, then she twig fed the majority of the time between 9
am. and 3 pm. Copulation occurred four times in the morning for an average of 16.0 i
5.7 min. Egg niche excavation and oviposition occurred for periods averaging 5.0 :1: 1.9

and 20.0 :i: 0.9 min, respectively, and occurred between 3 pm. and 6 pm. (Figure 1.3,

14

and Figure 1.4). The male bit Female 1 numerous times on the pronotum during
copulation, and she “ran away” from the male three different times. She also spent more
time apart from the male than the other females. From 6 pm. to 3 am, Female 1 either
walked or was stationary, except for a bout of twig feeding (Figure 1.3). This female
excavated an egg niche, then oviposited five times. Female 1 fed on twigs after every
oviposition event (Table 1.1). However, twig feeding preceded egg niche preparation in
only 2 of 5 cases. In 2 of the 5 oviposition bouts, Female 1 chewed another egg niche
immediately after laying an egg. C opulation was followed by twig feeding in 1 of the 2
copulations.

Female 2 was generally more active than Female 1 (Figure 1.3), although she was
mostly stationary until 8 am when she began twig feeding (Figure 1.3). Female 2
copulated and excavated egg niches in mid-moming, and copulated, excavated egg niches
and oviposited from noon until 3 pm. She spent 60 % of her time twig feeding from 3 to
6 pm. and twig fed for 27 :1: 9.90 min on average (Figure 1.3). Female 2 didn’t become
active again until 10 pm when bouts of copulation, egg niche excavation, and oviposition
began. Female 2 oviposited directly after egg niche excavation in only 2 out of 5 egg
niches and abandoned 7 egg niches without ovipositing in them (Table 1.1 and Figure
1.4). Mean duration of oviposition and egg niche excavation for this insect was 31.5 :1: 2.7
and 27.2 d: 4.4, respectively (Figure 1.4). Oviposition occurred after copulation in three
of five oviposition events, egg niche excavation followed copulation five out of eight
copulations and egg niche excavation followed ovipositon three out of eight times. The

beetle pair copulated seven times for 20 i 3.45 min on average (Figure 1.4).

15

Female 3 was stationary for a few h after the test began and then became very
active (Figure 1.3). Female 3 began twig feeding, copulating, and ovipositing in mid-
morning and continued twig feeding, chewing egg niches, ovipositing and copulating for
most of the day. At 6 pm, activity slowed and she spent 70 % of the next 2 h twig-
feeding. Female 3 copulated again around 8 pm. and then excavated egg niches and
oviposited until midnight. The female left the male and walked to the other olfactometer
arm containing wood then continued excavating egg niches and ovipositing from
midnight until 6 am. (Figure 1.3). This female also groomed more regularly than the
other two females. Since Female 3 actively exhibiting host selection behaviors
throughout most of the 24 hours, we saw more transitions between host selection
behaviors than for the other females. Female 3 oviposited in ten of the 12 egg niches she
excavated, but oviposited twice without excavating an egg niche in the bark. She
copulated immediately following egg niche excavation twice (Table 1.1) and chewed new
egg niches after 8 of the 14 oviposition events and after 5 of the 19 twig feeding events.
Twig feeding, copulation, egg niche excavation, and oviposition, lasted 17.6 d: 4.52, 17.5
:1: 6.54, 16.4 :1: 2.41, and 17.4 i 1.74 min on average, respectively (Figure 1.4).

Fifi‘y min bioassays. Overall, female time allocation to different parts of the
olfactometer varied considerably. Beetles spent 25 to 45 min of the 50 min observation
period in the central arena of the olfactometer and significantly less time in the chamber
arms or on the branch section (Figure 1.5) (P<0.05; d.f.=2). Forty-eight of 76 beetles
attempted escaping when introduced into the olfactometer, which included walking up
the central chamber walls and circling around the lid of the chamber. Beetles attempted

escaping and stayed in the center arena for an average of 39.2 :1: 5.1 min. Ten beetles

l6

attempted escaping when honeylocust was the alternate host, nine beetles for eastern
cottonwood, eight for northern red oak, nine for sycamore, seven for white oak, and five
for tulip poplar. It was likely that the beetles had not yet acclimated to the olfactometer
conditions after 50 min.

When any alternative host was present, beetles consistently spent less time on
sugar maple wood than when the beetles were given only sugar maple and blank (empty)
arms. When beetles left the central arena, there were significantly more visits to the
sugar maple arm and wood than to the alternative species arm and wood (P<0.05, d.f.=3).
Of the 92 total trials in the olfactometer, 60 beetles entered arms with sugar maple
sections and 31 beetles moved onto sugar maple wood (a second-level choice) (Table
1.3). In contrast, 15 beetles entered arms with an alternate host and six made a second
level choice (Table 1). Similarly, when beetles entered a chamber arm, they spent
significantly more time in sugar maple arms (P=0.004, d.f.=1) and on sugar maple wood
(P<0.001, d.f.=1) than on all other alternative species arms combined.

We found no differences in time allocated to the arms and wood of the
olfactometer when compared on a species by species basis, because relatively few beetles
selected arms with alternative host species. However, when northern red oak was the
alternative host, three beetles moved into the arms with red oak and two spent about 15
min on the wood (Table 1.2). In comparison, ten of the beetles moved into sugar maple
arms and the five that made a second level choice spent about the same amount of time
on the sugar maple wood as the beetles on the red oak wood (Table 1.2). Four beetles
selected arms with either white oak or eastern cottonwood, but the two beetles that made

a second level choice spent little time on the alternative host wood. Only two beetles

l7

made a first level choice for sycamore and one beetle made a second level choice, but
neither spent much time in these arms. No beetles made a second level choice for either
tulip poplar or honeylocust (Table 1.2).

Twig feeding was rare; four beetles fed on sugar maple twigs ranging from 2 to
10 min. There was no twig feeding on any alternative species. Seven beetles excavated
egg niches on sugar maple wood sections, when paired with an alternate host, a process
that ranged from 1 to 21 min. A single beetle excavated five egg niches on northern red
oak, with excavation activity persisting for 1 to 15 min per egg niche. This beetle
appeared to be positioning her body for oviposition when the 50 min bioassay period
ended. In addition, a single female excavated 4 niches on white oak lasting from 1 to 24
min. Only one beetle oviposited on sugar maple, for 13 min, and there was no
oviposition on wood sections of alternate hosts.

Host suitability. In 1999, number of egg niches, eggs, and survival of first instar
larvae varied widely among the host species tested (Table 1.3). Overall, egg hatch
ranged from 44.6 to 96.0% for the species tested. Significantly more egg niches were
created on hophombearn than on any other species tested, including sugar maple
(P<0.001, d.f.=4). Similarly, significantly more eggs were laid (P<0.001, d.f.=4) and
larvae recovered (P<0.001, d.f.=4) on hophombeam than for any other species tested.
Females caged on sugar maple laid 2 eggs per day, while females laid less than half as
many eggs on alternate species. Significantly more egg niches were created on sugar
maple than on ironwood (P<0.025), hackberry (P<0.025) but there were no differences

between sugar maple and basswood.

18

During the 2000 bioassays, the excavation of egg niches, number of eggs laid, and
survival of first instar larvae varied widely among the host species tested (Table 1.3).
The number of eggs per wood section ranged from 0 to 23 per section. Overall, 68% of
eggs hatched and those that did not hatch often appeared to be desiccated. Relatively
little phloem feeding had occurred at the time of dissection; by the end of the 21-d rearing
period, most first instar larvae had consumed an area slightly larger than their body size
(4-7 mm).

In Test 1, more eggs and larvae were recovered from sugar maple than from any
of the alternate species (Table 1.3) and the numbers were comparable to numbers of eggs
and larvae reared in other A. glabripennis projects at the Ansonia facility (Keena 2002).
In Test 2, however, relatively few eggs and larvae were recovered from the sugar maple
sections, perhaps because beetles assigned to sugar maple in Test 2 were older than those
used in Test 1. At the end of Test 2, the females were up to 73 days old, while at the end
of Test 1, beetles were no more than 66 days old. Female A. glabripenm's, however, are
fairly long-lived beetles and average adult female survival may exceed 100 days on some
hosts (Smith et al. 2002). Recent research has shown that egg production decreases as A.
glabripennis age, especially after the first mating, although the proportion of viable eggs
may increase (Keena 2000, Smith et al. 2002).

Northern red oak (Test 1) and white oak (Test 2) appeared to be more suitable
hosts for ovipositing A. glabripennis beetles than we originally expected. Female beetles
assigned to northern red oak sections usually excavated egg niches and oviposited around
branch nodes where the bark was relatively rough and thick. The total number of eggs

laid on northern red oak (23) was comparable to the number laid on sugar maple in Test 1

l9

(22), although only 39% of the eggs on northern red oak hatched compared with 82% of
the eggs on sugar maple. At least eight of the unhatched eggs on northern red oak
appeared desiccated at the time of dissection and the northern red oak appeared to dry out
more rapidly than sections of other species even though all cut ends of the branch
sections were waxed. White oak sections had significantly more egg niches excavated
than any other species in Test 2 (P<0.05, d.f.=3) (Table 1.3), but the shape of the niches
was unusual. We observed females repeatedly scraping the thick, corky bark on the
white oak sections, presumably to find suitable sites for oviposition, but they actually laid
eggs in less than half of the egg niches. Females laid significantly more eggs on white
oak than on other species in Test 2 (P<0.05, d.f.=3) and 87% of the eggs on white oak
sections hatched.

Honeylocust, sycamore, eastern cottonwood and tulip poplar appeared less
suitable for egg niche excavation and oviposition. The number of egg niches was
significantly lower on honeylocust than on northern red oak and sugar maple (P<0.05,
d.f.=3), but the number of eggs and larvae did not differ significantly between
honeylocust and northern red oak or sugar maple (Table 1.3). However, less than half as
many eggs were laid on honeylocust as on the sugar maple in Test 1, and only 50% of the
eggs on honeylocust hatched. Beetles did excavate a few egg niches on rough-barked
areas of cottonwood and sycamore sections, but no eggs were laid on any of these
sections. Tulip poplar appeared to be highly unsuitable as a host; there was no evidence

of any attempts to excavate egg niches or oviposit on these sections.

20

DISCUSSION

The goals of the 24 h bioassays were: to observe the temporal pattern of host
selection behaviors, to note when behaviors occurred, and the duration and sequence of
behaviors. Behavior of the three females varied considerably, suggesting that behavior
probably varies similarly in natural populations of A. glabripenm’s that will include an
even wider range of beetle age and size.

The data suggest that, overall, A. glabripenm‘s females display certain behavioral
sequences associated with oviposition. The typical sequence of behaviors we observed
was twig feeding and copulation succeeded by egg niche preparation then oviposition.
Adult Lamiinae are the only cerambycids that excavate egg niches prior to oviposition
(Hanks 1999). However, the sequences were not entirely consistent and at times this
sequence was disrupted by copulation or excavation of another egg niche. Congeners
Anoplophora macularia (Lee and Lo 1998), and Anoplophora chinensis (Wang et al.
1996) also chew an egg niche prior to ovipostion and exhibit similar sequences associated
with ovipositon site selection (Lee and Lo 1996, Wang et al. 1996). Inconsistencies in
the oviposition sequences described here could be explained by male interference. For
example, the egg niche excavation to oviposition sequence in our bioassays were
interrupted by copulation. Likewise, A. chinensis and M. scutellatus males are not always
' associated with females during oviposition (Hughes and Hughes 1982, Wang et al. 1996).
Futhermore, all females were somewhat active in the dark period between 10 pm. and 6

am.

21

Observations of the second stage of the host selection process indicate that
females constantly taped their palps and antennae along the surface of the wood while
walking to assess the suitability of the host for oviposition. Contact chemoreception has
been suggested to occur via drumming of the palps in Monochamous scutellatus (Say),
and Phoracantha semipunctata (F .) (Hanks 1999, Hanks et al. 1993, Hughes et al. 1992).

Differences in relative size of females and males may also affect beetle behavior.
For example, Female 3 was larger than her male mate and could effectively thwart his
mating attempts. She allocated more time to ovipostion than Female 1, who was smaller
than her mate. Female 1 allocated more time to walking, being stationary, and twig
feeding than to copulation, egg niches excavation, and oviposition combined, possibly
because of male interference. Female 2 was intermediate in size and exhibited fewer
behaviors than Female 3, but spent more time in each behavioral state. Here, the males
seemed to interfere with egg niche excavation by trying to initiate further copulations as
was found for M. scutellatus (Peddle 2000) but not for A. chinensis (Wang et al. 1996).

Anoplophora chinensis females consistently allocated separate periods of the day
to copulation and to egg laying and females weren’t receptive to males while laying eggs
(Wang et al. 1996). However, A. chinensis females mate in the tree canopy and move
down the stem for oviposition (Wang et al. 1996). In contrast, for Monochamus
scutellatus, another Lamiinae species, activity peaks were not seen for mating and
oviposition (Hughes 1979). Our females were confined in the olfactometer and could not
easily escape mating attempts by males unless they were large enough to physically
thwart them. In the US, males and females are typically seen feeding in tree canopies

. (Haack et al. 1997) but the extent of the role of males in oviposition is unkown. It would

22

be interesting to conduct further behavioral bioassays in a field setting or at least in larger
areas with bigger logs to quantify separations of mating and oviposition activity peaks.
Future studies of A. glabripermis ovipostion and mating behavior should control for
differences in body size. Wang et al. and Smith et al. are currently researching the above
topics with populations in China.

During the 50 min bioassays, females spent a large percentage of their time in
escape behavior while in the olfactometer and the 50 min may not have provided
adequate time to assess the second level host selection process. These beetles showed a
long acclimation period, sometimes remaining stationary for the duration of the test.
Directed movement was observed in two cases but mostly females entered arms and
seemed randomly locate the wood sections that were suspended in the olfactometer arms.
This suggests that A. glabripennis females may use visual as well as olfactory cues to
locate host plants during the first stage of the host selection process. Even though cells of
freshly cut wood remain attractive to primary wood-boring pests for a few days (Hanks
1999), virgin A. glabripenm’s females may need the presence of live foliage and a full
canOpy to readily elicit a feeding response, as with A. chinensis (Wang et al. 1996).
Additionally, twig feeding, egg niche excavation and oviposition occurred on sugar
maple twigs but no twig feeding occurred for alternate species and egg niche excavation
only occurred on the oak wood sections. In the presence of sugar maple, or other highly
preferred hosts, females may not attack less preferred hosts such as tulip poplar and
honeylocust in the field (Nowak et a1. 2000).

Host preference studies for A. glabripenm’s would be more realistic in a large

room or greenhouse where arrays of logs or young trees of test species would allow

23

females to fly and use both vision and olfactory cues to locate preferred hosts. Field
studies of host preference using North American tree species in China would also be
valuable. Host preference field studies have been conducted for other Lamiinae species
such as Monochamus scutellatus, Monochamus carolinensis (Olivier), and the
Phoracantha semipunctata (Hanks et al. 1993, Lazarus and McCullough in preparation,
Peddle 2000, Walsh and Linit 1984).

Based on the results from the 1999 and 2000 host suitability tests, it appears that
hophombeam, basswood, northern red oak and white oak could be acceptable hosts for
oviposition by A. glabripennis beetles, especially in situations where more preferred
hosts are not available. Development to the first instar suggests that four of the alternate
hosts tested may be at least somewhat suitable. In Lepidoptera, the major behavioral or
toxicological barrier to favorable host suitability is determined by the first instar (Scriber
1984). Females also laid a few eggs on honeylocust, ironwood, and hackberry while
females did not lay eggs on eastern cottonwood, sycamore and tulip poplar. The lack of
oviposition on eastern cottonwood is interesting given that some types of hybrid poplars
in China are readily attacked by A. glabripennis and are suspected of being the
dissemination source for A. glabripennis throughout the world in the form of solid wood
packing materials (Smith et al. 2001). Recently, egg niches were found on species in the
same genera as sycamore (Plantanus) and eastern cottonwood (Populus) trees in New
York (Nowak et al. 2001). The lack of eggs on sycamore is also notable because species
of Plantanus are preferred hosts in China (He and Huang 1993, Li and Wu 1993, Li et al.
1999) and egg niches were found on trees in New York (Nowak et a1 2001).

Additionally, females excavated egg niches on oak trees in New York and females

24

excavated egg niches on basswood in Chicago (N owak et al. 2001), but no exit holes
were found even though species in the same genus as basswood are considered to be
preferred hosts in China (He and Huang 1993, Li and Wu 1993, Li et al. 1999).

The suitability of sycamore as a larval host in the US. remains questionable
because no larvae developed in our tests or on trees in New York. Since larvae
developed to the first instar on small branch sections of white oak and Northern red oak,
further testing on live trees is needed to quantify development rates of later instars.
Moreover, larvae developed on Northern red oak and green ash for up to 90 days on
potted trees in a greenhouse (Ludwig et al. 2002). Females were able to oviposit on
seven tree species previously undocumented as susceptible hosts in the US. Likewise,
females oviposited on three species with no current records of oviposition in New York
and Chicago. This suggests that females may find many more hosts suitable for
oviposition than for larval development. Perhaps A. glabripenm's larvae have not yet had
sufficient time to acquire mechanisms to adapt to novel host chemicals before discovery
in 1996 and 1998. Host suitability classifications may change in subsequent generations
as has occurred with other insects (Thompson 198 8).

Our results should be considered preliminary, given that relatively few beetles
were available for our bioassays. Additional research is needed to further assess how A.
glabripennis responds to other species in genera where larvae developed to the first
instar. Studies are also needed to evaluate whether the response and behavior of female
beetles encountering live trees is similar to their response to cut sections of wood that we

used in this study.

25

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A. glabripennls females

 

Female 3

 

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Figure 1.2. Total percentage of time spent in specific behavioral states by three A.

glabripennis females over a 24 h period.

30

 

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Figure 1.3. Percentage of time spent by three A. glabripennis females in specific behaviors

during 3 hour increments of 24 h bioassays.

31

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Figurel.4. Duration and frequency (:leEM) of behavioral states for three A. glabripennis
females in an olfactometer over a 24 hour period.

32

 

 

 

Mean % time (min)
01
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Central Arena Sugar Maple Alternate Host

Choice level

Figure 1.5. Mean percentage of time (d: SEM) that A. glabripennis females spent in the
central arena, in an olfactometer baited with sugar maple or in the olfactometer arms that

were baited with one of the alternate tree species. Significant differences are designated
by different letters, 11 = 92.

33

CHAPTER 2

Host Preference and Larval Performance Of Monochamus scutellatus (Say)
(Coleoptera: Cerambycidae) and Competition for Phloem on Four Conifer Species

INTRODUCTION

The relationship between host preference of adult insects and suitability of hosts
for larval development is essential in understanding evolution of host plant associations

in herbivorous insects (F utuyma and Peterson 1985, Thompson 1988). Presumably,

 

adults females confronted with a variety of potential host plants should choose the one
that would confer increased fitness to their progeny. In some populations of insects,
however, host preference of females is associated with varying degrees of performance
by their progeny. Thompson (1988) concluded that adult host preference does not always
correlate with larval performance because some females oviposit on hosts unsuitable for
larval development. Characteristics used to evaluate larval performance on potential
hosts may include survival of immature stages, growth rate, pupal weight and the
subsequent adult fecundity and longevity. These characteristics are not always positively
correlated with each other, however, and evaluations of host suitability may be different
depending on the criteria of performance (Thompson 1988, Via 1986).

Oviposition choices by female wood-boring insects are critical because larvae
cannot disperse to alternate or more suitable hosts because most are legless (Hanks et al.
1993). Hosts of wood-boring beetles are ofien patchily distributed within forested areas,
and wood-borers must rely on olfaction to locate potential hosts (Haack and Slansky

1987).

34

The white-spotted sawyer, Monochamus scutellatus (Say) (Coleoptera:
Cerambycidae) (subfamily: Lamiinae), is a common insect throughout Michigan
(Parmalee 1941) and much of the northern US. and Canada (Rose 1957). Monochamus
scutellatus is an important insect in Michigan conifer forests from both an ecological and
economic perspective. Adults are thought to be polyphagous and larvae of M. scutellatus
can develop in a wide range of conifer hosts throughout its range, including species of
pine (Pinus), spruce (Picea), balsam fir (Abies balsamea (L.) Mill), Douglas-fir
(Pseudotsuga menziesii (Mirv.) Franco) and eastern larch (Larix laricina (Du Roi) K.
Koch). Monochamus scutellatus is a cosmopolitan species found transcontinentally from
Alaska throughout Canada (and the Northern United States) and southward to North
Carolina in the east and New Mexico in the west. There are five species of Monochamus
found in Michigan (Yanega 1996). Tunnels excavated by larvae of species of
Monochamus contribute to decomposition and nutrient cycling, but can also damage logs
that would otherwise be salvaged (Edmonds and Eglitis 1989, Rose 1957, Cerezcke 1975,
and Post and Werner 1988).

The life history of M. scutellatus has been studied by various researchers (Belyea
1952, Chenier and Philogene 1989, Dyer and Seabrook 1978, Hanks 1999, Hughes and
Hughes 1982, 1985 and Rose 195 7). Monochamus scutellatus adults require a 3-7 day
period of maturation feeding on conifer shoots before eggs mature (Rose 1957). Adults
are not known to produce long range attractants or sex pheromones to locate hosts or
mates, rather, they rely on olfactory detection of volatiles and individual tree
characteristics to locate hosts (Dyer and Seabrook 1978). Monochamus scurellatus

employs a resource-defense polygyny mating system whereby males compete for access

35

 

to the regions of the trees or logs most attractive to females (Hughes and Hughes 1982,
1985). In Michigan, adult beetles typically colonize stressed or dying trees and recently
cut or fallen logs from late May through early August (Dyer and Seabrook 1978). Male
and females arrive on the potential host and may be observed persistently palpitating the
host surface as they walk (Hughes and Hughes 1985, Peddle 2000). Males are thought to
use their long antennae to aide in mate location and will mate with receptive females after
antenna] contact (Hughes 1979, Hughes and Hughes 1982). After mating, females cut

egg niches into the bark, often near old branch stubs or around branch whorls (Hughes

 

and Hughes 1985, Peddle 2000, Rose 1957). Eggs hatch within two weeks and larvae
begin tunneling in the cambium and phloem (Rose 1957). After three to four weeks,
larval galleries are wide enough and deep enough to score the sapwood. By late summer,
larval tunnels extend deeply into the sapwood, where larvae overwinter. Feeding
resumes the following spring and in Michigan, most beetles complete development and
emerge in late May to August. Adult M. scutellatus are most active in sunny locations in
the forest (Post and Werner 1988, Rose 1957).

Potential host material for M. scutellatus is often highly localized, resulting from
windstorrns or logging events, or sporadic individual tree mortality. Monochamus
scutellatus beetles must be able to locate potential hosts, often from a distance when
walking or in flight, and secondarily decide if the material is a suitable host after landing
(Bemays and Chapman 1998). Monochamus scutellatus responds strongly to host-
specific volatiles from a distance (Peddle 2000), but whether females discriminate among
potentially acceptable hosts or consistently respond differently to pine and spruce species

is unknown. Female M. scutellatus arriving on potential hosts for oviposition use contact

36

chemoreception to detect host semiochemicals, terpenes, resin pressure, and moisture
levels (Bemays and Chapman 1994, Brattli et al. 1998, Dyer and Seabrook 1977, Hanks
1999, Hughes and Hughs 1992). The presence of eggs or larvae of conspecifics can also
influence oviposition choices of M. scutellatus by deterring female oviposition on host
material already occupied by eggs of conspecifics (Hughes and Hughes 1992, Peddle
2002).

After oviposition, survival of offspring is driven by host suitability rather than by
host preference, since larvae must feed and develop on the host chosen by the female
(Hanks 1999). In the most basic sense, a host can be considered to be suitable if the
larvae survive and develop into adults. However, for polyphagous species such as M.
scutellatus, host quality may vary. Survival to the adult stage is one measure of
suitability, but vigor and size of beetles has been shown to be correlated with fitness
(Hanks 1999, Kato et al. 2000). Therefore, large and vigorous progeny may indicate a
higher degree of suitability for M. scutellatus larval development. Adult preference was
correlated to performance for a leaf-mining fly (Via 1986). Host preferences did not
match up exactly with larval performance for many other insects (Cronin et al. 2001,
Karban and Courtney 1987, Larsson et al. 1995).

Relationships between preference and performance have rarely been studied in
wood-boring Cerambycidae (Hanks et a1 1995). Host preference studies with wood-
borers usually involve counting the number of egg niches (Walsh and Linit 1985), and
eggs (Donley 1978) or twig feeding during choice tests (Walsh and Linit 1984,
Kobaayashi et al. 1984). In the only evaluation of M. scutellatus host preference, Peddle

(2000) found that females chewed more niches on red pine than on white pine logs.

37

 

Studies with other wood-boring species on multiple hosts reported differences among
larval density (Hanks et al. 1993), development time (Hanks et al. 1995), abundance of
galleries and mortality (Kobayashi et al 1984), but suitability of potentially acceptable
conifer species has not been evaluated for M. scutellatus.

Competition and cannibalism may be an important source of mortality for M.
scutellatus and for other insects that utilize phloem. Early instar M. scutellatus are

cannibalistic at high densities (Peddle 2002, Rose 1957). When two or more species

 

occupy the same host species, some degree of resource partitioning often occurs. For

example, bark beetles, weevils and M. scutellatus all occupy phloem but partition

themselves phenologically or spatially on the host (Kennedy and McCullough 2002), and
scolytid bark beetles partitioned themselves spatially on hosts in British Colombia
(Poland and Borden 1994).

I postulated that female M. scutellatus adults would consistently exhibit
preferences when four potentially acceptable hosts were available for oviposition. I
further postulated that female host preference, or host selection behavior, would affect the
development and survival of all progeny and that the four conifers would differ in
suitability for M. scutellatus development. Additionally, I postulated that phloem feeders
would partition themselves spatially on the logs to minimize mortality from interspecific
competition. Jack pine, red pine, white pine and white spruce were selected as potential
hosts in this study because they are native conifers within the host range of M.
scutellatus, their attractiveness and suitability for M. scutellatus is unknown, with respect
to each other, and they were available at the study site. The specific objectives of this

study were to evaluate 1) host selection preference and behavior by M. scutellatus

38

females in the field; 2) the survival and development of M. scutellatus larvae on four
conifers; and 3) the impacts of intra- and inter-specific competition on M. scutellatus

larval development.

39

 

METHODS AND MATERIALS

Study sites: In May 2001, one compartment each of red pine (Pinus resinosa
Ait.), jack pine (Pinus banksiana Lamb), Norway spruce (Picea abies (L.) Karst) and
white spruce (Picea glauca (Moench) Voss) were selected in Michigan State University’s
W. K. Kellogg Experimental Forest in Kalamazoo County. Compartments were 0.3 ha F,
(0.8 acres), 2.8 ha (7 acres), 0.4 ha (1 acre) and 0.4 ha (1 acre) in size for jack pine, red
pine, Norway spruce and white spruce, respectively (Greg Kowalewski, pers. comm.)

Each stand had coarse woody debris from logging and windthrow with recent M.

 

scutellatus frass piles and/or entrance holes into the sapwood. Each compartment had *
gaps in the canOpy where overstory trees had been felled in previous years.
Compartments were even-aged and shrub and forb layers varied but mainly included
European buckthom (Rhamnus cathartica L.), glossy buckthom (Rhamnusfrangula L.),
black cherry (Prunus serotina L.), sugar maple (A cer saccharum Marsh), and sassafrass
(Sassafrass albidum (Nutt.) Ness).

Site characteristics varied among compartments. All stands had a medium site
index of 51-56, based on site index charts for Michigan pine and spruce stands, and so
were of similar quality for tree growth (Avery and Burkhart 1983) (Table l). Basal area
of the compartments, measured with a 10 BAF prism, ranged from 36 mz/ha (155
ftZ/acre) for jack pine to 40 mz/ha (175 ftz/acre) for Norway spruce (Table 1) (Greg
Kowalewski pers. comm). Tree height ranged from 17.8 m (53.5 ft) (jack pine
compartment) to 21.5 m (64.6 ft) (Norway spruce compartment).

Eight trees of each species, jack pine, red pine, white pine and white spruce, at

WK. Kellogg Forest were felled 1-3 May 2001 and the stems were cut into logs, 46 cm

40

(18 in) in length and 10 to 18 cm in diameter. Two log arrays were centered under two
canopy gaps in each compartment. Each log array consisted of logs from two trees each
of jack pine, red pine, white pine, and white spruce, placed end to end by increasing size
in the north and south sides of each compartment under gaps in the canopy. Each
essentially reassembled tree was placed parallel to logs of the next species, 3.3 m (10 it)
apart. I placed from 44 to 62, 59 to 90, 49 to 63, and 45 to 65 jack pine, red pine, white
pine and white spruce logs in compartments, respectively (Table 2.1). The order that
species were placed in each of the two arrays in each stand was randomized. The canopy

cover directly over the log array was measured once at mid-day with a convex spherical

 

13——

crown densiometer, and showed that sun exposure on all log arrays was consistent
because the surrounding dominant trees provided similar amounts of shade in all
compartments.
Host preference of adult females

Field observations: Compartments were visited at least three times each week
from 22 May to 14 August 2001. Each log array was observed for 25 min, four times a
day between 10 am. to 6 pm. Compartments were visited sequentially, with the initial
stand determined by random draw. Our observations involved carefully walking along
the sides of the logs and inspecting all areas, including shaded areas adjacent to the
ground, without disturbing the logs. Observations were made for log species
sequentially. We used focal sampling within the observational period for each
compartment location, so that the first M. scutellatus seen was observed for the full 25
(Martin and Bateson 1993). If the first M. scutellatus observed flew away or fell into the

leaf litter before the 25 min observational period was finished, observations for M.

41

scutellatus activity on the other logs were continued. The number of male and female M.
scutellatus seen on logs was quantified. The frequency and duration of behaviors of
interest including walking, stationary, egg niche excavation, oviposition, feeding and
copulation was also quantified.

Log Retrieval and dissections. I assessed host preference of adult M. scutellatus

I V VH1

females by quantifying the attack rates on each conifer species. The top of each log was
marked with paint before it was collected from the field sites. From 25 to 31 August
2001, every third log in each log array was collected and transported to the laboratory at

Michigan State University. The remaining logs were transported to a screened enclosure

 

on campus that exposed logs to ambient conditions but not precipitation. Logs were
stored over the winter to allow adults to develop and emerge the following spring. The
first third of the logs were brought directly to the laboratory were designated as Batch-1.
The other two-third of the logs that were stored overwinter were designated as Batch—2.
Batch-l logs were dissected in September 2001 to January 2002. Batch-2 logs were
dissected from April to June 2002. Female M. scutellatus host preference was again
assessed by counting attacks on Batch-1 and Batch-2 logs combined. Suitability for
larvae, however, was measured separately for Batch-l and Batch-2. Because logs
remained in the forest for four months, egg niches could not be accurately counted due to
natural weathering of the bark. Instead, host preference using the number of galleries per
log was estimated. While some eggs initially laid on hosts may not have hatched, we
assumed that this rate would be similar among species. The number of M. scutellatus

galleries found on each region of the log (top, left side, right side, and bottom) was

42

quantified. Galleries that covered more than one position were assigned to the one where
the majority of the gallery occurred.

Physical characteristics that could affect M. scutellatus host preference were
assessed for each log. On each log two length measurements, three diameter
measurements and four bark thickness measurements were made (measurements were
made on only the outer bark at the end of the study) and the mean of each variable for
each log was calculated. The number of curculionid and scolytid bark beetle exit holes
(distinguishable by size) was recorded on each log. Each log was debarked and a grid
system was used to quantify the number of M. scutellatus galleries and the area of larval
feeding on each log. A gridwriter apparatus was used to accurately draw seven
equidistant lines, 4.5 cm (1.75 in) apart, around the diameter of each log. Each log was
marked four times lengthwise corresponding to the t0p, sides and bottom of the log as it
lay during the field study, to assess the effect of microhabitat on M. scutellatus
oviposition site selection.

Host suitability

Log retrieval and dissections: I measured 72, 97, 72, 71 jack pine, red pine, white
pine and white spruce logs were measured, respectively in Batch-1. For Batch-2 logs
118, 147, 131, and 115 jack pine, red pine, white pine and white spruce logs were
measured, respectively. Host suitability was estimated by counting larval gallery area,
number of entrance holes and percentage early instar survival for both Batch-l and
Batch-2 logs. Suitability was also measured by counting the number of exit holes and
total percentage survival for only Batch-2 logs. The total area phloem consumed by M.

scutellatus larvae for each log was recorded by summing the percentage of area

43

consumed within each grid-square and then multiplying that by the surface area of each
log. Early instar survival was assessed by counting the number of galleries with no larva
or entrance hole into the sapwood (survival=# larvae that died prior to the third
instar)/total number of galleries. The number of entrance holes refers to the oval-shaped
tunnel larvae make as they chew into the sapwood at an angle and can be used to estimate
whether larvae survival to the fourth instar (Cerezke 1977). The exit hole refers to the
circular-shaped hole that is created as adults chew out of the sapwood during emergence.
Suitability was also measured by overall percentage survival of M. scutellatus, calculated
as the number of emerging progeny/the total galleries per log.

Rearing F1 generation from Batch-2 logs. Batch-2 logs were placed in
individual cardboard rearing containers consisting of a cardboard tube (4, 6, 8, and 10 in
diameter, 0.32 —0.64 cm wall, 71 cm overall length (Michigan Can and Tube, Inc.,
Saginaw, MD) with screened endcaps on both ends and left undisturbed until spring
2002. I removed 200 logs from the screened enclosure and transported them to a 4.4C
(40F) cold storage room on MSU campus in March 2002.

Groups of logs (120-200 logs) representing all log species from each
compartment were brought into the laboratory to rear out M. scutellatus on four separate
occasions; 11 April, 2 May, 21 May, and 10 June 2002. Logs were soaked in water for 1-
3 minutes until bark was thoroughly moistened to hasten F1 emergence. Logs were then
placed into 121.1 L (32 Gallon) Rubbermaid® trash cans in which the plastic covering
the tops of the trash can lids was cut out and replaced with fiberglass mesh screen so that
newly emerged beetles would crawl up to the light. In this way, emerging M. scutellatus

could be easily collected from the lid.

Adult M. scutellatus were collected from the rearing containers twice daily
beginning 23 April, 2002. Newly emerged beetles were weighed, assigned an
identification number and placed individually into a 1.9 L (2 qt) plastic container with a
screened lid. Every time a new M. scutellatus was discovered in a trash can, its
corresponding exit hole was found and the number of the log it emerged from was
recorded. Adult M. scutellatus were provided with fresh Scotch pine (Firms sylvestris L.)
shoots in 2 dram glass vials filled with water weekly. Scotch pine was fed to the progeny
because it was available and was not one of the conifers focused on in the experiments.
Intra- and Interspecific Competition

I recorded the number of times M. scutellatus larval galleries intersected each
other on each log as an estimate of potential intraspecific competition. Galleries were
traced with a red wax pen to assist in accurate visualization of the interceptions. To
assess inter-specific competition for phloem between M. scutellatus and scolytid bark
beetles or curculionids, I traced scolytid bark beetle egg galleries and curculionid
cocoons and feeding galleries. The number of times a M. scutellatus gallery intercepted a
scolytid bark beetle gallery or curculionid gallery or cocoon was recorded. The area of
phloem consumed by scolytids and curculionids for the top, sides and bottom of each log
was quantified using the grid method as previously described.

Statistical Analyses

All variables were tested for normality with the Shapiro-Wilk test (Shapiro and
Wilk 1965). Variables were non-normal even after transformations, except for the area of
phloem consumed by scolydid bark beetles per log and M. scutellatus progeny weight.

Differences in initial number of logs colonized, and the number of logs where at least one

45

M. scutellatus emerged per species were tested using the Mantel-Haenszel Chi-square
statistic in a 2 X 4 contingency table (Rayner and Best 2001, Stokes et al. 2000). The
Nonparametric Rank F test was used to determine if number of galleries differed among
the 4 conifers (N eter et al. 1996). The Nonparametric Rank F test was used to determine
if the number of entrance holes per log, area of phloem consumed per log, and percent
early instar mortality differed for Batch-1 and 2 logs. Additionally, the Nonparametric
Rank F test was used to test for differences in number of exit holes and total percentage
M. scutellatus mortality among conifers for Batch-2 logs. I could not test for an
interaction between conifer species and log position because area of M. scutellatus
phloem consumption was not normally distributed and violated assumptions of a split-
plot design. Analyses of differences in the area of phloem consumption on the four
conifers and of area of phloem consumed on the tops, sides and bottom of logs were
conducted at the P<0.025 level of significance (after Bonferroni adjustment) using the
Nonparametric F -test (Kuehl 2000). One-way analysis of variance (AN OVA), was used
to determine if progeny weight differed among conifer species (SAS Institute, Inc. 1989).
When results of the Nonparametric F-test or AN OVA were significant, a Nonparametric
Rank F test Multiple Pairwise Testing Procedure and Fischer’s protected least significant
difference (LSD) test was used to determine which conifer species differed, respectively
(N eter et al. 1996). The nonparametric Mann-Whitney test was used to determine
differences in area of phloem consumed by scolytid bark beetles and curculionids
between logs with and logs without M. scutellatus galleries (Mann and Witney 1947).
Spearman’s rank correlation (R5) was used to test for associations among log diameter,

bark thickness and the number of M. scutellatus galleries for the 4 conifers species.

46

 

Spearman’s Rank Correlation was also used to test for associations between M.
scutellatus and scolytid bark beetle phloem consumption and M. scutellatus and
Curculionids phloem consumption. Unless specified differently, analyses were
conducted at the P<0.05 level of significance, using SAS statistical software (SAS

Institute, Inc. 1989).

 

 

47

RESULTS
Host Preference

Observations of M. scutellatus at Kellogg Forest: A total of 16 female M.
scutellatus and 17 male M. scutellatus was observed from 22 May to 14 August 2001 to
identify behaviors associated with selection of oviposition sites. Figure 2.1 shows the
frequency of the observed behaviors. Females oviposited on red pine, white pine, and
white spruce logs, for an average of 12 it out of the 25 min observation period. Females
oviposited on the sides of the logs in three of the four ovipositional events and once on
the top of the log. Females excavated six egg niches and three were on white pine.
Females were observed in a stationary state six times; on the top and the sides of red pine,
white pine and white spruce logs. A female walked only once on a white pine log and
pair of M. scutellatus copulated one time on a white pine log.

An interesting behavioral event observed was an escape behavior designated as
dropping. Because beetles had to be approached very closely to monitor behavior, a
shadow or noise was often unavoidable. On two occasions, females dropped from a red
pine log to the leaf litter where they were hidden from sight. Monochamus scutellatus
“squeaked” as they dropped. Three different females flew from logs of red pine, white
pine and white spruce when approached too closely.

Male M. scutellatus were stationary during most of our observations. Two
stationary males were observed on red pine, ten males on white pine logs, one male on a
jack pine log, and one on a white spruce log. On average, males were stationary 20.2 i
3.5 min of the 25 min observation period. Males walked on red pine logs twice and once

on a white spruce log. A male was also observed feeding on the side of a red pine log

48

and one male fed on the top of a white pine log. Males were not observed in flight but
two males dropped from red pine logs and one male from a white pine log and they
squeaked as they dropped.

Two related species, M. notatus (Drury) and M. titillator (Fabricius), were
observed on log arrays on a few occasions. One female M. titillator walked on red pine,
one female chewed an egg niche on white pine and two stationary females were observed
on white pine logs. One stationary male M. notatus was seen on a white pine log.

Overall, 69.1% of experimental logs were colonized by M. scutellatus. Female

M. scutellatus colonized 64%, 67%, 68% and 76% of jack pine, red pine, white pine, and

I-u luv.‘-—-I._‘4.i-u ‘ . A:

white spruce logs, respectively. A significantly different percentage of logs were
colonized by M. scutellatus females (X2=24.26, P<0.001, d.f.=3). Only logs with M.
scutellatus attacks were included in further analyses of variables related to galleries and
survival. Host preference is represented in Table 2.2 by the number of galleries per log.
For Batch-l logs, the number of M. scutellatus galleries per log ranged from 1 to 11 for
jack pine, l to 23 for red pine, l to 17 for white pine and l to 21 for white spruce logs.
There were significantly more galleries on white pine and white spruce logs than on jack
pine and red pine logs (P=0.001, F=5.72, d.f.=3) (Table 2.2).

Results of correlation tests comparing associations between variables related to
log characteristics and the number of M. scutellatus galleries differed. Log diameter was
not significantly correlated to number of galleries when all logs across species were
pooled together (P=0.09, R,=-0.07). Log diameter was significantly correlated to the
number of M. scutellatus galleries when jack pine logs (P=0.04, R,=0.19) and white pine

logs (P=0.007, R,=-0.23) were analyzed separately, but not for red pine and white spruce

49

logs. However, bark thickness was negatively correlated with the number of galleries but
explained little of the variation (P=0.003, Rs=-0.15) when species were pooled.
Additionally, bark thickness was weakly and negatively correlated to the number of M.
scutellatus galleries for red pine logs (P=0.006, R,=-0.22), and white pine logs (P=0.004,
R,=-0.27) when conifer species were analyzed separately.
Host suitability

Batch-I logs: Suitability of the 4 conifer species for larval development was
assessed by the area of phloem consumed per gallery and per log, percentage early instar
survival, and progeny weight. On average, M. scutellatus larvae consumed 8.4 i 0.9, 9.9
:i: 0.9, 12.7 i 1.3 and 10.8 i 1.6 percent of the phloem on the Batch-l jack pine, red pine,
white pine, and white spruce logs, respectively (Tables 2.2, 2.3 and Figure 2.2). For
Batch-1 logs, dissected in fall 2001, the total area of phloem consumed per log ranged
from a low value of 2.1 cm2 on a jack pine log, to a high of 603.2 cm2 on a white pine
log. The area of phloem consumed per gallery ranged from minimum values of 2.1 (jack
pine) to 5.4 cm2 (red pine) while maximum values ranged from 71.9 cm2 (white spruce)
to 80.4 cm2 (red pine). However, for Batch-l logs, the log species did not significantly
influence M. scutellatus phloem consumption (P<0.254, F=15.74, d.f=3) (Table 2.2). but
significantly more phloem was consumed on the sides of logs than on the tops and
bottoms (P=0.003, F=6.88, d.f.=3) (Figure 2.3). Monochamus scutellatus consumed 15,
13, 14 percent of phloem on the tops, sides and bottoms of logs. Again, the number of
measurements for log sides was double that of the top and bottom position. The number
of larvae that entered the sapwood ranged from 0 to 9 per log for jack pine, 0 to 21 for

red pine, 0 to 14 for white pine and 0 to 20 for white spruce and did not significantly

50

differ among species (Table 2.2). The percentage of larvae that entered into the sapwood
was 65%, 88%, 89%, 89% for jack pine, red pine, white pine and white spruce,
respectively. Early instar survival was generally high, averaging nearly 80 % for all
conifer species (Table 2.2).

Batch-2 logs: For the logs dissected after M. scutellatus beetles emerged, total
phloem consumption ranged from 3.1 to 342 cm2 for jack pine, 23.9 to 753.2 cm2 for red
pine, 24.5 to 571.9 cm2 for white pine, and 12.8 to 637.2 cm2 for jack pine logs. On
average, larvae on jack pine, red pine, white pine and white spruce consumed 9.1, 14.5,
18.7, and 14.0 percent of the phloem, respectively. However, M. scutellatus larval
galleries were significantly larger on red pine and white pine than on white spruce logs
(P<0.001, F=4.85, d.f.=3). Gallery size was statistically similar among the pines and
between jack pine and white spruce. The number of entrance holes per log where late
instar larvae entered the sapwood ranged from 0 to 7, 0 to 12, 0 to 17, and 0 to 19 for jack
pine, red pine, white pine, and white spruce per log, respectively and the number of
entrance holes were statistically similar among the four conifers. The percentage of
larvae that entered the sapwood was 95%, 96%, 98%, and 97% for jack pine, red pine,
white pine, and white spruce, respectively. Early instar survival and total survival both
ranged from 0 to 100 percent for all conifer species. Total percentage of survival was
significantly lower for the white spruce logs than for any of the pine species (P=0.04,
F=8.29, d.f.=3), but early instar survival did not differ. The maximum number of exit
holes per log where F1 adults emerged was 2, 6, 6, and 7 for jack pine, red pine, white
pine and white spruce logs, respectively, and differences among the conifer species were

not significant (Table 2.3). Monochamus scutellatus consumed 15, 16, and 13 percent of

51

phloem on the tops, sides and bottoms of logs, respectively. There was no significant
difference in the area of phloem consumed for the log position (P<0.296, F=1 .24, d.f.=3)
(Figure 2.3).

Progeny: F1 Monochamus spp. adults emerged from 57.4% of the jack pine
logs, 73.9 % of the red pine logs, 66.3 % of the white pine logs, and 64.8 % of the white

spruce logs. The number of logs with at least one emerging beetle did not vary among

__.— r.

the conifers (X2 =0.24, P>0.05) (Table 2.3). Beetles emerging from white spruce were
significantly smaller than beetles emerging from all three pine species (P<0.001,

F=47.89, d.f.=3) (Table 2.2). Female and male weights were very similar for beetles

 

reared on the same host (Table 2.3). The largest beetles reared from each conifer species
were typically the first of the group to emerge for both males and females. However, the
first beetles emerged from white spruce 5-7 days later than beetles from the pines. The
ratio of males to females differed among conifer species (Table 2.3), with 42%, 53%,
58%, and 55% females for jack pine, red pine, white pine and white spruce, respectively.
Compartment effects: We could not statistically analyze differences among
compartments because compartments were not replicated. Characteristics of
compartments, however, may have influenced the rate of their colonization by M.
scutellatus. The mean number of galleries per log was consistently lower for logs placed
in the white spruce study areas and much higher for logs placed in the red pine study
areas. The percentage of colonization of logs in compartments was 70.4%, 84.2%,
70.2%, and 53.1% for jack pine, red pine, Norway spruce and white spruce
compartments, respectively. The number of galleries and area of phloem consumed for

conifers of Batch-2 were compared. The range of galleries per log was 1-8, 2-5, 2-3, and

52

0.5-3 placed in jack pine, red pine, Norway spruce, and white spruce compartments,
respectively. Also, when we reared progeny, the mean number of exit holes per log was
lower for logs that were placed in the white spruce study areas.
Intra- and Interspecific Competition

Intraspecific competition: Potential competition among early instar M.
scutellatus larvae was assessed by counting the number of times galleries intercepted
other M. scutellatus galleries in Batch-1 logs. For Batch-1 logs, the nrunber of logs with
M. scutellatus gallery interceptions ranged from 18 % of j ack pine logs to 43 % of white
pine logs. There were no statistically significant differences among the conifer species
(Table 2.3). The number of interceptions between M. scutellatus galleries per logs
ranged from 1 to 3 (jack pine) and 1 and 20 (white spruce), but did not differ significantly
among conifers. For Batch-2 logs, the mean number of logs where M. scutellatus gallery
interceptions occurred ranged from 6 % of j ack pine logs to 37 % of red pine logs, but
differences among conifers were not significant (Table 2.3). For Batch-2 logs, the
number of interceptions between M. scutellatus and M. scutellatus galleries per log
ranged from 1 interception (jack pine) to 17 (white spruce), but the number of M.
scutellatus gallery interceptions was not significantly different among conifers.

Interspeciflc competition. Potential competition between M. scutellatus and
other phloem feeders varied among conifers. All Batch-1 logs were colonized by scolytid
bark beetles, while curculionids colonized 90%, 93%, 97%, 37% of the jack pine, red
pine, white pine and white spruce log, respectively. Scolytid bark beetles and/or
curculionids colonized 31, 20, 23, and 22 percent of jack pine, red pine, white pine and

white spruce logs, respectively, in the absence of M. scutellatus. Scolytid bark beetles

53

consumed significantly more phloem on logs where M. scutellatus were absent (P<0.05,
27.51, d.f.=3). The area of phloem consumed by scolytid bark beetles was greater on
jack pine and white spruce than on red and white pine logs (P<0.05, F =15.64, d.f.=3)
(Figure 2.2). Scolytid bark beetles consumed 8, 11, and 9 percent of the phloem on the
tops, sides and bottoms of logs, respectively, and they consumed more area of phloem on
the top and bottoms than on the sides of logs (p<0.003, F =6.88, d.f.=3) (Figure 2.3). The
number of interceptions between M. scutellatus and scolytid bark beetle galleries ranged
from 1 to 10 (red pine) and l to 21 (white pine and white spruce) and significantly more
interceptions were found on white pine and white spruce logs than on jack pine or red
pine logs (P<0.001, F=7.67, d.f.=3). Curculionids consumed more phloem on logs with
no M. scutellatus galleries than on logs with at least one M. scutellatus gallery (P<0.05).
Curculionids consumed a greater area of phloem on pines than on white spruce (P<0.001,
if. = 3) (Table 2.4) (Figure 2.2). Curculionids consumed 5, 7, and 11 percent of phloem
on the tops, sides and bottoms of logs, respectively, and they consumed more phloem on
the bottoms of logs than on the tops or sides (P<0.001, d.f.=3) (Figure 2.3). The number
of M. scutellatus and curculionid interceptions ranged from 1 to 9 (jack pine) and 1 to 22
(white pine), but differences were not significant among conifers.

All of the Batch-2 jack pine and white pine logs were colonized by scolytid bark
beetles, and they colonized 94% red pine logs and 97% white spruce logs. Curculionids
colonized 92% of the jack pine logs, 100% of the red pine, 94% of the white pine, but
only 25% of white spruce logs. Like the Batch-1 logs, significantly more phloem was
consumed by scolytid bark beetles on logs with no M. scutellatus galleries than on logs

with one or more cerambycid galleries (P<0.05). Significantly more phloem was

54

consumed by scolytid bark beetles on the white spruce logs than on pines (P=0.002,
F=41.58, d.f.=3) (Figures 2.2, 2.3). Scolytid bark beetles consumed 4, 3, and 3 percent of
phloem on the top, sides, and bottoms of logs. Scolytid bark beetles consumed more
phloem on the top than on the sides of logs, but area of phloem consumed on the bottom
did not differ from the tops and the sides (P=0.021, d.f.=3). The number of interceptions
of M. scutellatus and scolytid bark beetle galleries ranged from 1 to 4 (red pine) to l to
17 (white spruce) and significantly more interceptions were found on white spruce logs
than on jack pine or red pine logs (P<0.005, F=12.67, d.f.=3). Significantly more phloem
was consumed by curculionids on logs with no M. scutellatus galleries than on logs with
at least one gallery (P<0.05). Again, curculionids consumed significantly more phloem
on the pines than on white spruce (P<0.001, F=67.99, d.f.=3) and more phloem was
consumed on the bottom of logs than for other positions on the logs (P<0.001, F=13.81,
d.f.=3). Curculionids consumed 3, 5, and 8 percent of the phloem on the tops, sides and
bottoms of logs. The number of M. scutellatus and curculionid gallery interceptions
ranged from 1 (white spruce) and l to 24 (red pine), and significantly more interceptions
were found on red pine, white pine and jack pine than on white spruce (P<0.003, F=5.57,
d.f.=3).

Galleries of M. scutellatus intercepted galleries of scolytid bark beetles on 62 %
of jack pine logs and 86 % of white spruce logs. The mean number of M. scutellatus bark
beetle gallery interceptions was significantly greater on white pine and white spruce than
on red pine and jack pine (Table 2.4). Galleries of M. scutellatus intercepted weevil
galleries on 12 % of white spruce to 63 % of white pine logs. Differences among conifer

species were not significant (Table 2.3).

55

For Batch-2 logs, galleries of M. scutellatus intercepted galleries of scolytid bark
beetles on 18 % of jack pine to 76 % of white spruce logs. The mean number of M.
scutellatus bark beetle gallery interceptions was significantly greater on white pine and
white spruce than on red pine and jack pine (Table 2.4). Galleries of M. scutellatus
intercepted weevil galleries on 20 % of white spruce to 82 % of white pine logs. The
number of M. scutellatus-curculionid galleries was significantly greater on the pines than
on white spruce logs (Table 2.4).

For Batch-1 logs, pooled across all species, the area of phloem consumed by M.
scutellatus was negatively correlated to the area of scolytid bark beetle feeding (P<.001,
Rs=-O.22), but not to the area of curculionid feeding. Phloem consumed by M. scutellatus
was positively correlated to weevil feeding (P<0.004, R,=0.37) when jack pine was
analyzed separately. The area of M. scutellatus feeding was negatively correlated to
scolytid bark beetle feeding (P<0.003, Rs=-0.39) and weevil feeding (P<0.03, R,=-0.29).

For Batch-2, pooled across species, the area of phloem consumed by M.
scutellatus was not correlated to the area of scolytid bark beetle feeding, or to the area of
curculionid feeding. The area of M. scutellatus feeding wasn’t correlated to bark beetle
and weevil feeding on jack pine, red pine, white pine or white spruce when conifer

species were analyzed separately.

56

DISCUSSION

Host Preference and Suitability

I attempted to assess whether M. scutellatus females in southwestern Michigan
demonstrate a preference hierarchy for four conifer species within their host range.
Female M. scutellatus prefer white pine and white spruce for oviposition over jack pine
and red pine. This conclusion was based on different numbers of galleries, the index of
preference in this study, and on the percentage of logs M. scutellatus oviposited on for
each species. In general, fewer beetles were observed visiting log arrays than originally
expected, but most of the beetles observed during the summer were on white pine and red
pine logs. Adult M. scutellatus are active throughout much of the day and beetles were
seen at varying times during observation periods Hughes (1979) and Peddle (2000). One
reason few beetles were observed was probably because harvesting occurred in
compartments adjacent to red pine, white spruce and Norway spruce compartments
during most of June. Adult M. scutellatus were frequently seen in other logged
compartments. Some M. scutellatus may have been attracted to the logged compartments
and passed over the log arrays because of the amount of terpenes and other volatiles
emitted from the logged area.

All of the conifer species presented to females were accepted for oviposition by at
least some M. scutellatus females, but females colonized jack pine logs less frequently
than the other pine species and white spruce logs. Many logs were not colonized by M.
scutellatus perhaps because of the abundance of cut pine in adjacent and nearby

compartments of Kellogg Forest. However, more jack pine logs were left uncolonized

57

than for other conifer species. A greater number of galleries was found on white pine and
white spruce logs than jack pine and white spruce and so were initially more preferred
than red pine and jack pine. In a related study, during 2-choice trials, females preferred
either white pine or white spruce over jack pine and red pine consistently, while similar
numbers of eggs and galleries were on white pine and white spruce (Lazarus and
McCullough in preparation). Preference for white pine by M. scutellatus has been also
shown by Peddle (2000). In addition, females were attracted to white spruce decks in
Alberta causing much loss in merchantable timber (Cerezke 1977), and attacked downed
white spruce more often than jack pine in northern Ontario (Gardiner 1975).

In general, log diameter did not influence female preference. There was a slight
correlation between diameter and number of M. scutellatus galleries for jack pine and
white pine logs. The logs in this study were all probably large enough to support a year-
long life cycle, but I would expect greater mortality on smaller branch-size wood. The
diameter or height of trees did influence host preference of Monochamus sutor L. when
diameters ranged from 6 cm to 27 cm (Zhang et al. 1993). In this study, the number of
galleries was slightly negatively correlated to bark thickness of red pine and white pine.
Females may have specifically chosen a certain bark thickness for oviposition (Hughes
and Hughes 1982).

Once females selected a log for oviposition, relative suitability of the four
conifers influenced the fate of the F1 generation. Overall, white pine and red pine were
most suitable for M. scutellatus development and white spruce was least suitable for a
number of reasons. Early instar larvae utilized more total phloem on the Batch-l white

pine and white spruce logs but differences among species were not apparent for the

58

Batch-2 logs that were dissected after the F 1 adults emerged. A study in Canada found
that M. scutellatus galleries were larger on white pine than on red pine logs, but jack pine
and white spruce were not evaluated (Peddle 2002). Galleries on our jack pine, red pine
and white pine logs were consistently larger than those in the Peddle (2002) study,
although the galleries were significantly smaller on our white spruce logs. Log diameters
were similar in the two studies, but phloem thickness was not measured.

Additionally, M. scutellatus preferred the sides of logs for oviposition. Females
allocate much time and energy to egg niche excavation and oviposition and egg
placement influences progeny survival in this species (Hughes and Hughes 1982). The
sides of the logs may be optimal for oviposition because phloem on the tops of the logs
may dry out faster than other areas, and the bottom of logs on the ground may be too
moist and cool. Larvae consumed more phloem on the sides of the Batch-2 logs,
regardless of conifer species, although differences were not significant for Batch—l logs.
In Canada, M. scutellatus preferentially oviposited on the sides of logs in white spruce
log decks (Post and Werner 1988, Cerezke 1977), but there were no differences in
number of egg niches or phloem consumption between the top and bottom halves of logs
that were on the ground (Peddle 2000). Female beetles, however, may have been more
restricted in regions of the log available for oviposition in log decks than when logs were
layed singly on the ground. Raske (1975) found the density of M. scutellatus galleries
was twice as high on scattered logs than on decked logs.

Total survival is one of the key indicators of suitability in this study.
Nevertheless, the survival of M. scutellatus on white pine and white spruce was quite

different when we compare all life stages on these two hosts. There were no differences

59

among conifer species in early larval instar survival for either Batch-1 or Batch-2 logs.
Once larvae hatched and began feeding, they generally survived long enough to begin
tunneling into the sapwood In white spruce logs in Alberta there was 29% M. scutellatus
total survival (Cerezke 1977), while 31% survived on white spruce (Table 2.2). Total
percentage survival was almost double on pines than on spruce in our study. Differences
in survival between the period of late instar/pupation and emergence may have been due
to differences in chemical composition between pine and spruce logs, or simply that some
dietary needs of M. scutellatus were not met by white spruce prior to the period between
overwintering and pupation of the beetles. Pine and spruce contain varying
concentrations of monoterpenes such as alpha-pinene, camphene, beta-pinene, myrcene
and lirnonene (Drew and Pylant 1966).

The final measure of suitability among the four conifer species was obtained
when progeny emerged from the host logs. Adult female size directly affects fecundity
of cerambycid females and also affects fitness of the beetles (Hanks 1999, Larsson and
Kustvall 1990). Pines were more suitable hosts for M. scutellatus offspring because the
F1 adults were larger than those that developed in spruce. Spruce-reared beetles were
small, but a similar number of beetles emerged as for pines. Competition may also
influence progeny size, but size varied for progeny that developed on equally preffered
white pine and white spruce. However, F; beetles began emerging from spruce 5-7 days
after those from the pines began emerging which is suggestive of delayed development in
spruce logs, especially since all logs were treated similarly in the field.

It is more difficult to discern if the three pine species differed in suitability for M.

scutellatus development because the percentage of progeny survival was similar among

60

the three pines. Jack pine may be the least suitable pine species because females
colonized fewer jack pine logs than other pines, and progeny were significantly smaller
than those reared on red pine and white pine. Interestingly, more female beetles emerged
from red pine, white pine, and white spruce, while more males emerged from jack pine.
A sex ratio favoring males may have implications for the abundance of M. scutellatus in
future generations. Jack pine-reared females often died within a week after emergence
compared with a life span of up to 70 days on other species. This suggests that
individuals that developed on jack pine had lower vigor or were sick.

Thompson (1988) stated that an ovipositional choice that confers greatest fitness
to offspring would seem to be favored evolutionarily, but preference and performance
often are not correlated. This is probably especially true for insects that cannot change
hosts while developing. Host preferences did not match up exactly with larval
performance in this study, as was found for other insects (Cronin et al. 2001, Karban and
Courtney 1987, Larsson et a1. 1995, Via 1986). Female M. scutellatus did not
demonstrate a neat hierarchical ranking of preferences, or one host that was preferred
over all others consistently. If preference is correlated to performance, than progeny
developing on white pine and white spruce should have performed better then those on
red pine and white spruce. I expected a tighter linkage between preference and
performance for wood-borers because oviposition site-selection influences offspring
survival. Explanations for inconsistencies between female preference for white spruce
and poor offspring performance on white spruce are unknown. Inconsistencies between
preference and performance may be explained by ecological conditions affecting

populations such as abundance of hosts from year to year or enemy-free space (Cronin et

61

al. 2001, Hanks et a1. 1993, Thompson 1988). Kellogg Forest is composed of
compartments of a number of different conifer species with varying amounts of host
material from conifers available every year. Perhaps the inconsistencies between
preference and performance found among conifers are due to the variability of available
hosts for colonization each generation. However, the survival on all hosts of differential
chemical composition is suggestive of adaptation to a more generalized diet, as was
found in gall-making flies (Cronin et al. 2001). Future studies on correlation of
preference and performance of wood-borers should address the importance of host
abundance to host suitability and implications of small body size and delayed
development on the fitness of wood-borers.

Intra- and Interspecific Competition

I postulated that intraspecific competition would vary among the four conifers if
females preferred one species over others. High densities of larvae may provide more
opportunities for contact interference or cannibalism if phloem is limited. Intraspecific
competition did not vary among the host species, even though the number of galleries
was greater on white pine and white spruce than on jack pine and red pine logs. Many
logs had more than one interception per log, indicating that galleries may be concentrated
on certain positions of the log, or that some logs were more attractive than others for
oviposition leading to a higher probability of competition among larvae for phloem on
specific logs. Larval galleries often came within a few millimeters of each other but
didn’t intersect on white pine and red pine logs in Ontario (Peddle 2000). Intraspecific
competition resulted in 60 to 80% mortality in other studies (Belyea 1952, Kobayashi et

a1. 1984, Rose 1957), much higher values than the 4 to 18% larval mortality we recorded,

62

but there were more larvae in those studies than this one. Mortality estimates in this
study are relatively low, however, because first and second instar competition was not
directly measured. Rose (1957) noted that small, early instar galleries often coalesced
around branch whorls. Additionally, M. scutellatus females preferentially ovipositied on
logs with no eggs rather than on logs with her own eggs or eggs of other females fl’eddle
2002). Females may have perceived occupied phloem and avoided ovipositing in those

regions of the log.

Additionally, I attempted to assess the effects of other phloem feeders on M.
scutellatus oviposition and larval gallery distribution among the top, bottom and sides of
the conifer logs. Since some scolytid bark beetles, such as T omicus pim'perda and Ips
pini, and curculionids colonize logs earlier than M. scutellatus, they may affect the
ovipositional site selection by female M. scutellatus, perhaps causing females to partition
egg laying differently or to choose less occupied areas of phloem.

Monochamus scutellatus probably reduced the density of scolytid bark beetles or
may have inhibited scolytid bark beetle oviposition on red pine and white pine logs
through interference competition when individuals were abundant on the logs. Host
preference of scolytid bark beetles for certain pines or spruce increased the probability of
interference competition with M. scutellatus larvae. Gallery interceptions among M.
scutellatus and Scolytidae occurred more often on the sides of logs. When interferences
were found for species that consume phloem concurrently with M. scutellatus larvae,
bark beetle feeding would be altered as was the case for Dendroctonusfrontalis by
foraging M. titillator (Coulson et al. 1976, 1980, Miller 1985). Scolytid bark beetles

were most prevalent on white spruce and white pine and we saw more of these

63

interceptions on white pine, the favored host of M. scutellatus. Additionally, negative
correlations between M. scutellatus and bark beetle feeding suggest bark beetle feeding is
influenced by the presence of M. scutellatus. In the absence of M. scutellatus larvae,
scolytid phloem consumption per log was greater. In the presence of M. scutellatus, we
found that scolytid bark beetle phloem consumption was generally greater on the tops and
the bottom of logs. Since M. scutellatus galleries were found most often on the sides of
the logs, and scolytid bark beetles preferentially colonize sides of the logs (Haack and
Lawrence 1995a), scolytid larval feeding was likely deleteriously affected if larvae
actively fed at the same time as M. scutellatus larvae.

Monochamus scutellatus probably reduced the density of curculionids. Gallery
interceptions did not occur as often as with scolytid bark beetles. However, Curculionids
feeding was greater on the bottom of logs, with or without M. scutellatus. More M.
scutellatus feeding occurred on the side regions of logs, thereby indicating that these
phloem feeders partitioned themselves on the log. Weevils were found more often on
pine logs than on spruce in our study and there were more interceptions on white pine.
Pissodes approximatus pupal cells were found most often on the cool, moist bottom
region of white and red pine logs in Canada (Peddle 2000, Phillips et al. 1987).

It is unlikely that M. scutellatus females avoid other phloem feeders when
locating host material, but locating unoccupied phloem for oviposition is probably more
important to females. It is likely that M. scutellatus will intercept galleries of other
phloem feeders if they are abundant on the log. Monochamus scutellatus used bark
beetle pheromones as kairomones to locate hosts in British Columbia (Allison et a1.

2001), suggesting that M. scutellatus employs a competitive strategy for utilizing phloem.

64

However, the tendency of greater M. scutellatus larval feeding on sides of logs, compared
to weevil feeding on log bottoms indicates some partitioning. Progeny of 1. pint may
suffer increased mortality because they have three generations a year and M. scutellatus
larvae are feeding in phloem for much of the second and third generations of I. pini.
Scolytid bark beetles may employ a strategy of occupying as many regions of the log as
possible thereby combating M. scutellatus induced mortality. Pissodes approximatus
were pupating at the time of Batch-1' log dissections and so populations were likely
reduced by M. scutellatus foraging. Future field studies addressing intra and inter-
specific competition should involve the removal of logs in smaller batches every week or
two after the first M. scutellatus females have been observed in study areas to count the

number of egg niches, eggs and egg placement on logs just after hatch.

65

 

 

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- Egg niche excavation
Oviposition
4 . _ Copulation
3 E Walking
3 - Stationary
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Jack pine Red pine White pine White spruce

Log species

Figure 2.1. Frequency of behaviors associated with female M. scutellatus host selection
observed at Kellogg Forest from May 22 to August 14, 2001 between 10 am. to 6 pm.

70

350

 

 

 

 

 

 

     

 

 

 

 

 

 

 

       

 

"E - M. scutellatus Ban" 1 I095
3 300 1 Scolytidae a
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a
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Jack pine Red pine White pine White spruce

Conifer species

Figure 2.2. Mean area of phloem (:i: SEM) consumed by M. scutellatus, scolytid bark beetles, and
curculid weevils on Batch 1 logs that were dissected in October to December, 2001 and Batch 2
logs that were dissected in June and July, 2002. Only logs with M. scutellatus galleries were used
in analyses. Batch 1 included 55,76, 54, and 51 for jack pine, red pine, white pine and white
spruce logs. Batch 2 included 17, 24, 22, and 25 Jack pine, red pine, white pine, and white spruce
logs, respectively, in analyses. Significant differences among log species are indicated by

different letters above columns for each type of phloem feeder.

71

 

 

 

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Log position

Figure 2.3. Mean area of phloem consumed by M. scutellatus larvae, scolytid bark beetles, and
weevils on the top, sides, and bottom of logs. Batch-l logs were dissected September to
December 2001 and Batch-2 logs were dissected summer 2002. Significant differences among
log species are indicated by different letters above the column for each type of phloem feeder.

72

CHAPTER 3

Effects Of Host Preference On Larval Performance Of Monochamus scutellatus

(Say) (Coleoptera: Cerambycidae)

INTRODUCTION

Understanding the relationship between adult host preference and relative
performance of offspring is essential to assess the evolution of host plant associations in
herbivorous insects (Futuyma and Peterson 1985, Thompson 1988). Presumably, adult
females confronted with a variety of potential host plants should choose the host that will
maximize larval fitness. However, in some populations of insects, host preference of
ovipositing females does not always correlate with larval survival or successful
development (Thompson 1988). Poor correspondence between ovipositional preference
and offspring performance may indicate that factors such as host abundance (Williams
1983), predation rates on larvae (Demo and Larsson 1990), and intraspecific competition
(Anbutsu and Togashi 1997, Hanks et al 1993) influence larval fitness.

Oviposition choices by adult female wood-borers are critical for offspring
survival because larvae cannot disperse to alternate or more adequate hosts (Hanks et al.
1993). Additionally, hosts of wood-borin g beetles are ofien patchily distributed within
the forest resource (Haack and Slansky 1987) and beetles must largely rely on chemical

communication to detect by potential hosts.

73

The white-spotted sawyer, Monochamous scutellatus (Say) (Coleoptera:
Cerambycidae), is a common insect in conifer forests throughout Michigan (Parmalee
1941), and much of the northern US. and Canada (Rose 1957). Adults are thought to be
polyphagous and larvae of M. scutellatus can develop in a wide range of conifer hosts,
including pine (Pinus spp.), spruce (Picea spp.), balsam fir (Abies balsamea), Douglas-fir
(Pseudotsuga mensiesii) and eastern larch (Larix laricina). Tunnels excavated by M.
scutellatus larvae contribute to decomposition and nutrient cycling, but can also damage
logs that would otherwise be salvaged (Cerezcke 1975, Edmonds and Eglitis 1989, Post
and Werner 1988, Rose 1957).

The life history of M. scutellatus has been studied by numerous researchers
(Belyea 1952, Chenier and Philogene 1989, Dyer and Seabrook 1978, Hanks 1999,
Hughes and Hughes 1982, Hughes and Hughes 1985, Rose 1957). After emergence,
adult M. scutellatus females require a 3—7 day period of maturation feeding on conifer
hosts before eggs mature (Rose 1957). Adults of M. alternatus Hope are the only species
of Monochamus known to produce pheromones to locate hosts (Kim et al. 1992), and M.
scutellatus adults are thought to rely on olfactory detection of host volatiles and
individual tree characteristics to locate hosts (Dyer and Seabrook 1978). Monochamus
scutellatus employs a resource-defense polygyny mating system whereby males compete
for access to the regions of the trees or logs that are most attractive to females (Hughes
and Hughes 1982, Hughes and Hughes 1985). Adult beetles typically colonize stressed
or dying trees and recently cut or fallen logs from late May through early August in
Michigan (Dyer and Seabrook 1978). Male and female adults arrive on the potential host

and may be observed persistently palpitating the host surface as they walk (Hughes and

74

Hughes 1985, Peddle 2000). After mating, females cut egg niches into the bark often
near old branch stubs or around branch whorls (Hughes and Hughes 1985, Peddle 2000,
Rose 1957). Eggs hatch within two weeks and larvae begin tunneling in the cambium
and phloem (Rose 1957). After two to three weeks, larval galleries are wide enough and
deep enough to score the sapwood. By late summer, larval tunnels extend deeply into the
sapwood, where larvae overwinter. Feeding resumes the following spring and in
Michigan, most beetles complete development and emerge in the summer. Interspecific
competition and cannibalism may be an important source of mortality for M. scutellatus
and for other insects that utilize phloem (Peddle 2000, Rose 1957).

Studies of preference versus performance for polyphagous cerambycid species
and species that utilize dead or dying woody hosts are rare and mainly focus on either
adult preference or larval performance, but not both. Host preference studies for wood-
borers are usually limited to counting the number of egg niches (Walsh and Linit 1985)
or eggs (Donley 197 8) found on logs in adult choice tests. Monochamus carolinensis
was shown to prefer feeding on Scotch pine (Pinus sylvestris L.) and jack pine (Pinus
banksiana Lamb.) in the lab (Walsh and Linit 1984) and M. alternatus consistently chose
Pinus elliottii Engelm. over Pinus densiflora Sieb. and Zucc. during choice tests
(Kobayashi et al 1984). A recent study of host preference of M. scutellatus found more
egg niches on white pine than on red pine logs (Peddle 2000). Studies comparing larval
performance of woodborers among multiple hosts have typically assessed only
differences in larval density (Hanks et al. 1993), abundance of galleries and late-instar

mortality (Kobayashi et al 1984). Other studies of M. scutellatus focused on oviposition

75

behavior, host quality, fecundity, and survival rates but were restricted to only a single
conifer species (Cerezke 1977, Dyer and Seabrook 1978, Peddle 2000 and Rose 1957).

Hopkin’s host-selection principle refers to the adaptation of insects to their host
plants or the idea that variation in host plant selection can be found (Craighead 1921).
Larval conditioning is an idea based upon Hopkin’s host-selection principle and refers to
the idea that many adult insects prefer host species on which they themselves developed
as larvae (Barron 2001, Craighead 1921). Craighead (1921) concluded that a
conditioning response of larvae to hosts over several generations was common for most
species of cerambycidae. Larval conditioning did not occur after one generation when M.
carolinensis was reared on Scotch pine (Walsh and Linit 1984). Futuyma and Peterson
(1985) concluded that there is little or no evidence for larval conditioning in
phytophagous insects. However, it is not known for certain whether conditioning affects
host selection by adult M. scutellatus.

In this study 1 evaluated whether ovipositing adult females consistently preferred
one conifer over another in two-choice tests. I evaluated potential differences in
suitability for M. scutellatus larval development. I assessed whether larval conditioning
affects host selection by adult females. I also evaluated whether the host preference of
ovipositing adult females is consistent with suitability for larval development. Jack pine,
red pine, white pine and white spruce were selected as potential hosts in this study
because they are native conifers within the host range of M. scutellatus and their

attractiveness and suitability for M. scutellatus is unknown, with respect to each other.

76

METHODS AND MATERIALS

In 2001, preliminary host choice tests were conducted to develop methodologies
for 2002 tests at the Tree Research Center, MSU campus. We collected newly emerged
M. scutellatus beetles from a one-year old Norway spruce (Picea abies) log deck at
Michigan State University’s WK. Kellogg Forest (Kalamazoo Co., MI) and gave them in
a choice of two conifer species in 1.8 m high x 3.7 m long x 1.8 m wide, tents made of 1
mm Lumite amber screen (Synthetic Industries, Inc., Gainsville, GA). In each tent we
placed two logs, 45.7 cm (18 in) long, of each test species (four logs total) and 1 to 6
pairs of sawyers. The number of M. scutellatus added to each tent depended on the
number of beetles collected from the log deck at Kellogg Forest the previous day. Black
cherry (Prunus serotz'na), red pine (Pinus rubra), white pine (Pinus alba), jack pine
(Pinus banksiana) and white spruce logs (Picea glauca) were cut to 46 cm (18 in) lengths
from trees felled at Kellogg Forest biweekly, beginning 15 May, 2001. Two logs were
placed 1 m from the other two logs, and 1 m from each other in the middle of the tent.

Two-choice tests were conducted during 2002 to assess host preference of four
conifer species. Prior to the choice tests in 2002, jack pine, red pine, white pine, and
white spruce logs from 2001 field study were brought into the lab to rear out adult
sawyers in batches of 100 to 150 at a time. Adult sawyers were provided with fresh
Scotch pine shoots in water-filled 2 dram glass vials for maturation feeding and were
placed in 21°C environmental, Michigan State University campus.

Adult M. scutellatus began to emerge on 1 April, 2002 and progeny were
collected twice daily until emergence ceased. Newly emerged beetles were weighed,

assigned an identification number and placed in a 1.9 L (2 qt) plastic container with

77

screening covering the lid. Every time a new adult M. scutellatus emerged its
corresponding exit hole was located and the identity of the log it emerged from was
recorded. Scotch pine was cut from trees on the Michigan State University campus.
Logs of j ack pine, red pine, white pine, and white spruce were cut in March 2002 to 38
cm (15 in) lengths at Kellogg Forest, and transported to the Michigan State University
campus, East Lansing, and logs were stored outdoors in three 2.43 x 4.87 m (8 x 16 it)
screen tents placed in the shade to exclude other phloem feeding insects.

Three-day, two-choice host tests were conducted to assess oviposition

preferences of females. We placed sixteen 1 m3 Lumite screen tents 20 meters apart from

 

each other in a zigzag pattern in a field at the Tree Research Center, Michigan State
University campus. Jack pine, red pine, white pine and white spruce shoots were
collected from nearby trees. A total of two logs, each a different conifer species, was
placed in each tent along with a previously unmated pair of beetles reared from logs used
in 2001 field study. Beetle ages ranged from 7 to 24 days post emergence. To assess
whether the larval host affects host preference of adult M. scutellatus we placed one log
of the larval host species in one corner of the tent and a log of a different conifer species
in the opposite corner. One shoot, with the current and last years growth, of the conifers
being tested was placed in a water pic and placed each on top of the corresponding log
species. Beetles were left undisturbed for 3 days, then we collected the logs and the
beetles from each tent and retm'ned them to the laboratory.

The number of egg niches on each log was counted immediately after each 3-day
choice test was completed. We measured the diameter, length and bark thickness of each

log and stored them in the lab. Mean log diameter ranged from 14.6 cm (jack pine) to

78

16.8 cm (red pine) (Table 3.1). Thirty days after the logs were removed from the tents
we removed the bark of each log to quantify the number of eggs and larvae on each log.
Each larva was weighed and body length was measured with calipers.

Female oviposition preferences was assessed by determining the mean number of
egg niches and eggs laid by each female during the 3-day trial, and the number of larvae
and galleries per log. Eggs, larvae and galleries were counted during bark dissections.
Host suitability for larval development was assessed based on the weight and length of
larvae, the length and width of larval galleries, the number of entrance hole per log and
percentage survival. Differences in larval grth rate were estimated by comparing the
larval weight and the number of entrance holes into the sapwood after 30 days of
development, since it is typically the third instar that begins to tunnel in sapwood.
Percentage survival was assessed by counting the number of live larvae/number of dead
eggs and larvae.

Statistical analyses.

All variables were tested for normality with the Shapiro-Wilk test (Shapiro and
Wilk 1965). When we pooled the data by log species for the 3-day host preference and
host suitability trials, the number of egg niches, eggs, galleries, larvae, entrance holes,
gallery width, gallery length, and percent survival for 3-day tests were not normal even
after transformations. These variables were analyzed using the Nonparametric Rank F
test, and the Nonparametric Rank F test mean comparison procedure was used to
determine which treatment pairs were significantly different (N eter et a1. 1996). Larval
weight and length were normalized by square-root transformations and were analyzed

using ANOVA to see if it differed among conifers. Variables associated with host

79

preference (the number of egg niches, eggs, larvae, and galleries) were analyzed for all
conifer species combinations using 2-sided T-tests or the Mann-Whitney test when data
was not normally distributed (Zar 1984). Variables associated with host suitability (larval
weight, larval length, gallery length, gallery width, and percentage survival) were
analyzed for all conifer species combinations using 2-sided T-tests or the Mann-Whitney
test when data were not normally distributed.

For the larval conditioning trials, the number of egg niches and eggs in each
conifer species combination were analyzed using a 2-sided T-test when normally
distributed and the nonparametric Mann-Whitney test when variables were not normal
even after transformation (Zar 1984). Results of the Mann-Whitney test were consistent
with the T-test results in all cases.

Spearman’s Rank correlation test was used to test for associations between
number of egg niches, number of eggs, number of larvae, and number of galleries (host
preference variables) and larval weight, larval length, and percentage survival (larval
performance variables) for each conifer species. Spearman’s Rank correlation test was
also used to test for associations between log diameter, bark thickness and the number of
egg niches, number of eggs, number of larvae, and number of galleries (host preference
variables) and larval weight, larval length, and percentage survival (larval performance
variables). All analyses were conducted at P<0.05 level of significance, using SAS

statistical software (SAS Institute, Inc. 1989).

80

RESULTS

Host Preference

Adult M. scutellatus preferred white pine and white spruce during the choice tests.
All the logs of each species were pooled across all conifer paired-combinations to test for
differences in the number of egg niches, eggs, larvae, and galleries (Table 3.2). Female
adults excavated significantly more egg niches and laid more eggs on white pine than on
the other conifers (Table 3.2). Females preferred to excavate niches on red pine and
white spruce over jack pine. Jack pine logs were the least preferred species. More than
twice as many egg niches were found on white pine than on jack pine and significantly
fewer eggs were laid on jack pine than on the other log species. Significantly more
larvae were recovered from white pine and red pine than from jack pine and white spruce.
The number of galleries were similar on red pine and white pine logs, but the number of
galleries on red pine was not significantly different from jack pine and white spruce.
Galleries were longer but not wider on red pine than on the other four conifers. Galleries
on white spruce were about half the size of galleries on the other species.

Female rank order of host preference was white pine, white spruce and red pine,
and jack pine for oviposition conifer paired-comparisons of four conifers to test for host
preferences. Table 3.3 shows results of 2-choice tests conducted to rank female
preferences between two conifers. White pine logs always had the most egg niches, eggs,
larvae and larval galleries, even though the number of eggs and egg niches wasn’t
significantly different from white spruce. Female M. scutellatus excavated almost twice

as many egg niches on white pine when paired with either red pine or jack pine than with

81

white spruce and twice as many larvae were recovered on white pine than on the other
pines. Females excavated significantly more niches, laid more eggs and more larvae
were recovered on red pine than on jack pine and about the same number of egg niches,
eggs, larvae and galleries were on white spruce. Jack pine always had the lowest number
of egg niches, eggs, larvae and galleries for all species combinations.
Host Suitability

Pooled variables measuring suitability for larval development yielded varying
results (Table 3.2). Overall, red pine and white pine were most suitable for M. scutellatus
larval development. Larvae found in red pine logs weighed at least twice as much and
were significantly longer than those found in the other conifers. Larvae developing on
red pine consistently weighed significantly more and were longer, however, than larvae
that developed on white pine logs. Larval galleries were longer on red pine than on white
pine logs, but gallery widths were not significantly different. Growth rate, assessed by
larval weight and number of entrance holes after 30 days of development, was different
for larvae on red and white pine logs because larvae on red pine were twice the size of
larvae on white pine. Larvae developing in white spruce logs weighed significantly less
and were smaller than those found in all the other conifers. Larvae found in white spruce
were actually only first or second instars possibly experiencing a delay in hatch. When
red pine and white pine logs were dissected most of the larvae were third instars, and
larvae of earlier instars were rarely encountered. When found, first and second instar
larvae were always on logs with multiple galleries and with other third instar larvae still
feeding on phloem. Significantly more entrance holes into the sapwood were found on

red pine and white pine logs than on jack pine and white pine, suggesting that larvae

82

grew faster on red and white pine. Early instar survival was not significantly different
among the four conifers, but survival was poorest on jack pine logs.

Results from the six different conifer paired-combinations revealed similar
patterns in larval performance on the pines and spruce (Table 3.3). There was no
significant difference in gallery length and width and early instar survival between jack
pine and red pine logs (Table 3.3). However, M. scutellatus larvae grew larger, weighed
more and were longer when they developed on red pine logs compared with jack pine
logs. They also developed faster; they were larger and more larvae excavated tunnels
into the sapwood after 30 days on red pine logs than on jack pine logs.

Again, suitability of white pine and jack pine logs for M. scutellatus development
varied depending on the variable considered (Table 3.3). Significantly more entrance
holes into the sapwood were on white pine logs than jack pine logs. No significant
differences in larval weight, length, or gallery length and width were found between
white pine and jack pine logs.

No significant differences in number of entrance holes into the sapwood, or
percentage of survival were found between jack pine and white spruce logs (Table 3.3).
However, larvae from jack pine logs weighed more and were longer than larvae reared
from white spruce logs. Larvae consistently consumed more phloem on jack pine than on
white spruce logs in 30 days.

Larvae weighed significantly more and their body was longer when they
developed on red pine than on white pine (Table 3.3). Significantly more larvae survived
on white pine than on red pine, even though they were smaller in body size. There were

about the same number of entrance holes into the sapwood after 30 days on red pine and

83

white pine. The smaller size of larvae developing on white pine didn’t likely affect their
ability to tunnel into the sapwood at the same rate as larvae on red pine.

Red pine was considered more suitable than white spruce logs because larvae that
developed on white spruce were only a small fraction of the size of those that developed
on red pine (Table 3.3). Larvae consumed less than half as much phloem on white spruce
logs than on red pine logs and they excavated significantly more entrance holes on red
pine, which makes sense because larvae were much larger on red pine. Larvae on white
spruce logs experienced significantly greater survival on logs in the red pine and white
spruce combinations than for other combinations for some reason.

When white pine and white spruce log combination was tested, no differences in
the numbers of recovered larvae, larval weight or mortality were found (Table 3.3).
However, larvae grew longer when they developed on white pine than on white spruce
and they consumed significantly more phloem on white pine logs. Additionally, larvae
developed faster on white pine logs because significantly more entrance holes were found
after 30 days.

Results of correlation tests comparing associations between variables related to
host preference and larval performance differed depending on the indices that were
tested. For jack pine, egg niches (P<0.01, R=O.47), eggs (P<0.001, R=O.60), larvae
(P<0.001, R=O.60) and galleries (P<0.012, R=O.59) were correlated to the number of
entrance holes. There were no significant correlations between variables that measure
host preference and host suitability on white spruce. For red pine, the number of egg
niches (P<0.001, R=O.59) eggs (P<0.001, R=O.73), larvae (P<0.001, R=O.80) and

galleries (P<0.001, R=O.81) were correlated to the number of entrance holes. For white

84

pine, egg niches were correlated to larval weight (P<0.04, R=O.43) and length (P<0.03,
R=O.45) and number of entrance holes (P<0.01, R=0.36). Additionally, the number of
eggs (P<0.001, R=O.58), larvae (P<0.001, R=O.54), and galleries (P<0.002, R=O.54) were
correlated to the number of entrance holes. Correlations between the four estimates of
host preference (number of egg niches, eggs, larvae, and galleries) and the two important
indicators of larval performance, larval weight and percentage larval survival, were not
significant for any of the conifer species.

Figure 3.1 provides a summary of the relationship between adult female M.
scutellatus host preference and larval performance on the four conifers. The preference
and performance of M. scutellatus was rated as strong, medium and poor for each of the
conifers. White pine was highly preferred for oviposition and for larval development.
Red pine was not preferred as much as white pine, but was a highly suitable host for
larval development. Jack pine was not preferred for oviposition and was rated as a
medium host for larval development. Females showed a strong oviposition preference
for white spruce, but it was a poor host for larval development.

Results of correlation tests comparing associations between variables related to
log characteristics and variables related to host preference and larval performance were
similar. Log diameter, bark thickness were not correlated to the number of egg niches,
eggs, larvae, or galleries. Log diameter and bark thickness were also not correlated to the
variables used to estimate larval performance: larval weight, larval length, number of

entrance holes, percentage survival or area of phloem consumed.

85

Larval conditioning

Females that were given only their larval host for oviposition over the 3-day test
layed a similar number of eggs over the 3-day test as for females given two logs to
choose between. Table 3.4 shows results of no choice tests to assess the effects of larval
conditioning on host preferences. As expected, the number of egg niches, eggs, and the
number of galleries did not differ significantly among red pine, white pine and white
spruce (Table 3.4). Jack pine could not be included in the statistical analysis because of
issues with healthy female availability. The control tests demonstrated that females
reared in red pine, white pine, and white spruce found their larval host acceptable for
ovipsition.

Larval conditioning does not appear to influence female M. scutellatus host
preference (Table 3.5). On average, females excavated 25.3 egg niches and laid 7 eggs
during the 3 day trials. The number of egg niches excavated by females, eggs laid and
number of galleries was lower for the jack pine versus white spruce combinations than
for any other jack pine combination. A higher percentage of egg niches, out of the total
excavated during the 3-day test, was consistently on the alternate host paired with jack
pine when we tested for conditioning effects of females that developed in jack pine.
Females that developed on jack pine also laid a significantly higher percentage of eggs on
the alternate host, white spruce. However, there were no differences in the percentage of
eggs laid, or in number of galleries between jack pine and red pine combinations and jack
pine and white pine combinations (Table 3.5).

Female M. scutellatus reared from red pine logs did not show a consistent

oviposition preference for red pine logs (Table 3.5). Females reared in red pine

86

excavated 31 egg niches and laid 10 eggs on average during 3 days. Females excavated
60% of the egg niches on red pine and 40% on jack pine, but females preferred to
excavate niches on white pine and white spruce over red pine, excavating about 60% of
the niches on these conifers. However, females laid twice as many eggs and twice as
many galleries were found on white pine than on red pine logs (P<0.05).

Females that were reared on white pine logs excavated 32.5 niches and laid 12
eggs on average over the 3-day test period (Table 3.5). Females from white pine logs
excavated two-thirds of the total egg niches and laid about two-thirds of her eggs on
white pine during white pine versus jack pine combinations (Table 3.5). Similarly,
females excavated about 66% of the egg niches and laid two-thirds of her eggs on white
pine during white pine versus red pine combinations, and the percentage of niches
excavated on logs in other species combinations was similar (Table 3.5).

Females that emerged from white spruce logs excavated 18.9 egg niches and laid
8 eggs on average on the two logs in the tents over a 3-day period. Females did not
consistently prefer white spruce over the other conifer species (Table 3.5). There was no
significant difference in the percentage of egg niches excavated, eggs laid, or number of

galleries found on white spruce and other conifers (Table 3.5).

87

DISCUSSION

One of the principal questions posed in this thesis is whether M. scutellatus
females prefer certain hosts when presented with several potentially suitable hosts for
oviposition. Thompson (1988) and Via (1986) show that preference is best expressed as
the proportion of eggs laid on each of the plant species offered in choice trials, or a
hierarchical ordering of species. The results suggest that, in general, M. scutellatus
females preferred to oviposit on white pine over other conifers. Females equally
preferred to oviposit on white spruce and red pine, while jack pine was the least preferred
conifer species. More than 50% of the female’s total egg niches and eggs over 3 days
were consistently on white pine over other Species in paired conifer comparisons. More
larvae developed on white pine and red pine than on white spruce and jack pine logs. In
a related study, females preferred white pine and white spruce for oviposition in the field
(Lazarus and McCullough, in preparation). In another study in Ontario M. scutellatus
excavated more niches and laid more eggs on white pine over red pine (Peddle 2000),
again suggesting that females have the ability to rank potentially acceptable host species
according to preference.

Another question addressed herein is whether the conifer species tested differed in
suitability for larval development. Overall, red pine and white pine were more suitable
hosts for M. scutellatus larval development than jack pine and white spruce.
Characteristics used to estimate larval performance could include survival of immature
stages, larval growth rate, pupal mass and the resulting adult fecundity and longevity
(Thompson 1988). The number of entrance holes, larval weight, larval length and

percentage survival did not consistently indicate larval performance in this study.

88

White pine was preferred for oviposition, but larvae performed well on red pine
also. Larvae that developed on red pine were consistently larger than those from the
other conifer species. Similar numbers of larvae developed on red pine and white pine
logs, thus differences in larval size could be attributable to either higher nutritional
quality of red pine or increased feeding rates by larvae on red pine. Larvae on red pine
and white pine appeared to develop faster than larvae on jack pine and white spruce
because more entrance holes into the sapwood were found on red pine and on white pine
logs. This was another indication that larval grth rate was greater for larvae developing
on red pine and white pine than on the other two conifers. Larvae consumed the most
phloem on red pine logs, followed by white pine, then jack pine and finally white spruce.
Percentage survival was very similar among the conifer species in our study. Other
studies have shown that mortality rate is high for first instar larvae (Dyer and Seabrook
1978, Rose 1957). Since logs were debarked after 30 days, most larvae on the pines were
second or third instar and we may not have been able to accurately record some of the
first instar mortality noted in other studies.

Phloem nutritional quality or different chemical composition between pines and
spruce was probably responsible for reduced performance of larvae in white spruce.
Larvae were smaller on white spruce than on other conifers. An explanation for the
discrepancy in development time and body size is unclear, but is most likely related to a
reduced ability to utilize spruce terpenoids or delaying egg hatch, since bark thickness
and diameter were not correlated to any of the variables that estimate host preference and
larval performance. Monoterpenes and resin acids found in pine and spruce differ, and

pine and spruce contain different percentages of monoterpenes such as alpha-pinene,

89

camphene, beta-pinene, myrcene, carene and limonene in oleoresin (Drew and Pylant
1966, Westfall, 1972). For example, 14 percent of white pine and white spruce oleoresin
is made of beta-pinene, but only 4 percent of Scotch pine oleoresin is made of beta-
pinene (Westfall 1972). In addition, white pine contains camphene, but white spruce
does not (Westfall 1972). Fungal associations (ectosymbiotic fungi) and micro-
organisms (endosymbionts) are known to have relationships with xylem-feeding insects,
but specific symbiotic associations with M. scutellatus that influence phloem quality and
suitability of jack pine, red pine, white pine and white spruce are not certain (Chapman
1998)

Indices or characteristics of host preference and host suitability are not always
positively correlated and evaluations of host preference may depend on the criteria of
interest (Thompson 1988). I tried to determine whether larval performance estimated by
survival of the immature stages, and growth rate (larval weight and number of entrance
holes) was correlated to adult host preference. Ovipositing females preferred white pine
and white spruce over jack pine and red pine, while the suitability of hosts for larval
development was not as easy to discern. Larval survival didn’t vary among conifers and
larvae were largest on red pine and then on white pine. There is a lack of correspondence
between M. scutellatus female host preference and larval performance since we found no
correlations between the indices of host preference and host suitability for any of the
conifers in this experiment. Our lack of correspondence between a white pine preference
and best performance on red pine larvae, using larval weight and percentage survival as
indicators of performance, is puzzling. Ultimately, there was high survival for larvae on

all conifer species even if larval grth rate on white spruce was slower. An explanation

90

for the inconsistencies found here may confer greater fitness to this polyphagous species.
The progeny reared on these logs may have been more adapted to Kellogg Forest where
females would have access to several conifer species every year, in varying abundance.
The lack of correspondence between preference and all of the indicators of performance
suggest that M. scutellatus females oviposit on some hosts that are moderately adequate
for survival, as was found in other studies (Singer 1983, Thompson 1988). Adult females
of polyphagous species may benefit by finding numerous hosts attractive for oviposition
in situations where host abundance is highly variable from year to year.

Finally, I hypothesized that the larval host species would influence adult host
preferences. If a conditioning effect existed there should have been a preference for the
larval host no matter what alternate log species was paired with it during the 3-day tests.
We did not find this to be the case for any females regardless of whether they were reared
in jack pine, red pine, white pine or white spruce. Females reared in jack pine always
excavated more niches and laid more eggs on the alternate species. Females reared in red
pine logs consistently laid more eggs on the alternate species, while females reared in
white spruce logs demonstrated no preference for either white spruce or alternative
species. The only exception was for females reared in white pine logs. Here, females did
consistently excavate more egg niches and oviposit most frequently on white pine no
matter what alternative host was available. This pattern likely reflects a preference of
ovipositing females for white pine rather than conditioning effects of the larval host
predisposing females to oviposit on white pine. Monochamus carolinensis, a congener of
M. scutellatus, was also not influenced by larval conditioning (Walsh and Linit 1985).

F utuyma and Peterson (1985) conclude that the Hopkins host selection principle, or the

91

idea that larval host influences the adult host preference, does not hold for most
phytophagous insects.

In this study, I attempted to understand patterns of host use within a population of
M. scutellatus studied by investigating the variability of behavior among ovipositing
adults and subsequent effects of their host selection on larval performance. Research
with butterflies has shown that individuals often oviposit on more than one host species
(Singer 1983), thereby showing that variations of preference exist even within the
individual. For this population of M. scutellatus there was much variability in the
preferences of females for the conifers studied. There is also phenotypic variability in
growth rates of individuals within a population of M. scutellatus depending on the host,
and an absence of a host preference-larval performance correlation. It would be
interesting to test for phenotypic and genetic variation in other populations of M.
scutellatus perhaps in other geographic locations to differences in the abundance and

diversity of conifer species.

92

 

 

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Figure 3.1. A graphic representation of the relationship between adult female M.
scutellatus host preference and larval performance on white pine, red pine, jack pine, and
white spruce logs from a 2-choice test experiment conducted summer 2002.

98

APPENDIX

99

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.

Voucher No.: 2003-01

 

Title of thesis or dissertation (or other research projects):
HOST PREFERENCE AND HOST SUITABILITY OF TEN HARDWOODS FOR THE
ASIAN LONGHORNED BEETLE, Anoplophora glabripennis (Motshulsky) AND OF

FOUR CONIFERS FOR THE WHITE-SPOTTED PINE SAWYER, Monochamus
scutellatus (Say)

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

Entomology Museum, Michigan State University (MSU)

Other Museums:

100

Appendix 1.1
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102

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103

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111