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Theses and Dissertations--Entomology Entomology

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A NON-NATIVE FOREST INVADER ALTERS FOREST STRUCTURE A NON-NATIVE FOREST INVADER ALTERS FOREST STRUCTURE

AND THE ASSOCIATED ARTHROPOD COMMUNITY AND THE ASSOCIATED ARTHROPOD COMMUNITY

Matthew B. Savage University of Kentucky, matthewsavage92@gmail.com Digital Object Identifier: https://doi.org/10.13023/ETD.2017.376

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Recommended Citation Recommended Citation Savage, Matthew B., "A NON-NATIVE FOREST INVADER ALTERS FOREST STRUCTURE AND THE ASSOCIATED ARTHROPOD COMMUNITY" (2017). Theses and Dissertations--Entomology. 41. https://uknowledge.uky.edu/entomology_etds/41

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Matthew B. Savage, Student

Dr. Lynne Rieske-Kinney, Major Professor

Dr. Charles Fox, Director of Graduate Studies

A NON-NATIVE FOREST INVADER ALTERS FOREST STRUCTURE AND THE ASSOCIATED ARTHROPOD COMMUNITY

THESIS

A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in the

College of Agriculture, Food and Environment at the University of Kentucky

By

Matthew B. Savage

Lexington, Kentucky

Director: Dr. Lynne Rieske-Kinney, Professor of Entomology

Lexington, Kentucky

2017

Copyright © Matthew B. Savage 2017

ABSTRACT OF THESIS

A NON-NATIVE FOREST INVADER ALTERS FOREST

STRUCTURE AND THE ASSOCIATED ARTHROPOD COMMUNITY The emerald ash borer (EAB, Agrilus planipennis Fairmaire) (Coleoptera:

Buprestidae) is a non-native wood boring beetle that is causing extensive ash (Fraxinus spp.) mortality in eastern North America, affecting both urban and wildland forests and drastically altering forest structure and composition. As EAB-induced ash mortality progresses, native arthropod associates of ash forests are impacted by the effects of rapid and broad scale tree mortality. These include loss of food source, increased canopy gap formation, alterations in litter inputs causing shifting temperature and moisture regimes on the forest floor, and significant accumulation of coarse woody debris.

I assessed the sub-canopy arthropod community in five forests, all in different stages of the invasion process, from introduction through impact. Additionally, I assessed the ground level arthropod community in a post EAB-invaded forest with 100% mature ash mortality. Arthropod communities were assessed at the ordinal level, and with a focus on coleopterans, they were further classified to families and trophic guilds to analyze abundance, richness, and diversity. Due to their overwhelming abundance, I identified scolytines collected in the post EAB-invaded forest to species to see if the EAB-invasion was part of a greater invasional meltdown. My results indicate that the EAB-invasion in North America is affecting the native coleopteran communities associated with these forests.

KEYWORDS: Agrilus planipennis, Fraxinus, invasive species, invasional meltdown, Scolytinae, trophic guild

Matthew B. Savage

July 12, 2017

A NON-NATIVE FOREST INVADER ALTERS FOREST

STRUCTURE AND THE ASSOCIATED ARTHROPOD COMMUNITY

By

Matthew B. Savage

Dr. Lynne Rieske-Kinney

Director of Thesis

Dr. Charles Fox

Director of Graduate Studies

July 12, 2017

Date

iii

ACKNOWLEDGEMENTS

This work has been possible thanks to the contributions of many individuals,

groups and institutions. I would first like to thank my advisor Dr. Lynne Rieske-Kinney

for believing in my potential and guiding me through my masters with her constant

support and advice. I would like to thank my past and current lab members, Abe Nielsen,

Ignazio Graziosi, Chris Strohm, Bill Davidson, Glenn Skiles, Sean Fenstemaker, and

Shouhui Sun for their friendship and all of their efforts in the lab and field. Thanks to our

undergraduate workers, Kyle Ritter, Jay Goldstein, Jeb Ayres, and Shane Stiles who

spent many long hours sorting through insect traps. Thanks to my research committee,

Dr. Mary Arthur and Dr. Lee Townsend, for their knowledgeable support to my research.

I would like to thank all of the landowners, Steve Martin, Lee Crawfort, Bonnie

Cecil, Taylorsville Lake State Park, Shelby County Parks and Recreation, and the

LFUCG staff at Raven Run Nature Sanctuary, for providing access to land for this

project. Thanks to Sarah Janse and Dr. Connie Wood for providing statistical support and

Dr. Eric Chapman for his expertise in beetle identification and his friendship. Funding

was provided by the USDA Forest Service Special Technology Development Program

and McIntire Stennis Funds from the Kentucky Agricultural Experiment Station.

Lastly, a special thanks to my parents, Kathy and Brian Savage, and Kelly

Jackson for their encouragement along the way.

iv

TABLE OF CONTENTS

ACKNOWLEDGEMENTS ............................................................................................... iii

TABLE OF CONTENTS ................................................................................................... iv

LIST OF TABLES ............................................................................................................ vii

LIST OF FIGURES ......................................................................................................... viii

CHAPTER 1: Introduction ................................................................................................. 1

CHAPTER 2: Coleopteran communities shift in response to emerald ash borer induced

ash decline in recently invaded forests ............................................................................... 5

Introduction ......................................................................................................... 5

Materials and Methods ........................................................................................ 6

Study Sites ...................................................................................................... 6

Forest Characteristics ...................................................................................... 7

Arthropod Monitoring ..................................................................................... 7

Analysis........................................................................................................... 8

Results and Discussion ....................................................................................... 9

Forest Characteristics ...................................................................................... 9

Arthropod Communities ............................................................................... 10

v

CHAPTER 3: Coarse woody debris accumulation associated with emerald ash borer-

induced ash mortality affects arthropod composition ....................................................... 20

Introduction ....................................................................................................... 20

Materials and Methods ...................................................................................... 22

Study Area .................................................................................................... 22

Coarse Woody Debris ................................................................................... 22

Arthropod Monitoring ................................................................................... 23

Analysis......................................................................................................... 24

Results ............................................................................................................... 26

Coarse Woody Debris ................................................................................... 26

Arthropod Monitoring ................................................................................... 26

Funnel traps ................................................................................................... 26

Pitfall traps .................................................................................................... 28

Pan traps ........................................................................................................ 29

Discussion ......................................................................................................... 30

APPENDICES .................................................................................................................. 45

APPENDIX A: Trophic guild designations ..................................................................... 46

vi

APPENDIX B: Temporal data by month.......................................................................... 48

APPENDIX C: CWD × Month treatment interactions ..................................................... 52

APPENDIX D: Scolytine abundance and origin .............................................................. 55

REFERENCES ................................................................................................................. 56

VITA ................................................................................................................................. 67

vii

LIST OF TABLES

Table 2.1, Forest characteristics and coleopteran abundance at five sites in north-central Kentucky used to evaluate changes in the colopteran community associated emerald ash borer-induced ash decline in 2014…14 Table 2.2, Relative abundance and richness of Coleopteran feeding guilds sampled from five sites affected by emerald ash borer ash decline……...15 Table 3.1, Effects of the presence of coarse woody debris and of season (Fdf/P) on coleopteran a) population parameters, b) feeding guild abundance, and c) family abundance sampled from funnel traps in EAB invaded forests of north-central Kentucky………………………………………..37 Table 3.2, Effects of the presence of coarse woody debris and of season (Fdf/P) on scolytinae species abundance (S.E.) sampled from funnel traps in EAB invaded forests of north-central Kentucky in 2015………………...39 Table 3.3, Effects of the presence of coarse woody debris and of season (Fdf/P) on coleopteran a) population parameters, b) feeding guild abundance, and c) family abundance sampled from pitfall traps in EAB invaded forests of north-central Kentucky………………………………………..41 Table 3.4, Effects of the presence of coarse woody debris and of season (Fdf/P) on coleopteran a) population parameters, b) feeding guild abundance, and c) family abundance sampled from pan traps in EAB invaded forests of north-central Kentucky………………………………………..43

viii

LIST OF FIGURES

Figure 2.1, Relative abundance of the ten numerically dominant coleopteran families found in forests under evaluation for emerald ash borer-induced ash mortality……………………………………………...16 Figure 2.2, Cumulative coleopteran family richness at five forested sites in north central Kentucky in 2014…………………………………………..17 Figure 2.3, Coleopteran feeding guild absolute abundance related to ash canopy decline (least to greatest) at five forested sites in north central Kentucky…………………………………………………………18 Figure 2.4, Correlation between Fraxinus canopy decline and absolute abundance of two coleopteran predators…………………………………19 Figure 3.1, Layout of plots, traps, and trap types used to evaluate the arthropod community associated with ash coarse woody debris in emerald ash borer-impacted forests……………………………………………….35 Figure 3.2, Volume (m3/0.1 ac.) of ash coarse woody debris in each decay class (mean (s.e.)) six years following invasion of emerald ash borer…..36

1

CHAPTER 1

Introduction

Non-native species’ invasions threaten forest ecosystem function (Ehrenfeld

2010) and native biodiversity (Wilcove et al. 1998, Byers et al. 2002), often with

widespread and devastating economic impacts (Pimentel et al. 2005, Aukema et al.

2011). Insect outbreaks can kill plants and alter the distribution of biomass over large

areas (Wilcove et al. 1998, MacLean 2002, Breshears et al. 2005, Kurz et al. 2008,

Brown et al. 2010), and also cause disturbance-induced changes in trophic interactions

(Schowalter 1985, Schowalter and Lowman 1999).

The emerald ash borer (Agrilus planipennis Fairmaire, EAB, Coleoptera:

Buprestidae) is a wood-boring beetle native to east Asia, which was first detected in

North America in 2002 near Detroit, MI (Haack et al. 2002). Larvae feed just below the

bark on the phloem of ash trees (Fraxinus spp.), forming serpentine galleries that destroy

vascular tissue and disrupt the translocation of water and nutrients to the canopy. This

ultimately girdles the tree, leading to rapid mortality (Cappaert et al. 2005, Flower et al.

2013). All North American Fraxinus, even healthy trees, are susceptible to EAB

(Cappaert et al. 2005), though blue ash, F. quadrangulata, demonstrates some putative

resistance (Tanis and McCullough 2012). Since its introduction, EAB has spread rapidly

through much of the eastern contiguous US and southeastern Canada (USDA APHIS

2017), and inflicted extensive ash mortality in affected regions. The EAB invasion in

North America is of unprecedented scope and magnitude (Herms and McCullough 2014),

and its effects are ultimately projected to be on a continental scale. The majority of EAB-

2

induced ash mortality (>85%) occurs within 3–5 years of an initial invasion (Poland and

McCullough 2006, Kashian and Witter 2011). However, following widespread

establishment residual populations may persist.

Ash trees are a consistent component of hardwood forests throughout the USA. In

addition to their economic value as a timber and landscape species, Fraxinus are

ecologically important as seed producers that provide valueable food resources for

wildlife (Kennedy 1990, Schlesinger 1990). Direct effects of the EAB invasion include

rapid ash mortality with subsequent alterations in ash-associated communities (Gandhi

and Herms 2010a). Specifically, ash tree mortality deprives many ash specialists of food

resources (Gandhi and Herms 2010b), though some species may thrive under the altered

conditions (Zhang and Liang 1995, Schowalter et al. 1999, Van Bael et al. 2004). The

indirect effects of rapid and broad scale tree mortality include increased canopy gap

formation, alterations in litter inputs causing shifting temperature and moisture regimes

on the forest floor, and significant accumulation of coarse woody debris (CWD) (Evans

2011, Perkins et al. 1987, Zhang and Liang 1995).

Increased coarse woody debris alters habitat heterogeneity, which can affect

arthropod communities through mechanisms such as increased exposure to predation and

parasitism (Shure and Wilson 1993, Van Bael et al. 2004). The increase in ash coarse

woody debris may have profound effects specifically on coleopteran community

associates, which are good indicators of disturbance (Schowalter and Ganio 2003) and

diversity (Hammond 1990), through altered habitats and resource distribution. These

habitat alterations could also facilitate invasions by other non–indigenous species,

creating an ‘invasional meltdown’ (sensu Simberloff and Holle 1999).

3

My research investigates the effects of EAB-induced ash mortality on arthropod

communities in invaded forests. I had two main objectives: (1) to evaluate arthropod

utilization of declining ash canopies during EAB invasions; (2) to evaluate arthropod

utilization of naturally occurring ash coarse woody debris post EAB invasion. I

completed my objectives through field experiments in six locations in east central

Kentucky over the course of two years.

In 2014, I monitored five locations in different stages of the invasion process from

introduction through impact. In chapter 2, I investigate the effects of EAB-induced ash

mortality on arthropod communities in invaded forests in different stages of the invasion

process. I predicted that EAB-induced ash mortality would lead to changes in forest

composition and structure resulting in shifts in arthropod communities. I compared the

arthropod activity across the five sites to determine any changes in arthropod

composition.

In 2015, I narrowed my focus to one county with three geographically distinct

study sites, where the EAB invasion had caused 100% ash mortality. In chapter 3, I

investigated arthropod utilization of naturally occurring ash coarse woody debris by

monitoring arthropods in plots with and without ash coarse woody debris. I predicted that

the influx of coarse woody debris due to EAB-induced ash mortality would create

additional habitat and resources resulting in changes in arthropod composition,

abundance, and richness.

Few studies have evaluated arthropod associates in EAB invaded forests, and a

clear understanding of the role of EAB induced ash mortality on forest arthropod

communities is lacking. By characterizing these effects on arthropod communities at

4

different stages of the EAB invasion, my research provides a greater understanding of the

impacts of the invasion and adds to the knowledge base surrounding arthropod

community structure following invasive species outbreaks. This is critical for conserving

native arthropod biodiversity following large-scale disturbance events.

5

CHAPTER 2

Coleopteran communities shift in response to emerald ash borer induced ash decline in

recently invaded forests

Introduction

Invasions by non-native invasive species pose significant threats to forest

ecosystem function (Ehrenfeld 2010) and native biodiversity (Wilcove et al. 1998, Byers

et al. 2002), and have widespread economic impacts (Pimentel et al. 2005, Aukema et al.

2011). Ash trees (Fraxinus spp.) are a consistent component of hardwood forests

throughout the United States (Kennedy 1990, Schlesinger 1990) whose prevalence and

persistence is threatened by the emerald ash borer (Agrilus planipennis Fairmaire, EAB,

Coleoptera: Buprestidae). This invasive wood-boring beetle is native to eastern Asia and

first detected in North America in 2002 near Detroit, MI (Haack et al. 2002). Since its

accidental introduction, EAB has rapidly spread through much of the eastern contiguous

United States and southeastern Canada (USDA APHIS 2016) inflicting extensive ash

mortality in affected regions. Larvae feed on phloem beneath the bark, forming

serpentine galleries and destroying the vascular tissue, disrupting translocation of water

and nutrients to the canopy, ultimately girdling the tree (Cappaert et al. 2005, Flower et

al. 2013). The majority of EAB-induced ash mortality (>85%) occurs within 3–5 years of

the initial invasion (Poland and McCullough 2006, Kashian and Witter 2011). All North

American Fraxinus species are susceptible to attack and EAB readily colonizes healthy

trees (Cappaert et al. 2005).

6

The direct effects of the EAB invasion include rapid ash mortality with

subsequent alterations in ash-associated communities (Gandhi and Herms 2010a). The

indirect effects of rapid and broad scale tree mortality include increased gap formation,

which alters light penetration to the forest floor, accumulation of coarse woody debris,

and qualitative and quantitative alterations in litter inputs causing shifting temperature

and moisture regimes on the forest floor (Perkins et al. 1987, Zhang and Liang 1995).

Such changes after EAB-induced ash mortality may affect native coleopteran community

associates of these invaded forests. My goal is to document how these native coleopteran

communities shifted in response to EAB-induced ash mortality in recently invaded areas

of the east central United States. Specifically I evaluated the extent to which coleopteran

trophic guild abundance and richness were affected by widespread EAB-induced ash

mortality.

Materials and Methods

Study Sites

Five forested study sites were established in north-central Kentucky along the

forefront of the expanding EAB infestation (Davidson and Rieske 2015), in Anderson,

Fayette, Henry, Shelby, and Spencer counties. Ash are historically a significant

component of the western mesophytic forests of the region (Wharton and Barbour 1973),

which thrive on the moist and fertile soils that predominate in this area (Campbell 1989).

At the onset of the study EAB was present at the Anderson, Henry, and Shelby sites

where ash decline was evident. EAB was first detected at Fayette and Spencer sites in

2014, but there were little to no signs of EAB-induced stress (Davidson and Rieske

2015). My sites were chosen to represent the full spectrum of ash decline associated with

7

the EAB invasion, including pre-invasion (Fayette, Spencer), peak invasion (Shelby), and

post-invasion (Henry, Anderson) forests.

Forest Characteristics

Forest vegetation was characterized at each site to determine pre-invasion

conditions and annually thereafter. For vegetation censusing, nine circular whole plots

(0.04 ha) were utilized to assess overstory and midstory vegetation (all trees ≥ 12.7 cm

diameter at 137 cm high; DBH), 0.004-ha subplots were used to assess saplings and

shrubs (< 12.7 cm DBH, > 137 cm high), and 0.0004-ha microplots were used to assess

seedlings and shrubs (< 137 cm height). One subplot and one microplot were positioned

at the whole plot center and in each cardinal direction, 7.7 m from the whole plot center.

Thus, a surveyed plot contained one 0.04-ha whole plot, five 0.004-ha subplots, and five

0.0004-ha microplots (Coleman et al. 2008). Measurements of vegetation and plot data

followed the Common Stand Exam protocol of the USDA Forest Service’s Natural

Resource Information System: Field Sampled Vegetation Module (USDA Forest Service

2009) and included tree height and DBH. Canopy dieback was visually assessed by a

single observer and each ash tree assigned a crown dieback rating from 0% (healthy) to

100% (dead).

Arthropod Monitoring

Native coleopteran communities in the sub-canopy strata were monitored using

12-unit Lindgren multi-funnel traps (one per plot, N=45) from 20 May to 12 September

2014. Traps were suspended over an ash branch (~4 m) and fitted with two 50 ml vials of

70% ethanol, a commonly used lure for xylophagous insects (Montgomery and Wargo

1983, Lindelöw et al. 1992, Bouget et al. 2009), hung from the funnel edge, and with a

8

dichlorvos strip (2 × 5 cm2) (AMVAC Chemical Corp., Los Angeles, CA) placed in each

trap bottom. Traps were monitored every 7-14 d; contents were rinsed and stored in 70%

EtOH in resealable plastic bags, and the lures replenished. In the laboratory samples were

sorted to order (Triplehorn and Johnson 2005); Coleoptera were further sorted and

identified to family using available keys (Triplehorn and Johnson 2005, Marshall 2006,

Evans 2014, BugGuide 2015), counted, and assigned to trophic guilds based on larval

feeding habits, including fungivore, predator, herbivore, xylophage, saprophage, or

parasitoid, (Hammond 1990). Ordinally the Coleoptera are trophically diverse, but more

or less trophically uniform within families (Hammond 1990, Hammond 1992), which

allows classifying families into feeding guilds that exploit resources in a similar manner

(Root 1967). In my study the carrion feeders, including some Silphidae, Staphylinidae,

Histeridae, Nitidulidae, and Lieodidae, were responding to the decaying trap contents

rather than the ethanol lure, which resulted in excessive fluctuations in abundance, and so

were excluded from my analyses.

Analysis

Ash decline, characterized by 2014 canopy dieback (Table 2.1), was compared

across all sites using a one-way analysis of variance (ANOVA), with post hoc analysis

performed using Tukey’s Honest Significant Difference (HSD) (Davidson and Rieske

2015). Ash canopy dieback ranging from low (Fayette) to high (Henry) was then used to

assess the influence of ash decline on coleopteran abundance and richness. Coleopteran

abundance measured with funnel traps (recorded as total Coleoptera trapped) was

calculated and richness (recorded as total number of coleopteran families trapped) was

derived by site. Evenness and diversity indices were not derived due to data gaps. Data

9

were tested for normality (PROC UNIVARIATE) and transformed when necessary.

Significance was determined at α = 0.05 unless stated otherwise. All analyses were

performed using SAS v9.3 (SAS Institute 2011).

Overall coleopteran abundance and cumulative richness by site were analyzed

using a repeated measures mixed linear model (PROC MIXED), with sample interval as

the repeated measure and individual plots (traps) as subjects. The difference of least

squares (Least Squares Means) was used to separate means for these population

parameters. Coleopteran feeding guild abundance and richness summed over the 16 week

sampling period were analyzed using a generalized linear mixed model (PROC GLM) to

compare guild × site interactions. Feeding guild abundance was transformed using a

square root transformation for total counts and arcsin transformation for percent

abundance. Feeding guild abundance (absolute and percent) was compared across all sites

where the difference of least squares was used to separate means and post-hoc analysis

was performed using pairwise T-Comparison if differences arose. Correlations between

the predator guild and ash canopy decline were analyzed (PROC CORR).

Results and Discussion

Forest Characteristics

Across my study sites ash relative stem density ranged from 12-26% for stems

>2.5 cm diameter, and ash mortality ranged from 0-50%. Ash canopy dieback differed

significantly across sites, and ranged from a low of ~7% at the most recently invaded site

to a high of 74% at the most degraded site (Table 2.1). Site level ash mortality and

canopy dieback were highly correlated with EAB abundance (Davidson and Rieske

2015).

10

Arthropod Communities

Funnel traps yielded 16,455 arthropods, including 11,786 coleopterans (>71%)

representing 57 families, excluding carrion feeders (Appendix 1). Elateridae was the most

abundant family (Fig. 2.1), with 16% of the total, followed by the Curculionidae and

Staphylinidae (13 and 12%, respectively); these three families comprised nearly 41% of

the coleopterans captured. The next most abundant families were the Ptilodactylidae

(9%), the Latridiidae (9%), and the Histeridae (8%); collectively they comprised almost

27% of the total coleopterans.

Coleopteran abundance (Table 2.1), but not cumulative family richness (Fig. 2.2)

differed among study sites; both tended to be lowest in pre- (Fayette) and post-invasion

(Henry) sites, and greatest at the site typifying peak invasion (Shelby). The increase in

abundance and cumulative richness of coleopteran associates at the site representing the

peak of the EAB invasion may be attributable to increases in habitat availability due to

newly forming snags and coarse woody debris and to volatiles released from dying trees

(Kimmerer and Kozlowski 1982, Montgomery and Wargo 1983, Harmon et al. 1986).

Coleopteran abundance was greatest among herbivores (4,207 individuals, 36%)

(Table 2.2), comprised primarily of the Elateridae, which feed on flowers, nectar, pollen,

and rotting fruit (Evans 2014). However, in spite of their abundance herbivores

comprised only 14% of the total with respect to family richness (8 families). In contrast,

coleopteran fungivore richness was highest at 40% (23 families), in spite of the fact that

abundance was relatively low (2,082 individuals, 17%) (Table 2.2). Fungivores are

dominated by the Latridiidae (Fig. 2.1), which typically feed on the reproductive

structures of fungi and are commonly found in plant debris (Evans 2014). Predators

11

comprised 26% of the total (3,050 individuals) and 19% of the coleopteran family

richness (11 families) (Table 2.2). Predators consisted primarily of the Staphylinidae

(Fig. 2.1), which are generalist predators, and the Histeridae, which has one subfamily

associated with bark beetle (Curculionidae) galleries, and another subfamily that feeds

principally on fly and beetle larvae associated with dung. Saprophages and xylophages

made up ~10% of the abundance, and similarly 10 and 12% of coleopteran family

richness, and consisted primarily of Ptilodactylidae and Scolytines (Fig. 2.1; Table 2.2).

Trophic guild abundance across sites varied (χ2 = 1,045.6; df = 20; P < 0.0001),

corresponding to levels of EAB-induced ash decline. Herbivore and saprophage

abundance was greatest at Shelby (Figs. 2.3a and 2.3b), which among the five sites

represents peak EAB invasion. Fungivore abundance (Fig. 2.3c) was positively

associated with ash decline and was greatest at Shelby, Anderson, and Henry, where the

EAB invasion is more advanced, and lowest at Fayette and Spencer, which were in the

very early stages of the invasion at the time of sampling. Xylophages were most abundant

at Spencer (Fig. 2.3d), again representing relatively early stages of EAB invasion.

Predator abundance (Fig. 2.3e) was lowest at Fayette and Henry, representing both pre-

and post-EAB invasion and highest at the sites where the invasion is nearer its peak.

Among the predators, Trogossitidae and Cleridae abundance (7 and 5% of total

predator abundance, respectively) were positively correlated with ash decline (α < 0.1)

(Fig. 2.4). Trogossitids consisted of 228 individuals of primarily Tenebroides sp.; these

are bark-gnawing beetles found beneath the bark of dead trees and are associated with

wood-boring beetles (Evans 2014). Clerids (144 individuals) consisted primarily of

Enoclerus sp.; these checkered beetles are associated with dead wood, and often found

12

predating larval Curculionidae, Cerambycidae, and Buprestidae (Evans 2014).

Interestingly, the parastic Passandridae, comprised entirely of Catogenus rufus

(Fabricius), were consistently present in low numbers, regardless of ash decline

(Appendix 1). Catogenus rufus (Fabricius) has been reported in association with EAB-

invaded forests and has been found as both larvae and adults in EAB galleries from dead

ash on these sites (Davidson and Rieske 2015); it was present in low numbers across sites

(Fig. 2.3f) and appeared unaffected by the stage of the EAB invasion or by the

corresponding decline in ash canopies. Its presence at all sites in similar abundance

suggests that it is capable of utilizing a variety of wood-boring hosts and is not

demonstrating a numerical response to the EAB invasion in these forests, although it may

be utilizing EAB as a prey resource. Tenebroides, Enoclerus and Catogenus have been

documented in association with EAB larvae and pupae near the epicenter of the EAB

invasion in North America (Bauer et al. 2004), suggesting that they may be playing a role

in population dynamics of this aggressive invader. Collectively, these data suggest that

native predators and parasites are being recruited to invading emerald ash borer

populations, and that these native natural enemies may be a viable component of post-

invasion EAB population dynamics in North American forests.

The reverberating effects of the emerald ash borer invasion in North America will

undoubtedly include effects on native arthropod associates. Changes in forest structure

and composition, alterations in light penetration to the forest floor, and inputs of coarse

woody debris create and eliminate habitats and affect resource availability. Ash-

dominated forests are predicted to have a loss of overall arthropod richness, are facing

cascading ecological impacts, and altered ecosystem processes (Gandhi and Herms

13

2010b). Of the 282 native arthropods associated with ash, 43 are monophagous; nine of

these monophages are coleopterans (Gandhi and Herms 2010b). These ash specialists will

undoubtedly be negatively affected and may experience localized extirpation.

The emerald ash borer devastates unprotected ash. Following depletion of the ash

resources, EAB populations sharply decline (Herms and McCullough 2014), greatly

reducing the pest pressure on regenerating seedlings and saplings. The decline in pest

pressure increases the chance of continued survival of young ash in North American

forests (Duan et al. 2015), providing essential resources for ash specialists. Ash forests

are changing, and a deeper understanding of how arthropod communities and trophic

guilds are responding will contribute to more proficient monitoring and protection.

Table 2.2. Forest characteristics and coleopteran abundance at five sites in north-central Kentucky used to evaluate changes in the

colopteran community associated emerald ash borer-induced ash decline in 2014.

Fraxinus spp.

Site

No. stems

(all spp.)

No. stems

%

% mortality

Canopy dieback

mean % (SE)

Coleoptera

abundance1

Henry 466 79 18.5

50.5 73.9 (4.6)a 1.30 (0.07)b

Anderson 402 53 18.8 18.9 56.9 (1.9)b 1.53 (0.07)ab

Shelby 385 121 26.0 7.4 27.4 (3.9)c 1.64 (0.06)a

Spencer 290 59 12.0 1.7 16.2 (2.8)cd 1.49 (0.11)ab

Fayette 267 42 13.1 0.0 7.4 (3.0)d 1.27 (0.08)b

F3, 350 58.6; P<0.001 F4, 527 2.1; P<0.02

1LS-means + se number of individuals per day captured in ethanol-baited funnel traps. Means separation on square root + 0.05

transformed data.

14

15

Table 2.2. Relative abundance and richness of Coleopteran feeding guilds sampled from

five sites affected by emerald ash borer ash decline.

Coleopteran family-level

Trophic guild Abundance (%) Richness (%)

Herbivore 36 14

Fungivore 17 40

Predator 26 19

Xylophage 10 12

Saprophage 10 10

Parasite <1 5

Unidentified <1 ---

Total 100 100

16

Figure 2.1. Relative abundance of the ten numerically dominant coleopteran families

found in forests under evaluation for emerald ash borer-induced ash mortality.

02468

101214161820

Col

eopt

eran

Fam

ily A

bund

ance

(%)

17

Figure 2.2. Cumulative coleopteran family richness at five forested sites in north central

Kentucky in 2014.

25

30

35

40

45

50

1 3 5 7 9 11 13 15

Cum

ulat

ive

Fam

ily R

ichn

ess

Weeks

FayetteSpencerShelbyAndersonHenry

May June July Aug. Sept. Oct.

18

Figure 2.3. Coleopteran feeding guild absolute abundance related to ash canopy decline

(least to greatest) at five forested sites in north central Kentucky, including a) herbivores,

b) saprophages, c) fungivores, d) xylophages, e) predators, and f) parasitoids. Means

followed by same letter do not differ (α=0.05).

19

Figure 2.4. Correlation between Fraxinus canopy decline and absolute abundance of two

coleopteran predators, including the Trogossitidae (Trogossitid abundance = 0.47 × (%

canopy decline) + 28.7, R2 = 0.84, P = 0.07) and the Cleridae (Clerid abundance = 0.45 ×

(% canopy decline) + 12.5, R2 = 0.86, P = 0.06).

0

10

20

30

40

50

60

70

80

0 10 20 30 40 50 60 70 80

Cole

opte

ran

Abso

lute

Abu

ndan

ce

Mean % Ash Canopy Decline

Trogossitidae AbundanceCleridae Abundance

20

CHAPTER 3

Coarse woody debris accumulation associated with emerald ash borer-induced ash

mortality affects arthropod composition

Introduction

Natural disturbances, including insect outbreaks, can extensively alter habitat

conditions and resource distribution across landscapes (White and Pickett 1985, Walker

1999, Schowalter 2012). Insect outbreaks can kill plants and alter the distribution of

biomass over large areas (Wilcove et al. 1998, MacLean 2002, Breshears et al. 2005,

Kurz et al. 2008, Brown et al. 2010), and also cause disturbance-induced changes in

trophic interactions (Schowalter 1985, Schowalter and Lowman 1999). In both cases,

non-native insect outbreaks pose significant threats to forest composition and structure,

forest ecosystem function (Ehrenfeld 2010) and native biodiversity (Wilcove et al. 1998,

Byers et al. 2002), and have widespread economic impacts (Pimentel et al. 2005, Aukema

et al. 2011). Each disturbance event is characterized by a combination of type,

magnitude, frequency, and extent that determines its effect on various organisms (White

and Pickett 1985, Walker 1999). Of particular concern in the USA are disturbance events

caused by invasion by the non-native emerald ash borer (Agrilus planipennis Fairmaire,

EAB, Coleoptera: Buprestidae).

Emerald ash borer is a wood-boring beetle native to east Asia and first detected in

North America in 2002 near Detroit, MI (Haack et al. 2002). Larvae feed on phloem of

ash trees (Fraxinus spp.), forming serpentine galleries which destroy the vascular tissue,

disrupting translocation of water and nutrients to the canopy, and ultimately girdling the

21

tree (Cappaert et al. 2005, Flower et al. 2013). Since its introduction EAB has spread

rapidly through much of the eastern contiguous US and southeastern Canada (USDA

APHIS 2016) inflicting extensive ash mortality in affected regions. Ash are a consistent

component of hardwood forests throughout the USA (Kennedy 1990, Schlesinger 1990),

and North American Fraxinus, even healthy trees, are susceptible to EAB (Cappaert et al.

2005). The EAB invasion in North America is of unprecedented scope and magnitude

(Herms and McCullough 2014), and its effects are projected to be on a continental scale.

The majority of EAB-induced ash mortality (>85%) occurs within 3–5 years of the initial

invasion (Poland and McCullough 2006, Kashian and Witter 2011). However, following

widespread establishment residual populations persist.

EAB-induced ash mortality affects insect communities both directly and

indirectly. Direct effects include rapid ash mortality which deprives many ash specialists

of food resources (Gandhi and Herms 2010b), though some species may thrive under the

altered conditions (Zhang and Liang 1995, Schowalter et al. 1999, Van Bael et al. 2004).

Indirect effects include increased gap formation which alters light penetration to the

forest floor, qualitative and quantitative alterations in litter inputs causing shifting

temperature and moisture regimes (Perkins et al. 1987, Zhang and Liang 1995), and

accumulations of coarse woody debris (CWD). The influx of ash coarse woody debris

following EAB invasions is significant (Evans 2011), affecting habitat heterogeneity of

the forest floor and thus resource availability and exposure to predation and parasitism

(Shure and Wilson 1993, Van Bael et al. 2004). The increase in ash coarse woody debris

may have profound effects on coleopteran community associates, which are good

indicators of disturbance (Schowalter and Ganio 2003) and diversity (Hammond 1990),

22

through altered habitats and resource distribution. These habitat alterations from the EAB

invasion could facilitate invasion by other non–indigenous species, creating an

‘invasional meltdown’ (sensu Simberloff and Holle 1999), whereby non-indigenous

species facilitate one another’s invasion success through increased likelihood of survival,

ecological impact, and magnitude of impact (Simberloff and Holle 1999). My goal was to

evaluate how this localized influx of ash coarse woody debris affects native coleopteran

communities in post EAB-invaded forests of the east central United States, and to

document evidence suggesting the occurrence of an EAB-induced invasional meltdown.

Materials and Methods

Study Area

My study area was a 13.8 ha forested tract in western Franklin County, KY,

which had been invaded by EAB since 2009. Historically ash were a significant

component of the western mesophytic forests of the region (Wharton and Barbour 1973),

which thrive on the moist and fertile soils that predominate in this area (Campbell 1989).

Initially ash comprised ~10 percent of the woody plant composition at the site, but at the

onset of my study in fall 2014 ash mortality was approaching 100% (Levin-Nielsen and

Rieske 2014). This EAB-induced ash mortality resulted in large accumulations of

naturally occurring coarse woody debris, woody material >2.5 cm in diameter, of varying

age, size, and state of decomposition.

Coarse Woody Debris

In October 2015 three discrete areas were designated containing six 0.04 ha whole

plots; three were designated as ash CWD present (CWD+), and three were designated as

ash CWD absent (CWD–). Within the CWD– plots, all ash CWD was carefully removed

23

and placed well outside the 0.04 ha plot boundary. Within the CWD+ plots the ash CWD

was carefully condensed into a central subplot and placed so as to maximize contact with

the ground. The length and diameter at both ends of all ash >2.5 cm diameter were

measured and assigned to a decay class (Vanderwel et al. 2006), which included (1) wood

hard, bark intact; (2) wood hard, bark beginning to slough; (3) wood softening, usually no

bark remaining; (4) wood substantially decayed, pieces sloughing off, inner heart wood

still intact; and (5) wood thoroughly decayed, texture powdery and resembles soil. To

calculate the amount of ash CWD present I recorded both small and large end diameter of

each piece to obtain an average diameter, then used the formula for a cylinder 𝑉𝑉 = 𝜋𝜋𝑟𝑟2ℎ

summed over all ash present to produce an ash CWD volume estimate, which is

expressed as m3 per plot. Ash CWD volume was also calculated for each decay class

(Fig. 3.2).

Arthropod Monitoring

To monitor associated EAB activity, green 12-unit multifunnel traps (N = 9)

affixed with manuka oil and leaf alcohol lures (Synergy, Brunbay, BC) were deployed

adjacent to study plots. Arthropod communities associated with the coarse woody debris

treatments were monitored from 21 March to 28 August 2014 using one 8-funnel trap (n

= 18) and two pairs of pitfall and pan traps (n = 36), spaced ~1 m apart equilaterally (Fig.

3.1). Funnel traps were mounted at a height of 1 m, and fitted with two 50 ml vials of

70% ethanol, a commonly used lure for xylophagous insects (Montgomery and Wargo

1983, Lindelöw et al. 1992, Zhang and Liang 1995, Bouget et al. 2009), hung from the

funnel edge, and with a dichlorvos strip (2 × 5 cm2) (AMVAC Chemical Corp., Los

Angeles, CA) placed in each trap cup. To sample surface and ground-dwelling

24

arthropods, pitfall traps were used consisting of two nested 473 ml plastic cups placed

into a hole flush with the soil surface and filled with ~100 ml of 1:1 70% ethanol:

ethylene glycol. A rain guard consisting of a clear 26 cm plastic plate was suspended ~3

cm above the trap using wire. To sample aerial arthropods and focusing on the

hymenoptera, pan traps were used consisting of two nested yellow 355 ml plastic bowls

(16 cm × 4 cm, Festive Occasion, East Providence, RI) with the internal bowl secured to

the outer bowl using binder clips and filled with ~75 ml soapy water. Traps were

mounted on 5 cm × 5 cm × 1 m wooden posts. Each trap type was deployed for 72 h

every 7 d. Following each deployment, contents were removed and strained through

paper filters (Rockline Industries, Sheboygan, WI), placed in sealable plastic bags with

70% ethanol, and stored at 4 °C until processing. In the laboratory arthropods were sorted

to order (Triplehorn and Johnson 2005) and counted. Coleopterans were further identified

to family (Marshall 2006, Evans 2014, BugGuide 2016). Coleopteran families are

relatively uniform trophically (Hammond 1990, Hammond 1992), which allows

classifying families into feeding guilds based on larval feeding behavior (Root 1967);

therefore I further assigned coleopteran families to trophic guilds based on larval feeding

habits, including fungivore, predator, herbivore, xylophage (including here xylomyceto-,

cambio-, and phloephagous members), saprophage, or parasitoid (Hammond 1990).

Scolytinae (Curculionidae) were further identified to species (Baker et al. 2009, Mercado

2010) and evaluated with respect to native vs non-native origin.

Analysis

The volume of CWD was compared across all decay classes in each plot using a

one-way analysis of variance (ANOVA), with a post hoc analysis using Tukey’s Honest

25

Significant Difference (HSD). For each monitoring approach, coleopteran abundance,

richness, diversity, and evenness were evaluated based on the presence of CWD.

Arthropod abundance measured with each trap type focused on coleopterans and was

analyzed by family and feeding guilds. Coleopteran diversity and richness were derived

for each sample approach (funnel, pitfall, and pan) within each plot. Richness is the total

number of taxa within a sample. Simpson’s diversity index utilizes the relative abundance

of each taxon and the total insect abundance within a sample (Magurran 2013), and is

calculated using 𝐷𝐷 = Σ𝑝𝑝𝑖𝑖2 𝑜𝑜𝑟𝑟 Σ 𝑛𝑛𝑖𝑖(𝑛𝑛𝑖𝑖−1)𝑁𝑁(𝑁𝑁−1)

. Shannon’s index, which utilizes both richness and

relative abundance and better represents my data consisting of many unique families with

low abundance, was also calculated using 𝐻𝐻′ = −Σ[(𝑝𝑝𝑖𝑖) ∗ 𝑙𝑙𝑙𝑙(𝑝𝑝𝑖𝑖)]. Evenness was

calculated using 𝐸𝐸 = 𝐻𝐻𝐻𝐻𝑚𝑚𝑚𝑚𝑚𝑚

𝑤𝑤ℎ𝑒𝑒𝑟𝑟𝑒𝑒 𝐻𝐻𝑚𝑚𝑚𝑚𝑚𝑚 = ln (𝑁𝑁) (Magurran 2013). Data over the 20-

week sampling period were grouped into five 4-week intervals (April, May, June, July,

and August) and summed. Data were tested for assumptions of normality (PROC

UNIVARIATE); abundance values were transformed using a square root transformation

to help reduce right skewness and because of many low and zero values. Significance

was determined at α = 0.05 unless stated otherwise. Arthropod abundance, richness,

Simpson’s and Shannon’s diversity indices, and Shannon’s evenness were analyzed using

a mixed linear model (PROC MIXED) to compare differences based on the presence or

absence of CWD and differences in time intervals (months) among coleopteran

population parameters, guilds, families, and scolytinae species (funnel traps only).

Sorenson’s index of similarity was computed to compare coleopteran family level

similarities between CWD+ and CWD–. All analyses were performed using SAS v9.3

(SAS Institute 2011).

26

Results

Coarse Woody Debris

The volume of ash coarse woody debris in the CWD+ plots averaged 0.7 m3,

whereas the volume in CWD– plots was 0. In CWD+ plots there were significant

differences in relative volume among decay classes (Fig. 3.2), with 89% (~0.62 m3) in

decay classes 1 and 2, and none present in decay class 5.

Arthropod Monitoring

All monitoring approaches collectively yielded 61,299 arthropods. Funnel traps

yielded 26,982 arthropods, including 24,107 coleopterans (89%) representing 54 families.

Pitfall traps yielded 16,878 arthropods, including 2,552 Coleoptera (15%) representing 30

families. Pan traps yielded 17,439 arthropods, including 2,864 Coleoptera (16%)

representing 46 families.

Funnel traps

In funnel traps there were no significant differences in overall coleopteran

abundance or evenness. However, coleopteran abundance was numerically greater in

association with ash CWD (11,560 in CWD+ vs. 8,663 in CWD–). Coleopteran family

richness and diversity (Shannon’s and Simpson’s) were significantly greater in the

presence of ash CWD (Table 3.1a). Similarly, there were significant temporal differences

in coleopteran population parameters, with a significant CWD × month interaction for

Simpson’s diversity index. Abundance was greatest in April and was driven by the sheer

number of xylophages (Appendix B.1(a)), which might also play a role in reducing

coleopteran evenness in April. Coleopteran family richness was greatest in June, and was

also significantly greater in the CWD+ treatment in June (p<0.01). Shannon’s diversity

27

was greatest in June and July, and Simpson’s diversity was greatest in May, June, and

July, and was significantly greater in the CWD+ treatment in August (p<0.01).

(Appendix C.1(a)).

There were significant differences in abundance between CWD treatments for

fungivores, herbivores, and saprophages captured in funnel traps; all three were

significantly more abundant in the presence of ash CWD, and the remaining feeding

guilds did not differ (Table 3.1b). Seasonal differences were evident for each guild (Table

3.1b). Fungivores were most abundant in April (Appendix B.1(b)). Herbivores were most

abundant in June and July, and saprophages in June. Saprophage abundance was also

significantly greater in association with ash CWD in June (p<0.01) (Appendix C.1(b))

Predator abundance followed a trend (p=0.07) of being greater in early spring and lower

at the end of the summer. Xylophages, consisting mainly of xylomycetophagous

scolytines, were significantly more abundant in April, with ~18,000 captured (Appendix

B.1(b)). Only seven parasitoids, represented by Catogenus rufus in the family

Passandridae, were captured in June and only in the funnel traps.

Of the 54 coleopteran families captured in funnel traps, 11 were significantly

more abundant in association with ash CWD and one was more abundant where ash

CWD was absent (Table 3.1c). Coleopteran abundance differed by month for most taxa

(Appendix B.1(c)), but only four taxa (Eucnemidae, Monotomidae, Ptilodactylidae, and

Silphidae) exhibited CWD × month interactions (Appendix C.1(c)). Sorenson’s index of

similarity of coleopteran families between CWD treatments equaled 0.90. The differences

among the coleopteran community in the CWD treatments included four unique families

in the CWD+ treatment, three of which were fungivores (Eucinetidae, Sphindidae, and

28

Throscidae) which feed on slime molds and basidiomycetes. There were six unique

families found in the absence of ash CWD; none were fungivores.

Of the 13 scolytinae species captured in funnel traps, none were significantly

different between CWD treatments (Table 3.2). This is likely because Scolytinae are

highly mobile; standing dead ash was scattered across the landscape and scolytines travel

large distances after emergence from infested trees. Scolytinae abundance differed by

month for most taxa (Table 3.2). Five taxa (Xyleborinus saxeseni, Ambrosiodmus

tachygraphus, Euwallacea validus, Monarthrum fasciatum, Xyleborus pelliculosus) were

significantly more abundant in April, and three taxa (Xyleborus ferrugineus, Xylosandrus

crassiuculus, Anisandrus sayi) were significantly more abundant in June (Appendix B.4).

Pitfall traps

Surprisingly, and in contrast to the funnel traps, Coleoptera activity density

measured by abundance in pitfall traps was significantly lower in association with ash

CWD (1,153 in CWD+ vs. 1,399 in CWD–), but there were no other differences in

population parameters between CWD treatments (Tables 3.4a). There were significant

temporal differences in coleopteran population parameters for all parameters, but there

were no significant CWD × month interactions. All population parameters were greatest

in June (Appendix B.2(a)).

Fungivore abundance in pitfall traps was significantly and unexpectedly lower in

association with ash CWD, and the remaining feeding guilds did not differ (Table 3.3b).

Seasonal differences were evident for each feeding guild. Fungivore abundance was

driven by the Nitidulidae, which were more abundant in the summer months.

29

Saphrophages were more abundant in June, predators in May and June, and xylophages

in April (Appendix B.2(b)).

Of the 30 coleopteran families captured in pitfall traps, Carabidae,

Chrysomelidae, and Nitidulidae were significantly more abundant where ash CWD was

absent. There was a weakly significant effect of CWD on Cryptophagidae and Elateridae;

these were more abundant in association with ash CWD present. (Table 3.3c).

Coleopteran abundance differed by month for most taxa and no taxa exhibited CWD ×

month interactions (Appendix C.2(c)). Over all coleopteran families, Sorenson’s index of

similarity was 0.87 between CWD+ and CWD- treatments. The difference consisted of

four unique families in the CWD+ treatment (Cryptophagidae, Ciidae, Mycetophagidae,

and Throscidae) which are all fungivores, and three families in the CWD– treatment

(Buprestidae, Geotrupide, and Ptinidae), two of which are fungivorous.

Pan traps

Coleoptera abundance in pan traps was significantly greater in association with

ash CWD (1,544 in CWD+ vs. 1,320 in CWD–) and coleopteran family richness and

diversity (Shannon’s and Simpson’s) were also greater in the presence of ash CWD

(Table 3.4a). Similarly, there were temporal differences in coleopteran population

parameters, and there were significant CWD × month interactions for Simpson’s diversity

index. Abundance, richness, and both diversity indices were greatest in June, and

evenness was greatest in May, June, and July (Appendix B.3(a)).

Herbivores, predators and saprophages captured in pan traps were significantly

more abundant in the presence of ash CWD, and the remaining feeding guilds did not

differ (Table 3.4b). Seasonal differences were evident for each feeding guild (Table

30

3.4b). The abundance of fungivores was greatest in June and in August. Herbivores,

saprophages and xylophages were most abundant in June. Predator abundance was

highest in April, May, and June. Xylophages were significantly greater in association

with ash CWD in April (p<0.01) (Appendix C.3(b)).

Of the 46 coleopteran families captured in pan traps, three were significantly

more abundant in association with ash CWD and three were more abundant where ash

CWD was absent (Table 3.4c). Coleopteran abundance differed by month for most taxa,

but only Scolytinae exhibited a CWD × month interaction (Table 3.4c), where abundance

in the CWD+ treatment was greatest in April (Appendix C.3(c)). Over all families,

Sorenson’s index of similarity equaled 0.84 between CWD treatments, with differences

consisting of eight unique families in the CWD+ treatment and five in the CWD–

treatment. Four of the unique families associated with ash CWD were fungivores (Ciidae,

Cryptophagidae, Ptillidae, and Zopheridae), and two unique fungivores (Cupedidae and

Synchroidae) found in the absence of ash CWD.

Discussion

EAB-induced ash mortality causes drastic changes in forest composition and

structure (Gandhi and Herms 2010a), resulting in significant inputs of ash woody

material and with substantial consequences for arthropod associates (Schowalter 2012). I

monitored arthropods in plots with significant downed ash and compared them to plots

lacking downed ash, and demonstrate discernable differences between coleopteran

communities associated with its presence or absence. The comparative approach

evaluating arthropods in the presence or absence of ash coarse woody debris provides a

31

means of estimating potential long-term changes in coleopteran community structure in

the wake of the emerald ash borer invasion.

I found that coleopteran family richness and diversity are associated with

increases in coarse woody debris in two of my three sampling approaches (funnel and pan

traps), and several coleopteran guilds, including fungivores, herbivores, saprophages, and

predators, appear to benefit from the influx of coarse woody debris to the forest floor.

Although not directly associated with the ash coarse woody debris, the appearance of the

parasitic passandrid, Catogenus rufus, in my study is not surprising given its previously

documented association with emerald ash borer invaded forests (Davidson and Rieske

2015). The increases in coarse woody debris associated with EAB-induced ash mortality

create and alter habitats, and local habitat characteristics influence arthropod activity

(Dauber et al. 2005). Thus, these differences in the coleopteran community I document

were anticipated given the varied microhabitats and differing roles occupied by different

feeding guilds. I used the coarser taxonomic resolution of family and guild in my analysis

because ‘taxonomic sufficiency’ recognizes that, within a community, changes at the

species level are often reflected at these coarser taxonomic levels (Ellis 1985, Birkhofer

et al., 2012); I found differences in the coarser taxonomic resolution based on the

presence or absence of ash coarse woody debris. However, even though coleopteran

families are relatively uniform trophically (Hammond 1990, Hammond 1992), the use of

coarser taxonomic identifications in ecological monitoring can be misleading (Longcore

2003), as not all genera in a given family function in the same trophic role.

In my study Sorenson’s index of similarity revealed that 84 – 90% of the

coleopteran community were similar regardless of the presence of ash coarse woody

32

debris, and plots associated with ash coarse woody debris had a similar number of unique

taxa (16) compared to plots with coarse woody debris absent (14). However, when

focusing on fungivores, the number of unique families associated with coarse woody

debris was substantially greater than in plots lacking coarse woody debris (11 vs 4

families). Fungivorous insect abundance is positively correlated with downed woody

material (Vanderwal et al. 2006). The increased abundance of fungivores in my study

suggests a potential functional change that may be occurring as coarse woody debris

volume increases due to EAB-induced ash mortality and all decay classes become

available.

The post-invasion forest plots supported an abundance of Scolytines utilizing the

stressed and standing dead ash, and fungivores utilizing the increasing volume and

variable state of decaying ash coarse woody debris. Bark and ambrosia beetles may play

a role in the initial process of decomposition of stressed and standing dead trees through

gallery construction and reproduction (Iidzuka et al. 2014), where the gallery entrances

can be a starting point for wood decomposition (Lindgren 1990). Their tunneling

activities further influence rates of decomposition by facilitating colonization by

microbes and other organisms, improving aeration, and promoting fragmentation

(Ulyshen 2014), potentially aiding in eventual community recovery (Schowalter 2012).

The tunneling activities of Scolytines can undermine the structural integrity of wood

(Blackman et al. 1924) hastening the fall of emerald ash borer-killed ash trees and

branches, which decay more quickly than standing or suspended wood (Swift et al. 1976).

Following the inputs of ash coarse woody debris, fungivores may then accelerate

33

decomposition of the downed ash resource, thus returning nutrients back to the system

more rapidly.

In my study, a single xylomycetophagous non-native Scolytinae, Xyleborinus

saxeseni (Ratzeburg), represented 89% of the Coleoptera captured in funnel traps (Table

3.6). Xyleborinus saxeseni is globally distributed and was among the first non-native

scolytids documented in North America (Rabaglia et al. 2006). It is also one of the most

damaging and potentially aggressive ambrosia beetles in the tribe Xyleborini in North

America (Rabaglia et al. 2006) feeding on ectosymbiotic fungi growing in wood and on

the wood itself (Deyrup and Atkinson 1987). My results suggest that EAB-induced ash

mortality at these heavily impacted sites has created and modified the forest, creating

optimal habitat and favoring subsequent colonization by Xyleborinus saxeseni. This

secondary invasion by a second non-native insect may be indicative of an ‘invasional

meltdown’ (Simberloff and Von Holle 1999), where each non-native invader enhances

the likelihood and success of subsequent non-native invaders. It’s possible that habitat

modifications by X. saxeseni and its fungal symbionts, through hastened tree fall and

decomposition, may favor the success of other non-native species, including their fungal

associates. If true, the influx of non-natives competing for the same resources as natives

could lead to displacement or local extinction of native insect associates, specifically

fungivores. Further research is needed to assess the impacts of X. saxeseni on these forest

systems following depletion of the ash resources.

My findings have conservation implications. Species richness of ground beetles is

positively correlated with downed woody material on a local scale (Gunnarsson et al.

2004), and 20 m3/ha of downed woody material is suggested to protect litter-dwelling

34

fauna (Kappes et al. 2009). The CWD+ plots contained 17.5 m3/ha of coarse woody

debris with many large diameter snags still standing, suggesting that post EAB-invaded

forests will attain or even surpass the suggested amount of coarse woody debris to

optimize habitat for ground-dwelling arthropods. Coarse woody material decomposes

much more slowly than foliage and fine woody material, creating more resilient habitat

and providing a long term source of nutrients (Harmon et al. 1986, Johnson and Curtis

2001, Greenberg 2002) that can be utilized by a variety of organisms (Hagan and Grove

1999, Åström et al. 2005). The increases in coarse woody debris following EAB

outbreaks could alter forest susceptibility to fire (Jenkins et al. 2008), however mesic

forests in the eastern U.S. are not typically prone to catastrophic fires (Evans 2011). After

evaluating biotic and abiotic risks following emerald ash borer invasion, conservationists

should optimize the amount of coarse woody debris retained to increase arthropod

richness and diversity, and to provide crucial habitat for a variety of wildlife (Evans

2011).

35

Fig. 3.1. Layout of plots, traps, and trap types used to evaluate the arthropod community

associated with ash coarse woody debris in emerald ash borer-impacted forests.

36

Fig. 3.2. Volume (m3/0.1 ac.) of ash coarse woody debris in each decay class (mean

(s.e.)) six years following invasion of emerald ash borer. Means followed by the same

letter are not significantly different at α = 0.05 (F3,29= 9.58; P< 0.01).

a

a

bb

0.00

0.10

0.20

0.30

0.40

0.50

1 2 3 4 5

CWD

Volu

me

/ plo

t (m

3/0.

1 ac

)

Decay Class

Table 3.1. Effects of the presence of coarse woody debris and of season (Fdf/P) on coleopteran a) population parameters, b) feeding

guild abundance, and c) family abundance sampled from funnel traps in EAB invaded forests of north-central Kentucky. Means

followed by the same letter do not significantly differ (P<0.05).

Funnel traps CWD CWD (mean (s.e.)) Month CWD × Month

F1,342/P Present Absent F4,342/P F4,342/P

(a) Population parameters Abundance 2.36/0.13 6.38 (0.53)a 5.56 (0.53)a 90.83/<0.01 0.42/0.79

Richness 16.33/<0.01 6.19 (0.24)a 5.22 (0.24)b 43.33/<0.01 1.56/0.19 Shannon's diversity 11.27/<0.01 1.25 (0.04)a 1.10 (0.04)b 134.15/<0.01 1.00/0.41 Shannon's evenness 2.34/0.13 0.73 (0.03)a 0.70 (0.03)a 97.47/<0.01 2.27/0.07 Simpson's diversity 10.56/<0.01 0.58 (0.03)a 0.49 (0.03)b 77.65/<0.01 2.95/0.03 (b) Feeding guild

Fungivore 15.42/<0.01 3.24 (0.18)a 2.54 (0.18)b 22.3/<0.01 0.66/0.62 Herbivore 8.91/<0.01 3.50 (0.19)a 2.95 (0.19)b 23.2/<0.01 0.11/0.98 Saprophage 8.0/<0.01 2.36 (0.16)a 1.92 (0.16)b 157.8/<0.01 2.06/0.10 Xylophage 0.71/0.40 9.07 (1.06)a 8.17 (1.06)a 114.3/<0.01 0.64/0.63 Predator 3.29/0.07 3.38 (0.18)a 3.06 (0.18)a 17.21/<0.01 0.86/0.49 Parasitoid 0.03/0.86 0.07 (0.05)a 0.08 (0.05)a 7.72/<0.01 0.03/0.99 (c) Families

Aderidae 4.46/0.04 0.07 (0.08)b 0.23 (0.08)a 3.33/0.01 0.74/0.57

37

Table 3.1 (continued)

Funnel traps CWD CWD (mean (s.e.)) Month CWD × Month

F1,342/P Present Absent F4,342/P F4,342/P

Cerylonidae 7.77/<0.01 0.51 (0.10)a 0.24 (0.10)b 19.48/<0.01 1.95/0.11 Ciidae 6.25/0.01 0.28 (0.09)a 0.04 (0.09)b 1.23/0.31 0.36/0.84 Endomychidae 3.95/0.05 0.33 (0.09)a 0.15 (0.09)b 13.75/<0.01 1.06/0.38 Eucnemidae 6.58/0.01 0.47 (0.12)a 0.16 (0.12)b 7.83/<0.01 2.71/0.04 Monotomidae 4.78/0.03 0.15 (0.05)a 0.02 (0.05)b 3.27/0.02 2.26/0.07 Mordellidae 10.36/<0.01 1.42 (0.15)a 0.93 (0.15)b 31.42/<0.01 1.97/0.11 Ptilodactylidae 6.29/0.01 2.26 (0.15)a 1.89 (0.15)b 177.92/<0.01 2.42/0.06 Silphidae 5.99/0.02 0.13 (0.05)a 0.00 (0.05)b 2.36/0.06 2.36/0.06 Staphylinidae 4.03/0.05 3.08 (0.18)a 2.71 (0.18)b 10.92/<0.01 0.30/0.88 Melandryidae 3.13/0.08 0.37 (0.09)a 0.21 (0.09)b 10.16/<0.01 2.19/0.08 Synchroidae 3.10/0.08 0.30 (0.10)a 0.13 (0.10)b 3.30/0.02 0.56/0.69

38

Table 3.2. Effects of the presence of coarse woody debris and of season (Fdf/P) on scolytinae species abundance (S.E.) sampled from

funnel traps in EAB invaded forests of north-central Kentucky in 2015. Means followed by the same letter do not significantly differ

(P<0.05).

Scolytinae Species CWD CWD (s.e.) Month CWD × Month

Funnel traps F1,16/P Present Absent F4,64/P F4,64/P

Xyleborinus saxeseni 0.88/0.36 7.95 (0.82)a 6.86 (0.82)a 112.21/<0.01 0.61/0.66

Ambrosiodmus tachygraphus 0.00/0.96 0.26 (0.05)a 0.26 (0.05)a 60.98/<0.01 0.00/1.00

Euwallacea validus 2.73/0.12 0.56 (0.08)a 0.38 (0.08)a 37.76/<0.01 1.81/0.14

Hypothenemus dissimilis 0.00/1.00 0.02 (0.02)a 0.02 (0.02)a 0.75/0.56 1.25/0.30

Monarthrum fasciatum 0.18/0.68 1.53 (0.12)a 1.46 (0.12)a 229.47/<0.01 0.17/0.96

Monarthrum mali 1.00/0.33 0.02 (0.02)a 0.00 (0.02)a 1.00/0.41 1.00/0.41

Pityophthorus concentralis 1.00/0.33 0.02 (0.02)a 0.00 (0.02)a 1.00/0.41 1.00/0.41

Pseudothysannoes dislocatus 0.39/0.54 0.04 (0.04)a 0.08 (0.04)a 1.09/0.37 1.05/0.39

Xyleborus ferrugineus 0.36/0.55 0.04 (0.03)a 0.02 (0.03)a 3.27/0.02 0.36/0.83

Xyleborus pelliculosus 0.14/0.71 1.33 (0.18)a 1.23 (0.18)a 104.42/<0.01 0.26/0.90

Xylosandrus crassiusculus 1.23/0.28 0.49 (0.11)a 0.66 (0.11)a 10.45/<0.01 0.61/0.66

Xylosandrus germanus 0.86/0.37 0.14 (0.07)a 0.23 (0.07)a 0.77/0.55 0.16/0.96

39

Table 3.2 (continued)

Scolytinae Species CWD CWD (s.e.) Month CWD × Month

Funnel traps F1,16/P Present Absent F4,64/P F4,64/P

Anisandrus sayi 1.55/0.23 0.90 (0.14)a 1.16 (0.14)a 61.87/<0.01 0.59/0.67

40

Table 3.3. Effects of the presence of coarse woody debris and of season (Fdf/P) on coleopteran a) population parameters, b) feeding

guild abundance, and c) family abundance sampled from pitfall traps in EAB invaded forests of north-central Kentucky. Means

followed by the same letter do not significantly differ (P<0.05).

Pitfall traps CWD CWD (s.e) Month CWD × Month

F1,342/P Present Absent F4,342/P F4,342/P

(a) Population

Abundance 4.01/0.05 2.42 (0.13)b 2.68 (0.13)a 14.77/<0.01 1.01/0.41

Richness 0.39/0.53 2.68 (0.15)a 2.77 (0.15)a 41.19/<0.01 0.66/0.62 Shannon's diversity 0.06/0.81 0.72 (0.05)a 0.73 (0.05)a 31.18/<0.01 0.54/0.71 Shannon's evenness 1.23/0.27 0.64 (0.04)a 0.60 (0.04)a 17.84/<0.01 0.32/0.86 Simpson's diversity 0.70/0.41 0.40 (0.03)a 0.42 (0.03)a 23.41/<0.01 1.20/0.32 (b) Feeding guild

Fungivore 7.76/<0.01 3.02 (0.24)b 3.69 (0.24)a 36.76/<0.0 0.92/0.45 Herbivore 0.00/0.97 1.29 (0.16)a 1.30 (0.16)a 3.40/0.01 0.66/0.62 Saprophage 0.08/0.78 0.33 (0.15)a 0.38 (0.15)a 10.02/<0.0 0.35/0.84 Xylophage 0.42/0.52 1.23 (0.15)a 1.13 (0.15)a 36.34/<0.0 0.30/0.88 Predator 0.08/0.78 2.50 (0.17)a 2.55 (0.17)a 14.95/<0.0 0.88/0.48 Parasitoid N/A N/A N/A N/A N/A (c) Families

Carabidae 3.92/0.05 1.54 (0.18)b 1.90 (0.18)a 9.13/<0.01 0.50/0.74 Cryptophagidae 3.18/0.08 0.07 (0.04)a 0.00 (0.04)b 0.53/0.71 0.53/0.71

41

Table 3.3 (continued) Pitfall traps CWD CWD (s.e) Month CWD × Month

F1,342/P Present Absent F4,342/P F4,342/P

Chrysomelidae 7.48/<0.01 0.12 (0.09)b 0.36 (0.09)a 1.49/0.22 2.16/0.08 Elateridae 3.50/0.07 0.17 (0.06)a 0.05 (0.06)b 7.40/<0.01 1.14/0.35 Nitidulidae 8.68/<0.01 2.73 (0.24)b 3.43 (0.24)a 38.26/<0.01 0.77/0.55

42

Table 3.4. Effects of the presence of coarse woody debris and of season (Fdf/P) on coleopteran a) population parameters, b) feeding

guild abundance, and c) family abundance sampled from pan traps in EAB invaded forests of north-central Kentucky. Means followed

by the same letter do not significantly differ (P<0.05).

Pan traps CWD CWD (s.e) Month CWD × Month

F1,342/P Present Absent F4,342/P F4,342/P

(a) Population

Abundance 5.80/0.02 2.73 (0.09)a 2.51 (0.09)b 102.43/ <0.01 0.47/0.76

Richness 5.42/0.02 3.84 (0.19)a 3.41 (0.19)b 107.93/<0.01 0.17/0.95 Shannon's diversity 5.48/0.02 0.97 (0.05)a 0.86 (0.05)b 77.66/<0.01 0.41/0.80 Shannon's evenness 1.56/0.22 0.68 (0.03)a 0.63 (0.03)a 34.16/<0.01 0.58/0.68 Simpson's diversity 8.34/<0.01 0.49 (0.03)a 0.39 (0.03)b 36.49/<0.01 3.56/0.01 (b) Feeding guild

Fungivore 0.74/0.64 2.27 (0.17)a 2.42 (0.17)a 20.76/<0.01 0.84/0.51 Herbivore 5.56/0.02 3.43 (0.20)a 2.97 (0.20)b 89.67/<0.01 1.24/0.30 Saprophage 0.01/0.91 1.10 (0.12)a 1.11 (0.12)a 82.61/<0.01 0.33/0.86 Xylophage 7.17/0.01 1.23 (0.15)a 0.83 (0.15)b 29.57/<0.01 2.93/0.03 Predator 4.38/0.04 2.14 (0.13)a 1.86 (0.13)b 41.44/<0.01 0.26/0.90 Parasitoid N/A N/A N/A N/A N/A (c) Families

Chrysomelidae 13.40/<0.01 2.02 (0.15)a 1.46 (0.15)b 71.33/<0.01 2.24/0.07

43

Table 3.4 (continued)

Pan traps CWD CWD (s.e) Month CWD × Month

F1,342/P Present Absent F4,342/P F4,342/P

Cupedidae 3.21/0.08 0.00 (0.04)b 0.07 (0.04)a 1.43/0.23 1.43/0.23 Phalacridae 3.21/0.08 0.07 (0.04)a 0.00 (0.04)b 1.43/0.23 1.43/0.23 Ptinidae 2.96/0.09 0.17 (0.09)b 0.33 (0.09)a 8.75/<0.01 1.14/0.35 Scolytinae 6.87/0.01 0.74 (0.13)a 0.40 (0.13)b 19.63/<0.01 4.57/<0.01 Throscidae 2.78/0.10 0.12 (0.11)b 0.31 (0.11)a 4.27/<0.01 0.62/0.65

44

45

APPENDICES

46

APPENDIX A

Trophic guild designations

Coleopteran family abundance relative to ash canopy decline at five forested sites in

north central Kentucky with trophic guild designations including; herbivores (H),

fungivores (F), predators (P), saprophages (S), xylophages (X), and parasitoids (Pa).

Coleopteran Families

Trophic Guilds

Abundance Fayette Spencer Shelby Anderson Henry Absolute

Elateridae H 205 345 678 353 240 1,821 Chrysomelidae H 127 176 272 56 55 686 Curculionidae H 81 127 154 99 113 574 Tenebrionidae H 94 83 142 54 154 527 Mordellidae H 48 103 148 75 59 433 Scarabaeidae H 20 24 20 23 23 110 Phalacridae H 2 5 26 6 12 51 Attelabidae H 0 0 2 0 3 5 Latridiidae F 141 33 274 230 340 1,018 Corylophidae F 21 2 73 41 94 231 Ptinidae F 16 47 38 36 13 150 Eucnemidae F 16 50 33 39 8 146 Erotylidae F 14 14 8 16 17 69 Mycetophagidae F 22 5 16 6 10 59 Tetratomidae F 3 12 7 21 15 58 Nitidulidae F 7 13 11 11 13 55 Cerylonidae F 15 4 8 13 7 47 Zopheridae F 6 10 5 13 5 39 Silvanidae F 1 22 1 6 7 37 Melandryidae F 5 11 10 6 4 36 Synchroidae F 4 4 6 9 9 32 Endomychidae F 7 1 6 4 4 22 Leiodidae F 5 2 3 8 4 22 Cryptophagidae F 0 1 4 3 0 8 Laemophloeidae F 3 1 1 2 1 8 Anthribidae F 4 1 0 1 0 6 Cucujidae F 0 0 1 1 0 2 Pyrochoidae F 0 0 1 0 1 2 Sphindidae F 0 0 1 1 0 2 Throscidae F 0 1 0 0 1 2 Salpingidae F 1 0 0 0 0 1

47

Staphylinidae P 225 224 356 443 192 1,440 Histeridae P 188 316 141 204 136 985 Trogossitidae P 27 42 46 43 70 228 Carabidae P 16 20 63 43 34 176 Cleridae P 7 27 31 31 48 144 Lampyridae P 3 14 14 13 4 48 Coccinellidae P 5 7 4 0 2 18 Melyridae P 1 0 1 0 4 6 Cantharidae P 0 2 1 0 0 3 Hydrophilidae P 0 1 0 0 0 1 Lycidae P 0 0 0 1 0 1 Ptilodactylidae S 139 131 480 266 40 1,056 Dermestidae S 7 26 1 6 7 47 Monotomidae S 2 6 0 0 0 8 Scirtidae S 0 1 0 3 0 4 Hybosoridae S 2 0 0 1 0 3 Silphidae S 0 Scolytinae X 168 465 107 117 134 991 Scraptiidae X 1 1 8 24 53 87 Cerambycidae X 6 13 12 19 24 74 Bostrichidae X 1 5 2 0 3 11 Buprestidae X 0 0 2 2 5 9 Lucanidae X 0 0 0 1 0 1 Lymexylidae X 0 0 0 1 0 1 Passandridae Pa 21 13 14 16 14 78 Rhipiceridae Pa 1 0 0 1 2 4 Bothrideridae Pa 0 3 0 0 0 3 Unidentified --- 33 22 9 21 15 100 Total 1,721 2,436 3,241 2,389 1,999 11,786

APPENDIX B

Temporal data by month

Temporal effects on the coleopteran community (mean (s.e.)) for coleopteran a) population parameters, b) feeding guild abundance,

and c) family abundance (for families where CWD treatment interaction α=0.1) sampled from all trap types in EAB invaded forests of

north-central Kentucky. Means followed by the same letter do not significantly differ (P<0.05).

Month

April May June July August

(I) FUNNEL TRAPS (a) Population parameters Abundance 15.9 (0.6)a 3.7 (0.6)bc 4.9 (0.6)b 3.3 (0.6)bc 2.0 (0.6)c Richness 5.7 (0.3)b 5.6 (0.3)b 8.2 (0.3)a 5.8 (0.3)b 3.2 (0.3)c Shannon's diversity 0.3 (0.1)d 1.3 (0.1)b 1.7 (0.1)a 1.5 (0.1)a 1.0 (0.3)c Simpson's diversity 0.1 (0.0)d 0.6 (0.0)b 0.8 (0.0)a 0.8 (0.0)ab 0.4 (0.0)c Shannon's evenness 0.2 (0.0)b 0.8 (0.0)a 0.8 (0.0)a 0.9 (0.0)a 0.8 (0.0)a (b) Feeding guild Fungivore 4.5 (0.2)a 2.8 (0.2)b 2.6 (0.2)b 2.3 (0.2)b 2.3 (0.2)b Herbivore 2.3 (0.2)bc 3.0 (0.2)b 4.5 (0.2)a 4.0 (0.2)a 2.2 (0.2)c Saprophage 0.2 (0.2)d 1.8 (0.2)c 5.4 (0.2)a 2.9 (0.2)b 0.3 (0.2)d Xylophage 31.1 (1.2)a 4.7 (1.2)b 4.7 (1.2)b 1.8 (1.2)b 0.9 (1.2)b Predator 4.2 (0.2)a 3.3 (0.2)b 3.5 (0.2)ab 2.9 (0.2)b 2.1 (0.2)c Parasitoid 0.0 (0.1)b 0.0 (0.1)b 0.4 (0.1)a 0.0 (0.1)b 0.0 (0.1)b

48

(c) Families Aderidae 0.0 (0.1)b 0.1 (0.1)ab 0.2 (0.1)ab 0.4 (0.1)a 0.0 (0.1)b Cerylonidae 1.2 (0.1)a 0.3 (0.1)b 0.4 (0.1)b 0.0 (0.1)b 0.0 (0.1)b Ciidae 0.2 (0.1)a 0.1 (0.1)a 0.1 (0.1)a 0.3 (0.1)a 0.1 (0.1)a Endomychidae 0.9 (0.1)a 0.1 (0.1)b 0.1 (0.1)b 0.1 (0.1)b 0.0 (0.1)b Eucnemidae 0.0 (0.1)b 0.4 (0.1)ab 0.9 (0.1)a 0.2 (0.1)b 0.1 (0.1)b Monotomidae 0.2 (0.1)a 0.0 (0.1)a 0.2 (0.1)a 0.0 (0.1)a 0.0 (0.1)a Mordellidae 0.0 (0.2)c 0.4 (0.2)b 2.1 (0.2)a 2.0 (0.2)a 1.5 (0.2)a Ptilodactylidae 0.0 (0.2)d 1.8 (0.2)c 5.4 (0.2)a 2.9 (0.2)b 0.3 (0.2)d Silphidae 0.0 (0.1)a 0.0 (0.1)a 0.1 (0.1)a 0.2 (0.1)a 0.0 (0.1)a Staphylinidae 3.7 (0.2)a 3.1 (0.2)ab 3.1 (0.2)ab 2.7 (0.2)b 1.9 (0.2)c Melandryidae 0.0 (0.1)c 0.5 (0.1)ab 0.0 (0.1)c 0.2 (0.1)bc 0.8 (0.1)a Synchroidae 0.0 (0.1)b 0.1 (0.1)ab 0.4 (0.1)a 0.2 (0.1)ab 0.4 (0.1)ab (II) PITFALL TRAPS (a) Population parameters Abundance 1.9 (0.1)c 2.4 (0.1)b 3.3 (0.1)a 2.4 (0.1)b 2.8 (0.1)b Richness 1.6 (0.2)c 2.8 (0.2)b 4.5 (0.2)a 2.4 (0.2)b 2.4 (0.2)b Shannon's diversity 0.4 (0.1)c 0.8 (0.1)b 1.2 (0.1)a 0.6 (0.1)b 0.6 (0.1)b Simpson's diversity 0.2 (0.0)c 0.4 (0.0)b 0.7 (0.0)a 0.4 (0.0)b 0.3 (0.0)bc Shannon's evenness 0.4 (0.0)d 0.7 (0.0)ab 0.8 (0.0)a 0.6 (0.0)c 0.6 (0.0)bc (b) Feeding guild Fungivore 0.7 (0.3)c 3.0 (0.3)b 4.6 (0.3)a 3.8 (0.3)ab 4.6 (0.3)a Herbivore 1.0 (0.2)b 1.0 (0.2)ab 1.6 (0.2)ab 1.3 (0.2)ab 1.7 (0.2)a Saprophage 0.0 (0.2)b 0.3 (0.2)b 1.3 (0.2)a 0.2 (0.2)b 0.0 (0.2)b Xylophage 2.7 (0.2)a 0.9 (0.2)c 1.7 (0.2)b 0.3 (0.2)c 0.4 (0.2)c Predator 1.8 (0.2)c 2.9 (0.2)ab 3.6 (0.2)a 2.0 (0.2)c 2.2 (0.2)bc (c) Families

49

Carabidae 0.8 (0.2)c 1.8 (0.2)ab 2.5 (0.2)a 1.6 (0.2)b 1.8 (0.2)ab Cryptophagidae 0.1 (0.0)a 0.1 (0.0)a 0.1 (0.0)a 0.0 (0.0)a 0.0 (0.0)a Chrysomelidae 0.2 (0.1)a 0.1 (0.1)a 0.4 (0.1)a 0.2 (0.1)a 0.2 (0.1)a Elateridae 0.0 (0.1)b 0.1 (0.1)b 0.5 (0.1)a 0.1 (0.1)b 0.0 (0.1)b Nitidulidae 0.5 (0.3)c 2.4 (0.3)b 4.2 (0.3)a 3.7 (0.3)a 4.5 (0.3)a (III) PAN TRAPS (a) Population parameters Abundance 2.2 (0.1)bc 2.0 (0.1)c 4.4 (0.1)a 2.6 (0.1)b 1.9 (0.1)c Richness 2.0 (0.2)d 2.9 (0.2)c 7.3 (0.2)a 3.9 (0.2)b 2.1 (0.2)d Shannon's diversity 0.4 (0.1)c 0.9 (0.1)b 1.6 (0.1)a 1.0 (0.1)b 0.6 (0.1)c Simpson's diversity 0.2 (0.0)c 0.4 (0.0)b 0.8 (0.0)a 0.5 (0.0)b 0.3 (0.0)b Shannon's evenness 0.3 (0.0)c 0.8 (0.0)a 0.8 (0.0)a 0.8 (0.0)a 0.6 (0.0)b (b) Feeding guild Fungivore 1.0 (0.2)c 2.3 (0.2)b 3.3 (0.2)a 2.3 (0.2)b 2.9 (0.2)ab Herbivore 2.1 (0.2)c 1.5 (0.2)c 6.5 (0.2)a 4.0 (0.2)b 2.0 (0.2)c Saprophage 0.1 (0.1)d 0.7 (0.1)c 3.1 (0.1)a 1.3 (0.1)b 0.4 (0.1)cd Xylophage 2.0 (0.2)a 0.5 (0.2)b 2.0 (0.2)a 0.5 (0.2)b 0.1 (0.2)b Predator 2.8 (0.2)a 2.4 (0.2)a 2.8 (0.2)a 1.5 (0.2)b 0.6 (0.2)c (c) Families Chrysomelidae 1.1 (0.2)c 0.4 (0.2)d 4.1 (0.2)a 2.1 (0.2)b 1.1 (0.2)c Cupedidae 0.0 (0.0)a 0.0 (0.0)a 0.1 (0.0)a 0.1 (0.0)a 0.0 (0.0)a Phalacridae 0.1 (0.0)a 0.1 (0.0)a 0.0 (0.0)a 0.0 (0.0)a 0.0 (0.0)a Ptinidae 0.0 (0.1)b 0.1 (0.1)b 0.7 (0.1)a 0.4 (0.1)ab 0.1 (0.1)b Scolytinae 1.7 (0.1)a 0.2 (0.1)b 0.6 (0.1)b 0.3 (0.1)b 0.1 (0.1)b Throscidae 0.0 (0.1)b 0.5 (0.1)a 0.5 (0.1)ab 0.1 (0.1)ab 0.0 (0.1)b (IV) Scolytines Xyleborinus saxeseni 29.35 (1.17)a 4.13 (1.17)b 2.83 (1.17)b 0.30 (1.17)b 0.41 (1.17)b

50

Ambrosiodmus tachygraphus 1.29 (0.07)a 0.00 (0.07))b 0.00 (0.07)b 0.00 (0.07)b 0.00 (0.07)b Euwallacea validus 1.70 (0.12)a 0.17 (0.12)bc 0.47 (0.12)b 0.00 (0.12)c 0.00 (0.12)c Hypothenemus dissimilis 0.06 (0.04)a 0.00 (0.04)a 0.00 (0.04)a 0.00 (0.04)a 0.06 (0.04)a Monarthrum fasciatum 6.72 (0.19)a 0.59 (0.19)b 0.11 (0.19)b 0.06 (0.19)b 0.00 (0.19)b Monarthrum mali 0.06 (0.02)a 0.00 (0.02)a 0.00 (0.02)a 0.00 (0.02)a 0.00 (0.02)a Pityophthorus concentralis 0.06 (0.02)a 0.00 (0.02)a 0.00 (0.02)a 0.00 (0.02)a 0.00 (0.02)a Pseudothysannoes dislocatus 0.00 (0.06)a 0.11 (0.06)a 0.13 (0.06)a 0.06 (0.06)a 0.00 (0.06)a Xyleborus ferrugineus 0.00 (0.04)b 0.00 (0.04)b 0.17 (0.04)a 0.00 (0.04)b 0.00 (0.04)b Xyleborus pelliculosus 6.33 (0.28)a 0.00 (0.28)b 0.00 (0.28)b 0.00 (0.28)b 0.06 (0.28)b Xylosandrus crassiusculus 0.64 (0.14)bc 0.69 (0.14)ab 1.14 (0.14)a 0.17 (0.14)c 0.22 (0.14)c Xylosandrus germanus 0.17 (0.10)a 0.06 (0.10)a 0.25 (0.10)a 0.28 (0.10)a 0.19 (0.10)a Anisandrus sayi 0.25 (0.17)c 0.81 (0.17)b 3.01 (0.17)a 1.11 (0.17)b 0.00 (0.17)c

51

APPENDIX C

CWD × Month treatment interactions

Effects of the presence of coarse woody debris (CWD+ mean (s.e.); CWD- mean (s.e)) by month on coleopteran a) population

parameters, b) feeding guild abundance, and c) family abundance (for families where CWD treatment interaction α=0.1) sampled from

all trap types in EAB invaded forests of north-central Kentucky. Asterisks following a value indicate a significant CWD × Month

interaction for that parameter.

Month

April May June July August

(I) FUNNEL TRAPS (a) Population parameters Abundance 17.0(0.8); 14.8(0.8) 3.9(0.8); 3.5(0.8) 5.2(0.8); 4.5(0.8) 3.4(0.8); 3.1(0.8) 2.3(0.8); 1.8(0.8) Richness 5.8(0.4); 5.6(0.4) 6.2(0.4); 4.9(0.4) 9.2(0.4); 7.3(0.4)* 6.1(0.4); 5.5(0.4) 3.7(0.4); 2.9(0.4) Shannon's diversity 0.3(0.1); 0.3(0.1) 1.4(0.1); 1.2(0.1) 1.8(0.1); 1.6(0.1) 1.6(0.1); 1.5(0.1) 1.2(0.1); 0.9(0.1) Simpson's diversity 0.2(0.0); 0.2(0.0) 0.8(0.0); 0.8(0.0) 0.8(0.0); 0.8(0.0) 0.9(0.0); 0.9(0.0) 0.9(0.0); 0.9(0.0) Shannon's evenness 0.1(0.1); 0.1(0.1) 0.7(0.1); 0.6(0.1) 0.8(0.1); 0.8(0.1) 0.8(0.1); 0.7(0.1) 0.5(0.1); 0.2(0.1)* (b) Feeding guild Fungivore 5.0(0.3); 4.1(0.3) 3.1(0.3); 2.5(0.3) 3.1(0.3); 2.0(0.4) 2.4(0.3); 2.2(0.3) 2.6(0.3); 1.9(0.3) Herbivore 2.7(0.3); 2.0(0.3) 3.3(0.3); 2.8(0.3) 4.8(0.3); 4.2(0.3) 4.2(0.3); 3.8(0.3) 2.5(0.3); 2.0(0.3) Saprophage 0.3(0.3); 0.1(0.3) 1.8(0.3); 1.8(0.3) 6.2(0.3); 4.8(0.3)* 3.0(0.3); 2.9(0.3) 0.5(0.3); 0.0(0.3) Xylophage 33.2(1.7); 29.0(1.7) 5.0(1.7); 4.4(1.7) 4.7(1.7); 4.8(1.7) 1.6(1.7); 1.9(1.7) 0.9(1.7); 0.8(1.7) Predator 4.2(0.3); 4.4(0.3) 3.4(0.3); 3.2(0.3) 3.8(0.3); 3.3(0.3) 3.2(0.3); 2.5(0.3) 2.3(0.3); 1.9(0.3)

52

Parasitoid 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) 0.3(0.1); 0.4(0.1) 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) (c) Families Aderidae 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.3(0.1) 0.1(0.1); 0.4(0.1) 0.2(0.1); 0.5(0.1) 0.0(0.1); 0.0(0.1) Cerylonidae 1.3(0.2); 1.1(0.2) 0.6(0.2); 0.0(0.2) 0.7(0.2); 0.1(0.2) 0.0(0.2); 0.0(0.2) 0.0(0.2); 0.0(0.2) Ciidae 0.3(0.1); 0.1(0.1) 0.2(0.1); 0.0(0.1) 0.1(0.1); 0.0(0.1) 0.5(0.1); 0.1(0.1) 0.2(0.1); 0.0(0.1) Endomychidae 1.2(0.1); 0.7(0.1) 0.2(0.1); 0.1(0.1) 0.2(0.1); 0.0(0.1) 0.1(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) Eucnemidae 0.0(0.2); 0.0(0.2) 0.7(0.2); 0.1(0.2) 1.4(0.2); 0.5(0.2)* 0.2(0.2); 0.1(0.2) 0.0(0.2); 0.1(0.2) Monotomidae 0.3(0.1); 0.1(0.1) 0.0(0.1); 0.0(0.1) 0.4(0.1); 0.0(0.1)* 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) Mordellidae 0.0(0.2); 0.0(0.2) 0.3(0.2); 0.4(0.2) 2.5(0.2); 1.7(0.2) 2.3(0.2); 1.6(0.2) 1.9(0.2); 1.0(0.2) Ptilodactylidae 0.0(0.2); 0.0(0.2) 1.8(0.2); 1.8(0.2) 6.0(0.2); 4.8(0.2)* 3.0(0.2); 2.9(0.2) 0.5(0.2); 0.0(0.2) Silphidae 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) 0.3(0.1); 0.0(0.1) 0.4(0.1); 0.0(0.1)* 0.0(0.1); 0.0(0.1) Staphylinidae 3.8(0.3); 3.6(0.3) 3.2(0.3); 3.0(0.3) 3.3(0.3); 3.0(0.3) 3.1(0.3); 2.3(0.3) 2.0(0.3); 1.7(0.3) Melandryidae 0.0(0.2); 0.0(0.2) 0.7(0.2); 0.3(0.2) 0.0(0.2); 0.0(0.2) 0.1(0.2); 0.3(0.2) 1.1(0.2); 0.5(0.2) Synchroidae 0.0(0.2); 0.0(0.2) 0.1(0.2); 0.0(0.2) 0.7(0.2); 0.2(0.2) 0.3(0.2); 0.1(0.2) 0.4(0.2); 0.3(0.2) (II) PITFALL TRAPS (a) Population parameters Abundance 2.0(0.2); 1.8(0.2) 2.2(0.2); 2.6(0.2) 3.3(0.2); 3.4(0.2) 2.2(0.2); 2.6(0.2) 2.5(0.2); 3.0(0.2) Richness 1.6(0.2); 1.5(0.2) 2.6(0.2); 3.0(0.2) 4.6(0.2); 4.4(0.2) 2.3(0.2); 2.4(0.2) 2.3(0.2); 2.5(0.2) Shannon's diversity 0.4(0.1); 0.4(0.1) 0.8(0.1); 0.9(0.1) 1.2(0.1); 1.2(0.1) 0.6(0.1); 0.6(0.1) 0.6(0.1); 0.6(0.1) Simpson's diversity 0.4(0.1); 0.3(0.1) 0.7(0.1); 0.7(0.1) 0.8(0.1); 0.8(0.1) 0.6(0.1); 0.5(0.1) 0.6(0.1); 0.6(0.1) Shannon's evenness 0.2(0.1); 0.2(0.1) 0.4(0.1); 0.5(0.1) 0.7(0.1); 0.7(0.1) 0.4(0.1); 0.4(0.1) 0.3(0.1); 0.3(0.1) (b) Feeding guild Fungivore 0.7(0.4); 0.7(0.4) 2.6(0.4); 3.4(0.4) 4.4(0.4); 4.8(0.4) 3.3(0.4); 4.4(0.4) 4.0(0.4); 5.2(0.4) Herbivore 1.1(0.3); 0.8(0.3) 1.0(0.3); 0.9(0.3) 1.4(0.3); 1.7(0.3) 1.2(0.3); 1.5(0.3) 1.7(0.3); 1.6(0.3) Saprophage 0.0(0.2); 0.0(0.2) 0.2(0.2); 0.3(0.2) 1.1(0.2); 1.4(0.2) 0.3(0.2); 0.1(0.2) 0.0(0.2); 0.0(0.2) Xylophage 2.9(0.2); 2.6(0.2) 0.8(0.2); 0.9(0.2) 1.8(0.2); 1.5(0.2) 0.3(0.2); 0.2(0.2) 0.3(0.2); 0.4(0.2) Predator 1.9(0.3); 1.8(0.3) 2.7(0.3); 3.2(0.3) 3.9(0.3); 3.4(0.3) 2.0(0.3); 2.1(0.3) 2.1(0.3); 2.3(0.3)

53

(c) Families Carabidae 0.6(0.3); 1.0(0.3) 1.5(0.3); 2.2(0.3) 2.5(0.3); 2.5(0.3) 1.5(0.3); 1.8(0.3) 1.7(0.3); 2.0(0.3) Cryptophagidae 0.1(0.1); 0.0(0.1) 0.1(0.1); 0.0(0.1) 0.1(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) Chrysomelidae 0.0(0.1); 0.4(0.1) 0.0(0.1); 0.2(0.1) 0.2(0.1); 0.7(0.1) 0.1(0.1); 0.4(0.1) 0.3(0.1); 0.1(0.1) Elateridae 0.0(0.1); 0.0(0.1) 0.1(0.1); 0.0(0.1) 0.6(0.1); 0.3(0.1) 0.1(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) Nitidulidae 0.5(0.4); 0.5(0.4) 2.1(0.4); 2.8(0.4) 4.0(0.4); 4.5(0.4) 3.2(0.4); 4.2(0.4) 3.9(0.4); 5.1(0.4) (III) PAN TRAPS (a) Population parameters Abundance 2.3(0.1); 2.1(0.1) 2.1(0.1); 1.8(0.1) 4.6(0.1); 4.2(0.1) 2.7(0.1); 2.5(0.1) 1.9(0.1); 1.9(0.1) Richness 2.1(0.3); 1.9(0.3) 3.2(0.3); 2.6(0.3) 7.6(0.3); 7.0(0.3) 4.1(0.3); 3.6(0.3) 2.3(0.3); 1.9(0.3) Shannon's diversity 0.5(0.1); 0.4(0.1) 1.0(0.1); 0.8(0.1) 1.7(0.1); 1.6(0.1) 1.1(0.1); 1.0(0.1) 0.7(0.1); 0.5(0.1) Simpson's diversity 0.3(0.1); 0.3(0.1) 0.8(0.1); 0.7(0.1) 0.8(0.1); 0.9(0.1) 0.8(0.1); 0.7(0.1) 0.6(0.1); 0.6(0.1) Shannon's evenness 0.2(0.1); 0.2(0.1) 0.5(0.1); 0.2(0.1)* 0.8(0.1); 0.8(0.1) 0.5(0.1); 0.5(0.1) 0.4(0.1); 0.3(0.1) (b) Feeding guild Fungivore 0.8(0.3); 1.1(0.3) 2.5(0.3); 2.1(0.3) 3.3(0.3); 3.3(0.3) 2.1(0.3); 2.4(0.3) 2.6(0.3); 3.1(0.3) Herbivore 2.0(0.3); 2.2(0.3) 1.7(0.3); 1.3(0.3) 6.9(0.3); 6.0(0.3) 4.2(0.3); 3.8(0.3) 2.5(0.3); 1.5(0.3) Saprophage 0.2(0.3); 0.0(0.3) 0.6(0.3); 0.8(0.3) 3.1(0.3); 3.1(0.3) 1.3(0.3); 1.2(0.3) 0.3(0.3); 0.5(0.3) Xylophage 2.6(0.2); 1.4(0.2)* 0.8(0.2); 0.3(0.2) 2.3(0.2); 1.7(0.2) 0.5(0.2); 0.5(0.2) 0.0(0.2); 0.2(0.2) Predator 3.0(0.2); 2.7(0.2) 2.6(0.2); 2.2(0.2) 2.8(0.2); 2.7(0.2) 1.7(0.2); 1.2(0.2) 0.7(0.2); 0.5(0.2) (c) Families Chrysomelidae 0.9(0.2); 1.2(0.2) 0.6(0.2); 0.1(0.2) 4.6(0.2); 3.5(0.2) 2.6(0.2); 1.7(0.2) 1.4(0.2); 0.8(0.2) Cupedidae 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.1(0.1) 0.0(0.1); 0.2(0.1) 0.0(0.1); 0.0(0.1) Phalacridae 0.1(0.1); 0.0(0.1) 0.2(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.0(0.1) Ptinidae 0.0(0.1); 0.0(0.1) 0.0(0.1); 0.1(0.1) 0.8(0.1); 0.7(0.1) 0.1(0.1); 0.6(0.1) 0.0(0.1); 0.2(0.1) Scolytinae 2.4(0.2); 1.0(0.2)* 0.3(0.2); 0.1(0.2) 0.8(0.2); 0.4(0.2) 0.2(0.2); 0.3(0.2) 0.0(0.2); 0.2(0.2) Throscidae 0.0(0.2); 0.0(0.2) 0.3(0.2); 0.7(0.2) 0.3(0.2); 0.7(0.2) 0.0(0.2); 0.2(0.2) 0.0(0.2); 0.0(0.2)

54

55

APPENDIX D

Scolytine abundance and origin

Abundance of scolytine species captured in funnel traps in 2015 for CWD+ and CWD-

treatments. Known origins also listed.

Scolytinae Species CWD + CWD - Origin

Xyleborinus saxeseni 10,404 7,615 Non-native

Ambrosiodmus tachygraphus 20 18 Native

Euwallacea validus 55 24 Non-native

Hypothenemus dissimilis 1 1 Non-native

Monarthrum fasciatum 450 424 Native

Monarthrum mali 1 0 Uncertain

Pityophthorus concentralis 1 0 Uncertain

Pseudothysannoes dislocatus 2 4 Uncertain

Xyleborus ferrugineus 2 1 Native

Xyleborus pelliculosus 440 395 Non-native

Xylosandrus crassiusculus 30 39 Non-native

Xylosandrus germanus 7 11 Non-native

Anisandrus sayi 103 142 Native

56

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67

VITA

Matthew Bryant Savage

Education:

• University of Kentucky, B.S. in Forestry, May 2014

Employment:

• Exotic Plant Management Team (Crew Leader) – Supervisor: Steven Bekedam, Yellowstone National Park, May 2017 – Present

• Field Research Technicain – Supervisor: Eric Chapman, Ministry of Agriculture and Fisheries in Muscat, Oman, February – March 2017

• Biological Science Technician (Plants) – Supervisor: Brian Hoefling, Uncompahgre National Forest, May – October 2016

• Undergraduate Field/Lab Technician – Supervisor: Dr. Lynne Rieske-Kinney, Department of Entomology, University of Kentucky, August 2013 – May 2014

• Forestry Technician – Supervisor: Ron Hollis, White River National Wildlife Refuge, May – August 2012 and May – August 2013

Awards:

• Graduate Student Grant – Kentucky Native Plant Society, October 2015 • Graduate Student Oral Presentation (2nd Place) – Southern Forest Insect Work

Conference, July 2015

Oral Presentaations:

• University of Kentucky, Department of Entomology, MS Exit Seminar, March 2016

• Southern Appalachian Forest Entomology and Pathology Symposium, March 2016

• 5th Annual Sustainability Forum Poster Competition: Tracy Farmer Institute for Sustainability of the Environment, December 2015

• Southern Forest Insect Work Conference Graduate Student Competition, July 2015

• University of Kentucky, Department of Entomology, MS Proposal Seminar, January 2015

• Ohio Valley Entomological Association Graduate Student Competition, October 2014