The Impact of Non-Native Woody Plants on the Native

THE IMPACT OF NON-NATIVE WOODY PLANTS

ON THE NATIVE HERBIVOROUS INSECT

COMMUNITY OF NORTHERN DELAWARE

Marion E. Zuefle

A thesis submitted to the Faculty of the University of Delaware in partial fulfillment of the requirements for the degree of Master of Science in Entomology and Applied Ecology

Spring 2006

UMI Number: 1435847

UMI Microform 1435847 Copyright 2006 by ProQuest Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code.

ProQuest Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, MI 48106-1346 THE IMPACT OF NON-NATIVE WOODY PLANTS

ON THE NATIVE HERBIVOROUS INSECT

COMMUNITY OF NORTHERN DELAWARE

Marion E. Zuefle

Approved: ______Douglas W. Tallamy, Ph.D. Professor in charge of thesis on behalf of the Advisory Committee

Approved: ______Douglas W. Tallamy, Ph.D. Chair of the Department of Entomology and Wildlife Ecology

Approved: ______Robin W. Morgan, Ph.D. Dean of the College of Agriculture and Natural Resources

Approved: ______Conrado M. Gempesaw II, Ph.D. Vice Provost for Academic and International Programs ACKNOWLEDGMENTS

First I would like to thank Bill Brown and Brutus for their constant support throughout this process. Bill also provided assistance with plot construction, plot maintenance, statistical analysis, and provided helpful comments on several earlier versions of this thesis. I would like to thank Jennie Witmer for her help with the first year’s data collection and continuing help and advice for the remainder of this project. I would also like to thank my family for all their support and encouragement. Thanks also to my committee members; Vince D’Amico, Judy Hough-Goldstein, and Doug Tallamy. I would like to acknowledge John Pesek for his help with statistical analysis. Several additional people helped with planting or data collection for which I am very grateful. The Northern Research Station of the Forest Service provided funding for this project and the Kranz family as well as White Clay Creek state park, provided the site for my study plot.

iii

TABLE OF CONTENTS

LIST OF FIGURES ...... vi LIST OF TABLES...... vii ABSTRACT...... viii

Chapter

1 INTRODUCTION ...... 1

2 METHODS ...... 5 Study Site...... 5 Plant Selection ...... 6 Experimental Design...... 7 Insect Sampling...... 7 Leaf Biomass ...... 9 Insect Identification and Biomass...... 9 Data Conversions...... 10 Statistical Analysis...... 11 Specialists/Generalists ...... 12 Biomass...... 12 Simpson's Diversity Index ...... 13 Species Richness...... 13 Faunal Overlap...... 14

3 RESULTS ...... 15 Specialists/Generalists ...... 15 Biomass...... 16 Simpson's Diversity Index ...... 17 Species Richness...... 18 Faunal Overlap...... 19

4 DISCUSSION...... 20

FIGURES...... 25 TABLES ...... 29

Appendix

A The total number of leaves used to determine mean biomass for one leaf for each plant species...... 34 B Total insect biomass collected (2004) from nine unique individuals of 45 plant species...... 35 C Total insect biomass collected (2005) from nine unique individuals of 45 plant species...... 39 D Native plant species and the insects collected (see Methods) from them in 2004 and 2005...... 43 E Non-native Congener plant species and the insects collected (see Methods) from them in 2004 and 2005...... 49 F Alien plant species and the insects collected (see Methods) from them in 2004 and 2005...... 55 G Simpson's diversity (probability that two randomly chosen individuals from a community belong to the same species) for each plant species for 2004, 2005, and both years combined ...... 61 H Species richness (total number of insect species found on a plant species) for each plant species for 2004, 2005, and both years combined ...... 62

REFERENCES ...... 63

LIST OF FIGURES

Figure 1 Aerial photograph (1997) of study site (red) within White Clay Creek State Park, Newark, Delaware...... 25

Figure 2 Diagram of the plot used in this study...... 26

Figure 3 The mean Simpson’s diversity index for Natives, Non-native Congeners, and Aliens for 2004, 2005, and both years combined...... 27

Figure 4 The mean Species richness for Natives, Non-native Congeners, and Aliens for 2004, 2005, and both years combined...... 28

LIST OF TABLES

Table 1 Family, scientific name, and plant code for the fifteen Native, fifteen Non-native Congeners, and fifteen Alien plant species used in this study...... 29

Table 2 Results of the repeated measures ANOVA on insect biomass per leaf biomass differences between congneric pairs for 2004 and 2005...... 30

Table 3 Results of the repeated measures ANOVA on differences in Simpson's diversity between congneric pairs for 2004 and 2005...... 31

Table 4 Results of the repeated measures ANOVA on differences in species richness difference between congneric pairs for 2004 and 2005...... 32

Table 5 Sorensen coefficient (degree of faunal overlap) between the congeneric pairs for 2004 and 2005...... 33

vii

ABSTRACT

Plants have been introduced to the United States for hundreds of years and are hypothesized to negatively impact native herbivorous insects. I tested this hypothesis by conducting a 2-year experiment in which native insect biomass production, native insect diversity, and native insect species richness was determined for native and non-native plants. A plot containing 45 species of woody plants with ten replicates each was set up in White Clay Creek state park for the purpose of this study. The plants consisted of 15 native species (Natives), 15 non-native species, which were congeners of the natives

(Non-native Congeners), and 15 non-native species that did not have a native congener present in the United States (Aliens). Herbivorous insects were sampled three times during the summer of 2004 and 2005, identified, and weighed.

In 2004, I found a difference in insect biomass per plant biomass production among the three groupings; Native, Non-native Congener, and Alien, but was unable to determine differences between the groups. In 2005 again there were differences among the three groupings and this time I could determine differences between the groupings.

Native plants produced more insect biomass per plant biomass than Non-native

Congeners or Aliens. However, insect biomass per plant biomass production between native and non-native congeneric pair members varied depending on the comparison: either the native member had more insect biomass per plant biomass, the non-native

viii member had more insect biomass per plant biomass, or there was no difference between the two species. In 2004 insect diversity tended to be greater on Natives than on Aliens, but there was no difference between Natives and Non-native Congeners or between Non- native Congeners and Aliens. Insect diversity was greater on Native species than on

Alien species in 2005, but there was no difference between Natives and Non-native

Congeners or Non-native Congeners and Aliens for that year. There was no difference in species richness among Native, Non-native Congener, and Alien plant species for either

2004 or 2005.

Overall, non-native plants had a negative impact on the native insect community

(insect biomass production and insect diversity), but not all non-natives were detrimental to the insects. The relatedness of non-natives to native plants (whether they are in the same genus or not) also had an impact on how strongly the herbivorous insect community was impacted.

Chapter 1

INTRODUCTION

Non-native plants have been introduced both intentionally and accidentally to the

United States since the arrival of the earliest European colonists. Intentional

introductions include crops such as corn (Zea mays), wheat (Triticum aestivum), and rice

(Oryza sativa); fiber plants such as eucalyptus (Eucalyptus grandis) and hemp (Cannabis

sativa); medicinal plants such as St. Johnswort (Hypericum perforatum) and foxglove

(Digitalis purpurea); and landscape plants such as multiflora rose (Rosa multiflora) and

melaleuca (Melaleuca quinquenervia). Landscape plants, defined by Reichard (1996) to

include plants used as ornamentals and those for soil protection or wildlife habitat, make

up 85% of all non-native plant introductions. The remaining non-native plants have

made their way here accidentally in contaminated soil, cotton or wool, grain, machinery

or in ship ballast (Randall 1996). Some non-natives, whether introduced intentionally or

not, have entered natural ecosystems and proliferated. Such plants are considered

“invasive” (Randall 1996).

Approximately 5,000 of the non-native plants introduced to the United States are

invasive (Pimentel et al. 2000). Invasive species are second only to habitat loss in their negative effect on biodiversity (Pimm and Gilpin 1989). Invasive plants can threaten natural ecosystems by out-competing native species and changing ecosystem functions

1 such as the fire regime or hydrological cycling (Blossey 1999). Others increase soil erosion or alter nutrient availability (Randall 1996), while others, such as white mulberry

(Morus alba) and oriental bittersweet (Celastrus orbiculatus) hybridize with native congeners, which can lead to the loss of native strains (Randall 1996).

Invasive plants possess many of the same characteristics worldwide that allow them to out-compete native plants. They are highly adaptable and as a group are tolerant of different climatic conditions, fast growing, disperse easily, produce numerous seeds early in life, reproduce vegetatively, have a long reproductive life, and germinate without any special requirements (Staples et al. 2000). An additional factor predicted to encourage invasive behavior in some plants is the inability of native insects to eat such plants (Elton 1958, Lodge 1993, Yela and Lawton 1997).

There are many non-native plants that do not become invasive, however, such as many of the ornamentals frequently planted in suburbs. Non-native plants, whether invasive or not, may still impact ecosystems. Ornamental plants have historically been chosen because they lacked insect herbivores (Dirr 1998), but the impact non-native plants have on the native insect community is one that has received little empirical attention (Tallamy 2004).

This lack of insect herbivores on non-native plants is predicted by the “enemy release” hypothesis (Keane and Crawley 2002). This hypothesis assumes that insect specialists of non-native plants are not present in the introduced range, that host switching by native specialists onto non-native plants is rare, and that generalist herbivores have a greater negative impact on native plants than on the non-native plants.

2 Several recent studies support the assumption that non-native plants have fewer insect herbivores in their introduced range (Goeden 1974, Strong et al. 1984, Andow and Imura

1994, Fenner and Lee 2001, Wolfe 2002), but few studies have looked at differences in herbivory between non-native species and a native congener (Agrawal and Kotanen 2003,

Vila and Weiner 2004, Agrawal et al. 2005). Such a comparison is necessary because insect herbivores are predicted to discriminate among non-native plants based on how closely they are related to the insect’s native food source.

Non-native plants are predicted to have a greater percentage of their insect herbivore load comprised of generalists than their native congeners, which are predicted to have a higher percentage of specialized herbivores (Goeden 1974, Strong et al. 1984,

Andow and Imura 1994, Fenner and Lee 2001). The reason is that generalist insects may accept novel plant hosts more readily than specialist herbivores. Specialist insects have evolved adaptations to overcome specific plant defenses and therefore have a limited ability to overcome novel plant defenses (Ehrlich and Raven 1965). Of the phytophagous insects, it is estimated that 90% restrict their feeding to host plants in three or fewer families (Bernays and Graham 1988). For this reason, escape from enemies is predicted to be even stronger for non-natives without a native congener in the introduced range than for non-natives with a native congener (Rejmanek 1999).

My objective was to determine if host use by native insect herbivores was influenced by plant origin (native or non-native). I determined differences in production of native insect biomass, native insect species richness, and native insect diversity on native plant species (Natives), introduced congeners of native species (Non-native

3 Congeners), and introduced plants without a native congener (Aliens). I also determined if native insect biomass production, native insect species richness, and native insect diversity differed between members of congeneric pairings of native and non-native plants. Congeneric comparisons constitute the most conservative measure of the ability of non-native plants to support native insect communities.

Specifically, I addressed the following questions:

1. Was there a difference in the number of specialist native insects occurring on Native, Non-native Congener, and Alien plant species?

2. Was there a difference in native insect biomass per leaf biomass production among Native, Non-native Congener, and Alien plant species?

3. Was there a difference in native insect biomass per leaf biomass production between the members of congeneric pairs?

4. Was there a difference in native insect diversity among Native, Non-native Congener, and Alien plant species?

5. Was there a difference in native insect diversity between the members of congeneric pairs?

6. Was there a difference in native insect species richness among Native, Non-native Congener, and Alien plant species?

7. Was there a difference in native insect species richness between the members of congeneric pairs?

8. To what degree did the insect fauna overlap between the members of congeneric pairs?

4 Chapter 2

METHODS

Study site

The study site was located in White Clay Creek State Park (WCCSP), Newark,

Delaware. WCCSP is a 1,369 ha park located in New Castle County, Delaware, and borders an additional 494 ha belonging to the White Clay Creek Preserve in Chester

County, Pennsylvania. The site for this study was acquired by WCCSP in 2000. Prior to its acquisition it was used as a Christmas tree farm for approximately 30 years (1969 to mid 1990’s) and then as hayfields for the last ten years. It continues to be hayed two to three times a season.

The study plot was established at the edge of one of the hayfields. It was bordered by mature forest and a hedgerow (Figure 1) and measured 42 m x 31.5 m (0.13 ha). A

2.13 m plastic mesh deer fence and five electrified strands of 12-gauge wire surrounded the plot. The plot was separated from the forest by approximately 12 m to allow the perimeter of the hayfield to be mowed. The soil of the plot site was uniformly composed of Chester Loam (ChB2) (Matthews and Lavoie 1970).

5 Plant Selection

Native plants (Natives) were defined as those plants native to Delaware (McAvoy and Bennett 2001). A native species was selected for study only if it could be found within the boundaries of WCCSP (pers. obs.) and also had a non-native congener present in Delaware. Non-native Congeners, defined as not having an evolutionary history in the

Mid-Atlantic region of North America but having a native congener present in WCCSP, were selected based on the USDA plants database (USDA, NRCS 2003). Plants were chosen if they were commonly planted as ornamentals in the surrounding area or considered invasive (McAvoy and Bennett 2001). Finally, Aliens, defined here as non- native to the United States (US) and having no native congener within the US, were chosen based on the USDA plants database (USDA, NRCS 2003). Aliens were selected for study if they were frequently planted in the surrounding landscape or considered invasive (McAvoy and Bennett 2001).

Two additional criteria were used to determine which plants were chosen. First, all plants had to be available through the nursery trade at a reasonable cost (<$100/plant).

Second, all plants had to be woody so as to control for life form, which is known to influence the herbivorous insect community (Ward et al. 1995). From these criteria three groupings consisting of 15 Natives, 15 Non-native Congeners, and 15 Aliens were selected for a total of 45 species in 30 genera (Table 1).

6 Experimental Design

The plot was divided into ten blocks. Each block contained all 45 species

randomly placed but with the congeneric pair (Native and Non-native Congener) always

occurring next to each other (Figure 2). The plants were placed 1.5 meters apart, and

were planted early April 2004. Even though plants were placed in a randomized block

design, sampling did not follow the design and therefore analyses were not based on a

randomized block design.

Insect Sampling

Insects were collected in mid-June, -July, and -August of 2004 and 2005. Three

replicates of the fifteen congeneric pairings and three replicates of the fifteen Alien plant

species were randomly selected for insect sampling, for a total of 135 plants sampled per

collection. No plant was sampled more than once a year. Approximately three days prior

to collecting, the entire plot was mowed and all plants were weeded. This was done to

insure that insects collected came from sampled plants and not from the surrounding

vegetation.

My sampling protocol for 2004 was as follows. A sheet was placed underneath

the plant to be sampled, then a knock down spray of Prentox® ExciteRTM containing 6%

Pyrethrins and 60% Piperonyl Butoxide was sprayed on all surfaces of the plant. After the spray was applied, dead insects were collected from the sheet and placed into labeled vials containing 80% ethyl alcohol (ETOH). Sampling was done between 5:30 and 7:30 am on June 19-21, July 19-21, and August 19-21. All three collections sampled 135

7 plants (three replicates of 45 species) except for the June collection in which

Koelreutaria paniculata was not sampled. All ten replicates of K. paniculata died after the first planting and were reordered; therefore, the first samples from K. paniculata were from the July sample.

The sampling protocol for 2005 differed from 2004. Instead of using a knock down spray to collect insects, I used an inverted leaf blower. The reason for this change was that spraying left very small dead insects (i.e. aphids and cicadellids) stuck to the leaf. Based on experimental trials (D. Tallamy, pers. comm.) it was determined that the leaf blower avoided this problem. Again, three replicates of each plant were sampled per collection and plants were randomly selected as in 2004. The plot was mowed and weeded several days prior to collecting. Sampling lasted only one day as this technique was more efficient than spraying. Samples were taken on June 16th, July 14th, and August

23rd, 2005. To collect insects, a paint strainer bag was placed at the end of the leaf blower and insects were ‘vacuumed’ from each plant. After collecting from a given plant, the paint strainer bag was tied shut with the corresponding plant number inside the bag. The bags were then placed into coffee tins containing ethyl acetate to kill the insects. Once the ethyl acetate took effect I removed the insects and placed them into labeled vials containing 80% ETOH.

As all plants were sampled using the same technique within a given year, and no comparisons between years were made, the effect of sampling protocol on results was not an issue.

8 Leaf Biomass

To determine leaf biomass, approximately 100 leaves were collected from each

plant species throughout the 2004 growing season (ten leaves from each plant). These

leaves were weighed on a Mettler AE100 scale to the nearest 0.01 g, providing an

average biomass per leaf for each species (Appendix A). The total number of leaves for

each plant was counted within a few days of sampling to provide an estimate of total leaf

biomass for each plant at the time of sampling (Appendices B (2004) and C (2005)).

Insect Identification and Biomass

Each insect morpho-species was given a reference number, identified to family,

dried, and weighed on a Mettler AE100 scale to the nearest 0.0001 g. A reference

collection was created containing several representatives of each referenced insect, when

available. This reference collection was then used to determine insect species or

“OTU’s” (operational taxonomic units) if species identification was not possible

(Futuyma and Gould 1979). Species determinations were based on Delong (1949),

Slater and Baranowski (1978), Hamilton (1982), White (1983), Stehr (1987), Arnett et al.

(2002), Wagner (2005), and the University of Delaware insect reference collection.

Only native herbivorous insects were considered in this study. The assumption was made that each plant on which an herbivorous insect was found was indeed a host plant. All identified insects were classified as either native to the US or non-native. As I was interested in the impact non-native plants have on native insect biomass, all non- native insects were removed from analysis. Insects that were classified only as OTU’s

9 were assumed to be native and used in analyses, providing a conservative estimate of the impact non-native plants have on the native insect community (Appendices D, E, and F give a list of insects found on each plant species for 2004 and 2005 combined).

Data Conversion

Prior to analysis, insect biomass data were scaled to leaf biomass data and log transformed using the following equation:

6 LOG 10(((x+1)/y)*10 ) where x is the total insect biomass collected from each plant and y is the total leaf biomass for each plant (Appendices B (2004) and C (2005) for total insect biomass collected from each plant and total leaf biomass for each plant). As some plants had no insect biomass collected from them, I added 1.0 to insect biomass totals for each plant prior to transformation, thereby retaining them in analyses. This transformed ratio is referred to hereafter as insect biomass per leaf biomass.

Simpson’s diversity index, which gives the probability that any two individuals chosen at random from the community belong to different species, was determined by the following formula:

D = 1 - Σ (n/N) 2 where n is the total number of individuals of a particular species and N is the total number of individuals of all species (Krebs 1999). A higher index represents greater diversity (greater “evenness”). I determined the Simpson’s diversity index for each plant species for 2004, 2005, and both years combined (Appendix G).

10 Species richness for each plant was determined as the total number of insect species (or OTU’s) found on each plant species (Krebs 1999) for 2004, 2005, and both years combined (Appendix H).

Finally, the degree of faunal overlap between congeneric pairs was determined using the Sorensen coefficient (Krebs 1999):

S = 2a/(2a+b+c) where a is the total number of species shared by both samples, b is the number of species found only on the first sample, and c is the total number of species found only on the second sample. This results in a number ranging from 0 (no overlap) to 1.0 (complete overlap). I determined the Sorensen coefficient for each congeneric pair for 2004, 2005, and both years combined (Table 5).

Statistical Analysis

Three different tests were used to analyze these data. I used a repeated measures

Analysis of Variance design (PROC MIXED, SAS Institute Inc., Cary, NC, 2004) in two different ways depending on which comparisons were being made. These were both

‘repeated measures’ because of the three sampling periods (June, July, and August). The first approach tested for differences among the means (see specific dependent variables specified below) of Native, Non-native Congener, and Alien plants, referred to here as

‘ANOVA means’. The second repeated measures design tested whether the difference between the means (see specific dependent variables below) of the congeneric pair was different from zero, referred to here as ‘ANOVA diff’. The final test I used was least

11 squares means regression (LS-means) (PROC GLM in SAS). LS-means was used to

determine if there were differences (see specific dependent variables below) between plant species when all replicates of a plant species were combined to give an overall insect species richness and Simpson’s diversity for each plant species (therefore no repeated measure involved).

Specialists/Generalists

To answer my first question of whether there were differences in the number of specialist insects on Natives, Non-natives Congeners, and Aliens, I followed the methods of Futuyma and Gould (1979). All insects found on more than three plant families were classified as generalists. The remaining insects were classified only if the species had at least ten individuals, fewer than ten were removed from consideration because to estimate whether an insect is a specialist or a generalist the number of individuals most be large enough to determine feeding preference (Futuyma and Gould 1979). If a species met this requirement and occurred on three or fewer families, it was classified a specialist.

Biomass

To determine if there were differences in insect biomass per leaf biomass

(question 2) among Natives, Non-native Congeners, and Aliens in 2004 and 2005, I used

‘ANOVA means’. This test was used because the mean insect biomass per leaf biomass among all three groupings (Native, Non-Native Congener and Alien) was being compared.

12 To answer question three, differences in insect biomass per leaf biomass production between members of congeneric pairs in 2004 and 2005, I used ‘ANOVA diff’. This test was used because Natives and Non-native Congeners were paired and therefore I could take the difference between their mean insect biomass per leaf biomass and test whether it was different from zero.

Simpson’s Diversity Index

To determine if there were differences in Simpson’s diversity index (question 4) among Natives, Non-native Congeners, and Aliens, I used LS-means. LS-means was used to test the difference in Simpson’s diversity index among each species in 2004,

2005, and the combined years.

To answer question five, are there differences in Simpson’s diversity index between members of congeneric pairs in 2004 and 2005, I used ‘ANOVA diff’. This test was used because Natives and Non-native Congeners were paired and therefore I could take the difference between their Simpson’s diversity index and test whether it was different from zero.

Species Richness

To determine if there were differences in species richness (question 6) among

Native, Non-native Congeners, and Alien, I used LS-means. LS-means was used to test the difference in species richness among each species in 2004, 2005, and the combined years.

13 To answer question seven, are there differences in species richness between members of congeneric pairs in 2004 and 2005, I used ‘ANOVA diff’. This test was used because Natives and Non-native Congeners were paired and therefore I could take the difference between their species richness and test whether it was different from zero.

Faunal Overlap

The degree of faunal overlap (question 8) between members of the congeneric pairs was determined for 2004, 2005 and the combined years using the Sorensen coefficient. A high Sorensen coefficient means greater faunal overlap between the congeneric pairs members.

14 Chapter 3

RESULTS

I collected a total of 4,866 individual insects comprising 164 native herbivorous

insect species or OTU’s for 2004 and 2005 combined. The 164 insect species or OTU’s

belonged to six insect orders (Orthoptera, Hemiptera, Homoptera, Coleoptera,

Lepidoptera, and Hymenoptera) and 45 insect families. For a list of insects found on

each plant species see Appendices D (Natives), E (Non-native Congeners), and F

(Aliens).

Specialists/Generalists

Question 1. Was there a difference in the number of specialist native insects among

Native, Non-native Congeners, and Alien plant species?

A total of 164 different native insect species were collected from 2004 and 2005 combined. Of this total, 106 species were not classified because there were fewer then 10 individuals collected, 51 species were generalists (found on more than three plant

families), and seven were specialist (more than 10 individuals found on three or fewer

families). Of the 51 generalists, 44 were found on Natives, 49 were found on Non-

natives, and 47 were found on Aliens. Of the seven specialists, four were found on

Natives, six were found on Non-native Congeners, and four were found on Aliens. There

15 was no difference in the number of specialist and generalist insects found on Native,

Non-native Congener, and Alien plant species. See Appendices D, E, and F for list of

insects and the plants on which they were found for 2004 and 2005 combined.

Biomass

Question 2. Was there a difference in native insect biomass per leaf biomass production among Native, Non-native Congeners, and Alien plant species?

Insect biomass per leaf biomass among Natives, Non-natives Congeners, and

Aliens differed in 2004 (F = 26.64, P <0.0001). The difference between Natives and

Aliens as well as Non-native Congeners and Aliens was non-estimable. Natives had more insect biomass per leaf biomass than Non-native Congeners (Estimate Native = 4.37,

Estimate Non-native Congener = 4.23, t = 4.15, P <0.0001). See Appendix B (2004) for the

untransformed insect biomass and total leaf biomass for each plant.

Insect biomass per leaf biomass among Natives, Non-native Congeners, and

Aliens also differed in 2005 (F = 9.54, P = <0.0001) and for this year I was able to

estimate pair-wise differences among all three groups. The insect biomass per leaf

biomass on Natives was greater than on Aliens (Estimate Native = 4.07, Estimate Alien =

4.00, t = -2.22, P = 0.03) and Natives were also greater than Non-native Congeners

(Estimate Native = 4.07, Estimate Non-native Congener = 3.93, t = 4.37, P <0.0001). Aliens, however, had greater insect biomass per leaf biomass than Non-natives (Estimate Non-native

Congener = 3.93, Estimate Alien = 4.00, t = 2.14, P = 0.03). See Appendix C (2005) for the

untransformed insect biomass and total leaf biomass for each plant.

16 Question 3. Was there a difference in native insect biomass per leaf biomass production between members of congeneric pairs?

Insect biomass per leaf biomass differences between the fifteen congeneric comparisons are given in Table 2. Values in bold are significant; the direction of significance (greater insect biomass per leaf biomass on the Native or the Non-native

Congener) depends on the estimate. If the estimate is positive then the Native had more insect biomass per leaf biomass than the Non-native Congener. If the estimate is negative then the Non-native Congener had significantly more insect biomass per leaf biomass than the Native.

Of the 15 comparisons in 2004, seven Natives had greater insect biomass per leaf biomass, four Non-native Congeners had greater insect biomass per leaf biomass, and four showed no difference. For 2005 the results were similar. Seven Natives had greater insect biomass per leaf biomass, three Non-native Congeners had greater insect biomass per leaf biomass, and five showed no difference between the two. The results differed between the two years because native Fagus and non-native Hamamelis, which had greater insect biomass per leaf biomass in 2004, showed no difference in 2005, and Tilia, which showed no difference in 2004, had greater insect biomass per leaf biomass on the native in 2005.

17 Simpson’s Diversity Index

Question 4. Was there a difference in native insect diversity among Native, Non-native

Congeners, and Alien plant species?

Natives had greater Simpson’s diversity than Aliens in 2005 (P = 0.03) and for

both years combined (P = 0.01) (Figure 3). There was no difference in Simpson’s

diversity among Non-native Congeners and Aliens for 2004, 2005, or both years

combined. See Appendix G for Simpson’s Diversity Index for each plant for 2004, 2005,

and both years combined.

Question 5. Was there a difference in native insect diversity between members of congeneric pairs?

Simpson’s diversity differences between the fifteen congeneric comparisons are given in Table 3. Values in bold are significant (direction of significance as before).

There were no differences in Simpson’s diversity for any of the 15 comparisons in 2004.

In 2005 two of the Natives had greater diversity and one of the Non-natives had greater diversity. For several of the comparisons I was not able to determine an estimate because either the model did not converge or the model had an infinite likelihood (McCullagh and

Nedler 1989).

Species Richness

Question 6. Was there a difference in native insect species richness among Native, Non- native Congener, and Alien plant species?

18 There was no difference in species richness among Native, Non-native Congener,

and Alien plant species for either year or both years combined (Figure 4).

See Appendix H for species richness for each plant for 2004, 2005, and both years

combined.

Question 7. Was there a difference in native insect species richness between members of congeneric pairs?

Species richness differences between the fifteen congeneric comparisons are given in Table 4. Values in bold are significant (the direction of significance as before).

Most of the congeners did not differ in species richness. In 2004 only one of the 15 comparisons had greater richness in the Non-native than its Native Congener. In 2005 one of the Natives had greater richness than its Non-native Congener. For several of the comparisons I was unable to determine an estimate because either the model did not converge or the model had an infinite likelihood.

Faunal Overlap

Question 8. To what degree did the insect fauna overlap between members of congeneric pairs?

The degree of faunal overlap (Sorensen Coefficient) between the fifteen congeneric comparisons is given in Table 5. A higher value means greater faunal overlap within the comparison. The faunal overlap for all of the comparisons was fairly low (≤

0.5) with the exception of the Salix comparisons, which were 0.63 in 2005 and 0.58 in

both years combined.

19 Chapter 4

DISCUSSION

The findings from this study generally support the enemy release hypothesis

(Keane and Crawley 2002). Native plants produced greater insect biomass per leaf

biomass than non-natives (Non-native Congener and Alien). It is theorized that when

non-native plants enter a new environment they often leave their native herbivores and

diseases behind (Keane and Crawley 2002). This is one of the many reasons given for

why non-native plants become so successful and often invasive in their new habitats.

However, my results did not support the prediction that non-natives without a native

congener (Aliens) produced the least insect biomass per leaf biomass. The results from my study also did not support the often-cited assumption that phytophagous insects are predominantly specialists (Eastop 1973, Futuyma and Gould 1979, Chapman 1982, Price

1983, Bernays and Graham 1988).

Of the 58 insect species that I could categorize, 88% were found to be generalists and 12% were specialists. This is in contrast to other studies that have found that less than 10% of all phytophagous insects are generalists and 90% are specialists (see references in Bernays and Graham 1988). The age of my study site might explain some of the results. Specialist insects will be more concentrated around their resource than in areas lacking their resource (Root 1973). As the plants in my plot were first sampled two

20 months after planting in 2004 and then again a year later, it is probable that specialist

insects were underrepresented in my plot. My samples also focused on folivores and

ignored internal feeders such as gall-makers, root feeders, and stem borers, all of which

tend to be more specialized (Fenner and Lee 2001).

Biomass results support the enemy release hypothesis. However, Aliens produced

more insect biomass per plant biomass than Non-native Congeners. This is contrary to

the assumption that non-native genera sustain a limited number of native insect

herbivores (Rejmánek 1999). Not only were Aliens in different genera than non-natives,

they also belonged to nine unique plant families not present in the congeneric pairs (see

Table 1 for list of plant families). Non-native plants that are taxonomically isolated are

thought to possess chemical defenses that native herbivores have not encountered and are

therefore unable to eat. This assumption is contrary to the “increased susceptibility”

hypothesis, which states that novel hosts might lack adequate defenses and would

therefore be more susceptible to native insect herbivores (Hokkanen and Pimentel 1989).

Depending on the plant species and its chemical composition, either one of these could be

true and lead to the observed results.

Time since introduction also influences the observed number of native insects

found on a plant as it takes time for native insects to switch onto a novel host.

Approximately 100 years are needed for a generalist insect to switch onto a new host

(Southwood 1984) and between 500 and 10,000 years for a specialist insect to switch hosts (Strong et al. 1984). In addition to time, the abundance of a non-native plant will affect the rate at which insects colonize it (Southwood 1961, Strong 1984, Auerbach and

21 Simberloff 1988). Both time since introduction and abundance were not known for my plants but could have contributed to the results.

Some non-native plants might have more insect herbivores due to a reallocation of resources, from defense to growth. Blossey and Nötzold (1995) state that a shift in resources, from defense to growth, might be observed in non-native plants in the absence of herbivores. This shift would favor genotypes with increased growth rates, yet could

also leave such plants vulnerable to herbivory when a host switch occurs. It is not

possible, however to determine which, if any, of my plant species had undergone a

genetic shift in resource allocation since introduction.

Insect biomass per leaf biomass production differed between congeneric pair

members as well. In general, the native representative produced greater insect biomass per leaf biomass but this was not always the case. Again, the enemy release hypothesis, increased susceptibility hypothesis, reallocation of resources, time since introduction, and plant abundance could all have had an impact on the result.

For species diversity, I found that Natives had greater diversity than Alien species but that there was no difference in diversity between Native and Non-native Congener species. These results were expected for two reasons. First non-natives have been found to reduce species diversity in other studies (Maron and Vila 2001) and are predicted to support fewer native herbivores because they lack an evolutionary history with these insects (Ehrlich and Raven 1965). Second, many native herbivores may possess behavioral and physiological adaptations that enable them to exploit close relatives of their native hosts. The fact that this was not always the case suggests that even congeners

22 can differ in traits such as leaf chemistry, trichome numbers, cuticular odors, or leaf

toughness. These are all factors that can influence herbivore success.

My results for insect species richness showed no difference among Natives, Non-

native Congeners, and Aliens and only one congeneric pair in each year showed a

difference in species richness. Species richness is directly related to the abundance of the

plant, the plant’s geographic spread (Strong et al. 1984), and ‘plant architecture’, which

includes size and structural variety (Strong et al. 1984). Plant architecture significantly

affects insect species richness. That is, when plant size increases, so does insect species

richness. I did not determine the abundance and geographic range for my study plants,

nor did I determine plant architecture beyond leaf biomass.

Faunal overlap, the degree to which two plants species share the same insect

species, was only determined for the congeneric pairs. In general the degree of faunal overlap was low, even though the diversity and species richness for congeneric pairs were similar. These results would be expected if there were a lot of specialists; however, I found only seven specialists and six of these were found on non-natives.

Conclusion

Non-native plants have been replacing native plants in the United States by the millions of hectares (Pimentel et al. 2000). This does not include the many non-native landscape plants associated with suburban sprawl. In Delaware alone 17% of the state’s total land area is developed (including residential (13%), urban, industrial, transportation, and commercial) (DOSPC 1999). These figures are for 1997 and have surely increased.

23 In fact, estimates of area converted to suburban/urban land use in the U.S. are as high as

54 % (Rosenzweig 2003). This shift from native to non-native vegetation will negatively impact the native insect community and therefore the higher trophic levels that depend on them.

Overall, insect biomass production was greater for native plants than for non- natives. This has to be considered when planting suburban landscapes. From the results of this study, we can determine which of the fifteen commonly planted non-natives would have an impact on native insect biomass production. If the goal is to increase native insect biomass production, then native congeners of non-native ornamentals should be planted with the exception of Betula, Morus, Viburnum, and possibly Hamamelis.

Figure 1. Aerial photograph (1997) of study site (red) within White Clay Creek State Park, Newark, Delaware.

25 42m

112635 4 1932122738 4 1939 1 1642 9 2435142939 6 2137102532 116401530315204514293615303872231217385203492431 132836 2 1733142943 5 2040 2 1737132845 4 1945132844153036 7 22 34 3 18 44 10 25 39 8 23 44 10 25 43 12 27 34 12 27 40 7 22 41 8 23 43 92442621378234111264162132318333183511642112633 122737 9 2441102531142939 4 1945 1 1638112631142936122745 3 1842 1 1638 2 1743122735102540 2 1743 5 2034153037 9 2432 41940153032621357224231844520343184162133132835 31.5m 11 26 39 7 22 34 14 29 45 15 30 41 8 23 36 11 26 32 4 19 38 8 23 44 1 16 40 5 2036132833 8 2344132831 6 2133 9 2437102539 7 2242 2 1743 Fence 52039153040924361163912273841933 11263762141318333184510253121737 1164213283872244520408233492435 1.5 meters between plants and fence 2 1734142931 8 2335153042 7 2236 6 2141 26 4 1945102532122743112643142932132844 6 2144102541132833 3 1845 6 2137 4 1938 3 18 43 2 17 32 15 30 31 2 17 43 15 30 35 13 28 44 112638 5 2039122742112639102531 9 2440 8 2337142936 4 1935122734 7 2242 5 2032 9244511640722341429338233611641 Gate

Native 123456789101112131415 ACRU BENI CACA COFL FAGR HAVI JUNI MORU PRSE RHPE ROCA SANI TIAM ULAM VIDE Non-native 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 Congener ACPL BEPE CABE COKO FASY HAMO JURE MOAL PRSU RHMU ROMU SABA TICO ULPA VIDI Alien 313233343536373839404142434445 AKQU ALJU COLU CYSC FOSU GIBI HEHE KOPA LAIN LIVU PATO PHAM POTR PYPA SYVU

Figure 2. Diagram of the plot used in this study. Plot was divided into ten blocks, with each block containing a random distribution of all 45 plant species (see Methods). Below the plot diagram is a list plant codes (see Table 1) with the corresponding number.

1.2 Native Non-native Congener 1 Alien

0.8 Index

0.6 27

0.4 Simpson's Diversity

0.2

0 2004 2005 2004-2005 Status by Year

Figure 3. The mean Simpson’s diversity index for Natives, Non-native Congeners, and Aliens for 2004, 2005, and both years combined. Natives had greater diversity than Aliens for 2005 (P = 0.03) and the combined years (P = 0.01).

30 Native

Non-native Congener 25 Alien

15 28 Species Richness 10

0 2004 2005 2004-2005 Status by Year

Figure 4. The mean Species richness for Natives, Non-native Congeners, and Aliens for 2004, 2005, and both years combined. There were no differences among the groups for either year or the years combined.

Table 1. Family, scientific name, and plant code for the fifteen Native, fifteen Non-native Congeners, and fifteen Alien plant species used in this study. See Methods for an explanation of the three plant groupings.

Family Native Code Non-native Congener Code Family Alien Code Aceraceae Acer rubrum ACRU Acer platanoides ACPL Lardizabalaceae Akebia quinata AKQU Betulaceae Betula nigra BENI Betula pendula BEPE Fabaceae Albizia julibrissin ALJU Betulaceae Carpinus caroliniana CACA Carpinus betulus CABE Rosaceae Cotoneaster lucidus COLU Cornaceae Cornus florida COFL Cornus kousa COKO Fabaceae Cytisus scoparius CYSC Fagaceae Fagus grandifolia FAGR Fagus sylvatica FASY Oleaceae Forsythia suspensa FOSU Hamamelidaceae Hamamelis virginiana HAVI Hamamelis mollis HAMO Ginkgoaceae Ginkgo biloba GIBI Juglandaceae Juglans nigra JUNI Juglans regia JURE Araliaceae Hedera helix HEHE Moraceae Morus rubra MORU Morus alba MOAL Sapindaceae Koelreuteria paniculata KOPA Rosaceae Prunus serotina PRSE Prunus serrulata PRSU Lythraceae Lagerstroemia indica LAIN

29 Ericaceae Rhodo. periclymenoides RHPE Rhodo. mucronatum RHMU Oleaceae Ligustrum vulgare LIVU Rosaceae Rosa carolina ROCA Rosa multiflora ROMU Scrophulariaceae Paulownia tomentosa PATO Salicaceae Salix nigra SANI Salix babylonica SABA Rutaceae Phellodendron amurens PHAM Tiliaceae Tilia americana TIAM Tilia cordata TICO Rutaceae Poncirus trifoliata POTR Ulmaceae Ulmus americana ULAM Ulmus parvifolia ULPA Rosaceae Pyrus pashia PYPA Caprifoliaceae Viburnum dentatum VIDE Viburnum dilatatum VIDI Oleaceae Syringa vulgaris SYVU

Table 2. Results of the repeated measures ANOVA on insect biomass per leaf biomass differences between congeneric pairs for 2004 and 2005. If the estimate was positive then the Native had more insect biomass per leaf biomass, and if the estimate was negative then the Non-native Congener had more insect biomass per leaf biomass. See Table 1 for plant codes and Appendices B and C for untransformed data.

Year Native Non-native N Estimate Std Err t -value P-value 2004 ACRU ACPL 9 0.58 0.08 7.27 0.002 BENI BEPE 9 -0.45 0.07 -6.22 0.002 CACA CABE 9 -0.35 0.17 -2.11 0.111 COFL COKU 9 0.08 0.12 0.63 0.564 FAGR FASY 9 0.33 0.06 5.59 0.007 HAVI HAMO 9 -0.22 0.07 -3.05 0.036 JUNI JURE 9 0.40 0.10 4.22 0.011 MORU MOAL 9 -0.76 0.16 -4.89 0.012 PRSE PRSU 9 0.63 0.13 4.89 0.004 RHPE RHMU 9 0.37 0.03 10.84 0.001 ROCA ROMU 9 0.19 0.12 1.52 0.180 SANI SABA 9 0.90 0.10 8.94 0.001 TIAM TICO 9 0.23 0.14 1.62 0.171 ULAM ULPA 9 0.42 0.05 9.30 0.002 VIDE VIDI 9 -0.34 0.08 -4.25 0.044 2005 ACRU ACPL 9 0.44 0.14 3.28 0.031 BENI BEPE 9 -0.69 0.12 -5.60 0.021 CACA CABE 9 0.07 0.13 0.54 0.612 COFL COKU 9 0.10 0.15 0.68 0.537 FAGR FASY 9 0.13 0.06 2.12 0.115 HAVI HAMO 9 -0.16 0.11 -1.38 0.234 JUNI JURE 9 0.28 0.07 4.01 0.014 MORU MOAL 9 -0.46 0.12 -3.85 0.016 PRSE PRSU 9 0.97 0.17 5.70 0.003 RHPE RHMU 9 0.25 0.04 6.29 0.001 ROCA ROMU 9 0.25 0.16 1.53 0.210 SANI SABA 9 0.92 0.22 4.08 0.014 TIAM TICO 9 0.04 0.09 0.48 0.659 ULAM ULPA 9 0.32 0.05 7.01 0.002 VIDE VIDI 9 -0.04 0.05 -8.23 0.010

30 Table 3. Results of the repeated measures ANOVA on differences in Simpson’s diversity between congeneric pairs for 2004 and 2005. If the estimate was positive then the Native had greater diversity and if the estimate was negative then the Non-native Congener had greater diversity. See Table 1 for plant codes and Appendix G for insect species diversity found on each plant species.

Year Native Non-native N Estimate Std Err t -value P-value 2004 ACRU ACPL 9 -0.24 0.11 -2.23 0.07 BENI BEPE 9 0.06 0.10 0.58 0.58 CACA CABE 9 0.23 0.14 1.62 0.21 COFL COKU 9 * * * * FAGR FASY 9 * * * * HAVI HAMO 9 0.11 0.12 0.87 0.43 JUNI JURE 9 * * * * MORU MOAL 9 * * * * PRSE PRSU 9 * * * * RHPE RHMU 9 0.18 0.10 1.75 0.13 ROCA ROMU 9 -0.01 0.13 -0.08 0.94 SANI SABA 9 * * * * TIAM TICO 9 * * * * ULAM ULPA 9 * * * * VIDE VIDI 9 -0.11 0.13 -0.84 0.44

2005 ACRU ACPL 9 -0.12 0.10 -1.26 0.28 BENI BEPE 9 -0.01 0.11 -0.12 0.91 CACA CABE 9 0.26 0.10 2.61 0.05 COFL COKU 9 0.04 0.08 0.47 0.66 FAGR FASY 9 * * * * HAVI HAMO 9 0.02 0.10 0.16 0.88 JUNI JURE 9 -0.07 0.13 -0.51 0.63 MORU MOAL 9 * * * * PRSE PRSU 9 -0.30 0.15 -2.07 0.12 RHPE RHMU 9 -0.31 0.12 -2.64 0.04 ROCA ROMU 9 0.07 0.09 0.83 0.47 SANI SABA 9 0.29 0.11 2.57 0.05 TIAM TICO 9 -0.22 0.09 -2.33 0.07 ULAM ULPA 9 -0.17 0.09 -1.97 0.12 VIDE VIDI 9 0.05 0.15 0.33 0.76 * non-estimable

31 Table 4. Results of the repeated measures ANOVA on the species richness difference between congeneric pairs for 2004 and 2005. If the estimate was positive then the Native had greater species richness and if the estimate was negative then the Non-native Congener had greater species richness. See Table 1 for plant codes and Appendix H for insect species richness for each plant species.

Year Native Non-native N Estimate Std Err t- value P-value 2004 ACRU ACPL 9 -0.78 0.51 -1.53 0.22 BENI BEPE 9 0.22 0.29 0.76 0.48 CACA CABE 9 0.78 0.64 1.22 0.33 COFLCOKU9**** FAGR FASY 9 -0.22 0.19 -1.15 0.29 HAVI HAMO 9 0.56 0.47 1.18 0.31 JUNI JURE 9 -0.22 0.31 -0.71 0.51 MORUMOAL9**** PRSE PRSU 9 0.22 0.35 0.63 0.55 RHPE RHMU 9 0.78 0.42 1.87 0.12 ROCA ROMU 9 0.33 0.76 0.44 0.68 SANI SABA 9 0.44 0.72 0.62 0.57 TIAM TICO 9 -0.44 0.51 -0.87 0.43 ULAM ULPA 9 -0.78 0.38 -2.02 0.11 VIDE VIDI 9 -1.56 0.56 -2.80 0.04

2005 ACRU ACPL 9 **** BENI BEPE 9 0.11 0.66 0.17 0.87 CACA CABE 9 1.56 0.37 4.22 0.01 COFL COKU 9 0.33 0.43 0.77 0.47 FAGR FASY 9 -0.56 0.25 -2.24 0.08 HAVI HAMO 9 -0.11 0.37 -0.30 0.78 JUNI JURE 9 -0.44 0.53 -0.83 0.45 MORU MOAL 9 0.00 0.25 0.00 1.00 PRSE PRSU 9 -0.78 0.87 -0.90 0.42 RHPE RHMU 9 -1.22 1.02 -1.19 0.31 ROCA ROMU 9 -0.22 0.78 -0.29 0.79 SANI SABA 9 0.67 0.46 1.46 0.20 TIAM TICO 9 -1.11 0.52 -2.13 0.09 ULAM ULPA 9 -1.11 0.53 -2.09 0.10 VIDE VIDI 9 -0.33 0.66 -0.51 0.64 * non-estimable

32 Table 5. Sorensen coefficient (degree of faunal overlap) between the congeneric pairs for 2004, 2005, and for both years combined. If Sorensen is zero then there is no faunal overlap, if Sorensen is one then there is complete faunal overlap. See Table 1 for plant codes.

Year Native Non-Native Sorensen Year Native Non-Native Sorensen Year Native Non-Native Sorensen 2004 ACRU ACPL 0.00 2005 ACRU ACPL 0.42 Both Years ACRU ACPL 0.31 BENI BEPE 0.29 BENI BEPE 0.43 BENI BEPE 0.38 CACA CABE 0.27 CACA CABE 0.43 CACA CABE 0.50 COFL COKO 0.00 COFL COKO 0.50 COFL COKO 0.48 FAGR FASY 0.00 FAGR FASY 0.20 FAGR FASY 0.13 HAVI HAMO 0.00 HAVI HAMO 0.40 HAVI HAMO 0.31 JUNI JURE 0.25 JUNI JURE 0.35 JUNI JURE 0.26 MORU MOAL 0.00 MORU MOAL 0.22 MORU MOAL 0.33

33 PRSE PRSU 0.00 PRSE PRSU 0.37 PRSE PRSU 0.33 RHPE RHMU 0.12 RHPE RHMU 0.46 RHPE RHMU 0.32 ROCA ROMU 0.34 ROCA ROMU 0.41 ROCA ROMU 0.41 SANI SABA 0.31 SANI SABA 0.63 SANI SABA 0.58 TIAM TICO 0.17 TIAM TICO 0.42 TIAM TICO 0.30 ULAM ULPA 0.22 ULAM ULPA 0.38 ULAM ULPA 0.40 VIDE VIDI 0.29 VIDE VIDI 0.34 VIDE VIDI 0.33

Appendix A. The total number of leaves used to determine mean biomass for one leaf for each plant species. The mean biomass for one leaf was then used to convert the total leaf count into total leaf biomass for each plant species for 2004 and 2005. See Table 1 for plant codes and Appendices B (2004) and C (2005) for leaf biomass for each plant replicate.

Plant Leaves Mean biomass for Leaf Count Leaf Biomass Leaf Count Leaf Biomass Species Collected one leaf (g) 2004 2004 (g) 2005 2005 (g) Native ACRU 60 0.362 275 99.65 702 254.37 BENI 100 0.148 6204 915.71 19278 2845.43 CACA 100 0.113 4457 501.86 6290 708.25 COFL 100 0.584 333 194.32 1428 833.31 FAGR 60 0.393 428 168.20 601 236.19 HAVI 70 0.446 920 410.19 2511 1119.55 JUNI 100 0.424 1032 437.96 1068 453.24 MORU 100 0.258 3172 817.77 5469 1409.96 PRSE 100 0.189 1499 283.19 3433 648.55 RHPE 100 0.115 9382 1080.40 8063 928.51 ROCA 100 0.039 26841 1057.54 59372 2339.26 SANI 100 0.035 4188 144.51 11557 398.77 TIAM 100 0.451 370 166.92 692 312.18 ULAM 100 0.325 1378 448.09 5164 1679.19 VIDE 100 0.362 9095 3296.41 28749 10419.85 Non-native ACPL 70 0.579 615 355.93 1009 583.95 Congener BEPE 100 0.130 2629 340.98 4943 641.11 CABE 100 0.139 2269 315.84 6645 924.98 COKO 100 0.318 931 296.14 3138 998.14 FASY 100 0.217 1735 376.14 1380 299.17 HAMO 60 0.610 393 239.60 1389 846.83 JURE 80 0.682 1586 1081.25 1185 807.87 MOAL 100 0.188 875 164.14 2301 431.63 PRSU 100 0.615 1676 1030.40 6495 3993.11 RHMU 100 0.051 48398 2463.46 33291 1694.51 ROMU 100 0.091 19584 1772.39 54147 4900.41 SABA 100 0.149 6185 919.20 26992 4011.47 TICO 100 0.155 1824 282.66 2313 358.44 ULPA 100 0.031 39917 1245.41 110854 3458.64 VIDI 100 0.438 4183 1833.83 10587 4641.34 Alien AKQU 100 0.033 4599 153.61 30630 1023.04 ALJU 100 0.003 79514 214.69 687402 1855.99 COLU 100 0.048 37550 1813.67 43506 2101.34 CYSC 100 0.002 24717 54.38 209745 461.44 FOSU 100 0.151 1224 184.46 6334 954.53 GIBI 100 0.360 1169 420.26 2640 949.08 HEHE 100 0.328 5758 1888.62 10717 3515.18 KOPA 100 0.050 1669 84.12 6259 315.45 LAIN 100 0.115 2988 342.72 9532 1093.32 LIVU 100 0.075 15168 1133.05 34806 2600.01 PATO 53 4.716 188 886.65 764 3603.20 PHAM 100 0.234 1276 298.46 2645 618.67 POTR 100 0.061 3103 190.21 3236 198.37 PYPA 100 0.076 5141 392.54 21249 1622.47 SYVU 100 0.269 2423 652.51 3208 863.91

34 Appendix B. Total insect biomass collected (2004) from nine unique individuals of 45 plant species. Samples were taken throughout the summer of 2004, three from June (sample 1), three from July (sample 2) and three from August (sample 3). Leaf biomass for each of the nine plant replicates is also given. See Table 1 for plant code. Native Non-native Congener Alien Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass ACRU 1 6.52 0.0000 ACPL 1 44.56 0.0034 AKQU 1 14.33 0.0092 ACRU 1 5.44 0.0000 ACPL 1 24.31 0.0597 AKQU 1 14.03 0.0028 ACRU 1 4.35 0.0000 ACPL 1 28.94 0.0000 AKQU 1 18.87 0.0000 ACRU 2 9.78 0.0000 ACPL 2 46.88 0.0019 AKQU 2 14.53 0.0000 ACRU 2 18.12 0.0000 ACPL 2 45.72 0.0000 AKQU 2 19.84 0.0000 ACRU 2 16.31 0.0005 ACPL 2 34.72 0.1415 AKQU 2 17.94 0.0000 ACRU 3 19.93 0.0011 ACPL 3 34.15 0.0000 AKQU 3 25.05 0.0000 ACRU 3 7.61 0.0001 ACPL 3 59.61 0.0000 AKQU 3 18.00 0.0000 ACRU 3 11.60 0.0000 ACPL 3 37.04 0.0571 AKQU 3 11.02 0.0000 BENI 1 104.06 0.0000 BEPE 1 29.18 0.0000 ALJU 1 5.41 0.0000 BENI 1 95.50 0.0000 BEPE 1 56.81 0.0015 ALJU 1 9.43 0.0008 BENI 1 79.56 0.0000 BEPE 1 35.80 0.0089 ALJU 1 8.06 0.0000 BENI 2 142.14 0.0029 BEPE 2 59.14 0.0057 ALJU 2 15.35 0.0000 BENI 2 113.21 0.0005 BEPE 2 22.05 0.0474 ALJU 2 23.33 0.0000 35 BENI 2 124.43 0.1607 BEPE 2 11.67 0.0000 ALJU 2 26.54 0.0006 BENI 3 83.25 0.0009 BEPE 3 39.95 0.0000 ALJU 3 0.13 0.0000 BENI 3 115.57 0.0008 BEPE 3 41.11 0.0000 ALJU 3 71.80 0.0000 BENI 3 58.01 0.0000 BEPE 3 45.27 0.0000 ALJU 3 54.65 0.0000 CACA 1 35.02 0.0019 CABE 1 90.62 0.0561 COLU 1 290.04 0.0401 CACA 1 49.77 0.0008 CABE 1 8.91 0.0052 COLU 1 243.19 0.1408 CACA 1 87.60 0.0171 CABE 1 10.16 0.0000 COLU 1 100.27 0.0047 CACA 2 81.52 0.0010 CABE 2 12.81 0.0141 COLU 2 161.32 0.0547 CACA 2 81.52 0.0131 CABE 2 9.74 0.0008 COLU 2 301.49 0.0300 CACA 2 81.52 0.0736 CABE 2 13.36 0.0009 COLU 2 178.13 0.0481 CACA 3 16.33 0.0004 CABE 3 71.83 0.0002 COLU 3 191.17 0.4580 CACA 3 29.16 0.0053 CABE 3 23.66 0.6618 COLU 3 227.20 0.0000 CACA 3 39.41 0.0635 CABE 3 74.75 0.1174 COLU 3 120.85 0.1591 COFL 1 26.26 0.0000 COKO 1 46.12 0.0000 CYSC 1 2.27 0.0769 COFL 1 17.51 0.0000 COKO 1 24.81 0.0000 CYSC 1 7.70 0.0067 COFL 1 20.42 0.0000 COKO 1 20.68 0.0000 CYSC 1 18.92 0.0027 COFL 2 14.01 0.0000 COKO 2 9.54 0.0000 CYSC 2 6.55 0.0000 COFL 2 29.18 0.0000 COKO 2 7.95 0.0000 CYSC 2 9.72 0.0000 COFL 2 26.26 0.0000 COKO 2 59.16 0.0260 CYSC 2 2.64 0.0000 COFL 3 25.68 0.0000 COKO 3 15.27 0.0000 CYSC 3 0.97 0.0006 COFL 3 22.17 0.0000 COKO 3 84.93 0.0000 CYSC 3 5.50 0.0000 COFL 3 12.84 0.0000 COKO 3 27.67 0.0000 CYSC 3 0.12 0.0000 FAGR 1 21.62 0.0000 FASY 1 63.74 0.0004 FOSU 1 12.21 0.0000 FAGR 1 20.44 0.0000 FASY 1 35.55 0.0000 FOSU 1 12.06 0.0000

35 Appendix B. cont.

36 HAVI 3 39.68 0.1237 HAMO 3 67.06 0.0000 GIBI 3 35.59 0.0000 JUNI 1 89.12 0.0000 JURE 1 82.49 0.0022 HEHE 1 273.22 0.0188 JUNI 1 63.66 0.0008 JURE 1 225.66 0.0000 HEHE 1 322.42 0.0145 JUNI 1 59.41 0.0008 JURE 1 100.90 0.0150 HEHE 1 247.97 0.0088 JUNI 2 40.32 0.0000 JURE 2 151.35 0.0046 HEHE 2 171.22 0.0061 JUNI 2 30.13 0.0003 JURE 2 105.67 0.0000 HEHE 2 223.04 0.0093 JUNI 2 58.99 0.0000 JURE 2 118.62 0.0000 HEHE 2 213.53 0.0023 JUNI 3 41.17 0.0000 JURE 3 91.35 0.0000 HEHE 3 147.93 0.0000 JUNI 3 17.40 0.0000 JURE 3 143.17 0.0000 HEHE 3 171.87 0.0202 JUNI 3 37.77 0.0000 JURE 3 62.04 0.0054 HEHE 3 117.42 0.0186 MORU 1 79.41 0.0004 MOAL 1 18.95 0.0000 KOPA 2 5.80 0.0000 MORU 1 85.59 0.0013 MOAL 1 30.58 0.0016 KOPA 2 12.60 0.0000 MORU 1 106.22 0.0018 MOAL 1 2.06 0.0000 KOPA 2 14.97 0.0000 MORU 2 97.71 0.0000 MOAL 2 18.20 0.0000 KOPA 3 38.51 0.0000 MORU 2 34.55 0.0000 MOAL 2 16.13 0.0000 KOPA 3 6.45 0.0000 MORU 2 117.30 0.0000 MOAL 2 26.64 0.0000 KOPA 3 5.80 0.0000 MORU 3 84.30 0.0000 MOAL 3 29.64 0.0000 LAIN 1 18.24 0.0000 MORU 3 149.53 0.0004 MOAL 3 9.75 0.0000 LAIN 1 17.32 0.0000 MORU 3 63.16 0.0000 MOAL 3 12.19 0.0000 LAIN 1 27.41 0.0000 PRSE 1 41.75 0.0007 PRSU 1 165.38 0.0000 LAIN 2 37.05 0.0000 PRSE 2 25.31 0.0500 PRSU 1 121.73 0.0000 LAIN 2 37.39 0.0000 PRSE 3 59.13 0.0074 PRSU 1 135.26 0.0007 LAIN 2 46.45 0.0003 PRSE 1 17.76 0.0000 PRSU 2 46.72 0.0000 LAIN 3 62.05 0.0000

Appendix B. cont.

Native Non-native Congener Alien Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass PRSE 1 28.53 0.0000 PRSU 2 82.38 0.0001 LAIN 3 39.34 0.0025 PRSE 2 8.31 0.0000 PRSU 2 149.40 0.0036 LAIN 3 57.46 0.0000 PRSE 2 68.95 0.0000 PRSU 3 146.94 0.0000 LIVU 1 78.73 0.0000 PRSE 3 11.71 0.0000 PRSU 3 98.37 0.0000 LIVU 1 156.05 0.0037 PRSE 3 21.73 0.0000 PRSU 3 84.23 0.0000 LIVU 1 143.20 0.0000 RHPE 1 136.23 0.0022 RHMU 1 240.76 0.0022 LIVU 2 251.44 0.0001 RHPE 1 124.37 0.0016 RHMU 1 262.59 0.0000 LIVU 2 182.72 0.0000 RHPE 1 152.47 0.0010 RHMU 1 271.35 0.0000 LIVU 2 93.30 0.0000 RHPE 2 161.80 0.0000 RHMU 2 258.93 0.0007 LIVU 3 23.68 0.0000 RHPE 2 79.11 0.0006 RHMU 2 228.54 0.0000 LIVU 3 109.44 0.0077 RHPE 2 104.91 0.0022 RHMU 2 291.96 0.0009 LIVU 3 94.50 0.0000 RHPE 3 85.79 0.0005 RHMU 3 303.97 0.0000 PATO 1 66.03 0.0002 RHPE 3 123.56 0.0018 RHMU 3 332.12 0.0110 PATO 1 56.59 0.0000 RHPE 3 112.16 0.0014 RHMU 3 273.23 0.0165 PATO 1 80.18 0.0000 ROCA 1 72.89 0.0064 ROMU 1 75.30 0.0000 PATO 2 80.18 0.0007 37 ROCA 1 43.22 0.0054 ROMU 1 211.87 0.0118 PATO 2 66.03 0.0006 ROCA 1 143.69 0.0093 ROMU 1 145.35 0.0000 PATO 2 113.19 0.0052 ROCA 2 118.79 0.0339 ROMU 2 303.91 0.0823 PATO 3 141.49 0.1449 ROCA 2 142.19 0.0083 ROMU 2 100.10 0.0021 PATO 3 150.92 0.0183 ROCA 2 48.34 0.0000 ROMU 2 145.71 0.0106 PATO 3 132.05 0.1798 ROCA 3 138.69 0.0054 ROMU 3 65.61 0.0010 PHAM 1 48.89 0.0000 ROCA 3 183.76 0.0112 ROMU 3 251.60 0.0341 PHAM 1 24.56 0.0000 ROCA 3 165.95 0.0025 ROMU 3 472.96 0.0000 PHAM 1 25.26 0.0000 SANI 1 17.74 0.0031 SABA 1 85.16 0.0049 PHAM 2 56.37 0.0000 SANI 1 3.24 0.0000 SABA 1 70.74 0.0000 PHAM 2 29.94 0.0000 SANI 1 4.11 0.0069 SABA 1 78.47 0.0000 PHAM 2 19.88 0.0000 SANI 2 48.89 0.0193 SABA 2 94.82 0.0000 PHAM 3 32.51 0.0000 SANI 2 10.04 0.0003 SABA 2 79.96 0.0000 PHAM 3 25.50 0.0055 SANI 2 16.80 0.0000 SABA 2 154.56 0.0445 PHAM 3 35.55 0.0000 SANI 3 15.70 0.0007 SABA 3 94.22 0.0000 POTR 1 25.38 0.0004 SANI 3 17.63 0.0000 SABA 3 176.71 0.0000 POTR 1 21.09 0.0000 SANI 3 10.35 0.0000 SABA 3 84.56 0.0000 POTR 1 20.41 0.0000 TIAM 1 17.14 0.0000 TICO 1 21.08 0.0011 POTR 2 25.93 0.0000 TIAM 1 10.83 0.0000 TICO 1 39.83 0.0000 POTR 2 18.76 0.0000 TIAM 1 18.05 0.0000 TICO 1 63.85 0.0055 POTR 2 12.75 0.0000 TIAM 2 11.28 0.0000 TICO 2 40.14 0.0000 POTR 3 20.05 0.0000 TIAM 2 23.01 0.0000 TICO 2 12.86 0.0007 POTR 3 20.54 0.0000 TIAM 2 27.52 0.0014 TICO 2 34.87 0.0000 POTR 3 25.32 0.0000

Appendix B. cont.

Native Non-native Congener Alien Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass TIAM 3 9.92 0.6380 TICO 3 37.35 0.0000 PYPA 1 17.03 0.0043 TIAM 3 17.14 0.0003 TICO 3 14.88 0.0048 PYPA 1 68.03 0.0015 TIAM 3 32.03 0.0009 TICO 3 17.82 0.0000 PYPA 1 18.40 0.0042 ULAM 1 48.78 0.0003 ULPA 1 119.37 0.0009 PYPA 2 84.14 0.0049 ULAM 1 45.85 0.0000 ULPA 1 132.88 0.0026 PYPA 2 12.14 0.0003 ULAM 1 45.52 0.0036 ULPA 1 173.78 0.0002 PYPA 2 42.91 0.0112 ULAM 2 39.02 0.0003 ULPA 2 51.04 0.1586 PYPA 3 31.31 0.0000 ULAM 2 54.63 0.0000 ULPA 2 157.87 0.0055 PYPA 3 92.47 0.0000 ULAM 2 71.54 0.0119 ULPA 2 117.50 0.0080 PYPA 3 26.11 0.0010 ULAM 3 45.52 0.0000 ULPA 3 181.58 0.0050 SYVU 1 14.81 0.0000 ULAM 3 63.73 0.0000 ULPA 3 193.69 0.0217 SYVU 1 47.67 0.0000 ULAM 3 33.49 0.0160 ULPA 3 117.69 0.0049 SYVU 1 55.21 0.0027 VIDE 1 395.06 0.0155 VIDI 1 272.68 0.0340 SYVU 2 44.97 0.0004 VIDE 1 509.23 0.0014 VIDI 1 56.99 0.0006 SYVU 2 94.79 0.0000 VIDE 1 326.56 0.0134 VIDI 1 141.16 0.0178 SYVU 2 180.97 0.0042 38 VIDE 2 406.30 0.0002 VIDI 2 339.32 0.7160 SYVU 3 57.63 0.0000 VIDE 2 316.77 0.0074 VIDI 2 139.41 0.0513 SYVU 3 32.32 0.0000 VIDE 2 365.34 0.0020 VIDI 2 198.16 0.3363 SYVU 3 124.15 0.0011 VIDE 3 288.50 0.0000 VIDI 3 234.54 0.0000 VIDE 3 328.74 0.0365 VIDI 3 225.78 0.0000 VIDE 3 359.91 0.0000 VIDI 3 225.78 0.0866

Appendix C. Total insect biomass collected (2005) from nine unique individuals of 45 plant species. Samples were taken throughout the summer of 2005, three from June (sample 1), three from July (sample 2) and three from August (sample 3). Leaf biomass for each of the nine plant replicates is also given. See Table 1 for plant codes. Native Non-native Congener Alien Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass ACRU 1 4.35 0.0000 ACPL 1 70.03 0.0000 AKQU 1 242.99 0.0015 ACRU 1 63.05 0.0000 ACPL 1 86.81 0.0000 AKQU 1 142.79 0.0099 ACRU 1 41.31 0.0013 ACPL 1 92.60 0.0000 AKQU 1 62.63 0.0004 ACRU 2 22.47 0.0012 ACPL 2 40.51 0.0167 AKQU 2 101.87 0.0042 ACRU 2 7.61 0.0015 ACPL 2 52.67 0.0027 AKQU 2 46.93 0.0020 ACRU 2 33.34 0.0000 ACPL 2 46.88 0.0089 AKQU 2 77.15 0.0013 ACRU 3 23.19 0.0005 ACPL 3 88.55 0.0191 AKQU 3 104.88 0.0578 ACRU 3 23.55 0.0037 ACPL 3 38.78 0.0011 AKQU 3 130.59 0.0066 ACRU 3 35.51 0.0004 ACPL 3 67.13 0.0051 AKQU 3 113.23 0.0000 BENI 1 222.88 0.0017 BEPE 1 96.37 0.0002 ALJU 1 285.04 0.0012 BENI 1 336.23 0.0039 BEPE 1 14.27 0.0031 ALJU 1 19.83 0.0000 BENI 1 200.15 0.0025 BEPE 1 115.69 0.0025 ALJU 1 76.04 0.0013 BENI 2 215.20 0.0037 BEPE 2 76.26 0.0035 ALJU 2 285.39 0.0009 BENI 2 509.96 0.0424 BEPE 2 104.02 0.0039 ALJU 2 146.45 0.0009 39 BENI 2 287.08 0.0015 BEPE 2 97.40 0.0019 ALJU 2 233.09 0.0013 BENI 3 349.52 0.1264 BEPE 3 58.11 0.0868 ALJU 3 199.82 0.0083 BENI 3 424.50 0.0003 BEPE 3 39.95 0.0006 ALJU 3 355.02 0.0000 BENI 3 299.92 0.0185 BEPE 3 39.04 0.0216 ALJU 3 255.31 0.0005 CACA 1 81.86 0.0004 CABE 1 27.84 0.0000 COLU 1 379.64 0.0031 CACA 1 160.00 0.0486 CABE 1 86.30 0.0033 COLU 1 389.10 0.0159 CACA 1 51.46 0.0004 CABE 1 42.04 0.0000 COLU 1 335.01 0.0134 CACA 2 59.68 0.0103 CABE 2 169.41 0.0049 COLU 2 163.01 0.0316 CACA 2 106.74 0.0077 CABE 2 99.53 0.0045 COLU 2 161.18 0.0210 CACA 2 109.90 0.0109 CABE 2 32.71 0.0026 COLU 2 132.15 0.0164 CACA 3 23.53 0.0098 CABE 3 234.69 0.0086 COLU 3 190.40 0.0248 CACA 3 59.45 0.0117 CABE 3 158.41 0.0011 COLU 3 187.40 0.0217 CACA 3 55.62 0.0024 CABE 3 74.05 0.0001 COLU 3 163.45 0.0198 COFL 1 114.96 0.0000 COKO 1 47.39 0.0017 CYSC 1 50.29 0.0091 COFL 1 28.59 0.0000 COKO 1 212.16 0.0011 CYSC 1 71.11 0.0270 COFL 1 83.45 0.0026 COKO 1 93.52 0.0000 CYSC 1 124.03 0.0420 COFL 2 95.12 0.0019 COKO 2 87.79 0.0000 CYSC 2 8.73 0.0075 COFL 2 109.71 0.0004 COKO 2 124.69 0.0032 CYSC 2 25.19 0.0351 COFL 2 39.68 0.0002 COKO 2 80.79 0.0000 CYSC 2 112.50 0.0213 COFL 3 151.14 0.0023 COKO 3 144.41 0.0380 CYSC 3 19.61 0.0061 COFL 3 49.60 0.0025 COKO 3 134.55 0.0039 CYSC 3 46.51 0.0183 COFL 3 161.06 0.0052 COKO 3 72.84 0.0005 CYSC 3 3.48 0.0033

Appendix C. cont.

Native Non-native Congener Alien Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass FAGR 1 25.94 0.0000 FASY 1 16.04 0.0003 FOSU 1 180.99 0.0013 FAGR 1 28.30 0.0008 FASY 1 16.48 0.0000 FOSU 1 158.54 0.0038 FAGR 1 62.09 0.0007 FASY 1 25.15 0.0008 FOSU 1 57.27 0.0021 FAGR 2 27.51 0.0000 FASY 2 34.25 0.0017 FOSU 2 146.48 0.0048 FAGR 2 40.09 0.0006 FASY 2 82.60 0.0014 FOSU 2 120.26 0.0011 FAGR 2 16.90 0.0018 FASY 2 35.12 0.0023 FOSU 2 70.68 0.0006 FAGR 3 3.93 0.0000 FASY 3 11.27 0.0003 FOSU 3 56.66 0.0000 FAGR 3 22.40 0.0000 FASY 3 31.00 0.0000 FOSU 3 91.63 0.0112 FAGR 3 9.04 0.0000 FASY 3 47.26 0.0000 FOSU 3 72.03 0.0005 HAVI 1 85.60 0.0000 HAMO 1 65.84 0.0045 GIBI 1 77.29 0.0007 HAVI 1 126.18 0.0001 HAMO 1 167.66 0.0008 GIBI 1 117.20 0.0003 HAVI 1 120.38 0.0001 HAMO 1 163.39 0.0017 GIBI 1 124.03 0.0005 HAVI 2 105.22 0.0263 HAMO 2 113.40 0.0047 GIBI 2 125.83 0.0013 HAVI 2 212.67 0.0062 HAMO 2 32.92 0.0058 GIBI 2 92.75 0.0024 HAVI 2 215.79 0.0291 HAMO 2 112.18 0.0218 GIBI 2 52.49 0.0006 40 HAVI 3 144.01 0.0005 HAMO 3 67.06 0.0005 GIBI 3 210.67 0.0038 HAVI 3 51.72 0.0005 HAMO 3 20.12 0.0000 GIBI 3 139.85 0.0004 HAVI 3 57.96 0.0000 HAMO 3 104.25 0.0000 GIBI 3 8.99 0.0009 JUNI 1 93.36 0.0000 JURE 1 36.81 0.0000 HEHE 1 558.91 0.0008 JUNI 1 81.91 0.0020 JURE 1 64.08 0.0014 HEHE 1 427.71 0.0059 JUNI 1 62.81 0.0000 JURE 1 94.76 0.0013 HEHE 1 329.97 0.0022 JUNI 2 45.41 0.0010 JURE 2 88.63 0.0020 HEHE 2 406.39 0.0080 JUNI 2 59.41 0.0038 JURE 2 131.58 0.0063 HEHE 2 588.43 0.0093 JUNI 2 31.83 0.0022 JURE 2 113.85 0.0114 HEHE 2 487.41 0.0388 JUNI 3 17.82 0.0005 JURE 3 49.09 0.0000 HEHE 3 199.42 0.1462 JUNI 3 42.86 0.0026 JURE 3 126.12 0.0049 HEHE 3 255.18 0.0586 JUNI 3 17.82 0.0000 JURE 3 102.94 0.0001 HEHE 3 261.74 0.0590 MORU 1 113.18 0.0002 MOAL 1 47.08 0.0000 KOPA 1 17.39 0.0000 MORU 1 215.53 0.0000 MOAL 1 68.28 0.0000 KOPA 1 21.07 0.0000 MORU 1 100.80 0.0024 MOAL 1 62.84 0.0002 KOPA 1 18.09 0.0033 MORU 2 526.71 0.0009 MOAL 2 36.20 0.0000 KOPA 2 29.79 0.0022 MORU 2 114.21 0.0017 MOAL 2 59.46 0.0008 KOPA 2 67.33 0.0016 MORU 2 145.15 0.0000 MOAL 2 15.19 0.0008 KOPA 2 7.96 0.0044 MORU 3 64.71 0.0000 MOAL 3 87.79 0.0006 KOPA 3 101.35 0.0049 MORU 3 80.69 0.0000 MOAL 3 18.76 0.0002 KOPA 3 23.69 0.0000 MORU 3 48.98 0.0091 MOAL 3 36.02 0.0026 KOPA 3 28.78 0.0054

Appendix C. cont.

Native Non-native Congener Alien Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass PRSE 1 4.53 0.0000 PRSU 1 374.41 0.0021 LAIN 1 93.71 0.0019 PRSE 1 47.98 0.0000 PRSU 1 433.43 0.0116 LAIN 1 91.76 0.0007 PRSE 1 43.64 0.0000 PRSU 1 415.60 0.0039 LAIN 1 100.59 0.0025 PRSE 2 181.55 0.2536 PRSU 2 641.85 0.0121 LAIN 2 63.89 0.0144 PRSE 2 96.54 0.0109 PRSU 2 447.57 0.0006 LAIN 2 122.38 0.0029 PRSE 2 45.34 0.0009 PRSU 2 520.73 0.0029 LAIN 2 261.06 0.0051 PRSE 3 13.04 0.0000 PRSU 3 513.36 0.0011 LAIN 3 154.62 0.0042 PRSE 3 88.22 0.0003 PRSU 3 369.49 0.0138 LAIN 3 69.74 0.0006 PRSE 3 127.71 0.0174 PRSU 3 276.66 0.0045 LAIN 3 135.58 0.0055 RHPE 1 115.04 0.0055 RHMU 1 206.81 0.0065 LIVU 1 362.52 0.0025 RHPE 1 98.34 0.0000 RHMU 1 147.20 0.0073 LIVU 1 287.74 0.0056 RHPE 1 135.54 0.0011 RHMU 1 145.37 0.0118 LIVU 1 133.71 0.0000 RHPE 2 121.03 0.0124 RHMU 2 191.23 0.0209 LIVU 2 549.57 0.0004 RHPE 2 132.09 0.0045 RHMU 2 277.71 0.0107 LIVU 2 359.76 0.0070 RHPE 2 85.68 0.0017 RHMU 2 95.84 0.0070 LIVU 2 435.65 0.0048 41 RHPE 3 95.23 0.0104 RHMU 3 221.31 0.0056 LIVU 3 280.57 0.0477 RHPE 3 71.86 0.0001 RHMU 3 241.88 0.0034 LIVU 3 63.87 0.0015 RHPE 3 73.70 0.0008 RHMU 3 167.16 0.0851 LIVU 3 126.62 0.0673 ROCA 1 276.59 0.0059 ROMU 1 1505.68 0.0070 PATO 1 636.69 0.0007 ROCA 1 436.75 0.0004 ROMU 1 748.54 0.0002 PATO 1 240.53 0.0006 ROCA 1 362.87 0.0000 ROMU 1 626.73 0.0023 PATO 1 410.31 0.0006 ROCA 2 451.76 0.0010 ROMU 2 112.40 0.0017 PATO 2 702.72 0.0158 ROCA 2 119.74 0.0048 ROMU 2 154.85 0.0061 PATO 2 141.49 0.0028 ROCA 2 106.06 0.0059 ROMU 2 750.17 0.0083 PATO 2 462.19 0.0023 ROCA 3 271.86 0.0510 ROMU 3 173.94 0.0033 PATO 3 301.84 0.0065 ROCA 3 177.62 0.0169 ROMU 3 433.96 0.0125 PATO 3 353.72 0.0057 ROCA 3 136.01 0.0276 ROMU 3 394.14 0.0009 PATO 3 353.72 0.0249 SANI 1 30.74 0.0022 SABA 1 400.37 0.0005 PHAM 1 43.97 0.0000 SANI 1 169.35 0.0098 SABA 1 119.19 0.0053 PHAM 1 65.26 0.0000 SANI 1 46.17 0.0133 SABA 1 1584.26 0.0024 PHAM 1 80.70 0.0047 SANI 2 33.81 0.0009 SABA 2 114.73 0.0015 PHAM 2 170.98 0.0016 SANI 2 27.26 0.0338 SABA 2 267.96 0.4790 PHAM 2 58.71 0.0000 SANI 2 11.52 0.0015 SABA 2 1007.62 0.0004 PHAM 2 16.37 0.0004 SANI 3 28.02 0.0006 SABA 3 153.52 0.0615 PHAM 3 60.11 0.0000 SANI 3 43.03 0.0147 SABA 3 258.59 0.2595 PHAM 3 79.53 0.0000 SANI 3 8.87 0.0000 SABA 3 105.22 0.0245 PHAM 3 43.04 0.0000

Appendix C. cont.

Native Non-native Congener Alien Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass Plant Code Sample Leaf biomass Insect biomass TIAM 1 69.47 0.0009 TICO 1 67.57 0.0024 POTR 1 6.25 0.0000 TIAM 1 41.50 0.0005 TICO 1 32.39 0.0000 POTR 1 5.39 0.0005 TIAM 1 40.15 0.0012 TICO 1 57.49 0.0003 POTR 1 34.57 0.0005 TIAM 2 38.35 0.0006 TICO 2 72.21 0.0111 POTR 2 36.11 0.0031 TIAM 2 41.05 0.0031 TICO 2 19.84 0.0014 POTR 2 14.53 0.0015 TIAM 2 13.53 0.0017 TICO 2 39.52 0.0141 POTR 2 18.94 0.0050 TIAM 3 22.11 0.0000 TICO 3 33.94 0.0096 POTR 3 41.19 0.0000 TIAM 3 26.62 0.0012 TICO 3 23.25 0.0113 POTR 3 18.21 0.0000 TIAM 3 19.40 0.0037 TICO 3 12.24 0.0022 POTR 3 23.17 0.0000 ULAM 1 242.58 0.0032 ULPA 1 375.90 0.0011 PYPA 1 403.61 0.0001 ULAM 1 161.61 0.0000 ULPA 1 210.91 0.0000 PYPA 1 144.92 0.0019 ULAM 1 234.45 0.0031 ULPA 1 537.14 0.0088 PYPA 1 304.58 0.0013 ULAM 2 239.65 0.0061 ULPA 2 503.38 0.0107 PYPA 2 229.83 0.0584 ULAM 2 179.17 0.0033 ULPA 2 487.09 0.0243 PYPA 2 112.70 0.0082 42 ULAM 2 102.43 0.0006 ULPA 2 264.33 0.0024 PYPA 2 67.73 0.0022 ULAM 3 231.52 0.0000 ULPA 3 354.18 0.0000 PYPA 3 52.00 0.0010 ULAM 3 198.03 0.0035 ULPA 3 408.56 0.0360 PYPA 3 201.27 0.0040 ULAM 3 89.75 0.0213 ULPA 3 317.15 0.0000 PYPA 3 105.83 0.0166 VIDE 1 1519.36 0.0008 VIDI 1 523.01 0.0013 SYVU 1 23.97 0.0000 VIDE 1 1161.26 0.0034 VIDI 1 428.32 0.0004 SYVU 1 98.03 0.0010 VIDE 1 1082.98 0.0031 VIDI 1 345.90 0.0005 SYVU 1 78.10 0.0002 VIDE 2 1145.32 0.0181 VIDI 2 967.99 0.0107 SYVU 2 67.06 0.0009 VIDE 2 1010.49 0.0064 VIDI 2 713.72 0.0582 SYVU 2 47.67 0.0020 VIDE 2 756.42 0.0020 VIDI 2 527.83 0.0041 SYVU 2 109.87 0.0646 VIDE 3 1186.64 0.0002 VIDI 3 177.11 0.0053 SYVU 3 95.06 0.0856 VIDE 3 971.35 0.0025 VIDI 3 399.82 0.0125 SYVU 3 139.50 0.0939 VIDE 3 1586.05 0.0039 VIDI 3 557.64 0.0003 SYVU 3 204.67 0.0481

Appendix D. Native plant species and the insects collected (see Methods) from them in 2004 and 2005. Insects were identified to species or Operational Taxonomic Unit (OTU). Also included are the total OTU’s for each plant species.

Native Plant Code Order/Family Insect species or (OTU) a ACRU BENI CACA COFL FAGR HAVI JUNI MORU PRSE RHPE ROCA SANI TIAM ULAM VIDE ORTHOPTERA Acrididae Camnula pellucida 1 Encoptolophus sordidus 1 Melanoplus sanquinipes Tettigoniidae Amblycorypha spp. Conocephalus brevipennis 12 1 2211 Conocephalus spp. Neoconocephalus spp. 1 Gryllidae Allonemobius fasciatus 1 Cyrtoxipha columbiana Gryllus pennsylvanicus 121 Orocharis saltator 1 HEMIPTERA Tingidae Corythucha cydoniae 1 43 Miridae Adelphocoris lineolatus Chlamydatus associatus 11 1 Cylapus spp. 1 Halticus bractatus Lygus lineolaris 85 Berytidae Jalysus spp. Jalysus spp. Lygaeidae Hypogeocoris spp. Lygaeus kalmii 1 25 1 Coreidae Euthochtha galeator 76 2 Pentatomidae Euschistus servus euschistoides 11 5 12 Euschistus tristigmus 1 Euthyrhynchus floridanus Mormidea lugens 14 81 1 102 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix D. cont.

Native Plant Code Order/Family Insect species or (OTU) a ACRU BENI CACA COFL FAGR HAVI JUNI MORU PRSE RHPE ROCA SANI TIAM ULAM VIDE HOMOPTERA Membracidae Micrutalis acutalis Cercopidae 24 2 1 1 1 Philaenus spumaris 11 1 11 121 203 Cicadellidae Aceratogallia spp. 1 Agallia constricta 6327472476535116567 Agallia deleta 12 41 Chloanthanus spp. Chloanthanus spp. Chlorotettix fallax Chlorotettix galbanatus 11 Chlorotettix spp. 1 Chlorotettix spp. 12 1 1 Chlorotettix viridius 44 Deltocephalus flavicosta 1111 31 Doleranus longulus 1 Draeculacephala spp. 11 11 Elymana spp. Elymana spp. 1 Empoasca spp. 1 1 Empoasca spp. 2 3 Empoasca spp. 11 1 3 4 Graphocephala coccinea Graphocephala versuta 113 1 Gyponana spp. Idiocerus spp. Jikradia olitorius 241 1 16 11 Osbornellus jucundus 1 Paraphlepsius spp. Paraphlepsius spp. 1 Paraphlepsius spp. Prairiana spp. 2 Texananus spp. 41 1 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix D. cont.

Native Plant Code Order/Family Insect species or (OTU) a ACRU BENI CACA COFL FAGR HAVI JUNI MORU PRSE RHPE ROCA SANI TIAM ULAM VIDE HOMOPTERA Cicadellidae Texananus spp. 22 3 2 113 155 1 1 163 16 2 1 1 2 2 5 200 3 1 1 202 11 Cixiidae Oliarus sablensis 2 4672118 Flatidae Anormenis chloris 221 205 11 Issidae Thionia simplex 1 Aphidae 9 33 46 3 61 1 98 1 14 45 174 1 COLEOPTERA Scarabaeidae Euphoria herbacea 1 Macrodactylus subspinosus Phyllophaga spp. 94 1 Elateridae Monocrepidius lividus 1 1 Cantharidae Chauliognathus marginatus 1 Podabrus rugulosus Silis percombis Nitidulidae Pallodes spp. 1 Cucujidae Telephanus velox 1 Phalacridae Phalacrus spp. Lathridiidae Malanophthalma distinguenda 1 Mordellidae Mordella marginata Cerambycidae Acmaeops directus Chrysomelidae Brachypnea punticollis 21113 Chaetocnema cribata 11 Colaspis brunnea 347 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix D. cont.

Native Plant Code Order/Family Insect species or (OTU) a ACRU BENI CACA COFL FAGR HAVI JUNI MORU PRSE RHPE ROCA SANI TIAM ULAM VIDE COLEOPTERA Chrysomelidae Diabolia borealis 1 111 Diabrotica undecimpunctata 121 Mantura chrysanthemi 1 Minota spp. 1 Neochlamisus spp. 1 Odontota dorsalis Paria quadrinotata 21 91 1 Curculionidae Acalyptus spp. Hypera punctata 1 Myllocerus helleri 15 Panscopus spp. Pseudoneorhinus bifasciatus 11 3 3 Rhynchaenus niger 46 149 1 Scolytidae Hylurgopinus rufipes LEPIDOPTERA Psychidae Thyridopteryx ephemeraeforms 62 4 3 21 Lyonetiidae 195 9 Gracillariidae 193 2 Gelechiidae 199 2 Yponomeutidae 183 Tortricidae 184 187 11 192 11 194 1 Pyralidae 186 1 197 2 Lycaenidae 185 1 Nymphalidae Limenitis archippus Geometridae Errannis tilaria 1 Iridopsis humaria 2 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix D. cont.

Native Plant Code Order/Family Insect species or (OTU) a ACRU BENI CACA COFL FAGR HAVI JUNI MORU PRSE RHPE ROCA SANI TIAM ULAM VIDE LEPIDOPTERA Geometridae 15 1 165 1 1 2 1 1 166 3 3 1 167 11 168 169 1 170 172 173 191 Saturniidae Automeris io Sphingidae Hemaris thysbe Notodontidae Clostera inclusa 2 47 Schizura concinna 1 Schizura ipomoeae Schizura unicornis 11 2 196 Lymantridae Orgyia leucostigma 1 1 Arctiidae Halysidota tessellaris 11 11 Lophocampa caryae 1 1 Spilosoma virginica 1 Noctuidae Acronicta funeralis 1 Lithophane laticinerea Morrisonia confusa 2 Morrisonia latex 11 Palthis angularis Plathypena scabra Simyra henrici 1 Spodoptera ornithogalli Trichordestra legitima 111 188 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix D. cont. Native Plant Code Order/Family Insect species or (OTU) a ACRU BENI CACA COFL FAGR HAVI JUNI MORU PRSE RHPE ROCA SANI TIAM ULAM VIDE HYMENOPTERA Tenthridinidae Caliroa cerasi Cladius difformis 6 Macrophya spp. 1 161 6 Total # OTU's 10252810 5 1910 7 18162519101327 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005 48

Appendix E. Non-native Congeners and the insects collected (see Methods) from them in 2004 and 2005. Insects identified to species or Operational Taxonomic Unit (OTU). Also included are the total OTU’s for each plant species.

Non-native Congener Plant Code Order/Family Insect species or (OTU) a ACPL BEPE CABE COKO FASY HAMO JURE MOAL PRSU RHMU ROMU SABA TICO ULPA VIDI ORTHOPTERA Acrididae Camnula pellucida Encoptolophus sordidus Melanoplus sanquinipes 111 Tettigoniidae Amblycorypha spp. 1 Conocephalus brevipennis 11 1814 Conocephalus spp. Neoconocephalus spp. Gryllidae Allonemobius fasciatus 1 Cyrtoxipha columbiana 1 Gryllus pennsylvanicus 1 Orocharis saltator 1 HEMIPTERA

49 Tingidae Corythucha cydoniae 34 21 Miridae Adelphocoris lineolatus 1 Chlamydatus associatus 11 Cylapus spp. Halticus bractatus Lygus lineolaris 1 85 Berytidae Jalysus spp. Jalysus spp. 1 Lygaeidae Hypogeocoris spp. Lygaeus kalmii 25 Coreidae Euthochtha galeator 1 76 Pentatomidae Euschistus servus euschistoides 11 2 92 1 4 Euschistus tristigmus 1 1 Euthyrhynchus floridanus Mormidea lugens 1 14 11 81 102 1 1 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix E. cont.

Non-native Congener Plant Code Order/Family Insect species or (OTU) a ACPL BEPE CABE COKO FASY HAMO JURE MOAL PRSU RHMU ROMU SABA TICO ULPA VIDI HOMOPTERA Membracidae Micrutalis acutalis 11 Cercopidae 24 1 1 1 1 Philaenus spumaris 12141121 203 Cicadellidae Aceratogallia spp. Agallia constricta 9 2 9163101951247185344612 Agallia deleta 2111 Chloanthanus spp. 1 Chloanthanus spp. 11 1 Chlorotettix fallax 1 Chlorotettix galbanatus 11 Chlorotettix spp. Chlorotettix spp. 2 3 Chlorotettix viridius 50 Deltocephalus flavicosta 113 15 Doleranus longulus Draeculacephala spp. 11 Elymana spp. Elymana spp. 1 Empoasca spp. Empoasca spp. 2 411 33 Empoasca spp. 33 21 3 3 Graphocephala coccinea Graphocephala versuta 11 12 1 Gyponana spp. 1 Idiocerus spp. Jikradia olitorius 1321 7 19 672 Osbornellus jucundus 1 1 Paraphlepsius spp. Paraphlepsius spp. 1 Paraphlepsius spp. 1 Prairiana spp. 11 Texananus spp. 621 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix E. cont.

Non-native Congener Plant Code Order/Family Insect species or (OTU) a ACPL BEPE CABE COKO FASY HAMO JURE MOAL PRSU RHMU ROMU SABA TICO ULPA VIDI HOMOPTERA Cicadellidae Texananus spp. 31 6 155 24 163 8 3 2 3 1 3 4 200 1 1 202 1 Cixiidae Oliarus sablensis 3232214272214 Flatidae Anormenis chloris 21152 205 2 Issidae Thionia simplex Aphidae 9 33 587 46 188 61 98 51 174 COLEOPTERA Scarabaeidae Euphoria herbacea Macrodactylus subspinosus Phyllophaga spp. 94 1 Elateridae Monocrepidius lividus 1 Cantharidae Chauliognathus marginatus Podabrus rugulosus Silis percombis 1 Nitidulidae Pallodes spp. Cucujidae Telephanus velox Phalacridae Phalacrus spp. Lathridiidae Malanophthalma distinguenda Mordellidae Mordella marginata 1 Cerambycidae Acmaeops directus Chrysomelidae Brachypnea punticollis 1111326 Chaetocnema cribata 11 Colaspis brunnea 121 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix E. cont.

Non-native Congener Plant Code Order/Family Insect species or (OTU) a ACPL BEPE CABE COKO FASY HAMO JURE MOAL PRSU RHMU ROMU SABA TICO ULPA VIDI COLEOPTERA Chrysomelidae Diabolia borealis 112 1 Diabrotica undecimpunctata 1111 Mantura chrysanthemi Minota spp. Neochlamisus spp. Odontota dorsalis 11 Paria quadrinotata 1 91 Curculionidae Acalyptus spp. 1 Hypera punctata Myllocerus helleri 13 Panscopus spp. 1 Pseudoneorhinus bifasciatus 11 1121 5 Rhynchaenus niger 1 52 149 Scolytidae Hylurgopinus rufipes 1 LEPIDOPTERA Psychidae Thyridopteryx ephemeraeforms 466 12 2 214 Lyonetiidae 195 Gracillariidae 193 Gelechiidae 199 Yponomeutidae 183 Tortricidae 184 1 187 192 194 Pyralidae 186 197 Lycaenidae 185 1 Nymphalidae Limenitis archippus 2 Geometridae Errannis tilaria 1 Iridopsis humaria 1 21 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix E. cont.

Non-native Congener Plant Code Order/Family Insect species or (OTU) a ACPL BEPE CABE COKO FASY HAMO JURE MOAL PRSU RHMU ROMU SABA TICO ULPA VIDI LEPIDOPTERA Geometridae 15 1 1 165 2 1 1 2 166 1 1 1 1 167 1 168 1 169 170 172 1 173 1 191 1 Saturniidae Automeris io 14 1 12 Sphingidae Hemaris thysbe 1 53 Notodontidae Clostera inclusa 281 Schizura concinna Schizura ipomoeae Schizura unicornis 21 1 1 1 3 196 1 Lymantridae Orgyia leucostigma 311111 Arctiidae Halysidota tessellaris 111 111 Lophocampa caryae Spilosoma virginica 11 Noctuidae Acronicta funeralis Lithophane laticinerea 1 Morrisonia confusa 12 Morrisonia latex Palthis angularis 1 Plathypena scabra 1 Simyra henrici Spodoptera ornithogalli 1 Trichordestra legitima 111 188 1 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix E. cont.

Non-native Congener Plant Code Order/Family Insect species or (OTU) a ACPL BEPE CABE COKO FASY HAMO JURE MOAL PRSU RHMU ROMU SABA TICO ULPA VIDI HYMENOPTERA Tenthridinidae Caliroa cerasi Cladius difformis 2 Macrophya spp. 161 2 Total # OTU's 16 22 16 11 11 13 13 5 24 21 33 12 17 22 31 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005 54

54 Appendix F. Alien plant species and the insects collected (see Methods) from them in 2004 and 2005. Insects identified to species or Operational Taxonomic Unit (OTU). Also included are the total OTU’s for each plant species. Alien Plant Code Order/Family Insect species or (OTU) a AKQU ALJU COLU CYSC FOSU GIBI HEHE KOPA LAIN LIVU PATO PHAM POTR PYPA SYVU ORTHOPTERA Acrididae Camnula pellucida Encoptolophus sordidus 1 Melanoplus sanquinipes 112 Tettigoniidae Amblycorypha spp. Conocephalus brevipennis 101211 Conocephalus spp. 1 Neoconocephalus spp. Gryllidae Allonemobius fasciatus 1 Cyrtoxipha columbiana Gryllus pennsylvanicus Orocharis saltator HEMIPTERA

55 Tingidae Corythucha cydoniae 1188 1 3 Miridae Adelphocoris lineolatus 1 Chlamydatus associatus 112 Cylapus spp. Halticus bractatus 11 Lygus lineolaris 113 85 1 Berytidae Jalysus spp. 9 Jalysus spp. 13 Lygaeidae Hypogeocoris spp. 11 Lygaeus kalmii 25 Coreidae Euthochtha galeator 76 1 Pentatomidae Euschistus servus euschistoides 2 423172 415 28 Euschistus tristigmus 23 Euthyrhynchus floridanus 11 Mormidea lugens 14 1 81 102 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix F. cont.

Alien Plant Code Order/Family Insect species or (OTU) a AKQU ALJU COLU CYSC FOSU GIBI HEHE KOPA LAIN LIVU PATO PHAM POTR PYPA SYVU HOMOPTERA Membracidae Micrutalis acutalis Cercopidae 24 1 1 Philaenus spumaris 224 111 11 203 1 Cicadellidae Aceratogallia spp. 11 Agallia constricta 15 4 14 8 9 143219601033 4 192616 Agallia deleta 11 3 6 Chloanthanus spp. 11 1 Chloanthanus spp. 3 Chlorotettix fallax 2 Chlorotettix galbanatus 211 1 Chlorotettix spp. 1 Chlorotettix spp. 33

56 Chlorotettix viridius 1 Deltocephalus flavicosta 221 Doleranus longulus Draeculacephala spp. 31 Elymana spp. 1 Elymana spp. 1 Empoasca spp. Empoasca spp. 452 2 1 Empoasca spp. 13 4 31 1 Graphocephala coccinea 1 Graphocephala versuta 1123 Gyponana spp. Idiocerus spp. 1 Jikradia olitorius 114 23 1 Osbornellus jucundus 114 1 Paraphlepsius spp. 1 Paraphlepsius spp. 7 Paraphlepsius spp. Prairiana spp. 11 3 Texananus spp. 313 13 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix F. cont.

Alien Plant Code Order/Family Insect species or (OTU) a AKQU ALJU COLU CYSC FOSU GIBI HEHE KOPA LAIN LIVU PATO PHAM POTR PYPA SYVU HOMOPTERA Cicadellidae Texananus spp. 13 2 1 3 3 155 1 163 72 3 11 11 200 1 2 202 Cixiidae Oliarus sablensis 9516910161561 141 Flatidae Anormenis chloris 121 1 205 1 Issidae Thionia simplex 1 Aphidae 9 2 33 1 941 2 46 61 98 1 57 174 1 COLEOPTERA Scarabaeidae Euphoria herbacea Macrodactylus subspinosus 1 Phyllophaga spp. 1 94 Elateridae Monocrepidius lividus 2 Cantharidae Chauliognathus marginatus 1 Podabrus rugulosus 1 Silis percombis Nitidulidae Pallodes spp. Cucujidae Telephanus velox Phalacridae Phalacrus spp. 1 Lathridiidae Malanophthalma distinguenda 1 Mordellidae Mordella marginata 1 Cerambycidae Acmaeops directus 1 Chrysomelidae Brachypnea punticollis 143 Chaetocnema cribata 11 2 13 11 Colaspis brunnea 26 1 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix F. cont. Alien Plant Code Order/Family Insect species or (OTU) a AKQU ALJU COLU CYSC FOSU GIBI HEHE KOPA LAIN LIVU PATO PHAM POTR PYPA SYVU COLEOPTERA Chrysomelidae Diabolia borealis 32 1 1 1 2 Diabrotica undecimpunctata 1111 Mantura chrysanthemi Minota spp. 1 Neochlamisus spp. Odontota dorsalis 1 Paria quadrinotata 91 Curculionidae Acalyptus spp. 11 1 1 1 Hypera punctata Myllocerus helleri 37 2 Panscopus spp. Pseudoneorhinus bifasciatus 31 1 Rhynchaenus niger 58 149 Scolytidae Hylurgopinus rufipes LEPIDOPTERA Psychidae Thyridopteryx ephemeraeforms 3 Lyonetiidae 195 Gracillariidae 193 Gelechiidae 199 Yponomeutidae 183 1 Tortricidae 184 1 187 1 192 194 Pyralidae 186 197 Lycaenidae 185 Nymphalidae Limenitis archippus Geometridae Errannis tilaria Iridopsis humaria 2 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix F. cont.

Alien Plant Code Order/Family Insect species or (OTU) a AKQU ALJU COLU CYSC FOSU GIBI HEHE KOPA LAIN LIVU PATO PHAM POTR PYPA SYVU LEPIDOPTERA Geometridae 15 165 1 1 1 166 3 1 167 168 1 169 170 1 172 173 191 Saturniidae Automeris io 51 Sphingidae Hemaris thysbe

59 Notodontidae Clostera inclusa Schizura concinna Schizura ipomoeae 2 Schizura unicornis 51 196 Lymantridae Orgyia leucostigma 11 1 1 Arctiidae Halysidota tessellaris 1 Lophocampa caryae Spilosoma virginica 12 Noctuidae Acronicta funeralis Lithophane laticinerea Morrisonia confusa Morrisonia latex Palthis angularis Plathypena scabra Simyra henrici Spodoptera ornithogalli Trichordestra legitima 188 1 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005

Appendix F. cont.

Alien Plant Code Order/Family Insect species or (OTU) a AKQU ALJU COLU CYSC FOSU GIBI HEHE KOPA LAIN LIVU PATO PHAM POTR PYPA SYVU HYMENOPTERA Tenthridinidae Caliroa cerasi 1 Cladius difformis Macrophya spp. 161 Total # OTU's 16103627 8 9 26 8 151719 3 7 2314 a Total number collected on nine plants, 3 in June, 3 in July, and 3 in August in 2004 and 2005 60

Appendix G. Simpson’s diversity (probability that two randomly chosen individuals from a community belong to the same species) for each plant species for 2004, 2005, and both years combined. See Table 1 for plant codes. Native Non-native Congener Alien Plant Code Diversity Plant Code Diversity Plant Code Diversity 2004 ACRU 0.62 ACPL 0.86 AKQU 0.66 BENI 0.84 BEPE 0.7 ALJU 0.5 CACA 0.88 CABE 0.75 COLU 0.12 COFL 0 COKO 0 CYSC 0.76 FAGR 0.5 FASY 0.75 FOSU 0.5 HAVI 0.88 HAMO 0.75 GIBI 0.16 JUNI 0.22 JURE 0.8 HEHE 0.81 MORU 0.56 MOAL 0 KOPA 0 PRSE 0.8 PRSU 0.66 LAIN 0.5 RHPE 0.88 RHMU 0.81 LIVU 0.75 ROCA 0.87 ROMU 0.92 PATO 0.8 SANI 0.9 SABA 0.81 PHAM 0 TIAM 0.75 TICO 0.87 POTR 0 ULAM 0.83 ULPA 0.75 PYPA 0.89 VIDE 0.91 VIDI 0.43 SYVU 0.83 2005 ACRU 0.79 ACPL 0.8 AKQU 0.79 BENI 0.84 BEPE 0.9 ALJU 0.87 CACA 0.71 CABE 0.75 COLU 0.56 COFL 0.87 COKO 0.68 CYSC 0.21 FAGR 0.37 FASY 0.8 FOSU 0.73 HAVI 0.58 HAMO 0.79 GIBI 0.38 JUNI 0.79 JURE 0.63 HEHE 0.75 MORU 0.69 MOAL 0.56 KOPA 0.54 PRSE 0.88 PRSU 0.81 LAIN 0.44 RHPE 0.52 RHMU 0.74 LIVU 0.85 ROCA 0.86 ROMU 0.86 PATO 0.57 SANI 0.88 SABA 0.09 PHAM 0.32 TIAM 0.77 TICO 0.59 POTR 0.36 ULAM 0.61 ULPA 0.62 PYPA 0.77 VIDE 0.88 VIDI 0.81 SYVU 0.63 Both Years ACRU 0.82 ACPL 0.87 AKQU 0.81 BENI 0.88 BEPE 0.91 ALJU 0.85 CACA 0.83 CABE 0.87 COLU 0.44 COFL 0.87 COKO 0.75 CYSC 0.24 FAGR 0.56 FASY 0.88 FOSU 0.79 HAVI 0.69 HAMO 0.81 GIBI 0.48 JUNI 0.77 JURE 0.71 HEHE 0.86 MORU 0.75 MOAL 0.64 KOPA 0.54 PRSE 0.89 PRSU 0.9 LAIN 0.46 RHPE 0.66 RHMU 0.76 LIVU 0.87 ROCA 0.92 ROMU 0.9 PATO 0.8 SANI 0.93 SABA 0.13 PHAM 0.5 TIAM 0.84 TICO 0.68 POTR 0.41 ULAM 0.85 ULPA 0.77 PYPA 0.83 VIDE 0.93 VIDI 0.47 SYVU 0.74

61 Appendix H. Species richness (total number of insect species found on a plant species) for each plant species for 2004, 2005, and both years combined. See Table 1 for plant codes. Native Non-native Congener Alien Plant Code Richness Plant Code Richness Plant Code Richness

2004 ACRU3ACPL7AKQU3 BENI 8 BEPE 6 ALJU 2 CACA 14 CABE 8 COLU 20 COFL 0 COKO 1 CYSC 6 FAGR 2 FASY 4FOSU2 HAVI 9 HAMO 4 GIBI 4 JUNI 3 JURE 5 HEHE 13 MORU 3 MOAL 1 KOPA 0 PRSE 5 PRSU 3 LAIN 2 RHPE 11 RHMU 6 LIVU 4 ROCA 14 ROMU 15 PATO 10 SANI 10 SABA 6PHAM1 TIAM 4 TICO 8 POTR 1 ULAM 6 ULPA 12 PYPA 11 VIDE 14 VIDI 21 SYVU 6 2005 ACRU 8 ACPL 11 AKQU 13 BENI 19 BEPE 18 ALJU 10 CACA 18 CABE 10 COLU 24 COFL 10 COKO 10 CYSC 24 FAGR 3 FASY 7FOSU8 HAVI 11 HAMO 9 GIBI 5 JUNI 8 JURE 9 HEHE 16 MORU 5 MOAL 4 KOPA 8 PRSE 15 PRSU 23 LAIN 14 RHPE 9 RHMU 17 LIVU 15 ROCA 15 ROMU 24 PATO 14 SANI 12 SABA 7PHAM2 TIAM 8 TICO 11 POTR 6 ULAM 7 ULPA 14 PYPA 15 VIDE 15 VIDI 14 SYVU 8 Both Years ACRU 10 ACPL 16 AKQU 16 BENI 25 BEPE 22 ALJU 10 CACA 28 CABE 16 COLU 36 COFL 10 COKO 11 CYSC 27 FAGR 5 FASY 11 FOSU 8 HAVI 19 HAMO 13 GIBI 9 JUNI 10 JURE 13 HEHE 26 MORU 7 MOAL 5 KOPA 8 PRSE 18 PRSU 25 LAIN 15 RHPE 16 RHMU 21 LIVU 17 ROCA 25 ROMU 33 PATO 20 SANI 19 SABA 12 PHAM 3 TIAM 10 TICO 17 POTR 7 ULAM 13 ULPA 22 PYPA 24 VIDE 28 VIDI 32 SYVU 14

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