Classical Weed Biological Control Outcomes:

A Catalogue-based Analysis of Success Rates

and Their Correlates

Submitted by: Mike Barbetta

Supervisors: Dr. Jay Malcolm and Dr. Sandy Smith

Final Capstone Submission for FOR3008H

April 12th

, 2018

Abstract

Classical weed biological control (hereafter CWBC) is an important and effective

management tool that may see an expanded role as more problematic weeds emerge worldwide,

more countries begin to adopt this approach, and broader applications of the practice are

explored. However, success is never certain in CWBC, and predicting success remains an

elusive goal. Here, a series of catalogue-based analyses of CWBC outcomes are conducted in

order to quantify success rates and to explore potential factors that may be associated with

success. Statistical analyses utilizing chi-squared tests were utilized to search for correlations

between CWBC efficacy and various host and agent characteristics. Multiple such relationships

were identified, including previously identified correlations between CWBC outcomes, host

habitat types, and agent orders. Significant differences in CWBC outcomes are also correlated to

agent feeding guilds, which represents a novel association revealed by this study that warrants

further investigation. For example, defoliating agents were associated with lower establishment

rates, and root boring agents were associated with higher control efficacy. Success rates derived

from the most recent catalogue were also delineated by region and time period, and management

implications are discussed.

INTRODUCTION

The deliberate introduction of exotic agents to control problematic invasive species; that

is, classical biological control, is an important tool in the management of weeds worldwide. In

many scenarios, use of classical biological control for weed control (hereafter CWBC) is touted

as the only and best option because conventional control techniques are often ineffective against

problematic weed species, and because CWBC offers the potential for significant, cost-effective

management impacts in the form of self perpetuating, permanent, landscape-wide control

mechanisms (Van Driesche et al. 2010). To date, over 450 insect agents have been released

against over 150 plant species in over 90 countries and regions worldwide (Winston et al. 2014),

with many notable successes (McFadyen 2000). Although the practice does entail the risk of

non-target impacts occurring (Myers and Bazely 2003), the ongoing and potentially expanded

use of this practice is encouraged by a number of factors: significant agent impacts on non-target

hosts are rare (Suckling and Sforza 2014), the number of problematic invasive weed species is

expected to increase worldwide (Seastedt 2015), the goals and applications of CWBC (including

relatively unexplored forestry applications) continue to evolve and expand (Van Driesche et al.

2010; MacQuarrie et al. 2016), and previously reluctant European jurisdictions appear willing to

make use of biological control (Shaw et al. 2016; Varia et al. 2016).

However, the efficacy of CWBC as a weed management tool is plagued by several

pervasive issues. Success is uncertain, widely variable, and is often inadequately monitored so as

to make accurate evaluation difficult (Suckling 2013; Seastedt 2015). Success prediction and

improvement in control agent selection and release is a complex and challenging task that,

despite recent attention and efforts, remains an elusive goal (Seastedt 2015). This is problematic

given the high costs of initiating and carrying out new biological control programs (Myers and

Bazely 2003), and the significant cost increases (Paynter et al. 2015) and inherent risks (Delfosse

2005) that are incurred with each additional agent that is required for any given program.

Determining success rates in CWBC can be difficult. Depending on the metric of success

being used (e.g., if the success of each individual agent is being evaluated, or if the net success of

a set of agents deployed against a specific host plant is being evaluated), reported success rates

can vary significantly (McFadyen 1998; Fowler et al. 2008). Furthermore, there is tendency in

CWBC publications (even those published very recently) to cite dated articles for this

information despite much more up to date information being available (Fowler et al. 2008;

DiTomaso et al. 2017). Attempting to establish generalized ‘rules-of-thumb’ to guide CWBC

releases is also a contested issue, with context-specific factors often cited as more important

determinants of success in CWBC programs (McFadyen 1998; Center et al. 2014; Seastedt

2015). Nevertheless, several authors have explored or suggested basic host and agent

characteristics as potential indicators of success in CWBC (Boughton and Pemberton 2008;

Clewey et al. 2012; Paynter et al. 2012) and the potential merits of post-hoc analysis can only

increase with time as more data is accumulated. An exhaustive and regularly updated catalogue

of worldwide CWBC releases and their results is available (Winston et al. 2014) and there may

be value in analyzing the information within this catalogue to identify and explore trends in the

successes and failures of CWBC worldwide.

Several reviews of the results of CWBC releases in relation to potential explanatory

variables have been conducted. Paynter et al. (2012) discovered that CWBC programs deployed

against plant hosts that resided in aquatic habitats, reproduced vegetatively, and/or were not

major weeds within their native ranges were much more effective than programs deployed

against plant hosts that lacked any of these characteristics. Using 61 published CWBC studies,

Clewey et al. (2012) discovered that Coleopteran agents from Chrysomelidae and Curculionidae

were more effective CWBC agents than other insect types. While both of these studies utilized

an analysis of precise quantitative data, it is still possible to take a similar approach using the

more generalized information available in the Winston et al. (2014) catalogue. Doing so involves

an important trade-off: the information within the catalogue consists of broad, generalized

categories, but it also much more comprehensive, readily available, and therefore easily applied

to evaluations of CWBC efficacy.

While Boughton and Pemeberton (2008) utilized a catalogue-based approach to analyze

establishment success rates of previous CWBC agents across different insect orders and

lepidopteran families, further catalogue analyses of CWBC outcomes has been limited.

The purpose of the current paper is to build on this approach through a series of post-hoc

catalogue-based analyses. My first objective was to quantify success rates of CWBC releases in

general, by major practicing jurisdiction, and in respect to how these rates may be changing over

time. Crawley (1989) indicated that success in CWBC was decreasing over time, whereas more

recently improved rates of success have been reported in certain regions (Hayes 2000; Fowler et

al. 2008). There is a need to update the information concerning CWBC success rates and to

present in a way that is useful for informing management decision making, avoids bias, and

clearly differentiates the different evaluation criteria that is used to assess success rates.

A second objective was to assess CWBC release outcomes in relation to pertinent host

and agent characteristics. Catalogue analysis with up to date information may allow the

confirmation of previously identified correlations (e.g., improved outcomes when utilizing

coleopteran agents (Crawley 1989; Boughton and Pemberton 2008; Clewley et al. 2012) or

against aquatic hosts (Crawley 1989; Paynter et al. 2012)), as well the exploration of other

possible trends. Myers and Bezely (2003) suggest that certain agent feeding guilds (e.g., seed

predation) may act as less effective control mechanisms than others. Therefore, the evaluation of

CWBC release outcomes in relation to agent feeding guilds will also be conducted to in order

provide an initial cursory evaluation of these previously unexplored relationships.

The results of this meta-analysis may be significant in a number of ways. If the results

obtained from this catalogue meta-analysis are found to emulate results from non-catalogue

based analysis (i.e., better CWBC against aquatic hosts, and when utilizing coleopteran agents),

this will help support the legitimacy of the catalogue-based approach. As such, the results

obtained from the novel relationships being explored (i.e. agent efficacy by feeding guilds) may

be more confidently used to help inform agent prioritization, or identify areas of interest for more

robust investigation. Furthermore, the investigation of the overall efficacy of CWBC in general,

by jurisdiction, and over time will evaluate how often and to what extent this weed management

strategy has been successful in the past, where it has been most successful, and whether or not

the efficacy of the practice has stagnated, improved, or deteriorated over time. This will assist in

informing management expectations and strategies when deciding upon options for the

management of problematic weeds.

METHODS

Data compilation of catalogue and supplementary information

Catalogue information, which contains the date, location, basic host (i.e., family and

species name) and agent characteristics (i.e., species name and classification), and outcome of

every CWBC release worldwide up until 2012 (Winston et al. 2014), was utilized as the primary

source for analysis. Information from the catalogue website (http://www.ibiocontrol.org/catalog)

was used to update project results where appropriate when additional information had become

available since the most recent catalogue publication date (date of most recent access: February

2017). Results were evaluated according to the establishment and general impact information

included for each release. Establishment information is categorized as either ‘Yes’ or ‘No’

depending on whether or not evidence of a self-perpetuating agent population had been

discovered within the release jurisdiction following their release. Impact information is

categorized according to five impact outcomes for each release where establishment occurred

(i.e., ‘None’, ‘Slight’, ‘Variable’, ‘Medium’, or ‘High’), as determined by the subjective

estimates of the catalogue editors (Winston et al. 2014).

Host habitat type was determined primarily by utilizing CABI weed datasheets (accessed

at cabi.org), with a small number requiring reference to other sources when the CABI datasheets

were incomplete or unclear. Each of 139 host weeds used in the analysis was classified as

‘aquatic’ or ‘terrestrial’. This classification followed the rational of Paynter et al (2012), who

assigned an aquatic classification to weeds which were described as being either aquatic (i.e.

emergent, floating, or submerged) or wetland plants which experience regular seasonal flooding.

See Appendix 1 for a complete list of host weed species and their habitat classifications. Agent

feeding guilds were determined with reference to available academic sources on the over 388

different CWBC agents included in the analysis. Reviews of CWBC programs which included

agent lists and their associated feeding guilds were deferred to when possible (e.g., Klein 2011),

otherwise individual, agent-specific sources were utilized. See Appendix 2 for a complete list of

insect agents and their feeding guild classifications.

Data Analysis

Due to the delays which often occur before successes in CWBC releases can be reliably

assessed, and in accordance with the recommendations of the Catalogue editors (Winston et al.

2014), data from release programs less than ten years old were not be included in the analysis

(i.e., any releases which occurred post-2007). Furthermore, only release information from the six

countries (Hawaii and the continental USA, treated separately) that have most frequently utilized

CWBC were used in the analysis. While this represents a small set of the 94 countries

represented in the catalogue, approximately 65% of the releases reported in the catalogue

occurred within these 6 top countries. Parsing the data in this manner allowed for appropriate

sample sizes for analyzing CWBC outcomes across different countries, while still providing a

representative picture of global CWBC outcomes. Furthermore, it also helped to avoid problems

that might be associated with including release data from countries which have minimal

involvement in CWBC (e.g., Cuba, with a single release of unknown outcome conducted in

1995).

For the purposes of this study, agent impact data taken from the catalogue was condensed

from five to three categories, leaving the three control impact categories as “None/Slight”,

“Medium/Variable”, and “High”. This was necessary in order to establish adequate sample sizes

for the analyses being conducted. In my estimation, having familiarized myself at length with

the case-by-case agent impact values and the accompanying notes describing these impacts, this

is the most appropriate way to combine these categories because the control outcomes of the

merged categories appear to be most strongly correlated to each other. That is, agents classified

with ‘Slight’ impacts are typically described as having negligible potential control benefits (and

can thus be paired with ‘None’); agents classified with ‘Medium’ or ‘Variable’ impacts are

typically described as only providing partial control benefits or control in certain contexts

(habitats, etc.) and will require additional measures or agents before satisfactory control can be

attained; and agents classified with ‘High’ control impacts are typically described as providing

substantial to complete control benefits with minimal or no need for further control measures.

Results for both establishment and impact levels were evaluated according to the best

outcome attained across all releases within the parameters being investigated. For agent

establishment, an agent was counted as successfully established so long as there was a single

successful establishment recorded, regardless of how many failed releases preceded (or

followed) that success. For agent impact, the highest impact level attained across all releases was

used for evaluation, regardless of whether other releases attained lower impacts. Host impacts

were evaluated according to the highest agent impact level attained across all releases of all

agents released to target that host.

Evaluating success rates

Success rates were evaluated according to agent establishment and the general impact of

agents once established. Rates were calculated on an agent by agent basis as well as on a host by

host basis in order to evaluate how often and to what degree individual CWBC agents were

successful, and how often and to what degree CWBC programs targeting a specific weed host

were successful. In addition to this, individual success rates for each jurisdiction were evaluated

as well as rates before/after 1989, the median year for the number of CWBC agent releases being

investigated.

Release outcomes by agent/host characteristics

Following the results obtained by Paynter et al (2012), and in accordance with the long

standing observation that CWBC has generally been more successful when targeting aquatic

hosts (McFadyen 1998), agent and host program success rates were evaluated for host habitat

type.

Agent success rates were evaluated according to major insect orders. This is similar to the

catalogue meta-analysis conducted by Boughton and Pemberton (2008) for establishment rates,

but has also been extended to include the control impact of established agents.

Agent success rates in relation to feeding guilds were evaluated for the feeding guilds that

were represented by a minimum of at least twenty agents. Guilds represented by less than twenty

agents were excluded from analysis. In order to include additional agents excluded from this

initial analysis, another analysis was conducted which grouped guild information into the area of

the host attacked. This was done under the assumption that although the feeding mechanism may

vary, agents which attack the same location of a plant (e.g., the roots, leaves, etc.) will have a

comparably similar impact on plant fitness.

Statistical Analysis

Contingency table analyses (chi-square) were used to test for deviations from random

expectations.

RESULTS

Release and program success rates

Seventy-one percent of all 388 assessed CWBC agents successfully established in at least

one instance across all release attempts across all jurisdictions for any given agent (Fig. 1).

Investigation of establishment success by region (Fig. 3) revealed no significant difference

among regions although some variation occurred, with the continental USA representing the best

result (79% establishment rate) and Australia and Hawaii representing the worst results (66%

establishment rates). Of the 262 established agents for which impact information was available,

the best result across all releases and regions for each agent was reported as ‘none’ or ‘slight’

45% of the time, ‘medium’ or ‘variable’ 31% of the time, and ‘high’ 24% of the time.

Controlling for agent impact success by region (Fig. 3) revealed no significant agent impact

difference among regions although some regions did have noticeably higher frequencies of low

impact agents and lower frequencies of high impact agents (53%/15%, 53%/13%, 54%/8%, for

Canada, Hawaii, and New Zealand, respectively) in comparison to others (39%/27%, 33%/29%,

for USA, and South Africa, respectively). Controlling for agent establishment and impact by

before or after the median year of 1989 yielded no significance, as results were very similar for

both metrics in each time period (Fig. 2).

Figure 1. General success rates of CWBC agents and programs through 2007. Figures in

percentage, with the sample size in each category at the top of each column.

Figure 2. Success rates of CWBC agents and programs by before/after median year of 1989.

Figures and sample sizes as above. p-values derived from Chi-square analysis are shown for

each plot.

Of the CWBC programs initiated against the139 target weeds assessed, the best result

across all releases and regions for each host were ‘no establishment’ in 9% of cases, ‘none’ or

‘slight’ in 20% of cases ‘medium’ or ‘variable’ in 30% of cases, and ‘high’ in ‘40%’ of cases

(Fig. 1). Controlling for region revealed considerable variation in program success rates by

region (Fig. 3), and while significance was indicated (p=0.0206), low cell counts may have

rendered the chi-square test suspect. South Africa was host to the best results, with 80% of

targeted weeds being host to at least one agent with a mid or high level impact. Conversely, New

Zealand was host to the worst results, with only 38% of targeted weeds being host to an agent

with mid or high level control impact. This result is consistent with the differences in CWBC

agent efficacy by region as noted above, and adds further corroboration to CWBC successes

being correlated to certain regions (McFadyen 1998). Controlling for program impact by before

or after the median year of 1989 revealed no significance, as each period produced nearly

identical program outcomes by best agent impact (Fig. 2).

Figure 3. Success rates of CWBC agents and programs by region. Figures, samples sizes and p-

values as in Figures 1 and 2.

Figure 4. Success rates of CWBC agents and programs against either terrestrial or

aquatic/wetland hosts. Figures, sample sizes, and p-values as in Figures 1 and 2.

Results by host habitat

Agent establishment rates by host habitat (terrestrial or aquatic/wetland) were not

significantly different (71% and 70% respectively, p=0.8535) (Fig. 4). However, differences in

agent impact rates by host habitat were highly significant (p=0.0008) (Fig. 4), with agents

released against aquatic or wetland hosts being generally much more effective than those

released against terrestrial hosts. Evaluating program outcome by host habitat type also revealed

highly significant results (p=0.0009; Fig. 4), with CWBC programs against aquatic hosts being

much more frequently successful in establishing a high impact agent compared to programs

targeting terrestrial hosts. Although the small sample size of aquatic hosts (n = 13) may render

the chi-square test somewhat suspect, it should be noted that virtually all aquatic hosts targeted

by CWBC have been successfully paired with an insect agent offering a high degree of control.

The one ‘not established’ exception in the current analysis refers to a single unsuccessful release

attempt from 2004 made against Old World climbing fern (Lygodium microphyllum) in the USA.

Since that time, successful establishment of insect CWBC agents have occurred against this host

(Winston et al. 2014), but as they occurred past the cut-off date of 2007 for this analysis, these

results were not included.

Figure 5. Success rates of CWBC agents by insect order. Figures, sample sizes and p-

values as above.

Results by agent order

Agent establishment by the four major insect orders evaluated yielded no significant

difference (p=0.3157; Fig. 5), with hemipteran agents exhibiting the highest establishment rates

(77%), and lepidopteran agents the lowest (64%). Agent impact by insect order revealed a nearly

significant difference (p=0.056), with hemipteran and coleopteran agents most frequently having

a mid or high level impact (27%/42% and 33%/33%, respectively), and dipteran agents having

the least (28%/8%).

Results by feeding location/guild

Agent impact by feeding location revealed a highly significant difference (p=0.0021; Fig.

6) with agents that attack roots most frequently having a mid or high impact, and flower

attacking agents having the least impact with only 37% of agents having a mid or high level

impact, and only 1 of the 19 evaluated agents having a high impact. Impacts for specific feeding

guilds showed similar results for root boring, seed predating and flower feeding agents when

compared to their analogous location types. However, a noticeable difference emerged between

the two leaf feeding guilds, with defoliating agents having high impacts more frequently and leaf

mining agents having mid or high impacts less frequently than the evaluation for general location

type analysis revealed. Similarly, stem boring insects were revealed to have a high impact more

frequently than the more generalized location analysis revealed.

Agent establishment by feeding location revealed no significant difference between

location types, whereas establishment by specific feeding guilds yielded a significant result

(p=0.019; Fig. 6) with leaf miners, stem borers, and seed predators being the most frequently

established agents (85-86%) and defoliators being the least frequently established (61%). This

was the only evaluation that yielded a significant difference in establishment rates.

Figure 6. Success rates of CWBC agents by feeding location and guild. Figures, sample

sizes and p-values as above.

Discussion

Release and program success rates

The results obtained by the current analysis show global averages of 71% of agents

establishing, 55% of established agents providing some control, and 24% of established agents

providing a high degree of control. Although the current findings show higher success rates for

agent establishment and control impact when compared to other sources (e.g., both McFadyen

(1998) and Myers and Bazely (2003) cite a 60% general rate for agent establishment and cite

control success rates of established agents at 33%, and 10-35%, respectively). The higher success

rates reported here is likely due to the success bias of the tabulation method utilized and should

not be interpreted as a recent change in success rates of CWBC. While the current analysis has

utilized more recent catalogue data, a change in success rates would be evident in the by-period

analysis through higher success rates in the more recent period, which was not the case. The by-

period analysis shows very similar rates for CWBC agent establishment and impact regardless of

whether or not releases were conducted before or after the median year of 1989.

Host impact rates show a roughly mirror-opposite outcome when compared to agent

impact rates, with a high impact agent becoming established on a target host being the most

common outcome of CWBC programs followed by medium and then low. This makes sense

when considering the way CWBC releases are conducted: since the optimal outcome of a CWBC

release is the establishment of an agent or agents with high control impacts, and since multiple

agents are typically available for release against any given host, the release process can be

repeated until the desired outcome is attained. So, in spite of the fact that roughly 60% of CWBC

agents can be said to ‘fail’ due to either an inability to successfully establish or to have a

meaningful control impact, 70% of CWBC programs against a target host have been successful

in attaining a control impact of some sort. Since from a management perspective the program

outcome is the more relevant concern as opposed to the individual outcome of agent releases, it

is more appropriate to use this metric evaluating overall rates of success in CWBC efforts

worldwide (McFadyen 1998).

By country

Of relevance to the limitations mentioned above is the fact that there is variation in

CWBC success rates when the region in which releases have occurred is taken into

consideration, with the success rates of certain regions such as New Zealand and Canada being

lower than the estimated global averages. This issue highlights one of the difficulties involved at

attempting to arrive at a meaningful overall estimates of how often CWBC agents and programs

are successful, and calls into question the purpose or utility of attempting to derive such figures

as anything other than a crude estimation. Therefore, when seeking to assess rates of success of

CWBC in general it is appropriate and much more useful to control for some broad context such

as the country in which the releases and programs occurred. Country-specific reviews of CWBC

practices such as those conducted by Klein (2011) and Suckling(2013) are more relevant to

consider when attempting to address the question of how often CWBC is successful, as opposed

to the entirely generalized estimates produced by the current analysis.

Understanding why the success of CWBC varies by region may be of interest for further

investigation. Potential explanatory variables might include the presence of favourable target

hosts in certain regions or broadly applicable climactic factors. Regarding the former, there is the

example of the Cactoblastis cactorum moth, which has been used to effectively control a number

of cactus hosts in a large number of regions (Winston et al. 2014), including the three regions

which exhibit the best successes with CWBC in the current analysis (i.e. Australia, USA, and

South Africa).

Regarding regional effects, range restrictions related to colder temperatures with insect

CWBC agents have been frequently hypothesized (McClay and Hughes 1995; Mangan and

Baars 2013; Sun et al. 2017). It may very well be the case that colder regions pose a general

challenge for the establishment and success of insect CWBC agents, although temperature is

only one of many climactic variables which can play a determining role in CWBC agent

successes within their introduced ranges (Sun et al. 2017).

Habitat

The finding of significantly higher success rates in CWBC agents and programs which

target aquatic hosts is consistent with the findings of Paynter et al. (2012) who correlated aquatic

hosts with CWBC success through a quantitative meta-analysis. This successful emulation of a

previously established result gives credibility to the approach and methodology used for this

analysis. In turn, this adds some degree of legitimacy to other significant associations, such as

agent establishment and impacts by feeding guild, that have been established using the current

catalogue based approach.

Order

Analysis of agent establishment by order produced similar results to Boughton and

Pemberton (2008) who took a similar approach using the previous version of the global

catalogue. That is, of the major insects orders investigated, lepidopteran agents established least

frequently, although there were no significant differences in establishment rates across orders.

Extending this approach to also evaluate impact levels by insect order revealed a significant

difference across orders, with coleopteran and hemipteran agents being much more frequently

effective than their lepidopteran and dipteran counterparts. This result is consistent with the

meta-analysis conducted by Clewey et al. (2012), which revealed Coleopteran CWBC agents to

have greater host impacts when compared to other taxa. While this result may be relevant for

agent prioritization considerations, it should be noted that several highly effective agents were

present in all orders. As such, any generalizations regarding potential CWBC agent effectiveness

by order will run the risk of overlooking potentially valuable exceptions (Cullen 1995). In either

case, the current results may warrant further investigation into these correlations. Towards this

end, one possibility is differing levels of parasitism across insect orders (McFadyen and Jacob

2004).

Feeding Guilds/Location

The results of the current analysis suggests that certain guilds may be more advantageous

than others in terms of both successfully establishing within introduced ranges, and effective

control of target hosts. For example, in the current analysis defoliating agents appear to be at a

disadvantage for successful establishment (with a 39% failure rate compared to the 29%

average), and root boring insects appear to be at an advantage for providing a high control

impact on their target hosts (with a 41% rate of high impact compared to the 24% average value),

while agents from other guilds such as flower feeders and leaf miners appear to be generally

ineffective at controlling their target hosts. This may provide some basis for looking at feeding

guilds for agent prioritization in CWBC, although given the cursory nature of the current

analysis, these relationships need to be explored further.

Explanations as to why certain feeding guilds may be correlated to greater CWBC agent

success and efficacy could explore a number of different avenues. Similar to insect order,

parasitism may play a role. This hypothesis was put forward by McFadyen and Jacob (2004),

who, following their own catalogue analysis, noted that there were no reported instances of root

feeding CWBC agents being subject to parasitic attack within their introduced ranges. In terms

of establishment, factors such as acute rainfall events have been implicated in the establishment

failure of CWBC agents due to agents simply being washed off of plants (Norris et al. 2002), so

whether the CWBC agent feeding guilds represent external or internal feeding may play some

role in determining establishment likelihood. In terms of better and more frequent control

efficacy across different guilds, it may be the case that while every target host is different, there

are commonly occurring traits across invasive weed biology (e.g., tolerance of defoliation) that

put certain feeding guilds at a disadvantage for providing effective control. As an example of

this, pale and black swallowwort (Vincetoxicum spp.), two invasive weeds introduced to North

America, have demonstrated extraordinary resilience in the face of heavy and sustained

defoliation treatments, which calls into question the potential efficacy of any defoliating CWBC

agent utilized for their control (Milbrath and Biazzo 2016; Milbrath et al. 2016).

Implications

The significant correlations identified by the current analysis – both those that have been

previously identified (i.e., better CWBC results against aquatic hosts, and when utilizing

Coleopteran agents), and those that are newly discovered (i.e., differing establishment success

and control efficacy across agent feeding guilds) – may provide promising grounds for informing

host and agent prioritization in management decisions involving CWBC. However, any

inclination to rely on these correlations as so-called ‘rules of thumb’ for informing success

predictions in CWBC must be measured against the facts that (1) the current analysis is cursory

in nature and the correlations identified require further study before meaningful conclusions can

drawn, and (2) even if such correlations become better established and defined in the future,

there appears to be important exceptions (e.g. all feeding guilds investigated contain at least one

agent with a high control impact).

This speaks to two important issues in the practice of CWBC; namely, that (1) CWBC

practitioners can scarcely afford to risk overlooking exceptions, as success in this field is entirely

contingent upon the existence of a limited number of suitable CWBC agents (most of which

either fail to establish or fail to have a significant control impact) and (2) context-specific

considerations are often stressed as more important than generalized rules in predicting success

in CWBC (Center et al. 2014).

As such, any generalizations derived from this or similar analyses must not become

overgeneralizations, must provide for exceptions, and must never supersede relevant context

specific information. Certainly, it seems safe to conclude that correlations can be identified

using CWBC catalogue-based analyses, and that correlations identified in the current analysis

(particular those between agent feeding guilds and CWBC outcomes) warrant further

investigation.

In terms of the evaluation of overall success rates in CWBC, management expectations

should be informed by the success rates associated with overall program outcomes and not by

individual agent outcomes, as the former indicates the control impact on the target weed, which

is primary management objective of CWBC. Furthermore, since rates of success in program

outcomes vary by region, it may also be appropriate to calibrate management expectations to the

success rates that have occurred in that region. In any case, while program outcomes represent

the higher success rates (70% with a control impact of some sort), and while the most common

program outcome is a high control impact (40%), there is a substantial percentage (30%) which

either fail to attain significant control impacts or fail to attain successful agent establishment

altogether. When this observation is coupled with the fact that under the current analysis,

program success rates have not improved over time, it is difficult to avoid the conclusion that

success is highly uncertain in CWBC and that such uncertainty may be an inherent feature of

CWBC practice. Management expectations should reflect this. However, despite this uncertainty,

it should be understood that while most agents fail, most groups of agents targeting a specific

host eventually succeed, and that while some CWBC programs fail entirely, highly successful

CWBC programs are common, and most programs succeed in attaining a significant control

outcome.

Conclusions

In conclusion, several useful insights have been produced by this study: (1) meaningful

correlations between CWBC outcomes and host/agent variables can be discovered through post-

hoc catalogue analysis. As the catalogue data continues to grow over time, the potential for

discovering new correlations can only increase; (2) evidence has been found to suggest a

correlation between CWBC outcomes across feeding guilds, and these potential relationships

should be investigated further; and (3) despite using the most recently available catalogue data,

success rates in CWBC appear stagnant, and still entail a significant amount of uncertainty and

failure. This information could be used for managing expectations of success in CWBC

practitioners, but it should also be used to stress the importance of figuring out new ways to

improve the efficacy of this important management tool.

Acknowledgements

I would like to thank the University of Toronto Faculty of Forestry for their continuous

support throughout the completion of this capstone. In particular, Dr. Jay Malcolm, for his

ongoing guidance, encouragement, and technical assistance with SAS and statistics, and Lukas

Seehausen and Dr. Sandy Smith, for contributing their knowledge of entomology and biological

control practices.

LITERATURE CITED

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Austromusotima camptozonale, against Lygodium microphyllum in Florida, considered in

the light of a retrospective review of establishment success of weed biocontrol agents

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APPENDIX 1. CWBC AGENT FEEDING GUILDS AND LOCATIONS

SPECIES ORDER FAMILY FEEDING GUILD FEEDING LOCATION

ACERIA CHONDRILLAE ACARI ERIOPHYIDAE BUD GALLER BUD

ACERIA GENISTAE ACARI ERIOPHYIDAE BUD GALLER BUD

ACERIA LANTANAE ACARI ERIOPHYIDAE LEAF GALLER LEAF

ACERIA MALHERBAE ACARI ERIOPHYIDAE LEAF GALLER LEAF

CECIDOPHYES ROUHOLLAHI ACARI ERIOPHYIDAE LEAF GALLER LEAF

FLORACARUS PERREPAE ACARI ERIOPHYIDAE LEAF GALLER LEAF

ACERIA SP. ACARI ERIOPHYIDAE UNKNOWN UNKNOWN

ACULUS HYPERICI ACARI ERIOPHYIDAE UNKNOWN UNKNOWN

TETRANYCHUS LINTEARIUS ACARI TETRANYCHIDAE UNKNOWN UNKNOWN

APION SP. A COLEOPTERA BRENTIDAE FLOWER FEEDER FLOWER

COELOCEPHALAPION ACULEATUM COLEOPTERA BRENTIDAE FLOWER FEEDER FLOWER

COELOCEPHALAPION PIGRAE COLEOPTERA BRENTIDAE FLOWER FEEDER FLOWER

COELOCEPHALAPION CAMARAE COLEOPTERA BRENTIDAE PETIOLE GALLER PETIOLE

APION FRUMENTARIUM COLEOPTERA BRENTIDAE SEED FEEDER SEED

APION SP. B COLEOPTERA BRENTIDAE SEED FEEDER SEED

EXAPION FUSCIROSTRE COLEOPTERA BRENTIDAE SEED FEEDER SEED

EXAPION ULICIS COLEOPTERA BRENTIDAE SEED FEEDER SEED

OMPHALAPION HOOKERORUM COLEOPTERA BRENTIDAE SEED FEEDER SEED

STENOPTERAPION SCUTELLARE COLEOPTERA BRENTIDAE SEED GALLER SEED

APION SP. COLEOPTERA BRENTIDAE UNKNOWN UNKNOWN

CERATAPION ONOPORDI COLEOPTERA BRENTIDAE UNKNOWN UNKNOWN

PERAPION ANTIQUUM COLEOPTERA BRENTIDAE UNKNOWN UNKNOWN

PERAPION NEOFALLAX COLEOPTERA BRENTIDAE UNKNOWN UNKNOWN

PERAPION VIOLACEUM COLEOPTERA BRENTIDAE UNKNOWN UNKNOWN

HYLAEOGENA JURECEKI COLEOPTERA BUPRESTIDAE DEFOLIATOR LEAF

LIUS POSEIDON COLEOPTERA BUPRESTIDAE LEAF MINER LEAF

AGRILUS HYPERICI COLEOPTERA BUPRESTIDAE ROOT BORER ROOT

SPHENOPTERA JUGOSLAVICA COLEOPTERA BUPRESTIDAE ROOT GALLER ROOT

MONEILEMA ARMATUM COLEOPTERA CERAMBYCIDAE CLADODE AND ROOT BORER MULTIPLE

MONEILEMA BLAPSIDES COLEOPTERA CERAMBYCIDAE CLADODE AND ROOT BORER MULTIPLE

MONEILEMA VARIOLARE COLEOPTERA CERAMBYCIDAE CLADODE AND ROOT BORER MULTIPLE

PAREVANDER XANTHOMELAS COLEOPTERA CERAMBYCIDAE ROOT FEEDER ROOT

PLAGIOHAMMUS SPINIPENNIS COLEOPTERA CERAMBYCIDAE MULTIPLE MULTIPLE

OBEREA ERYTHROCEPHALA COLEOPTERA CERAMBYCIDAE MULTIPLE MULTIPLE

AERENICOPSIS CHAMPIONI COLEOPTERA CERAMBYCIDAE STEM BORER STEM

APHANASIUM AUSTRALE COLEOPTERA CERAMBYCIDAE STEM BORER STEM

LAGOCHEIRUS FUNESTUS COLEOPTERA CERAMBYCIDAE STEM BORER STEM

MECAS CANA SUBSP. SATURNINA COLEOPTERA CERAMBYCIDAE STEM BORER STEM

MEGACYLLENE MELLYI COLEOPTERA CERAMBYCIDAE STEM BORER STEM

NEALCIDION CEREICOLA COLEOPTERA CERAMBYCIDAE STEM BORER STEM

NUPSERHA VEXATOR COLEOPTERA CERAMBYCIDAE STEM BORER STEM

OPSILIA COERULESCENS COLEOPTERA CERAMBYCIDAE STEM BORER STEM

AGASICLES HYGROPHILA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

ALAGOASA PARANA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

ALTICA CARDUORUM COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

ANACASSIS FUSCATA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

ANACASSIS PHAEOPODA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CALLIGRAPHA PANTHERINA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CASSIDA AZUREA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CASSIDA RUBIGINOSA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CASSIDA SP. 3 COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHARIDOTIS AUROGUTTATA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHARIDOTIS PYGMAEA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHLAMISUS GIBBOSA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHLAMISUS MIMOSAE COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHRYSOLINA BRUNSVICENSIS COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHRYSOLINA FASCIATA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHRYSOLINA HYPERICI COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHRYSOLINA QUADRIGEMINA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHRYSOLINA SCOTTI COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHRYSOLINA SP. B COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CHRYSOMELA VARIANS COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

CRIOCERIS SP. COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

DEUTEROCAMPTA QUADRIJUGA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

DIORHABDA CARINATA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

DIORHABDA CARINULATA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

DIORHABDA ELONGATA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

DIORHABDA SUBLINEATA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

DISONYCHA ARGENTINENSIS COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

GALERUCELLA CALMARIENSIS COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

GALERUCELLA PUSILLA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

GONIOCTENA OLIVACEA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

GRATIANA BOLIVIANA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

GRATIANA SPADICEA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

HOMICHLODA BARKERI COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

LEMA CYANELLA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

LEPTINOTARSA DEFECTA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

LEPTINOTARSA TEXANA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

LILIOCERIS CHENI COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

LIOPLACIS ELLIPTICA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

LOCHMAEA SUTURALIS COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

LYSATHIA SP. COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

METALLACTUS NIGROFASCIATUS COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

METALLACTUS PATAGONICUS COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

NEOLEMA OGLOBLINI COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

PHENRICA GUERINI COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

PLECTONYCHA CORRENTINA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

TRIRHABDA BACHARIDIS COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

ZYGOGRAMMA BICOLORATA COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

ZYGOGRAMMA SUTURALIS COLEOPTERA CHRYSOMELIDAE DEFOLIATOR LEAF

LEMA BASICOSTATA COLEOPTERA CHRYSOMELIDAE MULTIPLE MULTIPLE

OCTOTOMA CHAMPIONI COLEOPTERA CHRYSOMELIDAE LEAF MINER LEAF

OCTOTOMA GUNDLACHI COLEOPTERA CHRYSOMELIDAE LEAF MINER LEAF

OCTOTOMA SCABRIPENNIS COLEOPTERA CHRYSOMELIDAE LEAF MINER LEAF

UROPLATA FULVOPUSTULATA COLEOPTERA CHRYSOMELIDAE LEAF MINER LEAF

UROPLATA GIRARDI COLEOPTERA CHRYSOMELIDAE LEAF MINER LEAF

UROPLATA LANTANAE COLEOPTERA CHRYSOMELIDAE LEAF MINER LEAF

LONGITARSUS ECHII COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

APHTHONA ABDOMINALIS COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

APHTHONA CYPARISSIAE COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

APHTHONA CZWALINAI COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

APHTHONA FLAVA COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

APHTHONA LACERTOSA COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

APHTHONA NIGRISCUTIS COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

LONGITARSUS FLAVICORNIS COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

LONGITARSUS JACOBAEAE COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

LONGITARSUS QUADRIGUTTATUS COLEOPTERA CHRYSOMELIDAE ROOT BORER ROOT

LONGITARSUS BETHAE COLEOPTERA CHRYSOMELIDAE ROOT FEEDER ROOT

ACANTHOSCELIDES MACROPHTHALMUS COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

ACANTHOSCELIDES PUNICEUS COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

ACANTHOSCELIDES QUADRIDENTATUS COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

ALGAROBIUS BOTTIMERI COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

ALGAROBIUS PROSOPIS COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

BRUCHIDIUS SAHLBERGI COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

BRUCHIDIUS VILLOSUS COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

LITHRAEUS ATRONOTATUS COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

MIMOSESTES ULKEI COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

PENTHOBRUCHUS GERMAINI COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

SULCOBRUCHUS SUBSUTURALIS COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

NELTUMIUS ARIZONENSIS COLEOPTERA CHRYSOMELIDAE SEED FEEDER SEED

NEOLEMA ABBREVIATA COLEOPTERA CHRYSOMELIDAE STEM BORER STEM

LONGITARSUS AENEUS COLEOPTERA CHRYSOMELIDAE UNKNOWN UNKNOWN

LONGITARSUS ALBINEUS COLEOPTERA CHRYSOMELIDAE UNKNOWN UNKNOWN

MALACORHINUS IRREGULARIS COLEOPTERA CHRYSOMELIDAE UNKNOWN UNKNOWN

NESAECREPIDA INFUSCATA COLEOPTERA CHRYSOMELIDAE UNKNOWN UNKNOWN

PSYLLIODES CHALCOMERA COLEOPTERA CHRYSOMELIDAE UNKNOWN UNKNOWN

MADA POLLUTA COLEOPTERA COCCINELLIDAE DEFOLIATOR LEAF

ERIOCEREOPHAGA HUMERIDENS COLEOPTERA CURCULIONIDAE CACTUS FEEDER CLADODE

CLEOPUS JAPONICUS COLEOPTERA CURCULIONIDAE DEFOLIATOR LEAF

EUTINOBOTHRUS PILOSELLUS COLEOPTERA CURCULIONIDAE DEFOLIATOR LEAF

PHRYDIUCHUS SPILMANI COLEOPTERA CURCULIONIDAE DEFOLIATOR LEAF

PHRYDIUCHUS TAU COLEOPTERA CURCULIONIDAE DEFOLIATOR LEAF

RHINONCOMIMUS LATIPES COLEOPTERA CURCULIONIDAE DEFOLIATOR LEAF

ANTHONOMUS SANTACRUZI COLEOPTERA CURCULIONIDAE FLOWER FEEDER FLOWER

DICOMADA RUFA COLEOPTERA CURCULIONIDAE FLOWER FEEDER FLOWER

LARINUS CARLINAE COLEOPTERA CURCULIONIDAE FLOWER FEEDER FLOWER

LARINUS LATUS COLEOPTERA CURCULIONIDAE FLOWER FEEDER FLOWER

ACYTHOPEUS BURKHARTORUM COLEOPTERA CURCULIONIDAE GALL INDUCER UNKNOWN

NEOHYDRONOMUS AFFINIS COLEOPTERA CURCULIONIDAE MULTIPLE MULTIPLE

ACYTHOPEUS COCCINIAE COLEOPTERA CURCULIONIDAE LEAF MINER LEAF

SIBINIA FASTIGIATA COLEOPTERA CURCULIONIDAE MULTIPLE MULTIPLE

EUSTENOPUS VILLOSUS COLEOPTERA CURCULIONIDAE MULTIPLE MULTIPLE

RHINOCYLLUS CONICUS COLEOPTERA CURCULIONIDAE MULTIPLE RECEPTACLE

HEILIPODUS VENTRALIS COLEOPTERA CURCULIONIDAE ROOT BORER ROOT

CYPHOCLEONUS ACHATES COLEOPTERA CURCULIONIDAE ROOT BORER ROOT

HYLOBIUS TRANSVERSOVITTATUS COLEOPTERA CURCULIONIDAE ROOT BORER ROOT

MOGULONES CRUCIFER COLEOPTERA CURCULIONIDAE ROOT BORER ROOT

TRICHOSIROCALUS MORTADELO COLEOPTERA CURCULIONIDAE ROOT FEEDER ROOT

MOGULONES GEOGRAPHICUS COLEOPTERA CURCULIONIDAE ROOT FEEDER ROOT

MOGULONES LARVATUS COLEOPTERA CURCULIONIDAE ROOT FEEDER ROOT

RHINUSA LINARIAE COLEOPTERA CURCULIONIDAE ROOT GALLER ROOT

TRICHOSIROCALUS BRIESEI COLEOPTERA CURCULIONIDAE ROSETTE FEEDER ROSETTE

TRICHOSIROCALUS HORRIDUS COLEOPTERA CURCULIONIDAE ROSETTE FEEDER ROSETTE

BANGASTERNUS FAUSTI COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

BANGASTERNUS ORIENTALIS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

CHALCODERMUS SERRIPES COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

CISSOANTHONOMUS TUBERCULIPENNIS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

ERYTENNA CONSPUTA COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

LARINUS CURTUS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

LARINUS MINUTUS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

LARINUS OBTUSUS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

MELANTERIUS ACACIAE COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

MELANTERIUS COMPACTUS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

MELANTERIUS MACULATUS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

MELANTERIUS SERVULUS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

MELANTERIUS VENTRALIS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

MICROLARINUS LAREYNII COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

RHINUSA ANTIRRHINI COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

RHYSSOMATUS MARGINATUS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

SMICRONYX LUTULENTUS COLEOPTERA CURCULIONIDAE SEED FEEDER SEED

CYDMAEA BINOTATA COLEOPTERA CURCULIONIDAE SHOOT BORER SHOOT

OXYOPS VITIOSA COLEOPTERA CURCULIONIDAE SHOOT FEEDER SHOOT

RHINUSA PILOSA COLEOPTERA CURCULIONIDAE SHOOT GALLER SHOOT

HADROPLONTUS LITURA COLEOPTERA CURCULIONIDAE MULTIPLE MULTIPLE

HEILIPODUS INTRICATUS COLEOPTERA CURCULIONIDAE STEM BORER STEM

LISTRONOTUS SETOSIPENNIS COLEOPTERA CURCULIONIDAE STEM BORER STEM

LIXUS AEMULUS COLEOPTERA CURCULIONIDAE STEM BORER STEM

LIXUS CARDUI COLEOPTERA CURCULIONIDAE STEM BORER STEM

LIXUS LINEARIS COLEOPTERA CURCULIONIDAE STEM BORER STEM

MICROLARINUS LYPRIFORMIS COLEOPTERA CURCULIONIDAE STEM BORER STEM

NEODIPLOGRAMMUS QUADRIVITTATUS COLEOPTERA CURCULIONIDAE STEM BORER STEM

MECINUS JANTHINIFORMIS COLEOPTERA CURCULIONIDAE STEM BORER STEM

MECINUS JANTHINUS COLEOPTERA CURCULIONIDAE STEM BORER STEM

MICROPLONTUS EDENTULUS COLEOPTERA CURCULIONIDAE STEM BORER STEM

EUTINOBOTHRUS SP COLEOPTERA CURCULIONIDAE STEM FEEDER STEM

CONOTRACHELUS ALBOCINEREUS COLEOPTERA CURCULIONIDAE STEM GALLER STEM

BAGOUS HYDRILLAE COLEOPTERA CURCULIONIDAE MULTIPLE MULTIPLE

ATHESAPEUTA CYPERI COLEOPTERA CURCULIONIDAE TUBER FEEDER TUBER

BAGOUS AFFINIS COLEOPTERA CURCULIONIDAE TUBER FEEDER TUBER

METAMASIUS SPINOLAE COLEOPTERA DRYOPHTHORIDAE STEM BORER STEM

STENOPELMUS RUFINASUS COLEOPTERA ERIRHINIDAE DEFOLIATOR LEAF

CYRTOBAGOUS SALVINIAE COLEOPTERA ERIRHINIDAE RHIZOME FEEDER RHIZOME

NEOCHETINA BRUCHI COLEOPTERA ERIRHINIDAE UNKNOWN UNKNOWN

NEOCHETINA EICHHORNIAE COLEOPTERA ERIRHINIDAE UNKNOWN UNKNOWN

BRACHYPTEROLUS PULICARIUS COLEOPTERA KATERIDAE FLOWER FEEDER FLOWER

NANOPHYES MARMORATUS COLEOPTERA NANOPHYIDAE FLOWER FEEDER FLOWER

MELIGETHES PLANIUSCULUS COLEOPTERA NITIDULIDAE FLOWER FEEDER FLOWER

CALYCOMYZA EUPATORIVORA DIPTERA AGROMYZIDAE LEAF MINER LEAF

CALYCOMYZA LANTANAE DIPTERA AGROMYZIDAE LEAF MINER LEAF

LIRIOMYZA SONCHI DIPTERA AGROMYZIDAE LEAF MINER LEAF

OPHIOMYIA CAMARAE DIPTERA AGROMYZIDAE LEAF MINER LEAF

PHYTOMYZA VITALBAE DIPTERA AGROMYZIDAE LEAF MINER LEAF

OPHIOMYIA LANTANAE DIPTERA AGROMYZIDAE SEED FEEDER SEED

BOTANOPHILA SENECIELLA DIPTERA ANTHOMYIIDAE MULTIPLE MULTIPLE

BOTANOPHILA SPINOSA DIPTERA ANTHOMYIIDAE ROSETTE FEEDER ROSETTE

BOTANOPHILA JACOBAEAE DIPTERA ANTHOMYIIDAE SEED FEEDER SEED

PEGOMYA CURTICORNIS DIPTERA ANTHOMYIIDAE STEM GALLER STEM

PEGOMYA EUPHORBIAE DIPTERA ANTHOMYIIDAE STEM GALLER STEM

DASINEURA STROBILA DIPTERA CECIDOMYIIDAE BUD GALLER BUD

RHOPALOMYIA CALIFORNICA DIPTERA CECIDOMYIIDAE BUD GALLER BUD

RHOPALOMYIA TRIPLEUROSPERMI DIPTERA CECIDOMYIIDAE BUD GALLER BUD

DASINEURA DIELSI DIPTERA CECIDOMYIIDAE FLOWER GALLER FLOWER

CYSTIPHORA SONCHI DIPTERA CECIDOMYIIDAE LEAF GALLER LEAF

ZEUXIDIPLOSIS GIARDI DIPTERA CECIDOMYIIDAE LEAF GALLER LEAF

MACROLABIS PILOSELLAE DIPTERA CECIDOMYIIDAE MULTIPLE MULTIPLE

JAAPIELLA IVANNIKOVI DIPTERA CECIDOMYIIDAE SHOOT GALLER SHOOT

SPURGIA CAPITIGENA DIPTERA CECIDOMYIIDAE SHOOT TIP GALLER SHOOT

SPURGIA ESULAE DIPTERA CECIDOMYIIDAE SHOOT TIP GALLER SHOOT

CYSTIPHORA SCHMIDTI DIPTERA CECIDOMYIIDAE MULTIPLE MULTIPLE

LOPHODIPLOSIS TRIFIDA DIPTERA CECIDOMYIIDAE STEM GALLER STEM

HYDRELLIA BALCIUNASI DIPTERA EPHYDRIDAE LEAF MINER LEAF

HYDRELLIA PAKISTANAE DIPTERA EPHYDRIDAE LEAF MINER LEAF

FERGUSONINA TURNERI DIPTERA FERGUSONINIDAE BUD GALLER BUD

CHEILOSIA URBANA DIPTERA SYRPHIDAE ROOT FEEDER ROOT

CHEILOSIA GROSSA DIPTERA SYRPHIDAE MULTIPLE MULTIPLE

CHEILOSIA PSILOPHTHALMA DIPTERA SYRPHIDAE UNKNOWN UNKNOWN

TEPHRITIS DILACERATA DIPTERA TEPHRITIDAE BUD GALLER BUD

UROPHORA JACULATA DIPTERA TEPHRITIDAE CAPITULUM GALLER FLOWER

UROPHORA SIRUNASEVA DIPTERA TEPHRITIDAE CAPITULUM GALLER FLOWER

UROPHORA TEREBRANS DIPTERA TEPHRITIDAE CAPITULUM GALLER FLOWER

ACINIA PICTURATA DIPTERA TEPHRITIDAE FLOWER FEEDER FLOWER

TEPHRITIS POSTICA DIPTERA TEPHRITIDAE FLOWER FEEDER FLOWER

UROPHORA QUADRIFASCIATA DIPTERA TEPHRITIDAE OVARY GALLER FLOWER

UROPHORA AFFINIS DIPTERA TEPHRITIDAE RECEPTACLE GALLER RECEPTACLE

UROPHORA SOLSTITIALIS DIPTERA TEPHRITIDAE RECEPTACLE GALLER RECEPTACLE

UROPHORA STYLATA DIPTERA TEPHRITIDAE RECEPTACLE GALLER RECEPTACLE

CHAETORELLIA ACROLOPHI DIPTERA TEPHRITIDAE SEED FEEDER SEED

CHAETORELLIA AUSTRALIS DIPTERA TEPHRITIDAE SEED FEEDER SEED

EUARESTA AEQUALIS DIPTERA TEPHRITIDAE SEED FEEDER SEED

MESOCLANIS MAGNIPALPIS DIPTERA TEPHRITIDAE SEED FEEDER SEED

MESOCLANIS POLANA DIPTERA TEPHRITIDAE SEED FEEDER SEED

TERELLIA VIRENS DIPTERA TEPHRITIDAE SEED FEEDER SEED

TETRAEUARESTA OBSCURIVENTRIS DIPTERA TEPHRITIDAE SEED FEEDER SEED

XANTHACIURA CONNEXIONIS DIPTERA TEPHRITIDAE SEED FEEDER SEED

EUTRETA XANTHOCHAETA DIPTERA TEPHRITIDAE STEM GALLER STEM

PROCECIDOCHARES ALANI DIPTERA TEPHRITIDAE STEM GALLER STEM

PROCECIDOCHARES UTILIS DIPTERA TEPHRITIDAE STEM GALLER STEM

UROPHORA CARDUI DIPTERA TEPHRITIDAE STEM GALLER STEM

APHIS CHLORIS HEMIPTERA APHIDIDAE MULTIPLE MULTIPLE

TRIBE ERYTHRONEURINI HEMIPTERA CICADELLIDAE DEFOLIATOR LEAF

SCAMURIUS SP. HEMIPTERA COREIDAE LEAF TIPS LEAF

CHELINIDEA TABULATA HEMIPTERA COREIDAE UNKNOWN UNKNOWN

CHELINIDEA VITTIGER HEMIPTERA COREIDAE UNKNOWN UNKNOWN

CATORHINTHA SCHAFFNERI HEMIPTERA COREIDAE STEM WILTER STEM

DACTYLOPIUS AUSTRINUS HEMIPTERA DACTYLOPIIDAE CLADODE SUCKER CLADODE

DACTYLOPIUS CEYLONICUS HEMIPTERA DACTYLOPIIDAE CLADODE SUCKER CLADODE

DACTYLOPIUS COCCUS HEMIPTERA DACTYLOPIIDAE CLADODE SUCKER CLADODE

DACTYLOPIUS CONFUSUS HEMIPTERA DACTYLOPIIDAE CLADODE SUCKER CLADODE

DACTYLOPIUS OPUNTIAE HEMIPTERA DACTYLOPIIDAE CLADODE SUCKER CLADODE

DACTYLOPIUS TOMENTOSUS HEMIPTERA DACTYLOPIIDAE CLADODE SUCKER CLADODE

STOBAERA CONCINNA HEMIPTERA DELPHACIDAE STEM FEEDER STEM

MEGAMELUS SCUTELLARIS HEMIPTERA DELPHACIDAE SAP SUCKER

RHIZASPIDIOTUS DONACIS HEMIPTERA DIASPIDIDAE STEM FEEDER STEM

ACONOPHORA COMPRESSA HEMIPTERA MEMBRACIDAE STEM SUCKER STEM

ECCRITOTARSUS CATARINENSIS HEMIPTERA MIRIDAE SAP SUCKER

FALCONIA INTERMEDIA HEMIPTERA MIRIDAE SAP SUCKER

RHINACLOA CALLICRATES HEMIPTERA MIRIDAE UNKNOWN UNKNOWN

ORTHEZIA INSIGNIS HEMIPTERA ORTHEZIIDAE SAP SUCKER

HYPOGEOCOCCUS FESTERIANUS HEMIPTERA PSEUDOCOCCIDAE STEM SUCKER STEM

ARYTAINILLA SPARTIOPHILA HEMIPTERA PSYLLIDAE DEFOLIATOR LEAF

HETEROPSYLLA SPINULOSA HEMIPTERA PSYLLIDAE DEFOLIATOR LEAF

BOREIOGLYCASPIS MELALEUCAE HEMIPTERA PSYLLIDAE SAP SUCKER

PROSOPIDOPSYLLA FLAVA HEMIPTERA PSYLLIDAE SAP SUCKER

APHALARA ITADORI HEMIPTERA PSYLLIDAE UNKNOWN UNKNOWN

AGONOSOMA TRILINEATUM HEMIPTERA SCUTELLERIDAE SEED FEEDER SEED

CARVALHOTINGIS HOLLANDI HEMIPTERA TINGIDAE LEAF SUCKER LEAF

CARVALHOTINGIS VISENDA HEMIPTERA TINGIDAE LEAF SUCKER LEAF

GARGAPHIA DECORIS HEMIPTERA TINGIDAE LEAF SUCKER LEAF

TELEONEMIA ELATA HEMIPTERA TINGIDAE LEAF SUCKER LEAF

TELEONEMIA HARLEYI HEMIPTERA TINGIDAE LEAF SUCKER LEAF

TELEONEMIA PROLIXA HEMIPTERA TINGIDAE LEAF SUCKER LEAF

TELEONEMIA SCRUPULOSA HEMIPTERA TINGIDAE LEAF SUCKER LEAF

LEPTOBYRSA DECORA HEMIPTERA TINGIDAE LEAF SUCKER LEAF

AULACIDEA ACROPTILONICA HYMENOPTERA CYNIPIDAE MULTIPLE MULTIPLE

AULACIDEA SUBTERMINALIS HYMENOPTERA CYNIPIDAE STEM GALLER STEM

TETRAMESA ROMANA HYMENOPTERA EURYTOMIDAE SHOOT GALLER SHOOT

TRICHILOGASTER ACACIAELONGIFOLIAE HYMENOPTERA PTEROMALIDAE BUD GALLER BUD

TRICHILOGASTER SIGNIVENTRIS HYMENOPTERA PTEROMALIDAE BUD GALLER BUD

MONOPHADNUS SPINOLAE HYMENOPTERA TENTHREDINIDAE DEFOLIATOR LEAF

PRIOPHORUS MORIO HYMENOPTERA TENTHREDINIDAE DEFOLIATOR LEAF

UCONA ACAENAE HYMENOPTERA TENTHREDINIDAE DEFOLIATOR LEAF

BUCCULATRIX IVELLA LEPIDOPTERA BUCCULATRICIDAE LEAF MINER LEAF

BUCCULATRIX PARTHENICA LEPIDOPTERA BUCCULATRICIDAE LEAF MINER LEAF

CARPOSINA BULLATA LEPIDOPTERA CARPOSINIDAE FLOWER FEEDER FLOWER

CARPOSINA AUTOLOGA LEPIDOPTERA CARPOSINIDAE SEED FEEDER SEED

COLEOPHORA KLIMESCHIELLA LEPIDOPTERA COLEOPHORIDAE LEAF MINER LEAF

COLEOPHORA PARTHENICA LEPIDOPTERA COLEOPHORIDAE STEM BORER STEM

ETEOBALEA INTERMEDIELLA LEPIDOPTERA COSMOPTERIGIDAE ROOT BORER ROOT

ETEOBALEA SERRATELLA LEPIDOPTERA COSMOPTERIGIDAE ROOT BORER ROOT

LOXOMORPHA FLAVIDISSIMALIS LEPIDOPTERA CRAMBIDAE CACTUS FEEDER CLADODE

MIMORISTA PULCHELLALIS LEPIDOPTERA CRAMBIDAE CLADODE BORER CLADODE

AUSTROMUSOTIMA CAMPTOZONALE LEPIDOPTERA CRAMBIDAE DEFOLIATOR LEAF

EUCLASTA WHALLEYI LEPIDOPTERA CRAMBIDAE DEFOLIATOR LEAF

NEOMUSOTIMA CONSPURCATALIS LEPIDOPTERA CRAMBIDAE DEFOLIATOR LEAF

PSEUDOPYRAUSTA SANTATALIS LEPIDOPTERA CRAMBIDAE DEFOLIATOR LEAF

SALBIA HAEMORRHOIDALIS LEPIDOPTERA CRAMBIDAE DEFOLIATOR LEAF

SAMEA MULTIPLICALIS LEPIDOPTERA CRAMBIDAE DEFOLIATOR LEAF

PYRAUSTA PERELEGANS LEPIDOPTERA CRAMBIDAE FLOWER FEEDER FLOWER

ATEGUMIA ADIPALIS LEPIDOPTERA CRAMBIDAE LEAF ROLLER LEAF

ATEGUMIA FATUALIS LEPIDOPTERA CRAMBIDAE LEAF ROLLER LEAF

ATEGUMIA MATUTINALIS LEPIDOPTERA CRAMBIDAE LEAF ROLLER LEAF

XUBIDA INFUSELLA LEPIDOPTERA CRAMBIDAE PETIOLE MINER PETIOLE

NIPHOGRAPTA ALBIGUTTALIS LEPIDOPTERA CRAMBIDAE STEM BORER STEM

ANTIBLEMMA ACCLINALIS LEPIDOPTERA EREBIDAE DEFOLIATOR LEAF

COMETASTER PYRULA LEPIDOPTERA EREBIDAE DEFOLIATOR LEAF

HYPENA LACERATALIS LEPIDOPTERA EREBIDAE DEFOLIATOR LEAF

HYPENA OPULENTA LEPIDOPTERA EREBIDAE DEFOLIATOR LEAF

PAREUCHAETES AURATA AURATA LEPIDOPTERA EREBIDAE DEFOLIATOR LEAF

PAREUCHAETES INSULATA LEPIDOPTERA EREBIDAE DEFOLIATOR LEAF

PAREUCHAETES PSEUDOINSULATA LEPIDOPTERA EREBIDAE DEFOLIATOR LEAF

SECUSIO EXTENSA LEPIDOPTERA EREBIDAE DEFOLIATOR LEAF

TYRIA JACOBAEAE LEPIDOPTERA EREBIDAE DEFOLIATOR LEAF

EUBLEMMA AMOENA LEPIDOPTERA EREBIDAE PETIOLE MINER PETIOLE

ARISTOTELIA IVAE LEPIDOPTERA GELECHIIDAE DEFOLIATOR LEAF

FRUMENTA NEPHELOMICTA LEPIDOPTERA GELECHIIDAE FRUIT GALLER FRUIT

FRUMENTA SP. LEPIDOPTERA GELECHIIDAE FRUIT GALLER FRUIT

DICHOMERIS AENIGMATICA LEPIDOPTERA GELECHIIDAE LEAF ROLLER LEAF

EVIPPE SP. #1 LEPIDOPTERA GELECHIIDAE LEAF ROLLER LEAF

METZNERIA PAUCIPUNCTELLA LEPIDOPTERA GELECHIIDAE MULTIPLE MULTIPLE

CRASIMORPHA INFUSCATA LEPIDOPTERA GELECHIIDAE STEM GALLER STEM

APLOCERA PLAGIATA LEPIDOPTERA GEOMETRIDAE DEFOLIATOR LEAF

CHIASMIA ASSIMILIS LEPIDOPTERA GEOMETRIDAE DEFOLIATOR LEAF

CHIASMIA INCONSPICUA LEPIDOPTERA GEOMETRIDAE DEFOLIATOR LEAF

LEUCIRIS FIMBRIARIA LEPIDOPTERA GEOMETRIDAE DEFOLIATOR LEAF

MACARIA PALLIDATA LEPIDOPTERA GEOMETRIDAE DEFOLIATOR LEAF

MINOA MURINATA LEPIDOPTERA GEOMETRIDAE DEFOLIATOR LEAF

COMOSTOLOPSIS GERMANA LEPIDOPTERA GEOMETRIDAE SHOOT TIP FEEDER SHOOT

APLOCERA EFFORMATA LEPIDOPTERA GEOMETRIDAE UNKNOWN UNKNOWN

EUEUPITHECIA CISPLATENSIS LEPIDOPTERA GEOMETRIDAE DEFOLIATOR LEAF

ARISTAEA THALASSIAS LEPIDOPTERA GRACILLARIIDAE LEAF MINER LEAF

CALOPTILIA CORUSCANS LEPIDOPTERA GRACILLARIIDAE LEAF MINER LEAF

CREMASTOBOMBYCIA LANTANELLA LEPIDOPTERA GRACILLARIIDAE LEAF MINER LEAF

DIALECTICA SCALARIELLA LEPIDOPTERA GRACILLARIIDAE LEAF MINER LEAF

CUPHODES PROFLUENS LEPIDOPTERA GRACILLARIIDAE SHOOT BORER SHOOT

NEUROSTROTA GUNNIELLA LEPIDOPTERA GRACILLARIIDAE STEM BORER STEM

HEPIALUS SP. LEPIDOPTERA HEPIALIDAE UNKNOWN UNKNOWN

STRYMON BAZOCHII LEPIDOPTERA LYCAENIDAE FLOWER FEEDER FLOWER

TMOLUS ECHION LEPIDOPTERA LYCAENIDAE FLOWER FEEDER FLOWER

LEUCOPTERA SPARTIFOLIELLA LEPIDOPTERA LYONETIIDAE STEM BORER STEM

MOMPHA TRITHALAMA LEPIDOPTERA MOMPHIDAE SEED FEEDER SEED

ACTINOTIA HYPERICI LEPIDOPTERA NOCTUIDAE DEFOLIATOR LEAF

AUTOPLUSIA ILLUSTRATA LEPIDOPTERA NOCTUIDAE DEFOLIATOR LEAF

CALOPHASIA LUNULA LEPIDOPTERA NOCTUIDAE DEFOLIATOR LEAF

DIASTEMA TIGRIS LEPIDOPTERA NOCTUIDAE DEFOLIATOR LEAF

NEOGALEA SUNIA LEPIDOPTERA NOCTUIDAE DEFOLIATOR LEAF

SPODOPTERA PECTINICORNIS LEPIDOPTERA NOCTUIDAE DEFOLIATOR LEAF

TYTA LUCTUOSA LEPIDOPTERA NOCTUIDAE DEFOLIATOR LEAF

RHYNCHOPALPUS BRUNELLUS LEPIDOPTERA NOLIDAE UNKNOWN UNKNOWN

SCEA NECYRIA LEPIDOPTERA NOTODONTIDAE DEFOLIATOR LEAF

AGONOPTERIX ASSIMILELLA LEPIDOPTERA OECOPHORIDAE DEFOLIATOR LEAF

AGONOPTERIX UMBELLANA LEPIDOPTERA OECOPHORIDAE DEFOLIATOR LEAF

ECTAGA GARCIA LEPIDOPTERA OECOPHORIDAE DEFOLIATOR LEAF

PTEROLONCHE INSPERSA LEPIDOPTERA PTEROLONCHIDAE ROOT BORER ROOT

OXYPTILUS PILOSELLAE LEPIDOPTERA PTEROPHORIDAE MULTIPLE MULTIPLE

LANTANOPHAGA PUSILLIDACTYLA LEPIDOPTERA PTEROPHORIDAE FLOWER FEEDER FLOWER

WHEELERIA SPILODACTYLUS LEPIDOPTERA PTEROPHORIDAE MULTIPLE MULTIPLE

PLATYPTILIA ISODACTYLA LEPIDOPTERA PTEROPHORIDAE MULTIPLE MULTIPLE

HELLINSIA BALANOTES LEPIDOPTERA PTEROPHORIDAE STEM BORER STEM

OIDAEMATOPHORUS BENEFICUS LEPIDOPTERA PTEROPHORIDAE UNKNOWN UNKNOWN

MELITARA DENTATA LEPIDOPTERA PYRALIDAE CACTUS FEEDER CLADODE

CACTOBLASTIS CACTORUM LEPIDOPTERA PYRALIDAE CLADODE BORER CLADODE

CACTOBLASTIS DODDI LEPIDOPTERA PYRALIDAE CLADODE BORER CLADODE

MELITARA PRODENIALIS LEPIDOPTERA PYRALIDAE CLADODE BORER CLADODE

NANAIA SP LEPIDOPTERA PYRALIDAE CLADODE BORER CLADODE

OLYCELLA JUNCTOLINEELLA LEPIDOPTERA PYRALIDAE CLADODE BORER CLADODE

TUCUMANIA TAPIACOLA LEPIDOPTERA PYRALIDAE CLADODE BORER CLADODE

CACTOBLASTIS SP. NR DODDI LEPIDOPTERA PYRALIDAE CLADODE BORER CLADODE

PEMPELIA GENISTELLA LEPIDOPTERA PYRALIDAE DEFOLIATOR LEAF

HYPOCOSMIA PYROCHROMA LEPIDOPTERA PYRALIDAE LEAF ROLLER LEAF

BRADYRRHOA GILVEOLELLA LEPIDOPTERA PYRALIDAE ROOT FEEDER ROOT

ARCOLA MALLOI LEPIDOPTERA PYRALIDAE STEM FEEDER STEM

MELITARA SP. LEPIDOPTERA PYRALIDAE UNKNOWN UNKNOWN

SCHRECKENSTEINIA FESTALIELLA LEPIDOPTERA SCHRECKENSTEINIIDAE DEFOLIATOR LEAF

SCYTHRIS GRANDIPENNIS LEPIDOPTERA SCYTHRIDIDAE DEFOLIATOR LEAF

PENNISETIA MARGINATA LEPIDOPTERA SESIIDAE CROWN BORER CROWN

CHAMAESPHECIA ASTATIFORMIS LEPIDOPTERA SESIIDAE ROOT BORER ROOT

CHAMAESPHECIA CRASSICORNIS LEPIDOPTERA SESIIDAE ROOT BORER ROOT

CHAMAESPHECIA EMPIFORMIS LEPIDOPTERA SESIIDAE ROOT BORER ROOT

CHAMAESPHECIA HUNGARICA LEPIDOPTERA SESIIDAE ROOT BORER ROOT

CHAMAESPHECIA MYSINIFORMIS LEPIDOPTERA SESIIDAE ROOT BORER ROOT

CHAMAESPHECIA TENTHREDINIFORMIS LEPIDOPTERA SESIIDAE ROOT BORER ROOT

PYROPTERON DORYLIFORMIS LEPIDOPTERA SESIIDAE ROOT BORER ROOT

CARMENTA SP. NR ITHACAE LEPIDOPTERA SESIIDAE MULTIPLE MULTIPLE

CARMENTA MIMOSA LEPIDOPTERA SESIIDAE STEM BORER STEM

MELITTIA OEDIPUS LEPIDOPTERA SESIIDAE STEM BORER STEM

HYLES EUPHORBIAE LEPIDOPTERA SPHINGIDAE DEFOLIATOR LEAF

EPISIMUS UNGUICULUS LEPIDOPTERA TORTRICIDAE DEFOLIATOR LEAF

LOBESIA EUPHORBIANA LEPIDOPTERA TORTRICIDAE DEFOLIATOR LEAF

LORITA BACCHARIVORA LEPIDOPTERA TORTRICIDAE DEFOLIATOR LEAF

CROCIDOSEMA LANTANA LEPIDOPTERA TORTRICIDAE FLOWER FEEDER FLOWER

CROESIA ZIMMERMANI LEPIDOPTERA TORTRICIDAE LEAF ROLLER LEAF

STREPSICRATES SMITHIANA LEPIDOPTERA TORTRICIDAE LEAF ROLLER LEAF

TORTRIX S.L. SUBSP. CHRYSANTHEMOIDES LEPIDOPTERA TORTRICIDAE LEAF ROLLER LEAF

AGAPETA ZOEGANA LEPIDOPTERA TORTRICIDAE ROOT BORER ROOT

PELOCHRISTA MEDULLANA LEPIDOPTERA TORTRICIDAE ROOT BORER ROOT

CYDIA SUCCEDANA LEPIDOPTERA TORTRICIDAE SEEDPOD FEEDER SEEDPOD

COCHYLIS ATRICAPITANA LEPIDOPTERA TORTRICIDAE MULTIPLE MULTIPLE

BACTRA VENOSANA LEPIDOPTERA TORTRICIDAE STEM BORER STEM

PLATPHALONIDIA MYSTICA LEPIDOPTERA TORTRICIDAE STEM BORER STEM

EPIBLEMA STRENUANA LEPIDOPTERA TORTRICIDAE STEM GALLER STEM

DICHRORAMPHA ODORATA LEPIDOPTERA TORTRICIDAE STEM TIP GALLER STEM

CORNOPS AQUATICUM ORTHOPTERA ACRIDIDAE DEFOLIATOR LEAF

AMYNOTHRIPS ANDERSONI THYSANOPTERA PHLAEOTHRIPIDAE LEAF MERISTEM LEAF

LIOTHRIPS URICHI THYSANOPTERA PHLAEOTHRIPIDAE UNKNOWN UNKNOWN

SERICOTHRIPS STAPHYLINUS THYSANOPTERA THRIPIDAE SHOOT FEEDER SHOOT

LIOTHRIPS TRACTABILIS THYSANOPTERA THRIPIDAE STEM GALLER STEM

APPENDIX 2. HOST WEEDS AND HABITAT CLASSIFICATIONS

FAMILY SPECIES COMMON NAME HABITAT CLASSIFICATION

AMARANTHACEAE ALTERNANTHERA PHILOXEROIDES ALLIGATOR WEED AQUATIC/WETLAND

ARACEAE PISTIA STRATIOTES WATER LETTUCE AQUATIC/WETLAND

ASTERACEAE BACCHARIS HALIMIFOLIA SEA MYRTLE AQUATIC/WETLAND

AZOLLACEAE AZOLLA FILICULOIDES WATER FERN AQUATIC/WETLAND

FABACEAE MIMOSA PIGRA GIANT SENSITIVE PLANT AQUATIC/WETLAND

FABACEAE SESBANIA PUNICEA RED SESBANI AQUATIC/WETLAND

HALORAGACEAE MYRIOPHYLLUM AQUATICUM PARROT'S FEATHER AQUATIC/WETLAND

HYDROCHARITACEAE HYDRILLA VERTICILLATA HYDRILLA AQUATIC/WETLAND

LYGODIACEAE LYGODIUM MICROPHYLLUM OLD WORLD CLIMBING FERN AQUATIC/WETLAND

LYTHRACEAE LYTHRUM SALICARIA PURPLE LOOSESTRIFE AQUATIC/WETLAND

MYRTACEAE MELALEUCA QUINQUENERVIA MELALEUCA AQUATIC/WETLAND

POACEAE ARUNDO DONAX GIANT REED AQUATIC/WETLAND

PONTEDERIACEAE EICHHORNIA CRASSIPES WATER HYACINTH AQUATIC/WETLAND

SALVINIACEAE SALVINIA MOLESTA SALVINIA AQUATIC/WETLAND

ANACARDIACEAE SCHINUS TEREBINTHIFOLIUS BRAZILIAN PEPPERTREE TERRESTRIAL

APOCYNACEAE CRYPTOSTEGIA GRANDIFLORA RUBBER VINE TERRESTRIAL

APOCYNACEAE VINCETOXICUM ROSSICUM DOG-STRANGLING VINE TERRESTRIAL

ASPARAGACEAE ASPARAGUS ASPARAGOIDES BRIDAL CREEPER TERRESTRIAL

ASTERACEAE AGERATINA ADENOPHORA CROFTON WEED TERRESTRIAL

ASTERACEAE AGERATINA RIPARIA MISTFLOWER TERRESTRIAL

ASTERACEAE AMBROSIA ARTEMISIIFOLIA COMMON RAGWEED TERRESTRIAL

ASTERACEAE AMBROSIA PSILOSTACHYA WESTERN RAGWEED TERRESTRIAL

ASTERACEAE CAMPULOCLINIUM MACROCEPHALUM POMPOM WEED TERRESTRIAL

ASTERACEAE CARDUUS ACANTHOIDES SPINY PLUMELESS THISTLE TERRESTRIAL

ASTERACEAE CARDUUS NUTANS MUSK THISTLE TERRESTRIAL

ASTERACEAE CARDUUS NUTANS L. SUBSP. NUTANS NODDING THISTLE TERRESTRIAL

ASTERACEAE CARDUUS PYCNOCEPHALUS SLENDER WINGED THISTLE TERRESTRIAL

ASTERACEAE CARDUUS TENUIFLORUS WINGED THISTLE TERRESTRIAL

ASTERACEAE CENTAUREA CALCITRAPA PURPLE STARTHISTLE TERRESTRIAL

ASTERACEAE CENTAUREA CYANUS CORNFLOWER TERRESTRIAL

ASTERACEAE CENTAUREA DIFFUSA DIFFUSE KNAPWEED TERRESTRIAL

ASTERACEAE CENTAUREA IBERICA IBERIAN STARTHISTLE TERRESTRIAL

ASTERACEAE CENTAUREA JACEA L. NOTHOSUBSP. MEADOW KNAPWEED TERRESTRIAL

ASTERACEAE CENTAUREA JACEA L. SUBSP. JACEA BROWN KNAPWEED TERRESTRIAL

ASTERACEAE CENTAUREA JACEA L. SUBSP. NIGRA BLACK KNAPWEED TERRESTRIAL

ASTERACEAE CENTAUREA SOLSTITIALIS YELLOW STARTHISTLE TERRESTRIAL

ASTERACEAE CENTAUREA STOEBE L. SENS. LAT. SPOTTED KNAPWEED TERRESTRIAL

ASTERACEAE CENTAUREA VIRGATA LAM. SUBSP. SQUARROSA SQUARROSE KNAPWEED TERRESTRIAL

ASTERACEAE CHONDRILLA JUNCEA SKELETON WEED TERRESTRIAL

ASTERACEAE CHROMOLAENA ODORATA TRIFFID WEED TERRESTRIAL

ASTERACEAE CHRYSANTHEMOIDES MONILIFERA NORL. SUBSP. MONILIFERA BONESEED TERRESTRIAL

ASTERACEAE CHRYSANTHEMOIDES MONILIFERA NORL. SUBSP. ROTUNDATA BITOU BUSH TERRESTRIAL

ASTERACEAE CIRSIUM ARVENSE CANADA THISTLE TERRESTRIAL

ASTERACEAE CIRSIUM PALUSTRE MARSH THISTLE TERRESTRIAL

ASTERACEAE CIRSIUM VULGARE SPEAR THISTLE TERRESTRIAL

ASTERACEAE ELEPHANTOPUS MOLLIS ELEPHANT'S FOOT TERRESTRIAL

ASTERACEAE GUTIERREZIA SPP. SNAKEWEEDS TERRESTRIAL

ASTERACEAE JACOBAEA VULGARIS RAGWORT TERRESTRIAL

ASTERACEAE MIKANIA MICRANTHA BITTER VINE TERRESTRIAL

ASTERACEAE ONOPORDUM ACANTHIUM SCOTCH THISTLE TERRESTRIAL

ASTERACEAE ONOPORDUM ACAULON STEMLESS THISTLE TERRESTRIAL

ASTERACEAE ONOPORDUM SPP. SCOTCH/ILLYRIAN THISTLE TERRESTRIAL

ASTERACEAE PARTHENIUM HYSTEROPHORUS PARTHENIUM WEED TERRESTRIAL

ASTERACEAE PILOSELLA AURANTIACA ORANGE HAWKWEED TERRESTRIAL

ASTERACEAE PILOSELLA FLAGELLARIS WHIPLASH HAWKWEED TERRESTRIAL

ASTERACEAE PILOSELLA OFFICINARUM MOUSE-EAR HAWKWEED TERRESTRIAL

ASTERACEAE PLUCHEA CAROLINENSIS SOUR BUSH TERRESTRIAL

ASTERACEAE RHAPONTICUM REPENS RUSSIAN KNAPWEED TERRESTRIAL

ASTERACEAE SENECIO MADAGASCARIENSIS MADAGASCAR FIREWEED TERRESTRIAL

ASTERACEAE SILYBUM MARIANUM MILK THISTLE TERRESTRIAL

ASTERACEAE SONCHUS ARVENSIS PERENNIAL SOW-THISTLE TERRESTRIAL

ASTERACEAE TRIPLEUROSPERMUM INODORUM SCENTLESS CHAMOMILE TERRESTRIAL

ASTERACEAE XANTHIUM STRUMARIUM NOOGOORA BURR TERRESTRIAL

BALSAMINACEAE IMPATIENS GLANDULIFERA HIMALAYAN BALSAM TERRESTRIAL

BASELLACEAE ANREDERA CORDIFOLIA MADEIRA-VINE TERRESTRIAL

BIGNONIACEAE DOLICHANDRA UNGUIS-CATI CAT’S CLAW CREEPER TERRESTRIAL

BIGNONIACEAE TECOMA STANS YELLOW BELLS TERRESTRIAL

BORAGINACEAE CORDIA CURASSAVICA BLACK SAGE TERRESTRIAL

BORAGINACEAE CYNOGLOSSUM OFFICINALE HOUNDSTONGUE TERRESTRIAL

BORAGINACEAE ECHIUM PLANTAGINEUM PATERSON'S CURSE TERRESTRIAL

BORAGINACEAE HELIOTROPIUM AMPLEXICAULE BLUE HELIOTROPE TERRESTRIAL

BORAGINACEAE HELIOTROPIUM EUROPAEUM COMMON HELIOTROPE TERRESTRIAL

CACTACEAE ACANTHOCEREUS TETRAGONUS SWORD PEAR TERRESTRIAL

CACTACEAE CEREUS JAMACARU DC. SUBSP. JAMACARU QUEEN OF THE NIGHT TERRESTRIAL

CACTACEAE CYLINDROPUNTIA FULGIDA VAR. FULGIDA CHAIN-FRUIT CHOLLA TERRESTRIAL

CACTACEAE CYLINDROPUNTIA FULGIDA VAR. MAMILLATA BOXING GLOVE CACTUS TERRESTRIAL

CACTACEAE CYLINDROPUNTIA IMBRICATA IMBRICATE PRICKLY PEAR TERRESTRIAL

CACTACEAE CYLINDROPUNTIA LEPTOCAULIS PENCIL CACTUS TERRESTRIAL

CACTACEAE HARRISIA BALANSAE HARRISIA BALANSAE TERRESTRIAL

CACTACEAE HARRISIA MARTINII HARRISIA CACTUS TERRESTRIAL

CACTACEAE HARRISIA POMANENSIS MIDNIGHT LADY TERRESTRIAL

CACTACEAE HARRISIA REGELII HARRISIA CACTUS TERRESTRIAL

CACTACEAE HARRISIA TORTUOSA MILLMERRAN HARRISIA CACTUS TERRESTRIAL

CACTACEAE OPUNTIA AURANTIACA JOINTED CACTUS TERRESTRIAL

CACTACEAE OPUNTIA ELATIOR PRICKLY PEAR TERRESTRIAL

CACTACEAE OPUNTIA ENGELMANNII TEXAS PRICKLY PEAR TERRESTRIAL

CACTACEAE OPUNTIA FICUS-INDICA INDIAN FIG TERRESTRIAL

CACTACEAE OPUNTIA HUMIFUSA CREEPING PRICKLY PEAR TERRESTRIAL

CACTACEAE OPUNTIA LITTORALIS PRICKLY PEAR TERRESTRIAL

CACTACEAE OPUNTIA MONACANTHA PRICKLY PEAR TERRESTRIAL

CACTACEAE OPUNTIA ORICOLA PRICKLY PEAR TERRESTRIAL

CACTACEAE OPUNTIA SPP. PRICKLY PEAR TERRESTRIAL

CACTACEAE OPUNTIA STREPTACANTHA WESTWOOD PEAR TERRESTRIAL

CACTACEAE OPUNTIA STRICTA SPINY PEST PEAR TERRESTRIAL

CACTACEAE OPUNTIA TOMENTOSA VELVET OPUNTIA TERRESTRIAL

CACTACEAE OPUNTIA TRIACANTHA SUCKERS TERRESTRIAL

CACTACEAE OPUNTIA TUNA PRICKLY PEAR TERRESTRIAL

CACTACEAE PERESKIA ACULEATA BARBADOS GOOSEBERRY TERRESTRIAL

CARYOPHYLLACEAE SILENE VULGARIS BLADDER CAMPION TERRESTRIAL

CHENOPODIACEAE HALOGETON GLOMERATUS HALOGETON TERRESTRIAL

CHENOPODIACEAE SALSOLA TRAGUS RUSSIAN THISTLE TERRESTRIAL

COMMELINACEAE TRADESCANTIA FLUMINENSIS TRADESCANTIA TERRESTRIAL

CONVOLVULACEAE CALYSTEGIA SEPIUM HEDGE BINDWEED TERRESTRIAL

CONVOLVULACEAE CONVOLVULUS ARVENSIS FIELD BINDWEED TERRESTRIAL

CONVOLVULACEAE CUSCUTA AMERICANA LOVE VINE TERRESTRIAL

CONVOLVULACEAE CUSCUTA INDECORA LOVE VINE TERRESTRIAL

CONVOLVULACEAE CUSCUTA REFLEXA DODDER TERRESTRIAL

CUCURBITACEAE COCCINIA GRANDIS IVY GOURD TERRESTRIAL

CYPERACEAE CYPERUS ROTUNDUS NUT GRASS TERRESTRIAL

DIOSCOREACEAE DIOSCOREA BULBIFERA AIR POTATO TERRESTRIAL

ERICACEAE CALLUNA VULGARIS HEATHER TERRESTRIAL

EUPHORBIACEAE EUPHORBIA CYPARISSIAS CYPRESS SPURGE TERRESTRIAL

EUPHORBIACEAE EUPHORBIA ESULA LEAFY SPURGE TERRESTRIAL

EUPHORBIACEAE EUPHORBIA OBLONGATA OBLONG SPURGE TERRESTRIAL

EUPHORBIACEAE JATROPHA GOSSYPIIFOLIA BELLYACHE BUSH TERRESTRIAL

FABACEAE ACACIA BAILEYANA BAILEY'S WATTLE TERRESTRIAL

FABACEAE ACACIA CYCLOPS RED EYE TERRESTRIAL

FABACEAE ACACIA DEALBATA SILVER WATTLE TERRESTRIAL

FABACEAE ACACIA DECURRENS GREEN WATTLE TERRESTRIAL

FABACEAE ACACIA LONGIFOLIA LONG LEAVED WATTLE TERRESTRIAL

FABACEAE ACACIA MEARNSII BLACK WATTLE TERRESTRIAL

FABACEAE ACACIA MELANOXYLON AUSTRALIAN BLACKWOOD TERRESTRIAL

FABACEAE ACACIA PODALYRIIFOLIA PEARL ACACIA TERRESTRIAL

FABACEAE ACACIA PYCNANTHA GOLDEN WATTLE TERRESTRIAL

FABACEAE ACACIA SALIGNA PORT JACKSON WILLOW TERRESTRIAL

FABACEAE CAESALPINIA DECAPETALA MAURITIUS THORN TERRESTRIAL

FABACEAE CYTISUS SCOPARIUS SCOTCH BROOM TERRESTRIAL

FABACEAE GALEGA OFFICINALIS GOAT'S RUE TERRESTRIAL

FABACEAE LEUCAENA LEUCOCEPHALA LEUCAENA TERRESTRIAL

FABACEAE MIMOSA DIPLOTRICHA GIANT SENSITIVE PLANT TERRESTRIAL

FABACEAE PARASERIANTHES LOPHANTHA STINK BEAN TERRESTRIAL

FABACEAE PARKINSONIA ACULEATA RETAMA TERRESTRIAL

FABACEAE PROSOPIS JULIFLORA MEXICAN THORN TERRESTRIAL

FABACEAE PROSOPIS SPP. MESQUITE TERRESTRIAL

FABACEAE ULEX EUROPAEUS GORSE TERRESTRIAL

FABACEAE VACHELLIA NILOTICA SUBSP. INDICA PRICKLY ACACIA TERRESTRIAL

HYPERICACEAE HYPERICUM ANDROSAEMUM TUTSAN TERRESTRIAL

HYPERICACEAE HYPERICUM PERFORATUM ST JOHN'S WORT TERRESTRIAL

LAMIACEAE CLERODENDRUM CHINENSE HONOLULU ROSE TERRESTRIAL

LAMIACEAE MARRUBIUM VULGARE HOREHOUND TERRESTRIAL

LAMIACEAE SALVIA AETHIOPIS MEDITERRANEAN SAGE TERRESTRIAL

MALVACEAE SIDA ACUTA SPINYHEAD SIDA TERRESTRIAL

MALVACEAE SIDA RHOMBIFOLIA PADDY'S LUCERNE TERRESTRIAL

MELASTOMATACEAE CLIDEMIA HIRTA KOSTER'S CURSE TERRESTRIAL

MELASTOMATACEAE MICONIA CALVESCENS MICONIA TERRESTRIAL

MYRICACEAE MORELLA FAYA FIREBUSH TERRESTRIAL

MYRTACEAE LEPTOSPERMUM LAEVIGATUM AUSTRALIAN MYRTLE TERRESTRIAL

OROBANCHACEAE OROBANCHE MINOR BROOMRAPE TERRESTRIAL

OROBANCHACEAE PHELIPANCHE RAMOSA BRANCHED BROOMRAPE TERRESTRIAL

OROBANCHACEAE STRIGA HERMONTHICA PURPLE WITCHWEED TERRESTRIAL

PASSIFLORACEAE PASSIFLORA TARMINIANA BANANA POKA TERRESTRIAL

PLANTAGINACEAE LINARIA DALMATICA SUBSP. DALMATICA DALMATIAN TOADFLAX TERRESTRIAL

PLANTAGINACEAE LINARIA VULGARIS YELLOW TOADFLAX TERRESTRIAL

POLYGONACEAE EMEX AUSTRALIS THREE CORNERED JACKS TERRESTRIAL

POLYGONACEAE EMEX SPINOSA LESSER JACKS TERRESTRIAL

POLYGONACEAE FALLOPIA JAPONICA JAPANESE KNOTWEED TERRESTRIAL

POLYGONACEAE PERSICARIA PERFOLIATA MILE-A-MINUTE WEED TERRESTRIAL

POLYGONACEAE RUMEX SPP. DOCKS TERRESTRIAL

PROTEACEAE HAKEA GIBBOSA ROCK HAKEA TERRESTRIAL

PROTEACEAE HAKEA SERICEA SILKY HAKEA TERRESTRIAL

RANUNCULACEAE CLEMATIS VITALBA OLD MAN'S BEARD TERRESTRIAL

ROSACEAE ACAENA ANSERINIFOLIA PIRIPIRI TERRESTRIAL

ROSACEAE RUBUS ALCEIFOLIUS GIANT BRAMBLE TERRESTRIAL

ROSACEAE RUBUS ARGUTUS PRICKLY FLORIDA BLACKBERRY TERRESTRIAL

ROSACEAE RUBUS CONSTRICTUS BLACKBERRY TERRESTRIAL

ROSACEAE RUBUS FRUTICOSUS EUROPEAN BLACKBERRY TERRESTRIAL

ROSACEAE RUBUS ULMIFOLIUS ZARZAMORA TERRESTRIAL

RUBIACEAE GALIUM SPURIUM FALSE CLEAVERS TERRESTRIAL

SAPINDACEAE CARDIOSPERMUM GRANDIFLORUM BALLOON VINE TERRESTRIAL

SCROPHULARIACEAE BUDDLEJA DAVIDII BUDDLEIA TERRESTRIAL

SOLANACEAE SOLANUM ELAEAGNIFOLIUM SILVERLEAF NIGHTSHADE TERRESTRIAL

SOLANACEAE SOLANUM MAURITIANUM BUGWEED TERRESTRIAL

SOLANACEAE SOLANUM SISYMBRIIFOLIUM WILD TOMATO TERRESTRIAL

SOLANACEAE SOLANUM VIARUM TROPICAL SODA APPLE TERRESTRIAL

TAMARICACEAE TAMARIX SPP. SALT CEDAR TERRESTRIAL

VERBENACEAE LANTANA CAMARA LANTANA TERRESTRIAL

VERBENACEAE LANTANA MONTEVIDENSIS CREEPING LANTANA TERRESTRIAL

ZYGOPHYLLACEAE TRIBULUS CISTOIDES FALSE PUNCTUREVINE TERRESTRIAL

ZYGOPHYLLACEAE TRIBULUS TERRESTRIS PUNCTUREVINE TERRESTRIAL

LORANTHACEAE PHTHIRUSA STELIS BIRD VINE UNDETERMINED

MELASTOMATACEAE MELASTOMA SEPTEMNERVIUM ASIAN MELASTOME UNDETERMINED