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 and agent characteristics. Multiple such relationships were identified, including previously identified correlations between CWBC outcomes, host 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 ; 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 agents have been released against over 150 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 , 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 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 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 , which has been used to effectively control a number of 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 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.

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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 CLADODE AND ROOT ARMATUM COLEOPTERA CERAMBYCIDAE BORER MULTIPLE CLADODE AND ROOT MONEILEMA BLAPSIDES COLEOPTERA CERAMBYCIDAE BORER MULTIPLE CLADODE AND ROOT MONEILEMA VARIOLARE COLEOPTERA CERAMBYCIDAE 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 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 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 LEAF MINER LEAF BUCCULATRIX PARTHENICA LEPIDOPTERA BUCCULATRICIDAE LEAF MINER LEAF BULLATA LEPIDOPTERA 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 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 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 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 LEAF MINER LEAF 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 DEFOLIATOR LEAF ASSIMILELLA LEPIDOPTERA 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 DENTATA LEPIDOPTERA 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 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 CROWN BORER CROWN 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 DEFOLIATOR LEAF EPISIMUS UNGUICULUS LEPIDOPTERA DEFOLIATOR LEAF LOBESIA EUPHORBIANA LEPIDOPTERA TORTRICIDAE DEFOLIATOR LEAF LORITA BACCHARIVORA LEPIDOPTERA TORTRICIDAE DEFOLIATOR LEAF 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

HABITAT FAMILY SPECIES COMMON NAME 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 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 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 BALANSAE HARRISIA BALANSAE TERRESTRIAL CACTACEAE HARRISIA MARTINII HARRISIA CACTUS TERRESTRIAL CACTACEAE HARRISIA POMANENSIS MIDNIGHT LADY TERRESTRIAL CACTACEAE HARRISIA REGELII HARRISIA CACTUS TERRESTRIAL MILLMERRAN HARRISIA CACTACEAE HARRISIA TORTUOSA CACTUS TERRESTRIAL CACTACEAE 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 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 EUROPAEUS GORSE TERRESTRIAL FABACEAE VACHELLIA NILOTICA SUBSP. INDICA PRICKLY ACACIA TERRESTRIAL HYPERICACEAE HYPERICUM ANDROSAEMUM TUTSAN TERRESTRIAL HYPERICACEAE 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 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 PRICKLY FLORIDA ROSACEAE RUBUS ARGUTUS 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 SOLANUM ELAEAGNIFOLIUM SILVERLEAF NIGHTSHADE TERRESTRIAL SOLANACEAE SOLANUM MAURITIANUM BUGWEED TERRESTRIAL SOLANACEAE SOLANUM SISYMBRIIFOLIUM WILD TOMATO TERRESTRIAL SOLANACEAE 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