A Meta-analysis of Canada Thistle Cirsium arvense Management Authors: Stacy Davis, Jane Mangold, Fabian Menalled, Noelle Orloff, Zach Miller, and Erik Lehnhoff This is a pre-copyedited, author-produced PDF of an article accepted for publication in Weed Science following peer review. The version of record, see full citation below, is available online at: https://dx.doi.org/10.1017/wsc.2018.6. Davis, Stacy, Jane Mangold, Fabian Menalled, Noelle Orloff, Zach Miller, and Erik Lehnhoff. "A Meta-analysis of Canada Thistle Cirsium arvense Management." Weed Science 66, no. 4 (July 2018): 548-557. DOI:10.1017/wsc.2018.6. Made available through Montana State University’s ScholarWorks scholarworks.montana.edu 1 Short title for running footer: Management of Canada Thistle A Meta-Analysis of Canada Thistle (Cirsium arvense) Management Stacy Davis, Jane Mangold, Fabian Menalled, Noelle Orloff, Zach Miller, and Erik Lehnhoff Although stand-alone and integrated management techniques have been cited as viable approaches to managing Canada thistle [Cirsium arvense (L.) Scop.], it continues to impact annual cropping and perennial systems world-wide. We conducted meta-analyses assessing effectiveness of management techniques and herbicide mechanism of action groups for controlling Canada thistle using 55 studies conducted in annual cropping systems and 45 studies in perennial systems. Herbicide was the most studied technique in both types of systems and was effective at reducing Canada thistle. However, integrated multi-tactic techniques, with or without herbicides, were more effective than sole reliance on herbicides for long-term control in both annual cropping and perennial systems. A variety of management techniques such as biocontrol, crop diversification, mowing, and soil disturbance provided similar control as herbicide. Our results suggest that many management techniques aimed at reducing Canada thistle can also improve crop yield or abundance of desired plants. This study highlights the need to devote more research to non-chemical and integrated management approaches for Canada thistle control.  First, second, and third authors: Research Associate, Associate Professor, and Professor, Department of Land Resources and Environmental Sciences, Montana State University, Bozeman, MT 59717; Fourth author: Associate Extension Specialist, Schutter Diagnostic Lab, Montana State University, Bozeman, MT 59717; Fifth author: Assistant Professor and Superintendent, Western Agricultural Research Center, Montana State University, Corvallis, MT 59828; Sixth author: Assistant Professor, Department of Entomology, Plant Pathology, and Weed Science, New Mexico State University, Las Cruces, NM 88003. Corresponding author’s E-mail: stacy.davis1@montana.edu 2 Nomenclature: Canada thistle, Cirsium arvense (L.) CIRARV Key words: Noxious weeds, invasive plants, herbicide, integrated weed management, perennial weeds, herbicide mechanism of action. 3 The control of and impacts resulting from perennial, invasive plants are challenging in both annual cropping systems and perennial plant communities. For example, a recent review of invasive plant control publications found that 16 of the 20 most-studied species were perennials (Kettenring and Adams 2011) and 83% of the most commonly listed noxious weeds in the United States and Canada were perennials (Skinner et al. 2000). Many perennial species spread both by seed and vegetatively, enabling effective dispersal (Hakansson 2003a). Such perennial plants also have the ability to store carbohydrate reserves in their extensive root systems. Due to these biological characteristics, perennial weeds can be persistent, difficult to manage, and tolerant of certain management techniques, such as mechanical disturbance (Hakansson 2003b). Canada thistle (Cirsium arvense L.) is a perennial plant that is particularly difficult to control once established in both annual cropping and perennial systems (Tiley 2010). Canada thistle was first introduced to North America in the 1600’s from Europe via contaminated grain seed, hay, and ship’s ballast (Morishita 1999). It commonly invades croplands, natural areas, pastures, rangelands, and roadsides (Tiley 2010). Canada thistle was listed as a noxious weed in all but four U.S. states by 1957 (Tiley 2010), and as of 2000 it was the most frequently listed noxious weed in the United States and Canada (Skinner et al. 2000). Producers on certified organic land in the Pacific Northwest and Great Plains regions recently listed Canada thistle among the most problematic weeds (OAEC 2013; Tautges et al. 2016). This invasive plant is an effective competitor, with a tall growth form and efficient vegetative spread aiding its rapid colonization ability and suppression of other plants (Tiley 2010). Yield losses in a variety of annual crops including barley (O'Sullivan et al. 1982), rapeseed (O'Sullivan et al. 1985), and spring wheat (Donald and Khan 1996) are associated with increased Canada thistle density. In addition, Canada thistle infestations can cause further 4 economic losses due to contamination of seed, grain, or straw, which results in changes in product handling, processing, and quality (Tiley 2010). Although Canada thistle is regarded as a major weed in perennial systems, there is less information published on yield losses in perennial pastures compared to annual crops (Tiley 2010). Grekul and Bork (2004) demonstrated significant grass and forb yield losses associated with increasing Canada thistle density in perennial pastures in western Canada. Furthermore, Canada’s thistle’s prickly mature foliage can reduce pasture productivity by deterring livestock from grazing in infested areas (Tiley 2010). Although stand-alone and integrated management techniques have been cited as viable approaches to managing Canada thistle (Donald 1990), it continues to invade and persist in temperate regions of the world. Carefully reviewing and systematically summarizing results from previous studies may help refine management strategies for Canada thistle. A statistical tool useful for achieving this goal is meta-analysis, i.e., the systematic and quantitative review and synthesis of previous studies (Koricheva and Gurevitch 2014). For example, through a meta- analysis of 52 studies, Lutman et al. (2013) determined that mechanical cultivation and time of seeding were the most efficient non-chemical approaches to manage blackgrass (Alopecurus myosuroides Huds.). Additionally, a meta-analysis on downy brome (Bromus tectorum L.) management indicated that a variety of control methods reduced short-term abundance of downy brome, but only those that included herbicide or revegetation led to long-term control (Monaco et al. 2017). Finally, a recently completed meta-analysis of management of Canada thistle and field bindweed (Convolvulus arvensis L.) in organic cropping systems showed that integrating two or more management strategies generally caused greater reductions in weed abundance than any method used alone (N Orloff, personal communication). Meta-analyses can also identify knowledge gaps and potential ways to improve experimental approaches. For example, a recent 5 review found that most experimental approaches assessing weed control of invasive plants evaluated efficacy of management techniques on short time frames (<1 year) and rarely included impacts on desirable vegetation or evaluations of control costs (Kettenring and Adams 2011). We conducted a meta-analysis to review and summarize results from previously published studies involving Canada thistle management in annual cropping and perennial systems. Our objectives were to 1) assess short- and long-term effectiveness of management techniques for controlling Canada thistle, 2) compare short- and long-term effectiveness of different herbicide mechanism of action (MOA) groups for controlling Canada thistle, 3) determine if and how management techniques for Canada thistle control impact crop yield (annual cropping systems) or abundance of desired plants (perennial systems), and 4) identify knowledge gaps. These objectives were conducted separately for annual cropping systems (row crop and fallow fields) and perennial systems (pasture, rangeland, natural areas, etc.). Materials and Methods Literature Search and Study Inclusion. In December 2015 we conducted a literature search for Canada thistle using the Web of Science® (1864-2015) and Agricola® (1927-2015) databases. We used the keywords “Cirsium arvense,” “Carduus arvensis,” “Canada thistle,” “creeping thistle,” “Californian thistle,” and “field thistle.” We limited our search to articles written in English. Following guidelines by Koricheva et al. (2013), all references underwent the following filtering process for their inclusion into the meta-analysis: (1) duplicate references from the two databases were removed, (2) abstracts and titles of retrieved articles were examined and clearly irrelevant literature (e.g., patents, studies about ecology or biology with no control treatments, 6 medical topics, genetics studies, pollination studies) was removed, and (3) full text of selected articles was examined and studies were included that met our pre-established inclusion criteria. Specifically, we included replicated field studies that assessed the relative efficacy of stand-alone or integrated weed management techniques taking place in annual cropping or perennial systems. We limited annual cropping system studies to cooler, temperate climatic regions specified as those that grew crops listed by the US Department of Agriculture state agriculture overview for the Northern Great Plains states (defined as Montana, Nebraska, North Dakota, South Dakota, and Wyoming): wheat (Triticum aestivum L.), hay (Medicago sativa L.), barley (Hordeum vulgare L.), peas (Pisum sativum), potatoes (Solamum tuberosum), lentils (Lens culinaris), corn (Zea mays L.), beans (Phaseolus vulgaris), canola (Brassica napus L.), safflower (Carthamus tinctorius), oats (Avena spp.), sugar beets (Beta vulgaris), sunflower (Helianthus annuus L.), sorghum (Sorghum bicolor (L.) Moench ssp. bicolor), millet (Panicum miliaceum L.), legumes (Medicago sativa L.), mustard (Brassica nigra (L.) W.D.J. Koch; Sinapis alba L.), and flaxseed (Linum usitatissimum L.) (USDA 2017). Perennial systems included rangelands, pastures, lawns, alfalfa, hay fields, and natural areas world-wide. Studies utilizing herbicides were included only if the applied herbicide was approved for use according to Shaner (2014) and if it was applied within recommended rates (Greenbook 2017; Shaner 2014). We followed terminology from Shaner (2014) and did not consider herbicides that had an unclassified MOA (e.g., sodium chlorate). We included studies with control/treatment comparisons that published quantitative response measurements for aboveground density, cover, biomass, frequency, survival, or percent control (measured from 0- 100%) of Canada thistle. Responses needed to be from established Canada thistle populations as 7 opposed to populations established artificially for testing purposes. The filtering process was conducted by a single author (S. Davis). Data Extraction and Synthesis. Following Gurevitch and Hedges (2001), we recorded means, measures of variation, and sample sizes for both control and treatment plots from published tables, within the text, or derived from published figures using WebPlotDigitizer (Rohatgi c2010-2017). Means included quantitative response measurements for aboveground density, cover, biomass, frequency, survival, or percent control of Canada thistle. We extracted additional information on type of system (annual versus perennial), study duration, and details of the treatment applied (e.g., herbicide type and rate, herbicide MOA group). Herbicide MOA groups followed the classification used in Shaner (2014) and were as follows: 2 (Acetolactate Synthase (ALS) or Acetohydroxy Acid Synthase (AHAS) Inhibitors), 4 (Synthetic Auxins), 5 (Inhibitors of photosynthesis at photosystem II site A), 6 (Inhibitors of photosynthesis at photosystem II site B), 9 (Inhibitor of 5-enolypyruvyl-shikimate-3-phosphate synthase (EPSPS)), 11 (Inhibitors of carotenoid biosynthesis (unknown target)), 27 (Inhibitors of 4-hydroxyhenyl- pyruvatedioxygenase (4-HPPD)). We included a “mix” herbicide MOA group, which we defined as an herbicide application including two or more herbicides from different groups. We also extracted data, when available, on crop yield or abundance of desired plants to examine how Canada thistle management techniques impacted them. Following Gurevitch and Hedges (2001), we developed a series of criteria to systematically extract data from the literature. If there were multiple types of response measurements within a study (e.g., density and cover), we selected one response type using a ranking process to avoid issues of non-independence. We ranked response measurements based on a pre-determined order of importance: 1) biomass, 2) cover, 3) density, 4) frequency, 5) 8 survival, and 6) percent control. Oftentimes, percent control response measurements did not include means for non-treated plots; therefore we assumed 0% control in non-treated plots. Additionally, percent control was transformed to the same scale as weed abundance measurements (i.e., biomass, cover, density, frequency, or survival) by subtracting percent control from 100. In this metric, lower numbers indicated more successful management. For each published article, if more than one site or treatment were assessed, data were extracted from each situation being tested. For multifactorial studies, we considered the response to each treatment as an independent data point (Gurevitch and Hedges 2001). For example, if an article compared five different types of herbicide active ingredients applied at two different rates, we extracted data for each herbicide active ingredient and rate combination, for a total of 10 data points. Although this meant the same control group mean was included for more than one data point, it allowed us to maximize use of existing valuable data on effectiveness of management techniques (Gurevitch and Hedges 2001). When multiple articles reported results on the same study across varying years, we used the data only once by extracting data from the latest set of observations. Many land managers apply management techniques every year for multiple years in a row, but in order to accurately compare short- and long-term control of Canada thistle, we only used response measurements that considered a single year of treatment. If responses to a single year of treatment were measured over multiple dates, we extracted data from two defined time periods (less than one year after treatment and one year or more after treatment) when possible and conducted separate analyses to compare short- versus long-term control. If there were repeated measures within our defined time periods, we only used the response from the longest time period since treatment (Gurevitch and Hedges 2001). 9 Data Analysis. For each data point, the effect size of a treatment was calculated as the log response ratio (lnR), where lnR = ln(XE/XC) = lnXE – lnXC [1] and XE and XC are means of experimental (treated) and control (non-treated) groups, respectively (Hedges et al. 1999). This variable quantifies the proportionate change that results from an experimental treatment and represents a meaningful approach to summarize and combine results of different studies (Hedges et al. 1999). We selected the response ratio for our analysis because it can be estimated without knowledge of sample sizes or variances (Adams et al. 1997). An example of an effect size measurement that utilizes measures of variance includes the standardized difference in means (e.g., Hedge’s D) (Koricheva et al. 2013). However, since many of our studies did not report measures of variation, we used the response ratio as our effect size measurement. Only 22% of data points from annual cropping systems and 2% of data points from perennial systems reported measures of variation. The response ratio cannot be calculated when data points have response measurements equal to zero because one cannot take a logarithm of a zero value (Koricheva et al. 2013). Therefore, 10 data points from annual cropping systems and 11 data points from perennial systems were excluded from the analysis (<3% of data for each system type). Although we could not include these data points, using the response ratio as our effect size allowed us to include a large amount of data that did not report measures of variation. We paired our response ratio with a nonparametric bootstrapping approach using a simplified weighting scheme. Meta-analyses using Hedge’s D as the effect size metric weight each effect size based on their relative sensitivity to measures of variance (Koricheva et al. 10 2013). However, since most of our studies lacked information about variance, we weighted each response ratio using sample sizes with the function FN, where FN= (nE x nC)/(nE + nC) [2] and nE and nC represent the number of replicates for the experimental (treated) and control (non- treated) groups, respectively (Adams et al. 1997). We used bootstrapping methods to calculate 95% confidence intervals around the pooled effect size mean with 1,000 iterations for individual management techniques or herbicide MOA groups (Adams et al. 1997). Individual management techniques or herbicide MOA groups were considered effective at managing Canada thistle if the mean response ratio was negative and the 95% confidence interval did not overlap zero (Adams et al. 1997; Gurevitch et al. 1992). For example, a 50% reduction in Canada thistle relative to a control group is equivalent to an effect size of -0.7. Mean response ratios from different management techniques or herbicide MOA groups were considered to be different from one another if their 95% confidence intervals did not overlap (de Graaff et al. 2006; Ferreira et al. 2015). Management techniques or herbicide MOA groups that had only one data point were included in figures to note knowledge gaps and should not be compared statistically to other management techniques or herbicide MOA groups because confidence intervals could not be calculated. All summaries and analyses were conducted in R statistical software (version 3.3.2), including the “plyr”, “ggplot2”, and “cowplot” packages (R Core Team 2016). We conducted separate meta-analyses corresponding to each objective. First, we examined management techniques used for Canada thistle control in annual cropping and perennial systems. Management techniques included biocontrol, burn, competition, crop diversification, fertilizer, herbicide, herbicide integrated, mowing, mulch, non-herbicide integrated, soil disturbance, and water manipulation (Table 1). We included the categories of 11 herbicide integrated and non-herbicide integrated to examine the effectiveness of integrated multi-tactic techniques, with or without herbicides. For our analyses in annual cropping and perennial systems, we examined management at different time periods by conducting two meta- analyses for each system, one for responses measured <1 year after treatment and another for responses measured ≥1 year after treatment. Next, we examined efficacy of different herbicide MOA groups (<1 year after treatment and ≥1 year after treatment). We did not compare additional specifics of individual management techniques (e.g., timing, types of biocontrol agents, herbicide rates) because this level of detail was outside the scope of our questions of interest and there was insufficient replication of specific practices within management techniques to adequately compare them. Finally, we compared the effect of management techniques on crop yield (annual) or abundance of desired plants (perennial). A positive response ratio indicated an increase in yield or abundance with treatment, while 95% confidence intervals overlapping zero indicated the management technique had no effect (Gurevitch et al. 1992). Complete bibliographies of the articles used in our annual cropping and perennial system analyses are given in Supplemental Appendices 1 and 2. Information from each article used in our meta-analysis, including study location, system description, management techniques, herbicide MOA groups (if applicable), and study duration grouping, is shown in Supplemental Tables 1 and 2. Results and Discussion Canada Thistle Management in Annual Cropping Systems. We extracted data from 55 articles published between 1957 and 2015, resulting in 650 total data points (Figure 1). The majority of studies took place in the United States (33 articles) and Canada (9 articles), while the 12 remaining took place in Denmark, England, Germany, Hungary, India, Iran, New Zealand, Norway, Poland, Serbia, and Sweden. Nearly three quarters (74%) of data points evaluated short- term efficacy of Canada thistle management (<1 year). All management techniques studied were effective at reducing Canada thistle in annual cropping systems when measured <1 year after treatment (Figure 2a). However, herbicide integrated management techniques were most effective for short-term control (Figure 2a). Herbicide, herbicide integrated, and soil disturbance were more effective than biocontrol and crop diversification (Figure 2a). Many management techniques were also effective ≥1 year after treatment. Biocontrol, crop diversification, herbicide, herbicide integrated, mowing, non- herbicide integrated, and soil disturbance all reduced Canada thistle ≥1 year after treatment (Figure 2b). Similar to effects <1 year after treatment, herbicide integrated management techniques were more effective than herbicide alone ≥1 year after treatment (Figure 2b). Additionally, herbicide integrated management techniques were similar to non-herbicide integrated management techniques ≥1 year after treatment (Figure 2b). Fertilizer had no effect on Canada thistle ≥1 year after treatment (Figure 2b). The availability of data for competition as a management technique in annual cropping systems was insufficient, with only one data point reporting treatment effects both <1 and ≥1 year after treatment. Although herbicide was the most studied management technique (79% of data points), other less studied management techniques were equally or more effective at controlling Canada thistle in annual cropping systems. For example, soil disturbance was as effective as herbicide <1 year after treatment but has not been studied to the extent that control via herbicide has. Furthermore, biocontrol, crop diversification, mowing, and soil disturbance all had the same level of effectiveness as herbicide ≥1 year after treatment. Only two data points in our meta- 13 analysis evaluated the long-term effectiveness of non-herbicide integrated management techniques, but these techniques resulted in improved control of Canada thistle versus herbicide alone (Figure 2b), highlighting the potential benefits of non-chemical multi-tactic strategies for Canada thistle control. Our meta-analysis also indicated that herbicide integrated management techniques resulted in greater reductions in Canada thistle than herbicide applied alone both <1 year and ≥1 year after treatment (Figure 2a, 2b). Herbicide integrated management techniques examined across both time periods included herbicide + fertilizer (3 data points), herbicide + mowing (14 data points), herbicide + soil disturbance (40 data points), and herbicide + soil disturbance + fertilizer (3 data points). Compared to herbicide used as a stand-alone technique, integrating herbicides with additional management techniques can help reduce crop injury and decrease the selective pressure towards the selection of herbicide resistance, while providing control of invasive perennial weed species (Miller 2016). All herbicide MOA groups reduced Canada thistle when measured <1 year after treatment (Figure 3a), while only half of the MOA groups tested ≥1 year after treatment were effective (Figure 3b). Less than one year after treatment, herbicide MOA groups 4, 6, 9, 11, 27, or mixes of MOA groups reduced Canada thistle more than herbicide MOA group 2 (Figure 3a). One year or more after treatment, herbicide MOA groups 4, 5 and 9 were effective at reducing Canada thistle (Figure 3b). However, herbicide MOA group 9 was slightly more effective than MOA group 4 (Figure 3b). Herbicide MOA group 4 (i.e., synthetic auxins) was most frequently tested ≥1 year after treatment (n=37) and included 2,4-D (32%), picloram (26%), and clopyralid (16%). Other MOA group 4 herbicides included dicamba, quinclorac, and MCPA (26%). Herbicide MOA group 5 (i.e., inhibitors of photosynthesis at photosystem II site A) assessed ≥1 14 year after treatment all consisted of atrazine in corn fields (n=6) (Parochetti 1974). Herbicide MOA group 9 (i.e., inhibitors of EPSPS, glyphosate) was used in 24 data points evaluated ≥1 year after treatment. Herbicide MOA groups 2, 11, or mixes of MOA groups did not provide Canada thistle control ≥1 year after treatment. Two of ten data points for the mix treatment had a positive effect size (i.e., increased Canada thistle). These treatments were bromoxynil + MCPA and glyphosate + bromoxynil + MCPA used in spring wheat (Carlson and Donald 1988). Overall, results from short-term studies suggest that all herbicide MOA groups studied have similar effectiveness. However, if the goal of the land manager is to reduce the frequency of herbicide applications, herbicide MOA groups 4, 5, and 9 are promising options for longer-term control of Canada thistle in annual cropping systems. Canada thistle causes significant yield losses in the northern part of North America (Tiley 2010), but our results suggest that management options can help reduce its impacts. In our meta- analysis, 90 data points reported on how various management techniques for Canada thistle control were associated with improved crop yield of barley, canola, corn, rapeseed, spring wheat, sugar beets, and winter wheat (Figure 4). These techniques included biocontrol, crop diversification, herbicide, herbicide integrated, non-herbicide integrated, and soil disturbance. Biocontrol, herbicide, and herbicide integrated management techniques were similarly associated with increased crop yield (Figure 4). Crop diversification and non-herbicide integrated management techniques were associated with increased crop yield more than biocontrol and herbicide (Figure 4). However, herbicide was the only management technique that had more than three data points recording crop yield, emphasizing the need to include measurements of crop yield in study design. The availability of data for fertilizer as a management technique was 15 insufficient to make comparisons with other techniques, with only one data point reporting treatment effects. Canada Thistle Management in Perennial Systems. We extracted data from 45 articles published between 1958 and 2015, resulting in 376 total data points (Figure 1). The majority of these studies took place in the United States (28 articles) and New Zealand (11 articles), while the remaining took place in Australia (1 article), Canada (2 articles), Czech Republic (1 article), Turkey (1 article), and the United Kingdom (1 article). Perennial systems studied included alfalfa fields, grass for seed, natural areas, pastures, rangelands, and roadsides. More than half of data points (58%) evaluated short-term efficacy of Canada thistle management (<1 year). Our meta-analysis revealed that biocontrol, competition, herbicide, herbicide integrated, mowing, mulch, and non-herbicide integrated management techniques reduced Canada thistle <1 year after treatment (Figure 5a). Herbicide and herbicide integrated management techniques were most effective at reducing Canada thistle, compared to all other techniques (Figure 5a). Fertilizer had no effect on Canada thistle control, whereas water manipulation increased the density of Canada thistle <1 year after treatment (Figure 5a). The three water manipulation data points were from low, medium, and high levels of irrigation on plots with forage species; all levels of irrigation resulted in an increase in density of Canada thistle (Thrasher et al. 1963). Only one data point existed for the impact of burning on Canada thistle, so comparisons to other management techniques should not be made. Many management techniques that were effective at reducing Canada thistle <1 year after treatment were also effective ≥1 year after treatment. These included biocontrol, herbicide, herbicide integrated, and mowing (Figure 5b). Although herbicide was equally effective as herbicide integrated <1 year after treatment, herbicide integrated was more effective than 16 herbicide alone ≥1 year after treatment (Figure 5b). Mowing was as effective as herbicide in controlling Canada thistle ≥1 year after treatment (Figure 5b). Mowing techniques included removing all vegetation to 5 cm using a sickle bar when Canada thistle was at early bud stage in a pasture (Grekul and Bork 2007), mowing alfalfa fields twice a year for hay (Hodgson 1958), and using a rotary mower set at 5 cm when Canada thistle was at bud stage in a pasture (Amor and Harris 1977). Even though competition decreased Canada thistle <1 year after treatment (Figure 5a), it had no effect on Canada thistle ≥1 year after treatment (Figure 5b). Similar to our annual cropping system results and the findings of other meta-analyses (Kettenring and Adams 2011), herbicide was the most studied management technique in perennial systems (61%). However, other management techniques were equally or more effective than herbicide for long-term control. Specifically, mowing was as effective as herbicide while herbicide integrated management techniques were more effective than herbicide applied alone ≥1 year after treatment. Herbicide integrated management techniques that resulted in long- term control included herbicide + burn (1 data point), herbicide + competition (2 data points), herbicide + soil disturbance (1 data point), herbicide + competition + mowing (1 data point), and herbicide + competition + soil disturbance (1 data point). While herbicides are the primary method of weed control in most rangelands, this study emphasizes the need to develop integrated weed management programs to achieve long-term control of weeds and healthy plant communities (DiTomaso 2000). All herbicide MOA groups reduced Canada thistle <1 year and ≥1 year after treatment (Figure 6a, 6b). Several herbicide MOA groups showed similar effectiveness in controlling Canada thistle. In particular, herbicide MOA groups 2, 4, and mixes of MOA groups were similar in effectiveness <1 year after treatment (Figure 6a). Herbicide MOA group 4 and mixes 17 of MOA groups showed higher effectiveness than herbicide MOA groups 5 and 6 <1 year after treatment (Figure 6a). Herbicide mixes included imazamox + bentazon, imazethapyr + bentazon, MCPB + bentazon, clopyralid + chlorsulfuron, and 2,4-D + chlorsulfuron. One year or more after treatment, herbicide MOA groups 2, 4, 9, and mixes of MOA groups were similar in effectiveness (Figure 6b). Herbicide mixes included dicamba + diflufenzopyr, 2,4-D + chlorsulfuron, and 2,4-D + metsulfuron. Herbicide MOA group 9 was represented in our meta- analysis by only six data points that measured control ≥1 year after treatment. This group is likely seldom used in pastures and rangelands because it includes non-selective products which could injure desired perennial vegetation. Data for herbicide MOA group 6 ≥1 year after treatment and MOA group 11 in both time periods was insufficient to make comparisons with other groups, with only one data point for each MOA group reporting treatment effects. Similar to our results in annual cropping systems, herbicide MOA group 4 was the most studied in perennial systems, with 81% of data points using this group across both time periods. Additionally, the majority of studies using mixes of MOA groups had group 4 as one of the ingredients (73% of data points). Herbicide MOA group 4 contains herbicides such as 2,4-D, dicamba, clopyralid, MCPA, and picloram that are considered important tools for managing perennial invasive species in rangelands and pastures (DiTomaso 2000; Morishita 1999). Common MOA group 4 herbicides used in studies in our meta-analysis included clopyralid (19%), 2,4-D (18%), dicamba (17%), a mix of two or more MOA group 4 herbicides (17%), and picloram (15%). Other MOA group 4 herbicides making up 15% of use in studies included aminopyralid, MCPB, and MCPA. In spite of the emphasis on MOA group 4 in past research, other MOA groups were just as effective at reducing Canada thistle ≥1 year after treatment (e.g., groups 2 and 9). 18 Both competition and herbicide were similarly associated with increased abundance of desired plants in perennial systems (competition mean lnR= 0.45, 95% CI: 0.21 to 0.67, n=5; herbicide mean lnR= 0.20, 95% CI: 0.08 to 0.32, n=27), as demonstrated by thirty-two data points measuring alfalfa yield, seed yield, and grass biomass. For example, decreasing row spacing of alfalfa was associated with an increase in alfalfa yield and a reduction in Canada thistle density (Celebi et al. 2010). A total of three articles that utilized herbicides also recorded abundance of desired plants with contrasting results. Mesbah and Miller (2005) found an increase in alfalfa seed yield after applying a variety of herbicides (18 data points), and Gallagher and Vandenborn (1976) observed both increases and decreases in creeping red fescue (Festuca rubra L.) seed yield after applying herbicides (seven data points). In contrast, Krueger- Mangold et al. (2002) observed a decrease in desired grass biomass after treating natural areas in the fall with glyphosate, a non-selective herbicide (two data points). It is important to note that other plant community components and ecosystem services may also change as a result of management efforts. For example, abundance of desired forbs may decrease as a result of broadleaf herbicide use (Ortega and Pearson 2010). Although no data points in our meta-analysis examined the response of native forbs to Canada thistle management techniques, the potential non-target effects on native forbs with broadleaf herbicides is an important concern. Incorporating herbicide use with other weed management strategies may help minimize such non-target impacts in perennial systems (Crone et al. 2009). General Management Recommendations. Our meta-analysis provides several general management recommendations for Canada thistle. First, in both annual cropping and perennial systems, land managers should consider integrating management techniques for enhanced long- term control of Canada thistle, as this approach proved to be more effective than solely applying 19 herbicides. Second, despite herbicide being the most studied management technique, a variety of other management techniques resulted in similar control of Canada thistle in the short- and long- term in annual cropping and perennial systems. This emphasizes the need to re-focus weed science research priorities by investigating alternative and integrated control methods more often. Third, herbicide MOA groups 4, 5, or 9 can be used for long-term control of Canada thistle in annual cropping systems. Finally, a variety of herbicide MOA groups can be used for long-term control of Canada thistle in perennial systems including groups 2, 4, 5, 9, and mixes of MOA groups. Future Research. First, our meta-analysis had limited data on certain non-chemical management techniques, such as burning, mulch, and water manipulation. However, some of these management techniques had negative effect sizes, suggesting these could be promising techniques for Canada thistle control. Additional management techniques, such as grazing, were not examined due to lack of papers addressing these. Therefore, increasing the amount of research devoted to non-chemical and integrated management techniques in annual cropping and perennial systems may help provide a broader range of management recommendations for Canada thistle control. Second, because short-term studies could misrepresent the impact of a management technique on a perennial species, we encourage researchers to conduct long-term evaluations on approaches to control Canada thistle. Finally, our meta-analysis, along with others (Koricheva and Gurevitch 2014; Philibert et al. 2012), highlights the need to include such basic information on means, measures of variation, and sample sizes in published articles related to invasive species management (Gurevitch and Hedges 2001; Gurevitch et al. 1992). Additionally, researchers should consider the importance of measuring not only weed control but also crop yield and abundance of desired plants, as well as other ecosystem services. 20 Acknowledgements The authors would like to thank the Montana Noxious Weed Trust Fund and the Montana Wheat and Barley Committee for funding this research. 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Can J Plant Sci 56: 331-338 22 Greenbook (2017) Plant Protection Label Data. https://www.greenbook.net. Accessed: 2016 Grekul CW, Bork EW (2004) Herbage yield losses in perennial pasture due to Canada thistle (Cirsium arvense). Weed Technol 18: 784-794 Grekul CW, Bork EW (2007) Fertilization augments Canada thistle (Cirsium arvense L. Scop) control in temperate pastures with herbicides. Crop Prot 26: 668-676 Gurevitch J, Hedges LV (2001) Meta-analysis. Pages 347-369 in Scheiner SM, Gurevitch J, eds. Design and Analysis of Ecological Experiments. New York, New York: Oxford University Press Gurevitch J, Morrow LL, Wallace A, Walsh JS (1992) A meta-analysis of competition in field experiments. Am Nat 140: 539-572 Hakansson S (2003a) Classification of plants based on traits of ecological signficance. Pages 4-13 in Hakansson S, ed. Weeds and Weed Management on Arable Land: An Ecological Approach. UK: CABI Publishing Hakansson S (2003b) Weeds with diverse life forms in various types of crops. Pages 16-55 in Hakansson S, ed. Weeds and Weed Management on Arable Land: An Ecological Approach. UK: CABI Publishing Hedges LV, Gurevitch J, Curtis PS (1999) The meta-analysis of response ratios in experimental ecology. Ecology 80: 1150-1156 Hodgson JM (1958) Canada thistle (Cirsium arvense Scop.) control with cultivation, cropping, and chemical sprays. Weeds 6: 1-11 Kettenring KM, Adams CR (2011) Lessons learned from invasive plant control experiments: a systematic review and meta-analysis. J Appl Ecol 48: 970-979 Koricheva J, Gurevitch J (2014) Uses and misuses of meta-analysis in plant ecology. J Ecol 102: 828-844 Koricheva J, Gurevitch J, Mengersen K (2013) Handbook of Meta-Analysis in Ecology and Evolution. Princeton, New Jersey: Princeton University Press. Pp. 407-419 23 Krueger-Mangold J, Sheley RL, Roos BD (2002) Maintaining plant community diversity in a waterfowl production area by controlling Canada thistle (Cirsium arvense) using glyphosate. Weed Technol 16: 457-463 Lutman PJW, Moss SR, Cook S, Welham SJ, Kim D-S (2013) A review of the effects of crop agronomy on the management of Alopecurus myosuroides. Weed Res 53: 299-313 Mesbah AO, Miller SD (2005) Canada thistle (Cirsium arvense) control in established alfalfa (Medicago sativa) grown for seed production. Weed Technol 19: 1025-1029 Miller TW (2016) Integrated strategies for management of perennial weeds. Invasive Plant Sci Manag 9: 148-158 Monaco TA, Mangold JM, Mealor BA, Mealor RD, Brown CS (2017) Downy brome control and impacts on perennial grass abundance: A systematic review spanning 64 years. Rangeland Ecol Manag 70: 396-404 Morishita DW (1999) Canada thistle. Pages 162-174 in Sheley RL, Petroff JK, eds. 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Vienna, Austria: R Foundation for Statistical Computing Rohatgi A (c2010-2017) WebPlotDigitizer. http://arohatgi.info/WebPlotDigitizer. Accessed: 2016 Shaner DL, ed. (2014) Herbicide Handbook. Tenth edn. Lawrence, KS: Weed Science Society of America. 513 p Skinner K, Smith L, Rice P (2000) Using noxious weed lists to prioritize targets for developing weed management strategies. Weed Sci 48: 640-644 Tautges NE, Goldberger JR, Burke IC (2016) A survey of weed management in organic small grains and forage systems in the northwest United States. Weed Sci 64: 513-522 Thrasher FP, Cooper CS, Hodgson JM (1963) Competition of forage species with Canada thistle, as affected by irrigation and nitrogen levels. Weeds 11: 136-138 Tiley GED (2010) Biological flora of the British Isles: Cirsium arvense (L.) scop. J Ecol 98: 938-983 [USDA] US Department of Agriculture (2017) National Agricultural Statistics Service, State Agricultural Overview https://www.nass.usda.gov/Statistics_by_State/Ag_Overview/. Accessed March 15, 2015 25 Tables Table 1. Descriptions of management techniques used in articles included in the meta-analysis of Canada thistle. The number of data points associated with each type of system is indicated. Management technique Description Annual cropping Perennial Biocontrol Biological control using insects or pathogens 24 20 Burn Prescribed fire 0 1 Competition Any method attempting to increase competitive ability including manipulating row spacing or revegetation 2 48 Crop diversification Adding cover crops or increasing crop rotation to a cropping system 17 0 Fertilizer Soil amendments including fertilizer or manure applied 2 7 Herbicide Applying herbicides 512 228 Herbicide integrated Any combination of two or more management techniques with at least one method utilizing herbicides 60 30 Mowing Mechanical mowing or clipping 12 11 Mulch Use of either plastic or organic mulches 0 3 26 Non-herbicide integrated Combination of two or more management techniques, none including herbicides 2 25 Soil disturbance Mechanical control methods including tillage, cultivation, hoeing, or harrowing 19 0 Water manipulation Changing water availability through irrigation 0 3 27 Figures Figure 1. Flow diagram depicting criteria applied during literature screening portion of the meta- analysis of Canada thistle management. In each box, “n” is the number of articles described in that step. 28 Figure 2. Mean effect size (lnR) and 95% confidence intervals for Canada thistle abundance measured (a) <1 year or (b) ≥1 year after treatment in annual cropping systems as a function of management techniques. For each management technique, the number next to the confidence interval represents the number of data points that was used to calculate the mean. 29 Figure 3. Mean effect size (lnR) and 95% confidence intervals for Canada thistle abundance measured (a) <1 year or (b) ≥1 year after treatment in annual cropping systems as a function of herbicide mechanism of action groups. For each group, the number next to the confidence interval represents the number of data points that was used to calculate the mean. Groups are as follows: 2= ALS or AHAS inhibitors; 4= synthetic auxins; 5= inhibitors of photosynthesis at photosystem II site A; 6=inhibitors of photosynthesis at photosystem II site B; 9=inhibitor of EPSPS; 11=inhibitors of carotenoid biosynthesis; 27 = inhibitors of 4-hydroxyhenyl- pyruvatedioxygenase (4-HPPD); and mix= includes two or more herbicides from different groups. 30 Figure 4. Mean effect size (lnR) and 95% confidence intervals for crop yield in annual cropping systems as a function of Canada thistle management techniques. For each management technique, the number next to the confidence interval represents the number of data points that was used to calculate the mean. 31 Figure 5. Mean effect size (lnR) and 95% confidence intervals for Canada thistle abundance measured (a) <1 year or (b) ≥1 year after treatment in perennial systems as a function of management techniques. For each management technique, the number next to the confidence interval represents the number of data points that was used to calculate the mean. 32 Figure 6. Mean effect size (lnR) and 95% confidence intervals for Canada thistle abundance measured (a) <1 year or (b) ≥1 year after treatment in perennial systems as a function of herbicide mechanism of action groups. For each group, the number next to the confidence interval represents the number of data points that was used to calculate the mean. Groups are as follows: 2= ALS or AHAS inhibitors; 4= synthetic auxins; 5= inhibitors of photosynthesis at photosystem II site A; 6=inhibitors of photosynthesis at photosystem II site B; 9=inhibitor of EPSPS; 11=inhibitors of carotenoid biosynthesis; and mix= includes two or more herbicides from different groups. 33 Appendix S1. References used in meta-analysis of Canada thistle management in annual cropping systems: Armel GR, Hall GJ, Wilson HP, Cullen N (2005) Mesotrione plus atrazine mixtures for control of Canada thistle (Cirsium arvense). Weed Sci 53: 202-211 Asadi G, Ghorbani R, Karimi J, Bagheri A, Mueller-Schaerer H (2013) Host impact and specificity of tortoise beetle (Cassida rubiginosa) on Canada thistle (Cirsium arvense) in Iran. Weed Technol 27: 405-411 Bicksler AJ, Masiunas JB (2009) Canada thistle (Cirsium arvense) suppression with buckwheat or sudangrass cover crops and mowing. Weed Technol 23: 556-563 Blasko D, Nemeth I (2006) Efficiency and long-term effects of certain herbicides against Canada thistle (Cirsium arvense (L.) scop.). J Plant Dis Prot: 739-745 Bondarenko DD (1957) 3-amino-1,2,4-triazole as an herbicide on Canada thistle [Cirsium arvense (L.) Scop.] and its effect on soil microorganisms. Ph.D dissertation. Columbus, OH: The Ohio State University. 125 p Brosten BS, Sands DC (1986) Field trials of Sclerotinia sclerotiorum to control Canada thistle (Cirsium arvense). Weed Sci 34: 377-380 Carlson SJ, Donald WW (1988) Fall-applied glyphosate for Canada thistle (Cirsium arvense) control in spring wheat (Triticum aestivum). Weed Technol 2: 445-455 Carson AG, Bandeen JD (1975) Chemical control of Canada thistle. Weed Sci 23: 116-118 Causey M, Webb F (1990) Canada thistle control in field corn. Pages 82-83 in Proceedings of the Northeastern Weed Science Society 34 Curtis RE and Haagsma T (1986) Selective control of Canada thistle in cereals with 3,6- dichloropicolinic acid (clopyralid). Pages 159-166 in Proceedings of the Western Society of Weed Science Darwent AL, Kirkland K, Baig M, Lefkovitch L (1994) Preharvest applications of glyphosate for Canada thistle (Cirsium arvense) control. Weed Technol 8: 477-482 Davies CJ, Orson JH (1987) The control of Cirsium arvense (creeping thistle) by sulfonyl urea herbicides and a comparison of methods of assessing efficacy. Pages 453-460 in Proceedings of the British Crop Protection Conference. British Crop Protection Council Derscheid LA, Nash RL, Wicks GA (1961) Thistle control with cultivation, cropping and chemicals. Weeds 9: 90-102 Donald WW (1993) Retreatment with fall-applied herbicides for Canada thistle (Cirsium arvense) control. Weed Sci 41: 434-440 Donald WW, Prato T (1992a) Effectiveness and economics of repeated sequences of herbicides for Canada thistle (Cirsium arvense) control in reduced-till spring wheat (Triticum aestivum). Can J Plant Sci 72: 599-618 Donald WW, Prato T (1992b) Efficacy and economics of herbicides for Canada thistle (Cirsium arvense) control in no-till spring wheat (Triticum aestivum). Weed Sci 40: 233-240 Farvani M, Khalghani J (2004) Synchronized weed chemical control and wheat harvesting. J Food Agric Env 2: 202-204 Glenn S, Heimer L (1994) Canada thistle (Cirsium arvense) control in no-tillage corn (Zea mays). Weed Technol 8: 134-138 Graglia E, Melander B, Jensen RK (2006) Mechanical and cultural strategies to control Cirsium arvense in organic arable cropping systems. Weed Res 46: 304-312 35 Gronwald J, Plaisance K, Ide D, Wyse D (2002) Assessment of Pseudomonas syringae pv. tagetis as a biocontrol agent for Canada thistle. Weed Sci 50: 397-404 Hodgson JM (1958) Canada thistle (Cirsium arvense Scop.) control with cultivation, cropping, and chemical sprays. Weeds 6: 1-11 Hoeft E, Jordan N, Zhang J (2001) Integrated cultural and biological control of Canada thistle in conservation tillage soybean. Weed Sci 49: 642-646 Howatt KA, Endres GJ, Hendrickson PE, Aberle EZ, Lukach JR, Jenks BM, Riveland NR, Valenti SA, Rystedt CM (2006) Evaluation of glyphosate-resistant hard red spring wheat (Triticum aestivum). Weed Technol 20: 706-716 Kirkland K (1977) Glyphosate for control of Canada thistle on summer fallow. Can J Plant Sci 57: 1015-1017 Kluth S, Kruess A, Tscharntke T (2005) Effects of two pathogens on the performance of Cirsium arvense in a successional fallow. Weed Res 45: 261-269 Kwiatkowski C (2009) The consequent influence of crop rotation and six-year-long spring barley monoculture on yields and weed infestation of white mustard and oats. Acta Agrobot 62: 241-247 Lym RG, Deibert KJ (2005) Diflufenzopyr influences leafy spurge (Euphorbia esula) and Canada thistle (Cirsium arvense) control by herbicides. Weed Technol 19: 329-341 McKay HC (1959) Control Canada thistle for greater profits. University of Idaho Rep 321. 16 p McKone MB (1989) Canada thistle Cirsium arvense (L.) Scop. control with clopyralid + 2,4-D alone and tankmixed with metsulfuron. Pages 271-273 in Proceedings of the Western Society of Weed Science 36 Miller BR, Lym RG (1998) Using the rosette technique for Canada thistle (Cirsium arvense) control in row crops. Weed Technol 12: 699-706 Miller SD (1987) Canada thistle control in barley. Western Society of Weed Science, p 270 Miller SD, Dalrymple AW, Lauer J (1989) Canada thistle and volunteer alfalfa control in barley. Western Society of Weed Science, p 259-260 Miller SD, Mesbah A, Fornstrom KJ (1994) Canada thistle control and competition in sugarbeets. J Sugar Beet Res 31: 87-96 Naish RW (1975) Dowco 290 --a new growth regulator herbicide. Pages 177-180 in Proceedings of the New Zealand Weed and Pest Control Conference. New Zealand Plant Protection Society. O'Donovan J, Blackshaw R, Harker K, McAndrew D, Clayton G (2001) Canada thistle (Cirsium arvense) management in canola (Brassica rapa) and barley (Hordeum vulgare) rotations under zero tillage. Can J Plant Sci 81: 183-190 O'Sullivan P, Kossatz V (1982) Selective control of Canada thistle in rapeseed with 3,6- dichloropicolinic acid. Can J Plant Sci 62: 989-993 O'Sullivan PA (1982) Response of various broad-leaved weeds, and tolerance of cereals, to soil and foliar applications of DPX-4189. Can J Plant Sci 62: 715-724 O'Sullivan PA, Kossatz V (1984a) Canada thistle suppression and rapeseed tolerance with dicamba and picloram. Can J Plant Sci 64: 971-978 O'Sullivan PA, Kossatz V (1984b) Control of Canada thistle and tolerance of barley to 3,6- dichloropicolinic acid. Can J Plant Sci 64: 215-217 Parochetti JV (1974) Canada thistle control with atrazine. Weed Sci 22: 28-31 37 Rabcewicz J (1995) Mechanical weed control by shallow cultivation with three vertical - axis rotary implements. J Fruit Ornam Plant Res 3: 125-142 Renner KA (1991) Canada thistle (Cirsium arvense) control in sugarbeet with clopyralid. Weed Technol 5: 392-395 Selleck G, Baird D (1981) Antagonism with glyphosate and residual herbicide combinations. Weed Sci 29: 185-190 Singh S, Malik R (1992) Evaluation of clopyralid against Cirsium arvense. Tests Agrochem Cultiv 13: 46-47 Terry HJ, Wilson CW (1964) A field study of the factors affecting the herbicidal activity of ioxynil and bromoxynil and their tolerance by cereals. Weed Res 4: 196-215 Thomsen MG, Mangerud K, Riley H, Brandsæter LO (2015) Method, timing and duration of bare fallow for the control of Cirsium arvense and other creeping perennials. Crop Prot 77: 31-37 Tolimir M, Veskovic M, Komljenovic I, Djalovic I, Stipesevic B (2006) Influences of soil tillage and fertilization on maize yield and weed infestation. Cereal Res Commun 34: 323-326 Warden RL (1964) Tordon for the control of field bindweed and Canada thistle in the North Central United States. Down Earth 20: 6-10 Warnes DD (1974) Integrated systems for control of Canada thistle in corn. Pages 95-97 in Proceedings of the North Central Weed Control Conference. North Central Weed Science Society. Wedryk S, Cardina J (2012) Evaluation of tef as a smother crop during transition to organic management. Weed Technol 26: 102-109 38 Westra P, D'Amato T (1988) Canada thistle control prior to planting winter wheat. Western Society of Weed Science, p 9-10 Zimdahl R, Foster G (1993) Canada thistle (Cirsium arvense) control with disking and herbicides. Weed Technol 7: 146-149 Zimdahl RL, Foster JM (1974) Canada thistle control. Fort Collins, CO: Colorado Agricultural Experiment Station. 4 p Zimdahl RLZ, P.S. (1979) Canada thistle control. Fort Collins, CO: Colorado Agricultural Experiment Station. 3 p Zuris N, Wilson R, and Nelson L (1987) Effects of plant growth stage on chlorsulfuron suppression of Canada thistle (Cirsium arvense) shoots and roots. Weed Technol 1: 10-13 39 Appendix S2. References used in meta-analysis of Canada thistle management in perennial systems: Almquist TL, Lym RG (2010) Effect of aminopyralid on Canada thistle (Cirsium arvense) and the native plant community in a restored tallgrass prairie. Invasive Plant Sci Manag 3: 155-168 Amor RL, Harris RV (1977) Control of Cirsium arvense (L) Scop. by herbicides and mowing. Weed Res 17: 303-309 Beck KG (1988) Canada thistle control in a non-grazed Colorado pasture. Western Society of Weed Science, p 4-5 Beck KG, Hanson DE, Sebastian JR (1989) Canada thistle control with chlorflurenol, dicamba, and clopyralid in a Colorado pasture. Western Society of Weed Science, p 27-28 Biesboer DD, Koukkari WL, Darveaux B (1994) Controlling leafy spurge and Canada thistle by competitive species. Minnesota Department of Transportation. 90 p Bixler LL, Carrithers VF, Cooley AW (1991) Canada thistle control at two stages of plant growth with clopyralid. Pages 44-47 in Proceedings of the Western Society of Weed Science Boerboom C, Wyse D (1988) Response of Canada thistle (Cirsium arvense) and birdsfoot-trefoil (Lotus corniculatus) to bentazon. Weed Sci 36: 250-253 Bourdot G, Harvey I, Hurrell G, Alexander R (1993) An experimental mycoherbicide utilizing Sclerotinia sclerotiorum controls pasture populations of Cirsium arvense in Canterbury. Pages 251-256 in Proceedings of the New Zealand Plant Protection Conference. New Zealand Plant Protection Society 40 Bourdot G, Hurrell G, Saville D, Leathwick D (2006) Impacts of applied Sclerotinia sclerotiorum on the dynamics of a Cirsium arvense population. Weed Res 46: 61-72 Bourdot GW, Hurrell GA, Saville DJ (2004) Wounding of Cirsium arvense enhances the efficacy of Sclerotinia sclerotiorum as a mycoherbicide. N Z Plant Protect 57: 292-297 Bultsma PM, Lamming F, Whitson TD (1992) Comparison of several herbicides applied at different growth stages for control of Canada thistle (Cirsium arvense) and musk thistle (Carduus nutans). Western Society of Weed Science, p I-9 Celebi SZ, Kaya I, Korhan Sahar A, Yergin R (2010) Effects of the weed density on grass yield of alfalfa (Medicago sativa L.) in different row spacing applications. Afr J Biotechnol 9: 6867-6872 Clements LJ, Salter AM, Banks CJ, Poppy GM (2012) The usability of digestate in organic farming. Water Sci Technol 66: 1864-1870 Derscheid LA, Nash RL, Wicks GA (1961) Thistle control with cultivation, cropping and chemicals. Weeds 9: 90-102 Enloe SF, Lym RG, Wilson R, Westra P, Nissen S, Beck G, Moechnig M, Peterson V, Masters RA, Halstvedt M (2007) Canada thistle (Cirsium arvense) control with aminopyralid in range, pasture, and noncrop areas. Weed Technol 21: 890-894 Foote L, Kill D, Williams C (1970) Canada thistle control on roadsides. Weed Sci 18: 307-310 Gaisler J, Pavlu V, Hejcman M (2008) Effect of different defoliation practices on weeds in an upland meadow. J Plant Dis Prot XXI: 541-546 Gallagher A, Vandenborn W (1976) Tolerance of creeping red fescue and timothy to herbicides used to control Canada thistle. Can J Plant Sci 56: 331-338 41 Gramig GG, Ganguli AC (2015) Managing Canada thistle (Cirsium arvense) in a constructed grassland with aminopyralid and prescribed fire. Invasive Plant Sci Manag 8: 243-249 Grekul CW, Bork EW (2007) Fertilization augments Canada thistle (Cirsium arvense L. Scop) control in temperate pastures with herbicides. Crop Prot 26: 668-676 Harrington TB, Peter DH, Devine WD (2014) Two-year effects of aminopyralid on an invaded meadow in the Washington Cascades. Invasive Plant Sci Manag 7: 14-24 Hartley MJ, James TK (1979) Cost benefit of selective control of Californian thistle in pasture. Pages 245-249 in Proceedings of the New Zealand Weed and Pest Control Conference. New Zealand Plant Protection Society Hartley MJ, Thomson NA (1982) Effect and control of Californian thistle in dairy pasture. Pages 104-107 in Proceedings of the New Zealand Weed and Pest Control Conference. New Zealand Plant Protection Society Harvey I, Waipara N, Bourdot G (1993) Sclerotium populations after inundative application of Sclerotinia sclerotiorum to Californian thistle. Pages 265-269 in Proceedings of the New Zealand Plant Protection Conference. New Zealand Plant Protection Society Harvey IC, Bourdot GW, Saville DJ, Sands DC (1998) A comparison of auxotrophic and wild strains of Sclerotinia sclerotiorum used as a mycoherbicide against Californian thistle (Cirsium arvense). Biocontrol Sci Technol 8: 73-81 Hodgson JM (1958) Canada thistle (Cirsium arvense Scop.) control with cultivation, cropping, and chemical sprays. Weeds 6: 1-11 Hurrell G, Bourdot G (1996) Sclerotinia sclerotiorum and mowing independently reduce Californian thistle in a sheep pasture. Pages 225-228 in Proceedings of the New Zealand Plant Protection Conference. New Zealand Plant Protection Society 42 Hurrell GA, Bourdôt GW (2001) Wounding of weeds enhances Sclerotinia sclerotiorum as a mycoherbicide. Pages 137-138 in Proceedings of the XI Internal Sclerotinia Workshop. UK: British Society for Plant Pathology Krueger-Mangold J, Sheley RL, Roos BD (2002) Maintaining plant community diversity in a waterfowl production area by controlling Canada thistle (Cirsium arvense) using glyphosate. Weed Technol 16: 457-463 McKay HC (1959) Control Canada thistle for greater profits. University of Idaho Rep 321. 16 p Meeklah FA, Mitchell RB (1984) Evaluation of herbicides for control of Californian thistle. Pages 20-23 in Proceedings of the New Zealand Weed and Pest Control Conference. New Zealand Plant Protection Society Melichar MW, Stafford MP (1989) Control of Canada thistle and musk thistle on roadside rights- of-way with clopyralid and 2,4-D. Pages 72-73 in Proceedings of the Northeastern Weed Science Society Mesbah AO, Miller SD (2005) Canada thistle (Cirsium arvense) control in established alfalfa (Medicago sativa) grown for seed production. Weed Technol 19: 1025-1029 Peterson SC, Parochetti JV (1978) Canada thistle (Cirsium arvense) control in timothy (Phleum pratense) and red-clover (Trifolium pratense) sward. Weed Sci 26: 215-220 Samuel LW, Lym RG (2008) Aminopyralid effects on Canada thistle (Cirsium arvense) and native plant species. Invasive Plant Sci Manag 1: 265-278 Sebastian JR, Beck KG, Owsley CJ (1992) Canada thistle control with metsulfuron, picloram, 2,4-D, and split applications of 2,4-D and the sulfonylureas. Western Society of Weed Science. 2 p 43 Thrasher FP, Cooper CS, Hodgson JM (1963) Competition of forage species with Canada thistle, as affected by irrigation and nitrogen levels. Weeds 11: 136-138 Tipping PW (2001) Canada thistle (Cirsium arvense) control with hexazinone in crown vetch (Coronilla varia). Weed Technol 15: 559-563 Travnicek A, Lym R, Prosser C (2005) Fall-prescribed burn and spring-applied herbicide effects on Canada thistle control and soil seedbank in a northern mixed-grass prairie. Rangeland Ecol Manag 58: 413-422 Vantoor R (1994) Effect of applying glyphosate and clopyralid by rotary weed wiper on Californian thistle in Southland. Pages 91-92 in Proceedings of the New Zealand Plant Protection Conference. New Zealand Plant Protection Society Wedryk S, Cardina J (2012) Smother crop mixtures for Canada thistle (Cirsium arvense) suppression in organic transition. Weed Sci 60: 618-623 Whitson TD, Ferrell MA (1986) Evaluation of herbicides for Canada thistle control. Western Society of Weed Science. p 55 Wilson R, Kachman S (1999) Effect of perennial grasses on Canada thistle (Cirsium arvense) control. Weed Technol 13: 83-87 Wilson RG, Martin AR, Kachman SD (2006) Seasonal changes in carbohydrates in the root of Canada thistle (Cirsium arvense) and the disruption of these changes by herbicides. Weed Technol 20: 242-248 Wilson RG, Michiels A (2003) Fall herbicide treatments affect carbohydrate content in roots of Canada thistle (Cirsium arvense) and dandelion (Taraxacum officinale). Weed Sci 51: 299-304 44 Table S1. Article information used in meta-analysis of Canada thistle management in annual cropping systems. Articles that had 1 information on crop yield are indicated by an * next to the citation. 2 Study duration Citation Study location Annual cropping system description Management technique Herbicide groups <1 year ≥1 year Armel et al. 2005 Delaware, United States corn herbicide 4, 27, mix X Asadi et al. 2013 * Mashhad, Iran winter wheat biocontrol na X Bicksler and Masiunas 2009 Illinois, United States fallow, cover crop- buckwheat, sudangrass, or sudangrass- cowpea mix crop diversification na X X Blasko and Nemeth 2006 Hungary corn herbicide 2, mix X X Bondarenko 1957 Ohio, United States fallow herbicide 11 X herbicide integrated 11 X Brosten and Sands 1986 Montana, United States fallow biocontrol na X X Carlson and Donald 1988 North Dakota, United States spring wheat herbicide 9, mix X X Carson and Bandeen 1975 Ontario, Canada fallow herbicide 4, 5 X Causey and Webb 1990 Delaware, United States corn herbicide mix X 45 Curtis and Haagsma 1986 Alberta, Canada wheat, barley herbicide 4 X X Darwent et al. 1994 Alberta, Canada barley, canola-barley-barley, lentil-barley-barley, wheat- barley-barley, field pea-wheat herbicide 9 X Davies and Orson 1987 Norfolk and Essex, England winter wheat herbicide 2, 4 X Cambridge and March, England fallow herbicide 2, 4 X Derscheid et al. 1961 South Dakota, United States oats herbicide 4 X X herbicide integrated 4 X Donald 1993 North Dakota, United States fallow herbicide 4, 9 X Donald and Prato 1992a North Dakota, United States spring wheat herbicide 4, mix X Donald and Prato 1992b * North Dakota, United States spring wheat herbicide 2, 4 X Faravani and Khlaghani 2004 Khorasan, Iran wheat herbicide 9, 2 X Glenn and Heimer 1994 * Maryland, United States corn herbicide 2, 4, 9, mix X Graglia et al. 2006 Slagelse, Denmark grass/clover, spring barley mowing na X red clover, spring barley soil disturbance na X Gronwald et al. 2002 Minnesota, United States soybean biocontrol na X 46 Hodgson 1958 * Montana, United States spring wheat fertilizer na X fallow, spring wheat, peas herbicide 4 X spring wheat, potatoes and corn herbicide integrated 4 X Hoeft et al. 2001 Minnesota, United States soybean biocontrol na X herbicide 6 X Howatt et al. 2006 North Dakota, United States spring wheat herbicide 9, mix X Kirkland 1977 * Saskatchewan, Canada fallow herbicide 9 X herbicide integrated 9 X Kluth et al. 2005 Lower Saxony, Germany fallow biocontrol na X Kwiatkowski 2009 Lublin, Poland spring barley crop diversification na X Lym and Deibert 2005 North Dakota, United States fallow herbicide 4, mix X X McKay 1959 Idaho, United States potatoes competition na X spring wheat fertilizer na X spring wheat herbicide 4 X spring wheat-alfalfa herbicide integrated 4 X McKone 1989 Montana, United States barley herbicide 2, 4 X Miller 1987 * Wyoming, United States barley herbicide 2, 4, mix X 47 Miller and Lym 1998 North Dakota, United States corn, soybean herbicide 4, mix X soil disturbance na X Miller et al. 1989 * Wyoming, United States barley herbicide 4, 5, mix X Miller et al. 1994 * Wyoming, United States sugarbeets herbicide 4 X Naish 1975 New Zealand fallow herbicide 4 X O'Donovan et al. 2001 * Alberta, Canada barley-canola herbicide 4 X O'Sullivan 1982 Alberta, Canada barley and wheat herbicide 2 X O'Sullivan and Kossatz 1982 * Alberta, Canada rapeseed herbicide 4 X O'Sullivan and Kossatz 1984a * Alberta, Canada rapeseed herbicide 4 X O'Sullivan and Kossatz 1984b * Alberta, Canada barley herbicide 4 X Parochetti 1974 Maryland, United States corn herbicide 5, 11 X Rabcewicz 1995 Alnarp, Sweden fallow soil disturbance na X Renner 1991 Michigan, United States sugarbeets herbicide 4 X Selleck and Baird 1981 Pennsylvania, United States corn and soybeans herbicide 9, mix X Singh and Malik 1992 Hisar, India fallow herbicide 4 X Terry and Wilson 1964 Essex, England winter wheat herbicide 5 X 48 Thomsen et al. 2015 * As, Norway spring wheat crop diversification na X non-herbicide integrated na X Tolimir et al. 2006 * Belgrade, Serbia corn soil disturbance na X Warden 1964 South Dakota, Minnesota, and Wisconsin United States fallow herbicide 4 X Montana, United States winter wheat herbicide 4 X Minnesota, United States oats herbicide 4 X Warnes 1974 Minnesota, United States fallow-corn herbicide 4 X X herbicide integrated 4 X X soil disturbance na X Wedryk and Cardina 2012 Ohio, United States fallow competition na X Westra and D'Amato 1988 Colorado, United States winter wheat herbicide 2, 4, 9, mix X Zimdahl and Foster 1974 Colorado, United States fallow herbicide 4, 5, 9, mix X X Zimdahl and Foster 1979 Colorado, United States fallow herbicide 4, 5, 9, mix X Zimdahl and Foster 1993 Colorado, United States fallow herbicide 2, 4, 9 X 49 Zuris et al. 1987 Nebraska, United States winter wheat-fallow herbicide 2 X X 3 4 5 6 7 8 9 10 11 Table S2. Article information used in meta-analysis of Canada thistle management in perennial systems. Articles that had information 12 on desired plant yield are indicated by an * next to the citation. 13 Study duration Citation Study location Perennial system description Management technique Herbicide groups <1 year ≥1 year Almquist and Lym 2010 Minnesota, United States range herbicide 4 X X 50 Amor and Harris 1977 Victoria, Australia pasture herbicide 4, 9, 11 X X mowing na X X Beck 1988 Colorado, United States pasture herbicide 4, mix X X Beck et al. 1989 Colorado, United States pasture herbicide 4 X X Biesboer et al. 1994 Minnesota, United States natural area competition na X X Bixler et al. 1991 Washington, United States natural area, pasture herbicide 2, 4, mix X X Boerboom and Wyse 1988 Minnesota, United States pasture herbicide 6 X Bourdot et al. 1993 Canterbury, New Zealand pasture biocontrol na X X Bourdot et al. 2004 Canterbury, New Zealand pasture biocontrol na X Bourdot et al. 2006 Canterbury, New Zealand pasture biocontrol na X Bultsma et al. 1992 Wyoming, United States range herbicide 4 X 51 Celebi et al. 2010 * Van, Turkey alfalfa competition na X Clements et al. 2012 Southampto n, England pasture fertilizer na X Derscheid et al. 1961 South Dakota, United States pasture herbicide integrated 4 X non-herbicide integrated na X Enloe et al. 2007 North Dakota, Colorado, and Wyoming, United States range, pasture herbicide 4, mix X Foote et al. 1970 Minnesota, United States roadside herbicide 4 X Gaisler et al. 2008 Liberec, Czech Republic natural area mowing na X mulch na X Gallagher and Vanden Born 1976 * Alberta, Canada grass for seed herbicide 4 X X Gramig and North Dakota, range herbicide integrated 4 X 52 Ganguli 2015 United States Grekul and Bork 2007 Alberta, Canada pasture fertilizer na X herbicide 4 X herbicide integrated 4 X mowing na X non-herbicide integrated na X Harrington et al. 2014 Washington, United States meadow herbicide 4 X Hartley and James 1979 Hamilton, New Zealand pasture herbicide integrated 4 X non-herbicide integrated na X Hartley and Thomson 1982 Normany, New Zealand pasture herbicide 4 X mowing na X Harvey et al. 1993 Canterbury, New Zealand pasture biocontrol na X Harvey et al. 1998 Canterbury, New Zealand pasture biocontrol na X X Hodgson 1958 Montana, United States pasture, alfalfa herbicide integrated 4 X 53 mowing na X Hurrell and Bourdot 1996 Canterbury, New Zealand pasture biocontrol na X mowing na X non-herbicide integrated na X Hurrell and Bourdot 2001 Canterbury, New Zealand pasture biocontrol na X Krueger- Mangold 2002 et al. * Montana, United States natural area herbicide 9 X McKay 1959 Idaho, United States alfalfa competition na X herbicide integrated 4 X Meeklah and Mitchell 1984 Otago, New Zealand pasture herbicide 4 X X herbicide integrated 4 X mowing na X Melichar and Stafford 1989 Iowa, Kentucky, and Indiana, United States roadside herbicide 4 X 54 Mesbah and Miller 2005 * Wyoming, United States alfalfa herbicide 2, 4, 6, mix X Peterson and Parochetti 1978 Maryland, United States range herbicide 4 X Samuel and Lym 2008 North Dakota, United States range herbicide 4 X X Sebastian et al. 1992 Colorado, United States range herbicide 2, mix X Thrasher et al. 1963 Montana, United States range fertilizer na X water manipulation na X non-herbicide integrated na X Tipping 2001 Maryland, United States range herbicide 5 X Travnicek et al. 2005 North Dakota, United States range burn na X herbicide 4 X herbicide integrated 4 X Vantoor 1994 Southland, New Zealand pasture herbicide 9 X 55 Wedryk and Cardina 2012 Ohio, United States pasture competition na X Whitson and Ferrell 1986 Wyoming, United States range herbicide 4 X Wilson and Kachman 1999 Nebraska, United States pasture competition na X herbicide integrated 4, 9 X non-herbicide integrated na X Wilson and Michiels 2003 Nebraska, United States range herbicide 4 X Wilson et al. 2006 Nebraska, United States pasture herbicide 4 X 14