CHAPTER 1: GENERAL INTRODUCTION

1.1 Invasive species There are three categories of human-induced environmental disturbance: (1) adverse or inappropriate resource use; (2) pollution; and (3) the introduction of exotic organisms. While the first two categories pose a serious threat to the environment, they can be corrected by changing human behavior. On the other hand, even if exotic/invasive organisms were no longer introduced into new areas, the existing populations would remain and expand their range. It is unlikely that the introduction of exotic organisms is going to end. As globalization of the world has increased, the number of exotic plants introduced into new areas has increased dramatically (Coblentz 1990). This trend will likely continue (Cronk and Fuller 2001). Given this information, it is unlikely that the problems associated with exotic/invasive species will disappear in the near future. 1.1.1 Definitions There are multiple definitions for the term invasive species. The United States government put forth a definition in Executive Order 13112, as an alien species whose introduction does or is likely to cause economic or environmental harm or harm to human health (Clinton 1999). In 2006, the National Invasive Species Management Plan (NISMP) further clarified that definition by defining an invasive species as a species

2 that is non-native and whose introduction causes or is likely to cause economic or environmental harm or harm to human health (ISAC 2006, 1). While these definitions illustrate the potential problems with invasive species, they do not cover the mechanism behind their spread. The scientific community has offered definitions that more fully cover the ecosystemic processes and mechanisms behind the spread of invasive species. Cronk and Fuller (2001) give a definition of invasive plants as an alien plant spreading naturally (without the direct assistance of people) in natural or seminatural habitats, to produce significant change in terms of composition, structure or ecosystem processes (1). While this definition points to the problem of invasive plants spreading naturally, it gives no consideration to the scale of distribution. According to Richardson et al. (2000), the main difference between a naturalized plant and an invasive plant is the extent which the species can spread. They define invasive plants as naturalized plants that produce offspring, often in very large numbers, at considerable distances from parent plantsand thus have the potential to spread over a considerable area (Richardson et al. 2000, 98). While these definitions include the ecological problems and/or the dispersal methods of invasive plants, they do not cover the economic and social issues surrounding invasive plants as the government definitions do. For this paper the following definition for an invasive plant will be used. An invasive plant is a non-native plant, capable of spreading great distance and establishing itself naturally; that poses a serious threat to the economic, environmental, and/or social stability of an area.

3 1.1.2 Problems There are many problems associated with the spread of invasive species. The economic costs from the loss of agriculture productivity and the effort spent eradicating these species are great. According to the National Invasive Species Council (NISC), invasive species cost the United States $125 billion annually (Baker 2001). A study done by Mack et al. (2000) found that invasive species cost the nation $138 billion dollars every year. This is in line with a study done by Pimentel et al. (2000). They put the losses at $137 billion dollars per year, not including the potential monetary losses due to the degradation of ecosystem functions, possible species extinctions, and aesthetics issues. Exotic plants add up to $34 billion of that figure (Pimetel et al. 2000). An introduced blight species from China was responsible for the destruction of 1 billion American chestnut (Castanea dentata (Marsh.) Borkh.) trees in the early 1900s. This disease not only affected the aesthetics of many towns, it significantly affected the ecosystem functions of our forests (McNeely 2000). These are examples of the multiple problems associated with invasive species. There are many different ways invasive species affect the functioning of ecosystems. Competition from invasive species has been cited as the second most common factor that cause another species to become endangered (Wilcove et al. 1998). This intense competition can cause a diverse ecosystem to be replaced by a monoculture of an exotic plant. It can also cause native flora or fauna to be reduced to population levels too low to be sustainable (Cronk and Fuller 2001). Invasive species not only compete for resources, but they can also fundamentally alter the ecosystem so that native species cannot find the resources needed to remain alive. In a study on the effects of

4 invasive species with relation to the function of ecosystems in Florida, Gordon (1998) found that 39 to 64% of the 31 invasive species studied could potentially alter the geomorphology, hydrology, biogeochemistry, and disturbance regimes of their representative ecosystems. Additionally, invasive species have been found to alter the fire regime, nutrient cycling, and energy budgets of native ecosystems (Mack et al. 2000). While change in ecosystem functions is a normal process, the problem lies in the rate and direction of change that stems from invasive species introduction (Cronk and Fuller 2001).

1.2 Invasive Plant Species 1.2.1 History Invasive plant species have been both accidentally and deliberately introduced into the locations where they have become a problem. While a majority of the plants that have been introduced into North America have not caused problems, a small number of plants escaped and became invasive (White 2001). Since the 1800s, the rate of introduction has increased dramatically, causing the cumulative numbers of invasive species present to expand rapidly (Wilcove et al., 1998). Invasive plants have reached new areas in many different ways. They have found transport in the ballast of ships, as was the case with purple loosestrife (Lythrum salicaria L.) (Cox 1999). Some plants were brought to North America for erosion control. For instance, Kudzu (Pueraria montana (Lour.) Merr.) was first showcased at the 1876 United States Centennial Exposition. Soon thereafter it was in use for erosion control due to its ability to grow rapidly. The widespread planting by government agencies soon brought disaster, as it

5 began to grow uncontrollably, often covering trees and buildings (Meyers and Bazely 2003). Still other plants have made their way here through botanic gardens, the horticulture industry, and individuals (White 2001). In the United States, 235 woody plants have been introduced as ornamentals, and 82% of these are now considered invasive (Myers and Bazely 2003). The introduction of plants began on the eastern seaboard as people from other countries moved to the New World. The introduction of plants spread throughout the rest of the country as people built roads and railroads to move to new areas (Cox 2004). 1.2.2 Examples of Invasive Plant Species There many examples of invasive species that have been introduced into North America. In the 1600s Kentucky bluegrass (Poa pratensis L.), a native of Eurasia, was introduced into North America. It spreads mostly by rhizomes, but occasionally produces a large quantity of seed. It has since become an invader of prairies and montane meadows. It is still a popular turfgrass of lawn and pastures due to its tolerance of repeated mowing or grazing. This provides the plant with more pathways of introduction into new areas. Another aggressive invader of prairies, pastures, and cropland is Canada thistle (Cirsium arvense (L.) Scop.). It, like Kentucky bluegrass, is a native of Eurasia. It can out-compete native plants in these areas, often forming dense mats of vegetation. It has rapidly developed resistance to herbicides and is so prolific that eradication would be impossible (Cox 1999). Another invasive plant that was introduced to North America is the Melaleuca tree (Melaleuca quinquenervia (Cav.) Blake). It was planted in Florida by a forester in 1906 in an effort to reforest the Everglades. The seeds were even broadcast from a plane to speed up the expansion of its range. It rapidly invades swamps, dry and

6 wet flatwoods, and the transition zone between pine forests and cypress swamps (Cox 1999). It is currently on the Federal Noxious Weed List, as well as nine state noxious weed lists (USDA-NRCS 2007). 1.2.3 Ecological Processes Behind Plant Invasion Cronk and Fuller (2001) have divided the process of invasion into six steps: (1) introduction, the translocation of the plant to a new site; (2) naturalization, the establishment of the plant outside of the site of introduction; (3) facilitation, a genetic change or suitable propagule distributor that increases the likelihood of spread; (4) spread, the process of increasing habitat range; (5) interaction with animals and other plants, the method by which the plant incorporates into the new ecosystems; and (6) stabilization, the time when a species finds a new niche, whether it is as a monoculture, or as the main component of an ecosystem. The above steps are the ideal method for a plant to become established (Cronk and Fuller 2001). There are many constraints that can prevent a plant species from moving from one step to the next. For instance, an introduced plant can have reproductive issues. This would prevent the plant from naturalizing in an area. A naturalized plant could encounter a barrier in propagule dispersal, therefore it would remain only naturalized in one local area and would not be able to spread. A plant that is in the process of spreading could enter a habitat in which a predator or another aggressive plant exists and this could prevent the new plant from establishing itself in that habitat. These interactions could have the potential to stop the invasion of the plant, thus making the plant just a naturalized species, not an invasive one (Richardson et al. 2000).

7 1.2.4 Long-distance Dispersal Events Long-distance dispersal events and the effects of corridors can increase the likelihood that a plant will become invasive. Nubert and Caswell (2000) found that longdistance dispersal events, even when rare, were the major component of invasion speed. They suggest that more attention should be paid to the factors that govern long-distance dispersal when studying the invasiveness of plants (Nubert and Caswell 2000). Longdistance dispersal events are aided by corridors. There are four types of corridors: (1) line corridors; (2) strip corridors; (3) stream corridors; and (4) networks of corridors (Godron and Forman 1983). These corridors can increase the speed and range that plants invade by providing suitable habitat in which the plants can grow, reproduce, and spread. Roads are a widely used corridor for invasive plants due to our management methods of them. The frequent mowing and upkeep road right-of-ways provides open areas for invasive to establish themselves. Additionally, salt-tolerant invasive species can thrive in the conditions alongside roads due to winter maintenance (Myers and Bazely 2003). Traits of known invasive species, such as a tolerance to saline conditions, can be helpful in identifying which plant have the potential to become invasive. 1.2.5 Traits of Invasive Plants There are general traits of plants that can help determine if a plant will become invasive. Baker (1974) cited five traits of invasive plants. These are: (1) rapid initial growth rate; (2) ability to interfere with the growth rate of neighboring plants; (3) high seed output; (4) morphological similarity to native species; and (5) self-pollination and outcrossing (Baker 1974). While these traits can be found in many invasive plants, there are many studies that show that these characteristics do not always predict invasiveness

8 (Myers and Bazely 2003). It may not be possible to use general traits to predict the possibility of invasiveness across all habitat types. It might be more accurate to examine the traits of invasive species associated with particular habitat types (Cronk and Fuller 2001). Cronk and Fuller (2001) have divided the habitat types exploited by invasive species into four general categories: (1) aquatic habitats; (2) forest and open-woodland habitats; (3) fire-vulnerable habitats; and (4) open habitats. The average aquatic invasive plant can tolerate a wide variety of aquatic conditions. They can grow rapidly, mostly through an effective method of vegetative reproduction. They are free-floating or emergent herbaceous perennials. Plants that invade woodlands and forests habitats are often small shrubs or trees. They have a high seed production, with birds as their dispersal mechanism. Generally they grow rapidly and reach reproductive maturity early in their life. The typical invasive plant in fire-vulnerable habitats often promotes an increase in fire frequency. They are herbaceous plants with either high vegetative reproduction or high seed production rates. The seeds have mechanisms to survive fire and are usually light and wind dispersed. Plants that invade open habitats are generally herbaceous perennial herbs or small shrubs. They have high seed production and are wind-dispersed. They reach reproduction maturity early and can often reproduce vegetatively (Cronk and Fuller 2001). A study by Smith and Knapp (2001) found that invasive plants of tallgrass prairies do not differ from native plants with respect to resource utilization and carbon gain. This is contrary to many predictions about the characteristics of invasive plants in open habitats (Smith and Knapp 2001). Additionally, invasive plants of central grasslands of the United States have been found to prefer areas

9 rich in native plant diversity instead of areas with little to no plant diversity or cover. This is due to the favorable conditions in which native plants thrive (e.g. abundant nutrients, water, and light) are often the best conditions for invasive plants (Stohlgren et al. 2002). These unexpected results show that more long-term studies on predicting the relationship between invasives and their environment must be undertaken for prediction models to become fully developed (Committee on the Scientific Basis2002). 1.2.6 Predicting Invasions Predicting whether or not a plant will become invasive is an important aspect of managing natural resources. The Committee on the Scientific Basis for Predicting the Invasive Potential of Nonindigenous Plants and Plant Pests in the United States (2002) created a report outlining the pathways and processes behind the invasion of plants. They created four conclusions on the current state of knowledge about predicting plant invasions. The first conclusion was that the record of a plants ability to be invasive in other geographic areas is the most reliable predictor of invasiveness. The second conclusion was that there is no reliable procedure for identifying invasive plants. The third conclusion is that the inability to predict the invasive potential of plant stems from a lack of scientific knowledge. The last conclusion was that while some data on the natural history of invasive plants exist, these data need to be organized in a systematic way so that proper analysis can occur. To increase our ability to predict invasions they suggest that standardization of methods and the increasing the number of long-term studies on invasability must happen before any further steps are taken (Committee on the Scientific Basis2002).

10 1.3 Control Methods The problems that invasive species pose for the stability of the economical, social, and environmental aspects of an area dictate that control measures must be undertaken. Predicting invasions and preventing invasive species from being established is the most effective control method (Naylor 2000), but this is not always possible (Committee on the Scientific Basis 2002). Additionally, many invasive species are already established, therefore the time to prevent these species from becoming entrenched has already passed. When the control of invasive species is the only option, a plan tailored to the specific invasion must be developed (Cronk and Fuller 2001). 1.3.1 Planning for Control Planning before the implementation of control methods is a necessary measure in order to not only reduce the negative effects upon the areas under control, but to also reduce the chance that control efforts will fail (Cronk and Fuller 2001). Proper planning should include a broad idea of the scale and scope of the problem; this will reduce the possibility that funds or other resources will run out before the project is completed (Myers and Bazely 2003). Planning should begin by deciding what species is to be controlled, where it is going to be controlled, and the proper control methods for working at controlling that species (Cronk and Fuller 2001). The next decision in proper planning is to decide whether continual maintenance or complete eradication is the end goal. 1.3.2 Eradication vs. maintenance There are two strategies for the control of invasive species, eradication and maintenance (Mack et al. 2000). Eradication can be a feasible option, especially when the species is detected early and resources can be applied quickly (Simberloff 1997,

11 Hobbs and Humphries 1995). This is often possible right after an introduced species begins to naturalize, but before it begins to spread into other areas (Sakai et al. 2001). There are three key factors that contribute to the success of eradication. The first is that a proper method is used to remove the species. The second is that sufficient resources are allocated to the removal of the species. The last factor is that widespread support from both the agencies involved in the removal and the public at large is provided. Often though, detection of invasive species is hindered by the time and resource allocation it takes to effectively monitor areas. This causes invasive species to be detected when they have reached numbers too high for eradication to be feasible. At this point, maintenance control might be the only realistic option (Mack et al. 2000). 1.3.3 Control Method Options There are four options for the control of invasive species: physical, environmental management, biological, and chemical. These methods are often used in combination with each other (Cronk and Fuller 2001). For instance, cheatgrass (Bromus tectorum L.) can be controlled by using repeated herbicide treatments and intensive grazing (Whitson and Koch 1998). There are benefits and detriments associated with each method. Some invoke a negative public opinion, while some require large amounts of resources for implementation. There are also proper timings for each type of use, and to use them at the wrong time can actually hurt control efforts. In order to decide which method is the best for the situation, the pro and cons of each method must be known and weighed (Mack et al. 2000).

12 1.3.3.1 Physical Control Methods Physical control consists of pulling, cutting, digging, or mowing invasive plants. It is very labor-intensive and can be more effective when large groups are working together (Cronk and Fuller 2001). It is often employed with volunteer groups because of the lack of specialized training needed for implementation (Meyers and Bazely 2003). While physical methods are effective at controlling some species, they do not work on every species. Some species will resprout after cutting; others will send out rhizomes when mowed. Many aquatic weeds can never be completely removed and the disturbance associated with attempting physical removal can actually encourage the remaining plants to grow (Cronk and Fuller 2001). Physical methods can work well in isolated areas, but if propagules of the invasive species are somehow able to be reintroduced, this method would not be worthwhile due to the amount of resource allocation necessary for repeated applications (Meyers and Bazely 2003). Fire is another method of physical control that crosses into the category of environmental management, so it will be discussed in that later section. 1.3.3.2 Environmental Management Control Methods Environmental management control is a broad term that encompasses any changes to ecosystem properties that reduce the dominance of invasive species. This can include restoration of the hydrologic cycle, changes in nutrient availability, or alteration of the fire cycle (Cronk and Fuller 2001, Perry et al. 2004). The restoration of the hydrologic cycle in a salt-affected wetland in Kansas was shown to significantly increase the cover of some native species, as well as exotic species. Additionally, it significantly reduced the amount of bare ground and increased the amount of a native Eleocharis species in

13 playa lake communities (Kindscher et al. 2004). In a greenhouse experiment, changes in the availability of inorganic nitrogen in carbon-enriched wetland soils have been shown to reduce the competitive effect of Phalaris arundinacea L. (Perry et al. 2004). The alteration of the fire cycle can have differing effects on both native and invasive plant communities. Fire disturbance can both inhibit and promote growth of native and invasive plants (Grace et al. 2001). Burning has been shown to significantly reduce the number of garlic mustard (Alliaria petiolata (Bieb.) Cavara and Grande) plants in a woodland in Illinois (Nuzzo 1991). Additionally, invasive plants can suppress the ability of fire to burn, reducing its effects as a control method (Grace et al. 2001). All environmental management techniques require that managers have intimate knowledge of the interplay between invasive species, the native species, and their representative habitat, so that native plant communities are not harmed by the process (Cronk and Fuller 2001, Zavaleta et al. 2001). 1.3.3.3 Biological Control Methods Biological control (biocontrol) is essentially an aspect of ecological control, but because it specifically uses predator/prey relationships, it will be dealt with separately. Biocontrol consists of using the natural enemies of a species to reduce their number. For plants, the vector of control is typically insects, but mites, nematodes, fungi, bacteria, or viruses can be used as well (Coombs et al. 2004). A biocontrol project often takes many years to come to fruition due to permitting, screening for detrimental effects, and the amount of time it takes an introduced species to grow into a large enough population in order to have an effect on the target species (Myers and Bazely 2003, Coombs et al. 2004). The most critical aspect of the biological control of plants is the host-specificity

14 of the control vector. There have been instances where organisms were introduced with little regard to host-specificity. This had disastrous effects on some non-target species, but now there is extensive legislation to reduce the possibility that this will not happen again (Cronk and Fuller 2001). The classic approach to biological control is to introduce an organism into an area where it does not naturally occur and let the populations grow on their own, all the while attacking the target plant. One example of traditional biological control is the introduction of a weevil (Rhinocyllus conicus) [Coleoptera: Curculionidae] to control musk thistle (Carduus nutans L.). The weevil was introduced in Virginia between 1969 and 1972. A study on the effectiveness of this agent found that by 1975 it was attacking 90% of the thistle plants in the area, causing over 10% of the terminal flower heads to be aborted (Kok and Surles 1975). Another biocontrol approach that is not as widely used is the bioherbicidal method. In this method, biological agents are grown and applied to target species like an herbicide treatment. This method is not as cost-effective as the classical approach due to the cost of specialized equipment, but it takes less time to see results. In general, biocontrol is being used more often due to the relatively low costs to control species over a large area as compared to physical, chemical, and ecological methods, but is often overlooked due to the extensive planning associated with it (Coombs et al. 2004). 1.3.3.4 Chemical Control Methods Chemical control of invasive plants consists of using an herbicide to alter or inhibit the growth of the plant, causing the plant to die or become incapable of reproduction (Peterson et al. 2001). There are multiple ways to employ chemical control methods. Individual invasive species can be isolated and targeted for removal. Large

15 areas of mixed plant composition can be targeted to either kill every species present, or to favor some species over others (Cronk and Fuller 2001). Herbicides can also be used to cause a disturbance that shifts plant composition to a more desired state (KruegerMangold et al. 2006, Radosevich and Holt 1984). There are many potential issues associated with the use of herbicides. Targeting individual plants can be costly and time consuming; often multiple applications are necessary. Applying herbicides to large areas can cause non-target organisms to be affected, often hurting the species that are supposed to be helped (Cronk and Fuller 2001). Large-scale use of herbicides can lead to an increase in herbicidal-resistant plants that will then need to be controlled using another technique (Radosevich and Holt 1984). Additionally, if treatments do not completely eliminate the target invasive, the plant will often re-establish itself in the area (Cox 1999). If herbicides are used for plant control, an understanding of how they work is needed so that the proper herbicide is selected for use (Peterson et al. 2001). 1.3.3.4.1 Modes and Sites of Action of Herbicides Herbicides vary widely in their modes of action and their sites of action. The mode of action for herbicides is defined as all interactions between the herbicide and the target plant. This encompasses everything from application and absorption, to the physiological response of the plant to the herbicide. Herbicides can affect different processes of the target plant, but often they affect a process necessary for normal growth and development. The site of action is the specific area of a plant affected by the herbicide. Some will affect shoot growth, while others cause changes to normal hormone production. Herbicides can vary in selectivity, some affect only broadleaf plants or grasses, while others will kill any plant. Some herbicides are used before seedling

16 emergence, some after. Some herbicides are translocated within the plant to cause changes, while others cause effects by contact. Contact herbicides, as their name suggests, must come in contact with the site of action (Peterson et al. 2001). For instance, Scythe disrupts the normal plant processes in the leaves it comes in contact with, causing burndown, leaving the below ground plant parts intact (Mycogen Corporation 2004). Systemic herbicides can be applied to different plant surfaces and then are translocated to the site of action. Knowledge of where and how an herbicide works is necessary in order to select the proper herbicide and select the correct method of use (Peterson et al. 2001). 1.3.3.4.2 Herbicidal Methods of Application Herbicides can be applied to target plants in a multitude of ways. The application method chosen is dictated by the size and growth type of the species, as well as equipment availability. For low-growing herbaceous species, a foliar application is often used. This can range from a single person using a backpack sprayer in small land areas, to aerial spraying of larger areas to remove large swaths of pest plants. Foliar application can often result in non-target species being affected, due to herbicidal drift. This can be avoided by using a weed wand, a hollow pipe with a wick on the end. Woody species, especially larger specimens, require different approaches in order for control methods to be effective. Often foliar spraying will not kill woody species after a single application, making this method less efficient due to repetition of control measures. Cutting down the plants and applying herbicide to the cut stumps is one method of woody plant control. This is only effective if the herbicide is applied quickly to the living sapwood of the plant, otherwise the plant will not absorb the herbicide into the sapwood. Herbicide can

17 also be applied to living trees by ringing or notching the tree and applying the herbicide, or injecting the chemical straight into the sapwood. These methods ensure that the plant takes up the herbicide, unlike the cut-stump method (Cronk and Fuller 2001). 1.3.3.5 Integrated Control Methods Often it is necessary to use multiple methods of control in tandem to achieve the desired results. This is one aspect of Integrated Pest Management (IPM), a specific integrated control method. IPM uses a variety of cultural, biological, and chemical techniques to reduce pest problems (Krischik and Davidson 2004). This method of controlling pest species began as a way to protect plants from insects and pathogens (Kogan 1998), but it has been adopted as a method for controlling invasive plants (Britton 2004). A study done on the control of yellow starthistle (Centaurea solstitialis L.) found that burning the plant in the first year and then applying the herbicide clopyralid in the second year provided the best control (DiTomaso et al. 2006). Another study done in Wyoming found that cheatgrass (Bromus tectorum L.) can be controlled by using repeated herbicide treatments along with intensive grazing, while native grass is actually encouraged to grow (Whitson and Koch 1998). These examples illustrate the potential for integrated pest management to control invasive plants.

1.4 Description of Dipsacus fullonum L. 1.4.1 Nomenclature Dipsacus fullonum L. Fuller's teasel, common teasel, teasel. Kingdom: Plantae; Subkingdon: Tracheobionta; Superdivision: Spermatophyta; Division: Magnoliophyta;

18 Class: Magnoliopsida; Subclass: Asteridae; Order: Dipsacales; Family: Dipsacaceae (USDA-NRCS 2007). The name Dipsacus fullonum L. is the current binomial used by United States Department of Agriculture, Natural Resources Conservation Service. They also recognize three synonyms: (1) D. fullonum L. ssp. fullonum L.; (2) D. fullonum L. ssp. sylvestris (Huds.) Clapham; and (3) D. sylvestris Huds. (USDA-NRCS 2007). D. sylvestris Huds. is the most common binomial found in the literature, especially in North American publications (Werner 1975d). In order to remain consistent with the USDANRCS, the binomial D. fullonum L. will be used here. 1.4.2 Species Description Common teasel (Dipsacus fullonum L.) is a monocarpic perennial, meaning that upon reaching a certain size, it flowers once and then dies (Czarapata 2005). It is often mistaken for a biennial due to its ability to flower and die in two years. Actually, it can remain as a vegetative rosette for up to five years before flowering (Werner 1975b). It produces a low, almost prostrate vegetative rosette up to 60 cm in diameter. The rosette persists throughout winter. The leaves of the rosette are lanceolate to oblanceolate, entire or undulate, with rigid spines on the underside of the midrib and smaller spines on the surface of the leaf. It has a thick taproot that can exceed 75 cm in depth. Once a rosette diameter of 30 cm is reached and a subsequent over-wintering has occurred, the plant will form a flowering stem 0.5 to 2.5 m in height. The flowering stem is pithy or hollow with opposite branching, often forming multiple branches. The stems can persist for up to two years, long after the rosette has died. The leaves on the flowering stem are opposite, basally connate, and form a cup that catches rainwater (Dipsacus originates from a Greek

19 word meaning thirst). Each branch ends in a cylindrical flowering head (capitulum). Each capitulum is 2.5 - 10 cm long, surrounded by multiple bracts, arranged in groups of five. The bracts are spiny, upwardly curving, with the outermost set reaching above the capitulum. The receptacle bracts of the flowers are ovate to lanceolate, ending in a awn that extends beyond the flowers (Werner 1975d). The flowers resemble a white tube with four purple lobes at the end of the tube. There are four stamens that alternate with the lobes. A reduced calyx encircles the flower at the base and adheres to the inferior ovary (epigyny). The flowers are arranged in a low spiral on the capitulum, so that they appear to be arranged in diagonal rows. The flowers begin blooming in the middle of the capitulum and continue both upward and downward along the flower head, a unique trait of this genus (Jurica 1921).

Figure 1.1. Picture of common teasel rosettes (Dewey 2006).

20

Figure 1.2. Line drawing of vegetative growth, inflorescence, flowers, and achene of common teasel (Britton and Brown 1913).

21 1.4.2.1 Reproduction D. fullonum produces flowers from July to September. Each teasel capitulum can produce hundreds of flowers, each of which produces a single seed (achene). The achenes are 4 to 5 mm in length, slightly four-angled, and grayish brown in color. Since teasel plants have multiple flower heads, a single plant can produce up to 3,000 seeds (Werner 1975d). The seeds have no adaptations for wind or animal dispersal. Most fall passively from the plant, with over 99% of the seed falling within 1.5 m of the parent plant (Werner 1975a). This adaptation is especially useful because the new seedlings can take advantage of the bare ground left by the dead parents rosette (Solecki 1993, Werner 1977). Additionally, seeds have been shown to float on water, providing a method for long distance dispersal. Germination studies have shown that seeds floating in water after 16 days still had the ability to germinate. This accounts for the plants spread along waterways (Werner 1975d). Humans can also provide another mechanism for longdistance dispersal. The mowing of mature seedheads has been shown to throw the seed farther than they would have fallen naturally (Cheesman 1998). D. fullonum germinates from April to June. Teasel germination rates vary greatly between greenhouse studies and natural conditions. In one greenhouse study, fresh seed germination rates were 99.7% 0.6. Additionally, the time to 50% germination was four days for fresh seed and seven days for two-year old seed (Werner 1975d). The rates of germination in successional fields have been shown to be variable, based on the amount and type of surrounding plant matter. One field study showed that common teasel had a 28 to 86% germination rate within two years of seedling introduction. The field with 28% germination rate had a thick cover of quackgrass (Agropyron repens L.), while the

22 field with the 86% germination rate had sparse A. repens cover and large patches of bare ground (Werner 1972). This illustrates the fact that leaf litter presence affects germination. As shown from the above germination rates, A. repens plant litter can suppress germination of teasel seed greatly (Werner 1975c). Litter from other plants, such as Kentucky bluegrass (Poa pratensis L.), as well as many other forbs, has been shown to reduce germination up to 41% (Bosy and Reader 1995). Despite these modes of suppression, the high rate of germination and the high number of seeds per plant provide a recipe for invasiveness.

Figure 1.3. Picture of common teasel flowering (Alexander 2007).

23 1.4.2.2 Invasiveness and Competition D. fullonum is native to Eurasia and Northern Africa, having been brought to North America for use as a tool used in yarn production and as a dried ornamental plant in flower arrangements (Rodale 1984, Werner 1975d). It is distributed throughout much of the contiguous United States (Solecki 1993). Currently it is present in every contiguous state except Georgia, Florida, Louisiana, Maine, Minnesota, North Dakota, and South Carolina. It is listed as a noxious weed in Colorado, Missouri, Iowa, and New Mexico (USDA-NRCS 2007). Teasel is found on many different soil types, from sandy soil with abundant moisture, to heavy clay soils. It has a moderate-salinity tolerance, and can therefore thrive in roadside conditions. Both roadsides and cemeteries, where its presence is contributed to flower arrangements, act as depositories for teasel populations. From these refuges, it can invade natural areas (Solecki 1993). It has the capacity to cover prairies, sedge meadows, seeps, and savannas in the Midwestern United States, as well as waterways in the more arid climates of the Southwest United States (Solecki 1993, Glass 1991, Huenneke and Thomson 1995). As with all plants, teasel has effects on the growth and development of other plants living in close proximity; and these plants affect the growth and development of teasel. The number of established plants after germination will dictate the effects on surrounding community members. In one field experiment, Patricia Werner (1977) introduced D. fullonum into habitat types with varying amounts of grass cover, herbaceous dicot cover, and shade levels. She found that the growth rates of teasel plants in areas with moderate levels of both grasses and dicots and no shade cover were relatively quick, with plants reaching flowering stage in the 2nd and 3rd years. In areas

24 with low levels of grass, high levels of dicots, and no shade cover, teasel established quickly, but had slower growth rates, often taking up to four years to flower. In areas with dense grasses and no dicots or shade cover, and areas with low grasses, high dicot cover amounts, and high shade cover, teasel could not establish itself, with plants dying shortly after germination (Werner 1977). These levels of establishment point to a model that suggests that the growth rates of teasel populations can be reduced by shade and the success of the rest of the plant community (Werner and Caswell 1977). Werner (1976) showed that teasel can reduce the net primary production (NPP) of established herbaceous dicots, but not that of the established grasses. This can be attributed to the fact that the grasses tested had shallow roots, while the dicots and the teasel plants had deeper root systems. Once the roots of a teasel plant reach below those of the grasses, they are effectively out of competition for nutrients and water. The opposite holds true for the herbaceous dicots tested. Their roots grow to the same levels as the roots of teasel do, thus putting them in direct competition (Werner 1976). The ability of the roots to compete for water and nutrient resources is just one of the mechanisms of competition that allows teasel to thrive in certain conditions. Common teasel has many mechanisms that help it out-compete its neighbors. Its horizontally oriented leaves produce heavy shade beneath them. Its taproot extends deeper than many of the grasses around it, reaching deeper water sources. It produces a large amount of seed, up to 3,000 seeds per plant. Once a seedling has established itself, it has an increased chance of survival due to its hold on resources (Werner 1975d). Teasel colonies can completely cover an area, so much so that the leaves of the rosettes can be packed so tightly that they are forced to grow upwards instead of horizontally

25 (Werner 1977). These dense colonies can remain present in an area for an extended period of time. One population in Michigan has been present for over 25 years (Werner 1975d). The ability to remain established in a site stems from the fact that when the parent plant dies it provides an open site for germinating seedlings. If no teasel seeds are available, often short-lived winter annuals will inhabit the area, allowing the germination space to be opened again in the spring (Werner 1977). This mechanism of population growth allows colonies to expand, often forming large monocultures that exclude native vegetation (Glass 1991). 1.4.2.3 Long-distance Dispersal of Dipsacus fullonum As was illustrated in an earlier section, long-distance dispersal events can have a major effect on the invasiveness of a species. Teasel is no exception. There have been reports of birds eating seed, possibly being a vector for long-distance dispersal (Pohl and Sylwester 1963). Laboratory experiments have shown that teasel seeds can float for up to 16 days and still germinate, making water dispersal ideal for longer distances (Werner 1975d). Additionally, between 1877 and 1900 common teasel had migrated from Niagara Falls, Ontario to the east coast of North America. This corresponds to a movement of around 27 km/year (Nuebert and Caswell 2000). This rate is much faster than that calculated from population models done by Neubert and Caswell (2000). They state that while the probability of such long-distance dispersal events occurring is low, it is certainly not impossible. For instance, a seed would have to only float in a river flowing at 0.5 m/s to cover 21 km in 12 hours. They also suggest that multiple introductions of D. fullonum could be responsible for the quick movement rates (Nuebert and Caswell 2000).

26 1.4.2.4 Case Study on Invasiveness Huenneke and Thomson (1995) provide a study on the changes of teasel distribution in central New Mexico and how this is affecting a federally threatened native thistle (Cirsium vinaceum Woot. and Standl.). They examined habitat requirements of the two species by surveying existing populations in 1990 and 1993. They also looked at the outcome of competition between the two species using both greenhouse and field experiments. There was an increase in the majority of the existing teasel populations surveyed. Additionally, there was not a single case of an existing teasel population becoming extinct between the two survey times. Overall, teasel increased its representation in the thistle habitat. In the greenhouse experiments, they planted both species using a deWitt replacement series. They found that while teasel had a significant negative effect on the growth of C. vinaceum, the thistle had no effect on the growth of teasel. In the field experiments, they randomly created 0.25 m2 plots that contained a desired amount of thistle and a desired amount or more of teasel. Teasel numbers were thinned to reach the desired amounts of plants, but since the thistle is a threatened species, these plants could not be thinned. Replicate plots were established with: 100% thistle; 75% thistle, 25% teasel; 50% thistle, 50% teasel; 25% thistle, 75% teasel; and 100% teasel. Most of the tested variables had no significant effect. Teasel did show significantly lower growth rates when it was a minority, but the researchers cite low replication numbers for the inconclusiveness of the data. Despite this reduction on the growth of teasel, the authors state that there appears to be substantial potential for interference effects of teasel on the threatened Cirsium vinaceum (Huenneke and Thomson 1995, 423).

27 1.5 Control Methods of Common Teasel 1.5.1 Current Literature While there is some information available about the control of common teasel, what is available is anecdotal, with no scientific study supporting the findings. Glass (1991) uses knowledge from Illinois natural areas managers to put forth a few recommendations for control. The only specific recommendation for herbicide control is to use Roundup at 1.5% concentration during the late fall or early spring. He also states that the herbicides 2,4-D and triclopyr can be used for control, but he gives no concentration guidelines. Solecki (1993) recommends that periodic late spring burns could control isolated rosettes, though they could be unaffected if the fire does not reach a high enough temperature. Even a hot fire has no effect on large clumps of teasel because of the absence of dead plant material to burn (Glass 1991, Solecki 1993). Digging up teasel plants can be effective in areas of small infestation, but plants can grow back if not enough root is removed (Glass 1991). Werner (1975d) suggests that repeated cutting of flowering stems prior to flowering can reduce population effectively, but if the stems are not cut low enough, the plants can resprout flowering stems (Glass 1991). Additionally, seed from the seedbank, as well as imported seed from nearby plants, can cause new plants to germinate. For these reasons, stem cutting has to be closely monitored and may have to be repeated for several years for this method of control to work (Glass 1991). Mowing a teasel infested area can actually increase the amount of teasel plants by increasing potential germination sites, as well as spreading the seed farther than it would have normally reached (Cheeseman 1998). Additionally, mowing must be repeated many times due to the possibility of the flowering stems resprouting

28 (Glass 1991). While there are some recommendations for control in the literature, there has not been a quantitative study on the effects of herbicidal control of teasel. 1.5.2 Biological control of common teasel Recently researchers have been examining candidates for biological control of teasel species. Field surveys of the native range of teasel occurred with emphasis on insects or pathogens that attack the rosettes or seedheads of teasel plants. The surveys yielded 102 species of insects, 27 species of fungi, three mites, two viruses, and one nematode that are natural enemies of plants in the family Dipsacaceae (Rector et al. 2006). Preliminary experiments have shown that the leaf beetle, Galeruca pomonae (Coleoptera: Chrysomelidae), feeds on the leaf blades and the tips of the rosette. The amount of damage depends on the number of larvae per plant and the size of the rosette. Given a large enough population of insects, the beetles can cause whole mats of the rosettes to be defoliated. Additionally, choice and no-choice tests between teasel, radish, carrot, lettuce, turnip, and cabbage, showed that the beetle prefers to eat only teasel (Sforza 2004). There has also been a report that the powdery mildew, Sphaerotheca dipsacearum, has begun attacking common teasel in Washington State. This is the first reported instance of this occurring (Dugan and Glawe 2006). This could be another good prospect for biological control of teasel.

29 1.6 Herbicides and Adjuvants Tested The herbicides and adjuvants tested are listed below under their trade names. Lists of active ingredients can be found in Appendix B. 1.6.1 BK 800 BK 800 is a broadleaf specific herbicide. Its main ingredient is Isoctyl (2ethylhexyl) ester of 2,4-dichlorophenoxyacetic acid (2,4-D ester) (PBI/Gordon Corporation 2005). It mimics natural growth hormones in plants, causing the plants hormone balance to be upset (Peterson et al. 2001). Specifically, it mimics the hormone auxin. This disrupts several growth processes, causing the plant to grow uncontrollably and eventually die (Tu et al. 2001). When used on plants, they exhibit stem twisting, leaf malformations, stem callus formation, and stunted root growth (Peterson et al. 2001). BK 800 is an ester compound which is oil-soluble. Ester formulations tend to volatize readily and cause vapor drift. 2,4-D has a short half-life, about ten days in soil and less than ten days in water. There is no longer a patent on the product so it is a cheap, widely-used herbicide (Tu et al. 2001). 1.6.2 Glyphomax Glyphomax is a glyphosate-based non-selective herbicide (Dow AgroSciences 2004). It inhibits the Enolpyruvyl-shikimate-phosphate (EPSP) enzyme (Peterson et al. 2001). This enzyme is involved in the syntheses of aromatic amino acids, which are involved in protein synthesis (Tu et al. 2001). Plants affected by glyphosate will not show injury until three to five days after exposure. Symptoms include stunting, foliage discoloration, and slow death (Peterson et al. 2001). Glyphosate is generally non-toxic to mammals. It is strongly adsorbed to soil particles which can inhibit degradation, causing

30 the average half-life to be two months. It is no longer under patent, so generic brands are becoming widely available (Tu et al. 2001). 1.6.3 Transline Transline is a clopyralid-based broadleaf selective herbicide (Dow AgroSciences 2005). It, like 2,4-D, is an auxin mimic, causing the plant hormone balance to be upset (Peterson et al. 2001). For more information on the site and mode of action see section 1.6.1. It is more selective about which broadleaf species it affects. It has little effect on the mustard family (Brassicaceae). Its half-life in soil can be up to one year. It is almost completely degraded by microbes, but it does not degrade in sunlight. It does not bind to soil particles readily and might pose problems to water resources due to this mobility. It is generally non-toxic to mammals, birds, and fish (Tu et al. 2001). 1.6.4 Diesel Fuel Diesel fuel is listed as an adjuvant in the instructions of BK 800. It is listed as an acceptable adjuvant for foliar spraying, basal bark application, cut-stump application, and ring-cut stump application. Diesel fuel has been shown to not increase the degradation of 2,4-D herbicides (Norris and Greiner 1967). A study done on the plant yaupon (Ilex vomitoria Ait.), in Texas found that diesel alone killed 92% of yaupon while a 5% solution of triclopyr had a 96% kill rate (Cathey et al. 2006). 1.6.5 Nu-Film-IR Nu-Film-IR is a non-ionic surfactant created by Miller Chemical and Fertilizer Corporation. Non-ionic surfactants are the most commonly recommended herbicide adjuvant (Tu et al. 2001). It produces a sticky film that binds the herbicide to the plant surface and reduces the washing effects of rainfall. It also reduces degradation of the

31 herbicide due to ultra-violet light. It allows the herbicide to slowly invade the leaf surface by reducing leaf burn which can negatively affect translocation (Miller Chemical and Fertilizer Corporation 2001).

1.7 Site Description Cooper Farm (UTM coordinates: Zone 16; 631379E; 4454074N; elevation: 288 meters above sea level) is a field station owned by Ball State University and managed by the Field Station and Environmental Education Center staff. It consists of a 14 hectare (35 acre) wooded area and a 23 hectare (57 acre) open area undergoing various stages of succession. Esther L. Cooper and Dr. Robert H. Cooper donated the woodland to the university in 1969. The area was grazed by cattle and swine before 1951. In 1959 it was placed in United States Department of Agriculture soil reserve for five years. After donation to the university, wildlife habitat was planted by students. In 1993 trail clearing and pathway mowing in the natural area was greatly reduced to allow the habitat to undergo succession (LeBlanc 2007). The soils of Cooper Farm are Blount-Del Rey silt loams on 0 to 1 percent slopes, eroded Glynwood silt loam on 1 to 4 percent slopes, and Pewamo silty clay loam on 0 to 1 percent slopes (NRCS 2007). There are restored prairies sections, sections undergoing natural selection, forest areas, and wet swales. Due to micro-topography and the presence of drain tiles, the swales remain wet until July while other sections dry earlier in the year. Most notably, a 0.36 hectare swale runs through the southwestern corner of the property, directly south of the study site (Taylor 2004).

32 The study area is in the process of being restored to native prairie, with the wetter swale areas being restored to a wet meadow habitat type. Plantings of native species have occurred yearly since 2002. Different blocks are planted each year, causing the restored prairie to be at different stages of development. The large prairie areas have been planted with native grasses and wildflowers using a seed drill. The smaller wetter areas have been planted with plugs and hand-broadcast seeds. Each are was repeatedly sprayed with glyphosate and 2,4-D products prior to and during restoration (Taylor 2004). The area of study was treated with 1% BK800 broadleaf herbicide in August 2003 and September 2004. It was also treated with 2% glyphosate in April and May 2004. The area was then seeded with prairie grasses and a cover crop in May and June 2004. The September 2004 treatment of BK 800 damaged the native grass plantings due to their young age (Taylor 2004). By the time of plot delineation in the study area, teasel was observed to be the dominant plant. The study area had an average of 25 plants/m2, with a high density of 83 plants/m2 observed in one plot. Due to the previous years herbicidal treatment, all the plants were one to two year-old rosettes. After the study period, some flowering stalks were observed in areas not treated during the study.

33 1.8 Project Overview This research study was designed to provide baseline quantitative data on herbicidal control methods of common teasel. An area that was densely covered with first and second-year teasel rosettes were divided into 20 blocks containing 12 plots each. Three different concentrations of three widely-used herbicides were applied to a plot in each block. The adjuvants used with the herbicides were also applied to plots to see if they had any isolated effect. A Kruskal-Wallis test and post hoc Mann-Whitney U tests were used to uncover differences in herbicidal efficacy. It was expected that herbicides would reduce the number of teasel rosettes and that there would be differences in the efficacy of the herbicide concentrations tested. This study addresses three questions: (1) Do herbicide treatments have an effect on the control of common teasel? (2) What is the most efficient concentration of each herbicide tested for the control of common teasel? (3) What is the optimum herbicide treatment for controlling common teasel?

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