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7/30/2019 Literature Review of Teasel (Dipsacus fullonum)
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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
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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.
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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
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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
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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
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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).
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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 long-
distance 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). Long-
distance 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
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(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
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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).
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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,
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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 (Bromustectorum 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).
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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 nativeEleocharis species in
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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 ofPhalaris 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 (Alliariapetiolata (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
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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
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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 (Krueger-
Mangold 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
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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
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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 (Centaureasolstitialis
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 (Bromustectorum 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 ofDipsacus fullonum L.
1.4.1 Nomenclature
Dipsacus fullonum L.Fuller's teasel, common teasel, teasel. Kingdom: Plantae;
Subkingdon: Tracheobionta; Superdivision: Spermatophyta; Division:Magnoliophyta;
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Class:Magnoliopsida; Subclass:Asteridae; Order:Dipsacales; Family:Dipsacaceae
(USDA-NRCS 2007).
The nameDipsacus 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 USDA-
NRCS, the binomialD. 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
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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).
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Figure 1.2. Line drawing of vegetative growth, inflorescence, flowers, and achene of
common teasel (Britton and Brown 1913).
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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 long-
distance 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
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field with the 86% germination rate had sparseA. 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).
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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)
introducedD. 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
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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
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(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 ofD. fullonum could be responsible for the quick movement rates (Nuebert
and Caswell 2000).
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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 (Cirsiumvinaceum 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 ofC. 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 Cirsiumvinaceum (Huenneke and
Thomson 1995, 423).
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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
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(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.
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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 (2-
ethylhexyl) 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
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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
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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).
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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.
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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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1.9 References
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Bosy, J.L. and R.J. Reader. 1995. Methods underlying the suppression of forb seedlingemergence by grass (Poa pratensis) litter. Functional Ecology 9:635-639.
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Minnesota, USA.
Britton, N.L., and A. Brown. 1913. Illustrated flora of the northern states and Canada.Vol. 3: 289 in USDA-NRCS PLANTS Database .
Cheeseman, O.D. 1998. The impact of some field boundary management practices on thedevelopment ofDipsacus fullonum L. flowering stems, and implications forconservation. Agriculture, Ecosystems, and Environment 68:41-49.
Clinton, W.J. 1999. Executive order 13112. Council on Environmental Quality Online.Washington, D.C, USA. .
Coblentz, B.E. 1990. Exotic organisms: A dilemma for conservation biology.Conservation Biology 4(3):261-265.
Committee on the Scientific Basis for Predicting the Invasive Potential of NonindigenousPlants and Plant Pests in the United States. 2002. Predicting invasions ofnonindigenous plants and plant pests. National Academy Press, Washington,D.C., USA.
Coombs, E.M., J.K. Clark, G.L. Piper, and A F. Cofrancesco, Jr. 2004. Biological controlof invasive plants in the United States. Oregon State University Press, Corvallis,
Oregon, USA.
Cox, G.W. 1999. Alien species in North America and Hawaii: Impacts on naturalecosystems. Island Press, Washington D.C., USA.
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Cox, G.W. 2004. Alien species and evolution: The evolutionary ecology of exotic plants,animals, microbes, and interacting native species. Island Press, Washington D.C.,USA.
Cronk, Q.C.B. and J.L. Fuller. 2001. Plant invaders: The threat to natural ecosystems.
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Czarapata, E.J. 2005. Invasive plants of the upper Midwest: An illustrated guide to theiridentification and control.The University of Wisconsin Press, Madison,Wisconsin, USA.
Dewey, S. 2006. Image 1459703 Fullers teasel,Dipsacus fullonum (Dipsacales:Dipsacaceae.
DiTamaso, J.M., G.B. Kyser, J.R. Miller, S. Garcia, R.F. Smith, G. Nader, J.M. Connor,and S.B. Orloff. 2006. Integrating prescribed burning and clopyralid for the
management of yellow starthsitle (Centaurea solstitialis). Weed Science 54:757-767.
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Glass, W.D. 1991. Vegetation management guideline: Cut-leaved Teasel (Dipsacuslacinatus L.) and (Dipsacus sylvestris Huds.). Natural Areas Journal 11(4):213-214.
Godron, M, and R.T.T. Forman. 1983. Landscape modifications and changing ecologicalcharacteristics. Pp. 12-28 in Mooney, H.A. and M. Godron, eds., Disturbance andecosystems: Components of response. Springer-Verlag, Berlin, Germany.
Gordon, D.R. 1998. Effects of invasive, non-indigenous plant species on ecosystemprocesses: Lessons from Florida. Ecological Applications 8(4):975-989.
Grace, J.B., M.D. Smith, S.L. Grace, S.L. Collins, and T.J. Stohlgren. 2001. Interactionsbetween fire and invasive plants in temperate grasslands of North America. Pp.40-65 in Galley, K.E.M. and T.P. Wilson, eds. Proceedings of the invasive speciesworkshop: the role of fire in the control and spread of invasive species. TallTimbers Research Station, Tallahassee, Florida, USA.
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Hobbs, R.J. and S.E. Humphries. 1995. An integrated approach to the ecology andmanagement of plant invasions. Conservation Biology 9(4):761-770.
Huenneke, L.F. and J.K. Thomson. 1995. Potential interference between a threatenedendemic thistle and an invasive nonnative plant. Conservation Biology 9(2):416-
425.
ISAC (Invasive Species Advisory Committee). 2006. Invasive species definitionclarification and guidance white paper.
Jurica, H.S. 1921. Development of head and flower ofDipsacus sylvestris. BotanicalGazette 71:138-145.
Kindscher, K., T. Aschenbach, and S.M. Ashworth. 2004. Wetland vegetation response tothe restoration of sheet flow at Cheyenne Bottoms, Kansas. Restoration Ecology
12(3):368-375.
Kogan, M. 1998. Integrated pet management: Historical perspectives and contemporarydevelopments. Annual Review of Entomology 43:243-270.
Kok, L.T. and W.W. Surles. 1975. Successful biocontrol of musk thistle by an introducedweevil,Rhinicyllus conicus. Environmental Entomology 4:1025-1027.
Krischik, V.A. and J. Davidson. 2004. IPM (integrated pest management) of Midwestlandscapes. University of Minnesota, Minnesota Agricultural Experiment Station,St. Paul, MN
Krueger-Mangold, J.M., R.L. Sheley, and T.J. Svejcar. 2006. Toward ecologically-basedinvasive plant management on rangeland. Weed Science 54:597-605.
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Mack, M.N., Simberloff, W.M. Lonsdale, H. Evans, H. Clout, and F.A. Bazzaz. 2000.Biotic invasions: Causes, epidemiology, global consequences, and control.Ecological Applications 10(3):689-710.
McNeely, J.A. 2000. The future of alien species: Changing social views. Pp. 171-190 inMooney, H.A. and R.J. Hobbs, eds. Invasive species in a changing world. IslandPress, Washington, D.C., USA.
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Mycogen Corporation. 2004. Scythe.
Myers, J.H. and D.R. Bazely. 2003. Ecology and control of introduced plants. Cambridge
University Press, Cambridge, United Kingdom.
Naylor, R.L. 2000. The economics of alien species invasions. Pp. 241-260 in Mooney,H.A. and R.J. Hobbs, eds. Invasive species in a changing world. Island Press,Washington, D.C., USA.
Norris, L.A. and D. Greiner. 1967. The degradation of 2,4-D in forest litter. Bulletin ofEnvironmental Contamination and Toxicology 2(2)65-74.
NRCS. 2007. Web Soil Survey. .
Nubert, M.G. and H. Caswell. 2000. Demography and dispersal: Calculation andsensitivity analysis of invasion speed for structured populations. Ecology81(6):1613-1628.
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